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The life histories of all Pacific
salmon species begin with egg development and hatching in freshwater,
and continue with migration, feeding, and growth in the freshwater and
marine environments of the Pacific Northwest. After a period of time,
the salmon to begin to sexually mature, and they commence spawning runs
to their home streams. The timing of maturation is controlled by internal
genetic codes in combination with the effects of environmental conditions
throughout the life of the fish. The spawning process completes the life
cycle (ending in death) of Washington's five species of Pacific salmon.
The pattern of ages at which this occurs are specific to each species
of salmon, however, the age at which individual fish sexually mature and
return to spawn is not fixed, but can vary from fish to fish (termed an
"indeterminate semelparous" life history).
These "age at return" patterns
vary from simple (virtually all pink salmon return as 2-year old fish)
to the very complex (chum salmon can return as 2-, 3-, 4-, 5-, or 6-year
old fish). In the case of chum salmon, offspring from a single spawning
year (called brood or brood year) can mature and return to their natal
streams from 2 to 6 years later, although 2- and 6-year fish are very
rare. This creates complex biological assessment and management issues
for each of the Pacific salmon species that have variable ages at maturity.
To effectively manage salmon, it is important to estimate age compositions
of major stocks each year.
Aging of returning salmon is
typically accomplished by counting the annual growth bands on scales or
otoliths (small bones in the head) collected from fish caught in fisheries
operating specifically on the stocks to be aged. When there are low numbers
of returning fish, or protective harvest regulations as in recent years,
there may not be sufficient numbers of salmon caught in local fisheries
to provide the opportunity to sample scales in fisheries at acceptable
levels for age determinations. When this happens, scales and/or otoliths
can be collected from salmon carcasses sampled in the spawning streams
or at salmon hatcheries.
Stock Productivity - Probably one of the most important uses for age data in salmon management is to assess
the basic productivity of a population, a value that is often expressed
in terms of adult fish produced (recruits) per parent spawner. This process
is of course more difficult in salmon populations where the offspring
will return from a single parent spawning over the course of several subsequent
years. As an example, it may be necessary to figure out how many adult
chum salmon were produced from a hypothetical parent spawning escapement
of 50,000 chum salmon in 1995. Chum salmon commonly return at either age-3,
-4 or -5 years old. Scales would be collected from fish returning to spawn
in the years 1998 (3 years later), 1999 (4 years later), and 2000 (5 years
later) in order to do this analysis. Each year there is a commercial chum
fishery that harvests fish from our hypothetical chum salmon stock. Scales
taken from a portion of these harvested fish are analyzed to determine
the age composition of the catch. These samples will represent the annual
age composition of our hypothetical run of chum salmon. In the following
simplified approach, the result of scale data from sampling 1,000 fish
each year is as follows:
Return
year |
Number
age
3 |
Number
age
4 |
Number
age
5 |
Total
sample |
| 1998 |
300
(30 %) |
600
(60 %) |
100
(10 %) |
1,000 |
| 1999 |
700
(70 %) |
250
(25 %) |
50
(5 %) |
1,000 |
| 2000 |
400
(40 %) |
500
(50 %) |
100
(10 %) |
1,000 |
If we consider total fishery
catches plus the spawning escapement to represent the total runsize returning
from the ocean, then application of the age sampling data in the above
table will allow estimation of the total number of adults produced from
the 1995 spawning escapement.
Return
year |
Total
Runsize |
%
age
3 |
age 3
adults |
%
age
4 |
age 4
adults |
%
age
5 |
age 5
adults |
| 1998 |
100,000 |
30 |
30,000 |
60 |
60,000 |
10 |
10,000 |
| 1999 |
200,000 |
70 |
140,000 |
25 |
50,000 |
5 |
10,000 |
| 2000 |
150,000 |
40 |
60,000 |
50 |
75,000 |
10 |
15,000 |
| Note:
The highlighted numbers
represent the returns of 1995 brood fish in 1998, 1999, and
2000. |
The total number of adult chum
salmon produced from the 1995 parent escapement will therefore be the
1998 age-3 return + the 1999 age-4 return + the 2000 age-5 return from
the above table.
30,000
+ 50,000 + 15,000 = 95,000 fish
Since our hypothetical chum
salmon run had 50,000 parent spawners in 1995, the total adult production
(recruits) and recruits per spawner (R/S) from the 1995 brood can then
be derived by the following calculations:
Total age-3+4+5 fish returning ÷ parent
1995 escapement =
95,000
recruits ÷50,000 spawners =
1.9 recruits / spawner
This type of analysis is conducted
every year on many Washington State salmon populations. Over time, a basic
understanding of the productivity of each salmon run is developed.
Runsize Forecasts - Another use of salmon age data is to predict future returns of fish
so various fisheries can be managed to allow the desired numbers of spawners
to reach their spawning streams. There are many approaches to forecasting
salmon returns, however, most are variations of two methods, survival
rate and sibling forecasts. For chum salmon, both forecasting methods
require reliable age data.
A simple survival rate forecast
uses the average number of salmon that return in each age category from
each spawner in the parent year. Assume a salmon population that had an
average total return rate of 3.0 fish (recruits) per spawner: 1.0 recruit/spawner
(R/S) as age-3 fish, 1.75 recruits per spawner as age-4 fish, and 0.25
recruits per spawner as age-5 fish. This brood age composition ratio (1.00:1.75:0.25;
or 34% age-3, 58% age-4, and 8% age-5) is termed the "maturity schedule"
for this specific population of salmon. To predict the return in year
2001 the number of parent spawners in 1998 would be used to predict the
3-year old return (1998 spawners x 1.0 R/S), the 1997 parent spawners
would be used to forecast the age-4 return (1997 spawners x 1.75 R/S),
and the 1996 parent spawners would be used to forecast the age-5 return
(1996 spawners x 0.25 R/S). The estimates of returning age-3, age-4, and
age-5 fish would the be added together to predict the year 2001 return
for this particular salmon population.
The sibling forecast uses the
earliest returning fish from each spawning year (typically 2- or 3-year
old fish) to predict the subsequent returns of the older fish (typically
age-3 or older). To show how this works, assume a salmon population that
has an average age composition of 40% age-3 and 60% age-4 fish (based
on ages from the same broods). If the 4-year old fish were returning this
season, and 1,000 3-year old fish from the same spawning year had returned
the previous year, a sibling forecast for 4-year returns would be:
4-year
returns = [(1,000 x 0.6) ÷ 0.4] = 1,500 salmon
Or, a simpler way of looking
at the problem is that on average, there will be 1.5 times as many 4-year
fish as 3-year fish (0.6 ÷ 0.4) in this hypothetical salmon population.
The sibling forecast for 4-year returns could also be calculated:
4-year
returns = 1,000 age-3 recruits x 1.5 = 1,500 salmon
Risk Assessments - Based on the age compositions in the above sibling forecast example
(40% age-3 and 60% age-4 fish), the average age at return for this hypothetical
salmon population would be 3.6 years. The average return age can be used
for assessments of the risk of extinction faced by a stock of salmon.
A calculation of "total population size" can be made by multiplying the
average number of spawners in a population times the average return age.
Total population size is an estimate of the total number of spawners that
will result from all of the individual fish alive at any given time (representing
production from all current brood years). If this number drops too low,
the population can be at risk of extinction.
Evaluate Mortality
- Another important use of survival estimates derived from age
and return data is to measure the impacts of various limiting factors
on a salmon population. Factors like flooding, urbanization, and the effects
of dams or logging can be related to the survival of salmon populations,
and recovery efforts can be developed if negative impacts are identified.
Most chum salmon are aged by
analyzing scales collected from three sources: 1) fish caught in commercial
fisheries, 2) fish sampled at hatchery racks, or 3) from carcasses in
spawning streams. Otoliths (ear bones) are rarely used for aging chum
salmon; but have occasionally been used in programs designed to monitor
and evaluate fish returning to artificial production facilities.
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Scale selection area
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Scale Sampling - Specific methodologies are used by the state and tribes to collect salmon
scales as a part of annual biological sampling programs. Scales are collected
from a "preferred area" of the fish. The preferred area is on the side
of the fish from the second to the seventh rows of scales above the lateral
line and on a diagonal from the back of the dorsal fin to the front of
the anal fin. Two separate scales are collected from this area within
10 scales in front or behind the diagonal line. The scales are removed
from the fish with forceps, and are placed on a gummed scale card. Information
for each fish is also recorded on the card with the scales including:
date, fishing area, gear type, and the species, sex, and length of the
fish.
Analyzing the Scales - The scale cards hold the scales samples from 20 individual salmon. In
the age analysis lab, a combination of high pressure and heat are used
(with a hydraulic press) to create impressions of the scales in strips
of clear acetate. Multiple sets of scale impressions can be made. The
scale impressions on these plastic strips are an archival record of the
scale samples that can be freely handled and will not deteriorate over
time.
The process of determining
the age of a fish from scales is called "scale reading". The scale impressions
are magnified by a factor of 24X and are viewed on a rear projected screen
(using a microfiche reader). Salmon ages are determined by examining patterns
of rings (called circuli) that are laid down on the scale as
the fish grows. The development of these growth rings is similar to tree
rings; during periods of rapid growth the rings are widely spaced, and
when growth slows the rings are more tightly spaced. Salmon grow rapidly
in the summer months when water temperatures and food availability are
highest, and they experience slower growth during the colder winter months.
Salmon scales typically exhibit alternating bands of widely spaced (summer)
and narrowly spaced (winter) growth rings. The winter bands of circuli
are called annuli, because the end of each winter season generally
corresponds to the completion of another year of life.
Chum salmon are relatively
easy to age because they spend nearly their entire lives in marine waters,
and do not have the complicated freshwater growth patterns associated
with some other salmon species. Reading a chum salmon scale is simply
a matter of counting the bands of widely spaced circuli representing summer
growth. Using the age-3 chum salmon scale shown below as an example, the first summer is represented by the widely spaced circuli in the
center of the scale, the second summer by the band of wide circuli
spacing between the first and second annuli, and the third summer by the band of circuli outside of the second annulus. At the time this
chum salmon was sampled, it was nearing the end of its third summer of
life. If the fish was a maturing adult or was sampled on the spawning
grounds, it was completing its life cycle at the end of its third year.
Four and five year fish are aged in the same manner (see examples below).
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