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                  <text>Colorado Division of Parks and Wildlife
July 2015-June 2016

WILDLIFE RESEARCH REPORT
State of:
Cost Center:
Work Package:
Task No.:

Colorado
3420
1680
N/A

Federal Aid
Project No.

N/A

:
:
:
:

Division of Parks and Wildlife
Avian Research
Bird Conservation
Raptor data integration, species distribution, and
suggestions for monitoring

Period Covered: January 1, 2016 – October 31, 2016
Author: R. Yale Conrey
Principle Investigators: R. Yale Conrey, J. Gammonley, CPW; J. DeCoste, W. Kendall, Colorado State
University
Collaborators: U.S. Fish and Wildlife Service; Bird Conservancy of the Rockies; U.S. Forest Service;
Bureau of Land Management; National Park Service; Boulder County; other agencies who have
submitted nest data; Cornell Lab of Ornithology
CPW biologists, GIS group, Species Conservation staff, and Parks staff: especially R. Sacco (GIS Unit);
Senior Terrestrial Biologists, M. Sherman, A. Estep, M. Cowardin, N. Seward (Terrestrial); D. Klute, L.
Rossi (Species Conservation); J. Thompson (Resource Stewardship).
All information in this report is preliminary and subject to further evaluation. Information MAY
NOT BE PUBLISHED OR QUOTED without permission of the author. Manipulation of these data
beyond that contained in this report is discouraged.
EXTENDED ABSTRACT
Where they exist, raptor monitoring databases have generated important insights into various
aspects of raptor ecology, and can provide a sound foundation for specific management priorities for
individual species or within the larger context of managing targeted habitats and ecosystems (Greenwood
2007). CPW has a statewide raptor nest database, developed by Bob Sacco (GIS Unit), which currently
contains records for ~9,000 nest records going back to the 1970s. Currently, the nest database is primarily
being used by CPW at a site-specific scale in the oil and gas comment process (CO House Bill 1298) and
other local-scale land use input. The potential of this database to assess raptor populations at regional or
statewide scales, and the field protocols used to provide records for this database, have not been
thoroughly assessed. Other data sources have potential to contribute to our understanding of Colorado
raptors, including eBird, Breeding Bird Survey, and Colorado Breeding Bird Atlas. By exploring the
possibility of integrating these various datasets, we hope to generate a more comprehensive picture of
raptor abundance and occupancy across the state, with the eventual goal being to provide a more concrete
understanding of raptor population trends for management purposes.

1

�We are focusing on breeding populations of bald eagles (Haliaeetus leucocephalus: BAEA),
golden eagles (Aquila chrysaetos: GOEA), prairie falcons (Falco mexicanus: PRFA), and ferruginous
hawks (Buteo regalis: FEHA). GOEA are a Tier 1 Species of Greatest Conservation Need in Colorado,
while the other three species are on the Tier 2 list and BAEA and FEHA are species of Special Concern.
We are first focusing on GOEA and BAEA, partly because we have more nest records (GOEA) and better
quality data, with repeat visits to nests that allow nest survival analysis (BAEA). In addition, GOEA are a
species of conservation concern due to declining populations rangewide and their sensitivity to human
activity and energy development. BAEA are a species of management concern due to the recent attention
to nest take, contention over revised federal regulations, and the frequency with which BAEA nest in
areas with rapidly accelerating housing development.
Research objectives are to 1) Assess and improve the data available in CPW’s raptor nest
database; 2) Build distribution models for our highest priority raptor species, evaluating the importance of
ecological and anthropogenic covariates; 3) Estimate nest survival for bald eagles, evaluating the
importance of ecological and anthropogenic covariates; 4) Evaluate the potential for integrating other data
sources, such as eBird, Breeding Bird Survey, and Colorado Breeding Bird Atlas; 5) Make
recommendations for a state-wide raptor monitoring protocol.
The first step in this research project was to assess the data available in CPW’s raptor nest
database. This database contains locations and nest site occupancy information for 29 species of raptors
from 1975 to present. Most of the nest data have been collected opportunistically, and known nest sites
are resurveyed at a much higher rate than new areas are surveyed. For a nest site to be considered active
during CPW consultation for HB 1298, it must be known to have been occupied sometime within the past
5 years. Although some sites are visited yearly, others are therefore visited only when they have reached
the end of their 5-year window, and most nest sites have a listed status of undetermined or unknown
(many of these are historic nests that have not been visited within the past 5 years). More detailed
information is sometimes available for nests, but this information is typically summarized into annual
records by GIS staff who enter and maintain the data.
There has been a special effort to monitor BAEA through multiple visits per known nest location
per year, making these data suitable for modeling of daily nest survival. Aside from estimating daily and
annual nest survival rates, the goals of this model are to determine what ecological and anthropogenic
covariates are important predictors of nest survival and to provide a comparison of the outputs,
usefulness, and monitoring methods suitable for nest survival modeling versus distribution modeling. The
nesting cycle averages 18 weeks (or 126 days) from egg laying until fledging: incubation begins with the
first egg and lasts 5 weeks while the nestling period ranges from 8 – 14 weeks, averaging 12 – 13 weeks.
In Colorado, egg laying usually begins in late January to early February, hatching occurs in early March,
and young eagles can fledge anytime between early June and August.
Modeling of daily nest survival requires the following data from each nest: 1) date when the nest
was found; 2) the last day the nest was checked when alive; 3) the last day the nest was checked; and 4)
the fate of the nest (0=successful, 1=failed). For spatial covariates to be calculated, an accurate location
was also required. Covariates are listed in Table 1 along with the data source and citations supporting
their inclusion. J. DeCoste is calculating spatial covariates using ArcGIS spatial analysis tools and
running nest survival models in Program MARK,which uses maximum likelihood methods to estimate
nest survival as a known fate data type. Model selection will be based on AICc values. We expect to
complete the BAEA nest survival model by the end of 2016, but protocols and coding will be well
documented so that the model can be updated in future as more nest and covariate data become available.
The goal of distribution modeling is to determine what variables predict breeding locations and to
map areas with high to low probability of use for mitigation planning and future survey design. The next

2

�phase of this research project will include distribution modeling for GOEA and BAEA. For GOEA, we
will collaborate with U.S. Fish and Wildlife Service’s Western Golden Eagle Team (WGET), which has
completed some ecoregional models and is currently working on a High Plains model. Models for PRFA
and FEHA will follow, data permitting.
It is our objective to make recommendations for future monitoring guidelines that can be
consistently applied state-wide. We plan to use the distribution maps that result from this research to
stratify the state of Colorado into appropriate survey regions. Ideally, monitoring guidelines will include a
time of year for surveys, how often various regions will be visited, and a means to record survey effort
and both presence (new and existing nests) and absence data. In addition, we hope to provide some
guidance on the amount of effort that should be allocated to surveys for new nests, versus return visits to
known nest sites, and how to prioritize survey effort by raptor species and spatial location.
The CPW raptor nest database contained 9479 nest records for 8696 locations, as of 7 November
2016 (Table 2). This included 1477 active nests known to be occupied within the past 5 years. The
majority of nest locations (6127 nests, 70% of the total) have an unknown or undetermined status,
meaning that the site has not been visited in at least 5 years or that an observer was unable to determine
the status of the nest. There are 12 species with at least 100 nest records, and 27% of records are for nests
occupied by an unknown species. We made an agency-wide and inter-agency request for additional nest
data to maximize sample size prior to our analyses, which likely contributed to the 20% increase (1730
nests) over 2 years. Our bald eagle nest survival model currently has an input file containing 121 nest
attempts at 79 locations from 2012−16: 45 nest sites are on the Front Range and 34 are on the Western
Slope. An additional 105 nests were not included, and the most common reason for exclusion was that the
observer did not visit enough times or at the appropriate time to confirm nest fate.
Modeling efforts are still in progress. This was intended as a 2-year research project, ending 31
December 2017, although it will take additional years to implement and evaluate the monitoring
recommendations that result from our work and from discussions among CPW employees and our
partners working on raptors. However, some useful information has been gleaned from data quality
control, calculation of covariates, and initial runs of the bald eagle nest survival model. First, nest survival
models require more intensive survey data than do distribution models, because at least two, and
frequently more visits are required during the nesting season. One visit must occur during the incubation
or nestling phases, in which nesting activity can be confirmed by sighting eggs or nestlings, or at least by
sighting an adult sitting on the nest who is presumably incubating or brooding. One visit must occur
shortly before or after the presumed fledging date to confirm the fate of the nest, where successful nests
have at least one fully feathered juvenile leaving the nest and failed nests have none. For all purposes,
observers should reserve the value of zero to represent true zero, as opposed to a placeholder or indication
of an unknown or inapplicable value. Finally, 70% of nest locations in the database have a status of
unknown or undetermined (Table 2). These sites have much less value for modeling or HB 1298 purposes
than have nests with any of the other status designations, so some effort should be allocated to re-visiting
these sites.

3

�COLORADO PARKS AND WILDLIFE RESEARCH REPORT
RAPTOR DATA INTEGRATION, SPECIES DISTRIBUTION, AND SUGGESTIONS FOR
MONITORING
REESA C. YALE CONREY
INTRODUCTION
Raptors are key components of many ecosystems (Sergio et al. 2008), and numerous raptors in
Colorado are species of conservation concern. Monitoring raptor populations is challenging due to their
ecology and life history, such as expansive home ranges requiring a large spatial sampling effort (Fuller
and Mosher 1987). Other factors include nests which are often in high, hard to reach places, with adults
regularly perching high in the canopy or on massive cliffs. Further, their numbers are intrinsically low
(Bird and Bildstein 2007) with patchy distributions. Therefore, large-scale data sets of breeding raptor
populations are rare and frequently incomplete. Where they exist, monitoring databases have generated
important insights into various aspects of raptor ecology, and can provide a sound foundation for specific
management priorities for individual species or within the larger context of managing targeted habitats
and ecosystems (Greenwood 2007).
CPW has a statewide raptor nest database, developed by Bob Sacco (GIS Unit), which currently
contains records for ~9,000 nests going back to the 1970s. The database houses field records gathered by
Biologists, Wildlife Managers, Researchers and others, and is continually updated. Currently, the nest
database is primarily being used by CPW at a site-specific scale in the oil and gas comment process (CO
House Bill 1298) and other local-scale land use input. April Estep (Biologist) and Mindy Rice (former
Avian Researcher) recently used the database to model prairie falcon nest sites to target appropriate areas
for future nest searches in the Southeast Region. However, the potential of this database to assess raptor
populations at regional or statewide scales, and the field protocols used to provide records for this
database, have not been thoroughly assessed.
In addition, scientists and volunteers throughout Colorado have been gathering data on raptor
locations and abundance for decades using various survey techniques, including point counts, roost
surveys, and nest surveys and monitoring. Some of these data have been incorporated into CPW’s
database, but other data sources have potential to contribute to our understanding of Colorado raptors,
including eBird, Breeding Bird Survey, and Colorado Breeding Bird Atlas. By exploring the possibility of
integrating these various datasets, we hope to generate a more comprehensive picture of raptor abundance
and occupancy across the state, with the eventual goal being to provide a more concrete understanding of
raptor population trends for management purposes.
After consulting with Biologists, Species Conservation Coordinators, and others within CPW, we
decided to focus on breeding populations of bald eagles (Haliaeetus leucocephalus: BAEA), golden
eagles (Aquila chrysaetos: GOEA), prairie falcons (Falco mexicanus: PRFA), and ferruginous hawks
(Buteo regalis: FEHA). GOEA are a Tier 1 Species of Greatest Conservation Need in Colorado, while the
other three species are on the Tier 2 list and BAEA and FEHA are species of Special Concern. BAEA,
GOEA, and PRFA are distributed statewide, while FEHA are most common on the eastern plains. We are
first focusing on GOEA and BAEA, partly because we have more nest records (GOEA) and better quality
data, with repeat visits to nests that allow nest survival analysis (BAEA). In addition, GOEA are a species
of conservation concern due to declining populations rangewide and their sensitivity to human activity
and energy development. U.S. Fish and Wildlife Service’s Western Golden Eagle Team has been building
ecoregional distribution models for GOEA, and we hope to collaborate with them. BAEA are a species of
management concern due to the recent attention to nest take, contention over revised federal regulations,

4

�and the frequency with which BAEA nest in areas with rapidly accelerating housing development. PRFA
are threatened by alteration of grasslands and areas near cliff faces where they nest, as well as by a rapidly
recovering population of peregrine falcons (Falco peregrinus), which are displacing PRFA at nest sites
(A. Estep, pers. comm.). Aside from development pressure, FEHA may also be impacted by plague
(Yersinia pestis) epizootics in prairie dog (Cynomys sp.) colonies where they forage.
We began exploring interest in this project and requesting additional data within and outside our
agency in 2014. Funding was secured in 2015 and a research associate (J. DeCoste) started work on the
project in January 2016. Research objectives are to 1) Assess and improve the data available in CPW’s
raptor nest database; 2) Build distribution models for our highest priority raptor species, evaluating the
importance of ecological and anthropogenic covariates; 3) Estimate nest survival for bald eagles,
evaluating the importance of ecological and anthropogenic covariates; 4) Evaluate the potential for
integrating other data sources, such as eBird, Breeding Bird Survey, and Colorado Breeding Bird Atlas;
5) Make recommendations for a state-wide raptor monitoring protocol.
METHODS
Study Area
The study area encompasses the entire state of Colorado. Most of the nests in our database were
located and monitored opportunistically, but some have been studied as part of a local sampling protocol
over a limited number of years. For the BAEA nest survival model (the model that is closest to
completion), more than half the nests in our sample are located in the Front Range, with the remainder on
the Western Slope, and most are near water.
CPW Raptor Nest Database
The first step in this research project was to assess the data available in CPW’s raptor nest
database. We obtained data summaries from GIS staff (Robert Sacco) and made an agency-wide and
inter-agency request for additional nest data to maximize sample size prior to our analyses. This database
contains locations and nest site occupancy information for 29 species of raptors from 1975 to present.
Most of the nest data have been collected opportunistically, and known nest sites are resurveyed at a
much higher rate than new areas are surveyed. However, for a limited number of sites and years, nests
have been discovered and monitored within a sampling framework, with survey effort and absences (no
nest found) reported, but these details are not included in the current database structure. The database was
designed to meet the needs of CPW Biologists consulting on development proposals, as required by
Colorado House Bill 1298. The priority was to record coordinates for each nest site, species occupying
the site (if known), and occupancy status by adults presumed to be nesting at the site so that a 5-year
activity status could be calculated. For a nest site to be considered active during CPW consultation, it
must be known to have been occupied sometime within the past 5 years. Although some sites are visited
yearly, others are therefore visited only when they have reached the end of their 5-year window, and most
nest sites have a listed status of undetermined or unknown (many of these are historic nests that have not
been visited within the past 5 years).
More detailed information is sometimes available for nests, but this information is typically
summarized into annual records by GIS staff who enter and maintain the data. Therefore, the data for
individual nest visits that were required for the BAEA nest survival model were mostly obtained directly
from the personnel who submitted the data. The database contains the following information, whenever it
was provided by the person who submitted the data: nest ID code (assigned by GIS staff to unique
locations), site name, nest location (UTMs), substrate (codes for tree species, cliff, etc.), landowner,
survey date, observer, agency, study name, raptor species, nest status (destroyed, occupied – failed,
occupied – intact, undetermined, unoccupied – dilapidated, and unoccupied – intact), threats to the nest,
number of eggs, nestlings, and fledglings, ground/air survey, and comments. Raw data, spatial files, and a

5

�web map application are internally available to CPW employees but are not publicly available, although
other entities may request public lands data.
Bald Eagle Nest Survival Model
Several organizations with volunteer programs, including Bird Conservancy of the Rockies and
Colorado State Parks, make a special effort to monitor BAEA through multiple visits per known nest
location per year, making these data suitable for modeling of daily nest survival. Aside from estimating
daily and annual nest survival rates, the goals of this model are to determine what ecological and
anthropogenic covariates are important predictors of nest survival and to provide a comparison of the
outputs, usefulness, and monitoring methods suitable for nest survival modeling versus distribution
modeling. BAEA and peregrine falcons are the only raptor species for which this is possible across a
broad range of years at a regional or state-wide scale.
BAEA pairs will typically defend a nesting territory of about 1 mile in diameter (Garrett et al
1993). Defending food resources is the main goal of a BAEA pair, although effort of defense is tied
directly to resource availability (Swenson et al 1986). BAEA feed primarily on fish and waterfowl (Gende
et al 1997), although there is evidence that proximity to prairie dog colonies may be important in the
foraging ecology of BAEA in Colorado. BAEA will also scavenge on a wide range of food sources,
making some individuals less reliant on a nearby water sources than others (Saalfeld and Conway 2010).
BAEA nests are generally giant stick nests built near the top of “super canopy” trees less than 1 mile from
a major water source suitable for foraging (Mundahl et al. 2013, Watts 2015). In Colorado, bald eagles
primarily use cottonwood trees, although Douglas-fir and ponderosa pine are also used. If constructing a
new nest, a pair will typically build in tall, live trees that can support the weight of the massive nests.
Once built, the pair can use an existing nest for years, making repairs and replacing sticks at the start of
each nesting season. Often these nest trees will die, eventually causing the tree to blow over and the nest
to fail. Bald eagles will also use cliffs, man-made structures, or the ground if adequate trees are absent
from the environment. There is evidence that BAEA prefer nesting sites with some buffer to human
activity (Chandler et al 1995, Stevens and Brown 1997, USFWS 2007), although this varies widely across
their range.
The nesting cycle averages 18 weeks (or 126 days) from egg laying until fledging: incubation
begins with the first egg and lasts 5 weeks while the nestling period ranges from 8 – 14 weeks, averaging
12 – 13 weeks (Swenson et al 1986, Gende et al 1997, Buehler 2000, USFWS 2007, Saalfeld and Conway
2010, Mundahl et al. 2013). A female will usually lay 2 eggs, although 3 are not uncommon. In Colorado,
egg laying usually begins in late January to early February, hatching occurs in early March, and young
eagles can fledge anytime between early June and August. The post-fledging period, during which eaglets
remain partly dependent on adults, may last from 4 – 10 weeks.
Modeling of daily nest survival requires the following data from each nest: 1) date when the nest
was found; 2) the last day the nest was checked when alive; 3) the last day the nest was checked; and 4)
the fate of the nest (0=successful, 1=failed). For spatial covariates to be calculated, an accurate location
was also required. For at least the initial stages of modeling, analyses will be restricted to nests from the
past 5 years, because covariate data are more likely to be accurate for recent years.
Covariates are listed in Table 1 along with the data source and citations supporting their
inclusion. These include year, time in season, region (Front Range or West Slope), and several ecological
and anthropogenic variables. Dominant vegetation within 1 km will be categorized as developed, crop,
natural (forested), and natural (non-forested). Nest substrate will be categorized as live tree, dead tree, and
other. Water-related variables include distance to nearest major water body and total area of water within
3 km. Monthly weather variables will include average minimum temperature, average maximum
temperature, total precipitation, and number of days with severe storm events, using data from PRISM

6

�(www.prism.oregonstate.edu) and the National Oceanic and Atmospheric Administration storm events
database (http://www.ncdc.noaa. gov/stormevents). Monthly weather will be averaged (temperature) or
summed (precipitation) into seasonal values for the nesting (incubation + nestling period) and non-nesting
seasons. Anthropogenic covariates include total length of roads within 250 m and a binary variable for
whether or not human-caused disturbances were noted by the observer. J. DeCoste is calculating spatial
covariates using ArcGIS spatial analysis tools and running nest survival models in Program
MARK,which uses maximum likelihood methods to estimate nest survival as a known fate data type with
a logit link function. Model selection will be based on AICc values. We expect to complete the BAEA
nest survival model by the end of 2016, but protocols and coding will be well documented so that the
model can be updated in future as more nest and covariate data become available.
Distribution Models
The goal of distribution modeling is to determine what variables predict breeding locations and to
map areas with high to low probability of use for mitigation planning and future survey design. The next
phase of this research project will include distribution modeling for GOEA and BAEA. For GOEA, we
will collaborate with WGET, which has completed some ecoregional models and is currently working on
a High Plains model. Hopefully some aspects of this model structure, plus the covariates calculated for
the BAEA nest survival model, can be applied to the BAEA distribution model. Models for PRFA and
FEHA will follow, data permitting. Our database contains far fewer active nest records for these species,
compared to the eagles.
eBird
We are working with collaborators at Cornell to explore how eBird data can contribute to
distribution models. Similar data have been used by other researchers to model occupancy and
distribution. Data can be queried to assure some level of quality control, including restricting the output to
observations from complete checklists, if desired. We are still considering whether these data can be used
in model building versus model validation.
Additional Data Sources
We are exploring additional data sources, such as Breeding Bird Survey and the Colorado
Breeding Bird Atlas, for possible inclusion in or validation of future models. In addition, we may
investigate data collected during winter and migratory periods, as our four focal species are present in
Colorado outside of the nesting season. Furthermore, new nests are added to CPW’s raptor nest database
each year, so our goal is to produce well-documented modeling protocols that can be re-run as new data
become available.
Monitoring Guidelines
It is our objective to make recommendations for future monitoring guidelines that can be
consistently applied state-wide; we consider this just as important as the model outputs themselves. Any
suggested recommendations will be considered throughout the agency and particularly by the Terrestrial
Section, as CPW Biologists collect most of the nest data in our database, via their own surveys and by
acquiring and submitting data collected by partner organizations. We will also discuss model results and
monitoring guidelines with partner organizations, with the goal of producing high quality data in future,
collected in a consistent manner.
We plan to use the distribution maps that result from this research to stratify the state of Colorado
into appropriate survey regions. Ideally, monitoring guidelines will include a time of year for surveys,
how often various regions will be visited, and a means to record survey effort and both presence (new and
existing nests) and absence data. In addition, we hope to provide some guidance on the amount of effort
that should be allocated to surveys for new nests, versus return visits to known nest sites, and how to
prioritize survey effort by raptor species and spatial location.

7

�RESULTS
The CPW raptor nest database contained 9479 nest records for 8696 locations from the 1970s to
present, as of 7 November 2016 (Table 2). This included 1477 active nests known to be occupied within
the past 5 years. The majority of nest locations (6127 nests, 70% of the total) have an unknown or
undetermined status, meaning that the site has not been visited in at least 5 years or that an observer was
unable to determine the status of the nest. The only raptor species having more active nests than those of
unknown status are BAEA and osprey (Pandion haliaetus). There are 12 species with at least 100 nest
records; the remaining species have few nest records, and 27% of records are for nests occupied by an
unknown species, including those coded as stick nests, Buteo sp., and so forth. We made an agency-wide
and inter-agency request for additional nest data to maximize sample size prior to our analyses, which
likely contributed to the 20% increase (1730 nests) over 2 years. Our bald eagle nest survival model
currently has an input file containing 121 nest attempts at 79 locations from 2012−16: 45 nest sites are on
the Front Range and 34 are on the Western Slope. An additional 105 nests were not included, and the
most common reason for exclusion was that the observer did not visit enough times or at the appropriate
time to confirm nest fate.
Modeling efforts are still in progress. However, some useful information has been gleaned from
data quality control, calculation of covariates, and initial runs of the bald eagle nest survival model. First,
nest survival models require more intensive survey data than do distribution models, because at least two,
and frequently more visits are required during the nesting season. One visit must occur during the
incubation or nestling phases, in which nesting activity can be confirmed by sighting eggs or nestlings, or
at least by sighting an adult sitting on the nest who is presumably incubating or brooding. One visit must
occur shortly before or after the presumed fledging date to confirm the fate of the nest, where successful
nests have at least one fully feathered juvenile leaving the nest and failed nests have none (for BAEA, at
around 12 weeks post-hatch).
For all purposes, observers should reserve the value of zero to represent true zero, as opposed to a
placeholder or indication of an unknown or inapplicable value. The overly frequent entries of zero in the
current database have made it difficult to discern among these states, so in future, “zero”, “unknown”, and
“NA” should be entered with more specificity. Finally, 70% of nest locations in the database have a status
of unknown or undetermined (Table 2). These sites have much less value for modeling or HB 1298
purposes than have nests with any of the other status designations, so some effort should be allocated to
re-visiting these sites, which frequently turn out to be occupied (L. Rossi, pers. comm.).
DISCUSSION
We began exploring interest in this project and requesting additional data within and outside our
agency in 2014. Funding was secured in 2015 and a research associate started work on the project in
January 2016. Thus far we have worked on a nest survival model for BAEA, and we have met several
times with collaborators from WGET to discuss progress on a GOEA distribution model. Much of the
first half of 2016 was spent acquiring and proofing nest data and developing a set of covariate data layers.
We have also worked with collaborators at Cornell to explore the potential use of eBird data in
distribution models. The next steps will be distribution models on GOEA (with WGET) and BAEA. The
comparison of monitoring methods and model output for BAEA nest survival versus distribution models
should be especially instructive. Analyses of BAEA nest data have already provided some useful
recommendations for monitoring. This was intended as a 2-year research project, ending 31 December
2017, although it will take additional years to implement and evaluate the monitoring recommendations
that result from our work and from discussions among CPW employees and our partners working on
raptors.

8

�LITERATURE CITED
Bildstein, K.L. and D.M. Bird. 2007. Raptor research and management techniques (manual).
Buehler, D. A. 1995. A Geographic Information System to Identify Potential Bald Eagle Breeding Habitat
for Southeastern United States Rivers and Reservoirs. Proc. Annu. Conf. Southeast. Assoc. Fish
and Wildl. Agencies 49: 292-302.
Buehler, D. A. 2000. Bald Eagle (Haliaeetus leucocephalus), The Birds of North America (P. G.
Rodewald, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America:
https://birdsna.org/Species-Account/bna/species/baleag
Chandler, Sheri K., James D. Fraser, David a. Buehler, and Janis K. D. Seegar. 1995. Perch Trees and
Shoreline Development as Predictors of Bald Eagle Distribution on Chesapeake Bay. Journal of
Wildlife Management 59 (2): 325–32.
Daly, C., M. Halbleib, J. I. Smith, W. P. Gibson, M. K. Doggett, G. H. Taylor, J. Curtis, and P. P.
Pasteris. 2008. Physiographically sensitive mapping of climatological temperature and
precipitation across the con- terminous United States. International Journal of Climatology
28:2031– 2064.
Fleishman, Erica, James R Thomson, E L Kalies, B G Dickson, and D S Dobkin. 2014. Projecting
Current and Future Location , Quality , and Connectivity of Habitat for Breeding Birds in the
Great Basin. Ecosphere 5 (July): 1–29.
Fletcher Jr., Robert J, Shawn T McKinney, and Carl E Bock. 1999. Effects of Recreational Trails on
Wintering Diurnal Raptors along Riparian Corridors in a Colorado Grassland. Journal of Raptor
Research 33 (3): 233–39.
Fuller, M.R. and J.A. Mosher. 1987. Raptor survey techniques. Pages 37–65 in B. A. Giron, Pendleton, B.
A. Millsap, K. W. Cline, and D. M. Bird [EDS.], Raptor management techniques manual.
National Wildlife Federation, Washington, DC U.S.A.
Garrett, Monte G., James W. Watson, and Robert G. Anthony. 1993. Bald Eagle Home Range and Habitat
Use in the Columbia River Estuary. The Journal of Wildlife Management 57 (1): 19–27.
Gende, S M, M F Wilson, and M Jacobsen. 1997. Reproductive Success of Bald Eagles (Haliaeetus
leucocephalus) and Its Association with Habitat or Landscape Features and Weather in Southeast
Alaska.” Canadian Journal Of Zoology-Revue Canadienne De Zoologie 75 (10): 1595–1604.
Glenn, Elizabeth M., Robert G. Anthony, Eric D. Forsman, and Gail S. Olson. 2011. Reproduction of
Northern Spotted Owls: The Role of Local Weather and Regional Climate. Journal of Wildlife
Management 75 (6): 1279–94.
Greenwood, J.J.D. 2007. Citizens, science and bird conservation. Journal of Ornithology 148 (Suppl 1):
S77-S124.
Harmata, Alan R., and George J. Montopoli. 2001. Analysis of Bald Eagle Spatial Use of Linear Habitat.
Journal of Raptor Research 35 (3): 207–13.
Inselman, Will M, Shubham Datta, Jonathan A Jenks, Kent C Jensen, and W Grovenburg. 2015. Buteo
Nesting Ecology : Evaluating Nesting of Swainson’s Hawks in the Northern Great Plains,” 4–9.
Lopez-Lopez, Pascual, C. Garcia-Ripolles, Soutullo, L. Cadahia, and V. Urios. 2007. Identifying
Potentially Suitable Nesting Habitat for Golden Eagles Applied to ‘Important Bird Areas’ Design.
Animal Conservation 10 (2): 208–18.
López-López, Pascual, Clara García-Ripolĺs, José Miguel Aguilar, Francisco García-López, and José
Verdejo. 2006. “Modelling Breeding Habitat Preferences of Bonelli’s Eagle (Hieraaetus
fasciatus) in Relation to Topography, Disturbance, Climate and Land Use at Different Spatial
Scales.” Journal of Ornithology 147 (1): 97–106.
Mathieu, R, P Seddon, and J Leiendecker. 2006. Predicting the Distribution of Raptors Using Remote
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Zealand Falcon (Falco novaeseelandiae). New Zealand Journal of Zoology 33:1 73-84.

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�Mundahl, N.D., Anthony G. Bilyeu, and Lisa Maas. 2013. Bald Eagle Nesting Habitats in the Upper
Mississippi River National Wildlife and Fish Refuge. Journal of Fish and Wildlife Management 4
(2): 362–76.
Saalfeld, Sarah T, and Warren C Conway. 2010. Local and Landscape Habitat Selection of Nesting Bald
Eagles in East Texas. Southeastern Naturalist 9 (4): 731–42.
Sergio, F., T. Caro, D. Brown, B. Clucas, J. Hunter, J. Ketchum, K. McHugh and F. Hiraldo. 2008.Top
Predators as Conservation Tools: Ecological Rationale, Assumptions, and Efficacy. Annu. Rev.
Ecol. Evol. Syst. 39:1-19.
Stevens, Lawrence E., and Bryan T. Brown. 1997. Winter Bald Eagle Distribution Is Inversely Correlated
with Human Activity Along the Colorado River, Arizona. Journal of Raptor Research 31 (1): 7–
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Swenson, Jon E., Kurt L. Alt, and Robert L. Eng. 1986. Ecology of Bald Eagles in the Greater
Yellowstone Ecosystem. Wildlife Monographs, no. 111: 3–57.
Tack, Jason D., and Bradley C. Fedy. 2015. Landscapes for Energy and Wildlife: Conservation
Prioritization for Golden Eagles across Large Spatial Scales. PLoS ONE 10 (8): 1–18.
Theobald, D M, J R Miller, and N T Hobbs. 1997. Estimating the Cumulative Effects of Development on
Wildlife Habitat. Landscape and Urban Planning 39 (1): 25–36.
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10

�TABLES
COVARIATE
Time and Site
Year
Time in season

Site
Land Class
Dominant Vegetation

Anthropogenic
Roads 250

Disturbance

Nest Substrate
Substrate

DESCRIPTION

DATA SOURCE

REFERENCES

Categorical variable for dominant
class within 1000 m
•
Crop
•
Developed
•
Riparian Forest
•
Other Natural

Basinwide, LANDFIRE

Inselman et al. 2015, Lopez-Lopez 2007, Watts and
Duerr 2010, Saalfeld and Conway 2010, Harmata and
Montopoli 2001, Tack and Fedy 2015, Watts 2015

Total length of roads within 250 m

BASE.DBO.Roads_Tiger2008,
BASE.DBO.Roads_Tiger2010

Variable indicating whether or not
a human-caused disturbance was
reported by the observer

Original input datasheets

Buehler 1995, Mathieu et al. 2006, Mathieu et al. 2006,
Lopez-Lopez 2005, Lopez-Lopez 2007, Watts 2015,
Fletcher, McKinney, Bock 1999, Chandler, Faser,
Buehler et al. 1995, Tack and Fedy 2015, Theobald et
al. 1997, Lopez-Lopez et al. 2006, Wallace et al. 2016
Watts 2015, Steven and Brown 1997, Fletcher,
McKinney, Bock 1999, Chandler, Faser, Buehler et al.
1995, Tack and Fedy 2015, Theobald et al. 1997

Categorical variable for nesting
substrate
•
Live Tree
•
Dead Tree
•
Other

Original input datasheets

2012−2016
Integer value for date where day 1
is earliest date of incubation in
dataset
Front Range or West Slope

11

Buehler 1995, Wallace et al. 2016, Watts and Duerr
2010, Watts 2015, Mike Sherman, CPW Biologist,
pers. comm., 2016, Will Keeley, CBOSMP, pers.
comm., 2016

�Foraging Habitat
Water distance

Distance to nearest major water
body (lakes, reservoirs, rivers)

CDOW.DBO.MajorRivers100k,
CDPW.DBO.WaterbodiesNHD_USGS

Pdog distance
(unlikely to have data)

Distance to nearest prairie dog
colony

SAM.DBO.WTPrairieDogColonies,
need more data in the Front Range

Water area

Total area of water within 3 km
(%)

CDOW.DBO.MajorRivers100k,
CDPW.DBO.WaterbodiesNHD_USGS

Averaged monthly minimum/
maximum daily temperatures (°C)
used to obtain mean seasonal
values during incubation, nestling,
and non-nesting periods
Total monthly precipitation values
(mm) summed to obtain seasonal
values during incubation, nestling,
and non-nesting periods
Number of days with severe storm
events (hail, heavy rain, heavy
snow, high wind, thunderstorm
wind, winter storms) during
incubation, nestling, and nonnesting periods

PRISM (www.prism.oregonstate.edu)

Daly et al. 2008, Wallace et al. 2016, Tack and Fedy
2015, Fleishman et al 2014, Gende et al. 1997, Glenn
et al. 2001

PRISM (www.prism.oregonstate.edu)

Daly et al. 2008, Wallace et al. 2016, Tack and Fedy
2015, Fleishman et al 2016, Gende et al. 1997, Glenn
et al. 2011

National Oceanic and Atmospheric
Administration storm events
database(http://www.ncdc.noaa.
gov/stormevents)

Wallace et al. 2016, Gende et al. 1997, Glenn et al.
2011.

Weather
Temp min
Temp max

Precip

Storm

Gende et al. 1997, Swenson et al. 1986, Mundahl et al.
2013, Harmata and Montopoli 2001, Watts 2015, Mike
Sherman, CPW Biologist, pers. comm., 2016, Will
Keeley, CBOSMP, pers. comm., 2016
Gende et al. 1997, Swenson et al. 1986, Mundahl et al.
2013, Harmata and Montopoli 2001, Watts 2015, Mike
Sherman, CPW Biologist, pers. comm., 2016, Will
Keeley, CBOSMP, pers. comm., 2016
Gende et al. 1997, Swenson et al. 1986, Mundahl et al.
2013, Harmata and Montopoli 2001, Watts 2015, Mike
Sherman, CPW Biologist, pers. comm., 2016, Will
Keeley, CBOSMP, pers. comm., 2016

Table 1. Covariates used in models of daily nest survival for bald eagles in Colorado, 2012−2016
.

12

�SPECIES
Bald Eagle
Golden Eagle
Ferruginous Hawk
Prairie Falcon
Total All Species

Active
194
103
63
25
1477

NUMBER OF NESTS
Inactive Destroyed Unk/Undeter
18
56
152
71
80
1630
3
29
325
13
1
163
474
618
6127

Total*
437
1978
436
215
9479

Table 2. Nest counts for Colorado raptors included in the CPW raptor nest database as queried on 7
November 2016. Active nests are those known to be occupied within the past 5 years. Inactive nests are
those known to be unoccupied within the past 5 years. Destroyed nests are those known to be no longer
usable (e.g., tree or branch has fallen). Unknown nests are those that have not been visited within the past
5 years, excluding destroyed nests. Undetermined nests are those for which status could not be
determined by an observer within the past 5 years. Counts of active, inactive, destroyed, unknown, and
undetermined nests are for the last raptor species occupying a nest site. *Total nest numbers exceed the
summation of those in specific status categories due to nest sites that have changed species or have
missing data.

13

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