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PREFACE.

HARDLY any branch of natural history has been so
neglected as that which treats of the various modes by
which the different classes of organisms have become
dispersed over the surface of the globe. Scattered
observations have indeed been made by many writers,
but Lyell and Darwin were the first to gather together
the existing evidence on the subject, or to test by
actual experiment the effects of exposure to salt water
on the vitality of seeds and land-shells. Owing to this
neglect the idea has arisen that seas of very moderate
width serve as complete barriers to the dispersal of
most living things ; and it has been thought necessary
to postulate great and often repeated geographical
mutations, and even to bridge across the widest and
deepest oceans, in order to account for the actual dis-
tribution of mammals or reptiles, of plants, insects, or
terrestrial mollusca.

It was Darwin who first taught us that these assump-
tions of vast and repeated changes in the distribution
of sea and land were at once inadmissible and un-
necessary. By his original and masterly investigation

vi PREFACE.

into the phenomena presented by oceanic islands he
showed that these islands had never been connected
with the continents, as had been almost invariably
assumed by previous writers, and, consequently, that
their entire fauna and flora must have originated from
such species as could, in the course of ages, have reached
the islands by natural means of dispersal. Hence the
importance of studying what are the means of dispersal
of the various groups, and why it is that, with the two
absolute exceptions of mammals and amphibia, none of
the larger groups of animals or plants are invariably
absent from this class ofislands. Asacorollary from his
investigation he was led to conclude that the great
oceans were, broadly speaking, permanent features of
the earth’s surface, and that it was scientifically in-
admissible to bridge them over in various directions and
at various geological epochs in order to provide a
short and easy road for the passage of beetles or
snakes, snails or frogs, and thus save us the trouble of
solving the problem of their actual distribution by less
obvious and also by less heroic means.

Having myself devoted some time and research with
the object of showing that almost every anomaly in the
distribution of animals and plants may be explained
by a careful consideration of the various means of
dispersal which organisms possess, combined with the
climatic and geographical changes which are known to
have occurred during later geological times, and taking
into account the known distribution of the several
groups at remote epochs as proved by the discovery of

PREFACE. vil

fossils in regions far removed from the lands now in-
habited by their living representatives, I am especially
interested in Mr. Kew’s attempt to bring together all
that is known of the means of dispersal of one of the
groups as to which such information was most needed.
He has devoted to the task much labour and research,
and has brought together a mass of information of
great value. Many of the facts he adduces are so
curious and interesting that they will attract the atten-
tion of many classes of readers and thus lead, it is to be
hoped, to the accumulation of facts which are still
required to complete our knowledge of this important
subject.

I heartily congratulate the author on his choice of so
useful and interesting an inquiry for his first work, and
on the systematic and accurate manner in which he has
marshalled the facts he has collected. Many books of
far greater pretension, even though they should contain
descriptions of scores of new species and work out their
internal structure with the greatest accuracy, may yet
be of less interest to the philosophical: naturalist than
this unpretending little volume. In its pages we are
afforded a glimpse of what seem at first sight to be
but trifles and accidents in nature’s workshop, but which
are really the tools with which she produces some of her
most striking results. It is owing to such trifling occur-
rences as the occasional attachment of a living shell to
a beetle’s leg, or the conveyance of seeds in the mud
adhering to a bird’s foot, that many remote islands
have become stocked with life, and the range of species

Vill PREFACE.

extended or modified over the earth; while through
changes of the organic environment thus effected even
the origination or the extinction of species may have
been brought about.

ALFRED R. WALLACE,

PV TLODUCTORY NOTE.

THE distribution of fresh-water and land shells has been
looked upon as presenting special difficulties on the
theory of single birth-places or “centres of creation.”
Mr. Darwin, in characteristic letters, in 1856-7, spoke of
being driven mad by land molluscs, and fresh-water
kinds, he said, had been a horrid incubus.’ In the
“Origin of Species,” however, he was able to suggest
several possible means of dispersal both for terrestrial
and aquatic groups, and speaking of the latter he re-
marked that many facts throwing light on the subject
would doubtless be discovered.” Quite a number of
facts of the kind indicated have been recorded since the
publication of the “ Origin,”
together with some hitherto unpublished items—both
as regards fresh-water and land shells—is now given.

and a collection of these

In compiling this, and the essays on subsidiary subjects
also given, the writer has received constant help in the
way of notes, references, &c., from a number of friends
and correspondents whose courteous co-operation is

1 “ Life and Letters,” vol. ii. pp. 85, 93.

2

* “ Origin,” 1859, p. 385, and see ed. 6, pp. 344, 353.

xX INTRODUCTORY NOTE.

gratefully acknowledged; to Mr. Robert Standen and
Mr. J. Ray Hardy he is under obligation for many
striking facts and the loan of specimens, to Mr. G. K.
Gude and Mr. R. W. Goulding for the preparation of
several translations, and to Mr. C. T. Musson for per-
mission to use a manuscript work on the “Land and
Fresh-water Shells of Nottinghamshire,” containing
excellent essays on local distribution and means of
dispersal.

To Dr. Alfred Russel Wallace, who, with great kind-
ness, has looked over the proofs and contributed the
foregoing preface, the writer owes a special and
irredeemable debt of gratitude.

With a view to the collection of further facts illustrat-
ing the means of dispersal possessed by molluscs and
allied animals, Dr. Wallace has suggested that the
writer should invite “naturalists and sportsmen in all
parts of the world” to co-operate by furnishing notes
of, or references to, observations which they may have
made or recorded, or which they may be able to make
or record in the future. Many persons, both at home
and abroad, he imagines, would make observations “ if
they knew what was wanted, and had the address of some
one who would appreciate and use them.” It may be
mentioned that, amongst other things, the examination of
large numbers of floating trees, etc., encountered upon the
ocean, and of drift-timber and brushwood found stranded
upon the coast-line, would possibly be productive of
surprising results, as also would the careful and system-
atic inspection of the feet and feathers of birds shot

INTRODUCTORY NOTE. xi

on the wing. It is desirable, moreover, that search
should be made for living mollusca in the crops of birds
and in the stomachs of other shell-eating creatures.

It need hardly be added that communications
with which the writer may be favoured will be wel-
comed at the under-mentioned address, and carefully
acknowledged.

Fe Wo

5, Giesbach Road, Upper Holloway, London.
September, 1893.

re SaarH OY Ee thei Set ade i a
Saath aia ARMIN ARIS AES
, Sahn ae hy raat ” Petar eee ca! plervein ieee a i
i
° ‘ oe Ps owe rr a

Maley ti ee [hy ae ‘ote, Lae ere | y an
cil Le are - hy Pues | or) ae Oe io lisoe!

Bes *2

CORTEN 1S.

CHAPTER. I:
PAGE
FRESH-WATER SHELLS—ANOMALIES IN LOCAL DISTRIBU-
TION

GHAPTER. II:

MEANS OF DISPERSAL . : : : : . R =) 2
CHAPTER, ‘Il.

TRANSPLANTATION OF BIVALVES : ; : : =) 56
CHAPTER IV.

TRANSPLANTATION OF UNIVALVES . : : : » 8
CHAPTER V.

LAND SHELLS: THEIR TENACITY OF LIFE ‘ : « . 96
CHAPTER VI.

MEANS OF DISPERSAL . - : ‘ : : ; » TI5

GHAPTER. VII.

DISPERSAL OF SLUGS . : : : 7 : s eh67

XIV CONTENTS.

CHAPDER Vill.

DISPERSAL OF FRESH-WATER AND LAND MOLLUSCA BY
MAN

CHAPTER IX,

ON THE FRESH-WATER AND LAND MOLLUSCA INTRO-
DUCED INTO THE BRITISH ISLES BY HUMAN AGENCY

INDEX

PAGE

178

foe DISPERSAL, OF SHELLS.

CHAPTER, I,

FRESH-WATER SHELLS—ANOMALIES IN LOCAL
DISTRIBUTION.

MOST naturalists of recent years have agreed that “the
several species of the same genus, though now inhabiting
the most distant quarters of the world, must originally
have proceeded from the same source, as they are de-
scended from the same progenitor,” and also, of course,
that “the individuals of the same species, though now
inhabiting distant and isolated regions, must have pro-
ceeded from one spot, where their parents were first
produced,” ' and the acceptance of this view, now that it
is very generally admitted that the great oceans—the
most formidable of all barriers limiting the migrations
of non-marine animals—have occupied the same general
positions as they do now throughout all known geological
periods,’ has naturally lent much interest and import-

1 Darwin, “ Origin,” ed. 6, 1890, pp. 319-20.

2 Wallace, “Island Life,” 1880, p. 144 ; ed. 2, 1892, pp. 149-50;
Darwin, “ Origin,” p. 288. For a summary of the evidence on this
head, see “ Island Life,” pp. 101-2; ed. 2, pp. 103-5.

B

2 THE DISPERSAL OF SHELLS.

ance to the investigation and study of means of dispersal.
Obviously, we are not now authorized in explaining all
the difficulties which geographical distribution presents
by suggesting multiple centres of creation or former
radical changes in the relative positions of land and sea.

The wide ranges and singular distributions now en-
joyed by many organisms seem, in these circumstances,
well-nigh inexplicable ; the ranges of some genera, as
everyone is aware, are almost world-wide, some species
extend over immense areas, and, moreover, there are
cases in which the same or closely-allied forms occur at
isolated points in remote parts of the world. Highly
effectual means of dispersal of some kind or other must
certainly have been in operation, for it is clear that the
distribution, at least of many groups, cannot have
resulted from gradual migration by ordinary modes of
progression, and this is the more apparent, of course,
with pre-eminently slow-moving inland animals like
fresh-water and land mollusca. To such creatures
evidently, as far as voluntary migration is concerned,
even small arms of the sea, arid deserts, and elevated
mountain-chains must be almost, or, perhaps, absolutely,
impassable; but it has been remarked that such ob-
stacles are not likely to have endured so long as the
oceans.! It will not be forgotten, of course, that the
great changes of level known to have taken place
“within the period of existing organisms” certainly
remove many difficulties, and it should be borne in

1 See “Origin; p.317-

FRESH-WATER SHELLS. 3

mind, more especially in connection with discontinuous
ranges, that many types are of immense antiquity,
having survived numerous changes of climate and great
oscillations of level, so that the distant regions in which
they are now found may,in some cases, have formed
parts of former continuous and very wide ranges. It
ought also to be remarked, perhaps, that some eminent
biologists, as, for instance, Professor Semper, think it
possible that similar faunas in distant parts of the world
may sometimes have been brought about by the con-
vergence of formerly distinct types,’ and this, of course,
cuts the Gordian knot much in the same way as did a
formerly prevalent belief in multiple centres of special
creation. Fresh-water forms are said to have been
derived originally from the sea, and even now certain
marine animals in all probability are gradually adapting
themselves to fresh water, a process which may have been
going on all through the ages, but I do not see that this
much affects our present inquiry ; it should certainly be
taken into account, however, whenever the occurrence at
distant places of allied fresh-water forms having near
relatives in the sea is considered.

Fresh-water shells, it seems, if left to their own
endeavours, would generally find great difficulty in
extending their range beyond the limits of their own
river-basin, for the dry land obviously presents a for-
midable barrier to these animals, and, as Mr. Darwin
remarks, each river-system with all the pools and lakes

1 Semper, “ Animal Life,” Eng. transl., ed. 4, 1890, pp. 294 and
461.

B 2

4 THE DISPERSAL OF SHELLS;

in connection with it seems completely cut off from
every other river-system of the same country, and still
more complete, of course, is the separation between the
fresh-waters of distinct islands and continents... Some
of the univalves live much out of water, but, even with
these, powers of voluntary migration over land must be
very limited. I find, however, in Mr. Musson’s manu-
script, a statement that a pond-snail (Lzmenca truncatula)
has been seen travelling across dry fields. Bivalves
seem practically incapable of migrating voluntarily
beyond the limits of their own element and its imme-
diate surroundings; some kinds, as is well known, are
occasionally encountered in a living state, buried in
sand, or amongst moss, &c., at distances from water,’
but it is unlikely that they intentionally proceed to
such spots. Notwithstanding these disabilities, how-
ever, we find a wide distribution characterizing many of
the creatures, both univalves and bivalves, together
also with most other freshwater productions. Some
species range very widely, and, as Mr. Darwin observes,

1 Darwin, ‘“ Nature,” xvili. (1878), p. 120, and see “ Origin,”
Pp. 343-

2 See, for instance, W. Thompson, “ Ann.and Mag. Nat. Hist.,”
vi. (1841), p. 195; J. ‘G.. Jeffreys, “British ‘Conchology,”? an
(1862), p. 17; L. Reeve, “ Land and Fresh-water Mollusks,” 1863,
p: 236; -G.. Roberts,. “ Zoologist,” (3), 1x: (1885); p. 470 oes
Milne; “Journ. of “Conch.” vi. (1891), pp. 413-14, 4105 CC)
Simpson, ‘‘ Nautilus,” v. (1891), p. 16.

3 One of the pea-shells (P2stdium pusillum), it is true, is said to
have been seen creeping in damp moss 6—20 paces from water ;
D. Weinland, quoted in “ Zoological Record,” xiii. (1876), Moll,
P. 57:

FRESH-WATER SHELLS. 5

allied forms, which “‘ must have proceeded from a single
source,” prevail throughout the world.’

Much has to be explained, also, even in local distri-
bution. Every naturalist remembers having seen shells
in puddles formed after periods of excessive rain, in
pools in quarries and pits, in drinking-troughs and
water-butts, in tanks on the roofs of buildings, in newly-
formed reservoirs and artificial lakes, in ponds on open
pastures far away from the nearest streams, &c. As
the Rev. James Dalton long ago remarked, the creatures
seem to possess a “ mysterious faculty ” of finding their
way to the most unlikely habitations.” Dispersal by
some means must be constantly going on.

That almost every isolated cattle-pond which a farmer
digs perhaps near the middle of an upland pasture should
come to possess a molluscan fauna within a few years of
its formation, though a matter of common observation,
is certainly a surprising fact, and it is remarkable that
writers of local lists when recording the existence of
shells in such places have so generally passed over the
circumstance without comment. If multiple birthplaces
were possible it is evident, of course, that the spreading
of forms from their several centres would have to be
explained, for no believer in multiple centres, I imagine,
would go so far as to suppose that cattle-ponds are

" © Origin,” p. 344; but see Semper, ‘Animal Life,” p. 298.

2 J. Dalton, ‘ Zoologist,” xix. (1861), 7318-g. Early in 1890
Mr. W. H. Heathcote saw a specimen of Limnea truncatula
crawling on the top of the tower of St. James’ Church, Preston,
ninety feet or more from the ground !

6 THE DISPERSAL OF SHELLS.

stocked by special acts of creation; curiously enough,
when a boy, I believe before reading any books, I was
much puzzled at seeing some living thing in a newly-
formed pool, and remember concluding that it must
have been created there! Mr. Clement Reid, who has
attended to this subject, has been in the habit of noting
from time to time the species occurring in artificial
ponds and other isolated waters; by so doing, as he
puts it, “ one can see the accumulated results of many
years’ dispersal, and can get some idea as to the extent
to which it must be going on.” Asa rule, he tells me,
such waters only contain Lzmnee (generally L. peregra)
but the species aurzcularia, truncatula, stagnalis, and
palustris also occur, associated, sometimes, with the
smaller Planorbes (generally spzrorbis and vortex), and
Physa fontinalis. The presence of bivalves, he adds,
“is quite exceptional ; the smaller Pzszdza, especially
P, pusillum, occasionally occur, and I have once found
Spherium corneum, but never Unzo or Anodon ;” oper-
culate pond-snails are almost invariably absent. More
recently, this observer has published an able paper on
the “ Natural History of Isolated Ponds,” based, for the
most part, upon observations made on the South
Downs.’ These undulating chalk hills, it is stated,
constitute a pre-eminently dry district in which, in order
to provide water for the cattle, ponds have to be dug at
comparatively short distances from each other. When
rendered impervious by puddling with clay or chalk-

1 Clement Reid, Trans. Norfolk and Norwich Nat. Soc., v.
(1692), 272.

FRESH-=WATER SHELLS. 7

mud, or sometimes by a lining of concrete, such “ dew-
ponds” retain a store of water, except during droughts
of exceptional duration, derived from “the rain, dew,
and condensation of the mists which often hang on the
tops of the hills.’ Many occur at the distance of a mile
or even two or three miles from the nearest stream or
marsh, and, as the Downs rise to 800 feet, their average
height is fully 200 feet above other water, so that the
aquatic animals and plants found therein must certainly
have been “transported uphill as well as across un-
congenial tracts of dry grass,’ or be descended from
ancestors thus transported. Notwithstanding this ex-
treme isolation, however, the ponds are often found to
be inhabited by fresh-water shells. Two, at least, of the
three examples given by Mr. Reid were thus inhabited :
(1.) “Large dew-pond on the open Down nearly a
mile east-south-east of Amberley Station ; height, 310
feet above the sea, and 300 feet above the marshes of
the Arun; nearest water, the marshes of the Arun,
distant half a mile. Noted 15th November, 1884.
This pond has evidently been made several years, and
is now full of water-plants. The species observed, were
Juncus (not in flower or fruit, though very abundant),
Potamogeton densus (very common), Ranunculus aqua-
tilts, Chara, and a single small, but vigorous tuft of
Elodea canadensts. ... (I have apparently omitted to
note the mollusca from this pond, if there were any).
(2). “ Redlion Pond, on the open Down, three miles
south-east of Lewes ; height 540 feet above the sea, and
530 feet above the marshes of the Ouse ; nearest water,

8 THE DISPERSAL OF SHELLS.

the marshes of the Ouse, distant seven furlongs. Noted
June, 1890. The species observed were Limmnvea feregra,
Ranunculus aquatilts, and Elodea.

(3). ‘‘Small pond by the side of the high road, half a
mile west of Christ Church, Stansted (near the Hamp-
shire border) ; height, 312 feet above the sea, and 220
feet above the nearest stream ; nearest water a small
stream, distant about two miles to the south-south-west,
but apparently containing neither of the plants found in
the pond. Noted September 25th, 1891. The species
found were Limnca peregra, Planorbis spirorbis, Potamo-
gcton natans, Zannichellia palustris, all abundant; the
two plants in fruit, and carrying many eggs of Limuea.
Zannichellia occurs again in a pond in a brick-field, two
miles to the south-west, but does not fruit there ; the
nearest natural station for it seems to be in the slightly
brackish marshes near Emsworth, four miles to the south.
Potamogeton natans can probably be found within three
miles. This pond is overshadowed by a large oak, and
supplied by dew and rain off the road; neither of the
plants, however, is a species likely to be brought to the
locality by carts or on the hoofs of horses.”

The absence of bivalves from such ponds, it is true,
need hardly be regarded as surprising, but I am unable
to agree with Mr. Reid in thinking that the occurrence
of the smaller kinds in isolated waters is likely to be
altogether exceptional. The creatures may easily be
overlooked, and Mr. Reid tells us, in the paper just
quoted, that he “could only stay to note the common
and conspicuous animals and plants.” The catch-water

FRESH-WATER SHELLS. 9

or dew-ponds of the Lincolnshire Wolds, dug out of
the chalk and puddled with clay, though often or usually
dry in summer, do not essentially differ, I imagine, from
those of the Downs. Recently my brother showed me
a pond of this kind, the most isolated he could find
about Louth, situate on one of the highest of the rolling
chalk hills in the neighbourhood, at a great distance
from any other water, and in this were bivalves of two
kinds, Spherium lacustre and Pisidium pusillum, the
former in good numbers. In ponds on lower ground
and less remote from other water (but perfectly isolated
and out of the reach of floods), where the conditions of
existence are probably more congenial, small bivalves
certainly seem to occur with some frequency, Uncon-
genial conditions, Mr. Reid remarks, have probably
much to do with the poverty of the fauna and flora in
dew-ponds. Mr. Musson, who, also, has paid some
attention to the subject, speaks of Spherium lacustre as
a species commonly found in “upland ponds” [ponds
without inlet or outlet, dependent directly on rain-water
for their supply, as distinguished from such as are
affected by floods and rivers], in which also, as he adds,
Spherium corneum, more generally found in rivers,
canals, &c., occurs occasionally. A small “upland”
horse-pond, near the Hemlock Stone, near Nottingham,
examined by him during 1883 and 1884, was inhabited
by six kinds of molluscs, of which three were bivalves:
Spherium lacustre and two species of Prstdium ; and
three univalves: Planorb¢s nitidus, Pl. nautileus, and
Auncylus lacustris—a strange set of species, as he remarks,

IO THE. DISPERSAL OF SHELLS.

to have come together in one small pond. I will here
add particulars of the Lincolnshire catch-water pond
referred to, and of one or two other isolated ponds, &c.
Details, in such cases, must always be tedious, but the
subject, as it seems to me, is an interesting one.

Pond at Welton-le-Wold, Lincolnshire-—A small
pond, about nine yards in length, on the Wolds not far
from Louth, near the crown of Bunker’s Hill. <A great
distance from any other water, and far above the
nearest stream. Between arable fields, but with no
ditches near. Probably old, having existed, perhaps
when the wold was unenclosed ; remains of a rough
pump are to be seen on one of the banks. Water
derived solely from rain, &c., falling in the immediate
neighbourhood. No water-weeds are likely to have
been planted by man. Four species of shells occur:
Spherium lacustre, Pisidium pusillum, Limnea peregra,
and Limnea truncatula.

Pond at Riplingham, Yorkshire—Mr. ¥. W. Fierke,
of Hull, has favoured me with particulars of a somewhat
similar pond on the Yorkshire Wolds, at Riplingham,
scooped out of the chalk and laid over with clay, and

supplied with water by rain and surface drainage only.
The ever present Limnca peregra,’ among higher
organisms seemingly the only inhabitant, exists in
countless numbers, almost covering the bottom. There
are no water-weeds. Many ponds of this_ kind,

' Specimens were exhibited at the Conchological Society on
7th October, 1891 ; see “ Journ. of Conch.,” vi. (1891), 398.

FRESH-WATER SHELLS. II

Mr. Fierke states, are scattered about the district, but
are usually unproductive.

Pond at Tothby, Lincolnshire, dug in 1871.—Particulars
communicated by Mr. J. E. Mason and Mr. J. B. Davy.
Of special interest from the fact that the date of excava-
tion is known. Situate on the clay near the foot of the
Wold, in the corner of an arable field (T. C. Johnson’s)
at Tothby, on the borders of the parishes of Alford and
Rigsby, and said to be quite isolated from other water.
Two small fence-ditches, sometimes containing a little
water received from the under-drains, open upon the
pond; but Mr. Mason, in whose knowledge of the
locality I have the utmost confidence, expresses the
opinion that it is in the highest degree improbable, next
to impossible, that a flood should have caused a com-
munication with any other pond or water-course. Of
the plants, none are of kinds likely to have been
intentionally introduced ; duckweed and two Potamoge-
tons are conspicuous. Both the common newts occur,
and water-bugs and beetles abound, but Mr. Mason has
not seen a fish of any kind, although the nearest ponds
are well stocked with sticklebacks. Of molluscs,
Mr. Davy, in 1891, obtained two species, Lzmnca
peregra and Spherium lacustre, both in fair plenty.
The presence of the former, which, it is said, is generally
the first to appear in newly-formed ponds," is of no
special interest, but the establishment of a bivalve-shell

1 See G. Roberts, “‘ Topography and Nat. Hist. of Lofthouse,”
E552; p. 167.

12 THE DISPERSAL OF SHELLS.

(S. Zacustre) in an isolated pond dug only twenty years
before is certainly worthy of note.

Pond at Rigsby, Lincolnshire-——Mr. Mason and Mr.
Davy have also favoured me with particulars of another
pond, probably much older than the last, and remark-
able for possessing a surprisingly rich molluscan fauna.
Situate in the parish of Rigsby (in the “ nineteen acres,”
abutting in part on Rigsby Wood), of small size, sur-
rounded by arable land, and apparently perfectly
isolated. The nearest ditch, it is true, is within about
eleven yards, but the water in the pond is considerably
above the level of that in the ditch, and Mr. Mason
assures me that no possible flood can have reached the
pond; with the exception of the water carried in
through the pipe-tiles of a small wnxder-drazn, it gets
every drop from rain and surface drainage. It is
apparently very old, and was probably dug at a time
when the surrounding district was an unenclosed sheep-
walk, in which case, of course, it is just possible that
sheep or cattle may have been instrumental in intro-
ducing shells or ova adhering to their feet or legs.’
There are no plants likely to have been intentionally in-
troduced, duckweed (Lemna minor),a Potamogeton, some
grasses, &c., making up the principal vegetation. Mr.
Davy and Mr. E. Woodthorpe, who have searched for
shells, have discovered no less than four species of bi-
valves and five univalves, representing together five

' The same may be said of most ponds which appear to be

more ancient than the surrounding enclosures, as well as of those
situate on large unenclosed pastures.

FRESH-WATER SHELLS. BS

genera: Spherium corneum, Spherium tlacustre, Pisi-
dium fontinale, Pisidium pusillum var. obtusalis, Pla-
norbis nitidus, Planorbis nautileus, Limneea peregra,
Limnea stagnalis, and Ancylus lacustrts.

Ponds at Finchley, Middlesex—Near the middle of a
meadow at Finchley (by the side of the foot-path to
Hendon, and in the third field from the road at East-
end, Finchley)’ are two ponds, without inlet or outlet,
which lie near together upon a ridge slightly higher than
the surrounding ground, perfectly isolated from other
water, and which, it seems certain, can never have been
affected by floods; they are evidently artificial, but
probably very old, and neither is of large size, the larger
measuring about 29 x 10 yards, and the smaller about
14 x 7 yards. In the case of ponds near large towns
or cities having many shell collectors one can never be
absolutely certain that molluscs have not been intro-
duced intentionally or unintentionally by human agency,
but it is unlikely that this often happens, and I know
of no reasonable ground for such a suspicion in the
present case. Ivy-leaved duckweed (Lemna trisulca),
water-thyme (Elodea canadensis), frog-bit (Hydrocharis
morsus-ran@), and a Potamogeton are the principal
water-weeds, and there are none of kinds likely to have
been planted by man. Molluscs occur in abundance,
the bivalve Spherium corneum being plentiful in the
mud, and very fine. In the larger pond four species,
belonging to four genera, are found, all in good numbers :

1 This field, I find, is already being cut up for building
purposes,

I4 THE DISPERSAL OF SHELLS.

Spherium corneum, Planorbis carinatus, Limnea stag-
nalis, and Ancylus lacustris; while the smaller one,
about thirteen yards distant, contains Spherium
corneum, Planorbis carinatus, Limnea stagnalis, and
Limnea truncatula ; and some idea of their individual
abundance may be gathered from the following note
of four caddis-cases—selected from a number col-
lected in both ponds—bearing, together, no _ less
than forty-five shells. (1) Five shells attached, two
of Spherium, one of Limn@ea, and two of Axncylus ;
(2) ten shells attached, eight of Sph@erzum, one of
Planorbis, and one of Limne@a, (3) fourteen shells
attached, eleven of Sphe@rium, one of Planordis, and
two of Limne@a,; (4) sixteen shells attached, one of
Spherium, and fifteen of Planorirs,'

Pond near the Black Hills, Leeds—During a period
extending over some twenty-four years Mr. W. Nelson
has obtained Spherium lacustre, Pisidium pusillum,
Planorbis nautileus, Planorbis vortex, Planorbis carina-
tus, Planorbis corneus, Planorbis contortus, Physa fon-
tinalis, Limnea peregra, Limnea stagnalis, and Ancylus
lacustris from this pond, which is of small size, isolated,
as Mr. Taylor assures me, from other water, and with no
inlet except “a natural drainage from the field.” In
times of heavy rain there is an overflow of surplus
water, but this disappears in or spreads over the land

1 Mr. J. W. Taylor, of Leeds, has obligingly looked over the
shells from these ponds and those at Welton-le-Wold, Tothby,
and Rigsby, determining the P7s¢d/a, and confirming my naming
of the other kinds,

FRESH-WATER SHELLS. 15

and has never been seen to reach a small stream which
runs along at the bottom of the field, perhaps 150 yards
away. The shell-fauna, it will be seen, is a wonderfully
rich one, comprising, as it does, eleven species belonging
to six genera, including two bivalves and no less
than five kinds of coil-shells or Planorbes. But the
pond is additionally interesting from the fact that
a number of species seein to have been introduced
quite recently. During 1860-3, when Mr. Nelson was
in the habit of collecting there, it yielded only four
species, Spherium lacustre, Pisidium pusillum, Planor-
bis nautileus, and Limna@a peregra, so that since that
period its fauna seems to have received a surprising
number of additions. About 1873, Mr. Nelson re-
sumed his visits to the locality and found Planorbis
corneus, but no further kinds were obtained till 1883,
when during frequent visits, he found six others, Physa
fontinalis and Planorbis vortex in the spring, Planorbis
carinatus, Limnea stagnalis, and Ancylus lacustris a
little later, and finally, in June, this small, but prolific
pond yielded its fifth Planorbis, Pl. contortus. Mr.
Nelson, who has published these facts,’ is a collector of
very great experience and a naturalist of ability, so that
I dare not suggest that all the shells ultimately found
probably existed in the pond, overlooked, during 1860-3,
and Mr. Taylor and Mr. J. Beevers, who collected with
him, agree that the four species named were the only
forms in the pond at that time. The case is at least

MW. Welson,)** Journ. of Conch.,” iv..(1883)) 177:

16 THE DISPERSAL OF SHELLS.

interesting and curious and is certainly worth giving,
but I do not put it forward as of so much value for
us as some of those previously referred to (the dew-
ponds of the South Downs for instance), for I do not
think it seems so clearly to imply the existence of dis-
persal independently of man; on the Downs, day after
day, as Mr. Reid states, “one meets only the shepherd
tending the sheep, or the farmer making a short cut
across the open country,” and everything seems to
show that shells are not likely to have been carried
to the dew-ponds by human agency, but in Leeds,
on the other hand, quite a number of shell collectors
have resided for many years past, and it seems just
possible that someone, having collected a number of
molluscs for which he had no particular use, may have
thrown them into the pond in question, but this, of
course, is very unlikely ; it seems just possible, also,
I venture to think, that the conchologists who col-
lected there during the earlier years may have uncon-
sciously introduced some of the species adhering (per-
haps when very minute) to their scoops or collecting
nets, and it is at least noteworthy that P/anorbes,
so conspicuous among the additions, commonly thus
adhere.

The presence of Limnea humilis, “ by the hundreds,”
in a small artificial pond, fed by a windmill from a
well twenty feet deep, was noted in the West American
Scientist during 1885 ; the pond, situate in the vicinity
of Todos Santos Bay, Lower California, was only about
ten years old, a few inches deep, and about six feet

FRESH-WATER SHELLS. t7

across, and for miles there was no natural surface water.’
Limn@a macrostoma, according to Dr. Ingalls (who
communicated the fact to Mr. Binney in 1860), not seen
elsewhere within twenty miles, seems to have suddenly
appeared in a pool, about twenty feet in diameter
(entirely cut off from streams and fed by a spring),
which had for years been frequented by the observer for
Desmidia, &c.; this, he added, “‘ comes as near a case of
spontaneous generation as anything within my observa-
tion.”? The Rev. James Dalton, in 1861, mentioned
the finding of a dozen species of shells in a pond at
North Stainley, near Ripon, at least half a mile from
any other water, and two of these, Limncea stagnalis
and Cyclas caliculata,|= Spherium lacustre|, did not,
as far as was known, occur elsewhere in the neighbour-
hood.’ The occurrence of shells in the isolated waters
of pits, quarries, brick-yards, &c., is notorious; Mr.
Standen tells me, for instance, that some recently ex-
cavated “brick pits” at Cheetham Hill, near Man-
chester, though far from other ponds, swarm with
common molluscs, Spherium corneum, Pisidium fon-
tinale, Limnea peregra, Limnea truncatula, Linnea
stagnalis,&c. Thepresenceof Anodonta fluviatilzsinnum-
bers in a hole, south of Dover Plains, Eastern Duchess

1 “ West American Scientist” (C. R. Orcutt, editor), i. (1885),
74.
? WwW. G. Binney, “Land and Fresh-water Shells of North
America,” part 2 (1865), p. 37, “Smithsonian Miscellaneous Col-
leetions,” vii., 1867.

% J. Dalton, “ Zoologist,” xix. (1861), 7318-9.

18 THE DISPERSAL OF SHELLS.

Co., N. Y., formed by peat-digging some thirty or more
years previously, was noted in 1889, by Mr. W. S.
Weator

The presence of shells in reservoirs on high ground
has been referred to as indicating dispersal by animal
agencies, but in such cases, I suppose, there is often a
possibility that the animals may arrive, perhaps as fry
or ova, in the water with which the reservoirs are
supplied, and, doubtless, a similar explanation very
often applies in the case of shells found in artificial
lakes, tanks, cattle-troughs, &c. As illustrating the
liability of shells to be carried with water, we have a
statement by Mr. Jeffreys that the zebra mussel
(Dreissena polymorpha) has been found in the most
frequented streets of London after they have been
flushed with water from the New River; and Canon
Norman, quoted by the same author, saw immense
numbers, in a living state, lining some iron water-pipes
which had been taken up in Oxford Street ;* Mr. Dyson,
in 1850, mentioned its occurrence also in most of the
large water-pipes supplying Manchester from the water-
works at Beswick, in the reservoir at which place it

1 See ‘‘ Nautilus,” iil. (1889), 67-9; for a mention of the occur-
rence, in 1881, of Margaritana margaritifera (believed to have
been absent in 1861), at the island of Anticosti—presenting “a
problem in the distribution of fresh-water shells which only the
methods of Darwin can surmount ”—see A. F. Gray, “ American
Naturalist,” xvii. (1883), 325-6.

2 See Tate, “ Land and Fresh-water Mollusks,” 1866, p. 188.

3 “ British Conchology,” i. (1862), 48; and see also Wood-
ward’s “ Manual,” ed. 4, rep. 1890, p. 424.

FRESH-WATER SHELLS. 19

abounded,’ and similarly, in Birmingham, according to
Mr. Sherriff Tye, it abounds in the town supply pipes,
derived, no doubt, from the reservoirs at Aston;
Mr. J. Macgillivray, it may be added, once observed
great numbers of Nerztena fluviatilis and a Limnea in
some large water-pipes which had been taken up near
St. Luke’s Hospital in the City Road, and other species
were seen there by Mr. E. Newman, who also visited
the spot Mr, Stariden tells me that the. lake in
Alexandra Park, Manchester (fully seventy yards from
the nearest pond), dug out of the peat and underlying
clay in the open fields, lined with concrete, and destitute
of introduced plants—filled through pipes with Man-
chester supply water—was found to contain, within two
or three years after its excavation, Sperza and Prszdza,
together with Planorbis spirorbis, Planorbts vortex,
Planorbis carinatus, Limnea peregra, and Limnea
palustris, all in good numbers, and the great upper lake
at Belle Vue Gardens, Gorton, he says, similarly formed
and filled with water, now contains Spherium corneum,
Pisidium amnicum, Bythinia tentaculata, Valvata pis-
conalts, Planorbrs albus, Planorbis carinatus, Limnea
peregra, and Limnea palustris. Physa gibbosa [genus
Bulinus| was taken abundantly, in 1887, in an iron tank,
“supplied with city water,” on the roof of a sugar
refinery in Sydney ;* the water probably came, as
Mr. Musson tells me, either from the Botany Swamps

1 Dyson, as quoted by R. Standen, “ Naturalist,” 1887, pp. 159-60.
2 J. Macgillivray, “‘ Zoologist,” x. (1852), 3420.
Ss steel, “Proce: Lin, Sec» N.S.W:,” (2), 11< (1888); To6.

C2

me

20 THE DISPERSAL OF SHELLS.

or the Nepean River, in or near both of which the shell
is not at alluncommon. At Burnley, it is said, Planor-
bis dilatatus has occurred in thousands, together with
nine other molluscs, Spherium corneum, Spherium
lacustre, Valvata piscinalts, Bythinia tentaculata, Limn@a
peregra, Planorbzs albus, Planorbis corneus, Planorbts
nitidus, and Planorbis glaber, ina cistern on the top of
an engine-house, “about sixty feet above the canal
from which the water is pumped.”' Mr. Musson, on
the authority of a member of the Nottingham Naturalists’
Society, states that a water-butt on Blue Bell Hill,
Nottingham, fed, until about 1882, from the corporation
reservoir at the top of Elm Avenue, contained no shells,
but when the source of supply was changed from
Nottingham to Pappiewick, Limncea truncatula soon
made its appearance in tolerable abundance, and
Limnca peregra, also, was discovered.

Shells are quite commonly found in horse and cattle-
troughs, and their occurrence in such places has been
noted by many writers. Mr. Roebuck, for instance,
mentions having found Lzmuca peregra in a new water-
trough on the summit of the Great Orme’s Head.’
Mr. W. A. Marsh has referred to. tiie presence se
Limnea parva in a horse-trough at his barn-well (in
Mercer County, Illinois), nowhere near any pond or
slough; how they happened to be found there, he
adds, “remains a mystery.”* Mr. Reid remarks upon

» F. C. Long, “‘ Science Gossip,” xxiv..(1688),,.2605.
2 W. D. Roebuck, “ Journ. of Conch.,” iv. (1884), 209.
3 W.A. Marsh, “ Conchologists Exchange,” li. (1888), 110.

FRESH-WATER SHELLS. |

the fact that Lémnea truncatula “has a curious tend-
ency to turn up in horse-troughs and stone basins ;”
in one instance, he found a number in a raised stone
cattle-trough on the marshes of the Humber, where the
dykes contained water much too salt for cattle to drink,
and a supply had to be provided by deep borings, one
of which overflowed from the tube into the trough in
question; thus, as he states, “a small colony of Z.
truncatula was found living in a cattle-trough in the
middle of a salt-marsh, where the surrounding dykes
were too salt for this snail to live in.’' In 1890, Mr.
C. R. Orcutt recorded the finding of thousands of living
Physe in a tank (with a capacity of four thousand
gallons) which had been erected, not quite a year pre-
viously, by a mining company at Hanlon’s Ferry, south
of Ft. Yuma, on the west bank of the Colorado River,
California. This tank, it is stated, was supplied “ from
a six inch well,” and “no shells were found alive in the
Colorado River only a few hundred feet away.” ?

It will doubtless be admitted, as already suggested,
that shells found in situations of the present kind
(reservoirs, artificially-filled lakes, tanks, troughs, &c.)
are in many cases carried in with the supply-water, but
it would frequently be found difficult, in individual
cases, to ascertain whether a possibility that such
is the case really exists. Transportal in this way,
to say the least, seems improbable in the three cases

* C. Reid, “ Trans. Norfolk and Norwich Nat. Soc.,” v. (1892),
279.
= C. R. Orcutt, “ Nautilus,” iv, (1890), 67-8.

22 THE DISPERSAL: OF SHELLS.

last mentioned, and certainly there is no such possibility
in a further case which Mr. C. P. Gloyne has obligingly
communicated. In Jamaica (where Mr. Gloyne resided
for some years) is a district in which—owing to the rain
percolating through the porous limestone and escaping
subterraneously—there is no natural standing or running
water; a supply is obtained, however, from artificial
cemented-tanks, which receive the rain-water from the
“ barbecues” (sloping planes, also cemented, for drying
coffee, &c.) and from the houses, and in all these tanks,
having, as I am assured, no connection whatever with
each other, one and the same species of Physa, of which
Mr. Gloyne forgets the name, makes its appearance!
Hardly less remarkable, perhaps, is the presence of
pond-snails in the basins of both the fountains in Tra-
falgar Square, erected in 1845, and supplied from two
Artesian wells, one behind the National Gallery, and
the other immediately in front of it.'| Within nine or
ten years after their construction, the basins were known
to contain shells, for, about the year 1854, Mr. E. L.
Layard exhibited specimens of them at a meeting of
the Zoological Society, and, notwithstanding the fact
that the basins are said to be frequently cleaned out,
I found Limnea peregra still living there, in 1891, in
good numbers. A very small fish and a water-beetle,
no doubt an Acz/ius, were also seen, but there were no
weeds. Artesian wells have been known to send up
seeds, small fish, and shells of several kinds, including,

1 See “ Old and New London,” (Cassell), iii. 142; “ Builder,”
lil, (1844), 370, and iii., (1845), III.

FRESH-WATER SHELLS. 23

in one instance, a specimen of Planorbis marginatus
[= P/. complanatus| presumably dead, and living fish
are said to have come up from a depth of 175 feet,’ but
I am assured that no such underground transportal to
Trafalgar Square is likely to have taken place. Mr.
Layard has suggested that the Lzmu@e@ were brought
to the basins from the ornamental water in one of the
parks, “attached to the feathers of the sparrows who
bathed, first in one and then in the other.” > When the
fountains cease to play, as I was told by a policeman
on duty in the square, many birds—both sparrows and
pigeons—bathe in the small upper basins from which
water falls into the large basins in which the Limnuge
are seen. It seems just possible, however, as Mr. Reid
has remarked to me, that “some misguided naturalist
with a mania for ‘acclimatization’ may have placed
some snails in the basins, or one of the boys who will
try to catch fish there may have brought his line
straight from the park with the small snails or eggs
attached.”

Aquatic molluscs, as is well known, commonly live
in marshes and swamps which are dry during a great
part of the year, and they occur also in other non-
permanent water, in that collected in hollows after
rains, in roadside puddles, &c. When the water is
derived from the overflowing of rivers the presence of
the animals is easily understood, for they are known to
be carried from place to place with flood-water ; their

1 See “‘ Lyell’s Principles,” ed. 12, i. (1875), 390-1.
2 E. L. Layard, “ Nature,” xxxviii, (1888), 296.

24 THE DISPERSAL OF SHELLS.

occurrence in waters derived exclusively from the rain-
fall of the immediate neighbourhood, however, has often
been regarded as surprising and mysterious. But we
must not fall into the error of concluding that the
creatures are necessarily introduced, after the appear-
ance of the water, by accidental means of dispersal, for
it is stated that they are able to remain alive ina torpid
state for long periods buried in the mud, and that on
the return of water to the surface they are ready to
burst into activity with remarkable rapidity. Their
almost immediate appearance after the occurrence of
rain, in tropical countries, in swamps and hollows which
have long remained dry is, I understand, quite familiar
to the naturalists of those regions. Adanson states that
a minute fresh-water shell (which he calls Lu/zmus) is
to be seen only from September to January in the
marshes of Senegal which are formed by the rains of
June, July, August, and September; when the marshes
dry up the animals disappear, a few empty shells alone
remaining upon the surface, but they never fail to return
with the rainy season.’ Major Becher, in a paper on
the mollusca of the Maltese Islands, mentions the find-
ing of Limnea truncatula in small hollows, the mud of
which, in summer, “must be baked till it becomes
almost brick.” Mr. W. A. Marsh records the finding
of Limnea humilis (said by some to be the same as
L. truncatula) in ponds on his land, in Illinois, which

“Hist. Nat. du Sénégal,” p. 7, as quoted by Dr. Johnston in
‘‘Loudon’s Mag. Nat. Hist.,” vii. (1834), 115.
2 E. F.. Becher, “ Journ. of Conehv, iv: (1884); 238.

FRESH-WATER SHELLS. 25

had been dry for three years at a stretch; when the
water reappeared in the fourth year the little mollusc
seemed as abundant as ever. Lzmnc@a parva, also, was
seen by the same naturalist in considerable numbers
about the margins of small basins which had been dry
for three or four years.' Mr. W. Jeffery, in a paper on
the mollusca of Western Sussex, remarks upon the
presence of Lizmne@a truncatula wherever a roadside
stream trickles down in spring, no matter at what
elevation, and notwithstanding the fact that the water
is almost sure to be dried up in summer, and he
mentions having seen specimens in a marl-pit far from
any permanent water.” Mr. H.C. Leslie has recorded
the finding of about half-a-dozen full-grown specimens
of Limnca peregra in a puddle, containing some two or
three gallons of water, formed by the breaking out of a
small spring by the roadside on one of the hills near
Erith,? and the same species has been noted by Mr.
Roebuck as common in roadside puddles between the
Orme’s Heads, and elsewhere, in Wales.“ Sir C. Lyell,
in one of his published letters, states that “ Lymnea
truncatella,’ introduced unintentionally by the Por-
tuguese into Madeira, went all over the island in thirty
years, and was said to have appeared even in pools and
ruts in the roads; if this be so, he adds, the creature
has powers of spreading “which require investiga-

1 W. A. Marsh, “ Conchologists’ Exchange,” ii. (1888), 110.
* W. Jeffery, “ Journ. of Conch.,” iii. (1882), 311-12.

sb. €. Leslie, “ Science Gossip,” 1870, p: 137.

4 W. D. Roebuck, “Journ. of Conch.,” iv. (1884), 209.

26 THE DISPERSAL OF SHELLS.

tion.”! Some Limnee, &c., we must remember in
connection with these facts, are not altogether incapable
of migration on land, for they are nearly amphibious ;
Limnea peregra, for instance, Mr. Jeffreys states,’ as
its name imports, is “fond of wandering and seeing a
little of the world,” and Lzmn@a truncatula, it is even
said, is “more frequently met with out of the water
thanan it,”

1 “ Life, Letters, and Journals of Sir C. Lyell, Bart.”, ii. (1881),

209, 212, in letters dated 1856.
2 “ British Conchology,” 1. (1862), 107, 116.

CHAPTER. If
MEANS OF DISPERSAL.

THE manner in which fresh-water shells are distributed
over the globe, the wide ranges enjoyed by many, and
their local diffusion in isolated waters, as we have seen,
clearly imply the existence of means of dispersal. The
creatures must certainly have been carrzed from pool to
pool, from river-basin to river-basin, and from mainland
to island, even, indeed, to some of the most isolated
archipelagoes of the open ocean.

Objections to suggested means have sometimes been
raised on the ground that the creatures would probably
be unable to withstand the exposure to which they
would be subjected, but, as is pretty generally known,
many kinds, at least, are able to live for a time out of
water, and under the most adverse conditions. Of a
number of pond-snails (Lzmnca truncatula), for instance,
placed by Professor A. P. Thomas in an open vessel, in
a dry laboratory, where the sunshine fell upon them for
an hour or so daily, rather more than fifty per cent.
survived for twenty-six days, and some few were alive
after more than six weeks.' A specimen of Paludina

1 A. P. Thomas, “ Quart. Journ. Micro. Sci.,” (n.s.), xxiii. (1883),
ust.

28 THE DISPERSAL OF SHELLS.

vivipara, an operculate water-snail, as Mr. R. M. Christy
states, accidentally ejected from an aquarium and allowed
to remain for more than three weeks on the ground in
a field, on being restored to its fellows seemed to
have taken no harm.’ Some Ampullaria, placed ina
drawer by Mr. Laidlay, are even said to have survived, in
the warm climate of Calcutta, for five years!” A fresh-
water mussel (A zodonta) once arrived alive in France
after having been wrapped up in dry paper for eight
months during its voyage from Cochin-China,* and
an Australian Uxzo, resembling our “swollen fresh-water
mussel” (Unzo0 tumidus),’ having already survived in a
dry drawer for 231 days, packed up (after being tested
in water) and forwarded to England, reached South-
ampton in a living state 498 days after its capture, and
was subsequently “ restored to its element, with full vital
powers,” in the care of Dr. Baird, of the British Museum.’
This great tenacity of life, common to many kinds, of
which quite a number of instances might be given,°
must certainly have largely facilitated dispersal.

1 R. M. Christy, “ Zoologist,” (3), v. (1881), 181.

2 Woodward’s “ Manual,” ed. 4, rep., 1890, p. 14.

3 “Journ. de Conch.,” xxiii. (1875), 81-4, as quoted in “ Quart.
Journ. of Conch.,” i. p. 78, and ‘ Zoological Record,” xii. (1875),
136.

4 Probably, according to Mr. Musson, U. amdiguus or U.
australis.

5 'J..S. Gaskoin, “Proc. Zool.- Soc.,” 1350) —pp;, 243:4;,and?see
also Woodward’s “ Manual,” ed. 4, rep. 1890, pp. 13-14.

6 See, for example, J. L. Hawkins, on Limn@a_ stagnalis,
“Science Gossip,” xvii. (1881), 23; Jeffreys,on Spherium lacustre,
“ British Conchology,” i. (1862), 11-12 ; Baker Hudson, on Sf/e-

MEANS OF DISPERSAL. 29

But we must inquire as to the means by which the
creatures have been diffused. The problem, everyone is
aware, is an old one, and has been answered or partially
answered by many writers ; Sir C. Lyell attended to it,
as also did Mr. Darwin, and many naturalists have
followed in the footsteps of these distinguished authors.

In early life, collecting in the fresh waters of Brazil,
Mr. Darwin noted with surprise the similarity of the
fresh-water insects, shells, &c., and the dissimilarity of
the surrounding terrestrial forms of life, as compared
with those of this country ; and questions as to the
diffusion of the former seem to have been more or less
prominently before his mind during many years; a
letter on means of dispersal of bivalves, one of his latest
published writings, appeared in “ Nature” only thirteen
days before his death. In the “ Origin” he expressed
the belief that the wide ranging power of fresh-water
productions could be expiained, in most cases, “ by their
having become fitted, in a manner highly useful to them,
for short and frequent migrations from pond to pond,
or from stream to stream, within their own countries,”
and liability to wide dispersal, he remarked, “ would
follow from this capacity as an almost necessary con-

sequence.”' The way in which such migrations are

rium corneum, “Science Gossip,” xx. (1884), 91; C. T. Simpson,
on Unio, &c., “ Conchologists’ Exchange,” ii. (1887), 50; E. B.
Sanger, on an Australian fresh-water mussel, ‘American Nat.,”
xvil. (1883), 1184-5; and J. E. Gray, on Drezssena polymorpha,
‘Annals of Philosophy,” ix. (1825), 139.

AOricm,” p: 343.

30 THE DISPERSAL OF SHELLS.

probably effected was discussed in some detail, and
diversified and curious means of dispersal were shown
to bein operation ; as regards molluscs, facts were given
suggesting transportal, possibly over the sea, by aquatic
birds and insects ;' and subsequently, in 1882, when
publishing several additional facts indicating dispersal
by animal agencies, Darwin demurred to the belief,
which had been somewhat doubtfully expressed by Mr.
Jeffreys, that the diffusion of fresh-water shells “had a
different and very remote origin, and that it took place
before the present distribution of land and water.” ?

Much diffusion, of course, as Darwin admitted, must
be attributed to changes in the level of the land, “ within
the recent period, causing rivers to flow into each other.” ®
Streams belonging to distinct systems, Mr. Wallace has
remarked, often approach and might be made to change
their course from one to the other basin by very slight
alterations of level. During the glacial epoch, ice,
blocking up valleys, compelled streams to flow over
watersheds to find an outlet.‘

The carrying powers of floods, as well as of rivers in
their ordinary action, are doubtless very great, and must
have largely influenced distribution. Professor Ralph
Tate, indeed, long ago expressed the opinion that the

" “ Origin,” pp. 344-5.

* Darwin, “ Nature,” xviii. (1878), 120-1; xxv. (1882), 520-30)
Jeffreys, ‘‘ British Conchology,” 1. (1862), Ixxx.

2 ““Oriein spas.

"Wallace, “Island. Life,” (1880), pp. 74 and 324; ed. 2, (pp: Jo
and 344.

MEANS OF DISPERSAL. 3!

diffusion of fresh-water shells had been cézefly effected
by “streams and land-floods;”' but the influence of
these agencies, it should be remembered, must be con-
fined, to a great extent, within the limits of river-basins ;
in some parts of the world, however, floods are known
to connect low watersheds. Unusual rushes of water
must certainly wash away large numbers of shells and
carry them down stream, possibly to great distances.
Every shell collector knows that ditches and pools
lying near to rivers, and liable to be affected by over-
flows, are generally stocked with more species than
those on high ground, far from the larger water-courses.
In Nottinghamshire, for instance, Mr. Musson has
emarked, the greatest number of species can be
obtained from those parts of the Trent Valley which
are subject to flooding, and, in fact, nearly every species
recorded for the county can be collected in the meadows
between Colwick and Beeston, while on the other hand,
the species found in ponds unconnected with the
general drainage system are comparatively few. Many
facts illustrating transportal by floods might have
been got together. A variety of Azcylus fluviatilis was
noted at one time as very abundant on stones in shallow
places in the River Went, but a strong flood, it is said,
“swept nearly all the stones and shells away.”* Mr.
L. E. Adams has described a flood in the Thames which
brought down large quantities of shells, comprising

‘ Tate’s “ Land and Fresh-water Mollusks,” 1866, p. 188.
? J. Wilcock, as quoted by Mr. George Roberts, “ Zoologist,” (3),
ix. (1885), 475.

32 THE DISPERSAL OF SHELLS.

twenty-three fresh-water and twenty-four !and species ;
the greater part were “dead and worthless,” but some
of the aquatic kinds, it appears, were still alive.’ Limnca
truncatula, Mr. Jeffery has remarked, “has a habit of
following the flow of water during floods,” and often
gets left high and dry.” During the summer of 1881,
Professor Thomas, studying the life history of the liver-
fluke, and anxious to try infective experiments with
this snail, searched repeatedly for it in the neighbour-
hood of Oxford, trying the localities in which he had
formerly found it, and those given in Whiteaves’ list,
but only empty shells could be discovered. In 1882,
however, there were floods in July, and the creature
came down in vast multitudes with the waters of the
Isis; so numerous was it that, several times, a single
sweep of a small hand-net gave more than 500
examples, and this in a ditch where the year before not
a single specimen could be found; all along the margins
of the ditches it occurred in the greatest profusion, and
it was found in numbers on the land after the flood-
waters had retired? Mr. W. A. Marsh, in his valuable
notes on the land and fresh-water shells of Mercer Co.,
Illinois, mentions that Lzmnc@a reflexa (common in the
small lakes of the county and in some of the larger
ponds in the Mississippi River bottom) is carried out
of the lakes in times of very high water, and may then

1 L. E. Adams, “Science Gossip,” xvii. (1881), 118.

2 W. jeffery, “ Zoologist,” (3), ii. (1878), 181.

3 A. P. Thomas, “ Quart. Journ. Micro. Sci.,” (n.s.), xxiil. (1883),
105-6, 130-1.

MEANS OF DISPERSAL 33

be found in considerable numbers along the margins of
the river, clinging to limbs of trees and pieces of bark ;
vast numbers of Pleurocera subularis (which abounds in
the small lakes of the Bay Island, &c.), he also states,
are to be found in July and August along the margins
of the river, clinging to drift-wood, having been carried
by a sudden rise in the waters from the different lakes
in the Bay.’ Cooper’s Creek, Central Australia, ac-
cording to Mr. E. B. Sanger, yields, amongst other
things, gastropods of three genera, “ Physa, Paludina,
and Zvryonza,’ which (he believes) die when the water
dries up; each flood, however, is found to stock the
creek again by bringing down young ones, which, it is
important to notice, are actually to be seen, “in all
stages of growth, in the flood-water.”* As showing
that Unzontde are carried over land by floods, Mr.
C. T. Simpson instances the finding of a Uuzo in great
numbers in low places and drains in the piney woods
of South Florida, at quite a distance from any stream,
where not a drop of water is to be seen outside, perhaps,
of three months of the rainy season, and where, during
the remaining nine months of the year, the animals
must have lain dormant in slightly damp sand; he had
dug them out in such places during the dry season by
the bushel.® Similarly, Mr. Cuming is said to have
collected living specimens of Cyclas maculata while

' 'W. A. Marsh, “ Conchologists’ Exchange,” ii. (1887-8), 81,
103
Sanger, “‘ American Nat.,” xvii. (1883), 1184-5.

E...B.
C. T. Simpson, “ Nautilus,” v. (1891), 16.

3

D

34 THE DISPERSAL OF SHELLS.

searching for land-shells in a garden in Old Panama,
where, it is stated, “they had been left after the rainy
season,’ and the occasional occurrence of living Pzszdza
on land, at some little distance from water, is familiar to
many observers, A little observation, recently recorded,
on the transportal of a shell by a mountain torrent is
perhaps worth giving. A house in which an English
naturalist stayed in Trinidad had a bath-room con-
structed so that a rapid stream (under control) flowed
in at one end and out at the other, and here, it is said,
the bather was joined sometimes by a specimen or two
of the “ram’s-horn water-snail,” which “would occa-
sionally come toppling down the rushing waters to a
deeper and quieter portion of the channel below.” ?
Shells adhering to detached and floating water-plants
seem eminently liable to transmission by rivers, and
when swept away during floods may be carried occa-
sionally to ponds or other waters at a considerable
distance from the river-bed. It is interesting to note that
a handful of dead pond-weed (Potamogeton), which was -
floating in a ditch and might have been carried to a
great distance in some circumstances, was once found
by Mr. W. K. Bridgman to contain “about a table-
spoonful” of little nautilus coil-shells (Planorbis tm-
bricatus)|!* Live shells, it is well known, are frequently

* Reeve, ‘‘ Land and Fresh-water Mollusks,” 1863, p. 236.

7-H. H. Higgins, “Address to Liverpool Naturalists’ Field Club,”
Jan. 30, 1891, p. 17.

* = Planorbis nautileus; W. K. Bridgman, “ Zoologist,” ix.
(1851), 3302.

MEANS OF DISPERSAL. 35

found upon the cases of caddis-worms: on some of those
fromthe pondsat Finchley, before mentioned, were several
living specimens of Spherium corneum ; indeed, with
Limnephilus flavicornis, which makes most of the shell-
cases sO common in our fresh waters, as Mr. McLachlan
has observed, it matters little whether the shells are
empty or still contain the living animal.' After the fly
has emerged, the empty cases, in all probability, are
sometimes floated to great distances, but the molluscs
may not always survive until set free by the rotting
of the binding materials. In March, 1892, I saw a
number of empty cases, bearing shells of Spherium,
Pisidium, Bythinia, Valvata, Planorbis, Limnea,
Auncylus, and Succcnea, floating amongst débris at the
margin of an overflow by the Lea, but I could not
ascertain that any of the shells contained living animals.
It is notorious how often shells and shell-bearing caddis-
cases are found adhering to the under-sides of floating
trunks and branches of trees, or hidden away in their cre-
vices,and in such positions the creatures are often likely to
be carried to considerable distances, sometimes, perhaps,
for hundreds of miles. Those molluscs which habitually
float, foot uppermost, on the under-surface of the water
are sometimes drifted along by steady currents. Mr.
George Roberts has remarked that Limncea peregra
“frequently allows itself to drift on the surface of run-

1 R. McLachlan, “Science Gossip” for 1868, pp. 152-3; and
see “Science Gossip” for 1866, pp. 95 and 110; George
Roberts, ‘Topography and Nat. Hist. of Lofthouse,” 1882
p. 238.

D2

36 THE DISPERSAL OF SHELLS.

4]

ning water,’ ' and in an old canal north of Inchbroom,
Mr. Martin’ is said to have seen hundreds of snails of
this species sailing from one end of the canal to
the other, resembling ‘‘a_ fleet of herring-boats in
miniature.” In a pondin Yorkshire, Mr. Christy ob-
served that this snail often crawled on the top of the
duckweed in mild weather and was liable to be blown
to one end of the pond by the breeze in large
numbers.* The free-roving larve of Dvrezssena, for a
time, live chiefly near the surface of the water, and, as
Dr. Korschelt observes, are likely to be carried forward
for long distances by currents.’ Univalves frequently
attach their eggs to loose leaves and sticks in the
water, and these, as Sir C. Lyell states, “are liable
to be swept away, during floods, from tributaries to the
main streams, and from thence to all parts of the same
basin ;” a species, it is added, might thus migrate during
one season from the head-waters of the Mississippi, or
any other great river, to countries bordering the sea at
the distance of many thousands of miles. Figures given
by Lyell show the ova of an Ampullaria fixed to a
small sprig which had fallen into the water, those of a
Planorbis attached toa dead leaf, and those of a Lzmne@a
adhering to a dead stick.* Ina poolin Epping Forest
I recently saw “egg-jellies” of Lzmn@a auricularia

1 “ Topography and Nat. Hist. of Lofthouse,” 1882, p. 242.

2 Quoted by the Rev. G. Gordon, “‘ Zoologist,” xii. (1854), 4457.
3 R. M. Christy, “ Zoologist,” (3), v. (1881), 184.

4 E. Korschelt, “ Ann. and Mag. Nat. Hist ,” (6), ix. (1892), 166.
&™ Principles,” i. p, 300; i. maa.

MEANS OF DISPERSAL. 37

adhering to every available object ; a detached bracken-
frond lying in the water could only be described as
literally covered ; I tried to count the number of masses
upon it, but soon gave up the task; there were hun-
dreds, and possibly each contained from fifty to sixty
eggs.’ An object of this kind, one can readily imagine,
if near a stream which occasionally overflowed, might
easily be carried away with the flood-water, and, if it
happened to reach a pond or other water where the
conditions were favourable, the introduced species could
hardly fail to establish itself in the new home.

Mr. C. T. Simpson has even expressed the opinion
that in some parts of the world trees, &c., drifted down
by rivers and floods, may occasionally float fresh-water
molluscs in safety over the sea. In his own words ;

“Great numbers of trees are washed out and
lodged along the rivers and torrents of tropical
countries ; and in the crevices of the bark of these
trees many species of fresh-water shells find a home.
Others live among the roots of living trees which
are washed by the water of streams. In Florida I
have collected handfuls of Unzo fuscatus and other
species among the matted roots of trees just under the
surface of the water. Such trees, washed out and
carried down stream, would take some of their molluscan
inhabitants with them. Others, carried in floods, with
broken jagged limbs and loosened bark, would plow up
quantities of mud and shells from the bottom and carry

1 See “ Rimmer,” p. 62.

38 THE ‘DISPERSAL OF: SHELES. .

them out to sea. Some such trees might drift into bays
at the mouths of rivers in other countries, and I have
seen just such floating in the mouths of the Manatee
and Caloosahatchee Rivers in Florida. It must be
remembered that all these rivers, during the rainy
season, are only very slightly brackish, or even entirely
fresh, throughout their estuary portions.

“Trees carried into such places, and bearing fresh-
water shells, might be driven up and landed by tide and
winds, and a colony of living inhabitants established.
Many of the shallow bays along the coast of Florida
become perfectly fresh during the rainy season, as some
five feet of water falls there in three months, and the
same thing no doubt occurs in other parts of the tropics
where the rain-fall is much greater. In Florida these
bays at such times connect with ponds of water on the
flat lands, so that often for miles the whole country
from the shore far inland is nearly covered with fresh
water.

“ At the south-western extremity of the State are found
the Ten Thousand Islands, an innumerable group
scattered over a space of a hundred and fifty miles of
coast separated by brackish channels through which the
tide flows in and out, gradually becoming entirely fresh
in the region of the Everglades. So there would be no
trouble about landing a colony of fresh-water snails on
the Florida coast, and the only difficulty would be in
bringing them across. Could they stand the drying
and the salt water of the ocean ?”

[After showing that many kinds are able to live fora

MEANS OF DISPERSAL. 39

long time out of water, the author proceeds to remark,
by way of answering the latter half of his question, that]
“many fresh-water species will live in brackish water,
as is well known. WNerztina reclivata, found in Florida,
seems indifferent as to whether it lives in fresh water or
that which is more or less salty, and WV. dineolata and
macrostoma, which I found in Honduras in the mouths
of rivers, often extended into the sea. Planorbis
tumtidus was often found in slightly brackish water in
Florida, and the Lzmn@as in the Baltic and some
places on the British coasts mingle with the Lz¢torznas.

“Ampullaria caliginosa, a Mexican species, closely
related to A. depressa, if not identical with it, Planorbis
tumidus and havanensis (identified by Mr. H. A. Pilsbry,
who has made a special study of the genus Planordzs),
and a few other tropical species found to-day in the
Southern States, may have been introduced, I think,
in the manner of which I have spoken. Of course such
voyages of living snails, with a successful termination,
could only happen rarely, but it must be remembered
that countless ages have elapsed since the present
species have occupied the earth.’’’

That both adult shells and their ova are frequently
carried out to sea with drift timber, brushwood, &c.,
cannot be doubted for a moment, but it is not actually
known, I believe, that any mollusc from a truly fresh-
water habitat can endure much exposure to the
waters of the ocean. More direct experiments on the

' C. T. Simpson, “ Conchologists’ Exchange,” ii, (1887), 50-1

40 THE .DISPERSAL OF SHELLS.

point are probably required, but I find it stated both by
Darwin and Wallace that salt water is immediately
or almost immediately fatal both to ova and adults.’
It seems improbable, therefore, that the creatures are
transported for any considerable distance over the
open sea in the way indicated. Molluscs belonging to
fresh-water genera or species, but inhabiting water which
is salt or brackish, might almost certainly be thus
carried, and in a new home in the course of ages some
of their descendants might re-adapt themselves to fresh
water. But we cannot argue that, because certain
members of a genus live in salt or brackish water, other
members from fresh-water habitats are likely to with-
stand a voyage over the sea; nor in the case of fresh-
water species having individuals living in the sea or in
brackish estuaries, in certain spots, can it be supposed
that examples from fresh waters could pass through
the sea unharmed. Some of the fresh-water species of
Neritina,as Dr. H. B. Guppy remarks in his “ Solomon
Islands,” have been widely dispersed, Veritzna subsul-
cata and JV. cornea occurring both in the Solomon
Islands and in the Philippines, VV. macgidlivrayz and
N. petite inthe Fijiand Solomon Islands, and WV. porcata
in Samoa and Fiji, as well as in the opposite extremi-
ties of the Solomon group; yet it is doubtful whether
the animals have been drifted far over the sea, for
although possessing a stony and close-fitting operculum
they are probably unable to resist the action of salt

1 “Origin,” p. 344; Wallace, ‘“ Geographical Distribution,” i.
p: 31:

MEANS OF DISPERSAL. 4I

water for any length of time; one individual of 1.
subsulcata survived submersion for twelve hours, but
of a dozen kept in water, changed from time to
time, for five days, not one survived. It has been
pointed out, however, that these animals have calcare-
ous egg-capsules which are probably thick enough to
resist salt water, and these, as Mr. Edgar Smith has
remarked, if attached to floating timber, might be
carried to considerable distances.’

Something, no doubt, may be attributed to the
agency of floating ice, for various kinds of shells have
been known to revive after having been frozen up.
Dr. Binney states that a Succinea “has been frozen in
a solid block of ice and yet escaped unharmed.’?
Anodonta cygnea and Paludina vivipara, also, M. Joly
has observed, may be kept frozen up for some time
without being killed,’ and the latter has even produced
young after being thawed. In some thick ice which
Mr. W. A. Gain once removed from a stone trough or
“out-door aquarium,” and allowed to melt slowly, were

1 H.B. Guppy, “ Solomon Islands,” 1887, pp. 338-9 ; E. A. Smith,
“ Proc. Zool. Soc.,” 1885, p. 588.

7 A. Binney, “ Terrestrial Air-breathing Mollusks,” i. (1851),
196.

* On the other hand, however, it is stated that two specimens of
A. cygnea, exposed in the winter of 1890-1, in an open vessel, to
the entire severity of the frost, were killed, their shells splitting
from dorsal to ventral surface on one side (see “ Nature,” xliii,
(1891), 464),and I have found several mentions of the fatal effects
of severe frosts upon bivalves in a state of nature.

* “Comptes Rendus,” xvi. (1843), 460, as quoted in ‘ British
Conchology,” i, (1862), xliv.

42 THE DISPERSAL’ OF SHELLS.

many young examples of Limunea peregra and L, stag-
nalis, ready to crawl away as soon as they dropped
from the blocks, and Planorbis corneus and P. com-
planatus also occurred in quantity, mostly young, and
all living. The water had been frozen for six weeks,
and it is probable that most of the animals had
been at least a month in solid ice. A few specimens
of Pisedtum pustllum, also frozen up, had died." We
have here a means by which the animals may almost
certainly cross arms of the sea, for when enclosed in
ice they would be effectually protected from the in-
jurious effects of salt water. Dr. Guppy has suggested,
also, that molluscs burying themselves in winter in the
mud at the bottoms of rivers, &c., may possibly be
transported, even across the sea, in the frozen mud
buoyed up by ice. In the shallow waters of the Lea,
he has noticed that after a frost of some duration, when
the water was frozen to the bottom, the mud beneath
for an inch or two was also frozen, so that on lifting up
a slab of the ice, a layer of frozen mud, an inch or two
thick, formed the lower part of the mass. But it seems
hardly likely that the creatures will often be landed by
these means on foreign shores in localities suitable for
the establishment of new colonies. Quite possibly,
however, on the breaking up of frosts, shells thus
enclosed in ice or frozen mud may be safely carried,
within their own river-basins, to great distances.
Powerful whirlwinds, according to Sir C. Lyell, some-

1 W. A. Gain, “ Science Gossip,” xxvii. (1891), 118.

MEANS OF DISPERSAL. 43

times dry up lakes and ponds,’ and fishes still alive, as
Darwin states, are not very rarely dropped by them at
distant points, so that we are justified, perhaps, in
assuming that the distribution of shells has been
influenced by such means. There are many authentic
accounts, it is said, of the falling of fish from the
atmosphere ;* and frogs and other creatures, no doubt
taken up by whirlwinds or hurricanes, have been seen
to fall in “showers.” On the oth of February, 1859, for
instance, a shower of small living fish appears to have
fallen in the valley of Aberdare. A sawyer at work in
Messrs. Nixon and Co.’s yard (whose statements were
taken down on the spot by the Vicar of the parish),
while getting out a piece of timber for the saw, was
startled by something (which he found to be little fish)
falling all over him; and a long strip of ground, it is
said, soon became covered with the creatures, “ jumping
all about ;””? many were seen, also, on the top of a large
shed, etc. Numbers were gathered and thrown intoa
rain-pool, where some were to be seen when the evi-
dence was taken. The wind at the time of the shower
was not very strong, but it was “uncommon wet,” and
the fish came down with the rain “in a body like.”
Mr. R. Drane, of Cardiff, who investigated the case, is
said to have obtained convincing evidence, from other
sources, that a large number of fish actually descended
with the rain, and over a considerable tract of country ;
specimens procured from three persons resident some
Sask rineiples,” il.,p-.,302- 2) Origin,” 2.344.
3 Wallace, ‘ Geographical Distribution,” i. p. 29.

44 TIIE DISPERSAL OF SHELLS.

distance from each other were of two species, the com-
mon minnow and the three-spined stickleback.’ To
give another instance, the late Thomas Cooper (well
known to many through his delightful autobiography)
witnessed, when a boy—with others—a shower of frogs,
which, “jumping alive, fell on the pavement at our feet,
and came tumbling down the spouts from the tiles of

the houses into the water-tubs;’’ he was as sure of

what he had seen as of his own existence, and recorded
the fact because it had been stated in books that such
a sight was impossible.” Water-beetles, also, are said
to have fallen in showers,’ but I am not aware that
aquatic molluscs are actually known to have done so,*

1 J. Griffith, “ Zoologist,” xvii. (1859), 6493; R. Drane, p. 6564.

2 “Life of Thomas Cooper,” pp. 20-1, as quoted in “ Science
Gossip” for 1872, p. 167; and see on this subject Gosse’s
‘Romance of Natural History,” second series, 1861.

3 See “American Entomologist, (2), i. (1880), 248; “ American
Nat.,” xvi. (1882), 600.

4 A case has been recorded since the above was written. See
“ Nature,” xlvii. (1893), 278: “Das Wetter of December last con-
tains an account of a heavy thunderstorm which occurred at
Paderborn on August 9, 1892,in which a number of living pond
mussels were mixed with the rain. The observer, who is in con-
nection with the Berlin Meteorological Office, sent a detailed
account of the strange occurrence, and a specimen was forwarded
to the Museum at Berlin, which stated that it was the Anodonta
anatina (L.). A yellowish cloud attracted the attention of several
people, both from its colour and the rapidity of its motion, when
suddenly it burst, a torrential rain fell with a rattling sound, and
immediately afterwards the pavement was found to be covered
with hundreds of the mussels. Further details will be published
in the reports of the Berlin Office, but the only possible explana-
tion seems to be that the water of a river in the neighbourhood

MEANS OF DISPERSAL. A5

Animals of various kinds, it can hardly be doubted,
are actively engaged in dispersing shells. The “short
and frequent migrations from pond to pond or from
stream to stream,” to which Mr. Darwin has alluded,
have, almost certainly, been largely brought about by
birds and other creatures, and transportal by such
agency over considerable tracts of ocean, it has been
shown, is at least possible. Many birds, fish, amphibia,
etc., swallow large numbers of shells, which are sure, in
some cases, to survive for a time in their stomachs ;
thus, for instance, Mr. Baker Hudson has often found
living Pzszdia, with their valves tightly closed, in the
stomachs of frogs, and this, of course, suggests a possi-
bility that sudden deaths of shell-eating creatures,
brought about by birds and beasts of prey, may have
led, at rare intervals, to the setting free of living shells
in spots considerably removed from their original
homes ; and some birds, it is important to notice, on
being frightened or wounded, occasionally or habitually
cast up the contents of their crops. That shells are
voided in a living state with the excrement seems less
likely, for they are almost sure to be killed during their
passage through the digestive organs. No less a person
than Dr. Kobelt, however, has ventured to express the
opinion that small mussels and even sometimes opercu-
late water-snails, when swallowed alive by birds, may
possibly pass the intestines unhurt.’ Experiments on

was drawn up by a passing tornado, and afterwards deposited
its living burden at the place in question.”
W. Kobelt, “‘ Fauna der Nassauischen Mollusken,” 1871, p. 14.

46 THE DISPERSAL OF SHELLS.

the point appear to be required, but I can hardly think
that anything other than negative results will be
obtained.! Four warty-newts, placed in a vessel of
water by Mr. C. Robson, extruded a number of Pzszdiza,
the shells of most of which were open, with the animal
extracted ; four, however, were closed, and three of
these contained young; the adult animals had been
extracted even from these closed shells, but, in
some of the young, the animals were still within the
shells, yet it does not appear that they were actually
observed to be alive. As far as diffusion by animals is
concerned, it seems clear, I think, that the creatures are
chiefly carried while very young, adhering to the feet
and feathers of aquatic birds, or attached to plants thus
adhering. Adult and partly grown shells, however, are
liable to occasional transportal by creatures of various
kinds, and, through the agency of birds, adult bivalves,
even some of the larger kinds, may sometimes be carried
through considerable spaces; but before giving the
facts on this head we will consider the possible dispersal
of fry and ova.

Mr. Darwin has a statement at page 344 of the
“ Origin” that the eggs are not likely to be transported
by birds; but Mr. Wallace, Professor Tate, Lieut.-Col.
Godwin-Austen, and other writers have referred to

| The eggs of Limn@a auricularia, it has recently been stated,
have passed unharmed through the digestive system of swans.
See “ Zoological Record,” xxviii. (1891), Moll. p. 37, referring to
Pascal, ‘‘ Journ. de Conch.,” xxxi. pp. 9-15.

2 C. Robson, “ Science Gossip” for 1875, p. 220.

MEANS OF DISPERSAL. Ay.

such transportal as probable,’ and Canon Tristram,
in the winter of 1856-7, it is interesting to find, had the
good fortune to discover the eggs of some mollusc—
probably Swuccenea—attached by their glutinous coating
to one of the feet of a passing mallard shot by him in
the Sahara, a hundred miles from water ;* thus, he
remarks, such a bird “might easily carry a Succenea or
Physa from Europe to the lakes of Central Africa.”
It ought to be remembered, however, that eggs of some
kinds, thus exposed to the atmosphere until dry, would
become very firmly attached and so remain even when
again moistened,* so that they would not be easily dis-
lodged in anew locality ; but, as many birds probably
travel during gales at the rate of thirty-five miles an hour
or much faster,* eggs might certainly be carried before
drying to considerable distances, and then when again
dipped in water would soon become detached. Many
kinds attach their eggs to aquatic plants, fragments of
which, we shall see, are likely to be carried by birds.
Something may possibly have been done by insects, for
Mr. Standen informs me that he once saw, in the
Hollinwood canal, an egg-capsule of the river-limpet
(Ancylus fluviatilis) attached to one of the wing-cases of
an Aclzus, a strong flying water-beetle. Some eggs, it

i *¥sland Life,” 1380, p. 76; ed. 2,p.79; R. Tate, “Land and
Fresh-water Mollusks,’ 1866, p. 188; H. H. Godwin-Austen ,
“ Field,” Ixvi. (1885), 499.

2 See “ Zoologist,” (3), 1. (1877), 260—1.

8 This happened, at least, to eggs of Lzmuca auricularia, which

I exposed on a tin tray and on fragments of water-weeds.
+ *Onirin, p. 320:

48 THE DISPERSAL OF SHELLS.

should be noted, from the localities in which they are
deposited or the nature of the objects to which they are
usually attached, are not at all likely to be dispersed by
animals, and it will be remembered that some molluscs
are viviparous. The fry, probably, are transported more
frequently than ova. Those Unzonitde whose larve are
parasitic for a time upon fishes are eminently liable to
wide dispersal, at least, through waters in any way com-
municating with their habitats. The larve of Anodonta
swim by the flapping of the valves of their shells, trailing
at the same time a long byssal filament, and weaving
themselves together in masses which spread out like a net.
Little fishes, stirring up the masses, carry many larve
away, the creatures managing to effect a hold—chiefly to
the fins—by means of the toothed edges of their shells.
They then become encysted, and are nourished, it is
stated, by the exudations of the fish. Ultimately, on
the splitting of the cyst, they drop out and fall to the
bottom. In aquaria, Axodonte which have retained
their brood in long quiescence, have been observed to
let it out immediately on the introduction of fishes,
The larve of Unzo, differing in certain respects, are said
to be inhaled by fishes, and are usually found attached
to the gills.’ Whether these fry are likely to be carried

1 E, Ray Lankester, art. Mollusca, “Encyclopedia Britannica,”
ed. 9, xvi. (1883), 694 ; and see also Dr. Schierholtz’s memoir as to
the development of the Unzonide, Imperial Academy of Science,
Vienna, 1888, my knowledge of which is derived solely from a
leaflet printed by Mr. Standen, in explanation of specimens illus-

trating the life history of Azedonia, shown by him at one of the
Manchester Microscopical Society’s soirees.

MEANS OF DISPERSAL. 49

from one piece of water to another by birds, etc., I am
unable to say, for I have not seen it stated that they
are able to withstand exposure to the atmosphere. The
same remark applies to the free-roving pelagic larve of
Dreissena. The just-hatched young of certain univalves,
we know, are well suited for such transportal, for they
cling firmly, and can live out of water for hours. On
this head we have Mr. Darwin’s celebrated and often
quoted experiment with a duck’s feet in an aquarium :

“T suspended the feet of a duck in an aquarium,
where many ova of fresh-water shells ' were hatching ;
and I found that numbers of the extremely minute and
just-hatched shells crawled on the feet, and clung to
them so firmly that when taken out of the water they
could not be jarred off, though at a somewhat more
advanced age they would voluntarily drop off. These
just-hatched molluscs, though aquatic in their nature,
survived on the duck’s feet, in damp air, from twelve to
twenty hours; and in this length of time a duck or
heron might fly at least six or seven hundred miles
and if blown across the sea to an oceanic island, or to
any other distant point, would be sure to alight on a
pool or rivulet.’”

Unfortunately, however, I cannot find, after inquiring
of a number of experienced ornithologists and sports-
men, that the creatures have ever been seen adhering to
birds shot on the wing; Mr. Dresser, for instance,
though he has shot thousands of aquatic birds in

’ Univalves, as Mr. F. Darwin informs me.
2 Orin.” p.. 345.

50 THE DISPERSAL OF SHELLS.

various countries, has never seen shells attached, and
Mr. O. V. Aplin, the Rev. J. C. Atkinson, Mr. Abel
Chapman, Mr. Cordeaux, the Rev. H. A. Macpherson,
Mr. T. H. Nelson, and Mr. Howard Saunders have
also replied in the negative to my inquiries on the
point. But from general considerations, and especially
in view of Mr. Darwin’s experiment, it can hardly be
doubted but that very young molluscs do thus adhere
somewhat frequently, and perhaps the absence of obser-
vations made in the field is not surprising, for, as Mr.
Layard remarks to me, the creatures are hardly likely
to be noticed unless specially looked for, and this I
suppose is very rarely done; indeed Mr. Aplin, replying
that he had never seen young molluscs on the feet of
water-birds, carefully added that he had never looked
for them. Mr. F. Norgate, it is interesting to find,
once noticed a small grey leech clinging to the plumage
of a mallard shot by him. Mr. Macpherson remarks that
widgeon, etc., sitting all day long half awake in the wet
sedge, are very likely to carry shells on their feet or
feathers when they fly at night to their feeding-grounds,
and as Dr. Kobelt' has suggested, the creatures will
have a good chance of adhering also to the feet and
legs of herons and other birds which often stand motion-
less in water for hours. Even adult shells of the smaller
kinds may often attach themselves, but these, I suppose,
in most cases, are soon shaken off. Mammalia fre-
quenting fresh water, otters for instance, seem likely to

* W. Kobelt, “Fauna der Nassauischen Mollusken,” 1871,
p. 14.

MEANS OF DISPERSAL, . a

transport molluscs adhering to their coats, and many
other creatures which occasionally journey over land
have probably helped in the work; but their influence,
that of mammalia at least, is doubtless chiefly local,
and must always be confined to a given tract of land.
Birds, it will be remembered, commonly fly across arms
of the sea and are not unfrequently blown by gales over
wide stretches of ocean; flying water-insects, also,
which may sometimes carry fry as well as ova, are
occasionally blown to great distances.

It seems probable that young and small shells
may sometimes be transported along with the small
water-plants which occasionally adhere to birds. Mr.
Darwin, in removing a little duckweed from one
aquarium to another, unintentionally stocked the one
with fresh-water shells from the other, and he twice saw
duckweed adhering to the backs of ducks which had
suddenly emerged from a pond covered with these little
plants.’ It would be interesting to ascertain whether
small shells, and the fry of larger kinds, are often found
amongst such plants. A small tin cartridge-box full of
gibbous duckweed (Lemna gibba) skimmed from the
surface of one of the streams by the Lea in Sep-
tember, contained a number of little aquatic animals,
shrimps, leeches, a beetle, etc., but no shells. A similar
quantity of lesser duckweed (Lemna minor), however,
carefully picked (in August) from the surface of a pond
near Louth, yielded no less than forty-two shells,
apparently alive: four of Spherium corneum and thirty-

’ “Origin,” pp. 344-5,
E 2

52 THE DISPERSAL OF SHELLS.

one of Limne@a peregra, all young, together with seven
full-grown specimens of the little nautilus coil-shell
(Planorbis nautileus). Mr. Cordeaux once noticed
minute molluscs among duckweed within the bill of a
wild duck shot by him, the bird having risen, no doubt,
in the very act of feeding. Confervoid vegetation, also,
seems likely to cling to birds, and Mr. Bridgman men-
tions having found masses of it literally teeming with
Planorbis nitidus, in company witha species of Valvata ;
after two or three weeks, however, the creatures, having
probably come to maturity, disappeared.! It must not
be forgotten that parts of larger water-plants, to which
fry or ova may possibly be adhering, may occasionally
be carried by birds. The moor-hen, for instance, might
easily carry parts of plants in which its feet had be-
come entangled, and Mr. Norgate suggests that herons,
etc., are likely to carry them when flying off with
struggling eels hastily snatched from the mud and
weeds. Mr. Standen mentions the shooting of suddenly-
flushed snipe, moor-hens, and wild ducks with weed
clinging to their feet, and he once flushed a heron which
rose with so large a quantity dangling from one of
its legs that its flight was considerably impeded.
Transportal of fragments of water-plants in this manner
probably happens quite commonly, for many kinds, as
Mr. Clement Reid* has recently remarked, have ex-
tremely brittle stems, and finely divided or thin leaves
which on removal from the water collapse and cling

1 W. K. Bridgman, “ Zoologist,” ix. (1851), p. 3303.
2 “Trans. Norfolk and Norwich Nat. Soc.,” v. (1892), pp. 278,'283-4

MEANS OF DISPERSAL. 53

closely to any object they may touch. The plants
found in ponds of extreme isolation, according to this
observer, are not usually species with succulent fruits,
such as are known to be habitually eaten by birds,
and species with burrs or hooked fruits are by no means
abundant; most of them, in fact, have the fruits dry-
seeded and thin shelled, which, if eaten at all, would
probably be digested and have their vitality destroyed ;
even water-thyme (E/odea canadensis), which is fruitless
in this country, sometimes occurs in isolated dew-ponds.
It seems probable, therefore, that most of these plants

e “transported in fragments, which breaking, cling to
the feet of waders, to be washed off when the bird flies
to the next pond,” and this, it is added, “will also
account for the constant occurrence of the Limnezids,
which both adhere to the stems and attach their eggs
to them.” Large numbers of young or small molluscs
might be transported together in this way. On one
occasion, on examining j a small quantity of horned pond-
weed (Zannichellia), which had been taken home (after
having been well shaken to get rid of snails) from an
isolated pond on the South Downs, Mr. Reid found,
adhering to it, fully 150 specimens of Lzmn@a and
Planorbis, mostly very minute, and several clusters of
eggs! Some molluscs, it is true, live much in or on the
mud, and do not attach their eggs to plants, or are
viviparous, but most of these, I believe, at some period
of their existence, or during certain parts of the year,
are to be met with among the leaves and stems of the
water-weeds.

54 THE DISPERSAL OF SHELLS.

The beaks and feet of birds sometimes have earth
adhering to them,’ and this in the case of those
frequenting the muddy edges of ponds and streams
may sometimes have young molluscs, and even full-
grown Pzszdia and other small shells sticking to or em-
bedded in it. A top-knot pigeon (Lopholaimus antarc-
tica\, shot by Mr. Musson in New South Wales, had a
small ball of earth surrounding each leg, just clear of
the ground, “no doubt accumulated as the bird
wandered about on the muddy margin of some water-
hole,’ ? and Mr. Darwin once received a woodcock’s leg
from a friend with a little cake of earth, weighing nine
grains, attached to the shank.* Mr. Hardy and Mr.
Standen both call to mind instances of snipe shot
with clayey mud clinging to their feet and legs, in
sufficient quantity, they think, for the retention of
Pisidia, etc., and no doubt many observations of this
kind might be got together. As is well known, seeds
of plants have been found in the earth thus adhering,
but I have not heard that shells have been dis-
covered in it. It is worth mentioning, perhaps, that
Mr. Darwin once found a pebble, as large as the seed
of a vetch, and, therefore, as large as some full-grown
Pistdia, in a little earth removed from the foot of a
partridge.’

P - Origin,” ps 345:

7 C. T. Musson, “ Proc. Lin. Soc. N.S.W.,” (2), iv. (1889), 388 ; one
of the legs of the pigeon, and the accompanying ball of earth,
weighing nine grains, were exhibited at the Society’s meeting.

=“ Origin,” p. 328.

4 %Origin,” p. 328.

MEANS OF DISPERSAL. Be

The probable dispersal of bivalves while holding on
by closure of their shells to the toes, etc., of birds,
amphibia, aquatic insects, etc., remains to be dealt with
in the next chapter, and it will be seen that animals
have been secured, in a few cases, in the very act of
carrying the creatures over land. Dealing with
univalves in chapter iv., I shall have to suggest a
somewhat analogous mode of dispersal of operculata,
namely, by closure of the operculum so as to hold on
to insects, etc. ; and a few cases of the clinging of inoper-
culate univalves, by adhesion, to amphibia, insects, and
the like, will be added, but these latter might almost
have been included above, for young shells will have
been oftener thus carried than full-grown ones,

CHAPTER TL.
TRANSPLANTATION OF BIVALVES.

BIVALVES frequently lie with their shells slightly apart,
and, as is well known, quickly close upon objects which
happen to be introduced between the valves. Birds,
wading about at the muddy and sandy margins of
ponds or rivers, and aquatic or amphibious animals of
various kinds sometimes accidentally insert their toes,
and the mollusc, in such a case, closing quickly and
often holding on for a considerable time, is liable, if not
too heavy, to be carried away and to be set down, per-
haps, in anew home, possibly at a great distance from its
original habitat. Accidents of this kind, there is reason
to suppose, happen much more frequently than might
at first seem probable, for numerous instances have
been recorded, and probably not more than one in
a thousand comes under the notice of an observer, and
the number recorded must be small as compared with
the number actually observed.

A few facts serving in a general way to illustrate the
liability of bivalves to be carried away upon objects
chancing to come between the valves are perhaps
worth giving. Rural folk who make cream-skimmers

TRANSPLANTATION OF BIVALVES. 57

of the valves of the great pond-mussels (Anodonta
cygnea) procure them, according to Mr. Jeffreys, by
means of a long pointed stick, which is inserted between
the gaping shells. The animal closes upon the stick
and allows itself to be drawn up out of the water.’ Pearl-
mussels (Unzo margaritifer), as Professor Tate relates,
are dragged to shore by country boys in a similar man-
ner upon long slender rods.’ I recently experimented
by the Lea upon a number of Anodonte and Uniones,
most of which, I found, allowed themselves to be drawn
from the mud and out of the water upon inserted grass-
stems. A few fell almost immediately, but, of six which
were carried away suspended upon the grasses, four
(two of each genus) were still holding on when I
reached home after the space of an hour and a half, in-
cluding about ten minutes in a train; these were then
suspended from a shelf, and one Anodonéa (two anda half
inches long) still retained its hold fifty-one hours after
ithad been taken from its habitat, and on being placed
in water it extended its foot and ultimately became
detached. The Rev. E. A. Woodruffe-Peacock tells me
that he used to catch hundreds of mussels in his father’s
fish-pond in this way, drawing them to land upon stiff
straws, twigs, or fine wire, and the fact that the
creatures will allow themselves to be thus taken seems
to have been long known, for Sir Robert Redding, in a
letter dated in 1688, mentioned that the poor people in

1 “British Conchology,” i. (1862), Ixvili.
? Tate, “ Land and Fresh-water Mollusks,” 1866, p. 27, and see
also ‘Science Gossip” for 1870, pp. 265-6.

58 THE DISPERSAL OF SHELLS.

the North of Ireland fished for pearl-mussels some with
their toes, some with wooden tongs; and “some by
putting a sharpened stick into the opening of the shell
take them up.”' The Rev. J. W. Horsley, while trolling
with a dead fish for pike, once brought up a large
Unzo which had closed upon the bait. A friend of Mr.
W. D. Crick’s, as the latter told Mr. Darwin, often, while
fishing in rapid streams, caught small Unzones upon
the hook,and Mr. F. Darwin, when fishing off the shores
of North Wales, several times caught mussels in a
similar way.” According to Mr. D. Pidgeon, heart-
cockles (/socardia cor) have been known to close upon
the shanks of accidentally intrusive fish-hooks with such
force as to crush the edges of their shells against the
steel wire, and they permit themselves to be drawn in
with the line to which the hook is attached, many
having been thus taken by the long-line fishermen on
the Irish coast.* Marine bivalves, such as cockles,
mussels, etc., have several times been found clinging to
the toes or bills of birds of various kinds,‘ and the

1“ Philosophical Transactions,” xvii. (1693), 660.

2 Darwin, “ Nature,” xxv. (1882), 529-30.

3 D. Pidgeon, “ Nature,” xxv. (1882), 584.

* Instances of the capture by marine bivalves of fish, mice, a
rat, foxes, etc., have also been recorded: see as to fish, ‘‘ Popular
Science Monthly,” xvii. (1880), 111; oyster and mouse, “ Science
Gossip,” 1875, p.68 ; oyster and mouse, “ Daily Telegraph,” quoted
in the “ Field,” Ixvi. (1885), 499 ; oysters and mice, several cases,
“ Bell’s Weekly Messenger,” etc., quoted in Loudon’s “ Mag. Nat.
Hist.,” ii. (1829), 150; oyster and young rat, caught by the tail,
“ Life Lore,” ii. (1890), 216; mussel and fox, caught by the tongue,
Loudon’s “ Mag. Nat. Hist.,” viii. (1835), 227-8; oyster and fox,

TRANSPLANTATION OF BIVALVES. 59

observed instances, of which I am tempted to give
notes, are perhaps suggestive ; but birds thus entrapped
—whether by marine or fresh-water bivalves—must
often be unable to carry away the molluscs to any
considerable distance, especially when the creatures
happen to close upon their bills. A dunlin with a
small cockle about the size of a hazel-nut clinging to
its bill was once found, near the estuary of the Moy, by
Mr. Robert Warren. It was seen to be making frantic
efforts to get rid of the shell, rising two or three yards
into the air and falling again, and after shaking its head
until exhausted, it lay with outstretched wings panting
on the sands.! A bird of the same kind with a cockle
similarly attached, which had been picked up dead on
the Yorkshire coast, was forwarded to the offices of the
Field, in 1884, by Sir R. Payne-Gallwey.? Another
dunlin with a cockle upon its bill, which got up from
the observer’s feet and flew heavily away, was shot in
1891.2 A tern with a cockle fixed on the upper
mandible was once shot, on the sands at Morecambe
Bay, by Mr. Hancock, who has given an account, also,
of the capture, on Fenham Flats, of a peewit in a similar
plight, having a cockle firmly grasping its bill4 A

caught by the tongue, “ Daily News,” October 5, 1892 ; racoons and
other animals are also said to have been entrapped by shell-fish.

1 R. Warren, “ Field,” Ixiii. (1884), 447.

a Bield,” lx. (1882); 385.

3 “ Scottish Naturalist,” 1891, p. 94.

4 Hancock’s “Catalogue of the Birds of Northumberland and
Durham,” “Nat. Hist. Trans. of Northumberland and Durham,”
vi. (1874), 142.

60 THE DISPERSAL OF SHELLS.

sandpiper which had a large cockle upon one of its
claws and was unable to fly was seen by Mr. D.
McNabb, in 1889, on the coast of Queensland,’ and a
curlew sandpiper with a cockle hanging to one of its
toes was shot,a few years ago,as Mr. J. H. Gurney
tells me, by Mr. G. Hoare, at Cley, in Norfolk ; a snipe
with a large cockle attached in the same way is said to
have been shot on the wing in or about 1866,’ and the
shooting of a sanderling with a cockle thus attached
was recorded in 1872.2. Mr. Gurney tells me that a
tern, caught by the foot by a mussel, was found some
twenty-five years ago on the Hunstanton beach in
the Wash by Mr. F. Cresswell, and that a grey crow
with a mussel upon its bill was caught by Mr.
C. Springall, in 1888, on the beach at Brancaster. An
account, as related by an old hunter, of the finding of
a shoveller duck with an otter’s-shell (Zw¢raria) upon its
bill was given by Mr. J. K. Lord in 1865.4. Mr. Buck-
land (on the authority of Mr. F. Hill, of Helston) has
described the capture of a rail by an oyster ; the speci-
mens, of which a photograph was obtained, had been
mounted in a case.” Mr. Norgate tells me that he saw
a stuffed water-rail, with its bill in an oyster-shell, at
the National Fisheries Exhibition, at Norwich, in 1881.

’ D. McNabb, “ Nature,” xlii. (1890), 415.

7 j.B., “ Science Gossip,” 1866, p: 62:

s H.R. Leach, “ Zoologist,? (2), vies{is72 3414.

4 J. K. Lord, “ Science Gossip,” 1865, p. 79.

> “ Popular Science Monthly,” xvii. (1880), 111-4, copied from
“Land and Water.”

TRANSPLANTATION OF BIVALVES. 6I

The closure of an oyster upon one of the toes of a golden
plover was recorded in the Fze/d, in 1889,! and the
taking of a cormorant with an oyster upon its bill was
reported, in 1892, in the Dazly News.? Many other
similar records, no doubt, might be found; but we must
pass on to the consideration of facts which have a more
direct bearing upon the subject in hand.

INSECTS.

From the number of observations which have been
made, it seems that small fresh-water bivalves, such as
Spherium and Pisidium, frequently attach themselves
to aquatic insects.

FIG. I.

Spherium corneum upon the leg of a Dragon-fly-larva ; taken at Twenty Pits, near
Manchester, and now in the Manchester Museum.

The larva of a dragon-fly with a shell of Sphe-
rium corneum clinging to one of its legs was once
caught at Twenty Pits, near Manchester, and the
specimen preserved with the shell attached (Fig. 1) is

no Paeld,” Lextit."(1889), 308.
* “ Daily News,” October 4, 1892.

62 THE DISPERSAL OF SHELLS.

now to be seen in the Manchester Museum, where it
was deposited by Mr. Hardy. This occurrence, however,
is not of much importance for us, for] am not aware that
dragon-fly larve ever journey overland.

The clinging of bivalves to flying water-bugs and
-beetles is of more interest. Five individuals, at least, of
the water-scorpion (JVepa), a large flying bug, have been
caught with shells attached. A specimen with a small
Spherium corneum on one of its legs, obtained in 1879
from Mere Mere, Cheshire, by Mr. J. Walken, is now
exhibited in the Manchester Museum, where, also, is
another specimen (preserved in spirit) having attached
toita much larger shell of the same species ; two others,
collected by Mr. Hardy in 1889, each carrying a shell
of Prsidium fontinale upon a leg of the hind pair, are,
also possessed by the Museum. (Fig. 2.) A fifth

FIG. 2.

Pisidium fontinale wpon the leg of a water-scorpion (Vefa) ; now in the Manchester
Musenm,

instance has been observed by Mr, Standen, who caught
a specimen with P. /fontinale attached, in a pond near
Birch Hall, Manchester, on 16th May, 1890.

A number of water-bugs (JVotonecta) with “small

TRANSPLANTATION OF BIVALVES, 63

mollusks attached to their legs,” were exhibited by Mr.
Whitelegge at a meeting of the Linnean Society of New
South Wales in 1885,’ and I hear from Mr. C. Hedley
that in the vicinity of Sydney, Mr. Whitelegge has
frequently noticed hemipterous insects, both Motonecta
and Corzxa, laden with bivalves: three specimens of
Pisidium etheridget have been seen attached to one
insect, one on each fore-leg and one on the snout.

Quite a number of instances of the clinging of these
molluscs to water-beetles can be given. As stated by
Mr. Darwin in a letter published in “ Nature” in 1882,
a specimen of the “great water-beetle,” Dytéscus
margtnalis, with a shell of Cyclas cornea [= Spherium
corneum] clinging to one of its legs, was caught by Mr.
W. D. Crick, of Northampton, on 18th February of that
year :—

“The shell was ‘45 of an inch from end to end, ‘3
in depth, and weighed (as Mr. Crick informs me) ‘39
grams, or six grains. The valves clipped only the
extremity of the tarsus for a length of ‘1 ofan inch.
Nevertheless, the shell did not drop off, on the beetle
when caught shaking its leg violently. The specimen
was brought home in a handkerchief, and placed after
about three hours in water; and the shell remained
attached from February 18th to 23rd, when it dropped
off, being still alive, and so remained for about a
fortnight while in my possession.” ?

Ps Proc, Lin. Soc., N:S;W.,”. x: (1886), 760,
2 Darwin, “ Nature,” xxv. (1882), 529-30; and see also “ Life
and Letters,” i11, (1888), 252.

64 THE DISPERSAL OF SHELLS.

Other specimens of the same kind of water-beetle
with Spherium corneum attached have since been
obtained: Mr. C. Oldham found one, with a shell of
good size firmly clasping the extremity of one of the
front legs, in a pond at Woodford, Essex, in September,
1886 ; and I had the good fortune to catch a specimen

FIG. 3.

Spharium corneum upon the leg of a water-beetle (Dytiscus); taken at West Bark-
with, Lincolnshire, and now in the Manchester Museum,

with a shell upon the right front leg in a pond at West
Barkwith, Lincolnshire, in August, 1888 (Fig. 3).'. Mr.
W. H. Heathcote found another specimen with a
shell similarly attached at Farington, Lancashire, in
1889 ; and a further case, in which a small shell was
attached to one of the hind legs of the beetle, was
observed by Mr. Standen, in June, 1890, in a pond at
Birch, near Manchester.

Some large water-beetles, kept by Mr. Norgate in ~
an aquarium, Mr. Darwin states, frequently had one
foot caught by a small fresh-water bivalve (S,
corneum ?) *

1 Journ. of Conch.,” vi. (1888), 363 ; ‘Proc, Eat. soc., Conde

1888, p. Xxxv.
2 “ Nature,” xxv. (1882), 529-30.

TRANSPLANTATION OF BIVALVES. 65

A specimen of Dytescus marginalis with a shell of
Spherium lacustre clinging to one of the front legs was
caught by Mr. Standen, in a pond at Gorton, in 1890.
The shell had been slightly broken, probably from
having been dashed against some object by the insect, but,
being still alive, it held on tightly. A beetle belonging
to the allied genus Aczlzus with a shell of Pzsedium
fontinale on one of the legs of the second pair has been
presented to the Manchester Museum by Mr. Hardy,’
and I hear from Sydney that two water-beetles with
Pisidia attached were obtained by Mr. Whitelegge (now
of the Australian Museum) when collecting, years ago,
in Lancashire ; in one of the cases two shells were
clinging to the same insect, one on each side.

Two or three instances of the clinging of bivalves to
the antennz of water-beetles have been observed,’ and
Mr. A. J. Jenkins tells me that in a large bell-glass,
which he used as a sort of aquarium, a Pzszdium on one
occasion fastened its valves tightly upon one of the
maxillipedes of a shrimp.

Before coming to any conclusion as to whether water-
beetles, -bugs, etc., have much affected the ranges of
bivalve-molluscs and have helped in any perceptible
degree to stock newly formed and isolated ponds with

1 This and the other specimens here stated to have been
presented to the Manchester Museum were obligingly sent to me
in London for examination ; two of them, I regret to say, were
damaged in the post.

2 A case of this kind once came under the observation of Mr.

Hardy; another is recorded in “ Science Gossip,” 1873, p. 190, and
see also “‘ Nature,” xxv. (1882), 529-30.

F

66 THE DISPERSAL OF SHELLS,

these animals, it seems necessary to inquire as to their
flying habits, and I have tried, but without much
success, to ascertain how often they take wing, whether
they are in the habit of making long flights, etc. Canon
Fowler, of whom I inquired, is inclined to think that
Dytisci, as a rule, do not take to flight unless their pools
are drying up, his impression being that they simply
fly when driven by necessity, and then make for near
water ; and if such be the case it is obvious, of course,
that the facts concerning them just referred to have
but little significance. But on the other hand, Mr.
Darwin! ventured to observe that “the species of
Dytiscus often fly at night,’ and indeed he had several
times heard of their having dashed down upon glass
cucumber-frames, mistaking the glittering surface for
water, and Mr. Norgate informs me that he has actually
seen the insects upon the glass of such frames. A
specimen of D. marginalis once flew into a shop in Hull,
and from the nature of the locality, it is probable that it
had travelled some distance.’ Mr. Standen states that he
has several times caught the creatures on the wing when
out moth-hunting after dusk in the evening,’ and Mr.
Musson tells me that he has taken large water-beetles
at the electric light in the main street of Tamworth,
New South Wales. It is unfortunate that more definite
information is not forthcoming, but I believe we may
conclude that these beetles journey from pond to pond

' “ Nature,” xxv. (1882), 529-30.
2 E. Lamplough, “ Science Gossip,” xxii. (1887), 19.
2 R. Standen, “ Nat. Hist. Notes,” iii. (1883), 39.

-

TRANSPLANTATION OF BIVALVES. 67

in the evening or at night somewhat frequently, and it
is perhaps worth mentioning that at the Linnean
Society in 1852, John Curtis, the distinguished en-
tomologist, expressed the opinion that the larger aquatic
insects—especially the Dytiscide—might without doubt
be the means of conveying fish-spawn from one piece of
water to another,’ and Mr. Wallace, in like manner,
discussing the means of dispersal of fishes, observes that
water-beetles “ flying from one pond to another” may
occasionally carry eggs.” No doubt, as Mr. Darwin has
remarked, the creatures generally alight on any pool of
water they may see, for there is no reason to suppose,
as far as I know, that they are in the habit of “ homing”
or returning to the same pool from time to time. But
another difficulty has to be met. At the Entomological
Society in 1888, when I showed the West Barkwith
specimen of D. marginalis with the attached shell of
Spherium cerneum, and ventured to suggest that
occurrences of the kind indicated were probably of
significance when viewed in connection with the dis-
tribution of bivalves, Dr. David Sharp, the president,
objected that the weight of the mollusc would in all
probability prevent or greatly impede the beetle’s flight,
and this objection, no doubt, will have suggested itself to
many. Mr. Darwin, however, has expressed the opinion
that six grains (the weight of an individual of S.
corneum of average size) would not prevent so powerful

1 See J. Hogg, “ Trans. Tyneside Nat. Field Club,” iii. (1854-

8), 75°
2 Wallace, ‘‘ Geographical Distribution,” 1. (1876), 29.

F 2

68 THE DISPERSAL OF SHELLS.

an insect asa Dytzscus from taking flight, adding that in
any case the beetle could carry smaller shells.’ Mr.
Oldham induced his Woodford specimen of JD.
marginalis, which was encumbered with a good-sized
shell of S. corneum, to fly about in a room, and it did not
seem to be greatly impeded; and what is still more im-
portant for us, individuals of this species of beetle carry-
ing S. corneum have actually been caught ox the wing on
two occasions, the insect, in one case, having two shells
clinging to it! Both captures were made by Mr.
Standen while pursuing nocturnal Lepidoptera: the
first specimen, netted in 1883 with a full-grown shell
clinging to one of its legs, was slowly flying along in
Mill Lane, Goosnargh, about five hundred yards from
the nearest pond, but it might possibly have come
from a small ditch about one hundred yards distant ;
the second capture was made at Moor Side, Swinton,
near Manchester, in 1888, when Mr. Standen had the
good fortune to take, on the wing, a specimen which
was carrying /f¢wo shells, one being of good size
and the other small; the nearest pond, he thinks, was
about one hundred and fifty yards distant. These
cases are obviously of extreme interest and value, for
they not only corroborate Mr. Oldham’s observation
and render it quite clear that D. margznalis is strong

1 “Nature,” xxv. (1882), 529-30.

2 This occurrence was briefly referred to by Mr. Standen in
‘‘Nat. Hist. Notes,” iii. (1883), 39, and in “Science Gossip,” xxi.
(1885), 281, and was quoted by me in the “ Naturalists’ World,” iii.
(1886), 61.

TRANSPLANTATION OF BIVALVES. 69

enough to carry shells, at least as large as S. corneum,
but they also furnish the much required proof of actual
overland transportal. No doubt the water-bugs above
mentioned are able to carry Pisidza of the size of
P. fontinale, as well as young Spheria, but I have not
succeeded in obtaining any definite information respect-
ing their flying habits. JVepa is certainly a powerful
insect, and probably it frequently takes wing. Some of
our common “water-boatmen” (JVofonecta, etc.) are
probably even stronger, and I have once or twice seen
them alight upon the surface of ponds in the sunshine,
fold their wings, and disappear into the water.

From the facts now given, I think it may be safely
concluded that the J/oca/ distribution of the smaller
bivalves has been influenced in a marked degree by
aquatic insects; many isolated cattle-ponds, we may
feel sure, have been stocked with these molluscs by
chance visits from flying water-beetles and the like, and
as these occasionally stray out to sea or are blown to
great distances by gales, the facts have possibly a wider
significance: Mr. Darwin records that a Colymbetes, a
water-beetle belonging to the Dytiscidz, once flew on
board the Beagle when forty-five miles distant from the
nearest land.’

AMPHIBIA.

Newts, frogs, toads, etc., it seems, often have their
toes caught by small bivalves: quite a number of
instances can be given; a frog, in one case, had two

U* Origin,” p:34§;

746) . THE DISPERSAL OF SHELLS.

shells upon the toes of the same foot, two newts have
been seen each carrying four shells, and toads have
been dredged with as many as six shells upon their
toes.

The first record, as far as I have ascertained, of fresh-
water bivalves clinging by closure to other creatures is
contained in Knapp’s “Journal of a Naturalist,”
published in 1829, and has reference to the “common
newt (/acertus aquaticus)” :—

“T have seen the boys in the spring of the year draw it
up by their fishing lines, a very extraordinary figure, .
having a small shell-fish (¢el/na cornea) [= Spherium
corneum | attached to one or all of its feet; the toes of
the newt having been accidentally introduced into the
gaping shell, in its progress on the mud at the bottom
of the pool, or designedly put in for the purpose of
seizure, when the animal inhabitant closed the valves
and entrapped the toes, But from whatever cause these
shells became fixed, when the animal is drawn up
hanging and wriggling with its toes fettered all round, it
affords a very unusual and strange appearance.” !

In 1885 Mr. R. W. Goulding recorded the finding of
a newt with a Cyclas (probably S. cormeum) upon one of
its feet in a pond near Louth, Lincolnshire, and Mr.
Heathcote on two occasions in 1889 found shells of S.

1 J. L. Knapp, “Journal ofa Naturalist,” 1829, p. 305; Mr. F. J.
Rowbotham called attention to this passage in “ Nature,” xxv.
(1882), 605.

2 R.W. Goulding, “ Science Gossip,” xxi. (1885), 238-9, and see
also p. 249.

TRANSPLANTATION OF BIVALVES. 771i

corneum attached to the toes of newts at Farington,
Lancashire. The late Mr. W. Jeffery told me that
ever since he was a boy dabbling with aquaria he
had occasionally met with this bivalve clinging to
newts ; the same has been frequently observed by Mr.
L. E. Adams in ponds in the South of England, and Mr.
Hardy, during his long experience as a collector, has

FIG. 4.

Spherium corneum upon the foot of a newt. Preserved in the Manchester Museum,

observed many instances (Fig. 4), as also has Mr.
Standen, who informed me in 1890 that for many years
he had not missed taking either smooth or great warty
newts (Molge vulgaris or M. cristata) with shells of this
species upon their feet. In 1890, in ponds in the neigh-.
bourhood of Manchester, he met with four instances, three
with 17, vulgaris and one with M, cristata ; in one case,

72 THE DISPERSAL OF 'SHEEES.

in a pond at Birch, a newt had four shells clinging to it,
two on one hind foot, and one on each of the fore feet.
A. specimen with four shells attached had been pre-
viously taken in 1883 in a ditch at Goosnargh, the
creature being so much encumbered that it stumbled
along with difficulty." The entrapped toes, according
to this naturalist, are frequently much swollen, which
seems to show that the shells often remain attached
for a considerable time. In 1881, Mr. Norgate informed
Mr. Darwin that the newts in his aquarium frequently
had one foot caught by a small fresh-water bivalve (S.
corneum ?); this, he said, made them swim about in a
very restless manner, both day and night, for several
days, until the toe to which the shell was fixed was
completely severed.? Mr. Jenkins states that he also
has seen newts caught in this way in aquaria. The
taking of a newt, the lower jaw of which was firmly
clasped by the valves of a Pistdium, was recorded by
Mr. Heynemann in 1870.'

Mr. J. T. Riches, ina note published in 1877, mentioned
that he once received a living frog (which had been
found upon a bank by the side of a canal) with a full-
sized shell of Cyclas cornea |= S. corneum] upon one of
its toes. The shell remained attached until, after two
days, it was removed by the observer.’ A recently killed

’ R. Standen, “ Science Gossip,” xxi. (1885), 281.

? Darwin, “ Nature,” xxv. (1882), 529-30.

3D. F. Heynemann, “ Bericht tiber die Senckenbergische
naturforschende Gesellschaft,” 1870, p. 130.

4 J. T. Riches, “Science Gossip,” xiii. (1877), 93.

TRANSPLANTATION OF BIVALVES. 73

frog with a shell of the same species attached to the
outer toe of one of its hind legs was found by Mr. Crick,
im the spring of 1882; by the side of the pond,
presumably near Northampton, in which, a fortnight
before, he had taken the water-beetle and shell referred
to above. The frog’s leg was cut off, and the shell
continued to cling for two days, during which it was
kept in water, but, on being left in the air, the leg soon be-
came shrivelled, and the shell, being still alive, detached
itself.’ Mr. Goulding, in 1884, found a frog with the shell
of a Cyclas (probably S. corneum) upon one of its toes
in a pond near Louth.? Mr. Hudson tells me that he
once saw one, swimming in a pond at Redcar, with two
shells (which were found to be those of S. corneum) upon
the toes of its left hind foot; and another, having an
immature Spherium upon one of its toes, was seen by
the Rev. S. Spencer Pearce, in 1885, at low water on the
south bank of the Thames between Putney and
Hammersmith Bridges. Mr. Standen, in a letter pub-
lished in 1885, mentioned that he had often found these
amphibians with shells of S. corneum attached,’ and he
has favoured me with a note of a case observed sub-
sequently in a lake at Drinkwater Park, near Prestwich.
Mr. Hardy in the course of his collecting has observed

' See “ Nature,” xxv. (1882), 529-30.

2 R. W. Goulding, “ Science Gossip,” xxi. (1885), 238-9, and see
also p. 249; a statement by me, in the “ Naturalists’ World,” iii.
(1886), 61, that a newt and a frog had been noticed in the
neighbourhood of Louth each with a Sfherium attached is based
upon Mr. Goulding’s observations.

2 R. Standen, “ Science Gossip,” xxi. (1885), 281.

74 THE DISPERSAL OF SHELLS.

several similar instances. While writing (March) I
received a living frog from Lincolnshire (through the
kindness of Mr. Davy) together with a specimen of
S. corneum, which when packed up was clinging to one
of the toes, but had become detached during the
journey: the specimens had been obtained by Mr.
Woodthorpe, one of the members of the Naturalists’
Society at Alford, from a batch of frogs which were
spawning in a ditch in that parish; five or six other
frogs similarly encumbered had been seen in the ditch,
all the shells being attached to toes of the hind legs.

EiGe, 5.

Spherium corneum upon the toe of a Toad; taken from a pond on Hampstead Heath,
and now in the British Museum.

The Rev. R. C. Douglas, in 1851, recorded the finding
of a toad, in June, crouching on the marshy edge of a
fish-pond, with the middle toe of one of its hind feet
held between the valves of a “mollusk about half an
inch in diameter, Cyclas cornea, I think ;” locomotion on
the part of the toad, he says, was effectually impeded.!
Mr. J. Peers, who wrote from Warrington in 1865, while
dredging in April ina pond in which both toads and

* R. C. Douglas, “ Zoologist,” ix, (1851), 3210.

TRANSPLANTATION OF BIVALVES. 75

Cyclas cornea abounded, observed a number of the
shells clinging “with the greatest tenacity” to the
toads, some of which had no less than three shells on
each of the hind feet, while instances in which the toes
were entirely free were very rare. None of the shells
were attached to the fore feet." Mr. Standen, according
to his published note of 1885, has also seen the toes of
toads firmly grasped by these shells,?> and in April,
1892, when a number of toads were spawning in the
‘“leg-of-mutton ” pond on Hampstead Heath, I fished
out an individual with a fine shell of the same kind on
a toe of one of the hind legs (Fig. 5).

These cases, it will be seen, furnish no actual
evidence of dispersal, for the amphibians, I believe, were
all caught in water or in its immediate vicinity. It is
obvious, of course, that such animals can never carry
shells to great distances, and if we are to form an
opinion as to the extent to which they have affected
Jocal distribution, we ought to know something of their
habits, whether they often journey from one piece of
water to another, etc., but as to this I have little or
no definite information. Some edible frogs (Rana
esculenta), turned loose by Mr. Henry Doubleday near
a pond by his residence, it is said, “soon migrated to
another pond;”* Mr. Darwin gives a statement, on the
authority of Mr. Norgate, that newts migrate at night
from pond to pond, and can cross over obstacles which

1 J. Peers, “ Zoologist,” xxiii. (1865), 9697-8.
2 R. Standen, ‘‘ Science Gossip,” xxi. (1885), 281.
3 E. Newman, “ Zoologist,” vi. (1848), 2268.

76 THE DISPERSAL OF SHELLS.

would be thought to be considerable ;! and climbing
powers are possessed also, both by frogs and toads, which
can surmount small barriers of many kinds, but a shell
upon one of the toes, though not usually much impeding
ordinary progression, might seriously hinder the animals
when climbing.?, Mr. Peers remarked, in 1865, that a
bivalve by clinging to an amphibian “acquires a more
rapid and extended locomotion than it possesses of
itself, which perhaps may bea means of its distribution,” ®
and in all probability he was right, for, from general
considerations, it seems in the highest degree probable
that such creatures occasionally carry shells from pool
to pool, or from swamps and marshes adjoining rivers
to ponds more or less remote and isolated, but I have
not heard of their having been found, with shells
attached, at any great distance from water. Three
cases, which I have kept back, however, are of some
significance. In 1886, a newt, with a shell of Spherium
corneum upon its right fore foot, was discovered by Mr.
Hardy at the base of a wall at Dunham Massey,
Cheshire, between the grass and the wall, and about
ten yards from the water of a small pond, which it
appeared to have left: a frog, with a shell of the same
kind upon one of its toes, was once found by Mr.
Standen undera log in a damp ditch about thirty yards

* Darwin, “ Nature,” xxv. (1882), 529-30.

? As to the climbing powers of frogs and toads, see “ Science
Gossip,” 1867, p. 234; 1868, p. 94; xvi. (1880), 23, 64, and 165;
xvii. (1881), 69; xviii. (1882), 215; “ Zoologist,” xxi. (1863), 8861 ;
xxil. (1864), 8927 ; (2), iv. (1869), 1830; (3), i. (1877), 184.

* J. Peers, ‘‘ Zoologist,” xxiii, (1865), 9697-8.

TRANSPLANTATION OF BIVALVES. 77

from the nearest water, a pond in which Spheria
were abundant: and, as Mr. Standen has recently told
me, two of these shells were found by Mr. Joseph
Henshall, in September, 1891, upon the toes of a newt
which was making its way across a field, and apparently
proceeding from one pond to another ; when discovered
it was nearly mid-way between two ponds, which are
about one hundred and fifty paces apart. The shells
were shown by Mr. Henshall at a meeting of the
Manchester Conchological Society in January, 1892.

BIRDS.

Near White-house landing, on the Pamunky River,
Virginia, it is said to be impossible to raise ducks on
account of the fresh-water mussels (Uzzo), which catch
the ducklings at low water, and hold them until
drowned by the rising tide," and it may perhaps be
assumed that the closure of bivalves upon the bills of
birds of various kinds (I presume the ducklings were
generally thus caught) is by no means an uncommon
occurrence—it will be remembered that in the case of
marine bivalves several instances have been observed—
but such accidents can have little actual bearing upon
dispersal: even a large and strong bird, in these circum-
stances, would hardly be likely to fly to any considerable
distance.

In 1885, Lieut.-Col. Godwin-Austen published an
extract from a letter received from Mr. F. Tweedie,
respecting the finding of a small bird’s claw between

1} F. Mather, “Fresh-water mussels v. ducks,” American
Naturalist, xii. (1878), 695.

78 THE DISPERSAL OF SHELLS,

the valves of a fresh-water mussel. His informant’s
sons were fishing ina pond at Hole Park, at Rolven-
den, Kent, and the water being low, they were able to
obtain a number of mussels, in one of which they found
a claw which was regarded by the Editor of the Fzedd,
to whom it was submitted, as the “hind toe of some
species of TZurvdus,’ probably a blackbird. The
specimen is now preserved in the British Museum
(Natural History), Cromwell Road. It appeared that
the toe, the tendon of which protruded from the shell,
had been torn off by the bird in endeavouring to free
itself from the grasp of the mollusc. A larger bird, of
course, would have flown away with the shell. As
Godwin-Austen observes :—

“The great interest that surrounds this well authen-
ticated observation is its connection with the distribution
of species. Thevery slight divergence in the characters
of the genus Unzo and of Anodonta all over the world
is a very well-known fact. They are constant over
enormous areas, few groups are more so, and here we
find an admirable example of how they must, from
time to time, be carried from one piece of water, or
from one river system to another. A specimen full of
ova (which are particularly numerous), might in this
manner be conveyed many hundred miles ina single
night, when aquatic birds are on their migration, and
thus stock a new habitat.” ’

A large fresh-water mussel (4 xodonta), upon the foot

' H. H .Godwin-Austen, F.R.S., “ Bird captured by a Fresh-water
Mussel,” “ Field,” Ixvi. (1885), 499.

TRANSPLANTATION OF BIVALVES. 79

of a redshank sandpiper (Totanus calidris) which had
been taken alive with the shell attached, was exhibited
ata meeting in Berlin by Mr. Schaff, in 1888,' and a few
years ago, Mr. Ford, of Redhill, showed Mr. Gurney a
common sandpiper (Zotanus hypoleucos) said to have
been caught, some time previously, on a stream in that
neighbourhood, with a mussel (Azodonta or Unio ?)
firmly clinging to one of its feet: I hear from Mr.
Ford that the bird, when found, was on the bank of the
stream, about three feet from the water, and that the
shell-fish had caught hold of the frog of its foot.
Bivalves must almost certainly cling to birds in this
manner somewhat frequently, but only a few observed
instances (compared with the number noted under the
two previous heads) can be given; we are fortunate,
however, in having clear evidence of occasional over-
land transportal: a tame duck has been known to drag
a heavy Anodonta across two large fields, and, what is
much more important, four birds, a heron, a blue-winged
teal, a duck, and a snipe having shells clinging to their
toes, have been actually shot on the wing.

The case of the duck and Azodonta came under the
observation of Mr. Standen when a boy. A flock of
ducks belonging to his grandfather at Goosnargh used
to make excursions to some ponds at a distance from
the house, and he remembers going out one evening
with several other boys in search of one which had not
returned with the rest, and which was found, about half

1 “ Sitzungs-Bericht der Gesellschaft naturforschender Freunde
zu Berlin,” 1888, pp. 121-2.

80 THE DISPERSAL OF SHELLS.

way home, with a big Axodonta attached to one of its
feet; it was slowly and painfully dragging the shell
along, and had already crossed two large fields.

An account of the shooting of a heron with a large
fresh-water mussel upon one of its feet was once related
~to Mr. Standen by an old gamekeeper at Claughton ;
the bird, flying near the ground, was endeavouring to
shake off the shell—“a big horse-mussel’—which
dangled from its foot, and prevented it from stretching
out the leg in the usual manner. Some years ago a
blue-winged teal (Querquedula discors), with a shell of
Unzo complanatus clinging to one of its toes, was shot
on the wing, by Mr. H. L. Newcomb, near the Artichoke
River, at West Newbury, Massachusetts. The foot
and shell were given to Mr. Gray, of Danversport,
whose letter to Mr. Darwin describing the case was
published in WVature, in 1878, together with a sketch of
the specimens and a note by Darwin. The Uxzo, it is
said, had abraded the skin of the toe, and left quite an
impression.’ In 1884, Mr. J. W. Fewkes recorded the
shooting of a duck, on the wing, near the Sebec River,
Maine, with a “common clam” holding on to one of
its feet by the middle toe. The leg (with the clam
attached) was cut off, and after a day or more placed
in a basin of water, when the mollusc opened its shell
and released the toe. The shell had probably been

' C. Darwin (and A. H. Gray), “ Transplantation of Shells,”
Nature, xviii. (1878), 120-1; a statement by Professor R. E.
Call in the American Naturalist, xii. (1878), 473, seems also to
have reference to this case.

TRANSPLANTATION OF BIVALVES. SI

attached for some little time, for the toe had been
chafed, and the observer heard from boys in the
neighbourhood that the bird had been seen about for
several days, and it seems, therefore, that the shell
“had not released its grasp even when the duck lit
upon the water, as it must frequently have done in
the intervals of time between observation.”' In the
same year Mr. H. V. Chapman submitted to the Feld
the foot of a snipe, with a shell of Spherium corneum
attached to the hind toe, stating that the bird had
been shot by him, while “sailing over my head with
apparently a leg down.”* The foot and shell have
since been presented to the British Museum, and are
now exhibited in the British-room at Cromwell road ;
the bird is said to have been shot near Rye, Sussex.

It is clear, I think, from these cases that bivalves
may be occasionally carried to vast distances by birds,
which, of all highly organized animals, are the least
‘confined by geographical barriers; many annually
migrate over large tracts both of land and sea, and they
are occasionally blown far over the ocean by violent
gales. It may be objected, perhaps, that shells are
likely to drop off during long journeys, but I do not
think this will often happen, for when bivalves once
firmly close upon an object they generally hold on for
a considerable time. But it is certain, of course, that

1 J. W. Fewkes, “Ducks transporting fresh-water clamp.”
Auk, i. (1884), 195-6.
2 H. V. C [hapman], “ Accident to a snipe,” F7e/d, |xiv. (1884),
597, and see also p. 760,
G

82 THE DISPERSAL OF SHELLS.

they must often chance to be set down in unsuitable
spots ; it is true, as Mr. Darwin has remarked, that a
bird, such as a duck or heron, ‘‘if blown across the sea
to an oceanic island or to any other distant point, would
be sure to alight on a pool or rivulet,” ' but probably a
bivalve would generally remain attached for some time
after the bird had landed even if kept much under
water. Sooner or later, however, the valves would have
to be opened, and this is more likely to happen in
water than on land. It may be seriously objected, I
admit, that the transplantation of a shell full of fry or
ova must be a comparatively rare occurrence, and it
is even doubtful, perhaps, whether a single “individual in
this condition would have much chance of establishing
a colony in a new home. Occurrences of the kind
dealt with in this chapter are perhaps, after all, chzefly
significant when viewed in connection with anomalies
in local distribution, and it can scarcely be doubted but
that they go far towards explaining the almost mysteri- |
ous presence of bivalves in isolated ponds, between
which and other waters a more or less constant com-
munication is kept up by animals of many kinds,
especially by flying water-insects and aquatic birds.

Many creatures, other than those above referred to,
are doubtless occasionally entrapped by bivalves.
Three cases of which I have heard are perhaps worth
giving, but they are not of much importance for us.

1“ Origin,” p. 345.

TRANSPLANTATION OF BIVALVES,. 83

A dead water vole with one of its feet firmly held
between the valves of an Axodonia, four inches in length,
was once found by Mr. Hardy on the banks of Mere
Mere, Cheshire.

A snapping turtle (Chelydra), with a Unio (com-
planatus?), about three inches in length, clinging to
its lower jaw, was caught by Mr. J. E. Todd, in 1882,
while on an excursion along Rock River, near Beloit,
Wisconsin. The animals, which were out of the water
several rods from the river, were taken home and kept
in a box, and the reptile was seen to make frequent and
vigorous attempts to push off the clam with its fore
legs, but without success, for when it escaped from con-
finement after two or three days it carried away the
shell still attached to its jaw. The end of the jaw
probably reached to about the middle of the inside of
one of the valves, so that the mollusc would no doubt
be considerably injured, but, ultimately releasing its
erasp in a suitable place, it might possibly recover.’

In 1855-6 Professor Girard found numbers of small
bivalves attached to crayfishes (Astacus fluviatzlis) in
ponds in the environs of Brie-Comte-Robert, Seine-et-
Marne. Every crayfish taken from a pond called “la
mare a lAnglais” had shells upon its toes; another
pond, close by, also contained individuals similarly
encumbered, and a man living at Brunoy, who was in

1 J. E. Todd, “ Chelydra versus Unio,” American Naturalist,
xvii. (1883), 428; see also “ Nachrichtsblatt der Deutschen Mala-
kozoologischen Gesellschaft,” 1883, p. 93; and I am indebted to
Mr. Todd for having communicated some additional particulars,

G2

84 THE DISPERSAL OF SHELLS.

the habit of procuring these animals for the market,
informed the observer that he had noticed the same
phenomenon in some other ponds in the neighbourhood.
Sometimes every one of the eight ambulatory legs
had a shell clinging to it, so that the animal appeared
as if wearing clogs. The shells in question are spoken
of as “ Cyclas fontinalis,’ but as Mr. Heynemann, who
had the kindness to draw my attention to these

FIG. 6.

** Astacus fluviatilis & Cyclas fontinalis.” After Girard, ‘‘ Annales de la Société
entomologique de France,” (3), vii. (1859), pl. 4, fig. r.
observations, points out, M. Girard describes and
figures a much larger shell (Fig. 6).’

Professor Rossmassler? mentions that zebra mussels
(Dretssena polymorpha) have frequently been found
attached, by the byssus, to the tails of crayfishes.

1 Professor Girard, “Annales de la Société entomologique de
France,” (3), vii. (1859), 137-142.
? As quoted in “Ann, and Mag. Nat. Hist.,” (3), xviii. p. 494,

CHAPTER EV.
TRANSPLANTATION OF UNIVALVES.

OPERCULATE water-snails, it seems possible, may some-
times be transported while clinging by closure of the
operculum to the legs of aquatic creatures of certain
kinds. Water-beetles and other such animals, walking
amongst the branches of aquatic weeds or upon the
mud at the bottoms of pools and slow-flowing rivers,
must occasionally insert their legs into the mouths of
these shells, and, zf the operculum be quickly closed,
the mollusc may possibly cling firmly, and may be
carried about by a sufficiently strong animal for a
considerable time. A large and strong-flying water-
beetle, in such a case, might obviously carry a shell of
small size from one piece of water to another. One
little observation, more or less apposite, has been made.
the aquatic larva of a dragon-fly having been seen by
Mr. Hardy in May, 1890, with one foot firmly held
between the operculum and lip of a specimen of
Bythinia tentaculata, and it is interesting to find, also,
that a humble-bee has been seen with an operculate
land-shell holding on to one of its legs in a similar

86 THE DISPERSAL OF SHELLS.

manner.’ I cannot think, however, that the operculate
pond-snails often cling in this way, and, as already
mentioned, those with which we are familiar in this
country are known to be very generally absent
from perfectly isolated waters.? In July, 1891, I
experimented, with grass-stems, on a good number of
specimens of Paludina vivipara in the Lea marshes at
Tottenham, and it seemed hardly likely that animals
chancing to insert their toes into the mouths of these
shells would often be entrapped, for the opercula were
not firmly closed with sufficient suddenness. Most of
the specimens which were induced to hold on to the
inserted stems dropped after a minute or two; one,
however, did not fall until I had carried it, suspended
upon the stem, for more than half an hour, but a sharp
jerk would probably have caused it to drop almost
immediately.

The fresh-water limpets (Axcylus), inoperculate
univalves which generally adhere to stones and water-
plants in ponds and rivulets, were specially mentioned
by Reeve as having very limited facilities for migration.’
But they sometimes ride upon the backs of large
flying water-beetles! The taking of a great water-
beetle (Dytiscus) with an Ancylus firmly adhering to it

* F. W. T., “ Humble-bee trapped by snail,” Fize/d, lxv. (1885),
$43.

7 See Clement Reid, on the “ Natural history of isolated ponds,”
“Trans. Norfolk and Norwich Nat. Soc.,” v. (1892), 279.

> Lovell Reeve, “Land and Fresh-water Mollusks,” 1863,
Pp. 255:

TRANSPLANTATION OF UNIVALVES. 87

was mentioned by Darwin (on Lyell’s authority) in the
“Origin of Species” in 1859;' in 1876, Mr. E. Duprey
stated that he had more than once seen young speci-
mens of A. fluviatilis adhering to another common
water-beetle, Acilius sulcatus, and I hear that Mr.
Hardy once found a specimen of Dytiscus marginalis
with three shells of A. dacustris adhering to the wing-
cases, one on the left and two on the right side.
Carried by these insects “ from pond to pond,” as Lyell
and several authors have surmised, the creatures could
be quickly distributed, of course, over a large tract of
country. That they are occasionally thus carried
seems conclusively proved by an observation made by
Mr. Standen, who was fortunate enough to detect a
shell of A. fluviatilis upon one of the wing-cases of a
Dytiscus caught on the wing after dusk on the evening

' See “Origin,” 1859, pp. 385-6; ed. vi. p. 345; I have not
been able to ascertain by whom this beetle was taken, and the only
mention of the occurrence by Lyell himself, as far as I am aware,
is contained in a letter, dated in 1861, to his nephew, then a boy of
ten, who had been finding specimens of 4. fluviatilis :—“ Natural-
ists used to wonder how this Amcy/us got spread over the country
in separated lakes and streams, till someone found a young
Ancylus adhering to the elytra of one of those large boat-beetles,
Dytiscus marginalis, which you will see in the collection at Drum-
kilbo, and which fly about at night from pond to pond, and may
sometimes carry the Avcylus with them, if, like the Pate//a which
you saw high and dry on the rocks here, he can manage to do
without water for an hour or two, as most probably he can.”
“Life, Letters, and Journals of Sir C. Lyell,” ii. (1881), 347.

2 E. Duprey, “ Ann. and Mag. Nat. Hist.,” (4), xviii. (1876), 344;
and see also Rimmer’s “ Land and Fresh-water Shells,” (1880),
p. 70.

838 THE DISPERSAL OF SHELLS.

of May 8th, 1890, at Birch, near Manchester, about fifty
yards from the nearest water.

Ancylus fluviatilis and allied molluscs requtatle
adhere to the shells of the large fresh-water mussels,
Unio and Anodonta ; for instance, Mr. Baker Hudson,
who has collected large numbers of pearl mussels (Uxzo
margaritifer) from the Yorkshire Esk, has found
A. fluviatilis to occur on at least sixty per cent. of
them,’ and this suggests another possible mode of dis-
persal which may have come into operation once every
now and then in the course of ages, for a bird carrying
a large bivalve clinging by closure to its toe might
obviously transport these limpet-like shells also, for
short distances at any rate, so that it is just within the
range of possibility that a newly formed pool might be
simultaneously stocked by a bird with a univalve and a
bivalve, molluscs as widely dissimilar as are Ancylus
and Unzo or Anodonta.

Fresh-water limpets and inoperculate pond-snails of
various kinds are known to crawl occasionally upon the
backs or limbs of frogs and other aquatic or amphibious
creatures. Mr. W. Thompson, in 1841, mentioned
that he once saw numbers of Limncea peregra (a
common pond-snail) attached to the backs of some
turtles kept in a pond at Fort William, near Belfast,
and they appear to have held on with some firm-

1 And see A. M. Norman, “ Zoologist,” xi. (1853), 3763; Tate,
‘‘Land and Fresh-water Mollusks,” (1866), p. 207; H. Pollard,
“ Naturalist,” 1887, p. 138; H.A. Pilsbry, ‘‘ Nautilus,” iv. (1890),

48.

TRANSPLANTATION OF UNIVALVES. 89

ness, for the turtles were “swimming about with the
Limnee still keeping ‘their seats’ upon them.’* Mr.
Jenkins has seen molluscs of various kinds, both young
and adult, crawling upon the bodies of frogs, toads, and
newts kept by him from time to time. In such
positions, no doubt, the creatures are occasionally
carried for short distances overland from one piece of
water to another. Two observations bearing directly
on the point can be given. A small living shell of
Ancylus lacustris was detected by Mr. Baker Hudson
adhering to one of the legs of a frog caught, hopping
through the grass by the side of a footpath, in Cowpen
Marsh, County Durham, perhaps thirty yards from the
nearest water, the main drainage dyke of the marsh;
and a full-grown specimen of Lzmuc@a peregra was seen
by Mr. Standen, in 1883, upon the back of a toad
which was tramping leisurely along the road, in the
dusk of evening, at Goosnargh, about twenty yards from
the water of a roadside pond.* I am not aware that
any animal has ever been seen to enter the water
with a shell adhering; such an observation—which is
within the bounds of possibility—would certainly be
“of the nature of a great good fortune.”

1 W. Thompson, “Catalogue of the Land and Fresh-water
Mollusca of Ireland,” “ Ann. and Mag. Nat. Hist.,” vi. (1841), 119.

2 This observation was referred to by Mr. Standen in “ Science
Gossip,” xxi. (1885), 281, and quoted by me in the “ Naturalists’
World,” iii. (1886), 61.

CHARTER, WV.
LAND SHELLS: THEIR TENACITY OF LIFE.

AMONG land-shells some of the species are spread over
large areas: of this our own little white snail (Helix
pulchella), which independently of dissemination by
human agency seems to have had a very extensive range,
is a familiar example, and many others might be men-
tioned ; Professor Semper has remarked upon the fact
that the species of Zrochomorpha (a sub-genus of Helzx)
are extremely similar in appearance whether they come
from India, the Moluccas, the Philippines, or the islands
of the Pacific, and several, he says, are distributed
throughout this vast region almost without any varia-
tion in their shells... But as a general rule terrestrial
species are restricted in distribution, and often wonder-
fully so ; most islands, even very small ones, have some
peculiar species, and it is even found that single moun-
tains or valleys sometimes possess species or varieties
found nowhere else in the world.? It is notorious, for

1 K. Semper, “ Animal Life,” ed. 4, 1890, p. 288.
2 See Wallace, “ Geographical Distribution,” ii. (1876), 524;
‘Island Life,” 1880, p. 76; ed. 2, (1892), p. 78.

LAND SHELLS: THEIR TENACITY OF LIFE. gI

instance, that numbers of species are confined to
the Madeiras,! and the islands Madeira and Porto
Santo, each in sight of the other, possess assem-
blages of land-shells which, though representative, are
for the most part different or proper to each.” From
the Sandwich Islands between three and four hundred
species of land-shells have been described, and all, it is
said, are endemic! In the sub-family Achatinelline
(entirely confined to the islands) the average range
of a species, according to the observations of the Rev.
J. T. Gulick, is “five or six miles, while some are re-
stricted to but one or two square miles, and only very
few have the range of a whole island.” It is even said
that each valley, often each side of a valley, and
sometimes each ridge or peak possesses its own peculiar
species.*

Fresh-water species, in some groups at least, generally
range much more widely than terrestrial species, and
it might at first sight seem probable that the former
possess far greater facilities for overcoming natural
barriers—indeed the latter seem to have been regarded
by some writers as practically destitute of all means of
dispersal*—but it seems improbable that such is the
case, for certain terrestrial genera and higher groups

1 See R. B. Watson, “ Journ. of Conch.,” vii. (1892), 4-5.

2 Lyell, “ Antiquity of Man,” ed. 4, 1873, p. 496.

3 Wallace, “Island Life,” pp. 303-4; ed. 2, pp. 316-18; J.T. Gulick,
«Journ, Ling Sec.” 1873, p- 496, and. “ Proc. .Zool.:Soc.,” 1873,
p- 80, as quoted by Mr. Wallace.

4 See, for instance, H. H. Higgins, “Proc. Lit. and Philos.
Soc. Liverpool,” xxxvi. (1882), pp. xliv-xlv.

92 THE DISPERSAL OF SHELLS.

range very widely, some having species scattered at
random in various parts of the world,’ and almost all
oceanic-islands (even those in which fresh-water shells
are very scarce or altogether absent) are inhabited by
land-snails, often in great plenty and variety. Many
of the genera and higher groups have limited ranges,
but it is notorious that some of the families and genera
are almost universally distributed. The family Helzcide,
a group of immense extent, is described as absolutely
cosmopolitan in its range, being found in the most
barren deserts, and on the smallest islands, all over the
globe, and reaching to near the line of perpetual snow
on mountains, and to the limit of trees, or even con-
siderably beyond it, in the arctic regions: the genus
FHlelix also is said to be universal, having found its way
to every country and to almost every island in the
world ; several other genera with very wide ranges might
be named; e.g., Pupa, like Helix, inhabits each of the
six zoological regions.?, Mr. Wallace has maintained
that all the animals now inhabiting truly “ oceanic
islands ” must have reached them by crossing the ocean
or be the descendants of ancestors which did so, for
such islands have been produced in mid-ocean and
have never formed part of a continent; but in some
cases, of course, the creatures may possibly have
migrated from former and unknown lands lying nearer

1 See Wallace, ‘ Geographical Distribution,” ii. (1876), pp.
612-13, 622.

2 “Geographical Distribution,” ii. (1876), pp. 512-13, 522; “ Island
Life,’ ip. 76:5 ed 2p. 74:

LAND SHELLS: THEIR TENACITY OF LIFE. 93

to the islands than the nearest now existing countries.
The Azores or Western Islands, lying at a great distance
from the nearest continental land, and separated there-
from by an ocean of great depth, have been specially
referred to by Mr. Wallace as typical “ oceanic islands,”
and they possess land-shells in some plenty, sixty-nine
species having been recorded for the group, thirty-seven
of which are common either to Europe or the other At-
lanticislands, and thirty-two peculiar, but almost all allied
to European types ; on the other hand, though there are
streams, springs, and lakes, apparently presenting the
most favourable conditions for the existence of molluscs,
not a single fresh-water species has yet been discovered.!
The wholly volcanic and strictly oceanic island of St.
Helena, only about ten miles long by eight wide, and
so remarkable for its extreme isolation, situate as it is
nearly in the middle of the South Atlantic, 1100 miles
from Africa and 1800 from America, possessed twenty
truly indigenous species of land-shells (a large propor-
tion of which have unfortunately become extinct since
the destruction of the forests) ; and several others, pro-
bably recently introduced by man, now occur in the
island. Here, as in the Azores, fresh-water shells are
altogether absent, the streams and tanks of the interior,
the trickling rocks, waterfalls, and pools, being all un-
inhabited by molluscs of aquatic genera? The Sand-

* “Island Life,” pp. 236-7, 239-40, 247, and 293; ed. 2, pp. 245,
247-8, 256 and 305; H. B. Tristram, as quoted by Wollaston,
“ Testacea Atlantica,” 1878, p. 6.

* “Tsland Life,” pp. 281 and 293 ; ed. 2, pp. 292 and 304; Wollas-

94 + ‘THE DISPERSAL OF SHELES-.

wich Islands, wonderfully isolated in mid-ocean, are
inhabited, as we have just seen, by three or four hundred
land species, and possess only “a few fresh-water
shells.”’ It seems evident enough from facts of this
kind that effectual means for occasional transportal
are certainly possessed by land-molluscs, and even in
local distribution within a given country we see much
which in the absence of means for dispersal, at least
over land, would seem well-nigh inexplicable, for more
or less isolated colonies of certain species are not un-
frequently found at great distances from others of their
kind. Mr. Standen adverted to this subject in a com-
munication with which I was favoured in 1891, remarking
that he had often pondered over the question as to
“how and when certain colonies of land-shells had
become located in various queer places,” and referring,
for instance, to two singular colonies of Vertigo pygma@a
in which the creatures live in great abundance within
limited areas, and to the presence of the heath-snail,
Helix ericetorum, on a little bit of rough land in the
middle of a cultivated field, many miles from any other
habitat of the species. In Norfolk, according to

ton, “ Testacea Atlantica,” 1878, p. 6; more recently the total
number of land-shells truly indigenous to St. Helena has been
estimated by Mr. Edgar Smith as twenty-seven, of which, it is
said, seven only are now living on the island, the remainder
having been exterminated by the destruction of the forests: see
report of the Zoological Society’s meeting on 5th April, 1892, in
“ Nature,” xlv. (1892), p. 597.

1 “Island Life,” pp. 293, 298, and 303; ed. 2, pp. 305, 310, and
316.

LAND SHELLS: THEIR TENACITY OF LIFE. 95

Woodward, Cyclostoma elegans is found in the greatest
profusion on the bosses of chalk that appear among the
overlying Tertiary gravels and clays, and is not met
with in the intervening areas.’

Mr. Reeve, it will be remembered, took it for granted
that these creatures possess even “greater facilities of
migration ” than the fresh-water kinds,’ but this conclu-
sion was not based upon definite data and has but little
value ; one is certainly inclined to agree with Mr. Belt,
however, in thinking that the land species “have at
least equal means of dispersion, compared with the
sluggish, mud-loving, water-shells of our ponds and
ditches;”’* but; if this be so, how is it that the
species of the one group are generally confined to
small areas while those of the other are frequently
widely diffused ?

In the first place it may be remarked that the
wonderfully restricted distribution of many terrestrial
molluscs cannot be taken as implying an absence of
means for occasional transport, though, of course,
frequent or constant transportal might have largely
hindered divergence. It must not be supposed, Mr.
Darwin points out, that species which have the capacity
of crossing barriers will necessarily range widely, for
this implies, not only the power of crossing barriers,
but the more important power of being victorious in the

1S. P. Woodward, as cited by Tate, ‘“‘ Land and Fresh-water
Mollusks,” 1866, p. 222.

7 L. Reeve, “‘ Land and Fresh-water Mollusks,” 1863, p. 252.

* T, Belt, ‘‘ Naturalist in Nicaragua,” ed. 2, 1888, p. 334.

96 THE DISPERSAL OF SHELLS.

struggle for life with foreign associates." When in free
intercommunication closely-allied species are not always
able to invade each other’s territory ; two equally well
fitted for their own places seem likely to be able to
hold their separate habitats for almost any length of
time.2 The Achatinelline of the Sandwich Islands,
just referred to, so remarkable for limited specific areas,
have in all probability been occasionally, though
perhaps very rarely, carried by accidental means into
the midst of each other’s districts, but, as their distribu-
tion clearly indicates, they must generally have failed
to establish themselves in the new surroundings, being
unable, no doubt, to compete successfully with those
already in possession. Sir C. Lyell has expressed
surprise that “in the course of that vast lapse of ages
which has occurred since the Newer Pliocene period”
the shell-faunas of Madeira and the adjacent islet,
Porto Santo, have been so little interchanged ;* but in
all probability shells have been occasionally carried
from one island to the other by natural agencies; if
both are well stocked, however, and if the respective
faunas are equally well able to hold their own as
against the other, they are not likely to be much
affected by the occasional transportal of a few
individuals. Some of the characteristic Porto Santo
species live in crevices of stone, and it is probable that
they have often been transported to Madeira by human
agency with the large quantities of stone annually

2 Onigin,? ip, 358: 2 eOnigin 2’ jp." 356;
3 “ Principles,” ed. 12, il. (1875), p. 433.

AWD SHELVES: BHEIR TENACITY OF LIFE. 97

carried there from Porto Santo, but they have not
colonized the island; on the other hand, European
land-shells, believed to have been introduced by man,
are said to flourish both in Madeira and Porto Santo.’

It must be admitted that neither fresh-water nor land-
shells are really well furnished with means for dispersal ;
the transportal of a species of either group over a large
expanse of ocean, or to great distances on land, with
subsequent establishment, must be an extremely rare
and exceptional occurrence, and one which happens,
perhaps, only once or twice in many hundreds of years.
It can hardly be argued, therefore, that fresh-water
species have been kept uniform by constant or frequent
transportal. Numerous facts, it is true, apparently
indicating means of dispersal for fresh-water kinds
have been referred to, and these are doubtless of value
as helping us to understand how it is that many have
been able to wander so far from their several birth-places,
but it cannot be said that they explain the wide ranges
and consequent uniformity which obtain in fresh-water
as against more restricted ranges and greater variety on
land.

It seems evident, as Mr. Belt has remarked, that on
land there has been more variation or that the varieties
which have arisen in fresh-water have less frequently
been preserved. Mr. Darwin has shown that in fresh-
water competition will have been less severe than on land,
and consequently new forms will have been more slowly
produced ;” but this consideration, taken alone, is per-

? Onegin, ip? 357- Origin, pe 33.

Hi

98 THE DISPERSAL OF SHELLS.

haps insufficient to account for the difference in specific
distribution which the terrestrial and non-tnarine
aquatic groups of mollusca exhibit. The problem
presented is of importance, and a solution of it, which
seems likely to be the true one, has occurred to Mr. Belt,
who has pointed out that the variation of fresh-water
species of animals and plants has probably been con-
stantly checked by the want of continuity of lakes and
rivers in time and space :—

“In the great oscillations of the surface of the
earth, of which geologists find so many proofs, every
fresh-water area has again and again been de-
stroyed. It is not so with the ocean—it is continuous
—and as one part was elevated and laid dry, the
species could retreat to another. On the great
continents the land has probably never been totally
submerged at any one time; it also is continuous
over great areas, and as one part became uninhabit-
able, the land species could in most cases retreat to
another. But for the inhabitants of lakes and rivers
there was no retreat, and whenever the sea overflowed
the land, vast numbers of fresh-water species must
have been destroyed. A fresh-water fauna gave place
to a marine one, and the former was annihilated so far
as that area was concerned. When the land again rose
from below the sea, the marine fauna was not destroyed
—it simply retired farther back. There is every reason
to believe that the production of species is a slow
process, and if fresh-water areas have not continued as
a rule through long geological periods, we can see how

LAND SHELLS: THEIR TENACITY OF LIFE. 99

variation has been constantly checked by the destruc-
tion, first in one part, then in another, of all the fresh-
water species ; and on these places being again occupied
by fresh-water they would be colonized by forms from
other parts of the world. Thus species of restricted
range were always exposed to destruction because their
habitat was temporary and their retreat impossible,
and only families of wide distribution could be preserved.
Hence I believe it is that the types of fresh-water
productions are few and world-wide, whilst the sea has
molluscs innumerable, and the land great variety and
wealth of species. This variety is in the ratio of the
continuity of their habitats in time and space.”’'

Before proceeding, in Chapter VI., to a consideration
of possible means for the dispersal of land-shells, I will
venture to append here a small collection of facts on
their tenacity of life. As is well known, some of these
creatures possess in a remarkable degree the power of
retaining vitality under conditions of the most adverse
kind, being able to live for long periods, even for years,
- without either food or moisture; and this faculty must
doubtless be of the greatest use to them in their
involuntary migrations, during which they must often
suffer great hardships; and thus it seems likely that
they are fitted for transportal by means which can never
operate for frailer creatures. Even if they have had to
depend almost solely on voluntary migration, as some
writers seem to suppose, the ability to remain dormant

1 T, Belt, “ Naturalist in Nicaragua,” ed. 2, 1888, pp. 334-5.
HZ

100 THE DISPERSAL OF SHELLS.

for considerable periods must have been highly useful,
for we find that many kinds have learnt to evade the
cold of winter by hibernation and the excessive heat
and dryness of summer by estivation, and thus they
have been able to overcome climatic conditions which
in other circumstances might have been fatal, and some
kinds, having learnt to continue the latter process for
great lengths of time, have been able to penetrate the
dryest deserts. It is important, of course, to inquire
also as to what extent the creatures are able to with-
stand the notoriously harmful effects of contact with
sea-water, for on this the value of certain suggested
means of trans-oceanic dispersal obviously depends ;
but I know only one or two facts bearing upon the
point, and these can be conveniently referred to in the
next chapter.

One of the most remarkable cases of long-suspended
vitality I have anywhere seen recorded is given in
Bingley’s ‘“‘Animal Biography,” vol. i.-’ p.) 5743;
where some snails—on being immersed in water—are
said to have recovered and crept about after an uninter- |
rupted torpidity of more than fifteen years :—

“Mr. Stuckey Simon, a merchant of Dublin, whose
father, a Fellow of the Royal Society, and a lover of
natural history, left to him a small collection of fossils
and other curiosities, had among them the shells of
some snails. About fifteen years after his father’s death
(in whose possession they continued many years), he by

1 As quoted by G. J[ohnston]., Loudon’s * Mag. Nat. Hist.,” vii.
(1834), pp. 113-14.

EAND SHELLS :. FHEIR TENACITY OF LIFE. ~- IO!

chance gave his son, a child about ten years old, some
of these snail shells to play with. The boy put them
into a flower-pot, which he filled with water, and next
day into a basin. Having occasion to use this, Mr.
Simon observed that the animals had come out of their
shells. He examined the child, who assured him that
they were the same he had given him, and said he had
also a few more, which he brought. Mr. Simon put one
of these into water, and in an hour and a half observed
that it had put out its horns and body, which it moved
but slowly probably from weakness. Major Vallancey
and Dr. Span were afterwards present and saw one of
the snails crawl out, the others being dead, most pro-
bably from their having remained some days in the
water. Dr. Quin and Dr. Rutty also examined the
living snail several different times, and were greatly
pleased to see him come out of his solitary habitation
after so many years’ confinement, Dr. Macbride, anda
party of gentlemen at his house, were also witnesses of
this surprising phenomenon..... A few weeks after-
wards the shell was sent to Sir John Pringle, who
showed it at a meeting of the Royal Society ; but some of
the members imagining that Mr. Simon must have been
imposed upon by his son having substituted fresh shells
for those that had been given to him, the boy was re-
examined by Dr. Macbride on the subject, who declared
that he could find no reason to believe that the child
either did or could impose upon his father. Mr. Simon’s
living in the heart of the city rendered it almost impos-
sible for the boy (if he had been so disposed) to collect

102 THE DISPERSAL OF SHELLS.

fresh shells, being at that time confined to the house
with acold. Mr. Simon has also declared that he is
positive those were the shells he gave to him, having
in his cabinet many more of the same sort, and nearly
of the same size.”

Dr. Johnston, who quoted this account in 1834, agreed
with Mr. Bingley in regarding the facts as well authen-
ticated, but the period during which the snails are said
to have remained torpid is exceptionally lengthy, and
I do not think that the evidence is quite conclusive.

HELIX DESERTORUM.—More satisfactory than the
foregoing is Dr. Baird’s celebrated and often quoted
account of a desert snuil (A. desertorum) which re-
mained for four years fixed upon a tablet in the British
Museum, and subsequently revived and lived some time
in captivity. Two specimens which had been collected
in Egypt, it appears, were presented to the Museum in
March, 1845, and on the 25th of that month they were
fixed upon tablets and placed in the collection among
the other mollusca, where they remained till about the
15th of March, 1850, when owing to the fact that a
glassy-looking,and evidently recently-formed, epiphragm
was seen to have been formed in the mouth of one of
the shells, both were removed and placed in tepid
water, and, after the lapse of ten minutes, the animal of
one was seen to gradually come forth, and in a few
minutes more it was walking along the surface of the
basin in which it had been placed: next day it was
supplied with part of a cabbage-leaf of which it partook
readily though in small quantity, and on the 24th of

LAND SHELLS: THEIR TENACITY OF LIFE. 103

June, the date on which the account was written, it was
still alive and in apparent health, preferring cabbage-
leaf to lettuce or any other kind of food which had been
tried.' Dr. Baird’s notice was accompanied by a draw-
ing of the living animal, made by Miss Waterhouse,
which now forms figure 2 in Woodward’s “ Manual.”
The animal in the other shell was found to be dead.
About 1858, two other specimens of the same kind of
snail, believed to have been dormant for more than four
years, were sent to the Museum by Mr. B. M. Wright.
They are said to have been collected in May, 1854, from
a heap of thorn bushes, by Mr. Vernedi, on his journey
through the Desert of Egypt: the bushes were rather
thickly studded with snails, and fifteen or twenty speci-
mens which were picked off were carried home and
locked up in a drawer, where they remained undisturbed
till September, 1858, when two were given to Mr.
Wright, who tried with success the experiment of
reviving them, and afterwards Mr. Vernedi himself suc-
ceeded with two of the others. Theelder Binney relates
that specimens of this snail which had been collected in
Egypt and shipped to Smyrna, thence to Constanti-
nople, thence to Rio Janeiro, and finally to Boston,
occupying a period of about seven months, appeared in
good health when taken from the papers in which they
had been enveloped, and after having lain in a drawer
for three years, some of them still came out in tolerable

* According to a label now affixed to the specimen in the
Museum, the creature continued to live, after its revival, for two
years.

104. THE DISPERSAL OF SHELLS,

vigour. Mr. Jenner Weir has been good enough to
inform me of the resuscitation of a snail, believed to
belong to this species, which had been in his possession,
in a dormant state, for about a year.

HELIX LACTEA.—Concerning this snail a case of
much interest has been recorded by Mr. J.S. Gaskoin. A
specimen, which appears to have contained fertile eggs
when captured, and which had subsequently remained
torpid for more than four years, revived and lived for
some time, a/ome, under a bell-glass, where, notwith-
standing the long suspension of animation, it became
surrounded with a family of about thirty “points.” or
young ones. In April, 1849, four or five specimens, which
had been obtained from a dealer, were placed in water
to be cleaned for the cabinet, and in the course of an
hour or two one of them resuscitated and escaped from
the vessel. The shells had been selected from many
others, ail of which had been stored in a dry dusty
drawer in the dealer’s shop for more than two years,
and had been imported by a merchant of Mogador, in
whose possession they had been, in a similar condition,
for a still longer period. The test of submersion in
water was tried with the whole of the dealer’s stock, but
all seemed to be dead. The individual which had
revived was placed, quite alone, under a large bell-glass
on a tub of earth, and lived well on cucumbers, cabbage-

1 W. Baird, “ Ann. and Mag. Nat. Hist.,” (2), vi. (1850), pp. 68-0.
Woodward’s “ Manual,” ed. 4, rep. 18go, pp. 4and 14; S. P. Wood-
ward, *‘Ann, and Mag. Nat. Hist.” (3), 1i.. (1859), p. 445) 4
Binney, “ Terr. air-breathing Moll,,” i. (1851), pp. 196-7.

LAND SHELLS: THEIR. TENACITY OF LIFE, 105

leaves, etc., and at the end of the following October it
was found to be accompanied by about thirty minute
black helices, of the origin of which the observer was
at first doubtful, but the markings and form of 7. dactea
soon became distinguishable, and when the account was
written, a year later, some of them were nearly as large
as the parent, which was still alive in captivity. In
Africa, incertain parts of the deserts where there is a
constant heat of over 110° Fahr., and where no trace of
vegetation can be seen, the ground is sometimes covered
with H. dactea so as to seem whitened, and at the end
of 1858 Aucapitaine is said to have collected specimens
while passing through places, where, it was believed, no
rain had fallen for five years; these were packed away
in a box and forgotten until August, 1862, when they
were found, and placed in a basin of water to be cleaned,
and next morning, to his astonishment, the observer
found them, full of life, crawling about on the furniture
of his study."

HELIX APERTA.— Woodward placed a specimen of
this species in a giass-box in June, 1855, and it
remained quiescent, without food or change of air,
until November, 1856, when, on being placed in water,
it revived for a few hours, but returned, without
taking any food, to its former condition, and so con-
tinued until July, 1857, when it was required for
dissection. In February, 1885, Mr. Darbishire bought
a quantity of snails of this kind in the market at Nice,

1 J. S. Gaskoin, “ Proc. Zool. Soc.,” 1850,pp. 243-4 ; Aucapitaine,
as cited by Mr. Darbishire, “Journ. of Conch.,” vi, (1889), Tot.

106 THE DISPERSAL OF SHELLS.

of which a few, having their shells sealed up with
strong calcareous lids, were packed away in cotton-
wool and kept in a workshop until December, 1888, by
which time they seemed to have lost the weight and
coolness usually characteristic of life, and one, when
broken, showed the animal dried up like a bit of hard
horn ; but two others, laid on a pad of flannel under a
shade, and thus kept in a moist atmosphere, after a few
weeks showed signs of life, and were then removed to
a damp fern-case, where, in the following March, they
were seen walking out, somewhat feeble, but in good
colour and substance.'

HELIX CANDIDISSIMA AND H. VERMICULATA.—
Two specimens of A. candidisstma, placed in a box
by Woodward, in June, 1855, with the specimen of
Hl. aperta just referred to, remained dormant till
November, 1856, when they were immersed in water
and revived for a few hours; on being put back into
the box, however, they became dormant again without
feeding, and in July, 1857, when the //. aperta was
removed for dissection, another individual of 7. can-
didissima and one of A. vermicularis [? H. vermz-
culata| were put in with them, and all four remained
in a torpid state until Michaelmas, 1858, when the
observer conveyed the box in his pocket to the British
Museum, and the snails were all excited to activity by
the warmth and shaking they experienced. In April,
1859, when a note of the facts was written for publica-

1S. P. Woodward, “Ann. and Mag. Nat. Hist.,” (3), iil. (1859),
448; R. D. Darbishire, “ Journ. of Conch.,” vi. (1889), 101.

LAND SHELES? THEIR TENACITY OF LIFE. ‘107

tion, the creatures were still alive under a bell-glass,
along with several other foreign snails which had been
received at the Museum ina living state." Specimens
of HA. vermiculata and of a species of Leucochroa, it is
stated, have survived a confinement of two years ina
leaden case.”

HELIX HORTENSIS.—A specimen which had been
collected in August, 1843, and laid by and forgotten
until April, 1844, on being placed in a window, soon
crawled upon one of the panes, attached itself to the
glass, and remained there until the following October,
when it was broken down by accident, and soon after
died, having lived without food for fourteen months.’

HELIX ASPERSA.—A specimen of H. aspersa, observed
by Mr. J. Ward, survived in aclosed pot of earth for
about ten and a half months, and, as it is interesting to
note, subsequently produced fertile eggs. Two in-
dividuals were enclosed in the pot at the beginning of
July, 1878, immediately after copulation, and when
taken out for examination about the middle of May,
1879, one was found to be dead—not a vestige remain-
ing except the empty shell—but the other, though the
animal had shrunk to about a third of its former bulk,
on being moistened and supplied with food, revived
and fed, and after about two months deposited ova, the

' S. P. Woodward, “Ann. and Mag. Nat. Hist.,” (3), ili. (1859),
p. 448.

2 QO, Reinhardt, in “SB. nat. Fr.,” 1886, pp. 55-6, as quoted in
the ‘‘ Zoological Record,” xxiii. (1886), Moll., p. 102.

3-H. T. Harding, “ Zoologist,” ii. (1844), 800.

108 TIIE DISPERSAL OF SHELLS,

greater part of which hatched out, so that “a little
colony of vigorous young snails” was_ established.
According toa statement in the American Naturalist,
in 1880, another individual has been known to live
“without food” for thirteen months ; but two, enclosed
in a wooden box by Mr. Gude in August, 1891, were
both dead when examined after about thirteen and a
half months.'

HELIX PISANA.—A snail of this species, collected by
Mr. O. V. Aplin about the middle of August, 1881, and
overlooked, was found to be alive on 18th June, 1882.
On being placed in a jar of water—the epiphragm
having been previously broken away—it crawled out in
less than an hour and a quarter, and was apparently
none the worse for its ten months’ sleep.’

HELIX FRASERI.—Some examples of this snail—
which does not form an epiphragm—brought to Sydney
by a collector from the Richmond River, New South
Wales, in April, 1890, were kept by Mr. Musson, wrapped
in paper, in an open cigar-box on a shelf, for about ten
months, and on examination at the end of this period
were found to be alive.

HELIX VEATCHIL—A specimen of 7. veatchii, from
Cerros Island, is said to have “lived without food from
1859, the year when it was collected, to March, 1865, a
period of six years.” *

* J. Ward, “ Nature,” xx: (1879), 363; Lockwood, “ Am. Nat,”
xiv. (1880), p. 214, as quoted in the “ Zoological Record,” xviii.
(1881), Moll., p. 16.

2 O. V. Aplin, “ Midland Naturalist,” v. (1882), p. 210.

3 R, E, C, Stearns, “Am.: Nat.;” x (1877); 100, and “=f ace

LAND SHELLS: THEIR TENACITY OF LIFE. I09

HELIX UNDATA AND OTHER SNAILS FROM THE
MADEIRAS.—Mr. Gaskoin mentions having received
from Mr. Wollaston several species of living snails,
mostly Helices, indigenous to Madeira and its adjacent
rocks, which, though they had lain in a box in dry
canvas bags for a year and a half, had been revived by
immersion in water. They were put under glass shades
on flower-pots filled with mould, or in large glass cases,
and all fed freely; three specimens of the beautiful
f7.undata, Lowe, produced more than two hundred small
pearl-like eggs, and some of the other species also
oviposited. Mr. Pickering is said to have received
from Mr. Wollaston a basketful of Madeira snails (of
twenty or thirty different species) three-fourths of which
were revived “after several months’ fasting and cap-
tivity,” including a sea voyage. On the same subject a
communication of much interest was published by Mr.
Wollaston in 1850:

“During my residence in the island of Porto Santo,
from April 27th to May 4th, 1848, I collected a large
quantity of Helices peculiar to the spot, and having
placed a small set of each, as types, in separate pill-boxes
(for examination by Mr. Lowe on my return to
Madeira), the rest were killed. These types were named
the following week by Mr. Lowe, and as I had to leave
immediately for England, I had no time to kill the
specimens. On my return home the boxes were

California Academy of Sciences,” October 18th, 1875, as quoted
in “Ann. and Mag. Nat. Hist.,” (4), xix., pp. 355-6 and “ Quart.
Journ. Conch.,” 1., p. 218.

I1O THE DISPERSAL OF SHELLS.

placed in empty drawers of my insect cabinet, since
which, up to the present time (October 19th, 1850),
they had never been opened, or if opened, the speci-
mens had certainly never been taken out. I concluded
of course that they were dead long ago, thinking it
more than probable that they never survived the
voyage to England, and therefore, a fortiori, that two
years and a half in dry pill-boxes was quite sufficient
to remove all traces of existence. However, by immer-
sion in cold water, I find that many of them are
still alive; and though a large proportion have
perished in this long interval, yet I have fourteen
specimens now before me crawling about with the
greatest activity. Thirteen of these are of the same
species, viz.: Helix (Carocolla) papilio, Lowe, and the
other Helzx tectiformis, Lowe, both collected May Ist,
1848, on the Ilheo de Baixo, a small limestone island
off the south-western extremity of Porto Santo... . I
may also mention that I possess a whole bagful of the
beautiful little Helix turricula, Lowe, collected on the
Ilheo de Cima (another and smaller rock, off Porto
Santo), on the 24th of April, 1849, all of which, I find
by immersion, are alive, though the dry and dusty bag
in which they have been enclosed has never been opened
since they were placed there, exactly a year and a half
ago. The same may be said of Helix duplicata
[= H. bicarinata, Sow.| and paupercula, of Lowe, (col-
lected at the same time as the last); I have both in
large quantities, perfectly active, though only now for
the first time taken out of the boxes in which they were

LAND SHELLS: THEIR TENACITY OF LIFE. III

originally placed. I have also a few specimens of a
minute Madeira species, Helzx lentiginosa, Lowe, which
I have ascertained to be alive, although they are so
small that it is difficult to conceive how sufficient mois-
ture to support life can have been retained through this
long period.”!

Three specimens of Helix becarinata (Porto Santo),
recently obtained from an English dealer and kept by
Mr. Gude in a glass-topped box for fifteen or sixteen
months, on being tested in water, proved to be alive and
crawled freely about among damp moss.

BULIMUS PALLIDIOR is said to have survived confine-
ment in a box, without food, during a period exceeding
two years and two months: a living individual exhibited
by Dr. Stearns at a meeting of the California Academy
of Sciences in 1875 was stated to be one of nine which
had been collected in Lower California in March, 1873,
kept in a box undisturbed till June, 1875, and then
revived in a jar containing some vegetable food
and a small quantity of tepid water; subsequently,
however, all had died, except the one exhibited,

which, though not very active, seemed in pretty good
health.’

1 J. S. Gaskoin, “ Proc. Zool. Soc.,” 1850, pp. 243-4: Woodward
and Wollaston, “Ann. and Mag. Nat. Hist.,” (2), vi. (1850),
pp- 489-90; and see also Woodward’s “ Manual,” ed 4, rep. 1890,
pita

Pin Be Steams, Am: Nat.,” xi. (1877); .teo; and “ Proc.
California Academy of Sciences,” October 18th, 1875, as quoted in

* Ann. and Mag. Nat. Hist.,” (4), xix., pp. 355-6 and “ Quart. Journ.
Conch.,” 1.,\p. 218.

I1l2 THE DISPERSAL OF SHELLS.

BULIMUS ROSACEUS.—Four individuals of a large
Bulimus from Valparaiso, as related by Sir C. Lyell in
the “Principles,” were brought to England by Lieu-
tenant Graves, who accompanied Captain King in his
expedition to the Straits of Magellan; they had been
“packed up in a box, and enveloped in cotton, two for
a space of thirteen, one for seventeen, and a fourth for
upwards of twenty months,” but, when exposed by Mr.
Broderip to the warmth ofa fire in London, and provided
with tepid water, Lyell saw them revive and feed
sreedily on lettuce-leaves. 4. rosaceus seems to be the
species referred to, of which Captain King, in a paper
on the Mollusca, etc., collected by the “ officers of H.M.S.
Adventure and Beagle employed between the years 1826
and 1830 in surveying the southern coasts of South
America including the Straits of Magalhaens and
the coast of Tierra del Fuego,” gives the following
note:

“Soon after the return of the expedition, my friend,
Mr. Broderip, to whose inspection Lieutenant Graves
had submitted his collection, observing symptoms of
life in some of the shells of this species, took means for
reviving the inhabitants from their dormant state, and
succeeded. After they had protruded their bodies, they
were placed upon some green leaves, which they fastened
upon and ate greedily. These animals had been in this
state for seventeen or eighteen months, and five months
subsequently another was found alive in my collection,
so that this last had been nearly two years dormant.
These shells were all sent to Mr. Loddige’s nursery,

LAND SHELLS? .THEIR TENACITY OF LIFE. 113

where they lived for eight months, when they .. . all
died within a few days of each other.” '

BULIMUS EREMITA.—A specimen of this snail, from
Turkestan, said to have been kept dry for two anda
half years, revived and ate, but died after three or four
days.”

PUPA TRIDENS AND CLAUSILIA RUGOSA.—Some
snails, apparently belonging to these species, collected
in France and close-packed in a pill-box by Mr. John
Curtis in July, 1830, and subsequently kept in a dry
place without food for nine months, on being placed
on wet moss were seen to revive within twenty-four
hours.”

OPERCULATE SNAILS.—The Cyclostomas, according
to a statement in :Woodward’s “ Manual,’* are well
known to be able to survive imprisonments of many
months. Some foreign species, procured by Mr.
Pickering from a dealer, and kept by him for some
weeks, are said to have revived in water.2 Numerous
examples of C. artzculatum, collected in February, 1858,
by Madame Ida Pfeiffer, in the Island of Rodriguez,
and conveyed from thence to Mauritius, continued
active, without taking food, during a stay there of two

? Lyell, “ Principles,” ii. (1875), p. 377; P. P. King, “Zoological
Journal,” v. (1835), p. 342.

* Goldfuss, as quoted in the “ Zoological Record,” xxi. (1884),
Moll., p. 19.

= |s Curtis, “rans; Lin. Soc.,” xvi. (1833), 766-7.

= id)4, fep. 1890, p. TA.

5 S. P. Woodward, “Ann, and Mag. Nat, Hist.,” (2), vi. (1850),
489-90,

ri4 THE DISPERSAL OF SHELLS.

months, and three individuals remained alive after the
voyage to England, which occupied ten weeks. They
were brought over, packed in paper and rags, ina tin
box with a lid, and were not taken out until a fortnight
after their arrival. One lived for some months under a
bell-glass with moss and ferns.’ A specimen of Rha-
phaulus chrysalis, collected in the month of January by
Captain R. H. Sankey, remained closed in its shell until
27th of June, when it yielded slowly to the means em-
ployed to revive it, finally moving about and creeping
freely under an inverted glass. Cyclophorus znatcus, from
Bombay, was once received by Mr. Benson in a living
state, after a voyage round the Cape, occupying four
months.? According to a statement by Mr. John Curtis
in 1831, operculate molluscs have been reanimated after
having remained dormant in cabinets for very long
periods, “it has been said for forty years” ! *

1S. P. Woodward, “Ann. and Mag. Nat. Hist.,” (3), iv. (1859),

320.
2 W.H. Benson (on R. chrysalis and C. zmdicus), “ Ann. and

Mag. Nat. Hist.,” (3), iv. (1859), 93-4.
3 J. Curtis, “ Trans. Lin. Soc.,” xvi. (1833), 766-7.

CHAPTER: V1.
MEANS OF DISPERSAL.

A CLUE to the almost universal distribution of many
families and genera of land-shells is to be found,
Mr. Wallace observes, in their immense antiquity : in
the Pliocene and Miocene formations, he says, most of
the remains of these creatures are either identical with
or closely allied to living species, while even in the
Eocene almost all are of living genera; no true land-
shells have been found in the Secondary formations,
but they must certainly have abounded, for in the far
more ancient Palzozoic coal measures of Nova Scotia
two species of the living genera Pupa and Zonztes have
been discovered in considerable abundance. Types
having thus “survived all the revolutions the earth has
undergone since Paleozoic times,’ are hardly likely to
be confined by now existing arms of the sea, mountain
chains, and other similar barriers which have effectually
limited the ranges of many groups of higher animals.’
But it is obvious, of course, that antiquity in itself, how-
ever great, could have effected nothing without migra-

1 “ Geographical Distribution,” ii. p. 528; ‘‘ Island Life,” pp.
7a-7,\ed, 2p. 79.

2

116 THE DISPERSAL OF SHELLS.

tion and dispersal. With the great age of the genera
and higher groups in view, however, and admitting that
the creatures are able to migrate on land, even with
extreme slowness, and that they possess some means,
however rarely occurring, of crossing the sea, we need
hardly feel surprise at their having spread into all lands
ever connected with continents, and even into the most
remote and isolated islands of the open ocean,’
Something no doubt is to be attributed to unaided or
voluntary migration over land by gradual progression
generation after generation, for, as Mr. Wallace states,
there is a natural tendency among animals to roam in
every direction in search of fresh pastures ;? but snails
are proverbially slow, and their powers of voluntary
cispersal, therefore, must necessarily be very limited.’
It appears, however, that they are in a better position
in this respect than fresh-water kinds, many of which,
more especially the bivalves, (as far as their own
powers are concerned) seem to be more or less com-
pletely restrained within the limits of the river-basin in
which they happen to live. Dr. Binney has remarked that
snails are not instinctively restricted to particular local
habitations, having no regular places of breeding or of
shelter, and this, he thought, would favour their diffusion,
but it seems, from observations which have been made
since he wrote, that certain kinds return from time to

* See “Geographical Distribution,” ii. p. 526, &c.; “Island
Dales ip. 77.

* “ Geographical Distribution,” i. p. 10,

> “Island Life,” p. 76,

MEANS OF DISPERSAL. DIZ

time to fixed resting places or homes, and probably many
possess the same habit ;' it is doubtless true, however,
as Binney argued, that—although no individual can
have made any considerable progress—a species in the
course of the countless generations which have existed
may have wandered to vast distances from its original
birth-place.” Vast changes of climate, it will be re-
membered, have occurred from time to time, and, as
Mr. Darwin has remarked, a region now impassable
owing to the nature of its climate may have been “a
high-road for migration when the climate was different.” »
It will be remembered also that a great many changes
in the relative positions of land and sea have certainly
taken place during the existence of living genera, so
that uninterrupted highways may once have existed
where arms of the sea now absolutely preclude unaided
migration: Forbes held indeed, as Darwin puts it, that
“all the isiands in the Atlantic must have been recently
connected with Europe or Africa, and Europe likewise
with America,” and other authors, it is said, “have thus
hypothetically bridged over every ocean, and united
almost every island to some mainland;’ it is certain,
however, that the enormous geographical changes here
implied have never really occurred during the period of
recent organisms, but geologists are agreed that great
mutations of level have taken place within this period,

' On the “Faculty of homing in Gastropods,” see “ Naturalist,”
1890, pp. 307-18.

* A. Binney, “ Terrestrial air-breathing Mollusks,” i. (1851),
p. 105.

118 THE DISPERSAL OF SHELLS.

and no doubt, as Mr. Wallace remarks, the general
stability of continents and oceans has been accompanied
by constant changes of form.’ The building up of great
continental extensions for the purpose of explaining
facts in distribution, as Mr. Darwin more than once
observed, has doubtless tended to check the investiga-
tion and study of means of dispersal, and has been an
ill-service to science.’

It seems to have been supposed by some writers that
voluntary migration is the only means of dispersal
possessed by these creatures, but in view of the facts
connected with their distribution referred to in the last
chapter we are bound to admit that they have been
carried, occasionally at least, from one place to another,
even over considerable expanses of ocean, unless indeed,
we fall back upon imaginary extensions of land, or
take for granted—as Reeve did—* “the doctrine of a
plurality of progenitors for each species.” We are con-
fronted at once, however, with a statement by Mr.
Wallace (repeated as recently as 1887 by Professor

* © Origin,” p. 323 ; “ Island Eife,” p. 502, ed. 2, p: 534.

2 See letters to Lyell and Hooker, 1856, “ Life and Letters,”
ii.(1888), pp.78-82; and see also Wallace, “‘ Island Life,” p. 10, ed. 2,
same page: “If we once admit that continents and oceans may
have changed places over and over again (as many writers main-
tain), we lose all power of reasoning on the migrations of ancestral
forms of life,and are at the mercy of every wild theorist who
chooses to imagine the former existence of a now-submerged
continent to explain the existing distribution of a group of frogs
or a genus of beetles.”

> ** Land and fresh-water Mollusks,” 1863, p. 254.

MEANS OF DISPERSAL. 1@ fe)

Heilprin) that the exact mode in which the diffusion
of the creatures has been effected is not known.’ It
seems probable, however, from many considerations,
that they are carried from place to place by accidental
or occasional means of various kinds similar in many
respects to those which we have seen have almost
certainly operated for the dispersal of the fresh-water
groups, yet the very general absence of evidence on the
peint is certainly surprising: hitherto, as Sir C. Lyell
remarked in the “ Principles,” “the naturalist has not
witnessed the arrival of a new continental Me/zr on any
remote oceanic island, except by the aid of man,”’ and,
indeed, it may be said that we have little or no actual
evidence of precise modes of dispersal even for short
distances on land. We shall probably find reason to
believe, however, that transportal through wide spaces
is by no means so rare as might at first sight have been
supposed, but the ultimate establishment of a colony in
a new home, which will always depend on many
complex contingencies and must often be altogether
impossible, is quite another matter; and it should be
clearly understood that the colonization of a species on
an oceanic island, or at any other distant point, as the
result of the trans-oceanic dispersal of a few individuals,
must certainly be extremely rare and exceptional, and
is hardly likely to happen more than once, perhaps,

1 “Geographical Distribution,” li. p. 525; Heilprin, “ Geo-
graphical and Geological Distribution of Animals,” 1887, p.

53-
* “Principles,” ii. p. 434.

120 THE DISPERSAL OF SHELLS.

in the course of many thousands of years. I have
collected a few notes on various suggested means of
distribution, but, as Darwin and A. de Candolle
agreed,’ the putting together of merely fosszb/e means
is poor work; nothing more can be done at present,
however, and we are not likely to obtain much in the
way of direct evidence until some traveller-naturalist
gives his whole attention to the subject, and carries on
a series of careful investigations.

OCEAN CURRENTS.

Professor Semper regarded the currents of the ocean
as beyond doubt the most important means of trans-
portal for land mollusca; indeed, according to this
author, it is “more than probable that land-snails can
travel only in this way, and in no other, from one
island to another.”* But it will be remembered that
Mr. Darwin concluded that the creatures are not likely
to be often thus transported, for, as stated in the
“Origin,” they are known to be easily killed by sea-
water, and their eggs, at least such as were tried, sank
in it and were killed ; it was found by experiment,
however, that when hibernating and having a membran-
ous epiphragm over the mouth of the shell, several
species were able to withstand immersion for seven
days:

“One shell, the elzx pomatia, after having been thus

1 See “ Life and Letters,” ii. (1888), p. 82.
* K. Semper, “ Animal Life,” ed. 4, (1890), pp. 282, 296, 314.

MEANS OF DISPERSAL. Lad |

treated and again hibernating, was put into sea-water
for twenty days, and perfectly recovered. During this
length of time the shell might have been carried by a
marine current of average swiftness to a distance of 660
geographical miles. As this Ae/zr has a thick cal-
careous operculum [or epiphragm], I removed it, and
when it had formed a new membranous one, I again
immersed it for fourteen days in sea-water, and again
it recovered and crawled away. Baron Aucapitaine has
since tried similar experiments; he placed 100 land-
shells, belonging to ten species, in a box pierced with
holes, and immersed it for a fortnight in the sea. Out
of the hundred shells, twenty-seven recovered. The
presence of an operculum seems to have been of im-
portance, as out of twelve specimens of Cyclostoma
elegans, which is thus furnished, eleven revived, It is
remarkable, seeing how well the Helzx pomatia resisted
with me the salt-water, that not one of fifty-four
specimens belonging to four other species of Helezx
tried by Aucapitaine, recovered.” '

It is suggested that the creatures are likely to be
carried by currents while hiding in chinks of drift-timber,
and more rarely with icebergs, in the interstices of floating
pumice, etc. ; thus situated they may sometimes be pro-
tected, or partially protected, for a time from contact with
sea-water, and may possibly be safely carried during calm
weather to great distances, so that the arrival of shells,
still alive, on the shores of a foreign country or distant

Es Oripin,? pi 353:

222 THE DISPERSAL OF SHELLS.

island may not be a very rare event. It cannot be
supposed for a moment, however, that they will often be
landed in spots suitable for the establishment of new
colonies, for drift-timber, etc., is only likely to be thrown
well out of the reach of high tides by violent storms.
Even when this happens it is quite likely that an immi-
grant would be picked up at once by some hungry bird,
or it might find the soil, the climate, or the food-stuffs
altogether unsuitable, and even if it survived and bred
for a time, it might be unable ultimately to com-
pete with the molluscs already inhabiting the spot,
or to withstand the attacks of new enemies surrounding
it. Even under the most favourable conditions a few
imported individuals would very likely lose sight of
each other, and die, consequently, without leaving off-
spring ; and this fate would surely happen to solitary
specimens, unless they chanced to contain fertile
ova. Occasionally, however, in the course of ages,
colonies have almost certainly been thus established,
perhaps by solitary individuals. The facts above
referred to concerning the production of fertile eggs by
snails which had long remained dormant are of interest
in this connection ; one can well imagine, for instance,
that the thirty “points” with which the solitary Helix
observed by Mr. Gaskoin became surrounded might
have founded a colony at some distant pla¢é, even if the
accidents leading to the transportal of the parent had
obliged it to remain without food, in a torpid state, for
a long period, provided of course that the offspring of
the same parent can multiply inter se. It will be

MEANS OF DISPERSAL. r23

remembered that time, independently of fresh trans-
_ portals, will have tended to increase the richness of
insular faunas, for in the course of vast ages the de-
scendants of originally introduced forms will generally
have divided up into distinct varieties, and ultimately
in many cases into distinct species and genera ; indeed,
if such were not the case, we should almost have to
believe in the special creation for oceanic islands of the
endemic types which enter so largely into their faunas. —

Icebergs “covered with an alluvial soil, on which
pine-saplings and a variety of herbaceous plants are
seen growing,” according to Sir C. Lyell, are sometimes
drifted with the currents or blown along by the winds in
the arctic seas, and, Mr. Darwin says, they are “some-
times loaded with earth and stones, and have even
carried brushwood, bones, and the nest ofa land bird,” *
so that it seems just possible that terrestrial molluscs
or their ova may have been thus carried—though perhaps
very rarely—more especially during times when ice
extended over the now temperate regions, and the
safe landing of such creatures at some more or less
distant point, though certainly happening only with
extreme rarity, is perhaps not quite impossible. It has
been remarked, however, in a work edited by Dr.
R. Brown,? that it “seems like straining a point” to
class ice as one of “the agents concerned in the dis-
persal of plants, insects, szol/uscs, etc.” Icebergs, accord-
ing to this work, do not carry nearly so much debris as

Principles,” i..p. 304; “ Origin,” p. 328.

2 « Our Earth and its Story ” (Cassell: no date), ii. p. 310.

124 THE? DISPERSAL OF SHEELS.

is popularly supposed ; and before having travelled far
the “raft” is sure to be lashed by the waves, or melted |
at the base by warmer water or by the wear and tear
of its motion through the sea; “ pieces will calve or
break off, and before long the berg will capsize, owing
to its being top-heavy, and the whole burden it bears be
tumbled into the sea ;” in like manner, it is stated, ice-
fields, too, will break in pieces and discharge their loads.
Referring more especially to the dispersal of seeds, it is
also remarked that “the only possible chance a seed
carried by ice has to germinate in a new situation is
when the ice grounds on a low islet, or is dashed on a
lee shore by the action of the winds—an event which
rarely happens, Even when stranded ona low shore
it cannot approach near enough for its burden to be
thrown on land, as a low-lying coast has shallow water
off it ; or if deep off shore the cliffs are in general high,
so that this amounts to the same thing.” Seeds or
fruits, it might be remarked, when ultimately washed
upon the strand by the waves, might be picked up and
carried inland by birds, for some are well known to pass
the intestines unhurt ; but shells, similarly washed up in
a living state, if swallowed by birds, would generally
be killed and digested.

Floating pumice-stone, from its porous and sponge-
like texture, Mr. Bates and Sir C. Lyell have pointed
out, seems likely to be a vehicle for the transport of the
eggs of various animals, both terrestrial and aquatic.
According to Mr. Bates, pieces have been seen
floating on the surface of the main current of the

MEANS OF DISPERSAL. 125

Amazons, twelve hundred miles from the volcanoes of
the Andes from which they must have come; he once
received a large piece which had been found in the
middle of the stream about 900 miles further down, and
pieces having reached this distance, he remarks, would
be pretty sure of being carried out to sea, and thence
probably with the north-westerly Atlantic current to
shores many thousands of miles away. From the
rounded and water-worn appearance of the fragments,
it was concluded that they must have been rolled about
for a long time in the shallow streams near the sources
of the rivers at the feet of the volcanoes before they leapt
the waterfalls and embarked on the currents leading
direct for the Amazons, and “they may have been
originally cast on the land and afterwards carried to
the rivers by freshets, in which case the eggs and seeds
of land-insects and -plants might be accidentally intro-
duced, and safely enclosed with particles of earth in
their cavities.” It seems very likely also that the eggs of
molluscs and small snails in a state of hibernation may
sometimes be thus enclosed, and as the speed of the cur-
rent, in the Amazons, for instance, during the rainyseason,
is said to be from three to five miles an hour, they might
thus travel, unharmed, on fresh-water to great distances,
and might possibly survive for some days on the surface
of the ocean. I am not aware, however, that snails
or their eggs have ever been found in the interstices
of pumice. It was not until after his return to
England that Mr. Bates came to regard the fragments
he had seen as probable agents for dispersal, and he

126 THE DISPERSAL OF SHELLS.

then much regretted having neglected, whilst on the
spot, the opportunity of ascertaining whether they
contained organisms of any kind.'| Pumice is occa-
sionally washed up on the shores of many islands, and
vessels have encountered vast quantities on the ocean,
but it may often be derived from volcanoes quite near the
sea, or from submarine ones.

Trees are carried out to sea by rivers in vast num-
bers, and being often drifted by ocean-currents to
creat distances, are occasionally thrown up on foreign
shores, even, as Mr. Darwin states, on those of islands
in the midst of the widest oceans,’ and land-snails of
many kinds, conccaling themselves under the bark
and within the trunks of old and hollow trees or living
among the earth and débris at their roots, or depositing
their eggs in such situations, must almost certainly be
carried with such drift-wood with some frequency.
Volumes might probably be filled with accounts of the
floating of timber, etc., upon the sea and its stranding
on more or less distant coasts; most naturalists are
familiar with such facts ; it may perhaps be useful, how-
ever, to refer briefly to one or two observations on the
point. At a great distance from the coast of New
Guinea, about seventy miles north-east of Point
D’Urville, where the great Ambernoh River runs into
the sea, the Challenger, as Mr. Moseley states, found the
water blocked with drift-wood, disposed in long curved

1 H. W. Bates, “ Naturalist on the River Amazons,” ed. 5, 1884,
Pp. 247-9; see also “ Principles,” ii. p. 379.
73° Origin,” p: 326.

MEANS OF DISPERSAL. 127

lines at right angles to the direction in which the river’s
mouth lay, and the screw had to be constantly stopped
for fearit should be fouled. Most of the pieces—which
did not appear to have been very long in the sea—were of
small wood, branches and small stems ; among them, how-
ever, were many whole up-rooted trees. As the ship was
nearing Dobbo, the port of the Aru Islands, large quan-
tities of leaves, fruits, etc., and branches of trees were
encountered, drifting about mingled with a floating sea-
weed, and off Ke Islands, and also among the Moluccas,
similar drifts of land vegetation were met with. The
sea-birds, especially terns, Mr. Moseley says, habitually
resort to floating logs as resting places, and it is curious
to see them in the distance appearing as if standing on
the surface of the water, the logs themselves being
often invisible." Dr. Binney states that he once saw
Nantasket beach, at the mouth of Boston harbour,
strewn with logs, “driven from the rivers of Maine by
easterly winds of several days’ continuance,” and he refers
also to the frequent stranding of trunks of trees, frag-
ments of wood, seed-vessels, and numerous other objects,
some of unascertained origin, and others from the
Spanish Main, Cuba, etc.,on the shore of Key West,
and on the beach of Cape Florida and the shores and
islands to the north of it.2, Mr. C. T. Simpson mentions
having seen, in sheltered caves on the island of Utilla,

’ H. N. Moseley, “ Notes by a Naturalist on the Challenger,”

1879, pp. 307, 432-4.
* A. Binney, “ Terrestrial air-breathing Mollusks,” i. (1851), pp.

I 53 I 57:

128 THE DISPERSAL OF SHELLS.

etc., and on the shores of Florida, large numbers of
stranded trees, some submerged all but a few branches,
others at the tide line, and others thrown high and
dry by storms out of the reach of the sea at ordinary
times.’ Dr. Alford Nicholls, of Dominica, when in
Tobago in 1891, saw large trunks of trees stranded
along the coast; they had evidently come from the
Orinoco, and many of them had the bark and some
of the branches still attached. In the Radack Archi-
pelago, situate in the western part of the Pacific, as
mentioned in the Beagle Journal, palms and bamboos,
from somewhere in the torrid zone, and trunks of
northern firs, are known to have been washed on shore,
the latter, it is remarked, having necessarily come from
an immense distance. Drift-wood, bamboos, and canes,
according to Captain Collnett, are often washed on the
south-eastern shores of the islands of the Galapagos
Archipelago. At the Keeling or Cocos Islands, in the
Indian Ocean, trunks of the sago-palm, and large masses
of Java-teak and yellow-wood, the blue gum-wood of
New Holland, and immense trees of red and white cedar,
etc., are said to have been washed up; fishing-canoes,
also, apparently from Java, have come ashore at times.”
A South Sea islander’s canoe, Mr. Musson tells me, was
once thrown up on Curtis Island, Queensland. Vast

1 C, T. Simpson, “ Conchologists’ Exchange,” ii. (1887), p. 38.
* Darwin’s Beagle Journal, ed. 2, 1845, pp. 392 and 454-5,
quoting Chamisso, on the Radack Archipelago; Collnett, on

the Galapagos group: and Keating (“‘ Holman’s Travels”), on the
Keeling or Cocos Islands.

MEANS OF DISPERSAL. 129

quantities of earth, etc., are sometimes carried away
upon the roots of trees by large tropical rivers, where,
also, floating rafts or islands, often of considerable size,
are frequently formed, and occasionally carried out to
sea, to be drifted, perhaps, during calm weather, to great
distances. Sir C. Lyell collected some very interesting
notes on this subject, which J give in his own words:

“Within the tropics ... . there are floating islets of
matted trees, which are often borne along through con-
siderable spaces. These are sometimes seen sailing at
the distance of fifty or one hundred miles from the
mouth of the Ganges, with living trees standing erect
upon them. The Amazon, the Congo,and the Orinoco,
also produce these verdant rafts, [which are formed of
accumulations of floating trees, arrested in their pro-
gress by snags, islands, shoals, or other obstructions,
as had been] already described when speaking of the
great raft of the Atchafalaya, an arm of the Mississippi,
where a natural bridge of timber, ten miles long, and
more than two hundred yards wide, existed for more
than forty years, supporting a luxuriant vegetation,
and rising and sinking with the water which flowed
beneath it.

“On these green islets of the Mississippi, observes
Malte-Brun, young trees take root, and the pistia and
nuphar display their yellow flowers: serpents, birds, and
the cayman alligator, come to repose there, and all are
sometimes carried to the sea, and engulphed in its waters.

“« Spix and Martius relate that, during their travels in
Brazil, they were exposed to great danger while ascend-

K

130 THE DISPERSAL OF SHELLS.

ing the Amazon in a canoe, from the vast quantity of
drift-wood constantly propelled against them by the
current ; so much so, that their safety depended on the
crew being always on the alert to turn aside the trunks
of trees with long poles. The tops alone of some of
the trees appeared above water, others had their roots
attached to them with so much soil that they might be
compared to floating islets. On these, say the travellers,
we saw some very singular assemblages of animals,
pursuing peacefully their uncertain way in strange com-
panionship. On one raft were several grave-looking
storks, perched by the side of a party of monkeys, who
made comical gestures, and burst into loud cries on see-
ing the canoe. On another was seen a number of
ducks and divers, sitting by a group of squirrels. Next
came down, upon the stem ofa large rotten cedar tree,
an enormous crocodile, by the side of a tiger-cat, both
animals regarding each other with hostility and mistrust,
but the saurian being evidently most at his ease, as
conscious of his superior strength.

“Similar green rafts, principally composed of canes
and brushwood, are called ‘camelotes’ on the Parana in
South America ; and they are occasionally carried down
by inundations, bearing on them the tiger, cayman,
squirrels, and other quadrupeds, which are said to be
always terror-stricken on their floating habitation. No
less than four tigers (pumas) were landed in this manner
in one night at Monte. Video, Jat: .35°.S:,:tovthe yanean
alarm of the inhabitants, who found them prowling
about the streets in the morning.

MEANS OF. DISPERSAL. 131

“In a memoir lately published, a naval officer relates
that, as he returned from China by the eastern passage,
he fell in, among the Moluccas, with several small
floating islands of this kind, covered with mangrove
trees interwoven with underwood. The trees and
shrubs retained their verdure, receiving nourishment
from a stratum of soil which formed a white beach
round the margin of each raft, where it was exposed to
the washing of the waves and the rays of the sun. The
occurrence of soil in such situations may easily be
explained ; for all the natural bridges of timber which
occasionally connect the islands of the Ganges, Mis-
sissippi, and other rivers, with their banks, are exposed
to floods of water, densely charged with sediment.

“Captain W. H. Smyth informs me, that, when cruis-
ing in the Cornwallis amidst the Philippine Islands, he
has more than once seen, after those dreadful hurricanes
called typhoons, floating masses of wood, with trees
growing upon them ; and ships have sometimes been in
imminent peril, as often as these islands were mistaken
for -terra -firma, when, in fact, they were in rapid
motion.

“Tt is highly interesting to trace, in imagination, the
effects of the passage of these rafts from the mouth of a
large river to some archipelago, such as those in the
South Pacific, raised from the deep, in comparatively
modern times, by the operations of the volcano and
the earthquake, and the joint labours of coral-animals
and testacea. If a storm arise, and the frail vessel be
wrecked, still many a bird and insect may succeed in

K 2

132 THE DISPERSAL OF SHELLS.

gaining, by flight, some island of the newly-formed
group, while the seeds and berries of herbs and shrubs,
which fall into the waves, may be thrown upon the
strand. But if the surface of the deep be calm, and
the rafts are carried along by a current, or wafted by
some slight breath of air fanning the foliage of the
green trees, it may arrive, after a passage of several
weeks, at the bay of an island, into which its plants and
animals may be poured out as from an ark, and thus a
colony of several hundred new species may at once be
naturalized.” '

Many kinds of snails, from the nature of their habitats,
seem eminently liable to be carried with floating trees,
rafts, etc. Large numbers, every one knows, hide
themselves beneath the bark of trees; the Swzcccnea
arborea, to give an instance, lives, Mr. Musson tells me,
on the inner side of the bark of gum-trees in Australia,
and is difficult to remove without breaking the shell ;
and several of our own snails—C/lauwsili@, etc.—as every
conchologist knows, are often found in similar situations,
under the bark of old forest trees. Crevices within the
trunks of partially hollow trees are also favourite hiding
places for many kinds, and some are in the habit of
eating their way far into the decaying wood. The Rev.
L. Blomefield has described a pollard-elm pierced in all

1“ Principles,” ed. 9, 1853, pp. 640-2, quoting “System of Geo-
graphy,” vol. v. p. 157; Spix and Martius, “ Reise, etc.,” vol. iil.
pp. 1011, 1013; Sir W. Parish’s “Buenos Ayres; p. o7)
Robertson’s “ Letters on Paraguay,” p. 220; and “ United Service
Journal,” No, xxiv. p. 697.

MEANS OF DISPERSAL. 133

directions by rock-snails (Hel¢x lapicida), great numbers
of which, of various sizes, were exposed to view when
the bark and portions of the wood were torn away.’
Species in no sense arboreal may often take up their
abode in chinks in logs which happen to be lying on
the ground on the banks of water-courses, and which of
course are liable to be drifted off during floods.
Strictly ground species may occasionally be carried, in
numbers, in the earth, etc., at the roots of trees. It
seems probable also that many may be transported
with miscellaneous vegetable débris, cocoanut-husks,
etc., which must often be swept from the ground and
carried away to considerable distances, but snails
travelling in this way would be much exposed to the
evil influences of salt water. A few shells (one of
which was alive) of Helix annulus—a species discovered
during the Chevert Expedition near the mouth of
Katow River, New Guinea—were found, about forty
miles south from Katow, at Dungeness Island, Torres’
Straits, to which place they are likely to have been
carried, Mr. Brazier thinks, on the “drift-timber and
palm roots which are to be seen floating in the Straits
after the north-west monsoons: ”’ the species lives, in the
original locality, “in clusters inside of old cocoa-husks,”’
and these also, it would seem probable, may have
served as vehicles for its transportal. Mr. Liardet has
remarked that the numerous small terrestrial shells

‘ L. Jenyns (now Blomefield), ‘‘ Observations in Natural His-
tory,” p. 321, as quoted in the “ Zoologist,” (3), x. (1886), 452-3.
“)estavier, Proc: Lim. coc, N.5.W.;" 1. (1877); 100:

134 THE DISPERSAL OF’ SHELLS.

inhabiting the coast-lands of the Fiji group, and common
to most of the islands, living as they do, “sheltered by
bark on logs, and in the decaying husks of old cocoa-
nuts,” are probably frequently transported from island
to island.’ As indicating a possibility of actual ¢vans-
plantation by means of agencies of the present kind, it
is interesting to note that drifted trees are known to be
occasionally left by storms on island or foreign coasts,
high and dry, out of the reach of the ordinary tides.
Mr. C. T. Simpson, as just quoted, states that he
has seen trees thus stranded on the island of Utilla, and
on the shores of Florida. Charles Goodridge, who
stayed some time in the Crozet Islands in 1821-3, de-
scribed the discovery of several tree-trunks, lying on
the ground as if thrown up by the sea, more than a
mile from the reach of the tides. The wood was
evidently not fossil, and “ Goodridge concluded that it
was drift-wood thrown up so far during some volcanic
convulsion.” ° It will always be remembered, of course,
that complete protection from sea-water during long
periods will hardly ever be afforded by floating timber,
and thousands of inland molluscs must annually perish
in the sea; partial protection, however, both for adult
animals and ova, must often be afforded for a consider-
able time, and some snails, as we have seen, can with-
stand total immersion for short periods. Trees,

1 E. A. Liardet; * Prac: Zool. Soc.,” 1676, p--99.

2 C. M. Goodridge, “‘ Narrative of a Voyage to the South Seas,
etc.,” pp. 42-3, as quoted by Mr. Moseley, ‘‘ Naturalist on the
Challenger,” 1879, pp. 182-3.

MEANS OF DISPERSAL, 135

especially if loaded with stones, etc.,as Mr. Darwin has
remarked, seem likely in many cases to float beneath
the surface. In the channels of Tierra del Fuego large
quantities of drift-timber are cast upon the beach, yet it
is extremely rare to meet a log swimming on the water.’
Even in the case of timber which floats upon the surface
the water will gradually penetrate every crevice, and
most trees are almost sure to lose their bark before long ;
the bark of those seen by Mr. Moseley off the coast of
New Guinea was often detached, floating separately,
and much wood was held vertically in the water; but
on the other hand, as we have just seen, many of the
tree-trunks evidently from the Orinoco, observed by
Dr. Nicholls on the coast of Tobago had the bark still
attached. The pollard-elm described by Mr. Blomefield
is said to have been so weakened by the attacks of
Helix lapictda that a slight wind might have overturned
it,and one can well imagine that sucha tree, after having
fallen to the ground, might easily be swept off by the
sudden overflow of a river, and the snails contained
within it, if “in a torpid state ” with the mouths of their
shells stopped up “with a bung of sawdust and small
chips of wood cemented together,” as was the case with
some of those in the tree in question, would be well
fitted to resist salt water, but a tree of this kind when
launched upon the waves would almost immediately
become sodden and fall to pieces. Immense numbers of
land snails, especially of kinds frequenting damp places

* Darwin’s Journal, ed. 2, 1845, p,. 462,

136 THE DISPERSAL OF SHELLS,

and river margins, might possibly be carried with
floating islands or rafts, but it is obvious, of course,
that the landing of objects of this kind on foreign shores
in such a manner as to enable slow-moving terrestrial
creatures to disembark must be very rare indeed, yet
such occurrences, as Lyell maintained, notwithstanding
their extreme rarity, may possibly account “in tropical
countries for the extension of some species of mammalia,
birds, insects, /and-shel/s, and plants to lands which
without such aid they could never have reached.” !
Professor Semper has remarked that large land-shells,
and such as live in the highest branches of trees and
lay their eggs there, like all the species of Cochlostyla,
will obviously be far more difficult to transport than
small species which can creep into rifts in trees or be-
tween the roots ; and species belonging to groups, which
(like Hlelzx stmzlaris) live on the ground among stones
and earth, will, he observes, be almost as well protected
during a sea voyage as the operculate kinds. This he
shows is in harmony with certain known facts in dis-
tribution, most of the small species and of the operculate
species having a much wider range than the large
inoperculate forms. In the Philippines the typical
genera (or sub-genera) Cochlostyla, Rhysota, Chlorea,
and Oddzma, which principally live on trees, are
almost confined to that group of islands, while the
small genera, as Subulina, Trochomorpha, and Eunea,
among the Helicidz, and the operculate genera Cyclo-

, * Principles,“ ip. i307.

MEANS OF DISPERSAL. 137

phorus, Alyceus, Helicina, and Diplommatina, have a
very wide distribution.’ It seems probable, however,
that large arboreal snails, even those which live in the
highest branches, may possibly be carried from place to
place when several trees are drifted off together in the
form of a floating islet. Some kinds adhere firmly to
the branches when hibernating, and others shelter
themselves among the epiphytic vegetation. Aulimus
fasciatus, a large species which lives on the branches of
trees, attaches the mouth of the shell very strongly to
the bark by means of a thick opaque secretion which
hardens to the consistency of glue, and in tearing the
animal away the bark or the shell is fractured sooner
than the secretion. Pulimus undatus, also, another
large arboreal snail, attaches itself by means of a similar
clutinous substance, which, though exposed to wind
and rain, forms a perfect adhesion and protection to
the animal and only yields to its own solvent powers
on the approach of spring. These, and other species
with similar habits, may almost certainly, as already
suggested by Mr. Simpson, be carried in safety over
the sea on floating trees the higher branches of which are
borne above the crest of the waves, and it even seems
likely that they may float safely for moderate distances
in the sea itself adhering to detached branches, etc.
Both the species above mentioned are enumerated by
Dr. Binney among shells believed by him to have been
introduced by currents from Cuba to the southern part

1 « Animal Life,” ed. 4, 1890, pp. 287-8.

138 THE DISPERSAL OF SHELLS.

of the peninsula of Florida and the adjacent islands:
Mr. W. G. Binney, however, writing of B. undatus, has
suggested that the creature may have been a widely
distributed species of an ancient fauna, which has
survived at various points around the gulf of Mexico.!
Unfortunately I do not know that any observa-
tion clearly indicating the transportal of molluscs, or
their eggs, with drift-timber, etc., has ever been made.
The creatures have never been found, as far as I have
ascertained, in the crevices or under the bark either of
trees encountered upon the sea, or of those stranded on
foreign coasts. Important discoveries may possibly be
made, however, by some traveller who applies himself
to the careful examination of a large number of such
trees. Mr. Layard tells me that landing on one
occasion on ‘fa lone, sandy island in the Indian Ocean,
north of Madagascar,” he found on the shore a huge
tree-trunk, ‘just arrived from the African Coast,”’
swarming with ants, and containing in a crevice the
living eggs of a lizard, probably a gecko !

RIVERS AND FLOODS.

No doubt, as Mr. Wallace observes, terrestrial molluscs
have been “ widely scattered over land areas” by “ large
rivers and occasional floods.”? From considerations
above referred to it is obvious, of course, that both adults

+ A. Binney, “Terrestrial air-breathing Mollusks,” 1. (1851),
p- 153; ii. (1851), pp. 270.0274; W.G. Binney, v. (1878), p, 4oe);
C. T. Simpson, ‘* Conchologists’ Exchange,” ii, (1887), p. 38.

> © Geographical Distribution,” ii. p. 525.

MEANS OF DISPERSAL. 139

and ova must often be carried down stream, sometimes
to great distances, along with floating timber, vegetable
débris of many kinds, pumice, &c., which must fre-
quently be stranded on the low and shelving banks of
river-margins and on islands in mid-stream, but I do
not know that we have any actual evidence of dispersal
in this way. The landing of a tree in a condition
similar to that of the pollard-elm above mentioned,
however, would certainly be sufficient to account for
the presence say of Helix Jlapictda—quite unknown
perhaps in the surrounding neighbourhood—on some
such island or river-bank, for a whole colony of snails,
possibly from a locality several miles distant, might
thus be safely landed in a new home. Dr. R. Brown
(or some author edited by him), mentions having seen,
on his way up some American river, the huge trunk of
a tree “floating leisurely down stream with plants
blooming on its decayed surface,” and weeks afterwards,
on coming down the river, he has noticed, as he assures
us, the same tree “left high and dry on the banks, with
the same plants, still in flower or in seed, growing upon
it.” The pumice-stones floated down by the Amazons,
as Mr. Bates states, are sometimes stranded on the
banks in different parts of the river... Many of the
smaller snails may frequently be floated in the hollow
stems, or kexes, of large umbelliferous plants, for these,
often lying loosely upon the ground, are liable to be
swept away by floods, and molluscs are known to hide

* See “ Our Earth and its Story,” (Cassell : no date), p. 306.
* “ Naturalist on the River Amazons,” ed. 5, 1884, p. 248.

I40 THE DISPERSAL OF SHELLS.

within them in winter-time: Vertigo edentula, for in-
stance, as Mr. Jeffreys mentions, is then to be found
in such stems, together with Carychtum minimum, and
other minute shells.’ Thus concealed, the creatures
would be buoyed up and protected in some degree from
contact with the water. It seems probable also that
entirely unprotected individuals, when swept from river-
banks during floods, or chancing to fall into the water
at ordinary times, might occasionally be carried in
safety to considerable distances, for they sometimes
float on the surface and are never immediately drowned.
Kinds which I have tried’ have generally floated when
the animal was extended from the shell. A specimen
of Helix aspersa, in this condition, for instance, thrown
into the river Lud, floated freely with the current, but
I soon lost sight of it: Fifteen specimens of 2
hortensis, with the animals fully extended, all floated.
The creatures sink, however, as a rule, if tried when the
animal is entirely withdrawn ; thus, of ten specimens of
FT. rufescens, all sank with the exception of two in
which the animals were extended, and these, tried again
when they had retired into their shells, sank also. But
this is not invariably the case: of thirteen specimens
of H. hortensis, for instance, placed in water when the

1 “ British Conchology,” 1. (1862), pp. xxxiv., 269, 301; and see
J. E. Harting, “ Zoologist,” (3), ii. (1878), 125. In gardens near
Sydney, Mr. Musson tells me, Helzx aspfersa often takes up its
abode in the hollows of cut bamboos.

* Helix aspersa, hortensis, ‘arbustorum, rufescens, &c.; freshly
collected specimens for the most part.

MEANS OF DISPERSAL. 141

animals were within the shells, four floated. An
individual of H/. aspersa in the same condition, and
having the mouth closed with an epiphragm, also
floated ; and it is worth noting, perhaps, that when left
in water for four hours, though it slightly forced the
protecting epiphragm, it did not extend its body. The
reader will remember, of course, that individuals which
from their weight could not be floated by slowly-flowing
rivers in their ordinary action, might be carried to con-
siderable distances by streams with powerful currents,
and by the rushes of water which occasionally occur
in many slower streams in times of flood. Prodigious
numbers of shells are known to be carried down by
rivers, and they are frequently collected from alluvium
(as dead-shells) in great quantity. Very large numbers,
for instance, some evidently from a great distance,
were collected by Dr. Scharffin March and April, 1879,
after the subsidence of an _ over-flow of the river
Garonne, comprising no less than 148 specimens of
Pupa muscorum, and 352 of Vertigo pygmaea!* I
hoped to have found records of the discovery of living
shells in such situations, but have not done so,
and it is perhaps hardly likely that evidently river-
carried individuals should often be noticed, for unless
they happened to be ina state of hibernation they would
be sure to crawl away almost immediately from the
débris with which they were stranded. None of the
specimens from the Garonne were seen to be alive, but

* R. Scharff, “Journ. of Conch.,” ii. (1879), pp. 315-16.

142 THE; DISPERSAL (OF (SHELES.

Dr. Scharff tells me that he remembers having somewhere
seen an account of the finding of Helzx erzcetorum alive
in alluvium, In December, 1880, Mr. Ashford win-
nowed out from amongst broken reeds and grasses more
than two thousand shells of the young of Swccinea putris,
which had been swept from their winter moorings on
the vegetation bordering the Avon by a recent flood,
and left in the meadows near Christchurch ; the greater
part, if not all, however, it is said, had “fallen a prey
to tiny but voracious larve, probably of the smaller
coleopterous insects,” ’

As instancing possible or probable results of river-
transportal we may advert to statements given by Mr.
W. G. Binney in the second supplement to his famous
work, previously quoted, on the land-shells of the United
States and adjacent territories.” Helix strvigosa, it is
stated, a widely distributed Central Province group of
forms, ranging to the westward to the Sierra Nevada
and Cascade Mountains, passes the latter even to the
Pacific Ocean, but the author doubts whether it is “really
an inhabitant of the Pacific Region,” the specimens col-
lected from time to time west of the Cascades in
Washington Territory and Oregon being possibly “ in-
dividuals brought down by the Columbia River from
the east of the Cascades, or colonies descended from
such.”” Mr. Hemphill, he states, first called his atten-

' C. Ashford, “ Journ. of Conch.,” 111. (1881), pp. 195-6.

2“ Terrestrial air-breathing Mollusks,’ second supplement to
vol. v., pp. 26-8, 32, 39 : Bulletin Museum of Comparative Zoology,
Harvard College, vol. xiii. No. 2.

MEANS OF DISPERSAL. 143

tion to this explanation of the presence in the Pacific
Province of Central Province species, in a letter, con-
cerning A. strigosa, from which the following is an
extract. The same considerations, according to Mr.
Binney, apply to Helzx solitaria, the group of Helex
mullant, and felix ptychophora :

“ The Cascade range of mountains in Oregon is, as you
are aware, a continuation of the Sierra Nevada. It
crosses the Columbia River between the Dalles and
Portland, and continues its northerly course on the west
side of the Columbia. Numerous spurs, however, break
off from the main range, and pass north through East
Oregon into Utah and Idaho. One of these spurs,
called the Blue Mountains, shoots off the Cascades near
Mount Hood, and runs nearly parallel with the
Columbia, forming the eastern boundary of its valley,
and is about forty miles from the river, and terminates
about abreast of the mouth of Salmon River, Idaho,
and on the south side of Snake River. On the north
side of Snake River these mountains have local names,
but are known by the general name of Bitter Root
Mountains. . . . By tracing the course of Snake River
and its tributaries you will see it drains the northern
part of the great central basin, and when it cut its way
through these mountains it very likely drained the
sreat system of lakes that once covered a great part
of this central basin. Now the mountain ranges in
this portion, north-east, are the metropolis of strzgosa
so far as we know at present ; and z¢ zs not emprobable
that many individuals, and quite likely whole colontes,

144 THE DISPERSAL OF SHELLS.

of that species are sometimes carried into the streams by
rains and floods, and are borne away on the waters to-
wards the Pacific Coast. Occasionally some of the
specimens must find or make a lodgment along the
banks of the streams, and if the conditions are favourable
a colony will spring up and perhaps spread over the
neighbourhood. The banks of the Columbia between
the Dalles and the mouth of Snake River, a distance of
one hundred and fifty miles, are destitute of timber, and
are covered for several miles back with loose drifting
sand, quite unfavourable to the existence and spread of
land-shells. Zhe locality where I found the variety cas-
taneus was on the bank of the Columbia near Celilo, about
jifteen miles above the Dalles, on the cast side of the Cas-
cades, but on the west side of the Blue Mountains. This
colony must have sprung from specimens brought down the
stream by floods. At a subsequent visit it had dis-
appeared. ... Very likely the original stvzgosa |the
original specimen, so named, was found on or near the
Pacific Coast| may have come from some colony planted
in this way.”

Obviously, as was remarked by the Rev. H. H.
Higgins, animals can be carried by means of rivers to
lower localities only ; it is clear, of course, that lands
bordering on the higher waters of a river can never be

thus reached :!

it is almost equally clear, however, as
the elder Binney observed, that ‘a species having by
its own powers attained the summit of a range of moun-

H. H: Higgins, “ Proc. Lit. and Philos. Soc., Liverpool)” xxxvn
(1882), pp. xliv.-xlv.

MEANS OF DISPERSAL. 145

tains, may, when aided accidentally by the current of
rivers, be very rapidly diffused through the country on
the other side.” '

Rivers, with their occasional floods, it can hardly be
doubted, have largely helped to bring about the wide
distribution now enjoyed by many organisms, or groups
of organisms ; for ages, it will be remembered, they have
been constantly carrying objects of all kinds out to sea,
thus bringing them under the influence of the oceanic

currents.

WIND.

Mr. Wallace has suggested that the smaller land-
shells, and others in a young state, attached to leaves
and twigs, may be carried long distances by storms and
hurricanes, and doubtless their eggs, even those of large
kinds, may occasionally be transported by these means,’
but here again, as might be expected, nothing approach-
ing actual proof can be given. The hurricanes of
tropical regions, Sir C. Lyell observes, sweep along at
the rate of ninety miles an hour, rooting up trees and
throwing down buildings in their course, possibly
carrying, as he remarks, “even the heavier fruits and
seeds over friths and seas of considerable width,” * and

1 A. Binney, “ Terrestrial air-breathing Mollusks,” i. (1851), p.

131.
2 See “Geographical Distribution,” ii. p. 525; “Island Life,”
py 76) ed.2;-p.78:
* * Prnciples,” i. p- 392:

146 THE .DISPERSAT (OF. SHELLS.

it has been suggested ' that minute land-shells and other
animals may be carried, by similar means, across arms
of the sea.

Dead leaves, which are frequently caught up and
blown to distances, are often inhabited by small land-
shells: indeed, in order to procure specimens for the
cabinet, some collectors take home quantities of leaves,
from which, when dried, great numbers of shells are
sometimes sorted out; Dr. Turton told Mr. Jeffreys, for
instance, that he procured many specimens of Helix
pygmea by collecting a bagful of dead and rather moist
leaves and afterwards spreading them on paper to dry
On Oak Island, Dr. Gould found Cochlicopa lubrica in
such numbers that hundreds could be taken from the
cround with a single fallen leaf, and, as the moisture
evaporated, all, it is said, “disappeared beneath the
leaves.” * Only violent hurricanes, it will be remem-
bered, are likely to carry away moist and decaying
leaves, which are generally matted together and lie
heavily one upon another, and it may be doubted,
perhaps, whether shells are often very abundant among
loose dry leaves which alone are likely to be blown
about by wind-storms at ordinary times. In April, in
Epping Forest, I collected a large number of little shells
belonging to seven species—/Hyalinia fulva, H. pura

* “Our Earth and its Story,” iii. (Cassell: no date), p. 4o.
2 “ British Conchology,” i. (1862), pp. 223-4.

3 Dr. Gould, “Invertebrata of Massachusetts,” as quoted by
Reeve, ‘“ Land and fresh-water Mollusks,” 1863, p. 93, and see

Binney, “ Terrestrial air-breathing Mollusks,” i. (1851), 139.

MEANS OF DISPERSAL. 147

v. margaritacea, Helix aculeata, H. pygmea, Vertigo
substriata, V. edentula, and Carychium minimum—from
moist leaves, chiefly those of beech and holly ; but in
the same locality, in September, when the weather was
hot and dry, a careful examination of four handfuls of
leaves, which were quite dry and might have been
carried away by a strong wind, yielded only a solitary
Vertigo ; this, however, on being moistened, was found
to be alive.

In Germany, according to Mr. Daniel, the Helzxr acu-
leata, travelling high up into trees, particularly the alder,
reaches the ground in autumn on the falling leaves.
Flelix fusca, too, M. Bouchard-Chantereaux states
inhabits alders in the woods near Boulogne, hiding on
the undersides of the leaves, and falling with them in
September and October.' Snails with habits of this kind,
obviously, might be carried to considerable distances
when leaves, instead of falling directly to the ground,
happen to be blown off during autumn gales.

Many small snails are so light that they could cer-
tainly never impede the flight of leaves to which they
happened to be more or less securely attached ; twelve
“mature specimens” of Vertigo mzlium, for instance,
a species found gregariously under or among dead leaves,
and measuring only four-fifths of a millimetre in length,
are stated by Professor Adams to weigh less than six-
tenths of a grain.”

1 British Conchology,” i. (1862), p. 176: and p. 206.

* A. Binney, “ Terrestrial air-breathing Mollusks,” ii. (1851), p.

338,
i. 2

148 THE DISPERSAL OF SHELLS.

It is perhaps worth while in this connection to notice
an extraordinary fall of oak leaves, witnessed in 1889,
in Dumfriesshire, on a farm on which were only five
trees (two ash and three larch); the following account
of the occurrence, in the words of the observer, was
published in Mature in 1890:!

“IT was struck by a strange appearance in the at-
mosphere, which I at first mistook for a flock of birds,
but as I saw them falling to the earth my curiosity was
quickened. Fixing my eyes on one of the larger of them
and running about a hundred yards up the hill until
directly underneath, I awaited its arrival, when I found
it to be an oak leaf. Looking upwards the air was
thick with them, and as they descended in an almost
vertical direction, oscillating, and glittering in the sun-
shine, the spectacle was as beautiful as rare. The
wind was from the north, blowing a very gentle breeze,
and there were occasional showers of rain. On ex-
amination of the hills after the leaves had fallen, it was
found that they covered a tract of about a mile wide
and two miles long. The leaves were wholly those of the
oak. No oak trees grow in clumps together nearer than
eight miles. The aged shepherd, who has been on the
farm since 1826, never witnessed a similar occurrence.”

Mr. Musson, writing from Sydney, tells me that the
dry-weather molluscs of that region hide under twigs,
logs, and scraps of wood, estivating in many cases, so
that they are almost certainly transported with the tre-

1 J. Shaw, quoting a letter received from Mr. Wright, “ Nature,’’
xlii. (1890), p. 637.

MEANS OF DISPERSAL, 149

mendous dust storms, or “darling showers,’ which
sometimes blow over the country, carrying with them
twigs, leaves, dust, etc.; at times also, he says, more
frequently in dry séasons than in wet ones, dust-columns
are formed, which, travelling with a rotary motion,
whirl up to a height light articles of all kinds that may
happen to be within their reach. Mr. Belt, when in
Australia, saw scores of these dust-whirlwinds, many
rising to a height of over a hundred feet, and some
strong enough to tear off limbs of trees, and carry up
the tents of gold-diggers into the air. Many were
observed on a small plain near Maryborough, in Vic-
toria, where in calm sultry weather during the heat of
the day, two or more were often in action at once on
different parts of the plain; the dust and leaves carried
up rendered their spiral movement very conspicuous,
as, lasting several minutes, they slowly moved across
the plain like great pillars of smoke. From whirlwinds
of this sort, Mr. Belt remarks, there is a complete grada-
tion through larger whirlwinds and tornadoes to the
awful typhoons and cyclones of China and the West
Indies." As helping us to understand what may be
done by causes of the present kind, short accounts of
two rather powerful whirlwinds, one of which occurred
in'-our own islands, may be ‘given. Sir C. Lyell
writes :

“Dr. Franklin tells us, in one of his letters, that he
saw, in Maryland, a whirlwind which began by taking

* Thomas Belt, ‘ Naturalist in Nicaragua,” ed. 2, (1888), pp.
301-4.

150 THE: DISPERSAL OF SHELLS.

up the dust which lay in the road, in the form of a
sugar-loaf with the pointed end downwards, and soon
after grew to the height of forty or fifty feet, being
twenty or thirty in diameter. It advanced in a direc-
tion contrary to the wind; and although the rotatory
motion of the column was surprisingly rapid, its onward
progress was sufficiently slow to allow a man to keep
pace with it on foot. Franklin followed it on horse-
back, accompanied by his son, for three quarters of a
mile, and saw it enter a wood, where it twisted and
turned round large trees with surprising force. These
were carried up in a spiral line, and were seen flying in
the air, together with boughs and innumerable leaves,
which, from their height, appeared reduced to the size
of flies, 7+

The following account of a whirlwind in Ireland in
1872, was printed in Vaéure in that year:

“Ina letter to the Belfast News-Letter, Mr. C.J. Webb
descrites an extraordinary whirlwind which occurred in
the district around Randalstown, about six miles north-
west of Antrim, near the shores of Lough Neagh, on
the 25th of August last [1872]. The same phenomenon
was witnessed about an hour and a half earlier the
same evening at Banbridge, about seven miles south-
west of Dromore. It was first seen near Randalstown
about 5 p.m. between that place and Toome, moving
rapidly up Lough Neagh from the south, and presenting
the appearance of a defined column of spray and clouds,

1 Principles,” Wie p.- 302

MEANS OF DISPERSAL. I5!

whirling round and round, and not many yards in
breadth, while at its base the water was lashed into a
circle of white foam. It was next heard of in the
neighbourhood of Staffordstown, about a mile from the
lake, where it partially unroofed two houses, and
damaged any trees or crops which happened to be in its
course. From this point it travelled in a straight line
for Randalstown, about three miles distant. It passed
across a field close to Mr. Webb’s house, levelling eight
haystacks, and carried a considerable part of the hay up
into the air out of sight... . Everything it lapped up
was whirled round and round, and carried upwards in
the centre, while dense clouds seemed to be sucked
down on the outside, and came close to the earth... »
Mr. Webb next observed its track in a hollow, some
three hundred yards further on, where it knocked down
a haystack, and then plunged into a wood of fine old
Irish oaks. Here it tore numerous branches and limbs
from the trees, carrying some along with it, and throwing
others to the ground.... It next passed across a corner
of Shane’s Castle demesne. Some who were ata short
distance from this point describe its approach as causing
considerable alarm. . . . It crossed the valley over the
railway viaduct, close to Randalstown, fortunately
avoiding the village. It here presented the appearance
of a vast whirling column of leaves and_ branches,
mingled with clouds which looked like smoke. The
railway station next suffered, innumerable slates and
two and a half hundred-weight of lead being torn from
the roof in an instant. A great part of the railings

152 THE DISPERSAL OF SHELLS.

surrounding the gardens was torn up, and an iron bar,
one inch thick, belonging to the gate, was bent to an
angle of sixty degrees. A small shed at the rear of the
station was unroofed, rafters and slates being hurled to
the ground. What will give some idea of the excessive
pressure of the wind, is the fact that three boards of the
flooring of the waiting-room were forced up, owing to
the wind finding an entrance to a cellar underneath,
though the only aperture was a round hole about one
foot in diameter. All this was the work of a few
moments. The storm then passed away, leaving com-
parative calm behind. It next crossed an adjacent bog,
scattering the turf in all directions. The last place Mr.
Webb heard of its having visited was a farm house
about three miles from Randalstown, between Antrim
and Ballymena.”

As remarked by Sir C. Lyell, whirlwinds occur “at
different intervals of time throughout a great portion
of the earth’s surface,” and it seems reasonable to sup-
pose that many organisms may have been widely dis-
persed by them, as well as by the larger circular storms
which occur in certain parts of the world ; from time to
time these causes may carry plants, insects, and, as
Lyell says, ‘land testacea and their eggs, with many
other species of animals, to points which they could
never otherwise have reached, and from which they
may then begin to propagate themselves again as from
a new centre.”? Leaves and other vegetable debris,

+S" Nature,” vi. (1872), 1; and see Belt’s “ Nicaragua,’ ed.
2; (1888), p:-303. 2% Prmeciples;’ itsp.gge2:

MEANS OF DISPERSAL. Ee

when carried to great heights, Mr. Wallace observes,
may occasionally be drifted off by strong upper currents
and dropped at great distances, “and with them small
insects and mollusca or their eggs.” '

“On 27th July, 1875, a remarkable shower of small
pieces of hay occurred at Monkstown, near Dublin.
They appeared floating slowly down from a great
height, as if falling from a dark cloud which hung over-
head. The pieces picked up were wet, and varied from
single blades of grass to tufts weighing one or two
ounces. A similar shower occurred a few days earlier
in Denbighshire, and was observed to travel in a direc-
tion contrary to that of the wind in the lower atmo-
sphere.” ?

From what distances the hay had come in these
cases no one can tell; it was, however, in a_ position
to be conveyed, as Mr. Wallace remarks, to almost
any distance by a violent wind, had such occurred at the
time ; doubtless it was originally taken up by whirl-
winds, and this was the case also, in all probability, with
the leaves which fell in Dumfriesshire in 1889. Slight
whirlwinds, according to Lyell, may frequently be
observed in our fields, in summer, carrying up haycocks
into the air, and then letting fall small tufts of hay far
and wide over the country ;* the Irish whirlwind of
1872 as just noted carried a large quantity of hay into

= Island Life,” p. 274; edi 2, py 285.

* Wallace, “ Darwinism,” ed. 2, (1889), p. 362; quoting “ Nature,”
xli. (1875), pp. 279, 298.

a? Prnerples,” ii. Pp; 392:

154 THE DISPERSAL OF SHELLS.

the air out of sight, and that observed in Maryland
carried leaves to such a height that they appeared
reduced to the size of flies. It is clear enough, I think,
from these facts that molluscs—even large and heavy
kinds—particularly those living among vegetable débris,
dead leaves, and the like, may frequently be carried
from place to place during whirlwinds, and it has even
been suggested also that those inhabiting open plains
or pastures may similarly be caught up and conveyed.
The idea that snails—He/ex virgata, for instance —some-
times descend in showers, may have originated in some
cases, Mr. Jeffreys thinks, “ in a whirlwind having caught
up a number of them by sweeping along a grassy plain
and dropping its contents in a limited area.”

ANIMAL AGENCIES.

Animals of various sorts in all probability, as we
have seen, have been actively engaged in the dispersal
of fresh-water molluscs, and this remark holds good, no
doubt, for terrestrial kinds, Something, there is reason
to suppose, has been done even by insects. Under

1 “ British Conchology,” i. (1862), pp. 211-12. For a popular
allusion to wind-dispersal, see Dickens in the ‘‘ Chimes”: ‘* Toby
himself all aslant, and facing now in this direction, now in that,
would be so banged and buffeted, and touzled, and worried, and
hustled, and lifted off his feet, as to render it a state of things but
one degree removed from a positive miracle, that he wasn’t carried
up bodily into the air as a colony of frogs or szaz/s or other port-
able creatures sometimes are, and rained down again to the great
astonishment of the natives, on some strange corner of the world
where ticket-porters are unknown.”

MEANS OF DISPERSAL. Bes

some loose stones near Rockhampton, Queensland,
Mr. Musson recently found a beetle carrying a land-
snail of the genus Vztrina (or Helicarion) upon one of
its wing-cases; and it would not require a very great
stretch of imagination, as he observes, “to consider
that, could the insect have taken flight with this strange
companion as passenger, it might have been the means
by which distribution would have been aided, and thus
a new colony be started where possibly the species had
been before unknown.”! Some years ago, at Lambley
Dumbles, near Nottingham, the same observer saw one
of our common little snails, Helix rotundata, riding
about on the back of a woodlouse, and it is perhaps
worth mentioning that on another occasion, at Clifton,
Bristol, he saw a full-grown chrysalis-shell (Papa wmbilt-
cata), not yet aroused from torpor, securely fastened to
a specimen of Helix virgata—the zoned-snail—which
was crawling out after a shower; these facts, however,
are chiefly interesting for their curiosity. Most kinds
of animals, no doubt, have helped in the work in
some way or other; mammals, for instance, some of
which habitually feed on molluscs, often carrying the
creatures for short distances, must sometimes let them
drop by accident, or desert them on being suddenly at-
tacked or frightened. A rat, after running some paces
from home, was once observed to “climb the stalk of a
hollyhock, clear off several snails, bring them down inone
paw like an armful, and run with them on three legs into
his hole,” the inside of which was strewn for some distance

1 C.T. Musson, “ Proc. Lin. Soc., N.S.W.,” (2), iv. (1889), 388.

156 THE DISPERSAL OF SHELLS.

with broken shells... Miss Warren has suggested to me
that small shells, such as those living at the roots of
grass, May sometimes stick to the muddy clay on the
feet of cattle, and it seems probable that the range of
a species within a given country may occasionally be
extended in this way. But of all animals, with the
exception perhaps of man (who has done a great deal
in recent times) the ranges of land-shells have certainly
been most affected by birds. The waders, it will be
remembered, as Mr. Darwin remarked,’ are great
wanderers being “ occasionally found on the most remote
and barren islands of the open ocean ;” many sorts of
birds, it is hardly necessary to repeat, annually migrate
across considerable tracts both of land and sea, and
most kinds, as every one knows, are liable occasionally
to be blown by violent gales of wind to great distances
over the ocean; almost every year, for instance, Mr.
Darwin states, one or two land-birds are blown across
the whole Atlantic, from North America to the shores
of the British Isles, and many, to give another instance,
even the smaller land kinds, are constantly blown
from Europe to the Azores, a distance of nearly a
thousand miles; indeed, according to Mr. Godman,
scarcely a storm occurs there in spring or autumn with-
out bringing one or more species foreign to the islands.”

1 W. W. Attree, in Merrifield’s “Sketch of the Nat. Hist. of
Brighton,” p. 157, as quoted in the “ Zoologist,” (3), il. (1878), 9o.

2. ¥ Origin; ps 345.

3 Origin,” pp. 326, 328-9; “Island Life,” p. 73, ed. 2, p. 75;
‘Principles, ii. p. 368; F. Du Cane Godman, “ Ibis,” (n.s.), il.
(1866), p. 105.

MEANS OF DISPERSAL. 157

Sir C. Lyell, remarking on the wide range of
Succinea putris, a \and-shell which inhabits moist places
on the borders of pools and streams, suggested that
water-fowl might have distributed its ova entangled
among their feathers,’ and it seems quite likely that ova
of certain terrestrial kinds may be occasionally thus
carried, either in the feathers or on the feet of birds;
indeed, we have a near approach to proof of such
transportal, the Rev. Canon Tristram, as we have seen,
having once found ova, believed to be those of a Szccznea,
upon one of the feet of a mallard shot by him, on the
wing, in the desert of Sahara.’ It is doubtful, however,
whether Szccinee, from the nature of the localities they
often or usually inhabit, ought not, for the present pur-
pose, to be classed with fresh-water, rather than with
land-shells. Mr. Darwin suggested that the just-
hatched young, possibly, might sometimes crawl upon
the feet of ground-roosting birds, “and thus get trans-
ported ;”* and it certainly seems in the highest degree
probable that such is the case, but, as far as I know,
no observations in support of such a supposition have
yet been made. In one of his letters to Wallace,
Darwin mentions having been interrupted when begin-
ning to experiment on the point.* As was mentioned
in chapter ii., small quantities of earth are occasionally

1 « Principles,” ii. p. 377.

2 H. B. Tristram, “ Zoologist,” (3), i. (1877), pp. 260-1.

3 “Origin,” p. 353; and see “ Geographical Distribution,” i. pp.
31:2.

4 “ Life and Letters,” iil. (1888), p. 231.

158 THE DISPERSAL ‘OF ‘SHELES,

found upon birds’ feet, and this, perhaps, may some-
times cause small shells or eggs to adhere, or the crea-
tures may possibly become embedded in it. In one
case Mr. Darwin removed sixty-one grains, and in
another case twenty-two grains, of dry, argillaceous
earth from the foot of a partridge, and in the earth, as
already mentioned, was a pebble as large as a vetch-
seed.! As previously noted, also, a woodcock’s leg,
once received by Mr. Darwin from a friend, had a little
cake of earth weighing nine grains attached ; and a
pigeon, shot in New South Wales in 1887, was found to
have a small ball of earth surrounding each of its legs ;
suddenly-flushed birds, I believe, are not unfrequently
found to have mud or clay adhering to their feet. In
the Field, in 1881, Mr. Tegetmeier figured the remains
of a young partridge, found dead by a correspondent,
destroyed, obviously, owing to its inability to drag about
the mass of clay which had become attached to one of
its feet. The claws, it seemed, had been first encrusted,
layer after layer having been subsequently added until a
mass of considerable size was formed. In poultry yards
and pigeon houses similar occurrences, it is said—never
allowed to become extreme —are frequently observed.*
Professor Newton once sent to Mr. Darwin “the leg of a
red-legged partridge (Caccabis rufa) which had been
wounded and could not fly, with a ball of hard earth ad-
hering to it, and weighing six and a half ounces.” Mr.
Swaysland, as is more important for us to notice, several

i “Origin, p. 320.
W, B. Tegetmeier, “ Field,” lviii. (1881), p. 330.

MEANS OF DISPERSAL. 159

times found little cakes of earth adhering to the feet of
wagtails, wheatears, or whinchats, shot, as he assurediMr.
Darwin, on their first arrival on our shores, and before
they had alighted.’ Mr. Wallace, we find, dealing with
the origin of the land-shell fauna of the Azores, ex-
presses the opinion that some kinds may have reached
the islands “ with the earth that often sticks to the feet
of birds,”* and every one must admit that transportal
in this way is at least possible.

Another possible mode of transport, already hinted
at, is suggested by the following passage, from pages
326-7 of the “ Origin,” referring specially to the dis-
persal of seeds :-—

“The crops of birds do not secrete gastric juice, and
do not, as I_ know by trial, injure in the least the ger-
mination of seeds; now, after a bird has found and
devoured a large supply of food, it is positively asserted
that all the grains do not pass into the gizzard for
twelve or even eighteen hours. A bird in this interval
might easily be blown to the distance of 500 miles, and
hawks are known to look out for tired birds, and the con-
tents of their torn crops might thus readily get scattered.”

That the “sudden deaths to which great numbers of
frugivorous birds are annually exposed must not be
omitted as auxiliary to the transportation of seeds to
new habitations,” had already been remarked by Sir C.
Lyell, who suggested that birds with recently picked up
seeds still in their crops would now and then chance to

** Origin,” ps 328:

& Sisland Lite.” p,247..ed, 2, pr 256,

160 THE DISPERSAL OF SHELLS.

be seized by hawks or eagles, which, “ soaring across
hill and dale to a place of retreat,” might, after devour-
ing their prey, leave the seeds to spring up and flourish
in a new soil.! Now,as birds of many kinds are known
to feed largely upon snails, which, no doubt, often re-
main alive for some hours in their crops, it seems almost
certain that these causes may have operated, very
rarely, perhaps, for the dispersal of land molluscs ; and
even fresh-water kinds, though many are thin-shelled,
may also have been thus transplanted, but much less
frequently, for the restoration of an animal in this man-
ner to a fresh-water habitat is sure to be extremely rare
and exceptional. Mr. Cordeaux tells me that after a day’s
shooting he has frequently opened the crops of birds and
found them to contain both land and fresh-water snails,
but, like several other ornithologists who have informed
me of similar observations, he is unable to say positively
whether any of the creatures were alive. Twenty
specimens of a Swccinea, peculiarly packed together,
and four of Pupa muscorum were once found by Mr. W.
H. Dikes in the crop of a bearded titmouse (Parus
biarmicus) ; all the shells, it is said, were uninjured, but
it is not stated that any were observed to be alive.’
Many birds, it ought, perhaps, to be mentioned, are
careful to kill the animals upon which they feed ; thus,
for instance, Mr. O. V. Aplin has seen a tame magpie
breaking the shells of rufous-snails (felzx rufescens) —

1 “ Principles,” ed. 9, (1853), pp. 624-5.
2 W. H. Dikes, Loudon’s “ Mag. Nat. Hist.,” ili. (1830), p. 239 ;
and see D. Cooper, Charlesworth’s (n.s.), ii. (1838), p. 479.

MEANS OF DISPERSAL. I6I

of which it was very fond—before eating them. But
this habit is by no means common to all birds ; some
kinds are actually known to swallow living snails, and
Mr. Cordeaux has favoured me with his opinion that in
all probability the creatures frequently keep alive for
some little time after being swallowed ; and on this
point, fortunately, an observation of much interest has
been recorded, the accuracy of which, Mr. Roberts, of
Lofthouse, assures me, cannot be doubted, In Septem-
ber, 1875, it appears, Mr. John Ward, carpenter and
bird-stuffer, of Lofthouse, took thirteen wrinkled-snails
(Helix caperata), together with a quantity of tares, from
the stomach of a wood pigeon which had been shot
three days previously. Most of the snails were alive,
and “began creeping about on being placed in a
dish containing a little water.”’ It may be mentioned,
as a somewhat analogous case, that Mr. Abel Chap-
man has frequently shot curlews stuffed with live
cockles, which, as he remarks, might easily have been
scattered if the birds had chanced to have been
killed and carried off by peregrines or other birds of
prey. Molluscs thus remaining alive in birds’ crops,

1 Mr. Roberts informed me of this fact, without comment, in
1890, but recently, after drafting this chapter, I was much pleased
to find that when recording it, in 1882, he had suggested that
molluscs might possibly be “carried in the crops of birds con-
siderable distances, and thus be distributed and established in
new districts, or on islands, as the living shells might be ejected
from the crop, or the birds might be killed by birds of prey and
the contents of the stomach dislodged and scattered.”—‘“* Topo-
graphy and Nat. Hist. of Lofthouse,” 1882, p. 333.

M

162 THE DISPERSAL OF SHELLS.

as Mr. Darwin’s remarks suggest, may possibly be
transported, especially during gales, under exceptionally
favourable circumstances, to new homes perhaps several
hundreds of miles from their original habitats. At
ordinary times birds often fly a long way to feed—wood
pigeons, for instance, do so when necessary—and_ if
struck down by birds of prey when returning home, the
living contents of their crops might possibly be liberated
at considerable distances from the feeding-ground.
There is no actual proof, as far as I know, that shells
have ever been scattered from crops in the manner here
indicated, but an observation of much interest in this
connection has been made by Mr. Aplin, who tells me
that he once noticed a little heap of barley, evidently
carried from a distance, lying among the remains of two
wood pigeons which had been killed by a hawk, pro-
bably a peregrine. Even when a snail-eating bird hap-
pens to be killed on its feeding-ground, the bird of prey,
“soaring across hill and dale” to a hiding-place, and
carrying its victim with it, might ultimately scatter a
few snails miles from their original home; it may be
remarked, however,as Mr, Aplin points out, that except
in the breeding season when they have young, hawks,
etc., often or usually eat their kills on the spot, or close
to it; but it is undoubtedly probable, as Mr. Cordeaux
assures me, that both seeds and shells have been dis-
tributed, to some extent, by hawks, owls, etc. sometimes
devouring their victims at a distance from where they
were struck down. It seems possible, also, that snails
in the crops of dead birds may be floated, at rare

MEANS OF DISPERSAL, 163

intervals, on the surface of the ocean, to be ultimately
liberated, perhaps, on some distant shore. Large num-
bers of land-birds, I suppose, annually perish in the sea ;
their floating carcases, Darwin states, sometimes escape
being immediately devoured,’ and, as shown by Mr.
Ward’s observation, a He/zx can retain life in the crop
of a dead bird at least for three days. In such circum-
stances, of course, molluscs would be protected to some
extent from the evil influences of salt water, and, in
illustration of this point, it may be noted that a number
of seeds of peas and vetches (which are killed by a few
days’ immersion in sea-water) nearly all germinated
after having been enclosed in the crop of a pigeon
which had floated on artificial sea-water for thirty
days.”

Various kinds of birds, it will be remembered, have
been observed to eject the contents of their crops when
frightened or wounded, as gulls and terns do when
pursued by the skuas, and it seems quite possible, as
Mr. Roberts has already suggested, that snails, still
alive, may occasionally be thus disgorged, and set down
in places perhaps some considerable distance from the
spot where they were picked up and swallowed, Mr.
Clement Reid suggests that a good deal may have been
done, also, by birds gorging themselves after a long and
wearying migration, and afterwards casting up the con-
tents of their stomachs, little injured, and perhaps a con-
siderable distance away, and he states that he has

Origin,” p: 320: a Oniott,” p.. 320.
M 2

164 THE DISPERSAL OF SHELLS,

several times come across castings of this kind, con-
taining soft seeds, live worms, etc.

It must be borne in mind, of course, that the
transportal of shells by these agencies to distant
points over the ocean can only very rarely happen, but,
as Mr. Wallace has remarked, “nature can afford to wait,”
and if but once in a thousand years, he adds, “a single
bird should convey two or three minute snails to a
distant island, this is all that is required for us to find
that island well stocked with a great and varied

”]

population of land-shells. We must remember, how-
ever, that resulting colonization is much more rare than
mere transportal, though, of course, snails carried by
birds into the interior of new countries or islands, or to
some little distance inland, are more likely to be able to
establish themselves than are those stranded upon the
shores with drift-timber, etc. The creatures are carried,
we may feel sure, for comparatively short distances
within given tracts of land much more frequently, and
many apparent anomalies in local distribution no doubt
have thus arisen, but even here the ultimate establish-
ment of a new colony seems likely to be a rare event.
Additional ways in which short migrations may possibly
be brought about might be referred to. Small shells, such
as those of the genus Vertigo, and eggs of various kinds
must occasionally happen to be adhering to twigs, etc.,
carried off by birds when building, and these are some-
times collected at some little distance from the nest ;
every one, no doubt, remembers having seen rooks

' “ Geographical Distribution,” i. p. 32.

MEANS OF DISPERSAL. 165

flying overhead with twigs in their mouths. A living
specimen of Lzmuea truncatula —the “ ditch mud-shell ”
of Gray’s Turton—once found, as the reader will
remember, on the top of a church in Preston, had very
probably been carried there, as Mr. Heathcote suggests,
adhering to one of the sticks taken up by a jackdaw
building in the tower. Thrushes, red-wings, etc.—great
snail-eaters—as Mr. Cordeaux remarks to me, will carry
shells to considerable distances, and from being disturbed
or otherwise may sometimes drop them, or neglect
to break them against their favourite stones, and Mr,
Roberts, I hear, has actually seen a thrush drop a snail
while flying. Quite recently, I found—on a snow-
covered road in Lincolnshire—a hibernating snail,
Felix aspersa, which, having a pierced epiphragm, had
almost certainly been carried and dropped by a bird.
No stones on which shells had been broken were near,
but several were seen at some distance along the same
road. On being placed in tepid water the snail revived,
and crawled away.

Operculate land-shells, it seems probable, may
occasionally be carried in the manner already suggested
for operculate pond-snails in chapter iv., namcly by
closure of the operculum, so as to hold on to the toes of
insects, etc. As there mentioned, a land-shell has
been seen holding on to a humble-bee in this way.
The observation referred to was made in June,
fao5, and recorded in the /ze/d in that year." The

" F.W.T., “ Humble-bee trapped by snail,” “ Field,” Ixv. (1885),
p. 543.

166 THE DISPERSAL OF SHELLS.

observer, it appears, while walking along by the road-
side, saw a large orange-tailed humble-bee, “ vigorously
towing something heavy after him,” which, on inspection,
proved to be “a Clausilia [Cyclostoma ?] the little snail
which has an operculum-like plate on the lip;” the
latter had fast hold of one of the hind legs of the bee,
by which, no doubt, it had been accidentally trodden
upon. The smaller operculate-shells, it can hardly be
doubted, may possibly be carried in this manner to
considerable distances by the large and _ powerful
insects which abound in some parts of the world.

CHAPTER. VII.
DISPERSAL OF SLUGS.

SLUGS, obviously, are not so well fitted for dispersal as
snails, for many are quite naked, and though some
possess small external shells, the well-developed snail-
shell, into which the animal can retire for rest and
during periods of adversity, and which, often closable
by an epiphragm or operculum, has doubtless largely
facilitated the dispersal of shell-bearing kinds, is always
absent, and it is clear, therefore, that many of the con-
siderations referred to in the preceding chapter cannot
be looked upon as necessarily applicable to the slugs,
such creatures being sure to succumb to many of the
hardships from which snails may often have escaped in
safety. Many slugs, it is notorious, from containing
much water, cannot even bear exposure in a dry at-
mosphere for any length of time, but in many respects,
it should be remembered, animals of this kind are much
more tenacious of life than might at first be supposed,
and this is true of our ordinary absolutely naked sorts,
Limax, Arion, etc. The Testacell@, which have a small
ear-shaped shell near the extremity of their bodies,
“ snail-slugs ” of some authors, are able to protect them-

168 THE DISPERSAL OF SHELLS.

selves by forming a kind of cocoon “secreted from their
skin and often mixed with earthy and extraneous par-
ticles”: the “nest of earth” made by 7. maugez in
times of drought, it is said, reminds the observer of the
cocoon of the puss-moth; within, the animal lies
encysted until moisture, finding its way through the
walls, rouses it again to activity. While in the encysted
state “a thin white membrane (a development of the
mantle) is extended from beneath the shell and stretched
over the back and sides of the animal,” checking evapora-
tion, and forming an admirably-designed protective
shield.’ The Testacella’s power of retaining life under
adverse conditions is well illustrated by observations
made by Professor Poulton in 1886 :—

‘Between four and five months ago I found eleven
specimens of this slug | Zestacel/a, species not stated]
upon a low wall surrounding the garden of a house
near the Oxford University Parks, and on the following
day I captured eleven more in the same place.... A
few of the twenty-two specimens were killed and
hardened, and the remainder were put in a box con-
taining earth, in which they buried themselves. In the
press of other work the box was neglected, and re-
mained untouched in my laboratory until to-day
[October 19th], the earth having quickly dried into a
hard cake. To-day I emptied the box, and fully ex-
pected to find the slugs dried up dead, but to my
surprise I found twelve specimens alive, each encysted

" ** British Conchology,” i. (1862), pp. 143, 147-8.

DISPERSAL OF SLUGS. 169

in a thin transparent capsule formed of the hardened
mucous secretion of the animal’s skin. The body was
contracted, and oval in shape, but it had been so
completely protected from evaporation that there was
no noticeable reduction in bulk after these hottest
months of the year, during which water had been
entirely withheld. One or two specimens had died
almost immediately after capture, and a few escaped,
so that all those which had been exposed to the heat
and dryness in the box had become encysted, and sur-
vived in apparent health.” *

Slugs in such a condition at the roots of trees, etc.,
would be nearly as well fitted for dispersal as snails, at
least by fresh-water currents.

Slugs of most sorts and the eggs of some kinds must
certainly be carried to short distances within given land-
areas with some frequency ; various means, organic and
inorganic—probably of kinds already suggested—operate
no doubt to bring about short involuntary migrations.
River agencies must have widely scattered the creatures.
Some are arboreal, inhabiting old and decaying trees
which, when blown to ground, or washed out of river-
banks, must sometimes be carried off by floods. In our
own country we find slugs—the great grey kind, Lzmax
maximus, for instance—living under the bark of old
willows by river-sides. At times, too, we find them well
concealed in chinks and among débris in the interior of
hollow trees, and, like some snails, they probably eat

1 E. B. Poulton, “ Nature,” xxxiv. (1886), p. 618.

170 THE DISPERSAL OF SHELLS.

their way into the decaying wood, for several specimens
of ZL. maximus, found when removing rubbish and
breaking away parts of the wood from a cavity in the
trunk of an old hornbeam, subsequently extruded pale
yellow pellets composed entirely of “saw-dust.” In
these situations the creatures would often be well pro-
tected, so that they might be carried in safety with
floating trunks to great distances. Many kinds, it must
not be forgotten, live mostly on the ground, rarely or
never ascending trees, and these of course are not likely
to be carried with timber, unless they occasionally hide
themselves in logs lying upon the ground; they must
frequently be swept in numbers from their haunts on
grassy or muddy river-margins, but unless buoyed up by
some object, they are not likely to be much dispersed by
the currents, for I believe they always sink in water, and
when carried away by the rushing waters of torrents or
floods are hardly likely to be often landed again in a
living state. It seems probable, however, that they
may sometimes be safely floated upon vegetable-rafts
of many sorts. A few years ago, indeed, I had the
good fortune to witness the conveyance of a slug by
this means, upon a very fragile raft. Walking in the
evening from the Lincoinshire coast towards Louth
along the banks of the Eau, I saw a large black slug
(Arion ater) crawling with out-stretched tentacles upon
a mass of ‘ green-scum ” which came drifting down with
the current at a good way from the bank. This was at
no great distance from the sea, and in all probability the
animal was ultimately carried out and drowned, but pos-

DISPERSAL OF SLUGS. ri

sibly it might be landed on one of the banks or arrested
by some projecting object, so as to be able to crawl away
in safety. Previously I had noticed that the level of the
water had risen rapidly, and the scum, lying, no doubt,
earlier in the evening stranded upon the mud at the
water’s edge, had evidently be enfloated off with its living
burden by the “flood.’ Slugs of the same_ kind
were numerous along the muddy margins for a consider-
able distance. The smaller ground-slugs, no doubt,
like little shells of many kinds, often resort to the
hollow “ kexes” of umbellifers, which, as already men-
tioned, seem likely to serve as vehicles for transportal :
the marsh-slug (Z. /@vis) has been noticed in the stems
of these plants by Mr. B. Hudson,' and I have seen the
field-slug (ZL. agrestis) in such stems lying upon the
ground near a little water-course in Highgate Woods.
Kinds which habitually burrow into the ground and
spend a great part of their lives beneath the surface
seem likely, of course, to be occasionally transported,
at least in some regions, in the soil attached to the
roots of floating trees, but they are sure to escape, in a
sreat measure, the various accidental causes which
above ground bring about comparatively frequent in-
voluntary migrations; thus we find Dr. Simroth re-
garding the Zestacelle as having but little scope for
dispersal owing to their subterranean habits, and as
a consequence, he says, they have divided up into local
forms.”

1 “ Journ, of Conch.,” v. (1886), p. 48.
2-H. Simroth, ‘‘ Journ. of Conch.,” vi. (1891), D. 423.

E72 THE DISPERSAL OF SHELLS.

It cannot be supposed that slugs often cross the
sea, and trans-oceanic migration for long distances,
if ever occurring, is likely to be extremely rare. Dr.
Scharff has stated, indeed, that the sea, ‘ which
is the principal means of communication for other
animals, and plants, between mainland and island, forms
an almost impassable barrier for slugs, sea water being
deadly both to their eggs and themselves ;” I do not
think it necessarily safe, however, to assume, as that
author does, that “if we find the slugs of mainland and
island agreeing in anatomical characters, we may
generally conclude that the island must have had a land
connection with the mainland at some time or other,” !
for this seems to imply that the creatures are in-
capable of crossing the ocean, and it is a fact, as-J
hear from Dr. Scharff, that their chances of surviving
a sea passage are regarded by him as practically nil.
But what do we actually know of the ways in which the
various organisms are or have been distributed over the
globe? Unthought-of means, even for the dispersal of
sluys, may be in operation, or may have operated in
former times. I have not ascertained whether any of
the true oceanic islands, independently of man’s dispersal,
possessed slugs belonging to continental species or
genera, but such I presume was the case ; some kinds
certainly seem to enjoy fairly wide ranges, but in many
cases I suppose it would be difficult to ascertain to what
extent man has affected their distribution. Few land

! R. F. Scharff, “ Sci. Trans. Royal Dublin Soc.,” (2), iv. (1891),

p: 513.

DISPERSAL OF SLUGS. r73

molluscs, it is said, have attained so wide a range as our
own little marsh-slug (Limax levis), which, “under
different names by various authors,” has been recorded
from Europe, North and. South America, the West
Indies, Madagascar, Australia, New Zealand, and many
islands of the Pacific.’ Possibly or probably it has
been carried to some of these regions by man, but it is
interesting to find that in Australia, for instance, where
it occurs “in very out-of-the-way places, and far away
from the coastal cities,” it is regarded as truly indigenous.”
Very probably, as it appears to me, these creatures are
able in some way or other to journey over the sea—no
doubt rarely—for short distances at least, and it seems
possible also that accidents leading to transit over the
ocean for very considerable distances may have occurred
now and then in the course of the vast periods
during which these naked genera have inhabited the
earth, but we are unable, it is true, to refer to any
actually observed occurrence that can reasonably be
said to lend direct support to this view. The creatures’
eggs, it will be remembered, do not differ essentially
from the eggs of many snails. Those of certain kinds
are deposited in the trunks of hollow trees ; thus, for
instance, Dr. Scharff mentions having found ova of the
yellow-slug (Lzmar flavus) “in-an old tree trunk, near
Dublin,’ * and those of the tree-slug (LZ. arborum), I

€. Hedley, Proc. Lin. Soc., N:S:W.,” December, 1890,.as
quoted in the “ Nautilus,” v. (1891), 12.

2 C. T. Musson, “ Proc. Lin. Soc., N.S.W.,”-for 1890, (2), v. (1891),
p. 885.

3 “ Scj. Trans. Royal Dublin Soc.,” (2), iv. (1891), p. 522.

174 THE. DISPERSAL OF SHELLS,

suppose, are usually placed either in the hollows
or under the bark of trees. In such _ situations,
protected, or partially protected, for a time from con-
tact with sea water, ova might possibly be carried
with floating trees over small arms of the _ sea,
and perhaps even to islands, though rarely if ever to
very remote ones. The animals themselves thus hiding
might also be carried, I think, in a similar way. It
is just possible, also, that eggs may be occasionally
carried over the sea, at least for short distances, in the
cavities of pumice, or in earth at the roots of trees.
Large numbers, both adults and ova, must certainly be
carried out to sea by large tropical rivers with floating
rafts or islands, which have already been referred to, and
if such objects have ever been stranded upon more or
less distant shores so as to permit their inhabitants to
be “ poured out as from an ark,” which was regarded as
possible by Lyell,’ any number of slugs may have
been thus transported. Hurricanes and whirlwinds
may have carried the creatures and their eggs (some-
times laid amongst dead leaves) over straits and small
arms of the sea, and the just-hatched young are as
likely as those of snails to crawl upon the feet of
cround-roosting birds, and thus fosszbly they may be
carried over sea, provided, of course, that they adhered
firmly ; this may be the case at a very early age, but
later, I find, and more especially when young than
adult, the creatures often let themselves drop purposely,

" See “ Principles,” ii. p. 366.

DISPERSAL OF SLUGS. 175

either with or without a slime-thread, from small
objects upon which they may be placed, and young
slugs, it should not be overlooked, soon perish from
exposure, so that after a little time, even if they ad-
hered, they would be sure to dry up and die. Dr.
Scharff raises a further difficulty; supposing that a
couple of young slugs, he writes, happened to stick to
the feet of a bird and were carried a long distance
without being shaken or blown off—which in itself he
regards as unlikely enough—on being safely deposited
on the ground they would “ still have to keep together
until mature, and not lose sight of one another in order
to propagate their species,’ The difficulties constantly
attending the establishment of new colonies as the result
of all occasional means of dispersal, it must of course be
admitted, are very great, and this fact, as already
insisted, must never be lost sight of, but in connection
with the present point it may be useful to remember
that many kinds of birds migrate in immense numbers
year after year along the same lines. To give an
imaginary case: hundreds of migratory birds, let us
suppose, roost together among tufts of herbage in a
swamp, where slugs, hatching out from the egg, and
abounding on al! sides, adhere in numbers to their feet ;
proceeding on migration, after a few hours, over an arm of
the sea, they might deposit onthe opposite shore a number
of individuals, which, joining others similarly transported
in former years, might become firmly established in the
new home; or losing their way in a fog or otherwise,
the birds might chance to alight altogether on some

176 THE DISPERSAL OF SHELLS.

lone islet, at no great distance, all the moist surfaces of
which, previously uninhabited by slugs perhaps, might
possibly be found, after a few centuries, as some traveller
might record, swarming with a kind common to the
mainland. We are bound to conclude, however, I think,
that in all probability dispersal over sea, with subse-
quent colonization, even to short distances, is very rare,
and such dispersal over wide stretches, though perhaps
not impossible, can hardly be supposed to have
happened, by any known means, more than once or
twice in the course of vast ages. One other point
deserves attention. Slugs are known to be largely
eaten by many birds; we find, for instance, that young
chickens and ducks are sometimes employed with good
results for the purpose of keeping the creatures within
bounds in kitchen gardens ; when the former are allowed
the free range of a garden they may be seen, it is
said, at daybreak, regularly searching rows of cabbages
and broccoli, picking up slugs, etc.; plovers and gulls
also, thus employed, are stated to be very useful,
and a tame petrel is said to have proved valuable for
the same purpose.’ If the creatures can survive in
the crop for a few hours, it would certainly seem that
they may be transported, as already suggested for
snails, for short distances, and if they can survive say
for twelve or eighteen hours, they may be carried over
sea, in exceptional circumstances, according to an

1 « Garden,” x. (1876), p..435 3 xill. (1878), pps275, 3045. 350 ave
(1879), p. 391; “ Gardeners’ Chronicle,” 1878, p. 664; “Science
Gossip,” 1868, p. 46; etc.

DISPERSAL OF SLUGS. ya

estimate of Mr. Darwin’s, even for a distance of 500
miles, and remaining alive in the crops of dead birds,
they might be drifted on the surface of the ocean
possibly to a still greater distance. The late Mr.
J. W. G. Spicer, of Spye Park, Wilts, once recorded
the shooting of pheasants in the Holt Forest with their
crops completely full of a small white slug, but he gave
no indication as to whether any were alive,’ and perhaps
it is hardly likely that these shell-less creatures can
retain life for any considerable time in the warm crop
of a bird, and it is to be remarked, also, that they are
certainly more likely than snails to be killed or severely
injured when picked up and swallowed.

1 J. W. G. Spicer, ‘‘ Zoologist,” (3), v. (1881), p. 383.

CHAPTER Viti.

DISPERSAL OF FRESH-WATER AND LAND MOLLUSCA
BY MAN.

IN comparatively recent times many molluscs have
been widely scattered over the giobe by man. Indéed
it may be safely said that both snails and slugs, during
the last three centuries at least, have been more
dispersed by human agency than by all other causes
together. European species, for instance, thus dis-
seminated, as everyone knows, have become thoroughly
naturalized in many parts of the world, even in the
most remote colonies.’ Many facts illustrating the
extent to which certain faunas have been alloyed by
man might be given. All the species of Limar
described as native to Australasia are referable, accord-
ing to Mr. Hedley, either to Z. maximus, flavus,
gagates, agrestis, or l@vts, all, except the last, believed to
have been introduced by man from Europe.? The
Madeiran Islands, to give another instance, discovered

+ See Woodward’s “ Manual,” ed. 4, rep. 1890, p. 289.

2 C. Hedley, “ Ann. and Mag. Nat. Hist.,” (6), 1x. (1802), p.7om
C.T. Musson, “ Proc. ‘Lin. Soc., N.S.W.” for 1890 (2), “2 (aicemn
p. 885.

DISPERSAL BY MAN. 179

in 1418, sea-girt, and less than 400 square miles in
area, possess, according to the Rev. Dr. Watson, no less
than thirteen kinds of molluscs (six slugs and seven
land and fresh-water shells), believed to have been
introduced by man, and now well-naturalized, namely :—
Arion ater, Limax gagates, L. maximus, L. flavus, L.
agrestis, Testacella mauget, Hyalinta cellaria, Helix
piulchella, Bulimus decollatus, B. ventricosus, Cochlicopa
lubrica, Limnea acuta, and “ Auncylus striatus Q. and
G.” ; and there are also six other similarly introduced
kinds— Testacella haliotidea, Helix aspersa, H. rotundata,
Planorbis glaber, Physa acuta,and Hydrobia similis—
much more recently imported, and hardly to be
regarded as truly naturalized.’ Snails thus introduced,
as itis interesting to note, often increase with surprising
rapidity ; it is notoriously dangerous, indeed, to turn
out any creature ina new country, where its enemies are
possibly absent, and the truth of this has been sorrowfully
enforced in some of our own colonies. The European
Hyalinia cellaria, it is said, occurs literally in hundreds
in the space of a few square feet near a water-fall in St.
Helena; the British Cemetery at Buenos Ayres,
according to a recent report, is “overloaded with
Helix pomatia; ” our common garden snail (Helix
aspersa), introduced into the Cape apparently within the
memory of a living naturalist, is there astonishingly
prolific; Mr. J. S. Gibbons, in 1878, mentioned that he
had nowhere seen it in such plenty as near Cape

R. B. Watson, “ Journ of Conch.,” vii. (1892), pp. I-3.
WN: -Z

180 THE DISPERSAL OF SHELLS.

Town ;' in Australia, too, as Mr. Hedley tells me, when
it once obtains a footing it increases at an enormous
rate, until there are far more shells to the square yard
than are commonly seen in Europe ; in the Botanical
Gardens of Sydney, for instance, it absolutely swarms.
Some hundred living specimens of Helzx nemoralzs,
imported from England by Mr. W. G. Binney in 1857,
increased with great rapidity in his garden in Burling-
ton, New Jersey, and by 1878 the whole town was said
to be full of them !? At Lexington, Virginia, also, this
snail, first noticed in 1886, probably introduced
accidentally, appears to have multiplied at a wonderful
rate ; about 1889, Professor J. H. Morrison, who had
transplanted the creatures to several fresh places, is
said to have collected over four hundred specimens in
about an hour’s time within a radius of twenty-five
yards.* Slugs in similar circumstances sometimes
increase with equal rapidity: in Victoria, in 1849,
Mr. John Carson states, a slug found in his garden was
welcomed as an old friend and carefully restored to its
feeding-ground, but by the following season “ matters
had changed” and it was every night’s work to pick up
“pints full” ; the warm, moist autumn and mild winter
of the region proved specially congenial, and the
creatures became a plague, unequalled, it is said, by

" J. S. Gibbons, M.B., “Quart. Journ. Conch:,” i: p:/367 Wate.
Rush, “ Nautilus,” vi. (1892), p. 81.

> W. G. Binney, “ Terrestrial air-breathing Mollusks,” v. (1878),
P. 379.

3H. A. Pilsbry, “ Nautilus,” iii. (1889), pp. 51-2.

DISPERSAL BY MAN. ISI

anything ever experienced in England.’ Similarly
among fresh-water groups the notoriously rapid
multiplication and spread of the introduced zebra-
mussel (Drezssena polymorpha) in England and certain
parts of the continent of Europe may be instanced; a
statement by Sir C. Lyell that a Lzna@a introduced
unintentionally into Madeira by the Portuguese ran all
over the island in thirty years has already been referred
to. It will be borne in mind of course that molluscs, no
matter how quickly they increase, can never spread
with much rapidity of their own accord; obviously,
migration at a “snail’s pace” can hardly be rapid, and
those kinds which have quickly occupied large areas
must certainly have been largely helped by external
causes.

Man’s influence upon a fauna, it need hardly be
remarked, is not always by way of addition, for he
often drives away or exterminates many aboriginal
forms. The introduction of new and dominant kinds
must in itself be highly prejudicial, while the disastrous
results attending the destruction of forests and the
breaking up of land for cultivation are but too well
known. St. Helena furnishes a striking illustration.
When discovered in 1501, the little island is said to
have been clothed with a luxuriant forest vegetation, but
the woods have been almost wholly destroyed by man
and the animals introduced by him; and notwithstanding
the establishment of a large number of foreign plants,
its general aspect is now described as barren and

1 John Carson, “ Garden,” xiii. (1878), p. 273.

182 THE DISPERSAL OF SHELLS.

forbidding. Much of the rich upper soil, which could
only be retained on the steep slopes so long as it was
protected by vegetation, has been washed away by the
violent rains.’ Asa result, we find that of the twenty-
seven truly indigenous species of terrestrial mollusca
belonging to the island—and the fact has already been
mentioned—only seven now survive, the remainder
having been exterminated, Mr. Edgar Smith remarks,
“by the destruction of the primeval forests.’ Foreign
snails have been introduced, and are now living on the
island, imported probably, perhaps as ova, with “some of
the many introduced plants ;”? the presence of yalinza
cellaria, for instance, has just been referred to: these,
however, from a naturalist’s point of view at least, form
no real compensation for the loss of aboriginal forms,
the destruction of an inoffensive member of a native
fauna—more especially when highly peculiar—being an
irreparable calamity.

Shells have been intentionally dispersed by man to a
considerable extent. Both land and fresh-water kinds, as
everyone is aware, have frequently been “ planted,”
chiefly by shell-collectors, in localities where they were
previously unknown, and permanent additions to faunas,
it will be admitted, have thus been made. To some
persons the transplantation of foreign or local snails
to spots near their residences seems to have special
fascination, anda whole volume might probably be filled

* Wallace, “ Island Life,” 1880, pp. 281-4; ed. 2, 1892, pp. 292-5 ;
and see aiso Darwin’s “ Journal,” ed. 2, 1845, pp. 488-9.
* See Darwin’s “ Journal,” pp. 489-90.

DISPERSAL BY MAN. 183

with accounts of such doings, but we are not much con-
cerned with this kind of dispersal. Numerous unsuc-
cessful attempts at colonization have been recorded, and
these are of significance, I think, as helping us to under-
stand how very small must be the chance of the ultimate
establishment of a new colony as the result of trans-
portal—often, no doubt, to very unsuitable spots—of a
solitary specimen or a few individuals by accidental
means under nature. Man often carries considerable
numbers, turning them out in localities apparently well-
suited to the requirements of the particular species he
wishes to acclimatize, sometimes even watching over
them with care and shooting the birds which come to
pick them up, yet it often happens that the creatures
gradually decrease in numbers and finally die out. It
must be remembered, however, that when thus carried
by man they are generally put down in districts already
well-stocked, and the creatures in such cases are ob-
viously less likely to survive than those which happen
to be transported by natural means to poorly stocked
regions or to newly formed and unoccupied islands.
The few accounts of attempts at colonization which I
have noted down may possibly be worth giving. Un-
successful attempts, it should not be forgotten, are
probably less likely to be placed on record than suc-
cessful ones. 7

The zebra mussel (Drezssena polymorpha) is said to
- have been planted by Mr. Stuchbury of Bristol in some
of the waters near that place,’ and similarly, in America,

‘-H. E. Strickland, “ Mag. Nat. Hist.,” (n.s.), ii., (1838), p. 363.

184 THE DISPERSAL OF SHELLS.

Dr. James Lewis is stated to have put down several
species taken from the Mohawk River—Jelantho
entegra, M. rufa, Goniobasis niagarensis, Somatogyrus
wsogonus, etc.—at the outlet of Schuyler’s Lake (about
eighteen miles from Mohawk), but I do not know the
result of either of these experiments ; an attempt made
by Dr. Lewis to colonize “ Vzvzpara contectoides Binney
(Pal. vivipara Say)” in the Mohawk River and Erie
Canal, however, is said to have been eminently success-
ful, the species having spread widely and being now
‘firmly established in both the canal and river,” and
I find it stated also that Bythinia tentaculata, planted
in the Mohawk River by the same naturalist, has be-
come very abundant, especially in the bends of the
river where the water runs slowly.’ Planorbis corneus
and &. tentaculata, planted by Miss Esmark in Norway,
“increased rapidly ;” Zzmnea stagnalis intentionally
introduced, it is said, as food for trout, into the river
Avon at Christchurch, New Zealand, was said to be
abundant below the Acclimatization Gardens in 1881 ;*
the same species, from England, thrown down in an old
quarry and in deep drains at Possil Marsh near Glas-
gow, is known to have survived at least for some years,*
and Spherium lacustre, Paludina contecta, B. tentacu-

* W. B. Marshall, “ Nautilus,” v. (1892), pp. 133-4, quoting Dr.
Lewis, “ American Journ. of Conch.,” iv. (1868), p. 245.

2 A. F. Gray, “American Nat.,” xvii. (1883), p. 205.

3 F. W. Hutton, “ Trans. and Proc. New Zealand Inst.,” xiv.
(1881), p. 157; C. T. Musson, “ Proc. Lin. Soc:, N.S: W.. ten tess
(2), v. (1891), p. 889.

+ W..D. Roebuck,“ Scot. Nat.” 1891, pe03e:

DISPERSAL BY MAN. 185

lata, Planorbis carinatus, P. complanatus, P. corneus, L.
stagnalis, and L. glabra, from localities, likely to be de-
stroyed, in the neighbourhood of Manchester, deposited
by Mr. Heathcote in quiet ponds about Farrington,
were stated in 1887 to be increasing rapidly in their
new homes.’ <A good deal of colonization from pond
to pond and from a distance, Mr. R. D. Darbishire states,
used to be practised in the neighbourhood of Man-
chester; P/. corneus, for instance, believed to have been
absent from the district in 1841, ‘‘is now common in
the canal at Gorton—most likely a dealer’s colony ;” in
1844-6, it is said, “there was a great rush to Southport ”
for Paludina contecta which was colonized by collectors
and dealers in canals near Manchester, at Twenty Pits,
Greenheys, and cotton pits, Chorlton; P. vzvipara,
also, at Birch, is regarded as a probable introduction.’
On the other hand, Z. stagnalis, put down by Mr. W.
Baillie in astream near Brora, Sutherlandshire, probably
“picked by birds,” could never afterwards be found ; *
S. corneum, B, tentaculata, and L, stagnalis, introduced
into a dam near Greenock, seemed to have died out by
the following year ;* and of several living specimens of
L. stagnalis and P/. corneus, placed in a small pond on

hz otangen: “ Nat:,” 1887,.p. 176.

2 R. D. Darbishire, cited by Mr. Standen, “ Nat.,” 1887, pp. 160
and 164.

3 Neritina fluviatilis, B. tentaculata, and Pl. complanatus have
also been planted in small numbers by Mr. Baillie in the neigh-
bourhood of Brora.

* 1. seott, “Trans. Nat. Hist; Soc; Glasgow,” (n..s.), 1.:(1887),
Pp. 252:

186 THE DISPERSAL OF SHELLS.

Redcar sandhills in 1882, only a single half-grown
specimen of the former could be found in 1884-5.’

With land-shells much more seems to have been
done. The Roman or edible snail (Helzx pomatia) has
often been experimented upon. It is even said by some
to have been originally introduced into this country
from the continent, but this is a point to be considered
in the next chapter. Specimens from Stonesfield,
Oxfordshire, were introduced many years ago into the
Botanic Gardens at Oxford, where, it was stated in
1883, a large colony “is now thriving.” * Many years
ago also a few specimens are said to have been turned
out by Mr. Bean in Forge Valley, Scarborough, and
individuals, as reported in 1868, were subsequently
found there from time to time ; Mr. Ashford mentions
having seen a full-grown living shell, taken in the locality
in April, 1868, “ doubtless a descendant of the originally
transferred stock.”* Mr. T. D. A. Cockerell records ©
the liberation of four specimens near Chislehurst, two
of which were seen again after nearly two years.* Mr.
Thompson, in 1841, stated that the creature had “of
late years been introduced from England to different
localities in Ireland, as Dalkey Island, off the Dublin
coast, Youghal, etc. ;” in the autumn of 1834 a few had
been turned out by him, together with Cyclostoma elegans,
on the chalk near Belfast, but they did not increase, and

' B. Hudson, “Journ. of Conch.,” v. (1886), p. 48.

2 'S.'S. Pearce, *“Zoologist,” (3), vile qaoe3), . Sos:

3 C. Ashford, ‘‘ Science Gossip,” 1868, p. 160.

4°T. D. A. Cockerell, “ Journ? of Conch.,? iv. (Toe4), p24.

DISPERSAL BY MAN. 187

after a few months not one of cither species could be
found about the place. Specimens were introduced
into Scotland “by Pat. Neill, Esq., and placed in his
curious and most interesting garden at Cannonmills,”
but they did not prosper,and,as Dr. Johnston stated
in 1830, were believed to be gradually disappearing.”
Dr. Lukis tried to introduce the creature into Guernsey
but apparently without success.” When put down in
the neighbourhood of Christiania, Miss Esmark tells me,
it always vanishes after the first year: some trans-
planted from Sweden, for instance, all disappeared, as
did others brought from Denmark by Professor k.
Collett ; probably they were eaten by foxes or hedge-
hogs which live everywhere in the neighbourhood.*
Professor Collett several times tried to transplant the
creature in Denmark, only from one estate to another
close by, but it always died. According to the Rev. J.
Morton, who wrote in 1712, many specimens were in-
troduced by Lord Hatton into a coppice near his house
at Kirby, in Northamptonshire, “with Intentions that
they should breed there ; but in a short time they all
dy’d.”* Fifty or sixty specimens were once turned out
in the neighbourhood of Petersfield, but did not establish
themselves ; here, also, as suspected by Mr. Harting,

1 W. Thompson, “ Ann. and Mag. Nat. Hist.,” vi. (1841), p. 21.

? G. J[ohnston]., Loudon’s “ Mag. Nat. Hist.,” iii, (1830), p. 47.

3 A. H. Cooke and H.M. Gwatkin, “ Quart. Journ. Conch.,” i. p.
333.

4 B. Esmark, “ Journ. of Conch.,” v. (1886), p. IIo.

5 J. Morton, “Nat. Hist. of Northamptonshire,” (1712)5 as
quoted in “Journ. Northants. Nat. Hist. Sees) 11.1 (1S85)) psrg:

188 THE DISPERSAL OF SHELLS.

they were probably devoured by hedgehogs. About
1868-9 Mr. W. Jeffery is said to have turned out thirty
or forty Surrey specimens partly in his garden at
Ratham, Chichester, and partly on a bank of light soil
near, but of those on the bank nothing more was seen,
and in the garden—though they seemed to do pretty
well for a time, producing, at least, one brood of young,
some of which attained full size—the original stock had
all vanished previously to 1874, and during the prece-
ding summer only two young ones had been seen.!

Mr. Jeffreys, as he stated in 1862, twice endeavoured,
at different times of the year, with an interval of nearly
three years between each attempt, to colonize the
beautiful and local Helix pisana on the sandhills near
Swansea; a basketful of living specimens from the
well-known habitat at Tenby, only about thirty miles
distant, were spread over different parts of the Burrows,
where the soil and herbage appeared to be the same as
at Tenby, a more easterly aspect at Swansea being the
only noticeable difference in the localities, but, as
Jeffreys added, although they seemed “ at first to thrive
tolerably well in the new locality, they did not multiply,
and the birds soon ate up the immigrants.” ? He was
mistaken, however, in supposing that his endeavours
had been altogether unsuccessful, for on searching the
sandhills in 1874, Mr. Rimmer, to his astonishment,
found the shell living there in countless numbers and of

' J. E. Harting, “ Zoologist,” (3), ii. (1878), pp. 89-90; and see
* Rambles in Search of Shells,” 1875, p. 72.
* “ British Conchology,” i. (1862), p. 208.

DISPERSAL BY MAN. 189

all ages, a white variety apparently predominating.
Mrs. Dyer of Swansea, who is familiar with the species,
as Mr. Gude tells me, saw the creatures still living on
the sandhills on last visiting their habitat in 1884.
The original immigrants, it seems, as Mr. Rimmer re-
marks, did not long survive, “the sudden change, per-
haps, injuriously affecting their delicate constitution ;”
evidently, however, they left behind them a progeny,
which, being born in the new home, took more kindly
to the place.! Similarly, as stated by Bourguignat, a
large number of Helix Jactea from Valencia died during
the second winter at Castel-Roussillon, but some more
vigorous individuals born in the new locality appear to
have survived for some time ; finally, in this case, how-
ever, all perished.” In 1860, a son of Dr. Lukis, while
staying in Jersey, where HZ. pzsana is abundant, some-
times covering the thistles,’ collected a number and
carried them to Guernsey, establishing two colonies
there, one at Vazon Bay and the other at Vale Castle,
and sending a few to his sister, Mrs. Collings of Sark,
who turned them out in that island “on the right bank
going down to the Port du Moulin.” In 1877, Mrs.
Collings reported, however, that none had been seen in
Sark for some years; for a few years a shell or por-
tions of a shell had occasionally been found near a stone

1 R. Rimmer; “Quart. Journ. Conch.,” i. p. 267; and see his
“Land and Fresh-water Shells,” 1880, p. 133.

2M. J. R. Bourguignat, ‘“ Mollusques nouveaux, litigieux, ou
peu connus,” 1867, p. 234.

3 “ British Conchology,” i. (1862), p. 209.

190 THE DISPERSAL OF SHELLS.

where a thrush or blackbird had had a meal, and thus it
was feared all hadvanished. The Vale Castle colonists
were seen by Mr. Rimmer during the summer of 1873,
but they inhabited only a limited area, and numbered
probably not more than a few hundreds. At Vazon
Bay, the creatures seem to have done much better, for
in the summer of 1876, Mr. Sherriff Tye saw “ many
thousands ” in that locality, and the following year the
colony was reported by the Rev. A. H. Cooke and the
Rev. Professor Gwatkin to have extended over an area
at least half-a-mile in length, and I find it stated that
specimens were again collected in 1884 by Mr. Brockton
Tomlin. It is interesting to note, Mr. Cooke and Mr.
Gwatkin have pointed out, that the capabilities of these
colonies for sustaining life varied directly with the
exposed nature of their situation: “that at Sark, on
rather high and rocky ground, probably never took
root at all, that at Vale Castle, in «spite of its aiorth=
easterly outlook, was on far more congenial soil, and
consequently survived until 1873 or later, while that at
Vazon, planted on low-lying ground, and facing the
north-west, has lived and flourished.”’ Mr. Heathcote
is said to have planted this snail in considerable numbers
in seemingly favourable situations amongst the South-
port sandhills, but I do not know with what result ; it
is stated also that Mr. T. Rogers has often tried to

1 A, H. Cooke and H. M. Gwatkin, “ Quart. Journ. Conch.,” 1.
pp. 332-3; R. Rimmer, “ Quart. Journ. Conch.,” i. pp: 266-758G.
S. Tye, “Quart. Journ. Conch.,” 1... pp. 230-15) J. W. @ayles,
“ Journ. of Conch.,” iv. (1885), p. 275.

DISPERSAL BY MAN, IQI

colonize it, presumably in Lancashire, but has hitherto
failed, owing, he believes, to the creatures having been
picked up by birds.’ A few turned out in the north of
Scotland, near the coast, by Mr. W. Baillie of Brora,
are believed to have perished during the first winter.
Mr. Baillie has experimented with several species
near Brora, with considerable success. In 1883, he
mentioned that he had introduced into the lower part
of the basin of the Brora River colonies of Helix acuta
and Helzx virgata from Llandudno, H. ericetorum from
North Sutherland, and H. asfersa from Banff, the first
three of which seemed likely to hold their ground, but
H. aspersa had been lost sight of, “ probably killed out
by blackbirds.” Subsequently, in 1887, some of these
colonies were reported to have done well; Hel¢r
acuta and Helix virgata were living in hundreds, extend-
ing northwards from the mouth of the Brora River for
upwards of half-a-mile. Clauszlia parvula, which had
been introduced from Normandy, seemed to hold its
ground. Clauszlia laminata, Cyclostoma elegans, and a
few specimens of 7. fesana, H. cantiana,and H. rufescens
had also been turned out.” In October, 1891, Mr.
Baillie had the kindness to furnish me with a few
additional notes ; the //7. wrgata colony, he stated, was
the most promising, the creatures having increased
near ithe’-sea to thousands. .77. ecvfa; which had
prospered for some time, had apparently died out.

mane otaneen: SiNat.;” 1887) p. 176)
2 W. Baillie, “ Journ. of Conch.,” iv. (1884), p..1160; v. (1887),

p. 192.

192 THE. DISPERSAL, OF “SHELLS:

FH. ertcetorum had increased to a considerable extent ;
HT. aspersa, also, had survived and had become so
numerous in his garden that hundreds had been trans-
planted to different parts of the East Sutherlandshire
coast during the preceding summer. A. cantzana and
FT, rufescens had done well, and Claustlia parvula was
still alive, as also in all probability were CZ. /aminata and
Cyclostoma elegans. Living specimens of Clauszlia bipli-
cata from the banks of the Thames, CZlausilia solida
from France, and Cochlicopa tridens and Helix pomatia
had also been turned out in small numbers. A number
of Clausilia laminata and Clauszlia rolphit from Buriton,
Hants, were once turned out by the Rev. J. McMurtrie
ina glen at North Berwick, Haddingtonshire, but, as
was stated a few years afterwards, “it is not likely they
have withstood the climate.”’ Mr. G. B. Adami, it is
said, “has tried to introduce seventeen species of land
and fresh-water molluscs at Edolo, Val Camonica, from
other parts of Upper Italy,” of which four soon disap-
peared, the rest surviving” A number of land-shells
(Helix ?, several species) from Buenos Ayres and
Montevideo were once turned out in a garden in York-
shire, and after three or four years many were re-
ported to be still alive and apparently healthy. It is
interesting to find, as the observer remarked, that the
creatures were able to withstand the rigour of our
climate ; one of the winters through which they passed

" J. McMurtrie, “ Journ. of Conch.,” vi. (1889), p. 5.
? “* Moll. terr. e fluv. di Brescia e Bergamo,” 1876, p. 91, as
quoted in the “Zoological Record,” xiii. (1876), Moll. p. 18.

DISPERSAL BY MAN. 193

seems to have been exceptionally severe.’ The suc-
cessful transportal of Helix nemoralzs from England
to New Jersey, and the establishment of several new
colonies of the same species from an accidentally intro-
duced stock in Virginia, have just been mentioned ;
Helix appressa, sent from Illinois, became well estab-
lished in New Jersey in the same garden;* Hlelix
austriaca, according to a statement in the Zoological
Record for 1871,°” acclimatized seventeen years ago
near Stettin, thrives there very well and multiplies
considerably ;” Helzx villosa from Switzerland turned
out by the Rev. Canon Tristram on a moor in
co. Durham was known to be living in the same place
ten years later, and Helix lapicida from the south
of England planted by one of Mr. Tristram’s friends
on some rocks on the banks of the Wear, in the same
county, ultimately became one of the recognized
local shells. But Clauszlie from Africa and from
Syria and various species of He/zx from the latter
region also turned out in the same county, being
able to “protect themselves against heat better than
cold,’ are said to have perished during the first
winter ;* Helzx lapicida from England, and Aulimus
decollatus from Charleston, South Carolina, disappeared

1 G. Norman, “ Zoologist,” xii. (1854), p. 4435.
2 W. G. Binney, “ Terrestrial air-breathing Mollusks,” v. (1878),
pp. 29, 306.
3 viii. (1871), p. 130, referring to Dohrn, “ Nachr. malak. Ges.,”
ill, pp. 17-18.
4 H, B, Tristram, “ Zoologist,” (3), i. (1877), pp. 260-1-
O

194 THE DISPERSAL OF SHELLS.

at once in Mr. Binney’s garden ;' Helzx hortenszs from
the island of Martha’s Vineyard, “of course introduced
from Europe,’ could not be established by Mr. J. H.
Thomson in his garden, the young being devoured,’ it
was believed, by slugs, Lzmax, Arzon, etc. ;* thirty or forty
specimens of the same species once placed in a hedge-
row in this country, in the neighbourhood of Upton,
were never afterwards seen ;‘ a large number of Helzr
virgata from various localities turned out near Chisle-
hurst did not “seem to multiply,’ and many specimens
of Helix cartusiana from Sandwich liberated in the same
neighbourhood are believed to have died out.’

Slugs, I suppose, have not been much dispersed inten-
tionally. Half a dozen Testacelle from the Isle of
Wight once turned out by Mr. W. Jeffery in his garden
appear to have been lost sight of; a single quarter-
srown individual, however, is said to have been detected

1 W. G. Binney, “ Terrestrial air-breathing Mollusks,” v. (1878),
379:

? With reference to this statement it may be interesting to
notice that in Charleston, S.C., a foreign snail, believed to be
Helix nemoralis, formerly existing there in great numbers, was
completely exterminated by Audlimus decollatus, also a foreign
species, at least so the elder Binney was informed on what was
considered good authority, but, as he admitted, the destruction of
the one species, though subsequent to the introduction of the
other, might not have had any connection with it. A. Binney,
‘Terrestrial air-breathing Mollusks,” i. (1851), p. 134.

$ J: H. Thomson, “ Journ. of Conch;,” 1v: (1885), p? 373°

4°T. Byerley, “ Proc. Lit. and Philos. “Soc., ' Liverpool)sre54
Pp. 47.

* T, D, A. Cockerell, “ Journ, of Conch.,” iv. (1884), p. 238,

DISPERSAL BY MAN. 195

after about a year.’ Mr. Baillie tells me that he put
down a few specimens of Amalia marginata in his
garden—in Sutherlandshire—about nine years ago, but
nothing has been seen of them since.

We have now to inquire as to the means by which
molluscs may have been unintentionally dispersed by
man’s agency.

FRESH-WATER SHELLS.

Aquatic shells and their ova—no doubt less frequently
dispersed by man than terrestrial kinds—seem likely to
have been occasionally carried with cargoes and con-
signments of various sorts. One species now flourishing
in this country, as we shall see, is believed to have been
imported attached to timber in the holds of ships, and
another is supposed to have come from America in
cotton bales. Our Lzmnea stagnalis, it is suggested, was
introduced into Tasmania, I presume unintentionally,
with fish-ova.” Many kinds, both ova and adults, must
certainly have been carried from place to place, even to
great distances and from one country to another, along
with consignments of plants, but I am not aware that
the creatures are ever actually noticed by those whc
unpack such consignments; several facts, however,
clearly indicating transportal in this way might be given.
Thus, for instance, Mr. W. W. Westgate of Houston,
Texas, once found his water-lily tubs, in which plants
from Fayette County, Texas, and from Florida were

, W. Jettery,““Journ.,of Conch:,” ii. (1882), p: 373.
SW) Pettard,.~ Jaurn.iofConch:,” ii. (1879); p: 81.
O 2

196 THE DISPERSAL OF SHELLS.

crowing, “ filled with Physa gyrina Say” of a different
form from those found in his neighbourhood ;' and some
years ago, Dr. Alford Nicholls found fresh-water shells
of two species inhabiting tubs in which he was cultivat-
ing aquatic plants, gathered some time previously in a
pond in the island of Antigua, and carried from thence
to Dominica. The digging of canals, of course, as
Forbes long ago remarked, has largely facilitated the
spread of these animals.” In America and in other
large land areas, it will be remembered, “ the waters of
streams which differ widely in their faunz” are some-
times thus connected.°

LAND-SHELLS.

It seems likely that land kinds are frequently carried
in ships’ ballast, collected, as it often is, near the shores
of one country and discharged on those of another.
Dr. Binney in a discussion on the introduction of foreign
species into the United States referred to this cause as
a “common mode of introduction ;”* and at least three
species which have occurred alive in England are
believed to have been thus imported. Discoveries of
living shells in freshly discharged ballast would be
of much interest, but Iam not aware that any records
of the kind have been made. Miss Esmark mentions

1 W. W. Westgate, “ Conchologists’ Exchange,” ii. (1888), p. 98.

? Edward Forbes, “ Mem. Geol. Survey of Gt. Britain,” i. (1846),
p- 398.

3 W. B. Marshall, “ Nautilus,” v. (1892), pp. 133-4.

4 See “ Terrestrial air-breathing Mollusks,” i. (1851), p. 151.

DISPERSAL BY MAN. 197

the occurrence of “a well conserved shell of Helex
aspersa in some imported sand near Langesund.” A
shell of Helix ericetorum once found near Christiania,
this author suggests, may have been imported “among
sand brought with vessels from England or France.”’
It seems almost certain that the creatures are occa-
sionally carried many miles with railway ballast, with
chalk, etc., used in making sea defences and other
embankments, with earth and other substances trans-
ported for horticultural purposes, etc. Thus the Kentish
snail (Helix cantiana), abundant on railway-banks near
Hartlepool, “ may have been introduced in ballast ;” °
Felix virgata, found plentifully on embankments
between Pevensey and Bexhill, where the London,
Brighton and South-coast Railway runs close to the sea,
was probably brought, as the Rev. S. Spencer Pearce
states, with the chalk forming the embankments, which
is believed to have come from the Eastbourne cutting,
some eight or nine miles away ; the same snail, accord-
ing to another observer, noticed in several places on
railway-banks also in Sussex “was probably brought
with chalk ;’ a colony of Helzx cartuszana, also, known
to have existed for some years at Cowfold, in the
same county, and now extinct, is believed to have
been originally introduced with chalk Helzx virgata,

1 B. Esmark, “ Journ. of Conch.,” v. (1886), p. 106.

2 «“ Naturalist,” 1886, p. 148, quoting J. E. Robson, “ Young
Nat.,” v. (1884), p. 192.

3 W. Borrer, cited by J. E. Harting, “ Zoologist,” (3), 11. (1878),

p..92.

198 THE DISPERSAL OF SHELLS.

abundant along the promenade at Southport, and only
observed there in recent years, is “supposed to have
been imported with the building materials,’' and the
same species, to give another case, established in gar-
dens, now disused and weed-grown, around some of
the Martello Towers in the midst of the broad shingle-
beach, which, of course, is destitute of terrestrial mol-
luscs, between Eastbourne and Hastings was probably
introduced, Mr. Pearce suggests, with the chalky soil
of which the little plats were formed, or it may have
been carried thither with garden plants.

That the creatures have been widely dispersed along
with plants, shrubs, etc., now so frequently sent from one
country to another, cannot be doubted for a moment ;
unconsciously horticulturists have probably done more
than any other class of persons to mix the aboriginal
faunas of distant countries, and doubtless transportal in
this way for short distances is constantly taking place.
Wardian cases, as Mr. Layard remarks to me, must
certainly be great factors of dispersal, and living snails
are known to have been transported with quite small
consignments of plants, in the earth at their roots,
and even with “ dry-bulbs” transmitted with little or no
earthy matter. Foreign snails, and others evidently
from a distance, are well known to turn up every now
and then in greenhouses: during 1883, Mr. Musson pro-
cured three foreign species, probably South American,

‘ H.H. Higgins, “‘ Address to Liverpool Nat. Field Club, January
30th, 1891,” p. 24; the creature occurs also near Southport at
Moels Cop Road.

DISPERSAL BY MAN. 199

two of them occurring plentifully, from Mr. Thacker’s
orchid-house in Nottingham, and many similar finds
might be instanced ; one exotic species, in fact, is now
well established in greenhouses, cucumber-frames, pine-
beds, etc. in many parts of this island. Mr. Heath-
cote is said to have taken some splendid specimens of
Hyalinia draparnaldi—apparently unknown as an in-
digenous species in Lancashire—in his greenhouse at
Preston in that county. Mr. Standen mentions having
discovered one in his fernery at Swinton near Man-
chester ; it had probably been transported with some
ferns sent to him by Mr. Heathcote.’ It is interesting
to find that the creatures have been actually noticed
among imported plants: thus Mr. Gude recently showed
me a living shell of Helzctna amena, found by one of
the employés at Messrs. Williams’ nurseries, Upper
Holloway, in a case of orchids (Cattleya citrina) which
had just arrived in a dry state from Mexico, and it is
perhaps worth mentioning that a living snail (elex
nemoralis or hortenszs) has been received at the same
nurseries in a consignment of Sphagnum-moss from
Essex. To give another instance: a Bulimus-snail,
possibly Swbulina striatella, was once found in Madeira
by Senhor J. M. Moniz amongst some plants received
by him from the island of St. Thomas in the Gulf of
Guinea ; and on another occasion, three dead shells
believed to belong to the same species, foreign to the
Madeiran Archipelago, were found in an old bone,

* R, Standen, “ Naturalist,” 1887, p. 176.

200 THE DISPERSAL OF SHELLS.

in a garden near Funchal.’ Helix acuta is said by
Kobelt to have arrived alive in gardens at Frank-
fort on palm-trunks.2>, The Lexington colony of /Helzx
nemoralts, according to Professor Morrison, probably
sprang from specimens imported “in earth, in flower-
pots,” * and many facts proving such transportal are
likely to have been observed in different parts of
the world. Thus—to give the only instances with
which I am acquainted—during Sir C. Lyell’s stay in
Madeira no less than three species of Portuguese
snails (/Ze/ices) were found in the earth of a single
flower-pot in which a garden plant had been sent from
Lisbon ;* and several specimens of Pupa umbzlicata were
once found alive, together with a specimen of Clauszlza
rugosa, in the mould around some ferns received by Mr.
Musson at Nottingham from Derbyshire. The eggs of
many kinds, seemingly specially suited for dissemination
in these modes, are probably carried much more fre-
quently than the adult animals, but from being com-
paratively inconspicuous and often buried in soil or
concealed about the roots of plants they are not likely
to be often noticed.

* T. V. Wollaston, “‘ Testacea Atlantica,” 1878, p. 205.

* “Zoological Record,” vii. (1870), p. 130, referring to Kobelt,
** Nachr. mal. Ges.,” ii. p. 160.

> H.A. Pilsbry, ‘* Nautilus,” iii. (1889), pp. 51-2.

* “Principles,” ii. p. 427; and see a letter from Lyell to Bun-
bury, dated in 1856, referring to Madeira: “a friend of Wollaston
received a flower from Europe in a pot very lately, and found five
species [sic] of European helices alive, buried in the mould.”
“Life, Letters and Journals of Sir C. Lyell, Bart.,” ii. (1881),
p. 209.

DISPERSAL BY MAN, 201

Cargoes and consignments of vegetable productions,
other than those intended for horticultural purposes,
seem likely also to have served occasionally as means
for dissemination. Thus in a recent enumeration of
Morocco shells mention is made of the finding of Helzx
dehnit amongst gum arabic ;' Helix vermiculata, as we
shall see in the next chapter, has been found alive in a
parcel of chemist’s horehound ;’ and living specimens
of Helix caperata have been discovered amongst screen-
ings from barley intended for malting ; we find, too,
that several specimens of a foreign snail were once
discovered alive in a garden into which a quantity of
such screenings had been imported.’ Dead shells of
Helix alauda,a species belonging strictly to the fauna
of Cuba, have been found on one or two occasions
among bananas in Boston, Mass., and a living specimen
once found in a grove in Rhode Island was probably
introduced, as Mr. John Ford has suggested, with a
bunch of those fruits.“ The importation with dye-wood
from Brazil of two specimens of Lulimus largilliertc has
been noted somewhat recently in the “ Nachrichtsblatt
der deutschen malakozoologischen Gesellschaft ;°” and
living specimens of Bulimus undatus, adhering to
tropical timber, are said to have been imported into

* “Zoological Record,” xvii. (1880), Moll., p. 24, referring to
“J. de Conch.,” xxviii., pp. 1-83.

-setec. Conch: Soci, “ joum.-of'Conelt.,” vi. (1897), p: 393.

3 P. B. Mason, “ Journ. of Conch.,” ili. (1880-2), p. 118.

4 J. Ford, “ Conchologists’ Exchange,” 11. (1887), 71-2.

> ©. Boettger,. “ Nachr. mal. Ges.,” xviii., p. 58, as quoted in
“ Zoological Record,” xxiii. (1886), Moll., p. 61.

202 THE DISPERSAL OF SHELLS.

Liverpool ; indeed, Sir C. Lyell was even informed by
Mr. Broderip that the species was naturalized in the
woods near that city!’ Dr. Stearns has referred also
to the conveyance of Bu/imz of two species to San
Francisco in a cargo of dye-woods from the Gulf of
California ; he states:

“The dye-woods had been hauled from the place
where they were cut, and piled up near the embarcadero
onthe gulf shore,and afterwards transferred to the vessel.
These snails had crawled into the hollows and crevices
of the wood, and were discovered when the cargo was
unloaded and put on the wharf in San Francisco.
When the sticks were thrown ashore the rough handling
shook out the snail-shells ; many also were found in the
hold of the vessel after the cargo was discharged.”

Unfortunately in this case, however, none of the
several specimens obtained by Dr. Stearns were alive.”

Foreign shells, it is curious to find, have even been
shipped to this country with cattle-bone. Mr. G. Norman,
of Hull, it appears, once “ procured a great number of
living mollusca (apparently several species of Helix)
from a vessel recently arrived with cattle-bone from
Buenos Ayres and Montevideo, in the cargo of which
they abounded,” and many turned out in a garden, as
just mentioned, lived for several years.*

* “Principles,” ii., p. 3713 and. compare Gray’s “ Turtons1s57)
p: 292.

* R. E. C. Stearns, “West American Scientist,” ‘vii. (1891),
p- 108.

* G. Norman, “ Zoologist,” xii. (1854), p. 4435.

DISPERSAL BY MAN. 203

The use of some land molluscs for food and for medi-
cinal purposes has doubtless conduced to their dissemina-
tion. Thus, for instance, Bulimus (Placostylus) edwards-
janus, an outlier of the B. fidratus group, is believed
by Mr. Layard to have been introduced into New Cale-
donia from the Loyalty Islands during the native wars,
the animal having been used as food by the Loyalty
Islanders and probably carried over by them when they
landed to'fight the New Caledonians. European kinds,
as everyone knows, constitute an important part in the
live stock of many ships, between seven and eight thou-
sand of Helix asfersa, as Tate states, for instance, form-
ing part of the provisions of a ship leaving the port of
Bordeaux for a long voyage.' As is well known, also,
the creatures are sometimes exported to distant countries
as delicacies, and they travel well, according to Gray, in
old casks, fixing themselves on one another round the
circumference and leaving a vacant space in the centre.”
As stated by Dr. Binney, the larger European species,
and particularly Helix aspersa, are sometimes im-
ported into the United States, for use as food by
foreign residents.* Now and then, doubtless, from some
cause or other, a few manage to escape near sea-
ports, or after arriving at their destinations further
inland, and many colonies have almost certainly been
established in this way. Portuguese sailors, with whom

1 R. Tate, “ Land and Fresh-water Mollusks,” (1866), p. 116.

> Grays: Luxton,” 1657,.p. 105.

3 A. Binney, “Terrestrial air-breathing Mollusks,” i. (1851),
Pv agi-

204 THE DISPERSAL OF SHELLS.

snails are much in favour as “live sea stock,” according
to Woodward’s “ Manual,’ have naturalized “ Helzx
aspersa in Algeria, the Azores, and Brazil, and Helix
lactea at Teneriffe and Montevideo.” ! WH. lactea is said
to be extremely common in the markets at Buenos
Ayres.” Occasionally, as might be supposed, imported
edible snails are intentionally thrown out in the hope
that they may establish themselves. Helzx pomatia,
as mentioned in the next chapter, is said by some
authors to have been thus established in England by
the Romans, and others say that it-was subsequently
planted by our own countrymen who had imported it
either as a delicacy or for medicinal uses. Mr. Layard
assures me that [Helix aspersa, now abounding about
Cape Town, first obtained a footing in this manner.
The following extract from a charactcristic letter of his
—dealing also with the introduction of the creature into
the Loyalty Islands—will be read with interest :

“In 1854 (or 1855, I am not sure which) I was living
in the ‘Gardens,’ that part of Cape Town running up
the slope of Table Mountain. I one day noticed in the
vineyard attached to my house, on a wall, a cluster of
young /7. aspersa. Of course I knew at once they
must be an introduction. Soon complaints were made
of vines and vegetables being devoured by snails, and
as I was known to be a “snail collector,” the introduc-
tion of the marauder was set down to me. This I at
once repudiated, and set to work to find out who had

* Woodward’s “ Manual,” ed. 4, rep. 1890, p. 87.
* W. H. Rush, “ Nautilus,” vi. (1892), p. 82.

DISPERSAL BY MAN. 205

done it. I discovered that some months previously a
French man-of-war had been in the harbour and having
received much kindness and attention from the French
Consul, Mons. D., had requited it somewhat by pre-
senting him with a little barrel of “escargots” of which
he was very fond. Mons. D. was a wise man! He
reasoned within himself, “I can both have my cake, and
eat my cake; I'll eat the big ones and plant the little
ones.” So “whene’er he took his walks abroad” he
took a few little ones with him, and chucked them over
the garden walls; and they increased and multiplied
and filled the land, and in the words of the old writer
(slightly altered)

Mons. D. bred the bore
But E. L. L. got the ‘ gloire’ !

“In 1879, I was on a collecting trip to the Loyalty
Islands, I was living in the house of a trader named
Wright at Chepenché, in the island of Lifou. The
natives, of course, discussed me and my doings, how I
shot and skinned birds and picked up shells. One day
a native came, and addressing Wright, said he had
brought a shell that no native of the island had ever
seen before ; he had found it in his Yam patch, and hoped
I would give him a good reward for it. After this
oration he carefully untied the corner of his waist-cloth,
and produced a bundle of Yam leaves, carefully tied up ;
in the centre of these was a half-grown HZ. aspersa !
Wright examined it with great care. He had never
seen anything like it before (he was born in Australia)

206 THE DISPERSAL OF SHELLS.

and he knew the shells of the country pretty well.
Had I ever seen it? I told him I had, rather! and I
told him and the native the mischief they would do,
and the intruder, being condemned to death, was
promptly “crunched” under my foot. Then came the
question, how got it there? I told Wright how it got
to the Cape, and went off in a spirit of inquiry (sea-
soned with diplomacy) to Mons. Luguier the French
chief official, or Résident as he is called; a little
manoeuvring brought round the question, did he like
escargots? “Ah, yes!” and he had lately had a stroke
of luck; a French man-of-war had called in and given
him a nice lot of them; he had eaten the large ones
and had distributed the small ones about the place, and
when they grew bigger and had large families, he would
always have a dish at command. So thus the introduc-
tion of the pest into two widely remote countries is
most distinctly traced to French men-of-war.”

SLUGS.

The naked molluscs, as Mr. Hedley remarks to me,
seem to have roamed further and faster than shell-
snails ; in their race to the antipodes, at least, as recently
pointed out by him, European kinds “have far out-
stripped their shell-bearing relatives,” * and this is cer-
tainly a surprising result, for, as we have seen, the
creatures clearly seem less fitted than snails for acci-
dental dispersal under nature. In all probability, how-

1 C. Hedley, “Ann. and Mag, Nat. Hist.,” (6), ix. (1892), p. 170.

DISPERSAL BY MAN. 2O;7

ever, they have been widely scattered by man, being
carried, no doubt, for the most part, along with consign-
ments of plants, shrubs, etc. Many kinds, indeed,
seem eminently suited for such transportal ; of Testa-
cella haliotidea, for instance, Mr. Jeffreys writes :

“A usual habitat of this kind of Testacella is at the
roots of flower-plants, or under heaps of dead leaves in
gardens; and if a plant were imported into this country
from the botanic garden at Montpellier with the native
soil or a compost made of leaf-mould, either the snail-
slug or its eggs would perhaps accompany it.”’

The presence of 7: maugez in this country is usually
attributed to transportal with plants. Many kinds
must annually travel in safety from one country to
another in Wardian cases, etc. Observed facts proving
dispersal in this way, however, are probably scarce,
but this is explained perhaps by the fact that the crea-
tures, probably travelling chiefly as ova, are not likely
to be often noticed. Tasmanian specimens of the great
erey slug (Limax maximus) have been observed to be
infested with anacarus, and if this proves to be identical
with the European parasite, the fact will argue, as Mr.
Hedley remarks, that the animals migrated in the aduit
rather than in the egg state.” The United States National
Museum, it is interesting to find, once “received a
specimen of the peculiar slug Veronzicella which had
been found by the giver in a bunch of bananas ;”* and

* « British Conchology,” i. (1862), p. 146.
2 ©. Hedley, “ Ann. and, Mag. Nat. Hist.” (6), ix. (1892), p.. 170.
3 R, E. C. Stearns, “‘ West American Scientist,” vii. (1891), p. 108.

208 THE DISPERSAL OF SHELLS.

it is worthy of note also that a couple of “splendidly
srown Parmacella”—slugs foreign to our fauna—no
doubt introduced accidentally with plants, are said to
have been detected on one occasion in a garden near
Newcastle.’

1 H. B. Tristram, “ Zoologist,” (3), 1.. (1877); pp. 260-1.

CHAPTER, IX.

ON THE FRESH-WATER AND LAND MOLLUSCA IN-

TRODUCED) INTO THE BRITISH ISLES BY HUMAN
AGENCY.

MAN’S agency, as Forbes long ago remarked, “may
materially affect a fauna, and has affected that of
Britain.’' Of the forty-six fresh-water species included
in the Conchological Society’s list of 1883, however,
only two, the zebra mussel (Drezssena polymorpha) and
an American coil-shell (Planorbis dilatatus), can be
reasonably regarded as human importations ; and, as
far as I know, only one other, Spherium ovale, has ever
been looked upon as even doubtfully indigenous.

Spherium ovale, which is local in this country, happens
to occur in company with Planorbzs dilatatus in canals
at Gorton, Pendleton, etc., so that Mr. Jeffreys was led
to suggest, in 1869, that it might possibly be the S.
transversum of Say introduced, like P. dilatatus, from
America ; but he observed that it had long been known
in this country, and that he possessed a specimen which
was in Dr. Turton’s collection of British shells more

+ Edward Forbes, ‘‘ Report, 9th meeting, British Association,
1839,” (1840), p. 130.

i

ZO THE DISPERSAL OF SHELLS.

than forty years previously... It has been further
suggested that shells regarded as pertaining to a form
of Planorbis glaber Jeff., found associated with P.
dilatatus in a water-lodge at Burnley, may be Say’s P.
parvus also introduced from America ; P. glaber, how-
ever—often regarded as identical with P. parvus—
though local, ranges widely in Europe, extending in this
country ‘‘ from the Shetland Isles to Land’s End,” and
its right to rank as a native is not doubted.’

A short account of what is known of D. polymorpha and
P. dilatatus in Britain, and of the manner in which they
were probably introduced, may be of interest ; and it is
worth while, perhaps, to add a note on an exotic pond-
snail, Physa acuta, which, though not referred to in the
Conchological Society’s list, is known to live, under
artificial conditions, in two localities in England, and
one in Scotland, and has in all probability been intro-
duced unintentionally by man.

DREISSENA POLYMORPHA Pall. On 2nd November,
1824, the Linnean Society, as appears by an extract
from their minute-book, received from Mr. J. De C.
Sowerby, specimens of a fresh-water shell, “ probably
the Mytilus polymorphus Gmel., 3363,” which had been
found in abundance, attached to shells and timber, in

1 J. G. Jeffreys, Ann. and Mag. Nat. Hist.,” (4), iv. (1869), pp.
341-2; and see also W. C. Hey, ‘‘ Journ. of Conch.,” iii. (1882), p.
271; T. Rogers, “ Journ. of Conch.,” v. (1887), p. 220; R. Standen,
“Naturalist, 1837, p-:157-

2 T. Rogers, “Journ. of Conch.,” v. (1887), p. 219; Jeffreys,
“British Conchology,” i. (1862), p. 86; Rimmer, “Land and
Freshwater Shells,” 1880, p. 40.

DISPERSAL BY MAN. 211

the Commercial Docks, in the Thames, by James
Bryant, Esq., who was in the habit of using the animal
as a bait for perch-fishing, and this is the first mention
of the existence in Britain of the now common and
widely distributed D. polymorpha. Ina letter accom-
panying the shells, Mr. Sowerby suggested that the
species, which was known in the Danube and in the
rivers of Russia, had probably been imported into this
country with timber, and in the following year Mr. Gray
expressed a similar opinion, believing the creature to
have been brought with timber from the Volga, and
further pointing out that it was able to live for a long
time out of water, he having thus kept one in health for
three weeks, As some molluscs attach themselves to
the bottoms of ships, Sir C. Lyell was led to suggest
that the present species might have been introduced in
this way, and it was explained by Mr. Garner that by
keeping the valves constantly closed it was perhaps able
to survive immersion in salt-water during the voyage.
It is much more probable, however, that the creature
was imported with timber as suggested by Sowerby and
Gray, and this was afterwards admitted by Lyell him-
self. A friend of Gray’s is said to have seen the
animals sticking to Baltic timber, which had not been
unloaded, and was still in the ship’s hold.’

Pe tanse Ii! OGC.4 Xiv. (1825), p/585 3 *“Ann, Phil.” (1s))/'nx
(1825); p.226; “Zool. Journ.,” 1. (1825),,p: §$4-;-Jz E. Gray, “Ann.
Phil A(ies;))i<., (0825), ps 1395 Lyell, “Principles,” ed: 9, (1853),
p. 652; R. Garner, Charlesworth’s “‘ Mag. Nat. Hist.,” (n.s.), iii.
(1539), pp. 302-3; Gray’s “* Turton,” 1857; p. 262.

EZ

212 THE DISPERSAL OF SHELLS.

In the “ British Conchology,” in 1862, Mr. Jeffreys
expressly inclined to a belief that the creature ought to
be looked upon as a native of Britain, and of the whole
of the North of Europe, adding that if, as he believed,
“the indigenousness of the Drezsséna as regards this
country should hereafter be established, the ingenious
theories which have been put forward to account for the
mode of its transport across the seas will not require
further discussion.” The circumstance ofits not having
been noticed here before 1824, and then only in a
metropolitan locality, he remarked, did not preclude the
possibility of its having previously existed in some
other part of the country, its not having been previously
recorded as British rather proving a want of observation
or opportunity than its non-existence. As tending to
confirm this belief, he pointed out—that /e/zx cartuszana,
H. obvoluta and Clausilia rolphit, “all of which are con-
spicuous land-shells,” were unknown to the observant
Montagu, although not uncommon in some parts of this
country, and clearly indigenous, and many similar
instances might be cited, he said, on this point, as well
as with respect to the sudden and unaccountable
appearance and disappearance of certain species in
particular spots: that the Puna fluviatilis of Sander
was doubtless our Dyvezssena, which was therefore found
in the interior of Germany before 1780, inhabiting
streams which flowed into the Rhine: that in draining
the Lake of Haarlem, which apparently never com-
municated with any port or harbour, it was found in
abundance, and it had been discovered, also, in an

DISPERSAL BY MAN. 213

inland lake near Copenhagen ; that, finally, it had been
recorded by M. Charles D’Orbigny as one of the fossils
occurring in an upper tertiary deposit in the north of
France, and on this ground he (Jeffreys) was not with-
out hope that it might be discovered in the correspond-
ing strata inthis country.’ From various considerations,
however, this idea that the creature had always existed
in Britain, and had been accidentally overlooked by all
our naturalists previously to 1824, seems to me in the
highest degreeimprobable. The “ British Conchology ”
has long been a standard hand-book, and the views of its
author have largely influenced the minds of those who
have attended to the mollusca, but the very general
impression that D. polymorpha is a foreign importation
has not been dispelled. There must be good reason for
supposing it to have been introduced, otherwise I cannot
think that this view, put forward by Mr. Sowerby in
1824, would have been endorsed by Gray in 1825, by
Fleming in 1828, by Berkeley in 1836, by Alder and
Strickland in 1838, by Captain Brown in 1844, by
Forbes and Hanley in 1853, as well as by Woodward
and nearly all other writers, even including Reeve,
Tate, and Harting, whose works were issued sub-
sequently to the date of the volume in which Mr.
Jeffreys’ views were expressed. With regard to Helx
cartusiana, H. obvoluta, and Clausilia rolphiz, the three
“conspicuous land-shells” mentioned by Jeffreys as
clearly indigenous yet unknown to Montagu (who
published his “ Testacea Britannica ” in 1803), it must be
1 J. G. Jeftreys, “ British Conchology,” i. (1862), pp. 47-59.

214 THE DISPERSAL OF SHELLS.

remarked that they are all local species, and their
presence in this country can hardly be said to be con-
spicuous, for an observer, unguided by published
records, might well collect for years, or for a life-time,
without finding them, while on the other hand, D.
polymorpha is “so generally distributed that it cannot
fail to attract the attention of the collector.” ' Dr.
Morch, it is true, has argued that there is no reason for
supposing the creature’s non-existence in the west of
Europe previously to 1824, maintaining that it is
Sander’s Pinna fluviatilzs, but eminent conchologists have
denied that this is the case.” In the errata given at the
end of his volume, Jeffreys corrects the. statement
respecting the specimen referred to by D’Orbigny, which,
it appears, was recent, not fossil. The animal is known,
however, to have occurred anciently in North Germany,
in the diluvium of East and West Prussia, etc., but it is
there supposed to have been driven away, by unfavour-
able climatic conditions or otherwise, and to have
subsequently regained its former range by immigration.’

This species has frequently been referred to as illus-
trating the rapidity with which a mollusc may become
naturalized and diffused over a large area, and it is
even said to flourish in this country, under certain con-
ditions, greatly in excess of any native form.’ In

9

* Harting’s “ Rambles,” 1875, p. 41.

* See 0. A. L; Mérch, * Ann, and: Mas: Nat. Hist. (3s
(1867), pp. 82-4.

3 See “ Ann. and Mag. Nat. Hist.,” (6), ix»*(1892), p. 160.

* See H. H. Higgins, ‘‘ Address to Liverpool Nat, Field-club,
30 January, 1891,” p. 7.

DISPERSAL BY MAN. 215

1838, Mr. Strickland pointed out thet it multiplied
with astonishing rapidity, and, in a few years, he
thought, it would probably become one of our com-
monest molluscs. On visiting the Commercial Docks,
the original British habitat, in or about that year, he
saw a vast number of the shells in the depdt for
bonded timber ; at least one generation, evidently, had
passed away since the introduction, for dead-shells
were scattered over the bottom and served as points of
attachment for the byssi of the living; it would be
interesting, he remarked, to watch the gradual spread
of the species over the kingdom.! About the years
1831-3, we find, it was discovered in great numbers
at Goole, Yorkshire, in docks belonging to the Aire and
Calder Company, opened only three or four years pre-
viously. On the water being drawn off a few feet
lower than usual, the walls were seen to be ‘‘completely
_ covered” with shells, pecks of which might have been
procured. One of the docks was used entirely for the
bonding of foreign timber which frequently remained
there for some months before being re-shipped or floated
upon rivers and canals to various inland towns, and
during such periods, it is said, those pieces which ad-
joined the sides or touched the bottom were sure to
become covered with shells, which, thus, might easily be
carried to the various districts communicating with the
canals of the Aire and Calder Company.* About this

1H. E. Strickland, Charlesworth’s “ Mag. Nat. Hist.,” (n. s.), 11.

(1838), pp. 362-3. |
* R. J. Bell, “ Zoologist,” i. (1843), p, 253.

216 THE DISPERSAL OF SHELLS.

time, also, it was observed in Scotland, where, as men-
tioned by Dr. J. Scouler of Glasgow in 1833, it was
found living in vast abundance towards the eastern
extremity of the Clyde and Forth Canal; and, in the
following year, its discovery in the Union Canal, near
Edinburgh, was communicated to the Wernerian Society
by Mr. Stark.’ In 1836, the Rev. M. J. Berkeley found
the species in plenty on the piers of a bridge crossing
the Nen at Fotheringhay, Northamptonshire, and also,
a little higher up the same river, on the stones of a
small overfall at Tansor. He had formerly examined
the stream carefully, for several successive years, both
above and below these spots, without detecting the
creature, and had reason to believe that its establish-
ment was of recent date (subsequent to 1828) ; pro-
bably, he thought, it had been introduced from Wis-
beach on timber.” As to the date of its detection at
the last mentioned place I know nothing. In 1837,
Mr. Strickland discovered it in the Avon, near Evesham,
in Worcestershire. From 1828 to 1834 he was in the
habit of conchologizing frequently in that river, and
could scarcely have failed to detect the creature ifit had
then lived there, but none occurred. In 1837, however,
after an absence from England of two years, he was
greatly surprised on finding several of its shells among

1 J. Scouler, Loudon’s “ Mag. Nat. Hist.,” vi. (1833), p. 532;
Stark, as quoted by Berkeley, Loudon’s “ Mag. Nat. Hist.,” ix.
(1836), p. 572.

* M. J. Berkeley, Loudon’s “ Mag. Nat. Hist.,” ix. (1836),
Pp. 572-3.

DISPERSAL BY MAN. 217

refuse on the banks of the river, and on further search
he found that the animal had become completely estab-
lished on the gravel of the river-bed ; several hundred full-
grown specimens were collected in the course of an
hour; “there is, therefore, clear evidence of the recent
introduction of this mollusc into the Avon, and of the
rapidity with which it has reached maturity and multi-
plied.” In the same or the following year, Mr.
Strickland observed the species in the canal between
Warwick and Birmingham, and, in a paper dated in
1838, he stated that it had been found also in the
canals near Wednesbury, in Staffordshire, and in the
Leam at Leamington ; the latter water is not navigable,
but is near a canal from which the Drezssena was pro-
bably introduced. In the same paper Mr. Strickland
stated that the animal had been “ planted” in some of
the waters near Bristol. About 1838, also, it was
detected in the Lesser Nen, which runs through the
town of March,’ and about the same time a specimen
was obtained in Nottinghamshire from unnavigable
water—an old mill-dam at Toton, supplied by the
Erewash, a small shallow stream joining the River
Trent nearly a mile from the place—that river being
the nearest navigable water ; afterwards (before 1846)
numbers were found “adhering to stones underneath
the water-fall of a pond at Lenton” in the same county,
to which they must have gone up a very small brook

’ -H. E. Strickland, Charlesworth’s “Mag. Nat. Hist.,” (n.s.), 1.

(1838), pp. 361-3.
4S, Smith, “Science Gossip,” for 1868, p. 238.

218 THE DISPERSAL OF SHELLS.

fully a mile from a canal, where, however, though the
observer had frequently searched, he had never found
them.! The animal was known to occur in streams
near Exmouth, in 1842. By 1843 it is said to have
found its way to various canals and other waters in
Leicestershire, and it had been discovered in immense
numbers in the Grand Trunk Canal at Burton, Stafford-
shire.” Captain Brown, in 1844, stated that it was
common in the Bridgwater Canal, and also “in many
places in the canal betwixt Manchester and Hull,” and
in that year it was noticed, also, on timber in the Ouse,
near St. Neots, Huntingdonshire.* We find, therefore,
that within twenty years from the announcement of its
discovery D. polymorpha was known in two localities in
Scotland, and in England in the counties of Surrey,
York, Northampton, Worcester, Warwick, Stafford,
Cambridge, Nottingham, Devon, Leicester, Somerset,
Huntingdon, and perhaps also in Lancashire, and it is
of course probable that its range at that time may have
been much wider, for I have not made a thorough
search for records; indeed, if a statement by Brown
(1844) is to be relied upon, it was then to be “ met with
in most of our canals.” It would be interesting to
ascertain the creature’s range at the present time,
but to find, and search, all the local lists and notes

" G. Wolley, “ Zoologist,” iv. (1846), p. 1420.

? L. M. Pratten, “ Science Gossip,” for 1868, p. 189 ; E. Brown,
“ Zoologist,” 1. (1843), p. 255.

’ T. Brown, “ Illustrations,” ed. 2, (1844), 'p.. 765 C,. Preatiee:
“ Zoologist,” v. (1847), p. 1651.

DISPERSAL BY MAN. 219

which have been published would be a tedious under-
taking, and might occupy the greater part of a lifetime!
According to a ‘‘ census of the authenticated distribu-
tion of British land and fresh-water mollusca,”' pub-
lished in 1889, it appears that specimens collected in
the following English and Scottish counties, or vice-
counties, have passed under the examination of the
Conchological Society’s referees :

Somerset, North. Staffordshire.

Surrey. Shropshire.
Middlesex. Leicestershire.
Berkshire. Nottinghamshire.
Oxfordshire. Derbyshire.
Cambridgeshire. Cheshire.
Huntingdonshire. Lancashire, South.
Northamptonshire. Yorkshire, South-West.
Gloucestershire, East. Yorkshire, Mid-West.
Gloucestershire, West. Renfrewshire.
Worcestershire. Lanarkshire.
Warwickshire.

No Welsh or Irish habitats are given.

The Drezssena is perhaps better fitted for dissemina-
tion by man and subsequent establishment than any
other fresh-water shell ; tenacity of life, unusually rapid
propagation, the faculty of becoming attached by a
strong byssus to extraneous substances, and the power
of adapting itself to strange and altogether artificial sur-
roundings have combined to make it one of the most

1 By J. W. Taylor and W. D. Roebuck, forming chapter v. of
the volume on “ Land and Freshwater Shells,” in Swan Sonnen-
schein’s “ Young Collector” series, 1889.

220 THE DISPERSAL OF SHELLS.

successful molluscan colonists in the world. The floating
of rafts of timber for commercial purposes from place to
place upon rivers and canals has doubtless greatly
_ facilitated its dispersal, and it may frequently attach it-
self to the bottoms of canal-boats ; possibly, also, from
having free-roving larve which live on the surface for a
time, it possesses considerable facilities for dispersal in-
dependently of man, and, as just mentioned, it has
found its way,in this country, to unnavigable water ;
every naturalist, the Rev. J. Dalton has observed, can
“cite examples of Drezssena polymorpha turning up
profusely in waters where it had hitherto been un-
known, and whither its progress could not be traced.” '
The creature is able to seize quickly upon new and un-
occupied water, and this must have greatly aided it in the
struggle with other forms. Asalready noted it was found
in immense numbers in new docks at Goole within three
or four years after they were opened. Some time ago,
a canal near Northampton was diverted by a railway
company, and after three or four years the stones form-
ing the new banks were found to be entirely covered
with the shells of this species In the neighbour-
hood of Gloucester, also, it is said to have appeared
within a few years after the opening of the Glouces-
ter and Berkeley Canal, and to have subsequently
increased greatly, so as to almost line the banks, from
the edge of the water to a considerable depth, from

' J. Dalton, “ Zoologist,” xix. (1861), pp. 7318-9.
2 W. D. Crick, as cited by Taylor and Roebuck, “ Journ. North-
ants.’ Nat. Hust. Soc.,” 11. (1é85); p. 212. .

DISPERSAL BY MAN. 221

one end of the canal to the other, a distance of sixteen
miles.’

PLANORBIS DILATATUS Gould. This American mol-
lusc was first detected in the British Isles, and indeed in
Europe, during the summer of 1869, in the Bolton
Canal, at Pendleton, on the west side of Manchester,
and in the autumn of the same year it was discovered
also in the Gorton Canal, on the east side of the city,
the two localities being about five miles apart. In both
cases, the shells were found close to cotton-mills,
where the waste from the cleaning or blowing machines
was ejected over and about the canals—a good deal
finding its way into the water—and where the
warm water from the engines was discharged. Mr.
Thomas Rogers, the discoverer, and Mr. Jeffreys, who
determined the species, suggested that the animal, or
its eggs, had been imported from America with cotton,
and it was pointed out that the best cotton is culti-
vated in river-bottoms, and that the crop, when picked,
is spread out to dry. We have already seen that many
molluscs possess great tenacity of life, and, as mentioned
by Mr. Jeffreys, certain species of P/anorbis, in times of
adversity, close the mouths of their shells with an
epiphragm, and thus protected they keep alive out of
water for weeks and even months; so that it may be
assumed that in all probability living specimens of P.
dilatatus could survive a voyage from America to Eng-
land ; and after examining the waste from the machines,

! R. Tate, “Land and Fresh-water Mollusks,” 1866, p. 23.

222 THE DISPERSAL OF SHELLS.

Mr. Rogers thought it likely that they might pass the
blowing process undamaged. This view of the creature’s
introduction was questioned, however, by Mr. Anthony,
an eminent American conchologist, who was of opinion
that the species did not inhabit the cotton-growing
states. But Mr. Rogers further suggested that the
shells might have attached themselves to submerged
cotton used for defensive purposes—as barricades for
steam-boats or river defences—during the war between
the North and South, and that it might have been trans-
ported to this country during the cotton famine brought
about by the war, when Lancashire people, it is said,
were glad to get material in any condition, wet or dry,
blockade-run, or otherwise; and this theory seems to
have been generally accepted by British concho-
-logists. The animal seems to have multiplied consider-
ably in its new home: a few years after its discovery,
when one of the canals was run dry for repairs, Mr.
Rogers saw it, in countless numbers, extending about a
mile from where it originally occurred ; unfortunately,
however, nearly all these vast numbers were killed when
the brickwork of the waterway was repointed with
mortar containing a good quantity of lime. In the
Conchological Society’s list the name of the species was
enclosed in brackets, in order to denote that its
claim to rank as British was not considered to be
thoroughly established ; but Mr. Collier, of Manchester,
commenting on this in 1884, expressed the opinion that
the creature might be regarded as fully naturalized, and
the brackets have been removed, I find, in a new edition

DISPERSAL BY MAN. 223

of the list published in 1892.' Mr. Collier stated that
he had not tried for some years to get the creature at
Pendleton, for when last there the towing-path side of
the canal had been undergoing repair, a large quantity of
new stones having been put down at the place where
the shells used to occur, and the water “had become
much worse”; at Gorton, however, he had found it
repeatedly—some in 1883—but not in great quantity,
and not in the original habitat near the cotton-mill,
but about a quarter of a mile away where the water
was quite cold. In 1886, seventeen years after its
discovery at Manchester, the creature was found at
Burnley, in the same county, in the “paper-works
lodge” which has no connection whatever with any
cotton-blowing machinery, the nearest cotton-mill being
half a mile distant. In the following year, it was taken,
also, in another Burnley locality, this time under condi-
tions similar to those at Manchester, Mr. F. C. Long
being reported to have found it, in thousands, in the
canal opposite Temple’s factory, where cotton refuse is
blown into the water. It occurred along the canal for
over a mile’in the direction of Hapton, but was most
plentiful opposite the mill. Mr. Rogers thinks it im-
probable that this colony was derived from Manchester,
and inclines to the belief that it was founded by
shells or ova imported from America, in the same
manner, and at about the same time, as those introduced
into the Manchester canals, and he thinks it probable,

* “Journ. of Conch.,” vii. (1892), p. 55.

224 THE DISPERSAL OF SHELLS.

also, that the two colonies near that city were distinct
and separate introductions. The presence of the crea_
ture, in thousands, in a cistern on the top of an engine-
house at Burnley, “about sixty feet above the canal,”
has already been referred to. The shells had been
pumped from the canal, no doubt, with the water
supplying the cistern. As suggested by Mr. J. R.
Wildman, the creature was probably carried by birds to
the ‘‘ paper-works lodge.”

These are the only places in Britain, as far as I am
aware, in which P. d/atatus has been detected. It is
now nearly a quarter of a century since the first
specimens were found, and it is surprising that the
creature has not, by this time, extended its range beyond
the limits of Lancashire, the county to which it was
imported ; within that county, as we have seen, it has
been found in four separate localities.’

PHYSA ACUTA Drap. In the spring of 1860, speci-
mens of this pond-snail were found by Mr. A. Choules
in a water-tank in the Royal Gardens at Kew, whither
they are supposed to have been imported from abroad
with aquatic plants.” The shells, it is said, were un-

' J. G. Jeffreys, “ Ann. and Mag. Nat. Hist.,” (4), iv., (1869), pp.
341-2; T;} Rogers, “Science Gossip” for 1870, p.. 138; ~ Older.
Journ. Conch.,” i. (1875), p. 81; ‘“ Conchologists’ Exchange,” 1.
(1887), p. 63; “ Journ. of Conch.,” v. (1887), pp. 218-220; R. Rim-
mer, “ Land and Fresh-water Shells,” 1880, pp. 47-9; E. Collier,
“Journ. of Conch.,” iv. (1884), p. 287 ; J. Bates, “ Journ. of Conch. =
v. (1887), p. 221; J. R. Wildman, “ Science Gossip,” xxiv. (1888),
p. 210; F. C. Long, “ Science ‘Gossip,” xxiv: (1683) p27 Zam

? It is doubtful whether P. acuta was previously known in

DISPERSAL BY MAN. 225

distinguishable from specimens, in the British Museum,
collected in Cuba, St. Thomas, and St. Croix. The
Rev. A. H. Cooke, in 1882, mentioned having taken
the animal abundantly in the Victorza regia tank in the
Royal Gardens, and I hear from Mr. J. B. Davy that
Mr. Watson, the assistant curator, has seen it in “the
lily-house, formerly called the Vzctoria regia house.”
Probably the creature is not confined to any particular
water within the gardens. The tank in which it was
originally found is said to have been exposed in the open
air to the inclemency of our winters, and this is not the
case either with the lily or Vectoria regia tanks, both of
which are in warm houses. Mr. Cockerell, in 1885,
reported that the species was “still abundant ina
water-lily tank at Kew.” In 1890, specimens were
obtained by Mr. Grocock and Mr. Jenkins from the
Royal Botanic Society’s gardens, Regent’s Park. They

Britain. According to Canon Norman the description in the
Linnean Transactions of the “Bulla rivalis” of Maton and
Rackett, 1804,—said to have been found by Mr.James Hay in
Hampshire—sufficiently accords with P. acu¢a (to which shell in-
deed Moquin-Tandon had assigned it as a synonym), and he
remarks that Montagu, who was well aware that Bulla rivalis was
a common West Indian shell, did not question the discovery of
Mr. Hay. “Physa acuta” is given also in Captain Brown’s
“ Tilustrations,” 1844, as having been “found in Anglesea, Wales,
and first identified as British by J. Sowerby, Esq.,” but it is now
stated that the shells, in this case, ought to have been referred to
a variety of our common British species Physa fontinalis. See
Matow and: ikackett, “* Trans. Lin. Soc.,” viii: (1807), p. 126; T.
Brown, “ Illustrations,” ed. 2, (1844), p. 30; Forbes and Hanley,
“ British Mollusca,” iv. (1853), pp. 145-6; A. M. Norman, “ Ann,
and Mag. Nat. Hist.,” (3), vil. (1861), pp. 115-16.

Q

226 THE DISPERSAL OF SHELLS.

occurred, in houses, in a tank containing seedlings of
Victoria regia, and in glass vessels and tanks in which
Vallisneria spiralis and other aquatic plants were
srowing. Very possibly they were introduced from
Kew, for Mr. W. Sowerby tells me that the Royal
Botanic Society is constantly exchanging plants of all
kinds with the Royal Gardens. The creature has
occurred, also, in one locality in Scotland, having been
collected, in 1887, from three or four ponds in the
grounds of the Banner Mill Company at Aberdeen.
These ponds, containing warm water discharged from
the mill, are tenanted also by gold-fish, various aquatic
plants, and the common wandering pond-snail (Lzmu@a
peregra). Mr. W. D. Rae, who first collected specimens
in this locality, stated in 1891 that the foreman had
seen the creatures in the ponds ever since he came to
the mill thirteen or fourteen years previously ; sixty-
two living specimens were collected and forwarded to
Mr. Rae, in London, in 1890. Presumably the animal
was introduced either with aquatic plants or with the
gold-fish.'

A specimen of Lzthoglyphus naticoides Fér., a fresh-

" A. Choules, “ Zoologist,” xviii. (1860), pp. 72, 78-9; A. M.
Norman, “Ann. and Mag. Nat. Hist.,” (3), vii. (1861), pp. 114-5 ;
“* Zoologist,” xix. (1861), p. 7355; “ Ann. and Mag. Nat. Hist.,” (3),
ix. (1862), p. 186; J. G. Jeffreys, “ British Conchology,” i. (1862),
p. 100; A. H: Cooke, “ Journ. of Conch.,” iil. (1882), Baer
T. D. A. Cockerell, “ Zoologist,” (3), ix. (1885), p. 14; A. J. Jenkins,
Journ. of Conch.,” vi. (1890), pp. 270-1; W. D. Rae, “‘ Science
Gossip,” xxvii. (1891), pp. 89-90.

DISPERSAL BY MAN. 22 7f

water shell which inhabits the Danube, was once found,
according to Mr. Jeffreys, in the bottom of a fishing-
boat at Cullercoats, Northumberland.'

Of terrestrial species, the Conchological Society’s
list (1883) contains the names of eighty-six species, of
which but three, Zestacella mauget, Helix villosa, and
Bulimus goodallii can be reasonably looked upon as
introduced by human agency ; and two of these names
(Helix villosa and Bulimus goodallit) are included within
brackets, showing that the creatures’ claims to be re-
garded as British were not considered to be established,
and felix villosa is known to have been placed in the
catalogue on the faith of a colony which is now extinct.
That the terrestrial molluscan faunas of the world have
been much more influenced by man’s dispersal than
fresh-water faunas cannot be doubted, but this hardly
seems to have been the case in ourown country. It may
be remarked, however, that the indigenousness of at least
six species (in addition to the three above mentioned)
has been questioned by various authors, and other
authors have probably doubted the right of still other
species to rank as native. The evidence which can
be got together does not, as far as I am acquainted
with it, warrant the conclusion that any of these have
certainly been imported, but it is of course fosszble that
this may have been the case with some of them. Of
the three species generally regarded as introduced,

) Jeffreys, “British Conchology,” i. (1862), p. 70.
2

228 THE DISPERSAL OF SHELLS.

flelix villosa having died out need not be dealt with
at present, but it will be well to recall the facts con-
nected with the others, 7. maugez and B. goodalli ;
and I have also added a note on Felx terrestris
and Pupa cinerea, names which have appeared (in
brackets) in the Conchological Society’s new list of
1892.

TESTACELLA MAUGEI Fér.—a “snail-slug ”—which
seems to bea native of the south-west of Europe, and
occurs also in Madeira, the Canary Isles, etc., was first
noticed in England between the years 1812 and 1816,
in Messrs. Miller and Sweet’s nursery (now Garraway’s),
near Bristol, by Mr. T. Drummond; and tir i
Miller, in a paper published in 1822, expressed the
opinion that it had been introduced with foreign plants ;
but he added that it propagated freely in the open
ground, increasing much in rich soil, and could “no
longer be. considered, as an alien.” |; Mr. J. Berge
Sowerby, as appears by a note in Loudon’s Magazine in
1833, also regarded the animal as introduced, the
Original specimens having been imported, he thought,
probably by accident, along with plants from Teneriffe
or elsewhere ; Mr. Alder, in 1838, while regarding 7.
haliotidea as a native, also thought the case different
with the present species “which belongs to a warm
climate, and has evidently been introduced along with
exotic plants;” Gray (1840 and 1857), Forbes and
Hanley (1853), Jeffreys (1862), Reeve (1863), Tate
(1866), and other authors agree in this view, and I am
not aware that any writer has maintained that the

DISPERSAL BY MAN. 229

creature is a native.' Jeffreys (1862) remarked that it
was still to be found in considerable numbers in the
original habitat, and that it had been observed also in
other parts of Somersetshire, as well as at Plymouth,
Cork, and in his own garden at Norton, near Swansea,
which was occasionally supplied with plants from
Miller and Sweet’s nursery. The occurrence of the
animal zz fields near Devizes is noted in Woodward’s
“Manual,” but I finda statement, in 1867, that it is found
in that neighbourhood only when gardeners are preparing
their ground or digging up their crops. The Concho-
logical Society’s referees have seen specimens from
West Cornwall, North Somerset, Dorsetshire, Berkshire,
West Gloucestershire, Glamorganshire, Pembrokeshire,
Waterford County, and the Channel Isles, and without
doubt the creature has obtained a permanent footing.’
BULIMUS GOODALLII Miller—This snail, which has
now a wide range, is said to bea native of the West Indies,

1 Since writing I find, however, that Dr. Scharff, in a recent
paper on the distribution of the British land and fresh-water mol-
lusca, seems to have regarded this animal as indigenous to our
soil, and it is of course quite Josszb/e that such is the case. See
“ Conchologist,” ii. (1892), pp. 1-6.

2 J.S. Miller, “ Ann. Phil.,” (n. s.), iii. (1822), p. 380; J. Fleming,
“ British Animals,” 1828, p. 257; J. De C. Sowerby, as cited by
J. D[enson], Loudon’s “ Mag. Nat. Hist.,” vi. (1833), p. 45 ; J. Alder,
“Mag. Zool. and Bot.,” ii. (1838), p. 105; Gray’s “ Turton,” 1840,
p. 5; 1857, p. 292; “ Forbes and Hanley,” iv. (1853), p. 28; Jef-
freys, “ British Conchology,” i. (1862), pp. 147-8 ; Reeve, “‘ Land and
Fresh-water Mollusks,” 1863, p. 32; Tate, ‘‘ Landand Fresh-water
Mollusks,” 1866, p. 89 ; Woodward’s “ Manual,” ed. 2, (1866), pp.
298-9 ; J. J. Fox, “ Science Gossip” for 1867, p. 89.

230 THE DISPERSAL OF SHELLS.

and is believed to have been introduced into England
with pine-plants. It has been found in several widely
separated places in this country, but can hardly be
said to be acclimatized, for I believe it lives chiefly or
exclusively in greenhouses, stoves, pits, etc. ; Dr. Turton,
however, in 1826, considered it “to be naturalized as
much as the 7estacellus mauget.” Like that species it
was first detected in Miller and Sweet’s nursery, near
Bristol; at about the same time -or a little laterj;7e
during the years 1816-17. The species was originally
described from the British specimens, as Helzx goodalliz,
in a list of shells occurring in the environs of Bristol
published by Mr. Miller in 1822. The author men-
tioned that he found the creature, in 1817, on boards
lining a pine-bed. Mr. Drummond, in a letter to Dr.
Fleming, however, stated that the animal was first
pointed out by him in 1816, and it appears to have
occurred in some plenty, for, when a supply was required,
he merely placed a flat board upon the surface of the
tan, with a few dead worms beneath it, and this, in a
few days, became “covered” with shells. Mr. Jeffreys
(1862) stated that the creature was still very common
on the tan in the pineries in which it was discovered ;
he certainly erred, however, in saying that it had not
been noticed elsewhere in this country, for Mr. Alder
(1838) had mentioned having seen specimens from a
garden in the neighbourhood of Manchester, and in
Gray’s “Turton” of 1840 the animal was said to be
common in some places in the neighbourhood of
London, especially in Kensington Palace Garden. Mr.

DISPERSAL BY MAN. 231

W. Borrer tells me that he received specimens many
years ago from the Royal Gardens at Kew. I hear
from Mr. G. F, Wilson that a colony—first noticed, he
thinks, nearly thirty years ago—used to exist in a
cucumber-house in a garden at Wandsworth, from
whence specimens were submitted to the Scientific
Committee of the Royal Horticultural Society and
identified by Mr. Berkeley ; fosszbly the creature was
introduced with pines, for there was formerly a pine-
house in the garden. Weybridge has more than once
been quoted as a locality for this snail, which has been
erroneously stated—in the “Garden” newspaper for
instance—to have occurred in Mr. Wilson’s garden
there; the mistake arose, no doubt, from the fact that
while living at Wandsworth, Mr. Wilson had a garden
at Weybridge, well known in the horticultural world.
The snail seems to have occurred, in 1876,in large
numbers, in a cucumber-house-at Gunnersbury. In
1881, Mr. Ashford stated that it had flourished for many
years in the orchid-houses of Mr. Day of Tottenham,
from which locality he had procured living specimens
in 1867. Mr. F. W. Wotton mentions having received
it, in 1886, from Mr. Chapman’s greenhouse, presum-
ably at or near Cardiff ; and in one of the hot-houses in
a garden at Croydon it was recently found in such vast
numbers that all the soil in the house had to be collected
and burned. Possibly the animal is now very generally
distributed, in greenhouses, etc., throughout a great part
of the country.’

1 Turton, “ Zoological Journal,” ii. (1826), p. 565; J. S. Miller,

232 THE DISPERSAL OF SHELLS,

HELIX TERRESTRIS Penn.—In. the new list of 1892,
as just mentioned, the name of an additional He/zx has
appeared, the present species, the Helix elegans of
Gmelin, finding a place in the catalogue; its claim to
rank as British, however, is not considered to be
thoroughly established. The animal is common on the
shores of the Mediterranean, especially in the south of
France and Corsica, and is quoted also, according to
Mr. Edgar Smith, from various localities in Spain, the
Balearic Islands, Algeria, Italy, and Sicily, but it does
not appear to range northward. It has recently been
found, however, in the county of Kent, where what
appears to be a thriving and well-established colony
was discovered, in 1890, by Mrs. McDakin, in the
neighbourhood of Dover. The colony extends for
about half.a mile on a chalky bank, by the side of a road,
in a retired locality some miles from the town, and
far from houses or gardens. When discovered the
animals were on the tall grasses in some numbers;
and afterwards, when winter had set in, Mr. C. S. B.
Cox was able to procure many living specimens, with
their mouths closed with an epiphragm, lying almost

“Ann. Phil.,” (n. s.), iii. (1822), p.381; J. Fleming, “ British Animals,”
1828, p. 266; Jeffreys, i. (1862), pp. 239-40 ; J. Alder, “‘ Mag. Zool.
and Bot.,” ii. (1838), p. 110; Gray’s “ Turton,” 1840,p. 6; J. E.
Daniel, “ Garden,” x. (1876), p. 542, and editorial note ; “ Quart.
Journ. Conch.,” i. (1874-8), p.246; and see also “Garden,” x.
(1876), p. 525; C. Ashford, “‘ Journ. of Conch.,” iii. (1881), p. 240,
and ‘Science Gossip ” for 1868, p. 17; F. W. Wotton, “ Trans.
Cardiff Nat. Soc.,” xx. (1888), p.36; K. McKean, “ B. goodalliz at
Croydon,” “ Journ. of Conch.,” vi. (1889), p. 84.

DISPERSAL BY MAN. 233

invariably among lumps of frozen chalk, and sometimes
frozen to the ground ; many dead shells, of all ages,
and in every condition of preservation, were also found.
The position of the colony, in Mr. Cox’s opinion, is
suggestive of its having been intentionally introduced by
a naturalist’s hand, probably some few years ago. Mr.
Smith agrees that the species is doubtless an importa-
tion, and thinks it probable that specimens recorded
as having been found at Beauvais, in the north of
France, were also introduced from a more southern
locality. Mr. Carrington, writing in the Feld, also
agrees, remarking that there can be little doubt but
that our colony was introduced, “either purposely or
by accident.” A specimen of this shell was in Dr.
Turton’s collection of British shells marked “ Corn-
wall ;’ the birth-place, Mr. Jeffreys remarks, of many
spurious or exotic shells. The Zvochus terrestris of
Pennant is said to have been discovered in the moun-
tains of Cumberland, but it is perhaps doubtful whether
that author really intended to indicate the present
species, and I have used his name merely because the
compilers of the Conchological Society’s list have done
so. It is curious to find that a colony of H. derrestris
was discovered, in 1875, in a churchyard in Charleston,
South Carolina.’

1 C. S. B. Cox and E. A. Smith, “ Journ. of Conch.,” vi. (1891),
pp: -377-9; |. b> Carrington, “Field,” 29 August, 1891, p- 334;
“ Jeffreys,” i. (1862), pp. 215-16; Pennant, “ British Zoology,” iv.
(1777), p. 111; and see Gray’s “Turton,” 1840, p. 9; Mazyck,

“Proc. Acad. Nat. Sci. Phila.,” 1876, p. 127, quoted by Mr. Smith
as above.

234 THE DISPERSAL OF SHELLS.

PUPA CINEREA Drap. is also included, as doubtfully
British, in the new catalogue, “in the faint hope that its
claim to inclusion in the British list may be strengthened
by further observation.” Jeffreys, in 1831, stated that
a specimen had been found by Mr. Gray among the
rejectamenta of the Thames at Battersea, and he himself
had detected an imperfect specimen at the same place.
But, in 1840, Gray treated the species as having been
improperly regarded as British, stating that he had
introduced it into the fauna in error, a specimen having
been accidentally mixed by Dr. Leach with some other
shells collected at Battersea,and Mr. Jeffreys’ record,
also, he thought, was probably erroneous. More re-
cently, however, specimens all presumably dead—two
of which have been identified by Mr. J. W. Taylor—are
reported to have been taken, on four occasions during
twelve years, near Accrington, Lancashire. Their
presence may most probably be attributed to man’s
dispersal, but some naturalists think it unlikely that
they were thus introduced, for the locality is described as
“ well-wooded out-of-the-way.” *

As already stated, there are at least six other land-
shells in the British list which have been regarded by
authors as probable or possible importations. Very
likely all of them are indigenous, but, be this as it may,

1 “Journ. of Conch.,” vii. (1892), pp. 54-62 ; J. G. Jeffreys, “‘ Trans.
Lin. Soc.,” xvi. (1833), p. 514; Gray’s “ Turton,” (1640), p. 13 50
Wrigglesworth, “ Science Gossip,” xxv. (1889), p. 281 ; and see also
“ Naturalist,” 1892, p. 336.

DISPERSAL BY MAN. 235

it is certain that the contrary cannot now be positively
asserted; we are unable to distinguish our native
fauna with any degree of certainty.'. A few notes on the
species in question, beginning with the famous and much
discussed edible or Roman snail may be given.

HELIX POMATIA L. Merret and Lister, two very
early writers on the British fauna, do not seem to have
doubted the indigenousness of the edible snail to this
country ; the first-named author in his “ Pinax,” pub-
lished in 1667, mentioned the animal (according to
Gray) without any note, as found in Sussex ; and Lister
in his “ Historie Animalium Angliz,’ dated in 1678,
mentioned it, as “ Cochlea cinerea, maxima, edults, etc.,”
as one of the largest of the land-shells of our island,
and, as far as I am aware, he never suggested that it
might have been imported; it was plentiful, he said, in
Hertford and other places in the south of England, but
he had never found it in the north. Ninety-nine years
later, strangely enough, we find Pennant (in 1777)
naming the species “exotic,” and stating that it had
become naturalized in the southern counties, having been
introduced, it was believed, by the eccentric Sir Kenelm
Digby, either for medical purposes or as a food; “‘tra-
dition says that to cure his beloved wife of a decay
was the object.” In 1778, Emanuel Mendes da Costa
named the creature “ Italian,” and gave a curious and
interesting dissertation as to its introduction and subse-
quent dispersal :

1 See Rimmer’s “ Land and Fresh-water Shells,” (1880), p. 145.

236 THE DISPERSAL OF SHELLS.

“ This snail is not zva¢genous, or originally a xadtive of
these kzxgdoms, but a naturalized species, that has throve
so well as now to be found in very great quantities. It
was first imported to us from /¢a/y about the middle of
last century, by a scavorr vivre, or Epicure, as an an [sic]
article of food. Mr. Audrey informs us, it was a Charles
floward, Esg., of the Arundel family, who, on that
account, scattered and dispersed those snails all over
the downs, and in the woods, etc., at Albury, an antient
seat of that noble family, near Ashted, Boxlull, Darking,
and Ebdisham or Epsom, in Surry, where they have
thriven so much that all that part of the county, even to
the confines of Swssex, abounds with them; insomuch
that they are a nusance, and far surpass in numbers the
common, or any other species of English snails.

“ The Epecures, or scavotr vivre, of those days, followed
this luxurious folly, and the snails were scattered or
dispersed throughout the kingdom, but not with equal
success; neither have records transmitted to posterity
the fame of those qworthies equal to the Roman Fulvius
flirpinus, except of fio, the one Sir Kenelin Digby, who
dispersed them about Gothurst, the seat of that family
(now of the Wrights) near Newport Pagnel, in Bucking-
hamshire, where probably they did not thrive much, as
they are not very frequent thereabout: the other worthy
was a Lord Hatton, recorded by Mr. Morton, who scat-
tered them in the coppices at his seat at Kzrby, in
Northamptonshire, where they did not succeed.

“ Dr. Lester found them about Puckerzdge and Ware, in-
Flertfordshire ; and observes they are abundant in the

DISPERSAL BY MAN. 237,

southern parts, but are not found in the xorthern parts
of thes zsland.

“In Surrey, as before mentioned, they abound; in
several other counties they are not uncommon, as in
Oxfordshire, especially about Woodstock and Bladen ;
in Gloucestershire, in Chedworth parish, and about Frog
Mill; in Dorsetshire, etc., but I have never heard that
they are yet met with in any of the zorthern counties.”

Pulteney, in his Dorsetshire catalogue of 1799, gave
the species along with the other e/ices, and made no
statement as to whether he considered it indigenous or
introduced, but, after stating that it had been recorded
for Dorsetshire, added, “happily this kind does not
thrive well in England.” He neither followed Pennant
in naming the creature “exotic” nor Da Costa in calling
it “Italian,” giving instead the more appropriate
English name of “ Edible snail ;” but Donovan, following
in 1801, again called the animal “Italian or exotic,”
and agreed with Pennant and Da Costa in regarding it
as an importation. By whom it was first introduced,
he remarked, was uncertain. In 1803, Montagu, in his
“Testacea Britannica”—“ next to Miiller one of the
best works on land and fresh-water shells ”—wrote ina
similar strain, remarking that the creature was “first
introduced about the middle of the sixteenth century
either as an article of food or for medical purposes,”
and repeating the stories told by Pennant and Da Costa
respecting the reputed doings of Sir K. Digby and Mr.
Howard. Dr. Maton and the Rev. Mr. Rackett, in
their catalogue read at the Linnean Society in 1804,

238 THE DISPERSAL OF SHELLS.

also mentioned the creature’s supposed importation by
Mr. Howard; and they referred to its presence in the
additional counties of Wiltshire and Kent. In 1817,
Dillwyn gave England as one of the habitats of this
snail, but without any statement as to its introduction ;
and Turton, in his “Conchological Dictionary of the
British Islands,” in 1819, gave the “edible snail-shell,”
along with the other species of the genus, presumably asa
native, making no statement as to whether it was to be
regarded as indigenous or imported ; but, even in 1830,
we find the statements both as to Sir K. Digby and Mr.
Howard again repeated, this time by Dr. George John-
ston, and they have been further repeated no doubt
by other writers. Fleming, in 1828, was probably in-
clined to consider the animal as an indigene, but he
made no direct statement to this effect, simply observing
that some had conjectured that the species was intro-
duced into England by Mr. Howard about the middle
of the sixteenth century ; it is noteworthy, however,
that he did not give it as a “naturalized species,” like
B. goodallit, nor as a “straggler” like D. polymorpha.
Turton seems to have been the first to publish an ex-
press opinion that 7. pomatia was in all probability a
native species, which he did, in his “ Manual,” in 1831,
remarking that at one period the animals appeared to
have been admitted at our own tables, “as Lister in
his ‘ Historia Animalium Angliz,’ p. 111, tells us the
manner in which they were cooked in his time,”
and Ben Jonson, in “ Every man in his Humour,” men-
tions the dish as a delicacy—‘“ neither have I dressed

DISPERSAL BY MAN. 239

snails or mushrooms curiously before me,’—and these
circumstances, he added, “ suppose their long foreknown
establishment in this country, and together with their
general diffusion in certain soils, incline us to consider
them as indigenous, and not introduced by Sir Kenelm
Digby for medical purposes, nor, according to Da Costa,
by Mr. Howard as an article of food.” This conclu-
sion is probably a just one, but it may be observed
that Lister’s remarks do not at all imply that the
creature was admitted to English tables in early times ;
Da Costa understood Lister to say that the animals
were cooked, in the manner indicated by him, in con-
tinental Europe, and no doubt rightly, for the often
quoted remark, “ Coquuntur ex aqua fluviatili, et adjectis
oleo, sale et pipere, lautum ferculum przparant,”
although a separate paragraph, follows immediately
upon the statement, “In Gallia Narbonensi admodum
vulgo eduntur. Item Parisiis tempore quadragesimali
magna quantitate vzeneunt.”

Turton’s view received support, in 1839,-from Forbes,
who, in his “ Report on the distribution of Pulmoniferous
Mollusca in the British Isles,” expressed the opinion
that there were good grounds for regarding the creature
as indigenous: “when we consider the partiality shown
by that shell for the newer calcareous strata in all parts
of Europe, and the geological correspondence of its
British and continental habitats, I think there can be
but little question of its.indigenousness ;” Gray, in both
his editions of “ Turton” (1840 and 1857), followed the
view originally expressed by the author in the first

240 THE DISPERSAL OF SHELLS.

edition, and Captain Brown (1844) also regarded the
animal as a native, as did Forbes and Hanley in their
“ British Mollusca” (1853) ; and indeed, Iam not aware
that any post-Turtonian author, with the exception
perhaps of Reeve,' has reverted to the older belief of
Pennant, Da Costa, and their followers. Leach) i
seems, held the view that this snail (Pomata antz-
guorum he called it) was an indigenous species long
before the publication of Turton’s “ Manual,” for he re-
ferred to it as such in his “ Synopsis,” published by Gray
in 1852, which is said to have been in course of printing
when the author was prevented from completing it by
ill health in 1820, and to have been re-printed and issued
exactly as it was left; the creature’s general diffusion
in a certain soil, he remarked, seemed to refute the
notion, held by many, that it had been imported from
the continent for the use of invalids. Mr. Jeffreys
(1862) was of opinion that there was no reason to
suppose that the creature was imported from Italy in the
sixteenth century, and he regarded it as equally indi-
genous, probably, with 7H. aspersa, our common garden-
snail. Neither of these species, he added, had been found
in this country in any recognized stratum of the upper
tertiary formation. Further, there was no foundation,
he thought, for the idea, prevalent at one time, that Z.
pomatia had been introduced by the Romans, for though
found near several encampments, it had occurred neither

* Since writing I find that Mr. Musson has recently referred to

Hi. pomatia as having been “ taken to Britain by the Romans.” See
«Proc. Lin. Soc._N.S.W.,” for 1890, (2), v. (1891), pp. 93-4.

DISPERSAL BY MAN. 241

at Wroxeter nor at York, nor in many other parts of Eng-
land and Wales where the Romans built cities or had
important military stations, and in “all probability this
kind of snail was not known to them, as another species
(7. lucorum) takes its place in Central Italy.” Recur-
ring to the subject in Nature, in 1883, Mr. Jeffreys
added that among the débris of an extensive Roman
villa, discovered in Northamptonshire, in which the
shells of cockles, oysters, mussels and whelks abounded,
not one of H. pomatia occurred, although at Woodford,
a few miles distant, it is plentiful ina living state. The
Rev. L. Blomefield, writing from Bath in the same
journal, stated that he had neither found, nor heard of
the discovery of, a single specimen, either living or dead,
in the neighbourhood of that city, which the Romans
occupied for more than four hundred years. On the
other hand, Mr. D. Pidgeon, though believing the
supposed connection between Roman remains and /.
pomatia to be merely fanciful, has remarked that a fine
example of the Roman villa stands at a short distance
from the spot near Charlbury, Oxfordshire, where,
within somewhat narrow limits, the snail is now
abundant.

It is curious to find that Reeve (1863) differed from
his contemporaries, and stated that the present species
could hardly be said to be indigenous; he remarked,
however, that if not indigenous it had become fully
naturalized in our southern counties. Tate (1866) did
not express any opinion on the point, and Rimmer
(1880) merely quoted Jeffreys.

R

242 THE DISPERSAL OF SHELLS,

I have no means of indicating the exact range of H.
pomatia in this country ; it seems to be confined for the
most part to chalky soils. Jeffreys mentions its occur-
rence in woods, hedge-banks, and uncultivated places
in Surrey, Hertford, Kent, Oxfordshire, Wiltshire,
and Gloucestershire; the Leeds Census records it for
the additional counties of Hampshire, Sussex, and
Bedfordshire, and the creature, as we have seen, is said
to have been found in several other counties.

It seems useless, at this time, to offer a decided
opinion as to whether the animal is really indigenous to
our soil or not: the doctors have differed, and there
appears to be no actual evidence on either side; the
weight of opinion, however, as far as conchological
authors are concerned, has for a long time favoured the
creature’s indigenousness, and the absence of any sug-
gestion that it was foreign to our fauna in the works, now
more than two hundred years old, of Merret and Lister
certainly seems to support this view, which was held, we
have seen, by Leach, Turton, Gray, Forbes, Jeffreys, etc.
On the other hand, we have also seen that Pennant and
Da Costa, who wrote more than a century ago, both
stated positively that the creature was a naturalized
species, and they were followed in this view by
more than one writer, and even by the _ illustrious
Montagu. When a species originally introduced by
man establishes itself in a new country, subsequent
naturalists, in the absence of records, will always be
likely to mistake it for an aboriginal inhabitant of that
region, especially if the creature has happened to stray

DISPERSAL BY MAN. 243

beyond the confines of cultivation. In the absence of
information on the point, for instance, a person visiting
Cape Town would hardly regard HZ. aspersa, which he
might find in the neighbourhood in large numbers, as
a comparatively recent importation from Europe.
Supposing HY. pomatia to be really indigenous in
England, it seems somewhat difficult to account for
the wide-spread notion that such is not the case. The
belief that the creature was imported from abroad
by our own countrymen, we have seen, has prevailed for
a long time, and is perhaps more general than the idea
that it was introduced, previously, by the Romans. In
1863, we find Reeve stating, on the authority of a Mr.
Barlow, that “specimens had been transported from
Italy some thirty years before by an English nobleman,”
to the range of hills in the neighbourhood of Reigate
and Box Hill, in Surrey, “and as they had bred
abundantly Mr. Barlow was induced to take a house in
that locality,” in order that a diet of snails might be
administered to one of his sons, who was considered to be
in the last stage of consumption. A correspondent writing
to Nature, in 1883, mentioned that when collecting the
shells of this animal, many years ago, about the foot of
Box Hill, he was told by a farmer resident in that
neighbourhood that “the snails were brought from
Italy by Mr. Hope, of Deepdene, who was well known
in the early part of this century as a writer on the
medizval architecture of Italy.” Murray’s “ Hand-
book to Surrey” and Bevan’s “ Guide” both refer to
the creature as imported, the first-named stating that it
R 2

244 THE*DISPERSAL OF SHELLS.

is supposed to have been introduced near Guildford
by an Earl of Arundel. The correspondent to WVature,
before quoted, who lived from 1849 to 1852 within
two miles of Woodford, Northamptonshire, where this
species occurs in a small wood known as Woodford
Shrubbery, stated that it was commonly said in that
neighbourhood that the snails were originally brought
from abroad by General Arbuthnot (?) who had formed
the Shrubbery some thirty years before. It seems
likely that there is some foundation for such statements
as these, or some of them, and it can hardly be doubted
but that the animal, although possibly or probably truly
indigenous, has been imported from abroad from time
to time.'

» Merret, “ Pinax,” 1667, as quoted in Gray’s “ Turton,” 1840, pp.
35, 46; Lister, “ Historiz Animalium Angliz,” 1678, pp. II1I-12, as
translated for me by Mr. R. W. Goulding; Pennant “ British
Zoology,” iv. (1777), p. 117; Da Costa, “ Historia Naturalis Testa-
ceorum Britanniz,” 1778, pp. 67, 70-1; Pulteney, ‘‘ Dorsetshire
Catalogue,” 1799, p. 47; Donovan, “ Natural History of British
Shells,” iii. (r801), plate Ixxxiv. (pages not numbered); Montagu,
“'Testacea Britannica,” ii. (1803), p. 406; Maton and Rackett,
“Trans. Lin. Soc.,” viii. (1807), p. 202; Dillwyn, “ Descriptive
Catalogue of recent Shells,” ii. (1817), pp. 920-1; Turton, “ Con-
chological Dictionary,” 1819, p. 56; Fleming, “ British Animals,”
1828, p. 257; G. Johnston], -—Loudon’s *“ Mag. Nat. Hists7 smn
(1830), p. 47; Turton, “ Manual,” 1831, pp. 47-8; Edward Forbes,
“ Report 9th meeting, British Association, 1839,” (1840), pp. 130,
133; Thomas Brown, “ Illustrations,” ed. 2, (1844), p. 43; Gray’s
“Turton,” 1840, pp. 35, 137; 1857, pp. 113-14; Leach, “ Synopsis,”
1852, pp. 64-5; ‘Forbes and. Hanley,” iv: +(r853);9p eer
“‘ Jeffreys,” i. (1862), pp. 177-8; “Nature,” xxvii. (1883), po 5iie;
L. Blomefield, “ Nature,” xxvii. (1883), p. 553; D. Pidgeon, “ Quart.
Journ, Conch.,” i. (1875), pp. 54-6; “‘ Reeve,” 1863, pp. 60-2 ; “ Tate,”

DISPERSAL BY MAN. 245

HELIX CANTIANA Mont. The Kentish snail, Mr.
Gray remarked, in 1857, was once “supposed to be
almost confined to the four metropolitan counties, but
it is also found in Suffolk, near Bristol, and near Dublin.
It may have been introduced in these latter localities ;
for it has been within these few years, according to
Mr. Fryer, introduced with ballast by the colliers on
the banks of the Tyne, and is now rapidly spreading
itself in the hedges of that neighbourhood. These
circumstances would lead one to imagine that it might also
have been introduced into England from the Continent ;
but Férussac, who has compared it with the continental
species, regards it at least as a local variety of A.
carthustana of Draparnaud (not of Miiller), which is a
native of the south of France and Italy.” Professor
Tate, however, has stated that the creature is indigenous,
at least to certain parts of this country ; most writers,
indeed, have thus regarded it, and now that it is known
to have a much more extended range than that above
indicated (except that the Dublin record is perhaps
erroneous) it can hardly be looked upon even as a
possible importation."

Esoow Pe: 137 2. Rimmer,” 1860, p: 113+ J.C.,. “* Nature”? xxviii.
(1883), p. 31; W. C. Atkinson, p. 81; Murray’s ‘‘ Handbook to
Surrey,” p. 70; and Bevan’s “ Guide to Surrey,” p. I1I.,as quoted
by Mr. Atkinson.

* Gray’s “ Turton,” 1857, p. 130; “Tate,” 1866, pp. 126-7; and see
also, as to the probable dispersal of HY. cantiana with ballast,
K.. Forbes, ‘‘ Report, 9th meeting, British Association, 1839,” (1840),
Pausch; &.. Ludson, * Journ. of, Conch,” ive(1884), p: 171; and
J. E. Robson, quoted in the “ Naturalist,” 1886, p. 148.

246 THE DISPERSAL OF SHELLS.

HELIX CARTUSIANA Mill. This snail (which,
according to Mr. Jeffreys, is known to inhabit the
whole of France, the Rhine district, South Germany,
Switzerland, Dalmatia, Italy, and Greece, and has been
recorded as a Siberian species) was detected in Britain
in 1814. In this country, however, though plentiful
where it occurs, it has a very restricted range, being
confined, apparently, to the neighbourhood of the sea,
along the coasts of Kent and Sussex; and Jeffreys, it
appears, at one time thought that it had been naturalized
here by an importation from the opposite coast of
France; but at the time of the publication of the
“British Conchology” he regarded it as “clearly in-
digenous.”' A colony, known at one time on the east
moors near Cardiff on a small patch of raised ground
covered with a luxuriant growth of ballast plants, was
almost certainly introduced with ballast; but the
creature has now disappeared from that locality.’

HELIX PISANA Mill. Extremely local as a British
species, this snail is distributed at random in a few
widely separated places on or near the coast; in the
Channel Isles, it occurs in Jersey, in the south of
England in Cornwall, in Wales in Pembrokeshire, and
in Ireland in Meath and Dublin counties ; intentionally
introduced colonies are known to exist near Swansea,
in Guernsey, and possibly in other places. The
creature’s foreign distribution, Mr. Rimmer, has

. Jeffreys, ‘“‘ Trans. Lin. Soc.,” xvi. (1833), p. 509; “ British Con-
chology,” i. (1862), pp. 48-9, 193-4.
2 F. W. Wotton, “ Journ. of Conch.,” v. (1886),

DISPERSAL BY MAN. 2A

<—

observed, is by no means confined to the sea-board,
and in his opinion the fact of its range in this country
being limited to a few places on the coast is suggestive
of the idea that it may have been originally “ brought
over in ballast from the Continent, or perhaps from
Jersey:”*

HELIX OBVOLUTA Miill. This species (the cheese
snail of Gray’s “ Turton”), unknown as an inhabitant of
this country to the early writers on our conchology,
was first recorded as British in March, 1831, by Dr.
James Lindsay, who discovered it, in 1830, “apparently
indigenous,” amongst moss near the roots of trees in
Ditcham Wood, near Buriton, Hampshire, and it was
to be found, he added, for a considerable distance along
the chalk escarpment of the South Downs facing to the
north. He had collected more than twenty specimens.’
The creature inhabits central Europe, and occurs, also,
according to the “ British Conchology,” in the north of
France ; its indigenousness in Britain was doubted by
Mr. Jeffreys (1831), who observed that “its confined
locality and the circumstance of its having remained so
long unnoticed by British authors might warrant a
suspicion that it may be of the same recent and pre-
carious indizenousness in this country with the JZ.
carthusianella” (or cartustana) which he then regarded
as a possible importation from France.* Gray (1840)

1 “ Jeffreys,” i. (1862), pp. 208-9; “‘ Rimmer,” (1880), p. 133.

7? jo tandsay, “Trans, Lin. Soc.;” xvi. (1833); p.. 765.

3 J. G. Jeffreys, in the supplement to his “Synopsis” read in
fogt, Lrans: Lin. Soc.” xvi. (1833), p: 510.

248 THE DISPERSAL OF SHELLS.

suggested that the creature might be an escape, intro-
duced with foreign plants; Captain Brown (1844),
Forbes and Hanley (1853), and Reeve (1863), also
regarded it as imported or questionably indigenous.
The Rev. W. H. Hawker (1853), however, when record-
ing the finding of many specimens in the beech-hangers
covering Stoner Hill, Ashford, Hampshire, expressed
a belief that the species was a native. It was im-
possible, he thought, that the animal could have spread
or wandered over the low and flat country between the
Ditcham and Ashford Woods, the two habitats being
about six miles apart.’ It may be remarked, of course,
that there are various means by which the creature
might have been transported from one place to the
other; but it has since been found at points many
miles distant from the original habitat, and Jeffreys has
referred to it in the “ British Conchology” (1862) * as
clearly indigenous. It sometimes occurs in this country
in fair plenty, but its known range, it must be admitted,
is still very limited, being confined, I believe, to a

1 W-H. Hawker, “ Zoologist,” xi. (1853), pp. 3764-5; and see
also on the British habitats of A. obvoluta: J. E. Harting,
** Zoologist,” (2), ii. (1867), p. 760; W. Thomson, “ Zoologist,” (2),
li. (1867), p. 837 ; C. Griffith, ‘‘ Science Gossip,” for 1873, p. 276;
T. Godlee, *¢Quart. Journ. Conch.,” 1.. (1874-8), p: 68 je
Harting, ‘ Zoologist,” (3), li. (1878), p. 94; W. Jeffery, “ Journ. of
Conch.,” 11. (1882), pp. 316, 339; B. Tomlin, “Science Gossip,”
xix. (1883), pp. 67-8 ; C. Ashford, “ Science Gossip,” xix. (1883),
p. 89; T: D. A. Cockerell, “ Zoologist,” (3), ix. (1885); pigeee
S. J. Da Costa, on its occurrence (dead) in the woods of Norbury
Park, Surrey, “ Journ. of Conch.,” v. (1886), p. 81.

* pp. 48-9.

DISPERSAL BY MAN. 249

comparatively restricted area within the counties of
Hampshire and Sussex, and to a single locality in
Surrey ; in Wales, Ireland, Scotland, and the north of
England it appears to be unknown.

Mr. Clement Reid has recently worked out the
British distribution of the creature, and at the Linnean
Society in December, 1890, he exhibited a map showing
its range, together with specimens from “new localities
in Sussex.”’ In reply to an inquiry he has favoured
me with the following note:

“The point of my remarks at the Linnean Society
was this: I found that H. obvoluta, instead of being
confined to a small area on the borders of Hants. and
Sussex, extended eastward along the chalk escarpment
as far as suitable localities were to be found. The
species seems to be very particular as to its habitat; it
must have calcareous soil and plenty of shade, but the
ground must never be sodden. It seems also to be an
exceptionally sedentary species, for as far as I could
see it was confined to ancient woods, and was never to be
found in plantations, even if the trees were a hundred
years old.

“ Almost the only place where the necessary conditions
are combined is the chalk escarpment, for there we find
slopes too steep ever to have been cultivated, and on
these, consequently, are preserved many patches of the
ancient forest. Nearly all these scattered patches, as
far east as the River Arun, are full of Helzx obvoluta.

1 See “ Nature,” xliii. (1891), p. 264.

250 THE (DISPERSAL OF. SHELLS:

East of the Arun there are few traces of the old forest,
and I have not yet come across this snail. In the
extensive beech woods on the top of the Downs the
species is nearly always absent, but most of these woods
have been planted within the last hundred years, and
the few relics of old forest are on ground liable to
become sodden in the winter. Selzx obvoluta has
been recorded from Kingley Vale, on the south
slope of the Downs. I could not find it there myself ;
but it occurs in a copse about four miles to the
north.

“Everything seems to show that the creature is a
relic of our old woodland fauna, now nearly exter-
- minated through the destruction of the forests.”

TESTACELLA HALIOTIDEA Drap. Gray (1840)
coupled this snail-slug with Drezssena polymorpha as a
species supposed to have been introduced in modern
times, but he regarded it as well naturalized, and Zos-
sibly indigenous. Jeffreys (1862) thought it almost
impossible to say whether the creature was a native or
had been introduced, but Mr. Rimmer has, more recently,
expressed the opinion that there are “some grounds
for believing it to be indigenous;” he remarks, how-
ever, that it has doubtless been unintentionally imported,
from time to time, from abroad in soil at the roots of
shrubs and other plants. Mr. Alder, as long ago as
1838, thought it might be reasonably regarded as a
native. Zestacella scutulum Sby. which appears in the
new British list as a distinct species, has been included,
for a long time and until quite recently, under Z. hadzo-

DISPERSAL BY MAN. 251

tidea as a variety: the aggregate has a fairly wide
range in this country, and the creatures sometimes
occur in large numbers, but it appears that 7. Laliotidea
proper is less common than 7. scutulum, for, according
to the Leeds ‘“‘ Census” (1889), the latter has been seen
by the Conchological Society’s referees from thirteen
counties, and the former from only eight.’

The following molluscs, probably among many
others, all exotic, have been detected in Britain, either
in a living state or as dead-shells. Some have been
found alive in open places, like true members of our
fauna, and several have actually been admitted, at one
time or another, into the British lists. Most of them
have almost certainly been imported by man, no doubt
for the most part unintentionally.

HELIX LIMBATA Drap., the “ white-keeled snail” of
Gray’s “Turton,” a native of the south of France, etc.,
is said to have been found in the neighbourhood of
London by Mr. G. B. Sowerby, and was introduced
into our catalogues, in 1837-8, by Mr. Alder, who had
received specimens from Mr. Sowerby. Gray included
itin his “ Turton ” of 1840, stating, on Mr. Sowerby’s
authority, that it ved “in the hedges near London, on
the New North Road to Barnet, near Hampstead, on

Gray Ge. RUEOn, \1640,.p. 4:5 1557; \p.° 291; “Jefireys,” 1.
(1862), p. 146; “ Rimmer,” 1880, pp. 88-9 ; Joshua Alder, “ Mag.
Zool. and Bot.,” ii. (1838), p. 105; on the specific distinctness and
geographical distribution of 7: scutulum, see J. W. Taylor,
‘Journal of Conch.,” v. (1888), pp. 337-47.

252 THE DISPERSAL -OF SHELLS,

brambles.” He admitted, however, that the specimens
originally found had possibly been accidentally intro-
duced with some plant from the Continent, for after
considerable inquiries he had not been able to hear of
the finding of any other specimens, either in the same
locality, or elsewhere. Forbes and Hanley (1853) gave
the creature as a ‘‘ spurious” British species, and Gray
in the “ Turton” of 1857 referred to it as a shell intro-
duced into the fauna by mistake. The statement in
this edition that “a few living specimens were set free
on the New North Road, near Hampstead, by Mr. G.
B. Sowerby ; but they did not propagate themselves,”
is somewhat puzzling, and probably wrong. Mr. G. B.
Sowerby, grandson of the finder or liberator, is in-
clined to think that the statement in the edition
of 1840 is correct, but he has no note of the facts.
Jeffreys, we find, noticing the species in 1862, even
thought that it might possibly be “rediscovered in
this country and have its claim to admission as a
British species recognized,”

HELIX LACTEA Mill. A living specimen of this fine
Mediterranean snail was once found, by Mr. Kindon, in
a field near the railway at Pateley Bridge, Yorkshire,
and was forwarded, still alive, to Mr. Taylor of Leeds.
It had probably been carried, as Mr. Taylor supposes,
with shingle brought by rail from the coast, and the
finding by Mr. J. H. Salter in the following year of a

” Alder, “ Mag. Zool. Bot.,” ii. (1838), p. 106; Gray’s “ Turton,”
1840, pp. 34, 53, 143-4; 1857, pp. 293-4; “Forbes and
Hanley,” tv. (1853), p. 655 “ jefireys,’ 1: (1062); pxrg2:

DISPERSAL BY MAN. 253

shell of the same species, “ only recently dead,” on the
sands at Filey on the coast of the same county doubt-
less strengthens such a supposition. Both specimens
were probably imported with ships’ ballast. Another
shell of the same species, presumably dead, has been
found on the moors near Cardiff, on which ballast is
often deposited.’

HELIX CARTHAGINIENSIS Rossm. A specimen of
this Spanish snail, found alive on the Cardiff moors by
Mr. Wotton—probably introduced with ballast—was
shown at a meeting of the Conchological Society in
1883; with it, a few dead-shells of the same species had
also occurred.”

HELIX VILLOSA Drap., a native of central Europe,
has also been found at or near Cardiff, Mrs. David
Robertson, of Glasgow, having taken four living speci-
mens, in August, 1873, while searching for Ostracoda
in the ditches on the moors. In 1877, Jeffreys re-
corded the species, in the “Annals and Magazine,” as
“‘an addition to our mollusca,” on the strength of which
it appeared (marked as doubtfully British) in the
Conchological Society’s list of 1883. It is now clear,
however, that the animal has no claim whatever to be
regarded as native to this country. It has not even

1 J. W. Taylor, “ Journ. of Conch.,” v. (1886), p. 81 ; “ Naturalist,”
foco, p: 2515. W: E. Clarke and others; “ Naturalist,” 1886;
po 207. )|. Higoalter, “Nat. Hist. Journ,” 1x..(1885), p.: 187 5 EE.
Collier, “ Journ. of Conch.,” iv. (1884), pp. 151, 214.

= Rroc. Conch. soc,“ journ..of Conch.,” iv. (1884), p.. 951: E.
Collier, p. 214.

254 THE DISPERSAL OF SHELLS.

established itself in the above locality. The four
living specimens mentioned are the only ones (other
than those intentionally introduced or their de-
scendants) which have been detected in these islands,
and the name of the species has very properly been
omitted from the new list of 1892. Mr. Collier and
Mr. Rogers, as stated in 1884, visited Cardiff soon after
the finding of Mrs. Robertson’s specimens, but, although
they found the exact spot in which the creatures had
been picked up, they failed to discover other speci-
mens, and came to the conclusion that those originally
found were ballast shells, the moor being the “ place
where all the ballast is put.” Mr. Wotton, in 1886,
mentioned that the original shells were found near a
small patch of raised ground, covered with ballast-plants,
where the 77. cartuscana, no doubt accidentally intro-
duced, was formerly known to occur. He had searched
closely very many times for HZ. vzllosa, but without
success ; the creature had certainly been imported, he
thought, probably with ballast, or possibly with Esparto
crass, large quantities of which are brought into Cardiff
and stored on the moors. A brickyard, he added,
“now covers the spot where 7. vz//osa was taken, and
various works and docks are fast occupying the
ground.” About forty years ago, as already noticed,
Canon ‘Tristram turned loose; on a moor ineee
Durham, several specimens which he had _ brought
alive from Switzerland, and they or their posterity
were ascertained to be living in the same place ten
years afterwards, The Rev. J. W. Horsley, also, has

DISPERSAL BY MAN. - 255

recently liberated Swiss specimens in his garden at
Woolwich.

HELIX VERMICULATA Mill. This snail, a fine and
well-known Mediterranean species, has not (as far as I
know) been found living in Britain as an escape; it has
been accidentally imported, however, in a living state,
and this is more than can be positively said for the two
following species, 17. aperta and LZ. personata, both of
which have actually been put forward by eminent con-
chologists as members of our fauna. In 1891, Mr.
L. E. Adams sent for exhibition at the Conchological
Society a specimen of A. vermiculata which he had
received from Barnsley asa Kentish shell. In investigat-
ing the history of the specimen, he ascertained that it
arrived in Barnsley, in January, 1891, in a parcel of
horehound, consigned toa chemist in that town. The
horehound was said to have been grown in Kent, but
was procured in the ordinary way of business from a
tradesman in London who was known to deal largely
in foreign herbs, and as the horehound in question may
have been in his warehouse for several months, the
probability is, as Mr. Adams thinks, that the snail
- crawled amongst it from some foreign herb which was
also stored there. Mr. Adams tells me that the creature
was alive when it arrived in Barnsley.

1 J. G. Jeffreys, “Ann. and Mag. Nat. Hist.,” (4), xix. (1877),
pp. 199-200; E. Collier, “Journ. of Conch.,” iv. (1884), p. 214; F. W.
Wotton, “Journ. of Conch.,” v. (1886), p. 56; H. B. Tristram,
* Zoologist,” (3), 1. (1877), pp. 260-1.

jo rec, Conch: Soc. * Journ. of Conch.,” vi, (1891), p. 393.

256 THE DISPERSAL OF SHELLS.

[| HELIX APERTA Born, another Mediterranean species,
hardly ought to be mentioned here at all, for it has
never been found, as far as I know, either living or
dead, in any of the islands of the British archipelago ;
it was introduced into our fauna, however, in 1839, when
a specimen received from Forbes, as from Guernsey, is
said to have been placed in the British Museum by
Gray ; but Guernsey, of course, belongs geographically
to France. The species was included by Gray in the
“Turton” of 1840, where it is stated that the shele
which was a crushed specimen, was found in mud under
the side of a hedge, a situation similar to those in which
Forbes had found the snail in Provence; and Forbes
and Hanley included it in their celebrated work of 1853;
but, in the “ Turton ” of 1857, Gray excluded it, believing
that it had been previously included by mistake. Jeffreys
(1862) remarked upon the fact that the sole ground for
supposing the species to be a native was the discovery
of the shell above mentioned (which he described as a
dead specimen found in a cart-track), adding that Dr.
Lukis, then and at the time of the discovery a resident
in Guernsey, had frequently searched for the shell in
vain ; its shape, he significantly remarked, “ is not much
unlike that of the variety fexuzs of Helix aspersa which
is common in Guernsey and is frequently bandless
and without coloured markings.” He also stated
that Forbes’ shell could not be found for reference
in the British Museum, Dr. Baird (then in charge
of the shells) never having seen it. Quite possibly,
of course, as Jeffreys admitted, the shell in question

DISPERSAL BY MAN. 257

was the true Helix aperta imported and dropped by
some French sailor, or it may have been introduced
with plants. | ’

HELIX PERSONATA Lam. Jeffreys, who did good
service in helping to banish AH. aperta and other foreign
shells from our catalogues, was himself guilty of the
insertion of other species which are now known to
have no claim whatever to be regarded as British.
HT. villosa, we have just seen, was introduced by him in
1877, and previously, in the “ Annals and Magazine ” in
1870, he had introduced the central European Z.
personata ;—

“The tale of British land and fresh-water shells is not
yet told. A dead specimen of Helix personata has been
found by Mr. S. A. Stewart, of Glasgow, at Newcastle,
inco. Down; and it is now...in my possession. Last
year I examined Mr. Stewart’s collection of fossil shells
from the Post-Tertiary beds at Belfast and in that
neighbourhood, and I have since received several com-
munications from him on the same subject. Judging
from his accuracy in these matters, I have every reason
to believe that 17. personata is a native of Ireland, and
that his specimen was not accidentally introduced, as
was the specimen of . apgerta into the Channel Isles.”

In 1883, when a discussion arose in ‘‘ Science Gossip ”
as to the admission of certain land-shells into the British
list, Mr. Stewart wrote to corroborate a statement that
the claim of HY. personata as a British species rested on

” Gray’s “ Turton,” 1840, pp. 36, 53, 127-8; 1857, p. 293; “ Forbes

and Hanley,” iv. (1853), pp. 43-4; Jeffreys, 1. (1862), pp. 184-5.
S

258 THE DISPERSAL OF SHELLS.

the single dead-shell referred to. He had picked up the
specimen while collecting mosses on the Newcastle
sandhills, but only discovered it (at the roots of a tuft of
a Hypnum) when he proceeded to examine the con-
tents of his vasculum, at Belfast, in the evening; subse-
quent searches on the same sandhills had proved un-
successful, and therefore he did not regard the species
as British, and was of opinion that its name ought to be
excluded from our lists. It was improbable, he thought,
that the specimen had been imported with ballast, for
there were no ballast-heaps about the little port, and
the shell was found a long way from the quay; but, as
he added, “ Newcastle is one of the most popular
watering places in the North of Ireland, and it is no un-
usual occurrence for people to stop there for a time who
have also been at watering places on the continent.
Shell collecting is one of the amusements of such resorts,
and it is quite likely that some person, having foreign
shells, dropped the specimen about which I write.” !
BULIMUS EXILIS Gmel. A dead shell of this animal,
a West Indian species, found on the shores of Lough
Carra, county Mayo, Ireland, was recently exhibited at
the Conchological Society on behalf of Miss Warren.”
The specimen was gathered by one of her friends
from amongst quantities of dead shells cast up on
the shores, and its presence in such a locality is
certainly surprising, especially as Carra is an inland
1 J. G. Jeffreys, “Ann. and Mag. Nat. Hist.,” (4), vi. (1870), pp.

423-4; S. A. Stewart, “ Science Gossip,” xix. (1883), p. 159.
2 Proc. Conch, Soc., “ Journ, of Conch.,”’ vi. (1891), p. 387.

DISPERSAL BY MAN. 259

lake. Miss Warren suggests that it may have been
a collection-escape. No one residing in the neigh-
bourhood, however, is known to possess West Indian
shells, but, by one means or another, quite possibly, the
specimen may have been carried, perhaps by a stream,
from a considerable distance.

PULIMwUS UNDATUS tug. Sir C,. Lyell, in the
* Principles,” stated that Bb. uwndatus, a West Indian
shell of considerable size, had been imported, adhering
to tropical timber, into Liverpool, and, as mentioned in
the last chapter, he was informed by Mr. Broderip that
the creature had become “naturalized in the woods
near that town.” It can hardly be assumed, I think,
that the animal is, or ever was, really acclimatized here ;
Gray, however, in 1840, and again in 1857, speaking of
it as B. zebra, mentioned its importation with mahogany
logs, and remarked that it often lived for some time in
this country. I am not aware that specimens have
been found during recent years.’

BULIMUS DETRITUS, Mull. In 1880 or 1881, Dr. P.
B. Mason wrote to the “ Journal of Conchology” that
more than a dozen specimens of B. detritus, in various
stages of growth, most of which were alive when found,
had recently been brought to him as having been taken
from a rockery in a garden in the neighbourhood of
Burton-on-Trent. They occurred among a number of
dead littcral shells, gathered at Scarborough, with
which the owner of the garden was certain they had

ae amihciples sal. pa 370 5 Grays * Turton,” 1840;. p. 7 ; 1857,

p--202.
5 2

260 THE DISPERSAL OF SHELLS.

been introduced. But it is, of course, unlikely that this
was the case, seeing that the shell is a native of central
and southern Europe; and it seems that its presence
can be accounted for in another way, for Dr. Mason
ascertained that a quantity of light barley and other
seeds (screened out of some samples of barley intended
for malting purposes) had been brought into the garden
for the purpose of feeding poultry; and a sample of
screenings which he subsequently procured, as already
mentioned, contained living shells of Helix caperata.'
BULIMUS DECOLLATUS L., a Mediterranean and now
widely distributed species, once found a place in
British catalogues, having been imported, it can hardly
be doubted, with plants.- Turton, in his “ Conchological
Notices,” of 1826, stated that the creatures had been ob-
served to breed in great abundance for many successive
years at Watton, in the south of Devon, the seat of H.
Studdy, Esq. They were lodged in the earth under the
wood-work of a green-house, whence they wandered
abroad in summer; at last, however, when the wood-
work and the earth were removed, the colony was lost,
“and all that were preserved we owe to the care of Mrs.
Griffiths and Miss Hill.’ In the “Manual,” in 162m
Turton added that no foreign earth was ever known to
have been admitted into the green-house, and that the
animals were considered by the gardeners as natives.
Mr. Alder, in 1838, however, remarked that the species
could not be regarded as British, and Gray seems to have

1 Pp. B. Mason, “ Journ. of Conch.,” iii. (1880-2), p. 118.

DISPERSAL BY MAN, 261

finally excluded it from our lists in his “ Turton ” of 1840,
observing that it was not even naturalized or acclima-
tized, for it occurred only in hot-houses warmed with
artificial heat. '

Mr. W. Borrer tells me that many years ago the late
Sir W. Hooker gave him specimens of this mollusc
(together with B. goodallz?) from the Royal Gardens at
Kew.

BULIMUS OCTONUS Chemn., a West Indian and
Central American species, is said to occur in green-
houses in this country. It has been recorded from near
Manchester, etc.”

CLAUSILIA PAPILLARIS Miill., of southern Europe,
seems to have found a place among our native shells
at one time. As stated by Mr. Alder, however, a
manuscript copy of Laskey’s “ North British Testacea,”
consulted by Forbes, was found to “fully explain the
history of the British C. papellaris.’ The creature
occurred, it appears, in Granton Park, near Edinburgh,
to which place it had been imported from abroad in
moss round the roots of some exotics.’

CLAUSILIA SOLIDA Drap. Of this species, also a
native of southern Europe, a single specimen (figured
in the supplement to the “ British Conchology ’’) is said
to have been found at Stapleton, near Bristol, by Mr.

’ Turton, “Zoological Journal,” ii. (1826), p. 565; “ Manual,”
1831, p. 79; Alder, .“‘ Mag. Zool. Bot.,” ii. (1838), p. 110; Gray’s
“Turton, 1840, pi 184.

2 T. D. A. Cockerell, ‘‘ Science Gossip,” 1893, p. 26.

= J; Alder, * Mag. Zool. Bots,” n. (1838), 111:

262 THE DISPERSAL OF SHELLS.

Rich. There are extensive nursery grounds in the
locality, and it is probable, as Mr. Rimmer has suggested,
that the shell was brought from the Continent in the
roots of plants or among moss.’

CLAUSILIA~ PARVULA © ‘Studer. || Varieties Yof Som:
common C. rugosa have several times been incorrectly
referred to this species, which on this account long ago
received a place in our catalogues. Thetrue C. parvula,
it is stated in the “ British Conchology” (1862), “in-
habits the North of France, as well as every other part
of the Continent, and may be expected also to be found
in Great Britain,’ and in fulfilment of this expectation,
as appears by the supplement to the fifth volume
(1869), several specimens have been found by Mr.
Grant Allen, at Kinver, near Stourbridge ; these, how-
ever, it has been suggested, were, in all likelihood,
“accidentally or intentionally imported from the con-
tinent.” > Both this and the last species were excluded
from the Conchological Society’s list of 1883, as not
having the slightest claim to rank as British. Con-
tinental specimens of both have been turned out by
Mr. Baillie, near Brora, Sutherlandshire.

Many other foreign molluscs, no doubt, have occurred

“Jeffreys,” v. (1869), Supplement, p. 162, pl. xcix. fig. 2;
“Rimmer,” 1880, p. 178.

? Gray’s “ Turton,” 1857, pp. 186-8; Alder, “‘ Mag. Zool. Bot.,” il.
(1838), p.111; “ Jeffreys,” i. (1862), p. 280; v. (1869), Supplement,
p. 161, pl.xcix. fig.1; ‘“‘ Rimmer,” 1880, pp. 117-18; J. T. Marshall,
** Science Gossip,” xvill. (1882), p. 261.

DISPERSAL BY MAN. 263

from time to time, especially in gardens and green-
houses. The finding of two Parmacella-slugs in a
garden near Newcastle, of a living shell of Melcczna
amceena ina nursery at Holloway, and of three exotic
land-shells in an orchid-house in Nottingham was
referred to in the preceding chapter. No doubt they
occur, also, somewhat frequently in shops and ware-
houses, generally dead, but sometimes, as in the case of
Flelix vermiculata at Barnsley, in a living state. A
grocer at Louth once gave me dead shells of Helx
virgata (from amongst raisins) and Helix cespituim
(from a bag of nuts), and he afterwards found other
species. It is probable also that many kinds, brought
over from abroad, or received from foreign corre-
spondents, like the Helzces and Clausitlie above
referred to, or mentioned in the preceding chapter,
have been intentionally turned out. elex terrestris,
which may possibly obtain a permanent footing, was
very probably thus introduced, and it has been recently
transplanted by the Rev. J. W. Horsley to his garden
at Woolwich. Some South American snails, as already
noticed, turned out in an English garden, are known to
have survived at least for three or four years.

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INDEX.

A.
ABERDARE, shower of fish in
valley of, 43.
Accrington, Pupa cinerea near,
234.
Achatinelline, restricted
ranges in, 9I, 96.
Acilius, snail’s eggs attached to,

specific

47.
bivalve clinging to, 65.
Ancylus adhering to, 87.
Adami, G. B., colonization by,
192.
Adams, L. E., on a flood in the
Thames, 31.
on bivalves clinging to newts,
71.
on Helix vermiculata at
Barnsley, 255.
Adams, Professor, on weight of
Vertigo milium, 147.
Adanson, on shells in swamps,

24.

Additions to the fauna of a pond,
14.

“Estivation, climatic barriers over-
come by, 100.

dispersal during, 148.

Alder, J., on introduced or doubt-
fully indigenous shells in
Britain: on Drezssena poly-
morpha, 213.

on Zestacella maugei, 228.
on Bulimus goodallizt, 230.
on Zestacella haliotidea, 250.
on Helix limbata, 251.

on Clausilia papillaris, 201.

Alexandra Park, Manchester, shells
in lake at, 19.
Allen, Grant, on Claustla parvula
near Stourbridge, 262.
Alluvium, shells in, 141.
Amalia, attempted colonization of,
195.
Amazons, floating pumice of, 125,
139.
rafts of, 129, 130.
Amphibia, dispersal by, 69, $8.
wandering habits of, 75.
climbing powers of, 76.
And see frogs, newts,
toads.
Ampullaria, tenacity of life of, 28.
Ancylus, Reeve on the means of
dispersal of, 86.
adhering to water-beetles, 86.
possibly dispersed with mus-
sels, 88.
Ancylus fluviatilis, carried away
by a flood, 31.
adhering to water-beetles, 87.
on to mussels, 88.
ova of, attached to a water-
beetle, 47.
A, lacustris, adhering to a water-
beetle, 87.
adhering to a frog, 89.
Animals, dispersal by, 45, 154, 174,
178, 209.
living molluscs (or ova) pass-
ing the digestive systems of,
45, 46.
possibly scattered from the
stomachs of, by birds of

prey, 45, 159, 176.

and

266

or ejected from the crops of
frightened birds, 45, 163.
Anodonta, tenacity of life of, 28,
4I.
shower of, 44.
dispersal of fry of, 48.
clinging by closure to various
objects, 57.
to a redshank, 79.
to a duck, 79.
to a water-vole, 83.
Anodonta or Unio, clinging to a
sandpiper, 79.
to a heron, 80.
to a duck, 80.
Anodonta cygnea and frost, 41.
A. fluviatilis, in hole formed by

peat-digging, 18.

Anthony, Mr., on Llanorbis dila- |

tatus in Britain, 222.
Antiquity, 3, 115, 173.
Ants, on drift-wood, 138.
Aplin, O. V., shells not seen upon
birds’ feet by, 50.
on tenacity of life in a land-
snail, 108.
on a magpie killing snails
before swallowing them,
160.
on barley scattered from a
bird’s crop, 162.
Arboreal snails, dispersal of, 136.
Arbuthnot, General, said to have
imported Helix pcmaiia,

244.

Arion, tenacity of life of, 166.

Arion ater, carried by a river,
170.

Artesian wells, 22.

Arundel, Earl of, said to have im-
ported Helix pomatia, 244.

Ashford, C., on the transportal of

Succinea putris by floods,
142.

on colonization of Helix fo-
matia, 186.

on SLulimus
Britain, 231.

Astacus, bivalves clinging to by

closure of the valves, 83.

by the byssus, 84.

goodallit in

INDEX.

Atchafalaya, great raft of, 129.

Atkinson, J. C., shells not seen
upon birds’ feet by, 50.

Atlantic, birds blown across the,
156.

Aucapitaine, Baron, on tenacity of
life in a snail, 105.

experiments with snails and

sea-water, I2I.

Austen. See Godwin-Austen.

Australasia, introduced slugs in,

178, 180.
Australia, dust whirlwinds of, 149.
Limax levis in, 173, 178.
Helix aspersa in, 180.
Azores, land-shells of, 93.
absence of fresh-water shells
in, 93.

birds blown from Europe to,
156.

shells probably carried to, by
birds, 159.

Bb,

BAILLIE, W., colonization by, 185,
I9I, 195, 262.

Baird, Dr., on tenacity of life in
the desert snail, 102.

on the Guernsey specimen of

felix aperta, 256.

Ballast, dispersal with, 196, 197,
245; 246, 247; 252; 253>

254.
Baltic timber, dispersal with, 211.
Bamboos, snails in hollows of,
140.
Bananas, dispersal with, 201, 207.
Barley, scattered from the crop of
a bird, 162.
Barley for malting, snails amongst
screenings from, 201, 260.
Barriers, 1, 2, 100,415, tyes
Bates, H. W., on dispersal by
floating pumice, 124, 139.
Beagle, beetle flying on board the,
69.

Beasts of prey, dispersal by means
of, 45.
Becher, E. F., on

puddles, 24.

Limnuece mM

INDEX.

Bee, snail clinging to, 85, 165.
Beetles, showers of, 44.
snail’s eggs attached to, 47.
bivalves clinging to, 63.
flying habits of, 66, 69.
carrying powers of, 67.
caught on the wing with shells
attached, 68, 87.
dispersal of univalves by, 85,
86, 155.
And see Aczlius, Dytiscus, and
Colymbetes.
Beevers, J., on additions to the
fauna of a pond, I5.
Belt, T., on dispersal of land-shells,

95.
on the difference in the dis-
tribution of fresh-water and
land-shells, 97, 98.
on whirlwinds, 149.
Benson, W. H., on tenacity of life
in a Cyclophorus, 114.

Berkeley, M. J., on Drvyezssena
polymorpha in Britain, 213,
216.

Bingley, a case of long suspended
vitality recorded by, 100.
Binney, A., on a snail reviving
after being frozen, 41.
on tenacity of life in the desert-
snail, 103.’
on voluntary migration, 116.
on drift wood, 127.
on dispersal by currents, 137,
144.

on #ulimus decollatus de-

stroying Helzx mnemoralis, |

194.
on dispersal with ballast, 196.
on the importation of snails
into the United States for
food, 203.
Binney, W. G., on the distribution
of Bulimus undatus, 138.
on river-dispersal, 142.
colonization by, 180, 193,

194.
Bird, claw of, found between the

valves of a mussel, 77.
Birds, dispersal by, 45, 77, 156,
174, 224.

267

shells possibly scattered from
the crops of, 45, 159, 176. _

ejection of contents of crops of,
45, 163.

rate of flight of, 47, 49, 159,
177.

shot on the wing with shells
and ova attached, 47, 79.

dispersal with plants and earth
adhering to, 51, 54, 157-

marine bivalves clinging to,

5°.

snails killed before being swal-
lowed by, 160.

dead, float on the ocean
possibly with molluscs in
their crops, 163, 177.

Birds of prey, dispersal by means

of, 45, 159.

Birmingham, Dvezssena polymorpha
in water-pipes at, 19.
Bivalves, voluntary migration of, 4.

out of water, 4, 33.

in isolated waters, 6, 8, 12.

effects of frost upon, 41, 42.

dispersal of, while clinging by
closure of the valves, 56.

And see Molluscs.

Blackbird, claw of, between the
valves of a mussel, 78.
Blackbirds, probable destruction of
snail-colonies by, I9gI.

Black slug. See Avon ater.
Blomefield, L., on snails eating

Birds,

into a _ pollard-elm, 132,
1a BAAS
on Helix pomatia in Britain,

241.

Bones, shells amongst, 202.

Borrer, W., on Bulimus goodallit
and &. decollatus at Kew,
230.265.

Bouchard-Chantereaux on_ snails
falling from trees with leaves,
147.

Bourguignat, J. R., on colonization
of Helix lactea, 189.

Brackish water, fresh-water shells
living in, 39.

Brazier, J., on dispersal by currents,

133.

268

Brazil, observation by Darwin in, 29.

Brick-pits, shells in, 17.

Bridgman, W. K., Planorbis nauti-
Jeus found among floating
weed, 34.

P. nitidus and a Valvata
amongst confervoid vegeta-
tion, 52.

Bristol, Zestacella

228.
Bulimus goodalhii near, 230.
Claustlia solida near, 261.

British Isles, introduced molluscs
established in:

Dreissena polymorpha, 181, 209,
210.

Planorbis dilatatus, 209, 221.

Physa acuta, 224.

Testacella mauget, 288.

Bulimus goodaliit (in green-
houses), 229.

Helix terrestris, 232.

the occurrence of other species
foreign to the fauna of, 226,
234,251.

doubtfully indigenous species
in, 209, 234 ef seq.

specimens of the indigenous
flanorbis glaber perhaps
introduced into, 210.

British land and fresh-water shells,

Conchological Society’s list of,
209, 227.

British Museum, specimens illus-
trating means of dispersal
exhibited in, 74, 78, 81.

the Guernsey specimen of
Helix aperta not to be found
in, 256.

Broderip, Mr., snail revived by,

112,
on the supposed naturalization

maugee near,

of Sulimus undatus near
Liverpool, 202, 259.
Brown, Captain, on Drezssena
polymorpha in Britain, 213,
218.
on Helix pomatia in Britain,
240.

Brown, R., on dispersal by ice, 123.
by floating timber, 139.

INDEX,

Bryant, J.. Dvretssena polymorpha,
discovered in Britain by,
211.
Buckland, Mr., rail caught by an
oyster, 60.
Buenos Ayres, felix pomatia at,
179.
snails from, colonized in York-
shire, 192, 202.
Helix lactea in markets at,
204.
Building materials, dispersal with,
198.
Bulimi, carried with dye-woods,
201, 202.
Bulimus decollatus, attempted colo-
nization of, in New Jersey,
193.
destruction of snails by, 194.
in Britain, 260.
B. detritus, at Burton-on-Trent,
259.
B. edwardsianus, carried to New
Caledonia as an article of

food, 203.
B. eremita, tenacity of life of,
its,

B. exilis, in Ireland, 258.
B. fasciatus, probable dispersal of,
by currents, 137.
B. goodalliz, in Britain, 229.
B£. octonus, in Britain, 261.
B. pallidior, tenacity of life of, 111.
B. rosaceus, tenacity of life of,
112,
B. undatus, probable dispersal of,
by currents, 137.
Mr. Binney on the distribution
of, 138.
carried with tropical timber,
201.
its supposed naturalization
near Liverpool, 202, 259.
BL, zebra. See BL, undatus.
Bulinus. See Physa gibbosa.
Buriton, discovery of Helix obvoluta
near, 247.
Burnley, Planorbis dtlatatus at,
223.
Bythinia, clinging to the larva of a
dragon-fly, 85.

INDEX.

c:

CADDIS-CASES, shells upon, 14, 35.
dispersal of shells with empty
cases, 35.
Canals, dispersal by means of, 196,
215, 202.
Canoes, drifting on the ocean, 128.
Cape, Helix aspersa at the, 179, 204,
2

43-
Cardiff, imported snails ‘near, 246,
253.
Carrington, J. T.,on Hel¢x terrestris
in Britain, 233.
Carson, J., on introduced slugs in
Victoria, 180.
Carychium minimum, in kexes,
140.
amongst leaves, 147.
Cattle, dispersal by, 12, 156.
Cattle-bone, shells amongst, 202.
Cattle ponds. See ponds.
Cattle-troughs. See troughs.
Census of the distribution of British
land and fresh-water shells,
219.
Centres of ‘‘ creation,”’ single, 1.
multiple, 2, 5, 118, 123.
Chalk, dispersal with, 197.
Chapman, Abel, shells not seen
upon birds’ feet by, 50.
on living cockles in birds’
crops, 161.
Chapman, H. V., on a_ bivalve
carried by a snipe, 81.
Chelydra, mussel carried by, 83.
Chemist’s shop, foreign snail in,
201, 255.
Chichester, colonization near, 188,

tO |
Chislehurst, colonization at, 186,

194.

Choules, A., Physa acuta discovered
in Britain by, 224.

Christy, R. M., on tenacity of life
in Paludina, 28.

Church, pond-snail on the tower of
a, 5, 165.

Cistern, shells in, 20.

City Road, shells in water-pipes of,

19.

269

Clausilia (or Cyclostoma) clinging
to a bee, 166

Clausilia papillaris in Britain, 261.

C. garvula in Britain, 262.

C. rugosa, tenacity of life of, 113.

C. solida near Bristol, 261.

Claustheé living under the bark of
trees, 132.

Climate, changes of, 117.

Climatic barriers, how overcome,
100.

Coal measures of Nova Scotia, fossil
land-shells of, 115.

Cochlicopa lubrica amongst leaves,

146.
Cockerell, T. D. A., Helix pomatia
colonized by, 186.

on Physa acuta at Kew, 225.
Cockle, clinging to dunlins, 59.

to a tern, 59.

to a peewit, 59.

to sandpipers, 60.

to a snipe, 60.

to a sanderling, 60.

Cockles alive in birds’ crops, I61.

Cocoanut-husks, floating, dispersal
by, 133, 134.

Cocoons of Testacellze, 168.

Collectors, dispersal by, 13, 16, 258.

Collett, R., on colonization of Helix
pomatia, 187. -

Collier, E., on Planorbis dilatatus
and Helix villosa in Britain,
222, 254.

Collings, Mrs., on an attempt to
colonize Helix pisana, 189.

Colonies, difficulties attending the
establishment of, 82, 96, 119,
$22; 1645 175;

isolated, of land-shells, 94.

Colonization of molluscs by man,
23, 182, 205, 263.

Columbia river, probable dispersal
by, 142.

Colymbetes flying on board the
Beagle, 69.

Commercial docks, Thames, dis-
covery of Dretssena poly-
morpha in, 211, 215.

Conchological Society’s list, 209,
2277,

270
Confervoid — vegetation, shells
amongst, 52.
slug carried by a floating mass
of, 170.
Congo, rafts of, 129.
Continental extensions, hypotheti-
cal, 11S;
Continents, general permanence of,
118.
Cooke, A. H., on Helix pisana in
Guernsey, 190.
on Physa acuta at Kew, 225.
Cooper, Thomas, on a shower of
frogs, 44.
Cordeaux, J., shells not seen upon
birds’ feet, by 50.
shells noticed among
weed by, 52.
on shells in birds’ crops, 160,
161-2.
on dispersal by
165.
Corixa, bivalves clinging to, 63.
Cormorant caught by an oyster, 61.
Cotton, supposed dispersal with,
221.
Cox, C. S. B., on 2elx cerresires
in Britain, 232.
Crayfish, bivalves clinging to, 83,
8

duck-

thrushes,

4.
Creation, centres of, I, 2.
Cresswell, F., on a tern caught by

a mussel, 60.
Crick, W. D., on mussels clinging
by closure, 58.
ona Spherium clinging to a
water-beetle, 63.
to a frog, 73.
Crops of birds, molluscs carried in,
45, 159, 176.
Felix caperata alive in the crop
of a pigeon, 161, 163.
living cockles in the crops of
curlews, 161.
Crow caught by a mussel, 60,
Cucumber-house, ulimus goodallit
in; 231.

Cuming, Mr., on a Cyclas probably
carried by a flood, 33.
Curlew sandpiper, cockle clinging

to, 60.

INDEX.

Curlews, living cockles in crops of,
161.
Currents, fresh-water, dispersal by,
18,21; 235730) 375 224 eos
160, I'70, 224.
rate of, in Amazons, 125.
marine, dispersal by, 37, 120,
163, 174, 177+
rate of, I2I.
Curtis, J., on dispersal by water-
beetles, 67.
on tenacity of life in snails, 113,

114.
Cyclas probably carried by a flood,

iNet see Spherium.
Cyclones, 149.
Cyclophorus indicus, tenacity of life
of, 114.
Cyclostoma (or Clausilia) clinging
to a bee, 166.
Cyclostoma elegans, isolated colonies
of, 95.
reviving after submersion in
sea-water, I2I.
Cyclostomas, tenacity of life of,

113.

D,

Da Costa, E. M., on Helzx poma-
tza in Britain, 235.
Da Costa, S. J., on Helix obvoluta
in Surrey, 248.
Dalton, J., on the occurrence of
freshwater shells in unlikely
spots, 5.
on an isolated pond, 17.
on the spreading of Dretssena
polymorpha, 220.

Daniel, Mr., on snails falling from
trees with leaves, 147.
Darbishire, R. D., on tenacity of
life in a land-snail, 105.

on colonization near Man-
chester, 185.
Darwin, C., on single centres of
“* creation,’ T.
on land as a barrier to fresh-«
water shells, 3.

INDEX.

on the wide distribution of
freshwater shells, 4.

on the dispersal of freshwater
organisms, 29.

on the difference in the distri-
bution of freshwater and
land shells, 29, 97.

on changes of level, 30, 117.

on salt-water killing snails,
40.

on fish carried by whirlwinds,

43-

on the dispersal of eggs of
freshwater shells, 46.

experiment with a duck’s feet
in an aquarium, 49.

on shells carried with duck-
weed, 51.

on earth adhering to birds, 54,
158.

on bivalves clinging to water-
beetles, 63, 64.

on the flying habits of water-
beetles, 66, 67, 69.

on their carrying powers, 67.

on a Unzvo clinging to a teal,

O.
on dispersal by birds, 82, 156,
177.
on an Azcylus adhering to a
Dytiscus, 87.
on wide ranges, 95.
on changes of climate, 117.
on hypothetical continental
extensions, I18.
on dispersal by ocean currents,
120.
experiments with snails and
sea-water, 120.
on dispersal by icebergs, 123.
on floating timber, 126, 128,
135;
on dispersal of the young of
land-snails, 157.
on organisms carried in birds’
crops, 159, 162.
on dead birds floating on the
sea, 163.
Darwin, F., on C. Darwin’s expe-
riment with a duck’s feet in
an aquarium, 49.

27 1
on mussels clinging by closure,

58.

Davy, J. B., on isolated ponds, 11,
12.

on bivalves clinging to frogs,

on Physa acuta at Kew, 225.
Denbighshire, shower of hay in,
153.
Denmark, attempted colonization
of Helix pomatia in, 187.
Descent, monophyletic, 1.
polyphyletic, 3.
Dickens, an allusion to
dispersal by, 154.
Digby, Sir K., said to have im-
ported Helix pomatia, 235.
Dikes, W. H., shells in a bird’s
crop, 160.
Dispersal, of freshwater shells, 27.
of bivalves while clinging by
closure of their valves, 56.
of operculates while clinging
by closure of the operculum,
85, 165.
of freshwater univalves while
adhering to other animals,

wind-

of land-shells, 95, 115.

scarcity of evidence of, 119,
172:

habitats conducive to, 34, 37,
52, 132, 140, 146, 169, 171,
173:

of slugs, 167.

Dispersal by man, 156, 178, 209.
Distribution, local, 1, 5, 23, 82, 94,
164.

discontinusus
173.
freshwater shells absent from
some oceanic islands, 93.
reasons for the differences in
the distribution of freshwater
and land shells, 95.
small snails, and operculates,
range more widely than large
inoperculate snails, 136.
Distribution of freshwater species,
wide, 4.
of land species, restricted, go,

ranges, 2, 92:

2/2
of land genera sometimes wide,

92.
of British land and freshwater

shells, ‘‘ Census ” of, 219.

Donovan, on Helix pomatia in Bri-
tain, 237.

Doubleday, H., on wandering of
frogs, 75.

Douglas, R. C., on a bivalve cling-
ing to a toad, 74.

Dover, Helix terrestris near, 232.

Downs, isolated ponds of, 6.

Dragon-fly larva, Spherium cling-
ing to, 61.

Bythinia clinging to, 85.
Drane, R., on a shower of fish, 43.
Dretssena polymorpha carried with

water, 18.

found in London streets, 18.

in water-pipes, 18.

dispersal of fry of, 36, 49, 220.

sometimes attached to cray-

fishes, 84.
its rapid diffusion, 181, 214,
217, 210.

its British history, 210.

its tenacity of life, 211.

its occurrence in unnavigable

water, 217.

in new water, 220.

its range in Britain, 219.
Dresser, H. E., shells not seen on

birds’ feet by, 49.
Drift-wood. See Timber, floating.
Drummond, T., on TZestacella

maugetand Lulimus goodallit

in Britain, 228, 230.
Dry-bulbs, dispersal with, 198.
Duck, a snail’s eggs on the foot of,

47, 157.

experiment with the feet of, in

an aquarium, 49.

a leech clinging to, 50.

caught by an otter’s shell, 60.

by mussels, 77, 79, 80.
Ducklings, caught by Oniones,

77:
Duckweed, dispersal with, 51.
Dumfriesshire, shower of leaves in,
_ 148, 153.
Dunlins, cockles clinging to, 59.

INDEX.

Duprey, E., on Azcylus adhering .
to water-beetles, 87.
Durham, Helix villosa and Helix

lapicida in, 193.
Dye-woods, dispersal with, 201,
202.
Dyer, Mrs., on Helix pisana near
Swansea, 189.
Dyson, Mr., on Dyezssena fpoly-
morpha in water-pipes, 18.
Dytiscus, bivalves clinging to, 63
et Séq.
the flying habits of, 66.
the carrying powers of, 67.
flying with shells attached, 68,
8

7°
Ancylus adhering to, 86, 87.

E.

EAGLES, probable dispersal by
means of, 160.
Earth, dispersal with: by birds, 54,

157:
by floating trees, 129, 133, 136,
169; 371, Z74.
by cattle, 156.
by man, 198, 200, 207, 250.
Eastbourne, snails in the gardens of
the martello towers near,
198.
Edible snail. See Helix pomatia.
Edible snails, dispersal of, 203, 235,
236, 240, 257.
Eggs, deposited by snails after dor-
mant periods, 104, 107, 109,
122.
of Zzmax in a tree-trunk, 173.
dispersal of: of freshwater
molluscs, 36, 39, 41, 46,
124.
of land molluscs, 120, 123,
124, 126, 138, 139,045,052,
153, 157, 158, 164, 169, 172,
173, 174, 200, 207.
Erie Canal, molluscs colonized in,
184.
Esmark, Miss, molluscs colonized
by, 184, 187.
dispersal with ballast, 197.

INDEX.

Esparto-grass, possible dispersal
with, 254.
Establishment of new colonies,
difficulties attending, $2, 96,
TIQ, 122, 164, 175.
European snails, exported as delica-
cies, 203.
carried by ships as part of pro-
visions, 203.
widely dispersed by man, 178.

F,

FAUNAS, insular, richness of, in-
creased by time, 123.
Fewkes, J. W., on a bivalve cling-

ing to a duck, 80.
Field-slug. See Lzmax agrestis.
Fierke, F. W., on an isolated pond,

10.
Finchley, isolated ponds at, 13.
Fish, showers of, 43.

dispersal of fry of Unionidze
by, 48. :

bivalves closing upon, 58.

possible dispersal with, by

human agency, 226.
Fish-ova, dispersal with, 195.
Fleming, J., on Dvezssena fpoly=

morpha and Helix pomatia

in Britain, 213, 238.
Floating of certain land-shells in

water, 140.

Floating husks, dispersal by, 133,

134.
Floating ice, dispersal by, 41, 123.
Floating islands, 129, 136, 137,

174.

animals seen on, 130.
pumas landed by means of, 130.
Lyell on the landing of, 131,

136, 174.

Floating molluscs, dispersal of, 35.

Floating pumice, 142, 139, 174.

Floating timber. See Timber,
floating.

Flood-water, shells in, 33.

Floods. See Currents.

Flower-pots, dispersal in, 200.

273

Food, dispersal of molluscs for,
203, 235, 230, 240, 257.
Forbes, E., oceans bridged over by,
117.
on dispersal by means of canals,
196.
on dispersal by man, 209.
on Helix pomatia in Britain,
239.
Helix aperta found in Guern-
sey by, 256.
Forbes and Hanley, on Dverssena
polymorpha and Helix po-
matia in Britain, 213, 240.
John, on dispersal with
bananas, 201.
Ford, Mr. (Redhill), on a mussel
clinging to a sandpiper, 79.
Fountains in Trafalgar Square,
shells in, 22.
Fowler, Canon, on the flying habits
of water-beetles, 66.
Foxes, mussel and oyster closing
upon the tongues of, 58.
snail-colonies probably exter-
minated by, 187.
Frankfort, Helix acuta carried to,

Ford,

200.

Franklin, Dr., a whirlwind de-
scribed by, 149.

French men-of-war, dispersal of
* escargots ’’ by, 205, 206.

French sailor, the Guernsey speci-
men of Helzx aperta per-
haps dropped by a, 257.

Fresh-water limpets. See <Azcy-
lus.

Fresh-water mussels. See
donta and Unio.
Fresh-water organisms, derived from

the sea, 3.
destroyed by changes of level,

Ajto-

99.
Fresh-water shells. See Molluscs.
Fresh-waters, want of continuity of,

Frogs, showers of, 43, 44.
living bivalves in stomachs of,

45:
bivalves clinging to, 72, 76.
wandering habits of, 75.

. fi

274

climbing powers of, 76.
Ancylus adhering to, 89.
Frozen mud, dispersal in, 42.
Fry, dispersal of, 36, 46, 48, i57,
174,°220:
Fryer, Mr., on Helex cantiana, 245.

Ge

Gain, W. A., on snails reviving
after being frozen, 41.
Gallwey. See Payne-Gallwey.
Ganges, rafts of, 129.
Garden-snail. See Helix aspersa.
Garner, Mr., on the dispersal of
Dreissena polymorpha, 211.
Gaskoin, J. S., on tenacity of life in
land-snails, 104, 109.
on the production of eggs by
snails after dormant periods,
104, 109, 122.
Gecko. See Lizard.
Germany, Drezssena polymorpha in,
212, 214.
Gibbons, J. S., on Helix aspersa
near Cape Town, 179.
on Hyalnia cellaria in St.
Helena, 179.
Girard, Professor, on_ bivalves
clinging to crayfishes, 83.
Glacial epoch, dispersal during, 30,
123,
Gloyne, C. P., on shells in cemented
tanks, 22.
Godman, F. Du C., on birds blown
to the Azores, 156.
Godwin-Austen, H. H., on the dis-
persal of eggs of fresh-water
shells, 46.
on the finding of a bird’s claw
between the valves of a
mussel, 77.
Gold-fish, possible dispersal by man
with, 226.
Goodridge, C., on drift-wood, 134.
Gorton, shells in lake at, 19.
Gould, Dr., on shells amongst
leaves, 146.

"INDEX.

Goulding, R. W., on a_ bivalve
clinging to a newt, 70.
to a frog, 73.
Gray, A. H., on a Unzo clinging to
a teal, 80.
Gray, J. E., on the exportation of
snails as delicacies, 203.
on Dretssena polymorpha in
Britainy 207;
on Helix pomatia, H. cantiana,
and #. obvoluta in Britain,
239; 245, 247.
on the occurrence of exotic
shells in the British Isles,
251, 250, ‘250, 208.
Great water-beetle. See Dytzscas.
Greenhouses, foreign snails in, 198,
230, 260, 261.
Griffith, J., on a shower of fish,

44.
Grocer’s shop, foreign snails in,
263.
Grocock, Mr., on Physa acuta in
Britain, 225.
Gude, G. K., on tenacity of life in
snails, 108, ILI.
on the finding of a Helicina
among orchids, 199.
Guernsey, attempt to colonize
felix pomatia in, 187.
Helix pisana colonized in, 189.
felix aperta found in, 256.
Gulick, J. T., on restricted specific
ranges, 91.
Gum Arabic, shell amongst, 201.
Guppy, H. B., experiment with
Neritina and salt-water, 40.
on dispersal in frozen mud,
2.
Gurney, J. H., on bivalves clinging
to birds, 60, 79.
Gwatkin, H. M., on Helix pisana
in Guernsey, 190.

Hi.

HABITATS conducive to dispersal,
34, 37, 52, 132, 140, 146,
1605 370,174:

INDEX.

Hancock, Mr., on cockles clinging
to birds, 59.
Hardy, J. R., on earth adhering to
birds, 54.
on bivalves clinging to the
larva of a dragon-fly and to
a water-scorpion, 62.
to a water-beetle, 65.
to newts, 71, 76.
to frogs, 73.
to a water-vole, 83.
on an operculate water-snail
clinging to the larva of a
dragon-fly, 85.
on Ancylus adhering to Dytzs-
cus, 87.
Harting, J. E., on Hekx pomatia
at Petersfield, 187.
on Dretssena polymorpha in
Britain, 213.
Hatton, Lord, attempted colo-
nization of Helzx pomatia by,
187, 236.
Hawker, W. H., on British localities
for Helix obvoluta, 248.
Hawks, probable dispersal by means
of, 159, 161-2.

Hay, carried up by whirlwinds, 151, |

ee
showers of, 153.
Heart-cockles, clinging to fish-
hooks, 58.
Heathcote, W. H., a Zzwnca seen
on the tower of a church by,
5, 165.
on a bivalve clinging to a
beetle; 64," °°
to newts, 70.
colonization by, 185, 190.
on AHyalnia draparnaldi at
Preston, 199.
Hedgehogs, snail-colonies probably
exterminated by, 187, 188.
Hedley, C., on introduced slugs in
Australasia, 178.
on Helix aspersa in Australia,
180.
on wide dispersal of slugs,
206.
on Limax maximus in
mania, 207.

Tas- |

275

Heilprin, Professor, on diffusion of
land-shells, 119.
Hlelicarion, carried by a_ beetle,

rss}
flelictde, wide range of, 92.
Helicina amena, carried from
Brazil with orchids, 199,
263.
Helix, wide range of, 92.
killed by sea-water, 121.
carried with Sphagnum, 199.
with cattle-bone, 202.
on the floating or sinking of in
water, 140.
flelix aculeata, amongst leaves, 147.
falling from trees with leaves,
147.
f1. acuta, carried on palm-trunks,
200.
HZ. alauda, carried with bananas,
201.
fH, annulus, probable dispersal of,
by currents, 133.
H. aperta, tenacity of life of, 105.
in Guernsey, 256.
fT, appressa, colonization of, 193.
FH, aspersa, in Australia, 140, 180.
at the Cape, 179, 204, 243.
in Madeira, 179.
in the Loyalty Islands, 205.
suspended vitality in, 107.
production of eggs by, after a
dormant period, 107.
found in cut bamboos, 140.
carried by a bird, 165.
found in imported sand, 197.
carried by ships as part of pro-
visions, 203.
imported into the United States
as an article of food, 203.
fH. austriaca, colonization of, 193.
H. bicarinata, tenacity of life of,
FIO; L191;
H. candidissima, tenacity of life of,
106.
fH. cantiana, on its indigenousness
in Britain, 245.
carried with ballast, 197, 245.
fH. caperata, alive in the crop of a
bird, 161, 163.
amongst barley, 201, 260.

i)

276

H. carthagintensis at Cardiff, 253.

H. cartusiana, on its indigenousness
in Britain, 246.

H. cespitum in a bag of nuts, 263.

H. desertorum, tenacity of life of,
102.

Hi. elegans. See H. terrestris.

Hi. ericetorum, isolated colony of, 94.
alive in alluvium, 142.
near Christiania, 197.

H. fraseri, tenacity of life of, 108.

H., fusca, falling with alder leaves,

147.

H. hortensis, tenacity of life of, 107.
attempted colonization of, 194.

H. lactea, suspended vitality in,

104.

production of eggs by, after a
dormant period, 104.

Bourguignat on colonization of,
189.

in markets at Buenos Ayres,
204.

in britain, 252.

H. lapicida, eating into a pollard-

elm, 133, 135:

possible dispersal of, by currents,
139.

planted in co. Durham, 193.

in New Jersey, 193.

H. lentiginosa, tenacity of life of,

ie

H. limbata, near London, 251.

H. nemoralis, in New Jersey, 180,

193.
in Virginia, 180, 193, 200.
H. obvoluta, on its indigenousness in
Britain, 247.
H. papilio, tenacity of life of, 110.
H. paupercula, tenacity of life of,
TIO.

H.. personata, in Ireland, 257.

H. pisana, tenacity of life of, 108.
colonized near Swansea, 188.
in Guernsey, and put down in

Sark, 189.
put down at Southport, 190.
in Lancashire, 191.
in Sutherlandshire, 191.
on its indigenousness in Britain,

246.

INDEX.

Hi. pomatia, revival of, after submer-

sion in salt-water, 120.
at Buenos Ayres, 179.
colonization of, 186.
British history of, 235.

H. pulchella, wide range of, go.

H. pygmea amongst leaves, 146.

Hi. rotundata carried by a wood-
louse, 155.

H. rufescens, shells of, broken be-
fore being swallowed by a
magpie, 160.

H. simtlaris, dispersal of, 136.

H. strigosa, dispersal of, 142.

Hi. tectiformis, tenacity of life of,
110.

H. terrestris,in Britain, 232, 263.

in South Carolina, 233.
Hi, turricula, tenacity of life of, 110.
H. undata, tenacity of life of, 109.
production of eggs by, after a

dormant period, 109.

H. veatchit, tenacity of life of, 108.

H. vermiculata, tenacity of life of,
106.

amongst horehoundat Barnsley,

201, 255, 203.

H. villosa, in Britain, 193, 253.

H. virgata, possible dispersal of, by
wind, 154.

carrying a smaller snail, 155.
amongst raisins, 263.

Hemphill, Mr., on dispersal by
rivers, 142.

Henshall, J., on bivalves clinging
to a newt, 77.

Heron, probable dispersal by, 50,

2

a mussel carried by, 80.
Heynemann, D. F., on a _ bivalve
clinging to a newt, 72.

to crayfishes, 84.
Hibernation, has enabled snails to
overcome climatic barriers,
100.
sea-water resisted during, 120.
dispersal by birds during, 165.
Higgins, H. H., on dispersal by
rivers, 144.
Hoare, G., on a cockle clinging to
a sandpiper, 60.

INDEX.

Holloway, a Helictva found among
orchids in, 199, 263.

Homing, faculty of, 117.

Hope, Mr., said to have imported
felix pomatia, 243.

Horehound, snail ina parcel of, 201,

255.
Horse-troughs, shells in, 20.
Horsley, J. W., on a mussel clinging
by closure, 58.
colonization by, 254, 263.
Hot-houses. See Greenhouses.
Howard, C., said to have imported
Helix pomatia, 236.
Hudson, B., on living bivalves in
the stomachs of frogs, 45.
on bivalves clinging to frogs,

73:
on Aucy/us adhering to mussels,
toa frog, 89.

on slugs in kexes, 171.
Humble-bee, snail clinging to, 85,

165.
Hurricanes, dispersal by, 43, 145,
146, 174.

Hyalinia cellaria in St. Helena, 179.
H. draparnaldi in a greenhouse at
Preston, 199.

I.

Icg, floating, dispersal by, 41, 123.
Ingalls, Dr., on an isolated pond,

17.
Insects, dispersal by, 47, 61, 85,154,
165.

Insular faunas, richness of, increased
by time, 123.
Ireland, whirlwind in, 150.
shower of hay in, 153.
colonization of snails in, 186.
Helix personata and Lulimus
exiles in, 257, 258.
Islands, land-shells of, often ende-
mic, 90.
And see Oceanic islands.
Islands, floating. See Floating
islands.

277

Tsocardia cor clinging to fish-hooks,
$

50.
Isolated colonies of land-shells, 94.
Isolated ponds. See Ponds.
Isolated water, absence of opercu-
late snails from, 86.
Italy, colonization of molluscs in,
192.
Helix pomatia said to have
been imported from, 236, 240,
243.

Je

JACKDAW, a snail probably carried
by, 165.
Jamaica, shells in cemented tanks
ini, 22:
Jeffery, W., on shells in non-
permanent water, 25.
on dispersal by floods, 32.
on bivalves clinging to newts,
vie
colonization by, 188, 194.
Jeffreys, J. G., Dretssena fpoly-
morpha carried with water,
18.
on the diffusion of fresh-water
shells, 30.
on Axnodonte
closure, 57.
on shells in kexes, 140.
amongst leaves, 146.
on dispersal by wind, 154.
on Helix pisana near Swansea,
188.
on probable dispersal of Testa-
cellee by man, 207.
on Dretssena polymorpha and
Planorbis dilatatus in Britain,
252, 221.
on Zestacella maugei and Bult-
mus goodallzt in Britain, 229,
230.
on exotic shells found in the
British Isles, 227, 252, 253,
256, 257, 262.
on doubtfully indigenous
species, 209, 240, 246, 247,
250.

clinging by

278

Jenkins, A. J., on a bivalve cling-
ing to a shrimp, 65.
to newts, 72.
on snails adhering to amphibia,
89.
on Physa acuta in Britain, 225.
Jenyns, L. See Blomefield.
Johnston, G., on long suspended
vitality in a snail, 102.
on Helix pomatia in Britain,
238.
Joly, M., on snails reviving after
being frozen, 41.

Ke

Kent, Helix terrestris in, 232.

Kentish snail. See Heltx cantiana.

Kew, Physa acuta discovered in the
Royal gardens at, 224.

on the occurrence of Azlimuts
goodallit and B. decollatius in
the gardens, 231, 261.

Kexes, dispersal of shells in, 139,
171.

Kindon, Mr., on Helzx Jdactea in
Yorkshire, 252.

King, Captain, on tenacity of life in
a land-snail, 112.

Knapp, J. L., on bivalves clinging
to newts, 70.

Kobelt, Dr., on shells passing un-
hurt through the digestive
system of birds, 45.

on dispersal by herons, 50.
flelix acuta carried with palm-
trunks, 200.

Korschelt, Dr., on dispersal of the

fry of Drezssena, 36.

L.

LAIDLAY, Mr., on tenacity of life in
Ampullaria, 28.
Lakes, want of continuity of, 98.

Lancashire, colonization in, 185,
IQI.

Planorbis dilatatus confined to,
224.

INDEX.

Land, changes of level of, 2, 30, 98,
Ti }s
a barrier to fresh-water shells, 3.
Land-shells. See Molluscs.
Layard, E. L., on the Trafalgar
Square fountains, 22.
on dispersal by birds, 23, 50.
lizard’s eggs found ina crevice
of a drifted tree-trunk by,
138.
on dispersal in Wardian cases,
198.

a snail carried to New
Caledonia by the Loyalty
Islanders, 203.

on Helix aspersa at the Cape and
in the Loyalty Islands, 204.

Leach, Dr., on Helzx pomatia in
Britain, 240.

Leaves, dispersal with, 146, 152,

154, 174.
a shower of, 148, 153.
Leech, clinging to a bird, 50.
Leeds, an isolated pond near, 14.
Leslie, H. C., on Lemna living in
a puddle, 25.

Leucochroa, tenacity of life of, 107.

Lewis, J., molluscs colonized by,
184.

Liardet, E. A., on dispersal by
currents, 134.

Life, tenacity of. See Tenacity of

life.

Limax, tenacity of life of, 167.

introduced into Australasia,
178, 180.
Limax agrestis, in kexes, 171.
in the crop of a bird, 177.
L. flavus, eggs of, in the trunk of a
free, 173.
L. levis, in kexes, 171.
wide range of, 173.
in Australia, 173, 178.
L. maximus, living under the bark
of trees, 169.
eating into decaying wood, 170.
in Tasmania, 207.
Limne@a, in isolated waters, 6 ef
Seq.
voluntary migration of, 4, 26.
adhering to pond-weed, 53.

on

INDEX.

Limnea auricularia, eggs of, pass-

ing the digestive system of

swans, 46.
on dried eggs of, 47.
L. humilis in an artificial pond, 16.
in non-permanent water, 25.
L. macrostoma in an isolated pond,
17.
L. parva, in a horse-trough, 20.
in non-permanent water, 25.
L. peregra in a water-trough, 20.
in the Trafalgar Square foun-
tains, 22.
in puddles, 25.
wandering on land, 26.
dispersal of, while floating, 35.
reviving after being frozen, 42.
adhering to turtles, 88.
to atoad, 89.
L. reflexa, dispersal of, by floods, 32.
L. stagnalis reviving after being
frozen, 42.
in New Zealand, 184.
in Tasmania, 195.
L. truncatula migrating over land,
4, 26,
on the tower of a church, 5,
165.
in horse-troughs, 21.
in non-permanent water, 24, 25.
diffusion of, in Madeira, 25.
tenacity of life of, 27.
dispersal of, during floods, 32.
Limpet. See Aucylus.
Lincolnshire wolds, isolated ponds
of, 9.
Lindsay, J., Helix obvoluta dis-
covered in Britain by, 247.
Lister, on Helix pomatia in Britain,
2255230.) 230:
Lithoglyphus naticoides in Britain,
226.
Liverpool, Bulimus wndatus near,
202, 259.
Lizard’s eggs found in crevice of
drift-wood, 138.
Local distribution. See Distribu-
tion.
London, Dreissena polymorpha in
streets of, 18.
Felix limbata near, 251.

279

Long, F. C., on Planorbis dilatatus
at Burnley, 223.

Lord, J. K., on an otter’s shell cling-
ing to a shoveller-duck, 60.

Louth, an isolated pond near, 9, Io.

Loyalty Islanders, snail trans-
planted by the, 203.

Loyalty Islands, Helzx aspersa in,

205.

Dr., attempt to colonize
Helix pomatia in Guernsey,
187.

on the supposed occurrence of
Helix aperta in Guernsey,
256.
Lukis, Mr., colonization of Helix
pisana in Guernsey, 189.
Lutraria, shoveller-duck caught by,
60.
Lyell, Sir C., on an_ introduced
Limne@ain Madeira, 25, 181.
on the dispersal of shells and
their eggs, 29, 36.
on dispersal by whirlwinds and
hurricanes, 42, 145, 149, 152,
153:
on = Ancylus adhering to a
Dytiscus, 87.
on the shells of Madeira and
Porto Santo, 96.
on tenacity of lifeina Bzlzmus,
112.
on the scarcity of evidence as
to the means of dispersal of
land-shells, 119.
on dispersal by icebergs, 123.
by pumice, 124.
by birds, 157, 159.
on floating islands, 129, 136,
174.
on snails carried in flower-pots,
200.
on Bulimus undatus near Liver-
pool, 202, 259.
on the dispersal of Dvredssena
polymorpha, 211.

Lukis,

M.

McDAKIN, Mrs., Heltx terrestris
discovered in Kent by, 232.

280

Macgillivray, J., on shells in water-
pipes, 19.

McLachlan, R., on caddis-larvze
attaching living shells to their
cases, 35

McMuttrie, J., colonization by, 192. |
McNabb, D., on a cockle clinging |

to a sandpiper, 60.
Macpherson, H. A., on dispersal by
birds, 50.
Madeira, possesses many endemic
land-shells, 91.
shells from Porto Santo have
not established themselves in,
96.
introduced molluscs in, 97, 179,
181, 199, 200.
tenacity of life in snails of, 109.
Magpie killing snails beforeswallow-
ing them, 160.
Mallard. See Duck.
Mammalia, dispersal by, 50, 155.
Man, dispersal by, 23, 96, 156, 173,

178, 209.
extermination by, 181.
shells introduced into the

British Isles by, 209.
Man-of-war, French, dispersal of
‘‘escargots’’ by, 205, 206.
Manchester, Dyetssena polymorpha
in water-pipes of, 18.
shells in artificial lakes near,
19.
colonization near, 185.
Hyalinia draparnaldz in a fern-
ery near, 199.
Planorbts dilatatus discovered
near, 221.
Bulimus octonus found in green-
houses near, 261.
Manchester Museum, specimens
illustrating means of dis-
persal exhibited in, 62, 64,
65, 71.
Margaritana margaritifera,
Island of Anticosti, 18.
Marine bivalves clinging to other
animals by closure of the
valves, 58.
Marine currents. See Currents.
Marsh-slug. See Zzmax levis.

in

INDEX.

Marsh, W. A., on shells in a horse-

trough, 20.
in non-permanent water, 24.
on dispersal by floods, 32.

Marshes, dry in summer, shells in,
23.

Martello-towers near Eastbourne,
snails in disused gardens of,
198.

Martin, Mr.,on dispersal of Lzm2c@a
while floating, 36.

Maryland, whirlwind in, 149.

Mason, J. E., on isolated ponds, II,
12.

Mason, P. B., on Bulimus detritus
at Burton-on-Trent, 259.

Maton and Rackett, on Helix poma-
tia in Britain, 237.

Medicinal purposes, dispersal of
molluscs for, 203, 235, 243.

Mendes da Costa. See Da Costa,

.M.

Merret, on Helix pomatia in Britain,
235.

Migration, voluntary, 4, 26, 116,
144.

Miller, J. S., on Zestacella maugez
and BLulimus goodallét in
Britain, 228, 230.

Miller and Sweet’s nurseries, 7esta-
cella mauget and Lulimus
goodalliz in, 228, 230.

Miocene land-shells, 115.

Mississippi, rafts of, 129.

Mohawk River, molluscs colonized
in, 184.

Molge. See Newts.

Molluscs, rapid increase of intro-
duced species, 25, 179.

reasons for the differences in
the distribution of fresh-water
and land kinds, 95.

European species widely dis-
persed by man, 178.

colonization of, by man, 182.

Molluscs, fresh-water, wide distri-

bution of, 4.
tenacity of life of, 24, 27.
means of dispersal of, 27.
dispersal of floating kinds, 35.
in brackish water, 39.

INDEX.

alive after being frozen, 41.

experiment with sea-water and
Neritina, 41.

a shower of, 44,

dispersal of bivalves while
clinging to other animals by
closure of their valves, 56.

of univalves while clinging by
closure of the operculum, 85.

while clinging by adhesion, 86.

their absence from some oceanic
islands, 93.

want of continuity of habitats

of, 99.

Molluscs, land, specific areas some-
times wide but generally re-
stricted, 90.

kinds found in
endemic, 91.
some genera range widely, 92.
dispersal of, 95, I15.
tenacity of life of, 99, 120, 167.
antiquity of, I15.
sea-water resisted by, 120.
habitats of, conducive to dis-
persal, 132, 140, 146, 169,
E71, 173
small species, and operculate
species, range more widely
than large inoperculate kinds,
136.
the floating or sinking of, when
thrown into water, 140, 170.
their occurrence in alluvium,
I4I.
found alive in the crop of a
bird, 161.
dispersal of operculates, 165.
of slugs, 167.
Moluccas, floating islands among
the; 1303
J. M., snail carried to

Madeira, 199.

Monophyletic descent, I.

Montagu, on Helix pomatia in
Britain, 237.

Montevideo, pumas landed at, 130.

snails from, living in Yorkshire,
192, 202.

Moor-hen, probable dispersal by,

Rat

islands often

Moniz,

281

Morch, Dr., on Dyretssena foly-
morpha, 214.
Morrison, J. H., Helzx nemorales in
Virginia, 180, 200.
Morton, J., on colonization of Helex
pomatia, 187.
Moseley, Hi. N., on floating timber,
126, 135.
Moss, a Helix found in a consign-
ment of, 199.
Mountain torrent, dispersal by, 34.
Mouse, oyster closing upon, 58.
Mussel closing upon the tongue of a
fox, 58.
upon the foot of a tern, 60.
upon the bill of a crow, 60.
Mussels, fresh-water. See Anodonta
and Unzo.
Musson, C. T., on voluntary migra-
tion by a Lzmnea, 4.
on bivalves in isolated ponds, 9.
on shells in an iron tank, 19.
in a water-butt, 20.
on dispersal by floods, 31.
on a ball of earth on the leg of
a bird, 54.
on the flying habits of water-
beetles, 66.
on tenacity of life in a land-—
snail, 108.
on a drifted canoe, 128.
on Szccinea arborea living under
bark, 132.
on Helix aspersa living in cut
bamboos, 140.
on dispersal by wind, 148.
on dust-whirlwinds, 149.
on a beetle and a woodlouse
carrying snails, 155.
on a Helix carrying a smaller
snail, 155.
on foreign snails in an orchid-
house, 198.
on transportal of snails at the
roots of ferns, 200.
on Helix pomatia in Britain,
240.

N.

NATURE can afford to wait, 164.

282

Neill, Pat., colonization of Helix
pomatia by, 187.
Nelson, T. H., shells not seen upon
birds’ feet by, 50.
Nelson, W., on additions to the
fauna of a pond, 14.
Nvepa, bivalves clinging to, 62.
carrying powers of, 69.
Neritina, wide dispersal of, 40.
experiments with, in salt water,
41.
egg-capsules of, 41.
Nests, probable dispersal by birds
when building, 164.
New Caledonia, snail carried to, by
Loyalty Islanders, 203.
New Jersey, Helix nemoralis and
LH. appressa colonized in, 180,

193.
New Zealand, colonization in,
184.
Newcastle, Parmacella found near,
208, 263.
Newcomb, H. L., ona Ux70 clinging
to a teal, 80.
Newman, E., on shells in water-
pipes, 19.
Newton, Professor, on earth on the
leg of a partridge, 158.
Newts, shells extruded by, 46.
bivalves clinging to, 70, 76.
wandering habits of, 75.
carrying bivalves over land, 77.
snails adhering to, 89.
Nicholls, Dr., on drift timber, 128,
135.
on transportal of shells with
plants, 196.
Norgate, F., on a leech clinging to
a bird, 50.
on probable dispersal by herons,
52.
on a rail caught by an oyster,
60.
on bivalves clinging to water-
beetles and newts, 64, 72.
on the flying habits of water-
beetles, 66.
on the wandering habits of
newts, 75.

Norman, A. M., on Dreissena

INDEX.

polymorpha in water-pipes,
18

on Physa acuta, 225.
Norman, G., on foreign snails
colonized in Yorkshire, 193.
on transportal with cattle-bone,
202.
Northamptonshire, attempt to colo-
nize Helix pomatia in, 187.
Norway, colonization in, 184, 187.
Notonecta, bivalves clinging to, 62,
63.
carrying powers of, 69.
Nottingham, foreign snails in an
orchid-house at, 199, 263.
Nova Scotia, land-shells of coal-
measures of, I15.
Nuts, shell in a bag of, 263.

O.

OAK-LEAVES, shower of, 148, 153.
Ocean currents. See Currents.
Oceanic islands, endemic land-shells
in, 90 é¢ seq., 123.
almost all inhabited by land-
shells, 92.
on the stocking of, 92, 164,
172,17 Os
fresh-water shells
absent from, 93.
arrival of species not witnessed,
119.
richness of faunas of, increased
by time, 123.
Oceans, general permanence of, 1,
Li.
changes in, 2, 118.
Oldham, C., on a bivalve clinging
to a water-beetle, 64.
on the carrying powers of
water-beetles, 68.
Operculate snails, dispersal of, 85,
165. Fe
absence of, from isolated waters,
86

sometimes

tenacity of life of, 113.
distribution of, 136.
Operculum, molluscs clinging by
closure of, 85.

INDEX.

Orchid-houses, foreign snails in,
199, 231, 263.

Orchids, a He/iéctna found in a case
of, 199.

- Orcutt, C. R., on shells in a tank,
25:

Orinoco, rafts of, 129.

drift-timber from, at Tobago,

£255 ;1 35.
Otter’s-shell, shoveller-duck caught
. by, 60.
Otters, probable dispersal by, 50.
Ova. See Eggs.
Owls, probable dispersal by means
of, 162:

Oxford, Hex pomatia in Botanic
Gardens at, 186.

Oxford Street, Dyrezssena foly-
morpha in water-pipes in, 18.

Oysters, mice, rat, and fox caught
by, 58

rails, plover, and cormorant

caught by, 60, 61.

P:

PAL/ZOZOIC coal measures, land-
shells of, 115.

Palm-trunks, shells carried on, 200.

Faludina vivipara, tenacity of life
of, 28.

clinging by closure of oper-

culum, 86.

Parana, rafts of, 130.

Parmacella found near Newcastle,
208, 263.

Partridges, earth on feet of, 158.

Parus biarmicus, shells in crop of,
160.

Pascal, L., on a snail’s eggs passing
the digestive system of swans,

6

46.
Payne-Gallwey, Sir R., on a cockle
clinging to a dunlin, 59.
Peacock. See Woodruffe-Peacock.
Pearce, S. S., on a bivalve clinging
; to: a frog, 73:
on dispersal with chalk, 197.
with soil, 198.

283

Pearl-mussel. See O70 mars
garittfer.

Peers, J., on bivalves clinging to
toads, 74.

Peewit, cockle clinging to, 59.

Pennant, on felix pomatia in
Britain, 235.

Peregrines, probable dispersal by
means of, 161-2.

Petersfield, attempt to colonize
Helix pomatia at, 187.
Pfeiffer, Madame Ida, tenacity of

life in Cyclostoma, 113.
Pheasant, slugs in crop of, 177.
Philippines, floating timber amidst,

Baie
land-shells of, 136.
Physa, in tanks, 19, 21, 22.
Physa acuta, in Britain, 224.
P. gibbosa, in a tank, 19.
LP. gyrina, carried with plants, 196.
Pickering, Mr., on tenacity of life
in land-snails, 109, 113.
Pidgeon, D., on_ heart - cockles
clinging to fish-hooks, 58.
on Helix pomatia, 241.
Pigeon, balls of earth on legs of, 54,

158.
living snails in the crop of, 161,
163.
barley scattered from the crop
of, 162.
Pine-beds, Aulimus goodalliz in,
230.
Pine-plants, supposed dispersal with,

230.
Pisidium, on land, 4, 34.
killed by frost, 42.
in stomachs of frogs, 45.
extruded by newts, 46.
clinging to water-beetles, 65.
to a newt, 72.
Pisidium or Spherium, clinging to
crayfishes, 83.
Pisidium etheridge, clinging to
water-bugs, 63.
P. fontinale clinging to WVepa, 62.
to a water-beetle, 65.
Planorbis, adhering to collecting
nets, 16.
reviving after being frozen, 42.

284

adhering to pond-weed, 53.
tenacity of life of, 221.
Planorbis dilatatus in England,
209, 221.
found in a cistern, 20, 224.
P. glaber, specimens of, perhaps
introduced into England, 210.
P. nautileus amongst floating weed,

34-

Plants, dispersal with, 34, 51, 133,
139, 170-1, 182, 195, 198,
201, 207-8, 224, 226, 228,
231, 247, 250, 252; 254-5)
257, 260-2.

Pleurocera, dispersal of by floods,

33:
Pliocene land-shells, 115.
Plover caught by an oyster, 61.
Polyphyletic descent, 3.
Pond-weed, shells adhering to, 34,

53.
Ponds, shells in, after being long

dry, 25.

Ponds, isolated, 5 e¢ seg.

additions to the fauna of an
isolated pond near Leeds, 14.

probably stocked by water-
beetles, 69.

and birds, 82.

absence of operculate snails
from, 86.

Porto Santo, its land-shell-fauna
not common to Madeira, 9I.
shells from, probably carried

to Madeira with stone, have
not established themselves,
96.
European shells in, 97.
tenacity of life of snails of, 109,
TET.
Portuguese, introduction of a Lzy2-
n@a into Madeira by the, 25,
181.
dispersal of land-shells by the,
200, 203.
Poulton, E. B., on tenacity of life
in Zestacella, 168.
Preston, pond-snail on the tower of
a church at, 5, 165.
Hyalinia draparnaldi in a
greenhouse at, 199.

INDEX.

Puddles, shells in, 23, 25.

Pulteney, on Helix pomatia in
Britain, 237.

Pumas, landed by means of floating
rafts, 130.

Pumice, floating, probable dispersal
by, 124, 139, 174.

Pupa, wide range of, 92.

in Paleozoic coal measures of

Nova Scotia, 115.

Pupa cinerea, in Britain, 234.

P. muscorum in the crop of a bird,
160.

P. tridens, tenacity of life of, 113.

P. umbilicata carried by a larger
snail, 155.

OVERQUEDULA DIscors, mussel carried
by, 80.
1a

RACOONS entrapped by bivalves,
Ppp y

9.
Rae, a D., on Physa acuta in
Scotland, 226.
Rafts. See Floating Islands.
Rails caught by oysters, 60.
Railway ballast, dispersal with, 197,
202.
Raisins, shell amongst, 263.
Rat, oyster closing upon the tail of,
8

mussel closing upon the foot of,
83.

carrying snails, 155.
Redcar, colonization at, 186.
Redding, Sir R., mussels clinging
by closure, VE
Redshank, bivalve clinging to, 79.
Redwings, probable dispersal by,
16

5-
Reeve, L., on ‘‘ facilities of migra-
tion,” 86, 95.
on multiple specific centres,
118.
on Dreissena polymorpha and

INDEX. 285

Helix pomatia in Britain,
253, 241,
Reid, C., on isolated ponds, 6.

on shells in horse-troughs, 21.

on the Trafalgar Square foun-
tains, 23.

on transportal of plants and
shells by birds, 52.

on shells adhering to plants,

53:
on birds ejecting the contents
of their crops, 163.
on Helix obvoluta, 249.
Reservoirs, shells in, 18.
Rhaphaulus, tenacity of life of, 114.
Rhode Island, snail from Cuba
found in, 201.
Rich, Mr., Clauszlia solida found
near Bristol, 262.
Riches, J. T., on a bivalve clinging
to a frog, 72.
Rigsby, an isolated pond at, 12.
Rimmer, R., on establishment of
Helix pisana near Swansea,

188.

on its indigenousness in Britain,
246.

on the _ indigenousness of

Testacella haliotidea in Bri-
tain, 250.
on the occurrence of Clausilia
soltda near Bristol, 262.
Riplingham, an isolated pond at,

TO.
Rivers, want of continuity of, 98.
trees and pumice stranded o
the banks of, 139.
And see Currents.
Roberts, G., on dispersal of
Limnee while floating, 35.
on living snails in a bird’s crop,
161, 163.
ona thrush dropping a snail,

165.

Robertson, Mrs. D., Helzx vzllosa
discovered near Cardiff by,
253.

Robson, C., on newts extruding
shells, 46.

Roebuck, W. D., on shells in a
water-trough, 20.

on Lzwnee in puddles, 25.

Rogers, T., attempted colonization
of Helix pisana by, 191.

Planorbis dilatatus discovered
in Britain by, 221.

Roman snail. See Helix pomatia.

Romans, Helzx pomatia said to have
been carried to Britain by the,
240.

Rossmassler, Professor, on Dvezs-
sena polymorpha attaching
itself to crayfishes, 84.

Royal Gardens. See Kew.

SAHARA, a snail’s eggs found upon
the foot of a bird shot in the,

47-
Sailors, dispersal by, 203, 259.
St. Helena, its isolation, 93.
its land-shells, 93, 94.
absence of fresh-water shells
from, 93.
introduced molluscs
182.
extermination of native species
in, 181.
Salt-water, injurious effects of con-
tact with, 39, 41, 120, 121,
134, 172.
revival of snails after submer-
sion in, 41, 120, 134.
seeds and shells in birds’ crops
protected from, 163.
Salter, J. H., on Helzx lactea in
Yorkshire, 253.
Sanderling, cockle
60.
Sandpipers, cockles
60.
fresh-water mussels clinging to,

ily, 179;

clinging to,

clinging to,

79»
Sandwich Islands, shells of, 91, 94,

restricted specific areas in, 9I,

96.
isolated position of, 94.

286

Sanger, E. B., on shells in flood-
water, 33.

Sankey, R. H., on tenacity of life in
Rhaphaulus, 114.

Sark, attempted colonization in,
FSO." + *

Saunders, H., shells not seen upon
the feet of birds by, 50.
Scarborough, colonization of Helix

pomatia near, 186.
Schaff, Mr., bivalve clinging to a
redshank, 79.
Scharff, R. F., on shells in alluvium,
TAI, 142:
on the dispersal of slugs, 172,
175.
eggs of a slug found in a tree-
trunk by, 173.
on Zestacella maugez in Britain,
229.
Scotland, shower of leaves in, 148,
153.
attempted colonization of Helix
pomatia in, 187.
Dreissena polymorphain, 216.
Physa acuta in, 226.
Clausilia papillaris in, 261.
Scouler, J., on Dretssena polymorpha
in Scotland, 216.
Sea, fresh-water organisms derived
from the, 3.
Sea-water. See Salt-water.
Semper, K., on polyphyletic de-
scent, 3.
on the distribution of Zzocho-
morpha, 90.
on dispersal by ocean currents,
120, 136.
Sharp, D., on the carrying powers of
water-beetles, 67.
Sheep, probable dispersal by, 12.
Shell-collectors, probable dispersal
by; 03, 16,250:
And see Colonization.
Shells. See Molluscs.
Ships, snails carried by, as “ live
sea stock,”’ 203.
Ships’ ballast, dispersal with, 196,
245-7, 252-4.
Shops, foreign snails found in, 201,
; 255, 263.

INDEX.

Shoveller-duck, caught by an otter’s
shell, 60.
Showers of fish, 43.
of frogs, 43, 44.
of water-beetles, 44.
of fresh-water mussels, 44.
of leaves, 148.
of hay, 153.
of snails, 154.
Simpson, C. T., on dispersal by
floods, 33.
on dispersal of fresh-water
shells by marine currents, 37.
on drift-wood, 127, 134.
on dispersal of large tree-snails,

137;
Simroth, Dr.,on dispersal of Testa-
celle, 171.
Skuas cause other birds to eject
the contents of their crops,
163.
Slugs, dispersal of, 167, 207.
cannot bear long exposure, 167,
75.
tenacity of life of, 167.
wide ranges of, 172.
allow themselves to drop from
small objects, 174.
introduced species in Austral-
asia, 178, 180.
supposed destruction of snail-
colonies by, 194.
Smith, E. A., on the egg-capsules
of Neritina, 41.
on extermination of native
species in St. Helena, 94,

182.
on felix terrestris in Britain,
222.
Smyth, W. H., on floating timber,
13i.

3
Snails. See Molluscs.
Snapping-turtle, a mussel carried by,
8

Be
Snipe, earth adhering to, 54.
cockle clinging to, 60.
Spherium clinging to, 81.
Soil. See Earth.
South Downs, isolated ponds of, 6.
Southport, Helzx pesana planted at,
190.

INDEX.

Sowerby, G. B., Helix limbata
found near London by,

25%.

Sowerby, G. B. (grandson), on
Helix limbata, 252.

Sowerby, J. De C., on Dyrezssena
polymorpha and Testacella
mauget in Britain, 210, 228,

Sowerby, W., on Physa acuta inthe
Royal Botanic Society’s Gar-
dens, 226.

Sparrows, suggested dispersal by,
2

Z
Species, closely-allied, cannot estab-
lish themselves in each other’s
territories, 96.
Sphaium or Pisidium, clin ging to
crayfishes, 83.
Spherium corneum, clinging to the
larva of a dragon-fly, 61.
to Wepa, 62.
to Dytiscus, 63 et seq.
to newts, 70, 76,77.
to frogs, 72, 76.
to toads, 74.
carried over land by water-
beetles, 68.
by a newt, 77.
by asnipe, 81.
S. dacustre, in a new isolated pond,
£2;
clinging to a water-beetle, 65.
S. ovale, its indigenousness in Britain,
209.

Sphagnum, a Helix found in a
consignment of, 199.
Spicer, J. W. G., slugs found in the
crop of a bird by, 177.

Spix and Martius, on the drift-wood
of the Amazons, 129.
Springall, C., a crow caught by a

mussel, 60.
Standen, R., on shells in brick-pits,
17.
in artificial lakes, 19.
on a snail’s eggs attached to a
water-beetle, 47.
on plants and earth adhering to
birds, 52, 54.
on Piszdium clinging to WVefa,
2.

287

on Spherium clinging to water-
beetles, 64, 65.

on the flying habits of water-
beetles, 66.

beetles carrying shells caught
on the wing by, 68, 87.

on bivalves clinging to newts,
71.

to frogs, 73, 76.

to toads, 75.

on a mussel clinging toa duck,
79:

to a heron, 8o.

on an Azcylus adhering to a
water-beetle, 87.

on a Lzmnea adhering to a toad,

on isolated colonies of land-
shells, 94.
Hyalinia draparnaldi found in
a fernery near Manchester
by, 199.
Stark, Mr., on Dryeissena fpoly-
morpha in Scotland, 216,
Stearns, R. E. C., on tenacity of
life in a land-snail, 111.
on dispersal with dye-woods,
202.
Stettin, colonization near, 193.
Stewart, S. A., on the Irish speci-
men of Helix personata,

257.

Stone, probable dispersal with, 96.

Streams. See Currents.

Strickland, H. E., on Dryetssena
polymorpha in Britain, 213,
215; 216.

Stuchbury, Mr., Dredssena poly-
ee planted at Bristol by,
183.

Succinea, reviving after being frozen,

4I.
found in a bird’s crop, 160.
eggs (probably of a) found
upon the foot of a bird shot
on the wing, 47, 157.
Succinea arborea, living under bark,
ia2;
S, putres carried by floods, 142.
its ova probably carried by
birds, 157.

288

Sutherlandshire, colonization in,
185, 191, 195, 262.

Swamps, shells in, 23.

Swans, eggs of a snail passing the
digestive system of, 46.
Swansea, Helix pisana colonized

near, 188.
Swaysland, Mr., on earth on the
feet of birds, 159.

a.

TANKS, shells in, 19, 21, 22.
Tasmania, Limnea stagnalis in,
195.
Limax maximus in, 207.
Tate R., on dispersal by streams and
floods, 31.
on dispersal of eggs of fresh-
water shells, 46.
on mussels clinging by closure,

on snails carried by ships as
live sea stock, 203.

on Dreissena polymorpha and
Helix cantiana, 213, 245.

Taylor, J. W., on anisolated pond, |

14.
species determined by, 14, 234.
Taylor and Roebuck’s census of the
distribution of British land
and fresh-water shells, 219.
Teal, mussel carried by, 80.
Teator, W.C., on Azodonta in a
hole formed by peat-digging,
18.
Tegetmeier, W. B., on earth on the
feet of a partridge, 158.
Tenacity of life of fresh-water shells,
2A, 27, 41, 40, 215,221.
of land-shells, 99, 120.
of slugs, 167.
Terns, cockle and mussel clinging
to, 59, 60.
Testacella, tenacity of life of, 167.
cocvons formed by, 168.
dispersal of, 169, 171, 194,

207.
Testacella haliotidea, 250.

INDEX.

T. mauget, 228.
its nest of earth, 168.
T. scutulum, 250.
Thames, shells carried by floods in,

35,
discovery of Drvrezssena poly-
morpha in, 211, 215.
Thomas, A. P., on tenacity of life
in a pond-snail, 27.
on dispersal by floods, 32.
Thompsen, W., on pond-snails ad-
hering to turtles, 88.
on snails planted in Ireland,
186.
Thomson, J. H., attempted coloni-
zation by, 194.
Thrush, snail carried and dropped

by, 165.
probable destruction of colonies

by, 190.
Timber, dispersal with, 200, 201,

211, 259.

Timber, floating, dispersal with, 33,
35, 375.41, 122, 126 2 osege
139, 169-71, 174, 215, 220.
seen on the ocean, 126-7, 129,
E3%:
often floats under the surface,
135.
bark often lost in the sea, 135.
drift-timber, stranded on coasts,
127-8, 134-5.
on a river-bank, 139.
eggs of a lizard found in a cre-
vice of, 138.
Time, richness of faunas increased
by;.123.
Nature can afford to wait, 164.
Titmouse, shells found in the crop
of, 160.
Toad, bivalves clinging to, 74.
climbing powers of, 76.
snail carried by, 89.
Todd, J. E., on a turtle carrying a
mussel, 83.
Tornadoes, 149.
Totanus, bivalves clinging to, 79.
Tothby, isolated pond at, II.
Trafalgar Square fountains, shells
in, -22.
Tree-snails, dispersal of, 136.

INDEX.

Trees, molluscs living under the
bark and in hollows of, 132,
169, 173.
eating into the wood of, 133,
170.
And see Timber.
Trent Valley, Mr. Musson on the
abundance of shells in, 31.
Tristram, H. B., snail’s eggs found
on the foot of a bird by, 47,
157.
on colonization in co. Durham,
193, 254. ;
Trochomorpha, distribution of, go.
Troughs, shells in, 20.
Trout, a Zzmnea transplanted as
food for, 184.

Turtle, mussel carried by, 83.
pond-snails adhering to, 88.
Turton, Dr., on shells amongst

leaves, 146.
on Bulimus goodallz, B. decol-
latus and Helix pomatia in
Britain, 230, 238, 260.
Tweedie, F., on the finding of a
bird’s claw between the
valves of a mussel, 77.
Tye, G. S., on Dretssena fpoly-
morpha in water-pipes, 19.
on Helix pisana in Guernsey,
190.
Typhoons, 131, 149.

w:

UMBELLIFEROUS plants, molluscs
im the kexes of, 130, 171.
Unio or Anodonta, clinging to a
sandpiper, 79.
to a heron, 8o.
to a duck, 80.
Unio, tenacity of life of, 28.
living among tree-roots, 37.
dispersal of fry of, 48.
clinging by closure, 57, 58.
ducklings caught by, 77.
Unio complanatus carried by a teal,

O.
by a turtle, $3.

289

U. margaritifer, dragged to shore
upon rods, 57, 58.
Ancylus adhering to, 88.
Untonide, dispersal of, by floods,

33:
dispersal of fry of, 48.

United States, snails imported into,
as articles of food, 203.
United States National Museum,

slug found in a bunch of
bananas received by, 207.
Univalves, dispersal of, while cling-
ing by closure of the opercu-
lum, 85, 165.
And see Molluscs.
Upton, attempted colonization of
Flelix hortensts at, 194.

V.

VERNEDI, Mr., on tenacity of life
in the desert-snail, 103.
Veronicella, found among bananas,
207.
Vertigo edentula, in kexes, 140.
amongst leaves, 147.
V. mtlium, weight of, 147.
V. pygmea, colonies of, 94.
Victoria, dust whirlwinds of, 149.
introduced slugs in, 180.
Virginia, Helix nemoralis in, 180,

193.

Vitality, suspended. See Tenacity
of life.

Vitrina (fHelicarion) carried by a
beetle, 155.

Vole, trapped by a mussel, 83.

Voluntary migration, 4, 26, 116,

144.
W.

WAGTAILS, earth on the feet of,

159.
Walken, J., efa with a bivalve
attached caught by, 62.
Wallace, A. R., on the general per-
manence of oceans, I, 118.

U

290

on changes of level, 30.

on salt-water killing snails, 40.

on dispersal of eggs of fresh-
water shells, 46.

on dispersal by water-beetles,
6

on the stocking of oceanic
islands, 92, 164.
on the antiquity of land-shells,
LIS:
on voluntary migration, 116.
on hypothetical continents, 118.
on the want of evidence of
exact modes of diffusion, 118.
on dispersal by rivers and
floods, 138.
by wind, 145, 152.
by birds, 159.
Ward, James, on tenacity of life in
a snail, 107.
Ward, John, living shells found in a
bird’s crop by, 161, 163.
Wardian cases, dispersal in, 198,
207.
Warren, Miss, on dispersal by cattle,
156.
on Lulimus extlis in Ireland,
258.
Warren, R., on a cockle clinging to
a dunlin, 59.
Wars, dispersal during, 203.
Water, dispersal with. See Currents.
isolated. See Ponds.
Water-beetles. See Beetles.
Water-bugs. See Corixa, Nepa,
and Wotonecta.
Water-butt, shells in, 20.
Water-pipes, shells in, 18.
Water-rails caught by oysters, 60.
Water-scorpion. See /Vepa.
aia trapped by a mussel,
ae
Watson, R. B., on _ introduced
molluscs in Madeira, 179.
Watson, W., on Physa acuta at
Kew, 225.
Webb, C. J., whirlwind described
by, 150.
Weir, J. J., on tenacity of life in the
desert snail, 104.
Wells, Artesian, 22.

INDEX.

Welton-le-wold, isolated pond at,

Q, 10.

Westgate, W. W., on dispersal with
plants, 195.

Wheatears, earth on feet of, 159.

Whinchats, earth on feet of, 159.

Whirlwinds, 42, 149, 174.

White slug. See Limax agrestis.

Whitelegge, Mr., bivalves clinging
to water-bugs, 63.

to water-beetles, 65.

Widgeon, probable dispersal by, 50.
Wildman, J. R., on Planorbis dila-
tatus at Burnley, 224.
Wilson, G. F., on Bulimus goodalliz,

22%.

Wind, dispersal by, 42, 145, 174.

Wolds,isolated ponds of,in Lincoln-
shire, 9.

in Yorkshire, Io.

Wollaston, T. V., on the absence
of fresh-water shells from St.
Helena and the Azores, 93.

on tenacity of life in snails,
109.

Wood, dispersal with. See Timber.

Woodcock, cake of earth on the leg
of, 54, 158.

Woodlouse, a snail carried by, 155.

Wood-pigeon. See Pigeon.

Woodruffe-Peacock, E. <A., on
mussels clinging by closure,

57:
Woodthorpe, E., shells collected
from an isolated pond by,

52;
frogs with shells attached ob-
served by, 74.
Woodward, S. P., on isolated colo-
nies of land-shells, 95.
on tenacity of life, 105, 106,
113.
on dispersal by sailors, 203.
on Dreissena polymorpha and
Testacella maugez in Britain,
213, 2209.
Worms, living, ejected from the
crops of birds, 164.
Wotton, F. W., on Bulimus goods
alliz, 231.
on Helix carthaginiensis, 253.

INDEX. 291

on Helix villosa and H. cartu- Helix lactea found in, 252.
stana, 254. Young molluscs. See Fry.
Wright, B. M., dormant snails re-
vived by, 103.
Z.

Me
ZANNICHELLIA, shells adhering to, 53.
YORKSHIRE, isolated ponds on the | Zebra mussel. See Drezssena
wolds of, Io. polymorpha.
foreign snails living in a garden | Zonztesin Paleozoic coal measures
int, 192,263. of Nova Scotia, 115.

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Classified List of Publications. 21

English-Danish Dictionary. By S. Rosinc. Crown 8vo,
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22 Kegan Paul, Trench, Triibner, and Co.’s

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Classitied List of Publications. 23

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ee ee ee Se ee

Supernatural in Nature, The. A Verification by the Free
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Mystery of the Universe Our Common Faith. By Rev. J. W.
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World to Come, The. Immortality a Physical Fact. By Rev.

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Parables of our Lord, Notes on the. By ARCHBISHOP TRENCH.
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Synonyms of the New Testament. By ARCHBISHOP TRENCH.
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Sermons, New and Old. By ArcHBisHop TRENCH. Cvown
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26 Kegan Paul, Trench, Triibner, and Co.’s

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Classified List of Publications, 27

Authentic Gospel, The, and other Sermons. By GEORGE
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28 Kegan Paul, Trench, Triibner, and Co.’s

Doctrine of Annihilation in the Light of the Gospel of Love.
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Christ in Modern Life. By Rev. Sroprorp A. BROOKE.

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Death—and Afterwards. By Sir Epwin ARNOLD. Reprinted
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Classified List of Publications. 29

Jesus, the Carpenter of Nazareth. By A Layman. Third
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Hebrew Theism. By Francis WILLIAM NEWMAN. foyal
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30 Kegan Paul, Trench, Triibner, and Co.’s

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Classified List of Publications. 35

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36 Kegan Paul, Trench, Triibner, and Co.’s

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Geometrical Psychology ; or, The Science of Representation.
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WORKS ON THE PHYSICAL SCIENCES,
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Classified List of Publications. 59

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Classified List of Publications. 61

TECHNOLOGY.

Technological Dictionary of the Terms employed in the Arts
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Classified List of Publications. 63

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POETRY AND BELLES-LETTRES.

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Songs of Two Worlds. Thirteenth Edition.

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Classified List of Publications. 65°

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Concordance to Shakespeare’s Poems. By Mrs. FURNESS. 18s.
E

66 Kegan Paul, Trench, Triibner, and Co.’s

The Works of William Shakspere— Continued.

A New Variorum Edition of Shakespeare. Edited by HORACE
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Index to Shakespeare’s Works. By E. O’CONNoR. Cvowz 8vo, 55.

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Sonnets. Edited by MARK PATTISON. With Portrait. Zilzevir 8vo
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Chaucer's Canterbury Tales. Edited by A. W. PoLiarp-
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Poems of P. B. Shelley. Edited by RicHarD GARNETT.
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The Poetical Works of John Keats. Edited by W. T.
ARNOLD. Large crown 8vo. Choicely printed on hand-made paper.
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Selected Poems of Matthew Prior. Edited by AUSTIN
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Fables of John Gay. Edited by Austin Dosson. With
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ment, 12s. Cheap Edition. Crown 8vo, 4s. 6d.

Wordsworth Birthday Book. Edited by ADELAIDE and VIOLET
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Classified List of Publications, 67

Works of Sir Henry Taylor. 5 vols. crown 8vo, 30s.
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The Virgin Widow, etc. cap. 8v0, 35. 6d.

The Poems of Ebenezer Elliott. Edited by his son, the

Rev. EDWIN ELLIOTT, of St. John’s, Antigua. 2 vols. crown 8vo, 18s.
Poems by W. Cullen Bryant. Cheap Edition. Small 8vo, 35. 6d.

Poems of Edgar Allan Poe. Edited by ANDREW LANG.
With Frontispiece. Elzevir 8vo (Parchment Library). Vellum,
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The Raven: with Commentary by JOHN H. INGRAM. Crown 8vo,
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Poems by Archbishop Trench. TZenth Edition. cap. 8vo,
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Sacred Latin Poetry, chiefly Lyrical. By ARCHBISHOP TRENCH.
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Household Book of English Poetry. Edited by ARCHBISHOP
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Chaucer to Burns. Translations. Lyrics of the Nineteenth Century.
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Selected Poems of Robert Burns. With an Introduction by
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Lucile. By the late EARL or LyTToN. With 32 Illustrations.
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Rhymes from the Russian. By JOHN POLLEN. Transla-
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LElzevir 8vo (Parchment Library). Vellum, 7s. 6d.; Parchment-or
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Living English Poets. With Frontispiece. By WaLTER
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68 Kegan Paul, Trench, Triibner, and Co.’s

———

Sea Song and River Rhyme, from Chaucer to Tennyson.
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Birds and Babies. By ErHeL CoxHeapb. With 33 Lllustra-
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The Christian Year. By J.KesiEe. With Portrait. LZlzevir
8vo (Parchment Library). Vellum, 7s. 6d.; Parchment or cloth, 6s.
The Poems of Wilfrid Scawen Blunt.
The Wind and the Whirlwind. 8vo, Is. 6d.
The Love Sonnets of Proteus. /ifth Huition. Elzevir 8vo, 55.
In Vinculis. With Portrait. /zevir 8vo, 5s.
A New Pilgrimage ; and other Poems. Zizevir 8vo, 5s.
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the collection of Ballads, Sagas, Hymns, and other Pieces known as
the Shih Ching, metrically translated. 8vo, 16s.

Shadows of the Lake, and other Poems. By F. LEYTON.
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The Poems of Mrs. Hamilton King. The Disciples. Zenth
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A Book of Dreams. Third Edition. Crown 8vo, 35. 6d.

Sermon in the Hospital (from ‘The Disciples’). cap. 8vo, 1s. Cheap.
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Ballads of the North; and other Poems. Crown 8vo, 55.

Classified List of Publications. 69

A Lover’s Litanies, and other Poems. With Portrait of
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Goethe’s Faust. Translated from the German by JouN
ANSTER. With an Introduction by BURDETT Mason. With Illus-
trations (18 in Black and White, 10 in Colour), by FRANK
M. Gregory. Grand folio, £3, 35.

French Lyrics. Edited by Grorce Saintsspury. With
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Poems by Alfred Gurney. The Vision of the Eucharist, and

other Poems. Crown 8vo, 55.
A Christmas Faggot. Smal/ 8vo, 5s.
Voices from the Holy Sepulchre, and other Poems. Crowx 8vo, 5s.

Poems by Edmund Gosse. New Poems. Cvown 8vo, 7s. 6d.

Firdausi in Exile, and other Poems. Second E-dition. Elzevir 8vo,
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On Viol and Flute: Lyrical Poems. With Frontispiece by L. ALMA
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London Lyrics. By F. Locker. Tenth Edition. With
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English Comic Dramatists. Edited by OswaLD CRAWFURD.
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Poems by Toru Dutt. A Sheaf gleaned in French Fields.
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Ancient Ballads and Legends of Hindustan. With an Introductory
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St. Augustine’s Holiday, and other Poems. By WILLiam
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A Strange Tale of a Scarabeus, and other Poems. By A.
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A Song-Book of the Soul. By Marjory G. J. KINLOCH.

Crown 8vo, 55.

Lyrics and Other Poems. By Lapy Linpsay. Second Edition,

Elzevir 8vo, 55.
Psalms of the West. Sma// 8vo, 1s. 6d.

Louise de la Valliére, and other Poems. By KATHERINE
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70 Kegan Paul, Trench, Triibner, and Co.’s

Shamrocks. By KATHERINE Tynan. Smadl/ 8vo, 5s.
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Granite Dust. Fifty Poems. By RoNALD CAMPBELL MACFIE.
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Carmela; or, The Plague of Naples. Crown 8vo, 2s. 6d.
The Marriage of the Soul, and other Poems. By W. Scortt-

ELLIOT. Crown 8vo, 5s.

Beauty and the Beast; or, A Rough Outside with a Gentle
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In Hours of Leisure. By Ciirrorp Harrison. Second
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Verses Written in India. By Sir Atrrep LyaLt. L£izevir
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Analysis of Tennyson’s ‘In Memoriam.’ (Dedicated by per-
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India Revisited. By Str Epwin ArnoLp. With 32 Full
page Illustrations. Crown 8vo, 7s. 6d.

Milton’s Prose Writings. Edited by E. Myers. £/zevir
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Select Letters of Shelley. Edited by RicHarD GARNETT.
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Calderon.—Essay on the Life and Genius of Calderon.
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the World.” By ARCHBISHOP TRENCH.- Second Edition, revised
and improved. Extra fcap. 8vo, 5s. 6d.

Confessions of an English Opium Eater. By DE QUINCEY.
Edited by RICHARD GARNETT. L/zevir 8vo (Parchment Library).
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A Word for the Navy. By ALGERNON CHARLES SWINBURNE.
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Biglow Papers. By JamEs RussELL LOWELL. Edited by
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Robert Browning.—Studies in the Poetry of Robert Browning.
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Lost Leaders. By ANDREW Lanc. Cvown 8vo, 55.

Classified List of Publications. 71

Goldsmith’s Vicar of Wakefield. Edited by Austin Dos-

SON. izevir 8vo (Parchment Library). Vellum, 7s. 6d.; Parch-
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Eighteenth Century Essays. Edited by Austin Dosson.
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Four Centuries of English Letters: A Selection of 350
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Munchausen’s Travels and Surprising Adventures. Illus-
trated by ALFRED CROWQUILL. (Lotos Series), 35. 6d.

Specimens of English Prose Style from Malory to Macaulay,
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Macaulay's Essays on Men and Books: Lord Clive, Milton,
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The Countess of Pembroke’s Arcadia. By Sir PuILip
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Carlyle’s Sartor Resartus. Z/zevir 8vo (Parchment Library).
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Swift’s Letters and Journals. Edited by SranLtey Lane
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Swift's Prose Writings. Edited by Stantey LANE POooLe.
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Vagabunduli Libellus. By JoHN ADDINGTON SyMonpDs.
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Disraeli and His Day. By Sir WILLIAM FRASER, Bart.
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NOVELS AND WORKS OF FICTION.

Novels By George MacDonald.
Donal Grant. With Frontispiece. Crown 8v0, 6s. Cheap Edition,
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Home Again. With Frontispiece. Crow 8vo, 6s.

Castle Warlock. With Frontispiece. Crow 8vo, 6s. Cheap Edition,
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72 Kegan Paul, Trench, Triibner, and Co.’s

Novels by George Macdonald— Continued.

Malcolm. With Portrait of the Author engraved on Steel. Crown 8v0, 6s.
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The Marquis of Lossie. With Frontispiece. Crown 8vo, 6s. Cheap
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St. George and St. Michael. With Frontispiece. Crown 8v0, 6s.

What’s Mine’s Mine. With Frontispiece. Crown 8vo, 6s. Cheap
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Annals of a Quiet Neighbourhood. With Frontispiece. Crows 8vo, 6s.

The Seaboard Parish: a Sequel to ‘ Annals of a Quiet Neighbourhood.’
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Wilfrid Cumbermede: an Autobiographical Story. With Frontis-
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Thomas Wingfold, Curate. With Frontispiece. Crown 8v0, 6s.

Paul Faber, Surgeon. With Frontispiece. Crown 8v0, 6s.

The Elect Lady. With Frontispiece. Crown 8vo, 6s.

There and Back. With Frontispiece. Crown 8vo, 6s.

Flight of the Shadow. With Frontispiece. Cvowm 8vo, 6s.

Hawthorne’s Novels and Tales.—Works. By NATHANIEL
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Novels by Col. Meadows Taylor.
Seeta: A Novel. With Frontispiece. Cvowz 8vo, 6s.
Tippoo Sultaun: A Tale of the Mysore War. With Frontispiece.
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Ralph Darnell. With Frontispiece. Crowz 8vo, 6s.
A Noble Queen. With Frontispiece. Crow 8vo, 6s.
The Confessions of a Thug. With Frontispiece. Crows 8vo, 65.
Tara: A Mahratta Tale. With Frontispiece. Crows 8vo, 6s.

Novels by Heshba Stretton.

David Lloyd’s Last Will. With 4 Illustrations. Mew Ladition,
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Novels by Maxwell Gray.
In the Heart of the Storm. With Frontispiece. Cvowmn 8vo, 6s.
The Reproach of Annesley. With Frontispiece. Crown 8vo, 6s.
Silence of Dean Maitland. With Frontispiece. Crown 8vo, 6s.

Novels by Rowland Grey.
In Sunny Switzerland; A Tale of Six Weeks. Second Edition. Small
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Lindenblumen, and other Stories. Small 8vo, 5s.
By Virtue of His Office. Crown 8vo0, 6s.
Jacob’s Letter, and other Stories. Cvowz 8vo, 6s.

Classified List of Publications. 73

—saeer vee

Novels by ‘ Tasma.’
' A Sydney Sovereign, and other Tales. Crows 8vo, cloth, 6s.
In Her Earliest Youth. Cheap Edition. Crown 8vo, 6s.

Novels by Lucas Malet.
Colonel Enderby’s Wife. With Frontispiece. Crown 8vo, 6s.
A Counsel of Perfection. With Frontispiece. Crown 8vo, 6s.

Little Peter: a Christmas Morality for Children of any age. With
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Stories by Mrs. G. S. Reaney.

Waking and Working; or, From Girlhood to Womanhood. Mew and
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Blessing and Blessed: a Sketch of Girl Life. Mew and cheaper Edition.
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Rose Gurney’s Discovery: a Story for Girls. Dedicated to their
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English Girls: Their Place and Power. With Preface by the Rev. R.
W. DALE. Fifth Edition. Freap. 8vo, 2s. 6d.

Just Any One, and other Stories. With 3 LIllustrations. 160,
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Sunbeam Willie, and other Stories. With 3 Illustrations. 16770,
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Sunshine Jenny, and other Stories. With 3 Illustrations. 1670,
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The Prig.—Black is White; or, Continuity Continued. 3s. 62.

The Prigment: ‘The Life of a Prig,’ ‘ Prig’s Bede,’ ‘ How to make a
Saint,’ ‘ Black is White.’ Second Edition. In 1 vol. crown 8v0, 5s.

A Romance of the Recusants. Crown 8vo, 5s.
Sultan Stork, and other Stories and Sketches, 1829-44. By

W. M. TuHackeray. Now first collected, to which is added the
Bibliography of Thackeray. Large 8vo, Ios. 6d.

In the Wrong Paradise, and other Stories. By ANDREW
LANG. Crowz 8vo, 6s.

Riches or Ruin. A Story. By the Author of ‘The Prig-
ment.’ Crown 8vo, 35. 6d.

Egosophy. By the Author of ‘The Prigment.’ Cvown
8v0, 35. 6d.

Strange True Stories of Louisiana.. By G. W. CABLE. 8v0,
7s. 6d.

South Africa, An I. D. B. in South Africa. By LOUISE

VESCELIUS-SHELDON. Illustrated. Crown 8vo, 7s. 6d.

Zululand.—Yankee Girls in Zululand. By Louise VEs-
CELIUS-SHELDON. Illustrated. Crowz 8vo, 5s.

74 Kegan Paul, Trench, Triibner, and Co.’s

Tyll Owlglass’ Marvellous and Rare Conceits. Translated by
KENNETH MACKENZIE. Illustrated by ALFRED CROWQUILL.
(Lotos Series), 35. 6d.

The Fortunate Lovers : Twenty-seven Novels of the QUEEN
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A Professor of Alchemy. By Percy Ross. Crown 8vo, 35. 6d.
The Wing of Azrael. By Mona Cairn. Crown 8vo, 6s.
Off the Skelligs: A Novel. By JEAN INGELow. With Frontis-

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My Ducats and My Daughter. By Hay HunNTER and WALTER
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Plucky Fellows: A Book for Boys. By S. T. MacKEnna.
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Mosquito: A ‘Tale of the Mexican Frontier. By FRANCES
FRANCIS. Crown 8vo, 35. 6d.

The Pillar House. By FLORENCE SEVERNE. With Frontis-
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Sylvia Arden. By OswaLp CRAWFURD. With Frontispiece.
Crown 8vo, Is.
Within Sound of the Sea. With Frontispiece. Cvrozzn 8vo, 6s.

Castle Blair: A Story of Youthful Days. By Fiora L. SHaw.
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The Apothecary’s Daughters. Translated from the Danish
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Scot Free: A Novel. By C. G. Compton. Cvown 8vo, 6s.

Christy’s Inheritance: A London Story. By FLORENCE
WHITAKER. Illustrated. Royal 16mo, 1s. 6d.

A Lost Love. By Anna C. OGLE. Small crown 8vo, 25. 6d.
Through To-day. A Novel. Crown bvo, 5s.
The Story of a Friendship. By the Rev. ALFRED GURNEY.

Crown 8vo, 5s.

His Heritage. By LinDA GARDINER. With Frontispiece.
Crown 8vo, 6s.
Novels by Rosa Mulholland.

Marcella Grace: an Irish Novel. Crown 8vo, 6s.
A Fair Emigrant. With Frontispiece. Crown 8vo, 6s.

Classified List of Publications. 75

ne Ee

A Sensitive Plant: A Novel. By E. and D. GERARD.

Crown 8vo, 6s.

Ulli: The Story of a Neglected Girl. Translated from the

German of EMMA BILLER, by A. B. Daisy Rost. Crown 8vo, 6s.

God’s Providence House. By Mrs. G. Linn&us BANKS.
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At Sundry Times and in Divers Manners. By May
ELEANOR BENSON. With Portraitand Memoir. 2 vols. Crown 8vo,
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BIBLIOGRAPHY.

Periodical Literature, Index to. By W. F. Pootr. Zzird
Edition. Royal 8vo0, £3, 135. 6d.

Periodical Literature, Index to. First Supplement. By W.
F. Poor and W. I. FLETCHER. 1882 to 1887. Royal 8vo, £1, 16s.

American Literature, Triibner’s Bibliographical Guide to.
From 1817 to 1887. 8vo, half-bound, 18s.

Dictionaries and Grammars, Triibner’s Catalogue of Dic-
tionaries and Grammars of the Principal Languages and Dialects of
the World. Second Edition. 8vo, 5s.

Memoirs of Libraries, together with a Practical Handbook
of Library Economy. By EpwarpD Epwarpbs. Numerous Illustra-
tions. 2 vols. Royal 8vo, £2, 8s.

Libraries and Founders of Libraries. By Epwarp Ep-
WARDS. 8vo, 18s.

Free Town Libraries: Their Formation, Management, and
History, with brief notices of Book Collectors. By EDWARD
EDWARDS. 8v0, 21s.

Philobiblon. By RicHARD DE Bury. Edited by E. C.

Tuomas. Crown 8vo, tos. 6d.

Ezypt and the Soudan, The Literature of: A Bibliography,
comprising Printed Books, Periodical Writings, and Papers of
Learned Societies. Maps and Charts, Ancient Papyri Manuscripts,
Drawings, etc. By H. H. Prince IpRAHIM HILMy. 2 vols, demy
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