THE SEA-TROUT
A STUDY 1N
NATURAL HISTORY
HENRY LAMOND
The Sea- Trout
EMO V-alisy Av.
ie oeca-l rout
A Study in Natural History
BY
HENRY LAMOND
Secretary of the Loch Lomond Angling Improvement A ssociaticn
London
Sherratt & Hughes
Manchester: 34 Cross Street
1916
Contents.
PAGE
PREFACE - - - - - - - - . - 1
Cuap. I. Introductory—Trout and Sea-Trout - - 5
» ‘&. The Sea-Trout - . - - - - 45
» Ill. Eggs and Alevins - - - : - - 69
» lV. The Fry Stage - - - - - = 81
= V. Parr and Smolts - : - - - - 89
» VI. Whitling - - - - - - - - 117
» WII. Maturity - - - - - = = - 11837/
, WIII. The Spawning Period - - - = - 157
, IX. Artificial Propagation - = - - - 177
ns X. Concluding Remarks - - . - - 203
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List of Illustrations.
PLATE I. A Sea-Trout (in Colour) - - - - - Frontispiece
Face page
”
II. Development of the Salmon from the Egg to the Smolt
Stage (in Colour) - - - - - . -
Ill. Development of the Sea-Trout from the Egg to the Smolt
Stage (in Colour) - - - s 5 <
IV. A Trout, a Loch Leven Trout, a Sea-Trout, and a Salmon
as “‘ Yearlings”’ (in Colour) - -~ -
V. A Trout, a Loch Leven Trout, a Sea-Trout, and a Salmon
as ‘‘ Two Year Olds’? (in Colour) - - - -
VI. A Silvery Loch Lomond Trout (in Colour)
VII. A Young Salmon assuming the Smolt Dress (in Colour)
VIII. An Exceptionally Large Sea-Trout Smolt (in Colour)
IX. A Spawning Male Sea-Trout (in Colour)
‘Fresh Run from the Sea’ - . - - - . Face page
FIG
1. A Salmon, showing normal spots of the ‘‘ maiden ’’ fish - -
2. A Salmon, showing speckled appearance of a fish which has
previously spawned - - - - - - : :
3- Two Loch Lomond Sea-trout, showing typically distinctive
markings” - - - : - = - - - : 3
4. A male Sea-trout at spawning time - - - : - -
5- A “two year old”’ Trout (fario) with ‘‘ halo’? surrounding the
spots strongly accentuated - - - - - - - -
6. Two “ two year old ”’ Trout (fario) showing normal markings -
je NoLwerianigiver (rout (fario) ~- - - = «= -«
8. A Clyde river Trout (fario) — - : = : - : : -
g. A Loch Lomond Trout (fario) showing spots geometrically arranged
10. A Loch Lomond Sea-trout showing spots geometrically arranged
11. Diagram showing arrangement of the fins of Salmon, Sea-trout
and Trout -~ - - fy Sa es Pi
12. Diagram contrasting tails of Salmon and Sea-trout - - -
13. Diagram contrasting heads of male Salmon and Sea-trout
ix
“I
No
14.
iit)
LIST OF ILLUSTRATIONS
Fac page
Diagram contrasting heads of female Salmon and Sea-trout - .
Diagram of gill-covers of Salmon Trout, Grey Trout, and Salmon
(after Yarrell, reversed) - - - = - - - -
Scales of a Sea-trout and a Grilse of equal weights contrasted to
show different rates of growth - - - - : - -
Vomer of a Clyde river Trout showing dentition - - - -
Vomer of a Loch Lomond Sea-trout showing dentition - -
Diagram contrasting outlines of tail of a Tweed ‘ Bull Trout ”’
and of a Loch Lomond Sea-trout — - - - - - -
“Bull Trout’’ (3}1b.) from the River Aln_ - :
Scale of River Aln “ Bull Trout’’ (3}1b.) indicating growth
similar to that of a salmon grilse - - - - - -
“Bull Trout ”’ (8 lb.) from the River Aln - - - - -
Scale of River Aln “ Bull Trout’? (81lb.) indicating growth
similar to that of a small summer salmon - - - - -
A Norwegian ‘ Bull Trout ’’ (83 Ib.) - - - - - -
Sea-trout eggs - - = : : - 4 : : = 5
ae ’
Sea-trout ‘‘eyed ova”? - - - = : : : :
Sea-trout alevins hatching out - - = : : 5 5
Late eggs and early alevins” - - - : - : : Z
Sea-trout ‘* monstrosities ’’ (after Gemmill) - - -
Scale of a late-descending Sea-trout smolt - - - - -
Scale of Sea-trout indicating a 2-winters’ residence in fresh water
prior to migration - - - - - - - - -
Scale of Sea-trout indicating a 3-winters’ residence in fresh water
prior to migration - : - - - - - : -
Scale of Sea-trout indicating a 4-winters’ residence in fresh water
prior to migration - - - - - 5 : =
Scale of Sea-trout Smolt indicating a 4-winters’ residence in fresh
water prior to migration - - - : - : - -
Scale of Sea-trout indicating a 5-winters’ residence in fresh water
prior to migration - - - - : - : : E
Scale of Norwegian Sea-trout indicating a 5-winters’ residence in
fresh water prior to migration - - : : : :
A typical Whitling - - - : - - : 2 2
Scale of Sea-trout illustrating period spent in the sea after descent
as a smolt and prior to return as a whitling : : : :
59
59
61.
Plan of Luss Hatchery — - - : : : 5 3
LIST OF ILLUSTRATIONS
Face
Scale of a very small Norwegian Sea-trout — -
Seale of a small Norwegian Sea-trout — - - - - -
Scale of Sea-trout indicating that the fish spawned as a whitling -
Scale of Sea-trout indicating that the fish had spawned as a whitling
Scale of Sea-trout indicating that the fish had spawned asa whitling
Scale of Sea-trout indicating that the fish had not spawned as a
whitling z E Z e Z E Z 2 - i ~
Scale of Sea-trout indicating that the fish did not remain over
winter in the sea after its descent as a smolt = : = -
Scale of Sea-trout indicating that the fish passed 1 winter in the sea
after its descent as a smolt - - - - : : s
Scale of Sea-trout indicating that the fish passed 2 winters in the
sea after its descent as a smolt - - - - - - -
Scale of Sea-trout indicating that the fish passed 3 winters in the
sea after its descent as a smolt - - - - - - -
Scale of Sea-trout which, although 11 years old, weighed only 23 Ib.
A Salmon leaping at a fall - - - - - - - -
A male Salmon in spawning dress - . - - . - -
A female and male Salmon on the eve of spawning . - -
A male Sea-trout in spawning dress - - - - - -
A female kelt Sea-trout - - < : : : : = =
Scale of Sea-trout indicating spawning in 2 consecutive years -
Scale of Sea-trout indicating spawning in 3 consecutive years -
Scale of Sea-trout indicating spawning in 4 consecutive years -
Scale of Sea-trout indicating spawning in 5 consecutive years -
Scale of Norwegian Sea-trout indicating spawning in 7 consecutive
years - - - eh De ie 4s oe aaa ae
A Norwegian Sea-trout (11}Ib.) - - - . op
Scale of Sea-trout, recaptured when spawning in 3 consecutive years
Scale of Sea-trout indicating that the fish spent 3 consecutive years
in the sea after migration as a smolt - - - - -
xl
181
od
_— -) Se.
PREFACE
Ir may, I think, be gathered from the general conversation of one’s
friends who are anglers, and, indeed, from much that one finds in the
literature of angling, that considerably less is definitely known regarding
the sea-trout than is popularly known nowadays regarding the salmon.
The truth is, that the salmon has, until very recently, practically
monopolised the serious attention of investigators of the life-history of
our more sporting fishes.
I do not of course mean to say that the sea-trout has hitherto been
wholly neglected. Much useful and interesting information regarding
it may be found scattered here and there in the pages of books on fish
and fishing, but I do not know of any recent work which bears to be
primarily a study of the gamest of fishes.
Dr. Francis Day, in his “ British and Irish Salmonidz,” published
in 1887, gave us perhaps the most exhaustive study of the sea-trout that
has yet been written, and, more recently, Mr. C. Tate Regan, in his
excellent treatise on “ The Freshwater Fishes of the British Isles,”
published in 1911, emphasised some of the more striking of Dr. Day’s
views. But Dr. Day, when he wrote, was not in a position to take
any very great advantage of such evidence as we know can be
gleaned from a study of the scales of the Salmonide, and he had not
before him, when he wrote, such definite data as we now know can be
gathered from the systematic marking of individual fish. Both of these
lines of inquiry, as I hope to show, help to shed a clearer light upon
much that is obscure, or that rests merely upon theory, in the
life-history of the sea-trout, and it is, I think, on the evidence that will
be obtained from further investigation along these lines that a complete
I
2 THE SEA-TROUT
understanding of the life-history of the sea-trout—if not also of the
trout—will ultimately be arrived at.
These pages pretend to give only a bare outline of what is known
concerning the sea-trout’s career; only in a few instances have I
ventured, lightly and tentatively, to fill in the details of the picture.
As Mr. W. L. Calderwood says, “ We require rather fuller knowledge
of the habits of this minor salmon.” His note upon “ Sea-trout ” in the
most recent Report of the Fishery Board for Scotland (namely that for
the year 1914) leads one to hope that his careful and painstaking work
as Inspector of Salmon Fisheries for Scotland will in time bring us to
that fuller knowledge. A “ Life of the Sea-trout” from his pen would
be an appropriate and acceptable companion picture to his “ Life of
the Salmon.”
I think I have in most instances in the text made suitable acknow-
ledgment of such information as has been freely and courteously given
me; here, in a word, I may briefly and generally thank all those to whom
I am indebted for valuable assistance in the preparation of this book.
A more definite tribute, however, must be paid to my friend, Mr. J.’
Arthur Hutton, of Manchester, who has been so largely instrumental
in bringing the Wye to its present state of productivity. Mr. Hutton’s
experience in the reading of salmon scales is, it is safe to say, now
probably unequalled, and he has been so generous as to apply his
knowledge definitely on my behalf to the elucidation of a large number
of sea-trout scales. He was, in particular, good enough to examine
for me a series of scales of Loch Lomond fish with which I supplied
him during the season of 1914. Without the data so derived this book
would have been lacking both in interest and value. I may explain
that we jointly agreed as to the accuracy of the reading of each scale
examined, and if it would be not wholly fair to myself to say that all
my conclusions based upon such readings are merely Mr. Hutton’s
conclusions, it is at least fair to him to say that his greater experience
PREFACE 3
led him to suggest to me wherein scale reading might be of value in
confirming as fact what had otherwise remained very much in the region
of theory. For example, I do not think I could have realised how
protracted the young sea-trout’s residence in fresh water might be before
migration without Mr. Hutton’s valuable help. I am further indebted
to him and to his skill with the camera for all photographs of scales
and for most of the photographs of fish reproduced. To him, also,
and to the Rev. Alex. Slater Dunlop, B.D., Minister of Luss, I am
indebted for a revision of the proof sheets.
As to the other illustrations, failing any skill with the camera, I have
perforce had to fall back upon such drawings as I could myself accom-
plish with pen and brush. They are intended to be viewed rather as
diagrams than pictures, diagrams which can be conveniently compared
one with another.
In spite of its many defects I hope this monograph may help to
induce others to apply themselves to a study of one of the most
interesting of our British fishes.
Ho,
Grascow, ist February, 1916.
Introductory
Trout and Sea-Trout
CHalaleER I.
INTRODUCTORY—TROUT AND SEA-TROUT
THERE is one question—not quite so simple as it looks—that appears
to me to require some investigation before any inquiry into the life-
history of the sea-trout is undertaken, and that question is—What is a
sea-trout? It is, one must admit, a question which has not yet been
very satisfactorily answered by anybody, nor can I pretend to give a
very satisfactory answer myself; but several writers appear to me to
have sought for the answer in the right direction, and I shall endeavour
in this introductory chapter to indicate to the reader what that direction
is.
In a difficulty it is often prudent to consult a lawyer, and as our
question is one of admitted difficulty, I think I cannot do better at the
outset than take the opinion of an eminent Scottish legal authority who
gave this matter some consideration. In the well-known “ Treatise on
the Law of Scotland relating to the Rights of Fishing,” by Mr. Charles
Stewart, Advocate, where the extent of the Crown’s right at common
law to the ownership of salmon and salmon fishing is under discussion,
there occurs this note :—‘“ The Crown holds the right of fishing salmon,
but it may be questioned whether that right extends to all fish of the
salmon kind. In recent salmon legislation, the word ‘salmon’ is
expressly declared to possess the wider significance, but natural
history and the common interpretation scarcely warrant the construction.
Sea-trout and bull trout are perhaps ‘ fish of the salmon kind,’ and the
Salmon Acts expressly declare them to be included in their operation ;
but in a question of title, can the right to fish for them be held to be
inter regalia? The vationale of the Crown’s right to salmonis .... .
the value of the fish, an argument which does not apply with equal force
to bull trout, herling, etc. These fish, along with the salmonide of
7
8 THE SEA-TROUT
smaller size, inhabit the lesser rivers and streamlets in common with the
yellow trout, from which they are almost indistinguishable in size, and
it seems anomalous that to fish for the one a Crown charter should be
necessary, while the right of fishing for the other is an accessory of the
soil. If the whole race of salmonide, including bull trout, herlings,
etc., are to be considered as infer regalia, the best practical test of royal
property would be the migratory habits of the fish.”
It will be observed that this legal authority, even so recently as
1892, found himself faced with the difficulty of accounting in a rational
manner for the inclusion of “ sea-trout and bull trout” and “ herlings,
etc.,” by implication along with salmon in the Crown’s right of salmon
fishing. He seems to have doubted whether the implication could
really be justified, notwithstanding the migratory test, and he clearly
considered that ‘“‘ natural history and the common interpretation” did
not warrant the inclusion of the fishes he somewhat tentatively named
¢ >
within the term “salmon,” although “recent salmon legislation” so
included them.
His hesitation was commendable, because even yet the Crown’s rights
at common law, in Scotland, so far as the inclusion of the sea-trout is
concerned, do not seem to have been definitely established by a decision
of the Courts. Ina case which was decided by Lord Johnston, one of
the judges of the Court of Session, in 1907,’ the question at issue was
whether salmon fishings in Orkney were inter regalia, and it was held
that they were not. But the question was incidentally mooted whether
sea-trout are “salmon” according to the common law of Scotland. Lord
Johnston devoted some part of his judgment to the point (which,
however, for the purposes of the case it was unnecessary to decide), and
said that, according to his own impression, a right of salmon fishing did
embrace the right of sea-trout fishing. But at the same time he
remarked that it was not conclusive on the point that the whole migra-
1. Lord Advocate vy. Balfour, July 20, 1907. Session Cases, 1907, p. 1360,
INTRODUCTORY 9
tory Salmonide were protected by statute, although the migratory
element helped to influence the affirmative view which he expressed.
It was this same element which had already prevailed in fixing the
definition of the word “salmon” in the Scottish statutes, and one may
trace the history of that definition in a very few words.
Salmon fishing appears to have been, in early times in Scotland, a
favourite topic of legislation, for the subject is dealt with in very many
of the Acts of the old Scots Parliament. One does not find many
references, however, in these old Acts, to ‘‘ other fish of the salmon
kind” than salmon. The earliest reference apparently occurs in an
Act of 1318, in the Bruce’s reign, where regulations are made as to
various kinds of obstructions in tidal waters which prevent the ascent
or descent “ salmunculi vel smolti seu fria alterius generis piscitum maris
vel aque dulcis,” that is to say, young salmon or smolts or the fry of
“ other kinds” of sea or fresh-water fishes. Another Act, that of 1426,
refers to “ salmondis and uthir fische.” The Act of 1469, again, more
definite, refers to “fisch, salmon grilses and trowtis.” Beyond such
rather vague references, in a series of some twenty Statutes, I know of
nothing in the old Acts which would extend the whole scope of the laws
relating to the salmon and its young to other fish than salmon.
It is in the modern series of Statutes that a definite extension of the
meaning of the word “salmon” first appears. In the Act of 1828, the
‘
first of the series, the fish are grouped as “ salmon, grilse, sea-trout, or
other fish of the salmon kind.” Then the Act of 1844 applies to
“salmon, grilse, sea-trout, whitling, or other fish of the salmon kind,”
whitling being thus specifically mentioned. But the Act of 1862, in
the definition now in force, dropped the whitling, added bull trout,
smolts and parr, and introduced the term “ migratory.” Thus, since
1862, the word “salmon” has, in statutory law, meant and included
“ salmon, grilse, sea-trout, bull trout, smolts, parr, and other migratory
fish of the salmon kind.”
fo THE SEA-TROUT
My object is, of course, to emphasise the fact that our knowledge
of the natural history of the sea-trout, as reflected in the law of the land,
is of a somewhat vague and indeterminate kind, the only characteristic
of the fish relied upon to establish its identity being the migratory
habit—a habit shared in an equal degree by the salmon, and, as is well
known, ina less degree by the trout. It occurs to me to inquire whether
the importance which has been attached by the legislature to the idea
‘
involved in the term “ migratory,” and the consequent legal classifica-
8 y q g
tion of the fish as a “‘ salmon,” has not helped to some extent to obscure
our perception of the real nature of the sea-trout.
I do not deny that in some respects the legal definition has served
its purpose admirably, even if, as some have hinted, the rights of
salmon fishery proprietors have incidentally been greatly enhanced
thereby, but in many respects I think the sea-trout has suffered from
being classed as a kind of poor relation of the salmon. The sea-trout
fisheries of Scotland, whether alone or supplementary to salmon fisheries,
are of great commercial value, and yet in almost every particular,
as regards legislation, the interests of the sea-trout have been relegated
to a subordinate position—if they have not, indeed, sometimes been
wholly lost sight of—the assumption apparently being that any enact-
ment appropriate to the salmon must be equally appropriate to the sea-
trout. So much, hitherto, has the salmon dominated the situation, that
the Royal Commissioners of 1902, upon whose recommendations any
fresh salmon fishery legislation will in all likelihood be based, virtually
apologised for exceeding their commission in considering the case of
the sea-trout at all. Moreover, it was only in 1913 that any specific
information regarding sea-trout began to be returned to the Fishery
Board for Scotland by the Clerks of the District Fishery Boards in
their annual reports. If the change came about partly owing to
representations on the subject which I had made from time to time to
INTRODUCTORY Il
my friend Mr. W. L. Calderwood, H.M. Inspector of Salmon Fisheries
—as to which | have no information—lI shall be the better pleased.
In another direction also I think the sea-trout has suffered from the
idea that it is a sort of inferior salmon—an idea, still based, I have no
doubt, upon its migratory habit.
Every writer on salmon fishing—there is hardly any exception—
used to feel it incumbent upon him to refer to sea-trout fishing, and
apparently most of these writers imagined, as Mr. John Bickerdyke in
“The Book of the All-round Angler” does, “that to fish for large
fresh-run sea-trout is to fish for salmon on a small scale.” Mr. Francis
Francis, in devoting two pages of “A Book on Angling ” to sea-trout
c
fishing, concedes that “the sport is little inferior to the best grilse-
fishing.” Sir Herbert Maxwell himself is hardly more generous to
the gamest of fishes. The net result has generally been rather a
travesty of the character of the sea-trout and of the methods of fishing
for it.
I must not be understood to say that no writer who treats of salmon
and salmon fishing is to be debarred from treating of sea-trout and
sea-trout fishing at the same time, but I think that before doing so in a
perfunctory chapter at the end of his book, the writer should consider
fairly whether the sea-trout does not, if not on its own account, at least
on account of the relationship it bears to each, deserve some greater
share of the recognition which has been so freely lavished on the salmon
on the one hand, and on the trout on the other. There are signs, how-
ever, that this state of indifference regarding the character of the sea-
trout, both as a sporting fish and as a study in natural history, will not
long continue, for I think that many have recently had their interest
aroused and their curiosity awakened in the subject.
If lawyers, then, and, as I fear, many angling writers, by concen-
trating their attention too fixedly upon the “migratory ” habits of the
sea-trout—or in sheer ignorance—have tended to confuse the mental
12 THE SEA-TROUT
picture which we form of it, one may now inquire how far the student
of natural history has helped to bring the picture into proper focus.
Here again, at the very beginning, one who, like myself, is
accustomed to the practical rather than the scientific aspect of fish life,
will encounter a difficulty, and perhaps I can best explain it by quoting
from a letter on the subject which I have received from one of the
students aforesaid.
“Tf you look,” writes my friend, “at the Article ‘ Ichthyology’ in
the ‘Encyclopedia Britannica’ (last edition), you will see that it is
divided into two sections, one on the morphology of fishes and one on
the classification of fishes. Now this division is a simple expression
of the fact that the whole subject is so divided and that the workers
upon it are similarly divided. The two types of workers occupy
somewhat different standpoints towards such conceptions as variety,
species, family, and so on. The morphologist looks on them as merely
convenient groups into which to arrange animals, expressive of their
blood-relationship in an evolutionary sense. He is interested in the
features of the various groups in so far as they are evidences of this
blood-relationship. He does not admit that there exists in nature any
general grouping of animals agreeing with what we call genera, families,
and so on. He says that you can only form a conception of ‘ genus,’
or ‘ family,’ or ‘ order,’ if you restrict your view to one isolated group of
animals. Thus within the group ‘ Birds,’ you may have a fairly clear
idea of what you mean by each of these terms; similarly within the
group ‘ Dipterous Flies,’ or within any other group. But he does not
see any evidence that ‘family, or ‘genus,’ means the same thing in
these different groups. In each case it is simply a convenience evolved
by specialised systematists within the particular group. The sys-
tematist, on the other hand, tends to look on such groups as represent-
ing, so to speak, well-defined conceptions in nature. His interest is in
cataloguing animals and separating them into the various groups. He
INTRODUCTORY 13
works away at minute differences in detail which the morphologist is
apt to regard as relatively trivial. The whole outlook is different.”
Now it seems to me to be necessary, in the inquiry to which we have
addressed ourselves, namely, What is a sea-trout? to recognise that there
are, in fact, two schools holding such very divergent aims, because it
may quite conceivably be important that we should be able to distin-
guish at any time whether a particular writer is discussing a particular
specimen as a morphologist or as a systematist. I can readily
understand that a failure to recognise this distinction may have led to
a misunderstanding of a particular writer’s views, and | think that if a
writer has not made it clear in what capacity at the moment he is
discussing any specimen of the Salmonide a certain confusion of
thought may have unwittingly been engendered in the minds of his
readers.
It occurs to me, for example, that much of the uncertainty which
at present exists as to the identity of the “ bull trout’? with the sea-
trout—one party strongly contending for identity and another party as
strongly contending for variety—rests upon the point of view of the
disputant as a morphologist or a systematist. If the former, he will
consider the “ bull trout” as a sea-trout and of interest as being of the
Salmonide@, if the latter, he will discover a marked distinction between
the two fish on the alleged ground that in mature specimens of the
sea-trout the tail is square whereas the “bull trout” has a tail so
convex in its margin that the fish has, in consequence, in certain
©
localities, been popularly designated the “round tail.” The morpho-
logist will agree that the family Sa/monide@ is an interesting study; but
the systematist, intent on cataloguing his “ bull trout,” will classify it, if
not as a separate “species” of the “genus” Salmo, at least as the
b ‘
“variety” erviox of the “ species” ¢rutta.
To take now another example of even greater pertinency to our
subject. Viewing the family Sadmonide, the morphologist will, for his
14 THE SEA-TROUT
own convenience, accept the systematist’s division of the family into
four genera, and of the genus Sa/mo will perhaps readily agree to a
distinction being drawn between two species, namely, Salmo salar and
Salmo trutta; but it will not interest him further to take cognisance of
minor distinctions, and discuss or dispute the existence or non
existence of variations of the species ¢va/¢a which may lead one
systematist to classify certain specimens as different species of Salmo
and another systematist to classify certain other specimens as other
species of Salmo, nor will he be much interested in whether there are
varieties of such alleged species, migratory or non-migratory, round-
tailed or square-tailed, plain or speckled. The systematist, for his
part, is ridden by the necessity of his point of view, and he discovers
in the genus Salmo first one species and then another, and thereafter
proceeds to find varieties and even sub-varieties in each.
I think it is not improbable that, coupled with the confusion of
thought already alluded to, this industry of the systematist, in dividing
and sub-dividing the grouping of the subject of his study, has accounted
for a great deal of misunderstanding in the popular mind regarding the
sea-trout. For, consider, of the genus Salmo, while the authorities are
agreed that, in European waters at least, there are no varieties of the
species salar, in other words that there is only one salmon, yet of the
trout, ¢vwtta, there are endless varieties alleged, if they are not indeed
held to be other species of Salmo. And, besides the migratory sea-
trout, Salmo trutta, one finds constant reference being made to Salmo
eriox, Salmo cambricus, and Salmo albus as other migratory varieties
of trutta if not species of Salmo; and, besides Salmo fario, generally
and somewhat arbitrarily accepted as a non-migratory fish, one finds
everywhere references to Salmo ferox, Salmo levenensts, Salmo
nigripinnis and Salmo stomachicus as other non-migratory trout; and
even an estuarine trout has been differentiated by the name of Salmo
INTRODUCTORY 5
orcadensis, though whether it is claimed as a migratory or non-migratory
trout is not clear.
Now it long ago occurred to me that there was a welter of confusion
in all this differentiation which might possibly be accounted for by the
fact that scientific systematists, who had no great practical knowledge
of the actual ways of fish, had lent too credulous an ear to local popular
beliefs in exceptional types, while, on the other hand, practical men
who had no scientific training at all had acquired the habit of claiming
as the attributes of some new species what were really after all only
superficial differences in some known species.
The somewhat specialised nature of my work with the Loch Lomond
Angling Improvement Association, carried on for over fifteen years,
has made it possible for me to appreciate the efforts of those who have
made simplication, and not complication, of our subject their aim and
study.
Some few writers had already conceived of there being only one
species of our British trout, and perhaps Dr. Francis Day, the author
of “British and Irish Salmonid@,’ published in 1887, is the most
representative of those whose views tended in that direction. In his
preface he says :—‘“ It has been increasingly evident to me for some
years that one of the main reasons militating against the successful
cultivation of trout (and possibly char) in this kingdom by riparian
proprietors, has been the confusion into which these forms have been
thrown by naturalists, who, in order to give greater accuracy to their
descriptive treatises on Museum specimens, have subdivided them into
many species. The consequence of this has been, that fish-culturists
who have accepted the statements made, have been constantly attempt-
ing to introduce new species into their waters in order to improve the
native race. For the zoologists who have been most active in raising
local varieties to specific rank have been compelled to admit that they
all very commonly interbreed, but that the young revert to one of the
16 THE SEA-TROUT
original parents.” In spite, however, of Day’s painstaking and
exhaustive treatise, which every angler ought to make a point of reading
for the wealth of information it contains regarding the sporting
Salmonide, | do not think that many anglers were prepared to learn, as
they did from an excellent work on “ The Freshwater Fishes of the
British Isles,’ by Mr. C. Tate Regan, Assistant in the Zoological
Department of the British Museum, which was published in 1911, that
“In the British Isles there is only one species of Trout.”
This authoritative pronouncement came with something of a shock
upon all but those anglers who had devoted some little study to the more
scientific literature of their craft, or who, from their personal observa-
tions made upon fish life, had formed conclusions leading in the same
direction. Others were frankly sceptical, and I for one see no reason
to blame them, nor indeed need anyone hastily jump at conclusions.
The whole subject demands more complete investigation at the hands
of competent students. Meantime it has to be noted that Mr. Regan
regards the single species as a “variable species,’ and anglers who
recognise the fact that trout are well-nigh infinite in their markings and
vary extraordinarily in their general characteristics according to environ-
ment, will agree with his qualification, whether they accept his main
proposition or not.
Mr. P. D. Malloch, in his “ Life-History and Habits of the Salmon,
Sea-Trout, Trout, and other Freshwater Fish,” first published in 1910,
a work justly popular on account of the excellence of its illustrations,
had, of course, already prepared our minds to some extent. With
reference to the “ Sea-trout ” (Salmo trutta) he writes :—‘ Having made
a careful study of this subject for thirty years, and having collected all
the information I could bearing upon it, I have come to the conclusion
that the sea-trout is the same fish as that which is called white trout,
bull-trout, gray trout, peel, sewin, brith-dail, salmon-trout, and many
other names.” Then of “ The Brown Trout” (Salmo fario), which he
INTRODUCTORY 17
also designates “the common yellow trout,” he writes :—‘* Many
naturalists maintain that there are different species of trout in the British
Islands—Loch Leven trout, Gillaroo trout, tidal trout, and many others;
but from a close study of all these trout for the last forty years, I have
come to the conclusion that there is only one species of trout in Great
Britain, and that in the different varieties the differences are caused by
the nature of the water in which they are found and by the food they
”
eat.” These views are valuable as being those of a practical man, who,
had he studied the sea-trout as long as he had studied the trout, might
have simplified matters still further. It was left to Mr. Regan, as a
scientific man, to take the more advanced step. ‘Although the silvery
Sea-trout and non-migratory Brown Trout,” he writes, ‘“‘ differ so much
in habits and appearance, there are no structural differences, and the
young are indistinguishable.”
I may say here that in my account of the life-history of the sea-trout
which follows I shall have occasion from time to time to comment on
the main theory that in the British Isles there is only one species of
trout, and I hope that such observations as I am able to make will help
the reader to decide for himself how far the theory is in accordance
with fact. Without doing so in so many words, in connection with every
point discussed, I will, I trust with perfect fairness and apparent
treason, substantiate the theory where in my opinion it is capable of
being substantiated, and refute it with equal candour if I find it
necessary to refute it. But, in the meantime, standing thus as I have
quoted it, the theory of which Mr. Regan has been the most recent
authoritative exponent is so striking in itself that it will be interesting
to consider the views by which he supports it.
As a good systematist, Mr. Regan gives us first a scheme of classifi-
cation of our British freshwater fishes, and those who prefer order and
method in their approach to any subject will naturally wish it to be
outlined, however briefly. The reader who is deeply interested in such
18 THE SEA-TROUT
matters may compare with it, say, the scheme of classification which the
late Mr. H. Cholmondeley-Pennell, in 1863, adopted and set forth in
his well-known book, “ The Angler-Naturalist.’ I hope I do not
misrepresent Mr. Regan’s scheme, but I gather that of the great number
of British vertebrate fresh-water animals known as fishes there are two
classes, with the first of which, M/arsipobranchii, having one order, and
one family (represented by the lampreys), we have nothing to do. Of
the other class, Pisces, there are two sub-classes, and with the first of
these, Chondrostei, which has one order, and one family (represented
by the sturgeons), we have also nothing to do. The other sub-class,
Teleostei, is our concern. In this sub-class Mr. Regan predicates
seven orders, each of which comprises one or more families; but it 1s
only the first of these orders, /sospondyli, which interests us here, for
in it are comprised three families, the important one for us being the
family Salmonide, of which the salmon and trout are more particularly
the representatives. Mr. Regan has courteously furnished me with the
reprint of a Paper, of which he is the author, on “ The Systematic
d
Arrangement of the Fishes of the Family Sa/monid@,” which enables
us to carry the classification a step further.
To the family of the Salmonide, then, in its universal aspect, Mr.
Regan is able to refer four genera :—(1) Salmo, Linn., which includes,
he says, “all the fish commonly known as Salmon and Trout.” The
other three, though not to the purpose, may be stated :—(2) Salvelinus,
Nilss., including the char; (3) Hacho, Giinth., including the hucho of
the Danube, etc., and (4) Bzachymystax, Giinth., including a species of
Siberian hucho. In the synopsis of the four genera given in this paper,
Mr. Regan states their general characteristics thus :—
“ Parietals not meeting in middle line. Teeth well developed
in jaws, on vomer and palatines, and in a double series on tongue.
Scales small, 19 or more in a transverse series from origin of dorsal
fin to lateral line. Dorsal fin short, with not more than 16 rays,
12 or fewer branched ”’;
INTRODUCTORY 19
and particularly those of the genus Salmo :—
“A double or zigzag series of teeth along shaft of vomer,
sometimes deciduous in the adult.”
It is interesting to note in passing that from a study of their skeletons,
and in particular their skulls, Mr. Regan finds that no doubt is left that
the Pacific species of salmon and trout “‘ form a perfectly natural group
that differs in several characters from the Salmon and Trout of the
Atlantic.”
In restricting himself in his book, however, to the British members
of the family Salmonid@, Mr. Regan again distinguishes four, but a
different four, genera, namely: (1) Salmo (Salmon and Trout), (2)
Salvelinus (Char), (3) Coregonus (Whitefish), and (4) Thymallus
(Grayling), a fact which reminds one of the morphologist’s caution in
viewing systems of classification, and the special features of Salmo and
Salvelinus are given thus :—
“Mouth rather large, the maxillary extending at least to below
the middle of the eye; teeth well developed; scales small or
moderate; dorsal fin with 10 to 16 rays”;
and particularly those of the genus Salmo :—
“No depression behind the head of the vomer, and a double
or zigzag series of teeth present on the shaft of that bone, at least
in the young.”
It only remains to complete this classification by noting that the genus
Salmo embraces two species, and two species only, namely, (1) Salmo
salar, which is, of course, the salmon; and (2) Salmo trutta, which is the
trout—be that what it may, sea-trout or trout, one or other, or both.
Thus we arrive at the basis of the theory that there is only one species
of trout in these islands.
i: Except in so far as many, up to this point, may, like myself, have
been treading on somewhat unfamiliar ground, I see no room for
criticism, and we have certainly arrived at a bare systematic conception,
if not of the fish itself, at least of the limits within which further
particulars may be sought for, and these we may now proceed to seek.
20 THE SEA-TROUT
I have indicated that the mgvatory habits of certain fish have been
taken by lawyers as the test of their nature, and, further, that angling
writers, in their writings, have at least been strongly influenced by these
habits. Indeed, no more strikingly marked feature of distinction would
probably occur to a superficial observer. But Mr. Regan, discussing
c
the trout, tells us that there “is sufficient evidence that the migratory
and non-migratory fish are not distinct species, nor even races.” It is
necessary to examine the evidence which Mr. Regan adduces in support
of this assertion; and I think the reader will agree that the evidence, if
not conclusive, is at any rate extremely suggestive and interesting. I
shall take the various points in their order.
I.—THERE ARE NO STRUCTURAL DIFFERENCES BETWEEN SEA-TROUT
AND TROUT.
I cannot believe that any investigator would commit himself to so
sweeping a statement as this unless he had, from exact study of many
specimens, fully convinced himself of the fact. I am not competent
to dispute Mr. Regan’s accuracy of observation, and, indeed, would
prefer to credit its correctness rather than attach weight to the numerous
alleged points of difference which other observers have professed to
discover in specimens submitted to them.
Apart from superficial colouring (which is of course a structural
difference though on a microscopical scale), differences in the form of
the gill-covers, differences in the arrangement of the teeth, and
differences in the shape and size of the fins, have all been claimed at
different times as constituting typical and characteristic distinctions.
But even admitting these and similar points of difference to occur in
certain specimens with more or less frequency and constancy, it seems
to me possible to attach more weight to them than they deserve in the
discussion as to whether migratory and non-migratory habits are
conclusive as to species—or even race. Until a marked structural
difference, constantly occurring, is proved to exist, I think we must
INTRODUCTORY 21
agree that there are no known structural differences sufficiently
pronounced to warrant the placing of sea-trout and trout in distinct
categories.
2.—TueE Younc oF SEA-TROUT AND Trout ARE INDISTINGUISHABLE.
Mr. Regan does not here clearly indicate what he means by “ the
young,” whether eggs, alevins, fry, parr, or smolts, or all or any of
them; and if he means indistinguishable in structure I must repeat that
I am not in a position to question his accuracy. But if he means in
other and more superficial respects—which I assume he does not mean—
his statement is too sweeping. I shall later, when discussing each
stage in the life-history of the sea-trout, indicate various points wherein
I have observed apparently characteristic differences between the
young, in all stages, of migratory sea-trout and, presumably, ‘non-
migratory trout. Indeed, as I shall show later, it is rather in the mature
state that sea-trout and trout tend to become “ indistinguishable.” But
at any rate, as to the young, failing proved differences in structure, I
would not be disposed to accept variable superficial differences as
evidence of distinction of species, or race, as between migratory and
non-migratory trout. Here again, therefore, we may accept Mr. Regan
as being substantially sound in his views.
3.—SEA-TROUT, IF PREVENTED FROM GOING TO THE SEA, witt LIvE AND
BREED IN Fresu WATER.
This is very important. It is, of course, the normal habit of sea-
trout to live at least temporarily in fresh water and to breed there. It
would, in the present inquiry, be necessary to show either (r) that male
and female smolts of sea-trout, having been prevented from migrating
to the sea, had been so far contented with their fresh-water environment
as to thrive there and in due season propagate their species, or (2) that
mature fish, having ascended to fresh water and there spawned, had
heen prevented from returning to the sea and had thereafter spawned
Cc
22 THE SEA-TROUT
in the next succeeding spawning season, or at least some subsequent
spawning season, and (3) that the eggs in both instances had developed
into healthy fry which had thereafter thriven.
Mr. Regan, however, only states in his book, that the fact “ has been
shown experimentally,” the precise nature and results of the experiment,
or experiments, not being more definitely indicated. But he has kindly
informed me that the experiments he had in view were those mentioned
by Dr. Day in his book on the “ British and Irish Salmonid@,” already
referred to. In a footnote to page 146 of that work, I find it stated
that :—‘‘ In November, 1886, Sir James Maitland, at Howietoun, laid
down some eggs of the sea-trout in order to follow out their life-history
if kept in fresh-water ponds; 350 hatched in 1887.” Dr. Day’s book
was published in 1887, and I am not aware whether he traced the career
of these sea-trout in any subsequent publication; but, on my making
inquiry On my own account as to the result of these experiments, Mr.
John Thomson, who has been so long connected with the hatchery work
carried on at Howietoun, courteously sent me the following very
interesting statement :—
“My notes on this experiment and recollections of it are shortly as
follows :—The.parents were caught in a tributary of the River Forth,
brought to Howietoun and spawned on November 23, 1886. There
were about 450 ova laid down to hatch of which some 350 hatched out
successfully in February, 1887, and the fry (some 250) were shifted
from the hatching-house to one of our ponds in June of the same year
and there fed the same as other fry. The young fish were again
shifted into a larger pond in June, 1888, when the average size was
found to be about 3 inches.
“In August, 1889, some specimen fish, about 6 inches in length,
were taken from the pond by Dr. Day for examination and comparison
with common trout, S. favio, and I remember we were all agreed that
it was impossible to distinguish them by the eye from S. fazio. In
INTRODUCTORY
to
Ww
April, 1890, the fish were again moved to another pond, and I spawned
some of the females in November of the same year, crossing the ova
with milt from S. devenensis and S. fontinalis. A few fry of the former
were hatched out and reared but were afterwards mixed with other fry.
“The remainder of the parent sea-trout were afterwards, | think,
turned out into a reservoir when about five years old. They never
attained to any great size.”
The facts relevant to the discussion may, I think, be stated thus :-—
(1) That the parr of the original sea-trout lived in fresh-water
quarters for four years, namely, during 1887, 1888, 1889 and 1890,
without apparent discomfort;
(2) That in their third year neither an expert ichthyologist like Dr.
Day, nor a practical hatchery manager like Mr. Thomson, could
distinguish them from common trout;
(3) That in their fourth year the females developed ova capable of
being fertilised by male trout (S. /evenensis) but not by male char (S.
fontinalis) ;
(4) That the resulting “cross” breed was “ mixed with other fry”
as apparently being to all intents and purposes ordinary trout fry; and
(5) That the parent true sea-trout brood lived on beyond 1890 in
a reservoir although they “ never attained to any great size.”
In view of these facts, which I give thus fully because they have
an important bearing upon certain passages which will follow in my
life-history of the sea-trout, it would be difficult to dispute the proposi-
tion that “sea-trout, if prevented from going to the sea, will live and
breed in fresh water.”
I may add that I do not believe that Loch Leven trout are
descendants of “ land-locked ” sea-trout, and therefore a peculiar breed
of trout. In my view they are simply trout which enjoy a specially
fayourable environment, and their origin need be in no way different
24 THE SEA-TROUT
from the origin of other trout which people our waters. Therefore
the main inquiry has no occasion to turn aside to consider their case in
detail, although any contention that they are descendants of “ land-
locked” fish, if capable of proof, would go far to substantiate this
branch of Mr. Regan’s argument.
4.—CONVERSELY, TROUT EXPORTED TO NEw ZEALAND HAVE FOUND
THEIR WAY TO THE SEA AND HAVE GIVEN RISE TO AN ANADROMOUS
Race,
The term “ anadromous” means “ migratory,” but in the restricted
sense of migration upwards from the sea for the purpose of spawning—
as in the case of the salmon and sea-trout—just as “ katadromous ”
refers to migration downwards to the sea for the purpose of spawning—
as in the case of the eel. Mr. Regan is here on safe ground for the
case of the New Zealand fish is now common knowledge, mainly
through the instrumentality of Mr. Calderwood’s admirable book,
* The Life of the Salmon,” which was published in 1907. There he
tells us that “ Brown trout taken from Dorsetshire to New Zealand
quickly acquired a migratory habit and became large silvery fish,
inhabiting the sea for the most part, and ascending rivers to spawn.”
5.—EsTuaRINE TROUT ARE OFTEN INTERMEDIATE IN APPEARANCE AND
HABITS BETWEEN THE MiIGRATORY AND NON-MIGRATORY FIsH.
I have no great belief in this so-called “estuarine” trout, and I
am surprised to find Mr. Regan thus countenancing the idea of its
existence. He seems in this to follow Mr. Calderwood who, in the
work already cited, states :—“In localities such as Orkney and Shetland
and the Outer Hebrides we have the established tidal variety which
has been called S. orcadensis; and in the West of Ireland we have S.
estuarius, the so-called slob trout. / do not agree that there is any
specific distinction, any more than I agree that a ferox is not a brown
trout.” The italics, which are mine, emphasise what I think is Mr.
INTRODUCTORY
to
e
eas
Calderwood’s real view, namely, that the estuarine trout is a variety,
but not a species, of trout. He had just written, however, in adverting
to the estuarial habits of the sea-trout :—‘‘ Many common brown trout
are to be found under precisely similar conditions and feeding on a
purely marine diet; in the estuary of the Tay near the Tay Bridge, I
have found large silvery common brown trout with freshly swallowed
herring as well as much digested herring in their stomachs and intes-
tines.” It is not very clear whether Mr. Calderwood means to
‘
distinguish these “ common brown trout” from “ the established tidal
variety.”
The late Mr. Hamish Stuart, in his well-known book on ‘ Lochs
and Loch Fishing,” wrote much that was suggestive, and something
that has a bearing on this discussion. In the Howmore River in South
Uist he states that he distinguished the sea-trout and the bull trout.
* The very first cast I ever made in the river,” he wrote, “ I rose a large
bull trout, with my second cast I rose, hooked, and lost an eriox of
perhaps 12 lbs.” In the river he also found, he says, “a few small
yellow trout, of a particularly pretty colour, fine flavour, and shape,
peculiar to the river.” Then he adds, ‘“ There are two other kinds of
fish in this river, namely, the estuary, slob or tidal trout, and a remark-
ably beautiful hybrid—apparently either between the said tidal trout
and the sea-trout, or between the latter and the common trout... .
It reaches occasionally the weight of 1 1b., but is rarely over, and
generally under, } 1b. This fish does not appear to spawn, but is
numerous.” And of the Strome Dearg, the connecting semi-artificial,
semi-tidal link between Loch Hallan, also in South Uist, and the sea,
he writes :—“ Besides sea-trout this salt-water river is frequented by a
species of yellow trout with extremely large spots,” which he attributes
generally “to the slob or tidal variety.” He concludes :—‘‘As the
Strome is frequented by yellow trout originally from Loch Hallan and
other lochs, the peculiarities of its environment have practically
26 THE SEA-TROUT
succeeded in producing a new species—or at least a variety of the trout
of the loch referred to. This new species or variety attains a recorded
size of at least 6 lbs., or three times the size which the trout, from which
it is a specific environment-born off-shoot, is known to attain.” It is
interesting and suggestive to consider all this alleged wealth of variety
of fish life in these small, out-of-the-way corners of the Hebrides in
the light of the theory promulgated by Mr. Regan.
This subject is worth pursuing a little further. That praiseworthy
publication, “The Sportsman’s and Tourist’s Guide,” describes the
Dhu Loch, near Inveraray, as “an excellent loch for sea-trout and
yellow trout.” Moreover, it says, ““ The Dhu Loch is a tidal water,
within a few hundred yards of the sea, and an unusual species of plump
tidal trout are occasionally to be found in it,” thus recognising sea-trout,
yellow trout and tidal trout. It is common knowledge, again, that trout,
as well as sea-trout, are frequently caught in the salt-water basin of
Loch Etive; and, indeed, I imagine that there are few districts in
Scotland, when opportunities of observation are possible, where common
brown trout will not be found in estuarial waters.
Now I have never seen it anywhere recorded that these alleged
species, or varieties, of trout which have in so many places been
detected by angler-naturalists and angling writers in salt and brackish
water, have been seen on the spawning redds or that the progeny of
them, distinguished by features characteristic of the parents, have been
observed in the spawning streams. I for one am inclined to believe
that such progeny will never be discovered. The facts to my mind
rather point to this, that one and all of these alleged tidal trout are
individual specimens of the ordinary fresh-water trout of the district,
which, descending the river until they have come within tidal influence,
have gradually become acclimatised to a salt-water environment.
Moreover, I believe that the greatest number of these trout will be
found in those localities such as the Orkneys and Shetland, and some
INTRODUCTORY
to
™N
of the Hebrides, where the streams are so trifling, or elsewhere in the
Highlands, where the streams are such barren, rocky, mountain torrents,
that there is neither security nor sustenance for trout in them.
It is rather corroborative of this view than inimical to it, that trout
are to be found, as Mr. Calderwood found them, in the estuary of the
greatriver Tay. The fact merely proves that even in such rich streams
some trout will so descend to the sea. In the estuary of the Clyde,
below the junction of the river Leven with it, such trout have frequently
been captured in the salmon nets. I take it that these fish had
gradually dropped down stream from the upper reaches of the Leven,
or perhaps even from Loch Lomond, because I am informed by a local
boatman, of whose capabilities of judging I have personal knowledge,
that on one occasion he caught with rod and line in the loch a trout
which had tide-lice on it. I give this incident for what it may be
considered worth, for it suggests not only that a trout had descended
to salt water but that it had returned to fresh water again, as such trout
must do if they propose to spawn.
If it is objected that in all this I am only stating grounds for the
belief that estuarine trout are really ordinary non-migratory trout that
have become more or less migratory, it may be recalled that this is the
particular point in Mr. Regan’s thesis that is at the moment under
discussion. But to revert now to the main argument, I shall simply
confine myself to the suggestion that, having regard to the foregoing
facts, the offspring of these enterprising trout, given favourable circum-
stances, may easily become indistinguishable from migratory sea-trout,
though this savours not a little of taking matters for granted. The
suggestion, however, leads naturally to the discussion of Mr. Regan’s
sixth, and perhaps most interesting, proposition, which I shall now
submit.
28 THE SEA-TROUT
6.—THERE IS GOOD REASON TO BELIEVE THAT IN NATURE THE RANKS OF
THE SEA-TROUT ARE REINFORCED BY THE OFF-SPRING OF THE
RIVER-TROUT AND VICE-VERSA.
The previous passage suggests how the ranks of the sea-trout may
be reinforced “ by the off-spring of the River-trout,” and, subsequently,
throughout my account of the life-history of the fish, it will be seen how
at nearly every stage the influence of the sea-trout—not to put it any
more strongly—dominates the trout.
It remains now to examine how the ranks of the trout may be
reinforced by the off-spring of the sea-trout, which is a deeper and more
wide-reaching question. It is so deep, indeed, that I do not think its
depths will soon be plumbed, but the theories to which the question has
given rise have occasioned much instructive speculation.
Mr. Regan, then, asks us to consider the distribution of the trout.
“There are,” he writes, “no true fresh-water fishes—Roach, Perch,
etc.,—in the Hebrides, Orkneys, or Shetlands; yet in these islands
every river and loch is full of Brown Trout, which is only to be
explained by the supposition that the latter have been derived from the
Sea-trout, which have lost their migratory instinct in different places
and at different times.” In other words, Mr. Regan’s view is that the
trout is a sea-trout; not that the sea-trout is a trout. He ascribes to
the trout, in fact, a marine origin.
Now it seems fairly certain that in order to arrive at the origin of
any species we must travel back a considerable distance, possibly
through aeons of evolution, into the dim and distant past. To establish
his point, however, Mr. Regan does not go so far back as that, but,
taking it for granted that trout then existed in the shape we know them
to-day, he carries us back to the Glacial Epoch.
“Tt would be out of place here,” he writes, “ to enter into the causes
of this climatic change, but it seems clear that at a comparatively recent
date the whole of Northern Europe, including our islands except the
INTRODUCTORY 29
South of England, became covered with ice; then a submergence of
the land perhaps brought the sea right to the edge of the ice-sheets, so
that any true fresh-water fishes which may have been in our rivers
would have perished. At the end of this cold period, a gradual
elevation of the land took place, culminating in the union of our islands
with each other and with the Continent, and then a subsidence followed
which gave to our islands approximately their present outline.
“Speculations as to the exact time occupied by these changes
are futile, but it is probable that the whole duration of the Tertiary
Period should be expressed in tens of millions of years and that of the
Glacial Epoch in tens, or at most hundreds of thousands. Possibly
about 100,000 years may have elapsed since the end of the Glacial
Epoch, and our final separation from the Continent was of still more
recent date.
‘As soon as the ice-sheet had begun to disappear char must have-
commenced running up into the lakes which were formed, and as the
elevation of the land proceeded new lakes appeared and in turn became
inhabited by Char. All this time the climate was gradually getting
warmer, and the southern limit of anadromous Char was receding
northward; then the sea-trout reached our islands, and these in turn
began to form fluviatile colonies in our lakes and rivers.”’
It is clear from the foregoing that Mr. Regan asks us to believe,
and to me at least his reasoning is as plausible as it is interesting, that
all the trout which to-day throng our waters, trout great and small,
migratory and non-migratory, are the immediate lineal descendants of
certain enterprising trout which came up out of the prehistoric Atlantic
Ocean into our rivers to spawn when the great ice-sheet began to recede
before more genial climatic conditions. It seems to me, however,
relevant to observe that although Mr. Regan thus reasonably credits
our British trout with a marine origin, he has not thereby settled the
question of the primary and original environment of the invaders, and
30 THE SEA-TROUT
it still remains to be considered whether the trout was originally a fresh-
water or a Salt-water species.
The question which exercised the minds of the Sophists, namely,
whether eggs or birds were created first, seems to have been particularly
fatuous in respect that the answer—whatever it might be—held out
very little promise of practical utility. If our present question is a
degree less futile than the classical conundrum, it can, for us at the
moment, only be useful for such light as the inquiry may help to shed
upon the question whether the migratory sea-trout is distinct from, or
is of the same species, and even race, as the non-migratory trout. The
inquiry can hardly hope to solve the deeper problem raised by the
mysterious habits of the fish in migrating at all. Yet that problem is a
most attractive one, and Mr. Regan supports the view which he takes
of it thus :—
“The Salmonide,’ he writes, “are found in the Arctic and
temperate regions of the Northern Hemisphere, and may be regarded
as marine fishes which are establishing themselves in fresh water” ;
and again, “ Some writers look upon the Salmon and its relatives as
true fresh-water fishes which have acquired the habit of going to the
sea for food, and which return to their original home to spawn. Against
this it may be urged that whereas many marine fishes take to fresh water,
the reverse is a rare phenomenon.” Now I do not propose to quarrel
with Mr. Regan over his view of. this problem, which to me seems
¢
immaterial, but I suggest that the word “rare” implies that the
phenomenon is not impossible, and it may be fairly argued, or arguable,
that the facts equally point to fresh water as the original habitat of the
Salmonide. In particular, the fact that in salt water the spawn of
salmon and trout cannot come to fruition seems to me to be pertinent,
for if one casts back in imagination to the time before the fish had
acquired its migratory habit, it is easier to suppose that the eggs then
too hatched in fresh water than that, from being able only to hatch in
INTRODUCTORY 31
the sea, they gradually reached a stage when sea-water became a deadly
poison to them. Whereas, if we imagine the original habitat to have
been fresh water it is plain that the breed might easily be perpetuated
throughout the ages, as in fact it has been.
It is unnecessary to prove why some fish adopted the migratory
habit and not others of the same species, but at the same time it 1s
plausible to suppose that pursuit of food was the determining factor.
lt is at any rate plain, if we suppose the trout to have had a fresh-water
origin elsewhere than in Britain, and to have become migratory, that the
migratory habit, having once become established, accounts for the
trout’s wide distribution throughout the Arctic and temperate regions.
It was a simple matter then for it to spread from the river of its birth
to any other river it might chance upon in the course of its wanderings
in the sea. Indeed, in the passage quoted, Mr. Regan very convinc-
ingly shows that this is the method by which our British waters received
their stock of trout. From the sea they came, doubtless, to our shores,
but it is not unreasonable to assume that they came from rivers in other
parts of the Continent of Europe, or rivers elsewhere, devoid of ice,
where they had their origin. It seems to me, then, that equally with the
trout being derived from a marine sea-trout, the facts point to the sea-
trout being a derivative of a fresh-water trout; or, in other words, the
proposition may as fairly be stated that the migratory sea-trout is a
trout, as that the trout is a non-migratory sea-trout.
I understand that this is a question upon which science itself has
come to no common agreement, although I am unaware of the precise
weight of authority which may be ranged on either side. Mr. George
A. Boulenger, F.R.S., the learned and courteous successor of Dr.
Giinther at the British Museum, thinks alike with Mr. Regan, who is
his colleague and assistant. Dr. A. Noel Paton, as quoted by Mr.
Calderwood, considers that “ the Salmonid@ are originally fresh-water
fish.” Mr. Calderwood is himself for a marine origin, and I cannot do
to
THE SEA-TROUT
>
better than refer the reader to his clear discussion of the subject in the
chapter on “ Grilse ” in his book “ The Life of the Salmon.” As Mr.
Calderwood’s point of view, in any debatable question, is ever that of
the open-minded inquirer after truth, all the more weight may be
allowed to his conclusion :—‘ In my view the prevailing characteristics
of the group of fishes to which the Salmon belongs are those of marine
fish of plastic nature, capable of much local variation both of appearance
and habit, many of which enter fresh water freely.’ Dr. Day was of
a like opinion.
Anglers would appear to prefer for the Salmonid@ a fresh-water
origin. Sir Herbert Maxwell discusses the question in his book on
“Salmon and Sea-Trout” (in “ The Angler’s Library” series), and
arrives at a fresh-water conclusion; while the late Mr. Hamish Stuart
in another connection recognised a fact which “ seems to assign to the
common ancestor of the Salmonid@ a fresh-water origin.”
.
Presumably discovery made of the “common ancestor” and its
habitat will finally settle this matter, but that discovery is not yet.
That science is not without hopes may be inferred from this extract
from a letter which Dr. A. Smith Woodward, Geologist in the British
Museum, has written to my friend Dr. J. W. Gregory, Professor of
Geology in the University of Glasgow, the purport of which has been
kindly placed at my disposal. Dr. Smith Woodward writes :—‘ Prac-
tically nothing is known of the ancestry of the Salmonide@. A few
typical remains are found in the fresh water Tertiaries of the Continent
(see references in Catal, Foss, Fishes, B.M., pt. iv), but they merely
show how old is the family. Of all the modern fresh-water fishes we
lack ancestors connecting them with the marine forms. We shall find
them some day.” I have no authority to rank Dr. Smith Woodward
either as an exponent or as an opponent of the “ marine origin ” theory,
but his letter suggests that something at least may be said for those
who take the view of a fresh-water origin.
INTRODUCTORY 33
As I have stated, the dispute, or rather discussion, is not very
material to the purpose, the more so if we accept Mr. Regan’s account
of the manner in which our streams became peopled with trout, namely,
through their having been invaded from the sea by trout which were
de facto migratory, some of which gradually lost the extreme migratory
habit. Such acquiescence in Mr. Regan’s account would also involve
acquiescence in the theory “that the migratory and non-migratory fish
are not distinct species, nor even races,” unless it be maintained that the
invaders consisted of more than one species or race, or had since
diverged into such, in my opinion an improbable development.
The reader who has been patient enough to follow the argument is
as capable as I am of forming a judgment upon the problems which I
have thus presented to him. But probably the first observation which
occurs to him will be that, even accepting Mr. Regan’s explanation of
the original stocking of our waters with trout, he would like to have
some clear proof that the interchange of habits and environment which
Mr. Regan predicates actually takes place in our waters to-day. It is
no reflection upon the general soundness of Mr. Regan’s position that
he cannot furnish such proof as would convince a British jury, but I
think the theory which he expounds—for one hardly ventures to accept
it as fact—is worthy of the most respectful consideration. There can
be no doubt of this at least that the migratory and non-migratory trout
are sometimes so alike that cases often occur where it is practically
impossible to say definitely whether a particular fish is a sea-trout or a
trout. In such cases the fish quite conceivably may be a trout which
has taken on the colour of a salt-water environment or a sea-trout which
has found congenial quarters in fresh-water. |
The late Mr. H. Cholmondeley-Pennell, who, following Yarrell
and others, made the very most of “ distinctions” in “ The Angler-
Naturalist,’ said :—‘‘ The difference in colour between the two fish—the
one being silver and the other golden—is wszally too obvious to admit
34 THE SEA-TROUT
of doubt; but it occasionally happens, especially when confined for a
long time in a loch, that the Sea-trout acquires a colour not altogether
unlike that of the Yellow Trout.”
In “The Fishing Gazette” of September 6th, 1913, to take a
specific instance, I observed that a party of anglers fishing from a
Donegal hotel had been divided in opinion on this question with respect
to one of the fish caught. With commendable enterprise one of the
party sent some of the scales of the fish for identification to Mr. R. B.
Marston, who passed them on to Mr. J. Arthur Hutton, for his
opinion. Mr. Hutton, disclaiming any expert knowledge of trout
scales, “rather hesitated to express a decided opinion,” but, arguing
the case intelligently on the facts, came to this conclusion :—“ I am
inclined to think that your correspondent’s fish was a sea-trout which
had been a long time in fresh water.” I may add that several similar
cases of dubiety regarding trout from other waters were discussed in
“The Field” in the autumn of 1913.
With regard to trout assuming the superficial appearance of sea-
trout, we have already noted that Mr. Calderwood found in the estuary
of the Tay “large silvery common brown trout.”
It is possible that the reader may seek to find some evidence of the
theory here discussed in the waters with which he as an angler is
familiar. I have myself sought for some such evidence, and it occurs
to me that the theory may have some bearing upon the diversity of
appearance which marks the trout in the area drained by the river
Clyde, an area which, according to the Bathymetrical Survey, includes
the Loch Lomond and Leven drainage area. The diversity is strongly
marked between the trout which inhabit the river Clyde above the Falls
of Clyde and those which one finds in the Loch Lomond basin which
drains into the Clyde estuary.
Accepting Mr. Regan’s proposition that our British trout, non-
existent during the Ice Age, originally ascended from the sea, it is easy
INTRODUCTORY 35
to explain the presence of trout above the Falls of Clyde by assuming
that the stock ascended the river before subsequent elevation of the
land and denudation combined slowly formed the now impassable
barriers of Stonebyres, Corra Linn and Bonington Falls, and so isolated
the trout of the upper waters. The trout above these falls are
uniformly yellow, beautifully marked with brown spots and are, whether
old or young, as unlike any fresh-run, silvery sea-trout in general
superficial appearance as it is possible for any trout to be. On the other
hand, for thousands of years migratory trout must have had free access
to the Loch Lomond basin, which, in a comparatively recent geological
period, has been a sea-loch, and is now only 26.9 feet above mean
sea-level, being connected by the short even-flowing Leven with the
Clyde estuary. But in contrast with the trout of the upper reaches of
the Clyde, hardly any trout are so likely to be mistaken by the angler
for sea-trout as are the trout of Loch Lomond. Indeed, in the later
months, when sea-trout which have been in the loch for some weeks
have lost their silvery sheen, they approximate in appearance so closely
to the native non-migratory trout that it is difficult, even for experienced
anglers, to distinguish between them. In spring it is a practical
impossibility to distinguish young trout of two or three years old from
sea-trout parr of equal age. I have been able to discover no features,
beyond some small presumptive differences of colouration, by which
one kind of fish can be distinguished from the other, and boatmen with
life-long experience will seldom undertake to say with any assurance
which is which.
The inference would seem to be legitimate that once the barrier on
the Clyde reached a certain height, and trout, though many might
descend, were barred from returning, the stock in the upper waters began
to revert to the original type of fresh-water trout, or at least to develop
along lines of colouration adapted to its isolated fresh-water environ-
ment, and has for centuries been so developing; while, as regards the
36 THE SEA-TROUT
Loch Lomond trout, there having been an uninterrupted coming and
going for centuries from and to the sea, the stock has retained
all the marked outward characteristics of the migratory fish. But so
far as structure and outline are concerned the isolated trout from above
the Falls of Clyde differ in no respect that I can discover from the
trout or sea-trout below the Falls.
It may of course be objected that the strain of each distinct species
in the loch (assuming for the sake of argument that the migratory and
non-migratory fish are distinct species) has been affected by inter-
breeding over long ages, and that each has in consequence approached
closely in its general characteristics to the other without wholly losing
its proper individuality. But as to this matter of the crossing of breeds,
in his great work “On the Origin of Species,’ Darwin makes this
pregnant observation :—“ The possibility of making distinct races by
crossing has been greatly exaggerated . . . . Certainly, a breed inter-
mediate between ¢wo very distinct breeds could not be got without
extreme care and long-continued selection; nor can I find a single case
on record of a permanent race having been thus formed.” He also
states :—‘‘ It is difficult, perhaps impossible, to bring forward one case
of the hybrid offspring of two animals clearly distinct being themselves
perfectly fertile.’ We all know, moreover, his views as to the origin
of our various breeds of pigeon :—“Great as the differences are between
the breeds of pigeons, I am fully convinced that the common opinion
of naturalists is correct, namely, that all have descended from the rock-
pigeon (Columba livia),’ a passage which it is interesting to contrast
with another written two hundred years earlier, to this effect :—“And you
are to note, that there are several kinds of trouts: but these several
kinds are not considered but by very few men; for they go under the
general name of trouts: just as pigeons do, in most places.” The
reader may be referred, for the rest of the passage, to Izaak Walton,
his book. On the whole, it would seem to be more in accordance with
INTRODUCTOR x 37
reason to adopt at once Mr. Regan’s view that there is only one species
of trout involved of which some unknown proportion in the Loch
Lomond basin is migratory in its instincts and some unknown propor-
tion non-migratory at any given point of time.
That the native trout of Loch Lomond descend to the Clyde estuary
is proved by the fact already mentioned that they are sometimes caught
in the estuary salmon nets, and I gave one instance of a trout taken in
the loch which, having sea-lice on it, may be presumed to have so
descended and then returned. If the objection is made that the mere
fact of the migratory and non-migratory trout being distinguishable,
even with difficulty, is an argument for, and not against, there being
two distinct species or races involved, it may be answered that it has
never been pretended that trout habituated to a salt-water environment
do not differ greatly from trout habituated to a fresh-water environment
in their exterior aspect.
But it may be said that, after all, it is not enough to prove that one
sea-trout ceased to migrate and that one trout acquired the habit of
going to the sea; what has to be proved is that a particular non-
migratory trout at some time adopted the migratory habit which there-
after persisted in itself and its descendants, or, on the other hand, that
any particular migratory trout ceased to migrate and thenceforth with
its descendants remained satisfied with its fresh-water environment.
Such proof would no doubt be final, but I do not think that Mr. Regan’s
six points of evidence are conclusive. It seems to me, however, that
taken together they give grounds for a strong presumption that there
is no real distinction in structure or other material respect between
certain of the migratory and all of the non-migratory trout, and that some
of the offspring of any pair of either class may at any time develop the
migratory instinct in an extreme degree while the rest may remain
content with their environment. In the chapters which follow I think
D
38 THE SEA-TROUT
[ shall be able to submit facts which at least suggest that such variation
of habit is by no means impossible or even improbable.
This introductory chapter has already grown to an unconscionable
length, but there is one other vexed question upon which some general
comment must be made, and I crave the reader’s patience.
Mr. Regan’s dismissal of the migratory habit as an indication of
species, or even race, leads him to the conclusion that the trout is a
sea-trout. I have for my part suggested that certain sea-trout may just
as probably be trout, but whichever view is taken will not affect the
discussion which follows.
Mr. Regan states :—‘‘ Two races of Sea-trout may be recognised,
although in many cases it is impossible to say to which race an individual
may belong unless one knows beforehand where it comes from,” about
which statement there is a curiously Hibernian ring. “ However,” he
continues, “the Sewen (S. cambricus) of Wales, Ireland, and our
Western coasts often differ from the Sea-trout (S. a/bas) of the east
coast, in having a longer head, a larger mouth with stronger jaws, the
suboperculum projecting backwards beyond the operculum, and the
fins somewhat larger, the lobes of the caudal especially being more
produced. When typical examples of the two races, of the same size
and sex, are compared, these differences may be seen, but they are
slight, and not always apparent.”” And he elsewhere states :—‘‘ Some
trout appear to go farther out to sea and to grow more quickly than
others, and this is especially the case with the trout of certain rivers
such as the Coquet and the Tweed; in the latter, sea-trout more than
four feet in length and weighing nearly 50 Ibs. have been captured.”
Now Mr. Regan had no wish to detract from the weight of his main
thesis, “ that there is only one species of trout ” in the British Isles, but
he was puzzled to account for the “bull trout,” and so he seems to
ascribe it to a barely distinguishable race. Other systematists, in much
the same way but more elaborately, have drawn a distinction between
INTRODUCTORY 39
the broad type SS. ¢vw//a and a southern variety, S. ¢vutta (eriox) for the
migratory fish; and between S. fario (Gaimardi) and a southern variety,
S. fario (Ausonii), for the non-migratory trout. But the “bull trout”
is not yet, I think, thereby explained.
There is no reason to believe, nor is there any evidence, that since
the prehistoric glacial epoch changes have occurred in the types of
migratory trout which became localised in British waters. As the types
were then we may suppose them to be to-day. While it is no doubt
true that there existed great river basins in the European plateau,
whereof our islands are a fragment, and of which ancient rivers our
greatest streams are, it is believed, but the surviving upper tributaries,
or at least extensions of these tributaries, the plateau had a compara-
tively uniform sea-board on which the ancient rivers debouched. One
need not therefore ascribe to each ancient river specific races of trout.
But I think a fair analogy may be drawn between our modern trout and
those prehistoric fish in this respect. Just as to-day there is great
external difference between the trout of one locality and the trout of
another locality, to such a degree that one can hardly allege the trout
of one river area to be exactly like the trout of any other river area, so
the trout of the ancient Rhine which flowed northwards over land which
now forms the bed of the North Sea, the trout of the “ Irish Channel
River” which flowed southwards, and the trout of the “ English
Channel River” which flowed westwards. may quite well have had
distinctive external characteristics. It is not a very extravagant
speculation that some of the local characteristics of these prehistoric
fish may survive to this day, and that, for instance, the trout of the
Tweed and the Coquet, the Haddington Tyne and the Northumberland
Tyne, the Forth and the Aln, all, we may suppose, connected with the
ancient Rhine system, may still possess them.
But setting speculation aside for the moment, the best scientific
opinion does not appear to concede the “ bull trout’s” claim to be a
40 THE SEA-TROUT
distinct species of trout. Dr. Giinther, the great ichthyologist of the
British Museum—I quote the Rev. W. Houghton, who published a
book on “ British Fresh-water Fishes” in 1879—wrote thus :—‘ No
distinct species is designated by this name (Bull Trout); at all events
the name is applied to different species at different localities, and by
different persons. We have received numerous examples of ‘ Bull
Trout,’ each of them of a peculiar aspect, but without any characters
by which the species could be determined. It would appear that many
examples somewhat differing in general aspect from S. ¢va/¢a are named
Bull Trout.”
In the year 1910 | had the honour of receiving a communication on
this subject from Mr. Boulenger, already quoted in a former work of
mine.! ‘I do not admit the ‘ bull trout’ even to the rank of a variety,”
he wrote, “ From a scientific view, the only one with which I have any
right to speak, the designation ‘ bull trout’ should be entirely dropped.”
Dr. Day, in his “ British and Irish Salmonide@,” had already negatived
the idea of any specific distinction. Mr. Calderwood, however, con-
cedes the distinction of being at least a “ variety ” to the bull trout. In
his book on “ The Salmon Rivers and Lochs of Scotland,” published
in 1909, he incidentally remarks :—“The sea-trout of the Tweed is now
almost exclusively the bull trout or round-tail, S. ¢vutta var. eviox, the
same fish as ascends the Coquet,” and he had already, in a paper read
before the Royal Society of Edinburgh given a more particular account
of the fish.?
A great deal of confused and somewhat confusing “ evidence ” on
this subject was listened to by the Royal Commissioners of 1902, and
is to be found scattered throughout the volume of evidence appended
to their report. It is vague and contradictory in the extreme, and the
reader who is curious enough to examine it will find some entertainment
1. ‘The Gentle Art” (1911). Chap. XVI, ‘‘ Concerning Definitions,” p. 295.
2. “The Bull Trout of the Tay and of Tweed ” by W. L. Calderwood. Reprint from Proceed-
ings of the Royal Society of Edinburgh, Session 1903-4, Vol. XXV, Part I (1904).
‘* Fresh Run from the Sea.”’
INTRODUCTORY 41
in endeavouring to reconcile the statements of the different witnesses.
Perhaps the views of anglers should be received with caution, but
1 may submit this opinion concerning Tweed “ bull trout” given me by
an angler residing on Tweedside, who is himself a close student of fish
and their habits and from whose pen any work on the Salmonid@ would
be of great value. _ “ I am afraid I am not an authority on the subject,”
he modestly wrote me, “but I hold that sea-trout, salmon-trout, and
bull-trout are one and the same fish. They vary very much in size
only—from 4 lb. to, say, 25 lbs.” So, too, Mr. John James Hardy, of
Alnwick, wrote me when I inquired if he could furnish me with a
characteristic photograph of an Aln or a Coquet “ bull trout ” :—“* The
trouble is that my opinion is that the Coquet so-called bull trout is only
an overgrown sea-trout; that there is no difference between the bull
trout and the sea-trout, and that Sa/mo eriox exists only in imagination.”
It is difficult to get over the fact, however, that as far back as the time
of Izaak Walton, the “bull trout” was a noticeable fish. “ There is
also,” said his Piscator, “in Northumberland a trout called a bull trout,
of a much greater length and bigness than any in the southern parts ”’;
and Walton’s detailed description of the “ Fordidge ” trout differs in
no essentials from a modern description of the “ bull] trout,’ of which
it may generally be said, if one be caught, that “ ¢had trout bit not for
hunger but wantonness.”’
This subject is of course one which is frequently dealt with in
current angling literature, and in the pages of “ The Fishing Gazette ”
and “ The Field” one finds constant references to the “ bul] trout.”
I have often curiously examined accounts of the fish to see if there is
any recognisable distinction between them and sea-trout, but it is not
easy to find anything in their habits that distinguishes “bull trout”
from big sea-trout, and not much more in their appearance than a more
pronounced convexity of tail.
It seems to me, therefore, admitting the characteristic of the “ round
42 THE SEA-TROUT
tail” to be a more frequent feature of eastern sea-trout than western
sea-trout, that there is no more warrant for claiming the “ bull trout ”
as a Separate species, or even race, on account of its round tail alone,
than there is for making such a claim on behalf of the tailless trout of
Loch-na-Maorachan, in Islay, or the “bull-nosed” trout of which
some reservoirs can show a high percentage. Conclusive evidence of
distinction must be sought along other lines.
I have, personally, an open mind on this really important subject,
and would be glad if it could be scientifically settled once for all. It
seems to me that careful scale examination may help to solve the
difhculty, and later in these pages I shall give more precise reasons for
so thinking. In the meantime it is enough to say that there may quite
well be a distinct race, or even species, of sea-trout which has nothing
in common with ordinary /favio, except that, like the salmon, it spawns
in very much the same way in our rivers, while at the same time there
may be another race, or species of sea-trout, in reality ordinary /fazio,
of which some individuals retain and some lose the migratory habit, its
retention or loss being at any time entirely dependent upon the nature
of the fish’s environment. There is really nothing definite known of
the matter, but I can easily imagine that naturalists might yet come
to agree that there are sufficiently distinctive points of difference to
justify the grouping of three species, namely (1) the salmon (S. sadar),
(2) the migratory true sea-trout (S. ¢vwta)—in other words, the “ bull
trout ’—and (3) the trout (S. favio), which last may be indifferently
migratory or non-migratory.
I may conclude the whole discussion with Dr. Day’s remark :—“ If,
as seems probable, we merely possess one very plastic species subject
to an almost unlimited amount of variation, that its largest race is found
in the ocean, while in order to breed it ascends streams, but usually (to
which there are many exceptions) not so far as the salmon, unless it
permanently takes up its abode in the fresh waters, we at once obtain a
Fig. 1.—A Salmon, showing normal spots of the ‘‘ maiden ’’ fish.
<<
:
_—
aj
2. A Salmon, showing spec kled appearance of a fish which has
previously spawned.
INTRODUCTORY 43
clue to the characters of the various so-called species, and relegate
these different trout to a single form, in which numerous local races are
to be found.” This is, of course, a very easy way of evading all
difficulties, but it does not dispose of them. I think the facts rather
point to such a differentiation as I have suggested above.
What purpose would the solution of such problems as I have
here considered serve? One never can tell of course in what direction
definite knowledge may ultimately prove useful. Something at least
is gained by the mere fact that the knowledge is definite. But there is
this further consideration. It has for some years been apparent that
the stock of sea-trout in once well-stocked waters has markedly
declined. Even as long ago as 1904 “ The Spectator ” feared that the
bd
fish was “ doomed to a not very remote extinction ’’; and as recently as
August 1914, there appeared in “ The Salmon and Trout Magazine ”
a suggestive article on the decline of ‘‘ White trout ” fishing in Ireland
from the pen of that well-known writer “ Corrigeen.”” Mr. Calderwood
also notes a general decline in the stock.
If there is anything at all in the theory that there is any one, or only
one, species of trout in these islands whereof at any given point of time
a certain proportion is migratory and a certain proportion non-migratory
—the migratory habit per se in either case not being an indication of
“species” or even of “race’’—then it necessarily follows that the
innumerable causes which have brought about a depletion of the stock
of trout in our once prolific rivers must be added to the many other
causes which have made for depletion in the ranks of the sea-trout.
If trout cease to exist in their pristine numbers in the main streams and
upper tributaries, whence, it may be asked, are the ranks of the sea-
trout of the estuaries to be reinforced?
It is plain that such a question as this is far-reaching in its conse-
quences, for it at once opens out the further question whether our
fresh-water fishery laws dealing with trout have not been improperly
re THE SEA-TROUT
divorced from the laws concerning the migratory fish embodied in the
salmon fishery statutes; or, one might put it alternatively, whether the
laws concerning the sea-trout are not too exclusively restricted to the
region of salmon fishery legislation. I began with an examination of
the legal view of the sea-trout and I find myself incv:tably led back
to it. The truth is that if any species of fish is to survive in these
modern days, legislation concerning it must be based upon reason and
a full understanding of the life-history and habits of the species.
I have in this introduction made free use of the views of Mr. Regan
as set forth in his excellent book. I hope I have made it clear that |
have not been influenced, in discussing these views, by any spirit of
captious criticism. It will be quite apparent that I have no qualifica-
tions to criticise them beyond those which any thoughtful reader may
possess. His book is the most recent work with which I am acquainted
which can pretend to be of any scientific authority on the subject of
British fresh-water fishes, and such observations as he makes upon the
trout have been for me a convenient text upon which to discourse more
particularly concerning the sea-trout.
To conclude—while one may very well accept the theories which
guide scientists to the systematic classification of a species, there is no
occasion to rest a description of that species on scientific classification
alone. The angler will not perhaps concern himself overmuch whether
the migratory and non-migratory trout constitute different species or
no. He will concern himself chiefly with the appearance and habits
of the sea-trout as a migratory fish so long as it retains such appearance
and habits. These are in any case sufficiently marked to warrant a
separate description being given of them. It is necessary to add that
the description which I shall endeavour to give in the following chapters
applies more particularly to the type of sea-trout which is commonly
found in our Western waters rather than to the type which, as the “ bull
trout,” has earned for itself a certain notoriety in our Eastern streams.
sere 7 A ‘
Aidt re Le tlt ee
ae | ee BOY 5 | ee, Beas
r ’ ,
Fig. 3—Two Loch Lomond Sea-trout, showing typically distinctive markings.
Fig. 4.—A male Sea-trout at spawning time.
See page 50.
—— ml
- _ =
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- - . . _ | ra op
7 * _ ie te gy
sae
a Le
S 7 ie
- rt fea ~ ie . | th
ae
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a 7
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: a at
‘ wy :
5 oe a tt i ons
seme an lawer hs sols, 1a Baht Sere
7 = =
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Fig. 5—A ‘‘ two year old’’ Trout (fario), with ‘‘ halo ’’ surrounding the
spots strongly accentuated.
See page
os
¥ 9M Fy hoe
S pats we tas ake be:
% ot + Sse SI! a
4° oe PE Seas cae aD ne
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Fig. 8—A Clyde river Trout (fario).
See tage 51.
CHAPTER II.
THE SEA-TROUT.
“There are three beauties in death,’ runs the Gaelic saying, “a
sea-trout, a blackcock and a young child.” Of these, it will be
observed, the sea-trout is ranked first, with how much of justice will be
a matter of opinion. There can be no question, however, that an
adult sea-trout of three or four pounds in weight, killed when fresh run
from the sea, is one of the most beautiful objects in nature, suggesting,
as it does, in its general outline and appearance, the perfect adaptation
of physical form and harmonious colouring to habits and environment.
In beauty of shape alone the sea-trout perhaps stands midway
between the salmon on the one hand and the trout on the other. The
neater head and more slender tail of the salmon give to that fish a grace
which the sea-trout with its more sturdy build cannot rival, and yet the
latter as far excels most trout in its suggestion of compact elegance.
In colouration, too, while emulating each of the others, the sea-trout
occupies a middle place, for, if it cannot vie with the splendour of the
blue and silver mail of the salmon, it adds to its sheen the distinction
of more characteristically varied markings. On the other hand,
although few trout can approach the silvery brightness of the fresh-run
sea-trout’s scales, many of them far out-rival it in the colour and number
and variety of their spots. On the whole it may be said that the sea-
trout successfully unites in its person the brilliance of the salmon and
the more variegated beauty of the trout.
The sea-trout, like other members of the family Salmonide, is an
inhabitant of the waters of the Arctic, sub-Arctic and Temperate zones
of the Northern Hemisphere. Its exact range it would perhaps be not
easy to define, but one might say generally that wherever there are trout
47
48 THE SEA-TROUT
found in the streams of those countries lying northwards of the Straits
of Gibraltar, sea-trout will be found in their coastal waters also. But
the success of modern experiments in the transplanting of trout ova
has been so remarkable that the range of trout has been extended to
the Antipodes, India and Africa, and in New Zealand at least a race of
migratory trout has sprung from the transplanted stock which—as
many of the facts noted in this book will suggest—is to all intents and
purposes now a race of sea-trout.
In the British Islands, it is certain, the sea-trout is indigenous to all
our coasts, and particularly it exists in enormous shoals on the western
coasts of Ireland and round all the coasts of Scotland, both of the
mainland and of the islands, wherever access can be gained from the
sea or estuary to suitable inland spawning grounds. It is reasonable
to assume that the absence from our rivers of impassable obstructions
and serious pollutions is a condition necessary to the presence of sea-
trout in them and their estuaries. How far the destruction of trout in
our streams from one cause or another will affect the stock of sea-trout
is a problem which has already been touched upon.
It is not easy to state precisely what weight a sea-trout may attain
to. The size of the fish appears to vary greatly according to the type
of waters in which it has been bred and the type of coastal waters to
which it may migrate. But the factors of feeding and spawning largely
affect the rate of growth, and I think it would be difficult to discover
any fixed rule. I have not the least doubt, too, that the problem of
weight has been confused by some of those salmon which for brevity
may be designated “ Tay bull trout,” having been recorded as sea-trout
before their identity was so well established as it is now. Further, the
fish which has more claim to be called a “bull trout” grows to a
considerable size, and it is not always easy to discover which class of
sea-trout is in question when any record of weight is cited. Mr. Regan,
discussing trout generally, refers to them as attaining a weight of 4o lb.,
pesuriie AyjvoLnewoas sjods SUIMOYS ‘jNOI]-vaG puowoy Yyoo"] wo!
; Be = en we oe host hea
‘pasuviie ATPeoLawoas sods Ssuimoys ‘(o1n,
hens Feta
=- . = > -y
" ar, Boas rake WD a.
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bs
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“3
oI
my
52
See page
ib
GENERAL FEATURES 49
but no particulars of individual specimens are given by him. ‘Then a
Tweedside angler, who favoured me with his views on “ bull trout,”
puts the limit of weight of these Tweed fish at 25 lb. Mr. Malloch
usefully gives a photograph of a sea-trout of 19} |b., the “ largest ever
caught in the Tay.’ I have personally seen a sea-trout of 12 Ib. which
was caught with the rod in Loch Lomond, and specimens are sometimes
taken in the Clyde estuary nets which weigh as much as 15 |b. But I
do not know of any statistics which give reliably the weights of the
heaviest specimens known to have been caught in British waters."
The limit of age is even more difficult to arrive at. So far as I am
aware there are no authentic records in the case of sea-trout. It will be
seen that on a later page (page 143) I submit evidence from the scales
of a Loch Lomond fish, which weighed only 23 lb., that its age was
actually as much as eleven years. The fixing of any definite limit of
age with our present knowledge is probably yet a matter of pure
surmise. Mr. J. A. Harvie-Brown speaks to having seen a trout
nineteen years old. It had been kept in confinement for that time.?
There are some points of interest connected with the spots of
salmon and sea-trout which I may here conveniently indicate. The
dark spots of the salmon are, in fish which have not previously spawned
— “maiden” fish as they are commonly called—few and irregular in
shape. They are (Fig. 1) on the body, mostly scattered above the
lateral line with a few of intense black on the cheeks. In a fish which
returns to spawn a second time the spots are now recognised to have
undergone considerable change. They have become much more
numerous (Fig. 2), and suggest the appearance of having been lightly
sprinkled on the surface of the shoulders and sides. Until recent
years these peculiarly speckled fish were thought to be a kind of “ bull
”
trout,” and, from the fact of their being frequently seen in that river,
they were popularly known as “ Tay bull trout.’ The Tay fish,
1. Day instances a Tweed male bull-tront of 441). caught in 1868.
2. ‘The Wonderful Trout ” (1898). p. 16.
50 THE SEA-TROUT
weighing 42 lbs. and 4o lbs., referred to by Sir Herbert Maxwell on
page 194 of his book on “ Salmon and Sea Trout,” and accepted by
him as being “ bull-trout,” were no doubt really salmon of this class.
Mr. H. W. Johnston, Mr. W. L. Calderwood, and Mr. P. D. Malloch
have amongst them proved that these large speckled fish are true
salmon which, having spawned in a former season, are now returning
from the sea to spawn a second time. I have not observed that the
spots of sea-trout undergo any similar modification consequent on
spawning.
The variation of the sea-trout’s markings, in the size and pattern
of the spots, is endless. The two types shown in Fig. 3 are as
distinct as well can be. They are both male fish, weighing 24 1b. and
23 lb. respectively, and were caught by two angling friends on the
same day in Loch Lomond. The spots are generally, but not
always, more numerous than those of the salmon, while the dorsal fin
of the sea-trout is also commonly spotted, a distinction not shared
by the salmon. I have even seen the upper lobe of the tail of a heavy
male sea-trout spotted, but this was when it was in full spawning livery.
At normal times the tail seems to be more generally fringed with a dark
bar plainly noticeable in the swimming fish. The regularity of this bar
was a feature of the fish which I used as a model for the frontispiece of
this book (Plate I). The spots of a fresh run sea-trout are markedly
cruciform, or X-shaped (Fig. 3 (2)), but not invariably so, and many
occur below the lateral line, but as spawning time approaches, the spots
tend to become blurred and, in the male fish especially, other spots not
visible on the scales of the fresh run fish begin to show as the fish
changes colour. (Compare Fig. 4 with Fig. 3 (1)). This discoloration
in the sea-trout, as in the case of the salmon, may perhaps begin before
a late running fish leaves the estuary, but I have not personally proved
the fact.
The spots of trout are even more diversified than those of sea-trout,
GENERAL FEATURES 51
and almost every water boasts its own variety and sometimes several
varieties, of size, shape and pattern. A peculiarity in trout spots often
observable is that each spot has round it a pale purple-coloured halo,
a feature only observable in sea-trout parr and in the mature fish after
it has become discoloured by residence in fresh water. In small trout
(fario) this halo is often very clearly marked (Fig. 5). The specimen
figured is a “two year old” farvio which Mr. Jas. A. Muirhead kindly
sent me from the West of Scotland Fishery, at Bridge of Weir. It
was so boldly spotted as to be rather a remarkable specimen. The two
trout also figured (Fig. 6) show the more usual appearance of the “ two
year old.” The Norwegian river trout of 6 in. in length (Fig. 7), whose
photograph Mr. Abel Heywood, of Manchester, has kindly given me
for reproduction, may be usefully compared with the river trout 13 in.
in length also shown (Fig. 8), which was caught above the Falls of
Clyde in waters wholly inaccessible to sea-trout. The markings of
these fish are similar to and may be compared with those of the sea-
trout in Fig. 3 (1) and Fig. 4. As having a direct bearing upon much
that is discussed here and elsewhere in this book I have inserted two
plates reproducing coloured drawings of sea-trout, one fresh-run from
the sea (frontispiece) and the other a fish on the eve of spawning
(Plate IX). Asa rule it may be said that superficial markings have but
little scientific significance in determining species, though with reference
to the sea-trout’s external appearance one must take account of them in
a general description. It has, however, sometimes occurred to me that
a careful comparison made between the spots of the trout and sea-trout
respectively of various waters might help to shed a sidelight upon the
theory that there is only one species of trout in these islands.
It is generally supposed, and I think with justice, that the spots and
markings of trout may be attributed to protective colouration. This
will vary greatly with environment, although Dr. Ward in his interesting
book on the “ Marvels of Fish Life as revealed by the Camera ”’ points
52 THE SEA-TROUT
out that “food is an equally important factor in bringing about colour
changes,’ and I have no doubt that sexual selection temporarily
affects colour also; but I think we may neglect these latter factors
altogether here. The immediate point is that when the sea-trout is in the
sea his best protection will clearly be the unbroken mirror-like surface
of his body, but when he returns to fresh water a reversion to more trout-
like colouration will be his best protection if it serve no other purpose.
It may therefore be more than a mere coincidence that the trout-like
aspect becomes most marked as spawning time approaches when the
fish is in the smaller spawning tributaries. I have already shown in
my introduction that it is when a sea-trout has been some time in fresh
water that doubts arise in the angler’s mind as to whether it is a sea-
trout or a trout.
I understand that the “spot” is caused by local concentration of
the colour cells in the skin—and this may account for the almost colour-
less halo observable round the spots of trout—but such concentration
will be less necessary to the sea-trout while it is in the sea than while
it is in fresh water. As the spawning season approaches, the exterior
parts of the scales of sea-trout, as of salmon, disintegrate considerably,
especially in the case of male fish, while at the same time the skin
thickens so much that the scales become almost wholly embedded in it.
Therefore the spots are accentuated on the skin of the sea-trout at this
period and stand out boldly as they permanently do in the case of the
trout.
I now arrive at the point to which these observations have tended.
bf
One of the commonest “ patterns’ on Loch Lomond trout is a single
spot surrounded by others arranged in an exact pentagonal or hexagonal
diagram (Fig. 9), so exact that each might have been geometrically
drawn. Yet that is precisely the pattern of spots that commonly
appears on Loch Lomond sea-trout (Fig. 10), a pattern that appears
even halo-surrounded like a trout’s spots at spawning time. It would
GENERAL FEATURES
al
w
be very suggestive if the sea-trout of any other rivers were discovered
in their spots to reproduce at spawning time the typical pattern of the
spots of the non-migratory trout of the district. Such a fact, if
established, would suggest that the variety of marking of the sea-trout
is infinite just because the variety of marking of trout is infinite, or, in
other words, that the fish are really one species whereof some are
migratory and some non-migratory.
The sea-trout, in common with the salmon and trout, has what
scientists call a fusiform body “ moderately elongate and compressed,”
as Mr. Regan puts it, “ deepest at or in advance of the middle of the
length and tapering posteriorly.” It possesses the same number of
fins as salmon and trout do, and they are similarly placed in relation
to each other (Fig. 11).
The first dorsal fin projects from the ridge of the back midway
between head and tail. In direct line with it and midway towards the
tail is the second dorsal fin, generally called the adipose fin, the
distinctive badge of all the family Salmonid@. The caudal fin, or tail
fin, with practically equal lobes above and below the middle line,
completes vertically the posterior extremity of the body. The anal
fin projects vertically downwards and, in the sea-trout, backwards,
immediately behind the vent. The pelvic fins, which used more
commonly to be called the ventral fins—and perhaps might still be so
called with advantage—are set midway on the belly, as a pair, imme-
diately below the first dorsal fin, and springing beside each of these is
a little rudimentary, or, it may be, an aborted fin, making four fins in
this group. The pectoral fins, having their origin on each side of the
gullet behind the gill-covers, complete the number of ten fins which
salmon, sea-trout and trout possess in common.
One may revert to Mr. Regan’s scheme of classification for such
special structural features as characterise the genus Sa/mo, but in many
matters of unscientific import salmon, sea-trout and trout bear so much
FE
54 THE SEA-TROUT
of a general resemblance to one another that occasionally it happens
that doubts arise as to the species of an individual. It may therefore
be of some service to put on record here such marks of structural
distinction between the species as have been found to be generally
constant and more or less easily recognisable. To contrast, then,
S. salar with S. trutta :—
“ Of characters which distinguish Salmon from Trout,” Mr. Regan
says, differentiating broadly, it will be observed, between salary and
‘
trutla, ““ we may note that the dorsal fin has usually more rays (10,
exceptionally 9, to 12 branched rays in the Salmon, 8 to 10, excep-
tionally 11, in the Trout), and that the scales on the tail are larger, in
an oblique series from the posterior edge of the adipose fin downwards
and forwards to the lateral line numbering 10 to 13 in the Salmon, 13
(exceptionally 12) to 16 in the Trout.” These figures, Mr. Regan
explains, are based on an examination by him of more than a hundred
examples of each species, and he states that these numerical differences
are of special importance because they are not subject to change with
the growth of the fish. It may occur to the reader that if the salmon’s
branched rays range between g and 12 and the trout’s between 8 and 11,
any fish that possesses 10 may, for aught anybody can tell, be either a
salmon or a trout, especially if it has at the same time 13 scales in the
oblique line indicated, which, according to Mr. Regan, either species
may have. But of course there are other features which help to supply
corroborative evidence, in doubtful cases, for one opinion or the other.
“Tn a Salmon,” continues Mr. Regan, “ the tail is more constricted
at the base of the caudal fin than in a Trout, and consequently the
anterior caudal rays form more of a shoulder, so that a Salmon does not
slip through the fingers when grasped round the caudal peduncle, but
a Trout usually does; the caudal fin is more or less emarginate, or in
large specimens truncate, or even rounded, but is usually more distinctly
notched than in Trout of the same size.”
SECOND DORSAL, OR CAUDAL, OR
FIRST DORSAL FIN. ADIPOSE FIN. TAIL FIN
Om
Bioene:
(1) Tail of Salmon.
) Tail of Sea-Trout.
>
See page ss.
GENERAL FEATURES
un
wn
Every angler will recognise in the first part of this description the
“wrist” which, with the more spreading and fleshier lobes at the base
of the tail (Fig. 12 (1)), enables one to hold a salmon with comparative
ease. As to the marginal shape of the tail it is less easy to refer a
particular salmon or sea-trout, for purposes of identification, to any
constant abstract standard. In the young of both species the tail is
markedly forked or notched, and as both fish grow to maturity the notch
fills up tifl the margin of the tail in each becomes perfectly straight.
In old fish, in both cases, the tail tends to become even round in outline.
It is not probable that both fish would often tally weight for weight and
age for age at the same time, but in a case where a grilse of 4 lb. weight
and a sea-trout of equal weight were in contrast, the tail of the grilse
would still be forked while that of the sea-trout would be perfectly
straight or even rounded, in its outline.
Dealing with the fins Mr. Regan also points out that “In adult
Salmon the anal fin is less pointed than in Trout, so that when it is laid
back the last ray usually extends farther than the longest, the reverse
obtaining in the Trout.” A reference to the fish themselves will clearly
show the distinction indicated. I have endeavoured in the diagram
(Fig. 12) to show the main distinctions between the tails and anal fins
of salmon and sea-trout and to contrast their scales.
Lastly Mr. Regan points out that ‘‘ The maxillary extends to (in
Grilse or small Salmon) or a little beyond (in large fish) the vertical
from the posterior margin of the eye, being shorter than in the Trout.”
Put rather more crudely the trout has a longer jaw and consequently a
relatively bigger mouth than the salmon (Figs. 13 and 14).
I make no apology for thus stating Mr. Regan’s distinctions in his
own words. I myself profess to have no scientific knowledge, and in
this matter prefer to offer the reader the safe guidance of his expert
authority.
It is interesting that Mr. Malloch makes no allowance for variation
56 THE SEA-TROUT
in the number of scales in the oblique line mentioned by Mr. Regan.
There are 14, he states,“ from the adipose or dead fin to the lateral
line’ in the sea-trout, 10 in the salmon; but he does not indicate, as
Mr. Regan properly does, which line of scales should be counted. As
to another structural distinction, in a letter to “ The Fishing Gazette”
of 10 May, 1913, Mr. Malloch wrote :—“ The quickest way to tell a
sea-trout from a salmon is to lay a pencil along the upper jaw past the
eye. If the eye is all above the pencil it is a sea-trout. If the pencil
line is through the centre of the eye itis a salmon. The difference is so
great one can tell at a glance without the pencil. The same with the
smolts of the salmon and sea-trout.” An illustration of using the
pencil is given in the second edition of his book. It may be remarked
that the late Mr. Cholmondeley-Pennell, in ““ The Angler Naturalist,”
also drew attention to this distinction, and it is possible that both
observers were in this matter under obligation to Yarrell. In “A
History of British Fishes” at any rate one finds that Yarrell gives
contrasted diagrams of the gill-covers of the salmon, grey trout and
salmon trout, and states that “looking at the form of the three gill-
covers, it will be obvious that a line drawn from the front teeth of the
upper jaw to the longest backward projecting portion of the gill-cover,
in either species, will occupy a different situation in respect of the eye;
that the line will fall nearest the centre of the eye in the first, that of the
Salmon, and farthest below it in the second, that of the Grey Trout.”
I reproduce here, roughly, for reference (Fig. 15), Yarrell’s diagram of
the three gill-covers reversed.
Yarrell, of course, accepted the grey trout, or “bull trout,” as a
distinct species, and so he attached considerable weight to distinctions
in the form of the gill-covers, a distinction which some modern observers
have not wholly ratified. But it is clear that the position of the eye
seemed to him to be of importance in the distinguishing of the species,
and a reference to several specimens of salmon and sea-trout will satisfy
GENERAL FEATURES
on
“J
the reader that the point is in fact material as a mark of distinction, at
any rate between salmon and trout.
Such bearing as the gill-covers have upon the matter in hand may
best be explained by reference to the diagrams of the heads of salmon
and sea-trout (Figs. 13 and 14) here given.
In the salmon, as Yarrell noted, the posterior free edge of the
gill-cover formed by the operculum, suboperculum and interoperculum
is almost invariably of so true a curve as to form part of a circle. In
the sea-trout, and, one may say, in the trout also, the free edge, where
the junction of the operculum with the suboperculum occurs, projects
backwards as a rule beyond the line of the curve of a true circle. One
cannot say that the distinction is absolute or constant, for the curve in
each species may approximate sometimes so closely as to be indistin-
guishable; but in most cases the true curve of the gill-cover of the
salmon will be easily marked off from the more projecting curve of the
gill-cover of the sea-trout.
Something also falls to be stated regarding the teeth in this connec-
tion because, although the young salmon, sea-trout and trout, begin life
with equally well-armed jaws, most of the teeth of the salmon disappear
as the fish attains maturity. As a distinction in mature fish of the two
species we are considering the teeth therefore are of some importance.
Yarrell here again puts the matter very clearly. “I have observed,”
he says, “that some specimens of the migratory or Sea Trout carry
their vomerine teeth longer than the Salmon; and the Trout which do
not migrate appear to carry their vomerine teeth longer than those which
do migrate.” He remarks that the salmon loses a portion of the
vomerine teeth during the first visit to salt water, and states further :—
“The teeth on the vomer of the Salmon, when the fish is old, seldom
exceed two or three in number, sometimes only one, and that placed
on the most anterior part.” It may thus be inferred that any large fish
which discloses a fairly complete set of teeth on tongue, jaw and vomer
is not a salmon.
58 THE SEA-TROUT
I may add as a further structural distinction, on the suggestion of
Mr. J. Arthur Hutton, that microscopic examination of the scales
shows, in most cases where the “ bull trout” is not involved, whether
the fish in question is a salmon or a sea-trout, for, weight for weight, the
salmon almost invariably is the younger fish. For purposes of com-
parison I am able, through the kindness of Mr. Hutton, to show in one
plate (Fig. 16) the contrast between the scales of a grilse and those of a
sea-trout each of which fish weighed 4} 1b. The scales respectively
indicate that the grilse is three and a half, and the sea-trout seven and
a half years old. It is possible, as I have elsewhere shown, that the
scales of the “ bull trout ” approximate closely to those of the salmon
in its various stages.
Subject to the foregoing remarks it may perhaps be convenient to
tabulate thus :—
Tue DISTINCTIONS BETWEEN SALMON AND SEA-TROUT.
1. Weight for weight, the sea-trout is generally shorter and of more
sturdy build than the salmon.
2. The gill-covers of the salmon form a true curve, those of the
sea-trout project backwards more or less to an apex.
3. The eye of the salmon is set nearly in the medial line of head
and body, that of the sea-trout markedly above it. The pupil of the
salmon’s eye is curiously pear-shaped.
4. The mouth of the sea-trout is larger than that of the salmon,
extending backwards beyond the eye. Relatively, the maxillary bone
is longer than in the salmon.
5. The salmon has generally more branched rays in the dorsal fin
than the sea-trout.
6. The posterior margin of the anal fin in the salmon tends to
convexity while in the sea-trout it is rather concave and projects towards
the tail.
7. The tail rays of the salmon spring from a relatively broader
base than in the sea-trout giving the former a “ wrist’ by which it can
be securely grasped.
Fig, 13. Diagram contrasting heads of male Salmon and Sea-trout.
(1) Head of male salmon.
(2) Head of male sea-trout.
Fig. 14.—Diagram contrasting heads of female Salmon and Sea-trout.
(1) Head of female salmon.
(2) Head of female sea-trout.
eee
=
Fig. 15.—Diagram of gill-covers of Salmon-Trout, Grey Trout, and Salmon.
(after Yarrell, reversed).
See pages 55 and 56.
GENERAL FEATURES 59
8. In fish of equal weight the margin of the sea-trout’s tail is less
forked than that of the grilse.
g. The scales of the salmon are generally actually, and always
relatively, larger than those of the sea-trout.
10. In the oblique line of scales traced forwards from the adipose
fin to the lateral line, the salmon has, as a rule, to scales, the sea-trout
14.
11. The rings of annual growth shown on the scales of salmon and
sea-trout widely differ, those of the sea-trout, weight for weight,
generally indicating greater age.
_We may now turn to the question whether there are any structural
differences between sea-trout and trout, a question which I have warned
the reader would crop up again and again throughout these pages.
For the scientific answer we are forced back upon Mr. Regan’s dictum
that “there are no structural differences.”
I have already stated that, failing proof to the contrary, Mr. Regan’s
dictum must be accepted, and I for one have no conclusive rebutting
evidence to offer. I may say indeed that I have again and again, with
sea-trout and trout of equal weights before me, examined them minutely
to see whether I could detect any particular part, or even outline, which
could fairly be held to differentiate the one fish from the other.
Hitherto (with the doubtful exception of a different rate of scale growth)
I have been unable to do so. It must be said, however, that in general
outline the trout of some rivers have a clumsier appearance than the
sea-trout caught in the same stream, while fin and tail have the
appearance of being coarser in texture. But on the other hand river
trout differ as greatly from the better proportioned trout which can be
caught in almost any lake.
In the general case it will be found that with equally well-
proportioned fish it is the colouration alone that enables one to distin-
guish whether a particular fish is a migratory or a non-migratory trout.
60 THE SEA-TROUT
| may now state more precisely wherein the teeth of trout differ
from those of sea-trout. There are always, I think, two rows of teeth
on the vomer in the case of both fish in their earlier stages of growth, but
as the fish get older the teeth are more or less freely shed and they
disappear the more rapidly as the fish adapts itself to a salt-water
environment. As to this matter Dr. Day, I think, writes with consider-
able weight of authority. “‘ However,” he states, “ the dentition varies
excessively, while we find examples possessing the colours, form, etc.,
of the brook trout resident in brackish waters or even the sea, but mostly,
not invariably, possessing the limited number of vomerine teeth of the
anadromous forms. On the other hand there are anadromous forms
(in colour) in fresh water, with the teeth assuming that present in the
brook trout or retaining the par dentition. It has been asserted and
reasserted that brook trout invariably have a double row of teeth along
the body of the vomer, and some authors have gone so far as to insist
that these teeth are not deciduous. Doubtless it is not uncommon to
find trout, even up to 2 lb. weight or even more, with all the vomerine
teeth thus remaining intact when a double row is present, but it is by no
means rare to see only one irregularly placed row. While in very large
specimens these teeth (unless they have entirely disappeared) are always
in a single row, and the vomer may be found even toothless or with one
or two teeth at the hind edge of the head of that bone. Equally
incorrect is the statement that the teeth disappear differently in different
forms, for in all they first assume a single row and then fall out, first
commencing from behind. But in the rapidly growing sea-trout the
vomerine teeth are shed sooner than in the brook trout.” In illustration
of this passage from Dr. Day’s work the reader may refer to the drawing
which I give of the vomer of a trout weighing 13 0z., taken from above
the Falls of Clyde (Fig. 17). The spaces which the shed teeth occupied
are distinctly seen, and the double row of teeth is clearly in process of
becoming a single row. The diagram (Fig. 18) shows the dentition of
a sea-trout of 24 Ib.
GENERAL FEATURES 61
Nor can it be said that the test of the scales will prove final as a
distinction between sea-trout and trout, for the concentric rings of
growth are added to the scales of the sea-trout exactly in the same way
as in the case of the trout, while both fish remain in their earlier stages
of growth in fresh water. No doubt an immediate distinction takes
place as to the scales of the sea-trout when that fish migrates to the
sea, the rings of growth which are then added being comparatively wide
apart, so that the salt-water series of rings contrasts strongly with the
fresh-water series. But the same kind of distinction immediately
occurs in the rings of growth of the scale of a trout when by chance it
happens to change its quarters from an ill-provided and circumscribed
hill stream or tarn to a rich and extensive lake.
I am far from wishing to appear dogmatic on this still somewhat
theoretical question of the identity of the sea-trout and trout—or, |
should perhaps put it, certain sea-trout and trout—but the fact has
apparently to be faced that it is not very easy to discover any such
characteristic structural differences between the fish as would warrant
their systematic classification in distinct groups.
Turning now to another matter, it is not always easy, with fresh-run
sea-trout, to distinguish the male from the female. The distinction
becomes more marked as the spawning season approaches, for then the
male fish» becomes reddish coloured and the female, as a rule, a dingy
grey. But if, on firmly closing the mouth of any sea-trout, it is found
that the under jaw closes well within the upper, the fish may be
pronounced a female. If not, it is more probably a male, and at
spawning time, or, in the case of old fish, at any time, a more or less
well developed “ hook” will be found on the lower jaw (see Figs. 13
and 14) which emphasises its length relatively to the upper. An
autopsy will, of course, always disclose the sex.
There is still the question to be considered whether, in respect of
the migratory trout, there is sufficient warrant for making a distinction
62 THE SEA-TROUT
between the sea-trout proper and the alleged “bull trout.” I have
already considered the matter at some length in my introductory chapter
and there is the less occasion to labour the subject here. “Two points,
however, may be referred to.
We have seen that Mr. Regan agreed that a slight difference might
sometimes be recognisable between western and eastern sea-trout. In
his own words, “ the Sewen (S. cambricus) of Wales, Ireland, and our
Western coasts often differ from the Sea-trout (S. albus) of the east
coast, in having a longer head, a larger mouth with stronger jaws, the
suboperculum projecting backwards beyond the operculum, and the fins
somewhat larger, the lobes of the caudal especially being more pro-
duced.” Now it was partially in respect of the shape of the gill-covers
that Yarrell distinguished between the “ grey trout” of the Tweed, and
ether rivers, notably those of South Wales, “where it is called the
Sewin,” and other sea-trout. He found that “ The operculum is
larger; the free vertical margin much more straight; the inferior
posterior angle more elongated backwards; the line of union with the
suboperculum not so oblique, but nearly parallel with the axis of the
body of the fish; the inferior edge of the suboperculum parallel to the
line of union with the operculum; the interoperculum much deeper
vertically; the vertical edge of the preoperculum more sinuous.” The
reader who admires the zealous exactness of this description may trace
its various parts with reference to the diagram (Fig. 15) given at page
56. I shall merely note that Mr. Regan’s observation of “ the
suboperculum projecting backwards beyond the operculum” and
Yarrell’s of the operculum having its “inferior posterior angle more
elongated backwards ” both help to describe the fact that nearly always
the gill-covers of sea-trout, unlike those of salmon, project backwards
in a curve considerably more abrupt than the circumference of a true
circle, the free edge of the suboperculum almost invariably projecting
beyond the edge of the operculum. That there is any distinction in
rd year
2nd year,
Ist year
Centre.
| | | |
Nn —"
= a
ont +=
— =
2. = oe)
+2
9
(1) Sez
Fresh water life, 4 ye
Fresh water life,
tig. 16.—Scales of a Sea-trout and a Grilse of equal weights contrasted to show different rates of growth
See page 58
17.—Vomer of Clyde river-trout Fig, 18—Vomer of Clyde sea-trout
(13 oz.) showing dentition. (x 6.) (21 Ib.) showing dentition. (x 24.)
See page 60.
GENERAL FEATURES 63
this respect between the sea-trout of different rivers sufficiently constant
or pronounced to warrant them being classed as separate species, or
even races, neither Dr. Ginther nor his successor, Mr. George A.
Boulenger, would agree, and I see no good reason to differ from their
opinion. ‘Lhe fact, however, may have cumulative weight.
As with the head so with the tail. I have already shown that, as
the sea-trout becomes older, the posterior margin of its tail tends to
become convex in outline, or, in other words, the hitherto square-ended
tail becomes round. The more pronounced tendency in this direction
in the sea-trout of the Tweed and some other East Coast rivers, and
the fact that such sea-trout are in general of a larger average size than
those of other districts, might point to the conclusion that they are older
fish, the roundness of the tail in the larger specimens having had longer
time to become accentuated. Careful examination of scales will settle
the question of age. Also one might infer that these so-called “ bull
trout ” are old fish, or at least that they have very frequently spawned,
trom the fact that their flesh is always poor and white and very unlike
the rich condition of the flesh of the smaller and presumably younger
specimens. Scale examination will settle this point also.
In this connection I am able to give, for purposes of comparison,
the diagrammatic outline of the tail of a 34 lb. male sea-trout caught at
Luss in November, 1914, and of the tail of a 71 lb. male “ bull trout ”
caught in the Tweed in the same spawning season (Fig. 19). The first
outline I took myself, and the second was kindly taken for me by a
member of the watching staff of the Tweed. One must admit that,
even taking account of the relative weights of these fish, there is an
essential difference in the outlines of the tails. Possibly, in the case of
each district, larger fish would show even more definite variation.
I am not contending either way for differences discoverable
between these “ bull-trout”? of the Tweed and other rivers and the
sea-trout of other districts. But the facts point to differences
64 THE SEA-TROUT
superficially apparent, though it is doubtful if they are a structural
difference warranting special classification. In the matter of size,
for example—because it must be admitted that these East Coast
sea-trout attain a large average size—there may be an augmentation of
growth through the fish ranging further afield in the North Sea.
Hardly any of the East Coast rivers in which these “ bull trout” are said
to be numerous have any estuary. The Scottish Tyne, the Tweed, the
Northumberland Aln and Coquet, and the English Tyne have none of
them, except the last, a distinctive and extensive estuary, and the local
sea-trout may possibly have acquired habits like those of the salmon of
ranging far afield in the ocean, with a proportionate effect upon their
growth. In the regions of pure surmise also it might be argued that
they go far afield because they are descendants of fish which in earlier
geological times had to traverse great river-channels long since
submerged in the waters of the North Sea. Such speculations have a
certain attraction. But there may be this prosaic explanation of the
size to which these sea-trout attain that, as they run in greatest numbers
after the netting close-time has begun, there is very little toll taken of
the stock and that therefore a higher percentage, or a greater proportion,
of the older and larger fish survives. However this be, I think that Mr.
Regan, apart from his theory that there is only one species of trout in
these islands, has not conclusively established that there are sufficient
grounds for recognising either two species, or two races, in the sea-trout,
although the cumulative effect of the facts may help to establish that
there are two types of sea-trout, and, I believe, two species.
This is clearly one of the points connected with the sea-trout and its
life-history in regard to which more definite information is required, and
I think that scale reading, if systematically undertaken, will help to
throw considerable light on the subject.
I have not myself, I regret to say, had the opportunity yet to
make such a systematic examination of the scales of East Coast fish,
Fig. 20.—* Bull Trout” (34 1b.) from the River Aln.
ord year.
ee
WN
)
{
Centre
Fig. 21.—Scale of River Aln “ Bull trout ”’ (3} Ib.), indicating
growth similar to that of a salmon erilse,
Fresh water lite, 2 years; after migration, ly years. Age, 35 years 32).
GENERAL FEATURES 65
but I am able to give a reproduction (Fig. 20) of a photograph of a sea-
trout which was caught in the coastal nets off the mouth of the river Aln
on 27th July, 1915. The fish, a male, weighed 3} lb., and there is, so
far as I can see, nothing very distinctive in its appearance. Mr. J. J.
Hardy, of Alnwick, who kindly sent me the photograph, states that “ it
is quite a typical fish for the Aln and Coquet.” There is something to
be said about the scales, however, and their appearance must give one
pause. They are far more like the scales of a grilse than one would
expect to see in a sea-trout weighing as much as 3} lb. The reproduc-
tion of one of the scales here given (Fig. 21) clearly indicates, I think,
that the fish spent two years’ residence in fresh water before its descent
to the sea as a smolt; thereafter it spent its third year in the sea, and
was apparently caught when making its first return to fresh water to
spawn in its fourth year. Now this would be exactly the life-story of
a grilse if the fish belonged to the species Salmo salar. It is, however,
undoubtedly a sea-trout, and the scales point to its being a maiden fish
in its fourth year; but, if so, one would in ordinary circumstances expect
it to weigh hardly more than at most 14 1b., equivalent to the weight
which one is accustomed to recognise in those sea-trout which, failing
to ascend the river as whitling, ascend in the following year as maiden
fish. But this fish weighed 34 Ib.
Now, if one is to accept the foregoing scale reading as correct—and
there seems to me no good reason why one should not do so—some
interesting speculations suggest themselves. The fish may be an
ordinary sea-trout whose rate of growth has been abnormal, which is
unlikely. East Coast sea-trout, again, may all show an equally rapid
growth, which, in view of the data given by Mr. Malloch of Tay fish,
seems also unlikely. Finally it may, after all, be that the “ bull trout ”
is a distinct species, or race, whose rate of growth in the earlier stages
is practically equivalent to the rate of growth of the salmon grilse.
I also reproduce here (Fig. 22) another sea-trout, weighing 8 lb.,
66 THE SEA-TROUT
which was caught at Alnmouth on 6th September, 1915, and show
(Fig. 23) an enlargement of one of its scales. Attributing to this fish
two years of fresh-water residence, it is quite plain that it spent two
further consecutive years in the sea without spawning, and was,
when caught, presumably making for the river to spawn in its fifth
year. Now if the fish shown in Fig. 20 discloses a life-history corres-
ponding to that of a salmon grilse, the life-history of this second fish
bears exact resemblance to that of a small summer salmon. Yet it is
indubitably a sea-trout, and, as sea-trout go, a large one ; but—and this
is the point—an unexpectedly young one. The question therefore
arises whether these two fish do not, after all, represent a distinct
species, or race, of sea-trout, which in rate of growth and possibly wide
range (evidence of which may be found in the late seasonal ascent of
the fish) approximates more closely to the salmon than to the trout.
I concede at once that no conclusions of value can be drawn from
the examination of the scales of only two specimens. But I give my
observations upon them simply to show in what directions a careful and
extended examination of the scales of East Coast fish may be fruitful of
interesting and definite results in a matter as to which, frankly speaking,
nothing at all is known.
It is of considerable interest that in Norway also the “ bull trout”
is recognised as a distinctive type of sea-trout. My friend, Mr. G. H.
Ramsbottom, of Alderley Edge, informs me that when fishing the
Gloppen River, Nordfjord, in 1913, he caught with fly a fine specimen
of the type which weighed 84 1b. Local people actually used the term
“bull trout ” to describe the fish, but the term was doubtless borrowed
from some British angler. Be that as it may, the important point is
that apparently the two types of fish are easily distinguishable. It was
further said that the “bull trout” of the Gloppen River occupied a
certain tributary stream at spawning time to the total exclusion of the
other and commoner type of sea-trout. I am able to give a reproduc-
lo
Fig. 19.—Diagram contrasting outlines of tail of a Tweed “ Bull Trout ’’ and
of a Loch Lomond Sea-trout.
(1) Tail of Tweed “‘ bull trout ’’ (73 Ib.).
(2) Tail of Loch Lomond sea-trout (3+ Ib.).
See pa ge O-
GENERAL FEATURES 67
tion of a photograph of Mr. Ramsbottom’s fish (Fig. 24), but unfortu-
nately no scales of the fish were taken for examination.
As it may be necessary from time to time to refer to points of law
in relation to my subject it seems appropriate here to remind the reader,
while we are at any rate considering distinctions, that, according to
Scottish statutory law, no distinction exists between a salmon and a
sea-trout, as the definition given in the Acts of the former expressly
includes the latter.’ Hence every statutory prohibition or provision
which affects the salmon and its young applies with equal force in
Scotland to the sea-trout and its young, a fact which it is very necessary
that the angler should know. I understand that the Iaw of England
is substantially to the same effect. The law which applies to trout and
trout fishing depends upon an entirely different set of statutes and legal
principles.
The life-history of the sea-trout, to which it is time now to turn, will
perhaps be brought most clearly into focus for the reader if the
description proceeds ab ovo. This will give an opportunity of
discussing in proper sequence the various stages of development in
the growth of the fish and the various important questions connected
with each.
1. “*Salmon’ shall mean and include salmon, grilse, sea-trout, bull trout, smolts, parr, and
other migratory fish of the salmon kind.” Salmon Fisheries (Scotland) Act, 1862, 25 and 26 Vict.
ce. 97, Sec. IT.
4th year
3rd year.
2nd year.
—— ist year.
Centre.
j a {
w\
\
oY)
Fig. 23.—Scale of River Aln “ Bull trout” (8 lb.), indicating
growth similar to that of a small summer salmon.
(x 16.)
; after migration, 24 years Age, 44 years
Fresh water life, 2 years
Fig. 24.—A Norwegian “ Bull
Prout *’ (84 Ib.
ges and Alevins
CHAPTER: Ii;
EGGS {AND ALEVINS.
Sea-trout eggs and salmon eggs are so like each other in point of
size and colour that they cannot readily, if at all, be distinguished at a
glance. As it is the size of the individual specimen rather than the
species which, with these closely-related fish, determines the size of
the egg, the sea-trout eggs, like the sea-trout themselves, are generally
but not invariably the smaller. The same determining cause holding
good, it will be found that trout eggs in general are smaller than sea-
trout eggs. As a rough indication of size it may be said that four
salmon eggs placed in a row almost exactly occupy the space of one
inch, sea-trout eggs on the average rather less, and trout eggs again less,
although five of either of the latter will generally exceed the inch.
I have found that twenty-six salmon eggs laid in line occupy
exactly the space of six inches, but I have seen an equal space occupied
by the same number of eggs of a well-developed sea-trout of only
21 1b. weight. It has been said, though I can neither confirm nor
disprove the statement, that the eggs of spring salmon are larger at
spawning time than those of autumn fish. The point does not seem to
be material with respect to sea-trout eggs.
The colour of the eggs of salmon, and of most sea-trout, when they
are first shed, is a rich salmon pink, but on fertilisation they become
immediately slightly paler and more opalescent. In one respect
salmon eggs differ wholly from those of sea-trout and trout. The
colour of the former does not seem to vary at all according to locality
or the breed of the parents, but there is a wide range of distinctive
colouration in the eggs of sea-trout and trout. They may vary from a
pale yellow to a dark rich pink according to the locality of the breeding
muy
72 THE SEA-TROUT
stock and the age of the parent fish. It is probable that the feeding of
the parents accounts largely for this uniformity in the case of the salmon
and variation in the case of trout, because, while the feeding of salmon
in the sea all round our coasts is comparatively of a fairly uniform
degree of richness, it is common knowledge that the fresh water and
estuarial feeding of trout and sea-trout varies both in quality and
quantity to an extraordinary extent in different localities. But while
feeding conditions undoubtedly affect the colour of sea-trout eggs to
some extent, the age of the fish, most probably on account of the
frequency with which it has formerly spawned which age implies, affects
it more. The older a fish is, the paler are its eggs. They approximate
nearly to the generally paler colour of the flesh. The vast majority of
salmon caught are “ maiden ” fish; all large, or at least old, sea-trout, as
will be shown, have previously spawned.
It may be worth noticing that salmon eggs are uniformly round,
while the eggs of sea-trout and trout are very irregular in contour. In
Plate II I have endeavoured to show the development of the salmon
from the egg to the smolt stage, and in Plate III the parallel develop-
ment of the sea-trout. I need not, I think, refer to these plates
specifically in connection with each point hereafter discussed. The
reader will perhaps bear this general reference in mind and turn to the
plates for any point which, up to the smolt stage, is mentioned in the
text.
As to the number of eggs which each species may shed, hatchery
operators are inclined to calculate the yield of salmon ova relative to
the weight per pound of the parent fish at about 850; sea-trout ova at
about 800; and trout ova at about 750.
Inspired by the example of Frank Buckland J once amused myself
by counting the eggs of a salmon, a well-shaped fish of 9% lb. weight.
I counted individually 1,000 eggs, and weighed the remainder against
them, finding the total to be 9,000 less 234 eggs, leaving a balance of
PrATE} lit
AY x
. : Ate
Ae a NANyhnews. WA ONS Ke
ALMO!
EGGS AND ALEVINS 73
8,766. There may have been a loss of, say, 34 in handling the fish
and detaching the ovaries, which would make the exact total of eggs
8,800, equivalent to almost exactly goo eggs per pound weight of fish.
Making the calculation another way I found that 260 counted eggs
weighed exactly 1 0z., so that a pound weight of ova would contain
4,160 eggs. As the mass of ova yielded by this fish weighed actually
2lb. 20z. presumably the eggs it contained numbered 8,840,
equivalent almost to the first result. But 900 eggs per pound weight
of fish is slightly in excess of the usual hatchery allowance of 850, which
figure may, however, represent a fair average.
It is not so simple a matter to calculate the number of sea-trout eggs.
One can never be certain that the female fish has not already shed
some eggs, and the number of eggs varies greatly in different females
according to the condition of the fish. Females out of condition,
obviously ill-nourished, contain few eggs, and old fish contain relatively
fewer eggs than young fish. Possibly 800 per pound weight of fish is
a generous average allowance.
On 12th November, 1914, when collecting sea-trout ova for Luss
Hatchery, I looked out for an opportunity of getting a perfectly shaped
unspawned female in order that I might count the exact number of
eggs she contained. In the afternoon I got such a fish which had
apparently just arrived to spawn in a small tributary of Luss Water.
It was a beautifully shaped and prettily marked fish of exactly 21 lb.
in weight. We stripped her of her eggs and fertilised them with the
milt of a male in a special basin. On arriving at the hatchery the eggs
were arranged by themselves on the glass grilles of one hatching box.
They were full-sized eggs, as large as average-sized salmon eggs and
very rich in colour. As is stated later in the chapter devoted to
’
“Artificial Propagation,” each glass tube in Luss Hatchery is a shade
over six inches in length and each row of the grille contains comfortably
26 eggs, a fact which I verified in this instance by counting the eggs of
74 THE SEA-TROUT
several rows here and there. As the eggs occupied 78 rows of the go
in the grille, there were apparently altogether 2,028, but allowing 52
for any possibly unshed and for loose eggs here and there on the grille,
we get a total of 2,080 eggs for this fish of 2} 1b. in weight. This
gives us (with one egg out) exactly 924 eggs per pound weight of parent.
Assuming that these eggs, as in the case of the salmon eggs men-
tioned above, weighed 260 to the ounce, the mass of eggs in this sea-
trout weighed exactly half-a-pound. Of course 924 eggs is greatly in
excess of the usual allowance of 800, but this was an exceptionally fine
specimen fish, caught too before a single egg had been extruded.
Most of our knowledge regarding the development of the eggs of
salmon, sea-trout and trout has been acquired in hatcheries, but there is
no reason to assume that such development differs in any important
respect from the development of the ova under natural conditions.
Perhaps hatchery conditions may accelerate matters to some little
extent because in nature development is often retarded by recurring
periods of low temperature, a disadvantage which the hatchery eggs
escape as the temperature within doors is kept at a uniform level. But
any difference is immaterial to the present purpose.
One can imagine, then, that in the fertilised egg, hidden away in
some dark recess of the gravel, with the temperature of the water
varying between 40° and 45° F.., the embryo fish will, after thirty days
or so, have so far developed that its eyes will have become visible as
two little dark specks through the semi-transparent covering of the
egg. At this stage, when the eggs are technically called “eyed ova,”
they may with impunity be subjected to a good deal of handling and
disturbance, and it is when in this state that the eggs of trout are usually
transferred from hatcheries to purchasers for the purpose of stocking
and re-stocking waters.
With things going propitiously, in other 60 days, or in about 90
days altogether (trout taking a total of 70 days and salmon 120 days)
Fig. 25.—Sea-trout eggs.
Fig. 26.—Sea-trout “ eyed ova.”
a
a
e
*
Ee ~e
‘ig. 28.—Late eggs and early alevins.
Ig. Ss i
See page 75.
EGGS AND ALEVINS 75
the sea-trout eggs will begin to hatch out through the imprisoned
embryo having become strong enough to burst its containing envelope.
The shorter time taken to hatch out of sea-trout and trout eggs
respectively may possibly be due to the higher temperature of the water
when the eggs are shed and during the earlier stages of incubation; at
any rate, in all cases, cold weather greatly retards the hatching process.
Most of the little fish, which when clear of the shell are known as
“alevins,” hatch out tail first, and easily succeed in detaching them-
selves from the shell. Some few, however, but in what proportion is
not known, hatch out head first and these, unless released forthwith,
generally perish, but whether by strangulation or through some inherent
lack of vitality in the embryo I am unable to say. In Figs. 25, 26, 27
and 28 will be seen reproductions of several photographs of eggs and
alevins, in one instance (Fig. 27) of the alevins actually in the process
of hatching out.
The little alevin is very unlike a fish. It is barely an inch in length,
and has a blunt head of which the prominent black eyes are the most
striking feature. The body is transparent and through it the rudimen-
tary organs are clearly seen. The pectoral fins are separate and keep
up a constant fanning movement, but the other fins yet consist of a
continuous fringe round the margin of the body, a fringe that will not
evolve into the permanent fins for several weeks.
But the most remarkable peculiarity of the alevin is a curious
transparent appendage called the umbilical sac, or yolk-sac, which
hangs between throat and vent, at first equal in bulk to the true body of
the fish. This sac, in which little drops of oil may be seen, gives
nourishment to the alevin during such period as it is unable either to
swim about or to feed in a natural manner. In fact, the alevin shuns
the light and wriggles into the deepest crevices it can find, prompted
doubtless by the instinct of self-preservation. But the umbilical sac
becomes gradually absorbed, the fish becomes bolder, and in about 50
76 THE SEA-TROUT
days, the sac being wholly absorbed, the little fish, perfect in shape,
may be seen darting about amongst the gravel. This is the “ sea-trout
+)
fry.
It may be convenient here (reverting to Mr. Regan’s dictum already
quoted) to consider in one view various differences, no. doubt more or
_less superficial, which I have noted as occurring between “the young ”
of salmon, sea-trout and trout.
1. Salmon eggs are uniformly round in shape while those of sea-
trout and trout are very irregular.
2. The eggs of salmon are practically uniform in colour; those of
sea-trout and trout vary considerably.
3. Salmon eggs take longer to hatch out than sea-trout eggs and
still longer than trout eggs.
4. The colour of the respective alevins differs materially and does
not seem to be necessarily dependent on the colour of the
respective eggs. The salmon alevin is always pale flesh
coloured—‘“ like finely chopped carrot” as it was once
expressed to me; the sea-trout alevin is at first crimson-red
in tinge and rapidly develops a dark brown colour on the
back. Trout alevins are much greyer than the other two all
the time they are alevins.
5. The salmon alevin is appreciably larger than the other two on
hatching, its body being markedly longer, while the greatest
‘ development of the umbilical sac appears to be midway in
the body rather than near the throat as in the case of sea-trout
and trout.
6. On being hatched, salmon alevins lie, as it were, dormant for
about eight days, the same period of inertia with sea-trout
lasting only two days.
7. The eye of the salmon fry is appreciably larger than that of
the sea-trout or trout fry.
8. At the earliest stage, parr marks are most sharply defined on
the salmon, less so on the trout, and least on the sea-trout.
DEVELOPMENT OF THE SEA-TROUT FROM THE EGG TO THE SMOLT STAGE
EGGS AND ALEYINS 77
’
I do not say that these “ differences” will be found of constant
occurrence, but they are worth noticing. At Luss I have had the
advantage of studying the ova of fish from commercial hatcheries as
well as of fish native to the district, and upon the eggs and the resultant
alevins and fry I have made notes at various times.
I think that the facts noted regarding the earliest stages in their
respective careers point to a marked distinction between S. salar and
S. tvutta, and to a considerable variation between the various breeds of
trout irrespective of whether they happen to be migratory or non-
migratory.
Distinctions between the fish at later stages of growth will be pointed
out as these stages are being considered.
A considerable number of “ monsters,’ or deformed alevins, is
always discovered amongst sea-trout hatched under artificial conditions,
although oddly enough few are ever found in the salmon boxes. No
very accurate estimates have been made by which it can be deduced
that the number of monsters is variable or fairly constant. In all
probability much the same number appears amongst naturally hatched
fish, but under natural conditions few deformities have any great chance
of being seen, and doubtless few survive to be seen. Some interesting
notes on this point are given in a work on “ The Teratology of Fishes ”
by my friend Dr. James F. Gemmill, Lecturer on Embryology at
Glasgow University. He states that the following figures have been
found to apply to deformities amongst the Salmonide :—1 in 50, and
I in 280 (Rauber); 1 in 600, none in 600, and 68 in 900 (Schmitt); 1 in
200, and 1 in 350 (Gemmill); over 100 in 400,000 (Coste). He adds
this interesting statement :—‘ It is worthy of note that the frequency
with which double monstrosity appears in the eggs of fishes is not far
from corresponding with its frequency in those of the fowl.”
Sometimes sea-trout with truncated snouts, spinal curvature, or
stunted tails are caught, the largest “ deformity’ I have seen being
78 THE SEA-TROUT
one—an extreme case of spinal curvature—where the fish weighed
about 3 lb. It is often thought by anglers who catch these fish that
some early injury may have been the cause of the deformity, but
apparently the origin goes back to the unhatched embryo. Dr.
Gemmill’s view seems to be that the actual causation has not yet been
definitely ascertained.
But the most curious “ monsters” found in the hatching boxes are
those of the Siamese Twin type (Fig. 29), where the “ twins” may be
attached to each other in a variety of ways, the result being double-
headed fish, double-tailed fish, or even double-bodied fish, in which
extreme case the umbilical sac forms the connecting link. But of this
type of “monster” it is safe to say hardly any survives the alevin
stage. The late Mr. Cholmondeley-Pennell, however, in “The
Angler-Naturalist,” states that, “In the river Towey, Carmarthenshire,
a fine fish of the Salmon or Trout species was caught, with the net,
which had two heads and two tails—the heads being joined on to one
neck, and the tails meeting about the centre. The fish was preserved
for some time in a small pool at Llangattock, for the inspection of
visitors.’ But the facts thus given concerning this fish are vague
enough. The author merely paraphrases those stated by Yarrell, who
adduces some evidence, with the date 1829.
It is often asked, as a matter of interest, what proportion of eggs
deposited reach the fry stage; and sometimes the question is put thus,
how many eggs are required to produce one mature sea-trout? Con-
servators of fisheries would be only too glad to discover the answer.
Taking one year with another the proportion is doubtless fairly constant
and in the chapter on “Artificial Propagation” which follows I give
Mr. J. J. Armistead’s estimate regarding trout. But unfortunately
there are no means of making even the roughest computation, and
hatchery work ‘throws but little light on nature’s operations in this
matter.
Fig. 29.—Sea-trout ‘‘ monstrosities.’
(after Gemmill.
(1) A double-headed alevin.
(2) An alevin of the Siamese-twin type.
EGGS AND ALEVINS 79
It is easy enough to assign such loss as must occur in nature to its
various causes. Amongst these must be reckoned original want of
fertilisation of a proportion of the ova shed and the immediate
devouring of some of it by predatory fish in attendance on the spawners.
Then the vicissitudes of the weather must be reckoned with, for an
excessive drought may lay bare the shingle in which the eggs are
deposited and exceptional floods may sweep the gravel beds wholly
away. Then there is a host of creatures—larve, beetles, fish, and
birds—which levy a constant toll upon the eggs and alevins. Finally,
it has been discovered through hatchery work that a heavy death-rate
occurs amongst the alevins, due perhaps, as it has been said, to an
inflammation of the gills at the stage when the umbilical sac has become
almost absorbed and the little fry begin to forage for food. I have
suggested elsewhere that probably this particular mortality is increased
by artificial rearing, especially where hand feeding is resorted to. But
in any case it requires no great stretch of the imagination to assume a
considerable mortality at this critical transitional period even with fish
in their natural environment.
All that human ingenuity can do to mitigate loss, without resorting
to hatchery operations, is to secure that as large a stock as possible of
healthy fish deposit their spawn in peace, and that the eggs and alevins
are protected as far as may be from predatory enemies.
It is hardly worth while to quote such estimates as have been made
of the wastage that occurs, but Dr. Francis Ward, in his “ Marvels of
Fish Life,” remarks that “ in nature only a small percentage of the eggs
deposited result in the birth of an alevin.” If this is so, how few of the
eggs hatched must produce a fish destined to reach maturity. Yet the
numbers of sea-trout, in some seasons, are in fact enormous.
I have postponed till later consideration of the actual process of
spawning and the periods of spawning, but, to summarise here what is
contained in the foregoing pages, we may assume the sea-trout ova to
80 THE SEA-TROUT
have been deposited on November 15; and we may tabulate the various
stages of development of the little fish up to the fry stage thus :—
Ova deposited (say)... sa .... November 15
Ova eyed, 30 days ve ae ... December 15
Alevin hatched, 60 days ve ... February 15
Perfect fry, 50 days... ae i) Apis
Altogether 140 days, or nearly five months, have elapsed since the
shedding of the ova. It is a fact, worthy I think of special observation,
that nature has arranged that the little fish have become best able to
fend for themselves just at the time when the stream in which they were
hatched is likely to afford a sufficiency of food for their subsistence
The Fry Stage
CHAPTER IV.
THE FRY STAGE.
By the time the umbilical sac has entirely disappeared the young
sea-trout has already learned to quest for food. It is interesting to
see that these little fish, even at this early stage, have inherited all the
graceful activity of their parents, as well as their graceful form. The
fry are pretty little things, about an inch and a quarter in length, which
boldly enough forage amongst the gravel of the shallower water and
‘
even “rise
”
to floating objects, but at any sudden alarm they take
cover with surprising quickness. At one moment a shallow stream may
be seen to be alive with them and at the next may seem barren of all life
whatever, so completely has each concealed itself in the gravel.
The body of the fry is at first still almost transparent, but it is
already delicately spotted and the transverse bars, or “ parr marks,” on
the sides, which characterise alike the young of salmon, sea-trout and
trout, have already appeared. Eight or more of these may be counted
on each side of the young sea-trout or salmon and indeed of the trout
too. There is apparently no constancy in this matter as regards any
of the fish.
Once the fry have begun to feed in the open their growth is fairly
rapid for their appetite is insatiable and very little that is edible and
of a convenient size escapes them. From this point of time the rivalry
between the sea-trout and salmon fry and the young native non-
migratory trout of the stream becomes acute. There is of course only
a certain amount of food available for all, and of that quantity the
young sea-trout from their greater activity and strength secure the
lion’s share. Here is a vital handicap against the trout in the struggle
for existence, and it may easily be understood that when fry and parr
of both sea-trout and salmon throng any of our streams, the native
83
84 THE SEA-TROUT
trout have small opportunity of being over-fed. As a matter of fact in
such circumstances they remain undersized and are of the poorest
quality. This is a matter of great practical importance in the proper
maintenance of trout waters, and I shall have occasion to refer later to
other handicaps against the trout and in favour of the sea-trout. It-is
my opinion that the cumulative effect of these is to reduce the value, as
trout fishing waters, of all waters to which sea-trout have access. From
another standpoint it is possibly a mistake to endeavour to encourage
trout in salmon and sea-trout waters. One might perhaps even go
further, and, having the history of the river Coquet more particularly
in view, argue that sea-trout are not fish to be whole-heartedly welcomed
in salmon rivers. In my introductory chapter I pointed out that the
broad theory of there being only one species of trout in these islands
would meet us at every turn in the life-history of the sea-trout. We
touched upon it in the case of the spots; we now come more closely
upon it. The question arises how far sea-trout may communicate their
pronounced migratory habit to the native non-migratory trout of any
stream. No one can really tell, but it seems to me that it is not an
unreasonable speculation to imagine that they do so to a greater or less
extent, and that, under special circumstances where, for instance, the
spawning grounds are limited in extent, all the trout of a stream may
become fish of pronounced migratory habit, while in other streams, the
bolder and stronger fish becoming migratory, the non-migratory
specimens left behind to people the stream—as, it may be said, “native”
trout—will be trout of poor quality.
It is noticeable that the young fish of any brood of sea-trout keep
together and feed as a shoal, far more so than is the case with either
salmon or the non-migratory trout, and, indeed, my belief is that this
habit persists to a certain extent all through life so long as any
individuals of the shoal survive. It might be going too far to say that
the brood of each female forms an independent and distinctive shoal,
THE FRY STAGE 85
but I think that the young fish which have hatched out together in
neighbouring redds in the one little spawning tributary tend to remain
together. 1 would wish to make my meaning perfectly clear in regard
to this matter. I do not of course pretend that the sea-trout fry are
gregarious, as a shoal of minnows which cruises up and down the still
waters of a pool is gregarious, each individual minnow conforming in
its movements to the general movements of the shoal. All I suggest
is that the sea-trout fry hatched out in any stretch of shingle remain
together in that stretch of shingle, until the necessity of seeking other
quarters where richer feeding may be obtained arises. When the
necessity arises I am inclined to think that all the fish hatched out
together seek the richer feeding grounds in a body, dropping down
stream to the main river, or to a loch, if such is within immediate reach.
The Loch Lomond district, with its numberless little streams, many
of which have perhaps no more than a dozen yards or so of spawning
ground, offers peculiar opportunity for observation. Where one of
these streams is so small as to be obviously incapable of providing food
for the growing fry in it, these are known to desert the stream and seek
sustenance in the loch where the shoal can be seen on a calm day
cruising along the shore. I believe, further, that sooner or later shoals
of these young fish even cross deep water to the feeding banks of the
numerous islands where these are near the parent stream. The shoals
which thus take up their feeding ground spread over the banks as the
growth of each young fish demands for it a wider range.
But doubtless every district has its own type of feeding ground, in
river, loch or even brackish water, and such fry as are not driven by
force of bare necessity to seek a livelihood elsewhere scatter themselves
over the shallows in the neighbourhood of the redds in which they were
spawned, redds which are almost always in some inconsiderable
tributary of a greater stream. Here they spend an active existence in
summer in pursuit of food and in evading their natural enemies. By
6
86 THE SEA-TROUT
the autumn they will have obtained a length of two or three inches.
As the temperature of the water falls and food grows scarce they retire,
like Cesar, into winter quarters, and, seeking a safe shelter below some
stone, they practically hibernate until the following spring when they
again emerge for another active summer of fresh-water sojourn. With
numerous opportunities for observation, I have never seen the young
=
sea-trout “ feeding” in a stream in mid-winter, but I have disturbed
them under the ice of a frozen river. |
Mr. Malloch has happily described this habit of the parr of salmon,
and I take it that the habits of the young sea-trout, at least of such of
them as have not sought sanctuary in a loch, are not materially different.
“Very little feeding,’ Mr. Malloch writes, “takes place after the end
of September, and when the cold weather sets in, the parr leave the
shallow water to take up their abode under stones, where they remain
till March or April, and almost entirely cease to feed. During their
stay in winter quarters they become very black and fall off in condition.
Often, when collecting larve, I have lifted a flat stone quietly and
disclosed to view three or four parr. These did not swim away at first,
but remained motionless for some time, apparently in a dazed, sleepy
condition. When the water becomes warmer their winter abode is
forsaken for the quieter pools.” Mr. Malloch adds a curious statement
difficult, one would think, to substantiate. ‘‘ Strange to say,” he writes,
“they are now smaller than they were during the autumn.” [I think it
is possible that, taking into consideration the wide distribution of
salmon and sea-trout in our islands, Mr. Malloch’s periods of feeding
may be somewhat over-definitely stated. In southern rivers, at any
rate, with warm weather and in low water, salmon parr may be observed
feeding freely during the first half of October and possibly earlier than
even March. I fancy that young sea-trout in this respect approximate
most closely to trout, which feed more or less freely as weather
conditions give them the opportunity, and that it is only during the
THE FRY STAGE 87
coldest period of mid-winter that there will be any prolonged cessation
from feeding. At any rate, the fish in the streams feed less because
there is less to feed on. Here, perhaps, the “reading” of a scale, as it is
technically called, may serve to confirm what is stated in the text as the
result of more general observation. The nucleus of the scale becomes
first visible to the eye some little time after the alevin has wholly
absorbed the umbilical sac. It is now of course generally known that
the scales are not annually shed, but that, growing with the growth of
the fish, they—besides other changes—add each season a series of rings
to their circumference. It will be necessary to point to the condition
of the scales at various stages in the sea-trout’s growth, and here it is
appropriate to state that the rings of growth acquired during the winter
between the first summer and the second summer of the fish’s life are set
more closely together than the rings acquired during each successive
summer with the effect of forming a sort of band, or ring, of demarcation.
This points to the two inferences that the salmon parr feeds hardly at all
in winter and that it practically ceases to grow during the winter months,
but it hardly bears out Mr. Malloch’s statement that the fish grows
“ smaller,” else there would be no “ band ” or series of winter rings, but
only one ring marking shrinkage of the body and disintegration of the
scale. But when it is pointed out that the winter band in the sea-trout
scale (and in the trout scale also) is seldom so clearly defined as in the
salmon scale, one may infer that sea-trout parr tend to feed with more
freedom during the winter months than salmon parr, a fact which helps
to account for the sea-trout’s proportionately quicker growth than the
salmon during the period of residence of both fish in fresh water.
It is worth while noticing these facts as to the sea-trout scales now,
at their earliest manifestation, because throughout the life of the fish
the scale rings of summer and winter growth may, as in the salmon
scale, be differentiated and distinguished.
It is not my purpose here to describe the general theory and practical
88 THE SEA-TROUT
application of scale reading, but only to refer to the subject in so far as
in my opinion it helps to shed light upon the life-history of the sea-
trout. For an introduction to the subject I need only refer the reader
to certain very clear and admirable works which have already been
written with reference to the scales of salmon by both English and
Scottish authorities.
Parr and Smolts
>) ee ee Se
— re rr on ae a — -— +
=
i
.
..°*
’ ¢
—
CHAPTER. V.
PARR AND SMOLTS.
With the advent of the warmer weather of spring the shallower
waters of the streams become again peopled with the active fry, which,
it will be observed, by the beginning of April have now lived through
twelve months since ceasing to be alevins. ‘They are in fact “ year-
lings,” and having survived the fry stage are beginning now, as
“sea-trout parr,” their second year’s residence in fresh water. How
long they may remain parr I will discuss immediately.
These sea-trout parr are as active as ever the fry were in pursuit of
food and they are bottom feeders, mid-water feeders and surface
feeders indiscriminately as each particular kind of food is in evidence.
They are, like salmon parr, troublesome to the angler, for they take his
fly and worm with avidity and will even boldly attack an artificial
minnow. If they are thus noticeable in the river they are equally so in
a loch, for fishing with fly inshore on the feeding banks the angler will
raise and hook numbers of them. It is clear that these, as I noticed
_ happened in the case of some of the Loch Lomond streams, are fry
which have left the parent stream for the loch and, having occupied a
convenient feeding bank there, have not since changed their quarters.
Nor will they do so, I think, till they are disposed to descend to salt
water as smolts.
In any case, whether im river or loch, the sea-trout parr grow apace,
outstripping in this respect the salmon parr of which at hatching,
however, it must be remembered they had fully a month’s start in life.
When they go into winter quarters for the second time they will measure
from four to as much as eight inches in length according to the quantity
and quality of the food they have been able to obtain. There is a
marked difference, at any rate, in the general condition of the sea-trout
91
92 THE SEA-TROUT
parr of a stream flowing gently through rich agricultural land and of
those of a rocky barren stream which hurries tumultuously down a
Highland glen. This initial difference persists through life and marks
the well-conditioned or ill-conditioned types of sea-trout characteristic
of the respective types of stream.
On this subject of the relative growth of salmon, sea-trout and trout,
Mr. Knut Dahl has much that is interesting and suggestive to say
regarding Norwegian fish,’ and I cannot conceive that, were equally
full investigation to be made regarding British fish, any great
discrepancy would be discovered between the results obtained in
Scotland and those obtained by him in Norway.
In Scotland, as every angler is aware, it is illegal to take salmon
parr, or, as the Act expresses it, “any smolt or salmon fry,” and as
sea-trout are “salmon” in the sense of the Salmon Acts, the provisions
on this head apply to them also. The section of the particular Statute
which deals with this offence relates also to offences concerning spawn
and spawning fish, and therefore I insert it fully here in the accom-
panying note.”
While it is thus illegal to take sea-trout parr the difficulty that meets
the angler everywhere and at all times is to distinguish young salmon
and sea-trout from the young of common trout, because, as there is no
law yet in Scotland prohibiting the capture of trout however small, the
1. “ The Age and Growth of Salmon and Trout in Norway as shown by their Scales,’ Knut
Dahl (1910). Trans. by Ian Baillie. Ed. by J. Arthur Hutton and H. T. Sheringham for the
Salmon and Trout Association.
2. “Every person who shall wilfully take or destroy any smolt or salmon fry, or shall buy,
sell, or expose for sale, or have in his possession, the same, or shall place any device or engine for
the purpose of obstructing the passage of the same, or shall wilfully injure the same, or shall
wilfully injure or disturb any salmon spawn, or disturb any spawning bed, or any bank or shallow
in which the spawn of salmon may be, or during the annual close time shall obstruct or impede
salmon in their passage to any such bed, bank, or shallow, shall be liable to a penalty not exceeding
five pounds for every such offence, and shall forfeit every rod, line, net, device, or engine used in
committing any such offence, and shall forfeit any smolt or salmon fry that may be found in
his possession ; but nothing herein contained shall apply to acts done for the purpose of artificial
propagation of salmon or other scientific purpose, or in the course of cleaning or repairing any
dam or mill lade, or in the course of the exercise of rights of property in the bed of any river or
stream: Provided also, that the district board may, with the consent of all the proprietors of
salmon fisheries in any river or estuary, adopt such means as they think fit for preventing the
ingress of salmon into narrow streams in which they or the spawning beds are from the nature
of the channel liable to be destroyed, but always so that no water rights used or enjoyed for the
purposes of manufactures, or agricultural purposes or drainage, shall be interfered with thereby.
Salmon Fisheries (Scotland) Act, 1868, 31 and 32 Vict. c. 123, Sec. XIX.
Prane DV:
1. A TROUT 2. A LOCH LEVEN TROUT 3. A SEA-TROUT
AND 4. A SALMON AS “YEARLINGS
PARR AND SMOLTS 93
angler may—quite innocently mistaking one kind for the other—incur the
considerable odium of a conviction. ‘There are those who may contend
that he deserves to be punished for taking even trout of the size of
salmon parr and smolts, but there is in many districts no discredit in
taking trout of nine or ten inches in length, and I have seen sea-trout
parr and smolts of that size quite innocently, if unlawfully, captured by
anglers and put in the basket. Anything that will help to guide the
angler to a safe decision in cases of doubt will consequently be of
advantage.
I am not sure that any author in this country has dealt very satis-
factorily with this matter, although a Norwegian author, Mr. Hartig
Hiitfeldt-Kaas, has tabulated certain more or less constant distinctions
between the young of salmon and trout. To tell the truth, it is by no
means easy to give a verbal description of such differences as may be
recognised between the several fish at their various stages of growth.
For convenience of reference I insert here a plate showing approxt-
mately the colouring and markings, when the fish are yearlings, 43
inches in length, of (1) a trout, (2) a Loch Leven trout, (3) a sea-trout,
and (4) a salmon (Plate IV). As a first step, I shall try to indicate some
distinctions between a salmon parr and a young trout.
One may start from the broad principle that if the angler is fishing
in waters which he knows contain salmon he is bound to be upon his
guard against taking their young. If he has any conscience at all in
the matter he will, after catching one or two of the smaller fish, be able
quite easily to detect that some of those he catches differ materially
from others. He may not be able to say at once wherein the difference
lies but-he will be quite conscious that there is a difference. It will be
found, I think, to be one of colour.
Young salmon, when held slanting to the light, have very obviously
a general bluish tinge; young trout, when similarly held, appear to be
of a yellowish-brown colour. This difference is so marked that I
94 THE SEA-TROUT
imagine the colouring of “blue and silver” and “ brown and gold”
phantom minnows is intended to give effect to it.
If then, having perceived this contrast, the angler now compares
the two kinds of fish more minutely, still dealing with the superficial
colouring and markings, he will see that the salmon parr is somewhat
differently marked from the young trout. Both kinds of fish for at
least the first year have a distinct series of dark slate-coloured oval
marks along their sides, the ovals being bisected by the lateral line.
They are variously referred to as “ bands,” or “arches”’ or “ finger
marks,” but the term “ parr marks ” describes them sufficiently.
The parr marks are not, in either the young salmon or trout, of
uniform number, but generally there are from eight to ten distinct ovals
along the side of the salmon parr; but a curious break, as if an oval
were missing or not fully formed, sometimes occurs in the middle of
the series. The “ parr marks” of the young trout are not only more
numerous and less regular in their oval shape but the curious breaks
referred to occur with more frequency in the series as if some of the
ovals had been broken into upper and lower portions by the lateral line.
The parr marks persist in the young salmon in definite outline until the
fish is on the eve of migration when the silvery smolt scales effectually
conceal them, yet on the scales being removed they may still be distinctly
seen. The parr marks of the trout usually become irregular in shape
and indistinct in outline in the second year, and appear then rather as
darker patches of the body colour. I have said “ usually,” because in
some instances, notably in Loch Freisa in Mull—and I may instance
also Loch Tay and Loch Lomond—the trout retain their parr marks
until they are fish of nearly a pound in weight. But in the general case,
when a trout is about seven inches in length, the marks have become so
indeterminate as to enable the fish to be easily picked out from a number
of salmon parr on this account alone. In order that the reader may
have an idea of the contrast between yearling and two-year old fish, I
PLATE V.
a
%,
“eet a wert |
» ,
=
LA TROUT 2. A LOCH LEVEN TROUT 3. A SEA
AND 4. A SALMON AS “TWO YEAR OLDS
PARR AND SMOLTS 95
insert here a plate indicating in colour the general appearance and
markings of (1) a trout, (2) a Loch Leven trout, (3) a sea-trout, and (4) a
salmon, when at the “ two year old” stage of growth (Plate V).
Young salmon and trout are both freely speckled with dark and
crimson spots, and as to these something may be said. The dark spots
of the salmon parr are small and occur only above the lateral line,
while those of the young trout are generally bolder and are often
numerous below the lateral line as well as on the back and shoulders.
Each spot of the trout, too, as I have noted elsewhere, is usually
surrounded by a ring or halo paler in hue than the general body colour.
But a very distinctive feature of the salmon parr which one clearly
recognises when attention is drawn to it is the intense black of the spots,
generally one, two or three in number, on the operculum, that is, on the
cheek. It may be noted too that the dorsal fin of the salmon parr is
never very boldly spotted, while that of the trout almost invariably is.
Dealing with the colouring of the fins, never a very certain feature,
it may be stated that the pectoral fins of the salmon parr appear of a
watery paleness compared with the rich yellowish-brown of those of the
trout. So too the adipose fin of the young trout has nearly always, but
I think not always, a reddish fringe, that of the salmon parr being in
colour merely a dark continuation of the dark blue back.
Again viewing both fish broadly, but now in regard to shape, it will
be seen that the salmon parr has a more graceful outline and a general
air of greater delicacy than the young trout. Coming to details, its
head is neater and its mouth relatively smaller than that of the trout,
for, while the jaw of the salmon parr reaches back to a level with the
middle of the eye, that of the trout extends as far as, or even beyond,
the posterior margin of the fish’s eyeball. The eye of the salmon parr
is also relatively larger than that of the trout, and, as already noted, is
set more nearly in the middle line of the head when viewed in profile,
that of the trout being set rather above the middle line.
96 THE SEA-TROUT
The pectoral fins and tail of the salmon parr are more delicate and
pointed than the rounder and coarser fins of the trout, and the deeply
forked tail of the salmon parr is a very distinctive feature.
It may be repeated that the numerical difference in the scales
formerly noted also holds good with both fish in the parr stage.
So much for colouring and structure, but I may add the curious fact
noted by Mr. A. H. Chaytor in his excellent “ Letters to a Salmon
Fisher’s Sons” that “A young salmon or a parr or smolt of any other
kind, when lifted out of the water, kicks and wriggles vigorously until
actually secured by the hand, whereas a small trout lifted out on the
line almost always hangs quietly or at most gives a few kicks,” and he
adds, “it is an absolutely certain way of knowing a young trout from a
parr or from a smolt.”
Altogether, it will be seen that there are a good many points which
differentiate the salmon parr from the young trout.
I have thus contrasted first the salmon parr with the young trout to
save complication of description as far as may be, and I shall now try
to point out wherein specially the salmon parr differs from the sea-trout
parr.
Perhaps, first, it should be said that in respect of size the two fish
differ greatly. The Royal Commissioners of 1902 found that about
go per cent. of the salmon smolts, as seen at Fochabers on the Spey,
“were about 7 inches in length, and weighed a little over 3 ounces.”
Hence it may be taken that a salmon parr in its third year seldom
exceeds seven inches in length. On the other hand sea-trout parr may
actually grow to nine or ten inches in length before becoming smolts.
In respect of general colour the sea-trout parr approximates more
closely to the young salmon than to the young trout, and curiously
enough this is characteristic of the Loch Leven “ parr ” too, the marked
bluish tinge in it also predominating.
As to the “ parr marks,” I have elsewhere noted that in the early
PARR AND SMOLTS 97
fry stage the sea-trout has these less clearly defined in outline than
either the salmon or the trout. The parr marks of the young sea-trout
hardly remain visible after the first year, but they always appear with
more or less definiteness upon the skin when the scales are scraped off,
even after the fish has become a smolt.
The sea-trout parr has also both crimson and dark spots, but the
dark spots vary greatly in number and appearance, and are far more
like those of young trout than those of young salmon. The sea-trout’s
greenish-yellow tinted dorsal fin is usually boldly spotted.
The colouring of the fins of the sea-trout varies very much but
generally the pectorals are yellow, tending sometimes to reddish-orange.
Sometimes they are quite dark and of a greenish-grey.
Structurally, as regards head and jaw, the sea-trout parr differs
from the salmon parr just as the young trout differs from it, and both
as regards the lesser size and higher set position of the eye the sea-trout
parr and the young trout are alike in differing from the salmon parr.
So too the fins and tail of the sea-trout parr are more akin to those of
the young trout, but perhaps as a rule the tail of the former is the more
forked.
Again, counting in the oblique line of scales formerly mentioned
the “ 14-scale formula ” obtains in the young sea-trout in contrast with
>
the “ro-scale formula” of the salmon.
On the whole it will be recognised that while the salmon parr and
sea-trout parr have something in common, in detail the sea-trout parr
approximates perhaps more closely to the young trout.
As to any differentiation between sea-trout parr and young trout,
account must be taken of Mr. Regan’s dictum that “there are no
structural differences, and the young are indistinguishable.” But that
there are some superficial distinctions there can be no question, although
these are never very constant in consequence of the endless variety of
marking and colouring that characterises both kinds of fish.
98 THE SEA-TROUT
Although not strikingly blue in the general tint of the body colour
as the salmon parr is, the sea-trout parr has a tinge of blue in its
colouring which, with the troutish yellow or brown that also enters into
its composition, results in a greenish-blue, or at least a hue that marks
it off from the purer yellow or brown of the trout. The distinction
becomes more apparent after the fish have been dead for some hours
and when the skin has dried.
I have never been able to detect in the spots a characteristic
difference between a young sea-trout and a young trout; nor is the
colouring of the fins in either fish always constant or distinctive.
As to structure, the sea-trout parr seems to retain longer than the
trout an immature appearance of slender delicacy. With two fish each
of eight or nine inches in length the trout will seem of more robust
build, and its pectoral fins and tail will appear stronger and less delicate
than those of the sea-trout parr. The tail of the latter certainly retains
its forked shape much longer than does that of the trout.
It has puzzled me not a little to compress within small compass, for
convenience of comparison and reference, these various distinctions,
but the following table may—subject to what has been written above—
serve the purpose :—
[DISTINCTIONS
PARR
AND SMOLTS
ee
DISTINCTIONS BETWEEN SALMON Parr, SEA-TROUT PARR AND YOUNG
TROUT.
LIMIT OF SIZE,
BODY COLOUR.
PARR MARKS.
BLACK SPOTS.
RED SPOTS.
PECTORALS.
ADIPOSE FIN.
SHAPE,
HEAD.
MOUTH.
TEETH.
EYE.
FINS.
TAIL.
SCALES.
ACTION
(Chaytor).
Salmon Parr,
Sea-trout Parr.
Seven inches.
Blue.
Very distinct.
One or two very intense on
cheek, small and few on
body above lateral line.
No halo.
Not distinctive.
Dusky pale.
Dark.
Slender.
Small, neat.
Small.
Not distinctive.
Large, and set low in head.
Delicate and pointed.
Forked.
“ to-scale formula.”’
Agile on capture.
Possibly ten inches.
Greenish-blue.
Less distinct.
Various and more or less
numerous. Halo not con-
stant.
Not distinctive.
Pale to yellow or dark.
Dark.
fringe.
Slender.
Frequent reddish
Longer.
Large relatively.
Not distinctive.
Relatively smaller, and set
higher in head.
Less delicate and more
pointed.
Less forked.
*“14-scale formula.”
Agile on capture.
Young Trout.
No criterion.
Yellowish-brown.
Very distinct when year-
ling. Traces only later.
Various and more or less
numerous. Halo usually
constant.
Not distinctive.
Yellowish-brown.
Reddish fringe.
Robust.
Larger and blunter.
Large relatively.
Not distinctive.
Relatively smaller, and set
higher in head.
Stronger and more rounded.
Less forked, and
rounded in outline.
“ 14-scale formula.”’
more
Quickly inert.
100 THE SEA-TROUT
It is probable that some one or other of the distinctions thus
tabulated will help the reader to discriminate in most waters between
salmon, sea-trout and trout when in the “ parr” stage of growth. It
remains to point out the special characteristics of the smolts of salmon
and sea-trout.
One cannot well speak of the smolt of the common trout because,
while individuals of the type known as Salmo fario seek the sea
and in doing so assume a silveriness of scale beyond the normal,
their migration is never so marked or so periodic as to constitute a
feature of their career. But it is curious to note that trout are commonly
more silvery in spring than at other seasons of the year. How far this
may suggest preparation for a possible migration, and how often that
migration may actually be effected it is not very easy to decide. In
circumstances where environment specially favours migration it may,
as we shall see, very commonly occur. I apprehend that if and when
such a descent does occur the descending trout will be indistinguishable
from descending sea-trout smolts. In this connection I submit a plate
of a Loch Lomond trout (Plate VI) in which the first figure shows the
complete silvery dress while the second figure (of the same fish) shows
the silvery scales partially removed. The scales of this fish, which was
caught towards the end of May 1915, showed that it had just commenced
feeding freely in its third year. Assuming my drawing to be approxi-
mately accurate it may readily be conceived that the difficulty of
distinguishing a Loch Lomond trout from a sea-trout is extreme, but
my own opinion, based very much upon the appearance of the exposed
skin surface that this fish was actually a trout, was confirmed at the time
by the opinion of several of the most experienced Luss boatmen, and
of the manager of Luss Hatchery.
The salmon smolt can hardly be mistaken. It is seldom less than
six and a half-inches in length and seldom more than seven and a half
inches. It is of the most brilliant silvery sheen, the general body colour
PLATE ail.
ee
Ne eNews ve
As gai
5 Ni A
Speen ARMAS
(4
ie
‘ errs
ee
Ni ny Sea oa) a. “5s
Nt . J - eon oe?
; % Lg g J . 4 ' 4 ae ~ wh
we
A SILVERY LOCH LOMOND TROUT
~ i
: PLATE VII.
P ’ ba i
ee et om
ay Se , ea (ARES ' ne?
A YOUNG SALMON ASSUMING THE SMOLT DRESS
PARR AND SMOLTS IOI
being of a pronounced blue on the back shading to pure white on the
under parts. The scales come off very freely and adhere to the hand
when the fish is grasped. The fins are delicate and the tail js deeply
forked. The tail is of a peculiarly rich black, the pectorals are a dusky
grey colour, and the dorsal fin is unspotted. The black spots on the
body are hardly discernible, but the distinctive spots on the cheeks
remain. The crimson spots may sometimes still be seen here and
there along the lateral line. Otherwise all the parr markings are
concealed beneath the smolt dress of silver.
The sea-trout smolt in the main resembles the salmon smolt, being
like it in general colour, blue-backed and white-bellied, with the same
glistening silvery scales easily detachable. The distinctions between
the two fish chiefly lie, first, in the relative size which in the sea-trout
may be as much as ten inches; second, in the colour of the sea-trout’s
fins—especially the pectorals—which are often of a rich yellow; thirdly,
its dorsal fin is usually spotted, and, fourthly, the black spots of the
body, and sometimes the red spots along the lateral line, are more
conspicuous than in the salmon smolt.
When the silvery scales are scraped off the distinctive parr markings
of both fish are clearly seen on the skin beneath. It may interest the
reader to see a coloured presentment, as I give it in Plate VII, of (1) a
young salmon in the transition stage between the parr and the smolt,
(2) the smolt in full sea-going dress, and (3) a smolt with the silvery
scales partially removed. The idea of this drawing is of course not
original. In Scrope’s “ Days and Nights of Salmon Fishing in the
Tweed ” (“ The Sportsman’s Library,” edited by the Right Hon. Sir
‘Herbert Maxwell, Bart.. 1898), there is a coloured frontispiece of a
salmon smolt with the silvery scales partially removed to disclose the
parr marks. Mr. Malloch, too, has adopted this method of showing
the smolt dress in some of his excellent photographic illustrations.
I think it is of interest to note that the more prominent of the black
H
102 THE SEA-TROUT
spots on the skin of the sea-trout parr are repeated on the silvery
surface of the sea-trout smolt as if they were endeavouring to shine
through, but they are not now so conspicuous because they have already
begun to be broken up into the X-shape which so often characterises
the spots of the mature fish.
In supplement of the foregoing notes upon the various distinctions
which characterise the young of salmon, sea-trout and trout, I may add
a table of the distinctions which have been noted by a Norwegian
observer, Mr. Hartig Hiiitfeldt-Kaas, as occurring between the parr
and smolts of salmon and trout, 7.e., sea-trout.
The following rough
translation has been kindly made for me by Mr. Hutton :—
SALMON.
The body is comparatively more
delicately and more pointedly shaped
with a long and slender tail.
The head is set lower on the body
and more finely shaped. At the first
glance the head seems to be exception-
ally long, but this is due to the fact
that the front portion (viz., the nose
and mouth) is distinctly shorter than
the trout’s, so that the gill-covers are
longer in proportion.
The eye is comparatively large and
is placed well forward in the head. A
perpendicular line drawn downwards
from the posterior margin of the pupil
will as a rule not touch the posterior
end of the maxillary bone.
The mouth is comparatively small,
and the length (measured to the
posterior edge of the maxillary bone)
is equal to about 13 diameters of the
eye.
All the fins (with the exception of
the adipose fin) are longer than the
trout’s.
The tail fin is distinctly V-shaped,
and the extremities of each lobe
are either pointed or very slightly
rounded,
TROUT.
The body is comparatively clumsy
with a short thick tail.
The head is comparatively larger
and more clumsily shaped than the
parr’s. The front part is much
longer, and the nose is longer and
deeper, and the lower jaw heavier.
Consequently the mouth is larger,
and therefore the maxillary bone is
distinctly longer than the parr’s.
The eye is distinctly smaller.
Although it is set further back in the
head, a perpendicular line drawn
downwards from the posterior edge of
the pupil will either cut or touch the
maxillary bone.
The mouth is large, and the length
(measured to the posterior edge of the
maxillary bone) is about 2} to 3 times
the diameter of the eye.
All the fins (with the exception of
the adipose fin) are shorter than the
parr’s.
The back of the tail fin when
stretched out forms either a straight
line or is only slightly concave, and
the lobes are broader and more
rounded.
—$<$—_—__—, 4th Winter
———_ 3rd Winter
Fig. 30.—Scale of a late-descending Sea-trout Smolt. (x 40.)
The fish had spent four winters in fresh water. It was caught in Loch Lomond
on 13th August, 1914. Weight, 4b. Length, 94in. Age, over 4 years.
See page 105.
’
:
*
, a
» . _
-
s
Mii bes ity =
i he 7
= : t
ay d ta oe 7 >
fe Bun oe ;
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Pe eu Wes) La ‘
PARR AND SMOLTS
SALMON.
The extremity of the pectoral fin
when stretched flat against the side
will reach a line drawn from the
anterior edge of the dorsal fin.
The adipose fin has no red colour.
The scales are larger and less in
number than in the case of the
trout. Ina slanting line between the
posterior edge of the adipose fin and
the medial line there are usually 11-12
and exceptionally 13 scales.
Before the migratory colouring is
assumed, the colouring of the parr is
similar to that of small beck trout,
with one row of large, distinctly red
spots along the medial line, and often
with some smaller red spots above and
below this line. There are also a
number of small dark spots spread
over the whole body above the medial
line, with a lesser number below.
What is especially characteristic of
the parr is the regular row of large
blue-grey blotches, even in breadth,
about 8-13 in number, and about 1 to
2 diameters of the eye wide, which
contrast remarkably with the white
colour of the under portion of the
body. They resemble the smudges
which would be made by fingers
smeared with blue-grey paint.
The smolt dress differs from the
parr colouring only in the fact that a
silvery sheen covers over the original
colouring, which latter shows through
the semi-transparent silvery coat, and
little by little the original colouring is
less and less visible, but so long as the
smolt remains in fresh water the parr
colouring can still be distinguished,
and especially in certain angles of
light.
There are usually from 2 to 4 black
spots on the gill-covers.
103
TROUT.
The pectoral fins when stretched
out will not reach a perpendicular line
drawn downwards from the anterior
edge of the dorsal fin.
The end of the adipose fin is usually
tinged with red or orange.
The scales are smaller and more
numerous, Ina slanting line drawn
between the posterior edge of the
adipose fin and the medial line there
are usually 14-16 scales and occasion-
ally more.
The colouring before the migratory
dress is assumed is more variable than
with the parr. In addition to the red
spots on the sides, which are usually
in three rows, one generally finds a
large number (exceptionally a few)
small dark spots scattered over the
back and sides of the fish, with a
relatively large number below the
medial line.
In place of the characteristic ‘‘finger
marks’’ of the parr, on the trout there
are fewer blotches and they are much
more irregular in shape.
As far as the migratory dress is
concerned (in which condition sea
trout are to be found in large numbers
and of the same size as salmon smolts)
the trout is of a silvery white colour,
but it retains unchanged all the small
black spots both above and below the
medial line, but not the red spots.
This is the most distinctive difference
between salmon and sea trout in the
smolt stage.
The black spots on the gill-covers
are usually more than 4 in number.
The following points should be specially noticed: the shape of the head,
the size of the eye and the mouth in comparison with one another, the number
104 THE SEA-TROUT
of scales, the shape of the tail, and the number and arrangement of the small
black spots: it is most exceptional to find on salmon any black spots below
the medial line behind the posterior edge of the back fin.}
The actual description of a sea-trout smolt which was taken from
the river Leven on 1st March, 1913, is as follows :—Length, over all,
93 inches, of which the head occupied 2 inches and the caudal rays
ti inches; girth, at dorsal fin, 44 inches; weight 6 ounces. Body
colour, a pronounced blue on the back shading towards the lateral line
to a pale blue and thence to clear white on the under. parts. Dark
cruciform spots closely speckled on the skin both above and below the
lateral line. The silvery scales, counted in the oblique line formerly
described, clearly numbered 14. The scales were easily removed and
beneath them 9 parr marks were faintly seen on each side of the fish.
The fins shaded from a pale yellow in the pectorals to a still paler yellow
in the ventral and anal fins. The dorsal fin was a dusky yellow closely
spotted with dark spots, and the dusky adipose fin had a reddish tinge
on its margin. The dusky tail was deeply forked. I am able to give
a coloured drawing (Plate VIII) of one of these exceptionally big
smolts. The specimen figured measured 9 inches in length.
I have seen in other districts ascending whitling which were scarcely
larger than the smolt figured, but Loch Lomond whitling, which are
noted for their exceptional size and quality could hardly be so small.
The fish was unmistakably a sea-trout smolt engaged in a leisurely
descent to the Clyde estuary.
The “run,” as it is called, of descending smolts begins intermit-
tently as early as February—in the Leven a sea-trout smolt has actually
been got as early as January—and gradually increases during March,
the main body descending towards the end of April, after which the
run again becomes intermittent in May and practically ceases in June.
But I have evidence that smolts may descend very much later in the
1. From “ Hyorledes adskilles lakse-érret-og roieyngel fra "hinanden,” 'by Hartig Hiiitfeldt-
Kaas. Christiania (1905),
LIOW INOYL“VAS JOUV ATIVNOILddDx NV
*
’
‘2 ”
* a ePe bbe 3 | Wx
“e
i.
TTA aLv1d
PARR AND SMOLTS 105
season. On August 13, 1914, I caught in Loch Lomond a fish which
was clearly a-sea-trout smolt on its way to the estuary. It measured
94 inches in length, was 44 inches in girth, and its scales came off freely
in handling. When sending some of the scales for examination to Mr.
Hutton I noted briefly :—“A very interesting fish as being apparently
a very late-running smolt. Four parr marks still visible towards the
tail on the scales being removed.” | reproduce a photograph of one
of the scales (Fig. 30), which indicates that the fish had completed four
winters’ residence in fresh water before the migratory instinct asserted
itself in its fifth year.
Weather conditions affect considerably the time of descent of the
main body, and where the stock is large and distributed throughout a
great stretch of country inland, the run will be proportionately pro-
longed. It is, I think, always prolonged where an extensive estuary
receives the inland waters through long tidal reaches. The descent
under these circumstances is a very leisurely affair, or it might be said
that once in brackish water the smolt is in no haste to go further afield.
It is a curious fact, worthy of special observation, that the descent
of the main body of sea-trout smolts in the river Leven always precedes
the main descent of the salmon smolts by about a fortnight, and I
understand, for I have made inquiry, that this order of progression to
the sea obtains elsewhere.
So far as I am aware, no investigations equivalent to those under-
taken by Mr. Knut Dahl in Norway have been attempted in Britain
with a view to determining when the young sea-trout first migrate to
salt water as sea-trout smolts, whether in the first, second, third, or
even some subsequent year of their existence. This appears to me to
be a vital question in the life-history of the fish and one deserving of
the fullest and most careful investigation in many rivers. JI have not
myself made any such extensive investigation, and can only submit
1. See later as to this subject, page 110.
106 THE SEA-TROUT
such very imperfect data as I possess bearing upon the point—imperfect
as being gathered in a restricted field. But my data, so far as they go,
suggest speculations which seem to me to be reasonable, though not a
little surprising, and all the more interesting that they once more bring
us into touch with the theory that there is only ene species of trout in
the British Isles.
I think, then, that it is very improbable that any young sea-trout will
in its first year—that is, during the year in which it is a “‘sea-trout fry’ —
descend to the estuary or the sea, nor is proof of such a descent any-
where to be found.
It is quite possible, however, in such rivers as the Tay, the Tweed
and the Leven, where the tidal influence is felt for a considerable
distance above the actual mouths of the rivers, and in waters such as
are found in the Orkneys and the Hebrides, which are hardly distin-
guishable as salt or fresh, that some of the young fish may, in their
‘
second year—that is, as “sea-trout parr’”—drop down towards salt
6
water though never adventuring, as true “ sea-trout smolts,” that year
into water which is markedly salt. Mr. Calderwood, in one passage
in “ The Life of the Salmon,” makes a suggestive remark in this
connection besides incidentally showing that he has some belief in the
“shoal theory,” which I have already propounded. As to some small
salmon fry being found near salt water in the lower reaches of the Tay
he hazards this explanation :—“ It seems probable, however, that other
young fry, following the instinct to form into shoals which we know
parr possess, have joined themselves to companies of larger parr, or it
may be have been to some extent washed down by floods.” Much
more is this descent probable in the case of the sea-trout, a more
characteristically estuarial fish than the salmon; but here again I can
give no proof of such a descent in the second year.
The earliest marked descent of the sea-trout smolt occurs in its
third year, that is, after it has spent two winters in fresh water After
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PARR AND SMOLTS 107
the turn of the year, then, and, if the winter has been mild and open,
all the sooner, the various shoals up-stream begin to pack in earnest
and work down towards estuary or sea. By April of this year the
youngest of these fish will have had two full years of life since ceasing
to be alevins. In hatchery parlance they are “ two year olds,” and it is
at this age the earliest true descent is made. In any event, it is only
on the fish assuming the sea-going silvery scales prior to its descent,
whensoever that descent may take place, that the young fish should
properly be designated a “Asea-trout smolt.” That the period when
the young sea-trout thus becomes a smolt varies considerably I shall
now proceed to show.
I mav make first this general observation that, with the Salmonide,
size has very little co-relationship with age. One is apt unthinkingly
to take it very much for granted that a large salmon, for instance, is an
old salmon and that a small trout is a young trout. How far this is
erroneous may be seen when we consider that a Tay salmon weighing
over 50 lb. may be only six years old, while an Add fish of to lb. may
quite well be two years older. Even more strikingly, a young salmon
may be larger than an older salmon taken from the same river owing to
the latter having been handicapped in its growth by having visited fresh
water more frequently for the purpose of spawning. Similarly with
regard to trout, the stunted little occupant of the moorland burn or hill
loch weighing but a few ounces may very well be an older fish than the
pampered occupant of some rich reservoir weighing as many pounds.
One has therefore no real warrant for supposing that the sea-trout
smolt which is seen descending a river at any time is only as old as it
looks. On the contrary there are reasonable grounds for the belief
that it may very well be much older. Even allowing that sea-trout
parr, as has been noted, grow proportionately quicker than salmon parr,
[ had often puzzled myself to account for the remarkable size attained
by some of the Loch Lomond and Leven sea-trout parr, specimens of
108 THE SEA-TROUT
which I have seen measuring 10% inches in length. In the same way
some of the Loch Lomond whitling appeared to me to attain excep-
tional growth. It had occurred to me that the question of age, as well
as questions of diet, might have some bearing upon the matter, but it
was not till the autumn of 1914 that the matter was made fairly clear.
Early in 1914 Mr. J. Arthur Hutton was kind enough to say that if
I would collect for him a series of scales taken from Loch Lomond
sea-trout he would devote some of his leisure to making a special study
of them, as the facts derived from such a study might help to throw
some light upon the various problems which would certainly arise in
course of the inquiry which I was undertaking. I need hardly say
how valuable such an offer in the circumstances appeared to me to be
and I gratefully availed myself of it, collecting as I did for Mr. Hutton
during the season of 1914 a series of scales taken from 45 sea-trout in
various stages of growth and in different states of physical condition.
Mr. Hutton was further so kind as to place at my disposal such data
on this subject as could be gleaned from other sets of sea-trout scales
in his possession. Notwithstanding that he has cautioned me against
rating his experience of the reading of sea-trout scales too highly, I
myself have no real doubts as to the substantial accuracy of his scale
interpretations, more especially as these are borne out by the results
obtained by Mr. Knut Dahl with respect to Norwegian fish.
On the particular point, then, as to the time of descent of the sea-
trout smolts as indicated by the scales, Mr. Hutton finds—and in his
company I| have confirmed his “‘readings”—that of the 45 Loch Lomond
fish none descended after only one winter spent in fresh water. Mr.
Hutton further informs me that of all the sea-trout scales he has
examined, including scales of fish from several rivers in Norway, from
the Wye, and from South Uist, he has not come upon a single instance
of a sea-trout having migrated to the sea after only one year’s fresh-
water life. The periods of actual descent, following Mr. Knut Dahl’s
method of tabulating the facts, may be shown thus :—
PARR AND SMOLTS 109
ScoTLAND.
Of these the Following had Completed previous to Migration
Loch Lomond Fish
ixamined j >. === Se i ee
2 Winters 3 Winters | 4 Winters | 5 Winters 6 Winters
|
45 7 30 | 7 | I O
Tabulating now in the same way the data drawn from Mr. Hutton’s
collection of sea-trout scales on this particular point, the results are :—
SCOTLAND.
Of these the Following had Completed previous to Migration
South Uist Fish eee Be Peak wee 2s SE ee ee
Examined
2 Winters 3 Winters | 4 Winters 5 Winters 6 Winters
| = - 2
6 I 5 fe) ) )
— —_
ENGLAND.
Of these the Following had Completed previous to Migration
River Wye Fish
Examined | =
| 2 Winters 3 Winters 4 Winters 5 Winters 6 Winters
17 | Gj 10 fo) fo) fe)
Norway.
Of these the Following had Completed previous to Migration
River Aa Fish
Examined
2 Winters 3 Winters 4 Winters 5 Winters 6 Winters
22 5 17 fe) fe) oO
Of these the Following had Completed previous to Migration
River Osen Fish | ¥ =
Examined |
2 Winters 3 Winters 4 Winters 5 Winters 6 Winters
25 I 19 4
110 THE SEA-TROUT
Before making further comments upon these results I venture to
submit for purposes of comparison with every proper acknowledgment,
the results obtained by Mr. Knut Dahl in Norwegian waters :—
Norway.
Of these the Following had Completed previous to Migration
Norwegian Fish
Examined
2 Winters 3 Winters 4 Winters | 5 Winters 6 Winters
192 25 78 67 | 21 | I
Accumulating all these results, as I think may instructively be done,
we obtain the following data :—
Of these the Following had Completed previous to Migration
Fish
Examined
2 Winters 3 Winters 4 Winters 5 Winters 6 Winters
307 46 159 78 23 I
In order that the reader may judge for himself of the value of this
evidence drawn from scale reading, I submit reproductions of photo-
graphs taken by Mr. Hutton of (1) scales of Loch Lomond sea-trout
indicating (a) a 2 winters’, (>) a 3 winters’, (c) a 4 winters’, and (d) a 5
winters’ residence in fresh water, and (2) to compare with the last of
these, because the reading is doubtful, the scale of a Norwegian sea-
trout indicating a 5 winters’ residence in fresh water, prior to the
migration of these fish to the sea as sea-trout smolts (Figs. 31, 32, 33,
35 and 36).
In this connection I may be allowed to repeat the scale of the
descending smolt (Fig. 34) described on page 105, which shows a 4
winters’ residence of the fish in fresh water prior to its descent to the
sea in its fifth year, in order that the reader may compare it with the
scale of the whitling (Fig. 33) which had spent 4 winters in fresh water
‘suvad ¢ qsouye ‘ety ‘ul Pe] ‘ysueyT “ql % ‘aq71I0My “FIGT
“TAQUIGAON 49% ‘Areqnq14 puowoy yoy e ut qygnRs sea “SUIpFIM B “YS ayy,
Cob x) ‘uonessiu 0) 101d Jaze ysodj
‘saved F JOAO 9I}QIT B ‘ASW ul t6 “YISuET qf ‘QYSIEM “FIGT “Gsneny YET Ul 9UAapPISal SIOJUIM-F B SuNvoIpul yNoI-eas jo IJLPIG—CE “S17
‘puowloT yooT ur 4ysneo sea ‘y[ous Surpueosep-aqyey e Ayyueredde ‘ysy oy
(oF x) ‘uonersiw oO} 101d 1a}eM YSetj UI aoUepIsal
SIUIA-F B SuNvoIpul YOWS yoi-vag jo ajeos—+t! “Bry
“91}09Q -
(
\
\
“TOPUIA\ 4ST —
‘eiquag
“JOJUTM pug -—
“IOJUTAA 8T
WV \\
\\\ \\
\
“JOJUI A pug
Nave
“19401 pie
“O4UIM 4p
“9[0Ws SB UOTFBISIW
“Q[0UIS 8B UOIZRIZITT
See page 110.
— |
PARR AND SMOLTS POL
before it descended as a smolt. The two scales are shown side by side
for convenience of reference.
Now it has to be admitted that the British data are too limited in
extent for any certain principles to be deduced from them, but I think
they prove conchusively that there is considerable variation of the time
of descent of the sea-trout smolts even in any one particular district or
river. They also suggest, if they do not actually prove, that the period
of descent of the majority of the smolts, in Scotland and England as
well as in Norway, is after three winters’ residence in fresh water.
One Loch Lomond instance also proves that a sea-trout may voluntarily
delay its descent till after its fifth winter spent in fresh water, while a
single instance given by Mr. Dahl also proves that in Norway the delay
may be protracted even for another year. There can be no question,
however, that the general principle is established from these data that
one has no right to assume as regards any river that all the sea-trout
smolts which descend to the sea in any particular year are the produce
of any one particular preceding spawning season.
It is an interesting speculation—and I think a legitimate specula-
tion—that the occurrence of a lake in any river system may encourage
the young fish to protract their stay in fresh water. The data concern-
ing Loch Lomond fish are at least not adverse to that assumption, and
I am informed by Mr. Hutton that two considerable lakes interrupt the
uniform channel of the Osen river in Norway. A wider investigation
may establish this theory which on the face of it does not seem to me
to be unreasonable. [ shall return to this point later.!
Some connection with what I have called Mr. Regan’s theory—
though of course it is not alone his—may, to speculate further on these
data, be traced in this way.
“Tf salmon smolts,” writes Mr. Calderwood, “ are retained in fresh
water beyond their natural time of descent to the sea, they assume again
the trout-like appearance of the juvenile.” This “natural time of
1. See post, page 144.
112 THE SEA-TROUT
descent”? of the salmon smolt, though not absolutely constant, is, in
Scotland, in the great majority of instances in the spring months after
two full years’ residence in fresh water. But if the young sea-trout
voluntarily remains in fresh water, as the scales seem to indicate, for
three, four, five, or (in Norway) actually six years, how much more will
ee
it be likely to retain a “ trout-like appearance”? To all seeming it is
quite content to remain a trout for an indefinite period, or, to put it
another way, the migratory habit takes a somewhat indefinite number of
years to declare itself. While this instinct or habit remains dormant,
the sea-trout is to all intents and purposes rather a miserable specimen
of a trout which will not develop generous proportions till it gains the
richer feeding grounds and more extended range of the sea or estuary,
just as the puny occupant of a mountain tarn will never develop in such
a habitat, but will, if removed to richer feeding grounds in a wider
range of water, at once begin to gain in bulk and appearance. Most
readers will be aware of some instance where such a result has followed
upon the transplanting of trout. For a curious example noted by Mr.
Knut Dahl I may refer the reader to pp. 77 and 78 and Fig. 33
(Plate X) of his work already noticed.
In this connection I think that the Howietoun experiments described
in my introductory chapter have some relevancy, and I would ask the
reader again to study the account given of them with some care. The
points material to the history of the sea-trout—and to Mr. Regan’s
theory—may for convenience be here again noted :—
1. That the Howietoun sea-trout parr lived in fresh-water quarters
for four years without any apparent discomfort;
2. That in their third year critical observers could not distinguish
them from common trout;
3. That in their fourth year the females inter-bred with Loch Leven
males;
PARR AND SMOLTS 113
4. That the resulting “ cross” breed became indistinguishable from
other trout fry; and
5. That the parent true sea-trout brood survived in a reservoir even
if they did not attain any great size.
I think on the whole it is a fair and reasonable conclusion to draw
from the facts of the foregoing experiments, supplemented as these
are by the evidence of the scales of wild fish which I have just
submitted, that the young of the sea-trout may without discomfort
postpone their descent to the sea for a period of years more or less
prolonged according to the nature and circumstances of their fresh-
water environment. To carry the matter still further, it does not seem
to me to be extravagant to imagine, if indeed it is the case that the
sea-trout brood is ever indistinguishable from the brood of the common
trout of the district, that some of the young fish never develop the
migratory habit at all, or, in other words, that they remain trout.
With the approach of the sea-trout smolt to the whitling stage the
facts definitely known about the fish become even less well defined.
No systematic examination of scales, or marking of sea-trout, have to
my knowledge yet been carried out, as they have been in the case of
the salmon, and there is no reason to believe that when they are so any
less interesting facts will be discovered than have been discovered with
reference to the life-history of the salmon.
I have already referred to the fry and parr of sea-trout maintaining
an original shoal formation, and I am inclined to think that, as these
shoals descend from the upper waters, whether they have packed
together in one great body or not, they still maintain their separate
individuality, just as at a review of troops battalions, companies and
platoons may mass together at one time and separate at another. It
might perhaps be difficult to adduce direct evidence of this, but if one
accepts the theory of shoal feeding at all, there seems no very good
reason why the shoal which foraged for food in the fry and parr stages
114 THE SEA-TROUT
of the fish’s existence should not continue its foraging in the sea or
estuary when the fish have become smolts and indeed thenceforward,
at least, as I shall suggest, until they have become whitling.
On reaching the brackish water of an estuary the sea-trout smolts
show no great anxiety to push far afield. In this way they differ
essentially from the salmon smolts which, it must be remembered, are
descending towards salt water at and about the same time, many of
each kind being actually in company. Mr. Calderwood and Mr. Knut
Dahl have, with some degree of definiteness, proved that, estuary or
no estuary, the salmon smolt makes directly for the sea and does not
linger for any length of time in brackish water—it makes a bolt for the
blue. I shall venture to state here what Mr. Calderwood discovered
concerning salmon smolts in course of a special netting expedition
which he conducted in the Tay in 1903, for some valuable facts
regarding sea-trout were ascertained at the same time. The object of
the expedition was to trace the descent of salmon smolts down the
estuary.
“From the time we left the neighbourhood of Kinfauns,” Mr.
Calderwood writes—the passage is from his valuable work on “ The
Life of the Salmon ”—“ the smolts became fewer, and when we had
descended about two miles and a half, and had reached a point a short
distance below the mouth of the river Earn, where sea-weed begins to
make its appearance upon the shore, smolts could not be found at all.
We proceeded down the estuary, however, and, thanks to the courtesy
of the Tay Fisheries Company, who granted the use of their steam
vacht, completed a survey of all available fishing places, both on the
shores of the lower estuary and on the shallow banks in mid-stream
near the Tay Bridge, till eventually we reached Budden Ness and the
open sea, some twenty miles below our starting point. Not another
salmon smolt did we catch, however, although sea-trout smolts were
everywhere in evidence, as well as brown trout, herrings, flounders, a
‘sawod Fo ‘easy ‘ur gy “QZuary ‘ql & ays
‘savad fg ‘SW “OF QUSIOM FIG ‘tequieydeg YyygT ‘puowoy yoory ur qysneo sem “Surya wv ‘ysy oy,
‘FI6T ‘AINE YG UO WIRES URISeMION & ul 4Yysne seM ‘Surya eB “Yysy ey, (‘ob x ) “UOT]eLSTW O07 Jowid AST EAS YSo.lj ul
Cor x) ‘uonersiw 0} 011d 19}eM Ysary Ul aouapisal SUSPISAT |SIOJUIM-S B GUIVOIPUL “INOII-BIG JO ajrog— SE -s1y
Sa} uIM-S ve Sunvoiput }NOI-VAS ULISaMION jO a[ePaG— 9 “Oy
POU Cle)
"IOUT AA ST -
“199UTA\ puz
qoqurM 43g ——————
“TOQUT AR I
“POWs SB MOTyeIZI JOIM 4b
“TOVUI AM 14S
“g/OWs SB UOTZRIAT TW
See page 110.
eas
ol
==
>
h
-
a
.
be
4 7 = — -
fase =
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Ni Fas A en
= - v i —
. = = ¥ <= 7G i
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= 5 7 r «
PARR AND SMOLTS 115
young turbot, sand eels, pipe fish, and various marine shore forms. In
this we repeated the experience of Mr. Dahl in his attempts to follow
salmon smolts from the rivers of Norway down the fjords.” Then in
course of renewed operations with a boom-net in the following year,
Mr. Calderwood made further discoveries of which one in particular
is relevant to the present subject, namely : “All salmon smolts captured
were taken when the net was set so as to fish the ebbing tide. When
the net was set so as to fish the flood tide or incoming current sea-trout
were alone taken, and many of these were nearly twice the size of the
salmon smolts. Sea-trout were also taken during ebb tide.”
The relatively larger size of the sea-trout was illustrated in my
description of a Leven smolt given on a previous page. But the main
point which Mr. Calderwood’s investigation brings out is that the sea-
trout is essentially an estuary fish. It may be presumed, therefore,
that in such great estuaries as those of the Tay and the Clyde there is
no occasion for the sea-trout to leave even the narrower waters of the
estuary in its transitional stage between the smolt and the whitling, nor
in such environment does the whitling do so before it again ascends to
fresh water, as one finds them there in all stages of the transition.
But there are few estuaries so extensive and so rich in feeding
grounds as those of Tay and Clyde. Many sea-trout streams have
practically no estuary at all, and between both types the variety of
river formation in Scotland is endless. On the type of estuary, then,
and the feeding grounds afforded by it, and where there is barely an
estuary or none, then on the available feeding grounds in the sea in the
vicinity, will, I think, depend in great measure the habits of the local
sea-trout. For this reason alone I think it would be unwarrantable to
lay down any scheme of definite uniformity in the habits of the fish
although most of these habits, I imagine, will be found to be not
inconsistent with certain broad general principles.
Whitling
a i
q
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)
“SulpUy AA peordAy w—Le +S
fr
&.
Oe le Siete ae: # 44 the iran yt
TOR oom be
9
Il
See page
CHAPTER VI.
WHITLING.
In brackish or salt water, then, the young fish, once they have
descended as smolts, whenever that may be, continue their existence.
While merging insensibly from smolts to whitling they finally lose all
traces of the parr state, their scales become more firmly set—but not so
much so as in the mature state—and, if the scales are now removed, no
parr marks will remain visible on the skin. The fins and tail become
stronger and the colouring of the upper body of the fish becomes much
darker—the small dark head giving origin to the Loch Lomond name
of “ black-neb” (z.e., black nose) for the fish at this stage. I insert a
drawing of a typical whitling (Fig. 37) which I caught at Luss on 20th
September, 1915. The fish weighed 3.1b. and was 124 ins. long, and
6} ins. in girth.
The richer quality of marine feeding makes for rapid growth and
the smolt of seven, eight or nine inches in length in about three months
becomes a whitling measuring twelve, thirteen or fourteen inches. In
three months, at least, they are ready to undertake their first return to
fresh water. One can fix this period of three months with exactitude,
for, just as the main body of smolts practically everywhere descends in
April and May, so the main body of whitlings in most rivers ascends
in July and August. So general is their habit in this respect that the
sea-trout of this small class are popularly called Lammas trout.
Confirmation of the time of return is also obtained from the scales,
and I could show several scales of whitling clearly illustrating it. But
I shall content myself with giving the reproduction (Fig. 38) of a scale
of a fish weighing 11} 1b. which I caught in Loch Lomond on 4th
September, 1914. I had noted it at the time of capture as “a beautiful
clean whitling.” The fresh-water residence in this instance is almost
119
120 THE SEA-TROUT
perfectly contrasted with the marine residence, while towards the
margin of the scale will be seen the closing in of the rings of growth
representing the beginning of what is termed the winter band. These
are the more apparent that the fish had ascended comparatively late in
the year, a fact which its considerable weight of 14 lb. helps to establish.
But just as some shoals of smolts descend earlier and later than the
main body in even normal seasons so some of the whitling return to
fresh water earlier and later than the main run. The run of the main
body itself varies in different localities. In the Solway rivers, such as
the Nith and Annan, it is a comparatively late run; in Loch Lomond
one expects the main run to occur by the second week of August;
and in the Tay, Mr. Malloch puts it as “ about the end of June.” As
he writes somewhat confusedly on this matter it is necessary to quote
what he says. “I am of opinion,” he writes, “that the yellow-fins do
not go far to sea before returning as whitling about the end of June,”
and then immediately after:—“ The yellow-fins, then, which return
about the end of June, have only been three months in the sea.” Now
what are we to understand by “ yellow-fins ” and what by “ whitling ” ?
I have taken pains in tracing the life-history of the sea-trout up to
the present point to use such nomenclature that no confusion need arise
in the reader’s mind at any moment as to what stage in the life of the
fish is being dealt with. Mr. Malloch, too, refers to the confusion that
may be caused through the fish being known under different names at
the various stages of its existence, but I cannot commend his own
practice or the recommendations which he makes in the interests of
uniformity. “It would be a very simple matter,” he writes, “to call
them sea-trout, in the grilse stage whitling, and in the smolt stage
yellow-fin,” and elsewhere he amplifies this thus :—‘“‘Salmo salar should
be called try, parr, smolt, salmon; foul salmon in the spawning season,
and kelt salmon after spawning. Salmo trutta should be called fry,
parr, yellow-fin, sea-trout; and if a further distinction is wished, gvi/se
WHITLING 121
could be called young salmon and whilling young sea-trout.’ But
apart from the curious inversion which | have ventured to italicise, and
‘ ’ )
his confusion between “ yellow-fin” and “ whitling” noted above, it
will be observed that he gives no distinctive name to the sea-trout
‘
earlier than “ yellow-fin,” so that he is driven to use such a cumbrous
_ phrase as “ parr of the yellow-fin”” to denote a “ sea-trout parr.” Not
b
only so, but “ yellow-fin” seems to be a futile term, for he himself
says :—“ although yellow-fin is the name applied to the smolt stage of
the sea-trout, it must not be understood all have yellow fins, for many
have their fins of quite as dark a colour as those of a salmon smolt.”
In my opinion the proper nomenclature for the sea-trout, from the
time of hatching, is :—‘sea-trout alevin,” until the umbilical sac is
absorbed; thereafter, “ sea-trout fry,” during the first year of residence
‘
in fresh water; “ sea-trout parr,’ during the second, or it may be third,
or even fourth year of residence in fresh water; and “ sea-trout smolt,”
when the silvery scales are actually assumed in descent to the sea.
“Whitling ” conveniently identifies the fish on its earliest return to
fresh water, and “ sea-trout”’ imports a fish that has reached maturity.
’
For the continued use of “ yellow-fin” there is neither necessity nor
authority.
39 66
The terms “fry,” “ parr,” and “ smolt”’ are used specifically in the
Salmon Acts, and as “ whitling ” was also so used at one time I would
‘ ’
retain that term in preference to or to such local
‘sea-trout grilse,’
terms as “herling,’ used in the Solway district; “finnock” or
“phinock,” used on Speyside; and “black-neb,” used at Loch Lomond.
For want of some precision in this matter long and acrimonious discus-
sion has persisted in many districts regarding the identity of the
sea-trout in its different stages, even after test decisions have been
pronounced in the law courts; so the matter is more important than the
casual reader might think.
The whitling of different localities vary in average size to a
122 THE SEA-TROUT
remarkable extent by the time they are ready to ascend to fresh water.
In some districts they are barely more than quarter of a pound in
weight, in others they weigh more than a pound, and I think this
variation is due rather to the quality of feeding they have formerly
experienced in fresh water as fry and parr than to their subsequent
feeding in the sea or estuary. Clearly also it may be due, as we have
seen, to some fish being in -fact older than others. As going to show
how extremely small some whitling may be I show (Fig. 39) the
scale of one which was caught in the Little Osen, Hoidalsfjord, by
Mr. Hutton. It weighed only 1 ounce. Yet this fish was a shade over
three years old. The reader may check the interpretation of this scale
from the scale of another Norwegian whitling here shown (Fig. 40).
The first of the three years’ residence in fresh water in this case is
clearly indicated by the rings of growth subsequently acquired being
disconform to those of the first year. This fish weighed 4 ounces. I
have heard of no Scottish whitling which weighed so little as the
Norwegian fish first here noted.
What instinct is it, one may ask, which induces the whitling to run
atall? The question, I believe, has never been satisfactorily answered.
Size alone appears to have very little to do with it, for in the same
district those that are late runners are generally, as one would expect,
rather larger than the early runners, but many of the last to ascend are
extremely small. Nor, apparently, has their condition of nourishment
much influence in the matter, for I have seen them ascend while in the
poorest condition. Nor can it be that they are following some special
prey up the river else that would easily be ascertained. Finally, their
ascent cannot be due to the spawning instinct, as it almost certainly is
in the case of grilse of salmon, for, as few out of the large numbers of
whitling that ascend can be seen spawning, it is ocularly demonstrable
that all whitling do not spawn on their first ascent as salmon grilse do.
The two cases are hardly parallel, however, for, while some whitling
Beginning of
winter band. |
Marine residence
about 3 months
3 years.
Fresh water residence
Fig. 38.—Scale of Sea-trout, illustrating period spent in the sea
after descent as a smolt and prior to return as a whitling. (* 39.)
The fish, a whitling, was caught in Loch Lomond on 4th September, 1914.
Weight, 1) lb. Length, 144 in. Age, 34 years. The fish probably descended as
a smolt in late May, had spent June, July and August in salt water,
and returned early in September.
See page 119
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JO quowaocejdsip ayy ajon
(ot x)
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(ob x)
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“JNO1}-PIS URIdaM
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See page 122.
WHITLING 123
return in the same year as their descent—within three months in fact—
grilse do not return until they have spent in addition a full autumn,
winter and spring in the sea. To describe the whitling as the “ grilse ”
of the sea-trout is thus a little misleading.
I can hazard no more plausible conjecture for the ascent than this,
that the sea-trout being of marked estuarial habits, the shoals of young
fish count it immaterial for their winter residence whether they range
upwards wholly into fresh water, in which to tell the truth they never
range very far, or downwards wholly into salt water, or remain hanging
about in the brackish water between the two, their preference being
determined at any time by the conditions which prevail in regard to
temperature and the volume of fresh water, perhaps, that affects these
conditions. It may also be, as Mr. Calderwood surmised, that the
younger fish in this matter simply follow the leadership of the maturer
shoals which are running up about the same time. It is seldom that
the whitling run anywhere in numbers until the Lammas floods have
broken.
Another question now presents itself. Do all the whitling of any
year so ascend to fresh water? I cannot believe that they do. I have
already suggested that from the time of hatching the tendency of the
young sea-trout is to remain together in shoal formation and that they
so remain together in the estuary. I would now carry this further and
suggest that the shoal still retains its identity in ascending the river,
but no longer the shoal in its entirety. Part only ascend, I think, and
part remain behind.
It has been proved to demonstration by Mr. Calderwood, who in
this matter confirms the views of Mr. H. W. Johnston, that the salmon
smolts which descend in any particular year do not return all together
_to spawn as grilse, but that only a proportion does so, the rest remaining
_1. There are a few isolated instances of salmon returning to fresh water in the same year as
their descént as smolts. Cf. Fishery Board for Scotland —Fisheries, Scotland, Salmon Fish, 1914,
IIT. and IV. (April, 1915).
124 THE SEA-TROUT
in the sea over winter to return, some of them, as maiden spring fish in
the following year. In pursuing further the same line of inquiry, Mr.
Johnston held, and Mr. Calderwood confirmed the fact, that, not only
do the lingerers not all return as maiden fish in the year subsequent to
that autumn in which the grilse spawned, but many of them delay
their return actually for four or five years, a proportion only of the
original run of smolts returning each season as maiden salmon for the
first. time.
Whatever be the real import of this provision of nature—of its
practical advantages to man there can be no question—it seems to me
to suggest the possibility of at least a partially similar habit in the
sea-trout. To be more precise, I think that the habit manifests itself
at this whitling stage, and that now a first great division of the shoal
occurs. In other words I think that a proportion only of each shoal
of smolts returns as whitling in the year of descent, though for some
reason not definitely known, or at least not definitely known to me, the
proportion varies from year to year in an extraordinary degree. It is
common to hear it said in a district that when any particular year is a
poor whitling year the next year will in consequence be a poor sea-trout
year. In my experience no reasoning can be more fallacious, for no
account is taken of the numbers of whitling which may have remained
over winter in the sea. In illustration I may point to the phenomenally
dry season of 1911 when, at Loch Lomond, there was an exceptionally
poor run of whitling. Yet in the more normal season of 1912 there
was a remarkably heavy run of sea-trout, and shoal after shoal of them,
fish of from 14 lb. to 24 lb. in weight, had all the appearance of the
missing whitling of 1911 now returned to fresh water for the first time
as mature sea-trout. I shall at a later stage show that scale examination
throws some light upon this question of the divided run.
As for my suggestion that the constituent members of the proportion
of the shoal which ascends remain together, I shall with great
WHITLING 125
difidence give my reasons for making it, and to elucidate these reasons
I would make the further suggestion that not only do sea-trout return
to their parent river but that the shoals which have been reared in Loch
Lomond and its spawning streams, and in similar waters in Scotland,
actually return to their special haunts where they were reared and fed
as fry and parr.
In Loch Lomond, then, one can trace the shoal of travelling whitling
by seeing individuals of the shoal leaping. They appear to head
directly for some particular spot on the shores of an island or the
mainland, and one’s inference is always later confirmed by finding that
spot, which had hitherto yielded no sport to the rod, now alive with fish.
Further, in comparing notes of an evening when the whitling are
entering the loch, the boatmen may find that banks widely separated
are thronged while intermediate and equally good ground is barren.
The local shoals have reached the first, but have not yet arrived at the
other ground. Yet in a day or two the intermediate ground has received
its local stock. It seems improbable that the fish, entering so vast a
sheet of water as Loch Lomond is, would make directly for a particular
quarter of it unless they had that quarter particularly in view. This is
more or less surmise in the case of the whitling, but I shall submit at
this stage instances of mature marked fish having returned to the same
trifling spawning burn. I discuss these instances later in another
connection, but meantime I may tabulate the returns of marked sea-
trout, to show their homing propensities, thus :—
340 B. Marked, Luss Water... ape ee ce OGE. ~ T9G4.
Recaptured, Luss Water oe fe nie) Ot: “1906
928 B. Marked, Arn Burn aie Pa ee ... Nov. 1906
Recaptured, Arn Burn ... aot << ... Nov. 1907
8352 B. Marked, Finlas Water ... ak mee Sea NOV.-1O13
Recaptured, Finlas Water (1st time)... Oct. Ord
Recaptured, Finlas Water (2nd time)... oe Oct. Igrs
126 THE SEA-TROUT
9331 B. Marked, Altnagairoch Burn... fe 7 NOV) Tons
Recaptured, Altnagairoch Burn se ... Nov. 1914
8328 B. Marked, Altnagairoch Burn... ~. Nov. 1918
Recaptured, Altnagairoch Burn (ist Pao ... Nov. 1914
Recaptured, Altnagairoch Burn (2nd time) .... Nov. 1915
8346 B. Marked, Smiddy Burn(?)* _ ... 2 =3. INOV= Org
Recaptured, Altnagairoch Burn (?)*... ... Nov. 1914
8347 B. Marked, Smiddy Burn ... abe ae 2 DNOV. LOIg
Recaptured, Smiddy Burn re a ... Nov. 1914
* There may easily be an error here in confusing the place of marking or
recaplure, as fish taken in both burns are often carried from one to the other
by the hatchery men. The two burns enter Luss Water within a stone’s
throw of each other.
In addition, two marked Loch Lomond fish have been recaptured
in nets in the Clyde estuary near the Leven mouth, which suggests that
the fish were “ homing” towards Loch Lomond; and another (besides
928 B) previously marked in the Arn Burn, was caught two years later
by an angler in the loch in the immediate vicinity of that burn. It
should be stated that no fish marked in one tributary (with the doubtful
exception noted above) has been recaptured in any other tributary.
If it be thus proved, then, that mature sea-trout return to Loch
Lomond in two consecutive years and even in three consecutive years,
and not only so but are found again in the same trifling tributaries (the
Altnagairoch, Smiddy and Arn Burns), as well as in the larger streams
(Luss and Finlas) in which they were marked, there is surely nothing
surprising in the fact that the shoals of ascending whitling should
return to the locality whence they descended as smolts. That each
shoal is actuated by an impulse to travel in the one direction convinces
me that it is one shoal returning to known ground and not a fortuitous
gathering of individuals travelling at haphazard towards the unknown.
Another thing which led me to form the conclusion that particular
shoals of sea-trout retain cohesion was this. At one time the river
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See page 131.
WHITLING 127
Leven was, if possible, even more polluted than it is now, and during
hot weather, in most seasons, whole shoals of whitling (besides mature
fish) were simply annihilated in their ascent. Now it was observable
in such disastrous years that certain feeding banks of the loch might
one year be destitute of fish and that during some other year other banks,
formerly fully populated, might prove barren. I have little doubt
that the explanation of this fluctuation simply was that the shoals of
whitling which ought to have peopled these banks were shoals which
had perished in the river Leven.
It may be objected that all this may be very plausible as regards
Loch Lomond but need not necessarily represent the habits of the fish
elsewhere. But I think on consideration it will be admitted that the
“entity of the shoal ”—if I may so term it where I have just referred
to and presupposed a first great cleavage—is a principle broad enough
to be generally applied elsewhere, and that the circumstances of Loch
Lomond are peculiar only in this that they give rather special oppor-
tunities for observation and deduction.
I have thought it well to leave as they were originally written the
foregoing passages with reference to the “homing” proclivities of
whitling, because it is interesting, to me at least, that the conclusions at
which I had arrived on the evidence submitted above have since been
found to be justified by incontrovertible fact.
In a Blue Book issued by the Fishery Board for Scotland towards
the end of 1914, the official reference to which is “ Fisheries, Scotland,
Salmon Fish., 1914, I and IT (October, 1914),” there appeared two
papers, one of which, by Mr. Ian T. Nelson, the proprietor of the
Glenetive deer forest, dealt with “ Hatching Results at Glen Etive.”
I may briefly state that with a view to developing the stock of salmon
and sea-trout in the river Etive, Argyllshire, Mr. Nelson constructed
an artificial loch in Glen Etive, with increased semi-artificial spawning
grounds in connection with it, and erected a small hatchery to
128 THE SEA-TROUT
supplement the natural spawning operations of the fish. In 1908, there
were turned out 87 yearling sea-trout which had been artificially hatched
and had subsequently been reared in a pond and artificially fed.
Before being liberated these fish were each marked in the dorsal fin
J
with a small piece of silver wire. ‘‘ In the main river in 1910,” writes
Mr. Nelson, “a sea-trout marked with silver wire was caught, which
could only be one of the 87 yearlings so marked in 1908.” This is the
first sea-trout with, so to speak, a completely “ artificial” record which
has apparently ever been recaptured on its return to the vicinity of its
birthplace. It is not with any intention of disparaging the valuable
outcome of Mr. Nelson’s enterprise if I add that it is only the first
instance of a fish being so recaptured because probably it was the first
carefully conducted experiment of the kind. There need not now be
the slightest doubt that such a return of the whitling, as I had already
deduced from observation, constantly occurs under natural conditions.
Still following the fortunes of the ascending section of the shoal of
whitling I think it may be accepted that the whitling when they do run,
whether in numbers or not, never ascend a river very far.' They
certainly never push on to the limits of the furthest tributaries with the
perseverance which characterises the mature fish when seriously making
for the spawning redds. Failing direct proof, this fact alone might be
held sufficient evidence that the main body of whitling does not ascend
to fresh water for the purpose of spawning. Mr. Malloch tells us that
“ The greater number prefer the tidal water, and in the Tay very few
are caught more than ten miles above this.” Where a great lake like
Loch Lomond is within easy run of the sea the whitling scatter through-
out its extent, but they do not ascend the tributary streams in any
numbers. In any case these whitling whether in river or loch, and
whether they propose to spawn or not, intend to remain in fresh water
over the winter months.
_1, It is fair to state that a correspondent of “The Field” once referred to one instance of a
whitling being caught, in some river which he did not specify by name, 40 miles from the sea.
WHITLING 129
Whitling feed ravenously, or at any rate they take a lure with
avidity, during their ascent which is generally when the water begins to
clear after a spate. Gaudy artificial flies and silver Devon minnows
are then the most attractive lures, but the truth is they will take readily
any kind of bait of which the earth-worm is not the least attractive.
When they have settled down in their quarters in river or loch and the
waters have become normal, they will still rise freely to the fly and take
other lures, but they are more capricious in their tastes then than when
fresh-run. At the same time they feed with more regularity than the
mature sea-trout do in fresh water, and if it cannot be said that they
increase in plumpness it is at least certain that they do not lose flesh,
and apparently they even grow in length, if slowly, after leaving the
salt water.
Although they soon lose their first brilliancy of scale, whitling do
not—or at least by far the greater number do not—assume the spawning
colours of the mature male fish, nor do the females darken so much.
One might count this as another argument against the universal
spawning of whitling. But I shall discuss this vexed question
immediately.
I may here describe the appearance of two of these young fish after
they had spent the winter months in Loch Lomond—I caught them when
trolling on 3rd March, 1913. They were to all intents and purposes
identical, both being females of three-quarters of a pound in weight
and each measuring 14 inches over all. They were fat, plump and in
good condition. The back, speckled as usual with black spots, was
in colour a dull green with a transparent watery look, the sides and
underparts being of a dingy white. One fish had been feeding freely
on caddis, the stomach and alimentary canal being distended with the
disintegrated cases, though near the gullet some of the large caddis
flies were in a fairly perfect state. The other whitling was literally
gorged with shell fish, the common physa fontinalis, and yet both these
130 THE SEA-TROUT
young fish attacked a natural minnow. In each the ovaries were
considerably developed and had already attained a length of some two
inches, though the egg pellets were yet very minute. Clearly, then,
they had not spawned (else their ova, if visible at all, would have been
hardly apparent), and the appearance of the ovaries rather suggested
that they would spawn for the first time in autumn when they had
re-ascended from the estuary in one of the later months.
I shall now submit such evidence as I possess for my belief that
some whitling at least do spawn on this their earliest and first return
to fresh water, and in this matter I may remark I am also indebted to
the valuable co-operation of Mr. Hutton.
While engaged collecting sea-trout eggs in the spawning season of
1914 for the purpose of stocking Luss Hatchery I kept a close watch
to see if any fish engaged in spawning had the general characteristic
appearance of a whitling. Out of 247 fish actually handled very few
were so like whitling as to be unquestionably whitling, but, on 26th
November, one of the men who knew what I was searching for, handed
me a fish from the net as to which, it might be said, there was no
superficial doubt whatever. It was a female fish, 13} inches in length,
bright and silvery, from which the scales came off readily in handling
and were scraped off with ease. The fish was ripe and when stripped
of her eggs weighed 8 ounces. The eggs had a curiously immature
appearance, they were so small; but the fish was obviously on the redds
for the purpose of spawning and the eggs were as readily and success-
fully fertilised as the larger eggs of larger fish. They were also
apparently of fair average number. I sent some of the scales to Mr.
Hutton for examination. His report, which I have no reason to
question, showed that the fish had spent four years’ residence in fresh
water and had been about three months in the sea. In other words it had
descended as a smolt in spring and was now, as a whitling, spawning
on its first return to fresh water in the same year. The reproduction of
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WHITLING 131
one of the scales of this fish (Fig. 41) in my opinion puts the fact of its
having spawned as a whitling beyond doubt.
I think it may fairly be assumed that others of the fish handled, as
to which the superficial appearances were not so convincingly charac-
teristic of whitling as the one selected, were whitling also. At any rate
Mr. Hutton pointed out to me that his reading of the scales of another
sea-trout which I had sent him, namely, those of a male fish weighing
1} lb. and measuring 18} inches in length, suggested that it too had
spawned as a whitling. He interprets the scale (Fig. 42) as showing,
subsequent to four years of fresh-water life, a three months’ residence
in the sea after which a distinct spawning mark occurs: another period
of residence in the sea, with a subsequent spawning mark; still
another period of residence in the sea, with the capture of the fish
occurring on its again visiting the redds—a comprehensive record for
so small a fish, but small, possibly, on account of its comprehensive
record. This, of course, was an old fish, but the record of its youthful
days was clear, and I think that other scales of younger fish like those
in the first instance cited, would often show equally an early spawning
record during the whitling stage. As a matter of fact out of the series
of 45 scales of Loch Lomond fish examined by Mr. Hutton, 5 show
evidence of the fish having spawned as whitling.
If the reader will further turn to Chapter VIII he will find the
record of two fish hatched in the same year, one of which (Fig. 43)
spawned as a whitling, while the other (Fig. 44) remained over winter
very probably in the sea. Confirmation is not easily obtained from
scale reading of a divided run, but at any rate both fish resided for the
same 3 years in fresh water, though one spawned as a whitling while the
other did not. IT ask the reader to accept such scanty evidence with
caution ; but, so far as it goes, it confirms the facts which I have other-
wise deduced.
I confess to thinking that any prolonged period of prior residence
132 THE SEA-TROUT
in fresh water may have a bearing on the period of spawning, it being
more probable that a fish which has spent three, four, or, it may be, five
years in fresh water will spawn as a whitling than one which has only
spent two years in fresh water, but I can submit no evidence upon which
any definite theory can be based. In considering this matter it is worth
while perhaps to keep in view that so experienced an observer as Mr.
Malloch states regarding whitling :—‘‘ My opinion is they do not
spawn.” Possibly the great majority of the smaller Tay whitling do
not.
It may be interjected here that the capture of these young fish in
spring, if they have not spawned, and of course similar fish everywhere
in Scotland, is strictly legal provided it is not during the close time.
They cannot, if they have not spawned, reasonably be held to fall
within the statutory prohibition against the capture of “unclean or
unseasonable salmon” contained in Section XX. of the Salmon
Fisheries (Scotland) Act 1868, because, not having spawned, they are
’ 2
not “unclean,” or kelts; nor are they “ unseasonable,” in the sense of
being caught within the close time which is the interpretation of that
term adopted by the English Law Courts. It should be pointed out,
¢ >
however, that the word ‘unseasonable” has. been several times
interpreted in the Sheriff Courts of Scotland as meaning, in the some-
what loose Scottish sense, “not very good according to time and
circumstance,” and therefore an ill-conditioned fish of this class caught
before regaining the sea might expose the captor to risk of a prosecu-
tion. I think the definition of an “unclean or unseasonable salmon,”
which I have elsewhere formerly given,’ is explicit enough for all
practical purposes, namely, “an unclean. or unseasonable salmon is a
~ salmon which is on the eve of spawning, is in the act of spawning, or has
not fully recovered from the effects of spawning.” But I think the
taking of these young fish at all at this early period of the year is to be
strongly deprecated, if for no other reason than that their capture simply
1. “ The Gentle Art,” p. 248. See also post p. 166.
WHITLING 133
leads by a speedy transition to the taking in spring of all sea-trout,
whether large or small, clean or kelt, without inquiry, to the detriment
of the stock of fish and to the grievous lowering of the standard of sport
in a locality.
In rivers uninterrupted by loch basins the shoals of these whitling
whether “clean” or “kelt,” work down rapidly to the estuary or sea,
the descent beginning more or less about the turn of the year and
continuing intermittently for some few months. But they seem loth to
leave good quarters in a loch and are only finally driven from it by, one
may suppose, the rising temperature of the water. In Loch Lomond,
I have observed the last of them has disappeared by the end of May.
But in their descent, as during their ascent, these greedy young fish eat
lustily and their diet, from flies to shell-fish, is as varied as the circum-
stances allow.
Assuming these whitling to descend in April, with this their second
visit to the sea, they are entering upon the mature stage of their existence
as “ sea-trout.”
It is time now to return to that portion of the shoal of whitling which
did not ascend to fresh water, and at the outset it is fair to say that my
views as to this first separation of the shoal into two portions are largely
theoretical, though I believe they will be found to meet the facts of the
case in most localities.
The richer marine feeding, which it is to be supposed this section
of the shoal will have obtained throughout autumn, winter and spring,
will have proportionately encouraged the growth of these whitling
beyond that of their contemporaries which migrated. They will also
be in far better condition in the spring months, and I think it is worth
considering whether these fish, which will in spring vary in weight
between 1 Ib. and 2 1b. do not form what is called the “ spring run”
of sea-trout in many districts.
>
I do not think that this “ spring run” is a true run of ascent, as is
J
134 THE SEA-TROUT
the Lammas run of whitling, because, as a lawyer would put it, the fish
has no animus remanendi and returns again to the estuary before the
true autumn ascent takes place. It would be more correctly described
as a tentative movement by these maiden fish to the upper tidal reaches,
and it is noteworthy that this spring run, so far as my observation gees,
is confined to waters where there are long tidal reaches or where there
is an immediate entrance from the sea into a loch which is almost on
sea-level. In any event, as I have shown that the whitling which did
ascend could not unless they had spawned be classed as kelts on their
descent to tidal water, so now these fish which did not ascend are not
kelts either. Hence in those districts where the sea-trout fishing is
practically tidal water fishing there is no reason why the spring months
should not furnish legitimate sport with sea-trout of this class, and I
think it is with such fish that spring fishing is got in the Beauly, the
Ythan and the Tay, and elsewhere.
I confess that I can make no suggestion which seems satisfactory
to myself why it is that this movement of the fish to the upper tidal
reaches, and even into some western lochs, like Loch Baa in Mull and
some of the South Uist lochs, takes place in spring any more than I can
definitely state why it is that some of the shoal made a more permanent
ascent as whitling in the autumn previously. I cannot believe that it
is due to the pursuit of any special food, and would be more inclined to
account for the movement by some change of temperature as between
tiver-water and sea-water.
Turning to another matter it would be extravagant to assert that the
separated wings of any original shoal re-unite on the return to the
estuary of the detachment which had migrated to fresh water. I think
it likely, however, that each wing retains its own corporate entity. At
such places where sea-trout rise to the fly in salt water, it is not unusual
for the angler to pick up shoal after shoal of the feeding fish as he
drifts over the ground. There seems no good reason in fact, though
WHITLING 135
of course assumption is not proof, why the shoals should get broken up
whether in migrating upstream, or migrating down stream, or roving in
the estuary. Whether the two wings of the shoal ever re-unite or not,
there can be no question that they are again ranging about contem-
poraneously feeding in the estuary or the sea.
The young sea-trout now find a rich harvest, for they are big enough
and swift enough in their movements to prey upon fish of considerable
size. Besides small shell-fish and sand-hoppers and other small deer,
shrimps and sand-eels form a large part of their dietary. But without
question their staple food in the early summer months is herring fry.
In May and June enormous quantities of these little fish—they are
about three inches long—approach the shores and myriads of them
enter our sea-lochs and push up the estuaries as high as the tide will
carry them. Sea-trout then feed upon them voraciously, and I am
inclined to the belief that, when the sea-trout which enter fresh water
are observed to be of poor quality in any season, the explanation is that
there has been a deficiency of herring fry in the shore waters during
that year. The numbers of these fry that a sea-trout will consume in
a day must be very great. I have found in one fish of about a pound
weight as many as eleven not wholly digested, besides a mass of scales
and bones in the anal canal. It looked as if this sea-trout, while
feeding, could, and did, swallow and digest fresh supplies as rapidly
as it could pass the unassimilated matter through its body.
There is good reason, beyond the mere growth of the fish, for this
extreme voracity, because the sea-trout has now to lay up within its
tissues a reserve of energy-making fat sufficient to meet not only the
exhausting journey to the upper waters, but the more exhaustive process
of the development of the milt and ova. There is at least no question
that if the sea-trout ascend to fresh water in this, the second summer
since they descended as smolts, they will proceed to spawn before
returning to the sea again for they have now reached maturity.
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CHAPTER VII.
MATURITY.
I have already indicated my opinion, though I do not pretend to
have conclusively proved, that all the whitling of any year do not ascend
together but that only a proportion of them does so. ‘The question has
now to be considered whether all, or only some, of the sea-trout which
have reached maturity ascend to spawn during the same season.
Mr. W. H. Armistead, in his book on “ Trout Waters—Manage-
ment and Angling,” tells us that all the mature trout of any lake ascend
the tributary streams each season to spawn. Personally, I should
prefer more exhaustive proof than Mr. Armistead offers of this fact
though one hesitates to doubt the conclusions of so well-known an
expert. On the other hand it is quite clear, as we have seen, that a
comparatively small proportion of the stock of salmon at any time in
the sea ascends to spawn, and it is not unreasonable to assume in the
mature sea-trout a similar habit. Definite proof of this matter is, so
far as I know, not yet available, but something may be learned
inferentially, and I shall pursue with that object the supposed move-
ments of those whitling (now grown to sea-trout) which did not migrate
to fresh water in the autumn of the year in which they descended as
smolts.
It seems appropriate to follow these fish first because clearly they
will have come earliest into condition to make the ascent owing to their
uninterrupted spell of rich marine feeding. In fact we do find in some
rivers a fairly early run of such sea-trout. In the Leven, for instance,
we find them ascending to Loch Lomond, but in no great numbers, as
early as the middle of April—fish of from 14 1b. to 24 1b. in weight.
In 1915, on April 3, a clean run sea-trout, weighing # lb., was caught
139
140 THE SEA-TROUT
with worm bait in Loch Lomond—a very unusual capture. In “ The
Glasgow Herald” angling report of 23rd May, 1913, it was noted that
“a fine sea-trout weighing 2} 1b. was caught in the Border Esk,” and
that “ This is the second sea-trout landed with the rod. The previous
one weighed 231lb. The opening catches of sea-trout usually consist
of fish of 14 lb. or 14 lb.” A Tweed report bears that on 15th February
1915, ‘“‘a spring sea-trout scaling 2 1b.” was caught at Carham; on
4th March in the same year, “a sea-trout, 14 lb.,” was reported from the
Logierait Hotel water on the Tay; while from the Beauly it was
reported in 1915 that on 8th March “ on the tidal waters an angler had
’
4 sea-trout,’ on 1oth March, 26 sea-trout, and on 12th March, 2 sea-
trout, but the weights of these were not reported, although one is aware
that these early fish in the Beauly are of a small class.
Whatever else these small sea-trout may be they cannot be those
whitling which ran the previous summer and descended in spring ready
now to run again. Nor is it likely that they are mature fish which,
having spawned in the preceding autumn and descended as kelts after
January, are now ready to ascend again. They can in fact be nothing
else than either, first, members of the detachment of whitling which did
not run in the previous year now grown to maturity, or second, possibly
fish which have spent two or more years in the sea without spawning.
Mr. Malloch gives an oddly confused account of the fish at the
stage which we are considering. “After the whitling goes to sea,” he
ce
writes, “it remains there for three or four months, and comes up as
a sea-trout from 1 to 24 |bs., according to the length of time it remains
in the sea. In the earliest rivers they begin to run in January, although
only in small numbers, and continue till October, but of course the
seasons have a great deal to do with their running early or late. The
Tay, I dare say, is the earliest river in Great Britain, and one would
expect them in it as early as in any river. Every spring paragraphs
appear in the public press stating that most of the sea-trout caught in
MATURITY 141
early spring are kelts. I have the numbers caught in our nets on the
Tay for the last eight years.” ‘Then in giving statistics he continues,
“On the 5th February of this year (1908) we caught more sea-trout at
one station—Almond Mouth—in one day than we had caught in a
month in some other seasons. Since then they have greatly increased,
and we are now (24th April, 1908) getting from 100 to 120 daily, all in
perfect condition, averaging 14 lbs. in weight. None of these have
spawned, although all will spawn this season.’ His final dictum is :—
“These sea-trout are now a little over three years of age, and are on
their second return from the sea.” With deference I cannot conceive
how with this explanation they can be on a second return from the sea.
How Mr. Malloch can hasten the descent of the whitling, bring them
into condition by a trip of “ three or four months” out to sea, and have
them ready to ascend the river again by 5th February it is a little difficult
to comprehend. If it had occurred to him that the whitling do not all
ascend to fresh water at once, he would have seen, | think, that those
early running fish so caught in the Tay were really members of that
body of whitling which elected to remain over winter in the sea, possibly,
as I have suggested, along with some fish which had remained for two
or more years in the sea without spawning, and were consequently fish
on a first return from the sea.
Assuming that I am correct in my surmise that many of them are
maiden sea-trout now on a first ascent more and more of these fish will
continue to run, some of them right up to the spawning season. When
that process is over, and another winter has come and gone, the fish as
kelts, will drop down to the estuary in the spring months, the last of
them disappearing from fresh water some time in May.
Meantime the detachment of whitling that did run and thereafter
returned to the estuary proceeds to recuperate its energies by marine
feeding, but there can be little doubt that the individuals which comprise
the detachment are not in condition to re-ascend, as mature fish to
142 THE SEA-TROUT
spawn, or, it may be (as we have seen) to spawn a second time, much
before July. Indeed I think that it is these fish which largely constitute
the great run of small sea-trout that in most sea-trout rivers occurs
at and after the middle of July. All those that ascend will spawn and
they too will descend as kelts in the ensuing spring.
I have traced—I trust clearly—the supposed ascent and subsequent
descent of members of the separated detachments, and jt only remains
to add that in their descent these fish will pass some of their contem-
poraries now ascending the river. They will, I think, first of all, pass
maiden fish on a first return since their descent as smolts; secondly,
they will pass some others on a second return, also maiden fish, as not
having spawned when they first ascended as whitling, and I think it
possible that those descending latest as kelts may even meet some
which, having spawned along with themselves and returned early to the
estuary, may have again come early into condition to make a second
spawning ascent. Should a kelt sea-trout, for instance, have returned
early in January, there is no reason why it should not be in condition
again by May until which time some of its companion kelts may have
lingered in fresh water before descending. Finally they will also meet
that class of fish which I supposed might have remained two or more
years in the sea without spawning.
I do not imagine, however, that this apparently complex, but really
very simple, running and counter-running takes place in any but the
largest of our river systems, and particularly in those where a loch or
a series of lochs induces a lengthy stay of the fish in fresh water. In
lesser streams the habits of the fish are generally more regular, I mean,
the runs of the mature fish are more marked—the first run is in July,
spawning takes place in November and the kelts descend in spring.
But that the whole stock of contemporaries anywhere ascends ex masse
to spawn in one year I do not believe for an instant.
That the period of continuous residence of sea-trout in the sea after
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But I give (Fig. 49) the scale of a fish which spent 3 winters in the sea
without spawning, and although 11 years old weighed only 24 1b: Its
record is :—
144 THE SEA-TROUT
Fresh-water life ... otis ss 3 years
Sea life without spawning ... 3 years \11 years=24 lb.
Sea life with spawning ... oo. 5, years
It would naturally be of great interest if scales of old fish could
be found which would show a wide difference in the migratory habits of
fish of the same season’s hatching, but one can easily understand that
the likelihood of securing such scales is not great because such fish
are relgtively fewer in number; nor are the scales of old fish very
easy to read. ‘There is no reason, however, it seems to me, why such
variation should not occur. From the set of scales upon which I have
so largely drawn for confirmation (so far as it goes) of my various views,
I have already cited (Fig. 48) one fish which apparently spent 3 years
in fresh water, thereafter 3 years in the sea without spawning, and 2
more years in each of which it spawned, the fish being caught in its
ninth year. As a contrast I cite later (Fig. 58) another fish which
apparently spent 2 years in fresh water, thereafter 1 year in the sea
without spawning and 5 more years in each of which it spawned, a total
of 8 years.
It is unnecessary to follow any further in detail the individuals of
the shoal whose supposed movements we have been endeavouring to
trace. The same process of division and sub-division will I imagine
continue annually till few of those originally hatched together survive.
As the same process is moreover going on with respect to each year’s
crop of smolts as these descend to the sea it is plain that the fish which
at any time are ascending will be of all sizes from the smallest to the
greatest.
It would be interesting if one could discover the precise bearing
which the presence or absence of a loch, or a series of lochs, has upon
the migratory habits of the local sea-trout. I am inclined to think that
this matter may ultimately be found to be the key to many of the
problems that puzzle one in the life-history of the fish. It is at least
suggestive that enormous numbers of sea-trout are found in those
MATURITY 145
districts in which lochs are common, districts such as the north-western
mainland and islands of Scotland, certain parts of Ireland, and almost
all the coasts of Norway. It seems reasonable to suppose that the
character of each loch, its accessibility from the sea, and its level
relatively to sea level, will help largely to influence the movements of
the fish which frequent it, so that the deduction of any general principles
must necessarily be extremely difficult. There is a broad resemblance,
however, between many of our loch basins which is worth adverting to.
The most common type of loch with which one is familiar in
Scotland is of course the “ glen-lake.” The origin of these glen-lakes
has occasioned much controversy amongst geologists, but it is enough to
note here in passing that the once popular theory of excavation by ice
action is not generally accepted nowadays. Be that as it may these
glen-lakes are of all sizes, ranging from inconsiderable tarns to great
inland seas. But their most interestine feature—so far as regards our
immediate subject—is that. owing to an alternate lowering and raising
of the general land surface the waters of some of these lochs, within
an almost measurable period of time, have become fresh instead of
salt. It is possible that the sea has so far encroached upon others as to
make their waters salt or brackish instead of fresh. At any rate it is
certain that the character of al] those that lie near sea-level has changed
within recent geological times, and is now in process of gradual change.
They are all in different Stages of transition. Some of them, in-the
West of Scotland, lie wholly submerged and open to the sea as in the
case of Loch Fyne and Loch Long. Others are so nearly at tidal level
that the tides flow freely in and out of them through a narrow passage,
albeit with much commotion as in the case of the Gareloch, or over a
rocky barrier which at low tide cuts off the loch from the sea by a
considerable fall, as happens at Loch Etive. Others again, like the
Dhu Loch, at Inverary, and some small lochs in the Hebrides—Lochs
Stenness and Harray in the Orkneys are of the same character though
146 THE SEA-TROUT
not glen-lakes—are so nearly beyond tidal influence that their waters
are barely more than brackish. And, lastly, of others near sea-level,
many of our great lakes, like Loch Shiel, Loch Maree, Loch Morar,
Loch Lomond and Loch Eck, are now wholly beyond tidal influence,
although the fact that sea-shells may be gathered in the clay of certain
beaches in a state of almost perfect preservation in some of them proves
that their fresh-water environment is of no great antiquity.
Now, whether we attribute to trout a marine or a fresh-water origin,
it is quite easy to imagine that the trout of any fresh-water loch to which
the sea gradually obtained access would adapt themselves to the
changing conditions of their environment and become in time, to all
intents and purposes, sea-trout. They would still, as such, continue
to migrate to their accustomed spawning streams. Similarly the sea-
trout of any sea-loch which gradually became elevated above tidal
influence would not for that reason necessarilv abandon their habits of
frequenting it, though these habits might possibly change to some
extent. For instance, the tentative run to certain lochs in spring may
be the survival of some ancient habit which existed when the loch still
retained its marine character. At all events the habit of seeking the
loch for the purpose of spawning in its streams would almost certainly
persist unchanged. It would, one may suppose, be the last habit to be
definitely abandoned. My belief is, that it is practically immaterial to
the trout of those districts, where such conditions as I have indicated
prevail, whether they inhabit at one time or another the fresh-water,
brackish, or salt-water loch, whichever it may be, which gives character
to the river channel, and that the runs of fish differ locally just because
of the variety of local conditions.
But particularly I would suggest that such tidal, semi-tidal, and
almost tidal lochs serve as convenient gathering grounds for the sea-
trout shoals preparatory to their periodical migrations. For example,
it is quite plain that Loch Lomond is the natural gathering ground for
MATURITY 147
the sea-trout of the Clyde district, because only through it have the fish
got convenient access to a considerable number of spawning streams in
an estuary where accessible spawning streams are remarkably scarce.
By way of contrast Loch Awe holds hardly any sea-trout, and this is
possibly because not it, but Loch Etive, is the natural gathering ground
of the fish. Loch Etive is a tidal loch in which the fish find convenient
spawning streams without being driven to the necessity of making a
long and adventurous ascent up the river Awe to the distant Loch Awe
spawning tributaries. One might multiply examples, but anglers will
be able to see, each for himself, how far this theory will apply to those
districts with which he is familiar.
A loch then, fresh, brackish or salt, may be the original and
hereditary habitat of the fish: it may, if salt or brackish, be still their
actual feeding ground: and it may, in any event, be the gathering
ground in which they assemble preparatory to spawning and to which
they return as kelts. If we further assume a strongly marked shoal
habit in the sea-trout—a habit which I think is beyond question—the
loch, if wholly fresh, will afford better than any river will a sanctuary
and nursery for the fry and parr until such time as they descend to the
estuary as smolts.
It follows that wherever a fresh-water loch, almost at sea-level and
within easy access of the sea. serves the fish of a district as both
sanctuary and gathering ground, one may there expect to find sea-trout
in great numbers, and that is precisely what one does find. Mr.
Calderwood has stated that in three seasons 33,000 sea-trout were netted
in the Echaig, the stream of no great size which drains Loch Eck to the
sea. The numbers of sea-trout which throng the Hebridean and
Orkney lochs are extraordinarily great, and of course such lochs as
Loch Maree, Loch Shiel and Loch Lomond attract anglers from all
quarters for the sea-trout fishing they afford.
In the Loch Lomond waters with which I.am most familiar the large
148 THE SEA-TROUT
class of sea-trout, that which averages about 4 1b., runs in June, and
the average weight becomes less as the season advances. Mr. Hutton
informs me that the same graduation of the runs occurs in Ireland and
in Norway, and I assume it is general. In a broad general view this
decrease in the average weight may be attributed, perhaps, not so much
to the gradual elimination of the larger fish as to the increasing influx
of the smaller whitling class.
I have not overlooked the run of heavy sea-trout which occurs in
our East Coast rivers, a run which once again opens out the “ bull trout”
controversy. The Tweed netting close time, and indeed the Aln close
time also, begins on September 15, and the evidence is fairly conclusive
that the major part of the run of these large fish occurs after the net
close season has begun, with the result that the nets take a compara-
tively small toll of them and their numbers tend to increase to the
prejudice of more marketable fish. I cannot explain why there should
be this late run of heavy fish, but it seems to me that it is another pointer
to the conclusion (suggested, as I have shown, by the scales) that there
is a species of sea-trout on the east coast that, whatever it may be
classified as, is not identical with the sea-trout of other waters. I have
in my introductory chapter suggested what that sea-trout may ultimately
be found to be.?
We have already seen that the sea-trout which spawn in October,
November and December do not delay their ascent from the sea until
these months. In districts where the spawning grounds are close to the
tidal reaches many may do so, but in most rivers the heaviest run of
sea-trout—if not the run of heaviest sea-trout—is a summer run. As
we have seen, too, some fish may actually in some districts have
ascended with the intention of remaining till the spawning season as
early as April or May, but such early runs at best are always inter-
mittent. In any case, it requires a considerable freshet in the river to
induce the fish at any time to leave salt water.
1. See also post, Chapter X.
MATURITY 149
If there has been a spell of dry weather prior to the usual time of
any run, in cases where a river enters directly on the sea without any
great intermediate stretch of tidal water, the sea-trout congregate in
numbers at the river mouth. One may see them gambolling and
leaping then on any calm evening manifestly showing the keenest
anxiety to enter the river proper, but few of them venturing to traverse
the dangerous shallows at the river mouth. At such times they fall an
easy prey to the splash-netter and, in unprotected districts, the shoals
are sadly decimated by unscrupulous persons even now in Scotland,
although of late years the officers of the Fishmongers Company of
London have done much to stop this illegal traffic.
Where the water is of the normal saltness of the sea off the mouths
of such streams, sea-trout seldom take freely any lure the angler can
offer them. But in many places, such as narrow channels or sea-lochs,
where the water is brackish, they will often take fly or “ minnow,”
although it is not easy to say why in one place they should prefer fly
and in another minnow and vice versa, as they often do. Even in the
sea an ordinary earth-worm is sometimes an effective bait, but, in truth,
the feeding habits of the fish—or at least the lures with which they may
be caught—vary so surprisingly in different localities that no rule can
be laid down for angling for them in salt water. They will seldom
anywhere take at all when actually waiting at a river mouth for the
weather to break. When the weather does break the fish eagerly run
up with the first of the spate and so closely crowded on each other are
they sometimes that the rushing noise of their passage up the shallows
can be distinctly heard above the sound of the rising waters.
In greater rivers, where there are tidal reaches, even if the river be
under its normal flow, a proportion of fish will always risk the ascent
at the top of each tide. They pass up to the fresh-water pools during
the darker hours, seldom running at all in daylight. Some of the best
K
150 THE SEA-TROUT
sport to be got with sea-trout is often to be found in such tidal reaches
for the fish as a rule, when they take at all, will take the fly freely.
The shoals for the most part fall back with the receding tide, only
a few braving the ascent, but when the river rises they no longer fall
back but boldly push upstream. In some places where a bridge
spanning the river gives opportunity for observation, hundreds may be
seen strung out in a long procession, maintaining a kind of formation,
it would seem, under an accepted leadership. Mr. Malloch, it is worth
noting here, makes a curious statement about the running habits of
sea-trout and salmon. “No matter what its size may be,” he writes,
“the sea-trout always runs in a zig-zag fashion—first to the one side
and then to the other. The salmon, on the other hand, runs in a
straight direction.” I cannot confirm this, although I do not pretend
to dispute the accuracy of Mr. Malloch’s observation. The fact seems
of no great importance and is hardly a practical method, one would
think, of distinguishing a salmon from a sea-trout in which connection
Mr. Malloch adduces it.
The sensitiveness—or is it instinct?—which enables sea-trout to
anticipate a coming change in the weather is, I think, well illustrated
by the following brief extracts taken from a diary which was kept by
the late Mr. Alfred Brown of Luss, for the year 1897.
“June 25. Fine weather continues. River and Loch very
low. Large quantity sea-trout entered the former on 22nd and
are lying in the pools in the meadow. This is supposed to
indicate a coming flood.
“June 26. The sea-trout noted above have returned to the
loch. No rain has come—dry—bright sun—north wind—very fine.
“June 27. The spate came. Sea-trout ascending the river in
great numbers. The fish have thus only been wrong by a few
days.”
In pushing on to spawning redds there are few obstacles which a
sea-trout will not surmount. They show the most astonishing skill in
MATURITY 151
negotiating rocky cataracts, and even the bare breast-work of mill-
dams, in low water. I have seen them wriggling successfully up the
pitching of the apron of a dam when only a trickle of water ran between
the stones; and in one river, where a sloping wooden shoot is placed for
their convenience, I have seen them throw themselves on to the
boarding when it was no more than wet and scuttle up it by hugging the
angle at the side. They run, however, in greatest numbers in a falling
water after a spate, and I think they prefer to run then in sunlight.
I cannot say what is the maximum perpendicular height that a sea-
trout will clear in its leap. Probably five feet is an outside limit, but
I fancy the powers of leaping of salmon and sea-trout are very similar
in this respect that they are both usually over-estimated.
Many districts have a natural fall or a mill-dam where the fish can
be seen leaping in numbers at certain times of the year, and such places
are always an attraction to visitors, for there is something intensely
interesting in seeing the efforts of the fish to surmount the obstacle.
I reproduce here a Photograph taken of a salmon leaping at a fall
(Fig. 50). Sea-trout leap in exactly the same way but the fish are too
small to be easily photographed. I think I have to thank Mr. H. W.
Johnston (indirectly through Mr. Hutton) for the photograph which
was taken, I understand, on the river Tummel. The Pot of Gartness,
a pool on the river Endrick, which flows westward through Stirlingshire
into the lower end of Loch Lomond, lies below a natural rocky
obstruction, and this forms one of the best-known “ salmon leaps” in
the West of Scotland. Sea-trout and salmon both ascend it, but only
sea-trout ascend the mill-dam on Luss Water where there is often a
remarkable display of the leaping fish, many being in the air at one
time along the whole perpendicular face of the dam while the broken
water below is seething with others. I have spent many hours watching
the fish thus leaping, and have not the least doubt that before making
a leap at the obstacle sea-trout carefully take their bearings. If
152 THE SEA-TROUT
watched closely for some time a particular fish will be seen poising
itself amid the foam of the pool, sometimes the head alone, but often
the head and pectoral fins, being out of water, facing the fall. Then
the fish disappears for a few seconds and one may expect it, and not
another fish, to be the next to leap. If it fails to surmount the fall and
is swept back into the pool, it will swim down to slacker water at the
tail of the pool where it rests for a little before making another effort.
It is quite easy to identify particular fish even when there are numbers
leaping. There is always some difference in size, shape, or colour
which distinguishes one from another.
It is an open question whether salmon or sea-trout are the more
persevering travellers to the upper waters. I think that if there is a
sufficient volume of water the salmon will push furthest up the main
stream; but where there is no steady flow the sea-trout only will attain
the higher reaches. In the greater river systems sea-trout very often
turn aside into the first suitable tributaries they come to. I have heard
the theory promulgated that just as sea-trout will not proceed far out
to sea so they prefer not to travel far inland, and the theorist placed his
inland limit at 30 miles. How much of truth there may be in this I am
not prepared to say, but I cannot conceive of there being any fixed
natural law in such a matter."
In normal states of the river and in ordinary daylight I think sea-
trout are not fish that care to expose themselves in open water. They
certainly do not love to lie in the sun as salmon seem to do. Rather
do they seek the bottom of the deepest pools, or burrow under the
overhanging banks, or hide below fringing trees. At nightfall only
do they venture upon the shallows, and then one can hear them in the
dark plunging in the quieter waters and splashing in the streams. In
1. Mr. Calderwood has kindly informed me that he has himself got two records of sea-trout
being captured out at sea, in the one case, 40 miles off Lowestoft, and in the other 45 miles off
Montrose. I have already (p.128) noted a vague reference to a whitling being taken 40 miles inland,
and may add that when I was at Hampton Bishop in August, 1915, a sea-trout weighing 43 lb. was
caught in the Wye by Mr. Hatton, Junr., of Hereford, 60 miles from the ocean—a somewhat
unusual capture as I was told.
See page 151
yer smth Age
MATURITY 15
ios)
many rivers, except during a flood or a half spate, night-time is the only
time when sea-trout can be caught, or at least it is the only time when a
good basket can be obtained. Even in a loch this preference of the
sea-trout for deep dark water is observable, for, while salmon may be
caught in a foot or two of water close to shore and trout on every
shallow, sea-trout, in such lochs as the conditions will allow of their
doing so, will remain outside in the deeper water, and will only rise
freely in depths of from eight to fifteen feet or so, unless in the later
part of the season as the spawning time approaches. When the bottom
is just lost sight of in a loch is usually the best fishing depth for sea-
trout, while “ between the shallow and the deep” is usually the best
fishing depth for salmon and trout.
With every recurring spate the sea-trout move higher and higher
upstream, and by the time the spawning period is reached many will
be on the redds, or in the main stream in close proximity to the small
tributaries in which the females prefer to deposit their ova.
In Scottish waters the sea-trout spawns on the whole earlier than
the salmon does, the difference in time roughly amounting to one
month. The sea-trout in this respect (supporting Mr. Regan’s theories)
approximates more closely to the trout. These seek the redds no
doubt rather earlier than the sea-trout, but in many localities suitable
weather and water conditions will find both kinds occupying the redds
at the same time. When this occurs the sea-trout, being the stronger
fish, ousts the trout from the more favourable positions on the redds.
This is another of those serious handicaps already referred to which
the trout has to accept in its struggle for existence with the sea-trout,
but I shall show later, when discussing the actual process of spawning,
that the trout suffers an even more vital handicap.
The actual months over which the spawning period of trout, sea-
trout and salmon extends in Scotland are September, October,
November, December and January, of which September, October and
154 THE SEA-TROUT
November are more particularly favoured by trout; October, November
and December by sea-trout; and November, December and January
by salmon. Viewing Scotland as a whole | think it will be found that
the middle month of each of these triple groups is the one during which
the great majority of each kind of fish respectively spawns, namely,
trout in October, sea-trout in November, and salmon in December.
The annual reports of the clerks of the District Fishery Boards
made to the Fishery Board for Scotland throw less light than is
desirable upon this subject, mainly because the Board did not until
very recently ask the clerks of the local boards to discriminate between
salmon and sea-trout, with the result that the dates of “ fish ” spawning
returned for many years are valueless. Although the term “fish” is
quite ambiguous few of the district clerks apparently ever thought of
differentiating between salmon and sea-trout in their returns. In the
blue book for season 1911, for instance, out of thirty-six reports
received by the Fishery Board only four districts differentiated between
the two species as regards the question when fish were first noticed
spawning, and only one (and that not even a statutory board) as regards
the remaining queries as to when the greatest number spawned, and
when spawning ceased. The Alness District Board puts the earliest
sea-trout spawning date as at 25th October; the Tweed District Board
puts it as at 22nd October; the Balgay District Board as at 18th
October; and the Voluntary Association, which supervises the Clyde
and Leven district (the district where there is no statutory board), as
at gth October. Only from the latter district, too, was the information
sent to the Fishery Board that the great majority of sea-trout spawned
in November. It seems almost incredible, in view of the Fishery
Board’s apparent desire for more precise data regarding sea-trout—and
having regard to the fact that nearly all the District Fishery Boards
note a falling off in the stock of that fish—that in the latest report (for
the year 1914) only one out of the 34 statutory district reports sent in to
MATURITY I
headquarters states definitely when sea-trout were first observed
spawning in that year. The Clerk of the Nairn District Board deserves
special mention for taking an intelligent interest in his work, and, it
might be added, for having a proper appreciation of his duties. As a
suitable statutory close time for sea-trout fishing, both by net and rod,
but more particularly by rod, can only be satisfactorily arrived at from
a close study of precise official statistics extending over a series of
years, it seems to me important that the queries now put by the Fishery
Board regarding sea-trout should be answered with some care and
exactitude by the clerks of the district boards to bring out this
information."
It might be appropriate to place on record here my view that the
respective close times fixed for salmon are wholly inapplicable to
sea-trout, though statutorily they apply to both species of fish. As I
have already stated the sea-trout is by about a month the earlier spawner,
and in my opinion sea-trout ought not to be permitted to be caught at
all in Scotland in October, or at least the close time ought to begin
concurrently with the Scottish trout close time on 15th October, and
should be universal, just as the trout close time is.
1. Mr. Calderwood, in his latest (1914) Report made to the Fishery Board for Scotland,
emphasises the importance of a more accurate study of the life-history of the sea-trout, and in
his report he has included what one may term a preliminary sketch of what are generally believed
to be the main facts of the sea-trout’s career. One may express the hope that this preliminary
sketch is but the forerunner of a series of studies of the fish from his pen based, as Mr.
Calderwood’s studies of the salmon have invariably been, upon an unchallengable mass of carefully
compiled statistics. The clerks of the various Boards can give him much assistance—if they will
The Spawning Period
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THE SPAWNING PERIOD.
Given a half flood, then, with perhaps a touch of frost, though that
is not essential, any day in November, or, indeed, any night—for if
conditions are otherwise suitable the fish seize their opportunity
irrespective of daylight—the ripe fish move up out of the pools to the
stretch of gravel that may have been selected, females and males
together.
By this time the fish, some of which may have been in fresh water
since May, are very unlike the beautiful silvery creatures which
ascended from the sea. I have heard them at this time pronounced
to be repulsive—even loathsome—in appearance by unsympathetic
observers. The males (Plate 1X) have wholly changed colour; instead
of a silvery skin daintily sprinkled with black cruciform spots, the bluish
darkness of the back contrasting finely with the gleaming white of the
belly, they have now a general yellow tinge mottled all over with
reddish brown blotches—I have seen some as yellow as ochre—and
the erstwhile silvery scales have sunk deep into their pockets under a
coating of mucus while fins and tail alike have become thick and
leathery. If the general outline of the body has not yet changed much
the head seems to have become elongated, and the lower jaw, which
normally projects but slightly beyond the upper, has developed a
hooked gristly protuberance like that of the male salmon at spawning
time, though in the sea-trout this protuberance is never so prominent.
For the sake of the contrast now adverted to I showed on a previous
page in rough diagram the outline of the head of a male salmon
(Fig. 13 (1)) and the head of a male sea-trout (Fig. 13 (2)) as the spawn-
ing season approaches. The female, still shapely enough if a little
159
160 THE SEA-TROUT
“bagged” in outline, is black but uncomely, for her scales have
darkened and all her gleaming white underparts have become dark.
In any ordinarily flowing stream the red male fish is a conspicuous
object at spawning time, but I have found it generally less easy to pick
out the dark grey female in the current. I am able to give here (Fig.
51) a reproduction of a very perfect photograph, taken by Mr. Hutton,
of the head of a 30 lb. Wye salmon which I caught at Hampton Bishop
on August 14, 1915, the fish from which diagram Fig. 13 (1) was drawn.
The projection of the lower jaw is admirably shown in the photograph,
and I| think the spawning livery of the male fish is as clearly indicated
as it can possibly be indicated without the assistance of colour. I also
show (Fig. 52) a male and female salmon on the eve of the spawning
season. With the male fish may be contrasted the male sea-trout which
I show (Fig. 53).
There is no doubt I think that some of the sea-trout which ascend
from the estuary each season are barren fish and will not spawn. On
September 29, 1915, I caught one with fly in Loch Lomond, weighing
11 lb., of such a silvery brightness that, concluding it to be fresh run,
I looked to see if it had any sea-lice or traces of them on it, but none
was found. Subsequent autopsy proved it to be a male fish with the
testes wholly undeveloped. The measurements of length, 15%ins.,
and girth, 8 ins., proved it to be in the pink of condition. It could not
have already spawned, therefore, and could hardly have spawned later.
The questions arise, first, whether the silveriness of its scales was due
to the fact of its barrenness, and, secondly, whether this is so far proof
that the change of colour of fish at spawning time is directly attributable
to the exercise of the spawning function. Examination of the scales
showed that the fish was not a whitling but a sea-trout, because the scale
growth indicated that a period of nearly two years had elapsed since
the descent of this fish to salt water as a smolt.
The process of spawning has often been described in the case of
*
eS
‘i
LEV Oey aes
oe ,
Fig. 52.—A female and male Salmon on the eve of spawning.
(ws
THE SPAWNING PERIOD 161
the salmon though not often with unimpeachable accuracy. I think the
best description of the process which I have read is contained in Mr.
A. H. Chaytor’s “ Letters to a Salmon Fisher’s Sons,” one of the best
books on salmon angling which has been published for many years.
At any rate sea-trout go through much the same procedure. The
gravel they select is, however, of finer quality than that in which salmon
prefer to spawn, the difference perhaps being fairly represented by
mixed gravel, the stones of which, in the sea-trout’s case, vary from the
size of a hazel-nut to ordinary road metal, and, in the salmon’s case,
from ordinary road metal to the size of a brick. Objects under water
naturally lose much of their specific gravity, but it is surprising notwith-
standing how large are the stones which the fish are able to displace.
Having selected a spot for the proposed redd the female sea-trout
throws herself on her side and, with a quick spasmodic action of the
body, rapidly repeated several times, works a hollow in the gravel into
which the ova as they are extruded, being heavier bulk for bulk than
water, are carried by the flow of the stream. Working her way slowly
upstream she covers the eggs thus deposited with the freshly loosened
gravel. The male fish for his part is closely in attendance shedding
his milt freely, and this, even when, as it must be, greatly diluted by the
stream, is able to fertilise all the eggs with which it comes in contact.
Yet the artificial propagation of salmon and sea-trout has shown that all
the ova at any time are not necessarily fertilised, and it is reasonable to
suppose that even when male fish are not scarce—as they sometimes
are—some of the ova shed under natural conditions escape fertilisation.
It is impossible to estimate the numbers of these, and probably the net
total fertilised varies enormously according as the spawning conditions
are favourable or the reverse. At all events any unfertilised eggs
ultimately become a dead white and soon perish.
On 28th October, 1913, I had a specially favourable opportunity of
seeing the sea-trout spawn in the Finlas Water, one of the tributary
162 THE SEA-TROUT
streams of Loch Lomond. I watched one female of about 6 lb. weight
within five paces of me. Before turning on her side she raised her
head and opened her mouth wide, then, almost doubling up, she lashed
out vigorously with her tail, and with a sort of burrowing motion
sideways scattered the gravel and raised a cloud of sediment which
almost concealed the male fish swinging to and fro in the current behind
her. After repeating the burrowing action several times she would
rest, then heading a little upstream she would begin again, the male
always being in attendance. The cleansing of the upturned gravel
from all sediment makes the redd easily distinguishable in the channel.
I am not certain that the female makes a hollow—I have seen it
described as a rudimentary nest—of set purpose, but a hollow is at any
rate the outcome of her spasmodic actions, and I imagine that a
considerable proportion of the ova finds a resting-place in the loosened
gravel behind her, to be effectively covered as the upstream gravel
is in turn displaced.
I was at some pains to find out how the stream acted on the ova.
From the floating leaves I gauged that the current ran at the rate of
nearly three miles an hour, and into a similar stream at more conve-
nience later I flung a quantity of ova. I was greatly struck with the
speed with which the eggs found each a resting place. One had the
idea that they would be carried initially a considerable distance down
stream, but this is not the case; they find bottom almost immediately.
The reason, I take it, is that the eggs are compact and materially heavier
than water and so sink at once. Then it is surprising to see how
effectual an “ eddy ” is caused by the slightest inequality of the bottom.
The surface may appear to be even-flowing, but under the lee of the
smallest pebble the water pauses for an instant, and behind every stone
an egg may come to rest. Once it is at rest it is also matter for surprise
how quickly the egg disappears from view. Its semi-transparency
makes it a very inconspicuous object in the water, but I think its
THE SPAWNING PERIOD 163
equal compressibility at every part enables it to adapt its shape to any
crevice, and so it is “ worked ” quickly by the motion of the stream
into any niche that will contain it.
It is erroneous to suppose that the male fish takes any part in the
formation of the redd. His active efforts are confined to protecting his
chosen companion from the attention of other males, and to dispersing
the small trout which hover near intent upon snapping up such stray
pellets of ova as come their way. It has been alleged that in spite of
his domestic anxiety there are grounds for suspicion that the male fish
is not averse from levying on his own account a food tax on the ova,
but in all fairness, I must admit that I have not myself seen him do it.
The male sea-trout does not remain to watch the redds as the male
salmon is said to do, but bolts for the shelter of deeper water as soon
as his labours are over,
It may occur to the reader that when trout, sea-trout and even
salmon are spawning in company, simultaneously and more or less
promiscuously (as may easily happen in some districts), a certain amount
of cross-breeding will inevitably take place. I do not doubt that such
cross-breeding does occur in nature although few hybrids can be
detected with any degree of certainty amongst mature fish. I think the
explanation probably is that, the three species being so closely inter-
related, the attributes of the hybrid become indistinguishable from the
general features of the species which most strongly predominates in the
cross.
For instance, if salmon ova be fertilised by trout or sea-trout milt,
or trout or sea-trout ova by salmon milt, I expect—as the salmon is the
stronger fish—that the progeny of the cross would be so like the progeny
of true salmon parentage as to be indistinguishable from true salmon.
As for trout and sea-trout, however they may be crossed inter se, there
can be little doubt (even apart from Mr. Regan’s theory) that the
progeny in any appropriate environment develop into migratory séa-
164 THE SEA-TROUT
trout. I can adduce very little definite evidence in support of these
presumptions, but I am aware that at Luss Hatchery on one occasion
the ova of certain female sea-trout were fertilised with the milt of male
salmon, there being no male sea-trout at the moment available for the
purpose. These ova thus fertilised took precisely the same time to
hatch out as uncrossed salmon ova deposited in the hatchery on the
same day, and to all appearance, up to the time of distributing them as
fry in the streams, the alevins were just the same in colour, and differed
as much in their general aspect from the alevins hatched from pure
sea-trout eggs, as did the uncrossed salmon alevins. The manager of
the hatchery bears me out in the supposition that any cross between
sea-trout and trout would, at any rate in Loch Lomond, result in a
migratory fish indistinguishable from a sea-trout. He often uses the
one kind to serve the other without anything but sea-trout fry apparently
resulting, and I am inclined to think, as trout and sea-trout commonly
spawn together in nature, that this is the main factor which results in
streams which are favourite spawning haunts of sea-trout holding but
few common trout, and these of a poor class.
The reader will make up his own mind in view of all that I have
written in these pages as to whether there is only one species of trout
in British waters or not. I can only point to this vital matter of inter-
breeding as being the strongest possible proof of trout and sea-trout
(excluding the “bull trout”’) being so closely related as to be to all
intents and purposes indistinguishable the one from the other.
The whole period of. spawning, which is an intermittent process in
the case of the sea-trout, as it is in the case of the salmon, cannot be
calculated with any degree of certainty either for the male or female
fish. Mr. Malloch puts it for the individual sea-trout at “ from two to
four days,” and for the individual salmon. at “ from three to fourteen
days,” and one need not question his accuracy of observation. But in
the spawning season of 1914 a ripe male sea-trout, whose milt was used
Fig. 53.—A male Sea-trout in Spawning dress.
(Note.—See Fishe ry Board label in dorsal fin.)
Fig. 54.—A female kelt Sea-trout.
1
and
7 : - S 7 =
THE SPAWNING PERIOD 165
on 9th November to fertilise the ova of some females, and which was
released after being marked with a Fishery Board label, was recaptured
on 26th November in the same stream, after an intervening period of
drought, when his milt was again used to fertilise the ova of other
females. This fish was thus engaged in active spawning operations
which extended (intermittently no doubt) over a period of 17 days. I
have no similar data in regard to female sea-trout, but I fancy from
watching them in the streams, that they shed their ova with some
regularity over a few consecutive days.
It may be worth while here to interject that in Scotland it is an
offence against the Salmon Acts to disturb any “salmon” spawn or
any spawning bed, unless for the purpose of artificial propagation or
certain other specified purposes."
Badly out of condition as many of the fish are by the time they come
to spawn it does not appear that the exhaustive effects of spawning have
a fatal result. The male sea-trout, as is the case with salmon, suffer
most, but few as compared with male salmon fail ultimately to regain
salt water. The female sea-trout suffer very little (Fig. 54). In
many years I have never seen, in the Loch Lomond district, a spent
sea-trout, either male or female, whose death could be attributed solely
to exhaustion following upon the spawning period, and after careful
inquiry made throughout a relatively large district such a case has never
been brought under my notice. I do not gather, either, from what I
know of other districts that such fatalities are anywhere very numerous
in the case of the sea-trout. The inference must on the whole be
allowed, I think, that only a very negligible proportion, in spite of great
emaciation of the fish, fails to survive the spawning period.
In this connection the following extract from the Report of the
Tweed Commissioners for 1915 is interesting :— The record of
salmon, grilse, and sea-trout found dead or dying from disease and
1. See note ante, p. 92.
L
166 THE SEA-TROUT
taken out of the river and buried from November 1914; to May 1915,
gave the following particulars :—
Spawned. Unspawned. Total.
Salmon _... Pe OL Se SF vee
Grilse ae Mer QO +a... Oo te
Sea-trout ... Me 9284. 5.2) 7) a eeee
1,045 82 1,127
The number for the year 1913-14 was 1,689, and for the previous year
2,343. Since 1879, when the annual returns of diseased fish were
instituted, the numbers of fish buried up to date were :—salmon,
111,782; grilse, 17,515; sea-trout, 26,565—total, 155,862.” These
figures of course seem very alarming, and may be hastily thought to
negative my views expressed above, but a known death-rate of 284
spent sea-trout out of the enormous numbers that ascend the Tweed
each year is really not a serious matter taking local circumstances into
account. It is probable that the river Tweed is responsible for the
highest death-rate amongst spent fish of any river, but it is a question
how far disease and pollution together accentuate the risks of the
spawning period. I am of opinion that the loss due to the exhaustion
consequent on spawning alone may in general in Scotland be said to
be practically nil in the case of the sea-trout.
I have already referred incidentally to the illegal capture of kelt
sea-trout (see page 132). The law, it may be repeated here, is quite
conclusive in Scotland regarding the capture of spent fish whether in
fresh water, in the estuary, or in the sea, and whether during close time
or in the open season. The law is, moreover, of general application
1. Every person who shall wilfully take, fish for, or attempt to take, or aid or assist in taking,
fishing for, or attempting to take, any unclean or unseasonable salmon, or who shall buy, sell, or
expose for sale, or have in his possession, any unclean or unseasonable salmon, shall be liable to a
penalty not exceeding five pounds in respect of each such fish taken, sold, or exposed for sale, or
in his possession, and shall forfeit every such fish; but this section shall not apply to any person
who takes such fish accidentally, and forthwith returns the same to the water with the least
possible injury, or to any person who takes or is in possession of such fish for artificial propagation
or scientific purposes.—Salmon Fisheries (Scotland) Act, 1868, § XX.
THE SPAWNING PERIOD 167
in Scotland, for, although at one time the Tweed Acts permitted the
capture of such fish, a clause of the general Salmon Act of 1868
specially abrogated that permission. '
In some streams where the stock of sea-trout is large relays of fish
will occupy the redds in succession, and where good spawning ground
is limited the gravel may be trenched again and again. It would not
be easy to estimate with accuracy how far the eggs first deposited might
suffer from subsequent disturbance, but displacement of the gravel
must expose many eggs and cause them to fall a prey to the natural
enemies which are always on the outlook for them. It is easy to
conceive of a case where the stock of fish may be too great for the
accommodation offered by the spawning streams, and, indeed (as an
example of this) I can point to the river Fruin, a tributary of Loch
Lomond, in the autumn of season 1912, which was the most productive
of sport of recent years. The same redds there, in that year, were
trenched three or four times over.
When the spawning operations are completed the sea-trout begin
slowly to drop down-stream but they are never in any great hurry to
change their quarters. In the short and steep streams of the West
Coast it is probable that a succession of floods will carry the fish
comparatively quickly to salt water, but in greater streams the sea-trout,
though gradually falling back, linger in each deep pool they come to.
A curious example of the sea-trout’s excessive timidity occurs to me in
this connection. In very dry summers, when the river Leven is too low
to permit of the working of the ferry boats, temporary wooden bridges
are thrown across the stream for the convenience of the artisans going
to and from the various works on the banks. During one very dry
spring these bridges had to be used at a time when multitudes of kelts
were descending. Now, if the sun happened to be shining, not a single
1. This Act shall not extend to England or Treland ; and no part of this Act. except the thirteenth.
eighteenth, twentieth, and twenty-third sections thereof, shall apply to the River Tweed as defined
by the Tweed Fisheries Act, 1859, &c. Salmon Fisheries (Scotland) Act 1868, § XX,
168 THE SEA-TROUT
fish would venture to pass the dark shadow cast by the first of these
bridges which hung so closely over their heads, and above it the fish
would lie in a mass waiting apparently till darkness came on. One
could see them dropping down to the shadow, but they would invariably
turn back upstream; none would face the mysterious obstruction.
During their descent as kelts, sea-trout take freely and impartially
‘
whatever lure the angler offers to them, fly, worm or “ minnow,” and
they may often be seen plunging at such natural flies as are on the
water, the March brown being a special favourite with them as it is with
salmon. It is often alleged that they make great havoc amongst the
smolts which are migrating to the sea about the same time that they
are descending the rivers. Judging from the voracity with which they
attack a spoon-bait or artificial minnow, it is reasonable to suppose that
many a smolt must fall a victim, but I am disposed to think that the
damage done by them in this respect has been much exaggerated. It
would be quite easy for a kelt sea-trout to gorge itself on smolts, but I
have never seen one so gorged, although I have seen them, when well-
mended in the estuary, literally gorged with herring fry.
I do not think that the sea-trout in their progress seaward rapidly
regain condition while in fresh water, for, except in one respect, they
“mend” but little. They do not put on flesh to any great extent, but
they assume a vivid brilliancy of silvery scale which even the ascending
“clean” fish can hardly rival. It is, however, meretricious splendour
at best—as someone has well expressed it the difference is like that
between tin and silver—and the “ well-mended kelt” a few hours after
death has a general limp look of washy flabbiness which betrays it.
If the kelt sea-trout thus linger in the river pools it is a natural
supposition that they will still further protract their descent if any
considerable loch has to be traversed in course of the journey. That
they do so all anglers in Loch Ness, Loch Awe, Loch Lomond, and
such great lakes are aware. It seems to require a specially warm April
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THE SPAWNING PERIOD 169
to induce them to leave such lochs at all. In the lower basin of Loch
Lomond they congregate in great shoals between the lowest island,
Inch Murrin, and the outflowing Leven. I have caught one in Loch
Lomond, a fish of about 4 lb. in weight, as late as May 24th, and in
Loch Ness I once caught one, a small fish, on June 13th.
When the kelts finally reach the estuary or the sea, they rapidly
begin to regain their best condition. I have been able to judge of their
progress in this respect from the improvement in appearance, as seen
from day to day, of those taken in the estuary sweep-nets of the Clyde.
I have no proof that any individual fish has been taken and returned
day after day until such time as its condition warranted its being killed,
although the netsmen aver that such fish can be recognised. But it is
generally obvious that the class of fish got in the nets changes gradually
each day for the better through the yield of the net containing each day
a higher percentage of kelts which have approached more nearly to the
condition of takable fish.
It would be difficult without specially collated evidence from many
rivers to pronounce upon a further question, namely, whether the mature
fish, once they have ascended to spawn, return again each year for that
purpose. I cannot find any reliable opinion on the point, and I do not
pretend that the scanty evidence from marking which I possess is
conclusive. I shall recapitulate the evidence in detail.
In October 1904, the men, when stripping fish in the neighbouring
streams to stock Luss Hatchery, marked twenty-one sea-trout with the
official labels supplied by the Fishery Board. In October 1905, as
many as nine of these marked fish were seen again in the streams, but
through an unfortunate misunderstanding the labels were not removed
from the fish for purposes of identification. So far the facts point to
a return in two consecutive years. But in season 1906 three more of
these twenty-one marked fish were recovered as tabulated thus :—
170 THE SEA-TROUT
Date. Weight. Length.
326 B. Marked, Luss Water... ... Oct. 1904 23 lb. 22 in.
Recaptured, Clyde nets ... July 1906 4 lb. (?)
336 B. Marked, Arn Burn... ... Oct. 1904 2 |b 20 in.
Recaptured, Loch, near do. Aug. 1906 32 lb. 22 in.
340B. Marked, Luss Water... Oct. 1904 1} lb. 16} in.
Recaptured, Luss Water ... Oct. 1906 33 |b. 203 in.
Several alternatives are here possible. These three fish may have
been amongst the nine seen but not identified in October 1905, in which
case, if they had been so, a return in three consecutive seasons would
have been established. Also they might conceivably have been others
of those marked returned in season 1905 without having been observed,
which would have equally meant a return in three consecutive years.
If, however, they were not any of the nine fish observed, or were not
others which had returned unobserved, then a return in alternate
seasons is the only inference.
Again, five other fish were marked in the Arn Burn, near Luss, in
November 1906, of which one was recaptured in the following year,
thus :-—
Date. Weight. Length.
928 B. Marked, Arn Burn ... ... Nov. 1906 1} Ib. 17 in.
Recaptured, Arn Burn... Noy. 1907 2 Ib. 22 in.
This instance proves a return in two consecutive years.
Further marking was accomplished in the spawning season of 1913
when, amongst others elsewhere, to fish were marked in the Finlas
Water. Of these, two reappeared in season 1914, as follows :—
Date. Weight. Length.
8349 B. Marked, Finlas ... ... 10Nov. 1913 33 lb. 221 in.
Recaptured, Clyde nets... 8 July 1914 54 |b. 254 in.
8352 B.- Marked, Finlas ... ... 10Nov. 1913 1} lb. 17 in.
Recaptured, Finlas ae 25 Octarong 23 Ib. 18} in.
These instances, the second conclusively, prove a return in two
4th spawning.
1914.
3rd spawning.
1913,
2nd spawning.
1912.
Ist spawning.
1911
1910.
1909,
Fig. 57.—Scale of Sea-trout showing spawning in 4 consecutive
years. (xX 30.)
The fish, a male, was taken on 30th November, 1914, when spawning in one of
the Loch Lomond tributaries. Weight, 24lb. Length, 205in. Age, 84 years.
When captured this fish was found to be one which had been marked with a
label when spawning in November, 1913.
See page 172.
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THE SPAWNING PERIOD 171
consecutive years, the first fish, being recaptured in the Clyde estuary,
might quite conceivably not have ascended the Leven to the loch.
Of 24 fish marked in the Altnagairoch Burn, a small tributary of
Luss Water, in 1913, two were recaptured :—
Date. Weight Length.
8331 B. Marked, Altnagairoch Burn Nov. 1913 2 Ib. 18 in.
Recap., Altnagairoch Burn Nov. 1914 23 Ib. 20} in.
8328 B. Marked, Altnagairoch Burn Nov. 1913 13 Ib. 15 in.
Recap., Altnagairoch Burn Nov. 1914 2 |b. 18} in.
while, again, of 16 fish marked in the Smiddy Burn’ in 1913, two were
recaptured also in 1914 :—
Date. Weight. Length.
8346 B. Marked, Smiddy Burn (?)! Nov. 1913 1} lb. 17 in.
Recap., Al’gairoch Burn ( ?) Nov. 1914 13 lb. 18} in.
8347 B. Marked, Smiddy Burn... Nov. 1913 2 |b. 19 in.
Recapt’d, Smiddy Burn ... Nov. 1914 24 Ib. 20} in.
all four of which prove returns in two consecutive years.
It is fair comment, I think, that if fish of varying weights are each
found spawning in two consecutive years, there is at least a presump-
tion that all may really spawn over more than two consecutive years.
But what ought perhaps properly to be called secondary proof is
furnished by the scales of some of the recovered marked fish and of
others unmarked. I shall examine first such scales as I have of marked
fish—accepting as correct, as I confidently do, the readings agreed upon
between Mr. Hutton and myself—as in their case the evidence obtained
from marking is so far placed beyond question.
1. In Fig. 55, I reproduce the scale of the sea-trout, a male fish,
which is referred to above as having been marked with label No.
8331 B in the Altnagairoch Burn, Luss, in November 1913, and
recovered in the same burn when spawning again in November 1914.
The photograph shows how greatly the scales of male fish become
1. See note in text, p. 126.
172 THE SEA-TROUT
eroded at spawning time. We have here, first, the known fact
of the fish spawning in 1914, which fact is represented by the frayed
edge of the scale, and second, the known fact of spawning having taken
place in 1913, represented on the scale by the dark band of broken
lines occurring first within the margin of the scale. Beyond that
nothing in directly known, but, tracing the life-history of the fish, Mr.
Hutton recognises three years (1909, 1910 and 1911) spent in fresh
water; descent as smolt early in 1912, the winter of 1912-1913 spent
in the sea without spawning, then the first known spawning in the
autumn of 1913, and the second known spawning in autumn 1914, when
the fish was caught. In this case the evidence is, we may say, indirect
that the fish did not spawn in its first year of sea-life, but direct that it
did spawn in the following two consecutive years.
2. In Fig. 56, I give a reproduction of a scale of the sea-trout, also
a male, which was marked No. 8346 B in one of the Luss burns, in
November 1913, and recaptured in November 1914, when the fish was
again spawning. ‘This scale also shows great erosion but the history
of the fish is fairly clear. It is of the same age as the preceding fish
but its record is different. This fish spent three years (1909, 1910 and
1911) in fresh water; descended as a smolt early in 1912, and, returning
the same year, as the scale indicates, it spawned as a whitling. There-
after it spawned again, as we know, in 1913, and a third time, as we
also know, in 1914. The evidence here, partly direct and partly
indirect, shows three consecutive years’ spawning.
3. In Fig. 57 is shown a scale of the fish, also a male, which was
marked No. 8347 B when spawning in the Smiddy Burn, Luss, in
November 1913, and recaptured when spawning again in November
1914. The earlier stages of growth, as indicated on this scale, are not
beyond question plain, but the indications of spawning are, I think,
undoubted. My supposition is that the fish spent three years (1906,
1907 and 1908) in fresh water and descended as a smolt in spring 1909.
iy
——— 5th spawning.
1914,
4th spawning.
1913.
- Srd spawning.
1912.
2nd spawning.
1911.
lst spawning.
1910.
1909.
1908.
1907.
Fig. 58.—Scale of Sea-trout showing spawning in 5 consecutive
years. (x 28.)
This fish, a male, was caught in Loch Lomond on 11th August, 1914. Weight,
431b. Length, 23in. Age, 74 years. The scale shows clearly four spawning
marks, and the fish was evidently on the eve of spawning for the fifth time.
See page 173.
eee OS bee re
fEeT Gay Qe
THE SPAWNING PERIOD 173
It spent the winter of 1909-1910 in the sea, or rather, I believe, in the
estuarial waters of the Leven, without spawning, and also the winter
1910-1911 in the sea without spawning. It spawned first in autumn
1911, a second time in 1912, a third time in 1913 as we know, and a
fourth time in 1914, as we also know. The evidence here, partly
direct and partly indirect, points to spawning in four consecutive years.
4. In Fig. 58 is shown the scale of an old cock fish weighing 43 Ib.,
which was caught in Loch Lomond by my friend, Sir James Hayes-
Sadler, on August 11, 1914. He very kindly gave me some scales
which Mr. Hutton has interpreted for me and one of which photo-
graphed by him is here reproduced. The scale indicates, I think
without question, that the fish spawned in five consecutive seasons, or
at least spawned in four and was on its way to spawn for the fifth time.
5. In Fig. 59, as showing an extreme case, I am able through the
courtesy of Mr. Hutton, to submit the scale of a sea-trout which
weighed 114 lb., caught by him in the river Osen, Norway. This scale
makes it fairly clear that the fish had spawned in seven consecutive
seasons. I reproduce a photograph (Fig. 60) of the fish itself as it is
a specially beautiful specimen of a large sea-trout.
Taking now the whole series of 45 sets of scales of Loch Lomond
fish I find that 16 show evidence of repeated spawning thus :—
In 2 consecutive years... a.) 5 dish.
In 3 * Gis;
In 4 ; . 3) toe
In 5 ; : 2
Examining Mr. Hutton’s sets of scales from other waters I find :—
(1) In regard to 6 South Uist fish (Scotland) evidence of repeated
Spawning in 2 instances, thus :—
In 2 consecutive years .., se, 2: fish:
(2) In regard to 17 Wye fish (England) evidence of repeated
Spawning in 2 instances, thus :—
In 3 consecutive years... ney 2) d1Sh-
9 THE SEA-TROUT
(3) In regard to 22 Aa fish (Norway) evidence of repeated spawning
in 8 instances, thus :—
In 2 consecutive years 5 fish
In 3 x) ” 2 bed
In 4 So Es s aye (Qu aee
(4) In regard to 25 Osen fish (Norway) evidence of repeated
spawning in I instance, thus :—
In 7 consecutive years... sm tu esis
‘
In all these sets of scales the ‘“‘ remainder” fish had either not yet
spawned or had only spawned once.
The point which I wish to bring out is, that, so far as this admittedly
limited collection of scales shows—and I would urge a cautious
acceptance of the results—in no single instance does it appear that,
having once spawned, a sea-trout fails to spawn in the ensuing season.
In other words sea-trout when they once begin to spawn would appear
to be annual spawners.
But it will be agreed that the matter of annual spawning is further
confirmed by two recent captures of marked fish. No. 8352 B, a male
fish, was marked in the Finlas Water on roth November, 1913. It was
recaptured on 26th October, 1914. Being remarked No. 8383 B and
returned to the river on that date, it was a second time recaptured on
25th October, 1915. Unfortunately it did not occur to the men to
keep scales of this interesting fish whose record I give in tabular form
thus :-—
Date. Weight. Length.
8352)B) ‘Marked’ Finlas ~ <.. 2% 10 Nov. 1913 14 Ib. 17 in.
Recap., Finlas (1st time) 26 Oct. 1914 2} lb. 18} in.
8383 B. Recap., Finlas (2nd time) 25 Oct. 1915 23 Ib. 20 in.
It may be added that this was the only re-marked Finlas fish, so that
its recapture was a somewhat lucky occurrence. An equally interesting
recapture was made a few days later. No. 8328 B was first marked in
the little Altnagairoch Burn, Luss, on 5th November 1913. It was
7th spawning
~ 6th spawning
5th spawning
4th spawning
3rd spawning
2nd spawning
Ist spawning
——
Fresh water
residence
Fig. 59.-Scale of Norwegian Sea-trout, indicating spawning in
7 consecutive years. (x 24.)
The fish, a female, was caught in the river Osen, on 18th July, 1914. Weight.
115 1b. Length, 293 im. Age, over 9 years.
Fig. 60.—A Norwegian Sea-trout (114 lb.).
THE SPAWNING PERIOD 175
recaptured on 9th November, 1914. Being re-marked No. 8399 B and
returned to the stream it was a second time recaptured on 9th November,
1915. Its career is tabulated thus :—
Date. Weight. Length.
8328 B. Marked, Altnagairoch... ... 5 Nov. 1913 13 Ib. 15 in.
Recap., Al’gairoch (ist time) 9 Nov. 1914 2 Ib. 18} in.
8399 B. Recap., Al’gairoch (2ndtime) — 9 Nov. 1915 24 lb. 193 in.
This fish was killed, and I have obtained a photograph of one of its
scales showing clearly the 1913 and 1914 spawning marks (Fig. 61).
It has been shown conclusively that what used to be called a “ Tay
bull trout” is nothing but a salmon which has already spawned returning
to spawn again in a subsequent season. As Mr. Malloch states,
“ Their flesh is often whiter and not so well flavoured” as that of a
salmon on its first return from the sea, and they fetch an inferior price
in the markets. So also it is common knowledge that large sea-trout,
which one may generally infer with safety to be old sea-trout, whether
we call them “ bull trout” or not, command a poor price in comparison
with smaller fish chiefly because the dealer expects their flesh to cut
whiter and to be much coarser than that of smaller and, as he presumes,
younger fish. I can attribute the paler colour of the flesh of particular
fish to no other cause than the frequency with which they have spawned
since coming to maturity. Hence [ would infer that once a sea-trout
has spawned it will continue to spawn yearly for a series of years.
It may be contended that what has been formerly stated regarding
the divided migration of the shoals argues against a return in consecu-
tive years; that if only a proportion of the stock of sea-trout ascends
each year then sea-trout cannot spawn each year, or, alternatively, that
if they do spawn each year, then no others than the small and more or
less immature fish can remain in salt water over the spawning months.
But it is now perfectly certain that salmon may delay their return from
170 THE SEA-TROUT
the sea, after having descended as smolts, for several years, and I see
no reason to suppose that the spawning instinct in the sea-trout should
not equally lie dormant for one or more years. The scale repeated
here as Fig. 62 indicates at any rate a three years’ residence in the sea
before the spawning instinct asserted itself, and the fact that, in spite
of spawning twice thereafter in successive years, the fish weighed 11 Ib.
points to a protracted and unbroken early period of rich feeding in the
sea. On the other hand I have discovered no scale, out of a not
inconsiderable number examined, which indicates that a fish which has
once spawned has thereafter passed one or more winters in the sea
without spawning. All I would venture to suggest, on admittedly
imperfect evidence, is that when the sea-trout has once commenced to
spawn it will return each year for that purpose during an indefinite,
because unknown, period of sexual activity.
ig aAO ‘Ady “UlZE “yysue'y “q] IT
‘QudIE\\ = FIBI “oun pagz uo syou Arenqsa apayTO eq} Ul 4ysnRO sem Yysy ey, eid oN We "
(@% x) “yJows e se uoner Siw sajye vas ay} Ul SIvAA BATINOASUOD L SBA JI p UES
uads yst 2 Sunes {nO1-PaS JO afvoC—zQ
re-arranged under water by means of a stiff feather. A small glass
a a
ARTIFICIAL PROPAGATION 195
> ito the tube any egg for closer examination. If the slightest trace
. of white fungus is observed upon it, the egg should be at once destroyed,
_ for the fungus growth develops very rapidly if not checked Careful
of all utensils, and careful examination of the eggs, are the best
preventatives against the disease appearing or spreading.
A thermometer, several syphon tubes, two or three clear glass
wine-glasses, some enamelled cups, a tiny gauze landing-net, and a
bunch of seagull wing feathers should be always at hand.
The only occasion for real anxiety at this period is during a spate,
when a large proportion of matter held in suspension in the water may
be expected to be deposited, if not on the eggs yet in the boxes. Of
course the best means to prevent this is careful filtration, first at the
source of supply, and again, if convenient, within the hatchery. But,
notwithstanding, the boxes will still require occasionally to be cleaned
while in service. This can be done without injury to the eggs, but, as
great care is necessary in the moving, cleaning with clear water and
replacing them, not to mention the cleaning of the boxes, it is a trouble-
some enough business. The danger of a deposit on the eggs is that it
prevents the free absorption of oxygen from the water through the shell
of the egg, and the free giving off of the carbonic acid gas generated
by the growth within, resulting in the suffocation and death of the
embryo. Periodical inspection should also be made at this time of the
intake chamber of the water supply, and the rose of the main pipe
should be examined to see that no floating matter has got in to foul it.
Decayed leaves are often troublesome in this respect.
As presumably is also the case in nature some eggs will escape
fertilisation in the process of artificial propagation. They will very
soon turn a dead white colour. It is impossible to give any definitely
196 THE SEA-TROUT
known percentage of loss, but one may expect more barren eggs from
an old fish than from one in the prime of life. Of the 2,080 eggs shed
by the fish described on page 73, only 10 were found to be dead or
barren after two days. But from one cause or another, during the
whole period of incubation an egg here and there will be found opaque
and whitish when it should at once be. destroyed. With careful manage-
ment the loss in eggs is wholly negligible.
With ordinary care the eggs will duly hatch out at the appointed time
when the young fish, or alevins, appear on the scene.
I do not propose to reiterate the description of the alevin formerly
given, but some points in hatchery management must be referred to.
In those boxes containing grilles it will be observed that, as the
eggs hatch, the alevins drop through between the glass tubes to the
bottom of the box, and the syphon tube will now be in constant use
removing the empty shells. When the eggs are all hatched, the glass
grilles, and the frames which carry them, and the longitudinal runners
on which the frames rest, are removed, thoroughly cleaned, and care-
fully put away. The hatching box has now become a small rearing
pond in which the alevins live comfortably until they reach the “ fry ”
stage. Before they leave the hatching box for the outer world, even
before the umbilical sac is wholly absorbed, they may be seen all lying
head upstream to the inflowing water, and “rising”? now and then
inquisitively to some tiny floating speck. As with the grille boxes so
with the basket boxes. When all the eggs are hatched, the baskets and
supporting runners are taken out, cleaned, and put away safely. It is
convenient to keep all the alevins of one mass of ova in their own basket
till the whole box has hatched out.
During the period of absorption of the umbilical sac, at least in the
earlier stages, the instinct of the alevin, which can only wriggle
spasmodically on its side, drives it to hide in any minute crevice that it
can find. Now it is, therefore, that previous careful smoothing of all
ARTIFICIAL PROPAGATION 197
angles and exact fitting of all fixed and movable parts of the box will be
seen to be of the utmost importance. For into every corner and crevice
the little creature will attempt to wriggle, and success, or partial success,
will almost certainly have fatal results. The best protection for them
at this time, beyond careful construction of the boxes, will be absolute
darkness in the hatchery.
Not much need be said by way of hint or warning now. Care and
cleanliness are still essential. Risks of spate water must still be run
and guarded against. But the main thing is to see that the alevins do
not find out some new and original method of committing suicide.
Hence every joint and junction must be examined in the boxes, and no
flaw in the intake or outflow perforated zinc screens must be permitted
to remain. So, too, any dead or injured alevins should be removed at
once, as should also all “ deformities,” of which there are always a few.
These I have described in an earlier page.
It is again necessary to contemplate a loss of stock up to the stage
when the alevin becomes a fry, but for obvious reasons the loss will
differ in every hatchery as so much will depend upon the personal
carefulness and skill of the hatchery manager, the excellence of his
appliances, and possibly the stamina of the brood. There need be little
loss in average circumstances.
Two questions now arise. The first is, where are our sea-trout fry
to be put? The second, how are they to be conveyed to the spot?
I have not contemplated for our supposed hatchery any supple-
mentary system of rearing ponds. These would doubtless be of
enormous advantage, both for the fish and for the education of the
owner (as many instances have shown, and are daily showing), but they
are costly to construct and the stock is costly to feed and protect. Nor
do I think that the practical gain would entirely compensate the outlay,
though if money were no object I would certainly recommend their
construction.
N
198 THE SEA-TROUT
Failing their construction, then, the fry must be placed in some
small stream, the preferable characteristics of which are that it does not
run dry in summer (as so many hill burns do); that it has a brisk clear
flow over gravel, and that it provides shelter and food from a fairly
luxuriant marginal growth. Wherever there is gravel, running water,
and marginal growth, the fry will find both security and food and may
safely be left to themselves. They should also be placed in a tributary
of a main stream rather than in the main stream itself; and in the feeder
of a fresh-water loch rather than in the loch. In both cases they will
seek the greater waters in their own good time.
It is said that great loss of stock occurs in the transition stage
between the curbed existence of the alevin and the free life of the fry.
I have already discussed this matter in Chapter III, and need not refer
further to it than to say that loss incurred at this stage has been put as
high as 50 per cent. But it is easy to see that with carelessness the loss
might equally be a total loss. © What is not known is whether loss occurs
with fish hatched and so far developed under wholly natural conditions.
Mr. J. J. Armistead states of trout hatching that “ It is well known that
the greatest loss that occurs amongst trout in our streams happens
whilst they are in their early stages. Probably some seventy-five per
cent. of the ova deposited by wild fish never hatch, and of the remaining
twenty-five per cent., more than half are lost during the ‘ alevin’ stage.
If, therefore, we can care for them so as to reduce that loss to some-
thing less than five per cent., it will be seen that an enormous advantage
is gained.” This statement applies equally to sea-trout hatching, but
I think the loss is rather nearer three than five per cent. at Luss. As
to loss incurred during the transition stage I can only say that none is
apparent while the fry are in the hatching boxes; and, although careful
watch has been kept after the fry have been placed in the streams which
for some little time thenceforward will be their home, no single dead
fry has been discovered, where if any had perished they might with
ARTIFICIAL PROPAGATION 199
practical certainty have been expected to be seen. But even assuming
that there is some loss I cannot see that the hatchery-bred stock should
suffer a greater loss than the wild stock, the whole conditions, so far as
one can judge, being in all essentials apparently identical. Where the
hatchery bred stock may suffer more is when hand feeding is resorted
to in the hatching box and subsequently in a rearing pond. I under-
stand that, until the little fish become accustomed to the food provided
many suffer from an inflammatory affection of the gills.
As to the means of transport it will be found that the fry may be
conveyed safely and conveniently to their future home in large glass
wicker-covered carboys, such as are used in chemical works. The
carboys should be full of water to prevent jarring in transit, and not
crowded to excess with fish. Should there be tisk of delay anywhere
en route, or the journey be long, fresh water should at intervals be
introduced. With ordinary luck the whole consignment will arrive at
and be transferred to the stream without mishap. We have found at
Luss that, if no great distance has to be covered, the fry can be quite
conveniently conveyed to neighbouring streams in ordinary galvanised
iron pails.
I may conclude by saying that, while these notes are based upon
actual experience of hatching out sea-trout ova at Luss Hatchery
during some eleven seasons, Mr. Walter McDiarmid, the very intelligent
and competent manager of the hatchery, was at first indebted for much
of the success of his operations to “A Handy Guide to Fish Culture,”
by Mr. J. J. Armistead, which was published in 1897, at the extremely
moderate price of one shilling by The Angler Limited. Scarborough.
Though dealing specially with trout this little Guide will be found to
be equally serviceable for the hatching of sea-trout, one more proof,
if any be required, of the close relationship, if not actual identity, of
the fish.
In submitting the Plan of the Hatchery, and the original specifica-
200 THE SEA-TROUT
tion and actual initial cost of the work done in construction in 1903, it
is only just to state that the whole hatchery arrangements were designed
by Mr. Thomas Duff, I.M., Glasgow, a member of the Loch Lomond
Angling Improvement Association, who ungrudgingly placed his skill
and much of his leisure at the disposal of the committee. The actual
initial outlay on the hatchery was as follows :—
[cost
tN
Nn
ARTIFICIAL PROPAGATION
Cost OF ARTIFICER’S WORK OF HATCHERY AT LUSs.
I. The Hatchery.
. Removing turf and preparing site @ gs. E
§ preparing 9
. 5” x2" Sawn standards and runners of walls (red pine), 440 lin.
ft. @ 2d. - - : 3 = 2
- 21 Do stakes in ground under sole runner each 24” long, to be
charred and well tarred, and the runner spiked to do. @ 5d. -
. 14” White pine tongued and grooved flooring in 63” breadths,
on outside of walls well tarred on outside, 76 sq. yds. @ 1s. gd.
. 14” Beads at angles, 37 lin. ft. @ 2d. - - - -
. 3” White pine half-checked sarking on inside, 68 sq. yds. @ 1s.
7. 63" 23” White pine spars of roof, checked to runners at top
and bottom, 162 lin. ft. @ 2d. - - - -
. 6"x2" Do. dwangs betwixt do., 34 lin. ft. @ 2d. -
. 3’ White pine half-checked sarking on do., include doubling
fillets at sides, 40 sq. yds. @ Is. - -
. Tarred felt on sarking in two thicknesses, 40 sq. yds. @ Is.
. Packing with sawdust space between flooring and sarking of
walls 5” thick, 69 sq. yds. @ 3d. - - - -
. 13” Fixed sashes with astragals glazed with sheet glass, and
primed and painted, 2 coats (red pine), 26 sq. ft. @ 8d.
13. 3” Chamfered checks to do. (—do.—), 34 lin. ft. @ 2d.
14. 2” Bevelled weather plate over sashes (—do.—) 8 lin. ft. @ 3d. -
15. 13” Framed and lined door (—do.—), 20 sq. ft. @ Is.
16. 3” Chamfered checks (—do.—), 16 lin. ft. @ 2d. - -
17. Do. facings (—do.—), 17 lin. ft. @ 2d. - - - -
18. 1 Pair 6” d. j. hinges and screws @ 6d. - - -
21.
22,
23.
Il. Hatching Boxes.
. 1 Galvanised iron 6” rim lock with brass mounting and four
keys @ 5s. : - - - - -
. Fitting and hanging and onputting mounting of 1 door @ 2s. -
with white lead.
14” Dressed deal bottoms of boxes in boards, 6! 9! x1! 7",
64sq. ft. @ 8d. - - - - - -
Do. sides and ends of do. 6” deep, 50 sq. ft. @ 8d. -
2” x2" Double checked teak framing of grille sliders, scalloped
on under edge 4o lin. ft. 6in. @ 8d.
oO
oO
oO
201
9 O
NG £4
8 9g
gl. (6)
(wr
8 Oo
7
5 8
Oo Oo
Oo Oo
Dire
Life <4
Ci
DEO
(a) {6}
2 8
2 10
Oo 6
5 Oo
2 oO
All dovetailed at corners and raggled and jointed
~
8
+
44.
45:
46.
. 8”x1" Plain do., do., 81lin.ft.@ id. - - - -
. 14”x 1” Checked teak frame of grilles, 160 lin. ft. @ 2d. -
. 24” x" Bevelled (pine) fillet at end of box, 9 lin. ft.6in. @ 1d.
27. Fine wire gauze 6” broad tacked on to box, g lin. ft. 6in. @ Is.
- 3
=
. 8 Solder joins of branch supplies @ 1s. - - -
. 1 Brass 3” screw down stop-cock @ 5s. - - : -
. 10 Brass 3” screw down nose-cocks @ 5s. 6d. - -
. 2 Bends on do. @ 2s. : i : : : :
. 2 Eyes on do. @ 2s. - : : - E
. 1 Y. junction piece @ 2s. - - - : 2
. Connecting drain with lead waste ©@ 2s. 6d. - -
. Charring timber of boxes - - = - 2 .
THE SEA-TROUT
oO OWN
wo onnn
Note.—Superseded by perforated zinc screen.
"x2" Mill dressed framing under boxes, 162 lin. ft. @ 1}d. - o
" Deal boxes for overflow, 14 sq. ft. 6in, @ 6d. - - O
Ill. Water Supply.
. 3” Lead 7 lb. supply pipe and laying in track where required,
66 lin. ft. @ 7d. - : Z : ; L :
wn © © F
. Forming connection of lead pipe to 3” iron pipe, include boring
and tapping and brass ferrule @ 17s. 6d. - - - oO
. 2” Lead 5 lb. waste pipe, include laying in track where required,
60 lin. ft. @ Is. - - = : = 3
. 4 Lead flanges in bottom of boxes @ gd. - - - Oo
. 4” Fireclay drain pipe, include digging track and filling up,
12 lin. yds. @ 2s. - : : : 2 4
(s) {e) {©}. (©) (2)
. Executing all jobbings and removing rubbish at completion
@ 10s. - : - - - . - - O
Fotal Contract Price £44
IV. Glass Grille System.
2,500 Glass tubes 62” x }” (carriage 2s. 6d.) per 100 @ 4s. -
5 8 g p 4 5
1,200 Glass tubes 62” x 4” (carriage 2s.) per 100 @ 3s. 8d.
to
V. Basket System.
Cost of perforated zinc baskets - - - : =
Total Original Cost £58
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‘CHAPTER X.
CONCLUDING REMARKS.
I po not think that I can usefully add anything to the foregoing pages.
The reader who has traced the life-history of the sea-trout as | have
given it in these chapters will, if he is an angler, be able to test the
description of each stage from facts which he has himself personally
observed. If he is a student of fish life as well as an angler—the one
interest does not always necessarily involve the other—he will be able
to recognise how far I have followed accepted beliefs and how far I
have departed from them. He will, I am afraid, find no great store of
facts and ideas that can pretend to be original or even novel. But
there is this to be said in extenuation, that while it is comparatively
easy to air original views in regard to the more or less abstruse problems
which one encounters in the study of fish life, it is not so easy to
substantiate them by facts derived from actual investigation. I have
tried as far as possible to support my theories with facts, and the reader
will judge how far these are to be accepted as evidence. In any case
it has been my care to distinguish between matters which are certain,
those which are uncertain, and those again which are simply theoretical
in the life-history of the sea-trout.
There is one matter, however, as to which [ think a final word may
here be added. However greatly we may extend our present know-
ledge regarding the details of the sea-trout’s career, it seems to me that
no real progress can be made until it is determined with certainty just
exactly what a sea-trout is.
My views upon this subject must inevitably lack the authority which
an exact scientific training, if I could claim such, would have given
them, but, for whatever value they have, they may be brought within
the scope of the following brief summary.
205
206 THE SEA-TROUT
WHAT Is A SEA-TROUT?
(1) Legally. According to the statutory law of Scotland, and indeed
of Great Britain, a sea-trout is a salmon, because the Acts declare that
the word “ salmon” shall mean and include “ salmon, grilse, sea-trout,
bull trout, smolts, parr, and other migratory fish of the salmon kind.”
In case any person may suppose that the definition is wide enough to
include ordinary trout within its meaning it may be stated that another
series of Scottish statutes, affecting only fresh-water fish, is conceived
to apply to
supposed that the sea-trout is not a bull trout in the eye of the law, but
‘
‘common trout (Salmo favio).’ Inferentially it may be
both are “ migratory fish of the salmon kind,” and both are “ salmon ”
in the sense of the Acts, though in this case things which are equal to
the same thing are not necessarily equal to one another.
(2) Scientifically. Whatever scheme of classification the reader
may see fit to follow he will of a certainty encounter in it the family
Salmonide. \n this family there is comprised, amongst other accepted
genera, the genus Sa/mo. Some doubt exists, however, as to the
number of species that should properly fall within this genus. Mr.
Regan, as we have seen, admits two, namely (1) Salmo salar, and (2)
Salmo trutta. | may say at once that no difficulty arises with regard to
Salmo salar, because, so far as investigation has yet been carried, the
Atlantic salmon is the sole representative of the species. Whatever
difficulty there is arises with regard to Salmo trutta.
It is conceivable, as I have shown, that a claim may reasonably be
made for the inclusion in the genus Salmo of three species, namely
(1) Salmo salar, (2) Salmo trutta, and (3) Salmo fario. This is not by
any means an original, or even a novel claim. There are many who
have so sub-divided the genus Sa/mo, but they have done so, or rather
many accept the classification, mainly on the ground that they seek to
differentiate between ¢vwt/a as a migratory trout, and favio as a non-
migratory trout. But Mr. Regan, very properly as I think, dismisses
CONCLUDING REMARKS 207
the migratory habit as a test of the trout’s nature, because it may be
broadly said that all trout are migratory in a greater or less degree.
The suggested classification is, however, worthy of support on other
grounds, but these involve the recognition of two distinct and separate
kinds of sea-trout.
Mr. Regan, as we have seen, admits in his classification only one
species of trout, Salmo trutta, to which fish he ascribed a marine origin.
In other words he presumes that the trout is a sea-trout.
I may interject here two definite statements of fact :—
(1) Common Ancestor. The common ancestor of the salmon and
trout has not yet been discovered.
(2) Original Habitat. It is not yet proved whether salmon and
trout, or one or other, have a marine or a fresh-water origin.
Mr. Regan, however, in company with other recognised authorities,
finds that two “races” of sea-trout are distinguishable, though with
difficulty. In much the same way Mr. Calderwood distinguishes two
“ varieties ” of sea-trout. I have already given, I hope impartially, the
views of those who are most competent to speak on this subject which
of course involves the identity, and therefore the proper classification,
of the “bull trout.” The reader may take it that the problem of the
bull trout is not yet definitely solved. Such definite light as I have
been able to throw upon the subject is of the faintest, but I think that
some grounds have been given for the belief that further systematic
scale examination will help to guide us to more definite conclusions.
The scales which I have examined (they could hardly be fewer in
number) suggest a rate of growth in a sea-trout typical of certain rivers
which is far more akin to the characteristic growth of the salmon than
it is to the growth of that other type of sea-trout which is perhaps most
familiarly recognised in our rivers. Taking this fact—if it be a fact—
along with other characteristic differences noted by various authorities,
the “bull trout’? would seem to me to be so distinct from that other
208 THE SEA-TROUT
type of sea-trout to which I| have referred as to entitle it to rank rather
as a species per se than as a mere race, or variety, of another species.
In other words the bull trout may fairly claim to be the species Salmo
“ruita of the genus Salmo.
What, then, is that other type of sea-trout? I think the foregoing
life-history suggests that the fish which we familiarly call a sea-trout is
no other than the common trout of these islands which has developed
the migratory habit to an extreme extent. Carrying the matter to its
necessary conclusion, if we recognise as a distinct species “ the common
trout (Salmo fario)” we shall find within the species certain trout which
do not migrate at all, certain other trout which haunt estuarial waters,
and still other trout which frankly migrate to and from the sea. Well
may it be said that the trout is a variable and plastic species, but it is
mainly so as the species Salmo fario and in relationship only to the
environment it elects to frequent.
These are then briefly the grounds on which I would suggest for the
genus Salmo the three species of (1) Salmo salar, with its sole repre-
sentative the salmon, (2) Salmo trutta, with, as its sole representative,
the “bull trout,” and (3) Salmo fario, with, as its representatives, trout
both migratory and non-migratory. In suggesting this classification. I
would warn the reader that the whole subject still remains very much
in the region of theory.
If besides being an angler the reader happens to be practically
interested in the development of either a trout or a sea-trout fishery, he
will recognise the importance of having this matter definitely elucidated,
and no doubt he will wonder, as I do, why the matter has not long ago
been cleared up. My hope is that these chapters will develop a more
active interest in the subject, because none of us at present can say with
any degree of assurance just precisely what a trout is, and what is a
sea-trout.
INDEX
A
Aborted (or rudimentary) fins, 53
Acts of Parliament referred to—
Act of 1318, fry of fishes, 9
Act of 1426, ‘‘ uthir fische,”’ 9
Act of 1469, ‘‘ fisch,’”’ 9
Salmon Fisheries (Scotland) Act, 1828,
“ fish of the salmon kind,’’ 9
Salmon Fisheries (Scotland) Act, 1844,
“fish of the salmon kind,” 9
Salmon Fisheries (Scotland) Act, 1862,
Statutory definition of ‘‘ salmon,”’ 9,
67 (note), 206
Salmon Fisheries (Scotland) Act, 1868,
eggs and spawn, 92, 165
fry, 92
kelts, 132, 166, 167 (note)
smolts, 92
Tweed Fisheries Act, 1859, kelts, 167
Adipose fin, 53, 54, 95
Age and Growth of Salmon and Trout in
Norway as shown by their Scales,
Knut Dahl, 92 (note), 108-110
Alevins, 74-76, 79, 80, 121, 196-198
salmon, 76
sea-trout, description, 75
distinctions, salmon, sea-trout and
trout tabulated, 76
eye; 75
hatching out, 74-76, 80, 196
mortality, 79, 197, 198
nomenclature, 121
trout, 76
Anadromous habit, char, 29
salmon and sea-trout, 24, 29
Anal fin, 53, 55
Angler-Naturalist, The, H. Cholmondeley-
Pennell, 16, 33, 56, 78
Armistead, J. J., authority on hatchery
work, 199
mortality of trout eggs, etc., 78, 198
Armistead, W.H., spawning trout, 139
Artificial propagation, 179-200
alevins, 196
basket system, 185, 186
collection of ova, 188-190
cost of hatchery, 201, 202
cross breeding, 15, 23, 36, 164
Artificial propagation, dead eggs, 195
disposal of fry, 197, 198
fertilisation, 190, 191
filling the boxes, 191, 192
fish handled, table showing data, 193
glass grille system, 184, 186
handling ova, 194
incubation, 194
Luss Hatchery described, 181-187
mortality of eggs, etc., 195-1098
nets, 189
output of boxes, 186
spate water, 195, 107
stripping fish, 190
transport of fry, 199
value of hatchery work, 179, 180
B
Bickerdyke, John, sea-trout fishing, 11
“* Black-nebs,”’ 119, 121
Book of the All-Round Angler, The, John
Bickerdyke, 11
Book on Angling, A, Francis Francis, m
Boulenger, George A., bull-trout, 40, 63;
Salmonide, original habitat, 31
Brachymystax, genus, 18
British and Irish Salmonide, Dr. Francis
Day, 15, 22, 32, 40, 60
British Fresh-water Fishes,
Houghton, 40
Bull Trout, Aln fish, scales, 65, 66
classification, 13, 17-19, 42, 206, 208
distinct species suggested, 42, 208
one species with sea-trout (Malloch), 16
“race” of S. trutta (Regan), 38
variety S. eriox of S. trutta (Calder-
wood), 40
Crown’s rights in Scotland, 7
gill-covers, 56, 62
“grey trout ’’ (Yarrell), 56
Hebridean, 25
late ascent, 64, 148
Norwegian, 66
Salmo eriox, 14, 25, 39, 40, 41
scales, 42, 64-66, 207
sea-trout, distinguished from, 38, 40, 42,
56, 61-67, 175, 207
Calderwood, 40, 207
Rev. W.
Bull Trout, sea-trout, flesh, 63, 175
Regan, 38, 62, 207
scales, 42, 64-66, 207
tail, 41, 42, 63
Walton, 41
Yarrell, 56, 62
size, 64
species discussed, 38-43, 61-67
statutory definition, 9, 67 (note), 206
tail, 41, 42, 63
Tweed fish, 63, 148
weight, 49
Bull Trout of the Tay and of Tweed, The,
W. L. Calderwood, 40 (note)
(e:
Calderwood, W. L., ‘‘ bull trout or round
tail,’”’ 40, 40 (note), 207
“common brown trout’? in Tay estuary,
25, 27, 34
New Zealand trout, 24
parr, juvenile appearance, 111
shoal habit, 106
Salmo eriox ‘ variety’ of S. trutta, 4o,
207
salmon, divided run, 123, 124
Salmonide, original habitat, 31, 32
Scottish Fishery Board reports,
155 (note)
sea-trout, caught at sea, 152 (note)
decrease, 43
netted in Echaig, 147
smolts, estuarial habits, 114, 115
“Tay bull trout,’’ 50
whitling migration, 123
Caudal fin, 42, 53, 54
Char, anadromous, 29
Chaytor, A. H., spawning salmon, 161
vigour of parr, 96
Cholmondeley-Pennell,
classification, 18
salmon and sea-trout, distinction, 56
trout, deformity, 78
trout and sea-trout, colouration, 33
Chondrostei, sub-class, 18
Classification, fishes, 12
Salmonide, Cholmondeley-Pennell, 18
Regan, 18, 19
suggested, 42, 206-208
Close seasons, for sea-trout criticised, 155
Common ancestor, of Salmonid@, 32, 207
Compleat Angler, The, Izaak Walton, 36, 41
Coregonus, genus, 19
“ Corrigeen,’’ decrease of sea-trout, 43
Il
H., Salmonide,
INDEX
Cross-breeding, artificial, 23, 164
Darwin, 36
Day, 15
natural, 163
Walton, 36
D
Dahl, Knut, Salmonide, growth of, 92
smolts, estuarial habits, 114
migration, 105
scale evidence, tabulated, 110
trout, rapid development, 112
Darwin, Charles, cross-breeding, 36
Day, Dr. Francis, bull trout, 40, 49 (note)
Howietoun experiments, 22
Salmonide@, original habitat, 32
teeth, 60
trout, varieties, 15, 42
Days and Nights of Salmon Fishing in the
Tweed, William Scrope, tor
Deformities, trout, 42
sea-trout, 77, 78, 197
District Fishery Boards in Scotland, annual
reports defective, 154, 155
data desirable, 155 (note)
sea-trout returns, 10
Dorsal fin, 50, 53, 54, 95
E
Eggs, 71-76, 78-80, 130, 162, 179-200
salmon, 71-74, 76, 192, 194
sea-trout, artificial propagation (q.v.),
179-200
colour, 71, 72, 195
development, 74, 75, 194
tabulated, 80
distinctions, salmon, sea-trout, and
trout, tabulated, 76
“eyed ova,’’ 74, 194
mortality, hatchery, 195, 196
natural, 79
trout (Armistead), 198
number, 72-74, 192
shape, 72
size, 71
weight, 162
whitling, 130
trout, 71, 72, 74-76, 78, 79, 192, 194, 198
Encyclopedia Britannica, Art. Ichthyology,
12
Estuarine trout, 24-27
““ Eyed ova,” 74, 194
m,
INDEX
F
Field, The, 34, 41, 128
Finnock (or phinock), 121
Fishery Board for Scotland, Annual reports,
10, 154, 155, 155 (note)
Glenetive Hatching Results, paper on, 127,
128
grilse, 123 (note)
sea-trout, 10
Fishing Gazette, The, 34, 41, 56
Francis, Francis, sea-trout fishing, 11
Fresh-water Fishes of the British Isles, The,
C. Tate Regan, 16-20, 38, 53-56, 62, 206,
207
Fry, 76-87, 121, 196-199
salmon, 76, 83, 84, 86, 87
sea-trout, artificial propagation
196-199
eye, 76
feeding habits, 83, 85-87
growth, 86
hibernation, 86, 87
nomenclature, 121
“parr marks,”’ 76, 83
tivalry between salmon, sea-trout and
trout, 83, 84
salmon, sea-trout and trout, distinctions,
76
scales, 87
shoal habit, 84, 85
trout, 76, 78, 79, 83, 84, 198
(q-v-),
G
Gathering grounds, lochs as, 144-147
Gemmill, Dr. James F., Salmonide, de-
formities, 77
Gill-covers, 56, 62
Glacial epoch, 28, 29
** Glen-lakes,’’ 145
Grilse, early running, 123 (note)
maxillary, 55
scales, 58
statutory definition, 9, 67 (note), 206
tail, 55
Giinther, Dr., bull trout, 4o, 63
H
Handy Guide to Fish Culture, A, J. J.
Armistead, 78, 198, 199
Hardy, John James, bull trout, 41, 65
Harvie-Brown, J. A., age of trout, 49
0
213
Hatching Results
Nelson, 127, 128
Hebrides, bull trout, 25
lochs and streams, 27
tidal trout (S. orcadensis), 24
true fresh water fishes absent (Regan), 28
Herling, Crown’s rights in Scotland, 8
nomenclature, 121
at Glenetive, Ian T.
Herring, sea-trout food, 135
trout food, 25
History of British Fishes, A, William
Yarrell, 56, 57, 62, 78
“Homing ”’ instinct, sea-trout,
169-175
whitling, 125-128
Houghton, Rev. W., bull trout, 40
Howietoun Fishery, sea-trout experiments,
22.823 Loe te
Hucho, genus, 18
Hiiitfeldt-Kaas, Hartig, salmon and trout,
distinctions between young, 93, 102-104
Hutton, J. Arthur, acknowledgment of in-
debtedness, 2
lochs in river channels, 111
salmon, leaping, 151
spawning dress, 160
salmon and trout, translation of distinc-
tions by Hartig Hiiitfeldt-Kaas, 102-
104
sea-trout and grilse scales contrasted, 58
sea-trout, descent of smolts, evidence from
scales, 108-111
Norwegian specimen fish, 173
opinion as to a doubtful fish, 34
recaptures, scales examined, 171-176
residence in sea, evidence from scales,
143
run of fish of graduated weights, 148
scales, examination undertaken, 108
125-128
’
smolt, late running, evidence from
scales, 105
whitling, first return, evidence from
scales, 119
scales of small Norwegian fish, 122
spawning, evidence from scales, 130, 131
Hvorledes adskilles lakse-drret-og roieyngel
fra hinanden, Hartig Hiiitfeldt-Kaas,
93, 102-104
Hybrids, Hebridean (Stuart),
occurrence in nature, 163
produced artificially, 15, 23,
rs)
nn
36, 164
I
Tsospondyli, order, 18
214
|
Johnston, H. W., salmon, divided run, 123,
124
leaping, 151
“Tay bull trout,” 50
K
Katadromous habit, eels, 2
Keeper’s Book, The, P. Jeffrey Mackie, 181
Kelts, capture illegal, 132, 166 (note), 167
(note)
descent, 142, 167-169
description, 159, 160, 168
feeding habits, 168
recuperation, 169
timidity, 167, 168
‘unclean or wunseasonable,’’
expression interpreted, 132
whitling, 132, 133
statutory
I,
Lateral line, 54
Letters to a Salmon Fisher’s Sons, A. H.
Chaytor, 96, 161
Life-History and Habits of the Salmon, Sea-
Trout, Trout, and Other Fresh-water
Fish, P. D. Malloch, 16, 17, 50, 56, 86,
TOlGelZO mete leo, 132, TAO; LAI,
150, 164, 175.
Life of the Salmon, The, W. 1. Calderwood,
ZA 255 27s) SLs 325 345) 50, LOO, LID, Ir4;
123, 124, 207
Lochs referred to—
Awe, descending kelts, 168
scarcity of sea-trout, 147
Baa, sea-trout spring run, 134
Dhuloch, brackish waters, 145
tidal trout, 26
Eck, little above sea-level, 146
netted sea-trout, 147
Etive, sea-trout gathering ground, 147
tidal loch, 145
tidal trout, 26
Freisa, ‘‘ parr-marked ”’ trout, 94
Fyne, sea-loch, 145
Gareloch, sea-loch, 145
Hallan, trout become tidal (Stuart), 25
Harray, brackish waters, 145
Loch-na-Maorachan, tailless trout, 42
Lomond, “ black-nebs,’’ 119, 121
Clyde river basin, 34
little above sea-level, 146
Luss Hatchery, 181-187
INDEX
Lochs referred to (continued )—
Lomond, parr, exceptional size, 107
“ parr-marked ”’ trout, 94
sea-trout, descending kelts, 168, 169
gathering ground, 146
“homing ”’ fish, 125-127, 169-175
recaptures, 170-175
residence in sea, 143, 144
spawning annual, 170-176
spring run, 139
silvery trout, 100
smolts, exceptional size, 105
migration, 105, 108-111
spawning tributaries, 85, 161, 170-175
trout descend to sea, 27, 37
spots, 52
whitling, delayed ascent, 124
descent, 133
exceptional size, 104, 108
first return, 119
“homing ”’ fish, 125-127
main run, 120
shoal habit, 125-127
spawning, 130, 131
Long, sea-loch, 145
Maree, little above sea-level, 146
sea-trout fishing, 147
Morar, little above sea-level, 146
Ness, descending kelts, 168, 169
Shiel, little above sea-level, 146
sea-trout fishing, 147
South Uist lochs, sea-trout spring run, 134
Stenness, brackish waters, 145
Tay, ‘‘ parr-marked ”’ trout, 94
Lochs and Loch Fishing, Hamish Stuart, 25,
32
uss Hatchery, description, 181-187
M.
Mackie, P. Jeffrey, acknowledgment of in-
debtedness, 181
Maitland, Sir James, Howietoun Fishery, 22
Malloch, P. D., parr, hibernation, 86
salmon distinguished from sea-trout, 56,
150
sea-trout, nomenclature, 16, 120, 121
Salmo trutta, one species, 16
smolt dress, 1or
spawning period, duration, 164
“Tay bull trout,” 50
flesh, 175
trout, Salmo fario, one species, 17
whitling, ascent, 120, 128
do not spawn, 132
return as sea-trout, 140, 141
INDEX F
Marine origin of Salmonide, 30-32
Marsipobranchii, class, 18
Marvels of Fish Life as revealed by the
Camera, Dr. Francis Ward, 51, 79
Maxillary, of salmon and trout, 55, 97
Maxwell, Sir Herbert, Bart., Editor
Scrope, ior
heavy sea-trout, 50
Salmonide@, original habitat, 32
sea-trout fishing, 11
Migratory habit, as affecting legal rights,
8) nr
no test of species or race (Regan), 20-38
““ Monsters,’’ 77
Morphology of fishes, 12
Morphologists and systematists, 12-14
of
N
Nelson, Ian T., Glenetive Hatchery, 127, 128
sea-trout recaptures, 128
New Zealand trout, 24
Nomenclature, sea-trout (Malloch), 120
suggested, 121
whitling, localisms, 107-109
oO
Origin of Species, On the, Charles Darwin,
36
Orkney and Shetlands, sea-trout, 28
trout, 24, 26, 28
true fresh-water fishes absent (Regan), 28
1B
Pacific salmon and trout, 19
Parr, 9, 67 (note), 84, 86, QI-99, 102-104, 106,
II2, 113, 206
salmon, 91-99, 106, 113
sea-trout, capture illegal, 92
colouration, 96, 98
Dahl, g2
distinctions tabulated, 99, 102-104
eye, 07
feeding habits, 91
fins, 97, 98
growth, 91, 92
hibernation, 86, 91
Hititfeldt-Kaas, 93, 102-104
maxillary, 97
“parr marks,”’ 96, 97
salmon, distinctions, 96-98
scale formula, 97
shape, 98
shoal habit, 84, 106, 113
Size, 93, 96, 107
un
Parr, sea-trout, spots, 97, 98
Statutory definition, 9, 67 (note), 206
tail, 98
trout, distinctions, 97, 98
vigour (Chaytor), 96
Paton, Dr. A. Noel, Salmonid@, original
habitat, 31
Pectoral fins, 53, 75, 95, 97, 98
Pelvic (or ventral) fins, 53
Phinock (or finnock), 121
Pisces, class, 18
Prehistoric river channels, 39
Protective colouration, 51
R
Regan, C. Tate, migratory habit no test of
species or race, 20-38, 206
Salmonid@, classification, 18, 19
structural features, 53-56
sea-trout, two races, 38, 62, 207
young indistinguishable from young
trout, 17, 97
trout, one species, 16
young indistinguishable from young
sea-trout, 17, 97
Rivers referred to—
Aa, Norway, sea-trout, annual spawning
174
smolt migration, 109
Add, age and size of salmon, 107
Aln, bull trout, 39, 41, 148
estuary not extensive, 64
Alness, sea-trout spawning, 154
Altnagairoch Burn, recaptures,
sea-trout, 171, 175
sea-trout, “‘ homing,” 126
Annan, whitling, main run, 120
Arm Burn, recaptures, marked sea-trout,
170
sea-trout ‘‘ homing,” 125, 126
Awe, sea-trout ascent, 147
Balgay, sea-trout spawning, 154
Beauly, sea-trout, spring run, 134, 140
Clyde, marked sea-trout recaptured, 170
river system, 34
Sea-trout in estuary, 115
spawning sea-trout, 154
trout in estuary, 2
Coquet, bull trout, 38-42, 65
estuary not extensive, 64
sea-trout versus salmon, 84
Echaig, sea-trout netted, 147
Endrick, salmon leap, 151
Esk, sea-trout, spring run, 140
marked
216 INDEX :
Rivers referred to (continued)— Rivers referred to (continued)—
Etive, Hatchery results, 127, 128 Tyne (Scotland) estuary, 64
Finlas, recaptures, marked sea-trout, 170, trout, 39
174 Wye, sea-trout, ascent, 152 (note)
sea-trout ‘‘ homing,’ 125 annual spawning, 173
spawning sea-trout, 161, 162 smolt, migration, 109
Forth, bull trout, 39 Ythan, spring sea-trout, 134
sea-trout at Howietoun, 22 Royal Commission, 1902, bull trout, 40
Gloppen, Norway, bull trout, 66 sea-trout, 10
Hebridean rivers, tidal trout, 2 Rudimentary (or aborted) fins, 53
parr, 106 S
Howmore, bull trout, 25 Salmo, genus, 13, 14
Leven, parr, 106 albus, 14, 38, 62
pollution, 127 cambricus, 14, 38, 62
sea-trout, spring run, 139 eriox, 13, 14, 39, 40
smolt described, 104 estuarius, 24, 25
spawning sea-trout, 154 fario, 14, 39, 42, 100, 206, 208
Little Osen, Norway, 122 ferox, 14, 24
Luss, sea-trout ‘“ homing,” 125 fontinalis, 23
leaping, 151 levenensis, 14, 23
recaptures, marked sea-trout, 170 nigripinnis, 14
Nairn, sea-trout spawning, 155 orcadensis, 15, 24
Nith, whitling, main run, 120 salar, 14, 19, 42, 54, 77) 206, 208
Orkney rivers, tidal trout, 24, 26 stomachicus, 14
parr, 106 trutta, 14, 19, 39, 40, 42, 54, 77, 206, 208
Osen, Norway, sea-trout annual spawn- Salmon, alevins, 76
mg, 173 colouration, 47, 93, 100
smolt migration, 109 cross-breeding, 163, 164
prehistoric river channels, 39 eggs, 71-74, 76, 192, 104
Smiddy Burn, recaptures, marked sea- eye, 58, 76, 95
trout, 171 fins, 53, 54, 95, 96, 101
sea-trout “homing,” 126 fry, rivalry with sea-trout and trout, 83,
South Uist, tidal trout, 2 84
Strome Dearg, trout, 2 gill-covers, 56, 57
Tay, parr, 106 maxillary, 55, 95
salmon, age and size, 107 Pacific, 19
sea-trout spring run, 134, 140, I41
smolts in estuary, 115
““Tay bull trout,” 4o
trout in estuary, 25, 27, 34
whitling, ascent, limits of, 128
main run, 120
return as sea-trout, 140, 141
Towey, monstrosity, 78
parr, 91-99, 105, 113
“parr marks,”’ 94, 101
scales, 54, 96, 97, 101
sea-trout, distinguished from, 53-59, 76,
96-98, 99, 100-107, 150
shape, 53, 95
smolts, 100-107, III, 114, 115
spawning dress, 50, 159, 160
Tummel, salmon leaping, 151 spots, 49, 95, 101
Tweed, bull trout, 38-42, 62, 63, 148 statutory definition, 9, 67 (note), 206
close time, 148 tail, 54, 96
estuary, 64 “Tay bull trout,’ 40, 49
kelts destroyed, 166 teeth, 57
parr, 106 trout, distinguished from, 54, 55, 76,
sea-trout, spring run, 140 93-96, 99, 102-104
spawning sea-trout, 154 “unclean or unseasonable,’’ 132
Tyne (England) estuary, 64 Salmon and Sea-Trout, Sir Herbert Maxwell,
trout, 39 Bart.) ily 325950
INDEX 217
Salmon fishery legislation, 43, 44
Salmon fishing, common law, Scotland, 8
Crown’s rights in Scotland, 7
Salmon Rivers and Lochs of Scotland, The,
W. L. Calderwood, 4o, 147
Salmonide, classification, 13-15, 18, 19, 42,
206-208
common ancestor, 32, 207
distribution, 30
marine origin, 30-32
Salvelinus, genus, 18, 19
Scales, bull trout, Aln, 65, 66
erosion, 5, 171, 172
examination urged, 42, 64, 66, 113
fry, 87
growth, 87
parr, 87, 96, 97
salmon and trout distinguished, 54
scale formula, salmon, 54-56, 96, 97
sea-trout, 54-56, 97
sea-trout, annual spawning, 171-176
grilse, distinguished from, 58
marine residence, 143, 144
recaptures, 171-176
salmon, distinguished from, 59
trout, distinguished from, 61
smolt, 101, 105
exceptional size, 105
migration, 108-111
whitling, ascent, 119
small Norwegian, 122
spawning, 130, 131
Scrope, William, parr and smolts, 101
Sea-trout, age, 49
alevins (q.v.), 74-76, 79, 80, 121, 196-198
anadromous habit, 24, 29
artificial propagation (q-V.), 179-200
bull trout, distinguished from, 38, 40, 42,
56, 61-67, 175, 207
classification, 12, 17-19, 42, 206-208
one species (Malloch), 16
two races (Regan), 38, 62
two species suggested, 42, 208
two varieties (Calderwood), 40
close season criticised, 155
colouration, 33, 35, 47, 50, 51, 52, 50, 61
cross-breeding, 23, 163, 164
Crown’s rights in Scotland, 7
deformities, 77, 78, 197
distribution, 28, 30, 47, 48
eggs (q.V.), 71-76, 78-80, 130, 162, 179-200
eye, 58
feeding habits, 83, 85-87, QI, 129, 135, 149,
168
Sea-trout, fry, 83, 85-87
kelts, 168
parr, gt
sea-trout, 135, 149
whitling, 129, 135
fins, 50, 53
first return, 140-144
evidence from scales, 143, 144
“ Fordidge,’’ 39
fry (q.v.), 76-87, 121, 196-199
gathering grounds, 144-147
gill-covers, 56, 57
Hebridean, 25, 28
“homing ’’ instinct, 125-128, 169-175
Howietoun experiments, 22, 23, 112, 113
kelts (q.v.), 132, 133, 142, 159, 160, 167-169
land-locked, 21-24, 112, 113
leaping, 149, 151, 152
limits of ascent, 152
lochs as gathering grounds, 144-147
maxillary, 55, 97
“monsters,’’ 77, 78
Norwegian, 66, 102-104, 108-111, 122, 148,
173, 174
nuinbers decreasing, 43
original habitat, 29-32
Orkney and Shetland, 28
parr (q-v.), 9, 67 (note), 84, 86, 91-99,
102-104, 106, 112, 113, 206
running habits, 147-153
salmon, distinguished from, 53-59, 76,
96-98, 99, 100-107, 150
scales (q-v.), 42, 52, 54-56, 58, 59, 61,
64-66, 87, 96, 97, 101, 105, 108-111, rie
I1Q, 122, 130, 131, 171-176
sexes, 61
Shape, 47, 53, 59, 98, 159, 160
shoal habit, 84, 85,106, 113, 123-127, 147,
150
smolts (q.v.), 9, 67 (note), gI-116, 206
spawning (q.v.), 50, 61, 92 (note), 132, 148,
153-155, 159-176
spots, 49, 50, 51, 52, 53, 97, 98
spring run, 133, 134, 139, 140
statutory definition, 9, 67 (note), 206
tail, 50, 54, 63
teeth, 57, 60
timidity, 152, 153, 167, 168
trout, no structural differences from
(Regan), 17, 20, 59, 97
“young ”’ indistinguishable (Regan),
17, 21, 97
distinctions noted, 76, 98, 99
“unclean or unseasonable,"’ 132
218
Sea-trout, weather, 150
weight, 38, 48, 49, 50
whitling (q.v-), 9, 104,
II9Q-135, 139
Sea-trout fishing, common law of Scotland, 8
Crown’s rights in Scotland, 7
Sewen, 38, 62
Slob trout, 24, 25
Smolts, 9, 67 (note), 91-116, 206
salmon, 100-107, III, I14, I15
sea-trout, age at time of descent, 105-113
capture illegal, 92
colouration, 101, 104
Dahl, 92, 108-110
descent, 104, 105
description, 104
distinctions tabulated, 102-104
distinguished from salmon, 100, 101
estuarial habit, 114
fins, 101, 104
Hititfeldt-Kaas, 93, 102-104
lochs, influence of, 111, 144-147
migration, scale evidence, 105, 108-112
migratory instinct dormant, 112
*‘ parr marks,’’ ror, 104
scales, Io1, 104, 105, 108-112
shoal habit, 113
size, IOI, 104, 115
spots, 101, 104
statutory definition, 9, 67 (note), 206
tail, ror, 104
vigour (Chaytor), 96
Spawning period, 159-176
annual function, 169-176
ascent, 148
barren fish, 160
cross-breeding, 163, 164
dress, 50, 61, 159, 160
duration, 164, 165
effects on fish, 165
kelts, 132, 165-169
capture illegal, 132, 166
mortality, 165, 166
months, 153-155
ova, 161, 162
process described, 160-163
redds, 161, 163
disturbance illegal, 92 (note), 165
season, 148, 153-155
Splash nets, 149
Stuart, Hamish, Hebridean trout, 2
Salmonide@, original habitat, 32
Stewart, Charles, Scottish law of fishing, 7
TOS, EUs ELLs,
INDEX
Systematic Arrangement of the Fishes of
the Family Salmonid@, The, C. Tate
Regan, 18
Systematists and morphologists, 12-14
sik
Tail, bull trout, contrasted with sea-trout,
40-42, 63
parr, 96, 98
salmon contrasted with trout, 54
salmon contrasted with sea-trout, 54, 55
smolts, 55, 101, 104 i
Tailless trout, 42
“Tay bull trout,’ 40, 49, 175
Teeth, 57, 60
Teleostei, sub-class, 18
Teratology of Fishes, The, Dr. James F.
Gemmill, 77
Thomson, John, Howietoun experiments, 22
Thymallus, genus, 19 .
Tidal trout, 24-27, 34
Treatise on the Law of Scotland relating to
the Rights of Fishing, Charles Stewart,
y
7
Trout, age, 49
bull-nosed, 42
classification, 12, 17-19, 42, 206-208
Salmo fario, one species (Malloch), 17
Salmo trutta, one species (Regan), 16
Clyde, 34
colouration, 33, 34, 35, 50, 51, 52, 59, 93-96
cross-breeding, 15, 23, 36, 163, 164
deformities, 42
distribution, 28, 47
eggs, 71, 72, 74-76, 78, 79, 192, 194, 198
estuarine, 24-27, 34
fins, 53, 95
‘* Fordidge,’’ 41
fry, 76, 78, 79, 83, 84, 198
Hebridean, 24, 25, 27, 28
hybrid (Stuart), 2
Loch Lomond, 27, 34, 37, 52, 100
marine origin, 28, 2
maxillary, 55
New Zealand, 24
Orkney and Shetland, 24, 26, 28
Pacific, 19
“ parr-marked,”’ 94
salmon, distinguished from (Regan), 54,
55
scales, 54, 61
sea-trout, no structural differences from
(Regan), 17, 20, 59, 97
INDEX
219
Trout, sea trout “young’. indistinguishable Whitling, characteristics, 119, 129, 130
from young of (Regan), 17, 21, 97
distinctions noted, 76, 98, 99
shape, 53, 59, 95
silvery, 25, 100
slob, 24, 25
spots, 50, 51, 52, 53, 95
tailless, 42
teeth, 57, 60
tidal, 24-27, 34
transplanting, 24, 48
weight, 49, 49 (note)
Trout Waters: Management and Angling,
W. H. Armistead, 139
Tweed Acts, kelts, 167
Tweed Commissioners, Annual Report,
1915, diseased fish destroyed, 165, 166
U
Umbilical (or yolk) sac, 75
“Unclean or unseasonable,
pression discussed, 132
”
statutory ex-
Vv
Ventral (or pelvic) fins, 53
Vomer, 57, 60
W
Walton, Izaak, cross-breeding, 36
bull trout, 41
Ward, Dr. Francis, colouration, 51
mortality of eggs, etc., 79
Whitling, 9, 104, 108, 113, 115, 119-135, 139
age, 113
ascent, limits of, 128
time of, 119
Yarrell,
colouration, 119, 129
descent, 133
eggs, 130
divided run, 123
feeding habits, 115, 119, 122, 129, 130, 133,
135
fins, I19, 121
growth, 119, 133
“homing ’’ instinct, 125-128
legality of capture, 132
maturity, 133, 139
migration, 122-124
nomenclature, 119-121
Norwegian, 122
*“ parr marks,” 119
scales, 119, 130
shoal habit, 124
size, 104, 108, 119, I2I, 122, 129
spawning, 128-132
dress, 129
evidence from scales, 130, 131
Malloch’s opinion, 132
spring run, 133, 134
statutory definition, 9
Wonderful Trout, The, J. A. Harvie-Brown,
49 (note)
Woodward, Dr. A. ‘Smith, Salmonide,
common ancestor, 32
Y
William, salmon, sea-trout and
“‘orey trout ”’ distinguished, 56, 62
teeth, 57
trout deformity, 78
“ Yellow-fins,’’? 120
Yolk (or umbilical) sac, 75
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