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NEMERTEANS OF THE PACIFIC
COAST OF NORTH AMERICA
PART I
(xi)
The present paper was originally published in the Proceedings of
the Washington Academy of Sciences, vol. in, pp. i-no, March 26,
1901. It is here reprinted from the same electrotype plates, so that
it may be quoted exactly as if it were the original. The original
pagination is the same as that of the present volume; the original
numbers of the plates are also the same as in the present volume, so
that no change has been necessary. The running plate caption has
been changed from Proceedings of the Washington Academy of Sci-
ences, vol. in, to H. A. E., vol. xi. No other alterations of any
kind have been made, save the introduction of the illustration at the
top of the first page, in place of the running heading of the Academy's
Proceedings, and the change of the title, which was : Papers from
the Harriman Alaska Expedition, xx. The Nemerteans.
The author desires to record the following corrections and additions :
Page 3. A summary of the contents of a posthumous paper by Mr. B. B. Griffin
on the Nemerteans of Puget Sound and Alaska is given on pages 114-118 of
Part II. This paper antedates Part I of the present report and necessitates
the following changes in nomenclature :
" Carinella speciosa sp. nov." should read Carinella rubra Griffin', pp. 5, 7
11-14.
"Carinella dinema sp. nov." should read Carinella sexlineata Griffin; pp. 5, 8,
15, 16, 17.
" Carinoma griffini sp. nov." should read Carinoma mutabilis Griffin; pp. 5, 8,
20-23.
" Amphiporus leuciodus sp. nov." should read Amphiporus imparispinosus Grif-
fin; pp. 6, 9, 31, 51-53, 55.
"Amphiporus exilis sp. nov." should read Amphiporus formidabilis Griffin; pp.
6. S1-.1)6-
Page 8, third line from top, after " surface " add : -while a sixth longitudinal line
is often indicated along the mid-ventral surface.
Page 34, at end of second paragraph add : Stylets are spirally futed, and appear
as if braided. See footnote, p. 144.
Page 55, at end of last paragraph add : except in Geonemertcs, -where BiJhmig
(Zeits.f. iviss. Zool., Ixiv, p. 34, i8gS] has found 8 or 9 nephridiopores above
and i or 2 bcloiv the lateral netves.
Page 82, add : Further notes on coloration and distribution of Cerebratulus albi-
frons are given in Part II, p. 200.
EDITOR.
(xii)
THE NEMERTEANS OF THE
EXPEDITION
BY WESLEY R. COE, PH.D.
YALE UNIVERSITY
CONTENTS
Introduction
Species previously recorded
Species here recognized
Keys to groups and species
Systematic discussion of genera and species .
Index .
3
5
7
ii
84
THE Harriman Alaska Expedition, during the months of
June and July, 1899, afforded exceptional advantages for the
investigation and collection of the marine invertebrate fauna
of the Alaska coast south of Bering Sea.
During the course of the expedition collections of Nemer-
teans were made at Victoria (Vancouver Island) and Lowe In-
let in British Columbia ; at New Metlakahtla (Annette Island) ,
Cape Fox, Wrangell, Farragut Bay, Taku Harbor, Juneau,
Skagway, Glacier Bay, Sitka, and Hot Springs on the islands
and mainland of southeastern Alaska ; at Russell Fiord and
Yakutat Bay; at Orca and Virgin Bay in Prince William
Sound ; at Kadiak ; at Popof Island, Shumagin group ; at
Kukak Bay on the Alaska Peninsula ; and at Dutch Harbor,
Unalaska.
2 COE
Stops were made also on St. Paul, St. Matthew, and Hall
Islands in Bering Sea ; at Plover Bay, Siberia, and at Port
Clarence, Alaska, just south of Bering Strait. At these latter
localities, however, the shores are surrounded in winter by
pack ice, which prevents the existence of any considerable
amount of life between tides. No Nemerteans were found
here between tides in the limited time at our command, though
it is not unlikely that in deeper water an abundance of these
worms could have been obtained by the dredge. South of
Bering Sea, on the other hand, the shores are densely covered
nearly to high water mark with a luxuriant growth of nearly
all classes of invertebrates.
All along that part of the coast which borders the North Pa-
cific Ocean — from Dixon Entrance to the Aleutian Islands —
with the exception of areas subject to local disturbing elements,
such as the proximity of rivers and glaciers, invertebrates oc-
cur in remarkable abundance. Not only is there an enormous
number of individuals, but also a great variety of species.
And, furthermore, the unusually large size which many of the
species attain is quite as striking as the wealth of species and
individuals. The purity of the water, its freedom from con-
tamination from muddy streams, and the great rise and fall of
the tides with their consequent swift currents, which continu-
ously provide a rapidly changing supply of pure water, furnish
conditions especially favorable to the growth of marine inverte-
brate animals.
Exceptions to these favorable conditions were found at Ju-
neau, where the water is of slightly less than normal density,
and contains a considerable amount of sediment. At Skagway
these unfavorable conditions are still more pronounced, and
here very few Nemerteans could be found between tides. In
Russell Fiord the water was decidedly brackish, there was lit-
tle tide, and almost no shore collecting could be done. Local-
ities in the vicinity of glacier fronts were extremely barren of
marine life, as was to be expected.
Perhaps the locality which proved the most fruitful was
Dutch Harbor, Unalaska, although the stations in Prince
William Sound were but little inferior, and Sitka was found to
possess an extremely rich littoral fauna.
ALASKA NEMERTEANS 3
Of the abundance of Nemerteans collected, colored drawings
were made of most of the species, and many specimens were
carefully preserved for anatomical study, and have proved most
interesting. Serial sections of most of the species have been
prepared, and details of their anatomy will be found on subse-
quent pages.
Two other members of the expedition, Prof. William E.
Ritter, of the University of California, and Prof. Trevor Kin-
caid, of the University of Washington, have generously sub-
mitted to me the Nemerteans collected by them.
SPECIES PREVIOUSLY RECORDED.
The Nemertean fauna of the northwest coast of North America
up to the present time has been almost entirely neglected.
Several small collections have been made, but the only litera-
ture relating to the group in this region seems to be a paper by
Stimpson. This author1 gives brief Latin diagnoses of the
following species from the west coast of North America :
Emplectonema virtde Stimpson [= Nemertes gracilis John-
ston = Emplectonema gracilis Verrill]. Found by Stimpson
under stones between tides in San Francisco harbor. This spe-
cies occurs abundantly along the whole Pacific coast of Alaska.
Cosmocephala beringiana Stimpson [= Amphiporus angu-
latus\ . Found in Bering Strait in five fathoms. This species
also is abundant along the whole Pacific coast of Alaska.
Cerebratulus impressus Stimpson [ = Micrura tmpressa] .
Found in twenty fathoms in Bering Strait. This species was
not met with by the Harriman Expedition.
The investigation of the systematic position and anatomy of
this group of worms has been rendered much less difficult since
the publication of Burger's excellent monograph on the Nemer-
teans of the Gulf of Naples.2 Burger's classification is found to
be admirable, and his nomenclature has been closely followed in
the present work. It will be impossible, however, to retain the
generic names Eunemertes and Eupolia which Burger adopts,
for the reason that they have been antedated by Stimpson. In
. Acad. Nat. Sci. Philadelphia, pp. 159-165, 1857.
2 Fauna und Flora des Golfes von Neapel. Monogr., 22, 1895.
4 COE
1857 Stimpson1 published brief, though accurate, diagnoses of
these same genera under the names Emplectonema and T&ni-
osoma. Moreover, Stimpson gives as the type of Emplectonema,
E. camillea [= Borlasia camillea Quatr. (1846) = Amphiporus
neesii Oersted (1844) = Eunemertes neesii Vaillant (1890)].
Not only this species, but also Stimpson's other species, Em-
plectonema niride [= Nemertes gracilis Johnston, 1837], is a
typical member of the genus named by Vaillant more than
thirty years later. With these facts in mind it is obvious, as
has been already pointed out by Verrill,2 that Stimpson's name
must be retained.
The status in regard to Eupolia Hubrecht (1887) is similar.
Stimpson named Borlasia quinquelineata Quoy et Gaimard
(1833) as belonging to his new genus Tceniosoma, and described
as new species T. septemlineatum and T. aquale, which is
probably identical with quinquelineatum Quoy et Gaimard.
But both of these species are typical of Eupolia Hubrecht.
There can be no doubt, therefore, about the identity of the two
genera3 and, as in the case of Emplectonema, Stimpson's name,
Tceniosoma, must hold.
The brevity of some of Stimpson's generic diagnoses, as men-
tioned above, is justified by the citation of well known typical
species of those genera, and in such cases their validity cannot
be questioned. Of his specific descriptions, on the other hand,
it is more than likely that some will prove insufficient for an
indisputable determination of the species.
In the preservation of the Nemerteans, I found a 2- to 5-per-
cent solution of formalin in sea-water to yield most satisfactory
results. Most forms die well-extended if a few drops of for-
malin are added to the water in the vessel in which the living
worms are contained. The formalin solution preserves the
general anatomical and some of the histological features excel-
lently, and sharp nuclear stains are easily secured, especially
with the epithelial structures. This solution, however, works
1 Proc. Acad. Nat. Sci. Philadelphia, 1857.
2 Trans. Connecticut Acad., vni, p. 413, 1892 ; ibid., ix, p. 146, 1895.
3 Burger states distinctly in his great Monograph (p. 26) that " Tceniosoma
deckt sich mit Eupolia," but, for some reason, does not consider Stimpson's
diagnoses valid.
ALASKA NEMERTEANS 5
havoc with the nervous elements and with the connective tissues.
Specimens killed in strong alcohol proved valuable adjuncts
to the formalin preparations. In most cases the formalin speci-
mens have been eventually transferred to alcohol for permanent
preservation. Corrosive sublimate-acetic, Gilson's fluid, and,
for nervous elements, Flemming's solution are always to be
recommended.
For serial sections no stains have given more happy results
than Delafield's hasmatoxylin, followed by Orange G.
In the limited time at our disposal it was naturally impossible
to make an exhaustive collection or study of the Nemertean
fauna. Some thirty-two species were collected, however, and of
these twenty-seven proved to be new to science, and only two of
the remaining species have been recorded from the Pacific.
LIST OF SPECIES HERE RECOGNIZED.
Following is a list of the species recognized, with brief notes
on their general distribution so far as our collections go :
PROTONEMERTEA.
1 . Carinella speciosa sp. nov. Along the whole Pacific coast of
Alaska. Vancouver Island (Shearer). Not uncommon.
2. C. dinema sp. nov. Victoria, B. C. ; Sitka. Uncommon.
3. C. capistrata sp. nov. Prince William Sound. Common.
MESONEMERTEA.
4. Cephalothrix linearis (Rathke) Oersted. Pacific coast. Abun-
dant.
5. Carinoma grijfini sp. nov. Collected by Mr. Creswell Shearer
at Albert Head, Vancouver Island.
METANEMERTEA.
6. Emplectonema gracile (Johnston) Verrill. Pacific coast.
Abundant. San Francisco (Stimpson).
7. E. bilrgeri sp. nov. Glacier Bay ; Sitka. Not common.
8. Zygonemertes thalassina sp. nov. Sitka. Not uncommon.
9. Z. albida sp. nov. Victoria, B. C. Uncommon.
10. Paranemertes peregrina gen. et sp. nov. Pacific coast.
Vancouver Island (Shearer). Abundant.
1 1 . P. pallida sp. nov. Yakutat Bay ; Popof Island. Uncommon.
0 COE
12. P. carnea. sp. nov. Pacific coast. Vancouver Island (Shearer).
Common.
13. Amphiporus angulatus (Fabricius) Verrill. Pacific coast.
Abundant.
14. A. bimaculatus sp. nov. Southeast coast. Uncommon.
Puget Sound (T. Kincaid).
15. A. tigrimis sp. nov. Farragut Bay. Uncommon.
16. A. nebulosus sp. nov. Kukak Bay, Alaska Peninsula. Un-
common.
17. A. leuciodtis sp. nov. Southeast coast. Common.
18. A. exilis sp. nov. Pacific coast. Abundant.
19. Tetrastemma blcolor sp. nov. Kadiak. Not common.
20. T". aberrans sp. nov. Glacier Bay ; Prince William Sound.
Uncommon.
21. T. c<ecum sp. nov. Dredged by Dr. Ritter at Kadiak. Common.
HETERONEMERTEA.
22. Tceniosoma princeps sp. nov. Southeast coast. Uncommon.
23. Lineus viridis (Fabricius) Verrill. Annette Island. Com-
mon.
24. L. torquattis sp. nov. Prince William Sound. Common.
25. Micnira verrilli sp. nov. Prince William Sound. Not un-
common.
26. M. alaskensis sp. nov. Southeast coast. Common.
27. Cerebratulus herculeus sp. nov. Sitka. Not uncommon.
28. C. marginatus Renier. Sitka. Not uncommon.
29. C. occidentalis sp. nov. Yakutat ; Prince William Sound.
Vancouver Island (Shearer). Abundant.
30. C. longiceps sp. nov. Yakutat. Uncommon.
31. C. montgomeryi sp. nov. Puget Sound to Unalaska Island.
Common.
32. C. albifrons sp. nov. Near Sitka. One specimen.
It must be remembered that the above list represents but a
few weeks' collecting, and sometimes with only a few hours
at a locality ; too much confidence therefore should not be
placed on the distribution or comparative abundance of the
various species. Further research will undoubtedly add greatly
to the number of species, and judging from the number found
in so short a time it seems not unlikely that the list may eventu-
ally be more than doubled.
ALASKA NEMERTEANS 7
General considerations in regard to the anatomical peculiar-
ities of the Nemerteans described below are reserved for a
future paper. My reason for postponing this important duty in
connection with the study of the collection is that since this
article was written, extensive collections of Nemerteans from
the west and northwest coasts of America and the Pacific Ocean
have been placed in my hands. These collections will doubt-
less include other forms than those collected on the Harriman
Expedition, and it seems desirable that as many as possible be
studied before drawing any general conclusions in regard to the
anatomical peculiarities presented by the Alaska forms.
KEYS TO GROUPS AND SPECIES.
For convenience in determination, the species at present
known from the northwest coast of America may be arranged
in the following analytical table, based mainly on superficial
and easily distinguishable characters.
A. Proboscis without stylets. Mouth posterior to brain. Intestinal
caecum absent,
a. Lateral nerves external to musculature of body walls.
PROTONEMERTEA.
aa. Lateral nerves imbedded in or between muscular layers of body
walls b.
b. Musculature of body walls in two main layers, to which an
inner circular layer is sometimes added in the esophagal
region. Cerebral sense-organs absent MESONEMERTEA.
bb. Musculature of body walls in three main layers, of which the
inner is longitudinal. Cerebral sense-organs present.
HETERONEMERTEA.
AA. Proboscis with stylets. Mouth in front of brain, usually opening
with proboscis in a single terminal or subterminal pore. In-
testinal caecum present. Lateral nerves internal to muscular
layers of body walls METANEMERTEA.
ORDER PROTONEMERTEA.
Body remarkably long, soft, and fragile. Head distinctly marked
off from body, usually broader than neck, often flattened dorso-ven-
trally Carinella, p. 1 1 .
I. Body very soft, attaining a length of upwards of 2 meters, deep red or
bright vermilion C.sfeciosa, p. n
8 COE
2. Color brown or chocolate with numerous narrow, transverse white rings
and five parallel, longitudinal white lines, of which one is in the dorso-
median line, and two on each lateral surface C. dinema, p. 15.
3. Color brown with numerous narrow transverse rings and three parallel, lon-
gitudinal white lines of which one is in the dorso-median line and one just
below each lateral margin C. capistrata, p. 16.
ORDER MESONEMERTEA.
A. Body musculature consists of a thin outer circular layer and a
strong inner longitudinal layer — an inner circular layer is
usually wanting entirely, or, if present, is much reduced. Body
long and filiform, often coiled in a close spiral. Head sharply
pointed. Mouth placed very far behind the brain. Neph-
ridia not discovered Cephalothrix, p. 19.
i. Slender, filiform, pale yellow, sometimes with reddish, grayish or greenish
tinge. No ocelli in adult C. linea ris, p. 19.
AA. Body musculature consists of a thin outer circular layer, a
diagonal layer, a longitudinal layer, and, in the esophagal re-
gion, an additional, enormously developed inner circular layer.
Head broader than neck. Nephridia well developed.
Carinoma, p. 20.
l. Body rather stout, flattened posteriorly. Mouth close behind brain. Milk-
white with grayish or brownish mottlings, and darker intestinal lobes.
C- griffini, p. 2O.
ORDER METANEMERTEA.
A. Body long and slender. Proboscis sheath usually less than half
the length of body. Proboscis small ; central stylet usually
small Emplectonema, p. 23 .
1. Stylets slender, curved. Dark green or pale green above, and whitish or
very pale greenish below E. gracile, p. 23.
2. Stylets straight and short, with swollen bases. Basis of central stylet
swollen and rounded posteriorly. Dark brown above, flesh-color be-
neath E. burgeri, p. 25.
AA. Body moderately elongated. Proboscis sheath about ^ to ^ the
length of body. Central stylet well developed.
Paranemertes, p. 32.
1. Two or 4 pouches of accessory stylets. Color purplish brown, dark-brown,
or orange-brown above, on sides, and on lateral margins of ventral sur-
face; rest of ventral surface (often only the median third) white or
yellowish white P. peregrina, p. 33.
2. Four pouches of accessory stylets. Opaque white P, pallida, p. 36.
ALASKA NEMERTEANS. 9
3. Six to 12 pouches of accessory stylets. Whitish, pinkish or flesh-
color P. carnea, p. 37.
AAA. Body commonly rather short and thick. Proboscis sheath us-
ually reaches nearly or quite to posterior end of body. Pro-
boscis large ; central stylet well developed a
a. Ocelli numerous, extending along the lateral nerves beyond the
brain. Basis of central stylet massive, with truncate or con-
cave posterior end Zygonemertes, p. 28.
1. Each of the 2 pouches of accessory stylets contains about 5 stubby sty-
lets. Olive green above and below Z. thalassina, p. 29.
2. Each pouch of accessory stylets contains 2 or 3 moderately slender sty-
lets. White Z. albida, p. 31.
aa. Ocelli do not extend posteriorly beyond the brain. Basis of
central stylet usually rounded posteriorly b.
b. Body not very small. Ocelli usually numerous.
Amphiporus, p. 40.
1. Short and broad. Usually two pouches of accessory stylets with 5 to
7 rather slender stylets each. Dark purplish or chocolate-brown
above, with a triangular white spot on each side of the head ; ventral
surface pinkish or flesh-color A.angulatus, p. 41.
2. Basis of central stylet but half as long as the slender stylet itself.
Usually 4 pouches of accessory stylets. Dorsal surface deep brown-
ish orange with 2 conspicuous oval or dark brown spots on head;
ventral surface pale orange or flesh-color A. bimaculatus, p. 44.
3. Body rounded. Color of females yellowish orange, obscured in in-
testinal region (in breeding season) by dark-green ova. Males
yellowish, with white flecks A. tigrinus, p. 46.
4. Body rather broad and flat. Whitish, thickly mottled with dark brown
blotches and dots; ventral surface whitish A. nebulosus, p. 48.
5. Body slender, rather small. Ocelli usually 15-40. Usually 3 pouches
of accessory stylets, with 2 or 3 stylets each. White.
A. leuciodus, p. 51.
6. Body extremely elongated for the genus. Ocelli usually 60-250, or
more. Usually 6 to 12 (most commonly 8) pouches of accessory
stylets, with i or 2 stylets each A.exilis, p. 54.
bb. Body very small. Ocelli few; usually 4 well-developed
ocelli arranged in a rectangle ; but sometimes these 4 eyes
are double or are fragmented into 4 groups of ocelli ; occa-
sionally ocelli are wanting Tetrastemma^. 57.
1. Ocelli 4. Brownish-red above, with median, white stripe; ventral
surface white T. bicolor, p. 57.
2. Ocelli of 4 groups, each consisting of several pigment spots of irreg-
ular size. Pale yellow T, aberrans, p. 58.
3. Ocelli wanting. Hermaphroditic. Whitish T. ccecum, p. 59.
JO COE
ORDER HETERONEMERTEA.
A. Head without deep lateral, longitudinal furrows. Proboscis mus-
culature of two layers Tceniosoma, p. 6 1 .
i. Of very large size, reaching a length of 2 meters and a diameter of 18 mm.
Deep yellow, thickly sprinkled with small irregular, dark red spots.
T. princeps, p. 62.
AA. Head with deep, horizontal furrows. Proboscis usually with
three muscular layers and muscular crossings.
a. Caudal cirrus absent. Body long and slender, rounded or flat-
tened; very contractile Lineus, p. 64.
1. A single row of 4-6 (or sometimes 8) ocelli on each side of head. Dusky,
or brownish green, dark brown, or reddish brown; commonly paler
beneath L. viridis, p. 65.
2. No ocelli in adult. Dark, reddish brown or purple, with a narrow trans-
verse white band connecting posterior ends of cephalic furrows across
dorsal surface of neck Z,. torqiiatus, p. 66.
aa. Caudal cirrus present. Body rather firm ; incapable of swim-
ming Micrura, p. 68.
1. Dorsal surface deep purple or wine-color with 15-40 sharp, pure white,
transverse lines ; tip of snout with triangular, bright orange spot ;
ventral surface of body pure white M. verrilli, p. 68.
2. Body flattened, grayish brown above with narrow, colorless, transverse
lines ; with a small, flesh-colored, sub-truncate head, much narrower
than body, and with minute gray spots near antero-lateral margins.
M. impressa,1 p. 70.
3. Salmon or flesh-color (more rarely light brownish); lighter, with tinges
of brighter red, or nearly colorless anteriorly. Intestinal lobes more
deeply colored, sometimes chestnut-brown. A cream-colored stripe is
conspicuous in the ventral median line M. alaskensis, p. 71.
aaa. Caudal cirrus present in perfect specimens, but often missing.
Body long and flat, with thin lateral margins. Swims read-
ily. Dorso-ventral and diagonal muscles well developed.
Eyes usually absent Ccrebratulus, p. 74«
1. Very large and stout, becoming 2 meters or more in length, and 25 mm.
in width. Dark, reddish brown C. herculeus, p. 75.
2. Large and ribbon-like. Slaty brown to grayish green, with thin, white,
lateral margins C. marginatus, p. 75.
3. Rather short. Chestnut-brown or reddish brown in esophagal region,
chocolate-brown in intestinal region above ; brownish flesh-color to
dark brown beneath, with a median longitudinal ochre stripe along
ventral surface C. occidentalism p. 76.
1 Known only from Stimpson's original description — Proc. Acad. Nat. Sci.,
Philadelphia, p. 160, 1857.
ALASKA NEMERTEANS II
4. Head and anterior portions of body narrow and slender, with remarkably
long and deep cephalic furrows. Dark brown or purplish, paler
on borders of cephalic slits and tip of snout C. longiceps, p. 77.
5. Very long and ribbon-like, becoming 2 meters or more in length. Deep
blood-red except tip of snout, which is yellowish white both above and
below C. Montgomery i, p. So.
6. Reddish purple except head which is white on both dorsal and ventral
surfaces back to anterior border of mouth C. albifrons, p. 82.
SYSTEMATIC DISCUSSION OF GENERA AND SPECIES.
Carinella Johnston.
Mag. Nat. Hist. London, vi, p. 232, 1833.
The species belonging to this genus are characterized by a slender,
soft, rounded body, capable of extending and contracting to a remark-
able degree. Head distinctly marked off from body, usually much
broader than parts immediately following, often flattened dorso-ven-
trally, and disk-like.
On each side of body a transverse furrow separates the head from
the esophagal region. Proboscis opening subterminal ; mouth a small
round opening on ventral surface just back of lateral transverse furrows.
The lateral nerves lie outside the muscular layers of the body, and
just beneath the basement layer of the integument. The body walls are
made up of a very thick outer epithelium with abundant glands, a
basement layer, a circular muscular layer, and a longitudinal muscular
layer in the order named from without inwards. In addition to these
there is, in the esophagal region, an inner circular muscular layer
which often forms a dorsal, and sometimes a ventral crossing with the
fibers of the outer circular muscular layer.
Proboscis sheath usually not more than ^ the length of body.
Proboscis small and short. Ocelli wanting. Cerebral sense organs
usually represented simply by a pair of sensory pits in the epithelium,
although some species (cf. C. speciosa} possess a pair of well-devel-
oped sense organs with ciliated canal, ganglion cells and glands. A
pair of peculiar lateral sense organs is usually situated well back in
the esophagal region. Some of the species show elaborate markings
of fine white lines on a brownish body, others are homogeneous in
color. Most species are colored in some shade of brown or red;
some are bright vermilion.
i. CARINELLA SPECIOSA sp. nov.
pi. in, fig. 6; pi. ix, figs. 1-3; pi. x, figs, i, 2.
Body large, very long, flattened below, rounded above, and remark-
ably soft and pliable. This beautiful Nemertean closely resembles C.
12 COE
polymorpka in external appearance, but is more brightly colored, and
shows conspicuous peculiarities in its internal anatomy. The body is
even softer than that of C. polymorpha, and more distensible ; and the
species is easily distinguished by the presence of enormously developed
cephalic glands, by having very highly specialized and sharply defined
cerebral sense organs (with ciliated canal, lined with glandular and
sensory cells), by the peculiarities in the nephridia, and by many other
details of structure.
Head rounded in front, broader than parts immediately following,
from which it is marked off by lateral, transverse furrows. Proboscis
opening and mouth as in other species of the genus. Body variable in
thickness in its different portions according to state of contraction of
the particular part, but usually flattened below and rounded above,
with well-rounded lateral faces ; posterior portion of body much more
slender than anterior.
The worms are very sluggish. When coiled together the turns of
the body are irregular and angular.
Color. — This is the most brilliantly colored and unquestionably the
most striking Nemertean encountered on the expedition. The color of
the whole body was a deep, rich vermilion, becoming somewhat paler
and more yellowish towards the posterior end. In formalin or alcohol
the natural color quickly disappears, although a characteristic marking
still remains. This feature is peculiar to C. polymorpha as well.
About 25 to 40 mm. back from the tip of the head (in a large speci-
men) is a broad band of dark brown color reaching entirely around
the body. This band is commonly 10 to 30 mm. in width, is very
sharply marked off anteriorly, but fades out gradually behind. All
the rest of the body, both in front and behind, is yellowish or grayish.
This marking remains after imbedding in paraffin, and even after
sectioning and staining in haematoxylin there is a sharp line of distinc-
tion at the anterior end of the dark band. A single section, cut
obliquely, shows the line of demarkation perfectly. The difference
lies wholly in the integument and does not affect any of the layers
beneath. In the integument the dark band is characterized not only
by the presence of an abundance of minute pigment granules, but by a
decided change in the staining qualities of the closely packed gland
cells.
Size. — The body is very long, one specimen measuring — when sus-
pended by its middle portion, and therefore well extended — fully three
meters in length, and about 5 mm. in width. Most specimens, how-
ever, were not more than ^ as long.
ALASKA NEMERTEANS 13
The epithelium of the body is very thick, and filled with closely
packed gland cells. The two circular muscular layers in the esopha-
gal region show almost no indication of a mutual crossing of fibers
either above or below, although, as indicated in pi. ix, fig. 3, a few
fibers of connective tissue and fine nerve fibers pass at intervals
from the region of the median dorsal nerve to the internal circular
muscles.
Proboscis. — Attached to tissues of head just opposite the mouth,
and therefore a considerable distance back of the brain commisures.
Its posterior attachment is in the region of the efferent nephridial ducts.
The muscular layers are as in other species of the genus. Two large
nerves enter the latero-ventral portion of the proboscis at its attach-
ment, and pass backwards on opposite sides just internal to the cir-
cular muscular layer.
The proboscis sheath shows a homogeneous basement layer beneath
its internal lining of flattened epithelium. The musculature consists
wholly of circular fibers, except that there is a distinct layer of longi-
tudinal muscles between the circular muscles of the proboscis sheath
and the epithelium of the esophagus. Lying on the muscles of the
proboscis sheath is a small median dorsal nerve which runs parallel
with the median nerve outside the circular body-muscles and corre-
sponds to the inner median nerve of other species.
The esophagus is provided with a pair of nerves continuous with
the unusually large buccal nerves. The cerebral ganglia are situated
wide apart, as in other species. The anterior portion of the head
receives a considerable number of remarkably large nerves (pi. ix,
fig. i).
Cerebral sense organs. — These organs deserve special attention
because of their high degree of development. They are each pro-
vided with a narrow but distinct canal leading outward to the lateral
surface of the integument. The sense organs lie immediately external
to the dorsal ganglia, from which they are separated only by a few
fibers of connective tissue which here represent the integumental base-
ment membrane (pi. x, fig. 2). In size, position and structure these
organs are closely similar to those of C. annulata, as figured by
Biirger.1 Each consists of a large, rounded mass of nerve cells and
connective tissue surrounding a narrow, central canal lined with
ciliated and specialized sensory epithelium (pi. ix, fig. 2). Periph-
erally there is an abundance of peculiar glandular cells, and the
whole is separated from the surrounding integumental cells by a thin
1 Fauna und Flora des Golfes von Neapel, Monogr. 22, pi. xn, fig. 5.
14 COE
sheath of connective tissue. The central canal leads ventrally and
outwards to open through the epithelium of the lateral margin of the
head. The sense organs are innervated by several large nerves from
the adjacent dorsal ganglion (pi. x, fig. 2).
Cephalic glands. — Enormously developed, making up a large por-
tion of tissues of snout (pi. ix, fig. i). These glands are closely
packed throughout the tissues internal to the integument, and form
thick layers around the cephalic blood lacunae and the rhynchodaeum.
Nearer the brain region they become more scattered, and disappear
just in front of the ganglia, those about the rhynchodseum reaching
back a little farther than those situated external to the blood lacunae.
The secretion from the glands stains most intensely with hasmatoxylin.
Most of the anterior glands pour out their secretion on the tip of the
snout ; those farther back open along the whole circumference of the
body, but most abundantly near the lateral margins (pi. ix, fig. i).
Still farther back all the glands open laterally.
Nephridia. — Situated mainly in, and a little behind, middle third of
esophagal region. There is a single large longitudinal canal on each
side, lying in close contact with the dorsal walls of the large, lateral
blood lacunae (pi. ix, fig. 3). The nephridial canals reach forward
about as far as the anterior end of the broad band mentioned above as
becoming darkly colored in alcohol. Near its posterior end each
canal exhibits a peculiar relation with the outer integument. At sev-
eral points in each canal the dorsal wall of the canal becomes discon-
tinued and the inner ends of the integumental cells reach inward into
the lumen of the canal (pi. x, fig. i). Sometimes this infolding of
the integumental cells occupies a large part of the lumen of the ne-
phridial canal. Superficially these areas always show one or more
openings in the outer layers of the integument, but such openings do
not penetrate the nephridial canal. At the posterior end of each ne-
phridial canal there is one, or sometimes two, efferent ducts (pi. ix,
fig. 3) which lead to the dorso-lateral surface of the body, as in other
species.
Habitat. — The species was found at Hot Springs (near Sitka),
Virgin Bay in Prince William Sound, and at Dutch Harbor, Unalaska.1
Individuals were occasionally seen crawling about on the bottom be-
neath the water, and in such instances were extremely conspicuous,
because of their large size and brilliant vermilion color. They were
more commonly found under stones near low-water mark, but were
nowhere abundant.
1 Collected also by Mr. Shearer at Vancouver Island, B. C.
ALASKA NEMERTEANS 15
2. CARINELLA DINEMA sp. nov.
Pl. I, figS. 2, 3.
This species somewhat closely resembles C. superba in general color
of body and the thread-like markings thereon. It differs from this
and allied species in many details, and especially in having two lateral
longitudinal white lines on each side — hence the specific name.
Body long and slender, largest in esophagal region, rounded an-
teriorly, somewhat flattened on ventral surface posteriorly. Head
broad, flat, often emarginate in front ; lateral, transverse furrows back
of head deep and conspicuous. Proboscis-pore subterminal. Mouth
rather large for the genus, elongated and situated slightly in front of
second white ring.
Color. — General color dark brown with a more or less pronounced
yellowish tone, becoming more nearly yellow posteriorly. The mark-
ings of fine longitudinal and transverse white lines on this ground
color are very characteristic and stable. Five parallel, longitudinal
lines run nearly the whole of the length of the body, and there are a
great number of transverse lines. Of the five longitudinal lines, one
lies in the middle of the dorsal surface and extends from the most an-
terior transverse ring to the posterior end of the body. The other four
lie symmetrically on the sides — two almost on the lateral margins ; the
other two much nearer the ventral surface. Seen in section, the distance
between the dorsal median line and the upper of the two lateral lines
on each side is 90°. Two lateral lines on either side are separated by
about 45°, while the remaining 90° lies between the two lower lateral
lines. All the four lateral lines extend from the second transverse line
to about y^ the distance towards the posterior end of the body. Here
they become interrupted, and back of this they are usually indicated
only by short segments and scattered dots, though their course may be
traced nearly to the end of the body. The five longitudinal lines are
all very fine and sometimes consist of rows of fine white dots closely
placed together. In addition to these five longitudinal lines some in-
dividuals show an indication of a median ventral longitudinal line in
the form of a row of isolated fine white dots extending from behind
the mouth nearly to the fifth white ring.
The first transverse white marking lies near the tip of the snout. It
is somewhat wavy in outline and does not reach below the lateral
margin. The second white marking is broader than the first, but is
likewise limited to the dorsal surface. It is indicated, however, on
the lateral surface by two narrow and inconspicuous spots. The third
16 COE
marking is not represented on the lateral surfaces, but is sharp on the
dorsal surface, and is indicated on the ventral surface by a few scat-
tered white dots. The fourth marking is very sharp and extends as a
ring completely around the body. The fifth is broader than any of the
other rings. The sixth is usually merely indicated by a series of fine
dots. Posterior to the sixth, there are commonly as many as 150 more
or less distinct and perfect rings, arranged at fairly regular intervals
throughout the length of the body. They are much more conspicuous
on the dorsal than on the ventral surface. In fact a large number of
them are interrupted on the ventral surface and represented only by
isolated dots. When the rings are well developed each consists of two
very fine white rings lying side by side, with a fine brown ring between
them. Over the greater portion of the body are alternating wider
and narrower white rings, or double and single rings, but there are
always exceptions to the regularity of arrangement. The white lines,
both longitudinal and transverse, appear as if formed by a coating of
fine white particles on the surface of the body.
The region of the proboscis-pore is very pale. Extending outward
and backward on each side from a point just above the proboscis-pore
is a shallow, horizontal groove. Each of the grooves is marked by a
black line, and each extends backward as far as the first transverse
white line. They do not quite meet in front however. The borders
of the mouth are pale in color. The sides of the brain region are dark
reddish.
In alcohol the body is grayish as far back as the fourth white ring.
Here it abruptly changes to black, which fades into dark brown through-
out the remainder of the body. The markings are well preserved.
The '•side organs' are indicated as a pair of rounded pits lying just
anterior to the fifth white ring and immediately below the dorsal of
the two white, lateral, longitudinal lines on each side.
Habitat. — This species was found on the piles of a wharf at Vic-
toria, B. C., and was also found among hydroids, etc. at Sitka. The
worms were from 300 to 500 mm. long in extension, with a diameter
of about 2 to 3 mm. They inhabited grayish, fragile, parchment-
like tubes, which were commonly much twisted and coiled. But few
specimens were met with.
3. CARINELLA CAPISTRATA sp. nov.
pi. i, fig. i.
This, like the preceding species, resembles C. superba (Kolliker)
Burger somewhat closely in color and general appearance. A careful
ALASKA NEMERTEANS 1 7
examination of a number of individuals, however, shows that the
markings on the body present such constant differences that the two
species must be considered distinct. C. capistrata is likewise differ-
ent from any of the other related and described species. It may at
once be distinguished from C. superba by lacking all indications of
a median ventral white line. The markings on the head and the ar-
rangement of the anterior transverse white lines distinguish it easily
from C. annulata and C. nothus Burger ; from C. dinema it may be
separated by the presence of but one pair of lateral white lines, in-
stead of the two pairs found in C. dinema.
C. capistrata is a very large species, attaining a length of more
than a meter with a diameter of about 5 mm. near the anterior end,
and of 2 to 3 mm. farther back. The worms are therefore extremely
long and slender, and the diameter in the esophagal region is twice as
great as it is farther back. The body is strongly rounded on the dorsal
surface throughout its whole length ; the posterior end is very slender
and easily broken.
Head broad, flattened dorso-ventrally, rounded or emarginate in
front, sharply marked off from succeeding portions by a pair of deep,
lateral, transverse constrictions, or furrows. Proboscis-pore minute,
subterminal, bounded on each side by a shallow horizontal groove.
Mouth small, slightly elongated, situated immediately behind the
lateral furrows.
Color. — General color of body rich, deep brown, sometimes vary-
ing towards grayish black ; posteriorly becoming lighter, and of a
yellowish brown. On this ground color is a series of conspicuous, but
fine, longitudinal and transverse white lines (pi. i, fig. i). The lon-
gitudinal lines are three in number and parallel — one in the middle of
the dorsal surface, the others just ventral to the lateral margins. The
dorsal line extends without interruption from near the tip of the head
to the posterior end of the body. The lateral lines, on the other hand,
are much broken up in the anterior portions of the body, and in the
esophagal region are only indicated at intervals, though their course
can be followed nearly to the head ; throughout all the rest of the
length of the body they are sharp and distinct.
The transverse white lines, excepting the most anterior one, com-
pletely encircle the body. This first transverse line appears as a sharp
V-shaped marking a little back of the head. Its angle is directed
backward and its extremities reach but little below the lateral margins.
The median dorsal line passes through the angle of this V-shaped
marking, and at the point of intersection the white area is somewhat
l8 COE
enlarged to form a small oval spot. The second transverse marking is
situated some 20 to 30 mm. (in large individuals) from the first, and
completely encircles the body. This ring is always sharp and con-
spicuous, and in alcoholic specimens is still more distinct in that it
separates a paler anterior portion from the much darker succeeding
parts. The third transverse ring is also complete, and is commonly
separated from the second by about half the distance that this is separated
from the first. The fourth and fifth rings are sometimes interrupted
below, but are more usually complete, and are much nearer together
than the second and third. Back of the fourth or fifth ring we find a
regular succession of complete rings to the end of the body. It is
common to find as many as 200 such rings arranged at fairly regular
intervals and separated from each other, when the worm is contracted,
by an average distance of about the diameter of the body. Some of
the rings are wider than others, but nearly all are as narrow as fine
threads ; a few are more or less interrupted.
The anterior and lateral margins of the head are bordered by a nar-
row band of light color, as in several other species, but this is mainly
visible from the ventral surface.
The ' side organs' are indicated by a pair of small pits situated in
the third transverse white ring and just dorsal to the position of the
lateral lines. Their position is about the same, therefore, as in C.
super ba.
In alcoholic specimens the arrangement of the white longitudinal
and circular lines is still distinguishable, except near the anterior
end of the body, which, back to the second transverse marking
(first complete ring), is pale or grayish brown. Back of this the
color abruptly changes to very dark brown or black, which reaches
posteriorly 50 to 75 mm. (in large individuals) and then gradually
fades out into light brown, which continues to the end of the body.
The color is usually paler on the ventral than on the dorsal sur-
face.
Habitat. — This species was found in abundance under stones near
low-water mark at Orca and Virgin Bay in Prince William Sound, but
was not met with elsewhere. The worms lived in long, fragile, gray-
ish, paper-like tubes of about the same diameter as the body. These
tubes were usually twisted about horizontally beneath the stones under
which the animals live. They were also frequently occupied by a
species of polychastous annelid (Nereis), which was often associated
with the Nemertean. The worms are sluggish in their movements,
and break up posteriorly if roughly handled.
ALASKA NEMERTEANS 19
Cephalothrix Oersted.
Entwurf der Plattwiirmer, Kopenhagen, 1844.
This genus includes very long, slender, filiform species which show
a tendency to coil in a close spiral. Brain situated well behind tip of
snout ; mouth several times as far posteriorly. Head sharply pointed
in extension ; proboscis-pore on ventral side a little back of its extrem-
ity. Inner circular muscular layer very much reduced, or (commonly)
entirely absent, the body musculature consisting of a thin outer circu-
lar muscular layer and a strong inner longitudinal layer ; lateral nerves
situated in the longitudinal muscular layer ; cerebral sense organs and
cephalic furrows wanting. These worms resemble in external appear-
ance some of the slender Nematodes.
4. CEPHALOTHRIX LINEARIS (Rathke) Oersted.
Planaria linearis RATHKE, Skrivter af Naturhist. Selsk. Kjobenhavn, v, p.
84, 1799.
Cephalothrix linearis OERSTED, Entwurf der Plattwiirmer , p. 82, Kopenhagen,
1844.
A very slender thread-like species capable of great extension and
contraction. Individuals may be extended till they resemble a very
fine thread, but when disturbed commonly coil themselves into a closely
wound spiral. Body commonly rather thicker in the middle and taper-
ing toward both extremities. Head very long, acutely pointed when
extended. Proboscis-pore situated ventrally, some little distance back
from tip of snout. Mouth very far back ; commonly distant from tip
of snout 10 to 12 times the diameter of body; or it is perhaps 4 to 5
times as far back as is the brain.
The length is subject to the greatest variation. Alaska specimens
commonly extended themselves 3 to 6 inches or more, but could con-
tract to a small fraction of this length.
Ocelli. — Wanting in adults although the embryos are provided with
a single pair soon after leaving the egg.
Color. — Usually pale yellow throughout, but some specimens had a
distinct reddish tinge, and some were gray, greenish, or pale green.
A median paler line, due to the proboscis sheath, appears on the dorsal
surface in the esophagal region.
Habitat. — The species is very common well up toward high-
water mark under stones in muddy places, among decaying mussels,
etc. The worms were commonly found where the mud was black,
slimy, and very foul. Scores of specimens were sometimes found
under a single stone. They were often associated with slender red-
2O COE
dish Nematodes. Very abundant at New Metlakahtla, Glacier Bay,
Sitka, Orca and at other places. The species is also common on the
coast of New England, and is likewise found along the shores of
northern Europe and in the Mediterranean.
Carinoma Oudemans.
Circulatory and Nephridial Apparatus of the Nemertea, Quart. Journ. Micr.
Sci., xxv, Suppl., pp. 1-80, 1885.
Body usually slender, often thickened and rounded anteriorly, flat-
tened in intestinal region ; head usually wider than parts immediately
following; mouth situated immediately behind the brain; proboscis
pore subterminal. Lateral slits, cephalic groves, and cerebral sense
organs wanting. Intestine with paired, lateral diverticula.
Body musculature composed of two muscular layers throughout
length of body, and of localized supplementary layers. These consist
of a thick internal longitudinal layer and a thin external circular layer,
but in the esophagal region a second circular layer lies internal to the
longitudinal muscles, and just in front of the nephridial region be-
comes enormously thickened. In the anterior portions of the esoph-
agal region a double set of distinct diagonal muscles lies just internal
to the outer circular muscular layer.
The lateral nerves are situated within the longitudinal muscular
layer.
In the anterior portions of the esophagal region are three pairs of
longitudinal blood vessels, of which one pair represents the main lat-
eral vessels and lies beside the esophagus, a second pair lies beside the
proboscis sheath, and the third pair is situated internal to the ventral
wall of the proboscis sheath and projects freely into therhynchocoel.
5. CARINOMA GRIFFINI sp. nov.
Two1 species of this interesting genus are already known from
-other parts of the world — C. armandi Oudemans, which is found
1 Miss C. B. Thompson has very recently added a third species, C. tremapkoros
(Zool. Anz., Vol. xxni, No. 631, pp. 627-630, Dec., 1900, from a single speci-
men collected at Woods Hole, Mass. I have found this species rather abundantly
in a large pond at Falmouth, Mass., connected with Vineyard Sound by a very
narrow outlet, and consequently but little affected by the tides. The species
must be unusually hardy, for the worms lived just on the edge of the pond in
sand much blackened by decaying organic matter. They have moreover to en-
dure great changes in the salinity of the water due to irregularity in rainfall and
evaporation. Further notes in regard to the anatomy of this species will be
published later, together with colored figures of the living worms.
ALASKA NEMERTEANS
21
occasionally on the shores of England, and C. patagonica Burger, of
which a single specimen has been collected from the Straits of
Magellan.
This new species was found by Mr. Creswell Shearer at Albert
Head on Vancouver Island. It attains a length of upwards of 600
mm. and a diameter of 3-5 mm. Anterior portions of body cylin-
drical ; intestinal region much flattened. Head broader than neck,
and pointed, rounded or emarginate in front according to the state of
contraction (figs, i and 2).
No ocelli were found.
* '•••:••?
£&•.'•'•' V--r^$
*>3
':'-'•. -.-' .
•ft ;•
FIG. i. FIG. 2.
FIGS, i and 2. Carinoma griffini sp. nov. Anterior part of ventral surface,
i. Head extended and pointed. 2. Head contracted and emarginate. X §•
Color. — General color milk-white anteriorly, with brownish mot-
tlings farther back ; intestinal lobes darker. After preservation the
color is completely lost, and the body becomes nearly cylindrical
throughout.
Body 'walls. — In internal organization this species agrees closely
with the descriptions which Burger gives of C. armandi1 and of C.
patagonica? but presents a number of peculiarities, which may be
stated briefly as follows : Outer integument divided into a superficial
and a deeper layer of epithelial cells, separated by a network of con-
nective tissue fibers. Basement layer in esophagal region about equal
in thickness to the integument itself, but is much thinner farther back.
Beneath basement layer in esophagal region is a loose sheet of circular
muscles, and beneath this a double set of diagonal muscular fibers.
1 Fauna u. Flora des Golfes von Neapel, Monogr. 22, Nemertinen, 1895.
*Zeits. f. wiss. Zool., Vol. LXI, pp. 19-20, pi. 3, figs. 1-9, 1896.
22 COE
The main longitudinal muscular layer in most regions of the body
equals in thickness that of all the other muscular layers combined.
The lateral nerve cords lie imbedded in this layer. Internal to the
longitudinal muscles lies a small amount of gelatinous tissue or paren-
chyma of the body cavity. The inner circular muscular layer extends
from the mouth to the posterior end of the esophagal region. Its
fibers are continuous with those of the proboscis sheath. Anteriorly,
it is even thinner than the outer circular muscular layer, but towards
the posterior end of the esophagal region it increases so greatly in
massiveness that for a short distance it exceeds in thickness the other
muscular layers combined. Having reached its maximum develop-
ment (near the efferent nephridial ducts), it suddenly disappears en-
tirely, its dorsal portion remaining for a few sections as a semicircular
arch above the proboscis sheath.
Proboscis sheath. — This organ is more strongly developed than in
the other species of the genus, and possesses a single muscular layer
composed of circular fibers interlaced with longitudinal fibers in small
groups.
Proboscis. — The anterior end of the proboscis is attached to the tis-
tues of the head at about the middle of the brain region. It walls con-
sist of a thin outer fibrous layer, on which rest the superficial flattened
epithelial cells bathed in the fluid of the rhynchocoel. Beneath is the
thick layer of longitudinal muscles comprising nearly the whole of the
musculature. Internal to these are a few scattered circular fibers, then
a thin basement membrane, and, finally, the internal layer of columnar
epithelial cells. This inner epithelium is composed of a simple layer
of very long and closely pressed cells, a portion of which contain rod-
like masses of secretion. Their nuclei are closely packed together
peripherally, and comprise several layers as is usual in much crowded
columnar cells.
A pair of rather large nerves extend throughout the length of the
proboscis just internal to the circular muscular layer. These nerves
originate from the ventral commissure of the brain very much as in
Carinella. They arise from the anterior border of the commissure
near its origin from the ganglia, and pass dorsally to the point where
the proboscis is attached to the tissues of the head. They then enter
the proboscis, and take up lateral positions in its walls corresponding to
those of the lateral nerves in the walls of the body.
Nephridia. — The condition of the nephridial system is in many
respects intermediate between that of C. patagonica and C. armandi,
the mass of tubules constituting the so-called nephridial glands not ex-
ALASKA NEMERTEANS 23
tending into the cavity of the blood space nearly so far as in the former,
but are much more profusely branched than in the latter species. The
single pair of efferent ducts lies in the same region as the posterior
ends of the series of ' nephridial glands.' Posteriorly to this point a
single unbranched duct of large size runs backward on each side for
a considerable distance, and then bends sharply and runs forward to
the efferent duct, always lying close beside, and just external to that
limb of the duct which is passing backward. The efferent ducts open
on the dorso-lateral surfaces of the body at the posterior end of the
enormously thickened internal circular muscular layer.
The blood vascular system, brain, buccal nerves, lateral nerves, and
dorsal and ventral median nerves are much as in C. patagonica.
Specimens collected in August had just discharged their sexual
products. In one individual a very few genital sacks still retained
their mature ova.
Emplectonema Stimpson.
Emplectonema STIMPSON, Proc. Philadelphia Acad., p. 163, 1857.
Nemertes MclNTOSH, British Annelids, Part I, Nemerteans, Ray Society,
1872-1873.
Eunemertes VAILLANT, Hist. Nat. des Anneles, Tome 3, Paris, 1890.
Body very long and slender, varying greatly in thickness according
to state of contraction, but most commonly considerably flattened ;
often sharply bent and folded into an irregular mass ; integument pro-
vided with an unusual abundance of mucous secretion ; proboscis and
mouth opening together on subterminal portion of snout ; proboscis
sheath limited to anterior third of body ; proboscis slender and very
short, often not more than ^ the length of body ; a large number of
minute eyes usually present ; cerebral sense organs situated well in front
of brain, and usually very small. The species are of sluggish move-
ment, and many of them are found very near high water mark among
mussels, barnacles, rockweeds, etc.
Of the genus Emplectonema two species, of which only one (1?. gra-
cz7e) had previously been described, were found on the Harriman ex-
pedition.
6. EMPLECTONEMA GRACILE (Johnston) Verrill.
pi. vin, fig. 3.
Nemertes gracilis JOHNSTON, Mag. Zool. and Bot. London, vol. i, 1837-38.
Emplectonema viride STIMPSON, Proc. Philadelphia Acad., p. 163, 1857.
Eunemertes gracilis VAILLANT, Hist. Nat. des Anneles, Tome 3. Paris, 1890.
Eunemertes gracilis BURGER, Fauna u. Flora des Golfes von Neapel,
Monogr. 22, Nemertinen, p. 543, 1895.
Emplectonema gracilis VERRILL, Trans. Connecticut Acad., vni, p. 413, 1892;
ix, p. 146, 1895.
24
COE
Body very long and slender, somewhat flattened below, irregular and
ungraceful in form, and sluggish in movement. Head slightly broader
than rest of body ; snout blunt, posterior extremity of body tapering
gradually to a point.
The common opening of the mouth and proboscis lies ventrally, a
little back from the tip of the snout. The cerebral sense organs lie
far in front of the brain, and each communicates with the exterior by
a canal which opens on the ventro-lateral margin near the tip of the
snout.
Color. — Uniform dark green above and nearly white below ; some-
times grayish or yellowish green above, and very pale yellowish green
below. The color of Alaska specimens is as a rule paler and more
grayish green than in Mediterranean specimens. The head is bordered
with a narrow band of white, uniform with that of the lower surface.
Back of the head is an inconspicuous transverse band paler than the
rest of the dorsal surface. The pigment resides wholly in the integu-
ment, the muscular layers and other organs of the body being prac-
tically colorless.
Ocelli. — The eyes (fig. 3) are characteristically arranged in two
groups on each side of the head. The anterior group on each side
usually consists of 8 to 10 pigment-cups ar-
ranged in a single row near the antero-lateral
border of the head. These lie deeply imbedded
in the tissues of the head, and are much more
conspicuous from the ventral surface owing to
the presence of much pigment on the dorsal
surface above them in ordinary states of con-
traction. Each posterior group lies nearly
above the brain and consists of 10 to 20 ocelli
in an irregular cluster. The eyes of the pos-
terior groups are visible only from the dorsal
surface, and are smaller in size than those
of the anterior groups. The integument may be scraped off so as to
expose the eyes clearly.
Proboscis. — The stylet apparatus is especially characteristic of the
species. The basis of the central stylet is very long and slender, and
is twice to three times as long as the stylet itself. The posterior
end is sharply swollen into a flattened knob (pi. vin, fig. 3). In
front of the knob the basis decreases in diameter evenly towards the
anterior end, except for a slight constriction which occurs at about y$
the distance from the posterior end. The anterior portion is slightly
FIG. 3. Emplectonema
gracile. Dorsal view of
head to show arrange-
ment of ocelli. X 12-
ALASKA NEMERTEANS 25
curved. The central stylet is slender, extremely sharply pointed, and
gracefully curved like a sabre or scythe. There are two accessory
stylet pouches, and each commonly contains 5 to 7 slender stylets
curved like the central one, and of approximately the same size (pi.
vin, fig. 3).
Habitat. — This species was found in the greatest abundance at nearly
all the collecting stations between Victoria, B. C., and Dutch Harbor,
Unalaska. It occurred everywhere along the shore, and was most
plentiful near high water mark, crawling over the thick growth of
mussels and seaweeds. Often a number of individuals were found
coiled together in a single slimy mass, and on being disturbed would
crawl apart and move sluggishly about, but usually made no effort at
concealment. Similar masses were met with under stones in very
muddy localities, and often where the water was very brackish. This
is probably the most abundant species of Nemertean on the Alaska
coast, and is found nearer high water mark and in more brackish water
than almost any other species.
The species has previously been recorded from the coasts of Eng-
land, the northern shores of Germany and France, the Mediterranean,
and Madeira. It was also found by Stimpson under stones between
tides in San Francisco harbor and described as E. -viride.
An excellent and detailed account of the minute anatomy of this
species is given in Burger's Monograph of the Nemerteans of the Gulf
of Naples.1
Specimens collected in Alaska in June and July were filled with
nearly mature genital products, in closely packed, but irregularly
arranged pouches. The genital glands were often noticeable from the
exterior because of their pale color.
7. EMPLECTONEMA BCRGERI sp. nov.
Pl. ii, figs. 1,2; Pi. vni, fig. i ; Pi. xn, fig. 3.
Body long, flattened both above and below, ribbon-like, as thick
near the edges as in the median line ; head usually narrower than the
parts immediately following ; posterior extremity slender. The rhyn-
choda3um opens on the ventral side of the tip of the snout ; a pair of
shallow lateral slits occur just in front of the mouth.
The worms attain a length of more than a meter, and are 5 mm.
or more in width. Like other species of the genus the integument is
furnished with a vast amount of mucus.
1 Fauna u. Flora des Golfes von Neapel. Monogr. 22, 1895.
26
COE
Color. — The color varies considerably, as will be seen from the two
color varieties on pi. n, figs, i and 2. The commonest form is
dark velvety-brown above, with a paler median line, much paler and
slightly yellowish in front, and flesh-colored or creamy white below.
Other individuals have the whole dorsal and lateral surfaces of a mot-
tled reddish brown color, with a tinge of purple, while the ventral
surface is pale yellow or flesh-colored. Sometimes the head is nearly
colorless. The dorsal surface almost always has an appearance sug-
gestive of velvet. This color is superficial, and is easily removed by
rough handling.
Proboscis. — Proboscis small, very short and delicate, sometimes
scarcely more than J^- the length of body. The proboscis sheath
reaches well toward middle of body, but is very small toward its pos-
terior end ; in esophagal region it is well developed, with a thick outer
layer of circular muscles, and a thin, inner, longitudinal muscular
layer. The rhynchodffium passes backward a considerable distance
before the intestinal canal is separated from the proboscis opening.
The armature of the proboscis consists of a weak central stylet, and
a pair of pouches of accessory stylets (pi. vui, fig. i). The basis of
the central stylet is rather slender in front, slightly contracted near its
posterior third, from which point it swells out
suddenly into a large spherical bulb. The cen-
tral stylet is slightly shorter than its basis. Each
of the reserve pouches usually contains three
small stylets with swollen bases. The posterior
chamber of the proboscis is narrow. The pro-
boscis is provided with n distinct nerves.
Ocelli. — The ocelli are very numerous. On
each side of the tip of the snout is an elongated
cluster of minute eyes, all situated near the dor-
sal surface, and without regularity of arrange-
ment. Their number is commonly 60 or more
on each side (fig. 4). Back of these are a few
other eyes, likewise very minute, situated deep in the tissues of the
body, and seen only with difficulty. They are scattered irregularly
from near the lateral borders in front of the brain inward towards the
median line (fig. 4).
Cerebral sense organs. — Unusually large. They lie lateral to the
rhynchodaeum, and slightly in front of the brain. The ducts by which
they communicate with the exterior pass forward to open latero-ven-
trally.
FIG. 4. Emplecto-
nema biirgeri. Dor-
sal view of head to
show arrangement of
ocelli. X 8-
ALASKA NEMERTEANS 27
The pair of blood lacuna in the head are rather small, and pass
backward in numerous branches. The dorsal vessel in the proboscis
sheath has several communications with the lateral vessels in the
esophagal region.
In the region of the brain a great abundance of sub-muscular glands
appears, occupying the whole lateral aspects of the body and reaching
far backward. As usual, they are multicellular. Their ducts pierce
the muscles and integument of the body wall to open on the sides of
the body in enormous numbers. Farther back they are restricted to a
narrow region in each section just lateral to the nerve cords, and open
somewhat ventrally from the lateral edge. They continue in dimin-
ished numbers, but of large size, backward beyond the point where
the esophagus opens into the intestine (pi. xn, fig. 3).
Alimentary canal. — A pair of remarkably narrow intestinal caeca
reach forward well toward the brain. Their diameter for a long dis-
tance back is insignificant compared with that of the esophagus.
They occupy positions ventro-laterally to the proboscis sheath and
above the esophagus. Occasional pouches are sent off laterally into
the tissues above the nerve cords. Farther back they become larger
and extend laterally beyond the ventrally placed nerve cords. Towards
the middle of the esophagal region the pouches become paired with
considerable regularity. A well developed network of muscular fibers
and connective tissue reaches between the pouches from the muscular
layer above to that below the alimentary canal. The pouches lie close
together still farther back, and near the posterior end of the esophagal
region extend on each side below the esophagus and open together
from opposite sides. The two intestinal caeca are thus connected to-
gether. From this point backward the esophagus decreases rapidly in
size, the caecum becomes large and is divided into numerous pouches
by fibrous partitions extending between the muscular layers above and
below, and eventually the esophagus opens into the intestine by a
narrow slit in its dorsal wall (pi. xn, fig. 3).
Reproductive organs. — Far in front of the opening of the esoph-
agus into the intestine the reproductive glands make their first appear-
ance. The anterior pouches are scattered, and lie above the intestinal
caeca well towards the sides of the body. These open directly on the
dorso-lateral surfaces. Farther back similar ones appear below the in-
testinal canal, and these open ventrally (pi. xn, fig. 3). In the intes-
tinal region the glands are very numerous and are scattered just inside
the muscular layers all over the body — dorsally, ventrally and laterally.
As many as 20 to 30 glands in a male are met with in a single section.
28 COE
Their ducts lead directly to the surface of the body, and consequently
open at any point instead of in certain definite regions as in many
species.
Both lateral nerves and blood vessels join above the hind gut as in
most other Nemerteans — the union of the nerves lying ventral to that
of the blood vessels.
This species is named in honor of Prof. Dr. Otto Burger, of Got-
tingen, whose monograph on the Nemerteans of the Gulf of Naples
forms by far the most important contribution which has yet appeared
relating to this group of worms.
Habitat. — Several individuals are often found knotted together in a
seemingly inextricable mass. The body lies coiled in a mass, and is
bent and folded in sharp angles.
The species was found under mussels on rocks between tides at
Glacier Bay (W. E. Ritter) and at Sitka.
Zygonemertes Montgomery.
Zool. Jahrb., x, p. 2, 1897.
A species of Nemertean was found at Sitka which agrees very
closely with Verrill's description of Amphiporus virescens.1 The
detailed anatomical description given by Montgomery,3 however, shows
at once that the Alaska Nemertean is a distinct species. Mont-
gomery has created a new genus for A. virescens Verrill based on its
anatomical peculiarities, especially the structure of the proboscis and
proboscis sheath. This genus he named Zygonemertes, with the
following characters as its chief peculiarities :
(i) The proboscis sheath reaches to the end of the body, while (2)
the thickened proboscis is but half as long; (3) basis of central stylet
large, elongated ; flattened or slightly concave posteriorly ; (4) central
stylet massive, not half the length of its basis : (5) ten or eleven pro-
boscidial nerves ; (6) body contractile, shape like Amphiporus, not
as elongate as in Emplectonema ; (7) ocelli numerous and small, ex-
tending along the nerve cords posterior to the brain.
While I am of the opinion that these characters are mainly of spe-
cific rather than of generic rank, yet I am convinced that because of
the large number of species already in the genus Amphiporus the
establishment of this new genus will be of great practical convenience.
The most tangible characters of the group are : ( i ) the eyes extend
posteriorly beyond the brain along the lateral nerve cords, and (2) the
1 Trans. Connecticut Acad., vui, p. 20, 1892.
2 Zool. Jahrb., x, p. 2 to 4, 12, 1897.
ALASKA NEMERTEANS
basis of the central stylet is massive and has a concave or a truncated
posterior end, while the stylet itself is comparatively weak. Of course
the number of nerves in the proboscis cannot be considered a generic
character. There are two Alaska species, then, which may be placed
in the genus — Z. thalassina, which is closely allied to the type species,
and Z. albida, which is a minute, white form.
8. ZYGONEMERTES THALASSINA sp. nov.
pi. n, fig. 5 ; Pi. vn, fig. i ; pi. xni, fig. 2.
Zygonemertes thalassina differs from Z. virescens chiefly in the
following peculiarities : Color of former species much darker ; eyes
more numerous ; shape of basis and central stylet of proboscis differ-
ent ; usually 5 stubby stylets in each lateral pouch, and 1 2 nerves in
proboscis. There are minor differences in other anatomical details.
Zygonemertes thalassina has a slender, somewhat flattened body ;
head broad, not sharply marked off from body ; one or two pairs of
very inconspicuous oblique furrows on sides
of head. The worms are active, and are rest-
less in confinement.
Ocelli. — Ocelli very numerous ; arranged
in two or three longitudinal rows along sides
of head, and extending backward along lat-
eral nerve cords far behind brain. They are
smaller posteriorly than in front and are more
widely scattered. In front of the brain there
are sometimes 40 or more ocelli arranged in
two or three irregular rows (fig. 5) which
follow the general outline of the lateral mar-
gin of the head. Immediately in front of the
brain about a half dozen much smaller eyes
are seen, and lateral to the brain are com-
monly 10 to 12 ocelli of moderate size. Be-
hind these are usually 8 to 15 small ocelli
scattered along the nerve cords at irregular
intervals. These commonly reach nearly as
far behind the brain as the distance from the
brain to the tip of the snout in moderate ex-
tension. In sections (pi. xin, fig. 2) they
are found to lie quite internal to the body musculature and almost di-
rectly upon the lateral nerves.
Size and color. — The specimens obtained were from 30 to 60 mm.
FIG. 5. Zygonemertes
thalassina. Dorsal view
of head showing outline
of brain and lateral nerves,
and arrangement of ocelli.
30 COE
in length in moderate extension, and ratner slender. The color was
olive-green both above and below. A coating of brown particles was
sometimes scattered over the dorsal surface. The proboscis is pale.
Proboscis. — The proboscis sheath extends to posterior end of body ;
the proboscis extends only about half way to posterior extremity. The
central stylet is remarkably short and blunt. Its basis (pi. vn, fig. i)
is at least twice, and often three times, as long as the stylet itself, and
is massive in proportions. Its diameter is nearly constant throughout
its length, although it is narrowed in front and constricted slightly at
about three-fourths the distance towards its posterior end. The pos-
terior extremity is sharply truncate or concave and often shows serrated
edges (pi. vn, fig. i). There are two lateral pouches of accessory
stylets, and each usually contains five stylets similar in size and shape
to the central stylet. The character of these remarkably stubby stylets
is shown in pi. vn, figures la and ib.
The lateral stylet pouches are imbedded in a thickened glandular
wreath, yellowish in color, which lies around the circumference of the
proboscis in front of the stylet. The mass of muscular tissue surround-
ing the basis of the central stylet is unusually thickened, and this ne-
cessitates an unusually long canal leading from the posterior chamber
to the stylet region. In each of the two specimens sectioned the pro-
boscis was provided with 12 nerves.
Cerebral sense organs. — Situated immediately in front of brain,
but in the ventral portion of the head. The canals leading to the ex-
terior pass obliquely forward and downward, and open on the antero-
ventral surface near the tip of the snout.
Nephridia. — The nephridia extend from a point immediately in
front of the brain backward throughout nearly the whole length of the
esophagal region. There is a single pair of efferent ducts situated in
the region of the brain, and opening to the exterior on the ventro-
lateral aspects of the body. In the specimens examined one of these
ducts lay as far forward as the ventral commissure of the brain, while
the other was situated opposite the posterior end of the ventral brain-
lobe.
Montgomery1 mentions for Z. virescens that the superficial epithelium
of the body contains numerous yellowish, sickle-shaped bodies situated
among the epithelial cells. These also occur in Z. thalassina, but
here they are of various sizes, and differ greatly in shape. Some are
sickle-shaped, others are rod-like or irregular in form, and have every
appearance of having been formed in the gland cells of the epithelium.
1Zoo\. Jahrb., x, p. 2, 1897.
ALASKA NEMERTEANS
31
Somewhat similar bodies may be found in various other Metanemerteans,
notably in Amphiporus bimaculatus and A. leuciodus. Here too they
appear to originate as secretions in the glandular cells of the epithelium.
The lateral nerve cords unite above the posterior end of the alimen-
tary canal as usual.
The tissues of the head in front of the brain contain large numbers
of sub-muscular glands which open mainly on the ante rior extremity,
although a portion of them open directly outwards. These sub-mus-
cular glands do not extend back into the esophagal region as they do
in many other species.
The intestinal caecum is paired from its origin. Its two branches ex-
tend forward on each side of the esophagus until they reach nearly
to the brain. Here they bend dorsally (pi. xui, fig. 2) and terminate by
abutting closely against the posterior ends of the dorsal brain-lobes.
The ova are large and opaque. They appeared to be fully mature
in June.
Habitat. — This species was found only at Sitka, where it was not
uncommon among hydroids, broken shells, etc., in clear water.
9. ZYGONEMERTES ALBIDA sp. nov.
pi. in, fig. 2 ; pi. vin, fig. 5.
This is a small, moderately slender spe-
cies, very active in habits. It was met with
only at Victoria, B. C., on the piles of a
wharf. Sexually mature individuals were
not more than 25 mm. in length. It was
associated with Amphiporus leuciodus,
which it somewhat resembles.
Color. — There are no markings on the
body, the color being white with a tinge
of yellow both above and below.
Ocelli. — This species may be easily rec-
ognized from other described forms by the
arrangement of the eyes (fig. 6). These
are scattered somewhat irregularly on the
head in front of the brain, and extend back-
ward as a single row on each side along the FIG. 6. Zygonemertes al-
lateral nerves for about two-fifths the length b{da- Outline of anterior
, ,, , , . rrn 11- .1 portion of body to show
of the esophagal region. The ocelli on the , J ... .
arrangement ot ocelli, or,
head are roughly arranged in two irregular brain . B> lateral nerve . /5>
rows on each side. There is an outer row proboscis sheath. X 12.
32 COE
of 9 to 1 2 eyes near each lateral border of the snout, and another row
inside and somewhat posterior to the latter, containing about 6 small
ocelli on each side, and of these 3 commonly lie nearly above the
brain. Back of the brain 7 to 10 ocelli occur at widely separated
intervals above the lateral nerve on each side.
Proboscis. — Proboscis comparatively large ; central stylet moderately
slender ; basis dark, moderately elongated, of nearly uniform diameter
throughout, and sharply truncated posteriorly (pi. vm, fig. 5). Ac-
cessory stylet pouches two in number, each commonly containing two
or three moderately slender stylets.
Cerebral sense organs. — Large, and situated immediately in front
of brain.
The eggs of these worms are few in number, but are fully YJ, the
diameter of the body in size; consequently there can be but a single
row on each side. The sexual products are mature in June.
Paranernertes gen. nov.
Body of large size, rather stout, usually much rounded in the eso-
phagal region but flattened posteriorly. Head not marked off from
body, of variable form, in some states of contraction often emarginate
in front. There is commonly a pair of inconspicuous oblique furrows
back of head. The nerve cords and blood vessels join on the dorsal
side of the posterior end of the intestine.
The mouth opens into the rhynchodseum. The proboscis sheath
commonly reaches but little beyond the middle of the body — in P.
peregrina to ^ the distance towards the posterior extremity. The
proboscis is small (P. pallida}, of medium size (P. peregrina}, or
large (P. carnea}. There is a single central stylet in the proboscis,
and usually four or more pouches of accessory stylets. Some indi-
viduals of P. peregrina, however, have but two. Ocelli are numer-
ous and minute. The cerebral sense organs are rather small and lie in
front of the brain. Sub-muscular glands are usually well developed.
The species of this new genus show considerable resemblance to
those of Emplectonema Stimpson. They differ, however, in general
shape and appearance of body, never being very long or slender, and
individuals do not coil their bodies into a mass as those of Emplecto-
nema are so prone to do. The proboscis is much larger and the
central stylet is always well developed. The proboscis sheath is also
much longer. In many respects the genus resembles Amphiporus.
The body is much longer, however, and not nearly so contractile, the
proboscis is not nearly so large, and the proboscis sheath is not so long.
ALASKA NEMERTEANS 33
The armature of the proboscis resembles that in some species of Am-
phiporiis.
Paranemertes is represented on the coast of Alaska by at least
three species.
10. PARANEMERTES PEREGRINA sp. nov.
pi. n, fig. 6 ; pi. in, fig. 5 ; pi. vn, fig. 7.
Body moderately elongated, flattened below, rounded on dorsal sur-
face ; anterior portion slightly more slender than middle region ; pos-
terior tapering gradually to extremity.
Head very variable in shape, commonly wider than the portion of
body immediately following ; flattened, sometimes sharply demarcated
by lateral constrictions. Tip of snout pointed, rounded, or emargi-
nate according to the state of contraction. On each side of the head
is an inconspicuous V-shaped furrow, with the ends pointing ob-
liquely forward above and below. The upper limb of the furrow
reaches into the dark color of the dorsal surface, where it is sometimes
conspicuous from its light color.
Color. — The color varies considerably as may be seen from a com-
parison of pi. n, fig. 6, and pi. in, fig. 5, but is commonly homo-
geneous dark brown, orange brown, or purplish brown above and on
the sides, while the ventral surface is opaque white or whitish yellow.
In most specimens the dark purple of the dorsal surface encroaches
considerably on the ventral surface, shading gradually into whitish or
yellowish. Seen from the ventral surface therefore the worms appear
dull white or yellowish white, with a wide border of dark purple.
Oftentimes the whitish color occupies scarcely more than the median
third of the ventral surface. Anteriorly the whitish color covers the
whole ventral surface, and on the head it covers also the sides and
front. The head is dark purplish brown above, bordered in front and
laterally by the light color of the ventral surface. At the posterior
border of the head is a small angular spot on each side corresponding
in color with that of the ventral surface. Behind the head is a nar-
row, V-shaped, dorsal marking, usually of lighter color, with its ends
pointing forward and outward. In paler individuals the pinkish
color of the brain lobes can be distinguished. The natural color oi
the body is well retained in formalin or alcohol.
Size. — Individuals of all sizes from 20 to 400 mm. were met with,
but the most common size was about 150 mm. in extension. The
width was commonly about 5 mm.
34
COE
rangement of
ocelli. Dorsal
surface. X 6.
Ocelli. — Numerous minute eyes are arranged in two groups on each
side (fig. 7). Of these, an anterior group of 12 or more small pig-
ment spots are scattered along each side of the antero-lateral margin,
while about as many more occupy an irregular cluster on
each side just in front of the brain. These latter ocelli
are usually well separated from the anterior, or mar-
ginal groups.
Proboscis. — Proboscis of medium size (as in Amphi-
porus}, slightly yellowish, and usually everted when the
animal is killed. The extruded proboscis is short and
thick with an unusually slender posterior chamber. The
armature (pi. vii, fig. 7) consists of a small, slender,
sharply pointed central stylet, and with either 2 or 4
FIG. 7. Par- Couches of reserve stylets. The basis of the central
anemertes per- . . .. . .. . . . ,
O stylet is very small and slightly enlarged posteriorly ;
ifne of anterior tne reserve stylets are slender and sharp like the central
portion of body one, and commonly number from 6 to 10 in each pouch,
to show the ar- When four pouches are present the number of stylets in
each is as great as when there are only two pouches.
The mouth and proboscis open together, but the
rhynchodaeum is short. The proboscis sheath reaches
about three-fourths the length of the body, or sometimes more than
three-fourths. In each of four specimens sectioned there were 14 con-
spicuous nerves in the proboscis. The proboscis has a remarkably nar-
row ring of gland cells on the periphery near the posterior end of the
basis of the central stylet.
A crowded mass of multicellular glands occupies the anterior por-
tion of the head. The anterior ones open on the tip of the snout. In
the brain region they open mostly on the lateral surfaces of the body,
while farther back, and in the esophagal region, they assume the
character of sub-muscular glands. No difference in appearance is no-
ticeable between the cephalic glands and those in the esophagal region.
They are present in the esophagal region only about as far back as
the openings of the nephridia. Throughout their course they open to
the exterior (by innumerable ducts which pass through the muscular
and basement layers) on the ventro-lateral aspects of the body.
Alimentary canal. — A pair of slender branches of the intestinal
caecum reach forward nearly to the brain commissures and lie well
above the lateral nerve cords. At about the point of the nephridial
openings these branches pass ventrally and occupy a position beneath
the esophagus. Another pair, coming forward from behind, take
ALASKA NEMERTEANS 35
their places. The ventral branches join to form the main median
caecum, from which short branches pass obliquely forward and dor-
sally at intervals. In cross section two pairs of branches are usually seen
besides the main caecum. One pair of these lies above and one be-
low the lateral nerves. They are disposed in such a way that one pair
ends at about the point where the second pair anteriorly originates.
The branches are only irregularly arranged in pairs. The esophagus
opens into the dorsal wall of the main caecum.
Nephridia. — The nephridia occupy the anterior z/$ of the esoph-
agal region. They are large, with numerous branches lying above the
lateral nerves, and frequently passing internally to the nerves and be-
neath the esophagus. They reach forward nearly to the brain. At
about 1/3 of their distance posteriorly the main nephridial tubes, lying
above the lateral nerves increase greatly in size and a pair of remark-
ably large efferent ducts pass externally to the lateral nerves to open on
the lateral aspects of the body slightly below the lateral margins. The
main duct reaching posteriorly from this point is larger than that in
front.
Blood -vessels. — There is a pair of large blood lacunae in the head as
usual ; they join anteriorly by a broad anastomosis. The three longitu-
dinal vessels are well developed to the end of the body where they an-
astomose above the anus. Sometimes the dorsal vessel lies beside or
even above the proboscis sheath throughout a portion of its course,
instead of below the sheath as usual.
Nervous system and sense organs. — The cerebral sense organs lie
well in front of the brain, and external to the blood lacunas. They
open into a slight furrow on the lateral aspects of the head a little an-
terior to their own position. The lateral nerves form a commissure
above the anus as usual.
Reproductive organs. — The sexual products were nearly mature in
June and July. They are formed in numerous pouches which sur-
round the intestinal canal on all sides. In a male as many as twenty
sexual pouches were seen in a single section. They open directly to
the exterior, whatever be their position.
Habitat. — This is a restless Nemertean, and on cloudy days was fre-
quently met with crawling about over the stones on the beach between
tides — which peculiarity has suggested its specific name. It was found
abundantly at nearly all the collecting stations from Victoria, B. C., to
Unalaska Island, and is one of the commonest Nemerteans of the coast.
It occurs from low tide well up toward high water mark in every
variety of location — under stones, among seaweeds, barnacles, mus-
36 COE
sels, etc. The individuals are very voracious feeders, and were taken
not infrequently with partially swallowed Chaetopods. Their tenacity
of life is remarkable — they will live for days in a small quantity of
filthy water.
ii. PARANEMERTES PALLIDA sp. nov.
Pl. VII, fig. 3; Pl. XII, fig. I.
Body rather large, stout, rounded, and almost cylindrical anteriorly,
somewhat flattened behind when extended ; head variable in shape,
not sharply marked off from portions immediately following, at
certain states of contraction emarginate in front. A pair of incon-
spicuous oblique furrows back of head. When contracted the worms
are nearly cylindrical and of about the same diameter throughout, ex-
cept at the extremities, both of which are pointed.
Color. — The \vhole body, both above and below, is commonly uni-
form opaque white, sometimes showing traces of yellowish or reddish
tints, especially in the anterior portions.
Ocelli. — Ocelli minute and numerous. In ordinary states of con-
traction they are arranged in a pair of elongated, irregular clusters on
the antero-lateral margins of the head. The number of such ocelli is
sometimes 30 or more in each of the two clusters.
Proboscis. — Small, short, and unusually slender (pi. xn, fig. i). Its
armature consists of a moderately slender central stylet and usually 4
pouches of accessory stylets. The basis of the central stylet is mod-
erately slender, slightly constricted near its middle portion, rounded
behind, and of approximately equal length with the stylet (pi. vn, fig.
3). There are commonly two accessory stylets in each of the 4
pouches. The chambers posterior to the stylet apparatus are remark-
ably narrow.
The mouth opens into the rhynchodseum. The proboscis sheath
extends but little beyond the middle of the body, and sometimes not so
far as the middle. One specimen had 9 nerves in the proboscis ; an-
other had 10. These nerves do not all enter the proboscis from the
ventral side, as they do in Amphiporus angulatus, but those supply-
ing the dorsal portion enter direct from that side.
Closely packed sub-muscular glands are present on the right and
left sides of the body, and extend well inward towards the median line.
Their ducts pierce the musculature and other layers of the body walls
mainly on the latero-ventral aspects of the body. Twenty or more
are frequently met with in a single section. These glands occupy also
the region in front of the brain, and extend backward in decreasing
ALASKA NEMERTEANS 37
numbers to the commencement of the intestinal region. In front of
the brain is a large and irregular cluster of glands, which open ante-
riorly on the tip of the snout.
Cerebral sense organs. — Situated in front of brain, and near latero-
ventral margins of head. The ducts which place them in communi-
cation with the exterior run obliquely forward and downward, and
open immediately on the surface.
Nephridia. — The nephridial canals extend from near the brain
throughout almost the entire esophagal region. The main trunks lie
above the lateral nerve cords, and send off numerous branches both
ventral and dorsal to the nerves. There is a single pair of remarkably
large efferent ducts opening just below the lateral margins of the
body, and slightly anterior to the middle of the esophagal region. In
one series of sections these ducts are so precisely paired that both ap-
pear in a single section (pi. xn, fig. i). The efferent ducts pass
from above obliquely downwards and external to the nerve cords.
The blood- vascular system consists of cephalic lacunae, and anasto-
mosing longitudinal vessels, as in related genera.
The intestinal caecum does not reach forward nearly to the brain.
There are comparatively few lateral diverticula (pi. xn, fig. i).
The lateral nerve cords unite above the posterior end of the intestine.
Paranemertes pallida was found only at Yakutat and at Sand Point
on Popof Island, and few specimens were obtained. These were
about 150 mm. to 250 mm. in length, and perhaps 5 mm. in width.
They were found between tides under stones covered with algae.
12. PARANEMERTES CARNEA sp. nov.
pi. in, figs. 3, 4; pi. vn, fig. 4; pi. vin, fig. 7.
This species was found to be extremely variable in size, shape of
body, and head, and especially in the armature of the proboscis. As
shown in pi. in, figs. 3 and 4, the body is rather stout, rounded in
the esophagal region, flattened both above and below posteriorly, and
ending rather abruptly behind. The head is very variable in shape,
being pointed, rounded, broadened, or emarginate in front, according
to its state of contraction. It is most commonly a little broader than
the parts immediately following, and is not distinctly marked off,
although a slight oblique furrow on each side is sometimes seen behind
the brain. From the dorsal surface the furrows of the two sides give
the appearance of a very faint V-shaped marking with the angle pro-
jecting backward in the median line. A little farther forward, as
seen in pi. vin, fig. 7, a pair of shallow furrows occupies the sides of
COE
the head. These are also V-shaped with the angle projecting back-
ward on the lateral margins. The ventral limb of each V-shaped fur-
row reaches nearly to the opening of the rhynchodzeum ; on the dorsal
surface the ends of the dorsal limbs are separated by about y§ the
diameter of the body in ordinary states of contraction (fig. 8).
In microscopic sections the V-shaped grooves on each side of the
head are conspicuous, because of their differentiated epithelium. In
these shallow depressions the epithelial cells are of smaller size, more
slender in form, and seem to partake more of the nature of sensory
cells. In these grooves the ordinary glandular cells are wanting. In
the ventral limb of each V-shaped groove opens the tube which places
the cerebral sense organs in communication with the exterior.
Ocelli. — There are commonly 20 to 32 eyes on the head arranged
in 4 more or less distinct clusters, though their relative positions change
with the contraction of the head. The two anterior clusters contain
about 4 to 6 eyes each, and occupy the antero-lateral margins of the
head (figs. 8, 9). The posterior clusters lie directly back of these,
FIG. 8. FIG. 9.
FIGS. 8 and 9. Paranemertes carnea. 8. Outline of head from dorsal sur-
face showing arrangement of ocelli. The two pairs of lateral indentations indi-
cate the lateral and dorsal furrows. X 8. 9. Outline of the head when extended.
Dorsal surface, showing arrangement of ocelli. X 6.
and immediately in front of the brain. Each of these clusters contains
8 to 12 scattered ocelli. An individual from Taku Harbor had 6
ocelli in each anterior cluster, and 10 in each posterior one. Another
specimen had only 3 or 4 eyes in each of the four clusters, but in this
case the ocelli were larger. A specimen from Popof Island had 3
large eyes in each anterior, and 6 to 8 in each posterior cluster ; one
from Virgin Bay had 6 in each anterior, and 7 in each of the others,
ALASKA NEMERTEANS 39
and had 3 single eyes between the anterior and posterior clusters,
while one from Yakutat had 3 large ocelli in a row on the anterior
margin of the head, and a pair of scattered clusters of 8 to 12 eyes
each in front of the brain. When the head is contracted so that it is
emarginate in front, the arrangement of the eyes into clusters disap-
pears, and they are then all scattered irregularly on the antero-lateral
margins. Figs. 8 and 9 show the general arrangement of the eyes.
Color. — The whole body, both above and below, is a homogeneous,
very pale red or flesh-color (pi. in, figs. 3, 4), and is entirely with-
out markings except for a longitudinal median line of deeper red,
showing the position of the proboscis sheath, and the deeper color of
the intestinal lobes. In some specimens the reddish color was more
pronounced anteriorly, and some were very pale. The intestinal lobes
are seen through the clear tissues of the body walls as narrow, trans-
verse markings, slightly darker than the rest of the body. The reddish
color of the brain and lateral nerves is often conspicuous from the
dorsal surface (pi. in, fig. 3).
Size. — Individuals were seen which were 500 mm. long in greatest
extension, although the majority were less than half this length.
Proboscis. — Mouth and proboscis open together through a subter-
minal pore. The proboscis is large (pi. vni, fig. 7) and of moderate
length. The short proboscis sheath does not reach far beyond the
middle of the body. The armature of the proboscis shows marked
variations. The size and shape of the central stylet and its basis,
however, remain fairly constant. The central stylet is of the regular
Amphiporus type, is moderately slender, and rests on a moderately
slender basis. The basis is slightly narrower in front and is rounded
behind (pi. vn, fig. 4). The number of pouches of accessory stylets
is commonly from 6 to 12. One specimen had 12 of these pouches,
each with one or two slender stylets ; each of two others had six
pouches with two stylets in each pouch. Three specimens had each
twelve nerves in the proboscis, while a fourth specimen had but
eleven.
Cerebral sense-organs. — Unusually small and situated some distance
in front of brain-lobes. They occupy positions very close to the ven-
tro-lateral borders of the head, and beneath the cephalic blood lacunae.
Nephridia profusely branched, and extending throughout the greater
portion of esophagal region, though they do not reach the brain.
Their numerous branches ramify both above and below the lateral
nerve-cords, and several efferent ducts of small size lead to the exterior
from both the dorsal and ventral branches. There may be about five
40 COE
pairs of efferent ducts, all of which open in the immediate vicinity of
the lateral nerves. In one series of sections a small efferent duct from
one of the branches above the lateral nerve was followed only two sec-
tions farther back by a similar, though larger, duct from a branch below
the lateral nerve on the same side. The nephridia end posteriorly near
the point where the esophagus opens into the intestine.
Sub-muscular glands of limited number lie between the muscula-
ture and the intestine and proboscis sheath. These glands are closely
packed together in the head in front of the brain ; back of this point
they become widely scattered, although they do not cease entirely until
back of the esophagal region.
A short intestinal caecum with a few wide lateral diverticula extends
forward beneath the esophagus. This cascum is shorter than in most
species of the genus, and does not reach nearly to the brain. The
esophagus also is short and opens directly into the dorsal wall of the
intestine.
A pair of large blood lacunas occupies the anterior portion of the head
as usual.
The lateral nerves and longitudinal blood vessels join above the pos-
terior end of the alimentary canal, as in most species.
Reproductive glands in both male and female are very numerous,
and are situated both above and below the intestine. They open di-
rectly to the surface, as could be determined from their rudimentary
ducts, although the sexual products were very immature in June and July.
Habitat. — This species is conspicuous because of its clear, rosy or
flesh-like color, which is all the more striking in contrast with the black
mud in which it is usually found. It occurs between tides in muddy
locations over a large portion of the southern Alaska coast. It was
collected at Taku Harbor, Sitka, Yakutat, Prince William Sound, and
Popof Island, although only a few were found at each locality. Usually
but one or two specimens were found in several hours' digging.
Amphiporus Ehrenberg.
Symbolce Physical, Berlin, 1831.
This is by far the most common genus on the Alaska coast, and to it
belong six of the thirty species of Nemerteans collected.
The genus Amphiporus includes mostly rather stout, solid, often
flattened forms, usually of considerable size, which are capable of an
almost incredible amount of extension and contraction. A few forms,
however, are long and cylindrical, even when contracted, but others
ALASKA NEMERTEANS
41
can contract until the body becomes almost barrel-shaped. The worms
can neither swim nor roll up spirally.
Proboscis. — Provided with a single, well-developed central stylet,
with a cartridge-shaped basis, and with two or more pouches of acces-
sory stylets. The proboscis sheath usually reaches nearly or quite to
the end of the body.
Ocelli, — Usually present in very considerable numbers. A few
forms are without eyes, and a few others have but a single pair — there
are never 4. The eyes do not extend far behind the brain.
Cerebral sense organs. — Usually well developed. Their position
is most commonly in front of the brain, but they are sometimes beside
or even behind the ganglia.
13. AMPHIPORUS ANGULATUS (Fabr.) Verrill.
pi. vi, fig. 4 ; pi. vn, figs. 2, 2a ; pi. xi, fig. 2 ; pi. xm, fig. 3.
Fasciola angulata O. FABRICIUS, Miiller's Verm. Terrest. et Fluv., i, p. 58,
1774.
Omatoplea stimpsonii GIRARD, in Stimpson, Invert, of Grand Manan, Smith-
sonian Contributions to Knowledge, p. 28, 1853.
Nareda superba (?) GIRARD, loc. cit.
Cosmoccphala beringiana STIMPSON, Proc. Acad. Nat. Sci. Philadelphia, p.
165, 1857.
Ai'iphiporus angulatus (FABR.) VERRILL, Marine Nemerteans of New Eng-
land, Trans. Conn. Acad., p, 10, 1892.
" This large and conspicuous species is gen-
erally easily recognized by its clear dark purp-
lish or chocolate-brown color above, with pale
margins and a trapezoidal or triangular white
spot on each side of the head and usually with
a narrow white line across the neck ; and by the
pinkish or flesh-colored lower surface. Ocelli
in two or more rows in an elongated groove on
each antero-lateral margin of the head, and a
pair of small sub-dorsal clusters on the transverse
white nuchal band." (Verrill, loc. cit.) The
arrangement of the eyes and markings on the
head of the Alaska specimens are shown in iig.
10 and in pi. vi, fig. 4. In ordinary state of
contraction the body is rather short and stout.
When disturbed it can become so greatly thick-
ened anteriorly that its transverse diameter is fully y$ as great as its
length. In extension the body is but moderately elongated, and is
relatively broad and flat. It contracts very much as does a leech.
FIG. 10. Amphi-
porus angulatus. Dor-
sal view of anterior por-
tion of body showing
markings on the head
and the arrangement of
ocelli. X 8-
42 COE
The Alaska specimens are commonly larger than have been recorded
elsewhere, often measuring 200 mm. or more in length and 10 mm. in
width.
Proboscis. — The proboscis is large, thick, and pale reddish or
salmon in color. The smallest specimen collected had 17 nerves in
the proboscis; four other specimens examined had each 18 probos-
cidial nerves, one had 19 nerves, and two others had 20 each.1 This
shows more strikingly than has previously been pointed out that the
number of nerves in the proboscis is variable to a very considerable ex-
tent. Burger2 has shown that Drepanoporus crassus may have 19 or
20 nerves, and D. spectabilis 24 or 26. Nevertheless in the other
Alaska species the number has been found surprisingly constant. The
number of nerves in any particular proboscis remains perfectly con-
stant so far as I have observed from the anterior end back as far as the
stylet region. Here they break up into a plexus and lose their iden-
tity. The nerves in A. angulatus enter the proboscis at its anterior
attachment and in its ventral portion (pi. xi, fig. 2). They then
divide into their definite number of branches (usually 18) which pass
obliquely dorsally and arrange themselves symmetrically on the periph-
ery. The proboscis sheath extends within a few sections of the pos-
terior end of the body.
The armature of the proboscis is made up of a moderately slender
central stylet, and (usually) two pouches of accessory stylets. The
basis of the central stylet is about as long as the stylet itself. It is
moderately slender, constricted near its middle (pi. vn, figs. 2, 2«),
enlarged and rounded posteriorly. Each reserve pouch commonly
contains 5 to 7 rather slender stylets, similar in size and shape to the
central stylet.
Ocelli. — Numerous and characteristic in arrangement. The dark
pigment on the head, however, often renders them difficult of accurate
determination. Girard3 states for Omatoplea stimpsonii that there are
six or more minute eyes " situated in an oblique, simple row, on either
side of the head anteriorly." The same author (loc. cit.) describes
JOf eight specimens of this species from Eastport, Maine, one had but 17
nerves in the proboscis, six had 18 each, and one had 19 or 20. Other anatom-
ical details in the eastern form agree perfectly with those of specimens from
Alaska. I have recently examined a number from the original locality of Stimp-
son's C. beringiana (Bering Strait), and have no doubt as to the specific identity
of this form with that from southern Alaska and from Eastport, Maine.
2 Fauna u. Flora des Golfes von Neapel. Monogr. 22, Nemertinen, p. 372,
1895.
3 Marine Invert. Grand Manan, Smithsonian Contr. to Knowledge, p. 28, 1853.
ALASKA NEMERTEANS 43
Nareda superba as having but a single pair of rounded ocelli situated
wide apart on the transverse white band of the neck. It seems highly
probable, as Verrill suggests,1 that both of the species are identical ;
in the one case only the marginal eyes were seen, while in the other
the cerebral clusters were supposed to represent single eyes and the
marginal ones were overlooked. Verrill l describes the eyes correctly,
and his diagnosis of the species is so full and accurate that it is neces-
sary to describe here the internal anatomy only. A pair of elongated
clusters of ocelli lies on the antero-lateral margins of the head, and an-
other smaller cluster on, or near, the angular white spot on each side
of the head. As shown in fig. 10, each of the anterior clusters may
contain upwards of 20 ocelli arranged in two or more irregular rows
nearly parallel with the antero-lateral margin of the head, while the
posterior groups may consist of 8 to 15 similar ocelli. The posterior
groups are situated deep in the tissues of the head. Of course the
number of ocelli varies greatly in different individuals.
Cerebral sense organs. — Well developed. They lie a little in front
of the brain, beside the esophagus, and below the cephalic blood
lacunas. Each sense organ has a wide canal which leads a short dis-
tance anteriorly and opens to the exterior on the latero-ventral aspect
of the body. The brain itself is of large size, with a thick ventral
and narrow dorsal commissure (pi. xi, fig. 2).
Nephridia. — The nephridia extend from near the brain (pi. xi,
fig. 2) well backward in the esophagal region. In one specimen there
were two pairs of efferent ducts opening on the latero-ventral aspect
of the body ; in another only one pair.
Cephalic glands. — The cephalic glands open on the tip of the snout
and are well developed. Sub-muscular glands, likewise, are remark-
ably abundant. They reach from the brain region well back towards
the end of the esophagus. They are multicellular, each one being
composed of upwards of a score of large, vacuolated cells with small
nuclei situated on the side farthest from the lumen. Each gland has
a twisted duct leading through the muscular layers, basement membrane,
and integument, and opening to the exterior on the ventro-lateral aspects
of the body (pi. xi, fig. 2).
Beneath the esophagus a broad caecal appendage of the intestine
stretches forward well toward the brain region. This caecum consists
of a large median canal with pouch-like diverticula extending dorsally
above the lateral nerve cords.
There is the usual anastomosis of the three longitudinal vessels, and
1 Marine Nemerteans of New England, Trans. Conn. Acad., vm, p. 12, 1892.
44
COE
of the pair of lateral nerve cords (pi. xiu, fig. 3), above the hind gut
and slightly in front of the anus. As seen from the figure, the union
of the blood vessels is directly dorsal to that of the nerve cords.
The reproductive glands occur both above and below the alimentary
canal. Sexual products were not nearly mature in June and July.
Habitat. — The species is extremely abundant along the whole
Alaska coast as far west as Unalaska Island, and Stimpson records it
from Bering Strait. It is found under stones between tides in all sorts
of situations. Stimpson's specimens came from a depth of five
fathoms. The species is found on the Atlantic coast of North America
from Massachusetts Bay to Greenland (Verrill, loc. cit.*).
14. AMPHIPORUS BIMACULATUS sp. nov.
pi. i, fig. 4; pi. v, fig. 10 ; pi. vm, fig. 2 ; pi. xn, fig. 2.
Body rather short, broad, and flattened both above and below. Head
narrower than parts immediately following. Body of about the same
width and thickness throughout esophagal and intestinal regions.
Posterior extremity tapers rather abruptly to the pointed or rounded end.
Opening of rhynchodaeum situated on subterminal portion of snout.
From near this opening a pair of slits pass obliquely backward and
upward behind the eyes to the brain region.
Color. — The color of this species is very striking. The whole dor-
sal surface back of the head is deep brownish orange, somewhat paler
behind. The head is without color, or of a very much paler color
than the rest of the dorsal surface, and in the center of this pale area
two oval, black or very dark brown spots lie side by side. These are
very characteristic, and are conspicuous even in alcoholic specimens.
The black spots sometimes occupy a considerable portion of the pale
area, and are sometimes sharply angular in front (pi. i, fig. 4). In the
median line of the body the color is slightly paler than elsewhere and
in the center of this paler stripe is a dark, but inconspicuous, longi-
tudinal line. The pale stripe and dark line both fade out at a point
about y§ the distance towards the posterior end of the body. The
brain lobes appear as pinkish bodies just posterior to the black cephalic
spots. The whole ventral surface is of a homogeneous, pale orange
or flesh color, with the exception of pinkish spots marking the position
of the brain, and a slightly paler stripe below the anterior portion of
the proboscis sheath.
Ocelli. — The eyes are rather large, and number 25 to 30 or up-
wards on each side (fig. n). The majority lie in an irregular margi-
nal row beside and in front of each of the black cephalic spots. At
ALASKA NEMERTEANS 45
the posterior end of each marginal cluster the ocelli are more closely
and more irregularly placed, and often occupy several rows. In ad-
dition to these marginal clusters a closely set group of about a half
dozen smaller ocelli is situated in the light area lateral to the posterior
end of each of the dark cephalic spots. These ocelli lie deeper in the
tissues of the head, and near the brain (pi. i, fig. 4).
Proboscis. — The proboscis is remarkably large, and its constituent
layers are very sharply defined (pi. xn, fig. 2). The proboscis
sheath has a correspondingly massive development,
and reaches to the very extremity of the body. The
stylet apparatus of the proboscis is very characteris-
tic of the species, because of the extreme minuteness
of the basis of the central stylet. The central stylet
itself is very long and slender, while the length of
its basis is but half as great. The basis is con-
stricted in the middle, and is ^ as wide as long FJG TI
(pi. vm, fig. 2). There are usually four pouches porus bimaculatus.
of accessory stylets. These pouches are not evenly Outline of head to
distributed on the circumference, for two lie close snow position of
.i • j f ., 1-1 markings and ar-
together on one side of the proboscis, the other two ,
L rangement of ocelli.
on the opposite side. There are usually five to \/ 5.
seven slender stylets in each of the four pouches.
Most of the stylets are much smaller than the central stylet. Measure-
ments of the stylets of one individual about 100 mm. long are : central
stylet .12 mm. long, .015 mm. wide near base; basis of central stylet,
.06 mm. long, .04 mm. wide ; largest accessory stylet, less than .1 mm.
long. The proboscis is provided with 16 large nerves (pi. xn, fig. 2).
The mouth and proboscis open together. There are three large
communicating blood lacunae in the head, one on the right, one on the
left, and one dorsal to the rhynchodaeum.
Cerebral sense organs. — Remarkable for their large size, being
fully as large as either of the brain lobes. They lie lateral to the
brain, slightly behind the commissures, and in the angle between the
dorsal and ventral lobes. A large process from the posterior end of
the dorsal lobe furnishes the sense organs of the same side with an
abundant innervation. Their posterior ends extend backwards beyond
the dorsal brain lobes, against the posterior faces of which they are
closely pressed. Behind the dorsal brain lobe the sense organs lie
directly dorsal to the lateral nerve-cords and are bathed on their
internal borders by large blood lacunae. A section through this point
is not very different from a corresponding section of a Heteronemer-
46 COE
tean. Of the Alaska Metanemerteans here recorded this is the only
one in which the cerebral sense organs lie posterior to the brain com-
missures. The canal by which each sense organ communicates with
the exterior is of large size, runs anteriorly in front of the brain, and
opens on the ventro-lateral aspect of the head.
Nephridia. — The nephridia reach forward close to the posterior ends
of the cerebral sense organs. Anteriorly there is a tangle of small
vessels, but farther back these unite into a single large, branched
canal which runs close beside the blood vessels above the lateral nerve
on each side. From these canals a pair of large efferent ducts pass
above the lateral nerve cord, and open to the exterior of the body just
below the lateral margin.
Sub-muscular glands. — Present along anterior portion of esophagal
region, but not very abundant.
A broad and profusely branched intestinal ccecum runs forward
from the intestine well toward the brain region. The caecum lies well
beneath the esophagus and sends off numerous pouch-like branches
dorsally above the lateral nerve cords.
Reproductive glands. — These occur both above and below the intes-
tinal canal. Although the sexual products were very immature in one
of the specimens sectioned, yet the efferent ducts of the glands were
formed as far outward as the basement layer of the cutis. Here each
duct ended in a swollen chamber lined with cylindrical epithelial cells.
Size. — The individuals of this species varied from 40 to 150 mm. in
length. The largest were about 6 mm. wide and 2 mm. thick.
Habitat. — The species was collected at Victoria, B. C., on the
piles of the wharf; at Sitka among hydroids, etc., near low water (W.
E. Ritter), and a finely preserved specimen from Puget Sound, State
of Washington, was given me by Prof. Trevor Kincaid.
15. AMPHIPORUS TIGRINUS sp. nov.
pi. iv, figs. 5-8; pi. vin, fig. 4; pi. x, figs. 3, 4.
Body moderately slender, rounded throughout, head not marked off
from parts immediately following, rather narrow and pointed in front ;
posterior extremity of body narrow. On each side of the head is a
shallow, inconspicuous, oblique groove.
Color. — In June, at the time the specimens were collected, the
sexual products were fully mature, and the species showed marked
sexual color varieties. The prevailing color of the females was yel-
lowish orange both above and below, but except in the esophagal re-
gion, this color was to a great extent obscured by the dark olive green
ALASKA NEMERTEANS 47
color of the mature ova. These ova developed in large pouches on
each side of the body, and each pouch with its contents appeared as a
dark green spot. In many cases several adjacent pouches lie nearly
in contact, giving the external appearance of dark green blotches.
Seen directly from the dorsal surface the green spots appear on
each side in more or less regular pairs, those of the two sides being
separated by an interrupted, narrow, longitudinal, median band of
yellow. From the sides the ovaries appear as irregular transverse
stripes of dark green alternating with the yellow color of the body —
hence the specific name, tigrinus. The males are much less deeply
colored. They are pale yellowish with a slight tinge of green, and
the spermaries appear as innumerable cream-colored specks. Both
males and females have a narrow, longitudinal, median band of brown-
ish on the anterior dorsal portion of the body. After preservation in
alcohol both sexes assume a greenish color, which is retained even after
mounting in balsam.
Ocelli. — The eyes are numerous, and are arranged in two irregular,
and scarcely separated, clusters on each side of the head in front of
the brain. The individual ocelli are so irregular in shape, so variable
in size, and so closely massed together, that it is difficult to determine
their precise number. Many of them appear as ragged pigment
masses. Commonly, however, there are a dozen or more of such
ocelli in each of the anterior clusters, and perhaps 8 to 10 in each of
the posterior ones. The ocelli of the anterior clusters are scattered
through the tissues of the head from the dorsal to the ventral surface.
Some of the ocelli are three times as large as are others. In contraction all
the eyes of the same side form a single confused cluster. Because of
their variability of position no drawing is given of their arrangement.
Proboscis. — The long and well developed proboscis is provided
with a remarkably weak armature. This consists of a small central
stylet and two pouches of accessory stylets. The central stylet is
small and short, but is acutely pointed. Its massive basis, double the
length of the central stylet itself, is short, thickened, and rounded be-
hind (pi. vin, fig. 4). In a worm 75 mm. long, the central stylet
measured about .075 mm. in length; the basis was .15 mm. long and
.075 mm. in average diameter. The accessory stylets are, like the
central stylet, short, broad at the base, but sharply pointed. They
usually number about five to each pouch. The glandular wreath
about the stylet is well developed, and in the specimens examined is
deep green in color even after mounting in balsam. The proboscis
sheath reaches nearly to the posterior end of the body.
48 COE
Cerebral sense organs. — Smaller than in most species of the genus.
They are situated slightly in front of the brain, but are well separated
from it because they lie near the ventro-lateral border of the head.
The canals placing them in communication with the exterior are, con-
sequently, extremely short. Each canal opens into the shallow,
oblique furrow on the side of the head. The posterior ends of the
sense organs lie beneath the anterior borders of the ganglia, but much
nearer the ventral surface. A pair of large nerves given off from the
dorsal ganglia opposite their commissure connect with the sense organs.
The lateral nerve cords unite above the posterior end of the intestine
as usual.
The body cavity in the esophagal region is filled with an unusually
large amount of gelatinous tissue, which occupies the considerable
space between the muscular layers and the esophagus and proboscis
sheath (pi. x, fig. 4). In this gelatinous tissue the lateral nerves are
situated, and through it a complex system of blood vessels and nephri-
dial canals ramifies.
The intestinal caecum is very broad and has but short lateral diver-
ticula. It lies wholly beneath the esophagus, and the branches do
not extend above the lateral nerves. The caecum ends anteriorly far
behind the brain region. The esophagus becomes very small before
it empties into the dorsal wall of the broad intestine.
The attachment of the proboscis to the tissues of the head, the posi-
tion of the rhynchodaeum and its openings into the esophagus and pro-
boscidial cavity, the position of the dorsal and ventral brain commis-
sures, the cephalic glands and other organs are shown in pi. x, fig. 4.
The sexual products are mature in June. The whole body becomes
distended with the pouches of sexual elements, and the cavity of the
alimentary canal is much reduced in consequence. The ova are large
and deep olive-green.
The length of the specimens obtained, both males and females, was
about 75 to 100 mm. in extension.
Habitat. — This species was met with only at Farragut Bay, where
it occurred under stones in muddy locations at about half tide.
16. AMPHIPORUS NEBULOSUS sp. nov.
pi. iv, fig. i ; pi. vm, fig. 6; pi. xi, fig. i.
Body short, rather broad, and much flattened ; narrower anteriorly
than in the intestinal region, and tapering gradually posteriorly.
Mouth sub-terminal ; head pointed or expanded in front, according to
ALASKA NEMERTEANS 49
state of contraction. A V-shaped furrow is present on each side of
the head near the tip ; the angles of these furrows point obliquely
forward above and below.
Color. — Dull white or pale yellowish on dorsal surface; very
thickly mottled with confluent dark brown blotches and dots which
largely obscure the ground color. Margins of the head without spots.
There are faint indications of a pair of transverse
lines without color — one near the tip of the snout
and the other near the brain region, the latter some-
times becoming a shallow, irregular, V-shaped fur-
row. Ventral surface dull white or yellowish,
without markings other than deeper yellow spots
which indicate the positions of the genital sacs, pIG I2
and the darker color of the intestinal canal. riis nebulosus. Out-
Ocelli. — On each side of the head are from 18 line of the head to
to 25 ocelli, arranged in three irregular groups show arrangement of
/£. x /-,, ,, •• f ., ocelli. Dorsal sur-
(ng. 12). Close to the anterior border of the ,
snout are 4 or 5 large cup-shaped ocelli on each
side. Behind these and bordering each lateral margin are about 8
much smaller eyes in an irregular group, while 3 or 4 small eyes are
scattered between these and the anterior group. Behind each lateral
group, and not far in front of the brain, about 7 to 10 small ocelli lie
in an irregular cluster deeper in the substance of the head, and are
therefore less easily visible.
Size. — The specimens obtained measured 100 to 150 mm. in length,
and 5 mm. in width. The esophagal region is short, rounded above,
flattened below, and thicker than the intestinal region.
Proboscis. — The proboscis sheath reaches nearly to the extreme end
of the body. Proboscis thick, fairly large, and white. It is provided
with 17 nerves. Basis of central stylet very much broadened poste-
riorly (pi. vm, fig. 6), flat or even emarginate behind, narrow in front.
Central stylet as long as the basis, slender, acutely pointed. Accessory
stylets in two pouches ; similar to central stylet, but sometimes very
slightly curved ; commonly 3 in each pouch. The pouches lie well
behind the central stylet in ordinary extension (pi. vm, fig. 6) . Wreath
of gland-cells surrounding basis of central stylet moderately broad.
In the brain region (pi. xi, fig. i) and for some distance posteriorly
an abundance of large multicellular glands are thickly placed in the
ventro-lateral aspects of the body, and are mostly situated among the
fibers of the longitudinal muscular layer. Behind the brain these sub-
muscular glands become so closely packed together that the muscular
50 COE
layer is divided into an outer and an inner portion in the region where
the glands are situated. The ducts from the glands (pi. xi, fig. i,
smg) pass through the muscular layers and basement membrane to
open to the exterior among the epithelial cells of the integument.
These glands become smaller and more scattered near the region of the
nephridial openings, but do not disappear entirely until near the end
of the esophagal region.
Alimentary canal. — The mouth and proboscis open together into
the rather long rhynchodaeum. The esophagus is as usual in the genus.
A single pair of small intestinal caeca reach forward well toward the
brain region. They lie immediately below the esophagus, and near
the middle line. Farther back they join a median, broad, unpaired
caecum. This has wide, paired, lateral outgrowths which, still farther
back, alternate with clusters of reproductive glands. These lateral ap-
pendages of the caecum lie mostly above the reproductive glands, as
well as alternate with them. Much farther back the esophagus de-
creases greatly in size and opens into the intestine by a longitudinal
slit in the dorsal wall of the latter. The intestinal pouches are rather
deep ; the anus is subterminal.
Nephridia. — The nephridial canals reach anteriorly nearly to the
region of the brain, where there is a single branched longitudinal ves-
sel on each side. Throughout the greater part of its length, this main
canal lies above the lateral nerve cord, but sometimes lies internal to
it or above it. In the anterior third of the esophagal region the main
canal becomes very large ; here an efferent duct branches off, passes
outside the lateral nerve, and bends downward to open on the exterior
of the body on the ventro-lateral aspect. There is but one efferent
duct on each side, and the two are sometimes exactly paired. Back of
the efferent ducts, the nephridial canals decrease rapidly in size, and
disappear far in front of the anterior end of the intestinal region.
A pair of large blood lacunae lie in the anterior portion of the head
as usual. In the brain region they divide into numerous smaller ves-
sels (pi. xi, fig. i). The lateral vessels form a broad anastomosis with
the dorsal vessel above the anus.
Nervous system and sense organs. — The dorsal ganglia are closely
fused with the ventral. They are without distinct posterior lobes, and
sink gradually into the ventral ganglia (pi. xi, fig. i). The cerebral
sense organs are well developed. They lie in front of the brain,
some distance ventrally from the dorsal ganglia, and communicate with
the exterior by a canal which opens ventro-laterally. Each sense
organ is lobulated posteriorly and provided with a large nerve (son,
ALASKA NEMERTEANS 51
pl. xi, fig. i) which arises from the ventral side of the dorsal ganglion
near the ventral commissure. The union of the lateral nerve cords
above the anus lies in the same section as the anastomosis of the three
longitudinal blood vessels. A pair of small nerves from the lateral
cords continues backward beyond the commissure to the end of the
body.
Reproductive organs. — The sexual glands first make their appear-
ance in the esophagal region at the point where the unpaired intesti-
nal caecum receives its pair of anterior branches. Those sexual
glands which are situated most anteriorly lie below the alimentary
canal, internal to the lateral nerves, and open on the ventral surface of
the body. Farther back are commonly four or five reproductive
pouches in a single section. These lie mainly below the intestine,
but no matter what their position they all open to the surface of the
body below the lateral margins. Those lying farthest from the mid-
dle line and above the lateral nerves open ventrally between the nerve
cords and the lateral margins. Sexual products appear to be fully
mature in July.
Habitat. — Beneath stones near low water at Kukak Bay, Alaska
Peninsula (T. Kincaid).
17. AMPHIPORUS LEUCIODUS sp. nov.
pl. vii, fig. 6.
Body usually not more than 50 to 75 mm. in length, slender, elon-
gated, flattened posteriorly, not capable of great contraction. Head
narrower than parts immediately following. An inconspicuous V--
shaped furrow on dorsal surface back of head, seen only under favor-
able conditions.
This species resembles young individuals of A. exilis, with which
it is often associated. It is likewise similar in many respects to A.
lactifloreus (Johnston) Mclntosh, from which it differs widely in the
armature of the proboscis and in many other anatomical features.
Color. — Opaque white, sometimes with a pale reddish or yellowish
tinge. This color is commonly uniform throughout, though it is some-
what influenced by the internal organs which show through the body
walls. The brain is pinkish, the intestine often brownish.
Ocelli. — There are four irregular, but usually distinct groups of
minute ocelli on the anterior portion of the head. Bordering each
antero-lateral margin of the tip of the snout is an elongated cluster of
about 8 to 12 ocelli (fig. 13). Posterior to these marginal clusters,
COE
FIG. 13. Amphi-
p o rus leuciodus.
Outline of anterior
portion of the body
showing brain and
arrangement of the
ocelli. X I2-
and somewhat nearer the median line is a pair of clusters, each of
which likewise contains 8 to 12 ocelli. These posterior groups lie
immediately above the brain. Smaller and evi-
dently younger individuals have but 4 to 6 eyes in
each of the four groups. There is considerable
variation in the size of the ocelli.
Proboscis. — The proboscis sheath reaches nearly
to the posterior end of the body. The proboscis is
rather slender but may be contracted so that its di-
ameter is equal to more than half that of the body
itself. The proboscis is commonly attached to the
proboscis sheath at a point situated from ft to ^
the distance towards the posterior end of the body.
The armature is weaker than in A. exilis. The
central stylet is moderately slender and acutely
pointed. Its basis is somewhat conical in shape,
contracted slightly toward its middle portion (pi.
vn, fig. 6), and is rounded at its larger, posterior
end. It is slightly longer than the stylet itself. The number of
pouches of accessory stylets is commonly three, although there are
sometimes only two, and occasionally a specimen is found which has
four. There are usually two or three stylets in each pouch. In a few
instances, however, four and five were observed. Where three pouches
are present, as is usual, they are situated at nearly equal distances on
the circumference of the proboscis (pi. vn, fig. 6).
Cerebral sense organs, — Moderately small and situated well in front
of brain and on ventral side of head. They communicate with the
exterior by means of a pair of "canals which open on the latero-ven-
tral margins of the tip of the head.
Nephridia. — The nephridia reach forward to the brain region.
There are several pairs of efferent ducts, some of which open on the
ventral, and some on the dorsal surface of the body. In one of the
specimens sectioned the first pair of efferent ducts extended from the
internal side of the lateral nerve cords and opened directly to the
latero-ventral aspect of the body after passing on the -ventral side of
the nerve cords. A little farther back in the esophagal region were
two efferent ducts on one side and one on the other which passed above,
and externally to the nerve cords to open likewise below the lateral
margins. In the remainder of the esophagal region were three more
efferent ducts on each side. With one exception all of these passed
above the lateral nerves and opened on the dorso-lateral surfaces of the
ALASKA NEMERTEANS 53
body as in A. exilis. Another specimen had 7 efferent ducts on the
left side and 8 on the right. Of those opening on the left side the
four anterior ones passed dorsally to the nerve cord and then bent ven-
trally to open on the ventro-lateral aspect of the body ; the fifth one
opened very near the lateral margin, and the last two opened on the
dorso-lateral surface. On the right side the four anterior ducts opened
ventro-laterally, and the four posterior ones opened on the dorso-lateral
surface. The nephridia extend backward beyond the first few pairs
of reproductive glands. Here, then, we find the connecting links be-
tween the typical Amphiporus nephridium (which passes above arid
external to the lateral nerve and then bends downward to open on the
ventro-lateral aspect of the body), and the type of nephridium which
is characteristic of A., exilis and the Heteronemerteans, and which
opens directly on the dorso-lateral aspect of the body.
Sub-muscular glands are closely packed together in front of the
brain and in the brain region. They are not found farther posteriorly,
and in this respect the species differs markedly from A. exilis.
The intestinal ccecum reaches forward well toward the anterior end
of the esophagal region. The main caecal cavity, which lies directly
beneath the esophagus, sends off numerous lateral pouches above the
lateral nerves, and at its anterior end branches into lateral diver-
ticula which extend forward on each side as far as the brain re-
gion.
Reproductive glands. — The genital products mature in June in the
region of Victoria, B. C. The ova develop in sacs which are regu-
larly arranged, and extend from the posterior third of the esophagal
region to the posterior end of the body. The ovaries, in all cases
noticed, were situated immediately above the lateral nerves. In the
intestinal region they alternate with the intestinal lobes with a great
deal of regularity. Their efferent ducts occupy positions on the latero-
dorsal aspects of the body exactly corresponding to those of the poste-
rior efferent nephridial ducts. The ducts from the ovaries, however,
pierced only the longitudinal muscular layer, and did not penetrate the
circular muscular layer of the body wall. These rudimentary genital
ducts are further distinguished from the nephridial ducts by lacking a
conspicuous epithelial lining.
Habitat. — These slender whitish worms were found in great abun-
dance beneath barnacles and other growths on the piles of the wharf
at Victoria, B. C. They were found less abundantly under stones be-
tween tides at New Metlakahtla and in Glacier Bay, but were not
noticed farther northwest.
54
COE
1 8. AMPHIPORUS EXILIS sp. nov.
pi. in, fig. I ; Pi. vii, fig. 5 ; pi. xi, fig. 3.
Body extremely elongated for the genus, rounded throughout ; not
capable of the great contraction which characterizes many species of
the genus ; only moderately flattened ; posterior extremity slender.
Head usually narrower than esophagal region. The general shape of
the body resembles that of Emplectonema. In shape of body, as
well as in color and habits, this species, like the
last, recalls A. lactifloreus (Johnston) Mclntosh.
Its anatomical structures are, however, very differ-
ent, as will be seen from the following description.
Ocelli. — Exceedingly numerous and minute.
They are arranged on the head in four elongated
clusters (fig. 14). Two of these clusters lie on each
antero-lateral margin of the head, while the two
FIG. 14. Ampki- Other groups lie more posteriorly (just in front of
° the brain), and extend from near the middle line
obliquely outward and backward. Sometimes the
two posterior clusters are united in front into a
continuous V-shaped group. The number and dis-
tribution of these eyes is indicated by the following table, which shows
the numbers found in ten individuals :
sal view of head to
show arrangement
of ocelli. X 8.
No. of ocelli in
anterior clusters.
No. in posterior clusters.
Right.
Left.
Right. Left.
I.
12
ii
15 17
2.
16
17
26 28
3-
18
19
33 30
4-
23
23
32 31
5-
25
24
40 42
6.
25
26
52 46
7-
28
26
45 46
8.
35
37
54 45
9-
35
35
5i 53
10.
60
56
7i 73
In the specimens examined, therefore, the number of ocelli in the
front clusters varied from 1 1 to 60, that in the posterior clusters from
15 to 73. The average in the 10 individuals is about 28 in each an-
terior cluster, and 41 in each posterior group.
All the ocelli are minute, but nevertheless very irregular in size, some
being several times as large as others. From the ventral surface the
anterior marginal clusters only are seen.
ALASKA NEMERTEANS 55
Proboscis. — The proboscis sheath is long and slender. Even in this
elongated species it reaches within a few millimeters of the posterior end
of the body. The proboscis also is slender, though of moderately large
size. It reaches well backward in the body. The armature of the
proboscis is especially remarkable. The central stylet is moderately
slender, rather small, and rests on a moderately heavy basis. The
basis is somewhat conical in form, and rounded posteriorly (pi. vu, fig.
5). In addition to the central stylet there are usually 8 pouches of ac-
cessory stylets (pi. vu, fig. 5), though the number varies from 6 to
12. In each pouch are one or two slender stylets. Often there is a
single fully developed stylet, and a second, immature stylet in most of
the pouches. The proboscis is usually extruded when the animal is
killed.
Cerebral sense organs. — Situated far in front of brain — nearly at
end of snout when the head is contracted — and fairly well developed.
Their canals open on antero-lateral borders of tip of snout.
The blood-vascular system resembles that in other species of the
genus.
Nephridia. — The nephridial system shows peculiar deviations from
the arrangement usually found in the Metanemerteans. A pair of
main canals with numerous branches runs longitudinally above the
lateral nerve cords, as in other species. These reach forward well
toward the brain and extend posteriorly far into the intestinal region.
Their branches ramify both above and below the lateral nerve cords.
The number and position of the efferent ducts is remarkable — there are
commonly 20 or more on each side. The first is near the anterior end
of the main nephridial canal, and sometimes opens on the ventro-lateral
aspect of the body, as in other species of the genus. Back of this,
however, were counted nine other efferent ducts on each side in the
esophagal region, and at least as many more were present on each side
in the intestinal region. These ducts were mostly small and opened
on the dorsal aspect of the body, as in many Heteronemerteans. The
positions of the efferent ducts were sometimes immediately above the
lateral nerves, and sometimes but a little laterally from the proboscis
sheath. Most commonly, however, the ducts occupied positions be-
tween these two extremes, so that the majority of the nephridiopores
were situated on the dorsal surface about half way between the lateral
margin and the median line (pi. xi, fig. 3). As noted on page 52, an
approach to this condition is found in A. leuciodus. These appear to
be the only species of the genus, and indeed the only Metanemerteans,
in which the nephridiopores are situated on the dorsal surface of the
body.
56 COE
The intestinal exeunt is enormously developed. Its diverticula
reach forward on each side even to the anterior end of the brain. In
the brain region each of the cascal diverticula appears as a rounded lobe
on either side directly above, and closely approximating to, the dorsal
brain-lobe. Back of the brain there are several rather slender lobes
on each side. These lie mainly above the lateral nerves, but send off
branches below the nerves at frequent intervals. Somewhat farther
back in the esophagal region these lateral lobes join the main, un-
paired caecum, which lies immediately below the esophagus. This
csecum, throughout its course to the intestine proper, gives off nu-
merous lateral diverticula on each side, and these branch upward
above the lateral nerve cords (pi. xi, fig. 3).
Sub-muscular glands, — These occur abundantly in the brain region,
and are still more closely packed together in the anterior portion of
the esophagal region. They occupy positions, as in other species, in
the connective tissues beneath the musculature in the latero-ventral
regions of the body. Each gland is composed of a number of cells,
and each has a duct leading through the layers of the body wall to the
exterior on the latero-ventral aspect. A much smaller number open
on the dorsal surface. In the posterior end of the esophagal region
these glands have almost entirely disappeared, although a few are met
with in the anterior portion of the intestinal region.
Color, — The color of the individuals of this species is commonly a
homogeneous, opaque white, very pale flesh color, or pale yellowish-
white. This color is continuous throughout the length of the body,
both above and below, except where the internal organs show through.
The smaller specimens are pale, while the larger ones almost always
exhibit a brownish color, which indicates the position of the intestine.
Sometimes minute reddish-brown specks are distributed over the dorsal
surface. Occasionally a worm of this species is met with in which
the intestinal lobes are pale orange. The brain is plainly distinguish-
able in the living worm because of its pinkish coloration.
Habitat. — Amphiporus exilis is one of the most common, as well
as the most widely distributed species of nemerteans met with on the
expedition. It occurred abundantly at nearly all the collecting stations
from Victoria, B.C., to Dutch Harbor, Unalaska. The worms are
restless and are often seen crawling over stones between tides. They
live among barnacles, mussels, etc., from low water well up to high
water mark, and are found abundantly under stones in almost all sorts
of locations. The species is especially hardy.
ALASKA NEMERTEANS 57
Tetrastemma Ehrenberg.
Symbolae Physicae, Berlin, 1831.
This genus includes a group of very small, slender worms, seldom
more than 20 to 30 mm. long, with slightly flattened body, and usually
with four well-developed ocelli, which form a quadrangle on the head.
In a few species (cf. T. aberrans) these ocelli are each replaced by a
group of two or three smaller ones, and in other species ocelli are
wanting entirely. The anatomical structures are very similar to those
of Amphiporus, and the distinctions between the two genera are not
clearly defined. The mouth and proboscis open together. The cere-
bral sense organs lie close in front of the brain. The proboscis sheath
extends to the posterior end of the body, and the proboscis is well de-
veloped, armed with central stylet and pouches with accessory stylets,
and usually provided with ten nerves.
Only three species of the genus were met with on the expedition,
although it seems probable that a number of other forms of these minute
worms will be found later.
19. TETRASTEMMA BICOLOR sp. nov.
pi. i, fig. 6.
Body moderately slender, rounded both above and below ; much
larger and longer than most species of the genus, sometimes becoming
50 to 60 mm. in length in extension.
Color. — This species is bright brownish-red or orange the whole
length of the dorsal surface ; the whole ventral surface is pale gray or
whitish. The anterior border and lateral margins of the head, as well
as the lateral margins of the body for a short distance back of the head,
have the same whitish color as the ventral surface. A narrow, median,
white stripe, sharply marked off from the reddish color of the dorsal
surface, extends from the white, anterior border of the head nearly to
the posterior end of the body. Posteriorly the stripe becomes more
irregular and is usually lost near the posterior extremity.
Ocelli. — Four, rather large, rounded, arranged nearly in the form
of a square.
Proboscis. — Proboscis sheath and proboscis as in typical species of
the genus. Proboscis provided with a moderately heavy central stylet
about .075 mm. in length. Basis of central stylet somewhat conical
in shape, swollen behind, and about \y2 times as long as the stylet it-
self. There are two pouches of accessory stylets, with usually three
or four stylets in each pouch.
COE
The blood is dark red, and the blood vessels may be traced in the
living worm the whole length of the body.
Habitat. — The species was found only at Kadiak, where it was
dredged in about three fathoms. It slightly resembles some varieties
of T. vermiculus Quatrefages, but the longitudinal bands of dark
pigment between the two ocelli of the same side are lacking, and the
median white line is sharply demarkated, so that the two species are
undoubtedly specifically distinct.
20. TETRASTEMMA ABERRANS sp. nov.
This is a minute Nemertean, the specimens found not exceeding 1 2
mm. in length in greatest extension. Body moderately slender, slightly
flattened. A pair of slight vertical slits on sides of head.
Color. — Pale yellow throughout, both above and below.
Ocelli. — Of moderate size, or rather small, arranged in four groups
which form a rectangle, as do the single eyes of typical species of the
genus. Each of the four groups is composed of three to five ocelli of
variable size. The two anterior groups lie well toward the tip of the
snout, while the two posterior groups lie above or slightly in front of
the brain (fig. 15). The appearance of the eyes
is such as to give at once an impression that the
multiple nature of each group has arisen from the
fragmentation of single ocelli. This I consider to
be the case. At least one other species of Tetra-
stemma ( T. cruciatum Burger ) is known in
which the four ocelli are double, and I have
often noticed other species of the genus — notably
the fresh water T. rubrum (Leidy) — which
showed almost conclusively that one or more of
the six or seven eyes present had been derived
FIG. 15. Tetrastem- from a Spiitting of the primary ocelli. In one
ma aberrans. Dorsal . it , , , , . , ,
, . instance the fragmentation had been carried so far
view of anterior por-
tion of body, showing that no fewer than 2O to 3° pigment spots were
outline of brain and ar- present.
rangement of the four Proboscis sheath and proboscis as in other
groups of ocelli. X25- Species of the genus. Mouth and proboscis open
together ; proboscis sheath reaches the posterior end of the body. Pro-
boscis provided with a rather slender central stylet and basis ; the two
accessory stylet pouches each with two or three stylets of typical form.
The proboscis possesses twelve nerves, and in this respect again shows
a departure from the typical Tetrastemma, where there are usually
but ten proboscidial nerves.
ALASKA NEMERTEANS 59
Nephridia. — The nephridial canals are short, and reach forward to
the brain. Anteriorly each nephridium lies above the lateral nerve,
but farther back the branches ramify both above and below the nerve.
Near the anterior end of each of the main canals a large efferent duct
passes outside of the lateral nerve, and bends downward and outward
to open to the exterior on the lateral margin of the body. In one in-
stance a double efferent canal was observed.
The head is provided with large cephalic glands which open on the
tip of the snout. The cerebral sense organs are large. They lie
mainly in front of the brain, although their posterior ends extend
backward beside and lateral to the brain lobes. Each connects with
the exterior by a canal which runs anteriorly to open on the lateral
margin of the head.
A broad intestinal ccecum reaches into the anterior fourth of the
esophagal region. It lies below the esophagus, and sends off a few
wide lobes on each side, but these reach dorsally only a little above the
lateral nerve. The esophagus opens into the caecum far behind the
anterior sexual glands, and nearly as far back as the middle of the
body. The reproductive pouches lie both above and below the intes-
tine.
As will be seen from the above description, this species agrees
closely with the typical species of Tetrastemma in size, general appear-
ance, and in the details of the internal anatomy. It differs from
known species of the genus only in the fragmented nature of its eyes
and in the number of nerves in the proboscis. It seems extremely
probable, however, that when more of the described species have been
examined in this regard, some of them will be found to contain more
or less than ten proboscidial nerves. The eyes certainly resemble
those of Tetrastemma more closely than they do those of any de-
scribed species of Amphtporus.
Habitat. — Found among hydroids in about four fathoms in Glacier
Bay, and between tides at Orca, Prince William Sound. Not com-
mon.
21. TETRASTEMMA C^CUM sp. nov.
A small species which I shall refer provisionally to this genus was
found by Ritter in considerable numbers at Kadiak. The species is
especially remarkable, and quite aberrant from most other species of
Tetrastemma, both in lacking ocelli and in being hermaphroditic.
The body is rounded and of almost even diameter throughout. The
head has a pair of inconspicuous, oblique, lateral furrows.
60 COE
Color. — The worms are very pale, whitish or pale yellowish in color,
with dark intestinal lobes.
Size. — Very small, not usually more than 5 to 10 mm. long and
0.5 to i mm. in diameter when sexually mature.
Ocelli. — Wanting.
Proboscis. — The proboscis sheath reaches nearly to the end of the
body. The proboscis is remarkable for its enormous size as com-
pared with the size of the body — when everted its diameter is practically
equal to that of the body itself, although it is then comparatively short.
Its armature consists of a central stylet with rather slender basis, and
of two pouches of accessory stylets. The basis of the central stylet
is slightly swollen and somewhat sharply truncated posteriorly. Each
pouch contains 2 or 3 long, slender and delicate accessory stylets.
In cross section of the proboscis the inner and outer circular muscu-
lar layers appear as usual. The intervening layer of longitudinal mus-
cles, however, is divided into two secondary layers separated by a thick
sheet of nerves and connective tissue. The nerves appear to be more
or less confluent, and not separated into a definite number of longitudi-
nal cords as in most other species of Tetrastemma. This appearance
may be partially due, however, to the action of the formalin in which
the worms were preserved.
Sense Organs. — The cerebral sense organs are very well developed,
and unusually voluminous as compared with the other organs of the
head. They lie in front of the brain, and extend posteriorly on the
ventral side of the brain lobes as far as the ventral commissure. Each
sense organ communicates with the exterior by a small canal passing
obliquely forward to open on the lateral margin of the head.
The brain is of small diameter, but its extent antero-posteriorly is
comparatively great.
Reproductive organs. — The individuals are hermaphroditic, and
probably to some extent protandric although there is considerable varia-
tion in this respect. One of the individuals sectioned was filled with
ripe spermaries only, but all the others possessed enormous ova, with
the spermaries disposed irregularly. The mature ova were fully two-
thirds the diameter of the body, and hence were arranged at irregular
intervals in a single row. Where the ova were mature the spermaries
were smaller and contained much fewer spermatozoa than in those in-
dividuals which were without large ova. Many of the spermaries had
ducts which pierced the muscular layers to reach the dorso-lateral sur-
faces of the body. In some cases where these ducts were fully formed,
and the spermatozoa therefore mature, the ova in the same individual
were but half grown. The sexual products were mature in July.
ALASKA NEMERTEANS 6l
Taeniosoma Stimpson.
Tceniosoma STIMPSON, Proc. Acad. Nat. Sci. Philadelphia, p. 162, 1857.
Polio, DELLE CHIAJE, Mem. sulla storia e notomia degli anamali senza
vertebre del regno di Napoli, Naples, 1823-28.
Eupolia HUBRECHT, Report of Challenger Exped. Zool., xix, 1887.
Eupolia BURGER, Fauna u. Flora von Neapel, Monogr. 22, p. 598, 1895.
For the reasons given on page 4, it seems absolutely necessary to
adopt for this genus the name given by Stimpson in 1857 rather than
accept that of Hubrecht of 30 years later, even though most European
writers have ignored Stimpson's brief, but careful, diagnoses.
The species belonging to this genus show a remarkable specific vari-
ation in the general shape and size of the body. Some are character-
ized by extremely long, slender, flattened, and much twisted bodies,
while others are short, thick, and cylindrical. In all the species, how-
ever, the head in life is rounded in front and is sharply marked off
from the parts immediately following by lateral constrictions. Hori-
zontal furrows are wanting, but small, oblique or transverse grooves
may be present on the head. In strong contraction the esophagal
region becomes greatly swollen, the head is drawn in (pi. n, fig. 4),
so that the anterior end of the body is large and shortly truncated.
Proboscis sheath and proboscis short, seldom reaching more than
one-third the length of body. Proboscis opening subterminal, minute.
Mouth a small round opening on the ventral surface immediately be-
hind the ganglia.
Muscular layers of body composed of a thick outer longitudinal, a
circular, and a less thickened inner longitudinal layer. Outside the
muscular layers is a well developed cutis, composed of a thick inner
layer of connective tissue, and an outer layer of glandular tissue. The
external epithelium is thin, as compared with the other layers of the
body, though the fibrous layer separating it from the cutis is well de-
veloped. The musculature of the proboscis consists of an inner longi-
tudinal, and an outer circular muscular layer. Consequently there can
be no muscular crosses.
The cephalic glands are enormously developed. They stretch back-
ward on all sides beyond the brain, and even reach some distance into
the esophagal region.
The lateral nerves lie immediately outside the circular muscular
layer. There are three longitudinal blood vessels.
Ocelli are usually present in great numbers, though very small.
The worms are sluggish in their habits, are unable to swim, and
usually show great irregularities in the diameter of the body. They
62 COE
are prone to twist themselves in sharp coils, or in knots, and often lie
in lumps. They are usually capable of contracting and extending their
bodies to a remarkable degree.
22. TVENIOSOMA PRINCEPS sp. nov.
pi. n, figs. 3, 4.
Body of very large size, long, thick, largest in the esophagal region,
cylindrical in anterior portion, flattened on ventral surface posteriorly ;
in contraction nearly cylindrical throughout. Head sharply marked off
from body in extension, rounded in front, flattened dorso-ventrally ; in
contraction drawn almost entirely into the parts immediately following,
so that the anterior portion of the body is greatly swollen and sharply
truncated in front. In contraction the anterior end of the body is
thrown into massive folds, and the whole body is remarkably short
and thick (pi. n, fig. 4). A pair of inconspicuous, oblique grooves
lies on the antero-ventral surfaces of the head ; into these the canals
leading from the cerebral sense organs open. The esophagal region is
scarcely more than one-twelfth the length of the body. In alcoholic
specimens there is usually a median ridge on the dorsal surface run-
ning the length of the body, except in the head and esophagal regions.
Color. — The dorsal surface is deep ochre yellow, sometimes inclin-
ing to orange, and sometimes to brownish, and thickly strewn with
minute irregular dark red spots. The reddish markings are most con-
spicuous near the anterior end of the body, and in the dorsal, median
line. In many places a large number of the reddish dots become con-
fluent, and form an irregular patch of deeper color. These patches
commonly occur as broken longitudinal lines. Such lines are most
abundant on the middle of the dorsal surface where they form a median
longitudinal band of reddish-brown. The ventral surface is paler and
the reddish markings are wanting. Along the median line the color is
brighter yellow than elsewhere on the ventral surface. This is appar-
ently due to the absence in this position of the opaque intestinal lobes.
The ventral surface often exhibits a greenish tinge to the yellow ground
color. The posterior extremity is pointed and much paler than the
rest of the body.
Size. — T. princeps grows to a size greater than has previously
been described for any species of the genus. The individuals found
were from half a meter to two meters in length when extended ; when
contracted, but a small fraction of this length, and proportionately
thick (pi. n, fig. 4). After long standing in alcohol a section of the
ALASKA NEMERTEANS 63
body of one specimen still measured 15 x 18 mm. It is one of the
largest Nemerteans of the coast.
Ocelli. — There are many minute ocelli arranged in an irregular
group on each side of the tip of the head. The number of such ocelli
is commonly 40 or more in each of the two groups.
The mouth is a small rounded pore, and is situated several milli-
meters back from the tip of the head in large individuals. The pro-
boscis-pore is also minute, and is situated subterminally as usual.
Proboscis. — The proboscis sheath reaches some distance into the
esophagal region, but is very short when compared with the length of
the body. The proboscis is short and weak. Its muscular and epi-
thelial layers are as in other species of the genus.
The cephalic glands are enormously developed. They occupy a
large proportion of the area in the anterior portion of the head, sur-
round the brain on all sides, and extend still further backwards into
the anterior end of the esophagal region. Here they lie in the outer
longitudinal muscular layer around the whole circumference of the body.
Body 'walls. — The outer longitudinal muscular layer about equals in
thickness the other two muscular layers combined. In the intestinal
region the inner longitudinal muscular layer becomes extremely thin on
the lateral aspects of the body, and is much reduced dorsally. It is only
on the ventral side that this layer retains its comparative thickness.
The cutis is thinner than in most species of the genus, and especially
is this true of its inner, fibrous layer. This layer is, throughout most
of the body, reduced nearly to the condition of a thin membrane. It
is commonly not much thicker than the muscular layer beneath the
body epithelium. The epithelium itself is thin in comparison with the
massive muscular layers of the body.
The blood lacunce in the head lie directly above the brain, as in other
species of the genus. They are, however, remarkably large, and
are crossed in various directions by numerous bundles of muscular
tissue, which tend to subdivide the lacunae into numerous smaller
spaces. The longitudinal blood vessels are as usual. The dorsal vessel
passes out of the proboscis sheath early in its course.
Nephridia. — Situated in anterior and middle portions of esophagal
region. Several efferent ducts on each side. These are small in
diameter and pass above the nerve cords, opening on the lateral aspects
of the body dorsal to the lateral margins.
Cerebral sense organs. — The dorsal lobes of the brain greatly exceed
the ventral lobes in size, and lie somewhat lateral as well as above
them. The cerebral sense organs are voluminous. They extend for-
64 COE
ward on each side, external and ventral to the dorsal brain lobes, nearly
as far as the ventral commissure. Here, at the anterior extremity of
each sense organ, a canal passes obliquely downward and outward to
open into a shallow oblique furrow on the ventro-lateral aspect of the
head. The sense organs are closely united with the posterior ends of the
dorsal brain lobes. In the middle region of the brain, the anterior ends
of sense organs are triangular in section, and lie external and between
the dorsal and ventral ganglia. Farther back a glandular appendage
of the sense organ pushes itself in between the internal faces of the
brain lobes. This appendage fuses with the ventral portion of the
sense organ more posteriorly. The brain lies deeply buried in the
tissues of the head and nearly in the median line, but the lateral nerves
while still in the region of the cephalic sense organs bend sharply
outward, and occupy throughout the remainder of their course posi-
tions immediately external to the circular muscular layer of the body
wall.
The genital products were nearly mature in July. The oviducts
were in many instances preformed, and opened on the dorso-lateral
aspects of the body.
Habitat. — Only four individuals of this species came under my ob-
servation. One was collected at Cape Fox (Kincaid), two at
Yakutat, and the fourth at Orca in Prince William Sound. All were
found under stones in rather hard mud at low water. The individuals
contract strongly when handled, throwing the surface of the body into
wrinkles, and often coil the posterior portion of the body into a close
spiral, much as does Cephalothrix.
Lineus Sowerby.
The British Miscellany, London, p. 15, 1806.
Representatives of this genus are characterized by a slender, some-
times thread-like body, usually rounded throughout. The body is
commonly twisted and coiled into an irregular mass. The movements
are sluggish. The animals creep over objects and readily move about
on the surface of the water, but they are unable to swim. The body
is extremely contractile ; the head is often slightly wider than the body,
of oval shape, and is usually provided with numerous minute ocelli,
often arranged in a single row on each side of the head. A caudal
papilla or cirrus, a diagonal muscular layer, and neurochord cells are
all wanting. The proboscis sheath is often short in comparison with
the length of the body.
ALASKA NEMERTEANS
23. LINEUS VIRIDIS (Fabr.) Johnston.
Planaria viridis O. FABRICIUS in O. F. Miiller, Zool. Dan. Prod., 1776 ; Fauna
Groenlandica, p. 324, 1780.
Planaria gesserensis MULLER, Zool. Danica, u, p. 32, 1788.
Ntmertes obscura DESOR, Boston Journ. Nat. Hist., vi, pp. i to 12, 1848.
Lineus viridis JOHNSTON, Catalogue British Non-parasitical Worms, pp. 27,
296, London, 1865.
As stated by Verrill,1 there seems little doubt that the description of
this species sent by Fabricius to Miiller and published by him in the
Zoologica Danica, is entitled to retain priority in nomenclature instead
of Muller's gesserensis of later date, which has been adopted by most
recent European writers.
Characteristic individuals of this species were found under stones at
low water at New Metlakahtla, on Annette Island.
Body moderately slender, rounded throughout, but slightly flattened
posteriorly ; head slightly wider than the parts immediately following ;
cephalic slits long and deep, with pale margins above and below,
reaching anteriorly close to the proboscis pore. The anterior end of
the mouth does not reach quite so far forward as the posterior end of
the cephalic slits. Length usually 100 to 200 mm.
On each side of the head in front of the brain and
close to the lateral borders is a single row of minute
ocelli (fig. 16). The number of these is commonly
from four to six on each side, though some individuals
have as many as eight, and very young specimens but
a single pair.
Color. — The Alaska specimens were dusky or brown-
ish green, becoming dark brown anteriorly, and com-
monly paler on the ventral surface, especially posteriorly.
The head is very pale on lateral margins and in front.
The brain is large, reddish, and shows distinctly
through the pigment of the body. Cerebral sense or- line of anterior
gans paler but easily distinguished in life, with con- portion of body
spicuous canals leading to the posterior ends of the of Lincus viri-
cephalic slits (fig. 16). dfs showing ar-
Habitat. — This species, besides being found in Alas-
ocelli, and posi-
ka, is widely distributed in northern waters. On the tion of brain, ce-
east coast of America it occurs from Long Island Sound phalic sense or-
to Greenland. It is found on nearly all the coasts of gans and mouth,
northern Europe. It has also been found in the Medi- X 8.
'Trans. Connecticut Acad., vm, p. 421, 1892.
66 COE
terranean, though it is there comparatively rare and small. It is usu-
ally found between tides under stones in muddy localities.
24. LINEUS TORQUATUS sp. nov.
pi. v, figs. 8, 9.
Body rather thick and stout for the genus, somewhat flattened
throughout, but especially posteriorly and on the ventral surface.
Head short, pointed in front, somewhat narrowed behind, slightly flat-
tened. A slight annular constriction commonly marks off the head
region from that immediately following. Sometimes this constriction
is very conspicuous, and the head much narrower behind. Esophagal
region rounded above, flattened below. Intestinal region commonly
well flattened but without narrow margins. Posterior end slender.
Cephalic furrows rather short. In alcohol or formalin they join
the terminal proboscis pore in front, but in life they are separated from it.
Ocelli. — Absent, at least in mature individuals. Mouth a minute
pore or a large slit, according to state of contraction ; situated a little
behind posterior end of cephalic furrows.
Color. — The color is usually dark, reddish-brown, chocolate, or pur-
ple above ; paler and commonly more reddish beneath. The dorsal
surface is often flecked with irregular minute, inconspicuous whitish
specks. A narrow transverse white band passes across the dorsal sur-
face at the posterior ends of the cephalic furrows. This characteristic
marking reaches only to the lateral edges, and is not seen from the ven-
tral surface. The cephalic furrows are sometimes, but not always,
bordered above and below by a narrow band of white. Furthermore
a minute white spot occupies the region of the proboscis pore. The
white borders of the cephalic furrows commonly connect this white
spot with the white transverse dorsal band on the posterior portion of
the head. On the extreme tip of the snout — in the white area around
the proboscis pore — a pair of small pigment spots is sometimes pres-
ent, one on each side of the proboscis pore.
Size. — Length usually 200 to 400 mm. in extension, width about
5 mm.
Proboscis. — The proboscis is moderately slender and of medium
size. It is without color. The inner longitudinal muscular layer is
almost entirely wanting ; the crosses between the circular layers are
distinct, and the nervous plexus is exceptionally well developed. A
single pair of large nerves enters the proboscis at its point of attach-
ment at its anterior end. They originate from the ventral commissure
of the brain near the ventral ganglia, enter the proboscis from the ven-
ALASKA NEMERTEANS 67
tral side, pass backward for some distance on the right and left sides
respectively, and later spread out into a cylindrical plexus immediately
internal to the muscular layer, and separated from the inner epithelium
only by a few longitudinal muscular and connective tissue fibers.
Glands. — The cephalic glands are very well developed, and occupy
a large portion of the region of the head in front of the brain, both
above and below the rhynchodaeum. They do not extend posteriorly
quite so far as the brain.
Nerves. — The nervous system is far more easily made out than in
any of the related species with which I am acquainted. The individ-
ual nerves are large and are sharply defined in all cases. The nerves
extending from the brain toward the tip of the snout are numerous
and are all of large size. The esophagal nerves are also surprisingly
large and quite conspicuous. They have several transverse connecting
branches after their origin from the ventral ganglia. The most pos-
terior of these connecting branches occurs just in front of the mouth.
In the mouth region they communicate in several instances with the
lateral nerves by means of branches which pass through the nervous
plexus outside the circular muscular layer. These branches from the
lateral nerves follow the nervous plexus to the vicinity of the esophagal
nerves, which they join by passing directly through the circular mus-
cular layer. I do not know that such an anastomosis between the
lateral nerve cords and the esophagal nerves has been previously noted
in any species.
The median dorsal nerve, situated just outside the circular muscular
layer, is unusually conspicuous, and throughout a considerable portion of
the intestinal region is supplemented by a second median nerve lying
directly beneath the first, but in the midst of the internal longitudinal
muscular layer. Branches connect these two nerves at frequent inter-
vals.
Cerebral sense organs. — Voluminous. The canals leading to the
exterior open on the summit of a broad papilla situated at the posterior,
widened end of each of the cephalic slits.
Nephridia. — The nephridial canals are of much greater diameter
than I have observed in any other Nemertean. The canal on either
side is, throughout a portion of its length, equal to the lateral nerve
cord in cross section. The nephridia extend through the anterior half
of the esophagal region. The main tubule has a few very large
branches, and these lie in the walls of the blood spaces about the esoph-
agus. Anteriorly the branches lie mainly dorsal to the lateral
nerves, but towards the posterior ends of the nephridia the branches
68 COE
ramify more towards the ventral side of the esophagus. There is a sin-
gle pair of efferent ducts which are of enormous size as compared with
those of other species. These ducts are situated at about two-thirds the
distance towards the posterior ends of the nephridia, and open on the
dorso-lateral aspects of the body as usual.
Habitat. — This species is common in mud and under stones in
muddy localities at Orca and Virgin Bay in Prince William Sound,
but was not met with elsewhere on the expedition. The individuals
are hardy and of sluggish movements. They do not break up nor
contract excessively when thrown into killing fluid, and the proboscis
is not usually everted when the animal is killed. Some, especially
the smaller ones, when preserved are nearly cylindrical, but most indi-
viduals are flattened ventrally. The color is fairly permanent in for-
malin, and even in alcohol for some months.
Micrura Ehrenberg.
Symbolic Physicae, Berlin, 1831.
This genus includes mostly moderately small, slender forms, gener-
ally less rounded posteriorly, and of rather more active habits than
Lineus. Its most marked distinction from the latter genus is that the
posterior extremity of the body is provided with a slender, usually
colorless, muscular caudal cirrus. This is formed of a continuation
of the muscular tissues and integument beyond the posterior end of the
alimentary canal.
The species of Micrura are generally, though not always, more
brightly colored and have more distinct markings than those of
Lineus. The vast majority of the species are provided with numerous
ocelli, though some are blind. The head is slender, and not distinctly
separated from the rest of the body. The lateral faces of the body are
not provided with thin edges as in Cerebratulus, the intestinal region
is not so much flattened, neurochord cells are not present in those
species which have been studied, and none of the species are able to
swim as do all species of Cerebratulus. The mouth is usually smaller
than in Cerebratulus, and the intestinal lobes are not so deep.
The proboscis is usually slender and comparatively weak ; the pro-
boscis sheath is sometimes considerably shorter than the body.
25. MICRURA VERRILLI sp. nov.
Pl. V, figs. I, 2, 3.
Body moderately elongated, widest anteriorly, tapering to an acute
anterior extremity ; much more slender posteriorly. Ventral surface
ALASKA NEMERTEANS 69
flattened ; dorsal surface rounded. Head narrow in front. Cephalic
furrows long and deep ; at their posterior ends each is met by an ob-
lique, shallow depression above and below. Proboscis-pore exactly
terminal ; anterior ends of cephalic furrows well separated from pro-
boscis pore. Mouth small, situated as far back as posterior ends of
cephalic furrows. Caudal cirrus small, slender, easily broken off.
Color. — This is one of the handsomest and most striking of all the
described species of Nemerteans. The color of the dorsal surface is
deep purple or wine-color ; that of the ventral surface is purest white.
On the dorsal surface are usually 15 to 40 very sharp, pure white,
transverse bands or lines connecting with the white color of the ven-
tral surface. These transverse bands are nearly as narrow as pencil
marks, and are situated at fairly regular intervals throughout the length
of the body. They are sometimes more or less interrupted, but
always sharp and distinct. On the dorsal surface of the tip of the
snout is a small triangular marking, always very conspicuous because
of its bright orange color. Following behind this is a narrow trans-
verse white band, followed by broad purple and narrow white bands
in succession. The cephalic furrows lie within the white color of the
ventral surface, and are bordered above with a very narrow margin of
white. They reach posteriorly as far as the second white band.
After preservation in formalin or alcohol the worms are strongly
rounded below as well as above, and the color of the dorsal surface
changes from purple to red, while the orange spot on the tip of the
snout disappears.
Ocelli. — None were found.
Proboscis. — Colorless, and shorter than in many species of the
genus. Its microscopic structure shows distinctly the three muscular
layers and the muscular crosses characteristic of the family. The
inner circular muscular layer is very thin. The nervous plexus inside
the circular layer is usually well developed.
Nervous system. — The dorsal ganglion of the brain is divided poste-
riorly into two distinct lobes, of which the smaller, dorsal lobe ends
shortly, while the larger, ventral lobe continues into the cerebral sense
organ. Each of these sense organs is of about the same size as one of
the ventral ganglia, but considerably smaller than either dorsal ganglion.
The duct lying on the external border of each of the pair of cerebral
sense organs communicates, as usual, with the enlarged posterior end of
the cephalic furrow by means of a wide duct opening directly outwards.
The dorsal median nerve, lying just outside the circular muscular layer,
is remarkably conspicuous throughout the whole length of the body.
70 COE
A section through the esophagal region shows that the outer longi-
tudinal muscular layer of the body wall is particularly thick, and that
the esophagus is unusually small. The other layers are as in most
species of the genus.
The lateral blood lacunae break up in the esophagal region into
numerous thin- walled vessels, or spaces, which surround the lateral and
ventral walls of the esophagus.
Nephridia. — The nephridia extend through the anterior 2/i of the
esophagal region, as a single profusely branched tubule on each side.
All the branches of the main canal are small, and lie in close connec-
tion with the walls of the blood spaces about the esophagus. The
efferent ducts are very numerous, but minute. They open on the
dorso-lateral aspects of the body, usually not far above the lateral mar-
gins. In a single instance one of the efferent ducts was found to
open beneath the lateral margin, but this must be looked upon as ab-
normal. In several instances two efferent ducts lay close together on
the same side, but in such cases one originated much nearer the ven-
tral side of the esophagus than did the other. The actual number of
efferent ducts counted in one medium-sized individual was 17 on one
side, and 24 on the other.
Reproductive organs. — The pouches containing the sexual products
when mature become so voluminous as to occupy more than half the
entire space within the body walls. In July, when the genital prod-
ucts are fully ripe, the ducts leading to the exterior are completely
formed some little time before the elements are discharged. Each
pouch has a single duct opening into a funnel-shaped depression through
the epithelium on the dorso-lateral aspect of the body. The opening
into the cavity of the ovary is on the surface of a broad papilla formed
of long columnar cells, a portion of which bend inward to guard the
opening. The ducts often have a somewhat tortuous course, and are
likely to broaden out considerably in passing through the circular mus-
cular layer.
Size. — The length of the largest individual observed was about 300
mm., and its width 6 mm.
The species is named in honor of Prof. A. E. Verrill, of Yale
University, who, more than any one else, has helped to bring the
American species of Nemerteans into orderly arrangement.
Stimpson l briefly describes from Bering Strait a species of Cere-
bratulus ( C. impressus = Micrura impressa) which bears a superficial
resemblance to the above. Stimpson's species, however, was flattened,
1Proc. Philadelphia Acad. Nat. Sci., p. 160, 1857.
ALASKA NEMERTEANS 7 1
of a dusky gray color above, with narrow, colorless, transverse lines,
and with a flesh-colored subtruncate head, much narrower than the body.
The same author describes a somewhat similar species ( C. bellus =
Micrura bella) from Yezo Island, Japan. This had 10 bluish-
white, narrow transverse lines across the ashy-gray dorsal surface, and
a white ventral surface. The head was short, and of a vermilion color.
Both these species should be referred to the genus Micrura as here
defined.
Habitat. — This species was found only at Virgin Bay, in Prince
William Sound. Here it was not uncommon under stones at low
water, and was frequently met with in the parchment like tubes of
Carinella capistrata, with which it was associated.
26. MICRURA ALASKENSIS sp. nov.
pi. iv, fig. 2 ; pi. xin, fig. i.
Body long and slender, rounded in esophagal region, flattened
throughout the whole intestinal region. Head remarkably slender,
elongated, and sharply pointed ; cephalic furrows correspondingly
long, but their anterior ends do not reach the proboscis pore. Mouth
small, and well back from tip of snout ; its anterior end reaches about
as far forward as posterior ends of cephalic furrows.
No ocelli are present. The brain is reddish in color, and shows
conspicuously through the tissues external to it.
The esophagal region is well rounded, and narrower than the suc-
ceeding portion of the body. The intestinal region is much flattened
both above and below ; its lateral margins are rounded, however.
After preservation in alcohol the intestinal region is flattened or even
hollowed ventrally, while the dorsal surface is very convex. Posteri-
orly the body tapers gradually, and at its pointed extremity an unusu-
ally long caudal cirrus is present. This is quite colorless, and contracts
after preservation to but a fraction of its original size and length.
Color. — Two color varieties were met with. Most commonly the
general color was a salmon or flesh-color. The esophagal region was
pale salmon with tinges of brighter red, becoming lighter anteriorly ;
the head pale or nearly colorless ; the brain region distinctly red ; the
intestinal region pale salmon, with much more deeply colored intestinal
lobes. Running the whole length of the ventral side of the body —
from near the mouth to the caudal cirrus — is a characteristic, narrow,
cream or flesh-colored stripe in the median line. This stripe is con-
spicuous only in the intestinal region, though it may be traced forward
to the mouth, as stated. In alcoholic specimens it can still be detected
72 COE
in the intestinal region, though the natural color of the body has mostly
disappeared. A similar, though much less conspicuously marked, me-
dian stripe occurs on the dorsal surface.
The second color variety had chestnut brown intestinal lobes, brown-
ish esophagal region, slightly paler below ; snout and margins of head
colorless ; brain red. The paler, ventral, median stripe was even more
conspicuous than in the other variety.
Serial sections show that, while the cephalic furrows are unusually
long, yet they are not really as deep as in many related species. The
brain is very large ; the posterior end of each dorsal ganglion is bi-
lobed, the dorsal lobe ending freely, while the ventral lobe continues
directly into the cerebral sense organ. The canals from the sense
organs open into the very posterior ends of the cephalic furrows. The
buccal, or esophagal, nerves are larger than in most species.
Accessory buccal glands. — On each side of the mouth, and extend-
ing a short distance into the esophagal region, is a series of peculiar
glands — accessory buccal glands, they may be called. These glands
are similar in their nature to, and apparently supplement, the ordinary
buccal glands which line the mouth cavity. In the present instance,
however, these accessory glands lie imbedded in the outer longitudinal
muscular layer ventral to the lateral nerves. Here they greatly en-
croach upon the domain of the muscles, and occupy a large portion of
the space between the circular muscular layer and the cutis (pi. xm,
fig. i ) . The gland cells are large, and are distended with a clear,
granular secretion. These accessory glands appear to have originated
from ordinary buccal glands which have passed outward and taken up
a position outside the two inner muscular layers (pi. xm, fig. i).
They are arranged in clusters, and discharge their contents by irregular
ducts leading through the two inner muscular layers to the epithelium
of the buccal cavity and adjacent esophagal wall. A short distance
back of the mouth the two lateral series of these accessory buccal
glands unite beneath the esophagus, and are not found further poste-
riorly. The ordinary buccal glands are present as in related species.
Alimentary canal. — The esophagus is large in comparison with the
thickness of the body wall. The histological difference between the
epithelium lining its anterior portion and that of its posterior half is
much more marked than in most related species. The delicate layer
of circular and longitudinal muscular fibers which surrounds the epi-
thelial lining of the esophagus in most of the Heteronemerteans be-
comes remarkably developed in this species. At the very posterior
end of the esophagus — just anterior to the first intestinal pouches —
ALASKA NEMERTEANS
73
the circular muscles of the esophagus increase so greatly in number
that they form a most conspicuous layer. In the region of its maximum
development this layer becomes nearly half as thick as the circular
layer of the body walls in the same section. In no other species of
the Lineidce has this muscle been found of even approximately this
thickness. Its fibers connect in part with the circular layer of the body
walls, and to a lesser degree with the circular muscles of the proboscis
sheath. But few fibers lie on the dorsal wall of the esophagus, so
that this organ is largely bound up with the proboscis sheath in a con-
tinuous layer of muscles, and one cannot fail to see the striking resem-
blance between this circular layer and the inner circular muscles that are
so highly developed in precisely the same region in Carinoma (p. 22).
Body ivalls, — The external longitudinal muscular layer of the body
walls is especially weak when compared with the same layer in related
species. This is partly shown by the fact that the cutis glands extend
Y^ or more of the distance from the exterior to the circular muscular
layer. Nearer the intestinal region the muscular layers are thicker,
and the esophagus occupies correspondingly less space.
The three longitudinal blood vessels are very conspicuous through-
out their length. The dorsal vessel leaves the proboscis sheath near
the posterior ends of the nephridia, or at about ^3 the distance towards
the posterior end of the esophagal region.
Nephridia. — The nephridial system consists of a pair of unusually
large, longitudinal canals, which lie in the dorsal walls of the lateral
blood lacunas beside the esophagus. The nephridial canals send off
very few branches, except near their anterior ends, where they divide
into smaller branches. The main canals extend through about the
middle third of the esophagal region. Each of the pair of longitudinal
canals terminates posteriorly in a single, remarkably large efferent
duct which opens on the dorso-lateral aspect of the body. The ducts
sometimes lie exactly opposite, but in other specimens one lies some
distance farther back than the other. In such cases, of course, one of
the nephridia extends a corresponding distance farther posteriorly than
the other. Each of the efferent ducts spreads out as it passes through
the circular muscular layer into a broad sieve-like or filter-like struc-
ture with scores of small, nucleated cells. From this point a straight
and comparatively narrow duct leads directly to the surface.
Reproductive glands. — The sexual elements are fully mature in the
month of June, and the genital ducts at this time are fully formed.
In the females each duct communicates directly with the cavity of the
ovary by a funnel-shaped opening, and a similar funnel-shaped open-
74 COE
ing lies at the outer end of the oviduct on the dorso-lateral aspect of
the body. The bodies of many of these worms were so distended with
sexual products that they were very easily ruptured and could conse-
quently be preserved intact only with great care.
Size. — The length of this species is commonly about 150 mm. in
extension, although several individuals were found which were more
than 300 mm. long.
Habitat. — The species was found under stones between tides in
rather muddy localities at New Metlakahtla on Annette Island, at
Glacier Bay, Sitka, Yakutat, and at Orca and Virgin Bay in Prince
William Sound. At the four last-named localities the worms were
abundant. Nearly 50 individuals, some of them nearly 300 mm. long,
were found at Sitka massed together in a single cavity in coarse
gravel mixed with mud. All were filled with fully mature sexual
products, and the bodies of nearly all were in contact in a tangled
mass.
Cerebratulus Renier.
Prospetto della Classe dei Vermi (t. Burger), 1804.
The species of this genus are distinguished by long, flattened bodies,
the lateral margins of which are thin, adapted for swimming. Most
species are very active, swim readily with undulatory motion, can roll
up spirally and become twisted, but are only moderately contractile in
length, and do not draw together into a tangled mass. The dorso-
ventral musculature is highly developed, as are likewise the longi-
tudinal and oblique muscles.
Head usually pointed anteriorly, but very changeable in shape;
lateral slits long and deep ; proboscis pore terminal ; mouth large, and
situated behind the ganglia ; esophagal region mostly rounded ; intes-
tinal region broad and flat with thin lateral margins ; the posterior end
extremely flattened and provided with a delicate caudal cirrus, which
extends beyond the opening of the intestine, and in most species is easily
broken off and lost. Eyes usually wanting ; body commonly of a nearly
homogeneous color without distinct markings (such as longitudinal and
circular bands and rings) .
Proboscis sheath reaches to posterior end of body ; proboscis very
long and strong. Intestinal pouches deep and mostly forked peripher-
ally ; central intestinal canal narrow. Neurochord cells probably com-
monly present in brain and lateral nerve cords, though they have as yet
been found in but few species.
ALASKA NEMERTEANS 75
27. CEREBRATULUS HERCULEUS sp. nov.
pi. i, fig. 5.
Body very large, stout, thick, broad, attaining a length of 2 meters
or more, and a breadth of more than 25 mm. ; remarkably broad when
contracted, the body somewhat resembling that of a leech. One speci-
men, preserved in alcohol and strongly contracted, now measures 300
mm. in length, 25 mm. in width in intestinal region; 18 mm. in width
and 15 mm. in thickness in esophagal region. The length of the
esophagal region in this specimen is only about 55 mm. after preser-
vation.
Head very short, thick, and bluntly pointed. Cephalic slits short,
separated in front, reaching back in the alcoholic specimen but 7 mm.
Mouth large, situated as far back as the posterior end of the cephalic
furrows.
Esophagal region short and thick ; intestinal region very broad and
flat, with thin, wavy margins in life. Posterior extremity provided
with a pale caudal cirrus of comparatively small size. Proboscis
large, colorless, fully equal in size to that of a large specimen of C.
marginatus. No ocelli.
Color. — Color very dark brown or reddish brown throughout the
whole dorsal surface, rather paler and less bright ventrally.
This gigantic Nemertean is not clumsy in its movements, but is
active, and swims rapidly and gracefully. It was met with only at
Sitka, where it occurred in considerable numbers in mud at lovr water
mark, associated with C. marginatiis. The worms are fragile, and
break spontaneously unless carefully handled.
28. CEREBRATULUS MARGINATUS Renier.
Cerebratulus marginatus RENTER, Prospetto della Classe del Vermi (t. Burger),
1804.
C. angulatus MclNTOSH, British Annelids ; Pt. i, Nemerteans, p. 195, Ray
Society, 1872-73.
C. fuscus VERRILL, Trans. Connecticut Acad., vin, p. 438, 1892.
This large and active Nemertean was found in several localities
about Sitka, but not in great numbers. The individuals there found
possess the typical color of the species, with which they agree exter-
nally in every detail except the shape of the body. All the Alaska
specimens are much shorter and comparatively broader than those I
have seen in Naples, although careful study of microscopic sections
revealed no essential anatomical differences. It is therefore probable
that this is but a local peculiarity of a single widely distributed species.
76 COE
The worms were found near low water mark living in soft black
mud beneath a considerable growth of 'eel grass.' The mud contained
a great quantity of decomposed vegetable matter, and was saturated
with sulphuretted hydrogen. The worms are excellent swimmers, and
are very rapid in their movements in their underground burrows.
The specimens obtained measured up to 500 mm. or more in length,
and about 15 mm. in width.
The color above was slaty-brown, dark gray, or greenish-gray, with
white or colorless margins.
The species has been previously recorded from the Mediterranean,
the coasts of Great Britain, Madeira, the northeastern coast of Amer-
ica, Greenland, and from other localities.
29. CEREBRATULUS OCCIDENTALIS sp. nov.
Pl. VI, fig. 3.
Body 300 mm. or more in length in moderate extension ; rather slen-
der, rounded in front, flattened behind, and with very thin margins in
the intestinal region, as in other species of the genus. Head change-
able in shape, either rounded or acutely pointed, according to state of
contraction. Cephalic furrows rather short.
Color. — Individuals vary considerably in the general color of body,
but are most commonly chestnut brown or dull reddish anteriorly, and
light chocolate brown in the intestinal region. The ventral surface is
brownish flesh-colored, with a median, ochre ventral stripe. Some-
times the ventral surface is nearly chocolate throughout its entire length.
A darker median dorsal stripe is indicated.
Proboscis. — Remarkably small and slender, being many times
smaller than in most related species of equal size. It is colorless, and
is usually everted when the animal is killed.
In internal anatomy but few points need special mention. The
cephalic glands are unusually well developed, and continue backward
as the cutis glands. The brain is remarkably voluminous, the dorsal
ganglion bilobed, the lower lobe continuing into the cerebral sense
organ.
The mouth is very large, and is provided with an unusual abundance
of buccal glands. There are many diagonal fibers between the pro-
boscis sheath and the body musculature.
Nephridia. — The nephridia are of moderate length, and occupy
the middle third of the esophagal region. The main tubules lie in the
angles between the esophagus and the proboscis sheath. They are
profusely branched towards their anterior ends, each branch lying in
ALASKA NEMERTEANS 77
the wall of one of the esophagal blood lacunae. Farther back the
branches unite into a single tubule on each side. Their main canal
continues backward for a considerable distance in the wall of a lateral
blood lacuna, and is without branches. At its posterior extremity the
efferent duct bends to the dorso-lateral surface of the body, as usual.
The sexual products are fully mature in July.
Habitat. — This is a very active species. It was found in abundance
at Wrangell, Yakutat, Orca, and Virgin Bay. It inhabits the shore at
half tide and below, in muddy places and under stones. The small
size of the proboscis will distinguish it from related species.
30. CEREBRATULUS LONGICEPS sp. nov.
pi. v, figs. 4, 5, 6, 7 .
Body much flattened throughout its whole length ; anterior portion
remarkably narrow and slender, becoming wider posteriorly. Head
much narrower, more slender, and longer than in most species of the
genus, acutely pointed in front, much flattened dorso-ventrally, and with
the tip of the snout often slightly curved upward. A section through
the head is often concave both above and below, showing that the head
is thicker laterally than in the median line. The head is directly con-
tinuous with succeeding portions of the body. Frontal sense organs
very highly developed.
Cephalic furrows very long, and remarkably deep and wide. An-
teriorly they do not reach quite to tip of snout. They are, conse-
quently, well separated from the proboscis pore, which is situated sub-
terminally. The mouth is situated as far back as the posterior end of
the cephalic furrows.
From the narrow, pointed head the esophagal region gradually
widens as it passes backward, but it remains unusually flat throughout ;
the intestinal region is not sharply marked off from the esophagal
region, but widens gradually toward the posterior third of the body,
and narrows toward the posterior extremity. The intestinal region
is even flatter than the more anterior portion of the body. The caudal
cirrus was not observed.
Proboscis. — The proboscis sheath is reduced to a frail tubule in the
posterior third of the intestinal region. The proboscis is slender and
colorless ; it has the three muscular layers and the muscular crosses,
as usual in the genus.
Color. — Dorsal surface dark brownish-black or purplish, much paler
on tip of snout, and on borders of cephalic furrows. This paler border
is wider on the ventral than on the dorsal borders of the furrows. It
78 COE
is sometimes so much increased that it covers the whole ventral surface
in the region of the mouth, and gradually becomes darker toward the
tip of the snout, as well as posteriorly. The color of the ventral surface
is similar to that of the dorsal surface, but is commonly slightly paler,
especially anteriorly, and has a grayish tinge.
No eyes were found.
Size. — Length of largest specimens collected about 300 mm. ; width
in intestinal region 6 mm.
The body is fragile, and the specimens are often broken in killing.
After preservation in formalin or alcohol the body tapers gradually
toward the narrow head, is widest in the posterior third or near the
posterior end, and greatly flattened throughout. The head retains its
long, pointed appearance, the tip is recurved, and the cephalic furrows
in most specimens are deep and widely open.
Frontal sense organs. — On the exact tip of the head are three
rather deep and wide pits, easily overlooked in the entire animal, but
very conspicuous in sections. These are undoubtedly sensory in their
nature, and are lined with slender rod-like cells with especially large
cilia. Burger has described in detail such sense organs in Micrura
and Cerebratulus, and somewhat similar ones in other genera. These
'frontal organs,' as they are called, lie above the proboscis pore; one
of them is situated in the median line and the other two are placed
symmetrically on the antero-lateral margins.
The brain is remarkably large, and the dorsal ganglia are much
larger than the ventral. The posterior ends of the long lateral furrows
are greatly expanded.
Cerebral sense organs. — These also are very voluminous and are
continuous with the posterior ends of the dorsal ganglia. The canals
leading to the exterior are large, and open into the posterior ends of
the cephalic furrows. Internally each canal passes directly beneath
the dorsal ganglion to its internal ventral border, and then bends dorso-
laterally to the external border of the sense organ. These canals leave
the cephalic furrows far back of the cerebral commissure, and at about
the point where the ventral ganglia merge into the lateral cords.
Nephridia. — The nephridial system presents remarkable deviations
from the type usually found in the genus. The nephridial canals ex-
tend throughout the whole length of the esophagal region, and com-
municate with the exterior by upwards of sixty efferent ducts on each
side. In the region of the mouth, or directly behind it, are one or two
nephridial tubules on each side, quite independent of the rest of the
system. Each of these tubules consists of a coil of fine canals lying
ALASKA NEMERTEANS 79
on the dorsal side of the lateral blood lacunae, and projecting freely
into the cavity of the lacuna. From each coiled tubule an efferent
duct leads directly to the dorsal surface of the body. Back of these
anterior, isolated nephridia a continuous richly branched and much
coiled canal extends posteriorly on each side as far as the end of the
esophagal region. The branches of these canals are limited to the
dorsal and lateral surfaces of the lateral blood spaces, which occupy
the angles between the esophagus and the proboscis sheath. The
coiled tubules project freely into the blood spaces, and encroach con-
siderably upon their area. The very numerous efferent ducts, which
are given off from these tubules, pass directly outwards to the surface
of the body. Those which pass out from the most lateral of the ne-
phridial tubules, open on the dorsal surface near the lateral margins of
the body, while those which leave the tubules nearest to the proboscis
sheath open on the dorsal surface not far from the median line. The
greater portion of the nephridiopores, however, lie rather nearer to
the lateral margins than to the median dorsal line. Although there are
practically the same number of efferent ducts on each side, this does
not signify that these ducts are paired, but rather that the average
number on each side is identical. In several instances an unusually
large efferent duct on one side was accompanied by one of similar
size nearly opposite. These ducts vary greatly in size, some being
several times as large as others. Great irregularity likewise exists as
to their distribution along the course of the nephridial canal. In the
anterior third of the esophagal region they are well separated, but
more posteriorly, portions of two or three sometimes appear on the
same side in a single section.
Sexual glands. — The reproductive glands appear immediately be-
hind the nephridia, and in both sexes alternate with the intestinal
pouches. The sexual products were fully mature in July, and in both
sexes the genital ducts were fully formed. They were lined with a
special flattened epithelium, and opened on the dorso-lateral aspects of
the body in both sexes. The ova when immature are attached by a
narrow stalk to the wall of the ovary, and later break off and fall into
the ovarian cavity. The spermatozoa have short, oval heads instead
of the slender, pointed ones so common in the genus.
Habitat. — Yakutat ; under stones at low water ; not common.
8o COE
31. CEREBRATULUS MONTGOMERYI sp. nov.
pi. vi, figs, i and 2.
Body large, very long and ribbon-like when fully grown, and much
flattened, except in esophagal region. Head variable in shape, ac-
cording to state of contraction ; sometimes rounded and obtuse, at
other times much elongated, pear-shaped, and flattened dorso-ventrally.
The anterior portion of the body, back as far as the intestinal region,
is narrower and thicker than the posterior portions, and has rounded
margins ; in the intestinal region the body is very flat, and the lateral
margins are extended to very thin edges. In partial contraction the
dorso-ventral muscles, situated a little distance from the outer edge of
the margins, contract so strongly as to produce a lateral ridge on each
side both above and below the lateral edge. A transverse section con-
sequently shows that the lateral margin is thicker distally than it is
nearer the median line. These lateral ridges are often seen in speci-
mens preserved in alcohol or formalin. Posteriorly the body becomes
still thinner, and is narrower toward the extremity. The posterior end
is either obtuse or sharply pointed, according to the state of contrac-
tion. The posterior opening of the intestine is subterminal. In none
of the many specimens obtained was the caudal cirrus present, though
such an appendage is probably present in uninjured worms.
The cephalic slits are moderately elongated, and extend backward
as far as the anterior end of the mouth. Their posterior ends are
wide and deep, and in life their margins are thin and widely separated.
Anteriorly they do not reach quite to the proboscis pore, except when
the animal is strongly contracted.
Color. — In life the whole body, both above and below, except the
tip of the head, is bright blood red. The anterior extremity is tipped
with a narrow band of white or yellowish-white. This white tip ex-
tends both above and below, and is not usually more than a few milli-
meters wide. The proboscis is light red in color.
Size. — Individuals were most commonly from one to two meters in
length, and 8 to 10 mm. in width in the intestinal region. A few of
the specimens obtained measured as much as two and a half meters
when extended.
The individuals are generally hardy, and are not so prone to break
spontaneously into fragments as are those of many other species of the
genus. They contract much less violently than most other Nemerteans
when killed, and may consequently be easily preserved entire by kill-
ing in a dilute solution of formalin in sea water.
ALASKA NEMERTEANS 8 1
Proboscis. — The proboscis is pale red, and of moderate size for the
genus. Its musculature consists of an inner circular, a longitudinal,
and a very thin outer circular muscular layer. It is, therefore, differ-
ent from those of nearly all the species of the family Lineidcs^ in that
the inner longitudinal muscular layer is wanting. There are fibers
passing between the inner circular layer and the thin outer circular
layer, but these fibers do not form such distinct crosses as are char-
acteristic of most of the members of the family. The internal epi-
thelium is thick and highly glandular. The nerve plexus beneath this
layer is unusually conspicuous, and lies directly internal to the circular
muscular layer. The proboscis is very often retained in place after the
animal has been killed. The muscular layers of the proboscis sheath
are rather thick, and the circular muscular fibers often cross into the
circular muscles of the body in the median line.
In the anterior portion of the head a rich growth of glands pene-
trates the other tissue nearly to the central proboscis sheath. These
glands mostly open directly outward on the whole circumference of
the head, although a few open on the tip of the snout. Back of the
lateral slits the glands are restricted to the cutis. In the region of the
mouth the cutis glands are separated from the outer epithelium by a
layer of interlaced fibers of connective tissue, forming a basement layer
of double the thickness of the epithelium. In the intestinal region the
cutis glands are much reduced, and scattered. The outer muscular
layer of the body is about as thick as the two inner muscular layers
combined.
The mouth is much elongated, and its anterior end reaches forward
as far as the posterior ends of the cephalic slits. The intestinal pouches
are deep and narrow, and are forked distally.
Nephridia. — The nephridial canals are profusely branched, and lie
in contact with the blood spaces around the esophagus in front of the
middle esophagal region. Their extent longitudinally, however, is
short. The main nephridial canals lie in the angles between the
esophagus and the proboscis. Near their posterior extremities a large
efferent duct passes obliquely upward on each side and opens on the
latero-dorsal aspect of the body. As described by Burger1 in C.
marginatuS) one of the efferent ducts often lies far behind the other.
Posterior to the efferent ducts the nephridial canals are smaller, and
their branches end a short distance farther back. The blood lacunae
in the head in the specimens sectioned were very much reduced in size.
1 Fauna u. Flora Neapel, Monogr. 22, p. 622.
82 COE
The brain lobes are large, and the ventral commissure enormously
thick. The well developed cerebral sense organs abut closely against
the posterior ends of the dorsal ganglia. Their ciliated canals pass
obliquely inward from the posterior ends of the cephalic slits. In their
course these canals pass external and ventral to the dorsal ganglia. The
median dorsal nerve, lying just outside the circular muscular layer of
the body, is large and conspicuous throughout nearly the whole length
of the animal.
Ocelli are wanting in the adult worm.
In specimens collected in June and July the genital products had
evidently been recently discharged.
This species is named in honor of Dr. T. H. Montgomery, Jr., of
the University of Pennsylvania, whose studies on the Nemerteans form
most valuable contributions to the knowledge of the group.
Habitat. — This magnificent Nemertean was found abundantly under
stones in muddy places near low water mark at most of the collecting
stations between Sitka and Unalaska. At Orca and Virgin Bay, in
Prince William Sound, and at Dutch Harbor, Unalaska, it proved to
be one of the commonest species. Professor Kincaid has sent me a
large specimen from Puget Sound, in the State of Washington, where
it is said to be not uncommon. This would indicate that the species
may be found locally 'along the whole northwestern coast of North
America south of Bering Sea.
32. CEREBRATULUS ALBIFRONS sp. nov.
pi. iv, figs. 3, 4.
Body elongated, ribbon-like, flattened behind, rounded in front as in
typical species of the genus. Margins of intestinal region pale and
thin. Cephalic slits unusually long and deep, reaching well beyond
posterior end of mouth.
Color. — General color of body brownish purple. Anterior end of
head white, including both dorsal and ventral surfaces, and extending
backward about as far as anterior end of mouth, and sometimes reach-
ing along borders of cephalic slits to their ends. The esophagal re-
gion is dark smoky purple on dorsal surface ; the ventral surface is
similar but paler ; the intestinal region inclines more to reddish. A
darker line runs along the middle of the dorsal surface and the in-
testinal lobes appear more opaque.
Habitat. — Only a single specimen of this pretty species was se-
cured, and this, unfortunately, had its posterior extremity missing.
The specimen was about 150 mm. in length and of moderate propor-
ALASKA NEMERTEANS
tions. It was restless in confinement, and an active swimmer. It was
found under a stone near low water mark at Hot Springs, near Sitka.
EXPLANATION OF PLATES.
The colored plates (I to VI) are reduced from colored sketches
made, with a few exceptions, from the living animal. They were com-
pleted after the return of the expedition and have been retouched, and
in some instances entirely redrawn, by Mr. A. H. Verrill. Every
effort has been made to make the colors and the form of the body as
natural as possible.
In the figures on the plates the
ap, attachment of proboscis.
bg, buccal glands.
bg' , accessory buccal glands.
bl, blood lacuna.
ble, epithelium of same.
bin, basement layer.
br, brain.
bv, blood vessel.
bva, anastomosis of lateral vessels.
cc, ciliated canal of cerebral sense or-
gan.
cgl, cephalic glands.
cm, circular muscular layer.
cso, cerebral sense organ.
ct, connective tissue.
cugl, cutis glands.
dc, dorsal commissure of brain.
dg, dorsal ganglion.
dgl, secretion of cephalic glands.
dn, median dorsal nerve.
e, esophagus.
eep, epithelium of same.
f, fibrous layer of cutis.
gc, glandular cells of cerebral sense
organ.
gl, integumental glands.
gp, genital pouch.
*', integument.
ic, intestinal csecum.
ion, inner circular muscular layer.
following reference letters are used
ijl, inner fibrous layer.
ilm, inner longitudinal muscles.
in, intestine.
Im, longitudinal muscles.
In, lateral nerve-cord.
m, mouth.
mep, epithelium of same.
nc, nerve commissure.
nd, efferent nephridial duct.
nep, nephridial canal.
np, nerve plexus.
nv, nerve.
oc, ocellus.
ocm, outer circular muscles.
oep, outer epithelium of proboscis.
olm, outer longitudinal muscles.
ov, ova.
pern, circular muscles of proboscis.
plm, longitudinal muscles of proboscis.
pn, proboscis nerve.
ps, proboscis.
ps/i, proboscis sheath or its cavity.
rh, rhynchodseum.
ro, opening of same.
smg, submuscular glands.
SH, nerve to cerebral sense organ.
sop, opening of ciliated canal of sense
organ.
•vc, ventral commissure of brain.
•vg, ventral ganglion.
84
COE
INDEX TO GENERA AND SPECIES, PART I
Synonyms are in italics ; species new to science, and pages on which generic
or specific descriptions occur, are in black face type.
Amphiporus 40, 9, 28, 32, 33, 34
angulatus 41, 3, 6, 9, 36
bimaculatus 44, 6, 9, 31
exilis 54, 6, 9, 51, 52, 53
lactifloreus 5;, 54
leucioduasi, 6, 9, 31, 55
nebulosus 48, 6, 9
neesii 4
tigrinus 46, 6, 9
virescens 28
Borlasia camillea 4
quinqnelineata 4
Carinella u, 7
annulata, 13, 17
capistrata 16, 5, 8, 71
dinemais, 5, 8, 17
nothus 17
polymorpha 12
speciosa n, 5, 7
superba 15, 16, 17
Carinoma 20, 8, 73
armandi 20, 21, 22
griffini 20, 5, 8
patagonica 21, 22, 23
tremaphoros 20
Cephalothrix 19, 8
linearis 19, 5, 8
Cerebratulus 74, 10, 68
albifrons 82, 6, n
angulatus 75
bellus 71
fuscus 75
herculeus 75, 6, 10
impresses 70, 3
longiceps 77, 6, n
marginatus 75, 6, 10
montgomeryi 80, 6, 1 1
occidentals 76, 6, 10
Cosmocepkala beringiana 41, 3, 42
Emplectonema 23, 4, 8, 28, 32, 54
biirgeri 25, 5, 8
camillea 4
gracile 23. 3, 5, 8
Emplectonema viride 23, 3, 4, 25
Eunemertes 23, 3
gracilis 23
neesii ^
Eupolia 61, 3, 4
Fasciola angulata 41
Lineus 64, 10, 68
torquatus 65, 6, 10
viridis 65, 6, 10
Micrura 68, 10
alaskensis 71, 6, 10
bella 71
impressa 70, 3, 10
verrilli 68, 6, 10
Nareda superba 41, 43
Nemertes 23
gracilis 23, 3, 4
obscura 65
Omatoplca stimpsonii 41, 42
Paranemertes 32, S
carnea 37, 6, 9, 32
pallida 36, 5, 8, 32
peregrina 33, 5, 8, 32
Planar ia gesserensis 65
linearis 19
•viridis 65
/W/fl 6 1
Tseniosoma 61, 4, 10
cequale 4
princeps 62, 6, 10
quinquelineatum 4
septemlineatum 4
Tetrastemma 57, 9
aberrans 58, 6, 9, 57
bicolor 57, 6, 9
csecum 59, 6, 9
cruciatum 58
rubrum 58
vermiculus 58
Zygonemertes 28, 9
albida 31, 5, 9, 29
tbalassina 29, 5, 9
virescens 29, 30
PLATE I.
FIG. I. Carinella capistrata sp. nov. A large individual about natural size.
Virgin Bay, Prince William Sound.
2. Carinella dinema sp. nov. Victoria, B. C. Enlarged 2^ times.
3. Head of same species. Side view. Enlarged 4 times.
4. Amphiporus bimaculatus sp. nov. Dorsal view of head. Glacier Bay.
Enlarged 12 times.
5. Cerebratulus herculeus sp. nov. A large individual. Sitka. Half nat-
ural size.
6. Tetrastemma bicolor sp. nov. Kadiak. Three times natural size.
(86)
H. A . e. VOL Xi
PLATE
ALASKA NEMERTEANS
PLATE II.
FIG. i. Rmplectonema burgeri sp. nov. Anterior portion of an individual of
the pale variety. Glacier Bay. % natural size.
2. E. burgeri. A large individual of the dark variety. Glacier Bay. %
natural size.
3. Tczniosoma princeps sp. nov. Large individual. Yakutat. % natural
size.
4. T. princeps. A contracted specimen in formalin. Cape Fox. Nat-
ural size.
5. Zygonemertes thalassina sp. nov. Sitka. Twice natural size.
6. Paranemertes peregrina sp. nov. Brown variety. Victoria, B. C. A
small specimen. Natural size. Compare pi. in, fig. 5.
(83)
H. A. E. VOL. XI
PLATE
ALASKA NEMERTEANS
PLATE III.
FIG. i. Amphiporus exilis sp. nov. Glacier Bay. Slightly enlarged.
2. Zygonemcrtes albida sp. nov. Victoria, B. C. Enlarged ij£ times.
3. Paranemertes carnea sp. nov. Taku Harbor. Anterior portion of
body with emarginate head. Natural size.
4. P. carnea. Sitka. % natural size.
5. Paranemertes peregrina sp. nov. Virgin Bay, Prince William Sound.
% natural size.
6. Carinella speciosa sp. nov. Hot Springs, near Sitka. ^ natural size.
(90)
H. A. E. VOL. XI
PLATE III
ALASKA NEMERTEANS
PLATE IV.
FIG. i. Amfhiporus nebulosus sy.nov. Kukak Bay, Alaska Peninsula. Natural
size.
2. Micrura alaskensis sp. nov. Sitka. Enlarged i^ times.
3. Cerebratulus albifrons sp. nov. Near Sitka. Twice natural size.
4. Side view of head of same individual.
5. Amphiporiis tigrinus sp. nov. Farragut Bay. Mature male specimen,
slightly enlarged.
6. A. tigrinus. Head of male ; dorsal view. Twice natural size.
7. A. tigrinus. Farragut Bay. Female with ripe ova. Slightly en-
larged.
8. Head of female of same species. Dorsal view. Twice natural size.
(92)
H. A. E. VOL. XI
PLATE IV
s •r*'-
i
ALASKA NEMERTEAN:
8>isellithBoslon
PLATE V.
FIG. i. Micrura -verrilli sp. nov. Virgin Bay, Prince William Sound. Natu-
ral size.
2. Side view of head of same species.
3. The same ; contracted.
4. Cerebratulnslongiceps^.nov. Yakutat. Enlarged \y2 times.
5. 6, 7. Heads of same species, from lateral, dorsal, and ventral aspects re-
spectively.
8. Linens torquahts sp. nov. Orca, Prince William Sound. Natural size.
9. L. torquatus. Side view of head.
10. Amphiporus bimaculatus sp. nov. Victoria, B. C. Natural size.
(94)
H A . E. VOL XI
PLATE V
ALASKA NEMERTEANS
PLATE VI.
FIG. i. Cerebratulus montgomeryi sp. nov. Dutch Harbor, Unalaska. Natural
size.
2. Head of same species from ventral surface.
3. Cerebratulus occidentalis sp. nov. Yakutat. Natural size.
4. Ampkiporus angulatus (Fabr.) Verrill. Kadiak. Natural size of large
specimen.
H. A. E. VOL XI
PLATE VI
ALASKA NEMERTEANS
PLATE VII.
FIG. i. Zygonemertes thalassina. Outline of stylet apparatus of proboscis
( X 50). la, i£, several accessory stylets from two individuals (X 225).
Sitka.
2. Amphiporus angulatus. Central portion of proboscis (X4°)- Taku
Harbor. 2a, central stylet and basis of another specimen (X So).
3. Paranemertes pallida. Outline of middle region of proboscis ( X 60).
4. P. cornea. Extremity of everted proboscis, with six pouches of acces-
sory stylets, of which but four are shown ( X 35)- Sitka.
5. Amphiporus exilis. Middle portion of proboscis, indicating, besides
central stylet and basis, eight pouches of accessory stylets (X4°)«
Yakutat.
6. Amphiporus leuciodus. Stylet apparatus of proboscis showing the three
accessory stylet pouches ( X6o). Victoria, B. C.
7. Paranemertes peregrina. Outline of middle portion of proboscis
(XS°)- Virgin Bay.
(98)
H A. E. VOL. XI
PLATE VII
Coe de!
ALASKA NEMERTEANS
PLATE VIII.
FIG. i. Emplectonema burgeri . Outline of stylet apparatus (X35)- Glacier
Bay.
2. Amphiporus bimaculatus. Middle portion of proboscis, with four
pouches of accessory stylets ( X 35 )• Glacier Bay.
3. Emplectonema gracile. Stylet apparatus of proboscis (X5°)» Popof
Island.
4. Amphiporus tigrinus. Middle region of proboscis (X6o). Farragut
Bay. The deeply stained wreath of glands is indicated only in section.
5. Zygonemertes albida. Middle portion of proboscis ( X 4°) • Victoria,
B. C.
6. Amphiporus nebulosus. Outline of stylet apparatus of proboscis (X5°)«
Kukak Bay.
7. Paranemertes carnea. Side view of head, with partially everted probos-
cis. The position of the lateral oblique furrows, and the arrangement
of the ocelli are indicated ( X 5)- Taku Harbor.
(100)
H. A. E VOL. XI
PLATE
J
ALASKA NEMERTEANS
PLATE IX.
FIG. I. Carinella sfeciosa. Transverse section through head in front of brain
showing, particularly, position of enormously developed cephalic glands
( ££•/) and arrangement of cephalic nerves (nv}. The cephalic glands
are thickly placed both above and below the cephalic blood lacunas (bl},
as well as around the rhynchodzeum (rh). The secretion from many of
these glands is discharged near the lateral margins as indicated (dgl}.
Between the longitudinal and oblique muscles (/?«), which lie in the
deeper parts of the head, and the circular fibrous layer (cm}, composed of
muscles and connective tissue fibers underlying the integument (i ), are
numerous and very massive nerves (nv} which supply the head regions
in front of the brain. The broad indentation on the ventral margin has
no relation to the mouth, which lies much farther back. Other reference
letters are explained above (p. 83). A small individual. ( X45)-
2. Carinella speciosa. Transverse, but somewhat oblique, section through
head in region of brain. On the right of the section the brain (dg and vg}
lies directly beneath the circular fibers (cm}. External to the brain, and
lying among the bases of the integumental cells, is the highly specialized
cerebral sense organ (cso), connected with the dorsal ganglion (dg) by
several small nerves (s«). The cerebral sense organ shows a conspicu-
ous central canal. The rhynchodseum (rh} has lost its fringe of gland
cells, these being limited to the regions anterior to it. The outer por-
tion of the circular layer (cm} constitutes the basement membrane of the
integument. In the region of the brain the circular layer splits into two
sheets, one passing external to the brain and the other (y?) internal.
Other reference letters as above. A large specimen. (X25).
3. Carinella sfeciosa. Transverse section through nephridial region show-
ing positions, and one of the openings, of the nephridial canals. The
lateral blood lacuna (bv} and the nephridial canal (nef) lie embedded in
a band of loose connective tissue on each side. The opening of one of
the efferent nephridial ducts (nd) is seen on the right of the drawing.
Other reference letters as indicated above. (X25).
(102)
H. A. E. VOL. XI
PLATE IX
.-rh.
...Ml.
m—-fo
Coedel
ALASKA NEMERTEANS
PLATE X.
FIG. i. Carinella sfeciosa. Portion of transverse section through nephridial
region. The ciliated nephridial canal (nep) shows the peculiar in-
folding of the integumental cells on its dorsal border. These integu-
mental cells are here loosely arranged, and show several deep infoldings
(of) which, under certain circumstances, may possibly have a more or less
distinct communication with the nephridial canal. But a small portion
of the closely packed gland cells in the integument are indicated. Ref-
erence letters are explained on p. 83. (X 75)-
2. Carinella speciosa. Portion of a transverse section through the brain
region. The relation of the dorsal ganglion (dg) with the highly spec-
ialized cerebral sense organ (cso) is indicated. The ciliated canal (cc)
of the sense organ connects directly v/ith the exterior by a narrow tube
(sof) — shown in dotted lines in the drawing, because it lies mainly in
another section — opening on the lateral margin of the head. Several
nerves (sn) are seen to pass from the dorsal ganglion to the sense organ.
Other reference letters as above. Only a small portion of the integu-
mental gland cells are shown. (X 75 )•
3. AmpJiiponis tigrinus. Dorsal view of anterior portion of body, cleared
in cedar oil. In front of the brain the arrangement of the ocelli is
shown, and farther back the position of the ovaries (ov). The intesti-
nal cseca are not indicated. Farragut Bay. ( X 8).
4. Amphiporus tigrinus. Median sagittal section through the anterior por-
tion of the body. The cephalic glands ( cgl ) lie above the opening of the
rhyncodseum (ro). The mouth (m) separates from the proboscis
opening a little way back. The attachment (a/) of the proboscis to the
tissues of the head is seen to be well in front of the brain commissures
(dc and vc). The section shows the comparative size and arrangement of
the proboscis, blood vessels, esophagus, and other organs. Preference
letters as above. (X 3°)-
(104)
H. A. E. VOL XI
PLATE X
Oft.
nap. ct. bv bm 'In
6m .
^gsfe^i!
^
' ' •' v. it '•""••• • -• • y> " -^~~
\r.
m
Coe del
ALASKA NEMERTEANS
!5e. i
PLATE XI.
FIG. i. Amphiporus nebulosus. Transverse section through posterior portion of
ventral commissure of brain. The submuscular glands (sniff} fill up
a large portion of the tissues of the ventral half of the head. The
pair of nerves leading forward to the cerebral sense organs is seen to
originate (son] from the ventral side of the dorsal ganglia. Other
reference letters are explained on p. 83. (X 26).
2. Amphiporus angnlatiis. Transverse section through ventral commis-
sure of brain. The dorsal attachment of the proboscis to the tissues of
the head is shown. The proboscis nerves (/«) enter the ventral side of
the proboscis, and divide into a definite number of branches (usually
18), which farther back arrange themselves symmetrically about the
periphery. The roots of the dorsal brain commissure (rdc) are indi-
cated. In the right half of the section the anterior ends of the nephri-
dial tubules (ncp} are seen. Submuscular glands (smg] are not as
numerous as in A. nebulosus. Other reference letters as above. (X 26).
3. Amphiporus cxilt's. Transverse section through nephridial region to show
efferent nephridial ducts (nep] opening on dorso-lateral aspects of body.
This condition is extremely rare in Metanemerteans. Several diverti-
cula of the intestinal caecum (inc) are represented. Other reference
letters as above. (X32)-
(106)
H. A. E. VOL XI
PLATE XI
inc.
ALASKA NEMERTEANS
PLATE XII.
FIG. i. Paranemertes pallida. Transverse section through nephridial region.
In this case the single pair of efferent nephridial ducts (nef) was so
symmetrically placed that both were cut in a single section. The very
small size of the proboscis (P$} in the large proboscis sheath (psk) is
remarkable. The intestinal cascum (ic) shows lateral diverticula above
the lateral nerve cords. (X 18).
2. Amphiporus bimaculatus. Transverse section of proboscis. The 16
proboscis nerves (pn) are very sharply denned. The cylindrical plexus
(np) of nerve fibers and connective tissue serves to connect the nerves,
and divides the thick longitudinal muscular band into an inner (pint]
and an outer (pirn') layer. The inner longitudinal muscular layer,
found in most related species, is wanting — the basement layer (bm) of
the internal epithelium (pep) lying immediately beneath the circular
muscular layer. ( X 66) .
3. Emplectonema biirgeri. Transverse section through body, showing
manner in which the esophagus (e) opens into dorsal wall of intestine
(»'»). This section also shows the genital pouches (gp) lying both
above and below the intestinal lobes, and indicating that they open
respectively on the dorsal and ventral surfaces of the body. Of the
submuscular glands (smg), which extend throughout the esophagal re-
gion in great abundance but a few remain as far back as the position of
the section figured. (X20)-
(108)
H. A. E. VOL. XI
PLATE xu
h\\, 6
Coe del
ALASKA NEMERTEANS
PLATE XIII.
FIG. i. Micrura alaskensis. Transverse section through mouth region. The
peculiar accessory buccal glands (bgf) are seen to lie outside the cir-
cular muscular layer (COT), and amongst the fibers of the outer longi-
tudinal muscular layer (olm). Their secretion passes through the layer of
circular muscles, and mixes with that of the true buccal glands (bg),
which lie immediately beneath the epithelium of the mouth (mep). The
buccal nerves (bn) lie on the lateral borders of the mouth as usual.
Other reference letters are explained on p. 83. (X 3°)-
2. Zygonemertes thalassina. Transverse section immediately back of the
brain. The intestinal caeca (tc) reach forward to abut against the dor-
sal ganglia. The ocelli (oc) extend back of the brain, and occupy posi-
tions, as shown, immediately above and external to the lateral nerve
cords (In). (X 100).
3. Amphiportis angtilatus. Transverse section through posterior end of
body. The drawing is slightly diagrammatic, as it contains portions of
the two adjacent sections. The anastomosis of the three longitudinal
blood vessels (bva) is seen to lie immediately dorsal to the commissure
(nc) of the lateral nerves (In) ; the basement layer (bm) is remarkably
thick in this region. The posterior end of the intestine (in) opens
ventrally a few sections farther back. Other reference letters are ex-
plained above. (X So).
(1 10)
H. A. E. VOL. XI
PLATE XIII
^.,' 3
Coe del
ALASKA NEMERTEANS
NEMERTEANS OF THE PACIFIC
COAST OF NORTH AMERICA
PART II
(in)
NEMERTEANS OF THE PACIFIC COAST
OF NORTH AMERICA
PART II
BY WESLEY R. COE, PH.D.
CONTENTS
Introduction , 113
Alaska species found in California 118
Species new to west coast of North America 119
Distribution of all species known from region 121
Key to new species and species new to region 125
Systematic discussion of new species 129
Index to genera and species 219
INTRODUCTION
SINCE the publication of the report on the Nemerteans col-
lected on the Harriman Alaska Expedition,1 I have had an
opportunity of studying extensive collections of this group
of worms from Alaska and from other portions of the Pacific
coast of North America. The report on these collections is
soon to appear in the Bulletin of the Museum of Comparative
Zoology. I have also found opportunity to spend a summer
on the coast of California,2 where I obtained a considerable
1 Published in Proc. Wash. Acad. Sci., in, pp. i-no, pis. i-xm, March, 1901.
8To Dr. \Vm. E. Ritter, of the University of California, for the hospitality
of the Marine Laboratory at San Pedro, and to Dr. C. H. Gilbert, of Stanford
University, for similar privileges at the Hopkins Seaside Laboratory at Pacific
Grove, the writer wishes to express his most cordial thanks. The writer is also
indebted to Professor C. B. Wilson, of Westfield, Mass., for numerous speci-
mens and very valuable notes on California Nemerteans ; and to Mr. J. F. Abbott,
formerly of Stanford University, for numerous important notes and drawings
from specimens collected at Monterey Bay, Calif.
114 COE
number of species which were not collected on the Harriman
Expedition, and a large proportion of which have proved to be
new to science.
A study of the Nemerteans of the California coast reveals the
fact that a number of the Alaska species extend southward
throughout the whole length of the State, and that many others
occur as far south as Monterey Bay. We may thus expect that
many of those forms which I have more recently found in Cali-
fornia may range northward into Alaska, so that their incor-
poration in the report on the Alaska species can by no means
be out of place.
In the following pages I shall not attempt to describe all the
species which have come into my hands from the Pacific coast,
but shall here confine myself to those forms which I had an op-
portunity of collecting personally and studying while they were
still alive. The specific descriptions can thus be made far more
precise and exhaustive than when preserved material only is to
be had. In all cases, however, serial sections have been care-
fully studied to determine the anatomical peculiarities of each
species recorded.
The most strikingly colored forms were drawn as nearly as
possible in their natural colors, and an attempt was made to re-
produce the natural shape and characteristic position of the body
when alive.
A single species (A. -paulinus) has been described by Pun-
nett1 from the Pribilof Islands, Bering Sea, since the first por-
tion of this report appeared. A brief description of this species
will be found on p. 155.
At the time of publishing the first portion of the report, I was
unfortunately unaware that a preliminary paper by the late B.
B. Griffin on Some Marine Nemerteans of Puget Sound and
Alaska2 had appeared since the death of this enthusiastic young
investigator. Several of the species very briefly described by
Griffin were through this oversight redescribed by me in my
previous paper with names which must be now relegated to syn-
onymy. Griffin's drawings, notes, and collections have recently
1Proc. Zool. Soc. London, p. 92, 1901.
2 Ann. New York Acad. Sci., xi, pp. 193-217, 1898.
NEMERTEANS 1 15
been placed in my hands, so there can be no doubt as to the
identity of these forms. They are :
1 . Carinella dincma Coe = C. scxlincata Griffin.
2. C. spcciosa Coe == C. rubra Griffin.
3. Carinoma griffini Coe = C. mutabilis Griffin.
4. Amphiporus leuciodus Coe = A. imparispinosus Griffin.
5. A. exilis Coe = A. formidabilis Griffin.
Griffin's paper includes twelve named species besides two
forms which are not designated by specific names. The geo-
graphical distribution of these is as follows :
1. Carinella scxlincata Griffin. Puget Sound ; Sitka, Alaska.
2. C. rubra Griffin. Puget Sound; Sitka, Alaska.
3. Carinoma mutabilis Griffin. Puget Sound; Strait of Juan
de Fuca. Varieties : argillina, in hard blue clay ; vasculosa, in
sand between tides.
4. Emplcctoncma viridc Stimpson = E. gracilc (Johnston)
Verrill. Southern Alaska ; Puget Sound.
5. E. violaccum Griffin, (non Burger) = E. biirgeri Coe.
This form, which Griffin considers identical with Burger's
species from the coast of Chile,1 was found on piles at Port
Townsend, Puget Sound. Griffin's description is substantially
as follows : Body extremely flattened, ribbon-like ; head rounded
in front, directly continuous with body; eyes numerous. Color
somewhat variable, with fairly constant pattern on dorsal surface
which is densely flecked with purple or brown upon a pale yel-
lowish brown ground color ; ventral surface yellowish white.
Length probably 50 cm., although it was difficult of measurement
because the body remained coiled up in tangled knots in an enor-
mous amount of slime which the worm secretes. These char-
acters agree in the main with the brownish variety of E, burger tt
but differ widely from Burger's E. violaccum, which is dark
brownish violet above and pale rose-violet on ventral surface
even after preservation. Griffin considers the internal anatomy
to agree « more or less closely ' with Burger's E. violaccum.
My own preparations, however, prove conclusively that the two
species are specifically distinct, for they differ decidedly in the
only really specific anatomical character which Burger gives ;
JZool. Jahrb., Abth. d. Syst., ix, p. 272, 1896.
COE
viz., in E. vtolaceum the cerebral sense organs are very small,
and lie far in front of brain, while in E. burgeri they are
remarkably large for the genus and lie only slightly in front of
brain.1 The practical absence of cephalic glands is in accord
with most species of the genus. Neither Burger nor Griffin
makes any statements in regard to the proboscis, which usually
presents the most tangible specific characters.
6. Amphiporus imparisjrfnosus Griffin. Sitka, Alaska ; Puget
Sound.
7. A. formidabilis Griffin. Alaska and Puget Sound
8. A. brunncus Griffin. Port Townsend, Puget Sound.
Presents the following peculiarities : Length in alcohol 3. 3 cm. ;
width 5 mm. Color in life dark brown or smoky black dorsally,
greenish or yellowish white ventrally ; on each side of neck is a
pale, angular spot. Cephalic glands moderately developed;
cephalic sense organs considerably in front of brain. Intestinal
caeca reach nearly to brain. Basis of central stylet long ; two
lateral pouches with two (or three ?) stylets each.
In some respects this description agrees with the characters
of Paranemertes*peregrina,) which is common in Puget Sound,
but on the following page (p. 213) the species is represented as
bearing " a more or less general resemblance to A. angulatus"
There can hardly be said to be the slightest resemblance be-
tween P. pcregrina and A. angulatus, so that even with Griffin's
notes and material at hand, there being no specimens of this
form it is impossible to determine to which, if any, of the des-
cribed species this A. bnmneus belongs. For the present it is
necessary therefore to consider it as a distinct species.
9. A. angulatus (Fabr.) Verrill. Sitka and Redout Bay,
Alaska. Doubtfully referred to this species by Griffin ; several
varieties obtained. This species is abundant along the whole
coast, from Bering Strait to Puget Sound.
10. A. drepanophoroides Griffin. No locality given. Length
4-5 cm. or less ; form short and stout : color red above, white
beneath ; eyes numerous, in rows along antero-lateral margins of
head. Cephalic and submuscular glands prominent. Cerebral
sense organs large, situated beside brain and extending pos-
JCoe, Proc. Wash. Acad. Sci., in, p. 26, 1901.
NEMERTEANS 117
teriorly behind dorsal ganglia ; canals open in front of ventral
commissure. Differs from all the preceding species in small-
ness of rhynchocoel, which is enclosed in a thick muscular sheath
in which longitudinal and circular muscles are interwoven.
No intestinal c£ecum ; circular muscle-layer quite thick.
The species is known only from the above description, and is
truly remarkable because of the absence of the intestinal caecum.
No specimens or slides showing any such peculiarity were con-
tained in Griffin's collections, however, when they were turned
over to the writer.
11. Linens striatus Griffin. Puget Sound. Color notes and
drawings lost by shipwreck. " Color brownish red on dorsum,
sharply marked off laterally from the much lighter ventral por-
tion. Dorsum marked by numerous creamy white transverse
bands which cease at demarcation-line between the dorsal and
ventral coloring. Tip of head brilliant red. Length probably
not over 4 cm." Nephridia have numerous efferent ducts.
These characters resemble those of very small individuals of
M. verrilli in many respects, and the two species may possibly
be identical. Perhaps Griffin's form is more closely similar to
Stimpson's Ccrebratulus impressus (== Micrura impressd) from
Bering Strait.
12. Ccrebratulus marginatus Renier. Puget Sound.
13. Lmeus&p.l Puget Sound. Smoky black with greenish
tinge dorsally, grayish brown ventrally. Probably = Lineiis
•viridis (Fabr.) Johnston, which is recorded from southern Alaska
(Coe, loc. cit., p. 65).
14. Ccrebratulus sp. ? A very large, dark species with flesh-
colored margins ; fragments measuring nearly 20 mm. in diam-
eter after preservation. No locality given, but the species is in
all probability C. herculeus Coe, which is also recorded from
southern Alaska.
Careful comparison of Griffin's notes, drawings, and material
indicates that eight of the twelve above-named species were new
at time of publication. Five of the eight must be retained in
place of five of my own names, as stated above. Two other
new species (Amphiporus brunneus and A. drcpanophoroides]
cannot be referred to any forms which have come into my
COE
hands, and must stand as new for the present. One other spe-
cies (E, bnrgcri Coe), although undescribed at the time, was
incorrectly referred to E. violaccuin Burger. Three of the four
remaining forms were correctly identified with European spe-
cies, while the one species remaining, Lincus striatus, is pos-
sibly identical with M. impressa (Stimpson), as stated above.
ALASKA SPECIES FOUND ON THE COAST OF CALIFORNIA
Of the species recorded in the first part of this report (pp. i-
110), the following were collected in the summer of 1901 on
the California coast in the localities indicated below.
Paleonemertea.
1. Carinella capistrata Coe. Monterey Bay.
2. C. sexlincata Griffin (= C. dincma Coe) . Monterey Bay ;
San Pedro.
3. Carinoma mutabilis Griffin (= C. griffini Coe). San
Pedro.
4. Cephalothrix linearis (Rathke) Oersted. Monterey Bay ;
San Pedro ; San Diego.
Hoplonemertea.
5. Emplcctonemagracile(]o\ir\stor\)Vei~Y\\\. Monterey Bay.
6. Paranemertes ^peregrina Coe. Monterey Bay ; San
Pedro.
7. Amphvporus bhnaculatus Coe. Monterey Bay.
8. A. angulatus (Fabr.) Verrill. Monterey Bay ; San Pedro.
9. A. imparispinosus Griffin (= A. Icuciodus Coe). Mon-
terey Bay; San Pedro; San Diego.
10. A. formidabilis Griffin (= A. cxilis Coe). Monterey
Bay.
Heteronemertea.
11. Micrura verritti Coe. Monterey Bay.
12. M. alaskensis Coe. San Pedro; Monterey Bay.
13. Cercbratulus marginatns Renier. San Pedro.
14. C. albifrons Coe. San Pedro
In the first portion of this report 32 species were enumerated.
Nearly half of these, or the above 14 species, were found also
on the California coast during a single summer. Eleven of
NEMERTEANS 119
these forms, which are common both to the California coast and
to Alaska, were found at Monterey Bay, ten at San Pedro or in
the deep water in the vicinity, while only two were collected at
San Diego, and these were both common in the other two locali-
ties. At San Diego, however, the opportunities for collecting
were comparatively limited, and but a short time was spent at
that place. Of the ten Alaska species found at San Pedro, five
only were collected at Monterey Bay, although in all probability
the others will be found to occur there.
There can be no doubt that future collections will add mate-
rially to the number of forms whose range extends from Alaska
at least as far southward as Monterey Bay or even to Point
Conception. And while this is a considerable range geographi-
cally, yet the environmental conditions of the marine forms are
not greatly different between Monterey Bay, Puget Sound,
Sitka, and the eastern Aleutian Islands. The temperature of
the water is but a few degrees different, and in some seasons of
the year is actually warmer on the coast in portions of Alaska
than it is in Puget Sound or even in the deeper water off the
California coast.
SPECIES NEW TO WEST COAST OF NORTH AMERICA
In addition to the species recorded in the previous paper (pp.
i-no), the following forms have since been studied, and are
described in this article. In accord with Bergendal's recom-
mendation1 the orders Protonemertea and Mesonemertea of
Burger are placed together under the older order Paleonemertea
proposed by Hubrecht.
Paleonemertea.
1. Carinclla frcnata sp. nov. San Pedro, Calif. Not
common.
2. C. attocincta sp. nov. Off San Pedro, Calif., in 50-100
fms. Common.
3. C. cingulata sp. nov. Monterey Bay, Calif., in 14 fms.
Not common.
Hoplonemertea.
4. JVemertapsis gracilis sp. nov. Monterey Bay, Calif.
Not common.
1 Kongl. Vetenskaps-Akad. Forhandlingar, pp. 721-742, 1900.
I2O COE
5. Paranemertes calif ornica sp. nov. Southern coast of
California. Not uncommon.
6. Carcinonemertcs epialti Coe. Parasitic on the crab, JEpi-
altus productus. Monterey Bay, Calif.
7. Amphiporus cruentatus Verrill. San Pedro and Monterey
Bay, Calif. Rather common.
8. Tctrastcmma signifcr sp. nov. San Pedro, Calif. Not
very common.
9. T. nigrifrons sp. nov. Monterey Bay, Calif. Abundant.
10. Z1. bilineatum sp. nov. San Diego, Calif. Common.
11. T. quadrilineatum sp. nov. San Pedro, Calif. Rather
common.
12. T. (CErstcdia) dor sale (Abildgaard) Mclntosh. Mont-
erey Bay, Calif., in 20 fms. Not very common.
13. T. (CErstcdid) rcticulatum sp. nov. San Pedro, Calif.
Common.
Heteronemertea.
14. Taniosoma -ptinnetti sp. nov. Off San Pedro, 50 fms. ;
Monterey Bay, Calif., 5-20 fms. Common.
15. Zygcupolia littoralis C. B. Thompson. San Pedro, Calif.
Common.
16. Linens rubcsccns sp. nov. San Pedro and Monterey
Bay, Calif. Not common.
17. L> flavescens sp. nov. Low water to 50 or more fms.,
off San Pedro, Calif. Common.
18. L. pictifrons sp. nov. San Pedro, Calif. Rather
common.
19. L. albotineatus sp. nov. Off San Pedro and in Monterey
Bay, Calif. Not common.
20. L. ivilsoni sp. nov. Monterey Bay, Calif. Common.
21. Micrura nigrirostris sp. nov. San Pedro, Calif. Not
common.
Besides those mentioned in the above list, which includes
only species which have not been hitherto recorded from the
Pacific coast of North America, a considerable number of forms
described in the first part of this paper (pp. 1 1-84) as occurring in
Alaska have since been found in other localities, notably on the
NEMERTEANS 121
California coast, as mentioned above, and have thus had the
range of their distribution greatly extended.
DISTRIBUTION OF ALL SPECIES KNOWN FROM THE WEST COAST
OF NORTH AMERICA
Including the foregoing and those which are described on the
following pages, 57 species of Nemerteans are at present known
from the Pacific coast, and their distribution as far as recorded
(including my own observations during the summer of 1901
and the study of several other collections) is as follows :
Paleonemertea.
1. Car indict rubra Griffin. Whole Pacific coast of Alaska
to Vancouver Island, B. C.
2. C. sexlineata Griffin. Sitka, Alaska, to San Pedro, Calif.
3. C. capistrata Coe. Prince William Sound, Alaska, to
Monterey Bay, Calif.
4. C.frenata sp. nov. San Pedro, Calif.
5. C. albocincta sp. nov. Off San Pedro, Calif.
6. C. cingulata sp. nov. Monterey Bay, Calif.
7. Carinoma mutabilis Griffin. Vancouver Island, B. C., to
San Pedro and San Diego, Calif.
8. Ce-phalothrix linearis (Rathke) Oersted. Whole Pacific
coast of Alaska to southern coast of California; New Eng-
land; northern coasts of Europe; Mediterranean Sea.
Hoplonemertea.
9. JEmplectoncma gracilc (Johnston) Verrill. Whole Pacific
coast of Alaska to Monterey Bay, Calif. ; northern coasts of
Europe ; Mediterranean Sea ; Madeira.
10. E. burgcri Coe. Southern coast of Alaska to Puget
Sound.
11. Zygoncmertes thalassina Coe. Sitka, Alaska.
12. Z. albida Coe. British Columbia.
13. Ncmerto^sis gracilis sp. nov. Monterey Bay, Calif.
14. Parancmcrtcs -peregrina Coe. Whole Pacific coast of
Alaska to southern coast of California.
15. P. -pallida Coe. Pacific coast of Alaska.
122 COE
16. P. carnea Coe. Pacific coast of Alaska and British Co-
lumbia.
17. P. californica sp. nov. Southern coast of California.
18. Carcinonemertes epialti Coe. Parasitic on Epialtus.
Monterey Bay, Calif.
19. Amphiporus angulatus (Fabr.) Verrill. Bering Strait to
Puget Sound ; New England to Greenland.
20. A. bimaculatus Coe. Southern Alaska; Puget Sound to
Monterey Bay, Calif.
21. A. tigrinus Coe. British Columbia.
22. A. ncbulosus Coe. Pacific coast of Alaska Peninsula.
23. A. cruentatus Verrill. Monterey Bay and San Pedro,
Calif. ; southern coast of New England.
24. A. imparispinosus Griffin. Pacific coast of Alaska to
southern coast of California.
25. A. formidabilis Griffin. Aleutian Islands, Alaska, to
Monterey Bay, Calif.
26. A. pauttnus Punnett. Pribilof Islands, Bering Sea.
27. A. brunneus Griffin. Puget Sound.
28. A. drepanophoroides Griffin. Puget Sound.
29. Tctrastcmma signifer sp. nov. San Pedro, Calif.
30. T. nigrifrons sp. nov. Monterey Bay, Calif.
31. T. bicolor Coe. Kadiak, Alaska.
32. T. aberrans Coe. Pacific coast of Alaska.
33. T. ccecum Coe. Kadiak, Alaska.
34. T. bilineatnni sp. nov. San Diego, Calif.
35. T. quadrilincatum sp. nov. San Pedro, Calif.
36. T. (CErstedia) dor sale (Abildgaard). Monterey Bay,
Calif. ; northern coasts of Europe ; Mediterranean Sea ; east
coast of United States.
37. T. (CErstedia) reticulatum sp. nov. San Pedro, Calif.
Heteronemertea.
38. TcBniosoma -princess Coe. Southeastern coast of Alaska.
39. T. punnetti sp. nov. Monterey Bay and San Pedro,
Calif.
40. Zygeupolia ttttoralis C. B. Thompson. San Pedro, Calif. ;
New England (Woods Hole, Mass.).
NEMERTEANS 123
41. Lineus viridis (Fabr.) Verrill. Southeastern coast of
Alaska.
42. L. torquatus Coe. Pacific coast of Alaska.
43. L. rudesccns sp. nov. Monterey Bay and San Pedro, Calif.
44. L. flavesccns sp. nov. Southern coast of California.
45. L. -pictifrons sp. nov. San Pedro, Calif.
46. Z-. ivihoni sp. nov. Monterey Bay, Calif.
47. L. albolincatus sp. nov. Monterey Bay and San Pedro,
Calif.
48. Micrura nigrirostris sp. nov. San Pedro, Calif.
49. M. verritti Coe. Pacific coast of Alaska to Monterey
Bay, Calif.
50. M. tmpressa (Stimpson) Coe. Bering Strait; Puget
Sound (?)
51. M. alaskensis Coe. Pacific coast of Alaska.
52. Cerebratulus herculeus Coe. Southeast coast of Alaska ;
Puget Sound.
53. C. marginatus Renier. Southeastern coast of Alaska to
southern coast of California ; New England to Greenland ;
northern coasts of Europe ; Mediterranean Sea.
54. C. occidcntalis Coe. Pacific coast of Alaska to British
Columbia.
55. C. longiceps Coe. Yakutat Bay, Alaska.
56. C. montgomeryi C.QQ. Aleutian Islands, Alaska, to Puget
Sound.
57. C. albifrons Coe. Southeastern coast of Alaska to
southern coast of California.
Of the above 57 species known from the Pacific coast of
North America, only 7 have been recorded in other parts of the
world. Of these seven species three — Cephalothrix lincaris^
Tctrastcmma dorsale and Ccrcbratulus marginatus — occur also
both on the east coast of North America and in Europe ; three
others — Amphiporus angutatus, A. cruentatus and Zygeupolia
littoralis — are found in New England, but are not known from
Europe; while a single species — Emplectonema gracile — is
common in Europe, but has not been recorded from the east
coast of America.
124
COE
Although such a large proportion of the species are peculiar
to the Pacific coast, yet in general they belong to common
European genera, and the Nemertean fauna as a whole resem-
bles that of Europe and the Mediterranean far more closely
than it does that of the Atlantic coast of North America.
This has been observed in regard to the general invertebrate
fauna of Puget Sound by Harrington and Griffin,1 and Griffin
noted the same in regard to the Nemerteans. The abundance
of species of Carinetta, the presence of Nemertopsis and of
Emplectonema gracilc, and the close resemblance of a number
of species of Tctrastemma, Amphiporus, Lincus, Micrura and
Cerebratulus to closely related European forms, are instances of
this similarity of faunas.
Certain other collections which are being studied will doubt-
less yield further light on the distribution of the species and
their relationships, as well as the resemblance of the Nemertean
fauna of the Pacific coast to that of other parts of the world.
The distribution of Nemerteans on the Pacific coast of North
America so far as now known is represented in the following table :
KUMBER OF SPECIES KNOWN FROM VARIOUS LOCALITIES ON THE
PACIFIC COAST
Genus.
Alaska.
Pug-et
Sound.
Central
California.
Southern
California.
Total.
PALEONEMERTEA.
Carinella.
3
3
3
3
6
Carinoma.
i
i
i
i
Cephalothrix.
i
i
i
i
i
HOPLONEMERTEA.
Emplectonema.
2
2
i
-
2
ZygonemerteS)
I
I
-
—
2
Ncmertopsis.
-
-
i
—
I
Paranemertes.
3
2
2
2
4
Carcinonemertes.
—
I
—
i
Amphiporus.
6
7
4
2
10
Tetrastemma.
3
—
2
4
9
HETERONEMERTEA.
Tcentosoma.
i
_
I
i
2
Zygeupolia.
-
-
-
i
I
Lineus.
2
—
3
4
7
Micrura.
3
?
2
2
4
Cerebratulus.
6
5
2
2
6
Total.
3i
22
24
23
57
lTrans. New York Acad. Sci., p. 161, 1897.
NEMERTEANS 125
The total number of species already recorded from the region
covered by this report is surprisingly large, considering the few
attempts at their collection. And it should be remarked that
we find in this region not only a large number of species, but
also a surprising abundance of individuals of the species repre-
sented. It now seems probable that this coast will eventually
be found to possess more species of Nemerteans than any other
region of equal size on the globe. And certainly I know of no
locality where so large a proportion of the invertebrate animals
found in a miscellaneous collection belong to the Nemerteans
as on the northwest coast of North America.
KEY TO THE PACIFIC COAST NEMERTEANS DESCRIBED ON THE FOL-
LOWING PAGES
This key is arranged for use in connection with the one published
in the former portion of this report (pp. 7-11), and is likewise based
mainly on superficial and easily distinguishable characters.
Order Paleonemertea.1
Body remarkably long, soft, fragile. Head distinctly marked off
from body, usually broader than neck, often flattened dorso-ven-
trally Carinella.
1. Body large, rather firm, only moderately slender, attaining a
length of 50 cm. or more, pale yellowish or rosy, sometimes
with greenish tinge ; with three longitudinal velvety, blackish
lines and a series of narrow rings of similar color.
C. frenata, p. 129.
2. Body firm, rather stout, upwards of 30 cm. long, deep red with
a series of narrow white rings C. albocincta, p. 136.
3. Body slender, subcylindrical, 15 cm. or more in length, deep
brown, with a series of white rings and four longitudinal white
lines, of which two are lateral, while the other two divide dor-
sal surface of body into three equal parts. . . C. cingulata, p. 138.
Order Hoplonemertea.
I. Body very slender, almost filiform, somewhat flattened ; with four
large ocelli ; with central stylet and two pouches of accessory
stylets. Proboscis sheath about one-third as long as body.
Nemertopsis .
*As stated above, Burger's orders Protonemertea and Mcsonemcrtea are here
united into the older order Paleonemertea Hubrecht.
126 COE
i. Length 15 cm. or more ; very pale brown or whitish, with two
longitudinal dorsal stripes of deep brown ; proboscis provided
with eight nerves N.gracilis, p. 142.
II. Body moderately elongated ; proboscis sheath about one-half to
three-fourths the length of the body ; central stylet well devel-
oped Paranemertes.
I. Four or six pouches of accessory stylets ; proboscis with ten
large nerves. Translucent, pale orange anteriorly, flesh color,
grayish or very pale salmon posteriorly, color much obscured
by deep green of intestinal tract P. californica, p. 144.
III. Body very small and slender ; two ocelli ; proboscis but little
developed, central stylet minute ; no accessory stylets ; parasitic
on crabs Carcinonemertes .
i. Only 4-6 mm. long when sexually mature; color orange or
reddish ; parasitic on the egg masses of Epialtus and per-
haps other crabs C. epialti, p. 151.
IV. Body rather short and thick ; proboscis sheath reaches nearly or
quite to posterior end of body ; proboscis large, central stylet
well developed.
aa. Ocelli do not extend posteriorly beyond the brain ; basis of cen-
tral stylet usually rounded posteriorly.
b. Body not very small. Ocelli visually numerous.
Amphiporus .
1. Rather slender, 10-25 mm« in length; yellow; very con-
spicuous red blood vessels ; five to ten ocelli in single
row on each side of head ; basis very slender, about same
length as stylet A . cruentatus, p . 1 54.
2. Slender, 50-90 mm. in length; proboscis sheath about
six-sevenths the length of body ; proboscis with fifteen
nerves; basis of same length as stylet; ocelli numerous.
A. pazilimis, p. 155.
bb. Body very small. Ocelli few ; usually four well developed
ocelli arranged in a rectangle Tetrastemma.
1. Rather slender, 15-25 mm. in length, reddish-brown
both above and below, except head, which is white
with sharply marked wreath of deep brown color on
dorsal surface. Blood corpuscles red.
T. signifer, p. 156.
2. Usually 20 to 70 mm. in length; head provided with
two pairs of very conspicuous, lateral, oblique fur-
rows. Color very variable, except head which is
NEMERTEANS 127
white with shield-shaped or triangular marking of
deep brown color on dorsal surface. Body deep
purple, deep brown with white dorsal longitudinal
band, reddish with brown flecks, pale brownish or
buff on dorsal surface ; of similar color, but paler,
and often with white longitudinal band on ventral
surface. Blood corpuscles red.
T. nigrifrons, p. 159.
3. Only 5 to 10 mm. in length, flesh color, creamy or
grayish, with two deep brown longitudinal stripes
on dorsal surface T. bilineatum, p. 164.
4. Usually 8 to 12 mm. in length; whitish, with four
deep brown longitudinal stripes, of which two lie
on lateral margins and two on dorsal surface.
T. quadrilineatum, p. 166.
5. Firm, cylindrical, slender, body only 8 to 15 mm. in
length ; flesh color or yellowish, mottled, especially
on dorsal "surf ace, with brownish blotches and dots
of various shades, often mainly collected into a
series of transverse bands.
T. (Oerstedia) dor sale, p. 169.
6. Firm, cylindrical, slender, 8 to 15 mm. in length.
White, with large rectangular and longitudinal,
dark brown markings almost covering dorsal sur-
face. Often with sixteen pairs rectangular marks
and pair of lateral brown lines in addition to bi-
lobed marking on dorsal surface of head. In some
varieties markings fuse together.
T. (Oerstedia) reticulatum,^. 170.
Order Heteronemertea.
A. Head without deep, lateral, longitudinal furrows. Proboscis
musculature of two layers, an outer circular and inner longi-
tudinal layer, without muscular crossings Tcznwsoma.
i. Body large, soft, flabby, 40 to 60 cm. or more in length.
Color brownish red or mahogany ; tip of snout darker
brown with terminal white border ..T. ptinnetti, p. 173.
AA. Head without deep, lateral, longitudinal furrows. Cerebral
sense organs open into pit on margins of acutely pointed
head. Proboscis musculature of two layers, outer longi-
128 COE
tudinal and inner circular layer, with muscular crossings.
Caudal cirrus present Zygeupolia.
i. Rather slender, 5 to 8 cm. in length; head pure white,
acutely pointed ; esophageal region whitish, pale yellow
or flesh color ; intestinal region rose, pale yellow, light
brown or chocolate brown from color of intestinal canal.
Caudal cirrus slender, white. No retractor muscle to
proboscis Z. littoralis, p. 177.
AAA. Head with deep, horizontal furrows. Proboscis of three
muscular layers, outer longitudinal, circular, and inner
longitudinal layers, or of the two former only; muscular
crossings usually present.
a. Caudal cirrus absent. Body long and slender, rounded or
flattened ; very contractile Lineus.
1. Slender, small, usually 10 to 15 mm. long. A single
row of two to four (rarely six to eight) irregular
ocelli on each antero-lateral margin of head. Pink or
rosy flesh color, sometimes with tinge of blue ; deep
flesh color to purplish brown in intestinal region ;
head white L. rtibescens, p. 179.
2. Up to 12 cm. in length; usually three to seven irregu-
larly fragmented ocelli on each side of snout. Yellow,
sometimes with decided tinge of orange, deep ochre or
pale yellow L. flavescens, p. 184.
3. Large, soft and flabby, up to 50 cm. in length ; cephalic
furrows remarkably long. Ocelli wanting. Dark
brown of various shades of color, velvety ; a series of
narrow, pale yellow rings, expanded to diamond-shaped
spots on dorsal surface, usually encircle body, but may
be very inconspicuous. Dorsal surface usually corru-
gated longitudinally with series of very delicate, in-
conspicuous, longitudinal yellow lines. Tip of snout
white, usually with two small orange-colored spots
situated in an area of pale yellow on dorsal surface.
L. pictifrons, p. iSS.
4. Length zoto 15 cm. or more; head broad ; ocelli want-
ing. Dark brown with conspicuous median dorsal
white or light yellowish stripe which expands on head
to form a broad, pear-shaped marking. Usually a
faint reddish line extends along each lateral margin of
body L. albolineatus, p. 193.
NEMERTEANS
129
5. Body rather stout, flattened in intestinal region, fragile ;
length up to 15 cm. or more ; head long and slender,
with long cephalic furrows ; ocelli wanting. Brown-
ish, dark drab or purplish brown, with a series of
very fine white rings, often much obscured ; tip of
head and borders of cephalic furrows white.
L. ivilsoni, p. 195.
aa. Caudal cirrus present. Body rather firm, not provided
with thin lateral margins in intestinal region ; incapable
of swimming Micrura.
I. Of small size, 4 to 8 cm. in length; ocelli wanting.
Deep blood red, sometimes with tinge of purple ;
head of same color, with narrow, transverse, white
crescent near tip of snout which has a small terminal
black or dark brown spot surrounded by red continu-
ous with that of ventral surface.
M. nigrirostris, p. 198.
SYSTEMATIC DISCUSSION OF NEW SPECIES
In the following pages those species recently found on the Pacific
coast of North America which have proved to be new to science are
described in detail from a study of both living specimens and prepared
sections. Only those anatomical features are discussed, however,
which offer peculiarities not found in most related species or which
are of special interest from a morphological standpoint.
Included with the species new to science are certain forms which
have not previously been recorded from the Pacific coast of North
America. Of these only the most striking anatomical peculiarities are
mentioned.
Descriptions of only those genera which were not represented in
the collections described in Part I of this report are given place here.
For the remaining generic descriptions the reader is referred to Part I,
previous article.
i. CARINELLA FRENATA sp. nov.
pi. xv, figs. 5, 6; pi. xxii, figs. 2, 3.
In general shape of body this species resembles those which are
most typical of the genus, as C.superba, C. capistrata, C.sexlineata,
and which have rather firm, moderately slender bodies. Head much
broader than body, rounded or emarginate in front, much flattened
dorso-ventrally, sharply marked off from body by deep, lateral, trans-
130 COE
verse grooves. Head commonly quite as wide as long, but shape liable
to the greatest variation. Proboscis pore and mouth as in related spe-
cies. The color and markings on body are very characteristic and
widely different from those of any described species of the genus.
Size. — Length of body 50 cm. or more, width 2-3 mm.
Color. — General color of body grades from a yellow cream or ochre
in anterior third of body to a sage green in intestinal region. When
filled with ripe ova the greater portion of dorsal surface of intestinal
region becomes a rosy but opaque flesh color or dull rose color.
On this ground color is arranged a series of remarkably sharp, deep
brown, transverse and longitudinal lines and bands. There are three
longitudinal, very distinct, dark brown or black lines throughout the
length of the body, except on the head. One of these lies in the dorso-
median line, while the other two are symmetrically placed on, or a
little beneath, the lateral margins (pi. xv, fig. 5).
The color of the markings is beautiful and has a velvety luster ; in
some lights it appears somewhat iridescent and sometimes shows a
rich, dark blue reflection. Of the three longitudinal lines the median
dorsal line is much broader than the others. It commences on the ex-
treme tip of snout, where it joins a narrow transverse terminal line of
the same color. On the head it is broader than elsewhere and occu-
pies about one-fifth the diameter of head. It continues through the
transverse bands or rings, usually expanding a little where the rings are
joined. In many cases the line can be traced directly through the
transverse bands by a deeper color, as if the two markings had been
painted the one over the other. In the intestinal region the median
line occupies perhaps one-seventh the diameter of body. The two lat-
eral or marginal lines each commence at the broad neck band (the first
transverse band) .
These lateral lines are scarcely more than one-third as wide as the
median line ; they are very sharp and clear cut in the anterior portion
of body, but become more irregular in outline in the intestinal region.
They are never wholly interrupted however. They are cut more
sharply ventrally than dorsally, for on their dorsal side they sometimes
fade out gradually into the general color of body. Like the median
line, the lateral lines broaden out somewhat where they join the trans-
verse bands, and, as is also true of the median line, they often show
corresponding thickenings in places where the transverse bands are not
formed or are very imperfect. A thickening of one line is almost al-
ways accompanied by a corresponding thickening of the other two,
showing clearly where the transverse band would lie if it were present.
NEMERTEANS 131
I
On the anterior third of body these longitudinal lines are situated
directly in the yellow ground color, but in the intestinal region the
median line is separated from the rose-colored ground color of the
mature females by an irregular border of sage green thickly flecked
with whitish dots. This sage green color probably represents the gen-
eral ground color of the worms when the sexual products are absent ;
while the rose color, which seems to make up the general color of the
dorsal surface of intestinal region, is due to the thickly-placed sacs of
ova, which are of a pale rose color. In some regions these rose-colored
spots are separated by a continuous green ground color, as shown in
fig. 5, pi. xv.
The transverse markings are very numerous and are of various
widths from the first two bands, which are more than half as wide as
the body, down to the finest possible lines. Many are extremely fine,
and many others incomplete. As a rule, the wider bands are separated
by one or two much finer ones, and seldom, or never, are two of the
wider bands immediately adjoining. The first transverse marking
borders the extreme tip of head and is narrow and barely visible both
on ventral and on dorsal surface. It extends laterally from end of
median longitudinal line about halfway to the posterior border of head.
The second transverse marking occurs on the neck just back of the
lateral furrows which separate the head from the body. This mark-
ing is broad and shield-shaped on dorsal surface, but is narrower
laterally, while on the ventral surface it is interrupted by the mouth,
which lies exactly in the region which would be occupied by the band
if it were continued. The first band is about half as wide as body.
The second is separated from the first by a distance about equal to
twice the diameter of body, and is a little broader than the first on the
dorsal surface. It forms a continuous band around the whole body,
but on ventral surface it is not much more than half as broad as on
median dorsal surface. The third band is separated from the second
by a distance less than that between the first and second, and is some-
what narrower than either of these. The fourth is separated from the
third by a greater distance than in any other case ; it is as broad as the
second, and is as wide ventrally as on the dorsal surface. The fifth is
very narrow, the sixth broad, seventh narrow, eighth broad. Then
come two imperfect narrow bands, and then a fairly broad one, and so
on, through the remainder of the body, with a broad band usually fol-
lowed by one or two narrow or imperfect ones. An individual meas-
uring 50 cm. in length shows from seventy to one hundred of these
transverse bands. In general the bands decrease in width toward the
132
COE
posterior end of body. Many of the broader bands show a number
(three to twelve) of fine, pore-like, pale dots in midst of dark band
between the median and lateral longitudinal lines. Transverse bands
often broaden out as they join the longitudinal lines (pi. xv, fig. 5),
but a few are interrupted just ventral to the lateral longitudinal lines.
Ventral surface anteriorly same as dorsal surface in color, but with
a more conspicuous flecking of minute whitish dots. In intestinal
region the yellow color gradually assumes a more greenish tone, until
imperceptibly a shade of sage green, or very light olive green, is
reached, and this color extends through to the end of the body. The
green color is more or less tempered and obscured by an irregular coat-
ing of very fine whitish dots which cover ventral surface irregularly,
except near the median line ; similar white flecks are scattered over the
dorsal surface also.
After preservation in formalin or in alcohol the portion of the body
situated immediately posterior to the third black ring becomes deep
slaty blue or blackish in color. This dark color is sharply demarcated
anteriorly, but fades out gradually after extending about as far as the
ninth ring (pi. xv, fig. 6).
The pale ochre color with deep black rings and longitudinal lines is
retained even in cedar oil and in paraffin. The colors, markings and
shape of anterior end of body after preservation are strikingly sugges-
tive of the abdomen of the yellow-jacket wasp ( Vespa).
Proboscis pore is situated just below anterior end of median dorsal
black line. Mouth, as stated above, lies in first transverse band.
In midst of lateral, longitudinal black line, and at the anterior border
of the fourth transverse band (or in it) is a rather conspicuous, color-
less, oval pore, representing the lateral sense organ, or ' side organ.'
Intestinal canal is usually greenish in color — sometimes sage green.
Body often shows a tendency to become constricted through the
transverse black bands, and when broken the rupture takes place in
these bands, as has been noted in other species of the genus.
Cephalic glands but little developed, appearing only as scattered
gland cells lying beneath and beside the rhynchodaeum. No glands
whatever are to be found between the cephalic blood lacunae and the
basement layer of the integument. The integumental gland cells, on
the other hand, are very highly developed, and form an unusually mas-
sive layer beneath the ordinary superficial glandular and ciliated cells.
The secretion of these deeper glands of the integument is apparently
of the same nature as that of the true cephalic glands, which in all
probability serve merely to supplement the supply furnished by the in-
NEMERTEANS
133
tegument. The few glands about the rhynchodaeum do not extend
back as far as the brain.
Integument of body remarkably thick and closely packed with
glands. In the intestinal region the glands are vastly more abundant
on the dorsal than on the ventral surface. It is in this layer that the
pigment which gives the body its characteristic transverse and longi-
tudinal dark markings is situated. Unless stained too deeply the posi-
tion of the markings is distinctly seen in each transverse section.
Proboscis. — Of small size, yellowish or ochre in color. Muscular
and epithelial layers as in other species of genus ; fibrous layer, situated
externally, unusually strong ; proboscis nerves, beneath internal epithe-
-nf
st
FIG. 17. C. frenata. Transverse section of body in region of
nephridiopores. X 25-
Hum, remarkably large and conspicuous. Proboscis sheath well de-
veloped. In the exact region of the efferent nephridial ducts the cavity
of the proboscis sheath becomes divided. A very small chamber,
situated dorsally (text-fig. 17, re), passes backward for some distance,
and in this the retractor muscle of the proboscis extends to the posterior
attachment ; while the ventral chamber (re'}, which is very much the
larger, passes backward but for a short distance where it ends in a broad,
blind sac. The retractor muscle is attached to the dorsal wall of the
smaller dorsal chamber a little distance back in the intestinal region, and
134 COE
the proboscis sheath itself does not extend backward behind the ante-
rior third of the intestinal region.
Body musculature consists of the usual circular and longitudinal
layers, and presents few peculiarities save that the inner circular layer,
which is found in the esophageal region, is remarkably thin, and
does not exhibit distinct dorsal or ventral crossings with the outer cir-
cular layer, as is the case in several other species of the genus.
Immediately in front of the nephridial openings the inner circular
layer becomes several times as thick as it is anteriorly, while it disap-
pears almost entirely just at the beginning of the intestinal region. The
circular muscles of the proboscis sheath present a similar increase in
strength in the same region, and are directly continuous with the inner
circular muscles of the body wall (text-fig. 17, tern). These thick-
enings correspond in nature and position with the enormously thick-
ened internal circular muscles in Carinoma.
Blood System. — Lacunae in head and lateral vessels present no
peculiarities. The pair of vessels situated within the proboscis sheath
originate anteriorly just behind the mouth and extend back nearly
half way to the intestinal region, being connected with the lateral ves-
sels at frequent intervals. Their walls are thickened, much con-
voluted, and appear to be somewhat glandular in nature.
Nephridia. — The nephridial tubules are limited to about the third
quarter of the esophageal region. They do not extend forward quite
as far as the posterior ends of the proboscis sheath vessels. Their
anterior branches are numerous, but of small size. They project in-
ward from the lateral walls of the lateral blood vessels, very much as
in Carinoma. The main longitudinal canals are large, and are situ-
uated in the parenchyma above the lateral vessels, and slightly separated
from them. There is a single main longitudinal canal on each side,
and this is usually about half the diameter of the lateral vessel in the
same region. At its posterior end each of the longitudinal canals en-
larges somewhat, and as usual sends off a large efferent duct (nd, text-
fig. 17) to the dorso-lateral aspect of the body.
Nervous System and Sense Organs. — The cerebral sense organs
are well developed for the genus, although they are not distinctly sepa-
rated from the other nervous tissues, as in the case of C. rubra.1
Yet there is a distinct ciliated canal leading from a specialized lateral
furrow into the nervous tissues above and beside the dorsal brain lobe.
The inner end of this ciliated canal (pi. xxn, fig. 2, cc) passes within
'Coe, Proc. Wash. Acad. Sci., in, p. 13, pi. x, fig. 2, 1901; also previous
article, paged identically.
NEMERTEANS 135
the basement membrane and outer fibrous layer, and is thus well re-
moved from the integument. Here it lies in close contact with the
dorsal surface of the brain, and is closely surrounded by large nerves
(pi. xxn, fig. 2, sn~) from the dorsal ganglia. Externally it termi-
nates on a slight papilla in the middle of the distinct, though short,
lateral groove with which the side of the head is provided. The
epithelium of the groove is also unquestionably sensory in its nature,
and lies in close proximity to the brain, from which it is supplied with
numerous small nerves.
The lateral sense organs, or side-organs, are situated immediately
behind the efferent nephridial ducts, as in other species. They are
rather conspicuous in life, when they appear as colorless oval spaces
in the midst of the lateral longitudinal black lines, and at the anterior
border of the fourth transverse band. They are not very extensive,
but possess a remarkable degree of specialization. The sensory cells
are less than half as high as the neighboring cells of the integument,
so that each of these sense organs appears as a conspicuous oval de-
pression exactly on the lateral margin. The cells, too, are rendered
still more conspicuous from the fact that the secretions situated deep
among the cells assume a deep blue stain with haematoxylin and
orange, while the integumental cells always take on more or less of the
orange color, and are partially obscured by the dark body pigment.
The relation of the sense organ to the other tissues is shown in pi.
xxn, fig. 3.
The brain and lateral nerves present no striking peculiarities.
There are two dorso-median nerves as in several other species of the
genus. The upper nerve lies in the median line, just external to the
outer circular muscular layer, while the lower, or inner, lies directly
beneath the former and just outside the inner circular muscular
layer.
Reproductive Organs. — Sexual products are mature in August.
The eggs are opaque and rose-colored, and give the characteristic rose
coloring to the bodies of the females at this season. They develop in
pouches in the parenchyma above the lateral nerves, and each pouch,
containing from 20 to 50 or more ova, opens directly to the dor-
so-lateral surface of body. The oviducts are completely formed in
advance of the deposition of the eggs, and are in all cases lined with a
distinct layer of small, closely placed epithelial cells. Even before the
eggs are fully mature the small pouches in which the ova of the follow-
ing year are to develop are already established. Several ovarian
pouches are encountered in each transverse section.
136 COE
Habitat. — Several feet below low-water mark on piles of wharf,
San Pedro Harbor, Calif. ; not common. Only sexually mature females
were obtained.
2. CARINELLA ALBOCINCTA sp. nov.
pi. xvi, figs. 4, 5.
Body rather stout for genus, but can become much more elongated
than the figures indicate ; not much flattened, rather firm, less soft than
in C. rubra or C. frenata. Head of moderate size, of variable
shape, broader than neck, from which it is demarcated by an annular
constriction. When body is strongly contracted, however, the anterior
portions become much swollen and wrinkled, and the head withdrawn
into the body until it is partially hidden from view from above. A
pair of very shallow transverse grooves lie on the lateral margins of
head, just in front of neck. Esophageal region rounded, intestinal
region not much flattened, posterior extremity not slender.
Proboscis pore subterminal ; proboscis rather small. Mouth situ-
ated just back of annular constriction marking the neck, of variable
size according to state of contraction of body.
Color. — General color of body usually a beautiful cherry red, some-
times inclining to brick red, and sometimes to purplish red, with a series
of narrow, pure white rings. These rings are all very narrow, hardly
thicker than a thread, though some are much finer than others and are
merely indicated as very delicate hair lines. They are placed at vary-
ing intervals throughout the length of the worm, and most of them
completely encircle the body. There may be 50-100, or more, in a
large individual.
Tip of snout provided with a narrow, terminal border of white,
which reaches back along the lateral margins of head for a little dis-
tance. This terminal white border extends to ventral surface, where
it is fully as conspicuous as from above. When head is extended and
obtusely pointed the white marking is angular, and extends back on
the lateral margins for about half the length of the head. In the angle
of the marking, on the ventral surface, lies the proboscis pore. When
head is contracted strongly the terminal white border appears merely
as a short transverse marking on each side of the proboscis pore.
The first white ring lies on the constricted neck portion, and is
interrupted by the mouth (pi. xvi, fig. 5), so that it is incomplete ven-
trally. The second ring is commonly separated from the first by two
to three times the width of the body. This ring is short and complete,
but is thinner ventrally than on the dorsal surface. The third ring is
about half as far behind the second as the latter is from the first. Then
NEMERTEANS 137
follows a long series of similar rings separated from each other by an
average distance of a little less than the diameter of body in moderate
extension. Of course the separation of the rings depends largely on
the state of contraction of body, for when strongly contracted they are
less than a quarter as far apart as when extended. Nearly all the rings
are complete, but many are thinner ventrally than above. Some are
extremely fine, and others consist of double lines separated by a very
thin line of the red color of body.
The general color of body becomes gradually lighter in intestinal
region, which often has a slightly yellowish tinge. Ventral surface is
of a lighter shade than dorsal, and has a grayish tinge.
After preservation in formalin or in alcohol the body assumes a dull
reddish brown or purplish color, with very faint white rings. An
abrupt change in color usually occurs at the second white ring, the parts
anteriorly commonly being brownish, while those immediately behind
the ring are often deep purple. The white terminal border on the
head remains conspicuous when the worm is not strongly contracted.
Size. — Largest specimen observed was about 30 cm. long and 4
mm. in width, although others were less than half this size.
In internal anatomy the species presents few deviations from that
described for related species of the genus.
Proboscis of rather large size for genus, with muscular layers and
pair of large nerves as in other species.
Musculature. — Fibrous crossings between the external and internal
muscular layers of the body walls are but little developed.
Cephalic glands are voluminous, and occupy a great portion of the
tissues of the head in front of the brain.
Alimentary canal presents no striking peculiarities.
Blood System. — Cephalic blood lacunas of large size. Their
branches pass posteriorly as large lateral vessels, or lacunas, which
send off unusually numerous branches about the esophagus. These
esophageal lacunas pass ventrally well beneath the esophagus, and ex-
hibit abundant anastomoses as in some species of Cerebratulus^ The
rhynchoccel vessels originate some distance behind the mouth region,
though not as far posteriorly as in C. cingulata (p. 141). As in the
latter species, these vessels are short and of much less extent than in
many related forms.
Nephridia. — The excretory tubules are limited to about the third
and fourth fifths of the esophageal region. Anteriorly there are several
canals which branch profusely among the lateral and esophageal blood
1 Notably C. lacteus ; Coe, Trans. Connecticut Acad. Sci., ix, p. 493, 1895.
138 COE
vessels. Farther back these branches unite into about five to eight lon-
gitudinal canals on each side, which lie above the lateral blood lacunas
and do not join until they are near the efferent ducts. Here they unite
to form a rather large lacuna, as in C. cingulata, from the dorsal wall
of which the efferent duct leads to the dorso-lateral aspect of the body.
Nervous System and Sense Organs. — Brain and lateral nerves as
in other species. Cephalic nerves numerous and of large size. Me-
dian dorsal nerve small. Cerebral and lateral sense organs are less
well developed than in the other species described from the Pacific
coast.
Habitat. — Rather common in 50-100 fathoms between San Pedro
and Catalina Island, Calif. The worms live among red algae, having
almost exactly the same color, so that they are not easily discovered
among the contents of the trawl. They are found associated with
Tceniosoma punnetti and exhibit a similar tenacity of life.
3. CARINELLA CINGULATA sp. nov.
pi. xiv, figs. 2-4.
Body long, slender, subcylindrical, resembling C. superba in gen-
eral appearance, and not strikingly different from C. sexlineata and C.
capistrata, which are also found on the California coast. In general
color of body, as well as in being furnished with narrow longitudinal
and transverse white markings, there is considerable resemblance.
The markings on body, however, are arranged in a very characteristic
manner, and differ from those of any known species.
The head is considerably broader than neck, rounded, truncate or
emarginate in front, flattened dorso-ventrally. It is marked off from
body by a distinct annular constriction.
Body often shows constrictions in the annular white lines described
below. Proboscis pore subterminal. Mouth large, situated on the
constricted portion spoken of as the neck.
Length 15 cm. or more ; width about 3-4 mm.
Color. — General tone of body is deep brown, varying sometimes to
chocolate and sometimes to cinnamon brown. When filled with ripe
genital products the general effect of the intestinal region is only pale
brownish. Head much paler than body in color, with two narrow,
transverse, terminal, dark markings — one on either side of the tip of
the snout (pi. xiv, figs. 2-4). In certain states of contraction, these
markings almost meet just dorsally to the proboscis pore.
On the neck is a darker brown transverse marking about one-fourth
as wide as diameter of body dorsally, but which becomes narrower
NEMERTEANS 139
laterally. Below the lateral margin it is quite narrow, and continues
ventrally as a thin line which joins the one from the other side just at
the posterior border of the mouth. This dark nuchal band is bordered
posteriorly by a distinct white band of about the same diameter (pi.
xiv, figs. 2-4).
On the ground color of the body are four longitudinal white bands,
extending with more or less distinctness throughout the length of the
body. These are situated symmetrically, two very near the lateral
margins of body, and the other two dividing the dorsal surface into
three equal parts. The lines all terminate anteriorly in the first white
ring, situated just behind the dark nuchal band mentioned above.
Anteriorly the lines are narrow, but on the approach to the intestinal
region, broaden out very irregularly. Those on the dorsal surface en-
croach so greatly on the general brown ground color as to limit it to a
narrow, brown, median dorsal stripe, and two other narrow brown
stripes on lateral margins. The two lateral lines come to lie ventrally
to the lateral margins, and become so wide as to occupy almost the
whole ventral surface — the brown color being largely replaced by the
brownish white of the longitudinal bands. This may be true of the
worms only at the time when the sexual products are mature, for the
pouches of reproductive elements are light in color and partially obscure
the brown color of body. At other seasons the white lines would
doubtless appear narrower, and the brown color of intestinal region
would be more pronounced.
The body is divided transversely into unequal segments by a series
of narrow white rings, situated at irregular intervals from the head to
the posterior end of the body, as in C. superba. Most of these white
rings are very narrow, but are rendered more conspicuous by being
bordered, sometimes on both sides, and sometimes only on one, by fine
brown rings which are continuous with the general ground color, but
of a darker hue. These brown rings are often more conspicuous than
the narrow white ring beside them, so that some of the annular mark-
ings appear dark, rather than lighter in color (pi. xiv, figs. 2-4) . The
brown rings pass through and interrupt the longitudinal pale bands.
The color of the white markings, especially the longitudinal ones, ap-
pears to be superficial and applied in small, confluent spots, and not
homogeneously.
The segment included between the first and second and that between
the second and third white rings are each fully twice as great as any
other segments. The rings are otherwise fairly regular in position, but
are often interrupted or wanting on ventral surface. Prof. C. B. Wilson,
140 COE
to whom I am indebted for specimens or this and other species of
California nemerteans, states in his notes that " in some specimens
the alternate white rings extend only to the lateral white longitudinal
lines, and do not go entirely around the body."
After preservation in formalin, the region between the second and
third white rings is much darker in color, as commonly occurs in the
genus. As usual the contrast in color is very marked anteriorly, but
shades off gradually beyond the third white ring.
A rather conspicuous rounded pit, of much paler color, situated
exactly on the lateral margin of the body just dorsal to the longitudinal
white band, and on the anterior border of the third white ring, marks
the position of the lateral sense organ on each side.
The worms are prone to break up when captured, the fragmentation
taking place at the white rings, where the body often shows con-
strictions.
Proboscis. — Proboscis sheath extends only a comparatively short
distance into intestinal region, but is well developed throughout the
esophageal region. Just in the vicinity of the nephridiopores the rhyn-
chocoel becomes sharply divided into a smaller posterior cavity and an
enlarged anterior chamber. The anterior chamber continues ventrally
for a few sections as a blind sack beneath the smaller dorsal cavity
into which the posterior end of the proboscis continues. These fea-
tures are similar to those described for C.frenata (p. 133), but are less
pronounced. Proboscis becomes separated from its sheath in the region
of the mouth, and therefore some little distance behind the brain.
It is provided with two large nerves which arise from the ventral
ganglia at points of origin of the unusually large ventral commissure.
The nerves pass dorsally and immediately enter the proboscis sheath,
in the tissues of which they extend posteriorly fora few sections, when
they enter the proboscis at its attachment to the sheath. The arrange-
ment of the epithelial and muscular layers is as in related species.
Musculature and integument present no marked peculiarities.
The basement layer beneath the integument is unusually well devel-
oped. Dorsal crossing of fibers between the internal and external
muscular layers is much better developed than in any of the other
known species from the Pacific coast.
Cephalic Glands. — In addition to the highly developed, deep-
staining masses of glands in the integument of the head, a thick layer
of similar glands is found around the rhynchodseum. These extend
backward nearly to the brain, and are very conspicuous from their
secretions, which stain deep purple in haematoxylin. The condition
NEMERTEANS
in this species is intermediate between that described above for C.
frenata, where these glands occur in the integument only, and C.
rubra? where they occur not only in the integument and around the
rhynchodseum, but are thickly massed in the cephalic musculature as
well.
Blood and Nephridial Systems. — Cephalic blood lacunae and lat-
eral vessels as in other species. Rhynchoccel vessels peculiar in that
they do not appear in the anterior fourth of the esophageal region, and
extend for only a short distance, terminating posteriorly in front of the
anterior end of the nephridial system. During their short extent,
however, they show numerous connections with the lateral vessels.
The nephridia are of the normal type for the genus, with a main canal
above the lateral blood vessel on each side. Anteriorly the canal
branches out on the wall of the blood vessel as usual. The nephridia
are of very limited extent, and occupy less than the middle third
of the esophageal region. Posteriorly each of the main canals exhibits
a sac-like enlargement, with highly columnar epithelium, from the
dorsal wall of which the efferent duct leads to a dorso-lateral aspect
of the body as usual. The efferent duct is not an open tubule, how-
ever, but spreads out in the external circular muscular layer into a
broad, spongy meshwork, from which a small duct leads to the sur-
face.
Nervous System. — Ventral commissure of brain even more massive
than in most related species. Large and numerous cephalic nerves
extend throughout the tissues of the head. Dorsal and buccal nerves
are also of large size.
Cerebral Sense Organs. — Much less highly differentiated than in
any other species of the genus yet described from the region. Ciliated
canals wanting, the sense organ consisting simply of an oval area
with differentiated sensory cells of smaller size and with longer cilia
than elsewhere, and provided with a rather large nerve from the dorsal
ganglion which lies adjacent, and just beneath the well-developed base-
ment layer. The region is always conspicuous by the absence of the
deeply staining glands which are found elsewhere on the body. Thus
the cerebral sense organs differ but slightly in their histological features
from the lateral sense organs found in the vicinity of the nephridiopores.
Lateral Sense Organs. — Small, but very sharply defined. They
are situated on the lateral margins of the body immediately posterior
to the nephridiopores. The sensory epithelium is made up of slender
^oe, Proc. Wash. Acad. Sci., in, p. 14, pi. ix, fig. i, 1901; also previous
article, paged identically.
142
COE
cells, which present a sharp contrast to the neighboring cells of the
integument because of their comparative freedom from secretion. The
general appearance of the sense organ is similar to that figured for C.
frenata (pi. xxn, fig. 2).
Reproductive Organs. — Sexual products mature in September.
Sexual pouches are usually situated dorsally to the lateral blood vessel,
and open on the dorsal aspect of the body.
Habitat. — Monterey Bay, Calif. Shallow water; not common.
Dredged from soft bottom in fourteen fathoms off McAbee's Beach,
Monterey Bay, Calif., by C. B. Wilson, 1899.
Nemertopsis Burger
Fauna and Flora des Golfes von Neapel, Monogr. 22, p. 548, 1895.
Representatives of this genus are characterized by extremely long
thread-like bodies of firm consistency. They resemble Emplectonema
in form, habits and general internal anatomy, but differ in having only
four ocelli, which are symmetrically placed on the head, in armature
of proboscis, and in other anatomical details.
The worms are quite as slender as those of the genus Cephalothrix,
but they do not coil the body in a spiral.
Proboscis sheath very short, proboscis armed with well-developed
central stylet and basis, and with two pouches of accessory stylets ;
cerebral sense organs small, situated well in front of brain. Cephalic
glands usually well developed.
But two species of this genus have thus far been described, both
of which have been found in the Mediterranean. A third form, de-
scribed below, occurs on the Pacific coast of North America.
4. NEMERTOPSIS GRACILIS sp. nov.
pi. xv, fig. i ; pi. xx, figs. 10, 11.
This very slender species bears a close external resemblance in form
and color to N. peronea (Quatr.) Burger, but differs in several fea-
tures of internal organization, especially in the structure of the pro-
boscis armature and the extent of the intestinal caecum.
Like N. peronea the body is extremely long and slender, probably
more so than any other Nemertean found on the coast except species
of Cephalothrix. The body is commonly 10 to 15 cm. or more in
length, and usually less than i mm. in breadth. The head is slightly
broader than the body, which is somewhat flattened dorso-ventrally,
but of nearly equal width throughout its length. Mouth and proboscis
have a common opening.
NEMERTEANS 143
Color. — The color of the dorsal surface of the body is dull whitish
with a tinge of brown, or sometimes decidedly brownish, with two
narrow longitudinal bands of deep brown extending throughout the
length of the body. Each of these brown bands is perhaps one-eighth
as broad as the body. They lie near the median dorsal line, and are
separated from each other by about twice the width of either band.
On the head they lie just internal to the eyes, and do not extend quite
to the tip of the snout (pi. xv, fig. i). They are sharply marked
off from the much paler color between them, but show a tendency to
shade off laterally into the general pale brownish color of the dorsal
surface. Towards the lateral margins the brownish tinge becomes
very inconspicuous and gradually shades off into the whitish or pale
flesh color which covers the ventral surface.
Ocelli. — Four eyes of large size are arranged, as in N. peronea,
to form the corners of a square (pi. xv, fig. i).
Cephalic Glands. — Enormously developed cephalic glands occupy
the greater portion of the head and stretch far back into body, extend-
ing even as far back as the most anterior sexual pouches. In the
esophageal region these glands often occupy more space than the pro-
boscis sheath and esophagus together, and fill up the space usually
taken by the body parenchyma, which is in this species very much
more reduced than in N. peronea.
Proboscis. — The proboscis sheath is not much more than one-third
as long as the body. The proboscis is provided with eight large
nerves which reach back to the stylet apparatus ; the muscular and
epithelial layers are as in Amphiporus. Central stylet of proboscis
slender, provided with an elongated, slender basis, measuring (in a
single specimen) about 0.123 mm. in length by 0.02 mm. in average
diameter.1 The basis is peculiar in being of nearly the same diameter
throughout (pi. xx, figs. 10, u), and not swollen posteriorly. The
shape of the basis alone will readily serve to distinguish the present
species from N. peronea, which has a short conical basis. The central
stylet is rather slender, but my notes unfortunately do not indicate its
dimensions, nor whether it is shorter or longer than the basis. Each
of the two lateral pouches contains usually from four to six slender
stylets. In the preserved specimens there is no evidence that the stylets
have their heads lobed or five-parted as do those of N. peronea.
Alimentary Canal. — The intestinal caecum, which lies in the
median line directly beneath the esophagus, is very short indeed and is
1 But a single basis was measured, so that these dimensions may not represent
average measurements.
COE
without branches. It is much shorter than in TV. peronea and is sepa-
rated from the brain by several times its length, while in TV. peronea
it is described by Burger (loc. cit., p. 549) as reaching nearly to the
brain. In one series of sections of TV. gracilis the caecum itself ex-
tended through seventeen sections only, while there were about seventy-
five sections between its anterior end and the brain.
Nervous System and Sense Organs. — Brain and nervous system
present no important deviations from those of TV. peronea. Cerebral
sense organs small, much elongated, situated far in front of brain and
connected with exterior as usual.
The sexual products ripen late in summer. The ovaries and sperm-
aries lie directly above the lateral nerve cords, but when fully devel-
oped extend also internal to them.
Habitat. — Pacific Grove, Calif., among mussels and other growths
on rocks at low water. Not common. Collected in same locality in
1899 by C. B. Wilson.
Paranemertes Coe
Proc. Wash. Acad. Sci., in, p. 32, 1901.
Three species (P. carnea, P. peregrina, P. pattida) of this genus
have been described from Alaska (loc. cit.).1 A fourth species was
found abundantly at San Pedro and San Diego, California. This form
agrees with those previously described in general anatomical features,
but differs in regard to the number and arrangement of the ocelli, so
that the part of the generic diagnosis which refers to the ocelli (p. 32)
will have to be amended to read, " Ocelli minute, usually numerous,
but sometimes consisting of but a single pair."
5. PARANEMERTES CALIFORNICA sp. nov.
pi. xv, fig. 2 ; pi. xvni, figs. 1-5 ; pi. xxi, figs. 1-9.
Body long, moderately slender, rounded or cylindrical in the short
esophageal region, very much flattened farther back. Intestinal region
much wrinkled when contracted, but smooth when body is well ex-
tended. Head small and acutely pointed in ordinary states of contrac-
tion. The snout, with the ocelli, can be retracted to a very consider-
able extent into the tissues of the head, very much as in some species
of Tceniosoma. Intestinal region flat and ribbon-like with thin mar-
gins wrhich are sometimes bent towards the ventral surface. Posterior
extremity rounded.
1 Also previous article, identically paged. The peculiar fluted or braided ap-
pearance of the stylets (both central and accessory) of P. peregrina was not
described in the previous article but is represented on pi. xx, figs. 14, 15.
NEMERTEANS 145
Size. — Length of largest specimen obtained 45 cm.; width 4-6
mm. ; visual length 10-20 cm.
Color. — The colors are far less opaque than in most species, and
are of such a nature as to give the tissues a remarkably translucent ap-
pearance. Anterior portion of body pale orange inclining to flesh
color, sometimes more decidedly orange and sometimes pale flesh
color. Head distinctly orange, but usually of a pale tint. Brain re-
gion a little more reddish in color. General color of intestinal region
grayish flesh color, or very pale salmon, and somewhat translucent,
but this ground color is so much obscured by the dark green color of
the intestinal tract that in effect this region appears of a greenish tinge.
Commonly the green color extends as a pair of broad, irregular longi-
tudinal bands separated by a pale reddish median band (where the in-
testinal lobes do not show) , and bordered on the lateral margins by
pale, grayish salmon or occasionally by whitish.
The ventral surface is somewhat paler than the dorsal, and the
median reddish band is replaced by the greenish color of the intestine.
On the dorsal surface near the anterior end of the body are two
regions of slightly differentiated color, the one reaching back in the
median line nearly to the brain, the other extending about as far pos-
teriorly behind the brain as is this organ from the tip of the snout.
These show simply as regions of more orange color, and the posterior
region is separated from the flesh colored portion behind by a V-
shaped, orange colored groove. The apex of the groove lies in the
median line and points posteriorly, while its limbs extend obliquely
antero-laterally to the ventral surface where they come nearly into
contact in the median line. A similar, but very indistinct, V-shaped
groove lies just posterior to the anterior orange colored region.
On the under side of the head is a pale area on the tip of the snout
marked off posteriorly by an orange colored line from the still paler
triangular area found on each lateral side of the head.
About 10-15 mm. back of the head (in an individual about 20 cm.
lono") darker areas appear on the sides of the body and increase in
number back to the intestinal region. These indicate the pouches of
the intestinal caeca which stretch forward far into the esophageal region,
and which in the intestinal region are dark green in color. This color
is very permanent and remains after preservation in formalin or in
alcohol, and even after imbedding in paraffin. The green intestinal
lobes nearly all fork distally, and each fork is again divided, but there
is much irregularity in this respect.
The median dorsal band of pale reddish color is due to the color of
146 COE
the fluid in the proboscis sheath. This is not very conspicuous in the
esophageal region, where the proboscis fills most of the space in its
sheath, but in the intestinal region the proboscis sheath is seen to be
filled with a clear red fluid. The red color does not reside in the cor-
puscles, but exists in the fluid itself, while the corpuscles are nearly
colorless. Occasionally an individual has the intestinal region grayish
in color, and in such cases the proboscis sheath is very conspicuous as
a wavy longitudinal, blood-red band about one-fourth the diameter of
the body.
After preservation the esophageal region becomes almost colorless,
but the intestinal tract gives a decidedly greenish color to the rest of
the body.
Proboscis, — When extruded the proboscis appears of a reddish color
due to the red rhynchocoel fluid within it, but when this fluid is pressed
out the proboscis remains colorless. It is of fairly large size, and is
provided with four or six pouches of accessory stylets in addition to the
central stylet (pi. xvm, fig. 2; pi. xxi, fig. 3). The basis of the
latter is rather long and slender, slightly narrower anteriorly, but of
fairly even diameter throughout (pi. xvm, figs. 4, 5 ; pi. xxi, figs.
4-8). The stylet itself is moderately slender and about half as long
as the basis, or sometimes a little more than half as long. There are
usually two or three accessory stylets in each of the four or six pouches.
All the stylets, both central and accessory, show a peculiar darker or
more opaque portion about the head (pi. xxi, figs. 4-9). This
darker portion extends perhaps one-sixth the length of the stylet.
Measurements in a single specimen are : Basis of central stylet 0.36
mm. long, o.i in average width ; stylets about 0.17-0.2 mm. in length.
In each of three specimens sectioned the proboscis was provided
with ten large and distinct nerves. A fourth individual showed an
abnormal condition in that there were twelve distinct nerves in a short
region of the proboscis, although there was but the usual number
(ten) both anteriorly and posteriorly to this region. A fifth specimen
showed eleven nerves. The nerves are often one-half to three-fourths
as great in diameter as the thickness of the longitudinal muscular
layer. Muscular and epithelial layers as usual, but the basement
layer beneath the internal epithelium is so very much thickened that it
practically equals the circular muscular layer in thickness.
Proboscis sheath reaches fully one-half the entire length of the
body, but does not extend into the posterior one-third of the animal.
In this respect the present species agrees well with the other members
of the genus from Alaska. Muscular layers of proboscis sheath enor-
NEMERTEANS 147
mously developed as far posteriorly as the anterior portion of the intes-
tinal region. In few other Hoplonemerteans is the proboscis sheath
so powerful as in the present species (pi. xvm, fig. i).
Ocelli. — The snout when well extended is sharply pointed, but can
be withdrawn to a considerable extent into the tissues of the head.
For this reason the two very small ocelli which are situated near the
tip of the snout (pi. xxi, fig. i) are likely to be overlooked, and are
difficult to see in the living worm. They lie deep in the tissues of the
head and can usually be seen clearly only after the specimen has been
cleared in cedar oil or some other suitable medium. Sometimes instead
of two single ocelli, we find two groups each made up of two or more
minute pigment spots (pi. xxi, fig. 2).
Cerebral Sense Organs. — These are extremely small, measuring
scarcely more than one-tenth the diameter of head in same region, lie
some distance in front of brain, and connect with exterior on latero-
ventral margin of head near tip of snout.
Cephalic glands are voluminous, composing the greater portion of
the tissues of the head in front of brain. They are much interspersed
with connective tissue and muscle fibers, and do not extend posteriorly
to the brain in any considerable numbers. There are a few isolated
S2ibmuscular glands in the esophageal region.
Brain small as compared with the diameter of head, but of the
usual proportions.
Muscular System. — Just in front of brain, and in the region of the
attachment of proboscis to its sheath, a longitudinal muscular layer
arises quite independently and is distinct from the longitudinal muscles
of the body walls. These muscles surround the brain, esophagus, and
proboscis sheath. This secondary longitudinal muscular layer in-
creases greatly in size back of brain, but remains separated from the
musculature of the body walls by a thick layer of parenchyma. Back
of the brain this muscular layer (//«', pi. xvm, fig. i) becomes thicker
than the main longitudinal layer (//«) of the body walls, and its fibers
are larger, more closely placed in their bundles and stain more deeply.
It reaches a considerable distance into the esophageal region, but gradu-
ally the bundles comprising it become more and more separated from
each other by parenchymatous tissue, and gradually they approach
nearer the body walls. Some of the fibers attach themselves to the
proboscis sheath just outside the circular muscular layer. Eventually
they become arranged just internal to the longitudinal muscles of body
walls and form a portion of this layer. Their fibers are much larger,
however, and by an increase in number farther back give rise to the
148 COE
main portion of the longitudinal muscles of body walls. In no other
species, so far as I am aware, has any such condition been described,
though an approach to it is- met with in A. nebulosusj- where the
submuscular glands develop to such an extent as to form a distinct
layer which divides the longitudinal muscular layer into an outer and
an inner portion.
An unusual amount of parenchyma surrounds the muscles, nerves,
proboscis sheath, esophagus and other organs.
Alimentary Canal. — Especially remarkable is the short extent of
esophagus, which separates from rhynchodaeum just in front of brain,
and enlarges posterior to this organ, as usual. Instead of extending
far posteriorly, however, as in most related species, it is only about
twice as long as the distance from tip of snout to brain. It then enters
the dorsal wall of the intestine, the anterior portion of which in this
case corresponds in position and histological structure to the intestinal
caecum of other forms, although it does not end blindly. An ex-
tremely short caecum proper is, however, present and extends forward,
with a few pairs of lateral lobes, for a very short distance anterior to
the posterior opening of esophagus.
The portion of the canal posterior to the esophagus which corre-
sponds to the caecum of other forms extends backward for a long dis-
tance before merging into the intestine proper. This condition has
evidently arisen from the disappearance of the long, slender pylorus
of the typical Hoplonemertean, so that the esophagus opens very near
the anterior end of the long caecum, instead of far back, as in most
other members of the order. I shall therefore refer to the caecum all
that portion of the alimentary canal which lies anterior to the intestine
proper and exhibits lateral diverticula. This will include the short
caecum proper, together with the intestinal canal back as far as the in-
testine proper.
The character of the epithelial lining of esophagus agrees with that
in related species. Where the esophagus enters dorsal wall of caecum,
however, a marked change in the character of its epithelium appears,
as in other forms. Both the caecum and the caecum proper have
the same anatomical and histological peculiarities. Both send off
paired lateral diverticula, which are closely placed together, of rather
small size (pi. xvm, fig. 3), and extend laterally somewhat above the
lateral nerves.
The histological structure of the caecum is as in related species,
'Coe, Proc. Wash. Acad. Sci., in, p. 49, pi. xi, fig. i, 1901; also previous
article, paged identically.
NEMERTEANS 149
except that the cells in the lateral diverticula are provided with a
peculiar granular pigment which gives them a deep green color.
The same pigment occurs in the intestine proper, and is conspicuous
in life, giving the body a large portion of its characteristic coloring.
It is insoluble in alcohol, cedar oil, or the mounting media, and is far
more conspicuous in preserved specimens after they have been placed
in clearing oil. In the caecum the pigment is limited to the lateral
diverticula, and is not found in the central canal.
Toward the posterior end of the caecum the diverticula become
longer and the canal gradually takes on the character of the intestine
proper near the most anterior sexual glands. The intestinal diver-
ticula are very closely placed, and for the most part fork once or twice
into two or four similar branches. These are very conspicuous in
life or after clearing in oil, because of their deep green pigment, as
stated above. Other than in the presence of this peculiar pigment the
histological structure of intestine is as in related forms.
Blood and Nephridial Systems. — In the head, as well as posterior
to the brain, the blood vessels branch out into very numerous branches,
which ramify through the parenchyma, including both that which lies
internal to the inner longitudinal muscles and that between the two longi-
tudinal muscular layers which are described above. The vessels are all
of small size, and extend on all sides above proboscis sheath and beneath
esophagus, as well as laterally. Back toward the intestinal region
they form the usual pair of lateral vessels. The proboscis sheath ves-
sel is as in related species.
In regard to the nephridia, it is necessary to state that no well-
developed nephridial tubules were found, although several series of
sections were examined carefully back as far as the anterior sexual
glands. A number of very fine tubules in the esophageal region may
possibly represent the nephridial system, although they were scarcely
to be distinguished from blood vessels. There were some indications
that these tubules were connected with several very minute efferent
ducts opening to the surface laterally, but this could not be demon-
strated with certainty in any case.
Reproductive Organs. — Sexual products had evidently been re-
cently discharged from individuals collected near the end of July.
The ducts, which still remained open, connected with the dorso-lateral
surfaces of the body.
Habitat. — Individuals of this species are rather common in sand
at low water in San Diego Harbor. They were also obtained from
sandy locations on Dead Mans Island, San Pedro. A single specimen
I5O COE
was collected on a pile of wharf in San Diego Harbor. When handled
the worms exude an abundance of a milky mucus.
Carcinonemertes Coe
American Naturalist, xxxvi, p. 440, 1902.
Parasitic Nemerteans living on various species of Crustacea. Body
small, slender, often filiform, rounded, and of about the same diameter
throughout ; head without distinct lateral grooves, not demarcated from
body. Body not usually coiled or much twisted, but often folded sharply,
so that anterior portion of body lies parallel and in contact with pos-
terior portion. Mouth and proboscis open together ; esophagus ex-
tremely short, opening broadly into intestine through a large muscular
chamber situated immediately behind brain ; intestine broad, with short
lateral pouches which are but little developed in posterior portion of
body.
Proboscis. — Proboscis sheath without muscular walls, consisting
merely of a thin membrane closely applied to the small proboscis. Pro-
boscis but little developed, very small in size, and extremely short, with-
out lateral pouches of reserve stylets, but armed with central stylet and
basis only. Central stylet minute, basis small and slender. Stylet region
of proboscis can be withdrawn but little behind brain ; consequently
anterior chamber is very short, without distinct muscular layers, without
distinct nerves, and without a thickened glandular epithelium such as
occurs in almost all other Nemerteans. Chamber immediately behind
stylet small but muscular, and with a lining of flattened epithelium,
while posterior proboscidial cavity is very short, often almost spherical,
highly glandular, connected closely with the rudiments of the proboscis
sheath and embedded in the connective tissue which lies internal to the
body musculature.
Ocelli two. Cerebral sense organs probably wanting.
Cephalic glands massively developed ; a remarkable development of
submuscular glands extends throughout whole length of body, usually
forming a distinct layer internal to the muscular walls of body, and
often thicker than all other layers of body wall combined.
Body musculature consists of a thin, oblique or circular muscular
layer and a somewhat thicker, but yet weak, longitudinal layer internal
to the former.
Usually oviparous, though fertilization often takes place internally,
and sometimes a portion of the ova of an individual may be retained in
the body until after the development of free-swimming embryos. De-
velopment without complicated metamorphosis.
NEMERTEANS 151
6. CARCINONEMERTES EPIALTI Coe.
pi. xix, figs. 1-9.
American Naturalist, xxxvi, p. 442, 1902.
Body small, rounded, slender, of same diameter throughout ; sex-
ually mature individuals about 4-6 mm. in length and less than 0.5 mm.
in diameter ; head not demarcated from body ; lateral grooves and
cerebral sense organs very inconspicuous or wanting.
Color. — Bright orange, sometimes inclining more to reddish and
sometimes to yellowish. Head a little paler, for the color is largely
due to the intestinal lobes which extend forward to brain.
Ocelli. — A pair of ocelli of irregular outline, but sometimes cres-
cent-shaped, lie about half way between tip of snout and brain. Some-
times the ocelli are irregularly fragmented, and the pigment is arranged
in four irregular masses.
Proboscis. — Proboscis sheath greatly reduced, extending but little
posteriorly to brain, where it becomes united with posterior chamber
of proboscis (pi. xix, fig. 2). The sheath consists merely of few fibers
of connective tissue supporting a very thin flattened epithelium, and
can be seen only in favorable preparations.
Proboscis very minute and short, extending scarcely more than its
own diameter posteriorly to brain (pi. xix, fig. 5). Rhynchodaeum
(fig. 5, r) slender; esophagus separates from proboscis cavity just in
front of brain (fig. 5). Anterior chamber of proboscis (figs. 2, 3, ac)
very small, not as long as the diameter of a brain lobe, lined with thin,
scarcely glandular epithelium. Stylet region swollen (figs. 2-4) and
provided with large and abundant gland cells (^) which open both
into anterior chamber and into the narrow canal connecting this with
cavity behind stylet region.
Basis of central stylet slender, about three to five times as long as
broad (figs. 3-5), measuring about .027-. 033 mm. in length and
.005-. 008 mm. in diameter. Basis slightly larger posteriorly than
near attachment of the very minute stylet (figs. 2, 6). There is no
trace of accessory stylets.
The usual small oval middle chamber lies directly behind stylet
region and connects with anterior chamber by a canal (figs. 3, 4) which
passes close beside the basis of the central stylet and which, though
narrow, is broader than in many other Hoplonemerteans. Middle
chamber, behind the stylet, is highly muscular, lined with flattened
epithelium, and is often filled with fluid containing an abundance of
granules resembling hardened secretions (fig. 3). These apparently
originate in the posterior chamber, as described below.
152 COE
The proboscis now bends sharply on itself in ordinary states of con-
traction and ends in an oval chamber with small lumen and very
massive glandular walls (figs. 2-5, pc). The cells lining this chamber
are highly columnar, irregularly arranged in several layers, and are
thickly packed with secretions which have great affinity for ordinary
stains. Posterior chamber closely imbedded in the surrounding con-
nective tissue (figs. 4, 5), and this appears to be connected with the
muscular walls of esophagus. Its movements are doubtless to a great
extent dependent on the contractions of esophagus, which, as described
below, is converted into a sort of muscular pharynx.
Body Watts. — Outer epithelium as in other genera, and richly
provided with glands.
Muscular layers of body wall consist of a thin, external circular or
oblique layer of muscles and an internal longitudinal layer (figs. 7, 8),
somewhat thicker than the former, but yet thinner than in most related
genera. Lateral nerves occupy the usual places internal to longi-
tudinal muscular layer. In this species, however, they lie internal
also to the thick layer of submuscular glands (figs. 4, 7, 8), and
therefore nearer center of body than in other genera where these glands
are not so highly developed.
There is very little body parenchyma, the intestine filling most of
the space internal to the glandular layer, except at the time when
genital products are developing.
Cephalic Glands. — Throughout the head the tissues are crowded
with cephalic glands. Those situated more anteriorly open mainly on
tip of snout (text-fig. 18; pi. xix, fig. 5, eg), but farther back they
open directly outward on all sides of body.
Back of brain they pass gradually into submuscular glands which
extend as a distinct layer throughout entire body. The glandular
cells composing this layer open directly outward to the surface (figs.
4i 7» 8, sg) and are situated on the whole circumference of body im-
mediately internal to the longitudinal muscular layer. The glandular
layer is in most regions so massively developed that it exceeds in
thickness all other layers of body wall combined. The secretions of
these glands furnish the sticky mucus by means of which the worms
cling so tenaciously to the crab or to other objects.
Alimentary Canal. — The esophagus, which leaves the rhyncho-
daeum just in front of brain (pi. xix, fig. 5), passes beneath the ventral
commissure as a narrow tube lined with rather flat cells, as in other
genera. Just back of brain, however, it becomes enormously enlarged
with high, columnar, ciliated epithelium, richly provided with gland
NEMERTEANS
153
:4V 4-r -UK
cells. This portion of esophagus is highly muscular and somewhat
barrel-shaped (fig. 5, e) , projecting a little way backward into the broad
intestine which immediately follows posteriorly. Its posterior portion
is therefore surrounded by the intestine, indicating rudiments of the
intestinal caeca found in other genera. Intestinal canal broad, with
short lateral pouches
which become very
much reduced toward
posterior end of body.
Nervous System. — . , .,,,
The nervous system //.V''«.-«,»
» • / '* * *l I • '
shows few deviations / :/'•/'• J ', *• •
from that in related gen- ' '**-*' '* '
era. Brain fairly well
developed. From dor-
sal lobes a pair of large
nerves pass anteriorly to
eyes and anterior por-
tions of head. These
are easily seen in living
worms. No indications
of cerebral sense organs
were found either when
the specimens were
stained in toto or when
examined in sections.
Reproductive Or-
gans.— The pouches of FlG l8 c ^jaM. Horizontal section through
genital products become anterior portion of body ; somewhat diagrammatic ;
enormously developed eg, cephalic glands, ac, me, pc, anterior, middle and
and encroach greatly posterior chambers of proboscis. X 125.
upon the intestinal canal at time of sexual maturity (rl. xix, figs. 7,
8). Genital pouches extend farther forward than in almost any other
Nemertean, reaching very nearly to brain. Ovaries (fig. 7, ov} regu-
larly paired, with a single large pouch containing usually from twelve
to thirty ova between each pair of intestinal lobes. Spermaries, on
the other hand, far more numerous, surrounding intestinal canal on all
sides. As many as fifteen or more separate spermaries (fig. 8, t ) are
sometimes found in a single transverse section of the body. As in
most parasitic animals the abundance of sexual products is greatly in
excess of that in related nonparasitic forms.
154 COE
Habitat. — This is a much smaller and less slender species than C.
carcinopkila (Kolliker) Coe when sexually mature, and differs from
it in regard to size of posterior chamber of proboscis, in the stylet ap-
paratus, and in many other anatomical details, although the differ-
ences are not very considerable.
In general appearance, in color, arrangement of ocelli, esophagus,
intestine, and brain the two species are very similar. C. carcinophila
lives on the gills and among the egg masses of various species of crabs
on the Atlantic coast of North America and on the coasts of Europe.
C. epialti also lives when sexually mature among the egg masses of
a crab — in this case Epialtus productus, the common kelp crab of
the California coast.
Upwards of one hundred of these little worms were found among
the eggs of a single crab at Monterey, Calif., September 3, 1901. In
practically all, the sexual products were nearly mature, but no eggs
were laid in confinement. The worm lived only a few days in a dish
of sea water and appeared less hardy than the species on the Atlantic
coast. I was unable to determine whether the worms pass their early
life on the gills of the crab, as does C. carcinophila, but suspect that
this may be the case.
7. AMPHIPORUS CRUENTATUS Verrill
pi. xx, figs. 1-6.
Proc. U. S. Nat. Mus., n, p. 184, 1879.
Trans. Connecticut Acad., vm, p. 399, pi. xxxm, figs. 7, 8 ; pi. xxxv, fig.
3, 1892.
This species, which has previously been found only in New England,
is fairly common among various growths on the rocks of the break-
water at San Pedro, Calif. A number of specimens were also obtained
from piles at Monterey, Calif.
Body small, soft, rather slender, usually 10-25 mm- m length, of a
pale yellow, bright yellow, or sometimes flesh color, and having very
conspicuous vessels with deep red blood, the color of which resides in
the large, discoid corpuscles.
Head slender, with inconspicuous oblique furrows placed far back
from tip.
Ocelli usually five to ten on each lateral margin of head (pi. xx,
fig. 6; text fig. 19), usually well separated, irregular in size and posi-
tion ; anterior ocellus on each side distinctly the largest and situated
more superficially.
Proboscis very long and large, of a pale, slightly pinkish color.
NEMERTEANS
155
Central stylet slender and acutely pointed, having a very slender basis
of about the same length as stylet.
Basis peculiar in that it is no wider,
and is often narrower, posteriorly than
at attachment of stylet. It is often irreg-
ular in shape (pi. xx, figs. 1-5) and
about five or six times as long as its
average width. Measurements vary
from 0.07 to o.i mm. in length and
0.013 to 0.017 mm. in width. Two
pouches of accessory stylets contain
from two to four slender stylets each.
Proboscis sheath reaches very nearly to
posterior end of body. Retractor mus-
cle of proboscis composed of about six
strong fibers, attached in two groups
to wall of sheath.
Blood system as in related spe-
cies. In every individual of a large FIG. 19. A. cruentatus. Dia-
number of specimens the dorsal, or Sram of anterior portion of body,
proboscis sheath vessel originated ante- showinS ocdli 5 *'• brain 5 ™> ^
. , , , . , , , lv> cephalic, dorsal and lateral
norly from the right lateral vessel (text blood yessels> x^
fig. 19).
8. AMPHIPORUS PAULINUS Punnett '
Proc. Zool. Soc. London, p. 92, 1901.
Punnett describes this new species from several specimens collected
by Professor D'Arcy Thompson in the Pribilof Islands. This is de-
scribed as being a slender form, 50-90 mm. in length and up to 4 mm.
in greatest diameter. Color in life unknown, but after preservation
the worms assume a pale yellowish brown color dorsally, and are
almost white ventrally. Submuscular glands well developed, reaching
back to intestinal region. Intestinal cseca do not reach nearly to the
brain. Proboscis sheath extends only about six sevenths the length of
the body ; the proboscis is about three fourths as long as body and
contains fifteen nerves. Its armature consists of central stylet and
two pouches with four reserve stylets each. Basis is same length as
central stylet. A single efferent nephridial duct lies on each side.
Cerebral sense organs small, situated immediately in front of brain.
There are numerous ocelli.
1 For the sake of completeness this species is included here, although it has
not been studied by the writer.
COE
9. TETRASTEMMA SIGNIFER sp. nov.
pi. xiv, figs. 9-11 ; xxi, figs. 10-12.
Body of moderate proportions or somewhat elongated for the genus,
rounded throughout ; not very changeable in shape. Head of mod-
erate size, somewhat narrower than body, marked off from parts im-
mediately following by rather conspicuous oblique lateral grooves ; a
second pair of similar oblique grooves lies farther forward on head, as
shown in pi. xiv, fig. 1 1 .
Color. — General color of body deep reddish brown or purplish
throughout esophageal and intestinal regions. In the intestinal region
the color is rather more opaque than it is farther
forward. Ventral surface of the same general
color as dorsal, but of a duller tone, and often
much paler in the median line. This paler
median band is sometimes quite distinct in the
anterior esophageal region. A little back of
the posterior pair of oblique lateral furrows the
reddish color of body suddenly ceases, and the
whole head is white, or colorless, except for a
characteristic large dorsal marking. This mark-
ing is of a deeper brown and less reddish color,
and is usually situated just anterior to the poste-
rior pair of ocelli, although the relative position
of these parts is largely dependent on the state
of contraction of the head.
In shape the marking resembles a wreath in
FIG. 20. T. signtfer.
Outline of anterior por-
tion of body, showing heraldry (pi. xiv, figs. 9-11 ; text fig. 2o),hav-
shape of cephalic mark- ing a transverse, posterior or basal portion from
ing and arrangement of which two semicircular branches pass anteriorly,
ocelli. X35- but do not usually join. A fourth portion of the
figure passes forward in the median line from the basal portion to the
anterior ends of the semicircular lateral bands, but does not usually
join them. All these parts of the marking have irregular edges so
that the wreath-like effect is made still more striking. The general
effect is often that of an open wreath with vertical crossbar and sub-
stantial base, the whole figure appearing upon a white field. The
wreath is not always open, for one or both of its anterior ends may
join the anterior end of the median longitudinal bar. The whole
figure is surrounded by white, and is separated from the reddish
brown of the esophageal region by a fairly wide band of the same
NEMERTEANS 157
color. This white color also covers the whole ventral side of the
head, as is shown in pi. xiv, fig. 10.
After preservation, and even after imbedding in paraffin, the dark
brown color of body and the white anterior portions with the peculiar
cep halic marking remain almost as distinct as in life.
Blood vessels are distinguished easily because of their reddish color.
This color is resident in the oval or rounded discoid corpuscles, as in
several other species of the genus. The corpuscles are very flat, with
several small pigment bodies in each.
Ocelli, — The four ocelli are of rather large size, and arranged
nearly in the form of a square (text fig. 20) . They lie deep in the
tissues of the head, and occupy a variable position as regards the mark-
ing on the head, but in ordinary states of contraction the posterior pair
lies just behind the basal portion of the marking, while the anterior
pair is situated beneath the antero-lateral portions of the wreath, as
indicated in pi. xiv, figs. 9-11. As seen in sections, the eyes lie deep,
in the midst of the cephalic tissues.
Size. — Length about 15-25 mm. ; width commonly less than i mm.
Proboscis. — Of moderate size ; whitish or pale flesh color. Mus-
cular and other layers as in related species. There are ten proboscidial
nerves. Central stylet slender, acutely pointed; basis conical, opaque
and granular in posterior third (pi. xxi, figs. 10-12).
Body Walls. — Composed of the usual layers. The pigment which
gives the body its color is found in a thin, but very dense, layer be-
tween the integument and the basement layer, and to a less degree
among the bases of the epithelial cells of the integument. It is even
more dense in the cephalic marking than elsewhere. Nearly the whole
space enclosed by the body walls is occupied by the internal organs, so
that there is but very little body parenchyma.
Submusctilar glands are almost entirely wanting. Cephalic glands
are but little developed. Practically the only glands, therefore, which
open at the surface of the body are the integumental glands. It was
noticed in life that the worms of this species were remarkably free from
mucus. There are numerous forms in which an abundance of mucus
is associated with an unusual development of the submuscular glands
(in Carcinonemertes epialti, for example), so that it seems probable
that these glands furnish a supply of mucus accessory to that of the
integument.
Alimentary Canal. — A pair of very slender intestinal caeca extend
forward to abut against the posterior faces of the dorsal brain lobes.
For a considerable distance behind their anterior ends these caeca are
158 COE
less than half as great in diameter as are the lateral nerves, close beside
which they lie. Quite anteriorly they are situated above the nerves,
then gradually assume positions immediately internal to them, and farther
back, and after increasing somewhat in size, come to lie beneath, as well
as internal to, the nerves. At a point about one-fourth the distance from
the brain to the most anterior sexual glands the ca?ca of the two sides
join to form a single broad chamber beneath the esophagus. This
is provided with a few broad lateral pockets, which are but slightly
differentiated from the main chamber. At about two-thirds the dis-
tance from the brain to the most anterior sexual glands the narrow
esophagus empties through the dorsal wall of the broad intestinal cham-
ber, which shows but slightly developed lateral pouches in front of the
anterior sexual glands, although they are well developed farther back.
The anterior portion of the esophagus is lined with highly columnar
ciliated cells, closely packed together with massive gland cells. These
latter gradually disappear more posteriorly until there is only a single
layer of short cells filled with a clear cytoplasm and provided with
long cilia, which line the narrow posterior portion of the esophagus —
or the ' pylorus ' as it is called by Burger.
Nephridial and Blood Systems. — The nephridia are remarkably
limited in extent, but consist of unusually large tubules. They are
situated above the lateral nerves, beside the esophagus, and immedi-
ately behind the brain. Two large canals often occur on each side,
one of which lies above and the other below the slender intestinal
caecum. After a very short extent, they unite to form a remarkably
large efferent duct on each side, which passes above the lateral nerves
to open on, or immediately below, the lateral margin of the body as
usual. In no other species of Nemertean have the efferent ducts been
found to be of such great volume as compared with the size of the body.
The blood vessels show no remarkable deviations from those in
related species. Many of the larger vessels are filled with large, oval
corpuscles, with very distinct nuclei.
Sense Organs. — The cerebral sense organs are voluminous and
highly specialized. They are situated immediately in front of the
brain. The large duct with which each communicates with the exter-
ior passes anteriorly to the ventro-lateral margin of the head as usual.
The brain shows a correspondingly high degree of development, being
remarkably large for the size of the body.
Reproductive Organs. — Sexual products are mature in August.
Habitat. — San Pedro Harbor, Calif., on piles of wharf; off San
Pedro, in 3-6 fms. on hold-fasts of kelp ; not very common.
NEMERTEANS 1 59
10. TETRASTEMMA NIGRIFRONS sp. nov.
pi. xv, fig. 7; pi. xvi, figs. 6-9; pi. xvii, fig. i ; pi. xx, fig. 16;
pi. xxi, figs. 15-23.
Body of moderate proportions, or rather slender for genus, rounded
throughout. Blood corpuscles red in color. Head more variable in
size and shape than in many related species ; provided with two pairs
of lateral, oblique furrows of more than ordinary distinctness. In pi.
xv, fig. 7 and pi. xvi, fig. 6, the head is represented as being well ex-
tended, while in pi. xvi, fig. 8, and pi. xvii, fig. i, it is considerably con-
tracted. The anterior pair of furrows lies between the anterior and
posterior pairs of ocelli ; each furrow extends on dorsal surface from
lateral border obliquely backward toward the median line, but fades
out gradually before meeting its fellow ; on the ventral side the fur-
rows extend obliquely forward and join in the median line not far
behind the rhynchodaeum opening (pi. xvi, fig. 7). The posterior
pair of furrows is situated just behind the posterior pair of ocelli, and
serves to mark off the head from the body. Conspicuous lateral con-
strictions at this point often accentuate the position of these furrows.
They, too, pass obliquely backward on dorsal surface, but are not very
distinctly marked in most cases.
Color. — In peculiarities of color and markings this species presents
more variations than any other Nemertean described in this paper.
A superficial examination led me to believe that no less than three
species were represented by individuals which on more extended
study were found to present all degrees of intergradation. They all
agree in internal organization, and are all similar in having a whitish
or pale yellowish head provided with a dark, dorsal marking of vari-
able size and shape. The under side of head is whitish or very pale
in all varieties. All agree, moreover, in having a deep red color in
the blood corpuscles. The three more distinct color varieties suggest
the varietal designations purpurcttm, bicolor and pallidum.
Variety purpuretim (pi. xvii, fig. i). — Head opaque white, with
large, shield-like dorsal marking of very dark brown color. This mark-
ing is rounded in front, and deeply bilobed in the median line ; it is
somewhat wider posteriorly and commonly shows two conspicuous
indentations near its posterior end ; posterior border of marking
usually straight. The white color of head extends a short distance
back of the marking, or as far as the posterior oblique furrows, where
it abruptly changes to a deep rich purple, which color extends to pos-
terior end of body. The individual shown in pi. xvii, fig. i, is repre-
sented as somewhat contracted, and the white color behind the marking
160 COE
is consequently narrower than where the head is more fully extended.
The line separating the white color of head from the purple of body
is as sharp as possible. Ventral surface of same general color as the
back, but somewhat paler and with a more reddish tinge ; often with
a much paler median band anteriorly. Color of ventral side of head
whitish, separated abruptly from body color on a line corresponding
to a similar line of demarcation on dorsal surface. Some of the purple
individuals become reddish brown in intestinal region.
Variety bicolor (pi. xvi, figs. 8, 9). Head whitish with tinge of
brown, with broad, triangular dorsal marking of dark brown. This
marking is widest behind, with its obtuse apex in the median line
anteriorly. Behind the head the color of body abruptly changes to
deep rich brown, with a narrow median band of white. The brown
color is deepest along the borders of the median white band, and is
much paler laterally ; toward the lateral margins of body it becomes
pale brownish, and this color continues to the ventral surface, becom-
ing gradually paler toward the median line. The ventral surface is
therefore pale brownish, and this color continues on the ventral side of
the head.
The third variety, fallidum (pi. xv, fig. 7), is much less deeply
colored than those above described, and has a much narrower marking
on the head. The marking is of the same deep brown color as in the
other varieties, but is often less than one third as wide as the head.
It is commonly acutely triangular with its pointed apex in the median
line anteriorly. In this variety the ocelli are situated about half-way
between the narrow marking and the margins of the head. The
general color of head is whitish with a faint tinge of brownish. The
whole body back of the head is of a pale brownish or buff color, or it
is occasionally whitish with a faint tinge of brown. Oftentimes a
paler, or whitish, line extends longitudinally on the dorsal surface,
sometimes reaching posterior end of body. Ventral surface in this va-
riety is pale buff throughout. The red blood vessels are naturally
more conspicuous than in the darker varieties.
A variety which occurs on piles at San Pedro is deep flesh color
with tinge of orange dorsally, and with dark brown cephalic mark-
ing. Ventral surface is grayish.
Another variety, common on the piles at Monterey, is shown in pi.
xvi, figs. 6, 7. In this the head is opaque white with a moderately
large wedge-shaped dorsal marking of dark brown color on head.
Back of head and throughout body the color is reddish brown,
thickly sprinkled with minute dots of darker color. Ventral surface
NEMERTEANS l6l
of head white, and this white color extends as a narrow band back-
ward along ventral median line as far as the intestinal region (pi. xvi,
%• 7)-
In some individuals of the other varieties mentioned a much paler
band extends backward along the middle of the ventral surface in the
esopLageal region, and this band is often directly continuous with the
pale color of the ventral surface of the head. In most cases this ven-
tral band is not sharply demarcated, but shades off gradually into the
darker color toward the sides of the body.
Other specimens are pale brick red inesophageal region, and chest-
nut brown posteriorly, with ventral surface grayish or pale reddish.
In fact, there are all possible gradations between the most extreme
color varieties.
Pigment to which color of body is due resides among the basal por-
tions of the cells in the integument. The pigment of the cephalic
marking, on the other hand, is situated among the cephalic tissues
internal to the circular muscles. When the worms are kept for some
time in stale sea water these outer tissues, including the pigment, are
sloughed off, but the worms, which are very hardy, remain alive and
crawl about for several hours more in this condition. Such worms
are pale pinkish or flesh color with the ocelli and red blood vessels
showing very conspicuously.
The general brownish color on the dorsal surface of the worms is
retained after long preservation, and the cephalic marking is perfectly
distinct even after imbedding in paraffin.
Ocelli. — The four medium-sized ocelli lie deep in the tissues of the
head and occupy a variable position as regards the cephalic marking.
Their position also varies greatly according to the state of contraction
of anterior portion of body. When head is somewhat contracted the
ocelli form a square (pi. xvi, fig. 8), but when well extended (pi. xv,
fig. 7 and pi. xvi, fig. 6) the distance between anterior and posterior
pairs of ocelli is considerably greater than between the two ocelli of
the same pair. In an occasional specimen one or more of the ocelli
are double, owing, no doubt, to fragmentation of the normal ocellus.
Size. — Largest specimens found were about 70 mm. long, and
about 2 mm. in diameter, while the majority of sexually mature indi-
viduals were not more than 20-30 mm. long and i mm. wide. Many
immature individuals were of much smaller size.
Proboscis. — Proboscis sheath extends to posterior end of body.
Proboscis pale, often slightly pinkish. It is provided with ten con-
spicuous nerves. Basis of central stylet of the ordinary conical form,
l62 COE
with swollen, rounded posterior end. As shown in pi. xxi, figs. 16-23,
there is considerable variation in size and shape of basis. Figs. 22 and
23 were drawn from two specimens of the same color variety collected
at the same time, and yet the difference in size and shape of basis is
greater than commonly occurs between two related species. The
central stylet is of moderate proportions, not particularly sharp, and is
usually a little more than half as long as basis. There are commonly
three stylets in each of the two lateral pouches, although there are
sometimes four, or occasionally only two in one or both of the pouches.
Measurements of several specimens show the following variations :
Length of Basis. Diameter of Basis at Widest Part. Length of Stylet.
.17 mm. .10 mm. .08 mm.
.15 .08 .09
.14 .07 .08
.14 .06 .07
.12 .05 .08
.11 .05 .07
.11 .05 .06
The length of the longest basis measured is therefore more than
half as long again as the shortest, while the diameter of largest is just
twice that of smallest. The length of stylet, on the other hand, is
much less variable, the longest being only half as long again as the
shortest.
The posterior portion of basis is dark and granular (pi. xxi, figs.
1 6, 17), but gradually assumes the usual translucence in the anterior
half.
Body Watts. — Integument and musculature as in related species.
Pigment of dorsal surface situated among the basal portions of the
integument. Basement layer of body walls thicker than in most
species of the genus, and the species is also peculiar in having a rather
large amount of body parenchyma separating the organs of the body.
This parenchyma extends forward into the head in front of the brain.
Cephalic glands are fairly well developed and surround the rhyn-
chodaeum on all sides, but they do not extend back as far as the brain.
Submuscular glands are wanting.
Alimentary Canal. — The rhynchoccel separates from the esopha-
geal opening well in front of the brain, so that the rhynchodaeum is un-
usually short. When the proboscis is partially extruded, the esophageal
opening (mouth) appears almost like a separate aperture. A pair of
slender diverticula of the intestinal caecum reach forward to the dorsal
brain lobes. They extend backward above the lateral nerves for some
NEMERTEANS 163
distance, and then unite with the main caecum, situated beneath the
esophagus, as usual. Other pairs of slender diverticula are given off
at intervals from the main caecum, and these also pass forward to the
dorsal side of the lateral nerves and end blindly at their anterior ends.
The posterior portion of the esophagus is not very slender, and it unites
with the intestine in front of the most anterior sexual glands.
Blood System. — The blood vessels are often very conspicuous in
life, especially on the ventral surface of the body, because of their deep
red color (pi. xvi, fig. 9). The coloring matter resides in the cor-
puscles themselves, which are oval and discoid. The union of the
three longitudinal vessels at the posterior end of body is especially con-
spicuous, and corpuscles are seen to pass freely, but without regularity,
in either direction in the same vessel. Their movement, either back-
ward or forward, seems wholly dependent on the contraction of par-
ticular parts of the body, and is not controlled by any direct contrac-
tion of the walls of the vessels.
The blood vessels are also conspicuous in sections because of their
content of large corpuscles with deeply staining nuclei. In many cases
the vessels are so closely packed with these corpuscles that they are
more conspicuous in the stained sections than most of the other organs
of the body. The dorsal vessel lies within the rhynchocoel in the an-
terior portion of its course. The general relations of the vessels are as
in related species, and the presence of the corpuscles allows the course
of even the smaller vessels easily to be followed. Similar corpuscles
are found abundantly in the rhynchocoel.
Nephridia. — The nephridial tubules extend from the brain region
well back toward the end of the esophageal region. The tubules are
largest anteriorly, where they ramify in the body parenchyma both
above and below the lateral nerves, as well as beside, and a little in
front of, the brain. There is a single pair of large efferent ducts, and
usually one or more pairs of smaller ones. The large ducts are situ-
ated about on a level with the posterior ends of the dorsal ganglia, and
pass obliquely downwards to open somewhat below the lateral margins
of the head. Sometimes a second, smaller efferent duct lies near the
larger one. Well back in the esophageal region a pair of smaller
efferent ducts pass above the lateral nerves to open a little below the
lateral margins of the body as usual. These smaller accessory ducts
are not found in all individuals.
Cerebral Sense Organs. — These sense organs are situated some
little distance — about their own diameter — in front of the brain, and
somewhat nearer the ventral surface. The large ducts which com-
164 COE
municate with the exterior pass anteriorly and ventrally to open on the
latero-ventral aspects of the head, as usual.
Reproductive Organs. — Sexual products were found to be nearly
mature early in September, but none of the eggs could be made to de-
velop at this time by artificial fertilization. They are probably usually
discharged late in September or in October. The sexual glands do
not extend forward as far as the opening of the esophagus into the in-
testine, as they do in many related species. They are surrounded by
a considerable amount of parenchyma, except when fully mature.
The genital ducts were preformed as far as the basement layer of the
body walls during the latter part of August. These ducts all open on
the dorsal surface of the body, although in the male some of the sper-
maries are situated ventral to the lateral nerves. In such cases a long
duct from each spermary passes internally to the nerve to open through
the body walls above the lateral margin, as in the case of the glands
situated dorsally.
Habitat. — Among algae between tides, Pacific Grove; common.
On piles of wharf, Monterey ; abundant. In similar situations, San
Pedro Harbor ; not common. Collections of bryozoa and small alga?
from piles at Monterey always furnished many specimens of this con-
spicuous, but variously colored species when left standing a short time
in sea water. As noted above, these worms are very hardy, moving
about actively for several hours after the pigmented integument has
been sloughed off. When placed in formalin they do not break up
spontaneously, but usually die intact and well extended.
ii. TETRASTEMMA BILINEATUM sp. nov.
pi. xiv, fig. 6 ; pi. xxi, figs. 13, 14 ; pi. xxn, fig. 4.
A minute species, measuring when sexually mature only 5 to 10
mm. in length, and less than a millimeter in diameter. Body rounded
throughout, of moderate proportions, but rather shorter and stouter
than in many related species. Head of moderate size, provided with
two pairs of faintly-marked oblique grooves, but not distinctly sepa-
rated from the parts following.
Color. — General color of body flesh color, creamy, or grayish,
with two very conspicuous deep brown stripes extending along the
dorsal surface nearly the whole length of the body. The longitudinal
stripes are sometimes reddish brown and sometimes deep chocolate.
The ventral surface is mainly grayish. Anteriorly the stripes ter-
minate somewhat in front of the ocelli, but sometimes reach the very
tip of the snout. Posteriorly they extend nearly, though not quite, to
NEMERTEANS 165
the posterior extremity of the body, and are sharp and conspicuous
throughout. The diameter of each stripe is perhaps one-sixth the di-
ameter of body, and the two are separated about twice the diameter of
each. In ordinary states of contraction the stripes are not quite as
widely separated as are the ocelli, so that they pass medially to the
latter. In preserved specimens the stripes retain their rich brown color
even after imbedding in paraffin.
Ocelli, — The four ocelli are of moderate size, and, as in many re-
lated species, usually occupy the corners of a square (pi. xiv, fig. 6).
They are situated deep in the tissues of the head. There is no trace
of pigment between the two ocelli of the same side.
Proboscis. — Presents few deviations from the normal type, and, as
usual, is provided with ten nerves. Basis is of very small size, rather
slender, and but slightly enlarged posteriorly. Its average length is
only about .05-. 07 mm. and its width .02-. 025 mm. The central stylet
is rather slender (pi. xxi, figs. 13, 14) ; each of the two lateral pouches
contains two or three accessory stylets. Proboscis is attached a little
in front of brain ; proboscis sheath does not extend quite to posterior
end of body.
Body Walls. — The pigment which gives the brown color to the
longitudinal dorsal bands (pi. xxn, fig. A^pig) is conspicuous in every
transverse section of the body. It is not situated in the integument,
but occupies the inner portion of the longitudinal muscular layer.
Throughout the whole body, except in the head and most anterior
esophageal region, the anastomosing pigment cells largely obscure the
muscular fibers in the region where they are situated. This causes the
longitudinal muscular layer to appear interrupted by an elongated dark
mass on each side of the proboscis sheath. Just back of the brain,
where the muscular layer is thickest, the pigment masses lie on the
internal border of this layer, but farther back they occupy its whole
thickness (pi. xxn, fig. 4).
Submuscular glands appear only in anterior esophageal region, and
are but little developed.
Alimentary Canal. — Abroad intestinal caecum extends forward
through about half the length of esophageal region. It lies beneath and
beside the esophagus, which opens through its dorsal wall posteriorly.
Nephridia. — Situated in the middle portion of the esophageal re-
gion. A single pair of efferent ducts passes immediately above the
lateral nerves to open on the surface of the body just beneath the lat-
eral margins. The efferent ducts are situated only a few sections pos-
terior to the anterior end of the intestinal caecum.
166 COE
Nerves and Sense Organs. — Brain is of large size, with remark-
ably large ventral commissure. Cerebral sense organs remarkably
voluminous, situated immediately in front of brain, and extending
somewhat beside and beneath the ventral ganglion. Canal to exterior
is of large size, and extends well forward toward tip of head, to open
below lateral margin in a shallow oblique furrow.
Reproductive Organs. — Sexual products are mature in August.
The most anterior reproductive pouches in the males are situated well
forward in the esophageal region, or, in other words, well in front of
the opening of the esophagus into the intestine. The anterior sper-
maries lie beneath the lateral nerves, but in the intestinal region, where
several spermaries appear in each transverse section of body, they are
found in smaller numbers above the nerves. The ovaries occupy sim-
ilar positions in the female. The eggs when mature are very large in
proportion to size of body, being fully one-third its diameter.
Habitat. — Common among bryozoa and tunicates ( done) on piles
of wharf in the harbor of San Diego, Calif.
12. TETRASTEMMA QUADRILINEATUM sp. nov.
Pl. XIV, fig. 5; Pi. XX, figs. 12, 13.
Body short, broad and stout ; somewhat flattened, but with rounded
edges. Head usually narrower than body; provided with the usual
pair of lateral oblique grooves, which appear as slight constrictions
opposite the posterior pair of ocelli. Intestinal region broad and some-
what flattened. Intestinal caeca not much branched ; reaching nearly
to brain region.
Color. — General color of body whitish, with four longitudinal
deep brown stripes. Two of these stripes lie near lateral margins of
body, while the other two are situated symmetrically on the dorsal sur-
face. The two dorsal stripes are much the wider, and are each about
equal in width to the white median stripe which lies between them.
They are narrower on the head, and terminate anteriorly a little in
front of the anterior pair of ocelli. Posteriorly they extend to the end
of the body where they likewise become much narrower. These
broad dorsal stripes are smooth in outline, but the lateral stripes, which
are very much narrower, usually present a much broken and ragged
appearance. In some individuals the lateral stripes are fully three-
fourths as wide as the dorsal, but ordinarily they are less than half as
wide. The two lateral stripes are strictly marginal, and ordinarily do
not show from dorsal surface unless the animal is somewhat com-
pressed. They each terminate anteriorly in the vicinity of the lateral
NEMERTEANS 167
grooves on the head, or a little behind them. The white space sepa-
rating the lateral from dorsal stripes is nearly equal to the width of one
of the dorsal stripes. While the color of the stripes is always a deep
rich brown (pi. xiv, fig. 5), the whitish ground color is subject to con-
siderable variation in different individuals and in different states of
development of the sexual products.
The head is usually pure white except for the brown stripes ; esopha-
geal region commonly pure opaque white, although several mature
male specimens were pale yellowish or flesh-colored in this region.
Intestinal region commonly has a tinge of yellow or of pale salmon,
due to the color of the intestinal lobes which show through the other
tissues. When filled with mature ova, the intestinal region often has
a yellowish or very pale greenish tinge, while the males may be pale
flesh colored.
The ventral is of practically the same color as the dorsal surface,
but the effect of intestinal lobes and sexual products is here more
marked, so that in the intestinal region there is often a tinge of salmon
(from the intestinal lobes), of flesh color, or of greenish yellow (from
the sexual products). The color is also affected by the dorsal brown
stripes which show through the other tissues to some extent.
Proboscis whitish ; brain pale yellow in color.
The four dark brown stripes are quite as conspicuous after preser-
vation, and even after imbedding in paraffin, as they are in life.
Ocelli. — The four medium-sized ocelli are arranged nearly in the
form of a square in ordinary states of contraction of the head. Their
relative position to the brown dorsal stripes is very variable, be-
cause they lie so deep in the tissues of the head as to be unaffected
by the contraction of the superficial tissues in which the pigment
of the stripes resides. Commonly, however, they lie in or just lateral
to the brown stripes — the anterior pair lying near the anterior ends
of the stripes, and the posterior pair about on a level with the lateral
oblique grooves, as these appear on the margins of the head (pi. xiv,
%. 5)-
Size. — A small species, averaging only about 8-12 mm. long, and
less than I mm. in width, when sexually mature.
Proboscis, — Proboscis sheath extends to posterior end of body.
Basis of central stylet bell-shaped, short, much enlarged and rather ab-
ruptly truncated posteriorly (pi. xx, figs. 12, 13). Central stylet about
three-fourths as long as basis, rather small, but of typical proportions.
Each of the two lateral pouches usually contains two accessory stylets,
in size and shape like the central stylet. Basis measures about .o6-.o7
i 68 COE
mm. in length, and .04-. 05 mm. in diameter near base ; central stylet
.O45-.055 mm. long.
Retractor muscle of proboscis attached to the dorsal wall of the
sheath at about two-thirds the distance from head to posterior extremity
of body. Proboscis of large size, provided with ten large and con-
spicuous nerves.
Body Walls. — The pigment of the four dark longitudinal lines
appears perfectly black in mounted sections, and is always very con-
spicuous. It is situated in the basal portions of the integument.
Cephalic glands are well developed, and fill up a considerable 'por-
tion of the tissues of the head in front of the brain.
Alimentary Canal. — A pair of broad diverticula of the intestinal
csecum extends forward above, and a little in front of the dorsal brain
lobes. Shortly behind the brain the two diverticula join the broad
unpaired cascum which lies beneath the esophagus. This latter por-
tion of the alimentary canal swells out into a large chamber immedi-
ately behind the brain, while its slender posterior portion (pylorus)
does not empty into the dorsal wall of the intestine until well behind
the most anterior sexual glands. The broad intestinal caecum sends
off a few pairs of large diverticula. The most posterior of these ex-
tend between the anterior sexual glands, while the most anterior pair
terminates above the dorsal ganglia, as mentioned above.
Nephridia. — The nephridial system is remarkable for its short
extent and for the fact that the efferent ducts open on the head immedi-
ately beside the dorsal ganglia. The nephridial tubules lie above the
anterior portions of the lateral nerves, but do not extend more than a
very short distance behind the brain. A few branches also ramify
beside and a little in front of the dorsal ganglia. The efferent ducts
are rather large, and open -on the sides of the head about on a level
with the middle portions of the dorsal ganglia.
Cerebral Sense Organs. — These are of moderately large propor-
tions. They lie a little in front of the brain and somewhat nearer the
ventral surface. The ducts communicating with the exterior pass an-
teriorly to open on the ventro-lateral aspects of the head as usual.
The lateral nerves unite above the posterior end of the alimentary
canal.
Reproductive Organs. — The sexual products were found to be
nearly mature in August. As in most species of the genus, the sexual
glands are very voluminous, and occupy the greater portion of the body
in the intestinal region. As stated above, the most anterior sexual
pouches lie some distance in front of the opening of the esophagus into
NEMERTEANS
169
the intestine. The distance from the tip of the snout to the anterior
sexual pouches is therefore very short.
Habitat. — Among ascidians and other growths on piles of wharf
in San Pedro Harbor, Calif. ; fairly common. Sexually mature in
August. Ova large, opaque ; pale greenish in color.
This new species resembles both T. vittatum (Hubrecht) Burger l
and T. quadristriatum Langerhans 2 in general appearance, and in
having four longitudinal brown lines on dorsal surface. In the former
species, however, the four lines usually become confluent behind the
head in two quadrangular patches which send two fine lines between
the posterior pair of ocelli ; in the latter species the two median lines
extend to the tip of the head before uniting, while the lateral are inter-
rupted between the ocelli and do not unite on tip of head. In T.
qicadrilineatum, as described above, the median lines reach nearly to
tip of snout, but do not join each other, while the lateral lines usually
end behind the ocelli.
13. TETRASTEMMA ((ERSTEDIA) DORSALE (Abildgaard)
Mclntosh
Planaria dorsalis ABILDGAARD, Zool. Danic., iv, p. 25, 1806.
Tetrastemma dorsalis MclNTOSH, British Annelids, Pt. I, Nemerteans, Ray
Soc., p. 172, 1873.
Oerstedia dorsalis Burger, Fauna und Flora des Golfes von Neapel, Monogr.
22, p. 592, 1895.
This small, widely distributed species was collected by Mr. J. F.
Abbott in about 20 fms. in Monterey Bay, Calif.
The species may be recognized by its firm, slender, cylindrical
body, usually only 8-15 mm. in length, somewhat narrower toward
both extremities ; flesh color or pale yellowish, mottled on dorsal sur-
face with brownish blotches and dots of .various shades and with con-
siderable variation in distribution, often being mainly collected into a
series of transverse bands with a few scattered blotches between.
1 Hubrecht, A. A. W., Genera of European Nemerteans critically revised.
Notes from Leyden Museum, p. 229, 1879.
Hubrecht placed this species in the genus (Erstedia, but Burger refers it to
Tetrastemma. The specific name -vittatum is preoccupied in Tetrastemma, how-
ever, by Verrill, who described and figured a widely different species under this
name in 1874 (American Journal of Science, vn, p. 45). If Hubrecht's species
actually belongs to Tetrastemma, it is obvious that it must receive a new specific
name.
3 Langerhans, P., Die Wurmfauna von Madeira, Zeits. f. wiss. Zool., xxxiv,
p. 136-140, iSSo.
170
COE
The head is continuous with body and provided with four ocelli.
Proboscis armature as in other species of Tetrastemma.
Habitat. — On piles of wharves, on rocks, among algas, bryozoa,
ascidians and other growths. Widely distributed in Northern Hemi-
sphere, occurring on the northern coasts of Europe, in the Mediter-
ranean and on both the east and west coasts of North America.
14. TETRASTEMMA (CERSTEDIA) RETICULATUM
sp. nov.
pi. xrv, figs. 7, 8 ; pi. xx, fig. 7-9.
A minute species measuring but 8-15 mm. in length when sexually
mature, and less than 0.5 mm. in diameter. Body short, thick,
rounded; much resembling Oerstedia in form and movements, as
well as in firmness of body and in general appearance. The peculi-
arities of the species are not sufficiently pronounced,
however, to warrant its separation from the genus
Tetrastemma. Head usually somewhat narrower
than body, from which it is usually distinctly
marked off by a pair of oblique lateral grooves.
These lie opposite the posterior pair of ocelli (pi.
xiv, fig. 8), as in related species.
Color. — General color of body white, with rec-
tangular and longitudinal brown markings which
obscure most of the white color of dorsal surface.
Two varieties, presenting widely different arrange-
ment of the dorsal markings, were met with, and
these in extreme cases would sviggest two different
species. In both varieties the head, which is white
or colorless, is provided with a very similar trans-
FIG. 21. T. retic- verse deep brown marking between the anterior and
ulatum. Outline of posterior pairs of ocelli. This marking consists of
anterior portion of a transverse band of color extending laterally some-
body, showing post- wha(. b d the ocell^ and living a Bunded and
tion of ocelli and , ,,.,,, . /. . r- \ •*>.
shape and arrange- deeP!y bilobed anterior margin (text fig. 21). It
ment of markings, is often shaped like a dumb-bell, except that the
X4°- contour is straight, and not indented, posteriorly.
In the most common variety (pi. xiv, fig. 7) there are about sixteen
pairs of rectangular brown markings situated on the dorsal surface and
extending from the head to the posterior end of the body. These oc-
cupy the greater portion of the dorsal surface, leaving but little space
NEMERTEANS 17 1
between' and beside the rectangular marks for the white ground color.
They are often decidedly irregular in shape, but in ordinary states of
contraction of body average about one and a half times as long as
broad, and are separated medially by a space about half as great as
their transverse diameters. In addition to the sixteen pairs of rectangu-
lar marks, a pair of slender, irregular, brown lines are situated near
the lateral margins along the whole length of the body behind the head.
Anteriorly these slender brown lines join the more anterior rectangular
markings, but at about the third pair of rectangular markings become
separated, to join again at about the fourteenth pair. When the
animal is slightly compressed, a narrow white margin appears out-
side these lateral brown lines. The fusion between the first pair of
rectangular markings and the lateral lines is usually complete, while the
second pair is often joined by only narrow bridges of the brown color
(text fig. 21 ). Sometimes other pairs of markings are connected with
the lines by similar bridges of color, and toward the end of body this
is usually the case. At about the fourteenth pair of markings there is
often a complete fusion of brown color, both of the lateral lines and
of the rectangular markings of the same pair, and very commonly
only a narrow, transverse band of white serves to indicate the division
between the fourteenth and fifteenth, and between the fifteenth and
and sixteenth, or last, markings. Often even this narrow transverse
white band does not extend more than half way to the margins of the
body. This fusion of all the brown markings is commonly limited
to the three or four posterior pairs of markings, but in some individuals
involves the greater portion of the dorsal surface. When carried to
the extreme condition all the markings on the body back of the head
are united, and this fusion gives rise to the second and less common
variety.
In this second variety (pi. xiv, fig. 8) the whole dorsal surface back
of the head is of a deep brown color interrupted by about fifteen irreg-
ular, transverse, whitish bands reaching laterally toward margins of
body. These fifteen whitish bands indicate the spaces between the
sixteen pairs of rectangular markings described for the first variety.
The lateral lines have become fused and lost in the rectangular mark-
ings, and both markings of the same pair have completely lost their
individuality. Occasionally, however, a paler median line is indicated.
Laterally, too, the adjacent pairs of markings have fused completely.
Often the transverse whitish bands do not extend more than half way
to the lateral margins, as was mentioned in regard to the caudal region
of the first variety. They often become indistinct, and gradually shade
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into the brown color laterally. Sometimes, moreover, the fusion is so
complete that only a portion of the full number of transverse whitish
bands can be found.
The brown color of body varies from a rather light brown to deep,
chocolate brown or sometimes deep reddish brown. The whitish in-
termediate spaces are caused by absence of brown color, and are not
due to any special pigment.
The ventral surface is commonly dull grayish with a tinge of yellow-
ish in median line, but there are often numerous fine granules of
brownish pigment scattered over the surface. The colors of the dorsal
surface show through to some extent, and in the second variety the
brown color often encroaches on the ventral surface and shades off
gradually toward the median line. When full of ripe ova, the ventral
surface of the intestinal region shows a broad lateral band of dull
greenish tint due to the olive green color of the ova.
Ocelli. — The four ocelli are of medium size and, as usual, are
arranged nearly in the form of a square (text fig. 21), although in ordi-
nary states of contraction of head there is rather more space between
the two ocelli of the same side than between those of either the ante-
rior or posterior pair (pi. xiv, figs. 7, 8). Moreover, the ocelli of
the posterior pair often seem to be slightly farther apart than those of
the anterior pair.
Proboscis. — Proboscis sheath reaches end of body as in related
species. Proboscis pale flesh color, of large size as compared to body
of worm. Basis of central stylet about two and one-half times as long
as broad, somewhat elliptical, of nearly equal diameter throughout, and
only a little wider posteriorly than near the attachment of stylet (pi.
xx, figs. 7-9). Posterior half of basis appears granular and dark, but
becomes gradually lighter anteriorly until the anterior third is as trans-
lucent as in other species. Central stylet slender, about two-thirds as
long as the basis. Each of the two pouches usually contains two acces-
sory stylets. Measurements average about .05 mm. for the length of
the central and largest accessory stylets, while the basis is about .07
mm. long and .025 mm. in average diameter. The proboscis is usually
provided with nine nerves, although in a single specimen there was an
indication of a tenth nerve.
Internal Organization. — The dark pigment of the dorsal surface
of the body is situated among the epithelial cells of the integument,
and extends upward among the cells well toward the surface.
Voluminous cephalic glands situated both above and beneath the
rhynchodaeum reach back nearly to the brain.
NEMERTEANS 1 73
Alimentary Canal. — A pair of rather large diverticula of the in-
testinal caecum reach forward to the brain, and pass to the dorsal side
of the dorsal ganglia. They join the broad unpaired caecum shortly
behind the mouth. The caecum extends backward beneath the esopha-
gus, sending off a few broad diverticula. The narrow posterior por-
tion of the esophagus does not join the intestine until after the appear-
ance of several pairs of sexual glands. In this, and a number of
other features, the species very closely resembles T. qtiadrilineatum.
Nephridia. — The nephridial system is limited to the region near
the brain. Its tubules extend beside, as well as a very short distance
behind, the brain. The single pair of efferent ducts open laterally
just opposite, or a trifle behind, the posterior ends of the dorsal
ganglia.
Cerebral Sense Organs are remarkably large. They are situated
beside and beneath the ventral ganglia, and project somewhat in front
of them. Large ducts pass antero-ventrally to the surface of the head,
as usual.
Nervous System. — The brain is unusually voluminous ; otherwise
the nervous system agrees with that of related species.
Reproductive Organs. — Sexual products are mature in September.
The ova are very large (about .18 mm. in diameter), pale olive green
in color, arranged in a single row along each side of intestinal region.
The spermatic pouches are correspondingly voluminous in the males.
As stated above, the anterior sexual glands extend well in front of the
posterior end of the esophagus (pylorus), so that but a comparatively
short distance intervenes between them and the brain.
Habitat. — On piles of wharf in San Pedro Harbor, Calif., common.
15. TyENIOSOMA PUNNETTI1 sp. nov.
pi. xvi, figs. 1-3 ; pi. xvin, fig. 6.
Body large, soft, flabby, extremely contractile, rounded in esopha-
geal region, and much flattened posteriorly when extended ; much like
T. princeps* in shape, and, like that species, becoming very short and
subcylindrical when strongly contracted.
Head and anterior portions of body extremely contractile ; smooth
and somewhat flattened when moderately extended, but abruptly trun-
cated, cylindrical, and thrown into deep circular wrinkles when con-
^he species is named in honor of Mr. R. C. Punnett, of St. Andrews, Scot-
land, well known for his recent work on this group of worms.
2 Coe, Proc. Wash. Acad. Sci., in, pi. n figs. 3, 4, p. 62, 1901 ; also preceding
article, paged identically.
174
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tracted. Snout small, rounded or emarginate in front, flattened con-
siderably (pi. xvi, figs. 1,2), but may be almost completely withdrawn
into the swollen succeeding portions of head, as shown in fig. 3.
Oblique cephalic furrows lie on the ventro-lateral margins. As seen
from in front when strongly contracted, the snout presents an elliptical
outline, separated by a deep groove from the tissues of the succeeding
regions, which have been moved forward to surround the retracted head
(pi. xvi, fig. 3). This groove passes through the cephalic furrows,
and is continuous with them. The anterior portion of the esophageal
region also is deeply wrinkled with circular grooves. Proboscis pore
subterminal ; proboscis rather small, whitish in color. Mouth ex-
tremely variable in size, according to state of contraction of anterior
portions of body. When head is well extended the mouth is situated
well back from snout and represents a large, much elongated opening
(pi. xvi, fig. 2), but when head is strongly contracted the mouth as-
sumes the proportions of a small, rounded pore (pi. xvi, fig. 3) on the
ventral surface a little posterior to the retracted snout.
Esophageal region is often greatly swollen just behind the widely
opened mouth when body is well extended, very much as in Cere-
bratulus. There are many indications that in this species of Tcznio-
soma the esophagus is often filled with water, which is ejected at inter-
vals, and thus serves to some extent as a respiratory organ, as has often
been considered the case in Cerebratulus. The greater portion of
esophageal region is subcylindrical, becoming more flattened pos-
teriorly.
Intestinal region often very flat and ribbon-like (pi. xvi, fig. i),
usually much wrinkled and showing a tendency to roll up into an ir-
regular spiral, as has been noted in other species. This region may
contract to but a small fraction of its usual length, becoming at the
same time nearly as thick as broad. Posterior extremity not very
slender.
Color. — General color of body usually deep brownish red, or dark
red with only a tinge of brown ; sometimes of a deep mahogany color.
The body color often appears as if covered with a delicate whitish
bloom. Anterior portion of head is much deeper brown or almost
black, sharply marked off from a narrow white border which occupies
the terminal and lateral margins of snout. The dark brown or blackish
color is limited to the dorsal surface of snout, and usually consists of a
rather narrow transverse marking which shades off gradually into the
general body color posteriorly. It is bordered laterally by the white
margin mentioned.
NEMERTEANS 175
Ventral side of snout pure white, continuous with the white of the
margins of dorsal surface, but this color does not commonly extend
posteriorly to the snout. Occasionally, however, the white area ex-
tends back on the ventral side of the head proper, and may include
the mouth region also. Back of snout the ventral color usually be-
comes gradually reddish, so that a pale red area generally surrounds
the mouth (pi. xvi, fig. 2).
Ventral surface of esophageal region of a brownish red color similar
to that of dorsal surface, but of a much paler and more grayish shade.
A much paler grayish median band is often found throughout the
whole ventral surface. In intestinal region there is a similar pale
brownish red color much influenced by the grayish color of the median
intestinal canal and its lateral diverticula. These latter are conspic-
uous when the body is well extended, and add largely to the grayish
effect of the ventral side of the whole intestinal region. Sometimes
ventral surface is of the same brownish red color on the lateral mar-
gins as occurs on the upper side of body, but toward the middle line
becomes much grayer and paler, while a median line of pure gray
extends through the intestinal region.
The tissues of body in intestinal region show a general salmon or
orange color when cut.
Size, — A large species, individuals often measuring 40-60 cm. in
length and S-io mm. in width when extended. One of the preserved
specimens still measures 20 cm. in length, and 5-7 mm. in width in
intestinal region, and 12 mm. just behind mouth. When strongly con-
tracted the body of a large individual may become but 6—8 cm. long,
but is several times as thick as when extended.
Ocelli. — On each antero-lateral border of head is an irregular
cluster of numerous minute ocelli. When cleared in cedar oil these
appear distinctly as an irregular longitudinal row of forty to sixty or
more small pigment spots just ventral to the dark head shield (pi.
xvui, fig. 6).
Proboscis. — Proboscis sheath possesses an unusual length, extend-
ing backward throughout the greater portion of the intestinal region.
Proboscis of large size, with two well-developed muscular layers —
internal longitudinal and external circular — as usual.
Body Walls. — The three muscular layers are massively developed
throughout the body.
Cephalic glands are enormously developed, as in most related
species, and extend backward into the esophageal region. In front of
the brain they encroach largely upon the muscular and connective
1^6 COE
tissues surrounding the rhynchodseum and blood lacunae, but are sepa-
rated from the more superficial glands of the cutis by the rather thick
layer of fibrous tissue which constitutes its inner portion. This line
of separation is more marked on the dorsal than on the ventral side of
the head. In the brain region the cephalic glands occupy a large por-
tion of the longitudinal muscular layer on all sides of the body. In
the region of the mouth they become more scattered, and lie mainly on
the right and left sides, although they extend inward to the inner por-
tion of the outer longitudinal muscular layer. Posterior to the mouth
they are limited to the middle portions of the outer longitudinal mus-
cular layer ventral to the lateral nerves, and occur only sparingly.
They cease entirely a short distance behind the mouth.
The cutis is thick and, as in related species, consists of two con-
spicuous layers — an outer, glandular layer with closely packed, deeply
staining gland cells, and an inner, fibrous layer consisting largely of
interlaced connective tissue fibers. The cutis is several times as thick
as the integument, and between the two lies a rather conspicuous
layer of muscular and connective tissue fibers forming a basement layer
for the integument.
Alimentary Canal. — As stated above, the mouth is remarkably
distensible, appearing when contracted as a small round pore, but
when distended is a large and long slit with thickened lips. The
posterior end of the esophagus becomes much widened before it joins
the intestine, and is peculiar in that it does not pass smoothly into
the latter. The anterior end of the intestine proper is narrow, and opens
into the widened esophagus immediately beneath the proboscis sheath,
while the esophagus continues backward as a broad blind sac for
some little distance, surrounding the lateral and ventral walls of the
intestine. A section through this region, therefore, shows a rather
small intestinal canal, surrounded, except dorsally, by the thick
glandular walls of the broad, caecal portion of the esophagus.
Blood and Nephridial Systems. — Cephalic blood lacunas, lateral
vessels and esophageal lacuna? as usual. Proboscis sheath vessel con-
tinues within the rhynchocoel nearly the whole length of the esopha-
geal region. Nephridia remarkably extensive, originating a short dis-
tance posterior to the mouth and extending nearly the whole length
of the esophageal region. The efferent ducts are numerous, but of
small size. They are scattered at irregular intervals along the whole
length of the esophageal region, and, as usual, pass to the exterior im-
mediately above the lateral nerves. In the single specimen sectioned
there were eight to twelve of these ducts on each side.
NEMERTEANS 177
Nervous System and Sense Organs. — Brain and lateral cords as
in related species. Cephalic and buccal nerves large.
Cerebral sense organs highly specialized, and of rather large size.
They are much elongated, and their anterior ends are situated laterally
in the angle between the dorsal and ventral ganglia. After extending
back nearly to the posterior end of the dorsal ganglion, each sense
organ enlarges until it is nearly equal in diameter to the dorsal gang-
lion, and eventually fuses completely with the posterior surface of the
latter. At the narrow, anterior end of each sense organ a ciliated
canal leads outward to open into the rather conspicuous oblique
cephalic furrows.
Reproductive Organs. — Sexual products were found to be mature
in the specimens dredged off San Pedro in August, but those examined
at Monterey in September had neither ripe eggs nor sperm, and had
evidently recently discharged their sexual products. The mature eggs
are large and opaque, and when brought into sea water surround them-
selves with a jelly-like coating.
Habitat. — Dredged among red algae, of color closely approxi-
mating that of the worms, in 50 fms., between San Pedro and Santa
Catalina Island, Calif. This is a common species in that locality, and
the individuals are remarkably hardy, living for a day or more among
damp seaweeds. A number of large living specimens were obtained
from Chinese fishermen, who caught them on their hooks in rather
deep water (perhaps 10-20 fms.) in Monterey Bay, Calif. Just how
the worms come to be caught on these hooks, which are set for ' rock-
cod,' is difficult to imagine, unless they crawl about among the sea-
weeds on the bottom, and, as the hooks drag through, are caught in
their bodies. It is possible that the worms actually find the bait and
cling to it. The fact that the body is often entirely without injury
points to this conclusion. The fishermen offered no conclusive evi-
dence in the matter. The species must be abundant, or it would be
more rarely caught on hooks.
Individuals live for a long time in captivity, and do not break up
when roughly handled. They may be easily preserved without rup-
ture, or without excessive contraction if killed slowly. Natural colors
are well retained in formalin, but fade in alcohol.
16. ZYGEUPOLIA LITTORALIS C. B. Thompson
Zool. Anz., xxin, p. 151, 1900.
Proc. Acad. Nat. Sci. Philadelphia, 1901, p. 657-739, pi. XL-XLIV, 1902.
This very interesting species which has been so carefully studied
and fully described by Miss Thompson (1902) occurs rather commonly
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on the flats laid bare by the lowest tides in San Pedro Harbor. It has
previously been recorded only from the vicinity of Woods Hole, Mass.
The worms are found in sandy, and not in muddy locations.
The species may be recognized by the following characters : Body
rather slender, usually 5-8 cm. in length, rather translucent. Head
slender, when extended tapering to a fine point, without cephalic fur-
rows ; pure white in color. Esophageal region rounded ; white, very
FIG. 22. Z. littoralis. Diagram of anterior por-
tion of body, showing relation of cerebral sense organ
(cso) to ciliated pit (c/5) on side of head; r/i, rhyncho-
dseum ; br, brain ; /«, lateral nerve.
FIG. 23. Z. littoralis. Diagram of caudal cirrus
and posterior end of body ; In, lateral nerve, extend-
ing into caudal cirrus (In'); in, intestine; a, anus.
Both figures after C. B. Thompson.
FIG. 23.
pale yellowish or flesh color. Intestinal region flattened in life, but
rounded after preservation ; color varies from rose to pale yellow, light
brown or chocolate brown, being largely dependent on the amount and
character of the contents of alimentary canal. Posterior extremity pro-
vided with a conspicuous, although slender, caudal cirrus, white in
color (text fig. 23), with large central blood space.
In internal organization, the presence of an internal circular muscle
at the beginning of intestinal region, the division of the anterior por-
tion of alimentary canal into esophagus proper and stomach, the open-
ing of the ciliated canals leading from cerebral sense organs into shallow
lateral pits (text fig. 22, cp} in place of cephalic furrows, the absence
NEMERTEANS I 79
of any retractor muscle of proboscis, and the absence of the inner lon-
gitudinal muscle of proboscis, are among the more important peculiari-
ties of the species.
In the paper referred to will be found detailed histological descrip-
tions and figures of all the principal organs of the body.
17. LINEUS RUBESCENS sp. nov.
pi. xiv, fig. i ; pi. xv, figs. 3, 4; pi. xxn, fig. i.
Body very slender, rounded anteriorly, flattened in intestinal region ;
posterior extremity slender ; head long, rather broad ; cephalic furrows
correspondingly long, reaching posteriorly as far as the anterior end of
the mouth, which is situated well back from tip of snout.
Size. — A small species, the specimens obtained measuring only
10-15 mm. long, and less than I mm. in diameter.
Color. — Anterior portions of body, except tip of head, beautiful
pink, or rosy flesh color, occasionally bright pinkish red, sometimes
with tinge of blue. Tip of head, both above and below, whitish, al-
most colorless, sharply marked off from pink color behind. Intestinal
region deep flesh color or pale, purplish brown, or occasionally buff,
anteriorly, shading off to very pale pinkish towards posterior end of
body ; very pale posteriorly. Ventral surface paler, but of similar
color, pinkish in front, pinkish buff or pale flesh color posteriorly ;
often with still paler median line. Intestinal lobes flesh color or buff ;
occasionally brownish or purplish. Head can be so much contracted
that white tip will disappear. Brain appears pale flesh color. Space
between ocelli is grayish. A most striking peculiarity of the color in
the esophageal region is that it becomes purplish or, sometimes,
bright blue in formalin. I know of no other species of Nemertean
where the color changes from pinkish to bluish on preservation.
The bluish color is not permanent, however, and after remaining a few
weeks in formalin will have practically disappeared. But when such
specimens are cleared in cedar oil, a delicate bluish or bluish green color
reappears in the esophageal region, while the intestinal region remains
colorless, except the intestinal lobes, which are slightly brownish. The
nature and situation of the pigment are noted below.
Ocelli. — Situated in whitish area on tip of head is a row of two to
four (or, rarely, as many as six or eight) ocelli on each side (pi. xiv,
fig. i ; pi. xv, figs. 3, 4). These are irregular in shape, very dark
reddish or almost black in color, and closely placed in a single row.
The anterior ocellus on each side is usually the largest, but the ocelli
180 COE
are often irregularly joined together, so that the individual ocelli cannot
be distinguished, appearing rather as a row of scattered pigment masses
on each side. In microscopic sections the eyes appear deep blue in
color.
In internal organization the species shows many deviations from
most other members of the genus although it is evidently closely related
to L.jlavescens. It presents a number of interesting anatomical pecu-
liarities, which are noted below.
Body Watts. — At the point where the posterior esophageal cavity
enters intestine, the circular muscular layer abruptly becomes less than
half as thick as it is farther forward, and allows the outline of the body
to become much wider and more flattened. In the esophageal region a
cross section is but little wider horizontally than vertically, but in the
intestinal region it is nearly twice as wide. This is due to the reduc-
tion of the circular muscular layer.
Extensive cephalic glands (pi. xxn, fig. i, cgV) reach inward
almost to the blood lacunae in the anterior portions of the head, but do
not extend behind the brain except on the ventral side, where they reach
as far as the posterior ends of the cerebral sense organs. Beneath the
rhynchodaeum is a conspicuous canal (a) which passes forward and
joins the rhynchodasum immediately where this opens on the ventral
side of the tip of the snout. This canal is apparently the common duct
of the cephalic glands which lie beneath the brain, for it originates pos-
teriorly in a large cluster of these glands. Yet in no other species of
Nemertean has such a well-developed duct been observed. Unfortu-
nately the character of its lining is obscured by the secretions which it
carries, so that its precise nature is not absolutely certain.
The glands of the cutis (pi. xxn, fig. i, cugl} are extremely abun-
dant in the head. They extend inward to the circular muscular layer
throughout the whole intestinal region. Back of the mouth they sink
gradually through the cutis and deeper into the external longitudinal
muscular layer. In the region of the nephridiopores they have passed
completely through this layer in a narrow area on the dorsal and on
the ventral side, and have come in contact with the outer border of the
circular muscular layer. Farther back the region in which the glands
reach inward as far as the circular muscle becomes increased, until at
the beginning of the intestinal region it extends along the whole cir-
cumference except in the vicinity of the lateral nerves. Throughout
the length of the intestinal region these glands continue to border the
whole surface of the circular muscles, except in the immediate position
of the lateral nerves. Perhaps in no other member of the family
NEMERTEANS l8l
Lineiclae have these glands been found to be so extensive as in the
present species.
Pigment. — The peculiar bluish color which appears after the
worms have been preserved, and have consequently lost their original
pink or reddish color, is due to a vast number of minute granules situ-
ated in the nervous plexus which lies immediately external to the
circular muscular layer. The granules are present only in that portion
of the plexus which lies dorsal to the lateral nerves.
Proboscis. — The proboscis sheath is remarkably long for the genus,
extending very nearly to the posterior extremity of the body. The
proboscis is of moderate proportions. In its musculature it exhibits an
interesting departure from the type characteristic of the genus. The
circular and outer longitudinal muscular layers are of the usual propor-
tions, while the internal longitudinal musculature is represented by two
bands placed symmetrically on opposite sides of the proboscis. These
longitudinal bands occupy about one-sixth to one-eighth of the circum-
ference of the internal epithelial layer, which elsewhere borders the
inner face of the circular musculature. In thickness the two muscular
bands, which represent the inner longitudinal musculature found in
most related species, often equal that of the outer longitudinal layer in
their middle portions, but are much thinner toward their borders.
Back toward the middle portions of the proboscis they become gradually
thinner, and more posteriorly eventually disappear, allowing the inter-
nal epithelium to border the circular muscles without interruption ex-
cept from the nervous layer. Where the two longitudinal bands are pre-
sent the proboscis nerves are well developed as a single pair of flattened
cords which lie immediately external to those bands, but after the bands
disappear, the nerves spread out as a plexus lying between the circular
muscles and basement layer of the internal epithelium, as in related
species. This peculiar arrangement of the musculature presents a con-
dition intermediate between those members of the family Lineida? in
which the three layers are well developed, and those (such as Lineus
flavescens, for example) in which the internal longitudinal layer is com-
pletely wanting. In the posterior portions of the proboscis only longi-
tudinal fibers occur, and these are bounded externally by a thin fibrous
layer, and internally by a thin epithelium lining the proboscis cavity.
Alimentary Canal. — Esophagus divided into two well-marked
portions, separated by a constriction, and distinguished by differences
in the epithelial lining similar to those which Miss Thompson has
recently described for Zygezipolia (I9O2).1 The mouth, situated as
1 Proc. Acad. Nat. Sci. Philadelphia, 1901, p. 709.
l82 COE
usual, opens into the typical, broad esophagus with greatly convoluted
walls lined with highly columnar ciliated epithelium and provided
with an abundance of glands. Exactly in the region of the nephridio-
pores, however, or at about one -fifth the distance to the intestinal
region, a marked change occurs. The esophageal lumen becomes nar-
row and situated immediately beneath the proboscis sheath, while be-
neath and beside it appears a second broad canal — posterior esophageal
chamber or stomach, as it may be called. This is likewise devoid of
lateral diverticula. A few sections back of the anterior end of this
posterior chamber, the narrow, terminal portion of the esophagus
proper, or anterior esophageal chamber, opens. The epithelial lining
the posterior chamber differs conspicuously from that of the anterior
chamber. In the latter the ciliated cells are situated superficially and
their nuclei are not far removed from the surface, while the glandular
cells lie mainly at a lower level and have their nuclei farther from the
surface. In the posterior esophageal chamber, on the other hand, a
comparatively small proportion of the cells are ciliated, and the nuclei
of all are far removed from the surface. The cells are all very slender,
and are thickly packed with small granules of secretions. The free
surfaces of the cells appear to be irregular, and are covered with the
secretion which partially fills the lumen of the esophagus, so that it is
often difficult to determine exactly where the cells terminate. In this
respect they differ greatly from those in the anterior chamber, where
the ciliated cells always show a sharp and distinct free border.
The posterior chamber is fully four times as long as the anterior,
and at its posterior extremity enters the broad cavity of the intestine
proper, which is provided with the usual lateral diverticula, and ex-
hibits the usual histological features. The transition from the pos-
terior esophageal chamber to the intestine is very abrupt in both its
anatomical and histological features. The circular muscular layer of
the body walls becomes much thinner at this point, and the body is
flatter and wider, as described above. The intestine at its anterior end
becomes correspondingly wide and flat, with broad lateral diverticula.
The lumen, on the other hand, becomes much narrower, and the epi-
thelial lining several times as thick as in the posterior esophageal cham-
ber. As in other species, the cells are large, without distinct outlines,
and packed with peculiar spherical masses of deep-staining secretions.
The nuclei are pushed deep into the bases of the cells. The general
appearance, therefore, of these two portions of the alimentary canal is
vastly different, although not so sharply marked as in other forms
where the esophagus exhibits less specialized portions.
NEMERTEANS 183
The posterior esophageal cavity — 'stomach/ or gastric portion of
esophagus, as it may be called — is by no means a structure peculiar
to the species at hand, for indications of a specialized posterior portion
have often been described for the esophagus of other species. Yet in
no other Heteronemertean has this chamber been found to be so highly
differentiated. The transition is usually far more gradual, especially
between the stomach and the intestine, as is the case in Zygcupolia.
The general nature of the cavity resembles the intestinal caecum of
the Hoplonemerteans far more than it does the so-called ' stomach '
(Magendarm) of these forms.
Stood and Nephridial Systems. — Cephalic blood lacuna?, lateral
vessels, and anastomosing blood spaces about esophagus as in related
species. Rhynchocoel vessel short, leaving the cavity of the proboscis
sheath in the immediate vicinity of the nephridiopores, or at about
one-fifth the distance from mouth to intestinal region.
Nephridial system remarkably short, but with rather large branches.
It commences anteriorly a short distance behind the mouth and is
limited to the anterior fifth of the esophageal region. The main longi-
tudinal canal on each side is situated as usual in the angle between the
proboscis sheath and the esophagus, and the branches ramify about the
esophageal blood spaces. After an extent of less than one-fifth the
length of the esophageal region, each of the two longitudinal canals
swells out into a comparatively large chamber from which the rather
large efferent duct passes above the lateral nerve to the nephridiopores,
situated as usual on the dorso-lateral surface of the body. The nephrid-
ial canals do not extend farther posteriorly than the position of the
nephridiopores, which are strictly paired.
Nervous System and Sense Organs. — These structures present no
noteworthy peculiarities. Terminal, or frontal, sense organs appar-
ently wanting. Cerebral sense organs well developed and closely
united with posterior surfaces of dorsal ganglia. The canal by which
each of the sense organs communicates with the posterior end of the
corresponding cephalic furrow is broad and short.
Reproductive Organs. — No sexual products are present in speci-
mens collected in August and September.
Habitat. — A few specimens of this beautifully colored species
were found on piles in San Pedro Harbor, Calif. A single specimen
was found by Mrs. Cockerell on a kelp 'hold-fast' outside the same
harbor. The species also occurs sparingly on the piles at Monterey,
Caiif.
184 COE
1 8. LINEUS FLAVESCENS sp. nov.
pi. xvn, figs. 3, 4.
Body of moderate proportions for the genus ; head long and rather
slender, usually a little narrower than body, often pointed, and often
slightly emarginate in front ; cephalic slits longer than in most related
species ; esophageal region rounded ; intestinal region only moderately
flattened ; posterior extremity rather slender, without caudal cirrus ;
proboscis sheath extends nearly to posterior end of body ; proboscis
rather slender.
Color. — General color of body yellowish, but of a great variety of
shades in different portions of the body, and in different individuals.
A number of specimens from crevices of rocks at San Pedro Harbor
were pale yellow, sometimes with a tinge of orange, in esophageal
region, and deep ochre throughout the remainder of the body except
the head, which was ochre with a decided tinge of orange. Margins
of head are paler. The pale yellow of esophageal region changes
abruptly to the ochre of the intestinal region ; posterior extremity and
whole ventral surface of body of a duller, paler color.
Esophageal region often shows two distinct regions of different color,
due to the two divisions of the esophagus described below. The
anterior pear-shaped region lying next the mouth is more deeply
colored, corresponding with the thicker esophageal walls, while the
remainder of the region is usually paler (pi. xvn, fig. 4).
Several specimens from among annelid tubes on San Pedro break-
water were deep ochre anteriorly, with a sharp, median, dorsal, longi-
tudinal white line which extended through the anterior half of the
esophageal region. The remaining portion of esophageal region was
greenish ochre, while the whole intestinal region was dull orange (due
largely to the color of the intestinal canal), covered superficially by the
pale yellowish of the body walls. Ventrally the color was dull whitish
on head and in anterior esophageal region, then greenish ochre back as
far as the intestinal region, which was of the same orange color as dor-
sal surface, but somewhat duller. Lateral and anterior margins of
head colorless ; cephalic slits long, sharply separating the ochre or
yellowish orange of dorsal surface from whitish of ventral surface of
head. Smaller specimens were commonly much paler in color. Some
individuals were of duller colors, with dark buff intestinal regions.
Numerous large individuals taken in from 50-100 fathoms between
San Pedro and Santa Catalina Island were of a rich golden brown
anteriorly, with a tinge of yellowish orange on the head. Intestinal
NEMERTEANS 185
region varied from ochre through deep buff to olive brown, sometimes
showing a median dorsal longitudinal stripe of darker, more brownish
color. These were filled with mature sexual products.
In all these varieties the brain region was indicated by its more rosy
color, and this was sometimes quite conspicuous. The lateral and
anterior margins of head were always very pale or colorless. The
esophageal region showed clearly the two divisions of the esophageal
walls, as seen through the somewhat translucent body walls. The
ventral surface of head is much paler than dorsal surface, while
throughout remainder of body the ventral differs from the dorsal sur-
face in color mainly in having a duller and more whitish tinge. A paler
median dorsal line in intestinal region, indicating the position of the
proboscis sheath, is often present.
Proboscis straw color or slightly yellowish.
Size. — Small, pale-colored individuals were 8-15 mm. long;
others 20-40 mm., while those from deeper water were often 80-120
mm. in length, with a diameter of 2-3 mm.
Ocelli. — On each side of tip of head is a transverse row of irregular
pigment masses (pi. xvn, figs. 3,4). These are very variable in shape,
in size, and in number, and vary in color from blood red to purple or black.
There are commonly three to seven irregular groups of pigment granules
on each side, and of these the more anterior are the larger. Those of the
two sides are separated by a fairly wide pale area above the proboscis
pore, and extend as a single row along each antero-lateral margin of
the head. Commonly the ocellus lying most anteriorly is much larger
than any of the others, while the three or four most posterior ones are
represented by minute dots only. The number of ocelli may not be
the same on the two sides, and perhaps in the majority of cases it is
impossible to determine the exact number because the pigment granules
are so much scattered. Often there is an irregular row of scattered
pigment masses, grouped irregularly, but not arranged into definite
ocelli. These appear to have arisen from a fragmentation of ocelli
which have previously existed.
Whether the ocelli should be blood red, wine color, purple or black,
seemed to be an individual peculiarity, and was not always correlated
with the color of the body or the environment under which the animal
lived. It is to be noted, however, that most of the individuals dredged
among the red seaweeds had wine-colored or purple ocelli.
As seen in sections, only the anterior pair of ocelli usually possess a
well-developed visual apparatus, with distinct, cup-shaped retina hav-
ing long, spindle-shaped sensory cells. They lie very deep in the
186 COE
tissues of the head on either side of the anterior end of the rhyncho-
daeum. The retina faces ventrally. The remaining ocelli are much
simpler in structure.
In internal organization this species shows a close resemblance to
L. rubescens, and hence a considerable departure from the typical
species of the genus in many details of structure. The close similarity
in many anatomical features between these two species and Zygeu-
polia, as described by Miss Thompson (I9O2),1 is very striking, and
serves to unite this genus more closely to the other members of the
Lineidae.
Proboscis. — Proboscis sheath does not extend to end of body.
Proboscis attached to dorsal wall of sheath at the posterior end of the
esophageal region, and only a few sections in front of the intestine, by
a powerful retractor muscle. In this same region the circular muscles
of proboscis sheath extend ventrally to surround the esophagus with
a rather thick layer of circular muscles. The fibers are connected to
some extent with the main circular muscular layer of the body walls
by a crossing of fibers immediately above the proboscis sheath. This
recalls the great thickness of the inner circular muscles found in this
same region in Carinoma. A similar condition has been described
for Micrura alaskensis (Coe, 1901), 2 and Miss Thompson (1902)* has
recently described for Zygeupolia littoralis and Micrura cceca a well-
defined inner circular layer in this region, and has shown its apparent
homology with the inner circular muscular layer of Carinella, Carz-
noma and other Paleonemerteans.
This localized inner muscular layer doubtless serves the double pur-
pose of holding the proboscis sheath, with its attached proboscis,
firmly in place among the other tissues, and of providing a strong
annular constriction between the esophagus and intestine. By the
action of this sphincter, food materials, having entered the intestine,
may be prevented from returning to the esophagus. Furthermore,
the esophagus may under some circumstances act as a respiratory
organ, as described by Wilson (1900)* for Cerebratithts lacteus, and
as I have observed in other species, and in this case the muscular con-
striction will prevent the water from passing back into the intestine.
The muscular walls of the proboscis consist of the outer longitudinal
and inner circular layers only. The inner longitudinal layer, which is
1Proc. Acad. Nat. Sci. Philadelphia, 1901, pp. 657-732.
2 Preceding article, p. 72.
8 Loc. ct't., p. 667.
* Quart. Journ. Micr. Sci., XLin, p. 109.
NEMERTEANS 187
represented in L. rubescens merely by two bands of longitudinal fibers
(as described on p. 181), is here wanting completely. The two pro-
boscis nerves are conspicuous in the anterior portions, but farther back
spread out into a plexus as usual. The basement layer of the proboscis
epithelium is therefore separated from the circular muscular layer only
by the nerves or nervous plexus, as described on p. 181 for L,. rubescens.
A precisely similar condition has recently been described for Zygeu-
polia by Miss Thompson (I9O2),1 and is known in other species of
the Lineidse.
Cephalic glands are well developed, but extend backward only to
the anterior portion of the brain region.
Cutis Glands. — In the esophageal region the cutis glands form a
thick and distinct layer in the midst of the outer longitudinal muscles.
Anteriorly they lie well toward the periphery of this muscular layer,
but sink gradually deeper until in the nephridial region they lie in con-
tact with the circular muscles dorsally and ventrally. This condition
is retained in the intestinal region, as was described for L. rztbescens.
Cephalic Ftirrows. — These are deep, and broaden out dorso-
ventrally on their internal faces near their posterior ends. The canal
leading to the cerebral sense organ on each side joins the posterior,
broadened end of the corresponding cephalic furrow through a broad
papilla.
Alimentary Canal. — Mouth large, elongated, situated opposite
posterior ends of cephalic slits. As stated above, the esophagus is
seen in the living worm to be divided into two distinct regions. These
consist of an anterior, pear-shaped cavity (pi. xvn, fig. 4) immedi-
ately behind the mouth and a posterior tube, or posterior esophageal
chamber, connecting with the intestine. The anterior cavity is dis-
tinguished by its much thicker, convoluted walls and deeper color.
At its posterior end this chamber is widened considerably, and is then
abruptly constricted as it opens into the posterior esophageal chamber,
or stomach. This latter portion has thinner walls and paler color, and
likewise exhibits a constriction just before it opens into the intestine.
There are very marked histological differences between these two por-
tions of the alimentary canal, but the conditions are here so very
similar to those described for L. rubescens that the detailed descriptions
given on p. 182 apply equally well for the species at hand. Perfectly
similar conditions have recently been described for Zygeupolia and
other forms.
The remarkable development of circular muscles about the posterior
1Loc. cit., p. 693.
i88 COE
end of the esophagus forms a strong sphincter between the posterior
esophageal chamber (stomach) and the intestine, as described above.
Blood and Nephridial Systems. — Cephalic and esophageal lacunae
as in related species. Proboscis sheath vessel passes outside rhyn-
chocoel in the nephridial region, or at about the point where the ante-
rior esophageal cavity opens into the stomach.
The main nephridial canals are of large size, but of limited extent.
They are situated beside the posterior fourth of the anterior esophageal
chamber, and the anterior third of the posterior esophageal chamber,
or stomach. In one small specimen sectioned there was but a single
pair of large efferent ducts, while in a larger individual there were two
pairs. In the former case the ducts were exactly paired, while in the
latter both the ducts on one side were in front of the more anterior of
those on the other.
Nervous System. — Brain and nervous system show no marked
deviations from the condition typical of the genus. The frontal sense
organ appears to be represented by a crescent-shaped groove above the
Bubterminal proboscis pore.
Habitat. — In crevices of rocks between tides, Dead Man's Island,
San Pedro Harbor; among annelid tubes, breakwater, San Pedro,
Calif. ; not very abundant ; one specimen from a floating kelp ' hold-
fast'off San Pedro Harbor (Mrs. Cockerell) ; common among red
algaa in 5ofms. between San Pedro and Santa Catalina Island.
The worms belonging to this species are remarkably hardy and will
live for more than a day in damp seaweed at a temperature of 70—80°
F., and may be kept for a long time in confinement in a small quan-
tity of sea water. Sexual products appeared to be fully mature in
August, but in some individuals had been previously discharged.
19. LINEUS PICTIFRONS sp. nov.
pi. xvu, figs. 5, 6.
Body of large size, remarkably soft and flabby, very changeable in
shape, but usually somewhat flattened throughout. Body often snarled
and tied in knots ; much fluted longitudinally, and constricted trans-
versely when contracted. Head narrower than body, elongated, often
emarginate in front, narrower at posterior end of cephalic slits than in
its middle portion. Cephalic slits unusually long, rosy in color pos-
teriorly, sometimes bordered by a narrow line of white. When con-
tracted, body is thick, and not flattened, except near anterior end.
Posterior extremity rather slender, rounded, not flattened.
Color. — General color of body, both above and below, deep brown,
NEMERTEANS 189
chestnut, or slaty, with a tinge of green in reflected light, or of plumbago
or bluish when seen in shadow. Sometimes a reddish tinge is given to
the brown, and there is usually a soft, velvety sheen. Head has a rosy
or chestnut tinge beneath the brown. Tip of tail very pale in color.
Throughout the whole body is usually found a series of transverse
and longitudinal yellowish markings on dorsal surface. The trans-
verse markings are the more conspicuous and consist of a series of
lemon yellow rings. The more anterior of these commonly encircle
the whole body, while farther back they appear on dorsal surface only.
They are all much widened in the median line, and in some specimens
appear to consist only of a series of diamond-shaped median markings.
More commonly, however, each marking continues laterally as a fine
line which completely encircles the body, although many are inter-
rupted on the ventral surface. In the posterior half of the intestinal
region these markings do not usually appear on the ventral surface.
In some specimens the markings are very much obscured and occa-
sionally are almost completely wanting. The first transverse marking
occurs at the posterior end of the cephalic furrows, and this is usually
much larger and more conspicuous than any of the others (pi. xvn,
fig. 5) although it does not extend to the ventral surface. On a worm
of 10-20 cm. in length there are usually sixty to one hundred or more
of these transverse rings.
The dorsal surface is corrugated with longitudinal flutings, except
when the animal is fully extended. These flutings are accentuated by
very fine, hair-like longitudinal lines of ochre or orange color which
extend throughout most of the dorsal surface. They are not only very
fine, but are much interrupted and irregular, usually wavy, and often
consist of rows of elongated dots of color. On the head the lines are
much interrupted and consist mainly of rows of very minute dots.
On the ventral surface they are entirely wanting throughout the body.
On the dorsal surface there are commonly about seven to fifteen or
more of the fine lines, of which the one in the median dorsal line is
more distinct than the others, and connects the transverse, diamond-
shaped, pale yellow markings described above. On each side of the
median line the lines are sometimes arranged rather symmetrically, but
are sometimes almost entirely lacking. This is often the case in the
posterior half of the body. In many specimens the lines are seen only
when very carefully examined, and in some others appear to be en-
tirely lacking. Sometimes they are wanting anteriorly to the third or
fourth transverse yellow marking. The transverse markings are of a
much paler yellow than are the longitudinal lines.
COE
Especially characteristic is the peculiar coloring on the tip of the
head, and this has suggested the specific name. On the dorsal surface
near tip of snout is a narrow terminal border of white, and situated
within this white border are two oval, orange-colored spots imbedded
in an area of lemon yellow (pi. xvn, figs. 5, 6). Sometimes a few
isolated dark brown dots are scattered in the yellow color. In some
specimens the orange-colored spots are very conspicuous, while in
others they are very small and indistinct. Sometimes the yellow color
surrounding them is wanting, but the white terminal border is always
present on the tip of the snout, both above and below.
After preservation in formalin the head becomes much contracted
and wrinkled, the cephalic slits become short, and the body assumes
a dull black or slaty color, without indications of the characteristic
yellow markings seen in life.
Mouth large, with pale lips, reaching forward to the posterior ends
of the cephalic slits.
Proboscis salmon or flesh-colored, very long and slender.
Size. — Usually about 15 cm. in length and 3—4 mm. in width,
although individuals were found as small as 15 mm. long, while a
single specimen measured nearly 50 cm.
Proboscis. — Provided with two muscular layers only, the internal
longitudinal muscles being wanting. There is a well-marked crossing
of fibers between the circular muscles and the outer fibrous layer, as in
many related species.
Body Walls. — Structure of body walls as in other representatives
of genus. The pigment which gives the dark color to the body is
situated as a very dense layer of minute pigment granules immediately
beneath the epithelium throughout the whole length of the body. It
is even present as a less distinct layer beneath the epithelium of the
cephalic furrows, and is lacking only in a narrow area immediately
surrounding the proboscis pore. This area corresponds to the white
zone seen in life at the tip of the snout. So dense is the pigment layer
that if ocelli were situated in it they would be very difficult to dis-
tinguish. They are probably not present, however, for they do not
appear in any of the sections.
A rather thin layer of connective tissue is found beneath the cutis
glands, which indicates an approach to the condition of a true cutis.
The cutis glands are massed in a distinct layer just beneath the pigment
layer, and do not extend among the fibers of the outer longitudinal mus-
cles even in the intestinal region. They are usually distinctly, though
not widely, separated from these muscles by the thin fibrous layer.
NEMERTEANS I 9!
Cephalic glands form a voluminous and well-marked mass of deeply
staining glands which extends back both dorsally and ventrally nearly
to the brain. Their secretions are discharged anteriorly at the tip of
the snout as in the Tagniosomicke.
Alimentary Canal. — The esophageal region is remarkably short in
comparison with the total length of the body There is a very short
esophagus lined with the characteristic glandular and ciliated cells.
At the posterior end of this is a decided constriction, or sphincter,
which opens into a widened posterior chamber without lateral pouches,
but having a lining of epithelium not widely different from that of
the intestine. This posterior cavity is even shorter than the esophagus
proper, and gradually passes into the true intestine with its paired
lateral pouches. The esophagus proper is sometimes only as long as
the transverse diameter of the body, and hence, much shorter than in
most related species. The cavity behind this evidently corresponds to
the ' stomach,' as described by Miss Thompson ( I9O2)1 for Zygeupolia,
and is doubtless homologous with the posterior esophageal cavity of
Lineus rzibescens and L. Jlavescens, as described above. In these
latter species, however, the epithelial lining of this cavity is markedly
different from that of either the esophagus or intestine, while in L.
pictifrons, as in Zygetipolia, it resembles the intestinal epithelium
very closely.
Although the change from esophagus to ' stomach ' is very abrupt
both anatomically and histologically, yet it is quite impossible to deter-
mine exactly where the stomach ends and the intestine proper begins.
The anterior intestinal pouches are but slightly indicated and pass
gradually into the wavy outlines of the stomach, and the histological
features show a similar gradation. The cells of the axial cavity of the
intestine retain the appearance of the stomach cells for some distance
posteriorly as in Zygeupolia.
An internal circular muscular layer, such as is described above for
L. rubescens and L. flavescens just anterior to the intestine, is not
found in the present species.
Blood and Nephridial Systems. — There is the usual arrangement
of cephalic blood lacunae, lateral lacunas, rhynchocrcl vessel, etc.
Branches of the lateral lacunas surround the esophagus, and are gath*
ered into a single pair of vessels in the region of the stomach. In this
region there are five large vessels in a transverse section of the body —
the rhynchocoel vessel, the pair of lateral lacuna? situated in the angle
between stomach and proboscis sheath, and the pair of ventral vessels
JProc. Acad. Nat. Sci. Philadelphia, 1901, p. 709.
COE
situated latero-ventrally beneath the stomach. At the beginning of the
intestinal region the lateral lacunas pass ventrally to join the ventral
vessels, which continue to posterior end of body. The rhynchocoel
vessel passes beneath the proboscis sheath at the beginning of the intes-
tinal region.
The nephridia are limited to about the middle half of the very short
region of the esophagus proper, and are consequently much less ex-
tensive than usual. The main longitudinal canal on each side lies on
the lateral wall of the lateral blood lacuna and sends branches among
the esophageal lacunae. A single pair of efferent ducts pass to the ex-
terior above the lateral nerves as usual. In one individual, one of the
efferent ducts was peculiar in that it branched just outside the circular
muscular layer. Only one of its branches passed to the exterior while
the other penetrated only about two-fifths the distance through the
outer longitudinal muscular layer.
Nervous System and Sense Organs. — The species is remarkable
in having many of the smaller nerves far more conspicuous than in
most other species. The cephalic nerves, for instance, are unusually
large and numerous. Esophageal nerves are also large, with a distinct
commissure near their point of origin from the ventral brain lobes.
These nerves join in a distinct plexus beneath the esophagus a short
distance behind the mouth. The dorso-median nerve is clearly marked
and of remarkably large size. The internal dorso-median nerve,
found in many species directly beneath the former and internal to the
circular muscular layer, is likewise well developed in this species.
The brain is massive. The cerebral sense organs are correspond-
ingly voluminous, and as usual are bathed posteriorly in the large lateral
blood lacunae. The cephalic furrows are of moderate depth, and their
lining of regularly arranged epithelial cells with long cilia furnishes
strong evidence of their sensory functions.
Reproductive Organs. — Sexual products are fully mature in
August.
Habitat. — One specimen measuring nearly half a meter in length
was found by Miss Robertson, of the University of California, in the
crevices of a rock at Dead Man's Island, San Pedro, Calif. Other speci-
mens of much smaller size were found among worm tubes and under
stones in the same locality. The species is rather common in mud
among tunicates growing on the piles of wharves in San Pedro Harbor,
where the worms are usually from 15 mm. to 15 cm. in length.
Dredged in several localities off San Pedro in 2 to 20 fms.
NEMERTEANS 1 93
20. LINEUS ALBOLINEATUS sp. nov.
Pl. XVII, fig. 2.
Body of moderate proportions for genus, rounded in esophageal
region and flattened posteriorly. Esophageal region longer in propor-
tion to length of body than in related species. Head short, broad,
commonly a little wider than neck, but not distinctly demarcated,
often slightly emarginate in front, flattened moderately. Cephalic
furrows deep, of moderate length, and, in most states of contraction of
head, well separated anteriorly ; when strongly contracted, however,
reaching nearly to proboscis pore as usual. Mouth small, situated
about as far back as posterior ends of cephalic furrows. Proboscis
small, with peculiarities as described below.
Color. — General color of body deep chocolate brown or olive
brown, with very conspicuous, clearly marked, white or pale lemon
yellow stripe extending whole length of body in the median dorsal line.
On the head this median stripe widens out to form a broad, pear-shaped
white marking (pi. xvn, fig. 2) which is often two-thirds to three-
fourths as wide as head. It is broadest, and often slightly emarginate,
very near the tip of the snout, gradually becoming narrower through
about half the length of the head, where it is lost in the dorsal stripe
which continues throughout the body. The narrow terminal border
lying in front of the white marking on head is paler brown and more
reddish than the general color of body. In some individuals a faint
reddish line extends from each cephalic furrow backward along the
lateral margin of the body. Occasionally this line becomes quite
distinct.
Ventral of same color as dorsal surface. A more rosy coloring
marks the position of the brain. Cephalic furrows often marked by
a slightly paler, rosy color. Intestinal region inclining toward an
olive brown shade. White dorsal stripe very sharply marked, with-
out gradation into adjacent color. It averages about one eighth as wide
as body, but is rather more irregular and less conspicuous posteriorly.
Ventral side of head reddish brown ; tip of snout ventrally more
distinctly reddish ; mouth reddish inside, lips paler.
In formalin, and even after imbedding in paraffin, the worms retain
their dark brown color with the strongly marked dorsal stripe and
cephalic marking as in life. In addition, a pair of narrow lines of
light color appear on the lateral margins, corresponding to the faint,
reddish lines seen in life. They are continuous with the cephalic
furrows, and lie exactly on the lateral margins throughout the body.
i94
COE
Size. — Length 100—150 mm. in extension ; width 2 mm. or more.
Ocelli. — Apparently wanting, although some irregular masses of
reddish brown pigment on the sides of the head may perhaps be con-
cerned with light perception.
Proboscis. — The pair of proboscis nerves lying internal to the
circular muscular layer are remarkably conspicuous. Inner longitu-
dinal muscular layer almost completely wanting ; consequently the
pair of nerves appear to lie directly beneath the inner epithelial layer.
Fibrous crosses between the internal and external muscular layers are
very inconspicuous, although they are sometimes indicated. In many
respects, therefore, the structure of proboscis approaches the condition
found in Tccniosoma. Proboscis is attached posteriorly at the bound-
ary of esophageal and intestinal regions by a broad and powerful
muscle to the dorsal wall of the proboscis sheath. Posteriorly to this
point the cavity of the sheath is very small, and does not extend very
far back into the intestinal region.
Vascular System. — A very large and extensive unpaired blood
lacuna is situated in the head in front of the brain, and completely
surrounds the walls of the rhynchodaeum, except on the ventral side.
Just in front of the brain the lacuna becomes broken up into smaller
spaces, which unite about the cerebral sense organs into a single exten-
sive lacuna on each side. These two large spaces are separated only
by a thin strand of tissue. Back of the mouth each sends off a series
of large anastomosing blood spaces around the lateral and ventral
walls of the esophagus. These esophageal Iacuna3 extend back nearly
two-fifths of the length of the esophageal region, where they unite
again with the lateral vessels, which have continued in the angle
between the proboscis sheath and esophagus.
Nephridia. — The nephridial system is very short and is limited to
the second fifth of esophageal region. It is remarkable for the small
number and comparatively large size of its branches. The anterior
branches lie on the outer walls of the esophageal blood lacunas, while
the main longitudinal canal on each side lies above the esophagus and
ventral to the large lateral blood lacuna. After extending backward
for a distance of about 0.3 mm. (in an individual 100 mm. long) the
branches in the esophageal lacunae join the main nephridial trunk
which passes to the dorsal wall of the lateral blood lacuna. The blood
ressels around esophagus extend posteriorly only as far as the most
posterior of these branches. The main nephridial trunk on each side
then becomes situated in the connective tissue just above the lacuna and
extends backward in this position, and without branching, for a dis-
NEMERTEANS
195
tance fully equal to that occupied by its branches, or about 0.35 mm.
The efferent nephridial duct passes directly outward from the posterior
end of the longitudinal canal and opens externally a little dorsally to
the lateral margin of body.
Cephalic glands remarkably voluminous, reaching deep into tissues
of head — fully three-fourths the distance from exterior to rhyncho-
daeum — and extending posteriorly almost to the anterior end of the
brain region, where they cease abruptly and completely.
Body Walls. — The dark pigment which gives the deep color to
the body, and which is not soluble in alcohol, cedar oil or xylol, is
situated among the cutis glands in the outer longitudinal muscular
layer in small amount, but its chief position is in a conspicuous layer
just outside the circular muscles. In the head it is scattered through
the deeper muscles and connective tissues. It is especially abundant
between the cephalic furrows and the rrrynchodaaum.
In the anterior esophageal region the cutis glands reach inward
entirely through the outer longitudinal muscular layer, except in the
vicinity of the lateral margins.
Frontal sense organs not developed as special sensory pits.
Nervous system presents no marked peculiarities. Dorsal nerve is
fairly conspicuous immediately behind brain, but is very little developed
farther back.
Reproductive Organs. — From specimens collected in August, the
genital products had evidently been recently discharged.
Habitat. — Dredged in 30 fms. off Point Fermin, near San Pedro,
Calif. But few specimens were found, and these inhabited strong,
parchment-like tubes among broken shells. Found also in 20 fms. in
Monterey Bay, Calif. (J. F. Abbott.)
The species somewhat resembles the variety of Linezis bilineatus
figured by Burger in his Naples Monograph (pi. v, fig. 15) in regard
to the markings on the anterior portions of the body, but is much less
slender, and shows many anatomical differences.
21. LINEUS WILSONI sp. nov.
pi. xvi, figs. 10, ii.
Body only moderately slender, rounded anteriorly, flattened in in-
testinal region, but with rounded lateral margins. Body is sometimes
wider in the intestinal region than the figure indicates. Head long
and slender, not marked off from body, but somewhat narrower just
back of brain ; cephalic furrows correspondingly long. Intestinal
196 COE
region commonly much wrinkled and with numerous constrictions.
Posterior extremity not very slender.
Body fragile, often constricted at the white rings described below,
and it is through these rings that the rupture usually takes place.
Several individuals broke spontaneously at the third ring, while the
rings in front and behind remained intact. This third ring seemed to
be the usual position of the first rupture.
Mouth large, situated immediately behind brain.
Proboscis pore subterminal, near ventral margin of terminal white
border. Proboscis slender, color very pale, with a tinge of yellow.
Proboscis sheath extends very nearly to the posterior extremity of the
body. Ocelli wanting.
Color. — General color of dorsal surface deep chestnut brown, slaty
brown, purplish brown, or occasionally dark drab, the shade varying
considerably in different parts of body. Some individuals are choco-
late brown in esophageal region and are much paler brown posteriorly.
The under side of the body is sometimes dark brown like the dorsal
surface, but is usually paler, with a tinge of gray, and is occasionally
light drab. Those individuals which have the less deep coloring on
the dorsal surface have a correspondingly lighter tone on the under
side of the body. When the intestinal lobes show through they appear
to be still lighter in color.
Head bordered anteriorly by a narrow terminal band of white which
also extends back along the borders of the cephalic slits. The white
color extends back a little farther in the median- line than elsewhere,
except on the margins. Sometimes the white color extends backward
to the posterior ends of the slits, both above and below, so that when
the slits are open they appear white in color (pi. xvi, fig. 10) . White
terminal border is a little less broad on ventral than on dorsal surface
and is less conspicuous owing to the paler color of the ventral surface.
Head is often paler brown in front of brain, much deeper brown an-
teriorly next the white terminal border, and is brighter red in the brain
region (both above and below) , where the rosy coloring of this organ
shows through the superficial darker brown color.
A series of very fine white rings encircles the body at intervals
throughout most of its length. These rings occasionally show slight
thickenings in the dorsal median line, but this is not usually the case.
The first of these very narrow rings appears nearly as far behind the
brain as is this organ from the tip of snout. The succeeding rings are
commonly separated from each other by about the diameter of the body
in ordinary states of contraction.
NEMERTEANS 197
Anteriorly the rings usually encircle the whole body, but farther
back they are merely indicated on ventral surface by very fine grayish
lines of much less distinctness than on dorsal surface. In intestinal
region they are sometimes separated by more than twice the diameter
of body when moderately extended.
In some individuals the white rings are very indistinct, in others they
are merely indicated on dorsal surface and are not present at all below,
while they are wanting entirely in the posterior portions of the body.
The fact that fission usually takes place through these white rings indi-
cates that there must be some peculiarity of the body walls in these
regions other than a lack or differentiation of pigment. This is true
of other species of the genus and of other genera, notably Carinella.
Body is often constricted at these points previous to rupture.
After preservation the delicate white rings disappear, and the body
assumes a slaty black appearance, sometimes more grayish below, and
with the distinct terminal white border.
Size. — Length commonly 7—I5 cm.; width about 2-6 mm.
Body Walls. — Cephalic glands not well developed. Cutis glands
limited to a rather thin but dense layer external to the outer longi-
tudinal muscles. They do not encroach on this muscular layer to any
great extent even in the intestinal region, nor do they sink in among
the muscular fibers. The pigment to which the color of the body is
due resides in the connective tissue among the cutis glands.
Nephridial and Blood Systems. — The nephridia are well devel-
oped, and extend through more than half the esophageal region.
They reach anteriorly well toward the mouth, and send large branches
among the esophageal lacunae. In each of two specimens sectioned
there was a single pair of large efferent ducts a little in front of the
middle of the esophageal region. In another specimen a single acces-
sory duct was found on one side, situated a considerable distance behind
the normal pair and toward the posterior end of the nephridia. The
single pair of efferent ducts is usually situated somewhat anterior to
the middle of the nephridial region.
Cephalic and esophageal blood lacunae large; numerous large blood
lacunae also surround the mouth. Proboscis sheath vessel leaves the
rhynchocffil a little in front of the intestinal region.
Nervous system and sense organs show few deviations from those of
related species. There is a large commissure of the esophageal nerves
just in front of the mouth. The cephalic furrows are very deep and
long. Frontal sense organs are present and well developed. They are
situated in three well marked pits, of which one is situated above the
198 COE
proboscis pore and one on each side, as in many related species, but
are well separated from the proboscis pore.
Reproductive Organs. — Sexual products had evidently been re-
cently discharged from specimens collected in August.
Habitat. — Common at Monterey, Calif., among kelp 'hold-fasts'
attached to stones on sandy or rocky bottom in 2 fms. Pacific Grove,
in crevices of rocks and under stones at low water. Dredged in several
localities off San Pedro in 2 to 20 fms.
The species is named in honor of Prof. C. B. Wilson, of Westfield,
Mass., well known for his work on Nemertean development, to whom
I am indebted for several specimens of this and of other Nemerteans,
and for valuable notes on a number of the species described in this
paper.
22. MICRURA NIGRIROSTRIS sp. nov.
pi. xvii, figs. 7, 8.
Body of small size, only moderately slender, rounded anteriorly,
only slightly flattened in intestinal region; head commonly a little
wider than parts immediately following ; tip of snout rather narrow ;
cephalic slits of moderate length; mouth as usual, its anterior end
situated opposite posterior ends of cephalic slits ; proboscis long,
flesh-colored. Caudal cirrus was not found in the few living indi-
viduals examined, but is probably present in uninjured individuals.
Color. — Dorsal surface of esophageal region bright blood-red ; in-
testinal region of same color, but deeper, and sometimes with a tinge
of purplish. Ventral surface of same color, but usually paler and
duller in tone. Head of same blood-red color as esophageal region,
with a narrow, but very sharp and conspicuous, transverse band of
white near tip of snout. In ordinary states of contraction this white
band is crescentic or V-shaped, its convex side pointing backward in
the median line. It is a little wider laterally than in the middle, and
is limited to the dorsal surface, only its ends showing from below.
Immediately in front of this is a narrow, blood-red area bounded behind
and laterally by the narrow white band, and in the middle of the red
area, and situated on the exact tip of snout, is a small, rounded, dark
brown or black spot. On this dark terminal spot are scattered a few
minute whitish flecks. The brown or black spot is only slightly visible
on ventral surface, as it lies a little more toward the dorsal than the
ventral side of the exact tip of snout (pi. xvii, figs. 7, 8).
Color after preservation brownish, but the narrow white ring near tip
of snout and the terminal black or dark brown spot are still retained.
NEMERTEANS 199
Ocelli. — None.
Size. — Length of the few individuals found 40-80 mm. ; diameter
about 2-3 mm.
Proboscis sheath extends to the posterior end of the body, and is
well developed throughout. The circular muscles of the proboscis
sheath increase to a remarkable degree near the posterior end of the
stomach region, but the fibers do not extend beneath the alimentary
canal as in several species described above. After reaching a great
thickness at the posterior end of the stomach region they suddenly de-
crease to a very thin layer at the commencement of the intestinal
region. It will be observed that although the fibers do not surround
the alimentary canal, yet the thickening of the circular layer occurs in
exactly the same region as in L. rubescens and other species, and is
probably homologous with the inner circular muscles of those species.
Proboscis. — There are only two muscular layers present, the inner
longitudinal muscles being completely wanting. Both the dorsal and
ventral fibrous crossings between the circular muscles and the outer
fibrous layer are well developed. A rather thick basement layer is
situated between the nervous plexus and the inner epithelium. Another
peculiarity of the proboscis in the single specimen sectioned is that
there is a marked differentiation of the epithelium on one side through-
out its whole length. This narrow strip of differentiated cells is quite
conspicuous in each cross section, from the fact that in it the epithelium
is much higher than elsewhere and behaves very differently to stains
than do the other cells. This specialized area is composed of very
slender cells, strongly resembling the sensory cells found in various
parts of the body, and having numerous small nuclei among their bases.
Body Walls. — The cutis glands form a compact layer beneath the
integument, and do not sink inward among the fibers of the outer
longitudinal muscles as they do in many related species. Pigment
which gives the brownish color to the body after preservation is
situated in the connective tissue among the cutis glands.
Cephalic glands are well developed, and extend posteriorly nearly
to the brain. Cephalic furrows are narrow, and are not deep. Except
at their posterior ends, they reach less than half the distance from
surface to rhynchodaaum or brain.
Alimentary Canal. — The two divisions of the esophagus described
for Lineus rubescens, L. Jlavescens and other forms are also well
marked in the present species. The change from esophagus proper to
stomach is abrupt, and the two sections are separated by a conspicuous
sphincter of connective tissue. The position of this sphincter is at
2OO COE
about two-fifths the distance from snout to intestinal region, and is in
the immediate vicinity of the efferent nephridial ducts. The histo-
logical peculiarities of the various portions of the alimentary canal
have been described on previous pages for the two species mentioned
above. The change from stomach to intestine is gradual, and the true
intestinal pouches do not begin until after the appearance of a num-
ber of shallow pouches in the walls of the stomach.
Blood and Nephridial Systems. — The nephridia are very limited
in extent, being confined to the posterior half of the esophageal region
proper. Large nephridial tubules ramify among the esophageal blood
lacunae, and are collected into a main canal on each side. At the pos-
terior end of this canal a single efferent duct passes to the exterior
above the lateral nerves as usual. The nephridiopores are thus situ-
ated in the region where the esophagus opens into the stomach. The
proboscis sheath vessel leaves the rhynchocoel at the same place.
Here, too, the esophageal lacunae unite into a pair of ventro-lateral
vessels which pass back through the stomach region, and into which
the lateral vessels empty at the beginning of the intestinal region. In
the single specimen sectioned a pair of conspicuous longitudinal folds
appear in the stomach epithelium immediately internal to the ventro-
lateral vessels, and in many places the vessels themselves are situated
within these folds.
Nervous System. — Brain and principal nerves as in related species.
The median dorsal nerve is remarkably conspicuous throughout the
length of the body. The internal median nerve, which rests on the
dorsal side of the proboscis sheath, is also conspicuous. It is best de-
veloped in the stomach region.
Cerebral sense organs present no peculiarities.
Reproductive Organs. — Sexual products were not developed in
individuals collected in August, and had evidently been discharged
some time previously.
Habitat. — Among ' hold-fasts ' of kelp and other algae, low water
to ? fms., Dead Man's Island, San Pedro, Calif., not common; on
floating kelp 'hold-fasts' off San Pedro Harbor, one specimen.
23. CEREBRATULUS ALBIFRONS Coe
pi. xvn, fig. 9.
Proc. Wash. Acad. Sci., in, p. 82 ; pi. iv, figs. 3, 4, 1901 ; also preceding
article, which has identical paging.
Specimens collected in mud in San Pedro Harbor, California,
measured upward of 30 cm. in length. Similar specimens were
NEMERTEANS 2OI
dredged at several localities off San Pedro in 2 to 20 fms. In these
individuals the coloring of the body, and the extent of the white ter-
minal border on head were somewhat different from that described for
the specimen from Alaska. The rather small mouth with its grayish
lips was situated some little distance posterior to the white terminal
border of head, and as far back as the posterior ends of the cephalic
furrows.
Color of body was usually very dark, smoky brown, with a tinge of
purple, but was sometimes almost black. A more reddish median
line was indicated only when body was much extended. Lateral
margins were not pale. Both dorsal and ventral surface of head pure
white for about three-fourths the distance to posterior ends of cephalic
furrows. Proboscis pinkish in color.
A single specimen belonging to this species was dredged in 50 fms.
between San Pedro and Santa Catalina Island, Calif. This specimen
was jet black in color except on the head, which had the characteristic
white cephalic border exactly as here described.
The species has previously been recorded only from Sitka, Alaska
(p. 85 of previous article), where a single specimen was found under
a stone at low water. It is not uncommon in mud between tides in
San Pedro Harbor. Calif.
2O2
COE
INDEX TO GENERA AND SPECIES, PART II.
Synonyms are in italics ; species new to science, and pages on which generic
or specific descriptions occur, are in black face type.
Amphiporus 124, 126
angulatus 116, 118, 122, 123
bimaculatus 118, 122
brunneus 116, 118, 122
cruentatus 154, 120, 122, 123, 126
drepanophoroides 116, 118, 122
exilis 115, 118
formidabilis 115, 116, 118, 122
imparispinosus 115, 118, 122
leuciodus 115, 118
nebulosus 122, 148
paulinus 155, 114, 122, 126
tigrinus 122
Carcinonemertes 150, 124, 126
carcinophila 154
epialti 151, 120, 122, 126, 157
Carinella 124, 125
albocincta 136, 119, 121, 125
capistrata, 118, 121, 129, 138
Cingulata 138, 119, 121, 125, 137
dinema 115, 118
frenata 129, 119, 121, 125, 136, 140
141
rubra 115, 121, 136, 141
sexlineata 115, 118, 121, 129, 138
speciosa 115
superba 129, 138, 139
Carinoma 124, 134
griffini 115, 118
mutabilis 115, 118, 121
Cephalothrix 124, 142
linearis 118, 121, 123
Cerebratulus 124, 174
albifrons 200, 118, 123
herculeus 117, 123
impressus 117
lacteus 1 86
longiceps 123
marginatus 117, 118, 123
montgomeryi 123
occidentalis 123
Emplectonema 124
biirgeri 115, 121
gracile 115, 118, 121, 123
•violaceum 115
viride 115
Lineus 124, 128
albolineatus 193, 120, 123, 128
bilineatus 195
Lineus flavescena 184, 120, 123, 128,
181, 191
pictifrons 188, 120, 123, 128
rubescens 179, 120, 123, 128, 186,
187, 191
striatus 117
torquatus 123
viridis 117, 123
wilsoni 195, 120, 123, 128
Micrura 124, 128
alaskensis 118, 123, 186
caeca 186
impressa 117, 123
nigrirostris 198, 120, 123, 128
verrilli 117, 118, 123
Nemertopsis 142, 124, 125
gracilis 142, 119, 121, 126
peronea 142, 143, 144
Oerstedia 170
dorsale 169, 122, 127
reticulatum 170, 120, 122, 127
Paranemertes 144, 124, 126
californica 144, 120, 122, 126
carnea 122, 144
pallida 121, 144
peregrina 144, 116, 118, 121
Planaria dorsalis 169
Taeniosoma 124, 127
princeps 122
punnetti 173, 120, 122, 127
Tetrastemma 124, 126
aberrans 122
bicolor 122
fcilineatum 164, 120, 122, 127
caecum 122
dorsale 169, 120, 122, 123, 127
nigrifrons 159, 120, 122, 127
quadrilineatum 166, 120, 122, 127
quadristriatum 169
reticulatum 170, 120, 122, 127
signifer 156, 120, 122, 126
vittatum 169
Zygeupolia 124, 128
littoralis 177, 120, 122, 123, 128,
1 86
Zygonemertes 124
albida 121
thalassina 121
PLATE XIV
FIG. I. Linens rubcscens sp. nov. A large individual of the deep red variety.
Pacific Grove, Calif. Enlarged five times. Compare figs. 3, 4, pi.
xv.
2. CarineUa cingulata sp. nov. Mature female, containing ripe ova.
Monterey Bay, Calif. Twice natural size.
3. C. cingulata. Anterior portion of body from ventral surface. En-
larged three times.
4. C. cingulata. Lateral view of anterior portion of body. Enlarged
three times.
5. Tetrastemma quadrilincatum sp. nov. San Pedro, Calif. Ten times
natural size.
6. Tetrastemma bilineatum sp. nov. San Diego, Calif . Ten times natural
size.
7. Tetrastemma ( CErstedia] reticulatum sp. nov. San Pedro, Calif. Ten
times natural size.
8. T. reticulatum. Anterior portion of body of individual of the variety
with confluent markings. Twelve times natural size.
9. Tetrastemma signifer^. nov. San Pedro, Calif. Three times natural
size.
10. T. signifer. Side view of head, showing position and extent of char-
acteristic cephalic marking. Enlarged ten times.
11. T. signifer. Dorsal view of head, showing typical form of cephalic
marking. Enlarged ten times.
(204)
H A.E. VOLXI
PLATE XIV
-•
LITH BRITTON £.REY. 8 F
NEMERTEANS
PLATE XV
FIG. l. Nemerfopsis gracilis sp. nov. Pacific Grove, Calif. Enlarged three
and one-half times.
2. Paranemertes californica sp. nov. San Diego, Calif. Natural size.
3. Lineus rubescens sp. nov. Head of bright red variety with few ocelli.
Monterey, Calif. Enlarged ten times. Compare fig. i, pi. xiv.
4. L. rubescens. Head of pale, pinkish variety, with regularly arranged
ocelli. San Pedro, Calif. Enlarged eight times.
5. Carinella frenata sp. nov. Mature female. The rosy color of dorsal
surface in intestinal region is largely due to color of ova. San Pedro,
Calif. Natural size.
6. C. frenata. Anterior portion of body after preservation in formalin.
The body is strongly contracted, and shows the dark color which ap-
pears on body back of the third black ring after preservation. San
Pedro, Calif. Twice enlarged.
7. Tctrastemma nigrifrons sp. nov. Variety pallidum. Monterey, Calif.
Enlarged five times. Compare figs. 6-9, pi. xvi, and fig. i, rl. xvn.
(206)
H.AE. VOL. XI
PLATE XV
LFTH BRTT TDK & R£X B T
NEMERTEANS
PLATE XVI
FIG. I. Tceniosoma funnetti sp. nov. Caught on fishing-line by Chinamen,
Monterey Bay, Calif. Somewhat less than natural size.
2. T. punnetti. Ventral side of head. Natural size.
3. T. punnetti. Ventral side of head when strongly contracted. Twice
natural size.
4. Carinella albocincta sp. nov. Body somewhat contracted. Dredged
between San Pedro and Santa Catalina Island, Calif., in 50-100
fathoms. Nearly twice natural size.
5. C. albocincta. Ventral side of head. Twice natural size.
6. Tetrastemma nigrifrons sp. nov. Anterior portion of body of the red-
dish variety. Monterey, Calif. Enlarged seven times. Compare
fig. 7, pi. xiv, fig. i, pi. xvn.
7. T. nigrifrons. Ventral side of head. Enlarged seven times.
8. T. nigrifrons. Variety bicolor. Monterey, Calif. Enlarged four
times. Compare fig. 7, pi. xiv, fig. i, pi. xvn.
9. T. nigrifrons. Variety bicolor. Ventral side of head.
10. Lineus -wilsoni sp. nov. Pacific Grove, Calif. Slightly enlarged. In
the process of reproduction this drawing has made the body of the
worm to appear much too uneven and irregular both in outline and
color.
11. L. wilsoni. Side view of head. Twice natural size.
(208)
H AE. VOLXI
PLATE XVI
8
MTH BRITTaN ^RETx'R V
NEMERTEANS
PLATE XVII
FIG. I. Tetrastemma nigrifrons sp. nov. Variety purpureum. Dorsal side of
anterior portion of body; head nnich contracted. Monterey, Calif.
Enlarged five times. Compare fig. 7, pi. xiv, figs. 6-9, pi. xvi.
2. Linens albolineatus sp. nov. Dorsal side of anterior portion of body.
Dredged in thirty fathoms off Point Fermin, near San Pedro, Calif.
Enlarged four times.
3. Lineus flavescens sp. nov. Dredged in fifty fathoms between San Pedro
and Santa Catalina Island, Calif. Enlarged three times.
4. L. JJavescens. Dorsal view of anterior portion of body showing arrange-
ment of ocelli, position of ganglia, and the two portions of esophageal
region. Enlarged ten times.
5. Lineus pictifrons sp. nov. Large individual. San Pedro, Calif. En-
larged one and one-half times. The longitudinal yellow lines are ex-
tremely delicate (and sometimes entirely wanting) in life, and appear
much too distinct in this figure.
6. L. pictifrons. Dorsal surface of head. Enlarged three times.
7. Micrura nigrirostris sp. nov. San Pedro, Calif. Enlarged five times.
8. M. nigrirostris. Ventral side of anterior portion of body. Enlarged
five times.
9. Cerebratuhis albifrons Coe. Ventral side of anterior portion of body.
San Pedro Harbor, Calif. Natural size.
(210)
H AE. VOL XI
PLATE XVI!
LITH BRTTTON &RIT,- B r
NEMERTEANS
PLATE XVIII
FIG. i. Paranemertes calif ornica Coe. Transverse section of body a short
distance behind brain, showing the two distinct layers of longitudinal muscles
(Im and lm' ) of body walls separated by a thick layer of parenchyma (par).
Numerous anastomosing bloodvessels (bv) are imbedded in this layer of paren-
chyma, as well as in that lying internal to the inner longitudinal muscular layer ;
bm, basement layer ; In, lateral nerve ; e, esophagus ; /, integument ; re, rhyn-
choccel. X 3°-
FIG. 2. P. calif ornica. Transverse section of proboscis through basis of
central stylet. The section shows the six pouches of accessory stylets symmet-
rically arranged between the outer and inner longitudinal muscular layers (oltn
and z7;«). The space between the stylet pouches is closely packed with large
gland cells (gl) filled with deeply staining secretion. In the center of the pro-
boscis, surrounded by the radially disposed bundles of the inner longitudinal
muscles, is a section of the basis of the central stylet (a), and to the right of this
a section of the duct (x) leading from the middle to the anterior chamber of the
proboscis; oep, outer epithelium of proboscis. X9°-
FIG. 3. P. californica. Longitudinal, and nearly horizontal, section of body
walls in the region of the intestinal ca;cum, the lateral diverticula of which (ic)
are arranged with great regularity in the parenchyma immediately internal to
the lateral nerve (In}. A thick layer of parenchyma (far) lies between the
diverticula of the intestinal csecum and the muscular walls (lm ' and cm ') of the
proboscis sheath ; bv, blood vessels ; In, lateral nerve ; lm, longitudinal muscles ;
cm, circular muscles ; bm, basement layer ; /', integument. X 3°-
FIGS. 4, 5. P. californica. Central stylets and bases (see pi. xxi, figs. 3-9).
Xpo.
FIG. 6. Tjzniosoma punnetti Coe. Lateral view of anterior portion of body
after clearing in cedar oil, showing distribution of the numerous small ocelli
immediately ventral to dark marking on snout, which is marked off from suc-
ceeding portion of head by an annular constriction. The mouth (m) is indicated.
x*.
(212)
H. A. E. VOL. XI
PLATE XVIII
rc
COE. DEL.
NEMERTEANS
HELIOTYPE CO.
PLATE XIX
FIG. i. Carcinonemertes epialti Coe. Diagram of body, showing ocelli, brain
lobes, alimentary canal (in dotted lines) and genital glands. X J5-
FIG. 2. C. epialti. Optical section of proboscis removed from the worm ;
ac, me, fc, anterior, posterior, and middle chambers respectively; g, gland cells;
c, canal connecting anterior and middle chambers ; ps, remnants of proboscis
sheath attached to posterior chamber ; ct, connective tissue in which posterior
chamber is imbedded ; b, basis of central stylet. X 225-
FIG. 3. C. epialti. Horizontal section of proboscis lying between the brain
lobes (br) in its natural position, and showing the posterior chamber lying at
right angles to the general axis of the proboscis. Reference letters as in fig. 2.
X225-
FIG. 4. C. epialti. Transverse section of body immediately back of brain.
The posterior chamber of the proboscis ( pc) is firmly imbedded in the surround-
ing connective tissue. Three lobes of the very short intestinal caecum (ic) are
seen ; e, esophagus lined with cilia ; cm, Im, circular and longitudinal layers of
muscles; sg, submuscular glands; In, lateral nerve; *, integument. X 15°-
FIG. 5. C. epialti. Oblique section through anterior portion of body; r,
opening of rhynchodseum; eg, cephalic glands ; ac, pc, anterior and posterior
proboscis chambers ; sg, submuscular glands ; vc, ventral commissure of brain ;
e, esophagus ; ic, rudimentary intestinal caecum ; in, intestine. X I25-
FIG. 6. C. epialti. Several stylets with their bases, showing variations in
form and size. X4°°'
FIG. 7. Transverse section of body of C. epialti, showing the thick layer of
submuscular glands (sg) and the ovaries (ov) with large ova. The intestine (in)
is reduced to a narrow canal. Other reference letters as in fig. 4. X I5°-
FIG. 8. C. epialti. Transverse section of body, showing the large number of
spermaries (t) and their distribution through the body parenchyma. Reference
letters as in fig. 4. X I5°-
(214)
H. A. E. VOL. XI
PLATE XIX
1C
,, •:<£"
In- K<>*
•4?
HELIOTYPE CO.
NEMERTEANS
,:.*, ,
PLATE XX
FIG. i. Amphiporus cruentatus Verrill. Optical section of stylet apparatus
of proboscis after extrusion. To the right of the basis is seen duct leading for-
ward from middle to anterior proboscis chamber. X 22°-
FIGS. 2-5. A. cruentatus. Outlines of central stylets and bases, showing varia-
tion in size and form in different individuals. X 22°-
FIG. 6. A. cruentatus. Outline of head, showing comparative size and posi-
tion of ocelli. X 3°-
FIG. 7. Tetrastemma reticulatum Coe. Optical section of stylet apparatus
of proboscis. X 22°-
FIGS. 8, 9. T. reticulatum. Outlines of central stylets and bases in two indi-
viduals. X 22°-
FIGS. 10, ii. Nemertopsis gracilis Coe. Outlines of bases of central stylets
in two individuals. X 3°°-
FIGS. 12, 13. Tetrastemma quadrilineatum Coe. Outlines of central stylets
and bases in two individuals. X 43°-
FIGS. 14, 15. Paranemertes peregrina Coe. Outlines of central stylets and
bases, showing the peculiar braided appearance of stylet. Both central and ac-
cessory stylets present this peculiarity. The braided appearance is probably due
to a deep spiral fluting, the translucence of the stylets allowing the flutings of
both upper and lower surfaces to appear as if actually crossing. X 3°°.
FIG. 16. Tetrastemma nigrifrons Coe. Outline of central stylet and basis
(see pi. xxi, figs. 15-23). X 22°'
(216)
H. A. E. VOL. XI
PLATE XX
HELIOTYPE CO.
NEMERTEANS
PLATE XXI
FIG. I. Paranemertes californica Coe. Dorsal view of anterior portion of
bodj, showing position of the two small ocelli on tip of snout. Position of
cephalic furrows indicated by dotted lines ; br, brain ; /«, lateral nerve. X I2-
FIG. 2. P. calif ornica. Dorsal view of tip of snout, showing the two frag-
mented ocelli. X I2-
FIG. 3. P. californica. Stylet apparatus of proboscis, showing central stylet
and basis and four pouches of accessory stylets. X 45-
FIGS. 4-8. P. californica. Central stylets and bases from five individuals,
showing variations in size and shape. X 9°-
FIG. 9. P. californica. Accessory stylets more highly magnified, showing
the peculiarly striated basal portion of each. The central stylets in figs. 4-8
are perfectly similar to these. X 22°-
FIGS. 10-12. Tetrastemma signifer Coe. Central stylets and bases from three
individuals. X 22O>
FIGS. 13-14- T. bilincatum Coe. Outlines of bases of central stylets. X 380.
FIG. 15. T. nigrifrons Coe. Stylet apparatus of proboscis. X 5°-
FIGS. 16-17. T. nigrifrons Coe. Central stylets and bases, showing more
opaque posterior portions. X I8°-
FIGS. 18-19. T. nigrifrons. Outlines of accessory stylets. X 22o.
FIGS. 20-23. T. nigrifrons. Outlines of central stylets and bases, showing
variation in form and size. Figs. 20-22 from very small individuals ; fig. 23
from the largest specimen collected. X 22°-
(218)
H. A. E. VOL. XI
PLATE XXI
HELIOTYPE CO.
NEMERTEANS
PLATE XXII
FIG. i. Ltneus rubescens Coe. Transverse section of head in front of brain,
showing distribution of cutis glands (cugl}, the highly developed cephalic glands
(cgl*), and the duct (a) leading forward beneath the rhynchodaeum (rfi). This
duct is probably the common efferent canal for the secretions of the more pos-
terior cephalic glands ; cf, cephalic furrow ; bl, blood lacuna ; cm, circular muscles.
XSo.
FIG. 2. Carinella frenata Coe. Portion of transverse section of head, show-
ing the deep ciliated pit (cf) from which a ciliated sensory canal (cc) leads inward
to the brain region, where it becomes surrounded with sensory nerves (sn) from
the dorsal side of brain to form a highly specialized sense organ ; /', integument;
_/?, layer of fibrous connective tissue surrounding brain ; cm, circular muscles ;
br, fibrous core of brain, surrounded except internally by a thick layer of nerve
cells ; bm, basement membrane. X 60.
FIG. 3. Carinella frenata. Portion of transverse section of body through
lateral sense organ; /, integument, sharply contrasted with the layer of special-
ized sensory cells lining the sensory pit; /«, lateral nerve; Zv, lateral blood
vessel ; sep, epithelium of stomach ; icm and ocm, inner and outer circular mus-
cular layers ; Im, longitudinal muscles. X 60.
FIG. 4. Tetrastemma bih'neatum Coe. Portion of transverse section of body.
The two pigment bands (pig) which give the body its characteristic markings
are situated in the midst of the longitudinal muscles (Im), on either side of the
proboscis sheath (re) ; tne, intestinal epithelium ; other reference letters as in
fig- 2. X23°.
(220)
H. A. E. VOL. XI
PLATE XXII
t-yl
sp|ps?fit
NEMERTEANS
HARRIMAN ALASKA EXPEDITION
WITH COOPERATION OF WASHINGTON ACADEMY OF SCIENCES
THE NEMERTEANS
OF THE
EXPEDITION
BY
WESLEY R. COE, PH.D.
NEW YORK
DOUBLEDAY, PAGE & CO.
1904