Introduction

Mortality and
Survival. In our early lectures about life-history variations and
life tables, we included survivorship schedules as one
essential part of looking at year-to-year survival.
The exercises in Homework 1 were fairly
sophisticated, because survivorship wasn’t constant
K. Limburg lecture across age classes, resulting in variable recruitment.
notes, Fisheries
Science & Management So, what is recruitment, really?
It’s a term you will hear repeatedly in fisheries
Mortality in fisheries biology. biology and management…
We’ll define it and look at some ways to estimate it.

Recruitment…
• Means an individual fish survives into a defined
life-stage Biologists define recruitment more broadly, i.e., fish
surviving to a given life stage.
• First defined in a fisheries context, that is, when a
fish grows large enough to be vulnerable to a
particular gear – then it is said to have been recruited
to the gear.
• Similarly, one can say that a fish of a given size (or
age) recruits to a fishery – this assumes that similar
gear is used throughout the fishery
For example, many fisheries biologists concern themselves
with survival from egg  larval or larval  juvenile stages
(“larval recruitment,” “juvenile recruitment,” etc.)

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 As we saw earlier,
We speak of recruitment failures, when a given year- # of mortality is highest in
class has poor survival fish
the youngest, most
So, one of the key biological/ecological factors we sensitive stages.
study is mortality, and the sources thereof. Survival of a cohort
looks like this.
Identifying and quantifying mortality has occupied
Age or time
fisheries scientists for more than a century, and will
likely remain a major study.
If the population is at steady-state (births = deaths), then this
Here, we’ll get introduced to some of the basic pattern will hold true for the entire population.
ways to estimate mortality, and the flip-side,
survival. The curve above sort of looks like the time sequence of water
in a bath-tub when you pull out the plug…

As the water drains from
the tub, its rate is limited
by the size of the drain
hole, and by the mass of
water remaining to drain
out…
It’s just about the simplest differential equation
that there is:
Let N = the number of fish. Then, the rate of
change of fish over time is described most simply
by the equation

The simplest model to describe this “draining” is often
called the “bath-tub model” or the exponential decay
model.
…and you have seen it before…!
dN/dt = -z N
Remember that “d” means “a very small change,”
“dt” means “a very small increment of time”
z is the coefficient of instantaneous mortality.

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 This is easily solved by integration, as we did the t t

  z dt   z  dt
second week of class.
r.h.s. :
dN 1 0 0
  zN  dN   z dt
dt N   z  t  (  z  0)   zt
Integrating both sides within the limits [0, t], we get:
Put the left hand side and right hand sides together:
t
1
 N dN  ln( N ) ]
t
l.h.s. :
0 ln( N t )  ln( N 0 )   zt
0

 ln( N t )  ln( N 0 ) ln( N t )  ln( N 0 )  zt

Take antilogs:
What does all this
ln( N t )  ln( N 0 )  zt mathematical
manipulation mean?
eln( N t )  eln( N 0 )  e  zt
 N t  N 0  e  zt
Neil Ringler, Dean of
Research and former
Thus, if we Nt teacher of Fisheries
know Nt and  ln( ) Biology, had a nice way of
N0, we can N0 putting it:
calculate z : z
t

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Imagine a year being broken up into lots
of small time intervals…n of them.
Then, z/n would be the fraction of the
population that dies during the nth period
of the year.
During the next interval, (1/n)
99,950 fish x 0.0005 = 49.975 fish die,
and 99,900.3 fish survive.
And so on…
For example, z = 0.5, n = 1,000 intervals
During 1/1000th of a year, then,
0.5/1000 = 0.0005 of the # of fish
present die.
If we start with 100,000 fish, then
during the first interval (1/n),
100,000 x 0.0005 = 50 fish die
and 99,950 fish live.
If you do this 1000 times over the year,
it’s like compounding interest in a bank–
only this is like having a negative
interest rate!
# of survivors = (1-(z/n))n x N
= (1 – 0.0005)1000 x N = 60,645
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 A few more interesting facts here,
folks:
Let S = the fraction surviving that year S  (1 – (z/n))n
= 60,645/100,000 As n  
= 0.60645 (as you chop up the year into an infinite
number of time intervals),
The annual rate of total mortality is
S  (1 – z/(n))n
A = 1 – S  0.394
= e-z

S = e-z

Q: how do we estimate z from data???

Constructing a catch curve - raw data

We do this by constructing a catch curve. 240 We can see that the first 2 age-
classes are under-represented
In this method, we assume that mortality is 190 (not fully recruited to the fishing
gear)

Number in sample
relatively constant from one age class to the next.
140

(Note – this works best on post-juvenile fish)

90
We can determine the # of fish in each age-class
40
by scales, otoliths, opercular bones, etc.
Then we make a graph of the number of fish in -10
0 2 4 6 8 10 12
each age-class, vs. the age-classes: Age (years)

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 If we log-transform the #s of fish per age-class, and re- The usual caveats:
plot the graph as a semi-log plot, we see that the
numbers of fish per age-class form a more-or-less If the assumptions Unbiased
straight line: about the population
are wrong (i.e., not
Constructing a catch curve closed, unequal gear Larger fish
6.00
We can fit a line vulnerability, etc) then migrating away
5.00
through the you will under or
overestimate z and S
points (ignoring
LN(Number in sample)

4.00 Ages 1 and 2)

by linear
3.00
regression.
2.00 Then, the slope
1.00
of the

0.00
regression is -z Larger fish more
0 2 4 6 8 10 12 vulnerable to gear
Age (years)

3. Finally, if M is
Last bits of definitions: very important! known, and you have a
target instantaneous
1. For fish populations, the total
mortality level (z), then
instantaneous mortality, z, is defined as
you can set F to meet
z=F+M that target.
where F = Fishing mortality This is precisely what
fisheries managers
M = Natural mortality
spend so much time
2. The total annual mortality, A, is defined as doing – setting the level
of fishing effort to
A = 1 – S = 1 – e-z meet a given total
 If you know S, you can calculate A. If you mortality level…and it
know Z and M, you can estimate F. can be VERY
contentious! Photo: Nando Times

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 Age: 2 3 4 5 6
Some other common estimation techniques: Nx : 150 95 53 35 17 sum = 350
Code
Robson & Chapman's method for estimating survival (S^) 0 1 2 3 4
class:
1. List out fish abundance by age class:
With this numerical example:
Age: 2 3 4 5 6 Call the row
Nx : 150 95 53 35 17 sum “n” T = (0•150) + (1•95) + (2•53) + (3•35) + (4•17) = 374
2. Assign a code class to each age class: n = sum(Nx) = 350
Code Call this
class: 0 1 2 3 4 variable “x” S^ = 374/(350 + 374 – 1) = 0.52
Var(S^) = 0.52 (0.52 – (374-1)/(350-374-2) = 0.0018
T
Estimate of Sˆ  , where T   ( x  N x )
survival is: n  T 1 Another way to ( N1  2 N 2  3N 3  4 N 4 )
write this Sˆ 
T 1 equation is: ( N 0  2 N1  3N 2  4 N 3  5 N 4  1)
Var ( Sˆ )  Sˆ ( Sˆ  )
n T 2

Weighted Formulae: assume

• recruitment is equal from one year to the next Heinke’s formula – used when it’s hard to
determine the age of older fish (big
• equal survival rates -------- “ --------------- problem with scale aging)
• equal vulnerability to the sampling gear
• provides a weighting proportional to the most
abundant age classes
N all age classes  N youngest , fully recruited
S
ˆ
N III  N IV  NV  NVI N all age classes
Jackson’s Sˆ 
formula: N II  N III  N IV  NV  NVI

Start w/first fully

recruited age-class

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 Wrapping up mortality with a few words about
fish larvae, and maximum age
Early life stages of fish are pretty vulnerable…
- can starve
- can be eaten
- can be damaged by turbulent eddies
- can be advected out of the nursery area

Johan Hjort was an early fishery scientist who

realized the importance of larval survival

Figures from Jones et al (1978)

Hjort, J. 1914. Fluctuations in the great fisheries of

northern Europe viewed in the light of biological Match-mismatch hypothesis: Cushing (1974)
research. Rapports et Procès-Verbaux Des Réunions, suggested that the match, or mis-match, of larval
Conseil International pour l’Exploration de la Mer 20: fish occurring together with their food determined
1-228. whether they fed or starved
Proposed that recruitment variation was caused by
the transport of larvae away (advection) from nursery
grounds
In a variation of this, Lasker (1981) showed that
northern anchovy hatching off California starved
during unstable oceanic conditions  oceanic
stability hypothesis of larval survival
Figure from Jennings et al (2001)

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For eggs and larvae, size Here is an
plays a big role in empirical way
determining their to estimate
survival…this led to the mortality rate,
“bigger is better” hypothesis based on the
(Houde, 1987), that larger maximum age
larvae had a better chance of reached by a
avoiding predators. species…

But obviously there is a cost This was

to growing fast (finding published by
enough food, e.g.) John Hoenig in
1983.

Figure from Jennings et al (2001)