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vast desert region to the south and west, may enter into the
explanation of this sparse shell fauna, as well as of some of the
peculiarities of the Baikal faunula.

Southeast of the Stanovoi Range we find between the moun-
tains and the sea, the valley of the Amur and several smaller
valleys, such as the drainage basins of the Ud and the Tugar.
To the southwest the sources of the Amur emerge from the
deserts of Gobi and Dauria, and along the line of these water
courses has crept a certain number of molluscan forms inti-
mately related to or identical with those of Mongolia, China,
and the Orient. This forms the second element of the fauna
of northeast Siberia. The number of purely endemic species
is remarkably small, and a portion of those claimed to be of
this character are probably mere local mutations of widespread
Palearctic forms already known. Yet it would seem as if a
more thorough exploration must add largely to the species
now known, and it is almost incredible that the luxuriant
fertile valleys of Kamchatka and the innumerable streams
and lakes of that country should not be well populated with
mollusks.

There are few species which seem to be common to the shores
of Bering Sea, both Asiatic and American, such as Succinea
chrystS) Punctum conspectum and Anodonta beringiana. There



GENERAL DISCUSSION II

is one local species, Eulota iveyrichi, known only from Sak-
halin Island ; * and another, Helicigona sub^per sonata, from the
valley of the Ud. Three forms of Vivtpara (of which two are
probably variants of Chinese forms) are the only local species
of the vast Amur valley, or drainage, not known from other
regions. Nine specially Kamchatkan species have been de-
scribed, but about half of them are doubtfully distinct.

The total number of land and fresh water mollusks known
from the Amurland, Sakhalin, Kamchatka, the Chukchi Penin-
sula, and the Asiatic coast north of the Amur and east of the
Stanovoi Range, is only eighty-two.

Of these, thirteen are circumboreal species and twelve are
supposed to be locally peculiar. The remainder are distributed
as follows :

Percent

Europe and west Siberia 55

Northeast China 22

Common to America 13

Erratic species 10

Of these erratic species a few may be especially mentioned.
Margaritana margaritifera, as is well known, is absent from
the whole of the great northern central region of North America,
though it appears in the lower Saskatchewan, the sources of
the Missouri, and in eastern Canada, while on the Pacific it
ascends at least to latitude 56 N. In eastern Asia it is known
from Kamchatka, Sakhalin Island, the upper portion of the
Amur basin, and southern Mongolia, but I find no authoritative
record of it thence westward to northern and middle Europe.
Schrenck did not find it on the lower Amur.

Physa fontinalis is reported from the upper Amur and (in a
duck's crop) the desert of Dauria, but is not known from Siberia
proper, though common in Europe. There is an entire absence
of typical Physa throughout east Siberia, so far as reported ;
and only one species of Ancylus or Unto is known from east
of the Yenisei River of Siberia.

1 In a recent paper Hugh Fulton describes Eulota fllexibilis and E. (Euhadra}
fiscina n. sp. as "probably" from Sakhalin Island; but this seems to me very
doubtful when we consider the size of these shells and the fact that the warmest
part of Sakhalin has a mean annual temperature of only 33.4 F. and for six
months of the year the mean is below the freezing point. The shells are more
probably from Yesso.



12 LAND AND FRESH WATER MOLLUSKS

Aplexa hypnorum is known from northern Europe, western
Siberia, and the Chukchi Peninsula, but has not been reported
from eastern Siberia, or the Amur, though abundant in Alaska,
and reaching on the Taimyr Peninsula to 73 30' north latitude.

Zoogenites harpa is known from northern Scandinavia in
Europe ; from northeastern America, the Hudson Bay territory
and Southeastern Alaska, in America; but in Siberia it is re-
corded only from the easternmost margin, the Chukchi Penin-
sula, Bering Island, Kamchatka and the lower Amur. These
singularities of distribution must await much more extended
knowledge before they can be adequately discussed, but it is
believed that to some extent they are due to the transgression
of the sea, or of glacial ice, over part of the area in which a
species might naturally be expected to occur, thus delaying the
occupation of the entire region by the species concerned.

In the following table the distribution is indicated by the
headings of the six columns. Varieties are not included when
the typical form appears in the table.

Column ' Eur.' includes those forms recorded as found in Eu-
rope, including the whole of European Russia and the Caucasus.

Column ' Lena ' includes the drainage of the Lena and the
whole of Siberia from the Lena westward to the Ural Moun-
tains. It should be noted that a number of species which reach
the Lena from the west do not cross the Stanovoi Range.

Column * Amur ' includes the Amur drainage basin, the Island
of Sakhalin, and the smaller drainage areas between the Amur
and the Stanovoi Range.

Column ' China ' includes those forms which, having their
center of distribution in China or Japan, extend their range to
the drainage basin of the Amur, though often reaching only the
southern and eastern part of it.

Column ' Kam.' indicates species belonging to the area in-
cluded in the Kuril Islands, the Commander Islands, Kamchatka
proper, the Chukchi Peninsula, and northeastern Siberia east of
the Stanovoi Range and north of Aian.

Column 'Am.' includes those forms found in the Aleutian
Islands, northern and northwest America, which also occur on
the Asiatic side.



GENERAL DISCUSSION 13

The nomenclature is brought up to date as far as possible.
The absence from the list of certain names which appear in the
memoirs of Schrenck, Middendorff, and others, is only apparent ;
they are really present under their revised names. I have
accepted Simpson's determination of the Naiades, and retain, for
the variety of Unto pictorum which occurs in eastern Siberia, the
early name adopted by Rossmassler from Ziegler's MS., rather
than the very recent one which has been proposed by Wester-
lund. The list of Amurland mollusks in the Vega Expedition
report includes several which belong only to the Lena province
or western Siberia and do not occur on the Amur.

The material examined from which this and the preceding
tables have been prepared, has been derived from several
sources. The collections of the National Museum containing
the boreal shells upon which the work of W. G. Binney was
partly founded, have been of great help. I have also had the
kind cooperation of Dr. J. F. Whiteaves of the Geological and
Natural History Survey of Canada. My own collections from
1865 to 1899 in Kamchatka and Alaska have furnished much
material. I have also had interesting collections from Messrs.
Randolph, McGregor, Stoney, Hepburn, Arnheim, Krause,
Palmer and others who have visited Alaska for pleasure or in
the Government service. The collections actually made during
the Harriman Expedition were more interesting than extensive,
but have helped considerably, especially those due to the
energy of Prof. Trevor Kincaid, of Seattle, while engaged in
his entomological researches.



LAND AND FRESH WATER MOLLUSKS



TABLE III. DISTRIBUTION OF NORTHEAST ASIATIC LAND
AND FRESH WATER SHELLS.



Name of Species.


1


j


1


1

u


B

0)

M


e


Zoogenites harpa Say


o




o




o


o


Vallonia adela West


o




o




o




Vallonia pulchella Miiller


o


o


o






o


Vallonia costata Miiller


o


o


o






o


Helicigona subpersonata Midd






o








Hygromia hispida L . ..


o


o


o








Hygromia rufescens Penn


o


o


o








Hygromia stuxbergi West




o


o








Kulota arcasiana Crosse






o


o






Eulota maackii Gerstf.






o


o






Kulota middendorffii Gerstf.






o


o






Eulota ravida Benson






o


o






Eulota schrenckii Midd


o


o


o




o




Eulota selskii Gerstf.






o


o






Eulota weyrichii Schrenck






Sak.








Pupilla muscorum L


o


o


o






o


Vertigo alpestris Alder


o


o


o




o




Vertigo arctica Wall


o


o






o




Vertigo krauseana Reinh










o


o


Vertigo borealis Morel










o




Cochlicopa lubrica Miiller


o


o


o


o


o


o


Vitrina exilis Morel










o


o


Vitrina pellucida Miiller


o


o


o








Vitrea radiatula Alder


o


o


o




o


o


Euconulus trochiformis Montagu


o


o


o




o


o


Zonitoides arboreus Say






?


o


o


o


Limax agrestis L


o


o


o




o


o


Limax hyperboreus West


o


o


?




o


o


Arion hortensis Fer


o




o






o


Arion ater L


o


o


o








Incilaria bilineata Benson






o


o






Pyramidula ruderata Studer


o


o


o


o


o




Punctum conspectura Bland










o


o


Punctum ? floccula Morel










o




Sphyradium edentulum Drap


o


o






o


o


Succinea putris L


o


o


o


o






Succinea chrysis West










o


o


Lymnaea stagnalis L


o


o


o




o


o


Lymnaea peregra Miiller


o


o


o




o




Lymnaea auricular! a L


o


o


o


o


o




Lymnaea ovata Drap


o


o


o


o


o


o


Lymnaea kamchatica Midd










o




Lymnaea palustris Miiller





o


o




o


o


Lymnaea truncatula Miiller


o


o


o




o


o


Planorbis limophilus West


o


o


o








Planorbis nitidus Miiller


o


o


o








Planorbis contortus Miiller


o


o


o








Planorbis carinatus Miiller


o


o


o








Planorbis borealis (Loven) West


o


o


o




o


o


Planorbis kamchaticus West










o




Planorbis mollendorffii Dyb










o




Planorbis albus Miiller <


o


o


o




o


o


Physa fontinalis L


o




o


o





















GENERAL DISCUSSION



TABLE III. DISTRIBUTION OF NORTHEAST ASIATIC LAND
AND FRESH WATER SHELLS. Continued.



Name of Specie*.


1


3


8


I





8


Aplexa hypnorum L


o


o






Q


o


Carychium minimum Miiller


o




o






o












o


o




o


o


o




o




Valvata piscinalis Miiller


o


o


o


o
















o














o




Vivipara limnaeoides Schr






o














o


?












o


?






Kvthinia troscheli Paasch


o








o




Bytbinia kickxii Westend ,


o


o


o








Bythinia striatula Benson






o


o






Melania cancellata Benson






o


o






Sphaerium corneum L


o


o












o


o


o




o




Sphserium asiaticum Mts




o


o




o




Corneocyclas amnica Muller


o








o




Corneocyclas abdita Hald










o


o




o


o


o








Corneocyclas sequilateralis Pr










o


o


Corneocyclas sibirica Clessin




o


o






o


Cristaria herculea Midd






o


o






Cristaria plicata Leach






o


o










o


o




o


o


Anodonta woodiana Lea






o


o






Margaritana margaritifera L


o




o


o


o


o


Unio pictorum L. var. longirostris Rossmassler...,


o


o


o









IV. CONCLUSIONS IN REGARD TO THE ALASKAN FAUNA.

The fauna of Alaska, so far as the land and fresh water shells
considered in this paper enable us to judge, is composite. The
mollusks are characteristic especially of two, and to a much
smaller extent of two other, faunas. The former are limited
by topographic features. Thus the fauna of boreal Canada, in
constantly diminishing number of species, is extended to the
northwest, north of the Alaskan Range to Bering Sea on the
west and the Arctic Coast on the north.

In like manner the fauna of the northern part of the Pacific
States is extended west of the ranges which in the north repre-
sent the Rocky Mountains, and between them and the sea,
northward into British Columbia and thence westward into
Alaska, south of the Alaskan Range, until the last representa-



1 6 LAND AND FRESH WATER MOLLUSKS

tives of the fauna disappear among the islands of the Aleutian
chain. In British Columbia a few species belong to the valley
region between the Rockies and the Cascade Mountains, and
do not reach the sea coast, but these are too few to modify
perceptibly the general rule, and as a matter of fact they, like
the valleys themselves, soon disappear after crossing the 49th
parallel.

Very much the same thing is true of the birds, as I was able
to demonstrate some thirty five years ago ; and even the marine
mollusks of the Alaskan coast form a somewhat analogous
assembly.

The other two faunas concerned are those (i) of Asia, or
rather eastern Siberia, that part of Asia nearest to Alaska, and
(2) the Holarctic or circumboreal group of species which are
common to the entire boreal zone and characteristic of it, though
rather few in number.

In referring to the Canadian fauna it will of course be clearly
understood that the fauna of that part of Canada discussed in
this paper and not the entire fauna of the whole Dominion is
meant. With this reservation we may proceed to discuss the
matter from the statistical point of view.

One hundred and forty seven species or strongly marked
varieties are enumerated here from the Canadian region as
above limited, and fifteen other forms are mentioned which
though not known to cross the boundary yet in all proba-
bility will eventually be found to do so. By reference to
the preceding tables the extension of each species will be
found recorded, and the particular localities as far as discov-
erable are enumerated in the following text. Forty one spe-
cies are known from the Alaskan extension of this fauna
north of the Alaskan Mountains, or characteristic of that part
of the territory. Half of these are circumboreal or Holarctic
forms.

The fauna of British Columbia, or the British Columbian
extension of what I many years ago designated the Oregon-
ian fauna, comprises seventy five species, so far as known, to
which in all probability should be added some thirty eight which
are known to approach the parallel of 49 from the south and



GENERAL DISCUSSION 17

whicn probably cross it, making a probable total British Colum-
bian fauna of one hundred and thirteen forms. Considering
the very small area occupied by this assembly, when compared
with the vast expanse populated by the Canadian fauna, the
number is notable. Doubtless in both cases future exploration
will add a reasonable number not now enumerated or still
undescribed by naturalists.

The contributions from the Columbian assembly to the fauna
of Alaska south of the Alaskan and west of the Cascade Ranges
comprise thirty five known and six probable species, a total of
forty one forms probably inhabiting the area referred to. Some
of these, however, are common to northern Alaska also, making
the proper deduction for which we find sixty five species of land
and fresh water mollusks known to inhabit the territory of
Alaska, with six or seven more which are likely with further
exploration to be credited to it in addition to those now known,
even if no undescribed species turn up.

The vast unexplored areas, the uncertainties connected with
lists of obsolete names and doubtful identifications, the doubt
as to what may be considered specific limits in groups of noto-
rious variability, and especially the frequent absence of specimens
from which better deductions might have been drawn than were
possible from the extant literature, have all contributed to the
difficulties under which this memoir has been prepared. Those
who have done work on similar lines will understand, and will
view without undue severity, the imperfections which the author
only too well realizes, and yet which it was out of his power,
in the present state of our knowledge, to avoid. It is hoped,
however, that this summary will make the path somewhat easier
for those who follow him, and contribute a reasonable share to
the better appreciation of the facts of Nature of which it treats.
And if, among the hardy explorers of whom our neighbors of
Canada are justly proud, this paper serves to stimulate an
increased interest in the subject, the author will feel that his
endeavors are amply repaid.



SYSTEMATIC CATALOGUE OF LAND AND FRESH

WATER MOLLUSKS OF NORTH AMERICA

FROM THE REGION NORTH OF THE

FORTY-NINTH PARALLEL.

The following annotated catalogue is intended to contain a
list of all the species known to inhabit the designated region,
with the addition of a few which approach the boundary so closely
that it is highly probable that on further search their range will
be found to cross it. Names of species belonging to the latter
category are preceded by an asterisk.

It is intended that the synonymy which follows the name
shall exhibit references to the original description of the species,
to a good figure, and to the work in which the synonymy, if at
all complicated, may be found most fully set forth. The syn-
onymy of some of the genera mentioned seeming to be in great
need of elucidation, an attempt has been made to clear it up.
In other cases, where the work has elsewhere recently been
done, the generic name and authority alone are cited. For the
HclicidfB and associated forms I have depended upon the ar-
rangement of Dr. H. A. Pilsbry, the acknowledged master of
the subject ; and for the Naiades, in like manner, on the
* Synopsis ' of Mr. Chas. T. Simpson. Some of the other
groups I had previously worked up elsewhere, and have utilized
the results in this catalogue.

After the synonymy it has been attempted to state the range
of the species geographically, in general terms. This state-
ment is followed by a citation of special localities within the
designated region from which the species has been reported, and
in those cases in which the writer has verified the report by the
examination of specimens, the name of the locality is followed
by an exclamation mark.

These data are exemplified or explained by notes following
the details of geographical distribution in a separate paragraph.

The data in many cases have been taken from the literature,
a bibliography of which concludes this paper ; and it follows
that the resent writer assumes no responsibility for the identifi-

(19)



2O LAND AND FRESH WATER MOLLUSKS

cation of species so derived. Usually, however, there is no
particular reason for doubting the accuracy of these identifica-
tions. It has not seemed necessary, in most cases, to cite the
authority for the locality, a course which would have unduly
increased the bulk and diminished the clearness of the distribu-
tional statement. The authority, as a rule, can easily be found
by reference to the bibliography. In a few cases, however, it
has seemed desirable to include in parentheses the authority for
the locality cited, especially when the latter seemed unusual or
debatable.

Family HELICHXffi.
Genus Helix (L.) Pilsbry.

Helix (Cepaea) hortensis Miiller.

Helix hortensis MULLER, Verm. Terr, et Fluv., n, p. 52, 1774. BINNEY,
Land and Fw. Sh. N. Am., i, p. 181, figs. 317-320, 1869.

Helix subglobosa BINNEY, Boston Journ. Nat. Hist., I, p. 485, pi. xvi,
1867.

Range. Europe from Hungary to the Atlantic between middle
Scandinavia and the Pyrenees, northeastern border of North America.
Labrador ; Newfoundland ; Anticosti Island ! Barachois, Gaspe ;
Cape Breton Island ! * Halifax ! Casco Bay, Maine ! shore of Cape Ann
and adjacent islets, Mass.! Nantucket Island! Pleistocene deposits
near Portland, Maine !

A single specimen was once found in Greenland, but was doubtless
imported accidentally. The prevalent type is light yellow, without or
with only faint traces of bands. The former is Binney's H. subglobosa.
The wide distribution of the species, often on un-
inhabitable islets off a coast little frequented, and its
presence, which I have verified, in the glacial Pleis-
tocene of Maine, tend to confirm the view that it is
a prehistoric immigrant if an immigrant at all.
FIG. i. Helix I have seen most of the commoner varieties which
hortensts var. are prevalent in Europe, but it is obvious to the col-
subglobosa Bin- Jector thaf the brighter colored types with sna rply




define ddark bands form an insignificant proportion
of the American specimens ; while the shells as a whole seem smaller
than the average dimensions of European specimens.

1 The exclamation point indicates that specimens from this locality have been
seen by me and verified as correctly identified.



FAMILY HELICID^E 21

Helix (Arianta) arbustorum L. has been noted as an introduced
species, at St. John, Newfoundland, just outside of our region, by

Whiteaves.

Genus Epiphragmophora Doering.

Epiphragmophora fidelis Gray.

Helix fidelis GRAY, Proc. Zool. Soc. Lond., 1834, p. 67 ; Conch. Cab., 2d

ed., Mon. Helix, p. 321, pi. LVII, figs. 12, 13.
Helix nuttalliana LEA, Trans. Am. Phil. Soc., vi, p. 88, pi. xxm, fig. 74,

1839.
Epiphragmophora fidelis PILSBRY, Class. Cat. N. Am. Landsh., p. 4, 1897.

Range. Northern California to Sitka, Alaska.

Sumas Prairie, Fraser River valley, B. C. (common to 6,000 ft., J.
K. Lord) ; Chilliwak Lake, B. C. ; Victoria ! Nanaimo ! Comox ! on
Vancouver Island; Growler Cove, Broughton Strait; Union Bay!
False Bay, Lasqueti Id. ; Malaspina Inlet ; N. point Texada Island,
British Columbia ; Sitka, Alaska !

The Sitkan and Columbian specimens are apparently not markedly
different from those collected further south, and pass through the same
color variations. If there is any difference it is that the northern speci-
mens are a little smaller and exhibit no tendency to pilosity. The
two specimens obtained at Sitka were found near the Hot Springs.
There is no evidence in regard to the distribution of the species north
of Sitka, but it would not be surprising if it were eventually found to
extend on the outer islands as far north as Cross Sound.

Genus Zoogenites Morse.

This group has been united with the Acanthinula of Beck, of
which, it would seem, little is known but the shell, while we have,
thanks to Morse, a very satisfactory account of our mollusk. I prefer
therefore to defer any consolidation with Acanthinula until it is shown
to be necessary. The information to be had from Moquin Tandon in
regard to Acanthinula aculeata is unsatisfactory and insufficient.
Westerlund (1902) has proposed a genus Aulaca to contain both
(prior) genera !

Zoogenites harpa Say.

Helix harpa SAY, Rep, Long's Exped., II, p. 256, pi. xv, fig. I, 1824 ; Bin-

ney's Say, p. 29, pi. LXXIV, fig. i.

Pupa costulata MIGHELS, Proc. Boston Soc. Nat. Hist., I, p. 187, 1844.
Bulimus harpa PFEIFFER, Conch. Cat., ed. n, Bulimus, p. 305, pi. LX, figs.

17-19.
Helix amurensis GERSTFELDT, M6m. des. Sav. etr., ix, p. 517, pi. i, figs.

26, a-c, 1859.



22 LAND AND FRESH WATER MOLLUSKS

Zoogenites harpa MORSE, Journ. Portland Soc. Nat. Hist., I, p 32, pi. I, figs.

1-14, 1864; Am. Nat., i, p. 608, figs. 50-51, 1868.
Acanthinula harpa BINNEY, Land and Fw. Sh. N. Am., I, p. 156, figs. 267-9,

1869 ; Bull. U. S. Nat. Mus. No. 28, p. 185, figs. 181-184, 1885.
Zoogenetes harpa auct. plur.

Range. Northwestern Scandinavia, northeastern America, British
America near Hudson Bay, Southeastern Alaska, and the easternmost
margin of Siberia.

Konyam Bay, eastern Siberia ; Avacha Bay, Kamchatka ! Bering
Island, Commander group ; lower Amur River region. Klehini, Chil-
kat Inlet and valley, Alaska ; English River, Manitoba ! Moose
Factory ! Hudson Bay ; Minnesota ; Gaspe ; New England ; etc.

The peculiarities of the distribution of this curious little mollusk are
referred to in the general discussion of the fauna of northeastern Asia.

Genus Vallonia Risso.

Vallonia Risso, Hist. Eur. Men, iv, p. 101, 1826; V. rosalia Risso, pi. 3,

fig. 30, = Helix costata Miiller.
Zurama LEACH, Proofsheets, p. 108, 1819. TURTON, Man., p. 64, 1831 ;

Gray's Turton, p. 141, 1840. LEACH, Syn. Moll. Gt. Brit., p. 77, 1852 ;

H. pulchella Miiller.
Amplexis BROWN, 111. Conch. Gt. Brit., expl. pi. XLI, figs. 75-79, 1827; H.

pulchella Miiller.

Amplexus BROWN, op. tit., 2d ed., p. 45, 1844.
Chilostonta (sp.) FITZINGER, Syst. Verz., p. 98, 1833.
Circinaria (sp.) BECK, Index Moll., p. 23, 1837.
Glaphyra ALBERS, Heliceen, p. 87, 1850.
Lucena MOQUIN TANDON, Hist. Moll. Terr, et Fluv. France, n, pp. 98, 140,

1855 ; not of Oken, 1815, or Hartmann, 1821.


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