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Abstract.
Purpose of this article is to give insight in the Mayfield method of estimating the daily survival
rate. We find from literature that little is known of the statistical behaviour of the Mayfield
method and its use is mainly ground on experiences and simulations.
This article provides the mathematical framework for the Mayfield method. The concept of
observation interval plays a central role in the statistical model. During this interval we observe
a clutch. At the end of each observation interval we decide if the clutch is destroyed or not.
In terms of the statistical model we define two Mayfield estimators: F = 1 - B/A and G = A/(A
+B). We investigate two variants of each estimator. We show that the Mayfield estimator has
approximately a normal distribution. We give equations for the expectation and variance. We
analyse the behaviour of the Mayfield estimator and we compare the various variants. We also
give the exact probability distribution of the Mayfield estimator and equations to calculate the
exact expectation and variance. It turns out that the Mayfield estimator has a bias: on average
the result has a bias with regard to the real daily survival rate. Mostly the bias is negligible.
Especially for small daily survival rates the bias can have a significant influence on the
estimate.
The Mayfield method aims to estimate the nest success or hatch rate. For each Mayfield
variant we define an estimator for the hatch rate. These estimators also have approximately a
normal distribution. The bias of the daily survival rate implies a bias for estimating the hatch
rate. The bias is largest if the hatch rate is small.
We present an alternative estimator for the daily survival rate and for the hatch rate. This
estimator has approximately a normal distribution. The benefit with regard to the Mayfield
estimator is that this estimator is (asymptotically) unbiased. We find that this alternative
estimator performs better than the Mayfield estimator.
Key words.
Mayfield method, estimator, daily survival rate, nesting success, hatch rate, breeding bird,
reproduction, sample size, survivorship, central limit theorem, probability distribution,
expectation, variance, bias, confidence interval, statistics.
1. Introduction.
Ornithologists are interested in estimating the nest success. One of the methods for estimating nest
success is the Mayfield method. The daily survival rate plays a key role. (Mayfield 1961) formulates an
estimator for the daily survival rates of nests. This estimator is widely used because of its simplicity.
In this article we formulate characteristic properties of the Mayfield estimator. We present formulas for
the expectation and the variance of the (asymptotic) probability distribution of the Mayfield estimator. The
mathematical details are worked out in the appendix.
This article aims to give insight in the behaviour of the Mayfield estimator and in estimating nest success.
In practice one uses a number of variants of the Mayfield estimator. We compare four different variants
and give an alternative estimator.
2. Definitions.
2.1 Concepts.
A nest in the stage of first egg laying until the hatching of the first young we call a clutch. The nest
success or hatch rate of a clutch is the probability that the clutch is successful; that is the probability
that the first young leaves the egg. The hatch rate is dependent of the species and the influence of
predators. Also other factors such as disturbance or weather influence the hatch rate.
The daily survival rate is the probability that a clutch survives the period of one day. The Mayfield
method assumes that the daily survival rate is the same for all the days the bird is incubating. The
duration of incubation includes the laying stage.
If p is the daily survival rate, then the hatch rate is: pH = pL. The variable L is the duration of incubation.
The value of L is for every bird species roughly the same. It varies from 20 days for small birds until 40
days for large birds. Sometimes one includes the nestling stage; then L is larger.
We divide the observation intervals of a sample in groups of equal length. We use the following notation:
N: the number of groups (N≥1).
m[i]: the length of the observation interval (days) of group i (i = 1..N and 1≤ m[i] ≤ L)
n[i]: the number of observation intervals of group i (i = 1..N)
S[i]: the random variable of the number of observation intervals of group i for which we have found no
destruction (i = 1.. N and 0 ≤ S[i] ≤ n[i]).
We use bold capitals to indicate random variables.
S[i] has the binomial distribution ß(n[i];pm[i]); i = 1..N with expectation E(S[i]) = n[i].pm[i] and variance
Var(S[i]) = n[i].pm[i].(1− pm[i]).
A is the total number of survival days of the examined clutches and B is the number of clutches that are
destroyed. In terms of the preceding statistical model we can write:
N
(2.2) A = ∑ ( m[i] S[i] + r(m[i]) ( n[i] − S[i] ) )
i= 1
N
(2.3) B = ∑ ( n[i] − S[i] )
i= 1
The value r(m) of the function r is the estimate of the number of survival days of a destroyed clutch if m
is the length of the observation interval. The case r(m) = ½ m is called the “midpoint assumption”.
As an illustration we show the data of a fictitious research to the daily survival rate. The values of A and
B are calculated with the midpoint assumption. The result of F1 is 0.951 and the result of G1 is 0.954.
Table 1: Data of a fictitious research project to nest success. Number of groups: N=3.
The results of S[i] are random generated and based on a daily survival rate of 0.95.
Calculation with r(m) = 0.5m.
Design of experiment Random Calculation
i n[i] m[i] S[i] m[i]S[i]+0.5m[i](n[i]-S[i]) n[i]-S[i]
1 110 5 90 500 20
2 120 7 93 745.5 27
3 100 10 50 750 50
A = 1995.5 B = 97
3. Results.
We notice that the variance decreases if n increases. As we shall see also an increasing m-value leads
to a decreasing variance. The asymptotic expectations µF and µG are independent of n.
0 0 0 0
0 0.2 0.4 0.6 0.8 1 0 0.2 0.4 0.6 0.8 1 0 0.2 0.4 0.6 0.8 1 0 0.2 0.4 0.6 0.8 1
Figure 1:
Graphs of the asymptotic expectation µ of the Mayfield estimators F1, F2, G1 and G2 as a function of p.
Each graph shows three curves: for m=5, 10, 20 (the larger m the larger µ). Parameters: N=1; n=100.
To reduce the number of parameters, we have chosen N=1 in figure 1. We assume that this gives a
representative picture of the general case with N>1. Calculations show that this is correct.
Figure 1 shows the development of the expectations of the estimators F and G as a function of the real
daily survival rate. The difference with the straight sloping line is the bias with regard to the real daily
survival rate p. We see the bias is smaller according as p is closer to one. It attracts attention that as p is
smaller, the estimator differs more from the real p. The estimator reaches its minimum as a limit for p→0.
In spite of this one uses the Mayfield estimator because small daily survival rates occurs not frequently.
If the daily survival rate is 0.9 and the duration of incubation is 30 days, then the hatch rate is 0.9 30 =
0.042. Mostly this hatch rate is too small for a population of birds to survive. Yet we have to take into
account with small daily survival rates, for example if increasing predation takes place temporary or
local.
Because the estimator F is normal distributed (for n large enough), 95% of the results of F are inside the
interval: [µF - 1,96σF , µF + 1,96σF]. We can calculate the 95% upper and lower bounds with the equations
for µF and σF. In the same manner we calculate 95% upper and lower bounds for the results of the
estimator G.
Figure 2 gives an illustration of upper and lower bounds. We notice that the upper and lower bounds are
closer together according as the daily survival rate is closer to one. Figure 2 shows the bias in more
detail.
Important to know: how far are the 95% upper and lower bounds separated. In other words: how far are
the results of the estimator spread around the expectation. If the spread is small then the precision is
large. The standard deviation σ is a measure for the precision: the smaller σ, the larger the precision. So
the precision increases according as the upper and lower bounds are closer together.
Figure 2:
Graphs of the 95% upper and lower bounds of the asymptotic probability distribution of F1,F2,G1 and G2 as
a function of the real daily survival rate p. The broken line between the bounds is the asymptotic expectation.
The straight broken line is the real daily survival rate p. Parameters: m=20; n=100; N=1.
Figure 3 gives an illustration of upper and lower bounds of the estimator G1 as a function of the real daily
survival rate p. The length of the observation interval m varies for each graph.
Figure 3:
Graphs of the 95% upper and lower bounds of the asymptotic probability distribution of G1 as a function of
the real daily survival rate p. The broken line between the bounds is the asymptotic expectation. The straight
broken line is the real daily survival rate p. Parameters: m=5, 10, 15 and 20; n=100; N=1.
We notice in figure 3 that the precision increases if the length of the observation interval m increases.
The graphs in figure 3 are made for n=100 observation intervals. The precision increases if n increases.
It follows from the equations of the asymptotic variance of the Mayfield estimator that we can write (for n
large enough): σ2 = τ2/n, where τ2 is independent of n. So the standard deviation is inversely proportional
to the square root of n.
For example: if we choose the number of observation intervals 4 times as large, the standard deviation
becomes 2 times as small. The distance between the 95% upper and lower bounds becomes 2 times as
small.
In figure 3 we notice an other asymptotic property: the bias increases according as the length of the
observation interval m increases.
The interval [F - 1,96σF , F + 1,96σF] is a 95% confidence interval for the parameter µF (if n is large
enough). We replace the unknown p in the equation of σF by the result of F. This is not a 95% confidence
interval for p because µF ≠ p. Only if the bias is sufficient small, it is a confidence interval for p. This
argument applies also to the estimator G.
Table 2: Asymptotic expectation of the hatch rate µL of the estimators F1L, F2L, G1L and G2L. Also the
expectation of an unbiased estimator QL is calculated (=pL). Results are calculated for various values of p
and m with fixed values for L=30 and N=1.
p (m=5) p (m=15) p (m=20) p (m=30)
0.92 0.95 0.98 0.92 0.95 0.98 0.92 0.95 0.98 0.92 0.95 0.98
F1L 0.076 0.208 0.542 0.100 0.222 0.545 0.120 0.234 0.547 0.174 0.267 0.552
F2L 0.068 0.199 0.539 0.074 0.197 0.534 0.085 0.201 0.533 0.120 0.218 0.532
G1L 0.094 0.224 0.549 0.118 0.239 0.551 0.138 0.251 0.553 0.192 0.282 0.559
G2L 0.079 0.210 0.543 0.073 0.196 0.534 0.078 0.194 0.530 0.101 0.199 0.524
QL 0.082 0.215 0.545 0.082 0.215 0.545 0.082 0.215 0.545 0.082 0.215 0.545
The graphs are grouped as a matrix. Each row relates to one of the estimators for the hatch rate. Each
column relates to the length of the observation interval (m=3, 10, 20 and 30). The parameters n=200 and
L=30 are fixed. On top of each graph a code indicates which parameters are used. For example:
G2(10,200,30) means an estimate of G2L with parameters m=10, n=200 and L=30.
To compare we show the graphs of an alternative estimator QL. The estimator QL is asymptotically
unbiased. The definition of this estimator is given in appendix F.
The relative error on the vertical axis has a range from -1 until 3 (read: -100% until 300%).
The graphs in appendix A tell us much about the asymptotic behaviour of the Mayfield estimator.
From these graphs we can draw the following conclusions:
>1. The relative error (bias) of the hatch rate is largest for small hatch rates and decreases if the hatch
rate increases.
>2. It is desirable that the relative error is zero (broken zero line). This is only the case if the estimator is
asymptotically unbiased. See graphs of the estimator QL.
>3. In general we can conclude that the relative error for small hatch rates increases if the length of the
observation interval m increases.
>4. A remarkable but pleasant property applies to the graphs. The shape of the graph does not change
hardly if we increase or decrease L provide that we hold the proportion m/L constant. This is only true if L
is large enough: roughly L>15. For example: the graph for m=20 and L=30 has roughly the same shape
as the graph for m=40 and L=60. In this case m/L=0.6666. We will call the proportion m/L the relative
length of the observation interval. The columns of graphs in appendix A have a relative length of
respectively 0.1; 0.3333; 0.6666; 1.
>5. The precision in the estimate of the hatch rate increases if m increases. Because the standard
deviation decreases if m increases. The precision decreases if L increases while n and m remaining the
same.
>6. The precision of QL is somewhat smaller than those of the Mayfield estimator, especially for small
hatch rates. This advantage of the Mayfield estimator is cancelled by the large bias for small hatch rates.
>7. Whether the bias in estimating the hatch rate is acceptable depends of the desired precision. If the
precision is large we expect a proportionally smaller bias. If n increases then the precision increases
while the bias does not change. So with increasing n the influence of the bias becomes more important.
>8. The range of the horizontal axis in the graphs of appendix A (the real hatch rate) is limited by 0.5.
For values greater than 0.5 the trend continues: a larger precision and a smaller bias.
3.5 Estimating the hatch rate with a small number of observation intervals.
The equations which describe the asymptotic behaviour are useful to formulate properties of the Mayfield
estimator. In principle they apply only for infinite large n and they apply only approximately for finite n.
Only if n is large enough we have a good approximation of the asymptotic behaviour.
In appendix E (approximation criteria) we present equations to calculate the exact distribution function,
expectation and variance of the Mayfield estimator for any number of observation intervals. Comparing
the exact distribution with the asymptotic distribution, we calculate minimum values of n for which the
approximation is acceptable (table 3 of appendix E).
Especially for small hatch rates n must be large to give a good approximation. This n is hardly to realise
in practise. So it is interesting to investigate the exact behaviour of the bias for small hatch rates and
small values of n. We don't use the asymptotic distribution but the exact distribution of the Mayfield
estimator.
The graphs in appendix B tell us much about the exact behaviour of the Mayfield estimator for small
values of n. The graphs are grouped as a matrix. Each row relates to one of the estimators for the hatch
rate. Each column relates to the number of observation intervals (n=5, 15, 25 and 35). Each graph
shows two curves which belongs to different lengths of the observation interval.
>1. See the conclusion 1 of the previous section with regard to the graphs in appendix A.
>2. Now for small values of n the estimator QL is not unbiased. The bias is largest for small hatch rates.
The bias decreases if n increases. For small hatch rates n must be very large to make the bias almost
zero. So we have the same problematic as for the Mayfield estimator: we can't prevent the bias for small
hatch rates. The difference is that the bias for Q decreases when n increases and becomes zero in the
limit. The Mayfield estimator has a bias that remains existing for increasing n.
>3. Each estimator has a large bias for small hatch rates if the number of observation intervals is small.
Especially the combination of a small number of observation intervals and a small length of the
observation interval causes a large bias. For example if n=5 and m=0.2L, then for a real hatch rate of 0.1
the relative error is more than 100%.
>4. See the conclusion 4 of the previous section with regard to the graphs in appendix A.
>5. Roughly we can derive from the graphs of appendix B that: the larger n the smaller the bias. So the
asymptotic behaviour is in a way an optimum state.
4. Discussion.
We have included in the definition of the Mayfield estimator the function r(m). We remark: having F with
some function r(m) is equivalent to having G with the function r(m)-1.
We have investigated some variants of r(m). It turns out that the choice r(m) = 0.33m for the estimator G
is more or less an optimum. So we can't diminish the bias with some other choice of r(m). We can proof
that it is not possible to choose a function r(m) such that the bias completely disappears. So we can't
make the Mayfield estimator (asymptotically) unbiased.
The graphs in the appendices A and B give a good survey of the quality of the various estimators.
Considering the bias for small hatch rates we can conclude that among the Mayfield variants G1 is the
poorest and G2 is the best choice. (Johnson 1979) advised to take F1 for small m (m<15) and for large
m to take F2. This choice can be explained if we compare the graphs for F1 and F2 in appendix A. But
from the graphs in appendix B we can conclude that F2 has always the smallest bias. With this kind of
discussions it is necessary to make use of the concept of "relative length" of an observation interval.
Johnson did not investigate the estimator G.
The structural bias inherent in using the Mayfield method is a tedious factor. Mostly the bias is
negligible. Especially for small daily survival rates the bias can have a significant influence on the
estimate. If we want to minimise the bias, we can consider to use the alternative estimator Q from
appendix F. It performs well if the number of observation intervals is large (see for example table 4 in
appendix F) and the length of the observation intervals is as large as possible.
0.9
0.6
0.01 0.01 0.01
0.3
0 0 0 0.3
0.8 0.85 0.9 0.95 1 0.8 0.85 0.9 0.95 1 0.8 0.85 0.9 0.95 1 0.8 0.85 0.9 0.95 1
Figure 4:
Left: the three utmost left graphs shows the SDJ (broken curve) and SDA (solid curve) as a function of the
real daily survival rate p. Parameters: N=1, n=100, m=5, 15 and 25.
Right: graph of the relative error (SDJ-SDA) / SDA as a function of the real daily survival rate p. Parameters:
N=1, n=100, m=5, 15 and 25 (the larger m the larger the relative error).
Abbreviations:
SDA: standard deviation σF of the asymptotic distribution of F1 (equation (3.1)).
SDJ: asymptotic expectation of the estimator for the standard deviation of F1 according to Johnson.
General conclusions.
In this article we have tried to give insight in the properties of the Mayfield estimator and we have
presented an alternative estimator. We can draw a number of general conclusions:
>1. We can now study the Mayfield method better and more precise thanks to the presented equations.
Especially the equations for the asymptotic behaviour are very useful. They only apply if the number of
observation intervals is large enough.
>2. All variants of the Mayfield estimator have a bias which is largest for small daily survival rates. The
bias decreases according as the daily survival rate is closer to one.
>3. The bias in the estimate of the daily survival rate implies a bias in the estimate of the hatch rate. The
bias is largest for small hatch rates.
>4. We can't eliminate the bias in the Mayfield estimator. We can arrange the Mayfield variants in
sequence of increasing bias as follows: G2, F2, F1 and G1.
>5. We can enlarge the precision of the Mayfield estimator by enlarging the number of observation
intervals or by enlarging the length of the observation intervals.
>6. The estimator Q is a better alternative for the Mayfield estimator. This estimator is asymptotically
unbiased and has a similar precision as the Mayfield estimator. Also its non-asymptotic behaviour
performs better. For small samples the estimator Q performs the best if we choose the length of the
observation intervals as large as possible.
>7. Until so far we have investigated only samples of type a. In appendix G we show that all estimators
perform badly for samples of type b. For mixed samples the bias is difficult to quantify. To minimize the
bias for mixed samples: try to keep the size of type b samples small and choose the length of the
observation interval small.
5. Acknowledgements.
I am grateful to Martijn van de Pol (Centre for Ecological and Evolutionary Studies, University of
Groningen) for a review in march 2006.
6. References.
[1] Bart J. & Robson D.S. 1982. Estimating survivership when the subjects are visited
periodically. Ecology 63:1078-1090.
[2] Beintema A. 1992. Mayfield moet: oefeningen in het berekenen van uitkomstsucces.
Limosa 65:155-162.
[3] Beintema A.J. 1996. Inferring nest success from old records. Ibis 138:568-570.
[4] Bickel P.J. & Doksum K.A. 1977. Mathematical Statistics: basic ideas and selected topics.
1.5:28-34. Holden Day Inc, San Francisco.
[5] Hensler G.L., Nichols J.D. 1981. The Mayfield method of estimating nesting success: a model,
estimators and simulations results. Wilson Bull. 93:42-53.
[6] Johnson D.H. 1979. Estimating nest success: the Mayfield method and an alternative.
Auk 96:651-661.
[7] Miller H.W. & Johnson D.H. 1978. Interpreting the results of nesting success. J.Wildl.Mgmt.
42:471-476.
[8] Mayfield H. 1961. Nesting success calculated from exposure. Wilson Bull. 73: 255 261.
2 2 2 2
1 1 1 1
0 0 0 0
1 1 1 1
0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5
2 2 2 2
1 1 1 1
0 0 0 0
1 1 1 1
0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5
2 2 2 2
1 1 1 1
0 0 0 0
1 1 1 1
0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5
2 2 2 2
1 1 1 1
0 0 0 0
1 1 1 1
0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5
2 2 2 2
1 1 1 1
0 0 0 0
1 1 1 1
0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5 0 0.1 0.2 0.3 0.4 0.5
Figure 5:
Graphs of the relative error of the asymptotic expectation of the estimated hatch rate with regard to the real hatch
rate pL as a function of pL. Including the relative error of the expected 95% upper and lower bounds. Each row
relates to one of the estimators F1, F2, G1, G2 and Q. Each column relates to the length of the observation interval
(m=3, 10, 20, 30). Code G2(10,200,30) means: G2 with m=10, n=200 and L=30. Parameters: N=1,n=200,L=30.
.
Figure 6:
Graphs of the relative error of the exact expectation of the estimated hatch rate with regard to the real hatch rate p L
as a function of pL. Each row relates to one of the estimators of F1, F2, G1, G2 and Q. Each column relates to the
number of observation intervals (n=5, 15, 25, 35). The broken curve is made with m=0.2L and the solid curve with
m=0.6L. Code G1(15) means: estimator G1 with n=15. Parameters: N=1 and L=30.
.
The next variant of the central limit theorem is a generalisation of theorem C1. It is the multidimensional
form of this theorem.
∑ ( h (μ))
N
2 σ[k] 2
(C.3) τ2 = /
k
k= 1 c[k]
Proof:
In an analogous manner as the proof of theorem C1. ▲
We proof the asymptotic properties for the estimator F; an analogous proof applies to the estimator G.
To investigate the asymptotic behaviour we define a row of estimators Fn. We define the integer n as
follows. Define the row of numbers c[1], ..., c[N] by:
n[i] = c[i].n, i = 1..N with n = Σi n[i] and 0 < c[i] ≤ 1. So: Σi c[i] = 1.
By rewriting the equations (2.2) and (2.3) we get the equations:
N S[i] S[i]
(D.1) An = n ∑ c[i] m[i]
n[i]
+ r(m[i]) 1 -
n[i]
i= 1
N
S[i]
(D.2) Bn = n 1−
∑ c[i]
n[i]
i= 1
Theorem D1:
There exists a function h: RN → R and a function τ: (0,1) → R, such that:
(D.7) n ( Fn − h(μD ) ) L
→ Z Z ~ N(0,1) n→ ∞
τ (p)
The values τ(p) and h(µD) are independent of n. See the proof for the equations of h and τ.
Proof:
We apply to the row of estimators Fn the multidimensional central limit theorem(theorem C2).
For that purpose we define the function h: RN → R such that:
S[1] S[N]
(D.8) Fn = h , ... ,
n[1] n[N]
So for n large enough it applies approximately that F ~ N( µF; σF2) and G ~ N( µG; σG2).
For the sake of a clear formulation of the asymptotic expectation and variance we define two functions
a, b :R→R as follows:
( m[i] p )
N
(D.14) a(p) = ∑ c[i] m[i]
+ r(m[i]) (1 - p m[i] )
i= 1
N
(D.15) b(p) = 1 − ∑ c[i] p m[i]
i= 1
Then the equations for the expectation and variance are:
b(p) ( m[k] - r(m[k]) ) + a(p) pm[k] (1− pm[k] )
2
b(p) 1 N
(D.16) μF = 1 − σ =
2
F ∑
n k= 1
a(p)2
c[k]
a(p)
Theorem D2:
If F ~ N(µF;σF2) approximately for n large enough.
Then FL ~ N(µFL ; (L µF L - 1)2σF2) approximately for n large enough.
Proof:
We apply theorem D1 to the composition (h1◦h2)(t) of the functions h1(t)=tL and h2(t)=h(t).
Then we use the chain rule to the partial differentation in equation D13. ▲
The question is: how can we apply the equations of the asymptotic probability distribution of the Mayfield
estimator as a approximation for the behaviour with a finite number of observation intervals n. Then we
have to define what we mean by a "good approximation". As an example we consider the estimator FL of
the hatch rate.
Let dfFe(x) the distribution function of the exact distribution of the estimator FL with a finite number of
observation intervals.
Let dfFa(x) the distribution function of the asymptotic (normal) distribution of the estimator FL.
Now we define that the exact distribution is a good approximation of the asymptotic distribution if it meets
three requirements:
>1. |dfFe(x) - dfFa(x)| < 0.1 for all x.
>2. the relative error of the exact expectation with regard to the asymptotic expectation is smaller than
0.5 times the asymptotic standard deviation.
>3. the relative error of the exact standard deviation with regard to the asymptotic standard deviation is
smaller than 0.1.
With this definition (which is subjective) we calculate the minimum n for which the exact distribution is a
good approximation of the asymptotic distribution. All values greater than the minimum n we call "large
enough". We have calculated the minimum values of n in table 3.
Table 3: The minimum number of observation intervals for which the exact distribution of F2L is
roughly equal to the asymptotic distribution of F2L. The values are calculated for various hatch rates u
and various relative length of the observation interval m.
The code ++ means: n>170 Number of groups: N=1.
m/L
u 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0
0.01 ++ ++ ++ ++ ++ ++ ++ ++ 100 110
0.05 ++ ++ ++ ++ ++ ++ ++ ++ 91 37
0.10 ++ ++ ++ ++ ++ ++ 158 90 49 30
0.15 ++ ++ ++ ++ 139 94 62 39 24 20
0.20 ++ ++ 125 83 57 40 27 18 15 15
0.30 127 56 33 21 20 15 15 15 15 15
0.40 60 35 25 20 15 15 15 10 10 10
0.50 75 35 30 25 15 15 15 10 10 10
0.60 80 50 30 30 20 20 15 15 15 15
0.70 120 65 40 30 20 20 20 15 15 15
0.80 130 85 45 35 30 25 20 20 20 20
0.90 ++ 135 75 55 45 40 35 30 25 25
0.99 ++ 125 85 65 50 45 40 35 30 30
[if h(s) ]
n[1]
(E.1) dfFe(x) = ∑ L
≤ x then P( S[1] = s ) otherwise 0 (N = 1)
s= 0
An analogous rule applies for N>2. The function h:RN → R is defined in appendix D. We calculate the
probabilities for S[i] with the binomial distribution ß(n[i];pm[i]); i = 1..N.
If L=1, then of course (E.1) is the distribution function of F.
To compare the moments of the exact distribution of FL with the moments of the asymptotic distribution
we give the equations for the exact moments:
[ (h(s)) ]
n[1]
(E.2) E(F L ) = ∑ L
P( S[1] = s ) (N = 1)
s= 0
[ h(s1, s2) ]
n[1] n[2]
E(FL ) = ∑ ∑ L
P( S[1] = s1) P( S[2] = s2 ) (N = 2)
s1 = 0 s2 = 0
Definition.
For the given statistical model we introduce the estimator Q as an alternative for the Mayfield estimator.
N
∑ w[k] U [k]
(F.1) k= 1 S [k]
Q= N
with U [k] = m[k] and w[k] = n[k] m[k]
n[k]
∑ w[k]
k= 1
So the estimator Q is the weighted average of the random variables U[k] with weight factors w[k].
In the same manner as with the Mayfield estimator, we can proof that Q has approximately a normal
distribution with expectation p (asymptotically unbiased). This applies in spite of the choice of the weight
factors. We use the freedom to choose the weight factor to make the variance as small as possible. The
used weight factor is found experimental. The equation for the asymptotic variance of Q is:
N
(
w[k] 2 1− pm[k] )
∑ n[k] m[k] 2 pm[k] − 2
(F.2) σ Q2 =
k= 1
2
N
∑ w[k]
k= 1
(Bart J. et al. 1982) describes an alternative estimator for the daily survival rate. It is the maximum
likelihood estimator of our statistical model. This estimator is asymptotically unbiased. Disadvantage is
the complex computation.
In the theory of estimation it is proofed that the maximum likelihood estimator of Bart & Robson has the
smallest (asymptotic) variance among all unbiased estimators for p with the given statistical model.
We use the term UMVU estimator (Uniformly Minimum Variance Unbiased). So the estimator Q has a
larger variance than that of Bart & Robson. Computations show that this difference between the
variances is small (a few percents). If N=1 then the variances are even equal. So the estimator Q is
approximately an UMVU estimator.
Table 4: The minimum number of observation intervals for which the exact distribution of QL is roughly
equal to the asymptotic distribution of QL. The values are calculated for various hatch rates u and
various relative length of the observation interval m.
The code ++ means: n>170 Number of groups: N=1.
m/L
u 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0
0.01 ++ ++ ++ ++ ++ ++ ++ ++ ++ 160
0.05 ++ ++ ++ ++ ++ ++ ++ ++ 74 60
0.10 ++ ++ ++ ++ ++ 133 75 35 25 55
0.15 ++ ++ ++ 114 71 43 24 15 15 30
0.20 ++ 152 85 50 31 19 15 15 15 30
0.30 114 48 26 20 15 15 15 15 15 15
0.40 65 35 25 20 15 15 15 15 15 15
0.50 70 40 25 20 15 15 15 15 15 15
0.60 85 40 35 25 15 15 15 15 15 15
0.70 100 50 40 30 25 25 20 15 15 15
0.80 115 65 45 35 30 25 25 20 20 15
0.90 ++ 135 75 55 45 40 35 30 25 25
0.99 ++ 125 85 65 50 45 40 35 30 30
[if h(s) ]
n[1]
(F.3) dfQe(x) = ∑ L
≤ x then P( S[1] = s ) otherwise 0 (N = 1)
s= 0
An analogous rule applies for N>2. We calculate the probabilities for S[i] with the binomial distribution
ß(n[i];pm[i]); i = 1..N. If L=1, then of course (F.3) is the distribution function of Q.
To compare the moments of the exact distribution of QL with the moments of the asymptotic distribution
we give the equations for the exact moments:
[ (h(s)) ]
n[1]
(F.5) L
E(Q ) = ∑ L
P( S[1] = s ) (N = 1)
s= 0
[ h(s1, s2) ]
n[1] n[2]
E(Q L ) = ∑ ∑ L
P( S[1] = s1) P( S[2] = s2 ) (N = 2)
s1 = 0 s2 = 0
We can calculate the exact expectation of Q more efficient with the equation:
N n[k]
s
∑ ∑ w[k] m[k]
n[k]
P( S[k] = s)
k = 1 s= 0
(F.6) E(Q ) = with w[k] = n[k] m[k]
N
∑ w[k]
k= 1
b(p) a(p)
(G.1) μF = 1− and μG =
a(p) a(p) + b(p)
Let us look at the (theoretical) case of samples of type b. So all clutches hatch within the observation
interval and therefore we let Sa=0 and Ta=0.
Table 5: Asymptotic expectation of the hatch rate µL of the estimators F1L, F2L, G1L and G2L using a
sample of type b. Also the real expected result pL is calculated. Results are calculated for various values
of p and m with fixed values for L=30 and N=1.
p (m=5) p (m=15) p (m=20) p (m=30)
0.92 0.95 0.98 0.92 0.95 0.98 0.92 0.95 0.98 0.92 0.95 0.98
F1L 0.096 0.228 0.550 0.147 0.271 0.567 0.174 0.293 0.575 0.232 0.335 0.590
F2L 0.087 0.219 0.547 0.120 0.247 0.557 0.138 0.262 0.563 0.180 0.293 0.573
G1L 0.113 0.244 0.557 0.165 0.286 0.573 0.192 0.307 0.581 0.248 0.348 0.595
G2L 0.098 0.230 0.551 0.119 0.247 0.557 0.131 0.256 0.560 0.161 0.277 0.566
pL 0.082 0.215 0.545 0.082 0.215 0.545 0.082 0.215 0.545 0.082 0.215 0.545
Table 5 shows the results of the asymptotic expectation of the Mayfield estimator using a sample of type
b. Comparing the results of table 5 with table 2, we conclude the bias for type b samples is larger than
the bias of type a samples. For example: If p=0.92 and m=0.5L then the real hatch rate is 0.082. The
expectation of the estimator F1L is 0.147 for type b samples (bias 80%) and 0.100 for type a samples
(bias 20%). The bias is still larger if α < ½ : hatch dates particularly at the first half of the observation
interval. If we have luck then α > ½ and the bias will be smaller.
So there is a great risk that the bias for type b samples is large. This is explicable. The number of
survival days of non destroyed clutches of type a is exactly the length of the observation interval. While
for non destroyed clutches of type b the number of survival days must be guessed because of the
unknown hatch date falling within the observation interval. A guess of q(m) = ½ m seems reasonable but
is not always the best choice. A guess of q(m) = 0 (omitting hatched clutches) is reasonable if clutches
hatch particularly at the start of the observation interval. This is mostly not the case. In fact we cannot
formulate a suitable rule for q(m).
In the case of mixed samples the effect of type b samples will depend on the size of the type b sample in
proportion to the total size of the sample.
Estimating the daily survival rate we conclude:
The bias of type b samples is on an average larger than the bias of type a samples.
The bias of type b samples is difficult to quantify because of the unknown hatch date.
If possible we had to keep the size of the type b samples small in order to minimize the bias. Another
way that helps to minimize the bias is to keep the length of observation intervalls small.
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