Demographics of the Golden-cheeked

Warbler (Dendroica chrysoparia) on Fort

Hood, Texas

by

Leslie A. Jett

Timothy J. Hayden

John D. Cornelius

The Golden-cheeked Warbler (Dendroica

chrysoparia) is a federally endangered

migratory passerine that has its breeding

range contained in the central Texas area that

includes Fort Hood. Its listing as endangered

in 1991 was a result of the rapid degradation

and increasing fragmentation of habitat,

causing a decline in their population. To assist

in conservation and recovery of the Goldencheeked Warbler and comply with the Endangered Species Act, environmental managers

need information on the demographic

parameters of the population on Fort Hood.

Researchers surveyed and documented the

Golden-checked Warbler on Fort Hood, Texas

between 1991 and 1996. The population remained relatively stable, with a slight decrease

in 1996. The average return rate for males

banded as adults was 45.5%; for males

banded as juveniles the average was 16.9%.

Dispersal distances for males banded as

adults averaged 223 m. Dispersal distances

for males banded as juveniles were significantly greater, with an average of 4,040 m.

Densities of territorial males remained relatively stable between 1992 and 1996, with a

peak density of 28 males per 100 ha (average

of 18 males per 100 ha). Productivity of males

within an intensive study area averaged 2.0

young per mated male.

US Army Corps

of Engineers

Construction Engineering

Research Laboratories

USACERL Technical Report 98/52

March 1998

Approved for public release; distribution is unlimited.
2 USACERL TR 98/52

Foreword

This study was conducted for Headquarters, III Corps and Fort Hood, TX, under

Military Interdepartmental Purchase Request (MIPR) 6FCER00439. The technical

monitor was John D. Cornelius.

The authors recognize the following people for their contributions. Howard Weinberg

provided insightful discussions concerning the status of the Golden-cheeked Warbler

on Fort Hood, and helped with early drafts of this manuscript. Robert Melton helped

with the statistics used in this study. Theresa Koloszar assisted in many aspects of

data management. Dean Keddy-Hector provided helpful discussions of differences

between populations around Austin, TX and Fort Hood. Several individuals were

responsible for collecting much of the field data. Mike Stake, Maria Elena Tolle,

Deborah Maas-Burleigh, John Styrsky, Edd Paradise, Danny Huffine, Wanda Eckrich,

Carrie Bemus, and Jeff Bolsinger conducted the field work during different times of

the study. Steve Mackie developed computer applications that facilitated data

collection and management, and helped to estimate the amount of Golden-cheeked

Warbler habitat present on Fort Hood. Randy Craft reviewed the manuscript.

Dennis Herbert, B.R. Jones, Tim Buchanan, Kevin Cagle, and Anne Hamilton of the

Natural Resources Branch at Fort Hood assisted in all aspects of data collection,

facilitated access, and helped coordinate field activities.

This research was supported in part by an appointment to the Research Participation

Program at the U.S. Army Construction Engineering Research Laboratory

administered by the Oak Ridge Institute for Science and Education through an

interagency agreement between the Department of Energy and USACERL.

This work was performed by the U.S. Army Construction Engineering Research

Laboratories, (USACERL) Natural Resources Assessment and Management Division

(LL-N). Dr. William D. Severinghaus is Operations Chief, CECER-LL. The

USACERL technical editor was Gloria J. Wienke, Technical Resources.

COL James A. Walter is Commander and Dr. Michael J. O’Connor is Director of

USACERL.
USACERL TR-98/52 3

Contents

Foreword............................................................................................................................ 2

List of Figures and Tables................................................................................................ 5

1 Introduction.............................................................................................................. 7

Background......................................................................................................................7

Objectives ........................................................................................................................9

Approach..........................................................................................................................9

Mode of Technology Transfer .........................................................................................10

2 Methods..................................................................................................................11

Data Collection...............................................................................................................11

Data Analysis.................................................................................................................15

3 Results....................................................................................................................16

Return rates ...................................................................................................................16

Dispersal Distances .......................................................................................................21

Detection of Territorial Males..........................................................................................23

Age Structures ...............................................................................................................24

Mated Status Within the Intensive Study Area...............................................................26

Productivity ....................................................................................................................27

Parasitism ......................................................................................................................30

4 Discussion .............................................................................................................32

Population Trends...........................................................................................................32

Age-related Differences .................................................................................................36

Relevance/Application to Predictive Models ..................................................................43

Comparison With Similar Species..................................................................................43

Incidence of Cowbird Parasitism....................................................................................45

5 Conclusions and Recommendations .................................................................46

References.......................................................................................................................47

Appendix A: List of all Adult Males Resighted on Fort Hood. .................................50
4 USACERL TR 98/52

Appendix B: List of all Adult Males Searched for and Not Found on Fort Hood...57

Appendix C: List of all Males Located Within the ISA Including Non-territorial

Birds........................................................................................................................63

Appendix D: List of all Adult Females Resighted on Fort Hood..............................71

Appendix E: List of all HY Males’ First Resightings on Fort Hood. .........................72

Appendix F: List of all HY Females Resighted on Fort Hood...................................73

Appendix G: Description of Palmer Drought Severity Index....................................74

Distribution
USACERL TR-98/52 5

List of Figures and Tables

Figures

1 Map of Fort Hood, TX, showing approximate extent of Golden-cheeked Warbler habitat,

including areas burned in February 1996 ......................................................................8

2 Map of Fort Hood, TX, showing approximate locations of intensive study area (ISA) and

areas burned in February 1996 fire................................................................................9

3 Map of the ISA (training area 13B) with Golden-cheeked Warbler habitat shown by a thick

white line ......................................................................................................................12

4 Annual return rates for newly banded ISA and non-ISA males on Fort Hood..............17

5 Return rates for adult males within the ISA, by age and year. .....................................18

6 Return rates for non-ISA adult males, by age class and year. .....................................20

7 Percentage of SY individuals in the ISA and non-ISA training areas. ..........................26

8 Percent of mated males within the ISA, by age class and year....................................27

9 Productivity for ISA SY males, by year.........................................................................29

10 Monthly averages of the Palmer Drought Severity Index (PDSI) for Texas, Division 7 (of

which Fort Hood is a part), from 1895 to 1996 ............................................................33

11 Map of all territorial males within the ISA during 1996.................................................38

12 Map of all territorial males within the ISA during 1995.................................................39

13 Map of all territorial males within the ISA during 1994.................................................40

14 Map of all territorial males within the ISA during 1993.................................................41

15 Map of all territorial males within the ISA during 1992.................................................42

Tables

1 Total number of HY Golden-cheeked Warblers captured yearly on Fort Hood, including the

ISA and non-ISA training areas....................................................................................13

2 Annual return rates for adult male Golden-cheeked Warblers. ....................................16

3 Return rates for ISA males present and territorial for 1-year periods between 1992 and 1996.

.....................................................................................................................................18

4 Return rates for searched-for males in non-ISA training areas for 1-year periods between

1992 and 1996. ............................................................................................................19

5 Dispersal distances for all female Golden-cheeked Warblers banded as adults..........20

6 Return rates for all hatch year (HY) Golden-cheeked Warblers...................................21

7 Dispersal distances summarized by age class for male Golden-cheeked Warblers banded as

adults on Fort Hood. ....................................................................................................22

8 Dispersal distances for adult males banded in the intensive study area (ISA), banded males

searched for, and banded males observed opportunistically (“chance-find”). .............22
6 USACERL TR 98/52

9 Dispersal distances for all male Golden-cheeked Warblers banded as HY on Fort Hood.

.....................................................................................................................................23

10 Dispersal distances for female Golden-cheeked Warblers banded as hatch year (HY).23

11 Number of territorial Golden-cheeked Warblers present in the ISA. ............................24

12 Number of male Golden-cheeked Warblers detected in all training areas (including the ISA),

and the total number of training areas in which males were detected. ........................24

13 Age structure of male Golden-cheeked Warblers present in ISA.................................25

14 Age structure of male Golden-cheeked Warblers observed outside the ISA...............25

15 Total number of males banded by age class on Fort Hood..........................................26

16 Percent of ISA males mated, by age class...................................................................27

17 Mean number of young fledged by successful ISA males, by age class and year.......28

18 Mean number of young fledged by territorial ISA males, by age class and year..........28

19 Mean number of young fledged by mated ISA males, by age class and year..............30

20 Success rate of ISA Golden-cheeked Warbler males. .................................................30

21 Summary of outcomes for all located Golden-cheeked Warbler nests on Fort Hood. .31

22 Mean daily temperatures and relative humidities for the month of March for Fort Hood from

1994-1996....................................................................................................................34
USACERL TR-98/52 7

1 Introduction

Background

The Golden-cheeked Warbler (Dendroica chrysoparia) is a federally endangered

migratory passerine that has its breeding range contained within the range of

Ashe juniper (Juniperus Ashei) on the Edward’s Plateau of central Texas. Its

listing as endangered in 1991 was a result of the rapid degradation and

increasing fragmentation of habitat, causing a decline in their population

(Benson 1990). Pulich (1976) estimated the breeding population to be

approximately 14,950 birds. Wahl, Diamond, and Shaw (1990) believed that

their estimation of 4,822 to 16,016 breeding pairs was “unrealistically” high. As

of 1995, the species was believed to breed in at least 24 central Texas counties, a

reduction of 23% since 1976 (31 counties; Pulich 1976; U.S. Fish and Wildlife

Service [USFWS] 1996).

The species is entirely dependent upon the mature Ashe juniper forests of

central Texas for its nesting and foraging substrates. The species builds its

nests almost entirely from shredded juniper bark at a height of 4 to 5 meters,

most commonly in Ashe juniper and occasionally in hardwoods common to the

region (e.g., live oak, Texas oak; Pulich 1976). Only mature Ashe juniper trees

shred their bark, limiting the Golden-cheeked Warbler to forests with a high

proportion of juniper trees older than 40 to 50 years (Pulich 1976). Warblers,

however, rarely exist in stands of pure juniper, as they also require hardwoods

for foraging substrate. Early in the season, they forage most frequently in

deciduous hardwood trees, primarily oaks, then switch to juniper as a principal

foraging substrate later in the season (Beardmore 1994). For a more detailed

description of the Golden-cheeked Warbler and its habitat, see Pulich (1976).

Fort Hood, Texas is located partially in both Bell and Coryell counties in east

central Texas. Vegetative cover on the installation varies with slope, aspect,

moisture, and previous use (e.g., cattle grazing and military activities), and

consists of grassland, open savannah, hardwood thickets, and dense juniper-oak

stands. For a more detailed habitat description of Fort Hood, see Tazik and

Cornelius, 1993. Within its 87,890 hectares (ha) is approximately 16,000 ha of

Golden-cheeked Warbler habitat.
8 USACERL TR 98/52

The Golden-cheeked Warbler was listed as endangered in 1991 under the

Endangered Species Act (ESA) of 1973, as amended. To assist in conservation

and recovery of the Golden-cheeked Warbler and comply with the ESA,

environmental managers need information on the demographic parameters of

the population on Fort Hood. In 1991, these demographics were not available.

Quality and amount of warbler habitat appeared stable on Fort Hood between

1991 and 1995 (Figure 1). However, in February of 1996, a large wildfire

destroyed approximately 4,300 ha (25%) of warbler habitat, mostly in the

northern and northeastern sections of Fort Hood (Figure 2). A separate study

was initiated to investigate the effects of this fire on warbler return rates and

dispersal distance; males affected by the fire were not reported in this paper,

and their numbers were not used in analyses of return rate or dispersal

distance.

Figure 1. Map of Fort Hood, TX, showing approximate extent of Golden-cheeked Warbler habitat,

including areas burned in February 1996. (1 cm ~ 2,900 meters.)
USACERL TR-98/52 9

= Intensive study area (ISA), 1992-1996

= All areas burned in February, 1996

= GCWA habitat not burned in February, 1996

= 1991 Intensive study area

Figure 2. Map of Fort Hood, TX, showing approximate locations of intensive study area

(ISA) and areas burned in February 1996 fire. (1 cm ~ 3,100 m.)

Objectives

The objectives of this work were to determine and document (1) demographic

parameters (return rates, dispersal distances, productivity, and mated status)

affecting population status and (2) occurrence of the Golden-cheeked Warbler on

Fort Hood, Texas.

Approach

Research on the Golden-cheeked Warblers was conducted on Fort Hood, Texas

between 1991 and 1996. Landsat MSS data were integrated with a Geographic
10 USACERL TR 98/52

Information System (GIS) application, Geographic Resources Analysis Support

System (GRASS), to suggest potential warbler habitat. Researchers then

surveyed these areas, documenting the presence or absence of habitat, and

numbers of birds seen or heard. Researchers also established an intensive study

area in 1991, banding Golden-cheeked Warblers, and determining territory,

mated status, and productivity. The research work consisted of systematically

surveying possible Golden-cheeked Warbler habitat, color-banding adults, and

monitoring an intensive study area to determine species productivity. Data from

these studies were compiled and analyzed, and recommendations were made

regarding the monitoring and management of the Golden-cheeked Warbler

population on Fort Hood, Texas. Study methods are described in Chapter 2 of

this report and in Hayden, Jett, and Weinberg (in prep.).

Mode of Technology Transfer

This research contributes to a fundamental understanding of the ecology of the

endangered Golden-cheeked Warbler, and serves as an example of a proactive

approach to endangered species management on Army lands. This and other

related reports are being transmitted to military, land, and wildlife managers at

Fort Hood, TX; Headquarters, U.S. Army Forces Command (HQ FORSCOM);

and the Department of the Army for use in ESA compliance efforts.
USACERL TR-98/52 11

2 Methods 

This chapter provides a brief overview of research methods between 1991 and

1996. For a detailed description of field methods used in this study, see Hayden,

Jett and Weinberg (in prep.).

Data Collection

The research focus changed in the years following the project’s initiation in

1991. The focus shifted from a survey of the entire population of Golden-cheeked

Warblers to documenting specific demographic and life history characteristics.

This change of focus allowed collection of detailed information regarding return

rates, dispersal distances, productivity, mated status, and age structures.

In 1991, a study area was established at the boundary of training areas 2 and

3A. It was necessary to change the location of the study area the following year

due to difficulties accessing the site. A new intensive study area (13B, hereafter

referred to as the Intensive Study Area [ISA], Figures 2 and 3) with an

approximate area of 178 ha was established in 1992 in the eastern side of

training area 13B. The 13B/ISA is relatively isolated from other patches of

warbler habitat. It is enclosed on the north and northeast by Lake Belton, on

the south by pasture, and on the southeast by savannah (open, partially wooded

grassland). To the west of the study area is a combination of grassland (west)

and stands of pure juniper (southwest) that were unoccupied by warblers. The

northwestern section of the study area was attached to more extensive warbler

habitat by a “corridor” of warbler habitat.

Golden-cheeked Warblers were captured using mistnets and playbacks of

recorded songs and calls. Individuals were banded, aged, and sexed according to

Pyle, et al. (1987) and Hayden, Jett, and Weinberg (in prep). For a review of

warbler-specific mistnetting techniques, see Hayden, Jett, and Weinberg (in

prep); and Weinberg, Jett, and Cornelius (1996). The methods used to age and

sex adult warblers during banding were refined during the course of this study.

In 1992, it was found that adult plumage characteristics could reliably be used

to age to the “second year” (SY) or “after-second-year” (ASY) age classes rather

than the general age class of “after-hatch-year” (AHY) used by Pyle, et al.

(1987).
12 USACERL TR 98/52
USACERL TR-98/52 13

Figure 3. Map of the ISA (training area 13B) with Golden-cheeked Warbler habitat

shown by a thick white line. (1 cm ~ 100 m.)

However, the AHY age class was still used on Fort Hood after June 15 since

most adults were in molt by then and could not be reliably aged. The majority of

males banded in a given year were aged more specifically than “AHY” (i.e., to SY

or ASY) beginning at 18% in 1992 and increasing to 96% in 1993, remaining

high for the rest of the study.

Sex determination of “hatch year” (HY) birds was not as reliable as it was for

adults, as individuals were captured at different stages of their sex-specific

plumage development. Males could be reliably sexed much earlier than females,

because at a young age males generally show the distinct beginning to the dark

throat and chest that is characteristic of adult males. If the HY bird was

captured late enough in the season (or late enough in the individual’s

development), it could be reliably sexed, regardless of its sex. Table 1 shows the

total number of HY birds captured between 1991 and 1996, separated by sex.

The ratio of banded, known-sex hatch year birds was approximately 1.86:1

(male:female), with 17% of the total HY capture population labeled as

“unknown-sex.” Since it is unknown if males and females return and disperse at

the same rates, only known-sex hatch year individuals were used to analyze

hatch year return rates and dispersal distances.

Return rates were estimated as the ratio of banded males that were found by

searching to the total of those searched for (including both found and not found).

Previously banded male Golden-cheeked Warblers were searched for by

returning to the last location the bird was sighted (or banded). Males resighted

by this method were categorized as having been “searched for.” If a banded male

was resighted opportunistically (e.g., was not being searched for at the time it

was found), the resighting was considered an “incidental” find. Dividing

returning birds into categories was necessary to evaluate bias in different

approaches to return rate estimations. The use of opportunistically resighted

males would bias the estimated return rate toward a higher-than-realistic

figure.

Table 1. Total number of HY Golden-cheeked Warblers captured yearly on Fort Hood,

including the ISA and non-ISA training areas.

Year Male Female Unknown Total Captured

1991 2 0 6 8

1992 27 8 20 55

1993 16 6 9 31
14 USACERL TR 98/52

1994 47 30 8 85

1995 56 34 7 97

1996 22 12 4 38

Total 170 90 54 314

A third resighting category consisted of those territorial males within the ISA.

The ISA was considered separately because of the increased detection rate due

to the intensive monitoring procedures. Therefore, the inclusion of the ISA in

the overall return rate estimation would also bias the results. All males

resighted are listed in Appendix A, which is sorted by resight type. All males

searched for and not found are listed in Appendix B.

Site fidelity was estimated by calculating dispersal distance between years.

Dispersal distance was determined by Pythagorean’s theorem, using the original

and resighting locations to obtain a straight-line distance. If a banded bird was

resighted in more than one year, the dispersal distance for the first resighting

was that between the original banding location and the resighting location; the

dispersal distance for the second (and subsequent) resighting was between the

first resighting location and the second (or third, etc.). Between 1992 and 1995,

researchers chose males to search for, based upon location. For example, field

technicians decided in which training area they were going to work on a

particular day, and looked at previous years’ maps of banding and resighting

locations within that training area to have a starting point for searching. In

1996, however, this protocol changed somewhat. A list was established prior to

the season, with band combinations and locations of certain males to be

searched for. This list included both older and newly banded males, in order to

obtain a broader mix of specific age-class return rates, rather than just for those

younger males banded in recent years.

Productivity within the intensive study area was estimated by documenting the

number of hatch year warblers being fed (by both males and females) on each

territory each year. In 1992, specific numbers of HY were not always recorded

for each territory within the ISA, as was standard protocol in later years. As

these methods differed from those in later years, these data were excluded when

analyzing overall and age-specific productivity. Productivity for 1992 was

gathered from Bolsinger and Hayden (1992). Also, data from the 1991 season

was not used for overall and age-specific productivity because birds weren’t aged

to specific age classes in 1991, different data were recorded in the baseline year,

and because it was in a different location (Training Areas 2 and 3A vs. the ISA).

Data recorded for observations of males outside the ISA generally included only

their location and band combination. For those males found within the ISA,
USACERL TR-98/52 15

however, more detailed demographic data were collected, including the

following: presence or absence of a female, number of HY birds observed (and

whether or not they were being fed), location of a nest (whenever possible), and

territory boundaries (determined by the males regularly visiting certain areas

and engaging in singing bouts and fights with neighboring males).

Data Analysis

Differences in return rates and mated status were analyzed by chi-square tests

of differences. Any chi-square test that violated Cochran’s rule of small sample

sizes had data lumped into adjacent categories until it no longer violated this

assumption. For all tests conducted, a chi-square test of heterogeneity was used

before attempting to lump years together. If the calculated value in the

heterogeneity test was not significant, the years were lumped, and were not

considered to be statistically different. If the calculated value in the

heterogeneity test was significant, the years were not lumped, as they were

considered statistically different.

Second year male percentages were calculated by dividing the number of banded

SY males in the population (including newly banded males and returning HYs)

by the total number of known-age banded adult males in the population (e.g.,

AHY, HY, and U-age males were excluded from these calculations).

When analyzing dispersal distances, the age classes of 3-year (3Y) vs. 4-year

(4Y) vs. “after 4-year” (A4Y+; which included A4Y, 5Y, A5Y, 6Y, and A6Y age

classes) were tested by a Kruskal-Wallis test to determine whether the age

classes dispersed differently. These age classes were chosen to reduce possible

overlap between “unknown” ages; for example, the ASY age class may contain

males in the 4Y, 5Y, etc. age classes. The Kruskal-Wallis test was also used to

determine significance in dispersal distances between the following groups: (1)

HY and adult males, (2) HY and adult females, and (3) ISA “chance” find and

searched-for males.

Return rates for 1992 were not calculated, as a separate list of males searched

for and not found was not maintained. It was also not noted in 1992 if resighted

males were found by “chance” or were searched for. “Annual return rate” refers

to males banded one year that were searched for the next year.

All tests were performed in Excel 5.0 and/or SPSS, version 6.1 for the PC.
16 USACERL TR 98/52

3 Results

Return rates

Annual Return Rates

Return rates were evaluated for the years 1992-93, 1993-94, 1994-95, and 1995-

96. Overall annual return rate of adult Golden-cheeked Warbler males in the

ISA was 51.8% (n = 83; Table 2, Figure 4; Appendices A, B, and C). Annual

return rates ranged between 31.3% (n = 32) in 1996 to 72.2% (n = 18) in 1995.

The return rate of 31.3% in the ISA in 1996 is considered low, as the return rate

for 1992 through 1995 was 64.7% (n = 51, Table 2). When tested by chi-square,

these differences were significant (2

 = 14.75, df = 3, P = 0.002).

Annual return rates of males in non-ISA training areas were consistently and

significantly lower than those within the ISA, with an overall rate of 42.7% (n =

185, 2 = 8.35, df = 3, P = 0.039; Table 2, Figure 4; Appendices A and B). The

highest annual return was 52.8% (n = 72) in 1994. The lowest rate was in 1996,

with 29.8% (n = 57).

Table 2. Annual return rates for adult male Golden-cheeked Warblers.*

Year

banded

Year

resighted

ISA return rate Non-ISA return rate Combined return rate

1992 1993 60% (20) 48.6% (37) 52.6% (57)

1993 1994 61.5% (13) 52.8% (72) 54.1% (85)

1994 1995 72.2% (18) 31.6% (19) 51.4% (37)

1995 1996 31.3% (32) 29.8% (57) 30.3% (89)

All years 51.8% (83) 42.7% (185) 45.5% (268)

* This table excludes AHY and HY males. (n) = Number of adult males banded that were searched for

in the following year.
USACERL TR-98/52 17

n = 32

n = 18

n = 13 n = 20

n = 19 n = 57

n = 72

n = 37

0

10

20

30

40

50

60

70

80

1992-

1993

1993-

1994

1994-

1995

1995-

1996

Year banded-year resighted

Percent resighted

ISA

non-ISA

Figure 4. Annual return rates for newly banded ISA and non-ISA males on Fort Hood.

While the overall annual return rate was lower for non-ISA training areas than

for the ISA (42.7%, n = 185, and 51.8%, n = 83, respectively), it was not

statistically different (2

 = 1.57, df = 1, P = 0.21). When the ISA and non-ISA

return rates were lumped by year (test of heterogeneity: 2

 = 6.12, df = 4, P =

0.19), it was found that the difference between years was significant, with the

1996 overall return rate the lowest at 30.3% (n = 89), and the 1994 overall

return rate the highest at 54.1% (n = 85, 2 = 12.48, df = 3, P = 0.006.

Overall, 228 males were resighted at least one time; 51 males at least two times;

17 males at least three times; and four males were resighted four times

(Appendix A).

Overall Return Rates

When looking at all adults present and territorial within the ISA one year that

returned the next, the Golden-cheeked Warbler male had an overall return rate

of 48% (n = 127), with a low of 30% (n = 50) in 1996 and a high of 65.6% (n = 32)

in 1995 (Table 3, Appendices A and C). This among year difference was

significant (2 = 12.03; df = 3, P = 0.007). When the age classes were tested

separately among years, SY and ASY return rates were both significantly

different (SY: 2

 = 7.85, df = 3, P = 0.049; ASY: 2 = 10.67, df = 3, P = 0.014; Table

3; Figure 5). SY males returned with the highest frequency in 1994, at 81.8%,

and the lowest in 1996, with 30%. All ASY males returned with the highest

frequency in 1995, at 76.5%, and the lowest in 1996 with 30%. Overall rates of

return, however, did not differ significantly when divided into separate age

classes: SY males returned at 49.1% (n = 53) whereas all ASY lumped together
18 USACERL TR 98/52

returned at 48.6% (n = 70, 2 = 1.017, df = 1, P = 0.90). When the age classes

were further divided into specific age groups (of SY, 3Y, ASY, A3Y, and 4Y+),

there were still no significant differences in overall return rates (P > 0.05). ASY

males returned most frequently at 57.7% (n = 26), and A3Y males returned the

least at 36.4% (n = 11, 2

 = 1.50, df = 4, P = 0.83).

Table 3. Return rates for ISA males present and territorial for 1-year periods between 1992 and

1996.*

Age Class 1992-1993 1993-1994 1994-1995 1995-1996 All years

SY 3/7 = 42.9% 9/11 = 81.8% 8/15 = 53.3% 6/20 = 30% 26/53 = 49.1%

All ASY 7/11 = 63.6% 5/12 = 41.7% 13/17 = 76.5% 9/30 = 30% 34/70 = 48.6%

 ASY 7/9 = 77.8% 1/2 = 50% 3/3 = 100% 4/12 = 33.3% 15/26 = 57.7%

 3Y 1/3 = 33.3% 6/8 = 75% 2/8 = 25% 9/19 = 47.7%

 A3Y 0/2 = 0% 3/6 = 50% 1/1 = 100% 0/2 = 0% 4/11 = 36.4%

 4Y+ 3/4 = 75% 3/8 = 38% 6/12 = 50%

 4Y 1/1 = 100% 2/5 = 40% 3/6 = 50%

 A4Y 2/3 = 66.7% 0/1 = 0% 2/4 = 50%

 5Y 1/1 = 100% 1/1 = 100%

 A5Y 0/1 = 0% 0/1 = 0%

AHY 1/4 = 25% 0/1 = 0% 1/1 = 100% 0/0 =0% 2/6 = 33.3%

All ages 11/22 = 50% 14/23 = 60.9% 21/32 = 65.6% 15/50 = 30% 61/127 = 48%

* Numbers indicate those found divided by those present and territorial previous year. “Age Class” refers to age at

banding or most recent observation. “ASY” in this table refers only to returning AHY males, and newly banded ASY

males. “All ASY” is a total of all banded males, excluding SY and AHY.

n = 20

n = 15

n = 11

n = 7

 n = 30

n = 17

 n = 12

n = 11

0

1 0

2 0

3 0

4 0

5 0

6 0

7 0

8 0

9 0

1993 1994 1995 1996

Year resig hted

Percent resighted

S Y

all A S Y

Figure 5. Return rates for adult males within the ISA, by age and year.
USACERL TR-98/52 19

As with the ISA, there was a significant difference among years in average

return rates for males outside the ISA (all ages combined, 2

 = 23.56, df = 3, P =

0.00003), with an average of 32.9% (n = 350), and a range of 19.2% (1996, n =

120) to 51.1% (1994, n = 88, Table 4, Figure 6). Unlike the ISA, however, there

was a significant difference in return rates between age classes outside the ISA

for all years combined; SYs returned more frequently (47.2%, n = 72) than all

ASYs combined (28%, n = 225, 2

 = 8.31, df = 1, P = 0.004). When all age classes

were considered separately (SY, 3Y, 4Y, ASY, A3Y, A4Y+), the difference was still

significant (2

 = 12.98, df = 5, P = 0.024).

Table 4. Return rates for searched-for males in non-ISA training areas for 1-year

periods between 1992 and 1996.*

Age class 1992-1993 1993-1994 1994-1995 1995-1996 All years

SY 4/7 = 57.1% 15/27 = 55.6% 6/12 = 50% 9/26 = 34.6% 34/72 = 47.2%

All ASY 8/28 = 28.6% 24/42 = 57.1% 18/62 = 29% 13/93 = 14% 63/225 = 28%

 ASY 5/17 = 29.4% 17/29 = 58.6% 5/26 = 19.2% 8/34 = 23.5% 35/106 = 33%

 3Y 0/2 = 0% 2/4 = 50% 4/8 = 50% 1/11 = 9.1% 7/25 = 28%

 A3Y 3/9 = 33.3% 4/8 = 50% 4/11 = 36.4% 0/19 = 0% 11/47 = 23.4%

 4Y 1/1 = 100% 0/1 = 0% 0/12 = 0% 1/14 = 7.1%

 A4Y+ 4/6 = 67% 4/17 = 24% 8/23 = 35%

 A4Y 2/3 = 66.7% 2/11 = 18.2% 4/14 = 28.6%

 5Y 0/1 = 0% 0/0 = 0% 0/1 = 0%

 A5Y 2/2 = 100% 2/4 = 50% 4/6 = 66.7%

 6Y 0/2 = 0% 0/2 = 0%

AHY 9/14 = 64.3% 8/19 = 42.1% 1/7 = 14.3% 1/1 = 100% 19/41 = 46.3%

U age 2/5 = 40% 2/5 = 40%

All ages 23/68 = 33.8% 45/88 = 51.1% 24/74 = 32.4% 23/120 = 19.2% 115/350 = 32.9%

* All males searched for, including those banded prior to previous season. Numbers listed are those found divided

by those searched for. “Age Class” refers to age at first banding or most recent observation. “All ASY” is a total of

all banded males, excluding SY, AHY, and those of unknown age at banding. “U age” refers to unknown age at

banding.
20 USACERL TR 98/52

n = 26

n = 12

n = 27 n = 7

n = 93

n =62

n = 42

n = 28

0

10

20

30

40

50

60

1993 1994 1995 1996

Year resighted

Percent resighted

SY

all ASY

Figure 6. Return rates for non-ISA adult males, by age class and year.

Adult female Golden-cheeked Warblers were resighted at 18.3% (n = 60 banded

between 1991 and 1995; Table 5 for dispersal distances; Appendix D). Of the

two females banded in 1991, one was resighted; the one female banded in 1992

was not resighted nor were any of the four banded in 1993. Three (15.8%) of the

19 females banded in 1994 were resighted, and 20.6% (n = 34) of those banded in

1995 were resighted. One female was resighted twice (Appendix D).

HY resighting rates were generally much lower than observed adult returns.

HY males were resighted at an average of 16.9% (n = 148, Table 6 and Appendix

E), ranging from 11.1% (n = 27) in 1993, to two of two HYs returning in 1992.

HY females were resighted at an average of 9% (n = 78, Table 6 and Appendix

F), with rates ranging from no returns in 1993 (n = 8) and 1994 (n = 6), to 11.8%

(n = 34) in 1996.

Table 5. Dispersal distances for all female Golden-cheeked Warblers banded as adults.*

Age at Number located at each distance

resight n

Mean

(m; S.D.)

Median

(m)

Range

(m) > 300 m > 1 km > 3 km

 3Y 1 316 316 316 1 (100%) 0 0

A3Y 4 135 (47) 141 100 - 200 0 0 0

A4Y 2 883 (172) 883 762 - 1005 2 (100%) 1 (50%) 0

ASY 4 231 (95) 253 100 - 316 1 (25%) 0 0

Total 11 322 (294) 224 100 - 1005 4 (36%) 1 (9%) 0

* (%) = Percent of that age class resighted at or beyond that distance from either their banding location

or observation in the previous year.
USACERL TR-98/52 21

Table 6. Return rates for all hatch year (HY) Golden-cheeked Warblers.*

Year banded Male Female

1991 2/2 = 100% 0/0

1992 3/27 = 11.1% 0/8

1993 3/16 = 18.8% 0/6

1994 10/47 = 21.3% 3/30 = 10%

1995 7/56 = 12.5% 4/34 = 11.8%

All years 25/148 = 16.9% 7/78 = 9.0%

* Numbers indicate those found divided by total number banded. Intensive study area-banded HY birds

are included in this list, regardless of where they were resighted. Unknown-sex HY birds are not

included in this list.

Dispersal Distances

Observed dispersal distances for all male Golden-cheeked Warblers banded as

adults (including the second resighting of four HY males that were resighted

twice (i.e., from 3Y to 4Y) ranged from 0 m to 3,523 m, with an average distance

of 223 m (n = 268, median = 141, S.D. = 307, Table 7). While the dispersal

distance differences for the 3Y, 4Y, and A4Y+ (A4Y, 5Y, A5Y, 6Y, A6Y) age classes

were not significant (Kruskal-Wallis test 2

 = 3.93, 1 df, P = 0.14), there was a

trend toward 3Y males being resighted farther (n = 80, average = 268 m, median

= 141, S.D. = 282) from their original banding locations than the 4Y males (n =

20, average = 162 m, median = 100, S.D. = 157). Although A4Y+ males were

resighted at an average distance of 262 m (n = 43, median = 121, S.D. = 552),

this may have been due to three males that were found at much greater

distances than the others (Appendix A). Excluding these three males, the

average resighting distance for A4Y+ males was 152 m (n = 40, median = 100,

S.D. = 120).

Dispersal distances were classified into three groups: ISA resightings, searchedfor males, and those males that were “chance” finds (Table 8 and Appendix A).

These three groups showed a significant difference in dispersal distance that

was most likely due to the methods used to find them (Kruskal-Wallis test 2

 =

14.16, df = 2, P = 0.0008). “Chance” find males were observed at the greatest

average distance (304 m, median = 200 m, S.D. = 480, n = 66); ISA males were

resighted at an average distance of 277 m (median = 200 m, S.D. = 310, n = 65);

searched-for males were resighted at the shortest average distance of 146 m

(median = 100 m, S.D. = 141, n = 137).
22 USACERL TR 98/52

Table 7. Dispersal distances summarized by age class for male Golden-cheeked Warblers

banded as adults on Fort Hood.*

Age at Number located at each distance

resight n

Mean

(m; S.D.)

Median

(m)

Range

(m) > 300 m > 1 km > 3 km

ASY 37 193 (191) 141 0 - 1000 9 (24%) 1 (3%) 0

3Y 80 262 (282) 141 0 - 1746 29 (36%) 2 (3%) 0

A3Y 86 167 (209) 100 0 - 1456 15 (17%) 1 (1%) 0

4Y 20 162 (157) 100 0 - 510 4 (20%) 0 0

A4Y+ 43 268 (552) 121 0 - 3523 8 (19%) 2 (5%) 1 (2%)

 A4Y 23 353 (734) 141 0 - 3523 5 (22%) 2 (9%) 1 (4%)

 5Y 5 125 (55) 100 100 - 224 0 0 0

 A5Y 9 126 (116) 100 0 - 316 2 (22%) 0 0

 6Y 1 0 0 0 0 0 0

 A6Y 5 277 (147) 224 200 - 539 1 (20%) 0 0

AHY 2 193 (191) 549 0 - 1000 2 (100%) 0 0

Overall 268 223 (307) 141 0 - 3523 67 (25%) 6 (2%) 1 (0.4%)

* Known dispersal distances only. This table also includes four HY “second” resightings incorporated into their

respective resight-age categories – i.e., they were resighted as SY, then as 3Y (or later). This table excludes 1996

fire-displaced males. (%) = % of that age class resighted at, or beyond that distance from a previous resighting

location, or from their original banding location (whichever occurred later).

Table 8. Dispersal distances for adult males banded in the intensive study area (ISA), banded

males searched for, and banded males observed opportunistically (“chance-find”).*

Resight Number located at each distance

type n

Mean

(m; S.D.)

Median

(m)

Range

(m) # > 300 m # > 1 km # > 3 km

ISA 65 277 (310) 200 0 – 1746 24 3 0

Chance 66 304 (480) 200 0 - 3523 22 3 1

Search 137 146 (141) 100 0 - 781 21 0 0

* This table excludes fire-displaced males and HY first resights. It includes only males with known dispersal

distances.

The average resighting distances for male Golden-cheeked Warblers banded as

HY (4,040 m, median = 3317, S.D. = 4,777, n = 25) was significantly greater

than for adult males (223 m, S.D. = 308, n = 284, Kruskal-Wallis test 2

 = 53.68,

df = 1, P = 0.000; Tables 7 and 9, and Appendix E). Nineteen (76%) of the

resighted HY males were observed 1 kilometer or more from their original

banding locations (n = 25) compared with only 2% of adult males (n = 268, Table

7). This difference in proportion dispersing farther than 1 km was significant (2

= 155.60, df = 1, P = 0.000).

The average resighting distance for adult females was 322 m (median = 224 m,

S.D. = 294, n = 11; Table 5 and Appendix D). The range of these resighted
USACERL TR-98/52 23

females was 100 m to 1,005 m. The average resighting distance for female

Golden-cheeked Warblers that were banded as HY was 1,525 m (n = 7, median =

856, S.D. = 1,388, range of 0 m to 3,448 m; Table 10 and Appendix F) from their

original banding locations. This difference in resighting distances between adult

and HY females was significant (Kruskal-Wallis test 2 = 5.62, df = 1, P = 0.018).

Detection of Territorial Males

Density within the ISA steadily increased from 1992 through 1995, peaking at

28.1 males per 100 ha in 1995 (50 territorial males within 178 ha, Table 11),

then dropping to 18.0 males per 100 ha in 1996 (32 territorial males). The

annual average between 1992-1996 was 18.8 males per 100 ha (S.D. = 6.91).

Numbers of territorial males detected postwide (including the ISA) also

fluctuated between 1991 and 1996, averaging 584 males (S.D. = 219.79, Table

12). The highest number detected was 915 males, in 1996; the lowest was 383

males, in 1992. The number of training areas in which males were detected also

fluctuated somewhat, with an average of 37.7 (S.D. = 4.46), and a range of 31

(1994) to 41 (1991 and 1996).

Table 9. Dispersal distances for all male Golden-cheeked Warblers banded as HY on Fort Hood.*

Age at 1st Number located at each distance

resight n

Mean

(m; S.D.)

Median

(m)

Range

(m) # > 300 m # > 1 km # > 3 km

SY 17 2461 1720 60 - 10004 16 (95%) 12 (71%) 7 (41%)

3Y 5 7864 (7982) 5906 906 - 20082 6 (100%) 5 (83%) 4 (67%)

4Y 1 5573 5573 1 (100%) 1 (100%) 1 (100%)

5Y 1 5557 5557 1 (100%) 1 (100%) 1 (100%)

Total 24 4040 (4777) 3317 60 - 20082 24 (96%) 19 (76%) 13 (52%)

* This table includes only initial resights. The 3 cases in which a male was originally banded as HY and was

resighted >1 time are listed in the table of adult male dispersal distances (Table 7). (%) = % of that age class

resighted at, or beyond that distance from their original banding location.

Table 10. Dispersal distances for female Golden-cheeked Warblers banded as hatch year (HY).*

Age at 1st

resight n

Mean

(m; S.D.)

Median

(m)

Range

(m)

Number located at each distance

# > 300 m # > 1 km # > 3 km

SY 6 1785 (1379) 906 0 - 3448 5 (83%) 2 (33%) 2 (33%)

3Y 1 224 0 0 0

Total 7 1525 (1388) 856 0 - 3448 5 (71%) 2 (29%) 2 (29%)

* (n) = % of that age class resighted at, or beyond that distance away from either a previous resighting location, or

their original banding location (whichever occurred later).
24 USACERL TR 98/52

Table 11. Number of territorial Golden-cheeked Warblers present in the ISA.

Year Number of territorial males (in 178 ha) Density per 100 ha

1992 24 13.5

1993 25 14.1

1994 36 20.2

1995 50 28.1

1996 32 18.0

Average 33.4 18.8

Table 12. Number of male Golden-cheeked Warblers detected in all training areas (including the

ISA), and the total number of training areas in which males were detected.

Year Number of males

Number of training areas in which males

were detected

1991 515 41

1992 383 40

1993 399 33

1994 499 31

1995 797 40

1996 915 41

Average 584.7 37.7

Age Structures

The percentage of SY males within the ISA averaged 41.7%, ranging from 30.3%

to 57.1%. The differences did not vary significantly among years (2

 = 4.27, df =

4, P = 0.37; Table 13). Observed percentage of SY males was consistently and

significantly higher within the ISA than in non-ISA training areas (ISA average

= 41.7%, non-ISA average = 27.5%, 2

 = 12.40, df = 1, P = 0.0004; Tables 13 and

14; Figure 7). Although rates in the non-ISA training areas did not vary as

widely among years as those within the ISA, the differences were statistically

significant (range: 18.5% to 34.7%, average of 27.5%, 2

 = 15.97, df = 5, P =

0.007). When the annual captures of the ISA and non-ISA training areas were

lumped together, SY males were captured at a significantly lower rate than ASY

males (2 = 13.54, df = 5, P = 0.019; Table 15).
USACERL TR-98/52 25

Table 13. Age structure of male Golden-cheeked Warblers present in ISA.*

Age class 1992 1993 1994 1995 1996 All years

SY (%SY) 6 (35.3%) 12 (57.1) 17 (44.7) 25 (42.4) 10 (30.3) 70 (41.7)

ASY 9 2 5 13 7 36

3Y 2 9 8 7 26

A3Y 2 5 1 3 4 15

4Y 1 6 2 9

A4Y 5 1 6

5Y 1 2 3

A5Y 2 2

6Y 1 1

AHY 4 1 1 1 7

HY 8 4 10 32 11 59

Total 17 21 38 59 33 168

* Includes all males identified within the intensive study area, regardless of territory status. Neither AHY nor HY are

included in % SY calculation.

Table 14. Age structure of male Golden-cheeked Warblers observed outside the ISA.*

Age class 1991 1992 1993 1994 1995 1996 All years

SY (%SY) 23 (33.3%) 23 (31.1) 33 (34.7) 32 (21.9) 72 (33.0) 29 (18.5) 212 (27.5)

ASY 46 34 48 64 97 69 358

3Y 5 5 17 10 22 59

A3Y 12 4 22 16 20 85

4Y 2 3 8 3 16

A4Y 3 6 10 3 22

5Y 1 2 3

A5Y 1 3 5 9

A6Y 2 3 5

7Y 1 1

AHY 15 24 43 20 13 13 128

HY 2 19 12 37 23 11 104

Total 69 74 95 146 218 157 770

* Includes both resighted and newly banded males. Neither AHY nor HY are included in % SY calculation. ISA

males are not included in this table. See Table 13 for age classes of ISA males.
26 USACERL TR 98/52

n = 33

n = 59

n = 38

n = 21

n = 17

n = 157

n = 218

n = 146

n = 95

n = 74

n = 69

0

10

20

30

40

50

60

1991 1992 1993 1994 1995 1996

Year

Percent SY

ISA

non-ISA

Figure 7. Percentage of SY individuals in the ISA and non-ISA training areas.

Table 15. Total number of males banded by age class on Fort Hood.*

Age class 1991 1992 1993 1994 1995 1996 Total

SY (%SY) 23 (33.3%) 29 (43.3) 44 (53.0) 45 (44.6) 91 (46.7) 32 (30.8) 264 (42.6)

ASY 46 38 39 56 104 72 355

AHY 15 28 42 22 13 14 134

HY 2 27 16 47 56 22 170

U 0 60 0 0 0 6

Total 86 128 141 170 264 140 929

* This table includes males captured within the ISA and in non-ISA training areas. “U” = unknown age at banding.

Mated Status Within the Intensive Study Area

Males within the intensive study area mated at an overall frequency of 89% (n =

167, Table 16, Figure 8). SY males mated consistently and significantly less

often, at 79% (n = 61, 1992-1996) than ASY males, at 97% (n = 88, 2

 = 14.20, df

= 1, P = 0.0005). SY mating percentage ranged from 70% in 1993 to 85% in 1995.

ASY mating percentage ranged from 82% in 1992 to 100% in 1993, 1995, and

1996. There was no significant heterogeneity among years, allowing the years to

be lumped (2

 = 2.75, df = 4, P = 0.60).
USACERL TR-98/52 27

n = 7 n = 10

n = 14

n = 20

n = 11 n = 10

n = 12

n = 16

n = 30 n = 19

0

10

20

30

40

50

60

70

80

90

100

1992 1993 1994 1995 1996

Year

Percent mated

SY

ASY

Figure 8. Percent of mated males within the ISA, by age class and year.

Table 16. Percent of ISA males mated, by age class.*

Age class 1992 1993 1994 1995 1996 All years

SY 71% (7) 70 (10) 79 (14) 85 (20) 80 (10) 79 (61)

All ASY 82 (11) 100 (12) 94 (16) 100 (30) 100 (19) 97 (88)

 ASY 78 (9) 100 (2) 100 (3) 100 (12) 100 (4) 93 (30)

 3Y 100 (3) 100 (8) 100 (8) 100 (7) 100 (26)

 A3Y 100 (2) 100 (7) 100 (1) 100 (2) 100 (16)

 4Y 100 (1) 100 (5) 100 (1) 100 (7)

 A4Y 67 (3) 100 (1) 75 (4)

 5Y 100 (1) 100 (2) 100 (3)

 A5Y 100 (1) 100 (1)

 6Y 100 (1) 100 (1)

AHY 100 (2) 100 (1) 100 (3)

UB 75 (4) 100 (3) 67 (6) 100 (2) 80 (15)

All ages 79 (24) 88 (25) 83 (36) 94 (50) 94 (32) 89 (167)

* (n) = total number of territorial males in that age class. “All ASY” is a total of all banded males, excluding SY and

AHY. “UB” = unbanded male.

Productivity

Fecundity within the ISA averaged 2.23 HY per successful male (n = 118

successful males including unknown-age and unbanded males, 263 fledglings;

excludes 1992), and ranged between 2.08 (1996, n = 25 successful males) to 2.29

(1995, n = 45 successful males; Table 17). Numbers of HY per territorial male
28 USACERL TR 98/52

averaged 1.84 HY per territorial male (n = 143 territorial males), ranging from

1.63 (1996, n = 32 territorial males) to 2.06 (1995, n = 50 territorial males, Table

18).

Table 17. Mean number of young fledged by successful ISA males, by age class and year.*

Age class 1992 1993 1994 1995 1996 All years

SY 2.83 (17) 2.00(20) 2.06 (16) 2.17 (13) 2.18 (83)

All ASY 2.00 (24) 2.36 (33) 2.41 (29) 2.12 (36) 2.26 (163)

 ASY 1.50 (3) 2.33(7) 2.33 (12) 1.75 (7) 2.14 (45)

 3Y 2.67 (8) 2.29 (16) 2.63 (8) 2.00 (10) 2.39 (55)

 A3Y 1.86 (13) 3.00 (3) 1.00 (1) 2.00 (8) 1.92 (25)

 4Y 3.00 (3) 2.20 (11) 2.00 (2) 2.29 (16)

 A4Y 2.00 (4) 4.00 (4) 2.67 (8)

 5Y 2.00 (2) 3.00 (6) 2.67 (8)

 A5Y 3.00 (3) 3.00 (3)

 6Y 3.00 (3) 3.00 (3)

AHY 2.00 (2) 2.00 (2)

UB 2.00 (4) 2.50 (10) 1.00 (1) 2.14 (15)

All ages 1.69 (27) 2.25 (45) 2.25 (63) 2.29 (103) 2.08 (52) 2.23 (263)

*(n) = total number of young fledged. “All years” column excludes 1992 data. “All ASY” is a total of all banded

males, excluding SY and AHY. “UB” = unbanded males.

Table 18. Mean number of young fledged by territorial ISA males, by age class and year.*

Age class 1992 1993 1994 1995 1996 All years

SY 1.70 (17) 1.43 (20) 1.65 (33) 1.30(13) 1.54 (83)

All ASY 2.00 (12) 2.06 (33) 2.33 (70) 1.89(36) 2.12 (163)

 ASY 1.50 (3) 2.33 (7) 2.33 (28) 1.75 (7) 2.14 (45)

 3Y 2.37 (8) 2.00 (16) 2.63 (21) 1.43 (10) 2.12 (55)

 A3Y 1.86 (13) 3.00 (3) 0.50 (1) 2.00 (8) 1.79 (25)

 4Y 3.00 (3) 2.20 (11) 2.00 (2) 2.29 (16)

 A4Y 1.33 (4) 4.00 (4) 2.00 (8)

 5Y 2.00 (2) 3.00 (6) 2.67 (8)

 A5Y 3.00 (3) 3.00 (3)

 6Y 3.00 (3) 3.00 (3)

AHY 2.00 (2) 2.00 (2)

UB 1.33 (4) 1.67 (10) 1.00(1) 1.36 (15)

All ages 1.13 (27) 1.80 (45) 1.75 (63) 2.06 (103) 1.63 (52) 1.84 (263)

* (n) = total number of young fledged. “All years” column excludes 1992 data. “All ASY” is a total of all banded

males, excluding SY and AHY.
USACERL TR-98/52 29

Productivity of mated males did not differ significantly between age classes.

The average fecundity for mated SY males was 1.93 young per mated SY (n =

43 mated SY; Table 19). The average for mated ASY males was 2.15 young per

mated ASY (n = 76 mated ASY excluding 1992, t = 0.77, df = 1, P = 0.50; Table

19). Productivity per successful male was also very similar between SY (2.18

HY per successful SY, n = 38 successful SY excluding 1992) and ASY males (2.26

HY per successful ASY, n = 72 successful ASY excluding 1992, t = 0.152, df = 1,

P = 0.89; Tables 17 and 20). The difference between age classes in productivity

per territorial male was statistically significant, however, with an average of

1.54 HY per territorial SY (n = 54 territorial SY excluding 1992; Tables 16 and

18) and 2.12 young per territorial ASY (n = 77 territorial ASY excluding 1992, t

= 6.45, df = 1, P = 0.008, Tables 16 and 18), due to the lower mating success of

SY males versus ASY males. Once mated, however, SY males appeared to be

equally successful as ASY males (see success rate information below). See

Figure 9 for SY productivity by year.

Although sample sizes were small, there was a slight trend toward males

producing more young as they aged. Third-year mated males’ productivity was

the same as all ASY mated males, with 2.12 HY per mated male (n = 26 3Y

males, Table 16). Fourth-year males produced 2.29 HY per mated male (n = 7

4Y males, Table 16), fifth-year males, 2.67 (n = 3 5Y males; Table 16), and the 1

sixth-year male present in the intensive study area produced 3 HY.

Figure 9. Productivity for ISA SY males, by year.

0

0.5

1

1.5

2

2.5

3

1993 1994 1995 1996

Year

Number HY per SY male

HYper successful SY

HY per mated SY

HY per territorial SY
30 USACERL TR 98/52

Table 19. Mean number of young fledged by mated ISA males, by age class and year.*

Age class 1992 1993 1994 1995 1996 All years

SY 2.43 (17) 1.82 (20) 1.94 (33) 1.63 (13) 1.93 (83)

All ASY 2.00 (24) 2.20 (33) 2.33 (70) 1.89 (36) 2.15 (163)

 ASY 1.50 (3) 2.33 (7) 2.33 (28) 1.75 (7) 2.14 (45)

 3Y 2.67 (8) 2.00 (16) 2.63 (21) 1.43 (10) 2.12 (55)

 A3Y 1.86 (13) 3.00 (3) 0.50 (2) 2.00 (8) 1.79 (25)

 4Y 3.00 (3) 2.20 (11) 2.00 (2) 2.29 (16)

 A4Y 2.00 (2) 4.00 (4) 2.67 (8)

 5Y 2.00 (2) 3.00 (6) 2.67 (8)

 A5Y 3.00 (3) 3.00 (3)

 6Y 3.00 (3) 3.00 (3)

AHY 2.00 (2) 2.00 (2)

UB 1.33 (4) 2.50 (10) 1.00 (1) 1.67 (15)

All ages 1.42 (27) 2.05 (45) 2.10 (63) 2.19 (103) 1.73 (52) 2.04 (263)

*(n) = total number of young fledged. “All years” column excludes data for 1992. “All ASY” is a total of all banded

males, excluding SY and AHY.

Table 20. Success rate of ISA Golden-cheeked Warbler males.*

Age class 1992 1993 1994 1995 1996 All years

SY 5/5 6/7 10/11 16/17 6/8 43/48

All ASY 8/9 12/12 14/15 29/30 17/19 80/85

AHY 0/2 0/0 0/0 0/0 1/1 1/3

UB 3/3 2/3 4/4 0/0 1/2 10/12

All ages 16/19 20/22 28/32 45/47 25/30 133/145

*Proportion of mated males that were also successful. “All years” column includes 1992 data.

Success rate (mated males that also reproduced) between the age classes was

very consistent. SY males had an overall success rate of 90% (n = 48 successful

SY males [including 1992 data], ranging from 75% in 1996 to 100% in 1992;

Table 20). ASY males had an overall success rate of 94% (n = 85 successful ASY

males [including 1992 data], ranging from 89% in 1992 and 1996 to 100% in

1993). This difference between SY and ASY male mating success was not

significant (χ2

 = 0.37, df = 1, P = 0.54).

Parasitism

Seven incidents of brown-headed cowbird (BHCO) parasitism of Golden-cheeked

Warbler nests were reported since 1991. In 1991, three of six Golden-cheeked

Warbler nests located were parasitized (at least two of which fledged cowbird
USACERL TR-98/52 31

young, Table 21), and an adult male was seen feeding a juvenile BHCO in a

cowbird trap. The one nest located in 1992 was not parasitized; in 1993, none of

the four nests whose contents were observed were parasitized. There was one

instance in 1993, however, of a pair of Golden-cheeked Warblers feeding one

BHCO young, as well as at least one (and possibly two) Golden-cheeked Warbler

young. In 1994, none of the ten nests were parasitized, and no feeding of BHCO

fledglings was observed. Similarly, in 1995, none of the seven nests found were

parasitized, and no feeding of BHCO fledglings was observed. In 1996, none of

the 11 nests found were parasitized, but there were two occasions during which

an adult Golden-cheeked Warbler was found in association with a juvenile

BHCO. In one instance, an adult female was seen within inches of the BHCO

HY, but did not feed it even though it was begging. In another instance, an

adult female was seen feeding a juvenile BHCO.

Table 21. Summary of outcomes for Golden-cheeked Warbler nests located on Fort Hood, 1991-

1996.

Year No. found

No. fledged

successfully

No. depredated

or abandoned

No.

parasitized

No. fate

unknown

1991 6 3 0 3 3

1992 1 1 0 0 0

1993 7 4 2 0 0

1994 10 10 0 0 0

1995 7 4 1 0 2

1996 11 5 2 0 4

Total 35 27 5 3 9
32 USACERL TR 98/52

4 Discussion

Population Trends

Results indicate that the Golden-cheeked Warbler population on Fort Hood,

Texas steadily increased through 1995, when most population measurements

were at their peak. Return rates were at their highest within the intensive

study area in 1995, and were average for non-ISA training areas. Territorial

males were at their greatest density within the ISA, and above average for all

training areas combined. The proportion of the population that was SY was

above average for both inside and outside the study area, indicating a young,

growing population. All measurements of productivity within the study area

were at their highest since the study began, also indicating a growing

population. However, 1996 showed decreases in many population indices.

Return rates and proportions of SY in the population were at their lowest for all

males, both within and outside the study area. Similarly, all measurements of

productivity, including success rate, were also at their lowest since 1992.

Because no baseline warbler demographic data was available before 1991, and

because of the steady increase between 1992 and 1995, followed by the sharp

decrease in 1996, no long term population trends can be discerned at this time.

Data collected in the years 1991 through 1996 on Fort Hood demonstrate a

capacity for high annual variability. While there were several possible reasons

for the observed decreases in population parameters in 1996, there was no

apparent cause for the increases between 1991 and 1995.

One possible explanation for the decreases of 1996 may be the stress caused by

severe weather in the spring and summer of 1996. According to the Palmer

Drought Severity Index, there was a “severe to extreme drought” in the spring

and summer of 1996 (Figure 10, Appendix G). When these drought conditions

were combined with the slightly cooler than average temperatures and

significantly lower than average relative humidity that were prevalent when the

Golden-cheeked Warblers were migrating north, and when they returned to Fort

Hood in March, a higher death rate might be expected due to stress caused by

food and water shortages (see Table 22 for weather data). This hypothesis,

though, is not supported, as the return rate in a Texas Parks and Wildlife

Department (TPWD) study less than 100 miles south of Fort Hood did not show

a similar decrease, although they generally had similar weather patterns
USACERL TR-98/52 33

PSDI Classifications for Dry and Wet Periods

4.00 or more Extremely wet

3.00 to 3.99 Very wet

2.00 to 2.99 Moderately wet

1.00 to 1.99 Slightly wet

0.50 to 0.99 Incipient wet

0.49 to –0.49 Near normal

-0.50 to –0.99 Incipient dry spell

-1.00 to –1.99 Mild drought

-2.00 to –2.99 Moderate drought

-3.00 to –3.99 Severe drought

-4.00 to –4.99 Extreme drought

Figure 10. Monthly averages of the Palmer Drought Severity Index (PDSI) for Texas,

Division 7 (of which Fort Hood is a part), from 1895 to 1996. For a detailed description

of this index, see Appendix G.
34 USACERL TR 98/52

Table 22. Mean daily temperatures and relative humidities for the month of March for

Fort Hood from 1994-1996.*

1994 1995 1996 Overall

Mean March daily

temperature

68.7° 64.0° 63.4° 65.4°

Temperature range 40.5° - 86.2°

S.D. = 13.43

33.1° - 89.4°

S.D. = 14.64

41.5° - 83.5°

S.D. = 14.37

33.1° - 89.4°

Mean March daily

Relative Humiditya

48.7% 58.5% 38.2% 48.5%

Relative Humidity

range

21% - 99%

S.D. = 21.5

22% - 91%

S.D. = 20.0

16% - 99%

S.D. = 19.7

16% - 99%

S.D. = 21.85 a

 = significant difference by ANOVA (F(2,92) = 7.714; P = 0.001). Data compliments of Natural Resources Branch,

Fort Hood (unpublished data).

(D. Keddy-Hector, Adjunct Professor, Austin Community College, Professional

Communication).

Another hypothesis concerns the stress and damage caused by the extensive fire

that destroyed nearly 4,300 ha of Golden-cheeked Warbler habitat

(approximately 25% of the warbler habitat on Fort Hood; Figures 1 and 2) in the

northeastern section of Fort Hood in February, 1996. Upon their return, many

birds would have found that their territories were either entirely or partially

burned. Locating and establishing new territories may have taken valuable

time and resources (i.e., fat stores) that the birds may not have had after their

migration. Some of the more dominant, newly displaced males that had

previously held territories within the area of the fire may have displaced less

dominant males that had held territories within nearby areas. All of these

sudden movements and displacements may have increased territory border

fights, and the need for males to increase “patrols” of their territory borders.

This, in turn, may have decreased foraging time and efforts to attract and

defend a mate and or care of nestlings and fledglings. Males that maintained

partially burned territories may have experienced a shortage of food. Any of

these scenarios could cause an increase in mortality.

This hypothesis, however, does not explain population dynamics observed in

other areas across Fort Hood. For example, there was no evidence that firedisplaced warblers affected demographic parameters in the ISA (a dispersal

distance of 3 to 4 kilometers). The percentage of SY males did not decrease

significantly within the study area due to the influx of older, more dominant

fire-displaced males (assuming that SYs were less dominant than older males).

While the percentage did decrease slightly, it was not significant (Table 13).

Secondly, some of the males within the fire region were color-banded in years

prior to the fire; if a significant number of these males moved south into the

study area and nearby regions, their chance of detection was high, and only one
USACERL TR-98/52 35

of these males was found there. (This male was found in habitat adjacent to the

study area. He was originally banded as an HY, however, so his movement may

be accounted for equally as well by HY dispersal patterns as by fire-related

dispersal). Thirdly, if fire-displaced males in turn displaced study area males, it

is assumed that these newly displaced males from within the study area would

also establish a territory in the nearest suitable habitat. Researchers did not

find this to be true, however, as all adjacent habitat was searched, and no males

previously territorial within the study area were found.

Productivity within the intensive study area remained relatively stable between

1992 and 1996, suggesting that fecundity may not have a strong effect on the

adult population of the immediate area, which varied during the same time

period. One possible explanation for this lack of strong effect may be caused by

the biology of the species, particularly the dispersal of the young away from

their natal territories. One possible hypothesis for their dispersal pattern is

inbreeding avoidance (Greenwood 1984). It may be inherent in many birds to

disperse away from their natal territories in order to not breed with close kin

(i.e., parents or siblings). Juveniles disperse much farther than do adults.

Therefore, local productivity likely does not affect adult populations of the natal

area in succeeding years. Judging by the long dispersal distances of juveniles

and high estimates of juvenile emigration (see Relevance/Application to

Predictive Models), there may be a significant proportion of Fort Hoodproduced young that may not return to Fort Hood.

The significant differences in adult return rate between the ISA and other

training areas can possibly be attributed to search effort and detection rate. The

monitoring of the ISA was extensive, with near-daily coverage; the detection rate

of males within this area probably approached 100%. Outside this area,

however, most males were searched for only once or twice. Many things may

have occurred during these search times to alter the probability of detection,

including circumstances of their breeding cycle, time of day, presence of other

singing males, females or HYs, weather effects, lighting, and canopy cover. It

may not be uncommon to search for a specific male and consider him “not found”

even though he may have been present and neither singing nor responding to

the playback tape. Due to the intensive monitoring effort within the ISA, many

of these factors were canceled, yielding the most reliable data.

While the cause of the decreased return rate within the intensive study area

was unknown, there was a possible explanation for the decrease in other

training areas. There was a slight change in searching protocol in 1996. Rather

than searching for males picked from all previously banded males (as had been

done in 1992 through 1995), it was decided that field technicians would search
36 USACERL TR 98/52

for pre-chosen males, which were, in general, older than those that would have

been chosen randomly (see Methods section). In general, passerine survival

rates level off after the first year, and remain essentially constant, but the

probability of surviving to a particular age decreases as the bird ages, due to

annual mortality rates (Gill 1990). This leads to a small sample size of older

birds, in turn decreasing the possibility of locating survivors.

There are two possible reasons for the fact that female band returns were

generally lower than that of males. First, adult females may disperse to a

greater extent than males (Tables 5 and 7; Gill 1990). Secondly, females are

generally very quiet and secretive in their behaviors, making it difficult to

observe and determine their identity.

Age-related Differences 

While return rates within the ISA did not differ between age classes when years

were lumped, those in other training areas did (Tables 3 and 4). This may be

partially explained by the differences in detection rates between the areas.

Field observations suggest that older males may sing less than younger males,

resulting in a lower detection rate for older males outside the ISA; older males

within the ISA had the same probability of detection as younger males due to

the intense monitoring regimen. Older males returning to non-ISA training

areas, therefore, may not have been detected by researchers.

Although not statistically significant, younger males (age 3Y) had the tendency

to disperse farther than older males (ages 4Y and A4Y+, Table 3). This idea also

had support from data gathered concerning territory boundaries of returning

males within the ISA. Three-year-old males seldom defended the same territory

as they did during their SY year (42% [10 of 24] of 3Y territories overlapped

their SY territory). Most of these SY-3Y territory connections were tenuous

though, as only 20% (n = 10) of the overlapping territories actually overlapped at

the center (see Figures 11 through 15). Returning ASY males, however,

generally defended the same territory as their previous year (85% [22 of 26] of

their territories overlap their previous territory). The majority (68%, n = 22) of

these connections between years overlapped at their centers. This difference in

the frequency of territory overlap by age class was statistically significant (SY

vs. ASY, territory overlap: 2

 = 8.21, df = 1, P = 0.0041, SY vs. ASY, centers

overlap: 2

 = 4.62, df = 1, P = 0.03).

Observed results indicate that SY males had the capacity to reproduce at the

same rate as ASY males. Productivity per successful SY was very similar to
USACERL TR-98/52 37

productivity per successful ASY male (Table 17). Similarly, success rate of

mated SY and ASY males was virtually identical (Table 20). The difference,

then, in SY fitness can be attributed to difficulty in finding, securing or

defending a mate. SY males mated significantly less often than did ASY males

(Table 16).
38 USACERL TR 98/52

Figure 11. Map of all territorial males within the ISA during 1996. For detailed information, see

Appendix C. “UB” = unbanded. “UK” = unknown individual. (1 cm ~ 100 m.)
USACERL TR-98/52 39

Figure 12. Map of all territorial males within the ISA during 1995. For detailed information, see

Appendix C. “UB” = unbanded. “UK” = unknown individual. (1 cm ~ 100 m.)
40 USACERL TR 98/52

Figure 13. Map of all territorial males within the ISA during 1994. For detailed information, see

Appendix C. “UB” = unbanded. “UK” = unknown individual. (1 cm ~ 100 m.)
USACERL TR-98/52 41

Figure 14. Map of all territorial males within the ISA during 1993. For detailed information, see

Appendix C. “UB” = unbanded. “UK” = unknown individual. (1 cm ~ 100 m.)
42 USACERL TR 98/52

Figure 15. Map of all territorial males within the ISA during 1992. For detailed information, see

Appendix C. “UB” = unbanded. “UK” = unknown individual. (1 cm ~ 100 m.)
USACERL TR-98/52 43

Relevance/Application to Predictive Models

Results from a regional population viability workshop (U.S. Fish and Wildlife

Service 1996) estimated that HY Golden-cheeked Warbler survival rate must be

above 50%, given observed annual fecundities of 0.8 and 1.01 HY per adult male

(SY and ASY, respectively), and an estimated average adult survival rate of 57%

(AHY) to maintain a stable population. In population viability analysis (PVA)

models simulating time until extinction, the use of any HY survival rate below

50% caused the model’s population to decline dramatically and become extinct

well within the 100 years of simulation (U.S. Fish and Wildlife Service 1996).

The observed return of HYs to their natal area in the ISA was 14.2% (Table 6).

Assuming HY survival of 50% to maintain a stable population, the observed HY

return rate in the ISA would indicate that at least 76% of surviving HYs must

emigrate from their natal area. The greater observed dispersal distance of HYs

compared with adult Golden-cheeked Warblers suggests a 76% HY emigration

rate is realistically possible. The average dispersal distance for returning HY

males was over 4 kilometers, with 41% of all HY resightings at a distance

greater than 3 km, and one actually being resighted 20 km from its original

banding location. This compares with a mean dispersal distance for adult males

of 217 m.

Comparison With Similar Species

Although the overall intensive study area return rate for the Golden-cheeked

Warbler (48%) was lower than that for the Kirtland’s Warbler (approximately

65%, Mayfield 1992), their ranges were similar. The Kirtland’s Warbler return

rate ranged between 29% and 82% (Mayfield 1992) whereas the Golden-cheeked

Warbler return rate has ranged from 30% to 65.5% within the study area. The

Golden-cheeked Warbler’s observed return rate was also lower than that of the

Black-throated Green Warbler (67%, Morse 1989), the Prairie Warbler (65%,

Nolan 1978), the Common Yellowthroat (54%, Roberts 1971) the Black and

White Warbler (71%, Roberts 1971) and the American Redstart (71%, Roberts

1971).

The observed Kirtland’s Warbler HY return rate of 2.7% (n = 296 as of 1968;

Berger and Radabaugh 1968) was much lower than the Golden-cheeked

Warbler’s (17%, overall HY male return rate). The Golden-cheeked Warbler HY

return rate was also high compared to that documented in other species (i.e.,

5.8% in the Indigo Bunting [Payne and Payne 1990], 5.0% in the Wood Thrush
44 USACERL TR 98/52

[Roth and Johnson 1993], and 0.6% in the American Redstart [Sherry and

Holmes 1992]).

There are few published studies concerning survival rates of first-year birds.

Loery, et al. (1987) reported an average 44% (range of 24% to 66%) HY survival

rate for the Black-capped Chickadee (Parus atricapillus), a figure that was

similar to the estimates for the Golden-cheeked Warbler. This study also stated

that the adult chickadee survival rate was, on average, 23% higher than that for

HYs, with an overall average of 69% (and a range of 45% to 100%) for 3Y

individuals. Loery, et al. (1987) attributed this difference in survival to

increased mortality within the first year and/or to increased dispersal. Ryel

(1981) calculated the Kirtland’s Warbler adult survival rate to be at least 35%.

These figures were similar to the warbler’s estimated 57% survival for adult

males (USFWS 1996).

Golden-cheeked Warbler productivity (2.04 HY per mated male within the ISA)

was similar to that documented in several other species. Mayfield (1992)

reported the Kirtland’s Warbler productivity to be 2.2 fledglings per nest

(without Brown-headed Cowbird parasitism). The Prairie Warbler had an

estimated productivity of 2.2 fledglings per territory (Nolan 1978), slightly

higher than Fort Hood’s Golden-cheeked Warbler average of 1.84 HY per

territory. Black-throated Green Warbler productivity was also higher, with

three (and sometimes four) fledglings per territory (Morse 1993).

Mated status for the Golden-cheeked Warblers within the ISA (89% of the

territorial males mated) was comparable to the 92% of the Kirtland's Warbler

within prime, wildfire-regrown areas, and 72% of Kirtland's Warblers within

plantations (Bocetti 1994).

Average Golden-cheeked Warbler density (0.13 pairs per ha) was much lower

than that documented for the Black-throated Green Warbler. Morse (1993)

reported that the maximum density was 2.0 pairs per ha in prime habitat. He

also reported, however, a minimum of 0.6 pairs per ha in a year of high

mortality. The Kirtland’s Warbler had a density lower than either species,

ranging from 0.048 pairs per ha to 0.07 pairs (Probst and Weinrich 1989, Bocetti

1994). Contrary to highly variable densities found within Fort Hood’s ISA

between 1992 and 1996, Morse (1989) stated that “Populations of most warblers

are surprisingly constant in density from year to year, although they experience

temporary and short-term depressions, probably in response to environmental

factors....” Holmes et al. (1986) also found that periods of stability were

punctuated by pulses in response to outbreaks of caterpillars.
USACERL TR-98/52 45

Incidence of Cowbird Parasitism

Rates of parasitism of the Golden-cheeked Warbler on Fort Hood may be

misleading, as cowbird control began in 1988, whereas Golden-cheeked Warbler

studies did not begin until 1991; no pre-control, baseline data is available. Even

though Pulich (1976) reports a very high parasitism rate (58%, 19 of 33 nests

parasitized), data suggest that cowbirds currently do not play a major role in

Fort Hood warbler nest success. There were seven incidents concerning Goldencheeked Warblers and cowbirds on Fort Hood since 1991, including three

parasitized nests (no parasitized nests since 1991, 9% nest parasitism rate;

Table 21).
46 USACERL TR 98/52

5 Conclusions and Recommendations

Fort Hood supports a substantial population of the endangered Golden-cheeked

Warbler (915 maximum documented males). This population is vital to the

survival of the species because it resides under a single land management

authority. This continuity of management will assist in the preservation of the

remaining habitat and the species as a whole.

A substantial conclusion stemming from this data analysis is the high variability

of the Golden-cheeked Warbler population on Fort Hood. During the first 4

years of the study, it appeared that the population was growing, at first slowly,

then more rapidly. During 1996, however, this trend reversed, and the

population appeared to decline. The long-term trends for this species cannot yet

be determined.

Currently, habitat availability is considered the primary limiting factor for

warbler populations on Fort Hood. Other potential limiting factors (e.g.,

cowbird parasitism or predation) appear to be of secondary importance at this

time. If cowbird control on the installation is discontinued in the future, this

relationship will need to be reevaluated.

It is recommended that demographic parameters are best estimated from data

obtained in intensively monitored areas.

It is recommended that these data be collected and updated annually to monitor

long-term population changes on Fort Hood. It is also recommended that

managers be cautious in making decisions regarding this species based on shortterm demographic variability.
USACERL TR-98/52 47

References

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1994. 59 pp.

Benson, R.H. 1990. Habitat area requirements of the Golden-cheeked Warbler on the Edwards

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the endangered Black-capped Vireo and Golden-cheeked Warbler, and the cowbird control

program on Fort Hood, Texas. Submitted to HQ III Corps and Fort Hood. 78 pp. Copies

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50 USACERL TR 98/52

Appendix A: List of all Adult Males

Resighted on Fort Hood

This list is presented by resighting type (“Search type”: ISA, incidental, searched

for), in order by band number and the number of times each was resighted. For

explanation of search types, see Methods section. “Distance (m)” is the distance

between observations, either between banding and resighting, or between

consecutive resightings.

Banding Information Resighting Information

Band

number

Search

Type

No. times

resighted Year Age East North Year Age East North

Distance

(m)

1290-90819 ISA 1 92 AHY 342 485 93 ASY 344 488 361

1290-90819 ISA 2 92 AHY 344 488 94 A3Y 341 485 424

1290-90819 ISA 3 92 AHY 341 485 95 A4Y 342 486 141

1290-90833 ISA 1 93 SY 346 480 94 3Y 343 482 361

1290-90834 ISA 1 93 SY 346 481 94 3Y 344 479 283

1290-90834 ISA 2 93 SY 344 479 95 4Y 343 479 100

1290-90836 ISA 1 93 SY 346 482 94 3Y 345 481 141

1290-90836 ISA 2 93 SY 345 481 95 4Y 345 481 0

1290-90837 ISA 1 93 SY 342 474 94 3Y 343 474 100

1290-90838 ISA 1 93 SY 341 473 94 3Y 341 476 300

1290-90838 ISA 2 93 SY 341 476 95 4Y 342 477 141

1290-90838 ISA 3 93 SY 342 477 96 5Y 342 476 100

1290-90846 ISA 2 93 HY 314 487 95 3Y 315 488 141

1290-90847 ISA 1 93 SY 340 489 94 3Y 338 491 283

1290-90847 ISA 2 93 SY 338 491 95 4Y 338 491 0

1290-90848 ISA 1 93 SY 340 489 94 3Y 340 489 0

1710-25228 ISA 1 92 ASY 340 477 93 A3Y 346 481 721

1710-25242 ISA 1 92 ASY 344 483 93 A3Y 341 484 316

1710-25242 ISA 2 92 ASY 341 484 94 A4Y 343 483 224

1710-25245 ISA 1 92 ASY 339 476 93 A3Y 339 476 0

1710-25245 ISA 2 92 ASY 339 476 94 A4Y 340 478 224

1710-25245 ISA 3 92 ASY 340 478 95 A5Y 341 478 100

1710-25246 ISA 1 92 SY 342 476 93 3Y 341 478 224

1710-25246 ISA 2 92 SY 341 478 94 4Y 341 479 100

1710-25246 ISA 3 92 SY 341 479 95 5Y 341 480 100

1710-25246 ISA 4 92 SY 341 480 96 6Y 341 480 0

1710-25265 ISA 1 92 ASY 344 486 93 A3Y 340 484 447

1710-25265 ISA 2 92 ASY 340 484 94 A4Y 341 474 1005

1710-25265 ISA 3 92 ASY 341 474 95 A5Y 341 473 100
USACERL TR-98/52 51

Banding Information Resighting Information

Band

number

Search

Type

No. times

resighted Year Age East North Year Age East North

Distance

(m)

1710-25268 ISA 1 92 ASY 344 479 93 A3Y 344 481 200

1710-25380 ISA 1 92 SY 339 472 93 3Y 340 478 608

1710-25381 ISA 1 92 ASY 343 475 93 A3Y 343 475 0

1710-25382 ISA 1 92 ASY 343 478 93 A3Y 340 476 361

1710-25384 ISA 1 92 SY 343 475 93 3Y 342 476 141

1710-25464 ISA 1 93 SY 337 483 94 3Y 339 481 283

1710-25464 ISA 2 93 SY 339 481 95 4Y 340 478 316

1710-25464 ISA 3 93 SY 340 478 96 5Y 339 478 100

1880-64770 ISA 1 91 ASY 344 479 92 A3Y 344 479 0

1880-64771 ISA 1 91 ASY 341 478 92 A3Y 343 478 200

1960-03104 ISA 1 94 ASY 339 489 95 A3Y 340 489 100

1960-03108 ISA 1 94 SY 343 483 95 3Y 343 483 0

1960-03110 ISA 1 94 SY 337 484 95 3Y 341 483 412

1960-03111 ISA 1 94 SY 345 485 95 3Y 339 484 608

1960-03115 ISA 1 94 SY 333 482 95 3Y 342 476 1082

1960-03116 ISA 1 94 ASY 343 487 95 A3Y 341 487 200

1960-03201 ISA 1 94 SY 344 479 95 3Y 336 480 806

1960-03203 ISA 1 94 SY 341 472 95 3Y 348 488 1746

1960-03203 ISA 2 94 SY 348 488 96 4Y 348 488 0

1960-03206 ISA 1 94 SY 338 477 95 3Y 337 473 412

1960-03206 ISA 2 94 SY 337 473 96 4Y 337 473 0

1960-03207 ISA 1 94 SY 345 475 95 3Y 344 475 100

1960-03208 ISA 1 94 SY 342 469 95 3Y 340 478 922

1960-03209 ISA 1 94 ASY 337 478 95 A3Y 338 478 100

1960-03238 ISA 1 94 AHY 343 483 95 ASY 343 483 0

1960-03410 ISA 1 95 SY 346 483 96 3Y 345 480 316

1960-03533 ISA 1 95 ASY 343 469 96 A3Y 341 472 361

1960-03553 ISA 1 95 ASY 346 469 96 A3Y 343 473 500

1960-03634 ISA 2 94 HY 341 477 96 3Y 338 480 424

1960-03709 ISA 1 95 ASY 341 475 96 A3Y 341 475 0

1960-03798 ISA 1 95 SY 343 478 96 3Y 340 481 424

1960-03815 ISA 1 95 SY 346 484 96 3Y 345 486 224

1960-03816 ISA 1 95 SY 347 485 96 3Y 344 485 300

1960-03821 ISA 1 95 ASY 340 485 96 A3Y 340 485 0

1960-03833 ISA 1 95 SY 344 488 96 3Y 345 488 100

1960-03838 ISA 1 95 SY 344 488 96 3Y 342 487 224

1290-90816 Incidental 1 92 AHY 360 556 93 ASY 360 566 1000

1290-90821 Incidental 1 92 AHY 343 483 95 A4Y 308 487 3523

1290-90854 Incidental 2 93 HY 298 486 95 3Y 298 486 0

1480-34250 Incidental 2 91 SY UK UK 93 4Y 306 487 UK

1480-34258 Incidental 1 91 ASY UK UK 94 A5Y 303 486 UK

1710-25238 Incidental 1 92 ASY 208 583 96 A6Y 206 588 539

1710-25254 Incidental 1 92 ASY 362 533 94 A4Y 362 531 200

1710-25258 Incidental 2 92 SY 339 560 94 4Y 339 561 100

1710-25401 Incidental 1 93 ASY 74 390 94 A3Y 80 388 632

1710-25401 Incidental 2 93 ASY 80 388 95 A4Y 73 392 806

1710-25402 Incidental 1 93 ASY 74 391 94 A3Y 75 393 224

1710-25402 Incidental 3 93 ASY 75 389 96 A5Y 74 392 316

1710-25403 Incidental 1 93 SY 183 643 94 3Y 184 640 316
52 USACERL TR 98/52

Banding Information Resighting Information

Band

number

Search

Type

No. times

resighted Year Age East North Year Age East North

Distance

(m)

1710-25415 Incidental 2 93 SY 317 563 95 4Y 316 567 412

1710-25420 Incidental 1 93 SY 310 537 95 4Y 315 536 510

1710-25425 Incidental 2 93 SY 312 539 95 4Y 312 540 100

1710-25431 Incidental 1 93 ASY 374 554 95 A4Y 374 554 0

1710-25436 Incidental 2 93 ASY 388 435 95 A4Y 387 435 100

1710-25450 Incidental 1 93 ASY 185 677 94 A3Y 184 678 141

1710-25450 Incidental 2 93 ASY 184 678 95 A4Y 183 678 100

1710-25452 Incidental 2 93 SY 335 491 95 4Y 334 490 141

1710-25452 Incidental 3 93 SY 334 490 96 5Y 333 492 224

1710-25460 Incidental 2 93 ASY 306 486 95 A4Y 305 487 141

1710-25466 Incidental 1 93 ASY 328 485 95 A4Y 328 485 0

1710-25466 Incidental 2 93 ASY 328 485 96 A5Y 327 485 100

1810-28929 Incidental 2 91 SY UK UK 93 4Y 242 482 UK

1810-28985 Incidental 2 91 AHY UK UK 94 A4Y 122 650 UK

1810-28985 Incidental 3 91 AHY 122 650 95 A5Y 122 650 0

1930-72771 Incidental 2 92 AHY 307 487 94 A3Y 309 488 224

1930-72773 Incidental 2 92 AHY 313 506 94 A3Y 313 506 0

1960-03031 Incidental 1 93 AHY 313 505 94 ASY 314 503 224

1960-03033 Incidental 1 93 AHY 122 649 94 ASY 122 651 200

1960-03049 Incidental 2 93 AHY 346 442 95 A3Y 346 442 0

1960-03060 Incidental 1 93 AHY 362 533 94 ASY 361 534 141

1960-03103 Incidental 1 94 ASY 73 393 96 A4Y 73 394 100

1960-03107 Incidental 1 94 ASY 78 378 95 A3Y 79 376 224

1960-03120 Incidental 1 94 SY 348 466 95 3Y 348 465 100

1960-03123 Incidental 1 94 SY 390 430 95 3Y 391 433 316

1960-03136 Incidental 1 94 ASY 81 377 95 A3Y 78 379 361

1960-03136 Incidental 2 94 ASY 78 379 96 A4Y 80 377 283

1960-03149 Incidental 1 94 AHY 309 486 95 ASY 308 487 141

1960-03153 Incidental 1 94 SY 385 517 95 3Y 385 517 0

1960-03155 Incidental 1 94 ASY 79 388 95 A3Y 77 387 224

1960-03180 Incidental 1 94 ASY 164 601 95 A3Y 164 600 100

1960-03191 Incidental 1 94 ASY 78 379 95 A3Y 74 393 1456

1960-03407 Incidental 1 95 SY 349 468 96 3Y 354 463 707

1960-03413 Incidental 1 95 ASY 342 565 96 A3Y 338 560 640

1960-03421 Incidental 1 95 SY 168 689 96 3Y 167 689 100

1960-03431 Incidental 1 95 SY 92 389 96 3Y 89 389 300

1960-03441 Incidental 1 95 ASY 82 388 96 A3Y 83 388 100

1960-03442 Incidental 1 95 SY 295 455 96 3Y 293 456 224

1960-03484 Incidental 1 95 SY 314 496 96 3Y 320 501 781

1960-03491 Incidental 1 95 ASY 363 522 96 A3Y 364 522 100

1960-03499 Incidental 1 95 SY 319 505 96 3Y 314 506 510

1960-03545 Incidental 1 95 ASY 339 555 96 A3Y 339 556 100

1960-03557 Incidental 1 95 AHY 74 387 96 ASY 75 389 224

1960-03572 Incidental 1 95 ASY 314 503 96 A3Y 313 502 141

1960-03599 Incidental 1 95 AHY 243 481 96 ASY 242 484 316

1960-03604 Incidental 1 94 ASY 159 603 95 A3Y 161 603 200

1960-03608 Incidental 1 94 ASY 168 611 95 A3Y 167 611 100

1960-03622 Incidental 1 94 ASY 169 598 95 A3Y 169 599 100

1960-03623 Incidental 1 94 ASY 175 613 95 A3Y 175 613 0
USACERL TR-98/52 53

Banding Information Resighting Information

Band

number

Search

Type

No. times

resighted Year Age East North Year Age East North

Distance

(m)

1960-03625 Incidental 1 94 ASY 164 608 95 A3Y 164 609 100

1960-03699 Incidental 1 94 AHY 316 537 95 ASY 315 534 316

1960-03704 Incidental 1 94 AHY 349 443 95 ASY 352 444 316

1960-03713 Incidental 1 95 ASY 331 490 96 A3Y 334 491 316

1960-03719 Incidental 1 95 ASY 328 524 96 A3Y 327 524 100

1960-03763 Incidental 1 95 ASY 156 706 96 A3Y 156 706 0

1960-03807 Incidental 1 95 SY 360 517 96 3Y 362 514 361

1960-03832 Incidental 1 95 SY 331 487 96 3Y 331 485 200

1290-90840 Search 1 93 ASY 319 536 94 A3Y 320 536 100

1290-90843 Search 1 93 SY 339 556 94 3Y 340 555 141

1290-90854 Search 3 93 HY 298 486 96 4Y 299 488 224

1290-90857 Search 1 93 AHY 315 560 95 A3Y 315 558 200

1480-34211 Search 1 91 ASY 331 556 92 A3Y UK UK UK

1480-34221 Search 1 91 ASY 338 562 92 A3Y UK UK 130

1480-34226 Search 1 91 ASY 321 499 93 A4Y 321 499 0

1480-34227 Search 1 91 SY 320 498 92 3Y UK UK 10

1480-34230 Search 1 91 ASY 233 605 92 A3Y UK UK 15

1480-34236 Search 1 91 ASY 306 442 92 A3Y UK UK 45

1480-34239 Search 1 91 ASY 386 522 92 A3Y UK UK 35

1480-34242 Search 1 91 ASY 316 537 92 A3Y UK UK 220

1480-34242 Search 2 91 ASY UK UK 93 A4Y 317 537 UK

1480-34242 Search 3 91 ASY 317 537 95 A6Y 316 539 224

1480-34243 Search 1 91 SY 315 539 92 3Y UK UK 60

1480-34248 Search 1 91 ASY UK UK 92 A3Y UK UK 120

1480-34250 Search 1 91 SY UK UK 92 3Y UK UK 490

1480-34258 Search 2 91 ASY 303 486 95 A6Y 301 486 200

1480-34261 Search 1 91 ASY UK UK 92 A3Y UK UK 85

1710-25235 Search 1 92 ASY 244 483 93 A3Y 243 482 141

1710-25235 Search 2 92 ASY 243 482 94 A4Y 243 482 0

1710-25235 Search 3 92 ASY 243 482 95 A5Y 244 482 100

1710-25248 Search 1 92 ASY 308 486 93 A3Y 308 487 100

1710-25249 Search 1 92 AHY 374 554 93 ASY 372 557 361

1710-25252 Search 1 92 SY 317 560 93 3Y 317 561 100

1710-25252 Search 2 92 SY 317 561 94 4Y 316 561 100

1710-25256 Search 1 92 SY 363 525 93 3Y 363 525 0

1710-25256 Search 2 92 SY 363 525 94 4Y 362 525 100

1710-25258 Search 1 92 SY 338 561 93 3Y 339 560 141

1710-25259 Search 1 92 SY 339 562 93 3Y 340 563 141

1710-25263 Search 1 92 ASY 363 514 93 A3Y 363 514 0

1710-25267 Search 1 92 ASY 350 442 94 A4Y 352 443 224

1710-25277 Search 1 92 AHY 83 388 93 ASY 82 388 100

1710-25277 Search 2 92 AHY 82 388 94 A3Y 82 388 0

1710-25277 Search 3 92 AHY 82 388 95 A4Y 81 388 100

1710-25277 Search 4 92 AHY 81 388 96 A5Y 81 388 0

1710-25280 Search 1 92 AHY 79 388 93 ASY 79 388 0

1710-25282 Search 1 92 AHY 89 386 93 ASY 88 387 141

1710-25282 Search 2 92 AHY 88 387 94 A3Y 88 388 100

1710-25370 Search 1 92 AHY 74 390 93 ASY 73 391 141

1710-25374 Search 1 92 AHY 317 561 93 ASY 317 561 0
54 USACERL TR 98/52

Banding Information Resighting Information

Band

number

Search

Type

No. times

resighted Year Age East North Year Age East North

Distance

(m)

1710-25402 Search 2 93 ASY 75 393 95 A4Y 75 389 400

1710-25404 Search 1 93 AHY 184 636 94 ASY 183 637 141

1710-25406 Search 1 93 ASY 201 634 94 A3Y 201 633 100

1710-25410 Search 1 93 SY 208 613 94 3Y 208 613 0

1710-25412 Search 1 93 ASY 317 561 94 A3Y 316 561 100

1710-25412 Search 2 93 ASY 316 561 95 A4Y 315 558 316

1710-25413 Search 1 93 SY 312 565 94 3Y 312 565 0

1710-25414 Search 1 93 SY 311 564 94 3Y 312 565 141

1710-25415 Search 1 93 SY 317 563 94 3Y 317 563 0

1710-25416 Search 1 93 ASY 317 537 94 A3Y 318 536 141

1710-25425 Search 1 93 SY 313 542 94 3Y 312 539 316

1710-25426 Search 1 93 ASY 351 559 94 A3Y 351 559 0

1710-25427 Search 1 93 SY 346 561 94 3Y 346 560 100

1710-25427 Search 2 93 SY 346 560 95 4Y 351 559 510

1710-25432 Search 1 93 SY 373 558 94 3Y 373 557 100

1710-25432 Search 2 93 SY 373 557 95 4Y 373 558 100

1710-25434 Search 1 93 ASY 385 421 94 A3Y 386 422 141

1710-25436 Search 1 93 ASY 388 434 94 A3Y 388 435 100

1710-25440 Search 1 93 SY 387 449 94 3Y 387 449 0

1710-25444 Search 1 93 ASY 122 649 94 A3Y 122 649 0

1710-25446 Search 1 93 ASY 119 654 94 A3Y 119 653 100

1710-25447 Search 1 93 SY 119 654 94 3Y 119 653 100

1710-25452 Search 1 93 SY 334 491 94 3Y 335 491 100

1710-25453 Search 1 93 SY 331 490 94 3Y 332 491 141

1710-25459 Search 1 93 SY 307 484 94 3Y 307 484 0

1710-25460 Search 1 93 ASY 305 486 94 A3Y 306 486 100

1710-25461 Search 1 93 ASY 304 486 94 A3Y 304 485 100

1710-25465 Search 1 93 SY 330 486 94 3Y 331 487 141

1710-25470 Search 1 93 ASY 333 584 94 A3Y 331 584 200

1710-25471 Search 1 93 ASY 333 584 94 A3Y 331 584 200

1710-25471 Search 2 93 ASY 331 584 96 A5Y 328 583 316

1710-25472 Search 1 93 ASY 331 586 94 A3Y 331 584 200

1710-25475 Search 1 93 SY 163 584 95 4Y 162 585 141

1710-25486 Search 1 93 ASY 354 568 94 A3Y 354 568 0

1710-25487 Search 1 93 SY 383 556 94 3Y 384 555 141

1710-25487 Search 2 93 SY 384 555 95 4Y 383 556 141

1710-25491 Search 1 93 AHY 344 563 94 ASY 346 564 224

1810-28925 Search 1 91 ASY 369 562 92 A3Y UK UK 400

1810-28926 Search 1 91 ASY 369 562 92 A3Y UK UK 20

1810-28929 Search 1 91 SY 245 484 92 3Y UK UK 135

1810-28929 Search 3 91 SY 242 482 94 5Y 243 482 100

1810-28947 Search 1 91 AHY 374 555 92 ASY UK UK 120

1810-28974 Search 1 91 ASY 316 561 92 A3Y UK UK 75

1810-28975 Search 1 91 ASY 317 560 92 A3Y UK UK 30

1810-28975 Search 2 91 ASY UK UK 93 A4Y 123 652 UK

1810-28980 Search 1 91 SY 308 543 92 3Y UK UK 340

1810-28985 Search 1 91 AHY 116 659 92 ASY UK UK 110

1810-28985 Search 4 91 AHY 122 650 96 A6Y 123 652 224

1810-28986 Search 1 91 AHY 370 557 92 ASY UK UK 25
USACERL TR-98/52 55

Banding Information Resighting Information

Band

number

Search

Type

No. times

resighted Year Age East North Year Age East North

Distance

(m)

1810-28988 Search 1 91 AHY 364 525 92 ASY UK UK 170

1880-64767 Search 1 91 AHY 304 547 92 ASY UK UK 15

1880-64769 Search 1 91 AHY 337 557 92 ASY UK UK 60

1930-72715 Search 1 92 ASY 242 482 93 A3Y 242 482 0

1930-72715 Search 2 92 ASY 242 482 94 A4Y 243 482 100

1930-72715 Search 3 92 ASY 243 482 95 A5Y 242 482 100

1930-72715 Search 4 92 ASY 242 482 96 A6Y 244 482 200

1930-72724 Search 1 92 ASY 378 558 94 A4Y 378 558 0

1930-72730 Search 1 92 U 345 564 93 AHY 339 559 781

1930-72756 Search 1 92 AHY 352 567 93 ASY 353 566 141

1930-72757 Search 1 92 U 354 568 93 AHY 353 565 316

1930-72771 Search 1 92 AHY 307 488 93 ASY 307 487 100

1930-72773 Search 1 92 AHY 315 505 93 ASY 313 506 224

1930-72773 Search 3 92 AHY 313 506 95 A4Y 314 505 141

1960-03020 Search 1 93 AHY 362 533 94 ASY 361 534 141

1960-03022 Search 1 93 AHY 363 528 94 ASY 363 528 0

1960-03033 Search 2 93 AHY 122 651 95 A3Y 122 652 100

1960-03044 Search 1 93 AHY 389 433 94 ASY 389 432 100

1960-03047 Search 1 93 AHY 349 443 94 ASY 352 443 300

1960-03048 Search 1 93 AHY 347 445 94 ASY 347 439 600

1960-03049 Search 1 93 AHY 346 442 94 ASY 346 442 0

1960-03106 Search 1 94 SY 77 382 95 3Y 77 381 100

1960-03128 Search 1 94 SY 351 559 95 3Y 351 558 100

1960-03137 Search 1 94 ASY 79 377 95 A3Y 79 377 0

1960-03144 Search 1 94 ASY 185 643 95 A3Y 184 643 100

1960-03149 Search 2 94 AHY 308 487 96 A3Y 308 488 100

1960-03174 Search 1 94 SY 313 542 95 3Y 313 542 0

1960-03190 Search 1 94 SY 374 554 95 3Y 374 554 0

1960-03405 Search 1 95 SY 153 322 96 3Y 159 319 671

1960-03416 Search 1 95 SY 156 707 96 3Y 155 705 224

1960-03451 Search 1 95 ASY 166 586 96 A3Y 165 587 141

1960-03480 Search 1 95 ASY 318 500 96 A3Y 318 500 0

1960-03627 Search 1 94 ASY 317 501 95 A3Y 317 501 0

1960-03631 Search 1 94 SY 321 500 95 3Y 321 499 100

1960-03632 Search 1 94 SY 317 497 95 3Y 319 496 224

1960-03640 Search 1 94 AHY 314 503 95 ASY 314 505 200

1960-03644 Search 1 94 AHY 363 527 95 ASY 363 528 100

1960-03760 Search 1 95 SY 165 593 96 3Y 162 594 316

1960-03772 Search 1 95 SY 165 593 96 3Y 165 592 100

1960-03775 Search 1 95 SY 141 649 96 3Y 142 648 141

1960-03777 Search 1 95 SY 156 648 96 3Y 157 649 141

1960-03778 Search 1 95 SY 153 647 96 3Y 156 648 316

1960-03796 Search 1 95 ASY 160 703 96 A3Y 163 701 361

1960-03811 Search 1 95 ASY 367 522 96 A3Y 366 520 224

1960-03823 Search 1 95 ASY 328 484 96 A3Y 330 484 200

1960-03831 Search 1 95 SY 331 485 96 3Y 332 489 412

1960-03841 Search 1 95 ASY 134 646 96 A3Y 134 646 0

1960-03851 Search 1 95 ASY 119 654 96 A3Y 122 655 316

2060-20811 Search 1 95 SY 300 483 96 3Y 297 486 424
56 USACERL TR 98/52

Banding Information Resighting Information

Band

number

Search

Type

No. times

resighted Year Age East North Year Age East North

Distance

(m)

3500-71102 Search 1 95 AHY 123 648 96 ASY 123 652 400
USACERL TR-98/52 57

Appendix B: List of all Adult Males

Searched for and Not Found on Fort

Hood

This list is presented by search type and band number. Also included at the end is a list of males that

were searched for and not found one year, but were found in a subsequent year.

Band number Search Type Year Banded Age at Banding Year Searched Age Searched

1290-90820 ISA 1992 AHY 1993 ASY

1290-90821 ISA 1992 AHY 1993 ASY

1290-90833 ISA 1993 SY 1995 5Y

1290-90834 ISA 1993 SY 1996 6Y

1290-90835 ISA 1993 SY 1994 3Y

1290-90836 ISA 1993 SY 1996 6Y

1290-90837 ISA 1993 SY 1996 6Y

1290-90847 ISA 1993 SY 1996 6Y

1290-90848 ISA 1993 SY 1995 5Y

1290-90856 ISA 1993 AHY 1994 ASY

1710-25228 ISA 1992 ASY 1994 A5Y

1710-25230 ISA 1992 SY 1993 3Y

1710-25233 ISA 1992 SY 1993 3Y

1710-25234 ISA 1992 ASY 1993 A3Y

1710-25242 ISA 1992 ASY 1995 A5Y

1710-25244 ISA 1992 SY 1993 3Y

1710-25245 ISA 1992 ASY 1996 A6Y

1710-25265 ISA 1992 ASY 1996 A6Y

1710-25268 ISA 1992 ASY 1994 A4Y

1710-25378 ISA 1992 ASY 1993 A3Y

1710-25380 ISA 1992 SY 1994 4Y

1710-25381 ISA 1992 ASY 1994 A4Y

1710-25382 ISA 1992 ASY 1994 A4Y

1710-25384 ISA 1992 SY 1994 4Y

1710-25386 ISA 1992 SY 1993 3Y

1710-25417 ISA 1993 ASY 1994 A3Y

1710-25418 ISA 1993 SY 1994 3Y

1710-25419 ISA 1993 SY 1994 3Y

1880-64770 ISA 1991 ASY 1993 A4Y

1880-64771 ISA 1991 ASY 1993 A4Y

1880-64773 ISA 1991 ASY 1992 A3Y

1880-64783 ISA 1991 ASY 1992 A3Y

1930-72772 ISA 1992 AHY 1993 ASY

1960-03104 ISA 1994 ASY 1996 A4Y

1960-03108 ISA 1994 SY 1996 4Y

1960-03109 ISA 1994 SY 1995 3Y
58 USACERL TR 98/52

Band number Search Type Year Banded Age at Banding Year Searched Age Searched

1960-03110 ISA 1994 SY 1996 4Y

1960-03111 ISA 1994 SY 1996 4Y

1960-03115 ISA 1994 SY 1996 4Y

1960-03116 ISA 1994 ASY 1996 A4Y

1960-03117 ISA 1994 SY 1995 3Y

1960-03132 ISA 1994 SY 1995 3Y

1960-03201 ISA 1994 SY 1996 4Y

1960-03202 ISA 1994 SY 1995 3Y

1960-03204 ISA 1994 SY 1995 3Y

1960-03207 ISA 1994 SY 1996 4Y

1960-03208 ISA 1994 SY 1996 4Y

1960-03209 ISA 1994 ASY 1996 A4Y

1960-03212 ISA 1994 HY 1996 3Y

1960-03229 ISA 1994 HY 1996 3Y

1960-03238 ISA 1994 AHY 1996 A3Y

1960-03407 ISA 1995 SY 1996 3Y

1960-03408 ISA 1995 SY 1996 3Y

1960-03529 ISA 1995 ASY 1996 A3Y

1960-03530 ISA 1995 ASY 1996 A3Y

1960-03531 ISA 1995 ASY 1996 A3Y

1960-03534 ISA 1995 ASY 1996 A3Y

1960-03535 ISA 1995 ASY 1996 A3Y

1960-03536 ISA 1995 SY 1996 3Y

1960-03537 ISA 1995 ASY 1996 A3Y

1960-03538 ISA 1995 SY 1996 3Y

1960-03547 ISA 1995 SY 1996 3Y

1960-03548 ISA 1995 SY 1996 3Y

1960-03708 ISA 1995 ASY 1996 A3Y

1960-03710 ISA 1995 ASY 1996 A3Y

1960-03801 ISA 1995 SY 1996 3Y

1960-03814 ISA 1995 SY 1996 3Y

1960-03819 ISA 1995 SY 1996 3Y

1960-03820 ISA 1995 SY 1996 3Y

1960-03822 ISA 1995 SY 1996 3Y

1960-03828 ISA 1995 SY 1996 3Y

1960-03829 ISA 1995 SY 1996 3Y

1960-03834 ISA 1995 SY 1996 3Y

1960-03901 ISA 1995 SY 1996 3Y

1290-90817 Search 1992 U 1993 AHY

1290-90840 Search 1993 ASY 1995 A4Y

1290-90841 Search 1993 ASY 1994 A3Y

1290-90842 Search 1993 SY 1994 3Y

1480-34227 Search 1991 SY 1993 4Y

1480-34238 Search 1991 SY 1996 7Y

1480-34252 Search 1991 ASY 1993 A4Y

1480-34253 Search 1991 SY 1993 4Y

1710-25235 Search 1992 ASY 1996 A6Y

1710-25236 Search 1992 SY 1994 4Y

1710-25243 Search 1992 ASY 1993 A3Y

1710-25247 Search 1992 SY 1993 3Y
USACERL TR-98/52 59

Band number Search Type Year Banded Age at Banding Year Searched Age Searched

1710-25252 Search 1992 SY 1995 5Y

1710-25253 Search 1992 SY 1994 4Y

1710-25255 Search 1992 ASY 1993 A3Y

1710-25257 Search 1992 ASY 1993 A3Y

1710-25260 Search 1992 ASY 1994 A4Y

1710-25262 Search 1992 SY 1993 3Y

1710-25263 Search 1992 ASY 1994 A4Y

1710-25278 Search 1992 ASY 1993 A3Y

1710-25279 Search 1992 ASY 1993 A3Y

1710-25282 Search 1992 AHY 1996 A5Y

1710-25283 Search 1992 AHY 1993 ASY

1710-25374 Search 1992 AHY 1994 A3Y

1710-25379 Search 1992 ASY 1993 A3Y

1710-25403 Search 1993 SY 1996 5Y

1710-25404 Search 1993 AHY 1996 A4Y

1710-25406 Search 1993 ASY 1996 A5Y

1710-25407 Search 1993 SY 1994 3Y

1710-25408 Search 1993 SY 1994 3Y

1710-25411 Search 1993 AHY 1994 ASY

1710-25413 Search 1993 SY 1995 4Y

1710-25416 Search 1993 ASY 1995 A4Y

1710-25421 Search 1993 ASY 1994 A3Y

1710-25423 Search 1993 SY 1996 5Y

1710-25424 Search 1993 SY 1994 3Y

1710-25426 Search 1993 ASY 1995 A4Y

1710-25427 Search 1993 SY 1996 5Y

1710-25428 Search 1993 ASY 1994 A3Y

1710-25429 Search 1993 SY 1994 3Y

1710-25430 Search 1993 SY 1996 5Y

1710-25432 Search 1993 SY 1996 5Y

1710-25433 Search 1993 SY 1994 3Y

1710-25435 Search 1993 SY 1994 3Y

1710-25437 Search 1993 ASY 1994 A3Y

1710-25438 Search 1993 SY 1994 3Y

1710-25439 Search 1993 AHY 1994 ASY

1710-25440 Search 1993 SY 1996 5Y

1710-25442 Search 1993 SY 1994 3Y

1710-25443 Search 1993 SY 1994 3Y

1710-25444 Search 1993 ASY 1995 A4Y

1710-25445 Search 1993 SY 1994 3Y

1710-25446 Search 1993 ASY 1995 A4Y

1710-25447 Search 1993 SY 1995 4Y

1710-25448 Search 1993 ASY 1994 A3Y

1710-25449 Search 1993 ASY 1996 A5Y

1710-25450 Search 1993 ASY 1996 A5Y

1710-25451 Search 1993 ASY 1994 A3Y

1710-25462 Search 1993 ASY 1994 A3Y

1710-25467 Search 1993 ASY 1994 A3Y

1710-25468 Search 1993 ASY 1995 A4Y

1710-25472 Search 1993 ASY 1996 A5Y
60 USACERL TR 98/52

Band number Search Type Year Banded Age at Banding Year Searched Age Searched

1710-25473 Search 1993 AHY 1994 ASY

1710-25476 Search 1993 SY 1995 4Y

1710-25480 Search 1993 ASY 1996 A5Y

1710-25482 Search 1993 ASY 1994 A3Y

1710-25484 Search 1993 AHY 1994 ASY

1710-25487 Search 1993 SY 1996 5Y

1710-25494 Search 1993 AHY 1995 A3Y

1710-25496 Search 1993 AHY 1995 A3Y

1810-28925 Search 1991 ASY 1993 A4Y

1810-28926 Search 1991 ASY 1993 A4Y

1810-28927 Search 1991 ASY 1993 A4Y

1810-28929 Search 1991 SY 1995 6Y

1810-28935 Search 1991 ASY 1993 A4Y

1810-28947 Search 1991 AHY 1993 A3Y

1880-64774 Search 1991 ASY 1993 A4Y

1930-72709 Search 1992 SY 1993 3Y

1930-72714 Search 1992 ASY 1993 A3Y

1930-72717 Search 1992 ASY 1993 A3Y

1930-72718 Search 1992 ASY 1993 A3Y

1930-72729 Search 1992 U 1993 AHY

1930-72751 Search 1992 AHY 1993 ASY

1930-72758 Search 1992 U 1993 AHY

1930-72759 Search 1992 AHY 1993 ASY

1930-72760 Search 1992 AHY 1993 ASY

1930-72761 Search 1992 AHY 1993 ASY

1930-72771 Search 1992 AHY 1996 A5Y

1960-03014 Search 1993 AHY 1994 ASY

1960-03016 Search 1993 AHY 1994 ASY

1960-03028 Search 1993 AHY 1994 ASY

1960-03031 Search 1993 AHY 1995 A3Y

1960-03032 Search 1993 AHY 1994 ASY

1960-03033 Search 1993 AHY 1996 A4Y

1960-03037 Search 1993 AHY 1996 A4Y

1960-03047 Search 1993 AHY 1995 A3Y

1960-03048 Search 1993 AHY 1995 A3Y

1960-03049 Search 1993 AHY 1996 A4Y

1960-03060 Search 1993 AHY 1995 A3Y

1960-03067 Search 1993 AHY 1994 ASY

1960-03068 Search 1993 AHY 1994 ASY

1960-03069 Search 1993 AHY 1994 ASY

1960-03107 Search 1994 ASY 1996 A4Y

1960-03118 Search 1994 ASY 1995 A3Y

1960-03122 Search 1994 ASY 1995 A3Y

1960-03124 Search 1994 ASY 1995 A3Y

1960-03125 Search 1994 ASY 1995 A3Y

1960-03126 Search 1994 SY 1995 3Y

1960-03134 Search 1994 ASY 1995 A3Y

1960-03137 Search 1994 ASY 1996 A4Y

1960-03139 Search 1994 SY 1995 3Y

1960-03142 Search 1994 ASY 1996 A4Y

1960-03143 Search 1994 ASY 1995 A3Y
USACERL TR-98/52 61

Band number Search Type Year Banded Age at Banding Year Searched Age Searched

1960-03145 Search 1994 ASY 1995 A3Y

1960-03147 Search 1994 SY 1996 4Y

1960-03150 Search 1994 SY 1996 4Y

1960-03151 Search 1994 SY 1996 4Y

1960-03154 Search 1994 ASY 1996 A4Y

1960-03161 Search 1994 SY 1995 3Y

1960-03162 Search 1994 SY 1996 4Y

1960-03163 Search 1994 ASY 1996 A4Y

1960-03165 Search 1994 SY 1996 4Y

1960-03166 Search 1994 ASY 1995 A3Y

1960-03167 Search 1994 ASY 1995 A3Y

1960-03168 Search 1994 ASY 1995 A3Y

1960-03169 Search 1994 ASY 1996 A4Y

1960-03170 Search 1994 ASY 1996 A4Y

1960-03171 Search 1994 SY 1996 4Y

1960-03181 Search 1994 ASY 1996 A4Y

1960-03187 Search 1994 ASY 1995 A3Y

1960-03188 Search 1994 ASY 1995 A3Y

1960-03190 Search 1994 SY 1996 4Y

1960-03191 Search 1994 ASY 1996 A4Y

1960-03192 Search 1994 SY 1995 3Y

1960-03224 Search 1994 AHY 1995 ASY

1960-03409 Search 1995 ASY 1996 A3Y

1960-03415 Search 1995 ASY 1996 A3Y

1960-03423 Search 1995 SY 1996 3Y

1960-03425 Search 1995 SY 1996 3Y

1960-03444 Search 1995 SY 1996 3Y

1960-03446 Search 1995 ASY 1996 A3Y

1960-03447 Search 1995 ASY 1996 A3Y

1960-03452 Search 1995 SY 1996 3Y

1960-03461 Search 1995 ASY 1996 A3Y

1960-03462 Search 1995 SY 1996 3Y

1960-03612 Search 1994 SY 1995 3Y

1960-03627 Search 1994 ASY 1996 A4Y

1960-03631 Search 1994 SY 1996 4Y

1960-03633 Search 1994 SY 1995 3Y

1960-03639 Search 1994 AHY 1995 ASY

1960-03667 Search 1994 AHY 1995 ASY

1960-03676 Search 1994 AHY 1995 ASY

1960-03703 Search 1994 AHY 1995 ASY

1960-03711 Search 1995 ASY 1996 A3Y

1960-03714 Search 1995 ASY 1996 A3Y

1960-03715 Search 1995 ASY 1996 A3Y

1960-03717 Search 1995 ASY 1996 A3Y

1960-03718 Search 1995 ASY 1996 A3Y

1960-03755 Search 1995 ASY 1996 A3Y

1960-03757 Search 1995 SY 1996 3Y

1960-03758 Search 1995 ASY 1996 A3Y

1960-03761 Search 1995 ASY 1996 A3Y

1960-03762 Search 1995 ASY 1996 A3Y

1960-03774 Search 1995 ASY 1996 A3Y
62 USACERL TR 98/52

Band number Search Type Year Banded Age at Banding Year Searched Age Searched

1960-03780 Search 1995 SY 1996 3Y

1960-03781 Search 1995 ASY 1996 A3Y

1960-03782 Search 1995 ASY 1996 A3Y

1960-03783 Search 1995 SY 1996 3Y

1960-03784 Search 1995 SY 1996 3Y

1960-03787 Search 1995 SY 1996 3Y

1960-03788 Search 1995 ASY 1996 A3Y

1960-03794 Search 1995 ASY 1996 A3Y

1960-03802 Search 1995 ASY 1996 A3Y

1960-03803 Search 1995 ASY 1996 A3Y

1960-03804 Search 1995 SY 1996 3Y

1960-03805 Search 1995 SY 1996 3Y

1960-03809 Search 1995 ASY 1996 A3Y

1960-03813 Search 1995 SY 1996 3Y

1960-03817 Search 1995 SY 1996 3Y

1960-03826 Search 1995 SY 1996 3Y

1960-03842 Search 1995 SY 1996 3Y

1960-03843 Search 1995 ASY 1996 A3Y

1960-03856 Search 1995 ASY 1996 A3Y

The following males were FOUND in a later year, after being searched for and NOT found:

Band number Search type Year Bd BdAge Year Search Age Search

1290-90819 ISA 1992 AHY 1996 A5Y

1710-25254 Search 1992 ASY 1993 A3Y

1710-25420 Search 1993 SY 1994 3Y

1710-25431 Search 1993 ASY 1994 A3Y

1710-25466 Search 1993 ASY 1994 A3Y

1930-72724 Search 1992 ASY 1993 A3Y
USACERL TR-98/52 63

Appendix C: List of all Males Located

Within the ISA Including Non-territorial

Birds

Territory numbers correspond to those listed in Figures 11-15. Two separate

numbers listed in the “No. HY” column indicate double brooding. “UK” =

unknown male or missing data. “UB” = unbanded male.

1996

Band

Combo

Band

Number

Age in

1996

Year

Banded

Age

Banded

1996 Terr. No. Mated in

1996?

No. HY

in 1996

BkPiGrSi 2060-20826 SY 1996 SY 1 Yes UK

BlPiPiSi 1960-03877 SY 1995 HY 2 Yes 1

BkSiWh 1960-03833 3Y 1995 SY 3 Yes 2

BkSiOrPi 1960-03815 3Y 1995 SY 4 Yes 0

RdSiPiYe 2060-20828 AHY 1996 AHY 5 Yes 2

GrPiMgSi 1960-03838 3Y 1995 SY 6 Yes 2

BkSiMgRd 2060-20602 ASY 1996 ASY 7 Yes 1

YeSiYeGr 1960-03821 A3Y 1995 ASY 8 Yes 2

BkWhMgSi 1960-03816 3Y 1995 SY 9 Yes 2

PiSiOrLa 3500-71111 SY 1995 HY 10 Yes 4&1

GrSiMgYe 2060-20823 SY 1996 SY 12 Yes 0

OrRdGrSi 2060-20912 ASY 1996 ASY 13 Yes 1

OrSiOrBk 1960-03798 3Y 1995 SY 14 Yes 2

BkSiBkBk 1930-03634 3Y 1994 HY 15 Yes 0

WhSi:Ye 1710-25246 6Y 1992 SY 16 Yes 3

WhSiYePi 1960-03410 3Y 1995 SY 17 Yes 2

RdSi:Ye 1710-25464 5Y 1993 SY 18 Yes 2

MgSi:Or 2060-20632 SY 1996 SY 19 Yes 1

BlOrRdSi 2060-20911 SY 1996 SY 20 Yes 2

GrLaYeSi 1960-03861 SY 1995 HY 21 Yes 2

OrRdPiSi 2060-20901 ASY 1996 ASY 22 Yes 2

Bl:WhSi 1290-90838 5Y 1993 SY 23 Yes 2&2

PiPiMgSi 2060-20906 SY 1996 SY 25 Yes 2

BlBlWhSi 1960-03709 A3Y 1995 ASY 26 Yes 1&1

BkSiPiBl 2060-20902 ASY 1996 ASY 27 Yes 3

YeBkMgSi 1960-03206 4Y 1994 SY 28 Yes 2

MgSiBkLa 1960-03553 A3Y 1995 ASY 29 Yes 2

LaBkOrSi 1960-03533 A3Y 1995 ASY 30 Yes 2

GrSiMgMg 2060-20824 SY 1996 SY 31 UK UK

Si:PiBl 1960-03883 SY 1995 HY 32 No --

UB 11 Yes UK

UB 24 Yes 1
64 USACERL TR 98/52

Band

Combo

Band

Number

Age in

1996

Year

Banded

Age Banded 1996 Terr.

No.

Mated in

1996?

No. HY in

1996

Non-territorial in 1996:

GrSiYeRd 1960-03203 4Y 1994 SY

WhSiOrRd 2060-20910 ASY 1996 ASY

LaSiRdRd 2060-20601 ASY 1996 ASY

BlSiOrRd 2060-20908 ASY 1996 ASY

LaSiYeBk? UK UK
USACERL TR-98/52 65

1995

Band

Combo

Band

Number

Age in

1995

Year

Banded

Age

Banded

1995 Terr

No.

Mated in

1995?

No. HY in

1995

Returned in 1996?

OrRdBlSi 1290-90847 4Y 1993 SY 1 Yes 2 No

LvSiMgPi 1960-03820 SY 1995 SY 2 Yes 2 No

BkSiMgYe 1960-03822 SY 1995 SY 3 Yes 2 No

LvRdPiSi 1960-03828 SY 1995 SY 4 Yes 2 No

GrSiBlBk 1960-03819 SY 1995 SY 5 Yes 2 No

BlSiPiRd 1960-03116 A3Y 1994 ASY 6 Yes 0 No

BkSi:Wh 1960-03833 SY 1995 SY 7 Yes 3 Yes

BkPi:Si 1960-03529 ASY 1995 ASY 8 Yes 3 No

BkWhMgSi 1960-03816 SY 1995 SY 9 Yes 2 Yes

GrLvWhSi 1960-03801 SY 1995 SY 10 Yes 2 No

GrRdOrSi 1290-90819 A4Y 1992 AHY 11 Yes 4 No

BlMgMgSi 1960-03111 3Y 1994 SY 12 Yes 1&2 No

YeSiYeGr 1960-03821 ASY 1995 ASY 13 Yes 1 Yes

OrSiGrMg 1960-03531 ASY 1995 ASY 14 Yes 1 No

OrSiRdGr 1960-03110 3Y 1994 SY 15 Yes 2 No

GrSiGrOr 1960-03108 3Y 1994 SY 16 Yes 3 No

MgSiBlBk 1960-03814 SY 1995 SY 17 Yes 2 No

BkSiOrPi 1960-03815 SY 1995 SY 18 Yes 2 Yes

WhSiYePi 1960-03410 SY 1995 SY 19 Yes 2 Yes

BkPiOrSi 1960-03534 ASY 1995 ASY 20 Yes 1 No

Rd:WhSi 1290-90836 4Y 1993 SY 21 Yes 2 No

RdBkWhSi 1960-03238 ASY 1994 AHY 22 Yes 3 No

YeBkYeSi 1960-03538 SY 1995 SY 23 Yes 2 No

MgGrMgSi 1960-03537 ASY 1995 ASY 24 Yes 2 No

WhRdMgSi 1960-03229 SY 1994 HY 26 Yes 2 No

Wh:YeSi 1710-25246 5Y 1992 SY 27 Yes 2 Yes

BkWhOrSi 1290-90834 4Y 1993 SY 28 Yes 2 No

YeSiLvBk 1960-03530 ASY 1995 ASY 29 Yes 2 No

YePiYeSi 1960-03675 SY 1994 HY 30 Yes 0 No

OrSiOrBk 1960-03798 SY 1995 SY 31 No -- Yes

RdSi:Ye 1710-25464 4Y 1993 SY 32 Yes 2 Yes

Bk:PiSi 1960-03536 SY 1995 SY 33 Yes 2 No

OrSiYeLv 1960-03209 A3Y 1994 ASY 34 Yes 1 No

LvPiPiSi 1960-03208 3Y 1994 SY 35 Yes 1&2 No

BlSiYeMg 1960-03212 SY 1994 HY 36 No -- No

RdYeBlSi 1710-25245 A5Y 1992 ASY 37 Yes 3 No

OrSiYeOr 1960-03115 3Y 1994 SY 38 Yes 2 No

Bl:WhSi 1290-90838 4Y 1993 SY 39 Yes 3 Yes

RdSiPiPi 1960-03548 SY 1995 SY 40 No -- No

BlBlWhSi 1960-03709 ASY 1995 ASY 41 Yes 4 Yes

BlSiGrMg 1960-03708 ASY 1995 ASY 42 Yes 3 No

WhSiYeOr 1960-03207 3Y 1994 SY 43 Yes 2 No

Si:LvLv 1960-03710 ASY 1995 ASY 44 Yes 3 No

GrSiYeRd 1960-03203 3Y 1994 SY 45 Yes 3 Yes

YeBkMgSi 1960-03206 3Y 1994 SY 46 Yes 3 Yes

LvBkOrSi 1960-03533 ASY 1995 ASY 47 Yes 3 Yes

MgSiBkLv 1960-03553 ASY 1995 ASY 48 Yes 2 Yes

MgSiRdBl 1960-03408 SY 1995 SY 49 Yes 3 No
66 USACERL TR 98/52

Band

Combo

Band

Number

Age in

1995

Year

Banded

Age

Banded

1995 Terr

No.

Mated in

1995?

No. HY in

1995

Returned in 1996?

PiSiRdBl 1960-03829 SY 1995 SY 50 Yes 1&1 No

LvSi:Or 1960-03834 SY 1995 SY 51 No -- No

GrPiMgSi 1960-03838 SY 1995 SY 52 Yes 1 Yes

Non-territorial in 1995:

YeSiYeWh 1960-03901 SY 1995 SY No

PiSiBlBk 1960-03407 SY 1995 SY No

OrSiRdBk 1960-03798 ASY 1995 ASY No

GrMgBkSi 1960-03547 SY 1995 SY No

BkSiBkBk 1960-03634 SY 1994 HY Yes

Or:YeSi 1710-25265 A5Y 1992 ASY No

Pi:RdSi 1960-03104 A3Y 1994 ASY No

WhGrBlSi 1290-90833 4Y 1993 SY No
USACERL TR-98/52 67

1994

Band Combo Band

Number

Age

1994

Year

Banded

Age

Banded

1994 Terr.

No.

Mated in

1994?

No. HY in

1994

Returned 1995?

OrRdBlSi 1290-90847 3Y 1993 SY 1 Yes 0 Yes

Pi:RdSi 1960-03104 ASY 1994 ASY 2 Yes 3 Possibly

BkBlOrSi 1290-90848 3Y 1993 SY 3 Yes 4 No

BlSiPiRd 1960-03116 ASY 1994 ASY 4 Yes 2 Yes

GrSiLvWh 1960-03132 SY 1994 SY 5 Yes 2 No

UB 6 Yes 4

BlMgMgSi 1960-03111 SY 1994 SY 7 Yes 2 Yes

GrRdOrSi 1290-90819 A3Y 1992 AHY 8 Yes 3 Yes

GrSiGrOr 1960-03108 SY 1994 SY 9 Yes 0 Yes

Wh:BlSi 1710-25242 A4Y 1992 ASY 10 Yes 2 No

OrSiRdGr 1960-03110 SY 1994 SY 11 Yes 1 Yes

BkSiMgOr 1960-03109 SY 1994 SY 12 Yes 2 No

LvYeYeSi 1960-03117 SY 1994 SY 13 Yes 1 No

WhOrBlSi 1290-90833 3Y 1993 SY 14 Yes 2 No

BkRdMgSi 1960-03201 SY 1994 SY 15 Yes 1 Yes

RdSi:Ye 1710-25464 3Y 1993 SY 16 Yes 1 Yes

Wh:YeSi 1710-25246 4Y 1992 SY 17 Yes 3 Yes

BkWhOrSi 1290-90834 3Y 1993 SY 18 Yes 2 Yes

Rd:WhSi 1290-90836 3Y 1993 SY 19 Yes 2 Yes

UB 20 Yes 1

UB 21 UK 0

LvWhOrSi 1960-03202 SY 1994 SY 22 Yes 3 No

OrSiYeLv 1960-03209 ASY 1994 ASY 23 Yes 2 Yes

RdYeBlSi 1710-25245 A4Y 1992 ASY 24 Yes 2 Yes

LvPiPiSi 1960-03208 SY 1994 SY 24 No -- Yes

MgBkBkSi 1960-03204 SY 1994 SY 25 Yes 3 No

YeBkMgSi 1960-03206 SY 1994 SY 26 No -- Yes

Bl:WhSi 1290-90838 3Y 1993 SY 27 Yes 2 Yes

UB 28 Yes 3

WhGrBlSi 1290-90837 3Y 1993 SY 29 Yes 3 No

WhSiYeOr 1960-03207 SY 1994 SY 30 No -- Yes

GrOrBlSi 1290-90845 SY 1993 HY 31 Yes 2 No

Or:YeSi 1710-25265 A4Y 1992 ASY 32 UK -- Yes

GrSiYeRd 1960-03203 SY 1994 SY 33 Yes 3 Yes

UB 35 Yes 2

UB 36 UK 0

Non-territorial in 1994:

OrSiYeOr 1960-03115 SY 1994 SY NT Yes?? -- Yes

RdBkWhSi 1960-03238 AHY 1994 AHY NT -- -- Yes
68 USACERL TR 98/52

1993

Band Combo Band

Number

Age

1993

Year

Banded

Age

Banded

1993 Terr.

No.

Mated in

1993?

No. HY in

1993

Returned

in 1994?

BkBlOrSi 1290-90848 SY 1993 SY 1 Yes 4 Yes

RdOrGrSi 1710-25417 ASY 1993 ASY 2 Yes 1 No

GrRdOrSi 1290-90819 ASY 1992 AHY 3 Yes 2 Yes

UB 4 Yes 0

YeYeGrSi 1710-25418 SY 1993 SY 5 No -- No

Or:YeSi 1710-25265 A3Y 1992 ASY 6 Yes 1 Yes

Wh:BlSi 1710-25242 A3Y 1992 ASY 7 Yes 2 Yes

RdSi:Ye 1710-25464 SY 1993 SY 8 Yes 2 Yes

RdBkGrSi 1710-25419 SY 1993 SY 9 UK -- No

UB 10 Yes 2

Rd:WhSi 1290-90836 SY 1993 SY 11 Yes 2 Yes

Ye:YeSi 1710-25228 A3Y 1992 ASY 12 Yes 1 No

WhOrBlSi 1290-90833 SY 1993 SY 13 Yes 0 Yes

RdGrYeSi 1710-25268 A3Y 1992 ASY 14 Yes 2 No

BkWhOrSi 1290-90834 SY 1993 SY 15 Yes 3 Yes

Wh:YeSi 1710-25246 3Y 1992 SY 16 Yes 2 Yes

OrWhRdSi 1710-25380 3Y 1992 SY 17 Yes 3 No

WhWhOrSi 1290-90835 SY 1993 SY 18 Yes 4 No

RdYeBlSi 1710-25245 A3Y 1992 ASY 19 Yes 3 Yes

WhOrBlSi 1290-90833 A3Y 1992 ASY 20 Yes 2 No

BlGrRdSi 1710-25384 3Y 1992 SY 21 Yes 3 No

YeWhRdSi 1710-25381 A3Y 1992 ASY 22 Yes 2 No

WhGrBlSi 1290-90837 SY 1993 SY 23 Yes 2 Yes

Bl:WhSi 1290-90838 SY 1993 SY 24 No -- Yes

UB 25 Yes 2

Non-territorial in 1993:

OrRdBlSi 1290-90847 SY 1993 SY -- -- Yes

WhRdBlSi 1290-90856 AHY 1993 AHY -- -- No
USACERL TR-98/52 69

1992

Band

Combo

Band

Number

Age 1992 Year

Banded

Age

Banded

1992 Terr.

No.

Mated in

1992?

HY present

in 1992?

Returned in

1993?

GrYeYeSi 1710-25230 SY 1992 SY 1 Yes Yes No

YeYeRdSi 1710-25378 ASY 1992 ASY 2 No -- No

BlRdOrSi 1930-72772 AHY 1992 AHY 3 Yes No No

UB 4 Yes Yes

GrRdOrSi 1290-90819 AHY 1992 AHY 5 Yes No Yes

Or:YeSi 1710-25265 ASY 1992 ASY 6 Yes Yes Yes

UB 7 Yes N

RdBkBlSi 1710-25234 ASY 1992 ASY 8 Yes Yes No

Wh:BlSi 1710-25242 ASY 1992 ASY 9 No -- Yes

GrWhYeSi 1710-25233 SY 1992 SY 10 No -- No

UB 11 Yes Yes

Wh:Si 1880-64771 A3Y 1991 ASY 12 Yes Yes No

RdGrYeSi 1710-25268 ASY 1992 ASY 13 Yes Yes Yes

BlRdRdSi 1880-64770 A3Y 1991 ASY 14 Yes Yes No

UB 15 Yes Yes

RdYeBlSi 1710-25245 ASY 1992 ASY 16 Yes No Yes

WhBlRdSi 1710-25382 ASY 1992 ASY 17 Yes Yes Yes

Ye:YeSi 1710-25228 ASY 1992 ASY 18 Yes Yes Yes

OrWhRdSi 1710-25380 SY 1992 SY 19 Yes Yes Yes

Bk:BlSi 1710-25244 SY 1992 SY 20 Yes Yes No

Wh:YeSi 1710-25246 SY 1992 SY 21 Yes Yes Yes

YeWhRdSi 1710-25381 ASY 1992 ASY 22 Yes Yes Yes

BlGrRdSi 1710-25384 SY 1992 SY 23 Yes Yes Yes

WhBkRdSi 1710-25386 SY 1992 SY 24 No -- No

Non-territorial in 1992:

Or:BkSi 1290-90820 AHY 1992 AHY No

Rd:BkSi 1290-90821 AHY 1992 AHY No
70 USACERL TR 98/52

1991

Band Combo Band

Number

Age 1991 Year

Banded

Age

Banded

Returned in

1992?

Bk:Si 1880-64773 ASY 1991 ASY No

BlRdRdSi 1880-64770 ASY 1991 ASY Yes

Si:Bl 1880-64783 ASY 1991 ASY No

Wh:Si 1880-64771 ASY 1991 ASY Yes
USACERL TR-98/52 71

Appendix D: List of all Adult Females

Resighted on Fort Hood

This list is presented by band number and the number of times each was resighted.

Banding Information Resighting Information

Band

number

No. times

resighted Year Age East North Year Age East North

Distance

(m).

1480-34256 1 1991 ASY UK UK 1993 A4Y 305 486 UK

1960-03186 1 1994 AHY 319 563 1995 ASY 318 565 224

1960-03481 1 1995 ASY 320 500 1996 A3Y 320 498 200

1960-03532 1 1995 ASY 340 486 1996 A3Y 341 485 141

1960-03558 1 1995 AHY 314 542 1996 ASY 317 543 316

1960-03620 1 1994 ASY 310 537 1995 A3Y 310 538 100

1960-03620 2 1994 ASY 310 537 1996 A4Y 317 540 762

1960-03620 3 1994 ASY 310 537 1996 A4Y 320 538 1005

1960-03795 1 1995 ASY 157 704 1996 A3Y 156 704 100

1960-03862 1 1995 AHY 341 487 1996 ASY 342 487 100

1960-03862 1 1995 AHY 341 487 1996 ASY 339 485 283

1960-03864 1 1995 SY 345 484 1996 3Y 342 483 316
72 USACERL TR 98/52

Appendix E: List of all HY Males’ First

Resightings on Fort Hood.

This list is presented by resight type (ISA, incidental, searched for), in order by band

number.

Band Banding Information Resighting Information

number

Search

Type Year East North Year Age East North Distance (m)

1290-90845 ISA 1993 346 471 1994 SY 336 473 1020

1290-90846 ISA 1993 346 471 1994 SY 314 487 3578

1930-72734 ISA 1992 342 478 1993 SY 319 502 3324

1960-03212 ISA 1994 342 473 1995 SY 341 477 412

1960-03229 ISA 1994 346 487 1995 SY 339 481 922

1960-03634 ISA 1994 294 454 1995 SY 341 477 5233

1960-03675 ISA 1994 306 487 1995 SY 345 478 4002

1960-03861 ISA 1995 338 490 1996 SY 345 479 1304

1960-03877 ISA 1995 345 487 1996 SY 342 488 316

1960-03883 ISA 1995 342 488 1996 SY 347 470 1868

3500-71111 ISA 1995 342 475 1996 SY 338 483 894

1290-90854 Incidental 1993 332 476 1994 SY 298 486 3544

1810-28984 Incidental 1991 303 562 1993 3Y 122 649 20082

1930-72731 Incidental 1992 345 564 1996 5Y 373 516 5557

1930-72753 Incidental 1992 243 482 1994 3Y 389 531 15400

1960-03618 Incidental 1994 316 541 1996 3Y 344 489 5906

1960-03642 Incidental 1994 364 524 1996 3Y 363 533 906

1960-03660 Incidental 1994 303 601 1996 3Y 333 587 3311

1960-03683 Incidental 1994 318 540 1996 4Y 327 485 5573

1960-03697 Incidental 1994 122 648 1995 SY 123 638 1005

1960-03888 Incidental 1995 342 470 1996 SY 332 484 1720

1960-03899 Incidental 1995 345 479 1996 SY 245 482 10004

1810-28979 Search 1991 312 562 1992 SY UK UK 60

1960-03552 Search 1995 341 477 1996 SY 315 500 3471
USACERL TR-98/52 73

Appendix F: List of all HY Females

Resighted on Fort Hood.

This list is presented by band number. All females listed in this table were resighted

once. * = Female was banded and recaptured by MAPS field crew. Exact location

unknown

Band Banding Information Resighting Information

Number Year East North Year Age East North

Distance

(m)

1960-03185 1994 308 485 1995 SY 341 475 3448

1960-03215 1994 341 477 1996 3Y 342 475 224

1960-03236 1994 343 483 1995 SY 339 478 640

1960-03581 1995 341 479 1996 SY 342 488 905

1960-03897 1995 339 482 1996 SY 363 462 3124

2060-20812 1995 310 497 1996 SY 314 504 806

2060-20816 1995 310 497 1996 SY 310 497 *0
74 USACERL TR 98/52

Appendix G: Description of Palmer

Drought Severity Index

Source: National Oceanic and Atmospheric Administration, National Climate

Data Center, Asheville, NC.

Palmer Drought Severity Index (PDSI)

In 1965, Palmer developed an index to “measure the departure of the moisture

supply” (Palmer 1965). Palmer based his index on the supply-and-demand

concept of the water balance equation, taking into account more than only the

precipitation deficit at specific locations. The objective of the Palmer Drought

Severity Index (PDSI), as this index is now called, was to provide a

measurement of moisture conditions that were “standardized” so that

comparisons using the index could be made between locations and between

months (Palmer 1965).

The PDSI is a “meteorological” drought index and responds to weather

conditions that have been abnormally dry or abnormally wet. When conditions

change from dry to normal or wet, for example, the drought measured by the

PDSI ends without taking into account streamflow, lake and reservoir levels,

and other longer-term hydrologic impacts (Karl and Knight 1985). The PDSI is

calculated based on precipitation and temperature data, as well as the local

Available Water Content (AWC) of the soil. From the inputs, all the basic terms

of the water balance equation can be determined, including evapotranspiration,

soil recharge, runoff, and moisture loss from the surface layer. Human impacts

on the water balance, such as irrigation, are not considered. Complete

descriptions of the equations can be found in the original study by Palmer (1965)

and in the more recent analysis by Alley (1984).

Palmer developed the PDSI to include the duration of a drought (or wet spell).

His motivation was as follows: an abnormally wet month in the middle of a

long-term drought should not have a major impact on the index, or a series of

months with near normal precipitation following a serious drought does not

mean that the drought is over. Therefore, Palmer developed criteria for

determining when a drought or a wet spell begins and ends, which adjust the

PDSI accordingly. Palmer (1965) described this effort and gave examples, and it
USACERL TR-98/52 75

is also described in detail by Alley (1984). In near-real time, Palmer’s index is

no longer a meteorological index but becomes a hydrological index referred to as

the Palmer Hydrological Drought Index (PHDI) because it is based on moisture

inflow (precipitation), outflow, and storage only, and does not take into account

the long-term trend (Karl and Knight 1985). In 1989, a modified method to

compute the PDSI was begun operationally (Heddinghaus and Sabol 1991). This

modified PDSI differs from the PDSI during transition periods between dry and

wet spells. Because of the similarities between these Palmer indices, the terms

“Palmer Index” and “Palmer Drought Index” have been used to describe general

characteristics of the indices.

The Palmer Index varies roughly between -6.0 and +6.0. Palmer arbitrarily

selected the classification scale of moisture conditions (Table 23) based on his

original study areas in central Iowa and western Kansas (Palmer 1965). Ideally,

the Palmer Index is designed so that a -4.0 in South Carolina has the same

meaning in terms of the moisture departure from a climatological normal as a -

4.0 in Idaho (Alley 1984). The Palmer Index has typically been calculated on a

monthly basis, and a long-term archive of the monthly PDSI values for every

Climate Division in the United States exists with the National Climatic Data

Center from 1895 through the present. In addition, weekly Palmer Index values

(actually modified PDSI values) are calculated for the Climate Divisions during

every growing season and are available in the Weekly Weather and Crop

Bulletin. These weekly Palmer Index maps are also available on the World Wide

Web from the Climate Prediction Center.

The Palmer Index is popular and has been widely used for a variety of

applications across the United States. It is most effective measuring impacts

sensitive to the soil moisture conditions, such as agriculture (Willeke et al.

1994). It has also been useful as a drought monitoring tool and has been used to

start or end drought contingency plans (Willeke et al. 1994). Alley (1984)

identified three positive characteristics of the Palmer Index that contribute to

its popularity: (1) it provides decision makers with a measurement of the

abnormality of recent weather for a region; (2) it provides an opportunity to

place current conditions in an historical perspective; and (3) it provides spatial

and temporal representations of historical droughts. Several states, including

New York, Colorado, Idaho, and Utah use the Palmer Index as one part of

drought monitoring systems.

There are considerable limitations when using the Palmer Index, and these are

described in detail by Alley (1984) and Karl and Knight (1985). Drawbacks of

the Palmer Index include:
76 USACERL TR 98/52

• The values quantifying the intensity of the drought and signaling the

beginning and end of a drought or wet spell were arbitrarily selected based on

Palmer’s study of central Iowa and western Kansas and have little scientific

meaning.

• The Palmer Index is sensitive to AWC of a soil type. Thus, applying the index

for a Climate Division may be too general.

• The two soil layers within the water balance computations are simplified and

may not be accurately representative for a location.

• Snowfall, snow cover, and frozen ground are not included in the index. All

precipitation is treated as rain, so that the timing of PDSI or PHDI values may

be inaccurate in the winter and spring months in regions where snow occurs.

• The natural lag between when precipitation falls and the resulting runoff is

not considered. In addition, no runoff is allowed to take place in the model until

the water capacity of the surface and subsurface soil layers is full, leading to an

underestimation of the runoff.

• Potential evapotranspiration is estimated using the Thornthwaite method.

This technique has wide acceptance, but it is still only an approximation.

Several other researchers have presented additional limitations of the Palmer

Index. McKee et al. (1995) suggested that the PDSI is designed for agriculture,

but does not accurately represent the hydrological impacts resulting from

droughts of longer time scales. The Palmer Index is also applied within the

United States and has little acceptance elsewhere (Kogan 1995). One

explanation for this is provided by Smith et al. (1993), who suggested that it

does not do well in regions where there are extremes in the variability of rainfall

or runoff. Examples in Australia and South Africa were given. Another

weakness in the Palmer Index is that the “extreme” and “severe” classifications

of drought occur with a greater frequency in some parts of the country than in

others (Willeke et al. 1994). “Extreme” droughts in the Great Plains occur with

a frequency greater than 10%. This limits the accuracy of comparing the

intensity of droughts between two regions, as well as planning response actions

based on a certain intensity more difficult.
USACERL TR-98/52 77

Table 23: PDSI classifications for Dry and Wet Periods

4.00 or more Extremely wet

3.00 to 3.99 Very wet

2.00 to 2.99 Moderately wet

1.00 to 1.99 Slightly wet

0.50 to 0.99 Incipient wet spell

0.49 to -0.49 Near normal

-0.50 to -0.99 Incipient dry spell

-1.00 to -1.99 Mild drought

-2.00 to -2.99 Moderate drought

-3.00 to -3.99 Severe drought

-4.00 or less Extreme drought
78 USACERL TR 98/52

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