Title: Reproductive behavior and performance of the female Florida wild turkey /
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Title: Reproductive behavior and performance of the female Florida wild turkey /
Physical Description: vi, 95 leaves : ill. : ; 28 cm.
Language: English
Creator: Williams, Lovett E
Publication Date: 1985
Copyright Date: 1985
 Subjects
Subject: Wild turkey -- Behavior -- Florida   ( lcsh )
Wild turkey -- Nests -- Florida   ( lcsh )
Wild turkey -- Reproduction -- Florida   ( lcsh )
Forest Resources and Conservation thesis Ph. D
Dissertations, Academic -- Forest Resources and Conservation -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: Thesis (Ph. D.)--University of Florida, 1985.
Bibliography: Includes bibliographical references (leaves 90-94).
Statement of Responsibility: by Lovett E. Williams, Jr.
General Note: Typescript.
General Note: Vita.
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Bibliographic ID: UF00099344
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000502958
oclc - 22739851
notis - ACS2692

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REPRODUCTIVE BEHAVIOR AND PERFORMANCE OF
THE FEMALE FLORIDA WILD TURKEY









BY



LOVETT E. WILLIAMS, JR.


A DISSERTATION SUBMITTED TO THE GRADUATE SCHOOL OF THE
UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE
REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY



UNIVERSITY OF FLORIDA















ACKNOWLEDGMENTS

This study was funded by the Florida Game and Fresh Water Fish

Commission and the Federal Aid in Wildlife Restoration

(Pittman-Robertson) Program. Former Commission Directors A. D.

Aldrich and 0. E. Frye, Jr., and former Wildlife Division Director J.

A. Powell were particularly supportive as were R. M. Brantly, H. E.

Wallace, F. W. Stanberry, A. L. Egbert, and T. H. Logan.

Most of the field work was conducted on Lykes Fisheating Creek

Wildlife Management Area with the support and cooperation of C. P.

Lykes and B. Swendsen of Lykes Bros., Inc. Thanks are due also

Owen-Illinois, Inc., and their tract manager W. Schlitsgus, for the

use of their property on Lochloosa Wildlife Management Area.

H. Webster generously gave his time and expertise in developing

and constructing the electronic equipment used to remotely monitor

turkey nests. B. L. Akey, L. H. Barwick, W. Bess, T. A. Breault, D.

Z. Caudill, N. F. Eichholz, M. J. Fogarty, D. J. Forrester, W. B.

Frankenberger, H. L. Hill, L. T. Hon, R. C. McCracken, B. Morris, S.

Osceola, J. H. Peoples, R. W. Phillips, W. J. Sadinski, Jr., J. S.

Scanders, A. J. Wilson, J. Wilson, L. Zimmerson, and the late H.

Haywood helped collect field data while they were involved in the

project. J. W. Hardy allowed me to use the bioacoustics laboratory at

the Florida State Museum; T. A. Webber assisted me in making the sound

spectrograms. L. D. Harris of the School of Forest Resources and

Conservation provided assistance with some of the mathematical









calculations, and A. G. Hyde of the U.S. Soil Conservation Service

provided information about the soil types on the study areas. D. D.

Wackerly, C. L. Abercrombie, III, and N. R. Fuller provided

statistical advice and assistance. T. F. Crossman of the U.S.

Agriculture Stabilization and Conservation Service assisted in

measuring habitat acreages of the study areas.

Secretaries W. Circy, M. Shawver, and L. S. Sanders were of much

help at the Fisheating Creek Study Area, as were secretaries M. A.

Lansberry, W. Dunbar, and L. L. Davisworth at the Wildlife Research

Laboratory in Gainesville. S. Street and T. Crown typed the

manuscript. The maps and graphs were prepared by M. Moffitt, B.

Harrison, and R. Harrison. W. A. Greer of the Commission, Office of

Informational Services, was helpful in photo processing.

I would like especially to thank my co-workers D. H. Austin and

the late T. E. Peoples, who participated in every aspect of the field

work and who were responsible for the collection of data during my

absence from the study area.

Assistant Professor M. W. Collopy, Professor D. J. Forrester,

Professor J. W. Hardy, and Assistant Professor G. W. Tanner, who

served on my graduate committee in the School of Forest Resources and

Conservation (Institute of Food and Agricultural Sciences) at the

University of Florida, deserve my sincere appreciation. Professor

R. F. Labisky, my Committee Chairman, provided much encouragement,

advice, and assistance during the preparation of this dissertation.














TABLE OF CONTENTS


ACKNOWLEDGMENTS

ABSTRACT.

INTRODUCTION


STUDY AREAS

Fisheating Creek
Lochloosa.

METHODS .

RESULTS AND DISCUSSION


Egg Covering and Nest Construction
Laying Posture .
Multiple Nesting
Egg Dropping .
Nest Attendance During the Laying
Clutch Size
Renesting. .
Nest Disturbance by Man
Incubation Behavior .
Nest Attendance During the Incubat
The Incubation Period.
Hatching Behavior
Egg Hatchability
The Imprinting Period.
Time of Nest Departure by Broods
Hatching Synchrony
Attendance of Infertile Eggs
Nesting Habitat.
Nesting Seasonality .
Nesting Success and Predation
Defensive Behavior


S 1

3

3
. 10


17

22


Period




ion Period


SUMMARY .

LITERATURE CITED

BIOGRAPHICAL SKETCH









Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy



REPRODUCTIVE BEHAVIOR AND PERFORMANCE OF
THE FEMALE FLORIDA WILD TURKEY

By

Lovett E. Williams, Jr.

May 1985
Chairman: Ronald F. Labisky
Major Department: Forest Resources and Conservation

Two hundred forty-eight nests of 202 radio-telemetered wild

turkeys (Meleagris gallopavo osceola) were monitored in Florida during

the period 1968-1982 to determine nesting phenology and habitats,

clutch size, activity patterns, sensitivity to disturbance, predation,

renesting tendencies, and nesting success.

Nest initiation began when hens laid eggs, usually in mid-day, in

depressions scratched in the soil, and covered them with dry leaves.

Incubation began gradually after the fifth egg was laid; first laid

eggs experienced about 25 hours more incubation than last eggs. The

incubation period was 27 1.3 days. Hatching synchronization seemed

to be poorly developed. Fifty-eight percent of the nests were located

in saw palmetto (Serenoa repens), 31% in cypress (Taxodium distichum)

woods, and 11% in various "other" habitats. Nest predation was

greatest in cypress woods and least in the habitat category "other."

Fifty-seven percent of the hens whose nesting was disrupted

during the laying period renested; only 28% of those disrupted while

incubating renested. Renesting rates were 44% for adults and 22% for

yearlings; nearly one-half chose a different habitat for renesting.









Hatching required more than 1 day. Egg hatchability was 89.2%.

Sixty-eight percent of the broods departed the nest during morning

hours after sunrise.

The earliest clutch was initiated on 6 March and the latest

clutch hatched on 2 July. Yearlings began nesting later in the season

but finished earlier than adults. Adults laid 10.5 (SE = 0.16) eggs

per clutch and yearlings laid ]0.0 (SE = 0.28); the mean was 10.3 (SE

= 0.14).

Fifty-one percent of the hens flushed from nests did not return.

Sixty percent of the incubating hens and 38% of the laying hens

returned; 56% of the adult hens returned but only 27% of the yearlings

returned. Thirty-nine percent of the hens flushed from nests in

cypress woods returned, whereas 59% flushed from nests in palmetto

returned.

Approximately 55% of the nests were depredated. When renesting

was taken into account, nesting success was 58.7% for nesting hens.

Laying behavior, incubating activities, nest attendance patterns,

hatching vocalizations, defensive behavior, and hatching behavior also

are discussed.














INTRODUCTION

The wild turkey is endemic to North America. The turkeys were

reclassified (American Ornithologists' Union 1982) recently into the

family Phasianidae. The only other living species in the turkey

subfamily Meleagridinae is the ocellated turkey (Meleagris ocellata)

of southern Mexico and Central America.

The Florida subspecies of wild turkey (M. j. osceola) was named

for the Seminole Indian, Osceola, by W. E. D. Scott (1890) from type

specimens taken near Tarpon Springs. It is a clinal subspecies that

intergrades with the eastern form (M. 1. silvestris) in a zone from

southern South Carolina, across southern Georgia, northern Florida,

southern Alabama, southern Mississippi, to eastern Louisiana (Aldrich

and Duvall 1955).

Florida had a sizable turkey population in early historic times

(Wright 1915), but the species was eliminated by unregulated hunting

from some parts of the state by 1948 (Newman and Griffin 1950). In

the statewide restocking program that was initiated in 1951 (Powell

1965), turkeys were trapped from protected Florida populations and

released in suitable range that was not inhabited by turkeys.

Completion of the restocking program in 1970 gave Florida the

distinction of being the first state to complete a successful

statewide turkey restoration program.

Estimates of the fall population in Florida topped 100,000 in

1964. In 1966 the population and harvest plumetted due to unknown









causes, which resulted in considerable concern. By 1969 turkey

populations had recovered to pre-die-off levels, although there was

evidence that populations on certain wildlife management areas were

suffering from over-harvest during fall, either-sex hunting seasons

(Williams et al. 1978).

Curtailment of hen harvest on wildlife management areas, by

termination of fall hunting and expansion of spring gobbler hunting,

was being contemplated. Because there had been no research on turkey

ecology in Florida, effects of spring gobbler hunting on nesting

processes were unknown. Consequently, this study on nesting ecology

was initiated with emphasis on aspects that relate to possible effects

of spring gobbler hunting on reproduction. Specific objectives were

to characterize the seasonality of nesting, identify nesting habitats,

determine activity patterns of nesting hens, measure the sensitivity

of nesting hens to human disturbance, measure renesting tendencies,

and generally elucidate the behavioral characteristics of the hen

during the reproductive period.


I















STUDY AREAS

Field work was conducted on Lykes Fisheating Creek Wildlife

Management Area (WMA) during 14 of the 15 years from 1968 to 1982 and

on Lochloosa WMA during 1969, 1970, 1973 and 1975. The hunting rights

on both privately owned areas are leased to the Florida Game and Fresh

Water Fish Commission for public hunting.

Fisheating Creek

The Fisheating Creek Study Area (Fig. 1) is located in Glades

County, about 20 km west of Lake Okeechobec. The terrain is flat,

ranging between 9 m and 17 m above mean sea level. Soils are

predominantly sandy. The major soil associations are Fresh Water

Marsh and Swamp along the creek to about 1 km outward, Myakka-Pomello-

Basinger on the higher areas between the creek and palmetto prairie,

and Oldsman-Wabasso-Felda in the palmetto prairie (Florida Department

of Administration 1974).

The climate is subtropical. At Ft. Myers, which is approximately

81 km to the southwest (the nearest reference point for complete

weather data), the mean maximum and minimum daily temperatures in

April are 28.9 C and 16.3 C, respectively. Mean maximum and minimum

daily temperatures in July are 32.8 C and 22.9 C, respectively. Mean

annual temperature at Moore Haven, about 30 km to the southeast, is

22.8 C. The mean date of the first subfreezing temperature at Moore

Haven is December 25 and the mean date of the last winter freeze is

January 23 (National Oceanic and Atmospheric Administration 1978).











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Mean annual precipitation at Moore Haven is 127.9 cm, and at LaBelle,

about 25 km to the southwest, 132.5 cm. More than 60% of the annual

precipitation comes from thunder showers during the summer months.

Six plant associations with narrow, ill-defined ecotones between

them, were delineated at Fisheating Creek. Cypress woods, which

comprised 51% of the study area, occurred along the creek (Fig. 2).

The creek was subject to shallow flooding at least once annually.

Understory shrubs and midstory trees, other than young cypress, were

sparse; the ground cover of carpet grass (Axonopus compressus),

smartweed (Polygonum sp.), mistflower (Eupatorium coelestinum) and

other annual herbs became dense by May (Fig. 3).

Live oak trees (Quercus virginiana), which comprised 12% of the

study area, occurred in stands called hammocks (Fig. 4). Most

hammocks contained small clumps of saw palmetto and cabbage palm

(Sabal palmetto). A few hammocks contained laurel oak (Q.

laurifolia), red mulberry (Morus rubra), and hackberry (Celtis

laevigata) also. Trees in the hammocks were draped with Spanish moss

(Tillandsia usneoides) and other epiphytes.

Glades paralleled the cypress woods, forming zones of short

grasses (mainly Axonopus compressus) that were dotted with isolated

live oak trees and small live oak hammocks (Fig. 5). These open areas

were grazed heavily by cattle and had the appearance of semi-improved

pastures. Approximately 17% of the study area was glades.

Broadleaf evergreen shrubs (e.g. Myrica cerifera), evergreen

trees (Gordonia lasianthus, Magnolia virginiana, and Persea spp.), and

vines (Vitis sp. and Smilax spp.) existed in bay heads and swamps

which comprised 2% of the study area.


























































Figure 2. Aerial view of cypress woods on the Fisheating Creek
Study Area in spring.

























































Figure 3. Inside the cypress woods at Fisheating Creek Study
Area in May.




































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Bayheads and swamps often contained surface water and supported dense,

woody, understory vegetation; the soils contained a relatively high

proportion of humus and litter.

Oak scrub, which covered 12% of the area, is an association of

small, mostly evergreen oaks (Q. chapmanii, Q. myrtifolia, Q.

geminata, and Q. inopina) and other short, woody vegetation (e.g.

Lvonia ferruginea, Befaria racemosa, Ilex opaca var. arenicola) on

sandy soils that lie about 13 m above mean sea level (Fig. 6). Sand

pine (Pinus clausa) was not present.

The prairie is a wide, flat expanse of saw palmetto and wire

grass (Aristida stricta), dotted with widely-spaced pine trees (Pinus

palustris and P. elliottii) and small islands of oak scrub. The

prairie extended for many miles to the north and south of the

woodlands along Fisheating Creek and was a travel barrier to turkeys.

It was not considered part of the study area except for the ecotones

between palmetto prairie and other plant associations. This ecotonal

zone was about i-km wide and comprised approximately 5% of the study

area.

Small patches of other plant associations, creeks, ponds,

sloughs, and marshes, improved pastures for cattle, and roads made up

the remaining 1% of the study area. The major plant associations

formed zones approximately parallel to Fisheating Creek (Fig. 7).

The owners ran cattle on the area and feral hogs were present.

No special land management practice for turkeys was used on the area.

Lochloosa

Lochloosa WMA is located principally in Alachua County of

northern Florida, about 350 km north of Fisheating Creek WMA. The




























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Figure 7. Aerial view of a typical cross-section of the Fisheating
Creek Study Area showing cypress woods (top), a glade parallel to
the creek, a live oak hammock adjacent to the glade, saw palmetto
prairie (left middle), and oak scrub (lower middle).
prairie (left middle), and oak scrub (lower middle).









major land owner, Owen-Illinois, Inc. (Forest Products Division),

managed the property primarily for pulpwood. The study was conducted

on a sector that comprised about 3,200 ha of the 12,500 ha WMA (Fig.

8).

The terrain is flat to slightly rolling, ranging 17 m to 30 m

above mean sea level. Soil types on the sand ridges are Millhopper,

Tavares, and Newnan sands. In the ponds and swamps, the main soils

are Monteocha loamy sand, Pomona sand, and Samsula muck, whereas the

main flatwoods soil is Pomona sand (U.S. Department of Agriculture

1982).

The mean annual temperature is 22.20 C at Gainesville, 15 km

northwest of the area. The mean date of the first and last frost is

December 6 and February 14, respectively. Mean annual rainfall is

133.2 cm, with more than 50% occurring in summer (National Oceanic and

Atmospheric Administration 1978).

The original upland plant associations were mainly longleaf pine

(Pinus palustris) and turkey oak (Quercus laevis), and slash pine (P.

elliotti) flatwoods (U.S. Department of Agricuture 1980). These

associations were replaced mostly by slash pine plantations during the

1950's. Areas planted were first chopped, burned, and then machine

planted at a density of 764 seedlings per ha. Pine woods comprised

approximately 50% of the area. About 9% of the study area was in

natural slash pine stands. Many of the large live oak hammocks had

been eliminated by commercial forest management practices. A hardwood

control program implemented during the 1950's was not entirely

successful; consequently, the slash pine plantations had nearly as












much young live oak and laurel oak as pine (Fig. 9). The planted pine

and oak association occupied about 15% of the study area.

Pinelands in the study area were logged, primarily by

clearcutting, at various times during the study making it difficult to

meaningfully catagorize the resulting associations and stages of plant

succession.

Recently clearcut pinelands covered approximately 7% of the area

and improved cattle pastures, 2%. Cypress heads and hardwood swamps

occupied 10% of the area and open ponds comprised about 1%. Approxi-

mately 2% of the area was in hardwood hammock; seasonally wet prairies

made up the remaining 4%. There were no palmetto flats or grazed

glades, as described for Fisheating Creek, but saw palmetto occurred

abundantly. The area was grazed by cattle; feral hogs occurred in

small numbers.

The wild turkey was eliminated by overhunting in the area that is

now in the Lochloosa WMA between 1900 and 1950. The area was

restocked with wild-trapped birds from Fisheating Creek WMA in the

early 1960's. No other turkey management, except enforcement of

hunting regulations, had been practiced. The area was open to

year-round public use. The spring gobbler season consisted of 16

half-days of hunting in late March during the study period.













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Figure 9. Thirty-year-old planted slash pine woods with oak
regeneration on Lochloosa Study Area, Alachua County, Florida,
1980.















METHODS

Hens were captured with cannon nets (Austin 1965), rocket nets,

orally administered tribromoethanol (Williams et al. 1973), and

alpha-chloralose (Williams 1966). Age classes of the hens were

determined by the configuration of the greater upper secondary coverts

(Williams 1961). All birds were weighed and banded prior to being

radio-instrumented. Turkey handling procedures have been described

previously (Austin et al. 1973).

Radio transmitters were attached to 414 hens, 35 on the Lochloosa

Study Area and 379 on the Fisheating Creek Study Area. Data were

obtained from 248 nests of 202 hens that nested one or more times.

Because of the small number of nests (N = 15) on the Lochloosa Study

Area, no attempt has been made to contrast the turkey populations on

the two study areas.

Transmitters were spaced across the frequency band (150.815 MHz

to 151.210 MHz) to provide 24 channels separated by 10 to 15 KHz.

Transmitters weighed from 65 g to 90 g, measured approximately 50 mm x

25 mm x 80 mm, and met or exceeded the performance requirements of 2-

km range and 6-month signal transmission without battery change.

Transmitters were fitted to the turkeys by underwing loops of

latex surgical tubing (3/32-inch wall, 1/16-inch inside diameter).

One loop was tied under each wing with a square knot or single beckett

bend. Tubing was attached to transmitters with fiberglass tape. The













Table 1. The number of Florida wild turkey hens instrumented and
monitored, Fisheating Creek and Lochloosa study areas, 1968-82.



Found dead Contact lost Monitored
before before during
Number nesting nesting nesting Nests
Year instrumented season season season found


1968 30 0 0 30 20

1969 26 0 4 22 14

1970 34 0 3 31 19

1971 33 5 4 24 14

1972 35 2 6 27 14

1973 30 1 0 29 20

1974 33 1 4 34 25

1975 25 1 2 22 20

1976 20 0 3 17 12

1978 18 0 1 13 12

1979 21 3 1 17 13

1980 43 5 8 30 28

1981 54 9 4 41 32

1982 12 1 3 8 5

Total 414 28 43 345 248

aNot including those found dead.









hens were not noticeably hindered by the transmitter package and those

recaptured showed no evidence of serious chafing.

Field monitoring was accomplished with 24-channel, crystal

controlled, portable radio receivers. Receiving antennas included

1/4-wave whips on trucks, hand-held yagis for work on foot, and large

multi-element directional antennas mounted on trucks. Nests were

found by radio signals when instrumented hens were present. Data were

collected at nests while the hens were absent.

Electronic devices were used to record nest attendance of the

hens. The low-power signal from the transmitter on a hen was

monitored by a battery-powered receiver hidden near the nest. This

signal was re-transmitted to the field station on a different

frequency by a directional antenna; maximum transmission was 8 km. The

sensitivity of the receiver at the nest was adjusted to restrict the

radius of signal reception to within about 1 m, which made the

equipment function as a proximity detector. At headquarters, the

incoming signal activated an electronic switch and was recorded on a

30-day time-calibrated Esterline-Angus event recorder, thereby making

a continuous record of the time the hen was on the nest. A maximum of

20 nests could be monitored simultaneously. Nests were monitored

approximately 8,000 hours by automatic recorders and about 400 hours

manually. Visual observations confirmed the reliability of the nest

monitors.

Behavior of hens was directly observed with spotting telescopes

and binoculars from portable cloth blinds located about 30 m from the

nests. Microphones were placed within 1 m of seven nests to monitor

and record the sounds of hatching on a 1/4-inch open reel Uher tape









recorder. Sound spectrograms of recorded hen and poult vocalizations

were made on a Sona-Graph model 7029A instrument (Kay Electric

Company).

All vegetation within 1.5 m of each of 57 nests found on the

Fisheating Creek Study.Area during the 5 years, 1968-1972, was

identified and its coverage estimated visually to the nearest 5%;

percent coverage of overhead vegetation within 2 m above each nest was

visually estimated. The habitat within 45 m of 236 nests found in the

study was classified by the most abundant vegetation present--

palmetto, cypress woods, or other. Habitat acreages were measured

from aerial photographs (scale 600 feet = 1 inch) with a Model

1211-H-1 Nemonics Corporation electronic digital planimeter.

The Mayfield (1961) method, which utilizers data from nests that

were observed during only part of the laying or incubation period, was

used to calculate nesting success. The Wilcoxon Rank Sum test was

used to compare clutch sizes of first, second, and third nests and of

nests in which incubation began before 1 May, between 1 May and 20

May, and after 20 May. Chi-square analyses were used to test for

differences in renesting tendencies of hens whose nests were disrupted

during the laying vs the incubating periods; differences in the

proportions of nests located in the three habitat types and the

success and predation rates in each habitat type; differences in

predation rates of clutches that hatched before 1 May, between 1 May

and 20 May, and after 20 May; differences in the tendencies of hens to

return to their nests when flushed; differences in return-after-

flushing tendencies of hens nesting in three habitat types; and

differences in the proportion of yearling vs adult hens that abandoned






21


their nests after being flushed. The t-test procedure was used to

compare renesting rates of adult and yearling hens and mean length of

recesses of incubating hens. The Kolmogorov-Smirnov Z test (Zar 1974)

was used to test for differences in the distribution of seasonal

nesting curves for adult and yearling hens. Fisher's least

significant difference test (SAS Institute Inc. 1982) was used to

compare length of recesses during four segments of the period of

continuous incubating behavior. The confidence interval on the

proportion of recesses hens took in the afternoon followed the form

x t c/2, dfSE(x)















RESULTS AND DISCUSSION

The mating system of the Florida turkey fits Oring's (1982)

description of male-dominance polygyny with intermediate dispersal.

Gobbling and strutting activities by males in early February, before

the hen flocks dissolved, indicated that gobblers were receptive to

mating earlier than hens. Hen flocks dissolved in March prior to the

onset of nesting. Hens, upon attainment of sexual receptivity,

visited the gobblers for mating.

Before nesting, hens established new home ranges and often

roosted alone in small, isolated hammocks and bay-heads where gobblers

and non-nesting hens seldom ventured and rarely roosted (Williams et

al. 1974). Hens frequently were seen traversing the edge of saw

palmetto prairies and the oak scrub-palmetto ecotone as though

searching for nesting sites. Tracking effort was concentrated in

these areas so that nests could be found early in the laying cycle.

Egg Covering and Nest Construction

Four hens were monitored by telemetry as they established their

nests and laid their first eggs. When a hen approached the nesting

area she would spend approximately 5-20 minutes moving in a restricted

area before becoming still. This behavior pattern suggests that she

was in the process of selecting a place for the nest.

The deposition of the first egg followed a definite pattern.

The hens scratched shallow depressions in the soil, laid the egg,

placed a few dried leaves over the egg, and departed. The mean length









of time spent on the nest while laying the first egg was 70 minutes

(N = 4, SE = 28.4). Freshly laid eggs were clean and chalky, with a

thin layer of sand adhering only to the side in contact with the

ground.

Five additional nests were found at the time that the second or

third egg was being laid. In two of these nests, shallow

scratched-out depressions were found within 10 m of the nests in what

appeared to be suitable nesting places, indicating that these hens had

scratched shallow depressions in more than one place before selecting

the place they would lay.

Evidently, hens did not transport nesting material to cover their

eggs, but rather used debris present at the site. Two hens observed

while laying covered their eggs with plant debris picked up from

beside their nests; no hen was observed carrying nesting material.

About a century ago, an observer reported to Bendire (1892) that

Florida turkey nests were lined with dead leaves and grass that were

so like the surrounding debris that he wondered whether the material

was placed there by the hen or was already present under the eggs.

Although numerous patches of bare ground were available in the

vicinity of most nests, only a single nest was established on such a

site. The availability of nearby dried plant material may be a factor

in nest site selection.

Twenty nests that were observed a total of 90 times during the

laying period were covered sparsely with dried leaves. Approximately

160 different nests were inspected at least once while the hens were

on recess during the incubating period; none of these nests was

covered. These observations indicate a tendency for hens to cover









their eggs while away during the laying period but to leave them

uncovered while on recess during the incubation period. Although many

writers (e.g., Audubon 1831, McIlhenny 1914, Mosby and Handley 1943,

Bailey et al. 1951) have stated that turkey hens cover their eggs with

leaves, they did not specify that this occurred only during the laying

period. Green (1982) reported that hens in Michigan did not cover

their eggs before taking incubation recesses.

Camouflage, rather than insulation, appears to be the function of

egg covering by the turkey. If insulation were required, it would

be needed also during the incubation period when embryos are more

vulnerable to chilling; however, turkeys do not cover the eggs during

incubation recesses. Furthermore, the fact that hens use only debris

from beside the nest to cover their eggs suggests that the function is

to blend the nest with the immediate surroundings.

Egg covering also is the means by which a turkey nest is

constructed. When the hen returns to the nest to lay each egg, she

does not uncover the nest before laying, which causes more debris to

accumulate with the laying of each egg. When the hen turns the eggs,

the debris settles to the bottom and sides of the nest and by the time

the last egg is laid, the nest depression is well lined with leaves.

A typical nest measures 2 cm deep, 20 cm wide, and 24 cm long

(Williams et al. 1968).

Laying Posture

Three hens were observed laying a total of 11 eggs. Hens sat on,

or crouched over, their nests before laying. The laying of the egg

was accomplished from a partially erect body position. They trembled









as they laid, with wings drooped slightly and tails raised. The back

feathers were ventilated. Eggs were laid on the ground beside the

other eggs and not on top of them. Of the more than 2,000 turkey eggs

examined, only two were cracked in a manner that would suggest one egg

had struck another while being laid.

Multiple Nesting

Some bird species, for example the ring-necked pheasant

(Phasianus colchicus), have very large clutches when more than one hen

lays in the same nest. Such multiple nesting has been reported for

the wild turkey (Bent 1932, Mosby and Handley 1943). However, the

only previous evidence of multiple nesting is Bendire's (1892) report

of a turkey hen seen on a nest while another hen was seen standing

close by and presumed to be waiting to lay in the same nest; Audubon

(1831) also mentioned three hens on the same nest.

In the present study, two hens were observed visiting the same

nest on three occasions and both were seen sitting on the nest

although never at the same time. Telemetry data indicated that these

hens did not associate with each other. More than one egg per day was

laid in the nest on two occasions and it seems almost certain that

both hens laid in this nest. The nest was deserted after the twelfth

egg was laid.

Another nest was photographed after the hen had been flushed

deliberately. When the nest was examined about 2 hours later, one egg

had been added. The setting hen had passed the twentieth day of

incubating behavior; therefore, the extra egg must have been laid by

another hen.









The potential for multiple nesting exists in Florida turkey

populations; however, it is probably not very common. If multiple

nesting depends upon chance encounters, it should be expected to be

more prevalent when nest density is higher, which would partly explain

the much higher incidence of multiple nesting observed in the

ring-necked pheasant (Labisky 1968).

Egg Dropping

Some birds occasionally lay single eggs where there is no nest.

Stoddard (1931) reported this phenomenon as common in the northern

bobwhite (Colinus virginianus). Single eggs that were never incubated

constituted a minimum of 8% of the annual egg production of

ring-necked pheasants in Illinois (Labisky 1968). In the present

study, only seven turkey eggs not associated with a nest were observed

in approximately 30,000 man-hours of field work during the nesting

season, indicating that egg-dropping is not prevalent among Florida

turkeys.

Nest Attendance During the Laying Period

Blakey (1937:7) provides the only reference about the pattern of

egg laying by the wild turkey by saying "Study of artificially

propagated wild turkey shows that, when . [egg laying] . .

becomes regular, one egg is laid daily, approximately one hour later

each day, beginning at about one hour after sunrise and continuing

until sundown terminates the cycle. Then the hen may skip a day and

begin over again at the early morning hour and repeat the cycle."

In four nests monitored from the deposition of the first egg, the

hens came to lay between 1120 hours and 1300 hours (x = 1234 hours).

In all laying events observed, only 8 of 74 eggs were laid before 1100









hours. Two of the four hens that were monitored from the laying of

the first egg skipped laying the day after laying the first egg; one

of these hens skipped laying also the day after laying the second egg.

The other two hens that were monitored from the laying of the first

egg skipped laying early in the laying period--one skipped the day

after the third egg and the other skipped after laying both the second

and third eggs. In a total of 103 laying events by 22 hens, the only

additional skipped days observed were after the first, fourth, and

ninth eggs. Thus, hens did not exhibit a regular pattern of skipping

a day in laying after the fifth egg of the clutch had been laid.

To test the hypothesis that wild turkeys laid approximately 1

hour later each day, the time of laying of 29 pairs of consecutively

laid eggs was compared. This comparison revealed that 50% were laid

within the same hour as the previous egg, 17% were laid at least 1

hour earlier, and only 24% were laid 1 hour later. Furthermore, in

65% of the laying events monitored, hens came to the nest to lay

between 1000 and 1500 hours. This pattern of laying in mid-day

compares closely with the pattern reported for the domestic turkey

(Stockton and Asmundson 1950) but not with the pattern reported for

wild turkeys by Blakey (1937).

In 74 observations of laying events in 22 nests, only 2 days were

skipped after the fourth egg was laid, and in neither of these cases

was laying resumed in the early morning of the following day. Thus

there was no evidence that early morning laying followed skipped days

late in the laying period.

In summary, the first egg of the Florida turkey is usually laid

in mid-day and is followed by a lapse day; the second egg is laid on









the third day and is sometimes followed by a lapse day. A few hens

skip laying one additional day after the third egg, but very few

lapses occur later in the laying period. The fourth through final

eggs are usually laid on consecutive days with a tendency to lay a few

minutes later and to remain on the nest longer each day. The clutch

is typically completed in late afternoon (Fig. 10). Hens failed to

lay only 4 of 76 times they were observed visiting their nests during

the laying period.

Hens remained on their nests an average of 55 minutes while

laying each of eggs one through five. The apparent decrease (X = -7.3

minutes, SE = 11.3) in mean attendance during the laying of each of

the first five eggs was not significant (P = 0.531) (Fig. 11). Hens

remained on their nests an average of 50 minutes (SE = 12.8) longer as

they laid each of eggs 5 through 12. Hens were remaining on the nest

about 348 minutes with the laying of the eleventh egg. Thus,

incubating behavior began gradually with the laying of the sixth egg

and the length of incubating sessions increased with the laying of

each successive egg. The first egg was subjected to about 25 hours of

incubation, on the average, before the twelfth, or last, egg was laid.

No hen began continuous incubating behavior with fewer than five

eggs. This finding, coupled with the observed tendency of hens to

begin incubating with the laying of the sixth egg, suggested that at

least five eggs are required to stimulate the gradual onset of

incubating behavior. This hypothesis was tested by manipulating eggs

in three nests during the laying period. In one nest, four eggs were

removed from a clutch of seven, and then one egg was removed each day

to maintain the clutch at three eggs. The hen abandoned this nest























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after laying the ninth egg. In another nest, three of six eggs were

removed at the time the nest was found; this clutch also was

maintained as a three-egg clutch. This hen abandoned the nest after

laying 12 eggs without initiating continuous incubating behavior. The

electronic monitor indicated that the hen did not exhibit the normal

pattern of gradually lengthening incubating behavior late in the

laying cycle; she remained at the nest no more than 34 minutes while

laying any egg, thereby demonstrating behavior typical of a hen in the

laying cycle. Another nest was found with four eggs and the clutch

was held at that number by daily removal of eggs during the remainder

of the laying period. This hen abandoned the nest after laying the

ninth egg. The abandonment of the three manipulated clutches supports

the hypothesis that approximately five eggs in the nest are required

to stimulate the gradual onset of incubating behavior.

If the turkey were a so-called "indeterminant" layer (Cole 1917),

these hens would have laid a greater number of eggs than represented

in normal clutches. The result indicates that the wild turkey is a

"determinant" layer and will not continue to lay indefinitely merely

because its eggs are removed as they are laid.

The gradual onset of incubating behavior is stimulated by five

eggs, probably through visual or tactile cues. Termination of laying

occurs several days after the onset of gradual incubation, probably

caused by increasing prolactin secretions as incubating behavior

progresses (Eisner 1958). The daily increase in the secretion of

prolactin would effectively determine the clutch size in the species

by leading to the termination of laying.









Clutch Size
A clutch is complete when no additional eggs are laid. Clutch

size averaged 10.3 eggs in 179 complete clutches (Table 2). Clutch

size of yearling hens, which averaged 10.0 differed significantly

(Wilcoxon two-sample test, S = 2248, Z = 1.6245, P = 0.104) from those

of adult hens, which averaged 10.5 eggs. Sixty-seven percent of the

complete clutches had 9, 10, or 11 eggs; the modal clutch size was 10

eggs (Fig. 12). No complete clutch contained fewer than five eggs.

Mean clutch size did not vary significantly among years (F = 1.50, df

= 178, P = 0.12).

The Wilcoxon Rank Sum test indicated that mean clutch size of the

first (x = 10.4, N = 150), second (x = 10.2, N = 26), and third (R =

9.8, N = 4) nests of the same hens within the the same year did not

differ significantly (first vs. second: Z = 0.088, P = 0.93; second

vs. third, Z = 0.751, P = 0.45; first vs. third: Z = 0.836, P =

0.40). Mean clutch size among nests in which incubating behavior

began before 1 May (x = 10.2, N = 99), between 1 May and 20 May (x =

10.0, N = 33), and after 20 May (x = 9.6, N = 8) did not differ

significantly (Kruskal-Wallis X2 = 1.25, df = 2, P = 0.54).

Mean clutch sizes of the wild turkey in other regions of the

United States indicated that clutches in Florida populations may be

smaller than in eastern wild turkey populations in Alabama,

Mississippi, Missouri and Virginia, but larger than those in

populations of the Rio Grande turkey in Texas (Fig. 13). Comparisons

using the Student's t-test indicated no significant difference in mean

clutch sizes between Florida and Mississippi (t = 1.15, df = 189, P >

0.01) or Florida and Texas (t = 0.73, df = 213, P > 0.01). The









33


















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clutch size reported in one of the Virginia studies (McDowell 1956)

was larger than in the present Florida study (t = 2.77, df = 211, P <

0.01). Mean clutch size in the turkey may be under genetic control

and may vary among populations, but the genetic component is

outweighed by the combined effects of experimental error and small

sample size in the data presently available from other regions.

Renesting

Hens carrying transmitters emitting weak signals and those that

moved away from the study areas were not monitored closely. Some of

these hens probably had their nests destroyed by predators without

the nests being detected in the study. Consequently, the renesting

statistics are minimum estimates.

Nearly fifty-seven percent (56.6%) of 30 hens that had their

nests disrupted during the laying period renested, whereas only 28% of

93 hens renested after their nests were disrupted during the period of

continuous incubating behavior (N = 123, X2 = 8.223, df = 1, P =

0.004) (Fig. 14). No hen renested after incubating longer than 18

days. Adult hens renested with greater frequency than yearlings; 44%

of 80 adults nested at least twice in the same year whereas only 22%

of 23 yearlings renested (t = 2.1, df 1, P < 0.05) (Fig. 14).

Nest Disturbance by Man

Sixty-two percent of 38 hens that were flushed from their nests

during the laying period did not return. Since 43% of 30 hens whose

nests were disrupted did not renest, the loss to the population of

potential reproduction was 0.27 (0.62 x 0.43) nests per hen flushed

during the laying period. During the incubation period, 40% of 38

hens that had been flushed abandoned their nests and 72% (N = 93) of























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them did not renest, representing a loss of approximately 0.29 (0.40 x

0.72) nests per hen flushed. Thus, it would make little difference

whether a hen were flushed during the laying or the incubation period

with respect to the net impact on annual reproduction. Five (2%) of

218 nests were deserted for no apparent reason.

Incubating Behavior

The term "incubation" as commonly used has two meanings, namely

1) the behavior of the hen sitting on or standing over the eggs and 2)

the embryonic developmental processes that take place inside the egg

shell. These components are independent--a hen can carry on

incubating behavior whether or not embryonic development takes place

(as with infertile or artificial eggs) and the embryo can develop

without a parent incubating it (as in artificial or natural

incubators). When the term incubation is applied to the behavior of

the hen, it involves a presumption that embryonic development is

occurring in the eggs, which may not be the case. Therefore, the

sitting of the hen would better be called "incubating behavior" to

distinguish it from the incubation process that occurs within the egg-

shell. In the Florida turkey, incubating behavior consists of

gradually lengthening afternoon incubating sessions after the fifth

egg is laid, followed by continuous day and night incubating sessions

upon completion of the clutch.

Twenty-four hens began incubating overnight on either the day

before laying the final egg, the same day of laying the final egg, or

the day after laying the final egg (Fig. 15). Two eggs were laid in

4 of the 24 nests after continuous incubating behavior began. Of

these eight additional eggs, only one was left unhatched despite





















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the fact that the last laid egg in each nest had been incubated at

least 2 days less than the rest of the clutch.

Stoddard (1931) stated that the northern bobwhite leaves its eggs

unincubated for as long as 1 week after the last egg is laid. No

turkey clutch went unincubated for as long as a whole day after the

last egg was laid. This behavioral trait of the turkey would minimize

the risk of nest predation by lessening the time the nest is exposed

to predators.

Approximately 400 man-hours were spent observing nine different

hens on their nests during continuous incubating behavior. While

sitting, the hens' body parts were positioned as in standing except

that their legs were folded at the upper tarsal joints forward under

the breast. While sleeping in the nest during daylight hours, the

head and neck were drawn in, the eyes were closed, and the wings

drooped along the sides.

When turning eggs, hens crouched by flexing the intertarsal

joints; they did not stand erect. Several eggs were usually

rearranged with a single motion of the head but the activity seemed to

be directed at only a single egg at a time. The motion of the head

not only turned the eggs, but also repositioned them in the nest so

that no egg remained in the same part of the nest for more than a few

hours at a time. Hens often arose and gazed down at their eggs for a

few seconds, and sometimes settled back without turning them. They

usually gazed intently before turning an egg--the gazing behavior

seemed to involve some cue that led to egg turning.









While standing in the nest, the bird's body posture was the same

as when it stood at other times. Hens settled back on the nest after

standing or crouching by moving the body forward with an upward

swinging motion that placed some of the breast feathers over the eggs

in front of the hen, thereby covering all the eggs.

As air temperatures increased during late morning, incubating

hens began to pant, with partly open bills, and sometimes ventilated

back feathers. Panting became faster, with the bill opened wider, as

mid-day temperatures increased. The sequence was reversed as

temperatures decreased in the afternoon. Hens would move their heads

or change positions to avoid spots of direct sunlight that penetrated

the overhead vegetation.

Two incubating hens retrieved single eggs that had rolled from

their nests; however, the retrival process was not observed. Two hens

deserted their nests when most of their eggs had been rolled out

intact by the rooting of armadillos. One hen continued to incubate a

clutch containing one broken egg; 12 hens deserted nests that

contained one or two broken eggs. It could not be determined whether

desertion was due to the presence of the broken eggs or to the

disturbance that caused the eggs to be broken.

Nest Attendance During the Incubation Period

Incubating hens recessed for 98 minutes at a mean interval of 1.9

days (Table 3). Four monitored hens remained continuously on their

nests for 3 days; the longest period of uninterrupted sitting was 4

days. A few hens recessed twice each day on several days. The period

of nest attendance immediately preceding hatching averaged 2.4 days (N

= 5).















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There was a tendency for a given hen to recess in morning or

afternoon on several consecutive days (Table 4). Consecutive daily

recesses were during the same morning, noon, or afternoon period 32

times and during a different period 30 times. None of the five hens

observed completed the-entire incubation period without changing

recess patterns from morning or noon to afternoon, or vice versa, at

least once. Thus hens do not recess at the same time of day

throughout the duration of the incubating activity.

Thirty-nine percent of 67 recesses by five hens began before noon

and 61% began after noon (Table 4), indicating a tendency for the hens

to take more recesses in the afternoon. Only 16% of the recesses of

these five hens included noontime (1200 hours).

Blakey (1937) reported that turkey hens tended to recess at

mid-day after the first few days of incubating, and Hillestad (1970)

stated that mid-day would be the best time for hens to recess because

air temperatures then most nearly approach required incubation

temperatures. The mid-day recess pattern was not predominant in this

Florida population. Only 10% (27) of 271 recesses occurred between

1130 and 1230 hours. A similar tendency for hens to avoid recessing

at mid-day was noted in a recent study in Michigan (Green 1982).

Recesses were less frequent in early afternoon than during late

afternoon (Fig. 16).

In Florida, the traditional closure of spring gobbler hunting

each day in early afternoon probably lessens the risk of incubating

hens on recess being shot by hunters.

The period of absence from the nest for hens that recessed at

noon averaged 137 minutes, whereas hens on recess in the morning









Table 4. Consecutive nest recesses of five Florida wild turkey hens
monitored during the period of continuous incubating behavior.
Fisheating Creek and Lochloosa study areas, 1968-82.



Length of Day of
Period of day recess incubating
of recess (minutes) Date behavior


BAND 255R


Afternoon
Afternoon
Noon
Morning
Morning
Afternoon
Afternoon
Afternoon
Noon
Afternoon
Morning
Morning
Morning


BAND 2875


April
April
April
April
April
April
May
May
May
May
May
May
May


Afternoon
Afternoon
Morning
Morning
Noon
Morning
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon
Noon
Afternoon



Afternoon
Afternoon
Noon
Morning
Noon
Afternoon
Noon


BAND 484R


19 April
20 April
21 April
22 April
23 April
24 April
26 April









Table 4, continued.



Length of Day of
Period of day recess incubating
of recess (minutes) Date behavior


Noon
Afternoon
Morning
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon



Afternoon
Morning
Afternoon
Morning
Morning
Afternoon
Noon
Afternoon
Afternoon
Afternoon
Morning
Morning
Afternoon



Noon
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon
Afternoon
Noon


BAND 485R


April
April
May
May
May
May
May
May



April
April
April
April
April
April
April
April
April
April
April
May
May


BAND 487R


aMorning recesses ended before noon; noon recesses began before and
ended after noon; afternoon recesses began after noon.


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stayed away 129 minutes; the difference was not statistically

significant (P >0.05). Mean noon recess period of 137 minutes was

greater than the afternoon period of 91 minutes (Z = 2.03, P = .021).

The mean length of recesses for 128 adult and 32 yearling hens of 106

and 103 minutes, respectively, did not differ significantly (t =

0.097, P > 0.50). Recesses averaged 109, 95, 84, and 140 minutes

during the first through fourth weeks of continuous incubating

behavior, respectively (Table 5). The percentage of time hens spent

off their nests was greatest (10%) during the final week of

incubation.

Green (1982) reported a mean recess time of 53 minutes for four

hens in Michigan (N = 41). The mean for 67 recesses of five Florida

hens was 95 minutes. The difference between the length of recesses in

Florida and Michigan was highly significant (t = 4.24, df = 113, P =

0.001). The Michigan study was conducted on a stocked population more

than 100 km north of the northern limit of the turkey's range in

Michigan; the present study was at the southern limit of the wild

turkey's range in the eastern U.S. The longer recesses in Florida may

be an adaptation to the warmer air and soil temperatures in Florida

where the eggs would not cool as rapidly in the hen's absence as they

might in Michigan.

The Incubation Period

The periods from the beginning of continuous incubating behavior

until the first poult hatched and until the brood left the nest

averaged 26 days (N = 7) and 27 days (N = 8), respectively (Table 6).

Healy et al. (1975) reported a mean incubation period of 28.6 days for

captive wild turkeys, but did not define the period or note the


















Table 5. Average length of Florida wild turkey hen nesting recesses
during the period of continuous incubating behavior, Fisheating Creek
and Lochloosa study areas, 1968-82.



Interval of
incubating Mean length Mean percentage
behavior Sample of recesses of time off
(days) size (minutes) nest


1- 7 38 109 8

8-14 35 95 7

15-21 48 84* 6

22+ 37 140* 10


*The difference between
significant difference
0.05).


these means was significant (Fisher's least
test, df = 154, t = 1.98, LSD = 55.66, P =





















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occurrence of gradual onset of incubating behavior. In artificial

incubators, domestic turkey eggs require 26.92 days to hatch and

another 0.33 day for the poults to dry, which totals 27.25 days

(Abbott and Craig 1960). This latter figure does not provide for the

several hours required.for imprinting in wild broods and therefore

would be more comparable to the 26-day period required for the first

poult to hatch in wild Florida turkey nests.

The 27-day period of continuous incubating behavior for Florida

turkey hens measured in the present study was briefer than the 28 days

reported by Mosby and Handley (1943) or the 27.3 days (Abbott and

Craig 1960) and 28 days (Marsden and Martin 1949) reported as the

incubation period of domestic turkey eggs in incubators. The reason

for the disparity in incubation periods is that Florida wild turkey

eggs are incubated for approximately 25 hours during the laying period

before continuous incubating behavior begins. No incubation occurs

during the laying period when artificial incubators are used because

the eggs are removed from the nests and stored at cool temperatures

until they are placed in the incubator.

Little published data exist concerning the time required for

whole clutches of wild turkey eggs to hatch after the first egg has

pipped. Most writers, such as Mosby and Handley (1943), state that

the hatching of the entire clutch requires about 24 hours. Healy et

al. (1975) reported that an artificially incubated wild turkey egg

hatches within an interval of 4 to 21 hours, that whole clutches hatch

in 12 to 48 hours and that 8 hours are required for a poult to become

dry. Cook (1972) reported that two wild clutches hatched in 23.3 and

26.2 hours. In the present study, the interval between the time the









first egg of a clutch was observed pipping and the time at least one

poult was seen hatched ranged from less than 1 hour to 52 hours (Table

7).

Hatching Behavior

Hatching behavior was monitored in 14 nests, 7 by direct

observation and 7 by electronic nest recorders. Eight of the 14 nests

were inspected during the hen's last recess prior to hatching. In two

nests, one egg was pipped; in another, two eggs were pipped; none was

pipped in the other five nests. None of the 14 hens recessed after

the pipping stage.

Vocalizations of hens and poults in three nests were recorded by

monitoring microphones during the hatching process. Brood hens made a

number of calls that are used by turkeys at other times; however, two

calls were restricted to the hatching period. One is a hatching yelp

(Williams 1984) that begins with single notes at 500 Hz spaced about

0.5 to 1.0 second apart (Fig. 17); it is sometimes accelerated to more

than four notes per second. Images in sonograms have a resemblance to

quarter notes in standard musical notation. The hatching yelp is

uttered by the hen immediately after the poult peeps. Another

previously undescribed call peculiar to the hatching period, termed

the hatching hoot (Fig. 18), consists of a prolonged, 600 Hz note that

is given irregularly throughout the hatching period.

While in the nest, poults give peeping calls of three types (Fig.

19): a single note that is repeated; a two-part ascending and

descending note; and a multi-syllable call resembling the lost

whistling call that poults use when they become separated from the

hen.





















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These five calls, which were heard from all seven nests monitored

by microphone, appear to be the basic vocal communications between the

hen and her poults. These are the sounds that enable the poults to

learn to distinguish the voice of their mother from other hens by the

second day of life (Ramsay 1951).

As the poults hatched and the nest became crowded, the hen moved

farther back, surrendering the front of the nest to the young. Poults

napped, mainly under the sides, tail, and drooped wings of the hen,

and sometimes ventured outside the nest during periods of activity.

When a poult ventured as far as 1 m from the nest, the hen would yelp

for it to return. Hens did not leave the nest or respond in any way

other than vocally when poults ventured out of the nest.

Egg Hatchability

Hatchability was 89% (SD = 0.14) of 839 eggs in 85 undisturbed

nests. McDowell (1956) reported a hatching rate of 97% in 13 nests in

Virginia, and Everett et al. (1980) reported a rate of 95% in 14 nests

in Alabama. Green (1982) reported only 72% hatchability in a Michigan

study in which the stock was descended from pen reared turkeys.

The Imprinting Period

Parental imprinting is a learning process by which young birds of

many species become socially attached to their parents and gain self

identity as members of their own species (Lorenz 1937). Imprinting

takes place during the first few hours of life and is crucial for wild

nidifugous birds. Imprinting is said to be irreversible in the turkey

(Schein 1963). Although there is some dispute about the









irreversibility of imprinting (Salzen and Meyer 1968), it is, at the

least, persistent and very difficult to reverse (Fabricius 1962).

Much of the research on imprinting has been conducted with the

mallard (Anas platyrhynchos). Ducklings remain in the nest for 24-62

hours after hatching (Hess 1972), which is 10-12 hours after the last

hatched duckling is dry (Kear 1965). Imprinting is compete (Bjarvall

1967) or nearly so (Fabricius and Boyd 1954) by the time of nest

departure. Both auditory and visual cues are involved in imprinting

(Ramsay 1951). Auditory cues strengthen with time, reaching their

peak effect several hours after visual cues. This is probably because

the need for audible communication between the hen and ducklings is

greatest after they leave the nest and travel in vegetation where

visibility is obscured (Fabricius 1964). Visual cues for imprinting

are enhanced and following behavior is strengthed when ducklings make

short excursions from the nest and can see the hen while she is

calling (Bjarvall 1967). Ducklings without visual reinforcement

probably would not follow as strongly upon nest departure.

The imprinting process in the wild turkey is similar to that of

the mallard. Observations of seven turkey broods indicated that the

broods remained in the nest for variable intervals of time after the

last-hatched poult was dry and active. Hens vocalized softly during

this time and the poults made short excursions out of the nest and

returned at the vocal command of the hen. During the 10-minute period

prior to nest departure, hens called more frequently than before.

Upon departure, hens arose abruptly, stepped out of the nest without

hesitating, and moved slowly away while continuing to yelp softly to

the poults.









In each of the seven brood departures, there was a gradient in

the strength of the following response among poults--some followed at

the hen's side whereas others straggled behind. In three cases, a

few poults failed to follow the hen farther than about 1 m from the

nest, which caused the hen to stop and yelp. In two cases, a few

poults remained in the nest, apparently more strongly attracted to it

than to the hen. They followed the hen only after she returned to the

nest and called to them.

Once an observation blind was erected too near a nest, which

caused the brood to depart prematurely about 6 hours after the last

poult had hatched. Three of the poults remained in the nest for 5

minutes during which time the hen called to them from a distance of

about 2 m. The hen had to return to within 1 m of the nest repeatedly

before the poults finally followed her.

Another brood hatched on 30 May and, due to investigative

disturbance, left the nest about 12 hours later. Four poults did not

follow the hen despite her continuous yelping. The observers left the

area so that the brood could reassemble; however, the next morning one

poult was found alone near the nest. Radio signals indicated that the

hen was still near. The observers again left the area. The following

morning, the four stray poults were found near the nest--the hen had

left the area without them. Three of the poults were captured

eventually by playing-back the sounds of hatching that had been

recorded at the nest 3 days earlier. The poults later became

parent-imprinted on humans, which indicated that they had not

imprinted on the brood hen. The foregoing observations suggest that









one of the reasons poults remain in the nest as long as they do after

hatching is to become adequately imprinted on the hen.

Time of Nest Departure by Broods

None of 28 broods departed the nest before sunrise or after

sunset (Fig. 20). Nineteen (68%) departed during morning hours.

Brood departure at mid-day was infrequent, corresponding to the

reluctance of incubating hens to recess at that time. Only one brood

departed after 1800 hours and it traveled only about 10 m before

roosting that night. No brood returned to its nest after departing.

Hatching Synchrony

Some birds with large clutches have adaptations that hasten the

hatching of the last laid eggs so that all the eggs hatch at nearly

the same time. Synchronized hatching is advantageous for nidifugous

birds with large clutches because it facilitates nearly simultaneous

parental imprinting and early nest departure of the brood. Hatching

synchronization is so well developed in the northern bobwhite that the

entire clutch usually hatches in less than 2 hours (Vince 1969).

If hatching is synchronized, the time required for hatching

should be less than the total incubating time experienced by the first

egg before the last egg is laid. The regression equation for the

onset of incubating behavior in the turkey (Fig. 11) shows that, on

average, the first eggs received approximately 25 hours of incubation

by the time the twelfth egg was laid. It would appear then, that in

taking more than 24 hours to hatch the clutch, the turkey exhibits

poorly developed synchronizing mechanisms.































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Attendance of Infertile Eggs

Two incubating hens attended infertile clutches for 35 and 64

days. The hen that incubated continuously for 64 days exceeded the

normal incubation period by 37 days, incubating about 2.5 times the

normal period. The northern bobwhite has been reported to sit on

unhatched eggs as long as 56 days, or 2.4 times its 23-day incubation

period (Stoddard 1931).

Nesting Habitat

There were too few nests on the Lochloosa Study Area to warrant a

detailed analysis of nesting habitat. Of 236 nests on Fisheating

Creek Study Area, 58% were in palmetto, 31% were in cypress woods, and

11% were in miscellaneous types of habitat (Table 8). The proportions

of the three habitats were 5%, 51%, and 44% for palmetto ecotone,

cypress woods, and miscellaneous types, respectively. Hens selected

nesting habitats in different proportions than they occurred on the

area (X2 = 1328, df = 2, P <0.001).

Cypress woods and palmetto habitats differed in plant species

composition (Table 9) and structure. The major structural difference

was a tree canopy in the cypress woods (Fig. 21), which was absent

from the palmetto ecotone (Fig. 22). Low vegetation in the cypress

woods habitat was fast-growing in contrast to the relatively

unchanging composition and structure of the woody vegetation that

predominated in the palmetto habitat.

Saw palmetto and wire grass occurred at all 44 nest sites in the

palmetto ecotone (Table 9). The foliar coverage of palmetto and wire

grass occupied more than 50% of the area within 1.5 m of 80% of the

nests and no less than 20% of the area near any nest site in this












Table 8. Number and proportion of 236 Florida wild turkey nests in
three habitat categories, Fisheating Creek and Lochloosa study areas,
1968-82.


Nesting habitat type
Year Palmetto Cypress woods Other

1968 16 (84%) 3 (16%) 0
1969 12 (86%) 2 (14%) 0
1970 15 (79%) 4 (21%) 0
1971 10 (71%) 1 ( 7%) 3 (21%)
1972 7 (50%) 7 (50%) 0
1973 12 (67%) 5 (28%) 1 ( 6%)
1974 12 (55%) 6 (27%) 4 (18%)
1975 11 (65%) 5 (29%) 1 ( 6%)
1976 7 (58%) 5 (42%) 0
1978 9 (75%) 0 3 (25%)
1979 5 (50%) 3 (30%) 2 (20%)
1980 8 (29%) 15 (54%) 5 (18%)
1981 12 (38%) 14 (44%) 6 (19%)
1982 1 (20%) 3 (60%) 1 (20%)
Total 137 (58%) 73 (31%) 26 (11%)






63


Table 9. Major plantsa occurring within 1.5 m of 63 Florida wild
turkey nest sites, Fisheating Creek Study Area, 1968-72.


Percentage occurrence in habitat


Palmetto Cypress woods Miscellaneous
Plant name (N = 44) (N = 15) (N = 4)


Trees

Taxodium distichum 100

Shrubs

Baccharis hamlimifolia 25
Callicarpa americana 20 25
Ilex glabra 25
Lyonia ferruginea 36
Lyonia lucida 73 25
Myrica cerifera 25
Quercus chapmanii 32
Quercus geminata 18
Serenoa repens 100 25

Vines

Smilax spp. 25 25
Rubus sp. 25
Vitis sp. 25

Herbs

Aristida stricta 100 25
Quercus minima 64
Andropogon sp. 32
Vaccinium myrsinites 25
Panicum sp. 36 25
Axonopus compressus 95 25
Hydrocotyle umbellata 33
Centella asiatica 47
Saururus cernuus 20
Polygonum sp. 40
Rhus radicans 47
Hypericum sp. 25
Iris savannarum 25
Eupatorium coelestinum 73
Eichhornia crassipes 33 25

a Nineteen additional plants, mostly immature seedlings, that did
not occur at more than two nest sites, are not listed.
















Al


Figure 21. Investigator standing beside turkey nest in cypress
woods habitat, Fisheating Creek Study Area, 1981.


























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habitat. Lyonia lucida was the second most prevalent shrub at

palmetto nests, occurring at 32 sites (73%).

In cypress woods, cypress trees occurred within 1.5 m of all 15

nests, and the grass Axonopus compressus occurred near 13 nests.

Nests in the cypress woods usually were in the densest ground cover

available, which often was in semi-aquatic vegetation such as Centella

asiatica, Saururus cqrnuus, and Polygonum spp. (Table 9).

Each of the four nest sites in the miscellaneous habitat category

was in distinctly different vegetation: a narrow, dry ditch; a flood

control dike; a high, isolated saw palmetto clump; and a wax myrtle

thicket along a fence row.

Ninety-five percent of the 20 cypress woods nests and 87% of the

62 nests in palmetto had 40% or more vegetative cover within 2 m above

them (Table 10). Only one nest (5%) in the cypress woods and two (3%)

in the palmetto ecotone had less than 10% overhead cover. Thirty-

eight percent of the nests in palmetto had 70% overhead cover, and 24%

had more than 90% overhead cover. None of the nests in cypress woods

had more than 90% overhead cover.

Saw palmetto was favored for first, second, or third seasonal

nesting attempts (Table 11). Nesting habitat was used in similar

proportions by adults and yearlings (N = 223, X2 = 0.68, df = 2, P =

0.712). A higher proportion (68%) of nests in saw palmetto was

successful than in cypress woods (45%) (N = 148, X2 = 8.043, df = 2, P

S0.018).

Forty-two percent of 36 hens that were first observed nesting in

the palmetto ecotone renested in another habitat; 33% first observed

nesting in the "other" habitat type renested elsewhere; and 22% first















Table 10. Cover within 2 m above 82 nests located in cypress woods
and palmetto nesting habitats, Fisheating Creek Study Area, 1968-72.


Number and percentage of nests
Percentagea overhead
cover Cypress woods Palmetto

< 30 1 ( 5%) 4 ( 6%)
31-40 2 (10%) 10 (17%)
51-70 6 (30%) 10 (17%)

> 70 11 (55%) 38 (61%)
Totals 20 (100%) 62 (100%)

visually estimated.

















Table 11. Proportions of first, second, and third nests of Florida
wild turkeys, established within a single season, by habitat type,
Fisheating Creek Study Area, 1968-82.



Percentage of nests


First Seconda Thirda All
nests nests nests nests
Habitat type (N = 191) (N = 38) (N = 7) (N = 236)


Palmetto 60 47 57 58

Cypress woods 30 37 14 31

Other 10 16 28 11

100 100 100 100

aThese were the first, second, or third nests observed. A few nests
were probably depredated during early laying stages; thus some nests
listed as second may have actually been third et cetera.









observed nesting in cypress woods renested in another habitat (Table

12). Although there seemed to be a preference for nesting in saw

palmetto cover, there was considerable variability in the nesting

habitat used in different years (Table 8). The tendency to change

freely from one habitat to another for renesting in the same year, and

between years, suggests that habitat imprinting may not be an

important factor in nesting habitat selection by the Florida turkey.

Nesting Seasonality

The earliest egg found in the study was laid on 6 March; the last

egg was laid on 6 June. The last clutch hatched on 2 July. The

median date of laying the first egg was 23 March (x = 25 March) and

the median date the last clutch hatched was 8 June (x = 10 June) (Fig.

23). Nests that were initiated after 1 May were probably the second

or third nests of the respective individual hens that season. The

molting pattern (Williams and Austin in press) of at least 3,000

juvenile turkeys examined during summer and fall on the Lochlossa and

Fisheating Creek WMAs substantiated that few poults were hatched after

early July.

Yearling hens began nesting about 2 weeks later and completed

nesting about 2 weeks earlier than adults (Fig. 24). The difference

was significant (N1 = 88, N2 = 25, Kolmogorov-Smirnov Z = 1.923,

maximum difference = 0.436, 2-tailed P = 0.001). Later initiation of

nesting by yearlings apparently occurs also in the Rio Grande turkey

as Smith (1977) reported that adult hens in Texas began copulating

before leaving their winter range whereas yearlings did not begin

copulating until after reaching their spring range.




































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Nesting Success and Predation

Sixty percent of the 171 nests that were not disturbed by

investigative activities were successful (Table 13). This rate of

nest success is not necessarily an accurate estimate, however, because

not all the nests were-observed from the time of clutch initiation, a

factor that would tend to bias nest discovery in favor of successful

nests. The Mayfield (1961) method, which provides an appropriate

adjustment for nests observed during only a portion of their full

terms, was used to calculate nesting success. During the 14-year

study, eight nests were lost to predators during 218 days of exposure

within the laying period (Table 14). Thus 0.0367 (8/218) nests were

lost per day during the laying period. The daily probability of

survival for any nest would be 0.963 (1-0.0367). Inasmuch as

approximately 12 days are required for a clutch of 10 eggs to be laid

(one egg per day plus two lapse days in a typical clutch), the entire

laying period would be 12 days and the probability of a nest surviving

the 12-day laying period would be 0.96312 or 0.638. The probability

of survival to hatching for each nest during the incubating period,

using 27 as the number of days in the incubation period, was 0.708

(38/2987 1.0 = 0.0.9873; 0.0987327 = 0.708). The probability of a

nest hatching would be 0.638 x 0.708 = 0.452. Thus, the proportion of

nests surviving was 63.8% during the laying period, 70.8% during the

period of continuous incubating behavior, and 45.2% during the entire

nesting period. As expected, the latter figure is lower than the 60%

hatching success recorded for all nests under observation (Table 13).
















Table 13. Fates of Florida wild turkey nests under observation,
Fisheating Creek and Lochloosa study areas, 1968-82.


All nests (N = 237)


Fate Number Percentage


Hatched 103 43

Predator 59 25

Flushed 42a 18

Deserted 31b 13

Flooded 2 1

alncludes 15 hens that were flush
hens deserted.
bIncludes 23 hens that may have b


Nests not disturbed by
investigators (N = 171)


Number Percentage


103 60

59 35


8 5

2 1


led intentionally; not all flushed


een disturbed by the investigators.










Table 14. Nest survival data used
successful nesting by Florida wild
Lochloosa study areas, 1968-82.


to calculate the probability of
turkey hens, Fisheating Creek and


Laying period Incubation period


Number of Days of Number of Days of
nests nest nests nest
Year depredated exposure depredated exposure


1968

1969

1970

1971

1972

1973

1974

1975

1976

1977

1978

1979

1980

1981

1982

Total


260

108

298

196

187

297

337

195

183

0

153

174

259

377

63

2987









It is possible to take renesting into account in estimating the

success of the nesting hen population by using the probabilities

calculated previously:

0.566 = the probability that a hen will renest if the nest is

lost during the laying period (see Renesting);

0.280 = the probability that a hen will renest if the nest is

lost during the incubation period (see Renesting);

0.638 = the probability that a laying hen will reach the

incubating stage;

0.708 = the probability that a clutch that reaches the stage of

being continuously incubated will hatch;

0.362 = (1-0.638) = the probability that any nest will be lost

during the laying period;

0.292 (1-0.708) = the probability that any nest will be lost dur

the incubation period.


-ing


In calculating nesting success of the population, it is assumed

that all hens will attempt to nest and will renest once if the nest is

disrupted during the incubating period and twice if disrupted during

the laying period. These calculations yield

(0.638)(0.708) = 0.452 as the proportion of the first time

nesters that will be successful,

(0.362)(0.566)(0.638)(0.708) = 0.093 as the proportion of hens

losing their nests during the laying period that will

successfully renest,

(0.638)(0.292)(0.280)(0.638)(0.708) = 0.024 as the proportion of

hens losing their nests during the incubating period

that will successfully renest, and









(0.362)(0.566)(0.362)(0.566)(0.638)(0.708) = 0.019. as the

proportion of hens losing their nests twice during the

laying period that will eventually be successful.

Overall nesting success would be the sum of the products:

0.452 + 0.093 + 0.024 + 0.019 = 0.588 or 58.8%.

The calculated rate of nesting success may not accurately reflect

nesting success of the entire population because hens that were not

known to nest are not included in the calculations. If some of them

did not attempt to nest, which is likely, the overall nesting success

rate would be lower. Turkey populations on the study areas increased

during the period of study despite the nest losses to predators.

Nest predation was distributed evenly throughout the incubation

period (Fig. 25). Predatory species responsible for nest losses could

not be ascertained in every case, but evidence at nest sites indicated

that the raccoon (Procyon lotor) was a prominent predator in the

cypress woods, and that the striped skunk (Mephitis mephitis) and

spotted skunk (Spilogale putorius) were major predators in oak scrub

and palmetto. Other predators included the opossum (Didelphis

virginianus), gray fox (Urocyon cinereoargenteus), bobcat (Felis

rufus), and domestic dog. The American crow (Corvus brachyrhynchos)

destroyed only a single nest.

Potential nest predators that were not known to take eggs

included the feral hog (Sus scrofa), which has been reported to

depredate turkey nests (Barkalow 1942, Blakey 1937), and the armadillo

(Dasypus novemcinctus), which is suspected by many laymen of taking









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turkey eggs. Although armadillos did not eat any turkey eggs, they

rooted through two nests, rolling out some of the eggs and causing the

hens to abandon the nests.

A higher proportion of nests was depredated in the cypress woods

(55%) than in saw palmetto (32%) or in the "other" (21%) habitats (N =

148, X2 = 8.043, df = 2, P = 0.018). The "other" habitat category was

heterogeneous in plant species composition and structure, whereas the

cypress woods and palmetto were relatively homogenous. The lower

predation rate in the heterogeneous "other" habitats supports the

experimental findings of Bowman and Harris (1980), who predicted that

predation rates would be higher in homogeneous habitats.

Predator success is reportedly high when prey is abundant and

increases with hunting experience of the predators (Tinbergen et al.

1967). If these factors were important in the present study, nest

losses should have been higher in mid- or late season nests than in

early nests. However, predation rates did not differ among

early-season (hatching before 1 May), mid-season (hatching 1 May

through 20 May) or late-season (hatching after 20 May) nests (N = 117,

X2 = 2.6, df = 2, P = 0.26).

Defensive Behavior

Predation is an important factor in the life and evolution of the

wild turkey. Nest losses to predators were 54.8% as previously

calculated and poult losses during the first 2 weeks were about 70%

(Williams and Austin in press). The annual turnover rate in turkey

populations has been estimated to range from 30% (Logan 1973) to 50%

(Smith 1977).









The major behavioral adaptations of the turkey hen to avoid nest

predation are 1) selecting a well-hidden nest site, 2) laying eggs in

mid-day when the common nest predators are inactive, 3) covering the

nest with debris before leaving it during the laying period, 4)

minimizing activity near the nest, 5) remaining on the nest when

predators approach, and 6) sometimes flying to and from the nest

rather than walking.

The plumage of the hen is cryptically marked with brown and

black. Natural colors of the ground litter at nesting sites tend to

optimize the camouflage effect. Nests are covered lightly with a few

dead leaves when left unattended during the laying period. Overhead

cover of nests is at least 40% and sunlight through the vegetation

creates a broken pattern of light and dark spots, which accentuates

the cryptic effect of the hen's speckled back pattern or the

camouflage of dried litter covering the eggs.

Hens lay in mid-day at which time the major nest predators

(raccoon, opossum, and skunks) are inactive. It is unlikely that

predators would see the hen approaching her nest at mid-day or would

track her by scent when they emerge to hunt several hours later.

Hens do not linger near their nests. Laying hens spend their

inactive daylight hours away from the nest and roost at least 0.5 km

from the nest (Williams et al. 1974). When approaching the nest to

lay, hens move steadily and enter the nest quickly. If a hen sees a

person as she returns to the nest, she delays her approach but returns

quickly to the nest when the person leaves the area.

Eight hens were monitored from observation blinds and by

telemetry as they left their nests during the laying period. They did









not dust, rest, or feed until they had moved at least 200 m from the

nest. Only two hens of the eight were known to visit their nests

except to lay or incubate and each did so only once.

Hens chose dense cover for nesting, and remained motionless if

approached on the nest. The turkey's strategy of holding tightly to

the nest is a general defense strategy of many prey animals (Edmunds

1974). An observer approached within 6 m of 13 nests located in

palmetto habitat a total of 19 times; in no case did the hen flush

unless the investigator passed closer than 2 m. In seven cases, hens

returning to their nests were frightened away by an investigator who

was within 5 m of their nests; none abandoned.

In approximately 40,000 man-hours of field work only one active

nest was found accidentally. This finding, coupled with the

observations on nest holding behavior, suggests that disturbance of

hens on their nests by humans in cover conditions such as those on the

study areas, is minimal.

Turkey hens in Alabama (Wheeler 1948) and in the Rocky Mountains

(Ligon 1946) have been observed flying to and from their nests. In

the present study, 20% (N = 140) of the hens flew from or to the nest

site. In flying to their nests, turkeys usually landed 3-15 m away

and walked the remaining distance; when flying from the nest, they

took two to five steps before flying, often taking wing within 3 m of

the nest.

Much has been written about nest abandonment as a result of human

disturbance of nesting hens (see Schorger 1966:265-266 for a review).

Logan (1973) reported that seven of nine hens in Oklahoma were flushed

repeatedly from their nests without abandoning. The two hens that did









abandon their nests were flushed during the laying period--the other

seven were flushed while incubating. Hens apparently have differing

tolerances to being flushed from the nest. Of the 38 hens flushed

from their nests a single time in the present study, 55% did not

return (Table 15).

It is widely believed that nesting hens are more likely to

abandon their nests if flushed during the laying period than if

flushed during the period of continuous incubating behavior (Schorger

1966). In this study, 60% of the hens that were flushed during the

period of continuous incubating behavior returned to their nests in

contrast to 38% of those flushed during the laying period (Table 15).

Although the observed behavior supports the consensus view on the

subject (Schorger 1966), the difference was not significant (X2 =

1.29, df = 1, P = 0.255).

There is evidence that a hen is more likely to return to her nest

if flushed late in the incubation period than during laying or early

in the incubation period. The return rate was 38% for hens flushed

during the laying period, 42% for hens that had incubated less than 8

days when flushed, and 60% for hens that had incubated longer than 8

days (Table 15); these differences, however, were not statistically

significant (X2 = 0.84, df = 2, P = 0.359).

Habitat type appeared not to influence nest abandonment.

Thirty-nine percent of the hens flushed from nests in the cypress

woods returned as compared to 59% returning to nests in palmetto (X2 =

1.61, df = 1, P = 0.446) (Table 16). However, a higher proportion of

yearling hens (73%) abandoned their nests after being flushed than did

adults (44%) (X2 = 2.751, df = 1, P = 0.097).


















Table 15. Number and percentage of Florida wild turkey hens returning
to their nests after being deliberately flushed a single time by an
observer, Fisheating Creek Study Area, 1968-82.



Hens returning
Number of
Nesting period hens flushed N %


Laying period 8 3 38


First week of incubation 19 8 42

After first week of 10 6 60
incubation

Incubation period 30 18 60

Entire nesting period 38 21 49


















Table 16. Number and percentage of Florida wild turkey hens by age
class and habitat type that returned to their nests after deliberately
being flushed by an observer during the laying and incubation period
combined, Fisheating Creek and Lochloosa study areas, 1968-82.



Hens returning
Habitat type or Number of
age class hens flushed N %


Palmetto 22 13 59

Cypress woods 18 7 39

Adult 32 18 56

Yearling 11 3 27















SUMMARY

Wild turkey hen flocks dissolved in spring as the hens

established new home ranges and began to visit nesting cover. Nests

were not prepared in advance of laying. Hens scratched away leaves

and soil to make shallow depressions and deposited their first eggs on

the bare soil in mid-day. Saw palmetto with wiregrass, in an ecotone

between oak scrub and saw palmetto prairie, was the favored nesting

habitat. A substantial proportion of the hens also nested in cypress

woods and in miscellaneous habitats. There was much variation in the

use of different nesting habitat from year to year and from one nest

to the next within the same season by the same hen.

Before a hen departed the nest after laying, she partially

covered the eggs with leaf litter picked up from beside the nest. Each

time she returned to lay another egg, the hen settled on the nest

without removing the litter. This habit camouflaged the nest and

produced the leafy lining characteristic of the completed nest.

Hens averaged about 1 hour at the nest while laying each of the

first 5 eggs, which were deposited from a standing position. Laying

was irregular at first--i to 3 days being skipped early in the laying

period before laying became a daily event. Hens rarely visited their

nests except to lay or incubate. After five eggs had been laid, hens

remained on the nest longer with each subsequent laying. This

behavior initiated incubating behavior gradually. By the time the

last egg was laid, hens were remaining on their nests from mid-day









until late afternoon. Most hens remained on the nest overnight for

the first time on the same day the last egg was laid, and continued to

incubate overnight except for daytime recesses during the remainder of

the incubation period. Incidents of more than one hen laying in the

same nest and of egg-dropping occurred only infrequently. The hens

exhibited characteristics of a determinant laying species.

Hens usually left their nests only once every 2 days although

some hens remained on the nest for a span of 4 days without a recess.

Recesses averaged 98 minutes. Most recesses were taken in the morning

or afternoon; mid-day recesses were infrequent. Successive recesses

were not routinely taken during the same time of day.

Hens did not cover their eggs when on recess during the

incubation period as they did during the laying period. Hens turned

their eggs during both the laying and incubation periods. Eggs rolled

intact from the nest were sometimes placed back in the nest by the

hen; however, hens usually abandoned nests containing a broken egg.

The first poult was hatched after about 26 days of continuous

incubating behavior. Although the gradual onset of incubating

behavior resulted in about 25 hours disparity in the amount of

incubation experienced by the first and last eggs, hatching

synchronization appeared to be poorly developed. The brood remained

in the nest for at least one-half day after the hatching of the last

poult. Parental imprinting of the poults was facilitated by frequent

voice communications between the hen and brood. The brood usually









departed the nest in the morning before the end of the twenty-seventh

day after the beginning of continuous incubating behavior.

Average clutch size was 10.3 eggs. Clutch size of adults and

yearlings differed, being 10.5 and 10.0 eggs, respectively. There was

no difference in the number of eggs in first, second, or third nests

or for early vs late nests in the same year. The two largest clutches

contained 17 eggs. Incubated clutches with fewer than five eggs were

not found, probably because hens are not adequately stimulated to

incubate fewer than five eggs. Egg hatchability averaged 89%.

Turkey nests were established in heavy ground cover and were well

camouflaged. Incubating hens held tightly to their nests when

approached by a human or potential predator. This behavior probably

increased nesting success inasmuch as flight by the hen usually would

reveal the location of the nest, and thereby increase the probability

of its destruction. About one-half of the nests were destroyed by

predators; however, when renesting was considered, nearly 60% of the

nesting hens were successful in hatching a brood. Major nest

predators were the raccoon, stripped skunk, spotted skunk, and

opossum. Predation incidents were distributed evenly throughout the

period of continuous incubating behavior. Laying hens renested more

readily than incubating hens; adults renested at about twice the rate

of yearlings.

The nesting season extended from early March, when the first

nests were established, to early July, when the last clutch hatched.

The median date of nest establishment was 23 March; the median date of

the last hatching was 8 June. Thus, the nesting season for the









Florida turkey is essentially restricted to the months of April, May,

and June.

Nesting Florida turkey hens are secure from serious levels of

disturbance associated with spring gobbler hunting or other human

presence in their habitat because of the dense nesting cover used and

the tenacity with which they hold to the nest in the presence of

humans. Even when disturbed, approximately half of the hens that were

flushed from their nests returned and continued normal nesting

activities. Furthermore, 39% of the flushed hens that abandoned their

nests renested.

Incubating hens leave their nests infrequently for very short

periods and would not likely be encountered by hunters. Furthermore,

more than half of their recesses are taken in afternoon during which

time spring gobbler hunting in Florida is not permitted. There would

be no advantage in setting the spring gobbler hunting season to

coincide with peak incubation activity for the purpose of protecting

nests from dispruption because the net impact on reproduction is the

same regardless of whether the hen is flushed while laying or while

incubating.

Although hens exhibited a strong preference for nesting in the

ecotone between oak scrub and saw palmetto prairie, and in cypress

woods, the highest rate of nest success occurred in habitats other

than palmetto or cypress. Because of the great variation in nesting

habitat used from year to year and the strong tendency of hens to

renest freely in a habitat different from that of the earlier nest,

manipulation of nesting cover is not warranted at this time. However,






89


it would probably be beneficial to preserve much of the palmetto-scrub

ecotone in habitat occupied by the turkey in Florida.















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