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POPULATION SIZE time to wait. The class determined if the capture and
survival rates were constant or time-dependent by
looking at 5-years of capture-recapture data. They also
recorded body condition index of all the chickadees to
Rachael E. Molidor
Rachael.Molidor@loras.edu
Molidor 1
Introduction
One method of determining population size of motile animals would be to count each
individual in the vicinity. This method, however, is flawed because there is no way to determine
which individuals have already been counted, especially during the recount/recapture period.
Another method in order to determine population size is via sampling, counting the amount of
individuals in a small area and then using this sample of the population to determine the
population of a larger area. An issue with this method is that the population can vary in different
areas of a larger location, therefore sampling is not a reliable method to determine a whole
population of an area.
The purpose of the capture and recapture of Black-clapped chickadees was to estimate
the population size of the resident chickadees at and around the EB Lyons Nature Center at the
Mines of Spain in Dubuque, Iowa to determine if supplemental feeding throughout the winter
months has an effect on the survival rates of the chickadees. To capture and mark, Black-capped
chickadees were caught in a net and then banded, later during the recapture the tagged
chickadees were recorded in order to determine the population size. The EcoBeaker computer
simulation was used to examine parameters that maximize the accuracy of prediction of the
population size estimate. The capture and mark-recapture data from the past five years was used
to estimate the survival rate of Black-capped chickadees and test for variation in survival
probability over time. The last objective met from this study is the testing of the hypothesis that
body condition in birds differs according to age between second-year birds and after second-year
birds. Loery et al. (1997) found that there was strong evidence in a lower survival rate of SY
birds in comparison to the survival rate of ASY birds. However, there was no conclusive data to
show that these birds are dying at a higher rate. Chickadees are faithful to their wintering
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areas which is the root cause of the higher mortality rates of older birds (Loery et al.). The
Loras class conducted their survey in late winter/early spring to determine if the same was true
for the Black-capped chickadees nesting in the Mines of Spain area and to deduce if
supplemental feeding was influencing the survival rates of the population as a whole.
Methods
On March 9, 2017 the Loras College conservation biology class went to the Mines of
Spain around the EB Lyons nature center at the Mines of Spain in Dubuque, Iowa to net and
band Black-capped chickadees. Small mesh nets were used near bird feeders in order to capture
the most birds possible. Two rows of nets were set up in different locations in order to maximize
the number of chickadees captured. Mark-recaptures were done on the 16, 19, and 21 of March
where non-tagged chickadees were tagged at that time as well while the birds that had been
captured on March 9 were recorded. The metal tags were placed around the chickadees leg and
the number on the tag was recorded on a spread sheet along with the length of the chickadees
wing, its weight, and whether it was a second year bird or an after second year bird. In order to
determine population size, we used the Lincoln Index: (n1/N) = (m2/n2), after resetting the
Before going to tag the chickadees, the Loras class did an EcoBeaker simulation in order
to determine what the best plan of action was to have the most successful capture and recapture
period. The EcoBeaker simulation was used to determine how many hours were necessary in
order to have successful capture and recapture to determine population size of pigeons as a guide
for the chickadee capture and mark-recapture. The simulation allowed the class to set the
number of individuals were possible for capture and the amount of time for capture, recapture,
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and the waiting interval in between. The simulation had pigeons flying randomly and each time
Once tagged and recorded, we used MARK software to fit four models to determine if the
5-year survival model for Black-capped chickadees tagged in the Mines of Spain survival-
capture probability were constant or time-dependent. The class also compared body condition
index between second-year birds and after second-year birds collected from the study to
determine if there was a difference between the two age groups body condition.
Results
On 09 March, we captured and banded 18 chickadees. The following week, March 16,
we captured and banded 12 new chickadees and recaptured 1. On March 19, 7 new chickadees
were captured and banded and 2 for the initial capture date were recaptured. On the last
recapture date, March 21, 9 new chickadees were captured and banded and 4 were recaptured (2
from the initial capture and 2 from the March 16 recapture). The results of the population of the
first recapture is N = ((18+1)*(13+1)) / (1+1)-1, this results in N= 132 with a 95% lower
confidence limit of 0.20 and an upper confidence limit of 263.8. Because these results were not
narrow enough, data from the following recaptures were taken in order to yield a more precise
population count. With the recaptures from the other two dates, the new equation become N =
((18+1)*(33+1)) / (5+1)-1 which yields N = 106.7 with the lower confidence limit being 46.8
March 16
21
(To determine if capture probability (p) and survival probability (phi or ) are dictated by both
being constant (.), time-dependent (t), or a combination of the two.) By using the MARK
simulation, we are able to concluded that capture and survival probability are both constant,
Image 1
The data shown in image 1 shows that Phi(.)p(.) yielded the model likelihood result of 1, while
the other possibilities: Phi(.)p(t), Phi(t)p(.), Phi(t)p(t) all had model likelihoods being less than 1,
showing that survival and capture probability were constant across the five-year period. By
keeping capture and survival constant throughout, we were able to estimate the survival rate at
53% (95% CL: 39-67%) and the capture probability at 35% (95% CL: 22-52%) that was
The last component that the data that was compiled through the capture of the Black-
capped chickadees that we attempted to determine was to test the null hypothesis that there is a
difference between body condition and age. By using the information collected during capture
and recaptures, the population of second year birds (SY) was 15 and after second year birds
(ASY) was 30. In this case, t43 = 2.44; p = 0.019 as shown in table 3. Because p = 1.9%, we are
Table 2
Group Statistics
Discussion
Based on the EcoBeaker simulation, there are a few adjustments that should have been
done to the capture and recapture in order to have a more successful estimate of the Black-
capped chickadee population. The hours for capture and recapture should have been drastically
increased compared to the actual capture and recapture. Due to class time constraints, there was
no way that the Loras class could capture for 80 hours and recapture for 200 hours with 300
hours in between the capture and recapture time periods. By increasing time for capture, time for
recapture, and the interval in between there is an increase in the accuracy and precision in
determining the population size estimate. Another key difference between the study and the
simulation was that the Black-capped chickadees in the area quickly realized that there were
traps surrounding the feeders, changing the routine patterns of the chickadees. This was not an
issue in the simulation because the pigeons moved at random and did not adapt to the presence of
the traps. In order to counteract this problem, the orientation of the nets could have been
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changed periodically, in order to yield a higher amount of captured chickadees. To increase the
amount of birds caught and marked, the chickadees could also have been marked with a safe dye
so that the Loras class could keep track of which chickadees had already been caught with no
In Loery, et al (1997) study of chickadee survival, they concluded that younger birds
have a higher mortality rate than older birds but was not able to distinguish between death or
emigration because he did not conduct his study in a closed population. Our study determined
that younger Black-capped chickadees have a better body condition index, we speculated this is
because they are more agile in their younger age. Neither study has confidence in the survival
In the winters between 1982 and 1985, Brittingham and Temple (1998) collected data
from 418 marked chickadees through three winters, between the months of October through
April. The experiment had a control group of 158 chickadees that were not given supplemental
food throughout the winter, while the rest were. The study found that the monthly survival rates
of the supplemental fed chickadees was 95% while the rate for the ones not given extra food was
87%. A larger difference was between the average overwinter survival rates, the experimental
group had a survival rate of 69% while the control groups rate was 37%. The data reveals that
through supplemental feeding, chickadees survival rate increase throughout the winter by more
than 30%. The findings of Brittingham and Temple (1998) helped us to determine that
supplemental feeding in the Mines of Spain area would increase the population and better the
survival rates of the chickadees in the area and, overall, there was a higher probability in
catching a bird that had been previously marked in the last five years.
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Gooch (2009) documented the effects of West Nile Virus and many species of birds,
including the Black-capped and Carolina chickadees. The expected impact on chickadees,
according to Gooch, was moderate. The study is from Ladeau, et al. (2007) and showed that the
number of regions that were studied in relation to chickadees was 4 while the regions that had a
significant impact due to the virus was 2 only 50% of regions that contained the chickadee
species had an impact from the West Nile Virus. The conclusion that Gooch comes to is that
Black-capped and Carolina chickadees, along with several other species were not significantly
affected by the West Nile Virus while other birds, such as the American Crow and Tufted
Titmouse suffered drastically from the outbreak. Our data shows that the Black-capped
chickadee population in the Mines of Spain was not affected by the West Nile Virus because the
survival-capture probability results were both constant over a 5-year period. This means that the
population over the course of 5-years did not decline drastically as it would have if the chickadee
We believed as a class that ASY birds body condition would be superior to SY birds
because they have more experience in survival and know how to find food the best. Smith
flocks. She found that there is a, in fact, a hierarchy within the male population of flocks that
coordinate with experience/age. The rank of females was found to be directly correlated with the
rank of her mate. This information led us to believe that after second-year birds would then be
the first allowed to feed and therefore have the best body condition overall. However, SY birds
have better body composition in comparison to ASY birds. A main reason for this difference
may be that the younger birds participate in more risk taking behavior in comparison to
experienced birds partaking in risk aversion behavior. Another difference between the young
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birds and experienced birds is that the second-year birds may not understand that they can limit
their food intake with the supplemental feeding. Their higher weight may be due to the fact that
they are just overeating because they do not have the experience to know that they will have a
meal next time they go to the feeders. The issue with this study is that this is the first winter for
many of the second-year birds. The study is potentially biased because after the first winter there
is usually high mortality rates among the second-year birds. The SY birds that were marked and
recorded could be considered the super group of survivors and therefore not an accurate
Literature Cited
Gooch, L. C. (2009). The Impact of West Nile Virus on Ohio Birds. The Ohio Cardinal 32(3-4),
176-194.
Loery, G., et al. (1987). Age-Specificity of Black-Capped Chickadee Survival Rates: Analysis of