THE SI S.
On
,
Observations on the Scales of the Chinook
Salmon (Oncorhynchus tschavytscha) with
Particular Reference to the Variations
which may affect the Determination of Age.
Submitted to the
(
REGON AGRICULTURAL
COLLEGE
==
=
In Partial Fulfillment of the R:equirements
For the Degree of
'LASTER
OF
SCIENCE
In
THE SCHOOL OF AGRIC-uLTURE
Department of Zoology
Redacted for Privacy
May 26, 1915
APP.: uT:D:
Redacted for Privacy
Professor
and Head of Department of Zoology
in charge of
Major
Redacted for Privacy
Dean of School of
Agriculture
Redacted for Privacy
Chairman of Committee of Graduate Students
and Advanced Degrees
Observations on the Scales of the Chinook Salmon
(Oncorhynchus tschawytscha) with Particular Reference to
the Variations which may affect the Determination of Age.
Introductory.
Shortly after the writer's arrival in Corvallis,
a call came to assist in determining if possible, the
etiology of an epidemic which caused a serious loss of
salmon fry in one of the State hatcheries.
From my work
in this connection (the results of which may be found in
the files of the departments of zoology and bacteriology
of the Oregon Agricultural College) sprung an acquaintance
with various other problems which faced the state hatchBut by the time my bacteriological investi-
ery workers.
gations were finished it was too late in the year to get
material for
El
start, and moreover my knowledge of fish
cultural practices and principles were still too scanty
to be of any assistance in carrying out any applied research work.
Accordingly the remainder of the year was
confined to gaining this required knowledge, and a more
thorough understanding of the problems at hand.
Perhaps the question naturally arises here as
to the necessity of artificial propagation.
People are
wont to remark that in other days the salmon succeeded in
reproducing their kind without human aid and that it is
not to be expected that they cannot do so to-day.
The
answer to this question is found in a study of conditions
which actually prevent the fish from ever depositing
-2-
their eggs in that portion of a river where instinct tells
the females their eggs are best cared for.
We know that
the females ascend the coastal streams great distances to
deposit their eggs in the fine gravel and cold clear medium of the headwaters.
But civilization has seen fit to
erect barriers such as power-dams and irrigation dams
which prevent this ascent.
Cities are built on the banks
of the rivers and the sludge from their sewers and the
distasteful and poisonous by-products of their manufactures are poured into the stream to sicken and devitalize
those individuals that escape the fishermen's nets, racks
and wheels.
Small wonder then that with the decreased
numbers which eventually reach the headwaters via man-made
fishways over dams almost impossible of ascent, and the
enormous loss due to predatory animals and fishes, that
those young which eventually mature come far from being a
fair numerical representation of their ancestral hosts.
Before the advent of a system of artificial
propagation of the various species involved, the annual
take of these food fishes in the Pacific Coast rivers had
begun to decrease at an alarming rate.
The same crisis
was faced in the British isles long before Columbus discovered our continent.
Records of the old Scottish courts
refer us to an article in the renal code of about 1432
wherein it was set forth that a man convicted for a third
time of poaching or over-fishing should forfeit his life
for the offense unless he be able to buy himself off.
But
the introduction of a working scheme for artificially
rearing salmon has -worked wonders with the supply of the
-3-
Oregon Coastal streams.
That this is no vainglorious
boast on the part of the hatchery superintendents is borne
out by the action of the Astoria cannery men, who during
the past summer (1914) subscribed 43,000 for the construction of artificial rearing ponds at the Bonneville State
Hatchery for the express purpose of rearing a high grade
of commercial salmon; viz, Spring, Chinook.
The financial support
thus gained
marked the
winning over of this group of business men who heretofore
were just the least bit skeptical about the project.
This change of sentiment was due largely to the efforts
of the present superintendent of hatcheries, Mr. R. E.
Clanton, who upon coming into his present position just
five years ago, in the summer of 1910, turned out during
his first year an unusually large number of
blueback variety (O. nerka).
salmon
of the
The normal life period for
the members of this genus has long been asumed to extend
over a period of four years (subject to individual digression).
If this be the case, it was to be expected that
four years from the date of liberation, that season would
be marked by a correspondingly large run of this species.
And this is exactly what occurred, demonstrating to the
fishermen of the lower Columbia the great value of the
salmon hatcheries.
It so happens, however, that the blueback salmon
is not the most profitable fish to handle, nor is it the
best liked by the consumers.
The species which is most
valuable from both standpoints is one commonly known as
the Chinook or King Salmon (U. tschawytscha) and which is
characterized by having two distinct spawning runs, one
in the spring, the other in the fall of the year.
Of the
two runs, the former is the more highly prized because of
its rich pink color, smooth flesh texture, high fat content and choice flavor.
The fall run is notably of a pal-
er tint, is dry and lacking in fat and quite flat to the
taste.
The cause of this difference is unknown at present
and will not be dealt with at greater length in this paper.
The subscription before mentioned was raised for the pur-
pose of increasing the output of the more palatable variety.
In the early days of artificial fish propagation
it was customary to release the fry as soon as the yolk
sack was nearly absorbed and the young began to feed for
themselves.
I am told by old hatchery employees that it
was a very noticeable fact that when these fry were liberated by dumping them from buckets into the river, that the
surface of the water literally swirled as the hungry trout
and other preying fish eagerly awaited the feast of these
unprotected fry.
Dow the practice has changed; after
hatching, the fry are placed in rearing ponds where they
are fed until they reach a length of two or three inches
or more; they are then able to take care of themselves,
consequently the return should be much greater.
However,
this practice also meets considerable criticism, some
claiming that the cost of building retaining ponds and
feeding over a stretch of several months entails an expense never met by the increase of fish taken commercially.
-5-
For a defense of the present day cultural prac-
tices we are at present obliged to rely on our convictions,
and such support as that
merchants.
advanced by the Astoria
But it is our hope to demonstrate the value of
the work experimentally and in a scientific manner overcome whatever objections still remain.
With this end in view a series of experiments
was begun last fall through the medium of which it is
hoped to obtain data which
shall
show what percentage of
the fingerlings liberated would eventually return as
adults.
Of prime importance in this connection is the
consideration of the spring and
fall
varieties.
Do the
eggs from spring and fall fish give rise to adults which
will migrate as spring and
fall runs
Can
respectively?
we establish without question a knowledge of the normal
life period for these
fish and so predict a
year when a
large return from any one brood may be expected to return?
To settle
this
latter point it is
necessary to
devise some scheme by which we can tell the age of a returning individual and perhaps check this record with some
mark placed on the fry at the time of liberation.
This
problem of age at maturity of our Pacific Coast siUmon
has a peculiar interest as concerns these species because
of the fact, now sufficiently demonstrated, that all
individuals perish when they attain sexual maturity.
They
spawn but once and then die, whatever the age or the size
at which
sexual
maturity has bcen reached and whatever
may be their physical condition at the
time
of spawning.
-6-
Gilbert has pointed out the rossible exception to this
rule in the case of certain male Zing salmon fingerlings
which mature precociously in the streams during their
first year, at a length of 3 to 7 inches; the fate of
these has as yet not been determined.
This peculiarity
in their life history renders the question of their age
at maturity an unusually important one, both from the
scientific fish cultural and the purely economic standpoints.
Many attempts have been made to solve the problem for the most important of our Pacific Coast species;
viz., O. tschawytscha and O. nerka, usually by marking
hatchery-reared fry with a metal tag or by merely clipping some one fin or a particular combination of fins.
Unfortunately these experiments have never been carried
through with detailed precision, the data gathered resulting in somewhat discordant and uncertain results.
For ex-
ample, we have partial data on an experiment carried on by
a Mr. Rutter in 1896 when he liberated 5,000 marked fry.
From this number 376 were reported to have returned in the
third year, but the records for ensuing years were not
kept.
Other fish have been liberated by Mr. Henry O'Mal-
ley acting on authority for the United States Government;
they were marked with a silver wire loop and are not due
to return until the fell of 1915.
I am given to understand
that Mr. O'Malley is of the opinion that this loop will
have become grown over and will be too deeply imbedded in
the body to be readily found.
-7-
In 1912
Lr. E. A. Clanton conceived the idea
of placing the mark on the operculum, a very tough structure which is seldom injured under normal conditions,
whereas fish frequently lose fin and tail rays and pass
through many natural barriers, on which such things as
metal loops and tags may be
easily
caught and torn out.
The scheme used by M. Clanton consisted in cutting a "V"
shaped piece from the operculum, using a sort of steel
punch which he had made for the purpose.
Some of these
cuts included more of the operculum than others, and during the last spawning season (1914) several fish with peculiar gill marks which seem to have been caused by the
method just described were taken.
it is a noticeable fact
that the clearness of the mark on the .adult depends on the
amount of operculum originally cut out; those incisions
reaching the preopercie
being much
more clearly defined
than those which went no farther than the sub-opercle or
the inter-opercle.
(See Plate
iI & III).
To my mind
this method offers the most feasible system for marking,
although there are still several investigators within my
acquaintance who have yet to be convinced of it.
The
best argument advanced in favor of this mark comes from
the men who do the actual fishing, and
fish
along the Columbia.
handling of
cannery
These men are schooled through
long years of service to observe peculiar marks on salmon
after the fashion of a cattleman trained in observing
brands, as the National Government has several times offer-
..8-
ed attractive rewards for the return of other marked fish.
Moreover this group of men are particularly willing to
co-operate in establishing new facts concerning the life
,
and habits of the salmon, this willingness amounting to a
pleasurable hobby.
Before the last spawning season (1914)
no fish bearing this mark had ever been observed by these
men; net injury to the gills and opercula being of an internal nature or confined to the extremities of the cleft
while the mark referred to is always median in position.
This marking experiment was merely to 'test its
value and now in view of the success obtained by its proper use, it is intended to mark a definite number of fingerlings in this manner and await their return.
This
should yield the data so long desired.
The experiments noted above have been supple-
mented by others in Tomales Bay, Cal., and in New Zealand
(according to Gilbert) where fry were planted in streams
not frequented by the species in question and the return
of the adults noted.
Also in the case of the Sockeye or
blueback (0. Nerka) there is the quadrennial increase in
the Frazer River, which has generally been accelted as
demonstrating a four year cycle for the species.
But these researches cast no light on the individuals of a run, which is without doubt composed of fish
exhibiting a great diveristy of age.
In the case of sev-
eral widely different genera, the markings of the scales
-9-
and otoliths have proved of decided value in classifying
various groups as well as giving an insight to the life
history and age of various types.
The works published since 1899 by Hoffbauer,
Heincke and Stuart Thompson seemed to show that
a classification of this kind was feasible in the case of
carp, plaice and several of the Gadidae.
In some cases
by means of the scales, and in others by investigation
the bones, it was demonstrated that there existed periodic forms of growth in summer and winter, and that this
difference was indicated by annual growth-rings in these
parts, similar to the rings found in trees.
In England
particularly, there has been a large
amount of material written on this problem, much of it,
however, of a
purely popular
nature.
has been some of this more popular
Unfortunately it
material
which has coma
to the attention of some of our untrained investigators,
and as it can not be regarded as the exposition of well
regulated
investigational work
it must be shunned as a
guide to important research.
The use of the scales of the
salmon as
a
means
for studying the age and other biological conditions of
the fish is due to Mr. H. W. Johnston.
In three p, pers
published in the annual report of the Scottish Board of
Fisheries for 1904, 1906, and 1907, he has succeeded in
establishing the fact that periodic growth takes place in
-10he
the scales of the Atlantic salmon (Salmo salar).
found that groups of rings are formed each year, and that
it is possible by counting the number of these to ascer-
tain the age of nearly every individual, and thus form an
opinion regarding its life history.
At
first
Johnston
relied on scales from fish taken at various seasons of
the year, but he succeeded in demonstrating by the study
of these scales that various growth zones on the scales
were formed at various times of the year.
Later he mark-
ed fish with a silver wire and, upon the recapture of
some of these, had fish of a known age to work with.
The work of
Johnston
was ably supplemented by
Al-
Knut Dahl for the Norwegian Fisheries Commission.
though Dahl worked with the
salmon frequenting the colder
waters of the Norwegian
coast and fjords,
great
within
value, as it kept
painstakingly done.
the work is of
the same species
and
was
In his researches Dahl observed the
growth zones to the season of the year when formed.
For the benefit of those who may desire to apply the principles of Johnston's & Dahl's results to our
Pacific Coast salmon, it seems wise to insert here is
some detail the results of Dahl's work, which are essentially the same as those of Johnston and are at the time
of writing the more available.
This extract is taken
from Dahl's "The Age and. Growth of the Salmon and Trout
of Norway as shown by their Scales," Published by
the
-11-
London Salmon & Trout Ass'n. in English:
.
.
.
"During the summer after hatching, rings
are added rapidly and increase with the length of the
fish. For example, the following table shows the development of the
rings on parr which were
1898, and killed as
hatched in March,
the dates indicate.
Wumber of rings!
1
Length in cm.i Date Killed
1
1
1
'I min.
Max.
Average
I
I May
1
1
1
3-5
1 June
2
5
4.2
4-5
1 July
4
6
4.8
5-5
1 Aug.
4
8
6.2
6
1 Sept.
6
8
7.4
12
19
10
1 Oct.
16
-12-
"Figure i (Plate i) shows a scale from the largest parr.
The inner growth-rings go regularly and prac-
tically without a break around the whole of the scale.
The ring formation is not broken until the three outer
rings are reached.
These latter rings are only perfect
on the anterior portion of the scale and elsewhere they
are merged in the edge of the scale.
Note that the fish
was killed late in the autumn.
"As a rule, summer rings are unbroken around
the circumference, although one may occasionally meet with
a scale with incomplete summer rings.
This probably ari-
ses from the fact that all growth does not proceed simultaneously and therefore the different portions of the
scale have not always the same uniform rate of growth in
length ang girth.
In autumn and winter the growth gen-
erally decreases or ceases entirely; such little growth
as does take place is shown in the production of narrow
rings close to the edge of the scale.
After the winter
when development begins again, the ring formation of the
new season does not exactly coincide with the last formed
rings of the preceeding season and the new growth is
shown by the formation again of entire and unbroken rings.
it should, therefore, be easy to recognize the winter
rings formed during the period of stagnation between the
two growth zones.
"The chief characters of this period of stag-
nation are the pronounced branching or ramification of
the rings and also the fact that they are narrower and
-13-
lie closer together so that a magnified representation
shows a narrow and more darkly shaded belt.
This nar-
row winter band has also a different optical refractive
power as the microscope shows when different methods of
illumination are used."
:Prom a government hatchery Dahl obtained sam-
ples of parr hatched in 14ril, 1907, and reared in the
hatchery until killed in the middle of December, 1908.
"These fish had, therefore, lived for two summers and a portion of the second winter; their scales
are shown in Figures 2 & 3,
(Plate I).
These correspond
with the seasons in which the fish lived.
The growth
zone in the center consisting of relatively unbroken rings
corresponds to the first summer.
Surrounding this is a
band of narrow branching rings depicting the slackened
rate of growth during the first winter.
Then again sur-
rounding this inner core are similar growth zones corresponding with the second summer and half of the second
winter, and which show a repetition of the characteristics of the first year's gro7!th.
"Figures 4,5 & 6 (Plate I) are scales of parr
from Softeland River in the Os district, taken in
June and October, and show clearly how the summer's growth
is indicated on the scales.
For example, in Fig. 4 we see
that the winter band formed during the second year of the
parr's existence lies close to the edge of the scale and
that only two new broad rings denoting summer growth had
been formed.
The June scale, Fig. 5, shows that these
broad summer rings have increased in number, whereas the
October example, Fig. 6, clearly indicates the fact that
the formation of broad slimmer rings had ceased, and the
narrow branching winter rings are present, showing the
commencement of the third winter of the fish's life."
Dahl presents further evidence to show that
this same condition continues to exist during the fish's
marine life by date taken4fish of known age which were
captured in the sea and placed in =salt water aquaria
for a time.
He also took fish from the sea at various
times of the year and demonstrated the gradual passage
from broad to narrow ringed areas as each season advanced.
In London, 1910, there was published a book entitled "The Life History and Habits of Salmon, Seatrout,
and other Fresh Water Fishes."
The author, Mr. P. J. Mal-
lock, attempted therein to set up a new system for the
determination of age for the salmon.
His studies led him
to believe that one had only to count the rings on a scale,
divide the total by sixteen, and the age to a month would
thus be told.
By the aid of several photographs he en-
deavors to show that each year there are formed sixteen
new rings, no more, no less, except in rare cases.
From
this information he makes the sweeping statement that this
furnishes the key to the entire life history, the length
of the time spent in fresh and salt water, the age at
-15-
migration, the type of fish, whether kelt, pike, small
spring fish, autumn fish, or large spring fish, weight
of the fish and number of times the individual has spawned.
In common with previous investigators he noted the varying
types of ring formation as caused by esturial or marine
life and the seasonal gradations in distance between rings.
That the yearly increase of scale rings is exant-
ly sixteen for the Atlantic salmon is open to serious question, as amply evidenced by Mallock's own observations,
for he points out the fact that many of these fish spend
as many as sixteen or seventeen months in the rivers prior
to spawning without feeding.
lose about
length.
25G)0
During this time the fish
of their weight and do not increase in
This being true, how can this long period be rep-
resented by an addition of a large number of rings?
The
study of scale development tells us that the number of
scales on a fish's body is constant throughout life and
that as the fish increases in size the scales also increase to cover the intervening spaces.
In face of this
fact, does it seem plausible that scale growth should con-
tinue regardless of bodily development, arrested as it is
by periods of starvation and emaciation?
Moreover, his
own observations show that during the spawning sojourns
the effect of the fresh water is to play havoc with the
edge of the scale, materially reducing its size and resulting in an area of irregular rings which give rise to the
so-called spawning mark when growth occurs anew.
Again,
16the dense bands formed in winter are characterized by
their irregularity and incompleteness, so that, even if
the first year's growth should be marked by the formation of just sixteen rings, what evidence can be brought
forward to prove that this
number is added each success-
ive year?
The problem as it faces as on the Pacific Coast
is still open to conclusive experiment.
In 1913 Dr. C. H.
Gilbert of Stanford jniversity published an article as
Document Dumber 767 of the U. S. Bureau of Fisheries, entitled "Age at Maturity of the Pacific Coast Salmon of
the Genus Oncorhynchus," in which he stated his observations on the scales of the adults of the five most important species of the racific salmon.
The results are
not supplemented by experimental evidence, but the conclusions drawn
take
their vases from the system as ad-
vanced by Johnston & Dahl,
difficult
and for
this reason
are rather
of interpretation unless one is familiar with
the writings of these two men.
However, the idea pre-
sents itself in a more logical and workable manner than
any other yet advanced.
On taking up my work with our state commission,
I was quite in ignorance of the methods available for the
solution of our problem, but the work to be done was all
mapped out and no difficulty was experienced on getting
it
underway.
Jar. M. J. Kinney of the Oregon State Fish
Commission was particularly
interested in the problem; as
- 17-
a retired cannery man himself,
he held an intimate
knowl-
edge of the fish in the lower river, as well as of a great
many of the problems which confront the hatchery-men.
He
recognized the importance of a workable age determination
scheme, and had searched for available literature on the
subject.
Jnfortanately, it seems to me, his search for
literature led him to discover Eallock's treatise on the
salmon and he was thus led to an erroneous conclusion.
The greatest point at fault in the application of Er. Mal-
lock's observations
to our Pacific coast salmon lies in
the fact that the two groups to be compared belong to
quite different genera.
Were this
one fact not suffi-
cient to question the applicability of Eallock's ring
count to the genus Oncorhynchus, there still remains the
wide difference in the life history and physiology of the
two, which of itself is entitled to great consideration.
But all of us who had part in the work at the beginning
were equally uninformed on
the
subject of
scale
growth
and age and so were willing to let the system in question
stand for the present.
evident to
As the work
went
on, it became
the writer that there were great possibilities
of error to be dealt with; many of the practical hatcherymen also began to doubt the value of the idea and to think
it advisable to find some other method for obtaining the
desired results.
.Towever, the work on hand was carried
forward on its original plan, the data for which is included in another part of this paper.
It
is hoped
that
-18-
something may be gained from the presentation, if no-
thing more than the statement of facts and principles
involved in such a problem, and a general consideration
of the many factors Thich enter in to influence the final
results.
At the very outset, therefore, let it be made
clear that this paper does not attempt to establish new
principles, overthrow previous theories, or greatly enlarge our knowledge of the subject in question.
The pur-
pose of the facts and data presented is to clear up in
the minds of those of us interested in the various phases
of the salmon fishing and propagatint, industry the sit-
uation as it stands today relative to one consideration
in the whole field, namely the determination of the age
of an individual salmon at any time when captured.
The
amount of literature dealing with this important subject
as applied to our Oregon fishes is pathetically small,
widely scattered and difficult to obtain.
It is hoped
that this paper will help to alleviate these difficulties
by compiling the salient facts from the most reliable
sources and placing them in a more readily available form
for the benefit of those who may desire to carry on further work in this state along similar lines.
There are included a few original observations,
some data and facts which should serve the purpose of
covering several of the preliminary steps which naturally
arise and must be disposed of before protracted research
-19 --
in this field can be attempted.
METHODS
It was not until after the two weeks spent on
the spawning grounds of the LcKenzie river that the
thought occurred to me that the material and data being
gathered there might be used as a thesis subject.
Our work on the McKenzie consisted in spawning
ripe female salmon and fertilizing the eggs by artificial
impregnation; that is to say, impregnation by the milt
of the male applied directly to the eggs as they fell into a bucket, as contrasted with the normal fertilization
as it occurs in the river.
On the eggs thus taken the
output of the hatchery is dependent, but the total number taken in a season amounts to a considerable figure;
in a single season it is not impossible to collect as
many as thirty millions of eggs, altho the number actually taken is considerably less as facilities for handling
so large a number are not to be had.
As stated before,no
data is available which will give us even an approximate
statement of the percentage of the fry from these eggs
which return as adults.
So this work was done for the
express purpose of answering this and one or two other im-
portant questions such as those following.
Do the eggs
from spring fish give rise to fry that return as spring
adults only or do some return with a fall ruh?
What per
cent of the total number liberated return after the expiration of three, four, five or more years?
20-
The idea was to take the females to be spawned,
examine the scales and separate out those which fell in-
to the four and five year classes, fertilize their eggs
with milt from males of the same age and keep the eggs
and fry separate thruout their development, without however,
giving
them any different treatment from that re-
ceived by the rest of the eggs in the hatchery.
The age
of the fish was considered to be determined by the Mallock ring-count system; table(I)gives the results of this
work.
More or Less complete data is given on eighty-
eight individuals; several more were examined and either
discarded or else the skin with the scales attached was
lost thru the ravages of mice.
Several of the males are
conspiciously lacking in accurate length and weight data
as there were but a few used
several females were fer-
tilized by one male, and each male was used as many as
three or four times on various days.
males and females were kept
in
When not in use the
large pens submerged in
the water from which they were removed by a dip net when
needed.
It so happened that the majority of the males
were still living when the
spawning operations for our
experiment were finished, so were left to be made use of
in the regular spawning work.
They were all killed a day
or so later,however, but at a time when the writer was at
the hatchery building working on the scales of the others
and was not present to measure or weigh them.
An attempt
was made to rescue them after butchering from the rest of
Table IA showing data gathered on fish regarded as 4 year
olds.
Length : Weight
inches
lbs.
No.Nings
Seined
No :
Sex
2
F
36
20
72
4
3
F
41
23
63
4
TT
TT
4
F
40
27
66
4
TT
TT
6
F
37
18
62
3
9
71
11
12
F
38
19
60f
3
91-
11
TT
13
F
40
21
70
4
4
H
'I
15
M
67-73
4-2to4-6
16
M
38
13
604-
3
21
M
40
25-
66
4
22
M
40
17
60
3
25
M
31
8
58-61
3-8to3-2.0
29
m
63-66
3-2'o3-4
n
IT
30
M
58-9
3
TT
TT
:
:
:
Yrs
:
Mos
6
9
9
8
:
9/7/14
9/8/14
IT
II
TT
TT
IT
IT
9/9/14
Spawned 9/9/14, yielding 31,000 eggs;. 6 females
an average of 5,166.66 eggs to the individual.
Table IB - 4 year-olds continued.
No
:
Sex
:
Length
inches
:
Weight
lbs.
:
Eo.Rings
:
Yrs
:
Mos
:
Seined
65-70
4to4
5
9/10/14
M
66
4
2
9/12/14
39
M
73
4
6
40
M
69-70
4
43
M
40
15
48
F
36
55
F
56
37
P
38
35
15
TT
TT
5
It
TV
II
If
16
3-9to4-3
2
66(av.5) 4
TT
11
36
16
64
4
3
9/11/14
F
36
17
72(av.3) 4
6
9/12/14
59
F
36
16
64-70
4
4-
63
F
33
12
63
4-
71
F
37
16
67
4
60-67
TT
TT
9/13/14
3
9/10/14
Spawned 9/13/14, yielding 21,000 eggs; 7 females
an average of 2,100 egp..s to each individual.
Table IC showing data gathered on fish regarded as 5 year
olds.
No.: Sex : Zen.
ins.
Wt
lbs.
:
No.
Rings
'Yrs.
:
Mos.
:
Seined
1
F
39
24
76
4
9
9/6/14
5
,,,
I,
42
27
76
4
9
9/7/14
7
F
38
20
BO
5
a
F
41
25
84
5
3
9
F
39
22
84
5
3
10
F
38
18
84
b
3
11
F
39
18
74
4
8
14
P
33
12
85
5
4
17
M
41
24
76
4
9
18
M
78
4
19
M
76
4
20
M
79
5
23
M
76-7
4
9
24
M
39
32
75-80
4
8
TT
TT
26
M
44
29
81-6
4
11
TY
27
M
TT
TT
28
M
5 to 5
5 to 5
4 to 5
11
IT
82-6
45
30
76-85
IT
Tt
II
TV
9/8/14
9/6/14
9/8/14
TT
IT
TT
lt
8
it
IT
9
7111
II
2 to 5
9 to 2
11
9/9/14
Spawned 9/9/14, yielding 28,000 eggs, 8 females,
an average of 3,500 eggs to the individual.
Table ID - 5 year-olds continued.
No :
Sex
:
Len.
ins.
:
Vt
No.
lbs.
Rings
Yrs.
Mos.
:
Seined
9/10/14
31
F
34
13
77
4
9
32
2
34
13
82-3
5
3+
TT
TT
33
F
40
23
72-4
4
6
II
t/-
34
F
23
10
80
5
."
"
35
F
33
11
83
5
IT
11
36
F
36
16
81-5
57-1 to 5-3
41
M
38
16
80-83
5 to 5
3
42
LI
38
15
82
5
2
44
M
36
12
73-85
4-6 to 5-3
45
F
41
25
83
5
3
IT
TT
46
'2
37
16
78
4
10
II
TT
47
F
37
15
76
4
9
IT
. It
49
F
36
16
88
5
6
IT
II
50
F
36
16
88
5
6
IT
If
51
F
38
16
76
4
9
TT
IT
53
F
36
16
87-8
5
5
3
IT
9/11/14
II
IT
?I
9/11/14
Spawned 9/11/14, 13 females, 47,000 eggs, average of 3,615.38+ to each.
Table 1E - 5 year-olds continued.
:
Sex
:
Len.
ins.
:o.
lbs.
Rings
:
Yrs.
10 8.
52
F
37
17
80-6
5
54
P
37
15
77-80
57
F
37
19
80-88
4-10 to :5
5 to 5 6
58
F
36
15
74
4
9
60
F
37
17
85
5
3
61
F
88
5
6
64
2
34
13
83(av)
5
2
65
F
34
15
79-80
5
6
F
35
14
82
54
70
F
34
12
72+
4
:
Seined
9/11/14
3
Ti
IT
9/12/14
Tr
tv
TT
9/13/14
TT
IT
IT
If
9/12/14
6+
IT
IT
Spawned 9/13/14, yielding 27,000 eggs, 10.
females, an average of 2,700 eggs to the individual.
Table 1F, 5 year-olds continued.
No.: Sex
:
Len.
ins.
:
Wt
lbs.
:
No.
Rings
:
Yrs.
62
F
35
16
77
4
66
F
33
8
80
5
67
F
36
15
86
5
68
F
38
17
84
72
F
37
17
79
73
F
32
9
74
M
Mos.
:
Seined
9/14/14
11
IT
5
IT
IT
78
5
17
IT
78
5
IT
IT
4
3
No
:
Sex :
7,en.
ins.
75
F
77
M
78
lvi
79
35
Wt
lbs.
11
:
No.
:
Yrs.
Mos.
72
4
6
77
4
9
72
4
M
81
54.
81
F
79
5
82
2
83
males, an
35
11
Seined
:
Rings
9/14/14
TT
37
17
85
5
3
35
12
76
4
9
Spawned 9/15/14, yielding 45,000 eggs, 10
average of 4,500 eggs to the individual.
n
r!
I/
IT
It
II
fe-
It will be noticed that there is a wide variation in the average number of eggs taken from the females.
An explanation of this difference is not to be had at
present, but it is probably due to the
variation in
cundity exhibited by the various individuals.
fe-
Then too
we must .onsider that there is a chance of having incor-
rectly
measuring the eggs, a very easy thing to do, as
the process is only intended to be approximate.
the pile, but it was found in most cases a hopeless
task as the marks used for identification (the clipping
of a fin or a combination of fins) were greatly altered
thru the degenerating action of fingers and other agents.
It was found most convenient to divide the work
among alternate days, the fish taken in the sein were
separated and marked during that day but were not spawnThen just before spawning
ed until the following day.
a new batch was taken and tied up for examination during the rest of th,t day.
Each fish as taken was placed
In a large common pen and from these were selected those
fish which were not too ripe to hold over a day or two,
or were not too badly infested with Saprolegnia.
These
fish, selected for no other considerations were then tied
by the tail, marked, and after giving up a section of skin
with the scales attached were allowed to lie in the water
near the pen until the scale rings were counted.
the fish were separated into pens according to
they approximated the four or five year class.
Then
whether
Great di-
versity was found to exist among the individuals and as
it would have been out of the question to obtain fish
reading exactly four or five years, they were divided
arbitrarily into two groups.
Those which had from 56 to
72 rings (between 3 yrs. 6 mos. and .4 yrs. 6 mos.) were
taken as four-year-olds; those which had from 76 to 88
rings (between 4 yrs. 9 mos. and 5 yrs. 6 mos. ) were
-22-
taken as five-year-olds.
The individuals of the two
classes were placed in their respective pans And on
the following day were killed, measured, weighed and
spawned.
They were killed by a blow on the head with
an axe; the total length was taken as the distance
from the tip of the upper jaw to the tip of the longest
dosal ray of the eandal fin; the
weight
was read by sus-
pending each fish by the gills from a large spring scale.
Up to this point only one operation necessitated much
care or thought and that was the determination of the
proper place to take the scale samples, and the best
method to employ.
It is a very noticeable fact that spawning
salmon and those taken just as they enter the river from
the ocean exhibit a great difference in the relative
position of the scales.
The scales of the latter are
readily seen as they overlap and cover the entire surface of the body and may be. readily scraped off.
It be-
comes quite impossible to remove them in this manner
from those on the spawning grounds, uand many times the
fishermen raised the questionas to why this was true.
At that time I was unable to answer the question or to
give any opinion, but now further experience seems to
indicate that the reason for the difference is caused
by a thickening of the integument which leaves the scales deeply imbedded in it.
It would be interesting to
see to what extent this supposition were borne out by
-23_
In taking the samples at first, a
histaojical study.
very sharp razor blade was used; this always resulted
in one or two deep gashes in the body at that point,
which were accompanied with a considerable flow of blood;
the entire thickness of the skin was removed by this
method, the scales being still imbedded and out of sight.
To remove them from this position it was necessary to direct the skin apart, a slow and trying process.
So a
better method was sought and finally we found that the
skin could
be split
with a knife that had a coarser edge,
1,r. 17.yckman the superintendent
like a sharp meat knife.
of the ..leabAx- Aation is to be credited with this dis-
covery; it marks an important point in the development
The operation con-
of the technique of scale removal.
sists merely of making a rapid stroke with the knife in
the direction of
the
head; with a little experience it
soon becomes possible to
skin with
the
separate
the upper layer of the
scales attached, from the lower, thus leav-
ing the scales in a position of complete exposure, and
the body at that point still protected by the under layer
of akin.
Those pieces of skin with the scales imbedded
were subjected to various methods of treatment in the
attempt to remove the scales after returning to the laboratory.
By this
time
the skin was quite dry and
and resisted all treatment
cellent results
while in
hard
this condition. Ex-
were obtained however, by soaking the
pieces of skin in a little 10% grain alchohol and following the soaking
with a short
steaming in the same fluid.
Dad-kr
used in the same manner resulted in a gluey mass that
was difficult to handle; moreover if allowed to stand
more than an hour or two would begin to show signs of
putrefaction especially if steamed.
alcohol greatly
reduced
the gluey
the decay after steaming.
The presence of the
condition and
prevented
The process waa always carried
on'in small'petri dishes, one for each piece of skin,
labelled to conform with the envelope in which each piece
was filed; the cover of the dish prevented the escape of
the alcohol which might be driven off during the warming
process.
The scales removed from each piece if skin we
filed away in small envelopes each
labelled to
with the field number of the individual fish.
correspond
The whole
process is one marked by much tedious work, the manipulation of the large fish, the removal of the skin, the
removing of scales, the counting of the rings in the
field, and the painstaking and close microscopic application to the material in the laboratory.
The importance of the selection of the proper
place from which to remove the samples of skin beedme
more evident as the work went on; but with us the position was guaged largely by the condition of each individual fish; particular effort being made to take the
samples from good bright fish; if the fish were at all
-25-
spotted with fungus, care was exercised to avoid infected regions.
In most cases it happened that the reg-
ion chosen was from the posterior half of the body, dorsal to the lateral line and at a point between the adipose fin and the median region of the dorsal fin.
Fur-
ther study is contemplated to determine if possible the
region of the least variability in scale growth as applied to the selecting of a propar region.
In this
paper will be included only the preliminary observations
on this important point.
While reviewing the literature
on the subject as a whole, conflicting opinions were found to exist on this very point, Calderwood claiming that
the scales should be taken from the "shoulder" region,
while Dahl chose practically the same region as the
present writer, but neither presented statistical argument to favor his choice.
It is hoped that some of the
facts and observations to be discussed later will be of
some value in straightening out the question.
A very convenient system for filing scales
consists of a series of small manila envelopes arranged
In any desired order but numbered so that reference can
be made from the data concerning the scales which may be
first placed in a small piece of paper folded to fit inside the envelope and numbered to correspond with it.
The envelopes are then put away in a box and the scales
are always handy for study, and when desired fro use can
be readily mounted in water or soft glycerine jelly; per-
-26-
,manent mounts are unnecessary and unsatisfactory as if
made in balsam, the scales become too highly cleared and
do not show the seasonal bands well refracted-
MATERIALS
From the very nature of our problems it became
necessary to start the work with adult salmon; moreover,
the degree of development reached by fish hatched and observed during the relatively short time allowed for the
work would be hardly sufficient to bring results.
This
whole subject as treated in this paper must then, be considered as
merely
preliminary work which it is quite im-
hand before final work can be started.
For study there was available for me a wide
portant to have in
variety of scales representing the various environmental
conditions met by the fish on its way from the
spawning grounds.
sea to
the
Thus I had scales taken from fish just
after leaving salt water, when they were playing around in
the brackish
water off
Astoria;
July 15 and August 15, 1913.
were taken from
time, Llay 15, June 15,
On July 30, 1913 samples
bright fish on the Rogue River, and dur-
a large number of
the spawning grounds of the McKenzie
ing the first two weeks of September
scales were taken on
River.
With the excertion of this latter set of scales,
all those mentioned so far were taken by some person oth-
er than
myself; these other scales are the property of
the Oregon State Fish Commission and were loaned me by
Mr. R. E. Clanton, Superintendent of Hatcheries.
About
-27-
the last of the
season, some
half a dozen fish were taken
which carried the sip:ns of a gill mark like that proposed
by rir
Clanton, and first used by him four years previous;
we have these opercula and scales from the same fish, but
the men who took them failed to gather the rest of the
data which seems quite important; e.g. length and weight.
The scales from hatchery fish may or may not be of help
to us but I have
examined some from fish of known ages
and I will include my observations along with other data.
In all but the spaw,ing fish and those from the
hatchery we have
but
one scale from each individual, but
in as much as the fish were in perfect condition when
taken they
are considered to be perfect.
STUDIES
As just mentioned above, the possession of but
a single scale from several individuals was not enough to
halt the work, but rather to give it the acid test. When
a scheme is ultimately found that will tell us the age of
the mature fish, it must be one which does not necessitate the examination of a large number of scales from the
same fish, but rather one that is constant for all
the
scales, making one as reliable as another except for occasional variations due to injury or arrested development
of one type
or another.
In carrying on the work following, only those
scales were used which were accompanied by complete data
-28-
weight and length. I
regarding other conditions such as
of the text:series of
am inserting through the first part
all the data available
tables which will place before us
with the idea in
on all the fish examined, and arranged
there might not be
mind of ascertaining if possible if
weight, or length and
some correlation between length and
If such a correlation does exist then
number of rings.
of a fish by merely
one can very nearly predict the age
So these tables are
knowing these two measurements.
each special
worked out for the fish from all sources,
population.
set of which I have chosen to regard as a
The Columbia Population (See Table II)
In this group are represented 197 individuals
and bearing from
ranging in length from 19 to 50 inches
and an
52 to 128 rings; an average length of 35.58 inches
equivalent to an averaverage ring rount of 96.73 rings,
mathematics
age age of 6 years. Without going into the
that with
of correlating this data it is quite evident
correlation
such a varied grour of individuals no direct
Examination of the
exists between these two characters.
is a wide variation
data rresented will show that there
by fish of the same
as to the number of rings retained
length.
For instance of the 23 specimens measuring 38
96, one 98,
inches, one had 74 rings, one 90, one 94, one
110, one 111,
one 100, three 102, three 104, two 106, one
Again,
one 114, two 116, two 118, one 120 and one 124.
14 individuals
are represented as 23 inches long, and
show a variation from 54 to 110 rings with the majority
however between 90 and 100.
As we exceed
the
average
length, it becomes noticeable that the greater part of
the scales have by far, more than the average number of
rings and less variation is exhibited.
This might take
on a different aspect if we had more material to work
with, both as far as individuals go and as
of scales from each.
to the
number
But it seems safe to say that as
far as the data available at present writing is concerned,
and with particular reference to the locality in ouestion,
the tendency seems to be to have a greater development of
the rings on those fish attaining a considerable length.
And after all, this is only wht might .naturally be
ex-
pected.
If we plot the characters in this table separately
the
curves at
once assume a certain
similarity,
which would no doubt be the more striking if the classes
in the
ring-plot
were reduced.
The polymodal
appearance
of most of the curves presented, at once suggests that
factors other than those considered are active.
A com-
plete analysis of the situation reveals several factors
which should without doubt be considered.
Probably the
most logical assumption is that in a population of this
kind, a
certain
diversity of age as well as individual
physiological and environmental conditions play an impor-
-30-
In as much as plenty of evidence has been ad-
tant role.
vanced to suplort the theory that certain individuals may
mature more or less rapidly than the majority it seems
quite within reason to assume that we are dealing with a
group of salmon which are of mixed age.
Doubtless
this
is true in the present instance, so that we have a run in
which three or more ages are represented, the over-lapping of the ages being caused by the maximum development
of those of a younger age over the minimum development of
those of a more advanced age.
The explanation of this condition may perhaps
be found in the study of the life history of the individuals.
But a close observance of the life of the members
of this species is quite out of the question at present.
Born in
the
headwaters of our coastal streams, the young
salmon remain there only until they are large enough to
assume the responsibility of life in
the
ocean; this they
are able to do usually at the end of the second year.
Once they reach the sea, all trace of them is lost until
they return to spawn, but it is supposed that the greater
part of their marine life is spent in the Japanese current
where the temperature of the surrounding medium shows
little variation through out the year.
members of the Salmonidae are
voracious
As a rule all
feeders, but it
is quite possible to concieve of certain members of a
single brood meeting with more favorable conditions than
-31-
others, the result being abnormal development, either
above or below the average.
would be
In either case the result
evident in the adult. If
this supposition be
given any weight in the face of our knowledge of the development of the scele, it becomes a relatively simple
matter to prove that
the
development of
the number of
rings within any given time is subject to such a wide
variation as to be practically inapplicable to
the prin-
ciple of age determination.
As far as the effect of this variation may influence the number of rings, the same principle does not
affect the system advocated by Johnston and Dahl.
The ex-
amination of the scales themselves gives ample proof of
this,
as it is readily seen that the growth during the
same season may valy in different years.
Occasionally
winter bands are quite absent, and two successive summer
bands may vary widely as to their extent, the completeness
of
the rings and the distance between them.
These
various rates of scale development are dependent on the
ra.de of body growth as determined by
physiological or ev-
vironmental conditions.
THE ROGUE EIVR POPULATION (See
Table III)
material from this locality was gathered
July 30, 1913, and like that of the Columbia River was
The
taken from fish on the cannery floor just after
being
brought in by the fishermen. Data is given on 63 individuals varying in length from 16 to 46 inches, and bearing
9
7
6
3
2
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-32-
from 68 to 120 rings.
Like those from the Columbia River,
but one scale for each fish was available, a condition
which really ought to be remedied until satisfactory
proof is gained that one scale is as gdod as another.
The average length in inches of this group of
fish is 28.6 while the average number of rings is 93.8.
Thus an interesting comparison is found between these two
sets of data; these from the Rogue RLver dealing with
fish noticeably smaller than those from the Columbia.
That the development of the area of the scale goes hand
in hand with the size of the fish seems to be well borne
out from these facts, for the scale rings show a tendency
to increase with the length of the fish tho by no means
is the correlation actually demonstrated mathematically.
The degree of variation in the ring count in
those fish grouped about the average length again becomes
quite noticeable, for among the 6 individuals 27 inches
in length, we get a variation of from 80 to 116 rings,
while again of the 11 fish 29 inches in length there is
found a variation of from 78 to 120 rings.
Here again, the curve is strikingly Dolymodal
(see curve 2) possibly somewhat over emphasized through
the relatively small amount of data presented.
The curv'e
for length variation seems to bear out the above state-
ment, as it is decidedly regular, being plotted for only
a few individuals, where as a larger selection would
doubtless have included many fish with widely varying.
-33-
length.
1}f
LcKEEZI: PT2ULAT.ION (See Table IV)
By introducing this series of data, a new factor is brought into consideration.
heretofore, all scales
have been taken from clean fish, fresh from the tidal
waters, now we are dealing with those on the spawning
grounds.
The members of this group have been a long time
without food, they have comrletely changed their environment and are open to the deleterious action of fungus
couled with the effects of a long, tedious journey.
The
external changes brought on by this great change are very
noticeable; the fish have fallen off in weight, their
skin has lost its bright lustre and in many cases, large
Patches of white fungus have developed on the body, especially in the head region.
The results of investigations with these fish
are of the utmost importance, because it is with these
that we have to deal in taking our eggs for artificial
propagation.
If
it is planned to carry on extensive
breeding experiments, it at once becomes of importance to
know the age of the parent fish so that there may be a
check on all the work.
If our interest were solely to
determine the age at maturity of the species in question
then the bright clean fish from the lower river would
answer the purpose very well, for then we could determine
the age at the time of capture and make an allowance for
-34-
the time elapsing before spawning; this would then be
accurate with a discripancy of no more than a month or
two.
Other things being equal, it seems quite reason-
able to suppose that the readings at the cannery and at
the spawning grounds should be practically the same, because during the interim no feeding takes place; hence it
is probable that the body does not increase over any dimension.
If this be true, then from what we knov, of the
development of the scale, there can be no addition to its
area.
As a matter of fact, the fish loose weight in their
passage up the river; this does not however, necessarily
mean that any body dimension is on the decrease, except
perhaps a comparison of the fish nt the two stations,
they being hard, plump and well rounded down river, but
soft and flabby on the grounds, the change being one sole-
ly to the absorption of stored fat from the body cells.
Investigation of the data available on spawning
fish reveals some interesting facts.
Of particular sig-
nificance is the comparison of the two sets of scale data,
one being the reading taken direct from the report on the
L1eKenzie spawning work, the other from further observa-
tions with scales from the same fish.
Occasionally, it was observed, while examining
the scales on the grounds to determine to which group a
fish belonged, there appeared a scale which was devoid of
-35-
rings over a la-re pertion of the center.
In this case,
the scale was discarded and a new one taken, generally
with the result that the second appeared perfect in every
way.
But as the work went on, the writer found himself
becoming more familiar with the scales, and ere
long
it
became evident that the variation in the ring count of
scales from the same fish would vary, so that an average
would have to be struck.
Or else, the number of scales
that presented the ringless center was so great that a
new patch of skin was chosen or the fish discarded entirely.
This made the work very trying as the difficul-
ties encountered in following out the rings; for counting are great and must be experienced to be appreciated.
That great error may easily be introduced into
the work at this end is borne out by the data presented.
For instance table IV shows the record of 59 individuals,
all within the age limits set by the experiments in question, the ring count varying from 60 to 88 and the length
from 23 to 42 inches.
Further record of the same type
may be found in the tabulation of the complete data taken
at the McKenzie hatchery, to be found in another part of
this paper.
Here, little variation iS to be found, for
as a matter of fact, the scales we-:e selected, those lack-
ing rings at the center being discarded because at first
the cause of this loss of rings was not understood.
)th-
er scales were found which lacked 1-ings at the edge and
I
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were likewise discarded.
As a check on these scales, ten individuals
were selected upon the return to the laboratory and all
the scales removed from one small area of about two
square inches were examined.
The variation existing over
so small an area was found to be quite alarming (see
table V).
Here we find scales from the same fish with
from 9 to 80 rings in one case, from 6 to 80 in another
and from 52 to 89 in a third.
It is evident that these
fish are quite typical of the entire catch and for such a
wide variation over a limited area, some explanation must
be found.
TABLE V.
Ring-Count from McKenzie Fish--Selected
Fish Ifo. Rin,g7Count
1.
43, 51, 55, 79, 79, 81, 82, 85, 85, 67, 87, 93
2.
20,
46, 51, 54, 63, 64, 66, 69, 70, 75, 75, 78,
80
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41, 41, 47
75,
77,
4.
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76,
80
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61,
62,
74, 78, 79,
5.
62, 65,
80
10, 39, 57, 59,
6: 65,
69,
71,
73,
77,
78,
84,
86
8.
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66,
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91,
92,
95,
97
9.
60,
60,
62,
66,
69, 71,
73,
75,
78,
10.
52,
54,
05,
69,
83,
6.
9,
7.
62,
84, 84,
75,
80
80
78
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-37-
This same condition was found to exist by Dahl
and Calderwood znd is attributed to the general condition
of degeneration exhibited by all spawning fish.
The imme-
diate cause of this degeneration is doubtless due to the
great change encountered by the fish in passing from the
sea to fresh water and the great emaciation and decay
found in all spawning salmon both males and females coupled with the deteriorating action of the fungus.
With the Atlantic salmon (Salmo sebago) which
spawns two or three times, scale growth again begins upon
the return to the sea, and the addition of new rings to
the frayed and torn edges results in a very noticeable
region of union known as the spawning mark.
Here again we face a condition of affairs from
which it becomes quite difficult to understand just how
Mallock would proceed in determining the age of a fish
that had spawned once or twice and had at that time lost
a number of marginal rings thus greatly reducing the total.
So great is this loss Lmong the different
scales of the same fish that many of the seasonal bands
described by Johnston, Dahl and 7-ilbert may become totally effaced.
So that, which ever system we employ, in
face of these changes, it becomes impossible to say to
any degree of accuracy how much of any one scale, no matter how perfect it may appear, remains for examination.
In as much as this loss is common to all spawning fish it
soon becomes evident that the scales from any one are so
-38-
liable to be in this same state as to make accurate determination of age quite
out
of the cuestion.
If this be true, must our researches stop here?
We trust not, because the nature of the experiments outlined previously demands that we have this data to start
with.
Perhaps a more comprehensive study along similar
lines as indicated here may do much toward perfecting our
knowledge of the present problem.
If this fails then
there remain other avenues of attack which should be investigated before Riving up the task as futile.
One
thing is certain, and has been amply proven by the experience of the present writer, that is the necessity of
having material of a known age as a
all studies.
matter of
check on
This was not appreciated until this work
was well under way, then it was too late to get such material this season.
Had this need been forseen, we could have instructed the fishermen to be on the look-out for specimens marked by Mr. Clanton's gill punch.
As it was,
eight of these fish wee taken; the marked epercula and
several of the scales from eLch were saved, but the men
who captured these fish were not informed of the desirability of taking the measurements and
weight of
we have nothing to strike a comparison with.
each, so
Moreover,
all these marked specimens were ripe for spawning and the
scales in poor shape as is shown by Table VI
which gives
-39-
the ring count from these individuals, as determined from
a few scales accompanying the marked opercula.
TABLE VI.
Ring-count from Fish with Marked Opercula
Fish
Ring-count
63
1.
43, 46, 48, 51,
56,
57,
63,
63,
2.
41, 47, 47, 49, 49,
51,
51,
56, 56, 58
3.
50, 51, 51, 53, 54, 54, 56, 57, 60, 63
4.
57, 67, 68, 70, 70, 71, 72, 72, 72, 75
5.
68,
72,
73, 74,
74,
74,
74,
79,
6.
65,
67,
70,
71,
74,
74,
76,
79
7.
50, 54, 55, 56,
58,
58,
59,
68, 70
8.
41, 56, 58,
bl,
05,
70,
76,
80,
84,
85,
89
85
In view of the fact that we hve no comparative
data to accompany the ring count from these fish as we
had in the case of all the others, it might seem at first
that the inclusion of the data at this point viere some-
what irrelevant.
But it seems to me that this very lack
of data offers an excellent o portunity for
testing the
value of the ring count on the spawning ground.
system reliable as a means for
spav.ning fish?
determining
Is this
the age of
Is the ring count an index to the size of
the fish either as to
length or weight?
Certainly the
evidence Es presented gives no
indication of the possibility of a correlation such as
-40-
indicated above.
The variation in ring count on the same
fish is in most cases strikingly great, so that is seems
at once possible to state that so far as spawning fish
are concerned the ring count is so liable to great variation on the same individual as to be of no value in
translating many of the unknown facts in the fish's life.
Of course allowance must be made here for the
possibility of the inadaptability of the gill mark in
auestion to this purpose.
It may be that some of the
specimens at hand were of different ages and that some of
the marks or scars on the opercula were misinterpreted.
In either event, the conclusions drawn seem to be still
viable.
The reader is here refered to Plates
I & III
where are presented photographs of twelve oporoula bearing scars claimed to be due to a gill punch; eight of
these opercula corresT;ond to the ring data presented, the
others were necessarily omitted because scales were not
taken with the opercula.
Those having the scales to cor-
respond are numbered accordingly; the arrangement on the
plates however is the least bit out of order from the
necessity of conserving space in arrangement.
One phase of our work not yet mentioned has
been carried on with a few specimens of hatchery reared
fish of a known age, for the express purpose of determining if possible (1), the amount of variation found in the
ring count over various portions of the lody; (2) the re-
-41-
gion or regions least liable to variation as a means to
determining the regions best suited for taking scale samples; and (3) the relation if any between size and scale
growth (ring count) as exhibited by hatchery fish.
For this purpose, the writer gained access to a
few young fish, some fourteen, some twenty-three months
old.
The younger of these, three innnumber, are spring
Chinook from eggs taken on the EcKenzie River; the larger
three are fall Chinook for which no record as to the locality of the eggs is available.
Both were hitched and
reared at the Klaskanine River hatchery, and were there
retained until late this last winter (1915) when they
were all liberated.
The six specimens used in this work
were all killed December 26, 1915, and arrived at the
laboratory in rather poor condition despite the fact that
they were well packed and promptly delivered.
Examina-
tion of the scales showed that a large portion were with.
out the nuclear rings, the total number thus being much
reduced; whether this was due to a partial decay or whether it was the common condition of all the hatchery fish
at that time it is now imporsible to say.
To facilitate the exLmination of the scales the
body
of
each fish WES marked o f into eight regions as
shown in uhe accompanyinc' figure. (Fig.1)
Nearly all the scales from each region were removed and it was planned at first to count a large pro-
portion of these but it later developed into a task of
such proportions that this proceodure was carried out only with the firs' 'J
ten scales from
removed,
lErge
fish.
each region,
were examined;
7,ith the other five, but
Lelected at random from those
the data for this work is to be
.
found in tables VII & VIII.
In the tables just referred to, it will be noticed that the scale readings are grouped according to
letters, these p:roups correspond to the body regions so
marked in the above figure.
The reason for thus dividing
ur the fish into various regions was to determine if possible which portion of the body is the most reliable for
gathering data on
the
scales.
It will be recalled that
Calderwood and Dahl chose widely different areas for
their work, each believing he had found the least variable area.
AS regards other fish the writer has been un-
able to find reference to any such possible variation but
he IL:a, been
told by Mr. W. I. Rich of the U.
S. Bureau of
Fisheries that in the case of the steel-head trout -1Salmo
-43-
gairdneri) the region of the lateral line was found to be
least subject to varistion., in scale growth.
Referring to Table VII, fish No. 1, 23 months
old, length 11.5 inches.
From this specimen are recorded
the readings on 273 scales as listed, regions "E", "F",
"G" and "H" showing by far more scales counted than "A",
"B", "0", and "D".
The reason for this is at once appar-
ent from the size of the areas in question, they being
nearly twice that of the others.
This same condition does
not hold throughout this table and the next, as the scope
of the work was materially changed at this point, and but
ten scales from each area were taken.
Throughout the work
an attempt was made to select for apparently perfect
scales; those with frayed edges, or with nuclear rings
missing were discarded.
Fish Lo. 1 shows the greatest range of variation in the ring count for the three larger specimens,
the count ranging from 35 to 56, but with the greatest
number of rings in the vicinity of 43 to 46.
Fish No. 2 has a variation of from 35 to 49,
with a slight majority about the count of 40 to 44, while
No. 3, the largest of the group, seems to hold its majority about 44 to 47.
It seems rather interesting to note the position
assumed by the scales from region "E" as this formed the
base of operations in the work of the present writer, al-
d cher9 FiSh
Ase
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-44-
so of Dahl.
In the first two cases especially the group-
ing of the rings of this area about the majority count
for the whole fish is cuite striking.
Another fact
brought out in examining the scales, but which is not evident from the tables presented is the relative degree of
perfection exhibited by those scales from region "E" as
Scales from
compared with that found over'other areas.
other regions particularly "F", "G"
and "H" were found
to be greatly affected with abnormalities and injuries
such as double nuclei, whorls in the body of the scale,
punctures and blank centers.
In some cases the predomi-
nance of such conditions was so marked as to offer con-
siderable difficulty in finding scales whole enough or
perfect enough to count.
With the exception of the disappearance of the
nuclear rings, a condition found auite generally over the
entire body of all the fish, the region designated as "E"
in all cases was found to be remarkably free from other
inequalities or abnormalities of growth.
This statement
the
is not made on the examination ofAfirst ten scales from
each area, but from a much larger number, varying in individual cases; sometimes as many as thirty scales were
examined before ten were chosen as perfect.
Doubtless
the loss of nuclear rings is due to the rapid degeneration of the integument -upon removal from the water.
It
may be that hatchery fish are never as perfect in this
respect as the naturally hatched specimens; but further
-45-
investiga ion only will settle this point.
Region "DP along the lateral line also shows
signs of adhering closely to the mean ring count for the
individual, but abnormalities and injuries are rLther
frequent; the other lateral line areas
arc more
variable.
After all might we not expect just this condition
to hold true
because of the relation of these vari-
ous areas to the contours of the body?
Thus region "F",
chosen by Calderwood in taking scales for study and called by him the "shoulder", is that which
naturally
first
comes in contact rith obstructions and which would tend
to show
the
effect of these forces.
Again, the ventral
portions "G" and "H" are continually at
the mercy of
the
rough character of the bed of the pond or stleam and
might be expected to respond to the wear and tear thus
encountered.
It is not necessary to determine the one best
region for taking scale samples, but rather to make certain of what regions .cannot be relied on for this purpose,
and what limits we may safely set and so avoid undue variations.
With
facts presented
this
idea in mind, and in face of the
it seems safe to say that ideal condi-
tions may be found
within the limits indicated.by regions
"A", "C" and !!E".
While working with the six
a very interesting
condition
hatchery raised fish
was found to
exist as regards
-46-
the relation of the
size of fish No. 4 to its ring count,
especially when this relation was compared with the same
data from No's. 5 and 6 on the other.
either 11.5 or 12 inches in length, the
and 8.5 inches respectively.
The former
latter
The ring-count of
are all
are 8.25 and
the
larger
three varies from 35 to 56, that of the smaller two from
19 to 34.
The fish in nuestion, No. 4 is 9.75 inches long
and presents a ring-count which zig-zags back and forth
between 29 and 50.
These facts would seemingly tend to
place this fish in a position intermediate between the
age
others as regards
to this factor.
if
the number
of rings is any index
But this seems quite out of the question
in-as-much as all the eggs of any one kind in a hatchery
have given up their fry before so great a difference of
time could elapse.
Moreover this iish was taken from a
pond which viLs supposed to contain only 14 months old
fish and the possibility of
getting
these mixed with oth-
ers is very slight if the work is painstakingly done,
which I believe to be true at the station under observation.
If, as the evidence seems to show, this fish is
really one of the older group, why does it not more
closely approximate their measurements?
the
In appearance
specimen is typically one of the younger fish, lack-
ing the
rounded body and well proportioned parts of the
more imture types.
It seems as though one of the older
fish placed in a pond with a number of others nearly a
-47year yuunger would
have such a decided advantage over its
fellows as to develop into an over father than an undersized individual.
Allowances must of course be made for
some factor which might influence this individual to dev-
elop
abnormally.
On the other hand, however, if the fish be of
the same age as those of the smaller size, the statement,
that as an index to
age, the
ring-count is praqtically
useless, seems to be well grounded.
The Seasonal Bands as an Index to Age.
So far the work has
lation
dealt
largely with the re-
between the ring-count and other factors as a means
of solving the problem of age determination.
Little or
nothing has been said of the application of the seasonal
bands of
writer
Johnston to this principle.
At the outset, the
feels bound to state thet but little has been done
as yet with this phase of the work owing chiefly to the
difficulties encountered.
Sometime, during the past winter, Mr. Rich was
kind enough to show me come very good photographs of salmon and steelhead trout scales which showed these bands
very plainly.
Excellent
photomicrographs may
also be
found in Dr. Gilbert's paper on this subject and quite unwittingly many of the illustrations in ljalloch's book af-
ford beautiful pictures of the baLding.
-48-
At this laboratory, however, considerable difficulty has been experienced in trying to demonstrate the
presence of the bands, due either to imlroper illumination or lack of experience in discerning
them.
This lat-
ter point is probably the main reason for the failure, as
Mr. Rich pointed out the fact that it did take considerable application and experience to be
able to observe
them readily.
however, observation on the hatchery fish has
shown that the majority of
the scales
narrow and broadly spaced rings.
exhibit hands of
The hatchery fish used
were all killed in December and those scales which are
not injured or malformed show a region of narrow spaced
rings about the periphery, a condition bearing considerable significance, cs it goes to show tha, these bands
may be formed at this time. (See Plates IV, V & VI)
But
we cannot say from the evidence at hand that the winter
season is the only factor which will cause a similar
growth.
Difficulty was expexienced in demonstrating two
such regions in the twenty-three months old fish, but
believe that
further work with material in better condi-
tion will result in more satisfactory findings.
If each
winter is to leave a band of narrow spaced rings to mark
the growth during that season, the Presence of tzo of
these bands may be reasonably expected to be present in
fish of sufficient age.
Perhaps, however the conditions
-49-
existing in the hatchery pons were so even throughout
the year, estecially as regards the food supply, that
growth proceeded so regularly as to preclude the necessity of retarding development and Tiving a chance for a
band of narrow rings to aprear.
What effect may be produced in the scales by
the various changes possible to create in the fishes' env;ronmentIcan only be determined experimentally.
Other Observations.
While the examination of the scales at hand proceeded, my attention wal
called frequently to abnormal
growths and signs of degeneration which might materially
affect the ease and precision of the 'work.
The most common variation of this nature was
found among the spawning fish and the hatchery fish and
c nsisted of the loss of the nuclear rings thus leaving
a large bare spot in the center of the scale.
In many
cases not more than two or three rings were left about
the periphery.
This is a very annoying feature and great-
ly reduces the rapidity of the work if the condition is
very common.
:lito-micrographs of
such scales may be
found on Plates IV, V, & VI.
Spawning fish also loae many of the peripheral
rings, the result being thatclear margin surrounds the
ringed portion. (See Plate VII).
-50-
Another condition peculiar to spawning fish is
found in the presence of a dense callous growth which
frequently obscures a large portion of the scale.
This
must not be confused with the raised line which traverses
the long axis of scales covering the lateral line organs.
The rings outside this dense area are usually distorted
and have lost their definite concentric arrangement.
The power of regeneration among the scales
seems to be quite marked.
Scales were repeatedly ol?serv-
ed which had evidently been punctured at a point outside
the nucleus.
In most cases this puncture was filled in by
a new growth showing distinct ring marks resembling a
secondary nucleus or the remainder of the scale.
Such a
condition was particularly noticeable in scales taken
from the 7shoulder" region, and may be found represented
on plate IV, figure 3.
All scales, except those from very young fish,
show that development does cot proceed by the addition of
rings about the entire periphery.
is readily seen thLt the
ring-count
With the long axis, it
varies from that of
the broad; this gives the scale its somewhat elongated
form.
It seems probable from this that the growth is the
more rapid over the long axis, time not
being
given for
the development of complete ring.
Among the more common of the malformations is
that of the development of a double nucleus about which
-51may be formed a double series of rings sometimes extend.-
ing to the periphery. Such a type is shown in plate IV,
figure 4.
-52Summary of 1-esults.
Lo direct correlation was found to exist
between
the ring-count of salmon scales and the length of
the individual from which removed.
spawning
The determination of age at the
grounds is subject to great error owing to the amount of
degeneration
undergone by
the entire organism.
The number of rings
found on a
single scale
or on a series of scales from the same individual is not
a direct index to the age of
the
fish.
In line with "3" above, the assessment of
age for our Pacific
halloch scheme of
Coast salmon cannot be based on the
sixteen annual rings.
110 proof has been found experimentally that
definite ring bands are formed daring the warm and cold
seasons of the year.
Accurate determination
of
age cannot be
based on the study of any one scale, because of the liability o: malformation or degeneration.
It is necessary to have
known age to serve as a check on all
adult
material of a
results.
-53-
Acknowledgments
The writer wishes to thank Prof. G. F. Sykes,
Prof. P. J. _:raus, Hr. H.
.
Clanton, 11r. h.
J. Kinney
nd Lr. H. I. lich for their many kind acts of assistance
and advice in the preparation of this paper.
He is also
indebted to Hr. A. J. Stover for invaluable guidance in
the preparation of the accompanying plates.
Literature Cited
Calderwood,
The Life
of the
Salmon. London, 1910
Cockerell, T. D. A.
Observations
on Fish Scales, in Doc. Eo 779
U. S. Bur. Fisheries, p.125
Ibid.
The Scales of Fresh-water Fishes.
Bulletin, Vol.
In Biol.
FX, Lc). 6.
Ibid.
The Scales of some American Cyprinidae. In Proc.
Biol.Soc. of Washington XXII, 157-164, July 28,
1909.
Dahl, Knut
of the Salmon and Trout of
Norway as shown Isy their scales. English transletion by Iaan Bail_e for the London Salmon
The Age and Growth
and Trout Association.
London, 1910.
Gilbert, C. H.
Age at Maturity of the Pacific Coast
the
Genus Oncorhynchus.
Salmon of
In Doc. No 767, U. S.
Bur. of Fisheries.
Ibid.
The Salmon of the Swiftsure
'River Sockeye Run
of 1912.
Bank and
the
Frazer,:
In Report of British
-55-
Columbia Commissioner of Fisheries, 1912.
Greene, C. W.
The Storage of Fat in the Muscular Tissue of
the King Salmon and its
"Aeabsorption During
the Fast of the Spawning Migration.
In Doc.
No. 799, U. S. Bur. of Fisheries.
Heincke, F.
Naturgeschichte des Herrings.
Abstract from
Abb. d. Deutch. Seefischereiverein 1897-98:
in BinfuhrlIng
in die Vererbungswissenschaft
(P. 82). H. Goldschmidt. Leipsig 1911.
Description of Plates
HATE I
Fig. 1,
Parr 10 c.M. Hatched 1898, killed
Nov. 8, 1898.
Pig. 2, Parr 13 c.m. Hatched. April 1907, killed
Dec., 1908
Fig. 3, 'Darr 13 c.m. Hatched
April
1907, killed
Dec., 1906
Fig
'Fig.
Fig.
4 Parr. 12 c.m. 6ofteland May 25, 1909.
5, Parr 10 c.m. June 11, 1908
6, Parr 12 c.m. Softeland, miAle of Oct.
1908.
From Dahl, H. W. H. del.
PLATES II & III
Opercula from spawning salmon, showing scars
left by wound claimed_ to have been caused by Mr. Clant-
on's gill punch.
PLATE IV
Fig. 1, ,,cale from Hatchery Fish .1,o. 1, Region
HATT
2, scale from Hatchery Fish 1,o. 1, Region
Fig.
3, Scale from Hatchery Fish Ho. 1, Region
Figs. 4, 5, 6, Scale from Hatchery Fish Eo 1
Region "F".
PLATE
Fig. 1, Scale from Hatchery Fish :o. 3, Region
"Hu.
Fig. 2, Scale from Hatchery Fish, Ho. 3,
Fig. 3, scale
Eegion
hatchery Fish No. 3,
Hegion
Fig. 4 Scale from Hatchery Fish No. 4,
EePAon
from
T11 TI
Fig. 5, Scale from
Hatchery Fish No. 4, Region
PLATE VI
Fig. 1, Scale from
11
B
Hatchery Fish Ho. 5, Region
I
ScLle from Hatchery Fish No. 5, Legion
21g.
3, Scale from Hatchery Fish Ho. 5,
Region
Fig. 4, Scale from Hatchery Fish No. 4,
Region
Fig. 5, Scale from Hatchery Fish iTo. 5, Region
Scale from
Hatoholy Fish Ho. 5,
Region
PLATE VII
Scales from sicawning fish shoring condition
exhibited. by the majority of the scales taken from the
fish with marked opercula.
tv,-(
PLATE I.
-
PLATE II.
NO .
7
NO
5
.
NO . 2
NO . 1
NO .4
PLATE III.
NO 3
4
No.6
mo .8
PLATE IV.
Fig. 2
Fig .1
Fig.
3
Fig,5
Fig.4
Fig. 6
PLATE V.
Fig.2
Fig.I
Fig. 3
Fig.4
Fig. 5
PLATE VI.
Pig.I
Fig.2
Fig.3
g . 4-
,
Fig.5
Fig
PLATE VII',
11?
Fig.2
rig. :t
Fig.3
4,- --
--%..
.,..,
-
.
.
,
1,
Pig .4
Fig.5