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The ecology and vocalizations of Scott's seaside sparrows

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Title:
The ecology and vocalizations of Scott's seaside sparrows (ammodramus maritimus peninsulae)
Creator:
McDonald, Mary Victoria, 1952-
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Language:
English
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xi, 145 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Bird nesting ( jstor )
Bird songs ( jstor )
Birds ( jstor )
Breeding seasons ( jstor )
Female animals ( jstor )
Mating behavior ( jstor )
Muting ( jstor )
Popular songs ( jstor )
Singing ( jstor )
Sparrows ( jstor )
Ammodramus -- Habitat ( lcsh )
Birds -- Florida ( lcsh )
Birdsongs ( lcsh )
Sparrows -- Habitat ( lcsh )
Seaside ( local )
Genre:
bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1986.
Bibliography:
Includes bibliographical references (leaves 136-144).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Mary Victoria McDonald.

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University of Florida
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University of Florida
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Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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AEN8908 ( NOTIS )
16109369 ( OCLC )

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THE ECOLOGY AND VOCALIZATIONS OF SCOTT'S SEASIDE SPARROWS
(Ammodramus maritimus peninsula)

















By

MARY VICTORIA MCDONALD


A DISSERTATION PRESENTED 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


1986
























































Copyright 1986

by

Mary Victoria McDonald

















ACKNOWLEDGEMENTS


Since this dissertation is a formal report of my doctoral research,

it would be inappropriate for me to describe all of my ideas and my

feelings over the years--false starts, hunches that paid off, my

occasional discouragement, confusion, and elation, and even the kinship

I felt working with "my" birds on "my" marsh. But these elements were

inextricably woven into my study. What saw me through was not just my

scientific approaches to interesting ornithological questions, but also

determination and good luck, and, most of all, invaluable help from

friends and advisors.

Dr. John William Hardy, my major advisor, steadfastly supported and

guided me through the years. His timely reviews of my work and my

requests for recommendations, his just criticisms, and his flexibility

in allowing me to work independently all significantly contributed to

the success of my graduate career. My other graduate committee members,

Drs. Pierce Brodkorb, John H. Kaufmann, Peter Feinsinger, and Thomas J.

Walker, thoughtfully and generously advised me throughout my project.

Many other faculty members in the Department of Zoology also helped me

greatly.

I am fortunate to have had valuable suggestions from three

ornithologists very knowledgeable about seaside sparrows--Herbert W.

Kale II, Jon S. Greenlaw, and William Post. William Post patiently

helped me get started in my field work at Cedar Key and allowed me to










incorporate some of his unpublished reproductive data and banding

records with my data, as presented in Chapter II of this dissertation.

The Department of Zoology generously supported me with teaching and

research assistantships, equipment, and vehicle use. The Florida State

Museum Bioacoustics Laboratory provided sound analysis equipment and

work space. Other financial support was provided with grants from

several sources: Sigma Xi Grants-In-Aid of Research (1982 and 1983),

Frank M. Chapman Memorial Fund Awards (1983 and 1984), Eastern Bird

Banding Association Research Award (1984), and Van Tyne Memorial Fund

Grant of the American Ornithologists' Union (1984).

John David Wood, Sr., graciously permitted me to conduct my project

on his property near Cedar Key, Florida. The Florida Department of

Natural Resources rangers of the Waccasassa Bay Station near Cedar Key

helped make my sometimes uncomfortable, and always wet and muddy field

work more bearable. Two occasional field assistants, Janine Russ and

David Specht, were genial and as well as adept companions. Thomas A.

Webber also helped me in the field with photography and sound recording.

Many, many other people helped me indirectly over the years. The

support of my parents, Carlyle A. McDonald and Margaret L. McDonald, was

invaluable. And likewise invaluable were the advice and support of many

fellow graduate students. Thomas A. Webber and Linda S. Fink deserve

special thanks--their suggestions, thoughtful and significant criticisms

of my work, and, most of all, their supportive friendships have been

essential parts of my graduate work and life.


















TABLE OF CONTENTS

Page
ACKNOWLEDGEMENTS . iii

LIST OF TABLES . vii

LIST OF FIGURES . viii

ABSTRACT . x

CHAPTERS

I GENERAL INTRODUCTION . 1

II NATURAL HISTORY AND BEHAVIORAL ECOLOGY OF SCOTT'S
SEASIDE SPARROWS: STUDY TECHNIQUES AND RESULTS 2

Taxonomy and Morphology . 2
Project History and General Methods 4
Study Site . 6
General Description . 6
Weather . 7
Marsh Fauna . 8
Vertebrate fauna . 8
Invertebrate fauna and seaside sparrow foraging 9
Marsh Flora . 10
Size and Demarcation of Study Sites 11
Reproduction and Reproductive Behavior .. 11
Overview of Annual Cycle 11
Annual Cycle of Reproductive Behavior 15
Territorial establishment and mating 15
Nests, eggs, and early development ... 18
Nesting behavior . 20
Post-breeding behavior 22
Productivity, Survival and Reproductive Success 23
Territories and Territorial Behavior 29
Definition and Methods of Determining Territories 29
Description of Territories at Cedar Key 29
Territorial Behavior ............... 30
Reactions to Other Species of Birds and to Humans 32
Discussion of Seaside Sparrow Territoriality 34
Territory types and variation 34
Territory quality and space use 35
Extension of territory definition and function 36










III VOCALIZATIONS OF SCOTT'S SEASIDE SPARROWS 40

Introduction . 40
General Methods of Data Collection and Analysis .. 41
Coverage ... 41
Sample Size and General Observation Methods 42
General Analysis of Notes and Recordings 43
Results and Discussion of Calls and Song ... 45
Description and Use of Vocalizations .. 45
Calls of Scott's seaside sparrows 47
Primary song 73
Subsong .o. 79
Countersinging and Repertoire Use .. 79
Introduction and comments .. 79
Methods of investigating repertoires .. 82
Results and discussion of repertoire use 83
Flight Songs: Description and Comparison to
Perch Songs . 84
Description . 84
Methods of investigating flight song activity 85
Results of singing activity analysis 87
Discussion of flight songs .. 88

IV FUNCTION OF SONG IN SCOTT'S SEASIDE SPARROWS .. 98

Introduction ... ... 98
Methods . 101
Experimental Design . 101
Muting Procedure . .. 106
Statistical Analysis . 108
Results .. . 109
Voice and Post-Operative Recovery .. 109
Mate Attraction and Retention 110
Territory Establishment, Retention, and Size Change 119
Behavioral Changes of Muted Birds 120
Reaction to Playback . 129
Voice Recovery and Subsequent Behavior .. 130
Discussion . 130

V CONCLUSIONS . 135

REFERENCES .. .. 136

BIOGRAPHICAL SKETCH . 145


















LIST OF TABLES


Table II-1.



Table III-1.


Table 111-2.

Table 111-3.



Table IV-1.

Table IV-2.


Table IV-3.



Table IV-4.


Table IV-5.


Table IV-6.


Summary profile of territoriality and reproduction
of Scott's seaside sparrows at Cedar Key, Florida,
1979-1984 . .

Vocalizations and behaviors measured during time
budget observations .

Vocalizations of Scott's seaside sparrows .

Characteristics of solo primary singing and
countersinging of 30 mated male Scott's seaside
sparrows during April and May .

Experimental design of muting experiments .

During songless comparison of territory ownership
and size changes for the Mid-Season mutings .

"After song regained" comparison of territory
ownership and size changes for the
Mid-Season mutings .

Behavior changes for individuals, from "Before"
to "During" muted .

Behavior differences comparing Muted, Sham, and
Undisturbed groups .

Sexual selection related to two main song types .





















Figure II-i.


Figure III-1.




Figrue III-2.


Figure 111-3.

Figure 111-4.

Figure 111-5.

Figure 111-6.


Figure 111-7.

Figure 111-3.

Figure 111-9.


Figure III-10.

Figure III-11.


Figure 111-12.


Figure 111-13.


Figure 111-14.


Figure IV-1.


LIST OF FIGURES

Breeding chronology of seaside sparrows at
Cedar Key, Florida, 1979-1984 .

(A) Audiospectrogram of Tuck call (short notes
8 kHz), and 1 interspersed Tsip call (vertical
(B) Audiospectrogram of Tsip call.
(C) Audiospectrogram of Seeep note .

(A) Audiospectrogram of male Primary song and
concurrent female Seeep note.
(B) Audiospectrogram of Tchi call .


Audiospectrogram of Whinny vocalization

Audiospectrogram of Zuck calls .

Audiospectrogram of Scree calls .

Audiospectrogram of Begging calls from
two nestlings, individuals A and B .

Audiospectrogram of Primary song .

Audiospectrogram of Subsong .

Audiospectrograms of Countersinging from
individuals A and B .

Audiospectrogram of Flight song .

Relationship between time of day and the
of Flight songs/Perch singing .


around
note).






* .


males,
* *


ratio
* o o


Relationship between wind velocity and the ratio
of Flight songs/Perch singing .

Relationship between temperature and the ratio
of Flight songs/Perch singing .

Number of Flight songs recorded in early 1983
given by birds of designated mated categories

Audiospectrogram of song of bird ABOR recorded
3 days prior to muting ..


viii


. .











Figure IV-2.


Figure IV-3.



Figure IV-4.


Audiospectrogram of "songs" of muted bird ABOR
recorded 5 days after muting .

Relative abilities of Muted versus Sham-Operated
and Undisturbed males to attract and retain
females 1983 and 1984 .

Behavioral changes of 21 muted birds Before and
During their songless periods .

















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

THE ECOLOGY AND VOCALIZATIONS OF SCOTT'S SEASIDE SPARROWS
Ammodramus maritimus peninsula

By

Mary Victoria McDonald

December 1986

Chairman: Dr. John William Hardy
Major Department: Zoology

I studied the ecology and vocalizations of Scott's seaside sparrows

(Ammodramus maritimus peninsula) in a Florida salt marsh from

1981-1985. I observed the behavior, vocalizations and reproductive

efforts of 20-30 pairs each year on the 30 ha gridded study site. These

monogamous birds defended all-purpose territories about 1,750 m2 in size

with song and short-distance agonistic behaviors. A high proportion

of males (0.79) returned each year to their previous territories, and

half (0.50) of these remated with their previous year's mates. Although

females initiated about 3.7 clutches of 3 eggs each season, productivity

was low (0.67 fledglings/female/year), because predators destroyed about

80% of the nests. Nevertheless, population numbers remained stable over

the years 1979-1985. The birds' vocal repertoires consisted of a

primary song, three modified primary songs (whispering song, flight

song, and subsong), and 10 calls. Only males sang; they had a song

type repertoire of 2-3 songs. I experimentally investigated the

function of song during the breeding seasons of 1983-1985 by temporarily











muting male birds in the field. I muted birds by rupturing the

interclavicular air sac. Birds remained songless for about 2 weeks but

could give all of their normal calls during this time. I administered

two rounds of mutings, each preceded and followed by time budget,

playback, and other observations on 3 treatment groups: Muted,

Sham-Operated, and Undisturbed birds. The Early muting round tested for

mate and territory acquisition. These Muted birds remained mateless

until they regained their voices; most eventually attracted a mate.

Muted birds acquired territories later than Sham-Operated and

Undisturbed birds. The Mid-Season muting round tested for mate and for

territory retention, and changes in behavior. All Muted birds lost their

mates; a few attracted new mates when they regained their singing

ability. Their territories either shrank or were lost, but new

territories were established (or the original re-expanded) when singing

ability returned. The intensity of close-range aggressive behaviors was

significantly greater for Muted birds than for Sham-Operated and

Undisturbed birds. There were no discernible differences between

Sham-Operated and Undisturbed birds for any of the attributes measured.
















CHAPTER I
GENERAL INTRODUCTION


I studied the ecology and vocalizations of Scott's seaside sparrows

(Ammodramus maritimus peninsula) on a salt marsh near Cedar Key,

Florida, for 5 years. William Post had studied the reproductive ecology

of this population from 1979-1980. In early 1981, I began my

investigations and continued my study through the spring of 1965. The

principal objective of my research was to determine the function of the

vocal repertoire of these birds. During the course of my field work,

however, I observed many aspects of their biology and also collected

reproductive and non-vocal behavioral data.

This dissertation summarizes both my observational and experimental

investigations of the study population. Chapter II is an overview of

the project history, study site, general methods, and the birds and

their biology. Chapter III describes the vocal repertoire and discusses

the probable function of these vocalizations. Chapter IV summarizes the

experiments I conducted in 1983 and 1984, whereby I determined the

function of song in Scott's seaside sparrows.

The information presented throughout most of this dissertation

portrays the behavior of birds I observed and tape recorded with a

minimum of disturbance. The exception, of course, is that Chapter IV

presents results of manipulative experiments. Thus I sometimes make the

distinction between "normal" and "experimentally manipulated" birds.















CHAPTER II
NATURAL HISTORY AND BEHAVIORAL ECOLOGY OF SCOTT'S SEASIDE SPARROWS:
STUDY TECHNIQUES AND RESULTS


Taxonomy and Morphology

Seaside sparrows (Ammodramus maritimus) belong to the "grassland

group" of Emberizine finches (subfamily Emberizinae), which also

includes the genera Ammospiza and Passerculus. The nominate form (A. m.

maritimus) was originally described by Wilson in 1811. Currently nine

subspecies of seaside sparrows are recognized (American Ornithologists'

Union [A.O.U.] 1957, 1973), including the nearly extinct dusky seaside

sparrow (A. m. nigrescens). Beecher (1955) postulated that formation of

the races of seaside sparrows was due primarily to geographic isolation

caused by post-glacial rise in sea level and the concurrent drowning of

river mouths forming bays (also see Funderburg & Quay 1983). All but the

southernmost populations of the nominate race are migratory. The other

races are generally considered to be non-migratory.

Scott's seaside sparrow (A. m. peninsula) was named in honor of W.

E. D. Scott and described in 1888 by J. A. Allen. The type specimen was

collected near Tarpon Springs (Pinellas County, Florida). A. m.

peninsula ranges from Old Tampa Bay (Hillsborough County, Florida)

north to Pepperfish Key (Dixie County, Florida) (A.O.U. 1957).

Austin (1983) and I (McDonald 1983a) have briefly discussed and

annotated the taxonomic history of the seaside sparrow assemblage. The

distinguishing plumage color characteristics of the seaside sparrow

subspecies are thoroughly described and compared by Funderburg and Quay










(1983). Three museums in the United States house good representative

collections of seaside sparrows that I have examined: The American

Museum of Natural History, the United States National Museum of Natural

History, and the Florida State Museum.

Seaside sparrows are generally recognized by their dark gray-brown

head and body (total length about 14 cm), narrow black streaks on the

breast and flanks, yellow lores and wrist spots, and long bills. The

tail is relatively short and narrow and the feet and legs are relatively

large, in proportion to the size of the bird. When flushed, seasides

characteristically fly short distances and then drop back into the marsh

vegetation. Unlike other sparrows, seaside sparrows eat many

crustaceans and insects, but few seeds.

In his original description, Wilson succinctly described seaside

sparrow habits when he said: "Amidst the recesses of these wet sea

marshes, [the bird seeKs the rankest growth of grass and sea weed, and

climbs along the stalks of the rushes with as much dexterity as it runs

along the ground, which is rather a singular circumstance, most of our

climbers being rather awkward at running" (Wilson 1811, p. 68).

Scott's seaside sparrows are among the darkest of the seaside

sparrows. As in other races a certain degree of individual color

variation exists. Some birds approach the lighter colored A. m.

maritimus in shading on the dorsum; others approach the darker A. m.

nigrescens. Early taxonomists made no mention of this color variation,

although Griscom (1944) recognized it. Austin states: "A gradual cline

is evident from the smaller, grayer populations in the south

peninsulaa) to the slightly larger, darker, and dorsally browner birds

in the northwest corner of the range in the Wakulla area (juncicola)"










(Austin 1968a, p. 838). However, W. Post and H. W. Kale (pers. comm.),

most familiar with the morphological variations in the southern

subspecies, agree that there are apparently no characteristics

consistently separating A. m. peninsula and A. m. juncicola (Wakulla

seaside sparrow). The range of juncicola is contiguous with peninsula

near Pepperfish Key, Florida and extends to southern Taylor County,

Florida. Thus, juncicola should probably be merged with peninsula,

which has taxonomic priority.

In all seaside sparrows the sexes are identically colored, but males

are slightly larger. In my study at Cedar Key I routinely took wing

length (wing chord) and weight measurements. For adult males and

females the mean wing chord measurements and 95%7 confidence intervals

were 58.9 (+0.4) mm and 54.0 (+ 0.3) mm; and mean weights 22.5 (+ 0.6) g

and 19.6 (+ 0.7) g, respectively (N=134 males and 78 females). Thus

females were 937o as large as males based on wing chord measurements and

89% as large based on weight measurements. Because the 99'% confidence

intervals for wing chord did not overlap, male and female adults handled

outside of the breeding season could be sexed using wing chord. During

the breeding season the age and sex of birds could more easily be

determined using other criteria. Females possessed a vascular, edematous

brood patch about 1.8 cm wide and males had a cloacal protruberance of

about 2 mm. Fledglings and older hatching-year birds had a

characteristic paler plumage pattern.



Project History and General Methods

The study site near the Gulf coast town of Cedar Key, Florida, was

established by William Post in December 1978. Post investigated the










habitat and reproductive biology of the birds for two years, under

contract with the Endangered Species Program of the Florida Game and

Fresh Water Fish Commission. Information and results obtained from his

project were to become part of a management plan for the rare and

endangered Atlantic coast population of A. m. nigrescens. Post gridded

the marsh with stakes, began color banding of birds, mapped vegetation,

sampled the invertebrate fauna, studied the interspecific influence of

fish crows (Corvus ossifragus) and rice rats (Oryzomys palustris) on

sparrow reproduction, and recorded other descriptive data pertaining to

the morphology and life history (Post 1980, 1981a, 1961b; Post &

Greenlaw 1982; Post et al. 1983; Greenlaw & Post 1965).

Post concluded his field work in the summer of 1980. I accompanied

him to the study site several times early in 1981, and then assumed

responsibility for the study site and birds later in the spring of 1981.

During the breeding seasons (March-June) of 1981-1982 I familiarized

myself with the marsh and the birds' behavior, and continued color

banding adults and some nestlings. I spent an average of 50 days in the

field each of these two years, observing and tape recording

vocalizations.

In 1983 and 1984 I continued comprehensive observations and color

banding of my Florida study population, each year averaging about 70

days in the field from March through June. Behavioral observations on a

field day consisted of 2-4 hours of watching, recording some of the

vocalizations, and describing behavior and territories. I recorded

spoken notes on a tape recorder, and included the location of birds and

activity time (measured with a stopwatch). I later transcribed tapes

and analyzed the data.










The major investigation of my study--temporary song muting--was

conducted during the breeding seasons of 1983-1984 and is described in

Chapter IV. In the spring of 1965 I spent 15 days in the field censusing

returning and new males and conducting playback experiments with female

seaside sparrow vocalizations.

Throughout this report the data designated as having been collected

in 1979-1980 are from Post's unpublished and published work and are used

with his permission. All other data were collected and analyzed by me.

I used non-parametric statistical tests (Siegel 1956; Conover 1980) for

most analyses.



Study Site

General Description

The 30-ha salt marsh study site was north of the Waccasassa Bay of

the Gulf of Mexico. The site was 6 km NNE of the town of Cedar Key, Levy

County, Florida, and within the Sumner quadrangle, USGS Map (29 11' N,

830 00' W). The physical boundaries of the marsh, exterior to the core

gridded study area, were Prodie Creek on the SW, Live Oak Key and

Waccasassa Bay on the S, and Dorset Creek on the NE. The property was

privately owned.

The boundary between the marsh and Waccasassa Bay was not sharply

defined. The shoreline was dotted with oyster bars and many small

islands and was cut by numerous tidal creeks. Except for these creeks,

which ranged from 0.5 to 1.5 m deep at mean high tide, the marsh was

flat. Tides averaged 0.8 m. Most years one or two flooding episodes

occurred during the breeding season, destroying any existing nests of

seaside sparrows. These floods were most often generated by prevailing










winds from the south, driving water in from Waccasassa Bay, combined

with a natural (high) spring tide event. The water level during these

floods usually rose about a meter above mean high tide. Average salinity

measured by Post (1980) at 32 stations on 24 April 1980 was 19.03 + 1.09

0/00.

Weather

Data on cloud cover, air temperature, and wind direction and

velocity were collected at about hourly intervals throughout each field

day. Phenological records such as flowering dates and spring migration

observations were also kept. More complete daily weather information

(maximum and minimum temperatures and rainfall) for all years was

recorded by a National Oceanographic and Atmospheric Administration

(NOAA) station near the town pier of Cedar Key.

My behavioral observations and the handling of birds were carried

out under mild weather conditions (18-33 C, wind <20 km/h) during the

months of February through June. During my 5-year study, however, I did

visit the marsh in all months of the year and under virtually every

weather condition. Temperatures I recorded ranged from 3 C (January) to

l40 C (August). On typical sunny days during the height of the breeding

season (April and May) the mid-morning temperatures averaged 26 C. For

those field days on which I took detailed behavioral observations, the

wind velocity usually ranged from about 2 km/h (just prior to sunrise)

to about 12 km/h by 0900, and then 16-24 km/h by early afternoon.

Prevailing winds were mostly from the southwest. A few sultry field

days were virtually windless. Winds approaching gale force (50+ km/h)

occurred occasionally in the spring and fall.







8


In September 1985, Hurricane Elena touched the Cedar Key area. My

surveys the following week and in 1986 indicated no discernible

population decline in the three bird species breeding on this marsh

(seaside sparrows, marsh wrens [Cistothorus palustris], and clapper

rails [Rallus longirostris]).

Marsh Fauna

Vertebrate fauna

The six-year average population densities of the three resident

bird species were seaside sparrows-2.5 + 0.1 birds/ha, N=253; marsh

wrens--0.6 +0.3 birds/ha, N=23; and clapper rails-0.5 + 0.1 birds/ha,

N=59 (Post 1981a; McDonald 1982, 1983b, 1984, and my unpublished surveys

for 1984 and 1985). In addition to these residents, migratory

sharp-tailed sparrows (Ammodramus c. caudacutus) wintered on the marsh.

They regularly arrived in the last week of September and left 'in the

second week of May. The population density of sharptails averaged about

2/ha. Other bird species seen foraging or resting on the study site

were those commonly found in Florida Gulf Coast salt marshes (Post

1981a; McDonald 1982, 1983b, 1984).

As described below in the section on reproduction, most seaside

sparrow nest losses were attributed to predation, primarily by fish

crows and rice rats. The rice rats dwelt on the marsh. Their density was

about 8/ha (Post 1981b). I also found evidence of predation by

transient racoons (Procyon lotor).

Other species of mammals were occasionally observed: river otters

(Lutra canadensis) played on the bank edges; Atlantic bottlenose

dolphins (Tursiops truncatus) and very rarely, manatees (Trichechus

manatus) swam in the larger tidal creeks. The only reptile I saw on the










study site was the Gulf Salt Marsh Snake (Nerodia fasciata clarki),

observed on three occasions.

Invertebrate fauna and seaside sparrow foraging.

The most obvious and abundant invertebrates on the marsh were

fiddler crabs (Uca rapex) and Gulf periwinkles (Littorina irrorata).

Genoni (1984) reported additional mud-inhabiting invertebrates from this

marsh. Terrestrial invertebrates were also censused previously by

sweep-netting the vegetation during the sparrows' breeding seasons (Post

et al. 1983). By percent of total sample, these were Tettigoniidae

(54.3%), Lycosidae (18.8%), other spiders (7.8%), and Lepidoptera

(Noctuidae and Pyralidae) (3.0%). Six additional invertebrate families

each represented less than 5% of the total sample. Post et al. (1983)

found that 85Y% (by volume) of the nestling's diet consisted of these 10

groups of arthropods.

The invertebrate food species consumed by adults on this marsh are

probably nearly the same as those eaten by Scott's and other subspecies

of seaside sparrows on similar marshes. Howell (1928, 1932), Oberholser

(1938) and Wilson (1811) examined stomach contents of adult Scott's,

Smyrna (A.m. pelanota), Louisiana (A. m. fisheri, and dusky (A. m.

nigrescens) seaside sparrows. In addition to the arthropod groups listed

above, which are known to be fed to nestlings at Cedar Key (and

presumably eaten by adults), these authors found adults had eaten marine

"worms," small crabs and other crustaceans, beetles, dragonflies, flies,

wasps, bivalves, gastropods, and some "weed" and grass seeds. (Audubon

[18311 said that a pie he made of seaside sparrows could not be eaten

due to its "fishy savour," surely attributable to the birds' having

consumed mostly salt marsh invertebrates!)










With the assistance of a biologist familiar with marsh

invertebrates, I examined the stomach contents of eight adult birds from

my study site. Surprisingly, three of the stomachs were virtually empty.

These males had died early in the morning during the breeding season,

perhaps prior to foraging. The other stomachs contained mostly

arthropod exoskeletons (fragments of crabs, insect adults and pupae, and

one small spider), snails (Pyramidellidae and Marginellidae), and

vegetation. Post et al. (1983) presented convincing evidence that food

was plentiful and accessible to the Cedar Key birds throughout the year.

I rarely observed birds drinking. Several times I saw then imbibe

dew from the vegetation, and once I saw a bird drinking and bathing at

the edge of a shallow tidal creek. Although no passerines have salt

glands, seaside sparrows do have the ability to concentrate urine when

consuming salt water (Poulson 1969).

Marsh Flora

Vascular plant species on the study site were described in detail by

Post (1980). In order of relative cover, the major plants were smooth

cordgrass (medium height) (Spartina alterniflora) 38%, black rush

(Juncus roemerianus) 26%o, seashore saltgrass (Distichlis spicata) 23%,

and perennial glasswort (Salicornia virginica) 8%7. There were no woody

plants within the gridded portion of this marsh. Also, there were no

pannes (bare mud areas devoid of vegetation) until September 1985, after

the passage of Hurricane Elena near the Cedar Key region. Apparently

some stands of Spartina were submerged for at least several days during

the storm; consequently these drowning-susceptible plants died.










Size and Demarcation of Study Sites

Post established two contiguous study areas, Cedar Key West ("CKW,"

10-ha in area) and Cedar Key East ("CKE," originally 20-ha in area ).

Although CKW and CKE were contiguous, the study populations of seaside

sparrows living on each site were separated by 200-400 m, and the

populations did not interact. Both study areas were gridded with 2-3 m

high wooden stakes placed at 25 m intervals. Stiff metal prongs on the

tops of the markers discouraged their use by predatory birds. Stakes

were alpha-numerically designated and could be read from at least 50 m.

Additional markers were placed around the peripheries of the study areas

for reference. I concentrated my preliminary observations on the CKW

birds in 1981, then switched to CKE study area (which I enlarged to

30-ha in 1954) for the remainder of my project.



Reproduction and Reproductive Behavior

Overview of Annual Cycle

Fig. II-1 graphically summarizes the reproductive cycle of Scott's

seaside sparrows at the Cedar Key study site. The cycle commenced in

early March with the onset of regular singing by males and subsequent

territorial establishment and mate acquisition. The peak of the breeding

season (as indicated by the number of newly completed clutches) for the

six years 1979-1984 occurred during the second and third weeks in May.

In a typical spring the first clutches were completed around the

beginning of April, and the last clutches by the end of June.

Most clutches (855) were completed by the end of May. Virtually all

breeding and territorial defense activities ceased by mid-June. Both







12


adults and juveniles entered a prolonged molting period during the early

fall and remained difficult to observe until the following early spring.




















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Annual Cycle of Reproductive Behavior

Territorial establishment and mating

Breeding behavior of Scott's seaside sparrows near Cedar Key began

in late winter (February) with sporadic singing of adult males on warm,

windless mornings. Young adult males, hatched the previous spring, also

began to sing imperfect subsongs. By early March all males were singing

typical primary songs, and females began to associate with males.

Females solicited male affiliation with the "whinny" and "tchi" calls

(Chapter III).

During these early weeks of the season, weather influenced the

intensity of singing and other observable breeding activities. Wind

velocities greater than 25 km/h accompanied by temperatures less than

200 C caused a nearly complete cessation of overt territorial and mating

behaviors. Mild weather, even in early February, greatly enhanced

regular singing activity and behavioral interactions. In addition to

this immediate influence of atypical weather, prolonged cool

temperatures delayed the onset of breeding within a given season, and

hot weather in late May caused an early cessation of breeding

activities. The early spring of 1963, for example, was unusually cold:

average daily temperatures for March were about 50 C below normal

throughout mid-Florida. The breeding activities of the study population

at Cedar Key were about 2 weeks delayed that year-I did not find a

completed clutch until 26 April. On the other hand, the unusually warm

February of 1985 certainly must have accounted for my finding fledged

seaside sparrows on 16 March and fledged marsh wrens on 23 February,

both record early nesting dates for these species for any known breeding

locality.










Most males re-established the same territories from year to year and

vigorously defended these during March and early April, the period when

the most fervent territorial disputes between first-year and older

adults and between residents and floaters broke out. By May territorial

ownership and boundary disputes were rare, seemingly occurring only when

floaters attempted (always unsuccessfully) to take up residence on an

occupied territory. The intensity of singing (as measured in minutes of

singing/hour) also declined gradually after a peak in early April. By

June there was only sporadic singing activity occurring daily from just

before sunrise to about one-half hour after sunrise.

Most birds paired by mid-March. The mates remained paired over the

remainder of the breeding season. I never observed a natural incidence

of divorce, although Werner (1975) reported several instances of

re-pairing in his Cape Sable birds in the same breeding season. Seven

percent of 97 non-experimental territorial males remained bachelors

throughout the breeding season in my study. Werner (1975) reported that

about 12% of his 111 Cape Sable seaside sparrows were bachelors.

Apparently the adult sex ratio of the breeding birds was slightly skewed

in favor of males. Although Post (1980) assumed that most unmated

territorial males were probably first year birds, my longer study of

these marked birds indicated no significant difference in the

proportions of unmated older territorial birds and unmated first-year

territorial birds (X2=0.4, N=10 bachelors, df=l, P>0.5).

My observations over the years also suggested that returning males

tended to re-mate with their female of the previous year, if she also

returned the second year. I documented mate retention using data for

non-experimental males and females present in both 1963 and 1984, the










years for which I had the most complete data. Of the 10 cases in which

both members of the pairs were present in succeeding years, there were

five re-mates. Mate retention from year-to-year also occurs in northern

seaside sparrows on Long Island, New York (Jon Greenlaw, pers. comm.).

Werner (1975) also stated qualitatively that there was some degree of

year to year mate retention in his Cape Sable seaside sparrow subjects.

Females solicited copulations with a slow, low-pitched variation of

the "whinny" vocalization accompanied by quivering bilateral wing

raises. This display was given at ground level with the bird concealed

in the marsh vegetation. A nearby male (usually the mate) was

immediately attracted to this enticing behavior and would quickly

copulate by mounting the female briefly with fluttering wings and then

bounce off, landing several meters away. The male sometimes gave one or

two songs immediately after copulation, whereas the female was almost

always silent afterwards.

Typical bouts of copulation involved three to five mounts during

time intervals of 0.5 to 1.5 hours. All of the copulation bouts I

observed occurred in the mornings. During this period the female

sporadically gave her solicitation whinny, while the male remained

within a 15 m radius of her and sometimes sang. Of the 96 copulation

bouts I observed under non-experimental conditions, all but three were

between birds I had previously known or subsequently knew to be mates.

On the remaining three occasions I observed neighbor or floater males

sneak in and copulate with a soliciting female while the mate was

vigorously singing or confronting other birds on the opposite side of

his territory.










Nests, eggs, and early development

Although this was primarily a behavioral study, nesting and other

reproductive data were routinely collected during the breeding seasons

of 1981-1904, and in some cases were consolidated with Post's

1979-1980 data.

Seaside sparrow nests are notoriously difficult to find in salt

marshes. My early efforts to find nests by dragging a chain through the

vegetation, by trying to flush nesting females by walking transects, and

even by using a bird dog were largely unsuccessful. I eventually

learned to locate most of the nests of subject birds by close

observation of the parents.

For all nests I found, I noted date and time of discovery, parents,

location relative to the grid system and territorial boundaries, and

vegetative microhabitat. Because I wished to minimize human

interference, I handled only a sample of the nests and nestlings that I

was aware of. The additional information taken for these sample nests

was form of nest cup (open or domed), height from ground to bottom of

cup, dimensions of cup (inside and outside diameter, inside and outside

depth), material used in constructing nest, number of eggs or

hatchlings, egg dimensions, hatchling weight (for 11 individuals), age

of hatchlings, and daily survival. I also weighed seven nests collected

after fledging.

The following summarizes my descriptive data on 194 nests. Two

nests had well-defined domes and entrances. I detected no preferred

directional entry pattern of attending parents for any of the nests,

although the nests of some subspecies of seaside sparrows are mostly

domed and have distinct entrances. About half of the nests were built










in Distichlis and the remaining roughly split between Salicornia and

Juncus microhabitats. All nests were constructed primarily with the

Distichlis grass; a few had small Spartina leaves woven in. The average

height of the nest (as measured from the ground to the bottom of the

nest cup) was 35 cm. Average nest size measurements were: outside

diameter of cup 9.3 cm; inside diameter of cup 5.4 cm; outside depth of

cup 5.3 cm; inside depth of cup 3.3 cm. Also in contrast to other

subspecies, at Cedar Key the average clutch size was smaller (3.02 +

0.17 eggs, N=194 nests), compared to about 3.65 in New York (Post et al.

1983).

Average egg dimensions were 20.6 mm X 15.7 mm (N=35), and egg

weights averaged about 2 g. While exhibiting minute individual

variation, the color pattern of the eggs was essentially the same as

described for seaside sparrows in detail elsewhere (Austin 1966a).

Egg and nest cup dimensions for Scott's seaside sparrows at Cedar

Key were within the ranges reported for other subspecies in the

literature (Austin 1968b; Norris 1968; Sprunt 1966; Trost 1966;

Woolfenden 1968; Werner 1975; Greenlaw pers. comm.) and in collections I

measured at the American Museum of Natural History, the United States

Museum of Natural History, and the Florida State Museum.

I categorized nest placement within a pair's territory as "edge" or

"center." An edge nest was so designated if it was built in the outer

fourth of the total area of the territory. I found no significant

difference in the birds' tendencies to locate nests at the territory

edge versus towards the center (X 2=0.7, N=194, df=l, P>0.4).

Only 116 nestlings were directly observed, but others were known to

be in active nests. The young remained in the nest 9-11 days. Their










weights increased from 2.2 g on Day 0 to 15.1 g on Day 10. The few

other morphological data I took on these birds agree with Werner's

(1975) extensive quantitative morphological data and photographs of

nestlings taken days 1-8 for Cape Sable seaside sparrows.

Norris (in Austin 1968a) gave a detailed description of nestling

Louisiana seaside sparrows (A. m. fisher) and the behavior of the young

and their parents. My observations of Scott's seaside sparrows at Cedar

Key are very similar to Norris's in Louisiana.

resting behavior

Nest building (by the female only) began on the day of or the day

after copulation bouts. Females did not re-use nests, but built a new

nest for each new clutch. The female usually completed building her

nest within a day and laid her clutch of three eggs (rarely four) over

the next five days. The eggs were laid one a day, at any time during the

day. Infrequently females that were building nests, laying, or

incubating, would copulate with their mates. Incubation was by the

female only, for 11-12 days.

Most years the first young of the season hatched around the second

week in April (Fig. II-1). All of the young hatched within 24 hours and

began to gape and utter soft "peeps" on the day of hatching. The eyes

were fully opened by day five and at about this time the begging

vocalizations changed from a "peep" to a "churp" and became more

intense. Some nestlings fledged and were able to run about as early as

Day 9. Most began short flights about Day 13.

The hatchlings remained in the nest for 9-10 days and were fed by

both parents. The combined average feeding rate was 3.2 visits per

hour. Parents also maintained a clean nest by removing fecal sacs. Post










et al. (1983) determined the diet of hatchlings older than 3 days by

placing pipe-cleaner ligatures around their necks and then removing the

food from the young or the nest cups after an hour had elapsed. The

most important food groups for the nestlings were found to be the

insects of the taxa Tettigoniidae and Lepidoptera and spiders of several

families. These invertebrates constituted 85% of the nestlings's diet.

Both parents continued to feed the young for 15-20 days after

fledging. If the breeding season had not progressed too far (i.e. prior

to about 15 May), a female with fledged young would begin to renest

about a week after the first brood had fledged and would leave the care

of fledglings to her mate. I noted five such cases in 84 pairs.

Since most nests were destroyed before young fledged, seldom did a

pair actually have the opportunity to rear a second brood within a

season. From late March through early June, unsuccessful parents would

try repeatedly to raise a family. The average number of nesting attempts

per season of 84 pairs (1981-1984) was 3.7, and the interval between

nest destruction and renesting was 5.7 days. In 1984 I observed one

pair attempt seven nests, all unsuccessful.

Fathers began to avoid their first brood after tending their

fledglings for about three weeks, or after their second successful

clutch hatched. They chased their fledglings (as well as any other

juveniles) off the parental territory and started to devote their

attention to their new brood. After nesting had stopped family groups of

fledged, dependent young and their parents were often observed during

the months of June and July. Adults without families were seldom seen

during this period, especially during the hot daytime hours.










Post-Breeding behavior

By August most juveniles were independent and began to form loose

groups of about three to eight. Sometimes adult males associated with

these groups, which persisted through October. Juvenile males gave

occasional bouts of imperfect subsong in August and September, and

rarely through the winter. Although present (as evidenced by occasional

mist net captures), adult females were rarely seen in summer.

From late September through October there was a resurgence of easily

observable but essentially non-vocal behavior, such as foraging and

territorial defense. From these observations and my intensive mist

netting in the fall of 1982 and 1983, I concluded that adult males and

females remained within a radius of about 250 m of their previous spring

territory throughout the year. Juveniles appeared to wander a great

deal more, perhaps as far as several km.

Little is known regarding the fate of the fledgling seaside sparrows

at Cedar Key or elsewhere. Of the total of 98 hatchlings and fledglings

Post and I banded at Cedar Key, I recaptured only 4 during my intensive

netting 1982-1984. Presumably the emigration and general mobility of

fledglings as well as of non-territorial older birds are mechanisms for

the birds' locating and colonizing any unoccupied territories and viable

habitat.

The Cedar Key population appeared stable (average density 2.5

birds/ha) throughout the 7 years 1979-1985. Adult birds with territories

were relatively long-lived. By the end of 1984, I had studied subject

males from 1 to 4 years. According to banding records, two males were at

least 5 years old, and one was at least 6 years old.










The adult post-nuptial molt (and only annual molt) of their nuptial

plumage began in early July, and the post-juvenile molt began in late

July. Both molts were completed by October.

The seasonal weight variation was slight but significant. Both

sexes were heavier in winter than in summer, but the difference was not

as great as in the nominate migratory seaside sparrows (Post 1980).

Productivity, Survival, and Reproductive Success

I summarized parameters of reproduction for seaside sparrows at

Cedar Key in Table II-1. The data combine my unpublished observations

(1981-1984) with Post's observations (1979-1980), as given in Post

1980 and Post et al. 1963.

Post (1981b) intensively studied seaside sparrow reproduction and

factors that affected productivity during his 1979-1980 project. He

also compared these characteristics of reproduction in A. m. peninsula

to another well-studied population, A. m. maritimus in New York (Post &

Greenlaw 1982; Post et al. 1983). I combined the two years of Post's

data with reproductive data I collected 1981-1984.

Studies of nest productivity should consider whether nest failures

are due to human disturbance. I did not test if my visitations affected

the nesting success I observed. Post (1980), however, did test for an

observer effect using the methods of Mayfield (1975) and Johnson (1979)

and found that the frequency of nest visitation did not affect nesting

success of Scott's seaside sparrows.










Table II-1. Summary profile of territoriality and reproduction
Scott's seaside sparrows at Cedar Key, Florida, 1979-1984


Territories, Mating, and Return Ratesa
Mean territory size
Ecological density of territorial males
Mean annual return of territorial males
Proportion of re-mates
Proportion of bachelor territorial males
Ratio unmated first year/older males0
Mean length of study of male in population

Productivity and Survivala,c
Mean clutch size (C)
Mean no. clutches initiated/female/yr (I)
Probability of egg producing fledgling (S)
Productivity (fledglings/female/yr)
Mean length of breeding season
Proportion of nests producing fledglings
hest success ratio of young/older breeders
Adult annual survival male return) rate
Juvenile survival rate
Production of fledglings->breeders next yrb


1759 + 242 m2
2.5 males/ha
0.78 males/yr
0.50
0.07
4/6 N.S.
2.3 yr


3.02 + 0.17
3.7
0.06
0.67
95 days
0.06
3/5 .S.
0.78
0.67
0.45/female/yr


Combined data from Post 1980; Post et al. 19b3; and author's
unpublished observations. Probability level: h.S.=P>0.05.
bSee text for explanation of calculations.


cBased on 194 nests










At Cedar Key the probability that a nest would produce fledglings

was remarkably low, but because females renested within 7-8 days of nest

destruction, the population seemed to have remained stable from 1979

through 1985. That is, bird density remained about the same over the

years we censused the study population (Post 1980, 1981a; McDonald 1982,

1983b, 1984, and my unpublished data).

Post et al. (1983) calculated probability of an egg's surviving 21

days (through the egg and nestling period) and producing a fledgling to

be only 0.06 (Post et al. 1983). This contrasts with an egg survival

rate of about 0.35 in New York.

Of a total (McDonald and Post) sample of 194 nests whose fates were

carefully documented, only 12 (6>7) produced at least one fledgling. The

major causes of nest mortality at Cedar Key were flooding, and predation

by fish crows and rice rats. Eight (4%) of the nests failed due to

desertion. Neither Post nor I ever suspected hatchling death due to

starvation. I found only a few instances of partial nest mortality.

Eleven (2%) of the 473 eggs I observed in the field failed to hatch and

were presumably infertile.

Destruction by flooding accounted for the mortality of 17 (9%) of

the nests. Normal tidal fluctuations in water levels seldom reached the

bottom of the nest cup (about 35 cm above the ground). However, in most

years at least one major unpredictable flooding of the marsh occurred.

For the seven years (1979-1985) there were nine such floods during the

breeding seasons.

Predation by fish crows destroyed 37 (19") of the total nests. Fish

crow predation was assumed when eggs disappeared without leaving shell

fragments or a disturbed nest (indications of mammalian predation). Fish










crows were about three times more likely to prey upon nests with

hatchlings than on those with eggs.

Post (1981b) concluded that nesting in Juncus should be preferred by

the birds because Juncus stands were less likely to be flooded and

because this vegetation provided cover protection from fish crow

predation. However, rice rats on the marsh apparently discouraged

seaside sparrows from nesting more in Juncus by destroying their nests

(Post 1981b). The birds were found to nest more frequently (857 of the

total nests) in less dense Distichlis and Salicornia, even though nest

failure was more likely to occur in these vegetation types, as compared

to failures in Juncus (Post 1981b).

The remaining 120 (624) of the total nests were destroyed by rice

rats and unknown predators (probably racoons).

Post (1961b) tested whether rice rats were actually destroying nests

by placing tin cylinders around nests. These were open at the top,

allowing entry by parents and aerial predators but excluding small

mammals. Forty-eight percent of 42 experimental nests with this

protection produced fledglings, whereas only 6%a of the 34 control

(unprotected) nests had young that fledged.

To summarize: about 80% of the nests at Cedar Key were destroyed by

predation. This contrasts with findings in New York where fewer seaside

sparrow nests were destroyed by predators and more losses (about 65% of

the total nest mortality) were due to flooding or rains, desertion,

hatching failure, and unknown causes (Post et al. 1983).

I report productivity as the number of fledglings/female/year.

Post's data (1980) were combined with mine: Post observed 109 clutches,

and I observed 85 clutches. Productivity was determined by the method










of Ricklefs and Bloom (1977): Productivity = Mean Clutch Size (3.02) X

Probability of Egg Success (0.06) X Number of Clutches Initiated by a

Female/Year (3.7). (The term "productivity" and its values are roughly

equivalent to the "realized specific natality rate" parameter that is

conventionally calculated using life table data.) The six year average

productivity for 1979-1984 was 0.67 fledglings/female/year. This value

contrasts with the much higher rate of 4.25 for the New York population.

New York females produced about six times as many young within their

average 75 day breeding season as the Florida birds did in their average

95 day season. The annual survival, however, was lower in New York

(Post et al. 1983).

I determined the adult annual survival rate at Cedar Key to be about

0.78, based on the return of 51 of 65 adult males over the years

1980-1983 and assuming a roughly equal sex ratio. This figure represents

a minimum value, since some birds may have emigrated. This is a

relatively high value for a small Passerine bird. Werner (1975)

estimated a 0.88 survival rate for the likewise non-migratory A. m.

mirabilis, based on a one year return sample. Adult survival is lower

for migratory A. m. maritimus--only about 0.45 (Post 1974).

I estimated the juvenile survival rate to be about 0.67. This figure

was determined using the method of Post et al. (1983) and using their

estimated ratio of juvenile/adult survival (0.85, as determined from

returns of banded nestlings and adults). Thus, I multiplied the adult

annual survival rate of 0.78 by the ratio of juvenile/adult survival of

0.85, giving 0.67.

Although I did not have enough data to directly calculate net

reproductive rate (R0) using life table parameters, I suggest that the










Cedar Key population of seaside sparrows is maintaining its numbers for

the following reasons: First, there was no decline in population

density over the years. Additionally, my experiments in 1983-1964

demonstrated that floaters (itinerant males without territories or

mates) were always available to take up residence on territories they

perceived as unoccupied. Secondly, the reproductive data I do have

suggest that enough juveniles were produced and survived to replace

non-returning adults. That is, using my calculated productivity figure

of 0.67 fledglings/female/year and Post's (1980) juvenile survival rate

of 0.67, each breeding female (pair) produced 0.45 first-year birds.

Assuming a sex ratio of about unity, each female produced 0.225

first-year female breeders. If the minimum adult annual survival rate

was 0.78 and applicable to both sexes, then the annual adult female

mortality rate was 0.22 (1.00 0.78 = 0.22) or less. Since 0.225 is

about the same as 0.22, these reproductive data do substantiate my

observation that the population size has been constant over the years.

Post et al. (1963) calculated a net reproductive rate of 2.72 for the

increasing New York population. The authors also state that the net

reproductive rate for the Cedar Key population was 1.11 (based on

1979-1980 data), but it is unclear how this figure was determined

without life table (i.e. survivorship) data.

These New York and Florida projects are the only major long-term

studies that report and summarize detailed reproductive data. Other

papers that contain some data on breeding success and productivity for

other subspecies of seaside sparrows are Nicholson (1946), Woolfenden

(1956), Stimson (1956), Norris (1968), Sprunt (1968), Stimson (1968),

Trost (1968), Woolfenden (1968), Worth (1972), and Werner (1975).













Territories and Territorial Behavior

Definition and Methods of Determining Territories

The working definition of "territory" I used in this study was "any

defended area" (Noble 1939, p. 267). My use of the term "defended"

implies that a bird exhibited aggressive behavior towards another, with

the seeming intention of driving or keeping the other out of his or her

territory. I considered vocalizations and visual displays directed at

another bird to be defensive behaviors. This study dealt primarily with

aspects of male territoriality, although female seaside sparrows also

exhibited territorial behavior. I considered a territory to be an area

in which a male invariably challenged intruders with singing, displays,

or other overt aggressive behaviors.

All of my behavioral observations were made relative to the grid

markers. I drew territory maps for the entire study population weekly,

by connecting the points of the most peripheral singing perches with

lines. I traced the territories thus delineated with a compensating

polar planimeter and then calculated the enclosed areas. When a map

indicated a boundary had moved by 15 m or more, I considered that to be

a territorial boundary shift. The summary information regarding

territory sizes and location reported in this chapter were those

measured for non-experimental birds during the first week of May unless

indicated otherwise.

Description of Territories at Cedar Key

Territories at Cedar Key were all purpose--used for mating, nesting,

and feeding. Mated pairs exclusively occupied territories throughout

the breeding season, although boundaries between territories fluctuated.










Only two instances of complete territory relocations were noted for 65

non-experimental males studied 1979-1982.

The mean yearly return rate for male territorial birds was 797%

(N=89). There was a strongly significant tendency for males to

re-establish territories on the same sites in succeeding years. (My

conservative null hypothesis was: Returning males are as likely to

establish a new territory as they are to re-establish their old

territory. My sample sizes and statistics were: 43 re-established

territories of 52 observed returnees; X2 one-sample test, X2=21, df=l,

P<0.001). Year-to-year retention of territories has also been reported

qualitatively in the few other studies of marked populations of seaside

sparrows (Worth 1972; Werner 1975; Greenlaw pers. comm.).

The territory sizes of non-experimental birds at Cedar Key ranged

from about 200 to 4000 m2. The mean size was 1759 m2 (SE=242 m2, N=65).

This average territory size for the population did not change

significantly over the years 1982-1984 (Kruskal-Wallis one-way ANOVA,

N=43 territories of non-experimental males, df=2, H=0.83, P>0.5). In

comparing the individual males' territories from year to year, I found

no significant increase or decrease in their territorial areas (Wilcoxon

matched-pairs signed-ranks test, N=23, T=91, P>0.05).

The boundaries of territories did not follow any apparent natural

landscape divisions, such as vegetation type borders or small tidal

creeks. However, larger tidal creeks (>20 m wide) did often separate

territories.

Territorial Behavior

Several studies of northern seaside sparrows (Post 1974; Post &

Greenlaw 1975) contain a few quantitative data describing territorial










behavior. Other authors (e.g., Audubon 1831; Nicholson 1946; Norris

1968) anecdotally portrayed territorial and breeding behavior of seaside

sparrows, some in detail and many from an anthropomorphic perspective.

Those reports that best characterize territorial birds and their social

interactions are Norris (1968), Worth (1972), and Werner (1975).

At Cedar Key males began to establish their territorial boundaries

in early March by singing regularly from song perches, usually clumps of

elevated vegetation such as Juncus tussocks located at the edges of

their territories. Overt aggressive encounters were also first observed

at this time and became most numerous in April. Behaviors of such

encounters, in order of frequency of occurrence, were chasing bouts

accompanied by strident "tchi," "zuck," and rapid "tuck" calls;

supplanting (a challenger flew towards a perched bird, and the perched

bird left without dispute); and short-range "facing off" displays,

wherein the participants engaged in wing raises, bobbing, bill

thrusting, and wing and tail flick displays, co-occurring with the

"tuck" and "tchi" calls and "whisper songs."

These Early Season territory-establishment behaviors, as well as

male-female pairing chases, decreased significantly in frequency and in

duration as the breeding season progressed into May. By the end of the

breeding season (June), territorial males seldom disputed intrusions by

neighbors and floaters.

Prior to experiments I conducted in 1983-1986, Post and I believed

that there were no excess males trying to establish territories and

consequentially compress existing territory sizes. I found, however,

that there were such-males, as discussed below in Chapter IV.










Females definitely displayed aggressive behavior towards both

intruding males and other females throughout the season, although I have

few quantitative data on the extent of their involvement in

territoriality. In contrast to males, which were more likely to give

short distance aggressive displays and calls, females were more likely

to supplant and chase intruders. Of the total 3,159 instances of overt

aggressive behavior I analyzed between seaside sparrows in March and

April, 22 percent involved females. Only 5 percent were known to be

female-female encounters. Observations of such encounters were

difficult to make, because females spent more time than males in dense

vegetation. The frequency of female chases of intruders from the

pairs's territory averaged 2.7 chases per hour in late March and early

April. These chases were usually accompanied by the "tchi"

vocalizations and sometimes by a fast rendition of the "whinny"

vocalization (female solicitation call).

Reactions to Other Species of Birds and to Humans

I observed no vocal or other behavioral interactions of seaside

sparrows with resident clapper rails or marsh wrens. Each of these

species seemed oblivious to the sounds emitted by the others. Calls and

songs often overlapped, even when given within close range (<15 m). This

is in contrast to other studies of bird communities, in which singing

characteristics of several species are influenced by each other (e.g.

Popp et al. 1985). Several times I did note that seasides alternated

songs in a definite pattern (countersang) with migrating red-winged

blackbirds (Ageliaus phoeniceus) and bobolinks (Dolichonyx oryzivorus).

Cruising fish crows (notable nest predators of seaside sparrows) almost

always caused seasides to stop singing and sometimes to give "si










twitter" alarm calls (23 observations of low-flying fish crows during

song, 18 instances of song ceasing).

Wintering sharptail sparrows incited aggressive behavior from

seasides, especially when the sharptails began to sing in early May. The

seaside's aggressive behavior was identical to that directed towards

conspecific intruders. Early in the breeding season the mostly silent

sharptails were tolerated and unchallenged. On 17 occasions I heard

distinct countersinging between seasides and sharptails, although their

songs are quite different.

My study population of seaside sparrows seemed oblivious to my

presence, except when I approached to within 5 m or handled them. The

birds were more wary when I was accompanied by field assistants. Other

researchers have made similar observations regarding the "tameness" of

their subject populations of seaside sparrows (Sprunt 1968; Post 1974).

At Cedar Key curious sparrows and wrens approached within 1 m of me when

I was still for 30 min or longer. Indeed, I once awoke from a nap on the

ground to find a sparrow staring at me about 20 cm from my face!

My close approach to nests (<10m), however, did provoke alarm and

sometimes distraction displays from the parents when they were in the

nest vicinity. If the female was on the nest, she usually ran about 3 m

then flew about 5 m farther away. Presumably these reactions are nearly

the same as those given in response to other "predators." Several times

I simulated identical predator-reaction behavior by placing a mounted

fish crow model 2 m from a seaside nest. The typical response by the

parents was high pitched "seeet calls" interspersed with the "tuck"

call, given while the birds rapidly flitted around in the vegetation

within 5 m of the nest. Three times I saw females give apparent










distraction displays by trailing their wings as they ran away from their

nests.

Discussion of Seaside Sparrow Territoriality

Territory types and variation

The seaside sparrow literature indicates that different races, and

populations within races (see Post 1974), may have functionally

different types of territories. These have been described variously as

colonies (Tompkins 1941; Sprunt 1966); areas with separately defended

nesting and feeding areas (Woolfenden 1956, 1968; Worth 1972); grouped

(defended) nesting territories with undefended remote foraging areas

(Post 1974); nest-centered activity spaces ("Type B" of Wilson 1975;

Post 1960); and all purpose breeding territories (mating, nesting, and

feeding; "Type A" of Wilson 1975, Werner and Woolfenden 1983). The

occasional references to seaside sparrows' being "colonial" may be

misleading in that this designation implies the birds had very small

territories, such as those of colonial shorebirds. Although the limited

nesting habitat and the concomitant foraging behavior of some

populations give the appearance of coloniality, seaside sparrow nesting

groups are perhaps better described simply as aggregations.

I have reviewed the literature describing seaside sparrow habits and

have observed populations from Florida to New York. I conclude that

nearly all subspecies and populations exhibit the "all purpose" type of

territoriality. A few populations (e.g. at Post's (1974) Gilgo Beach

study site) exhibited variations of the all purpose type, in that the

birds sometimes foraged out of their territories on undefended areas.

Territories at Cedar Key were of the all purpose type. Occasionally










individuals were seen foraging, unchallenged, on other known

territories.

Territory quality and space use

Post and co-workers (Post 1974, 1981b; Post & Greenlaw 1962; Post et

al. 1983; Greenlaw & Post 1985) comparatively studied the

interrelationships of territory size and quality, predation, food

availability, reproductive success, and mating systems of seaside

sparrows on Long Island, New York, and at Cedar Key, Florida. Their

studies reached several conclusions with which I generally concur: Food

seemed not to be a limiting factor in determining territory size;

"Space-use patterns" (i.e. maintainence of territory size, and foraging

on and off the territories) were more influenced by the numbers of birds

attempting to settle into a habitable area than by food availability.

In other studies Post (1974) and Werner (1975) contended that it is

unlikely that territorial spacing limits the population size of seaside

sparrows. They suggested that as the density of birds increases over

the years the territory sizes simply decrease in order to accommodate

more birds. The population size at Cedar Key remained stable throughout

ny study; thus, I did not have an opportunity to support or refute this

hypothesis under the circumstances of increasing population as described

by these authors. Territorial compression either did not occur at Cedar

Key, or if it did occur then the territories must have become maximally

compressed prior to ny study. I conclude this after considering the

findings of my 1963 and 1984 experiments on territorial birds (Chapter

IV). Because I found that non-territorial males did exist, I contend

that territory boundaries and sizes are not necessarily so flexible as

to accommodate all aspiring breeders, as suggested by Post and Werner.










A few bird studies have established an inverse relationship between

territory size and resources (Zimmerman 1971; Seastedt and MacLean

1979). Greenlaw & Post (1985), however, found that the sizes of the

territories they investigated were not correlated with their composite

index of territory quality (which included nesting, food and

cover/protection factors). Furthermore, territory size was not directly

related to volume of vegetation, area of vegetation, or amount of food

on the territory (Post 1980).

Several recurrent conclusions appear in all seaside sparrow

literature describing territorial and feeding behaviors. My

observations on birds at Cedar Key support most of these: Territory

sizes and probably quality vary considerably within and among

populations. Food is seldom if ever a limiting resource, as

convincingly argued by Post et al. (1983). Animals with small

"nest-centered" territories simply forage, unchallenged, off their

territories. Both migratory and non-migratory males tend to establish

their territories in the same locations year after year. Territorial

shifts within a season rarely occur. Birds in dense populations spend

more time involved in aggressive behavior but neither spend less time

feeding nor suffer lower reproductive success as a result of crowding

(Post 1974). Male sparrows are probably not defending areas for the

food value per se, but are rather advertising and defending territories

in order to increase their chances of mating and keeping mates.

Extension of territory definition and function

Kaufmann (1983) stated that territoriality is one form of social

dominance, dominance being defined as priority of access to critical

resources (e.g. food, mates) that increase the fitness of the dominant










individual. Kaufmann further described territoriality as "relative

dominance." Individual A may be dominant to a subordinate, B, while in

A's territory, but not dominant to B when outside of A's territory.

These interpretations of territoriality are more inclusive than the

restricted working definition I gave at the beginning of this section.

They synthesize that traditional, restricted definition of a territory

as "any defended area" with the broader concept of social dominance.

Kaufmann's characterization of territoriality as one manifestation of

social dominance is quite applicable to seaside sparrows at Cedar Key.

The territories were indeed vigorously defended areas, yet a male off

his territory was invariably subordinate in another bird's territory.

Males together on the few areas of the marsh not occupied by territories

did not evidence dominant-subordinate behavior: apparently shared were

food and the rare female affiliations that occurred outside of

territories. Thus no dominance hierarchy or absolute dominance seemed to

exist.

As Kaufmann (1963) pointed out, arguments about functions of

territoriality have mostly been concerned with whether a territory

serves as an area to enhance social stimulation, or as an area to allow

resource acquisition. Evidence in support of the former hypothesis,

proposed by Darling (1952), is that some birds clump their territories

even when nearby suitable habitat is available. Considering the

synchronized breeding behavior of seaside sparrows and the observation

that populations do indeed clump their territories, one may concur that

social stimulation is a function of territoriality in these birds.

However, I do not believe this to be the case in the birds I have

observed, for the following reasons.










First, I suggest that clumping is a response to many individuals

having the same gestalt perception of preferred habitat, which

researchers have not been able precisely to identify. Since food is

apparently not limiting, I suspect that the preferred habitat for a male

seaside sparrow selecting and setting up a territory is an area of the

marsh that allows maximum singing display advertisement (i.e. moderately

low vegetation with dispersed singing perches such as tussocks of

Juncus). Singing and being able to be heard while singing are vital to

the fitness of male seaside sparrows, as determined by experiments

described in Chapter IV. Thus, what may initially appear to be suitable

unoccupied habitat may in fact be unsatisfactory for establishing a

song-defended territory. Secondly, I do not believe that social

stimulation is a function of territoriality in seaside sparrows, because

their apparently synchronized breeding behavior can almost always be

directly attributed to weather and other environmental conditions, such

as flooding.

For seaside sparrows at Cedar Key, the possession of a territory

does seem unequivocally to allow the owner priority of access to the

resources contained therein, the most important of these probably being

singing perches, a mate, and a relatively undisturbed mating/nesting

area. Thus the territories of seaside sparrows easily fit the

conventional concept of bird territories' being "defended," areas

exclusively occupied by a relatively dominant male. Also applicable is

Kaufmann's broader characterization of a territory as "a fixed portion

of an individual's or group's range in which it has priority of access

to one or more critical resources over others which have priority

elsewhere or at another time. This priority of access must be achieved







39


through social interaction" (Kaufmann 1983, p. 9). Because this

definition includes temporal and relative dominance criteria, this

definition more comprehensively characterizes seaside sparrow

territoriality.
















CHAPTER III
VOCALIZATIONS OF SCOTT'S SEASIDE SPARROWS


Introduction

Seaside sparrows (Ammodramus maritimus) have a vocal repertoire of

distinctive calls and song types. Because these birds are cryptic and

tend to remain low in their densely-vegetated salt marsh habitat, vocal

communication is important in their social behavior. Intergradations and

varying contextual uses of sounds presumably convey different

information to other birds receiving the messages.

Most older seaside sparrow literature is descriptive, and

vocalizations are reported in general and anecdotal behavioral

observations (Kopman 1915; Sprunt 1924; Holt & Sutton 1926; Howell 1932;

Stone 1937; Tomkins 1941; Nicholson 1946; Woolfenden 1956; Norris 1968;

Trost 1963; Worth 1972; Lowery 1974). Relatively recent studies of the

northern seaside sparrow (A. m. maritimus) by Post & Greenlaw (1975)

and of the Cape Sable seaside sparrow (A. m. mirabilis) by Werner &

Woolfenden (1983) include more objective descriptions of calls as well

as songs and associated behaviors. Post & Greenlaw's (1975) detailed

description of the vocal and display repertoire of their Long Island,

New York, population is especially thorough.

In this chapter I describe and discuss the vocalizations of a

Florida population of Scott's seaside sparrow, A. m. peninsuale. I

include more quantitative and conclusive data than in my preliminary

report on Scott's seaside sparrow vocalizations (McDonald 1983c). This










chapter also compares the vocalizations of my study population to those

of northern seaside sparrows described by Post & Greenlaw (1975) and to

those of Cape Sable seaside sparrows described by Werner & Woolfenden

(1983).



General Methods of Data Collection and Analysis

The project history, study site, and subjects were described in

Chapter II.

Coverage

I recorded vocalizations and described the behavior of the study

population throughout the years 1981-1984. My most intensive field work

occurred during the breeding seasons (late February through mid-June)

when I went to the marsh at least 3 days/week. I also visited the marsh

about once every 10 days during the remainder of the year. During the

breeding season I made the majority of my behavioral observations and

recordings when the birds were most active--from just before sunrise

until about four hours after sunrise. The birds' activity level

increased again moderately about an hour before sunset. The few evening

behaviors and vocalizations I recorded seemed not to differ from early

morning observations, thus I only mention their occurrence in this

chapter. Unless otherwise noted, my quantitative descriptions of

vocalizations and behaviors are for observations taken on mild spring

mornings when the temperature ranged from 18-330 C, there was no

precipitation, and the wind ranged from 0-24 km/h.










Sample Size and General Observation Methods

The total number of marked birds (1981-1984) was 213, including

birds manipulated in 1983-1984. In generalizing about the use of

vocalizations, I considered only data for 91 mated and 11 unmated birds

that I had observed for at least 15 h per season. Detailed analyses

were based on fewer males, as described below. I studied each subject

male at least one hour per week throughout the breeding season. On a

typical morning I observed 1-3 focal males for about an hour (an hour's

"time budget" block), then watched another 1-3 males for the next hour,

and so on up until about 0900. I randomly determined the sequence of

these time budget observation blocks. Thus, all birds received

approximately equal coverage that was representative of the morning

hours. Females were difficult to observe; I described their behaviors

and vocalizations as parts of their mates' time budgets.

During the time budget I dictated all observed behaviors and the

birds' locations on cassette tape. I made most recordings (including

all used for sound analyses and sonograms in this section) with a Sony

TC 150 cassette tape recorder (Mineroff-modified), using a Bell and

Howell "Shotgun" unidirectional electret condenser microphone with

windscreen. I measured the amplitude (in dB) of a singing male 7 m from

me with a Bruel & Kjaer Sound Level Meter (Type 2219) on 5 Apr 1985.

I also used field data forms to record contextual information,

banding records, nest descriptions, territorial boundaries, and other

biological information. Territory maps for the entire population were

drawn in the field at least bi-weekly throughout the breeding seasons.










General Analysis of Notes and Recordings

I transcribed, tabulated, and summarized the behavioral data. I

determined characteristics of tape recorded sounds (e.g. frequency

ranges, length of song) with a Uniscan II FFT Real Time Sonogram

Spectral Display (Model 4600), and prepared sonograms (Fig. III-i

through Fig. III-10) with a Kay Elemetric 7029A Sono-graph using the

wide-band filter (300 Hz) and either the 80-8000 Hz or the 1600-16,000

Hz scale. The specific vocalizations and behaviors I analyzed are

listed in Table III-1.

Detailed quantitative descriptions of vocalizations and other

behaviors were determined from an analysis of 103 time budget hours of

30 mated and territorial best-studied males. I chose time budgets for

analysis based on comparability of day of year (Apr-May), time of day

(sunrise-0900), and the typical mild weather conditions described above.

I used non-parametric statistics (Siegel 1956; Conover 1980) for the

analyses.

My specific methods of investigating flight songs ("complex flight

vocalizations" of Post & Greenlaw 1975) and countersinging are discussed

in separate sections below.










Table III-1. Vocalizations and behaviors measured during time budget
observations (in min and sec duration, and frequency)


Singinga,b
Primary song
Whispering song
Matched countersinging
Not matched countersinging
Flight song

Other Vocalizationsa ,b
Tuck
Tsip
Si Twitter
Seeep
Tchi
Tchi flight
Whinny
Zuck and other calls

Position Changesab
When not singing
Singing, song type change
Singing, no song type change
Singing, stop singing


Observations Relating to Females


and Younga'b


Female present
Female solicitation calls, other calls
Copulations
Females building nests
Other behaviors related to females and young

Other Behaviora,b
Feeding, resting, and preening
Off territory: feeding; invading nearby territory

Reactions to Intrudersa,b
Frequency, duration, and location of intruder
Flight chases
Vocalizations: Displays:
Solo primary song Sham-preen
Countersong Grass-pick
Whispering song Bob and bill jab
Calls (as above) Contact

a Contextual information also noted: bird's identity, location, mated
status; presence of conspecifics, date, weather, and location of
observer.
bTerminology follows Post & Greenlaw 1975.












Results and Discussion of Calls and Song

Descriptions and Use of Vocalizations

Scott's seaside sparrows make 14 distinct vocalizations: a primary

song, three modifications of the primary song, and 10 calls. The

probable functions of their songs and calls are summarized in Table

III-2. I distinguish "songs" from "calls" as implicitly defined by most

ornithologists, and as specifically stated in Pettingill's working

definition: "Song is a vocal display in which one or more sounds are

consistently repeated in a specific pattern. It is produced mainly by

males, usually during the breeding season. All other bird vocalizations

are collectively termed call notes or, simply, calls" (Pettingill 1970,

p. 319). I use Post & Greenlaw's (1975) vocalization terminology

throughout, except for "whispering primary song," which is from Werner &

Woolfenden (1983).










Table 111-2. Vocalizations of Scott's seaside sparrows


Vocalization
a
Name-
Primary song


Whispering
song

Complex flight
vocalization

Subsong
(young birds)

Tuck


Tsip call and
Si twitter

Seeep note



Tchi
vocalization


Whinny
vocalization

Zuck call


Scree call

Begging and
Chup calls


Sex and
Context-
Male;
1, 2, 4

Male;


Male;
1, 2, 5

Male;
1, 4

Male and Female;
1-5


Male and Female;
1-3, 5

Male and Female;
1, 2, 5


Male and Female;
1-3, 5


Female;
1, 2, 4


Male and Female;
1, 5

Male and Female; 3

(given by young
only)


Probable Function(s)

Territory defense and
mate attraction

Territory defense
during intrusion

Territory defense and
mate attraction

Practice song


General purpose;
moderate aggression;
chase; nest defense

Heightened aggression;
attack; nest defense

Proclaim location of
female(?); fear(?);
Flocking (fall & winter)

Moderate aggression;
chase; female: nest area
distraction, proclaim sex


Attract males for
copulation


Heightened aggression;
chase; drive off young

Extreme distress call

Begging for food


aTerminology follows Post & Greenlaw (1975). This table is modified
from Post & Greenlaw's (1975) Table 3, which summarized "vocal
displays" of northern seaside sparrows.
bcontexts in which the vocalization were elicited: 1-in presence of
intruder or neighbor bird; 2-in presence of mate; 3-in presence of
predator, or human disturbance; 4-also given when bird apparently alone;
5-given in flight.











Calls of Scott's seaside sparrows

In this section I give sonograms, descriptions, and suggested

functions of 10 calls, presented in approximate order of their frequency

of occurrence during the breeding season.

Tuck call

The "tuck" (Fig. III-1A) was the most common of the calls. It was

given by males and females throughout the year. The short tuck covers a

wide frequency range and is probably the same call Werner & Woolfenden

(1983) labeled as the "chip" call for Cape Sable seaside sparrows. The

tuck appeared to be an all-purpose vocalization, but seemed to function

mostly as a general alarm and moderately intense aggressive call. It

was invariably given during short-distance aggressive interactions and

usually accompanied by wing and tail flicks, and by the "tsip" calls

(shown specifically in Fig. III-1A).

Males gave tuck calls at an average rate of 0.3 min/h when there was

no apparent disturbance on their territories. During intrusions,

however, the tuck rate increased to 13.5 min tucking/h intruder present.

This call was also given alone (or together with "tsips") by both

parents when a predator or human approached a nest within 5 m. The tuck

call was the only one I heard regularly during the non-breeding season.

I also heard fledglings and juvenile birds give this call.

Tsip and si twitter

Both males and females also gave the higher-pitched, short "tsip"

call (Fig. III-1B). When birds were highly agitated the "tsip" was

rapidly repeated, thus intergrading into the "si twitter" call. The

"tsip" was often interspersed with "tuck" calls, as mentioned above, and










was accompanied by wing and tail flicks. During bouts of calling, which

lasted 0.25 to 17.3 min, birds gave "tucks," "tsips," and combined

"tuck-tsips" at a rapid and usually regular rate of 30-145 calls/min.

As with the "tuck" call, "tsips" and "si twitters" generally signaled

alarm and aggression, but more often indicated a higher level of alarm.

My closest approach to nests (<2 m) and my placing of a stuffed fish

crow near nests nearly always elicited "si twitters." I never heard

"tsips" or "si twitters" from undisturbed birds. During intrusions the

average combined "tsip-si twitter" rate was 4.5 min tsip-si twitters/h

intruder present. I only rarely heard the "tsip" call during the

non-breeding season, and I never heard fledglings or juveniles give

either of these calls.

Seeep note

During the breeding season I occasionally heard females give a soft,

high-pitched "seeep" call Fig. III-1C). Most of the "seeeps" occurred

during bouts of copulation. Sometimes a female gave this call as a

nearby male began to sing relatively softly (Fig. III-2A). Post &

Greenlaw (1975) suggest that the "seeep" call conveys a weak fear

message. This call, or a very similar one, was the second most common

vocalization I heard outside of the breeding season. Males and females

of both Scott's seaside sparrows and wintering sharptail sparrows at

Cedar Key gave "seeeps," especially in the late fall and early winter. I

agree with Post & Greenlaw (1975), and with Werner & Woolfenden (1983),

that this call probably functions as a social and flocking vocalization

outside of the breeding season.










Tchi vocalization

The loud and relatively short "tchi" call (Fig. III-2B) was given by

both sexes and was most often heard during chases. About half of the

total thisi" I heard were given during or immediately following chases.

Usually the thisi" were rapidly repeated, and frequently they

intergraded with a slower, slurred "tyu." The "tyu" itself often slurred

into a "whinny"-like vocalization (described below). The "tchi" call

was probably the same as the "squeaz" call described by Werner &

Woolfenden (1983), and the same as the "jee-jee-jee-jee-jee-jeeeu-jeeeu"

call described by Norris (1968).

The primary function of this call seemed to be signalling general

aggression during chases. Frequently, however, I heard females give

rapid thisi" in flight ("tchi flight") as they left a nest they were

building or incubating, even though no threatening bird seemed to be in

the vicinity. These nest-departure thisi" were not given adjacent to

the nest, but rather at about 10-15 m from the nest. During a 1 h

observation of nest building in early May 1983, for example, I saw a

female depart and come into her nest 18 times. Fifteen of her

departures were accompanied by thisis" but only one arrival was

accompanied by this call. Jon Greenlaw (pers. comm.) reports that female

northern seaside sparrows behave in a similar manner. I suggest,

therefore, that the "tchi" also functioned as a distraction call by

drawing a potential predator's attention away from the nest as a female

departs. Jon Greenlaw (pers. comm.) also thinks that thisi" in the

nest vicinity function to let the resident male know that this departing

and arriving bird is a female who belongs, rather than an invading male.










Other than the females giving the "tchi" as described above, I

seldom heard this call when no intruders were present (0.11 male

tchis/non-intrusion hour). It was difficult to quantify the "tchi" rate

during intrusions. On the average, however, a territory owner spent

2.62 min chasing/h intruder present. I estimate that at least half of

these chases were accompanied by rapidly repeated thisis" most of these

calls given by the owner.

Whinny vocalization

I heard only females give the "whinny," with one unusual exception

described below. This whirling, quavering call varies in rate from 0.3

to 3/sec (Fig. 111-3). The whinny was the female's solicitation call;

it functioned to attract a male. Females gave this call more often when

a male was relatively close to her (within about 7 m), and especially if

the male was singing. Males were immediately attracted to the "whinny"

vocalization, and copulation usually followed. Even though females gave

the "whinny" from on or near the ground, males seemed to have had no

problem finding the vocalizing females. It is likely that some auditory

characteristics) of this vocalization made it easy to locate.

Females gave this call most often prior to, and during nest

building. Because females usually initiated up to 5 clutches per

breeding season, the overall intensity of "whinnies" heard for the

population remained the same from late March through May. The average

whinny rate heard on all territories during these months was 2.02 min

whinnies/h.

Post & Greenlaw (1975) describe the postures associated with the

"whinny" vocalization and copulation. Werner & Woolfenden (1983) do not

describe a female solicitation vocalization.











I only once heard a male give a distinct "whinny" (as opposed to the

"whinny"-like "tyu," described above). On 6 May 1984 a male invading an

experimentally muted bird's territory repeatedly gave whinnies while he

intruded.

Zuck call

The loud, raspy "zuck" (Fig. 111-4) was an aggressive call given

during intense fights. Most of the zucks I heard were given by males,

but sometimes females and juveniles gave this call. It may have been the

same as the "Shu-shu...shu" call of Cape Sable seaside sparrows that

Werner & Woolfenden (1983) mention but do not describe. In addition to

its functioning as a general, high intensity aggressive call, "zucks"

were the vocalizations most often given by fathers when they drove their

older, almost independent fledglings off their territories (about 2-3

weeks post-fledging). Other than these father-fledgling encounters, I

never heard this call given during non-intrusion circumstances. During

intrusions both territory owners and intruders gave "zucks" (often mixed

with thisis) at the rate of 0.7 min zuck/h intrusion.

Other calls

The "Scree" (Fig. 111-5) was a harsh distress call I heard only

(with a few rare exceptions) from a few birds trapped in mist nets and

being handled. Both sexes of adults and juveniles gave this call.

The monotonous and insistent "Begging Calls" (Fig. 111-6), given by

older nestlings and fledglings, were heard most often towards the end of

the breeding season. Nestlings gave "Peep" (Werner & Woolfenden 1983)

and "Chup" (Post & Greenlaw 1975) calls.

Post & Greenlaw (1975) describe one further call I have not

mentioned--the "Chew" call. To my knowledge, I never heard this call at







52


Cedar Key. Jon Greenlaw (pers. comm.) proposes that the "Chew" call is

probably a modified "tchi-whinny" vocalization.

















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Primary song

Audiospectrogram. The seaside sparrow primary song (Fig. 111-7) is

relatively simple. The introductory portion of rapid clicks (centering

around 6.5 kHz) is immediately followed by a buzzy trill (centered

around 3.8 kHz). These frequency ranges and the song duration of about

1.0 second were characteristic for the entire population.

General description of singing behavior. During my study, only

males sang. Although Post & Greenlaw (1975) describe "female songlike

vocalization" that resembles male song, I never heard a vocalization of

this type at Cedar Key. Aales sang to delineate territories and to

attract females. Experimental evidence for these functions of song is

given in Chapter IV.

Males usually sang from elevated perches, such as tussocks of

Juncus. Most birds had 3-7 favorite singing perches within their

territories from which they gave about 35% of their songs. From

1979-1984 only two males sang from grid marker stakes, although Post &

Greenlaw (pers. comimi.) found that birds in their dense study population

often sang from grid markers.

The singing pattern of seaside sparrows is similar to that of many

song birds (Table 111-3). Birds sang in "discontinuous" patterns of song

"bouts" (songs repeated continuously without a pause), of "eventual

variety" (song type change after three or more repetitions of the same

song type) (Hartshorne 1956; 1973). An average bout throughout the

season consisted of 19.2 songs and lasted 1-5 min (mean = 1.62). Highly

variable periods of silence (mean = 7.9 min) separated song bouts. Males

changed perches during 17% of these between-bout silences, before

resuming singing. Song rate within a bout ranged from 6-9 songs/min










Table 111-3. Characteristics of solo primary singing and countersinging
of 30 mated male Scott's seaside sparrows during April and May



Primary
Song (Solo) Countersong b
Characteristics Total Total Matched Not Matched


N (bouts) 391 223 143 80

Mean bouts/h obs. 3.80 2.17 1.39 0.73

Total min sing 635 631 438 192

Mean min sing/h obs. 6.16 6.13 4.26 1.86

Mean min song/bout 1.62 2.83 3.07 2.40

Total bouts edgec 198 123 30 43

Total bouts center 117 41 26 15

Total min edge 315 369 275 94

Total min center 193 110 66 44

Mean song type <0.01 0.02 0.02 <0.01
switches/bout

Mean song type 0.32 0.33 0.47 0.29
switches/h sing

Mean perch 0.97 0.33 0.31 0.35
changes/h sing

Mean perch changes/ 0.16 0.02 0.02 0.01
bout

Mean perch changes/ <0.01 <0.01 <0.01 <0.01
bout with song switch

Mean perch changes/ 0.03 0.02 0.02 0.01
bout without switch

Mean perch changes/ 0.13 0.05 0.01 0.02
bout, song cessation







75



Table 111-3, continued.

Characteristics of solo primary singing and countersinging determined
from analyses of 103 h of time budget observations, when no intruders
were present on focal birds' territories. Other conditions during time
budget observations are described in the text.
bRate values calculated for Matched and Not Matched columns based on
number of bouts and minutes of Matched and Not Matched singing,
respectively. Thus the sums of rates of Matched and Not Matched values
do not add up to the Total column for Countersong.
c
Edge defined as the outer-most 10 m of the bird's territory.

dCenter defined as the area within a 25 m radius of the estimated center
of the bird's territory.










(mean = 7.7), close to the 6.6 song rate Woolfenden (1956) reported for

New Jersey birds, and slightly lower than the 10.6 song rate reported by

Post & Greenlaw (1975).

Individual males had a repertoire of two to four similar song types

(mean = 2.41). When both solo singing and countersinging, birds seldom

switched song types within or even between song bouts. The mean rate of

song type switches during a bout was about 0.01/bout. Thus their

singing was extremely "monotonous," sensu Hartshorne (1956; 1973),

because their singing pattern was both continuous and of low

"versatility" (infrequent song type changes).

Most males shared their song types with nearby territorial birds. A

few of the birds had one highly distinctive song type within their

repertoire that enabled me to recognize individuals aurally from as far

away as 150 m. I did not detect any tendency for males to have a

preferred song type from within their small song repertoire.

Seaside sparrows at Cedar Key seldom began a new song type after

moving to a new perch while singing (Table 111-3). Of a total 67

incidents of perch changes during singing, there was only one concurrent

song type change. This constrasts with Krebs's (1977a, 1978) "Beau Geste

hypothesis" that territorial birds switch song types when changing

perches, thus possibly giving potential intruders the impression that

territories are saturated.

Influence of abiotic factors on singing behavior. Singing activity

varied throughout the year. It was most intense relatively early in the

breeding season (late March through mid-April), especially before and

just after birds were mated. The average time singing of 29 males

measured 1-15 Apr 1983 was 19.4 min singing/h. In the third week of May










their song rate decreased to 12.1 min singing/h, a significant

difference (two-tailed Wilcoxon matched-pairs signed-ranks test, T=2,

P<0.01).

Weather modified the intensity of singing, on both a daily as well

as a seasonal basis. Very early in the spring (February through mid

March) birds sang more as the day warmed slightly, up to about 200 C.

During the middle and especially towards the end of the breeding season,

birds sang less as the day warmed. Virtually all singing stopped at

temperatures above 320 C, regardless of the time of day. From the middle

of the breeding season till the end, birds sang more on cloudy, cool

mornings. Light rain also enhanced singing activity at this time of the

year, and even heavy rain and thunder and lightning dampened singing

activity only slightly. Brisk winds (>25 km/h), however, greatly

reduced singing activity, especially early in the season on cool days.

The intensity of singing activity also varied throughout the day.

Peak song activity was early in the morning, from sunrise to about 3

hours after sunrise. The earliest I heard singing was 20 min before

sunrise. Birds also sang in late afternoon until about one-half hour

after sunset. Compared to seaside sparrows on Long Island, New York,

that I observed in June 1982, birds at Cedar Key sang less in the

evenings. The first morning and last evening songs were similar to

songs given during peak morning activity, except the song rate

(song/min) within the bout was lower very early and late in the day.

Influence of other birds on singing behavior. Unmated males sang

significantly more bouts than mated males, and thus their total time

singing (21.4 min/h) was greater than that of mated males (12.3 min/h),

as measured in May (two-tailed Mann-Whitney U-test, N=30 and 11,











P<0.01). Singing rate also differed significantly according to whether

an intruding male seaside sparrow was present. After territories were

well established (by mid-April), intrusions were rare (mean = 0.7 min

intrusion/hour). When an intrusion did occur an owner's first reaction

was increased singing, and this was often successful in repelling the

intruder. If the intruder was not driven off by normal primary song,

territory owners switched to short-distance and more aggressive

vocalizations and behaviors--whispering primary songs, tuck and si

twitter calls, chases, quivering wing raises, and (rarely) contact

fights.

The "whispering primary song" (Werner & Woolfenden 1983) was a

slowly delivered (mean=3.5 song/min), low amplitude, and slightly lower

pitched modified primary song. It was usually given by a male perched

near the ground and confronting an intruder at 1-5 m. The whispering

primary song was delivered with an almost closed beak, in contrast to

the normal primary song where the bird threw back his head and opened

his beak wide as he sang. I do not have data describing reactions of

males that heard these whispering songs. Because I only heard it during

agonistic encounters, this vocalization may have functioned to transmit

a message of threat and likelihood of attack. Jarvi et al. (1980)

described a similar use of modified primary song in the willow warbler

(Phylloscopus trochilus) that transmitted a message that the singer was

likely to attack his opponent.

Because short-distance encounters were more often accompanied by

calls and displays, the combined normal and whispering primary song rate

during intrusions (2.4 min song/h intrusion) was much lower than the

song rate when no intruder was present (12.3 min song/h non-intrusion).










Calls of most other marsh birds, including the raucous clapper

rails, seemingly had no effect on the seaside sparrows' vocalizations or

other behaviors. Seaside sparrows did respond antagonistically toward

vocalizing sharp-tailed sparrows (Ammodramus c. caudacuta) when these

wintering migratory birds began singing prior to their departure the

first week in May.

Subsong

Young males sang rudimentary subsong late in July and in August

(Fig. 111-8). These songs consisted of garbled warbles and parts

(usually the terminal trill) of the adult primary song. Subsong was not

as loud as the primary song and was sung only sporadically instead of in

bouts. These first song attempts improved and came to resemble the

adult's primary song by late fall, when virtually all singing activity

stopped. Both yearling and adult males began to sing again in late

February, and songs of the first year birds could still be

differentiated from those of older birds. By mid-March, however, songs

given by first-year and older birds could no longer be distinguished.

Countersinging and Repertoire Use

Introduction and comments

Countersinging occurs when two, three, and sometimes four male birds

alternate their songs in a regular pattern. Countersinging is described

as "matched" when the same song type or types are sung by the

participating birds. The behavioral and evolutionary significance of

song repertoire sharing in general, and of song matching during

countersinging in particular, has been the subject of recent stimulating

studies (Krebs & Kroodsma 1980; McGregor et al. 1981; Payne 1982; Kramer

et al. 1985; Searcy et al. 1985).










Many relatively simple explanations for song matching exist:

Matching indicates high-intensity interactions (Lemon 1968b). Birds

match in order to monitor the positions of their neighbors (Lemon

1968a). Relative dominance between countersinging birds influences the

degree of matching (Kroodsma 1979). Time intervals between songs of

countersinging birds indicate which of the participants is dominant

(Smith & Norman 1979). Matching functions as a vocal threat that

regulates distance between birds (Todt 1981). Matching is a graded

signal used in territorial encounters (Krebs et al. 1981). Finally,

matching indicates readiness to interact aggressively (Horn & Falls

1986).

Further interpretations of song sharing that deal more generally

with repertoires and repertoire sharing (which may or may not involve

matched countersinging) are: Larger song repertoires make recognition

of neighbors more difficult (Falls & d'Agincourt 1981; Searcy et al.

1981). Larger repertoires are more attractive and stimulating to

females (Howard 1974; Searcy et al. 1982). Larger repertoires allow

resident birds to fool potential intruders through increasing the

perceived density of singing birds (Krebs 1976), and, young indigo

buntings (Passerina cyanea) mimic the repertoires of older territorial

males and thus deceive other males through mistaken identity (Payne

1982). Searcy et al. (1985) found no correlation between song repertoire

size and male "quality" in song sparrows (Melospiza melodia).

While these may or may not be adequate explanations for song sharing

in birds, some recent experiments on song sharing have become

increasingly sophisticated. These studies have incorporated more

objective methods of determining song similarity, the possible effect of










sound degradation from sender to receiver influencing behavior, the

birds' natural use of their repertoire, and their familiarity with

neighboring males (Falls et al. 1982; Schroeder & Wiley 1983b; Whitney &

Miller 1983).

Almost all investigations of song matching and repertoire use have

involved playback experiments and measures of response strength to the

tape recorded songs. Ideally, field studies to determine the occurrence

and significance of song sharing (without using controlled playback

experiments) would be on birds with only one or two songs in their

repertoires. Usually, though, this is not the case. Therefore a major

design consideration in natural observational investigations is first to

make comprehensive studies of normal singing behavior of undisturbed

birds. That is, the song repertoires (and the possible differential use

of songs from within the repertoires) of all subjects must be thoroughly

analyzed, and the probabilities that each bird will sing a given song in

different circumstances determined. Simply describing the composition of

the bird's repertoire is not adequate.

Studies of the birds' natural use of song types are complicated by

additional considerations. The observer's method of determining degree

of song similarity may greatly influence the results. Also, some birds

are known to use different song types according to: location within

their territory (Lemon 1968b; Lein 1978; Schroeder & Wiley 1983a),

whether a male or female is present (Smith et al. 1978; Schroeder &

Wiley 1983b), distance from rival conspecifics (Simpson 1985);

familiarity with the neighboring male (Wunderle 1973; Falls et al.

1982); similarity of rival male's song to self song (McArthur 1986),

rate of song type switching (Simpson 1985), acoustic degradation over











distance (Morton 1982; McGregor et al. 1983); overall aggressive

motivation (Lein 1978; Schroeder & Wiley 1983a); and stage in breeding

(Falls & Brooks 1975; Petronovitch et al. 1976). Furthermore, three

recent studies have shown that the intensities of agonistic interactions

were not correlated with the choice of song types per se, but rather

with the rate of song type switching alone (Kramer et al. 1985; Simpson

1985) or together with the degree of matching (Horn & Falls 1986).

Thus, what may appear to be unambiguous results from studies of how

birds use their repertoires, including song matching during

countersinging, should be carefully evaluated. Some recent studies

have, however, more scrupulously investigated countersinging and song

type use in general based on detailed analyses of repertoire composition

and contextual use (McGregor & Krebs 1982; Schroeder & Wiley 1983a,

1983b; Simpson 1984, 1985.; Lemon et al. 1985).

The purpose of these introductory remarks is several-fold. First, a

field study seeking to describe and interpret the function of repertoire

sharing and countersinging is likely to be more complicated than is

intuitively obvious. The design must consider many variables that may

be difficult, if not impossible to control. Secondly, I introduce this

section on the countersinging of Scott's seaside sparrows with these

remarks in order that the reader may realize why I present only

descriptive summaries rather than the testing of the significance of

repertoire sharing and matched countersinging.

Methods of investigating repertoires

When birds countersang during time budget observations, I noted

whether they matched or not, in addition to taking my standard singing

behavior measurements (bird's identity, presence or absence of










intruders, time singing, perch changes while singing, and incidence of

song type switching). As described in the "General Methods," I

tabulated in detail 103 time budgets (selected for comparability of

date, weather, absence of intruder, etc.) for 30 mated males observed

Apr-May 1981-1984.

Results and discussion of repertoire use

I measured parameters of solo primary and countersinging activity of

Scott's seaside sparrows (Table 111-3). The mean total singing activity

(solo singing and countersinging) for males was 12.29 min song/h.

Two to four males often participated in bouts of countersinging, the

alternation of songs separated by 3-8 sec (Fig. 111-9). The pattern of

song alternation was usually regular, but in 3% of the countersingng

bouts the alternating songs drifted out of phase and began to overlap.

Countersinging comprised about half of the total singing time.

Approximately two-thirds of the total countersinging time was matched

countersong (Table 111-3).

The mean song repertoire size of the 30 males was 2.41

songs/bird/season. I did not measure the proportional use of song types

within repertoires, nor did I measure the degree of repertoire sharing

within the population.

Some conclusions are apparent from the data presented in Table

111-3, even without statistical testing. First, apparently neither solo

singing nor countersinging was preferred in the several contexts

analyzed. That is, countersinging did not seem to be used

proportionately more often than solo singing at territory edge, nor did

perch and song type switches seem to be correlated more or less strongly

with countersinging. The same lack of differential patterns according










to context seemed to exist when comparing matched to non-matched

countersinging. A possible exception is that birds may have sung more

minutes of matched song than non-matched song at territory edges.

Of course more comprehensively analyzed data are needed before

statistically valid statements regarding repertoires and countersinging

in seaside sparrows can be made. I do, however, conclude the following

at this time. First, countersinging is a form of vocal "duelling" that

indicates a slightly elevated aggressive or aroused state. That is,

countersinging is a graded signal in seaside sparrows, but probably a

less strongly graded signal than in most birds. Second, song matching

while countersinging and repertoire sharing in general in these birds

are probably chance events, analogous to the "epiphenomena" of song

pattern variety suggested by Wiens (1982) for sage sparrows (Amphispiza

belli). Songs of seaside sparrows may be observed to match simply as a

consequence of their small repertoires of simple and similar songs,

their low population turnover rates, and their sedentary habits.

Flight Songs: Description and Comparison to Perch Songs

Description

An early Florida naturalist, Donald J. Nicholson aptly

characterized seaside sparrow singing: "Towards the end of March the

marshes are fairly buzzing with the purring, wheezing songs...He tries

one perch and seeks another fifty or seventy-five yards away flying low

over the rank growth. Every so often he fairly 'explodes' with passion

leaving his concealment to rise on fluttering wings sixty or seventy

feet above the marsh uttering his erratic little song as he goes up and

down dropping out of sight in the salicornia" (Nicholson 1946, p. 41).











The flight song (Fig. III-10), accompanying the flight song display,

is a combination of preliminary calls followed by the primary song. This

vocalization and display of about 3-4 sec is sporadically given by males

throughout the breeding season. It begins with introductory

high-pitched "si" calls, followed by lower-pitched "tuck" calls, and

ends with one or two condensed versions of the primary song. The buzzy

trill of the primary song portion is about 0.4 sec shorter than the

trill of the normal primary song. The introductory calls vary in

duration and type, but are always given on the ground and during the

bird's ascent to about 10-15 m. The primary song is uttered as the bird

descends. Woolfenden (1956), Trost (1968), Post & Greenlaw (1975), and

Werner & Woolfenden (1983) also describe singing and flight songs of

seaside sparrows.

Methods of investigating flight song activity

My observations of Scott's seaside sparrows during the breeding

seasons of 1981-1982 suggested that overall song activity seemed to be

affected by whether the birds were mated, and by time of day, time of

year, and weather. In most Passerines these factors do influence the

intensity of primary song singing from a perch (hereafter designated in

this section as "perch singing"). I measured and compared perch song

and flight song activities of my study population during the early

breeding season of 1982. Then I tested whether the above factors

differentially affected flight song activity (as compared to perch

singing) by comparing the relative frequencies of flight songs to perch

singing under specified conditions described below.

I sampled the overall singing activity of 28 males during 31 field

days from 12 Feb-26 Apr 1983. Throughout the day I monitored flight










song and perch song activity within one hour time blocks from pre-dawn

through mid-afternoon. The time blocks were designated in hours

relative to sunrise: the first hour block of a day began 15 min before

sunrise, the second hour block began 45 min after sunrise, and so on. I

randomly chose and timed the perch singing of a different male during

each hour block. Simultaneously, I counted all flight songs heard within

100 m during the hour. As I counted and timed songs during each hour

block, I also noted wind velocity, temperature, the singing bird's

location, and identity and mated status (if known).

In summarizing my data, I defined an hour's "flight song activity"

as the average number of flight songs heard within the 100 m radius

during the hour. I defined "perch song activity" as the average number

of minutes the randomly chosen birds sang during the hour within the

same 100 m. Then, for every level of increment of the variables being

investigated (e.g. for temperature category 5-9 C), I tabulated flight

song activity (FS), perch song activity (PS), and a calculated FS/PS

ratio. A statistical analysis using multiple regression would have been

the ideal method of exploring these data. Yet because some data were

ordinal rather than interval and normality was not assumed,

non-parametric statistical analyses were required. In order to validly

consider the effect of only one variable at a time, I chose subsets of

data, as described below for each analysis. I used Spearman rank

correlation (Siegel 1956) to determine if PS, FS, and FS/PS ratios were

correlated with day of year, time of day, temperature, and wind

velocity. I tested significance of the correlation coefficient at the

two-tailed P<0.05 level. I also compared FS and PS activities of mated
2
and unmated birds with a two-tailed X one-sample test (Siegel 1956).










Results of singing activity analysis

Males sang perch songs at an overall rate of 17.1 (SE=5.9) min

song/h. Flight song rate varied considerably among males. The mean rate

was 0.15 flight songs/male/h. The rates of perch songs, flight songs,

and FS/PS ratios did, however, vary significantly as detailed below.

Four variables correlated significantly with both perch singing and

FS/PS ratios: day of year, time of day, temperature, and wind velocity.

Mated and unmated birds differed significantly in the number of flight

songs given.

Day of year. I analyzed the following subset of data to determine

the correlation of singing activity with day of year: 88 h of

measurements for unmated birds taken on 19 days when the temperature was

15-24o C and the wind was <24km/h. I found that perch song activity

increased significantly (r=0.72), and FS/PS ratios (r=-0.81) decreased

significantly as the breeding season progressed, up until the last week

of April. Flight song activity considered alone, however, did not

correlate significantly with day of year (r=-0.01).

Time of day. I determined the effect of time of day by considering

the same subset of observations: 88 hours of measurements for unmated

birds noted when the temperature was 15-24 C and the wind was <24km/h.

I found perch song rate was greatest just after sunrise and declined

significantly (r=-0.95) as the day progressed (excluding evening

singing). Flight song activity remained about the same all day,

although the FS/PS ratio increased significantly (r=0.33) through the

day (Fig. III-ll).

Wind. I analyzed the following subset of data to determine the

effect of wind: 79 morning hours (hour blocks 1-5) of measurements for










unmated birds noted when temperatures were 15-24 C. I found that brisk

winds had a definite dampening effect on perch song activity (r=-0.97)

but no significant effect on flight song activity. The FS/PS ratio

increased significantly (r=0.88) as wind velocity increased (Fig.

111-12).

Temperature. To determine the influence of temperature, I

considered 77 hours of observations made on unmated birds, during hour

blocks 1-4, and when the wind was <24km/h. I found that cooler

temperatures significantly (r=0.88) decreased perch song activity but

had no significant effect on flight song activity. The FS/PS ratio did

significantly decrease (r=-0.83) as temperature rose (Fig. 111-13). In

other words, as with high winds, birds gave proportionally more flight

songs in cooler weather.

Mated status. I considered the singing activity of 23 males before

and after they were mated. The time elapsed between measurements was

8-19 days. I observed birds for 73 h during hour blocks 1-4, when

temperatures were 15-24o C and the wind was <24km/h. Males sang less

(14.9 min/h) after they were mated than before mated (19.7 min/h), but

this decrease in singing may have been at least partially due to the

seasonal decline in singing activity described above, or vice versa.

From a sample of 122 flight songs, 74 were given by birds of known

mated status. Unmated birds sang 84% of these 74 flight songs (Fig.

111-14). This difference was highly significant (two-tailed X2

one-sample test, X2=33.8, df=1, P<0.001).

Discussion of flight songs

Flight songs are characteristic of birds that live in open

grasslands or tundra. Many emberizine sparrows, including the







89


congeneric seaside, sharptail, Baird's (A. bairdii), and Le Conte's

sparrows (A. leconteii), have well-developed flight songs.

Why do seaside sparrows give flight songs? Although the flight song

is a highly conspicuous vocalization, obviously its energetic cost is

greater than that of perch singing. Post & Greenlaw (1975) report that

in their migratory northern seaside sparrows, flight songs are much

more common soon after females arrive on their male's territories. This,

as well as my findings that unmated Scott's seaside sparrows gave more

flight songs than mated birds, suggests that flight songs may be used in

mate attraction. All seaside sparrows, however, continue to give these

vocalizations throughout the breeding season.

A major portion of the flight song vocalization is the uttering of

one or two modified primary songs. Elsewhere (Chapter IV) I have shown

experimentally that the primary song is essential for both mate

attraction/retention and territory establishment/retention. Thus, it is

reasonable to assume that flight songs can effect the same critical

functions. I have presented evidence that birds give relatively more

flight songs (and sometimes they give only flight songs) during

inclement weather and when unmated. I therefore suggest that even

though they may be energetically costly, flight songs so effectively

magnify the message of the primary song that they are worth more to the

male, especially under otherwise adverse singing conditions.




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PAGE 158

81,9(56,7< 2) )/25,'$


THE ECOLOGY AND VOCALIZATIONS OF SCOTT'S SEASIDE SPARROWS
(Ammodramus maritlmus peninsulae)
By
MARY VICTORIA MCDONALD
A DISSERTATION PRESENTED 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
1986

Copyright 1986
by
Mary Victoria McDonald

ACKNOWLEDGEMENTS
Since this dissertation is a formal report of my doctoral research,
it would be inappropriate for me to describe all of my ideas and my
feelings over the years—false starts, hunches that paid off, my
occasional discouragement, confusion, and elation, and even the kinship
I felt working with "my” birds on "my" marsh. But these elements were
inextricably woven into my study. What saw me through was not just my
scientific approaches to interesting ornithological questions, but also
determination and good luck, and, most of all, invaluable help from
friends and advisors.
Dr. John William Hardy, my major advisor, steadfastly supported and
guided me through the years. His timely reviews of my work and my
requests for recommendations, his just criticisms, and his flexibility
in allowing me to work independently all significantly contributed to
the success of my graduate career. My other graduate committee members,
Drs. Pierce Brodkorb, John H. Kaufmann, Peter Feinsinger, and Thomas J.
Walker, thoughtfully and generously advised me throughout my project.
Many other faculty members in the Department of Zoology also helped me
greatly.
I am fortunate to have had valuable suggestions from three
ornithologists very knowledgeable about seaside sparrows—Herbert W.
Kale II, Jon S. Greenlaw, and William Post. William Post patiently
helped me get started in my field work at Cedar Key and allowed me to
ill

incorporate some of his unpublished reproductive data and banding
records with my data, as presented in Chapter II of this dissertation.
The Department of Zoology generously supported me with teaching and
research assistantships, equipment, and vehicle use. The Florida State
Museum Bioacoustics Laboratory provided sound analysis equipment and
work, space. Other financial support was provided with grants from
several sources: Sigma Xi Grants-In-Aid of Research (1982 and 1983),
Frank M. Chapman Memorial Fund Awards (1983 and 1984), Eastern Bird
Banding Association Research Award (1984), and Van Tyne Memorial Fund
Grant of the American Ornithologists' Union (1984).
John David Wood, Sr., graciously permitted me to conduct my project
on his property near Cedar Key, Florida. The Florida Department of
Natural Resources rangers of the Waccasassa Bay Station near Cedar Key
helped make my sometimes uncomfortable, and always wet and muddy field
work more bearable. Two occasional field assistants, Janine Russ and
David Specht, were genial and as well as adept companions. Thomas A.
Webber also helped me in the field with photography and sound recording.
Many, many other people helped me indirectly over the years. The
support of my parents, Carlyle A. McDonald and Margaret L. McDonald, was
invaluable. And likewise invaluable were the advice and support of many
fellow graduate students. Thomas A. Webber and Linda S. Fink deserve
special thanks—their suggestions, thoughtful and significant criticisms
of my work, and, most of all, their supportive friendships have been
essential parts of my graduate work and life.
iv

TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS iii
LIST OF TABLES vii
LIST OF FIGURES viii
ABSTRACT x
CHAPTERS
I GENERAL INTRODUCTION 1
II NATURAL HISTORY AND BEHAVIORAL ECOLOGY OF SCOTT'S
SEASIDE SPARROWS: STUDY TECHNIQUES AND RESULTS 2
Taxonomy and Morphology .. 2
Project History and General Methods 4
Study Site 6
General Description 6
Weather 7
Marsh Fauna 3
Vertebrate fauna 8
Invertebrate fauna and seaside sparrow foraging . . 9
Marsh Flora 10
Size and Demarcation of Study Sites 11
Reproduction and Reproductive Behavior 11
Overview of Annual Cycle 11
Annual Cycle of Reproductive Behavior 15
Territorial establishment and mating 15
Nests, eggs, and early development 13
Nesting behavior ..... 20
Post-breeding behavior 22
Productivity, Survival and Reproductive Success .... 23
Territories and Territorial Behavior 29
Definition and Methods of Determining Territories ... 29
Description of Territories at Cedar Key 29
Territorial Behavior 30
Reactions to Other Species of Birds and to Humans ... 32
Discussion of Seaside Sparrow Territoriality 34
Territory types and variation 34
Territory quality and space use 35
Extension of territory definition and function . . 36
v

IllVOCALIZATIONS OF SCOTT'S SEASIDE SPARROWS
40
Introduction 40
General Methods of Data Collection and Analysis 41
Coverage 41
Sample Size and General Observation Methods 42
General Analysis of Notes and Recordings 43
Results and Discussion of Calls and Song 45
Description and Use of Vocalizations 45
Calls of Scott's seaside sparrows 47
Primary song 73
Subsong 79
Countersinging and Repertoire Use 79
Introduction and comments 79
Methods of investigating repertoires 82
Results and discussion of repertoire use 83
Flight Songs: Description and Comparison to
Perch Songs 84
Description 84
Methods of investigating flight song activity . . 85
Results of singing activity analysis 37
Discussion of flight songs 88
IVFUNCTION OF SONG IN SCOTT'S SEASIDE SPARROWS 93
Introduction 93
Methods 101
Experimental Design 101
Muting Procedure 106
Statistical Analysis 103
Results 109
Voice and Post-Operative Recovery 109
Mate Attraction and Retention 110
Territory Establishment, Retention, and Size Change . . 119
Behavioral Changes of Muted Birds 120
Reaction to Playback 129
Voice Recovery and Subsequent Behavior 130
Discussion 130
VCONCLUSIONS 135
REFERENCES 136
BIOGRAPHICAL SKETCH 145
vi

LIST OF TABLES
Table II-l. Summary profile of territoriality and reproduction
of Scott's seaside sparrows at Cedar Key, Florida,
1979-1984 24
Table III-l. Vocalizations and behaviors measured during time
budget observations 44
Table III-2. Vocalizations of Scott's seaside sparrows 46
Table III-3. Characteristics of solo primary singing and
countersinging of 30 mated male Scott's seaside
sparrows during April and May 74
Table IV-1. Experimental design of muting experiments 102
Table IV-2. During songless comparison of territory ownership
and size changes for the Mid-Season mutings .... 117
Table IV-3. "After song regained" comparison of territory
ownership and size changes for the
Mid-Season mutings 113
Table IV-4. Behavior changes for Individuals, from "Before"
to "During" muted 121
Table IV-5. Behavior differences comparing Muted, Sham, and
Undisturbed groups 123
Table IV-6. Sexual selection related to two main song types . . 131
vii

LIST OF FIGURES
Figure II-l. Breeding chronology of seaside sparrows at
Cedar Rey, Florida, 1979-1984 14
Figure III-l. (A) Audiospectrograra of Tuck call (short notes around
8 kHz), and 1 interspersed Tsip call (vertical note).
(B) Audiospectrogram of Tsip call.
(C) Audiospectrogram of Seeep note 54
Figrue III-2. (A) Audiospectrogram of male Primary song and
concurrent female Seeep note.
(B) Audiospectrograra of Tchi call 56
Figure III-3. Audiospectrograra of Whinny vocalization 53
Figure III-4. Audiospectrogram of Zuck calls 60
Figure III-5. Audiospectrograra of Scree calls 62
Figure III-6. Audiospectrogram of Begging calls from
two nestlings, individuals A and B 64
Figure III-7. Audiospectrogram of Primary song 66
Figure III-3. Audiospectrograra of Subsong 63
Figure III-9. Audiospectrograms of Countersinging from two males,
individuals A and B 70
Figure III-10. Audiospectrogram of Flight song 72
Figure III-ll. Relationship between time of day and the ratio
of Flight songs/Perch singing 91
Figure III-12. Relationship between wind velocity and the ratio
of Flight songs/Perch singing 93
Figure III-13. Relationship between temperature and the ratio
of Flight songs/Perch singing 95
Figure III-14. Number of Flight songs recorded in early 1983
given by birds of designated mated categories . . 97
Figure IV-1. Audiospectrogram of song of bird ABOR recorded
3 days prior to muting 112
viii

Figure IV-2. Audiospectrogram of "songs" of muted bird ABOR
recorded 5 days after muting 114
Figure IV-3. Relative abilities of Muted versus Sham-Operated
and Undisturbed males to attract and retain
females 1983 and 1984 115
Figure IV-4. Behavioral changes of 21 muted birds Before and
During their songless periods 128
ix

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
THE ECOLOGY AND VOCALIZATIONS OF SCOTT'S SEASIDE SPARROWS
Ammodramus marltimus peninsulae
By
Mary Victoria McDonald
December 1986
Chairman: Dr. John William Hardy
Major Department: Zoology
I studied the ecology and vocalizations of Scott's seaside sparrows
(Ammodramus maritimus peninsulae) in a Florida salt marsh from
1981-1985. I observed the behavior, vocalizations and reproductive
efforts of 20-30 pairs each year on the 30 ha gridded study site. These
2
monogamous birds defended all-purpose territories about 1,750 m in size
with song and short-distance agonistic behaviors. A high proportion
of males (0.79) returned each year to their previous territories, and
half (0.50) of these remated with their previous year's mates. Although
females initiated about 3.7 clutches of 3 eggs each season, productivity
was low (0.67 fledglings/female/year), because predators destroyed about
80% of the nests. Nevertheless, population numbers remained stable over
the years 1979-1985. The birds' vocal repertoires consisted of a
primary song, three modified primary songs (whispering song, flight
song, and subsong), and 10 calls. Only males sang; they had a song
type repertoire of 2-3 songs. I experimentally investigated the
function of song during the breeding seasons of 1933-1985 by temporarily
x

muting male birds in the field. t muted birds by rupturing the
interclavicular air sac. Birds remained songless for about 2 weeks but
could give all of their normal calls during this time. 1 administered
two rounds of mutings, each preceded and followed by time budget,
playback, and other observations on 3 treatment groups: Muted,
Sham-Operated, and Undisturbed birds. The Early muting round tested for
mate and territory acquisition. These Muted birds remained mateless
until they regained their voices; most eventually attracted a mate.
Muted birds acquired territories later than Sham-Operated and
Undisturbed birds. The Mid-Season muting round tested for mate and for
territory retention, and changes in behavior. All Muted birds lost their
mates; a few attracted new mates when they regained their singing
ability. Their territories either shrank or were lost, but new
territories were established (or the original re-expanded) when singing
ability returned. The intensity of close-range aggressive behaviors was
significantly greater for Muted birds than for Sham-Operated and
Undisturbed birds. There were no discernible differences between
Sham-Operated and Undisturbed birds for any of the attributes measured.
xi

CHAPTER I
GENERAL INTRODUCTION
I studied the ecology and vocalizations of Scott's seaside sparrows
(Ammodramus maritimus peninsulae) on a salt marsh near Cedar Key,
Florida, for 5 years. William Post had studied the reproductive ecology
of this population from 1979-1960. In early 1981, I began my
investigations and continued my study through the spring of 1965. The
principal objective of my research was to determine the function of the
vocal repertoire of these birds. During the course of my field work,
however, I observed many aspects of their biology and also collected
reproductive and non-vocal behavioral data.
This dissertation summarizes both my observational and experimental
investigations of the study population. Chapter II is an overview of
the project history, study site, general methods, and the birds and
their biology. Chapter III describes the vocal repertoire and discusses
the probable function of these vocalizations. Chapter IV summarizes the
experiments I conducted in 1983 and 198*+, whereby I determined the
function of song in Scott's seaside sparrows.
The information presented throughout most of this dissertation
portrays the behavior of birds I observed and tape recorded with a
minimum of disturbance. The exception, of course, is that Chapter IV
presents results of manipulative experiments. Thus I sometimes make the
distinction between "normal" and "experimentally manipulated" birds.
1

CHAPTER II
NATURAL HISTORY AND BEHAVIORAL ECOLOGY OF SCOTT'S SEASIDE SPARROWS:
STUDY TECHNIQUES AND RESULTS
Taxonomy and Morphology
Seaside sparrows (Ammodramus maritimus) belong to the "grassland
group" of Emberizine finches (subfamily Emberizinae), which also
includes the genera Ammospiza and Passerculus. The nominate form (A. m.
maritimus) was originally described by Wilson in 1811. Currently nine
subspecies of seaside sparrows are recognized (American Ornithologists'
Union [A.O.U.] 1957, 1973), including the nearly extinct dusky seaside
sparrow (A. m. nigrescens). Beecher 1,1955) postulated that formation of
the races of seaside sparrows was due primarily to geographic isolation
caused by post-glacial rise in sea level and the concurrent drowning of
river mouths forming bays (also see Funderburg & Quay 1983). All but the
southernmost populations of the nominate race are migratory. The other
races are generally considered to be non-migratory.
Scott's seaside sparrow (A. m. peninsulae) was named in honor of W.
E. D. Scott and described in 1888 by J. A. Allen. The type specimen was
collected near Tarpon Springs (Pinellas County, Florida). A. m.
peninsulae ranges from Old Tampa Bay (Hillsborough County, Florida)
north to Pepperfish Key (Dixie County, Florida) (A.O.U. 1957)*
Austin (1983) and I (McDonald 1983a) have briefly discussed and
annotated the taxonomic history of the seaside sparrow assemblage. The
distinguishing plumage color characteristics of the seaside sparrow
subspecies are thoroughly described and compared by Funderburg and Quay
2

3
(1983). Three museums in the United States house good representative
collections of seaside sparrows that I have examined: The American
Museum of Natural History, the United States National Museum of Natural
History, and the Florida State Museum.
Seaside sparrows are generally recognized by their dark gray-brown
head and body (total length about 14 cm), narrow black streaks on the
breast and flanks, yellow lores and wrist spots, and long bills. The
tail is relatively short and narrow and the feet and legs are relatively
large, in proportion to the size of the bird. When flushed, seasides
characteristically fly short distances and then drop back into the marsh
vegetation. Unlike other sparrows, seaside sparrows eat many
crustaceans and insects, but few seeds.
In his original description, Wilson succinctly described seaside
sparrow habits when he said: "Amidst the recesses of these wet sea
marshes, [the bird] seexs the rankest growth of grass and sea weed, and
climbs along the stalks of the rushes with as much dexterity as it runs
along the ground, which is rather a singular circumstance, most of our
climbers being rather awkward at running" (Wilson l8ll, p. 68).
Scott's seaside sparrows are among the darkest of the seaside
sparrows. As in other races a certain degree of individual color
variation exists. Some birds approach the lighter colored A. m.
maritimus in shading on the dorsum; others approach the darker A. m.
nigrescens. Early taxonomists made no mention of this color variation,
although Griscom (1944) recognized it. Austin states: "a gradual cline
is evident from the smaller, grayer populations in the south
(peninsulae) to the slightly larger, darker, and dorsally browner birds
in the northwest corner of the range in the Wakulla area (juncicola)"

4
(Austin 1968a, p. 838). However, W. Post and H. W. Kale (pers. comm.),
most familiar with the morphological variations in the southern
subspecies, agree that there are apparently no characteristics
consistently separating A. m. peninsulae and A. m. Juncicola (Wakulla
seaside sparrow). The range of juncicola is contiguous with peninsulae
near Pepperfish Key, Florida and extends to southern Taylor County,
Florida. Thus, juncicola should probably be merged with peninsulae,
which has taxonomic priority.
In all seaside sparrows the sexes are identically colored, but males
are slightly larger. In my study at Cedar Key I routinely took wing
length (wing chord) and weight measurements. For adult males and
females the mean wing chord measurements and 957» confidence intervals
were 56.9 (+9.4) mm and 54.0 (_+ 0.3) mm; and mean weights 22.5 (_+ 0.6) g
and 19.6 (_+ 0.7) g, respectively (11=134 males and 78 females). Thus
females were 937» as large as males based on wing chord measurements and
897» as large based on weight measurements. Because the 997» confidence
intervals for wing chord did not overlap, male and female adults handled
outside of the breeding season could be sexed using wing chord. During
the breeding season the age and sex of birds could more easily be
determined using other criteria. Females possessed a vascular, edematous
brood patch about 1.8 cm wide and males had a cloacal protruberance of
about 2 mm. Fledglings and older hatching-year birds had a
characteristic paler plumage pattern.
Project History and General Methods
The study site near the Gulf coast town of Ceuar Key, Florida, was
established by William Post in December 1978. Post investigated the

5
habitat and reproductive biology of the birds for two years, under
contract with the Endangered Species Program of the Florida Game and
Fresh Water Fish Commission. Information and results obtained from his
project were to become part of a management plan for the rare and
endangered Atlantic coast population of ni. n-j grescens. Post gridded
the marsh with stakes, began color banding of birds, mapped vegetation,
sampled the invertebrate fauna, studied the interspecific influence of
fish crows (Corvus ossifragus) and rice rats (Oryzomys palustris) on
sparrow reproduction, and recorded other descriptive data pertaining to
the morphology and life history (Post 1980, 1981a, 1981b; Post &
Greenlaw 1982; Post et al. 1983; Greenlaw & Post 1985)»
Post concluded his field work in the summer of 1980. I accompanied
him to the study site several times early in 1981, and then assumed
responsibility for the study site and birds later in the spring of 1981.
During the breeding seasons (March-June) of 198I-I982 I familiarized
myself with the marsh and the birds' behavior, and continued color
banding adults and some nestlings. I spent an average of 50 days in the
field each of these two years, observing and tape recording
vocalizations.
In 1983 and 1984 I continued comprehensive observations and color
banding of my Florida study population, each year averaging about 70
days in the field from March through June. Behavioral observations on a
field day consisted of 2-4 hours of watching, recording some of the
vocalizations, and describing behavior and territories. I recorded
spoken notes on a tape recorder, and included the location of birds and
activity time (measured with a stopwatch). I later transcribed tapes
and analyzed the data.

6
The major investigation of my study—temporary song muting—was
conducted during the breeding seasons of 1983-1984 and is described in
Chapter IV. In the spring of 1985 I spent 15 days in the field censusing
returning and new males and conducting playback experiments with female
seaside sparrow vocalizations.
Throughout this report the data designated as having been collected
in 1979-1980 are from Post's unpublished and published work and are used
with his permission. All other data were collected and analyzed by me.
I used non-parametric statistical tests (Siegel 1956; Conover 1980) for
most analyses.
Study Site
General Description
The 30-ha salt marsh study site was north of the Waccasassa Bay of
the Gulf of Mexico. The site was 6 km NNE of the town of Cedar Key, Levy
County, Florida, and within the Sumner Quadrangle, USGS Map (29° 11' Li,
83° 00' W). The physical boundaries of the marsh, exterior to the core
gridded study area, were Prodie Creek on the SW, Live Oak Key and
Waccasassa Bay on the S, and Dorset Creek on the NE. The property was
privately owned.
The boundary between the marsh and Waccasassa Bay was not sharply
defined. The shoreline was dotted with oyster bars and many small
islands and was cut by numerous tidal creeks. Except for these creeks,
which ranged from 0.5 to 1.5 m deep at mean high tide, the marsh was
flat. Tides averaged 0.8 m. Most years one or two flooding episodes
occurred during the breeding season, destroying any existing nests of
seaside sparrows. These floods were most often generated by prevailing

7
winds from the south, driving water in from Waccasassa Bay, combined
with a natural (high) spring tide event. The water level during these
floods usually rose about a meter above mean high tide. Average salinity
measured by Post (1980) at 32 stations on 24 April 1980 was 19*03 1.09
0/00.
Weather
Data on cloud cover, air temperature, and wind direction and
velocity were collected at about hourly intervals throughout each field
day. Phenological records such as flowering dates and spring migration
observations were also kept. More complete daily weather information
(maximum and minimum temperatures and rainfall) for all years was
recorded by a National Oceanographic and Atmospheric Administration
(NOAA) station near the town pier of Cedar Key.
My behavioral observations and the handling of birds were carried
out under mild weather conditions (18-33° C, wind <20 km/h) during the
months of February through June. During my 5-year study, however, I did
visit the marsh in all months of the year and under virtually every
weather condition. Temperatures I recorded ranged from 3° C (January) to
1*1° C (August). On typical sunny days during the height of the breeding
season (April and May) the mid-morning temperatures averaged 26° C. For
those field days on which I took detailed behavioral observations, the
wind velocity usually ranged from about 2 km/h (just prior to sunrise)
to about 12 km/h by 0900, and then 16-24 km/h by early afternoon.
Prevailing winds were mostly from the southwest. A few sultry field
days were virtually windless. Winds approaching gale force (50+ km/h)
occurred occasionally in the spring and fall.

8
In September 1985, Hurricane Elena touched the Cedar Key area. %
surveys the following week and in 1986 indicated no discernible
population decline in the three bird species breeding on this marsh
(seaside sparrows, marsh wrens [Cistothorus palustris], and clapper
rails [Rallus longirostris]).
Marsh Fauna
Vertebrate fauna
The six-year average population densities of the three resident
bird species were seaside sparrows—2.5 +_ 0.1 birds/ha, N=253; marsh
wrens—0.6 +0.3 birds/ha, N=23; and clapper rails—0.5 0.1 birds/ha,
N=59 (Post 1981a; McDonald 1982, 1983b, 1984, and ay unpublished surveys
for 1984 and 1985)» In addition to these residents, migratory
sharp-tailed sparrows (Ammodramus c_. caudacutus) wintered on the marsh.
They regularly arrived in the last week of September and left in the
second week of May. The population density of sharptails averaged about
2/ha. Other bird species seen foraging or resting on the study site
were those commonly found in Florida Gulf Coast salt marshes (Post
1981a; McDonald 1982, 1983b, 1984).
As described below in the section on reproduction, most seaside
sparrow nest losses were attributed to predation, primarily by fish
crows and rice rats. The rice rats dwelt on the marsh. Their density was
about 8/ha (Post 1981b). I also found evidence of predation by
transient racoons (Procyon lotor).
Other species of mammals were occasionally observed: river otters
(Lutra canadensis) played on the bank edges; Atlantic bottlenose
dolphins (Tursiops truncatus) and very rarely, manatees (Trichechus
manatus) swam in the larger tidal creeks. The only reptile I saw on the

9
study site was the Gulf Salt Marsh Snake (Nerodia fasciata clarki),
observed on three occasions.
Invertebrate fauna and seaside sparrow foraging.
The most obvious and abundant invertebrates on the marsh were
fiddler crabs (Uca rapex) and Gulf periwinkles (Littorina irrorata).
Genoni (1984) reported additional mud-inhabiting invertebrates from this
marsh. Terrestrial invertebrates were also censused previously by
sweep-netting the vegetation during the sparrows' breeding seasons (Post
et al. 1983)» By percent of total sample, these were Tettigoniidae
(54.3%), Lycosidae (18.8%), other spiders (7.8%), and Lepidoptera
(Noctuidae and Pyralidae) (3-0%). Six additional invertebrate families
each represented less than 57» of the total sample. Post et al. (1983)
found that 857» (by volume) of the nestling's diet consisted of these 10
groups of arthropods.
The invertebrate food species consumed by adults on this marsh are
probably nearly the same as those eaten by Scott's and other subspecies
of seaside sparrows on similar marshes. Howell (1928, 1932), Oberholser
(1938) and Wilson (l8ll) examined stomach contents of adult Scott's,
Smyrna (A.m. pelanota), Louisiana (A. m. fisheri), and dusky (A. m.
nigrescens) seaside sparrows. In addition to the arthropod groups listed
above, which are known to be fed to nestlings at Cedar Key (and
presumably eaten by adults), these authors found adults had eaten marine
"worms," small crabs and other crustaceans, beetles, dragonflies, flies,
wasps, bivalves, gastropods, and some "weed" and grass seeds. (Audubon
[1831i said that a pie he made of seaside sparrows could not be eaten
due to its "fishy savour," surely attributable to the birds' having
consumed mostly salt marsh invertebrates!)

10
With the assistance of a biologist familiar with marsh
invertebrates, I examined the stomach contents of eight adult birds from
my study site. Surprisingly, three of the stomachs were virtually empty.
These males had died early in the morning during the breeding season,
perhaps prior to foraging. The other stomachs contained mostly
arthropod exoskeletons (fragments of crabs, insect adults and pupae, and
one small spider), snails (Pyramidellidae and Marginellidae), and
vegetation. Post et al. (19&3) presented convincing evidence that food
was plentiful and accessible to the Cedar Key birds throughout the year.
I rarely observed birds drinking. Several times I saw then imbibe
dew from the vegetation, and once I saw a bird drinking and bathing at
the edge of a shallow tidal creek. Although no passerines have salt
glands, seaside sparrows do have the ability to concentrate urine when
consuming salt water (Poulson 1969)-
Marsh Flora
Vascular plant species on the study site were described in detail by
Post (19Ü0). In order of relative cover, the major plants were smooth
cordgrass (medium height) (Spartina alterniflora) 38%, black rush
(Juncus roemerianus) 26)4, seashore saltgrass (Distichlis spicata) 237»,
and perennial glasswort (Salicornia virginica) 8%. There were no woody
plants within the gridded portion of this marsh. Also, there were no
pannes (bare mud areas devoid of vegetation) until September 1985, after
the passage of Hurricane Elena near the Cedar Key region. Apparently
some stands of Spartina were submerged for at least several days during
the storm; consequently these drowning-susceptible plants died.

il
Size and Demarcation of Study Sites
Post established two contiguous study areas, Cedar Key West ("CKW,"
10-ha in area) and Cedar Key East ("CKE," originally 20-ha in area ).
Although CKW and CKE were contiguous, the study populations of seaside
sparrows living on each site were separated by 200-400 m, and the
populations did not interact. Both study areas were gridded with 2-3 m
high wooden stakes placed at 25 m intervals. Stiff metal prongs on the
tops of the markers discouraged their use by predatory birds. Stakes
were alpha-numerically designated and could be read from at least 50 m.
Additional markers were placed around the peripheries of the study areas
for reference. I concentrated my preliminary observations on the CKW
birds in 1981, then switched to CKE study area (which I enlarged to
30-ha in 1984) for the remainder of my project.
Reproduction and Reproductive Behavior
Overview of Annual Cycle
Fig. II-l graphically summarizes the reproductive cycle of Scott's
seaside sparrows at the Cedar Key study site. The cycle commenced in
early March with the onset of regular singing by males and subsequent
territorial establishment and mate acquisition. The peak of the breeding
season (as indicated by the number of newly completed clutches) for the
six years 1979-1984 occurred during the second and third weeks in May.
In a typical spring the first clutches were completed around the
beginning of April, and the last clutches by the end of June.
Most clutches (85/0 were completed by the end of May. Virtually all
breeding and territorial defense activites ceased by mid-June. Both

12
adults and juveniles entered a prolonged molting period during the early
fall and remained difficult to observe until the following early spring.

Figure II-l.
19Ü0; Post et
sample of 293
breeding chronology of seaside sparrows at Cedar Key, Florida, 1979-19^* Data from Post
al. 1963; and author's unpublished observations, bars represent proportions of total
nest observed during indicated 2 week intervals.

PERCENT OF TOTAL COMPLETED CLUTCHES
MALES SING REGULARLY
40-
30
20 -
10 -
JAN
A
[MALES DEFEND TERRITORjES
[MATE ACQUISITION I
' ~ RETENTION *
I DEPENDENT YOUNG j
f
INDEPENDENT JUVENILES
/
/
¿i
V'y
^7
.
L
2
~7
A
FEB
MAR
APR
POST NUPTIAL MOLT
P
? 7
MAY
/
JZL
JUN
JUL
AUG
SEP
4 f
OCT
NOV
-1
DEC

15
Annual Cycle of Reproductive Behavior
Territorial establishment and mating
Breeding behavior of Scott's seaside sparrows near Cedar Key began
in late winter (February) with sporadic singing of adult males on warm,
windless mornings. Young adult males, hatched the previous spring, also
began to sing imperfect subsongs. By early March all males were singing
typical primary songs, and females began to associate with males.
Females solicited male affiliation with the "whinny" and "tcni" calls
(Chapter III).
During these early weeKs of the season, weather influenced the
intensity of singing and other observable breeding activities. Wind
velocities greater than 25 km/h accompanied by temperatures less than
20° C caused a nearly complete cessation of overt territorial and mating
behaviors. Mild weather, even in early February, greatly enhanced
regular singing activity and behavioral interactions. In addition to
this immediate influence of atypical weather, prolonged cool
temperatures delayed the onset of breeding within a given season, and
hot weather in late May caused an early cessation of breeding
activities. The early spring of 1983, for example, was unusually cold:
average daily temperatures for March were about 5° C below normal
throughout mid-Florida. The breeding activities of the study population
at Cedar Key were about 2 weeks delayed that year—I did not find a
completed clutch until 26 April. On the other hand, the unusually warm
February of ly85 certainly must have accounted for ny finding fledged
seaside sparrows on 16 March and fledged marsh wrens on 23 February,
both record early nesting dates for these species for any known breeding
locality

16
Most males re-established the same territories from year to year and
vigorously defended these during March and early April, the period when
the most fervent territorial disputes between first-year and older
adults and between residents and floaters broke out. 3y May territorial
ownership and boundary disputes were rare, seemingly occurring only when
floaters attempted (always unsuccessfully) to take up residence on an
occupied territory. The intensity of singing (as measured in minutes of
singing/hour) also declined gradually after a peam in early April. By
June there was only sporadic singing activity occurring daily from just
before sunrise to about one-half hour after sunrise.
Most birds paired by mid-March. The mates remained paired over the
remainder of the breeding season. I never observed a natural incidence
of divorce, although Werner (1975) reported several instances of
re-pairing in his Cape Sable birds in the same breeding season. Seven
percent of 97 non-experimental territorial males remained bachelors
throughout the breeding season in my study. Werner (1975) reported that
about 12> of his 111 Cape Sable seaside sparrows were bachelors.
Apparently the adult sex ratio of the breeding birds was slightly skewed
in favor of males. Although Post (i960) assumed that most unmated
territorial males were probably first year birds, my longer study of
these marked birds indicated no significant difference in the
proportions of unmated older territorial birds and unmated first-year
territorial birds (_X2=0.U, N=10 bachelors, df=l, £>0.5).
My observations over the years also suggested that returning males
tended to re-mate with their female of the previous year, if she also
returned the second year. I documented mate retention using data for
non-experimental males and females present in both 1963 and I96U, the

17
years for which I had the most complete data. Of the 10 cases in which
both members of the pairs were present in succeeding years, there were
five re-mates. Mate retention from year-to-year also occurs in northern
seaside sparrows on Long Island, New York (Jon Greenlaw, pers. comm.).
Werner (1975) also stated qualitatively that there was some degree of
year to year mate retention in his Cape Sable seaside sparrow subjects.
Females solicited copulations with a slow, low-pitched variation of
the "whinny" vocalization accompanied by quivering bilateral wing
raises. This display was given at ground level with the bird concealed
in the marsh vegetation. A nearby male (usually the mate) was
immediately attracted to this enticing behavior and would quickly
copulate by mounting the female briefly with fluttering wings and then
bounce off, landing several meters away. The male sometimes gave one or
two songs immediately after copulation, whereas the female was almost
always silent afterwards.
Typical bouts of copulation involved three to five mounts during
time intervals of 0.5 to 1.5 hours. All of the copulation bouts I
observed occurred in the mornings. During this period the female
sporadically gave her solicitation whinny, while the male remained
within a 15 m radius of her and sometimes sang. Of the 96 copulation
bouts I observed under non-experimental conditions, all but three were
between birds I had previously known or subsequently knew to be mates.
On the remaining three occasions I observed neighbor or floater males
sneak in and copulate with a soliciting female while the mate was
vigorously singing or confronting other birds on the opposite side of
his territory.

lti
Nests, eggs, and early development
Although this was primarily a behavioral study, nesting and other
reproductive data were routinely collected during the breeding seasons
of 19Ü1-1964, and in some cases were consolidated with Post's
1979-19ÃœO data.
Seaside sparrow nests are notoriously difficult to find in salt
marshes. My early efforts to find nests by dragging a chain through the
vegetation, by trying to flush nesting females by walking transects, and
even by using a bird dog were largely unsuccessful. I eventually
learned to locate most of the nests of subject birds by close
observation of the parents.
For all nests I found, I noted date and time of discovery, parents,
location relative to the grid system and territorial boundaries, and
vegetative microhabitat. Because I wished to minimize human
interference, I handled only a sample of the nests and nestlings that I
was aware of. The additional information taken for these sample nests
was form of nest cup (open or domed), height from ground to bottom of
cup, dimensions of cup (inside and outside diameter, inside and outside
depth), material used in constructing nest, number of eggs or
hatchlings, egg dimensions, hatchling weight (for 11 individuals), age
of hatchlings, and daily survival. I also weighed seven nests collected
after fledging.
The following summarizes ny descriptive data on 19^ nests. Two
nests had well-defined domes and entrances. I detected no preferred
directional entry pattern of attending parents for any of the nests,
although the nests of some subspecies of seaside sparrows are mostly
domed and have distinct entrances. About half of the nests were built

19
in Distichlis and the remaining roughly split between Salicornia and
Juncus microhabitats. All nests were constructed primarily with the
Distichlis grass; a few had small Spartina leaves woven in. The average
height of the nest (as measured from the ground to the bottom of the
nest cup) was 35 cm. Average nest size measurements were: outside
diameter of cup 9*3 cm; inside diameter of cup 5*4 cm; outside depth of
cup 5*3 cm; inside depth of cup 3*3 cm. Also in contrast to other
subspecies, at Cedar Key the average clutch size was smaller (3*02 +
0.17 eggs, N=194 nests), compared to about 3.65 in hew York (Post et al.
1983).
Average egg dimensions were 20.6 mm X 15.7 ami (N=35), and egg
weights averaged about 2 g. While exhibiting minute individual
variation, the color pattern of the eggs was essentially the same as
described for seaside sparrows in detail elsewhere (Austin 1966a).
Egg and nest cup dimensions for Scott's seaside sparrows at Cedar
Key were within the ranges reported for other subspecies in the
literature (Austin 1966b; Norris 1968; Sprunt 1968; Trost 1966;
Woolfenden 1968; Werner 1975i Greenlaw pers. comm.) and in collections I
measured at the American Museum of Natural History, the United States
Museum of Natural History, and the Florida State Museum.
I categorized nest placement within a pair's territory as "edge" or
"center." An edge nest was so designated if it was built in the outer
fourth of the total area of the territory. I found no significant
difference in the birds' tendencies to locate nests at the territory
edge versus towards the center (X^=0.7, N=194, df=1, P>0.4).
Only lib nestlings were directly observed, but others were known to
be in active nests. The young remained in the nest 9-11 days. Their

20
weights increased from 2.2 g on Day 0 to 15*1 g on Day 10. The few
other morphological data I took on these birds agree with kerner's
(1975) extensive quantitative morphological data and photographs of
nestlings taken days 1-8 for Cape Sable seaside sparrows.
Morris (in Austin 1968a) gave a detailed description of nestling
Louisiana seaside sparrows (A. m. fisheri) and the behavior of the young
and their parents. My observations of Scott's seaside sparrows at Cedar
Key are very similar to Morris's in Louisiana.
Nesting behavior
Nest building (by the female only) began on the day of or the day
after copulation bouts. Females did not re-use nests, but built a new
nest for each new clutch. The female usually completed building her
nest within a day and laid her clutch of three eggs (rarely four) over
the next five days. The eggs were laid one a day, at any time during the
day. Infrequently females that were building nests, laying, or
incubating, would copulate with their mates. Incubation was by the
female only, for 11-12 days.
Most years the first young of the season hatched around the second
week in April (Fig. II-l). All of the young hatched within 24 hours and
began to gape and utter soft "peeps" on the day of hatching. The eyes
were fully opened by day five and at about this time the begging
vocalizations changed from a "peep" to a "churp" and became more
intense. Some nestlings fledged and were able to min about as early as
Day 9. Most began short flights about Day 13»
The hatchlings remained in the nest for 9-10 days and were fed by
both parents. The combined average feeding rate was 3*2 visits per
hour. Parents also maintained a clean nest by removing fecal sacs. Post

21
et al. (1983) determined the diet of hatchlings older than 3 days by
placing pipe-cleaner ligatures around their necks and then removing the
food from the young or the nest cups after an hour had elapsed. The
most important food groups for the nestlings were found to be the
insects of the taxa Tettigoniidae and Lepidoptera and spiders of several
families. These invertebrates constituted 857» of the nestlings's diet.
Both parents continued to feed the young for 15-20 days after
fledging. If the breeding season had not progressed too far (i.e. prior
to about 15 May), a female with fledged young would begin to renest
about a week after the first brood had fledged and would leave the care
of fledglings to her mate. I noted five such cases in 8b pairs.
Since most nests were destroyed before young fledged, seldom did a
pair actually have the opportunity to rear a second brood within a
season. From late March through early June, unsuccessful parents would
try repeatedly to raise a family. The average number of nesting attempts
per season of 8b pairs (l98l-198b) was 3.7, and the interval between
nest destruction and renesting was 5*7 days. In 198b I observed one
pair attempt seven nests, all unsuccessful.
Fathers began to avoid their first brood after tending their
fledglings for about three weeks, or after their second successful
clutch hatched. They chased their fledglings (as well as any other
juveniles) off the parental territory and started to devote their
attention to their new brood. After nesting had stopped family groups of
fledged, dependent young and their parents were often observed during
the months of June and July. Adults without families were seldom seen
during this period, especially during the hot daytime hours.

22
Post-Breeding behavior
By August most juveniles were independent and began to form loose
groups of about three to eight. Sometimes adult males associated with
these groups, which persisted through October. Juvenile males gave
occasional bouts of imperfect subsong in August and September, and
rarely through the winter. Although present (as evidenced by occasional
mist net captures), adult females were rarely seen in summer.
From late September through October there was a resurgence of easily
observable but essentially non-vocal behavior, such as foraging and
territorial defense. From these observations and my intensive mist
netting in the fall of 1982 and 1983, I concluded that adult males and
females remained within a radius of about 250 m of their previous spring
territory throughout the year. Juveniles appeared to wander a great
deal more, pernaps as far as several Am.
Little is known regarding the fate of the fledgling seaside sparrows
at Cedar Key or elsewhere. Of the total of 98 hatchlings and fledglings
Post and I banded at Cedar Key, I recaptured only 4 during my intensive
netting 1982-1984. Presumably the emigration and general mobility of
fledglings as well as of non-territorial older birds are mechanisms for
the birds' locating and colonizing any unoccupied territories and viable
habitat.
The Cedar Key population appeared stable (average density 2*5
birds/ha) throughout the 7 years 1979-1985- Adult birds with territories
were relatively long-lived. By the end of 1984, I had studied subject
males from 1 to 4 years. According to banding records, two males were at
least 5 years old, and one was at least 6 years old.

23
The adult post-nuptial molt (and only annual molt) of their nuptial
plumage began in early July, and the post-juvenile molt began in late
July, both molts were completed by October.
The seasonal weight variation was slight but significant. Both
sexes were heavier in winter than in summer, but the difference was not
as great as in the nominate migratory seaside sparrows (Post 19Ü0).
Productivity, Survival, and Reproductive Success
I summarized parameters of reproduction for seaside sparrows at
Cedar Key in Table II-l. The data combine my unpublished observations
(1931-19^) with Post's observations (1979-1980), as given in Post
1980 and Post et al. 19Ó3*
Post (19dlb) intensively studied seaside sparrow reproduction and
factors that affected productivity during his 1979-1980 project. He
also compared these characteristics of reproduction in A. m. peninsulae
to another well-studied population, A. m. maritimus in New York (Post &
Greenlaw 1982; Post et al. 1983). I combined the two years of Post's
data with reproductive data I collected 1981-1984.
Studies of nest productivity should consider whether nest failures
are due to human disturbance. I did not test if my visitations affected
the nesting success I observed. Post (1980), however, did test for an
observer effect using the methods of Mayfield (1975) and Johnson (1979)
and found that the frequency of nest visitation did not affect nesting
success of Scott's seaside sparrows.

24
Table II-l. Summary profile of territoriality and reproduction of
Scott's seaside sparrows at Cedar Key, Florida, 1979-198*+
cL
Territories, Mating, and Return Rates
Mean territory size
Ecological density of territorial males
Mean annual return of territorial males
Proportion of re-mates
Proportion of bachelor territorial males
Ratio unmated first year/older males0
Mean length of study of male in population
Productivity and Survivala,c
Mean clutch size (C)
Mean no. clutches initiated/female/yr (I)
Probability of egg producing fledgling (S)
Productivity (fledglings/female/yr)°
Mean length of breeding season
Proportion of nests producing fledglings
Rest success ratio of young/older breeders
Adult annual survival ^male return) rate
Juvenile survival rate°
Proauction of fledglings->breeders next yr
1759 + 242 m2
2.5 males/ha
0.78 males/yr
0.50
0.07
4/6 N.S.
2.3 yr
3.02 + 0.17
3-7
0.06
0.67
95 days
0.06
3/5 a.S.
0.78
0.67
0.45/female/yr
aCombined data from Post 1980; Post et al. 1983; and author's
unpublished observations. Probability level: R.S.=P>0.05*
b
See text for explanation of calculations.
Based on 194 nests

25
At Cedar Key the probability that a nest would produce fledglings
was remarkably low, but because females renested within 7-8 days of nest
destruction, the population seemed to have remained stable from 1979
through 1985. That is, bird density remained about the same over the
years we censused the study population (Post 1980, 196la; McDonald 1962,
1963b, 1984, and my unpublished data).
Post et al. (1983) calculated probability of an egg's surviving 21
days (through the egg and nestling period) and producing a fledgling to
be only 0.06 (Post et al. 1983). This contrasts with an egg survival
rate of about 0.35 in New York.
Of a total (McDonald and Post) sample of 194 nests whose fates were
carefully documented, only 12 (6/0) produced at least one fledgling. The
major causes of nest mortality at Cedar Key were flooding, and predation
by fish crows and rice rats. Eight (4%) of the nests failed due to
desertion. Neither Post nor I ever suspected hatchling death due to
starvation. I found only a few instances of partial nest mortality.
Eleven (2%) of the 473 eggs I observed in the field failed to hatch and
were presumably infertile.
Destruction by flooding accounted for the mortality of 17 (9%) of
the nests. Normal tidal fluctuations in water levels seldom reached the
bottom of the nest cup (about 35 cm above the ground). However, in most
years at least one major unpredictable flooding of the marsh occurred.
For the seven years (1979-1985) there were nine such floods during the
breeding seasons.
Predation by fish crows destroyed 37 (19/0 of the total nests. Fish
crow predation was assumed when eggs disappeared without leaving shell
fragments or a disturbed nest (indications of mammalian predation). Fish

26
crows were about three times more likely to prey upon nests with
hatchlings than on those with eggs.
Post (1981b) concluded that nesting in Juncus should be preferred by
the birds because Juncus stands were less likely to be flooded and
because this vegetation provided cover protection from fish crow
predation. However, rice rats on the marsh apparently discouraged
seaside sparrows from nesting more in Juncus by destroying their nests
(Post 1981b). The birds were found to nest more frequently (85% of the
total nests) in less dense bistichlis and Salicornia, even though nest
failure was more likely to occur in these vegetation types, as compared
to failures in Juncus (Post 1981b).
The remaining 120 (62%) of the total nests were destroyed by rice
rats and unknown predators (probably racoons).
Post (1981b) tested whether rice rats were actually destroying nests
by placing tin cylinders around nests. These were open at the top,
allowing entry by parents and aerial predators but excluding small
mammals. Forty-eight percent of 42 experimental nests with this
protection produced fledglings, whereas only 6% of the 34 control
(unprotected) nests had young that fledged.
To summarize: about 80% of the nests at Cedar Key were destroyed by
predation. This contrasts with findings in Hew York where fewer seaside
sparrow nests were destroyed by predators and more losses (about 65% of
the total nest mortality) were due to flooding or rains, desertion,
hatching failure, and unknown causes (Post et al. 1983).
I report productivity as the number of fledglings/female/year.
Post's data (1980) were combined with mine: Post observed 109 clutches,
and I observed 85 clutches. Productivity was determined by the method

27
of Ricklefs and Bloom (1977): Productivity = Mean Clutch Size (3*02) X
Probability of Egg Success (0.06) X Number of Clutches Initiated by a
Female/Year (3*7)• (The term "productivity” and its values are roughly
equivalent to the "realized specific natality rate" parameter that is
conventionally calculated using life table data.) The six year average
productivity for 1979-1984 was 0.67 fledglings/female/year. This value
contrasts with the much higher rate of 4.25 for the New York population.
New York females produced about six times as many young within their
average 75 day breeding season as the Florida birds did in their average
95 day season. The annual survival, however, was lower in New York
(Post et al. 1983).
I determined the adult annual survival rate at Cedar Key to be about
0.78, based on the return of 51 of 65 adult males over the years
I98O-I983 and assuming a roughly equal sex ratio. This figure represents
a minimum value, since some birds may have emigrated. This is a
relatively high value for a small Passerine bird. Werner (1975)
estimated a 0.88 survival rate for the likewise non-migratory A. m.
mirabilis, based on a one year return sample. Adult survival is lower
for migratory A. m. maritimus—only about 0.45 (Post 1974).
I estimated the juvenile survival rate to be about 0.67- This figure
was determined using the method of Post et al. (1983) and using their
estimated ratio of juvenile/adult survival (0.85, as determined from
returns of banded nestlings and adults). Thus, I multiplied the adult
annual survival rate of O.78 by the ratio of juvenile/adult survival of
O.85, giving 0.Ó7.
Although I did not have enough data to directly calculate net
reproductive rate (R^) using life table parameters, I suggest that the

28
Cedar Key population of seaside sparrows is maintaining its numbers for
the following reasons: First, there was no decline in population
density over the years. Additionally, my experiments in 1983-1984
demonstrated that floaters (itinerant males without territories or
mates) were always available to take up residence on territories they
perceived as unoccupied. Secondly, the reproductive data I do have
suggest that enough juveniles were produced and survived to replace
non-returning adults. That is, using my calculated productivity figure
of 0.67 fledglings/female/year and Post's (1980) juvenile survival rate
of 0.67, each breeding female (pair) produced 0.45 first-year birds.
Assuming a sex ratio of about unity, each female produced 0.229
first-year female breeders. If the minimum adult annual survival rate
was 0.78 and applicable to both sexes, then the annual adult female
mortality rate was 0.22 (1.00 - 0.78 = 0.22) or less. Since 0.225 is
about the same as 0.22, these reproductive data do substantiate my
observation that the population size has been constant over the years.
Post et al. (1983) calculated a net reproductive rate of 2.72 for the
increasing New York population. The authors also state that the net
reproductive rate for the Cedar Key population was 1.11 (based on
1979-1980 data), but it is unclear how this figure was determined
without life table (i.e. survivorship) data.
These New York and Florida projects are the only major long-term
studies that report and summarize detailed reproductive data. Other
papers that contain some data on breeding success and productivity for
other subspecies of seaside sparrows are Nicholson (1946), Woolfenden
(1956), Stimson (1956), Norris (1968), Sprunt (1968), Stimson (1968),
Trost (1968), Woolfenden (1968), Worth (1972), and Werner (1975)*

29
Territories and Territorial Behavior
Definition and Methods of Determining Territories
The working definition of "territory" I used in this study was "any
defended area" (Noble 1939, p. 267)- My use of the tern "defended"
implies that a bird exhibited aggressive behavior towards another, with
the seeming intention of driving or keeping the other out of his or her
territory. I considered vocalizations and visual displays directed at
another bird to be defensive behaviors. This study dealt primarily with
aspects of male territoriality, although female seaside sparrows also
exhibited territorial behavior. I considered a territory to be an area
in which a male invariably challenged intruders with singing, displays,
or other overt aggressive behaviors.
All of my behavioral observations were made relative to the grid
markers. I drew territory maps for the entire study population weekly,
by connecting the points of the most peripheral singing perches with
lines. I traced the territories thus delineated with a compensating
polar planimeter and then calculated the enclosed areas. When a map
indicated a boundary had moved by 15 m or more, I considered that to be
a territorial boundary shift. The summary information regarding
territory sizes and location reported in this chapter were those
measured for non-experimental birds during the first week of May unless
indicated otherwise.
Description of Territories at Cedar Key
Territories at Cedar Key were all purpose—used for mating, nesting,
and feeding. Mated pairs exclusively occupied territories throughout
the breeding season, although boundaries between territories fluctuated.

30
Only two instances of complete territory relocations were noted for 65
non-experimental males studied 1979-19^2.
The mean yearly return rate for male territorial birds was 797»
(N=tí9)« There was a strongly significant tendency for males to
re-establish territories on the same sites in succeeding years. (My
conservative null hypothesis was: Returning males are as likely to
establish a new territory as they are to re-establish their old
territory. My sample sizes and statistics were: 43 re-established
2 2
territories of 52 observed returnees; _X one-sample test, X =21, df=l,
_P<0.001). Year-to-year retention of territories has also been reported
qualitatively in the few other studies of marked populations of seaside
sparrows (Worth 1972; Werner 1975; Greenlaw pers. comm.).
The territory sizes of non-experimental birds at Cedar Key ranged
from about 200 to 4000 m^. The mean size was 1759 m^ (S£=242 , N=65).
This average territory size for the population did not change
significantly over the years 19Ü2-19Ü4 (Kruskal-Wallis one-way aNOVA,
N=43 territories of non-experimental males, df=2, tí=O.Ü3, P>0.5). In
comparing the individual males' territories from year to year, I found
no significant increase or decrease in their territorial areas (Wilcoxon
matched-pairs signed-ranks test, N=23, T=9l, _P>0.05).
The boundaries of territories did not follow any apparent natural
landscape divisions, such as vegetation type borders or small tidal
creeks. However, larger tidal creeks (>20 m wide) did often separate
territories.
Territorial Behavior
Several studies of northern seaside sparrows (Post 1974; Post &
Greenlaw 1975) contain a few quantitative data describing territorial

31
behavior. Other authors (e.g., Audubon 1831; Nicholson 19^6; Norris
I960) anecdotally portrayed territorial and breeding behavior of seaside
sparrows, some in detail and many from an anthropomorphic perspective.
Those reports that best characterize territorial birds and their social
interactions are Norris (1968), Worth (1972), ana Werner (1976)»
At Cedar Key males began to establish their territorial boundaries
in early March by singing regularly from song perches, usually clumps of
elevated vegetation such as Juncus tussocks located at the edges of
their territories. Overt aggressive encounters were also first observed
at this time and became most numerous in April. Behaviors of such
encounters, in order of frequency of occurrence, were chasing bouts
accompanied by strident "tchi," "zuck," and rapid "tuck" calls;
supplanting (a challenger flew towards a perched bird, and the perched
bird left without dispute); and short-range "facing off" displays,
wherein the participants engaged in wing raises, bobbing, bill
thrusting, and wing and tail flick displays, co-occurring with the
"tuck" and "tchi" calls and "whisper songs."
These Early Season territory-establishment behaviors, as well as
male-female pairing chases, decreased significantly in frequency and in
duration as the breeding season progressed into May. By the end of the
breeding season (June), territorial males seldom disputed intrusions by
neighbors and floaters.
Prior to experiments I conducted in 1983-198U, Post and I believed
that there were no excess males trying to establish territories and
consequentially compress existing territory sizes. I found, however,
that there were such-males, as discussed below in Chapter IV.

32
Females definitely displayed aggressive behavior towards both
intruding males and other females throughout the season, although I have
few quantitative data on the extent of their involvement in
territoriality. In contrast to males, which were more likely to give
short distance aggressive displays and calls, females were more likely
to supplant and chase intruders. Of the total 3,159 instances of overt
aggressive behavior I analyzed between seaside sparrows in March and
April, 22 percent involved females. Only 5 percent were known to be
female-female encounters. Observations of such encounters were
difficult to make, because females spent more time than males in dense
vegetation. The frequency of female chases of intruders from the
pairs's territory averaged 2.J chases per hour in late March and early
April. These chases were usually accompanied by the "tchi"
vocalizations and sometimes by a fast rendition of the "whinny"
vocalization (female solicitation call).
Reactions to Other Species of Birds and to Humans
I observed no vocal or other behavioral interactions of seaside
sparrows with resident clapper rails or marsh wrens. Each of these
species seemed oblivious to the sounds emitted by the others. Calls and
songs often overlapped, even when given within close range (<15 m). This
is in contrast to other studies of bird communities, in which singing
characteristics of several species are influenced by each other (e.g.
Popp et al. 19b5). Several times I did note that seasides alternated
songs in a definite pattern (countersang) with migrating red-winged
blackbirds (Ageliaus phoeniceus) and bobolinks (Dolichonyx oryzivorus).
Cruising fish crows (notable nest predators of seaside sparrows) almost
always caused seasides to stop singing and sometimes to give "si

33
twitter" alarm calls (23 observations of low-flying fish crows during
song, 18 instances of song ceasing).
Wintering sharptail sparrows incited aggressive behavior from
seasides, especially when the sharptails began to sing in early May. The
seaside's aggressive behavior was identical to that directed towards
conspecific intruders. Early in the breeding season the mostly silent
sharptails were tolerated and unchallenged. On 17 occasions I neard
distinct countersinging between seasides and sharptails, although their
songs are quite different.
My study population of seaside sparrows seemed oblivious to my
presence, except when I approached to within 5 m or handled them. The
birds were more wary when I was accompanied by field assistants. Other
researchers have made similar observations regarding the "tameness" of
their subject populations of seaside sparrows (Sprunt 1968; Post 1974).
At Cedar Key curious sparrows and wrens approached within 1 m of me when
I was still for 30 min or longer. Indeed, I once awoke from a nap on the
ground to find a sparrow staring at me about 20 cm from my face!
My close approach to nests (<10m), however, did provoke alarm and
sometimes distraction displays from the parents when they were in the
nest vicinity. If the female was on the nest, she usually ran about 3 m
then flew about 5 m farther away. Presumably these reactions are nearly
the same as those given in response to other "predators." Several times
I simulated identical predator-reaction behavior by placing a mounted
fish crow model 2 m from a seaside nest. The typical response by the
parents was high pitched "seeet calls" interspersed with the "tuck"
call, given while the birds rapidly flitted around in the vegetation
within 5 m of the nest. Three times I saw females give apparent

34
distraction displays by trailing their wings as they ran away from their
nests.
Discussion of Seaside Sparrow Territoriality
Territory types and variation
The seaside sparrow literature indicates that different races, and
populations within races (see Post 1974), may have functionally
different types of territories. These have been described variously as
colonies (Tompkins 1941; Sprunt 1968); areas with separately defended
nesting and feeding areas (Woolfenden 1956, 1968; Worth 1972); grouped
(defended) nesting territories with undefended remote foraging areas
(Post 1974); nest-centered activity spaces ("Type B" of Wilson 1975;
Post 19Ü0); and all purpose breeding territories (mating, nesting, and
feeding; "Type A" of Wilson 1975» Werner and Woolfenden 1983). The
occasional references to seaside sparrows' being "colonial" may be
misleading in that this designation implies the birds had very small
territories, such as those of colonial shorebirds. Although the limited
nesting habitat and the concomitant foraging behavior of some
populations give the appearance of coloniality, seaside sparrow nesting
groups are perhaps better described simply as aggregations.
I have reviewed the literature describing seaside sparrow habits and
have observed populations from Florida to New York. I conclude that
nearly all subspecies and populations exhibit the "all purpose" type of
territoriality. A few populations (e.g. at Post's (1974) Gilgo Beach
study site) exhibited variations of the all purpose type, in that the
birds sometimes foraged out of their territories on undefended areas.
Territories at Cedar Key were of the all purpose type. Occasionally

35
individuals were seen foraging, unchallenged, on other known
territories.
Territory quality and space use
Post and co-workers (Post 1974, 19blb; Post & Greenlaw 19&2; Post et
al. 19b3; Greenlaw & Post 1985) comparatively studied the
interrelationships of territory size and quality, predation, food
availability, reproductive success, and mating systems of seaside
sparrows on Long Island, New York, and at Cedar Key, Florida. Their
studies reached several conclusions with which I generally concur: Food
seemed not to be a limiting factor in determining territory size;
"Space-use patterns" (i.e. maintainence of territory size, and foraging
on and off the territories) were more influenced by the numbers of birds
attempting to settle into a habitable area than by food availability.
In other studies Post (1974) and Werner (1975) contended that it is
unlikely that territorial spacing limits the population size of seaside
sparrows. They suggested that as the density of birds increases over
the years the territory sizes simply decrease in order to accommodate
more birds. The population size at Cedar Key remained stable throughout
ny study; thus, I did not have an opportunity to support or refute this
hypothesis under the circumstances of increasing population as described
by these authors. Territorial compression either did not occur at Cedar
Key, or if it did occur then the territories must have become maximally
compressed prior to my study. I conclude this after considering the
findings of my 1983 and 19^4 experiments on territorial birds (Chapter
IV). Because I found that non-territorial males did exist, I contend
that territory boundaries and sizes are not necessarily so flexible as
to accomodate all aspiring breeders, as suggested by Post and Werner.

36
A few bird studies have established an inverse relationship between
territory size and resources (Zimmerman 1971; Seastedt and MacLean
1979)* Greenlaw & Post (1965), however, found that the sizes of the
territories they investigated were not correlated with their composite
index of territory quality (which included nesting, food and
cover/protection factors). Furthermore, territory size was not directly-
related to volume of vegetation, area of vegetation, or amount of food
on the territory (Post 1980).
Several recurrent conclusions appear in all seaside sparrow
literature describing territorial and feeding behaviors. My
observations on birds at Cedar Key support most of these: Territory
sizes and probably quality vary considerably within and among
populations. Food is seldom if ever a limiting resource, as
convincingly argued by Post et al. (1963). Animals with small
"nest-centered" territories simply forage, unchallenged, off their
territories. Both migratory and non-migratory males tend to establish
their territories in the same locations year after year. Territorial
shifts within a season rarely occur. Birds in dense populations spend
more time involved in aggressive behavior but neither spend less time
feeding nor suffer lower reproductive success as a result of crowding
(Post 1974). Male sparrows are probably not defending areas for the
food value per se, but are rather advertising and defending territories
in order to increase their chances of mating and keeping mates.
Extension of territory definition and function
Kaufmann (1963) stated that territoriality is one form of social
dominance, dominance being defined as priority of access to critical
resources (e.g. food, mates) that increase the fitness of the dominant

37
individual. Kaufmann further described territoriality as "relative
dominance." Individual A may be dominant to a subordinate, B, while in
A's territory, but not dominant to B when outside of A's territory.
These interpretations of territoriality are more inclusive than the
restricted working definition I gave at the beginning of this section.
They synthesize that traditional, restricted definition of a territory
as "any defended area" with the broader concept of social dominance.
Kaufmann's characterization of territoriality as one manifestation of
social dominance is quite applicable to seaside sparrows at Cedar Key.
The territories were indeed vigorously defended areas, yet a male off
his territory was invariably subordinate in another bird's territory.
Males together on the few areas of the marsh not occupied by territories
did not evidence dominant-subordinate behavior: apparently shared were
food and the rare female affiliations that occurred outside of
territories. Thus no dominance hierarchy or absolute dominance seemed to
exist.
As Kaufmann (19^53) pointed out, arguments about functions of
territoriality have mostly been concerned with whether a territory
serves as an area to enhance social stimulation, or as an area to allow
resource acquisition. Evidence in support of the former hypothesis,
proposed by Darling (1952), is that some birds clump their territories
even when nearby suitable habitat is available. Considering the
synchronized breeding behavior of seaside sparrows and the observation
that populations do indeed clump their territories, one may concur that
social stimulation is a function of territoriality in these birds.
However, I do not believe this to be the case in the birds I have
observed, for the following reasons.

3b
First, I suggest that clumping is a response to many individuals
having the same gestalt perception of preferred habitat, which
researchers have not been able precisely to identify. Since food is
apparently not limiting, I suspect that the preferred habitat for a male
seaside sparrow selecting and setting up a territory is an area of the
marsh that allows maximum singing display advertisement (i.e. moderately
low vegetation with dispersed singing perches such as tussocks of
Juncus). Singing and being able to be heard while singing are vital to
the fitness of male seaside sparrows, as determined by experiments
described in Chapter IV. Thus, what may initially appear to be suitable
unoccupied habitat may in fact be unsatisfactory for establishing a
song-defended territory. Secondly, I do not believe that social
stimulation is a function of territoriality in seaside sparrows, because
their apparently synchronized breeding behavior can almost always be
directly attributed to weather and other environmental conditions, such
as flooding.
For seaside sparrows at Cedar Key, the possession of a territory
does seem unequivocally to allow the owner priority of access to the
resources contained therein, the most important of these probably being
singing perches, a mate, and a relatively undisturbed mating/nesting
area. Thus the territories of seaside sparrows easily fit the
conventional concept of bird territories' being "defended," areas
exclusively occupied by a relatively dominant male. Also applicable is
Kaufmann's broader characterization of a territory as "a fixed portion
of an individual's or group's range in which it has priority of access
to one or more critical resources over others which have priority
elsewhere or at another time. This priority of access must be achieved

39
through social interaction" (Kaufmann 1983, p. 9)* Because this
definition includes temporal and relative dominance criteria, this
definition more comprehensively characterizes seaside sparrow
territoriality.

CHAPTER III
VOCALIZATIONS OF SCOTT'S SEASIDE SPARROWS
Introduction
Seaside sparrows (Ammodramus marltimus) have a vocal repertoire of
distinctive calls and song types. Because these birds are cryptic and
tend to remain low in their densely-vegetated salt marsh habitat, vocal
communication is important in their social behavior. Intergradations and
varying contextual uses of sounds presumably convey different
information to other birds receiving the messages.
Most older seaside sparrow literature is descriptive, and
vocalizations are reported in general and anecdotal behavioral
observations (Kopman 1915; Sprunt 1924; Holt & Sutton 1926; Howell 1932;
Stone 1937; Tomkins 1941; Nicholson 1946; Woolfenden 1956; Norris 1968;
Trost 1963; Worth 1972; Lowery 1974). Relatively recent studies of the
northern seaside sparrow (A. m. maritimus) by Post & Greenlaw (1975)
and of the Cape Sable seaside sparrow (A^. _m. mirabilis) by Werner &
Woolfenden (1983) include more objective descriptions of calls as well
as songs and associated behaviors. Post & Greenlaw's (1975 ) detailed
description of the vocal and display repertoire of their Long Island,
New York, population is especially thorough.
In this chapter I describe and discuss the vocalizations of a
Florida population of Scott's seaside sparrow, A_. m. peninsuale. I
include more quantitative and conclusive data than in my preliminary
report on Scott's seaside sparrow vocalizations (McDonald 1983c). This
40

41
chapter also compares the vocalizations of my study population to those
of northern seaside sparrows described by Post & Greenlaw (1975) and to
those of Gape Sable seaside sparrows described by Werner & Woolfenden
(1983).
General Methods of Data Collection and Analysis
The project history, study site, and subjects were described in
Chapter II.
Coverage
I recorded vocalizations and described the behavior of the study
population throughout the years 1981-1984. My most intensive field work,
occurred during the breeding seasons (late February through mid-June)
when I went to the marsh at least 3 days/week. I also visited the marsh
about once every 10 days during the remainder of the year. During the
breeding season I made the majority of my behavioral observations and
recordings when the birds were most active—from just before sunrise
until about four hours after sunrise. The birds' activity level
increased again moderately about an hour before sunset. The few evening
behaviors and vocalizations I recorded seemed not to differ from early
morning observations, thus I only mention their occurrence in this
chapter. Unless otherwise noted, my quantitative descriptions of
vocalizations and behaviors are for observations taken on mild spring
mornings when the temperature ranged from 18-33° C, there was no
precipitation, and the wind ranged from 0-24 km/h.

42
Sample Size and General Observation Methods
The total number of marked birds (1981-1984) was 213, including
birds manipulated in 1983-1984. In generalizing about the use of
vocalizations, I considered only data for 91 mated and 11 unmated birds
that I had observed for at least 15 h per season. Detailed analyses
were based on fewer males, as described below. I studied each subject
male at least one hour per week throughout the breeding season. On a
typical morning I observed 1-3 focal males for about an hour (an hour's
"time budget" block), then watched another 1-3 males for the next hour,
and so on up until about 0900. I randomly determined the sequence of
these time budget observation blocks. Thus, all birds received
approximately equal coverage that was representative of the morning
hours. Females were difficult to observe; I described their behaviors
and vocalizations as parts of their mates' time budgets.
During the time budget I dictated all observed behaviors and the
birds' locations on cassette tape. I made most recordings (including
all used for sound analyses and sonograms in this section) with a Sony
TC 150 cassette tape recorder (Mineroff-modified), using a Bell and
Howell "Shotgun" unidirectional elec tret condenser microphone with
windscreen. I measured the amplitude (in dB) of a singing male 7 m from
me with a Bruel & Kjaer Sound Level Meter (Type 2219) on 5 Apr 1985.
I also used field data forms to record contextual information,
banding records, nest descriptions, territorial boundaries, and other
biological information. Territory maps for the entire population were
drawn in the field at least bi-weekly throughout the breeding seasons.

43
General Analysis of Notes and Recordings
I transcribed, tabulated, and summarized the behavioral data. I
determined characteristics of tape recorded sounds (e.g. frequency
ranges, length of song) with a Uniscan II FFT Real Time Sonogram
Spectral Display (Model 4600), and prepared sonograms (Fig. III-l
through Fig. III-10) with a Kay Elemetric 7029A Sono-graph using the
wide-band filter (300 Hz) and either the 30-8000 Hz or the 1600-16,000
Hz scale. The specific vocalizations and behaviors I analyzed are
listed in Table III-l.
Detailed quantitative descriptions of vocalizations and other
behaviors were determined from an analysis of 103 time budget hours of
30 mated and territorial best-studied males. I chose time budgets for
analysis based on comparability of day of year (Apr-May), time of day
(sunrise-0900), and the typical mild weather conditions described above.
I used non-parametrie statistics (Siegel 1956; Gonover 1980) for the
analyses.
My specific methods of investigating flight songs ("complex flight
vocalizations" of Post & Greenlaw 1975) and countersinging are discussed
in separate sections below.

44
Table III-l. Vocalizations and behaviors measured during time budget
observations (in min and sec duration, and frequency)
Singing3’
Primary song
Whispering song
Matched countersinging
Not matched countersinging
Flight song
o K
Other Vocalizations *
Tuck
Tsip
Si Twitter
Seeep
Tchi
Tchi flight
Whinny
Zuck and other calls
Position Changes3’D
When not singing
Singing, song type change
Singing, no song type change
Singing, stop singing
3 fo
Observations Relating to Females and Young ’
Female present
Female solicitation calls, other calls
Copula tions
Females building nests
Other behaviors related to females and young
Other Behavior3 ’
Feeding, resting, and preening
Off territory: feeding; invading nearby territory
o K
Reactions to Intruders *
Frequency, duration, and location of intruder
Flight chases
Vocalizations:
Solo primary song
Countersong
Whispering song
Calls (as above)
Displays:
Sham-preen
Grass-pick
Bob and bill jab
Con tac t
Contextual information also noted: bird's identity, location, mated
status; presence of conspecifics, date, weather, and location of
observer.
^Terminology follows Post & Greenlaw 1975.

45
Results and Discussion of Calls and Song
Descriptions and Use of Vocalizations
Scott's seaside sparrows make 14 distinct vocalizations: a primary
song, three modifications of the primary song, and 10 calls. The
probable functions of their songs and calls are summarized in Table
III-2. I distinguish "songs" from "calls" as implicitly defined by most
ornithologists, and as specifically stated in Pettingill's working
definition: "Song is a vocal display in which one or more sounds are
consistently repeated in a specific pattern. It is produced mainly by
males, usually during the breeding season. All other bird vocalizations
are collectively termed call notes or, simply, calls" (Pettingill 1970,
p. 319). I use Post & Greenlaw's (1975) vocalization terminology
throughout, except for "whispering primary song," which is from Werner Sc
Woolfenden (1983).

46
Table III-2. Vocalizations of Scott's seaside sparrows
Vocaliza tion
Name—
Sex and^
Context—
Probable Function(s)
Primary song
Male;
1, 2, 4
Territory defense and
mate attraction
Whispering
song
Male;
1
Territory defense
during intrusion
Complex flight
vocaliza tion
Male;
1, 2, 5
Territory defense and
mate attraction
Subsong
(young birds)
Male;
1, 4
Practice song
Tuck
Male and Female;
1-5
General purpose;
moderate aggression;
chase; nest defense
Tsip call and
Si twitter
Male and Female;
1-3, 5
Heightened aggression;
attack; nest defense
Seeep note
Male and Female;
1, 2, 5
Proclaim location of
female(?); fear(?);
Flocking (fall 6 winter)
Tchi
vocaliza tion
Male and Female;
1-3, 5
Moderate aggression;
chase; female: nest area
distraction, proclaim sex
Whinny
vocalization
Female;
1, 2, 4
Attract males for
copula tion
Zuck call
Male and Female;
1, 5
Heightened aggression;
chase; drive off young
Scree call
Male and Female; 3
Extreme distress call
Begging and
Chup calls
(given by young
only)
Begging for food
Terminology follows Post & Greenlaw (1975). This table is modified
from Post & Greenlaw's (1975) Table 3, which summarized "vocal
displays" of northern seaside sparrows.
^Contexts in which the vocalization were elicited: 1-in presence of
intruder or neighbor bird; 2-in presence of mate; 3-in presence of
predator, or human disturbance; 4-also given when bird apparently alone;
5-given in flight.

47
Calls of Scott's seaside sparrows
In this section I give sonograms, descriptions, and suggested
functions of 10 calls, presented in approximate order of their frequency
of occurrence during the breeding season.
Tuck call
The "tuck” (Fig. III-1A) was the most common of the calls. It was
given by males and females throughout the year. The short tuck covers a
wide frequency range and is probably the same call Werner & Woolfenden
(1983) labeled as the "chip" call for Cape Sable seaside sparrows. The
tuck appeared to be an all-purpose vocalization, but seemed to function
mostly as a general alarm and moderately intense aggressive call. It
was invariably given during short-distance aggressive interactions and
usually accompanied by wing and tail flicks, and by the "tsip" calls
(shown specifically in Fig. III-1A).
Males gave tuck calls at an average rate of 0.3 min/h when there was
no apparent disturbance on their territories. During intrusions,
however, the tuck rate increased to 13.5 min tucking/h intruder present.
This call was also given alone (or together with "tsips") by both
parents when a predator or human approached a nest within 5 m. The tuck
call was the only one I heard regularly during the non-breeding season.
I also heard fledglings and juvenile birds give this call.
Tsip and si twitter
Both males and females also gave the higher-pitched, short "tsip"
call (Fig. III-1B). When birds were highly agitated the "tsip" was
rapidly repeated, thus intergrading into the "si twitter" call. The
"tsip" was often interspersed with "tuck" calls, as mentioned above, and

43
was accompanied by wing and tail flicks. During bouts of calling, which
lasted 0.25 to 17.3 min, birds gave "tucks," "tsips," and combined
"tuck-tsips" at a rapid and usually regular rate of 30-145 calls/min.
As with the "tuck" call, "tsips" and "si twitters" generally signaled
alarm and aggression, but more often indicated a higher level of alarm.
My closest approach to nests (<2 m) and my placing of a stuffed fish
crow near nests nearly always elicited "si twitters." I never heard
"tsips" or "si twitters" from undisturbed birds. During intrusions the
average combined "tsip-si twitter" rate was 4.5 min tsip-si twitters/h
intruder present. I only rarely heard the "tsip" call during the
non-breeding season, and I never heard fledglings or juveniles give
either of these calls.
Seeep note
During the breeding season I occasionally heard females give a soft,
high-pitched "seeep" call Fig. III-1C). Most of the "seeeps” occurred
during bouts of copulation. Sometimes a female gave this call as a
nearby male began to sing relatively softly (Fig. IIL-2A). Post &
Greenlaw (1975) suggest that the "seeep" call conveys a weak fear
message. This call, or a very similar one, was the second most common
vocalization I heard outside of the breeding season. Males and females
of both Scott's seaside sparrows and wintering sharptail sparrows at
Cedar Key gave "seeeps," especially in the late fall and early winter. I
agree with Post & Greenlaw (1975), and with Werner & Woolfenden (1983),
that this call probably functions as a social and flocking vocalization
outside of the breeding season.

49
Tchi vocalization
The loud and relatively short "tchi" call (Fig. III-2B) was given by
both sexes and was most often heard during chases. About half of the
total "tchis" I heard were given during or immediately following chases.
Usually the "tchis” were rapidly repeated, and frequently they
intergraded with a slower, slurred "tyu." The "tyu" itself often slurred
into a "whinny"-like vocalization (described below). The "tchi" call
was probably the same as the "squeaz" call described by Werner &
Woolfenden (1983), and the same as the "jee-jee-jee-jee-jee-jeeeu-jeeeu”
call described by Norris (1963).
The primary function of this call seemed to be signalling general
aggression during chases. Frequently, however, I heard females give
rapid "tchis" in flight ("tchi flight") as they left a nest they were
building or incubating, even though no threatening bird seemed to be in
the vicinity. These nest-departure "tchis" were not given adjacent to
the nest, but rather at about 10-15 m from the nest. During a 1 h
observation of nest building in early May 1983, for example, I saw a
female depart and come into her nest 18 times. Fifteen of her
departures were accompanied by "tchis,” but only one arrival was
accompanied by this call. Jon Greenlaw (pers. comm.) reports that female
northern seaside sparrows behave in a similar manner. I suggest,
therefore, that the "tchi" also functioned as a distraction call by
drawing a potential predator's attention away from the nest as a female
departs. Jon Greenlaw (pers. comm.) also thinks that "tchis" in the
nest vicinity function to let the resident male know that this departing
and arriving bird is a female who belongs, rather than an invading male.

50
Other than the females giving the "tchi" as described above, I
seldom heard this call when no intruders were present (0.11 male
tchis/non-intrusion hour). It was difficult to quantify the "tchi" rate
during intrusions. On the average, however, a territory owner spent
2.62 min chasing/h intruder present. I estimate that at least half of
these chases were accompanied by rapidly repeated "tchis," most of these
calls given by the owner.
Whinny vocalization
I heard only females give the "whinny," with one unusual exception
described below. This whirling, quavering call varies in rate from 0.3
to 3/sec (Fig. III-3). The whinny was the female's solicitation call;
it functioned to attract a male. Females gave this call more often when
a male was relatively close to her (within about 7 m), and especially if
the male was singing. Males were immediately attracted to the "whinny"
vocalization, and copulation usually followed. Even though females gave
the "whinny" from on or near the ground, males seemed to have had no
problem finding the vocalizing females. It is likely that some auditory
characteristic(s) of this vocalization made it easy to locate.
Females gave this call most often prior to, and during nest
building. Because females usually initiated up to 5 clutches per
breeding season, the overall intensity of "whinnies" heard for the
population remained the same from late March through May. The average
whinny rate heard on all territories during these months was 2.02 min
whinnies/h.
Post & Greenlaw (1975) describe the postures associated with the
"whinny" vocalization and copulation. Werner & Woolfenden (1983) do not
describe a female solicitation vocalization.

51
I only once heard a male give a distinct "whinny" (as opposed to the
"whinny"-like "tyu," described above). On 6 May 1984 a male invading an
experimentally muted bird's territory repeatedly gave whinnies while he
intruded.
Zuck call
The loud, raspy "zuck" (Fig. III-4) was an aggressive call given
during intense fights. Most of the zucks I heard were given by males,
but sometimes females and juveniles gave this call. It may have been the
same as the "Shu-shu...shu" call of Cape Sable seaside sparrows that
Werner & Woolfenden (1983) mention but do not describe. In addition to
its functioning as a general, high intensity aggressive call, "zucks"
were the vocalizations most often given by fathers when they drove their
older, almost independent fledglings off their territories (about 2-3
weeks post-fledging). Other than these father-fledgling encounters, I
never heard this call given during non-intrusion circumstances. During
intrusions both territory owners and intruders gave "zucks" (often mixed
with "tchis") at the rate of 0.7 min zuck/h intrusion.
Other calls
The "Scree" (Fig. III-5) was a harsh distress call I heard only
(with a few rare exceptions) from a few birds trapped in mist nets and
being handled. Both sexes of adults and juveniles gave this call.
The monotonous and insistent "Begging Calls" (Fig. III-6), given by
older nestlings and fledglings, were heard most often towards the end of
the breeding season. Nestlings gave "Peep" (Werner & Woolfenden 1983)
and “Chup" (Post & Greenlaw 1975) calls.
Post & Greenlaw (1975) describe one further call I have not
mentioned—the "Chew" call. To my knowledge, I never heard this call at

52
Cedar Key. Jon Greenlaw (pers. comm.) proposes that the "Chew" call is
probably a modified "tchi-whinny" vocalization.

Figure III-l. (A)
(vertical note).
Audiospectrogram of Tuck call (short notes around 8 kHz), and 1 interspersed Tsip call
(B) Audiospectrogram of Tsip call. (C) Audiospectrogram of Seeep note.

SQNOD3S NI 3WI1

Figure III-2. (A) Audiospectrogram of Male Primary song and concurrent
female Seeep note (indicated with "X"). (B) Audiospectrogram of Tchi
call.

kHz kHz
56
8 -1
0
TIME IN SECONDS
1

Figure III-3. Audiospectrograra of Whinny vocalization.

N
5 4H
2 -
O
O
TIME IN SECONDS
1

Figure III-4.
Audiospectrogram of Zuck calls.

o
TIME IN SECONDS
On
O
T
1

Figure III-5. Audiospectrogram of Scree calls

TIME IN SECONDS

Figure III-o.
Audiospectrogram of Begging calls froin two nestlings, individuals A and B.

o
TIME IN SECONDS
1

Figure III-7. Audiospectrogram of Primary song.

TIME IN SECONDS
1

Figure III-8. Aucliospectrogram of Subsong.

TIME IN SECONDS
1

Figure III-9. Audiospectrograms of Countersinging from two males,
individuals A and B.

70
8 -
ó J
4
2 -
0 1
TIME IN SECONDS
TIME IN SECONDS

Figure III-10. Audiospectrogram of Flight song.

o
TIME IN SECONDS
2

73
Primary song
Audiospectrogram. The seaside sparrow primary song (Fig. III-7) is
relatively simple. The introductory portion of rapid clicks (centering
around 6.5 kHz) is immediately followed by a buzzy trill (centered
around 3.3 kHz). These frequency ranges and the song duration of about
1.0 second were characteristic for the entire population.
General description of singing behavior. During my study, only
males sang. Although Post & Greenlaw (1975) describe "female songlike
vocalization" that resembles male song, I never heard a vocalization of
this type at Cedar Key. Hales sang to delineate territories and to
attract females. Experimental evidence for these functions of song is
given in Chapter tV.
Males usually sang from elevated perches, such as tussocks of
Juncus. Most birds had 3-7 favorite singing perches within their
territories from which they gave about 35% of their songs. From
1979-1984 only two males sang from grid marker stakes, although Post &
Greenlaw (pers. comía.) found that birds in their dense study population
often sang from grid markers.
The singing pattern of seaside sparrows is similar to that of many
song birds (Table III-3). Birds sang in "discontinuous" patterns of song
"bouts" (songs repeated continuously without a pause), of "eventual
variety" (song type change after three or more repetitions of the same
song type) (Hartshorne 1956; 1973). An average bout throughout the
season consisted of 19.2 songs and lasted 1-5 min (mean = 1.62). Highly
variable periods of silence (mean = 7.9 min) separated song bouts. Males
changed perches during 17% of these between-bout silences, before
resuming singing. Song rate within a bout ranged from 6-9 songs/nin

74
Table III-3. Characteristics of solo primary singing and countersinging
of 30 mated male Scott's seaside sparrows during April and May
Primary
Song (Solo) Countersong
Characteristic3 Total Total Matched11 Not Matched3
N (bouts)
391
223
143
30
Mean bouts/h obs.
3.80
2.17
1.39
0.73
Total min sing
635
631
433
192
Mean min sing/h obs.
6.16
6.13
4.26
1.86
Mean min song/bout
1.62
2.33
3.07
2.40
Total bouts edge0
198
123
30
43
Total bouts center^
117
41
26
15
Total min edge
315
369
275
94
Total min center
193
110
6o
44
Mean song type
<0.01
6.02
0.02
<0.01
switches/bou t
Mean song type
0.32
0.33
0.47
0.29
switches/h sing
Mean perch
0.97
0.33
0.31
0.35
changes/h sing
Mean perch changes/
0.16
0.02
0.02
0.01
bout
Mean perch changes/ <0.01
bout with song switch
<0.01
<0.01
<0.01
Mean perch changes/
0.03
0.02
0.02
0.01
bout without switch
Mean perch changes/
bout, song cessation
0.13
i
0.05
0.01
0.02

75
Table III-3, continued.
Characteristics of solo primary singing and countersinging determined
from analyses of 103 h of time budget observations, when no intruders
were present on focal birds' territories. Other conditions during time
budget observations are described in the text.
^Rate values calculated for Matched and Not Matched columns based on
number of bouts and minutes of Matched and Not Matched singing,
respectively. Thus the sums of rates of Matched and Not Matched values
do not add up to the Total column for Countersong.
c
Edge defined as the outer-most 10 m of the bird's territory,
d
Center defined as the area within a 25 m radius of the estimated center
of the bird's territory.

76
(mean = 7.7), close to the 6.6 song rate Woolfenden (1956) reported for
New Jersey birds, and slightly lower than the 10.6 song rate reported by
Post & Greenlaw (1975).
Individual males had a repertoire of two to four similar song types
(mean = 2.41). When both solo singing and countersinging, birds seldom
switched song types within or even between song bouts. The mean rate of
song type switches during a bout was about 0.01/bout. Thus their
singing was extremely "monotonous," sensu Hartshorne (1956; 1973),
because their singing pattern was both continuous and of low
"versatility" (infrequent song type changes).
Most males shared their song types with nearby territorial birds. A
few of the birds had one highly distinctive song type within their
repertoire that enabled me to recognize individuals aurally from as far
away as 150 m. I did not detect any tendency for males to have a
preferred song type from within their small song repertoire.
Seaside sparrows at Cedar Key seldom began a new song type after
moving to a new perch while singing (Table III-3). Of a total 67
incidents of perch changes during singing, there was only one concurrent
song type change. This constrasts with Krebs's (1977a, 1973) "Beau Geste
hypothesis" that territorial birds switch song types when changing
perches, thus possibly giving potential intruders the impression that
territories are saturated.
Influence of abiotic factors on singing behavior. Singing activity
varied throughout the year. It was most intense relatively early in the
breeding season (late March through mid-April), especially before and
just after birds were mated. The average time singing of 29 males
measured 1-15 Apr 1983 was 19.4 min singing/h. In the third week of May

77
their song rate decreased to 12.1 min singing/h, a significant
difference (two-tailed Wilcoxon matched-pairs signed-ranks test, T=2,
P<0.01).
Weather modified the intensity of singing, on both a daily as well
as a seasonal basis. Very early in the spring (February through mid
March) birds sang more as the day warmed slightly, up to about 20° G.
During the middle and especially towards the end of the breeding season,
birds sang less as the day warmed. Virtually all singing stopped at
temperatures above 32° C, regardless of the time of day. From the middle
of the breeding season till the end, birds sang more on cloudy, cool
mornings. Light rain also enhanced singing activity at this time of the
year, and even heavy rain and thunder and lightning dampened singing
activity only slightly. Brisk winds (>25 km/h), however, greatly
reduced singing activity, especially early in the season on cool days.
The intensity of singing activity also varied throughout the day.
Peak song activity was early in the morning, from sunrise to about 3
hours after sunrise. The earliest I heard singing was 20 min before
sunrise. Birds also sang in late afternoon until about one-half hour
after sunset. Compared to seaside sparrows on Long Island, New York,
that I observed in June 1982, birds at Cedar Key sang less in the
evenings. The first morning and last evening songs were similar to
songs given during peak morning activity, except the song rate
(song/min) within the bout was lower very early and late in the day.
Influence of other birds on singing behavior. Unmated males sang
significantly more bouts than mated males, and thus their total time
singing (21.4 inin/h) was greater than that of mated males (12.3 min/h),
as measured in May (two-tailed Mann-Whitney U-test, N=30 and 11,

78
£<0.01). Singing rate also differed significantly according to whether
an intruding male seaside sparrow was present. After territories were
well established (by mid-April), intrusions were rare (mean =0.7 min
intrusion/hour). When an intrusion did occur an owner's first reaction
was increased singing, and this was often successful in repelling the
intruder. If the intruder was not driven off by normal primary song,
territory owners switched to short-distance and more aggressive
vocalizations and behaviors—whispering primary songs, tuck and si
twitter calls, chases, quivering wing raises, and (rarely) contact
fights.
The ’’whispering primary song” (Werner & Woolfenden 1933) was a
slowly delivered (mean=3.5 song/min), low amplitude, and slightly lower
pitched modified primary song. It was usually given by a male perched
near the ground and confronting an intruder at 1-5 m. The whispering
primary song was delivered with an almost closed beak, in contrast to
the normal primary song where the bird threw back his head and opened
his beak wide as he sang. I do not have data describing reactions of
males that heard these whispering songs. Because I only heard it during
agonistic encounters, this vocalization may have functioned to transmit
a message of threat and liklihood of attack. Jarvi et al. (1980)
described a similar use of modified primary song in the willow warbler
(Phylloscopus trochilus) that transmitted a message that the singer was
likely to attack his opponent.
Because short-distance encounters were more often accompanied by
calls and displays, the combined normal and whispering primary song rate
during intrusions (2.4 min song/h intrusion) was much lower than the
song rate when no intruder was present (12.3 min song/h non-intrusión).

79
Galls of most other marsh birds, including the raucous clapper
rails, seemingly had no effect on the seaside sparrows' vocalizations or
other behaviors. Seaside sparrows did respond antagonistically toward
vocalizing sharp-tailed sparrows (Ammodramus c. caudacuta) when these
wintering migratory birds began singing prior to their departure the
first week in May.
Subsong
Young males sang rudimentary subsong late in July and in August
(Fig. III-8). These songs consisted of garbled warbles and parts
(usually the terminal trill) of the adult primary song. Subsong was not
as loud as the primary song and was sung only sporadically instead of in
bouts. These first song attempts improved and came to resemble the
adult's primary song by late fall, when virtually all singing activity
stopped. Both yearling and adult males began to sing again in late
February, and songs of the first year birds could still be
differentiated from those of older birds. By mid-March, however, songs
given by first-year and older birds could no longer be distinguished.
Countersinging and Repertoire Use
Introduction and comments
Countersinging occurs when two, three, and sometimes four male birds
alternate their songs in a regular pattern. Countersinging is described
as "matched" when the same song type or types are sung by the
participating birds. The behavioral and evolutionary significance of
song repertoire sharing in general, and of song matching during
countersinging in particular, has been the subject of recent stimulating
studies (Krebs & Kroodsma 1980; McGregor et al. 1981; Payne 1982; Kramer
et al. 1985; Searcy et al. 1985).

30
Many relatively simple explanations for song matching exist:
Matching indicates high-intensity interactions (Lemon 1968b). Birds
match in order to monitor the positions of their neighbors (Lemon
1968a). Relative dominance between countersinging birds influences the
degree of matching (Kroodsma 1979). Time intervals between songs of
countersinging birds indicate which of the participants is dominant
(Smith & Norman 1979). Matching functions as a vocal threat that
regulates distance between birds (Todt 1931). Matching is a graded
signal used in territorial encounters (Krebs et al. 1981). Finally,
matching indicates readiness to interact aggressively (Horn 6 Falls
1986).
Further interpretations of song sharing that deal more generally
with repertoires and repertoire sharing (which may or may not involve
matched countersinging) are: Larger song repertoires make recognition
of neighbors more difficult (Falls & d'Agincourt 1981; Searcy et al.
1981). Larger repertoires are more attractive and stimulating to
females (Howard 1974; Searcy et al. 1982). Larger repertoires allow
resident birds to fool potential intruders through increasing the
perceived density of singing birds (Krebs 1976), and, young indigo
buntings (Passerina cyanea) mimic the repertoires of older territorial
males and thus deceive other males through mistaken identity (Payne
1982). Searcy et al. (1985) found no correlation between song repertoire
size and male "quality" in song sparrows (Melospiza melodia).
While these may or may not be adequate explanations for song sharing
in birds, some recent experiments on song sharing have become
increasingly sophisticated. These studies have incorporated more
objective methods of determining song similarity, the possible effect of

81
sound degradation from sender to receiver influencing behavior, the
birds' natural use of their repertoire, and their familiarity with
neighboring males (Falls et al. 1932; Schroeder & Wiley 1983b; Whitney &
Miller 1983).
Almost all investigations of song matching and repertoire use have
involved playback experiments and measures of response strength to the
tape recorded songs. Ideally, field studies to determine the occurrence
and significance of song sharing (without using controlled playback
experiments) would be on birds with only one or two songs in their
repertoires. Usually, though, this is not the case. Therefore a major
design consideration in natural observational investigations is first to
make comprehensive studies of normal singing behavior of undisturbed
birds. That is, the song repertoires (and the possible differential use
of songs from within the repertoires) of all subjects must be thoroughly
analyzed, and the probabilities that each bird will sing a given song in
different circumstances determined. Simply describing the composition of
the bird's repertoire is not adequate.
Studies of the birds' natural use of song types are complicated by
additional considerations. The observer's method of determining degree
of song similarity may greatly influence the results. Also, some birds
are known to use different song types according to: location within
their territory (Lemon 1968b; Lein 1973; Schroeder & Wiley 1983a),
whether a male or female is present (Smith et al. 1973; Schroeder &
Wiley 1983b), distance from rival conspecifics (Simpson 1985);
familiarity with the neighboring male (Wunderle 1973; Falls et al.
1982); similarity of rival male's song to self song (McArthur 1986),
rate of song type switching (Simpson 1985), acoustic degradation over

32
distance (Morton 1982; McGregor et al. 1983); overall aggressive
motivation (Lein 1973; Schroeder & Wiley 1933a); and stage in breeding
(Falls & Brooks 1975; Petronovitch et al. 1976). Furthermore, three
recent studies have shown that the intensities of agonistic interactions
were not correlated with the choice of song types per se, but rather
with the rate of song type switching alone (Kramer et al. 1985; Simpson
1985) or together with the degree of matching (Horn & Falls 1986).
Thus, what may appear to be unambiguous results from studies of how
birds use their repertoires, including song matching during
countersinging, should be carefully evaluated. Some recent studies
have, however, more scrupulously investigated countersinging and song
type use in general based on detailed analyses of repertoire composition
and contextual use (McGregor & Krebs 1982; Schroeder & Wiley 1983a,
1983b; Simpson 1984, 1985.; Lemon et al. 1985).
The purpose of these introductory remarks is several-fold. First, a
field study seeking to describe and interpret the function of repertoire
sharing and countersinging is likely to be more complicated than is
intuitively obvious. The design must consider many variables that may
be difficult, if not impossible to control. Secondly, I introduce this
section on the countersinging of Scott's seaside sparrows with these
remarks in order that the reader may realize why I present only
descriptive summaries rather than the testing of the significance of
repertoire sharing and matched countersinging.
Methods of investigating repertoires
When birds countersang during time budget observations, I noted
whether they matched or not, in addition to taking my standard singing
behavior measurements (bird's identity, presence or absence of

33
intruders, time singing, perch changes while singing, and incidence of
song type switching). As described in the "General Methods," I
tabulated in detail 103 time budgets (selected for comparability of
date, weather, absence of intruder, etc.) for 30 mated males observed
Apr-May 1981-1984.
Results and discussion of repertoire use
I measured parameters of solo primary and countersinging activity of
Scott's seaside sparrows (Table III-3). The mean total singing activity
(solo singing and countersinging) for males was 12.29 min song/h.
Two to four males often participated in bouts of countersinging, the
alternation of songs separated by 3-8 sec (Fig. III-9). The pattern of
song alternation was usually regular, but in 3% of the countersingng
bouts the alternating songs drifted out of phase and began to overlap.
Countersinging comprised about half of the total singing time.
Approximately two-thirds of the total countersinging time was matched
countersong (Table III-3).
The mean song repertoire size of the 30 males was 2.41
songs/bird/season. I did not measure the proportional use of song types
within repertoires, nor did I measure the degree of repertoire sharing
within the population.
Some conclusions are apparent from the data presented in Table
III-3, even without statistical testing. First, apparently neither solo
singing nor countersinging was preferred in the several contexts
analyzed. That is, countersinging did not seem to be used
proportionately more often than solo singing at territory edge, nor did
perch and song type switches seem to be correlated more or less strongly
with countersinging. The same lack of differential patterns according

84
to context seemed to exist when comparing matched to non-matched
countersinging. A possible exception is that birds may have sung more
minutes of matched song than non-matched song at territory edges.
Of course more comprehensively analyzed data are needed before
statistically valid statements regarding repertoires and countersinging
in seaside sparrows can be made. I do, however, conclude the following
at this time. First, countersinging is a form of vocal "duelling" that
indicates a slightly elevated aggressive or aroused state. That is,
countersinging is a graded signal in seaside sparrows, but probably a
less strongly graded signal than in most birds. Second, song matching
while countersinging and repertoire sharing in general in these birds
are probably chance events, analogous to the "epiphenomena" of song
pattern variety suggested by Wiens (1982) for sage sparrows (Amphispiza
belli). Songs of seaside sparrows may be observed to match simply as a
consequence of their small repertoires of simple and similar songs,
their low population turnover rates, and their sedentary habits.
Flight Songs: Description and Comparison to Perch Songs
Description
An early Florida naturalist, Donald J. Micholson aptly
characterized seaside sparrow singing: "Towards the end of March the
marshes are fairly buzzing with the purring, wheezing songs...He tries
one perch and seeks another fifty or seventy-five yards away flying low
over the rank growth. Every so often he fairly 'explodes' with passion
leaving his concealment to rise on fluttering wings sixty or seventy
feet above the marsh uttering his erratic little song as he goes up and
down dropping out of sight in the salicornia" (Nicholson 1946, p. 41).

85
The flight song (Fig. III-10), accompanying the flight song display,
is a combination of preliminary calls followed by the primary song. This
vocalization and display of about 3-4 sec is sporadically given by males
throughout the breeding season. It begins with introductory
high-pitched "si" calls, followed by lower-pitched "tuck" calls, and
ends with one or two condensed versions of the primary song. The buzzy
trill of the primary song portion is about 0.4 sec shorter than the
trill of the normal primary song. The introductory calls vary in
duration and type, but are always given on the ground and during the
bird's ascent to about 10-15 m. The primary song is uttered as the bird
descends. Woolfenden (1956), Trost (1968), Post & Greenlaw (1975), and
Werner & Woolfenden (1983) also describe singing and flight songs of
seaside sparrows.
Methods of investigating flight song activity
My observations of Scott's seaside sparrows during the breeding
seasons of 1981-1982 suggested that overall song activity seemed to be
affected by whether the birds were mated, and by time of day, time of
year, and weather. In most Passerines these factors do influence the
intensity of primary song singing from a perch (hereafter designated in
this section as "perch singing"). I measured and compared perch song
and flight song activities of my study population during the early
breeding season of 1982. Then I tested whether the above factors
differentially affected flight song activity (as compared to perch
singing) by comparing the relative frequencies of flight songs to perch
singing under specified conditions described below.
I sampled the overall singing activity of 28 males during 31 field
days from 12 Feb-26 Apr 1983. Throughout the day I monitored flight

8ó
song and perch song activity within one hour time blocks from pre-dawn
through mid-afternoon. The time blocks were designated in hours
relative to sunrise: the first hour block of a day began 15 min before
sunrise, the second hour block began 45 min after sunrise, and so on. I
randomly chose and timed the perch singing of a different male during
each hour block. Simultaneously, I counted all flight songs heard within
100 m during the hour. As I counted and timed songs during each hour
block, I also noted wind velocity, temperature, the singing bird's
location, and identity and mated status (if known).
In summarizing my data, I defined an hour's "flight song activity"
as the average number of flight songs heard within the 100 m radius
during the hour. I defined "perch song activity" as the average number
of minutes the randomly chosen birds sang during the hour within the
same 100 m. Then, for every level of increment of the variables being
investigated (e.g. for temperature category 5-9° C), I tabulated flight
song activity (FS), perch song activity (PS), and a calculated FS/PS
ratio. A statistical analysis using multiple regression would have been
the ideal method of exploring these data. Yet because some data were
ordinal rather than interval and normality was not assumed,
non-parametric statistical analyses were required. In order to validly
consider the effect of only one variable at a time, I chose subsets of
data, as described below for each analysis. I used Spearman rank
correlation (Siegel 1956) to determine if PS, FS, and FS/PS ratios were
correlated with day of year, time of day, temperature, and wind
velocity. I tested significance of the correlation coefficient at the
two-tailed P_<0.05 level. I also compared FS and PS activities of mated
2
and unmated birds with a two-tailed _X one-sample test (Siegel 1956).

87
Results of singing activity analysis
Males sang perch songs at an overall rate of 17.1 (SE=5.9) min
song/h. Flight song rate varied considerably among males. The mean rate
was 0.15 flight songs/male/h. The rates of perch songs, flight songs,
and FS/PS ratios did, however, vary significantly as detailed below.
Four variables correlated significantly with both perch singing and
FS/PS ratios: day of year, time of day, temperature, and wind velocity.
Mated and unmated birds differed significantly in the number of flight
songs given.
Day of year. I analyzed the following subset of data to determine
the correlation of singing activity with day of year: 88 h of
measurements for unmated birds taken on 19 days when the temperature was
15-24° C and the wind was <24km/h. I found that perch song activity
increased significantly (r=0.72), and FS/PS ratios (r=-0.31) decreased
significantly as the breeding season progressed, up until the last week
of April. Flight song activity considered alone, however, did not
correlate significantly with day of year (r=-0.01).
Time of day. I determined the effect of time of day by considering
the same subset of observations: 38 hours of measurements for unmated
birds noted when the temperature was 15-24° C and the wind was <24km/h.
I found perch song rate was greatest just after sunrise and declined
significantly (r=-0.95) as the day progressed (excluding evening
singing). Flight song activity remained about the same all day,
although the FS/PS ratio increased significantly (r=0.33) through the
day (Fig. III-ll).
Wind. I analyzed the following subset of data to determine the
effect of wind: 79 morning hours (hour blocks 1-5) of measuremets for

83
unmated birds noted when temperatures were 15-24° G. I found that brisk
winds had a definite dampening effect on perch song activity (r=-0.97)
but no significant effect on flight song activity. The FS/PS ratio
increased significantly (r=0.88) as wind velocity increased (Fig.
HI-12).
Tempera ture. To determine the influence of temperature, I
considered 77 hours of observations made on unmated birds, during hour
blocks 1-4, and when the wind was <24km/h. I found that cooler
temperatures significantly (r=0.88) decreased perch song activity but
had no significant effect on flight song activity. The FS/PS ratio did
significantly decrease (r=-0.83) as temperature rose (Fig. III-13). In
other words, as with high winds, birds gave proportionally more flight
songs in cooler weather.
Mated status. I considered the singing activity of 23 males before
and after they were mated. The time elapsed between measurements was
8-19 days. I observed birds for 73 h during hour blocks 1-4, when
temperatures were 15-24° C and the wind was <24km/h. Males sang less
(14.9 min/h) after they were mated than before mated (19.7 min/h), but
this decrease in singing may have been at least partially due to the
seasonal decline in singing activity described above, or vice versa.
From a sample of 122 flight songs, 74 were given by birds of known
mated status. Unmated birds sang 84% of these 74 flight songs (Fig.
2
III-14). This difference was highly significant (two-tailed )C
one-sample test, X2=33.3, df=l, £<0.001).
Discussion of flight songs
Flight songs are characteristic of birds that live in open
grasslands or tundra. Many emberizine sparrows, including the

89
congeneric seaside, sharptail, Baird's (A. bairdii), and Le Conte's
sparrows (A. leconteii), have well-developed flight songs.
Why do seaside sparrows give flight songs? Although the flight song
is a highly conspicuous vocalization, obviously its energetic cost is
greater than that of perch singing. Post & Greenlaw (1975) report that
in their migratory northern seaside sparrows, flight songs are much
more common soon after females arrive on their male's territories. This,
as well as my findings that unmated Scott's seaside sparrows gave more
flight songs than mated birds, suggests that flight songs may be used in
mate attraction. All seaside sparrows, however, continue to give these
vocalizations throughout the breeding season.
A major portion of the flight song vocalization is the uttering of
one or two modified primary songs. Elsewhere (Chapter IV) I have shown
experimentally that the primary song is essential for both mate
attraction/retention and territory establishment/retention. Thus, it is
reasonable to assume that flight songs can effect the same critical
functions. I have presented evidence that birds give relatively more
flight songs (and sometimes they give only flight songs) during
inclement weather and when unmated. 1 therefore suggest that even
though they may be energetically costly, flight songs so effectively
magnify the message of the primary song that they are worth more to the
male, especially under otherwise adverse singing conditions.

Figure IIL-11. Relationship between time of day and the ratio of Flight songs/Perch singing.

3S1ÜNHS (ajojeq u¡iu 90 y31dV SdCIOH
Sd/Sd
OllVd

Figure III-12. Relationship between wind velocity and the ratio of
Flight songs/Perch singing, as defined in text. The first bar indicates
the FS/PS ratio when there was no wind; the second bar indicates the
ratio when the wind was 1-3 km/h; the third, 9-16 km/h, and so on.

93
WIND VELOCITY (KPH)

Figure III-13. Relationship between temperature and the ratio of Flight
songs/Perch singing, as defined in text. The first bar indicates the
FS/PS ratio when the temperature was 5-9° G; the second bar indicates
the ratio when the temperature was 10-14° C, and so on.

95

Figure III-14. Number of Flight songs recorded in early 1983, given by
birds of designated mated categories.

97
70—
60-
50—,
40-
no.
F 3
30-
20-
10-
0
UN¬
MATED
MATED
UN¬
KNOWN

CHAPTER IV
FUNCTION OF SONG IN SCOTT'S SEASIDE SPARROW
Introduction
"Why do birds sing" is one of the oldest questions in ornithology.
Song meaning and function (Smith 1977) vary among bird species and
sometimes within species. Meaning is the combined product of signal and
context manifested as a change in behavior; function is the way in which
the behavioral change benefits the signaler by ultimately contributing
survival value and promoting reproductive success.
Functions of avian vocalizations, including calls, are reviewed by
Thielcke (1970). Song is used primarily for territorial establishment
and defense, mate attraction and maintenance of pair bond, coordination
of reproductive behavior between mates and within the population,
signaling motivation, and species and individual recognition (Beer
1982). The role of song in individual recognition is discussed by Falls
(1982), and the function of song in species recognition is reviewed in
Becker (1982).
The "dual function of song" hypothesis prevails in most explanations
of why birds sing: a male bird sings to establish and maintain a
territory and to attract and retain a mate. Early writers emphasized
the apparent dichotomy between mate attraction and territorial function
(reviewed in Armstrong 1973). These two proximate functions, however,
are not necessarily mutually exclusive; both can be accomplished
simultaneously. Tinbergen (1939) made this clear and proposed the term
98

99
"advertising song." The duality of song function and its ultimate role
in promoting reproductive success are reviewed in Catchpole (1982).
I directly tested the dual function of song hypothesis with field
experiments in which I temporarily muted male seaside sparrows. I chose
this approach for the following reasons. A significant design problem in
the study of bird song has always been how to separate the auditory
component of song from the visual stimulus of the vocalizing and
displaying bird. One can remove the bird and replace it with recorded
song, or remove the song from the bird. Using the first approach, many
excellent studies have indirectly tested the role of song with
combinations of male removals, models, and playback experiments
(Catchpole 1977; Krebs 1977b; Krebs et al. 1978, 1981; Yasukawa 1981;
Kramer et al. 1985). The second approach—experimentally muting birds
in the field—has been reported for only one species, red-winged
blackbirds.
Peek (1972) and Smith (1976) permanently muted male red-winged
blackbirds by sectioning the hypoglossal nerve. They found that muted
birds suffered heightened territorial losses and trespass rates,
compared to sham-operated control birds. For several reasons it was
difficult to determine to what extent the muted males were less
successful in attracting and retaining mates. Red-winged blackbirds are
polygynous, and females may discriminate among males on the basis of
territory quality (Smith 1972; Holm 1973; Lenington 1980). Only limited
anecdotal information on mate retention was reported for these muting
experiments. The authors also pointed out that the muted birds'
behaviors were influenced by the administration of anesthesia and by
respiratory impairment.

100
In 1976 and 1979 Smith described a different field muting technique
that overcame the complications of anesthesia and respiratory distress.
Without using anesthesia, he temporarily muted red-winged blackbirds by
cutting the membrane of the interclavicular air sac, one of the air sacs
that propels air into the syrinx. Although Smith used only one sham and
three experimental birds, his results suggested that there were more
intrusions on the territories of muted males, and that muted birds used
the "sidling" visual display more frequently than before they were
muted. As in the previous experiments, it was unclear whether females
reacted differently to muted versus non-mu ted males.
In designing a test of song function, I sought to avoid the
complications of previous field muting experiments. I used a
modification of Smith's temporary muting technique, with larger sample
sizes controlled for time of breeding season, and collected more
quantitative data. Furthermore, I used a species more representative of
songbirds in general. Extrapolating from the experiments on red-winged
backbirds is problematic because, as already mentioned, they are
polygynous, and also sexually dimorphic, actively displaying birds whose
visual behaviors are used in conjunction with their vocalizations. In
contrast to red-winged blackbirds, monogamous and cryptically-colored
seaside sparrows have few visual displays. They nest in dense
vegetation where visibility is reduced. Being songless should have an
especially marked effect on seaside sparrows, whose "displays" are
virtually all acoustic.
My muting experiments specifically tested the two predictions of the
dual function of song hypothesis: 1) Male seaside sparrows sing to

101
attract and retain mates; and 2) Male seaside sparrows sing to establish
and maintain territories.
Me thods
Project history, subjects, and study site were discussed in Chapter
II.
Experimental Design
Muting experiments simultaneously tested the predictions of the dual
function of song hypothesis: female attraction/retention and territory
acquisition/raaintenance. The experimental design is summarized in Table
IV-1.
I administered two rounds of temporary mutings (and sham-operations)
in 1983 and 1984. Each round was preceded and followed by time budget,
mated status, and other observations of the experimental (Muted),
control (Sham-Operated), and Undisturbed birds. The first round of
mutings (Early Season: 1-3 Apr 1983; 17-23 Mar 1984) allowed comparison
of the proportions of Muted birds acquiring mates and territories to the
proportions of Sham-Operated and Undisturbed birds acquiring mates and
territories. The second round of mutings (Mid-Season: 27 Apr-5 Jun
1983; 1-25 May 1984) permitted investigation of the role of song in mate
retention and in territorial integrity. These second round experiments
allowed measurement and comparison of: incidence of intrusion and
reaction to intruders; repulsion success; reaction to playback
(simulated intrusion); loss or retention of territory; territory
boundary change; mated status change; and time spent in singing and
other vocalizations and behaviors (Table IV-1). Some of these measured

102
Table IV-1. Experimental design of muting experiments
a b
Muting Event/Behavior Measured ’
Round
Early- Female attraction
Season Territory acquisition
Mid- Female retention
Season Territory retention
Territory size change (in )
Q
Behaviors Measured When No Intruder Present
Primary song (or attempt) time
Whisper song time
Countersong time (matched)
Countersong time (not matched)
Tuck call time
Tuck/Tsip call time
Si Twitter call time
Tchi/Tyu call time
Zuck call time
Tchi flight frequency
Flight song frequency
Female (mate) on territory time
Female (mate) aggressive call time
Female (mate) chase intruder time
Female (mate) solicitation call time
Associate/copulate with female time
Feeding, resting, and off territory time
Behaviors Measured While Intruder Present
Intrusion occurrence frequency
Repel intruder frequency
Not repel intruder frequency
Total intrusion time
Intrusion time >5 m between birds
Intrusion time 0.5-5 m between birds
Intrusion time <0.5 m and contact
Primary song (or attempt) time
Whisper song time
Countersong time (matched)
Countersong time (not matched)
Tuck call time
Tuck/tsip call time
Si twitter call time

103
Table IV-1—continued
Mid-
Season- Event/Behavior—continued
cont.
Behaviors Measured While Intruder Present——continued
Tchi/Tyu call time
Zuck call time
Tchi flight frequency
Wing raise time
Chase time
Grass pick time
Sham-preen time
Bob and bill jab time
Behaviors Measured During Playback Experiment
Primary song (or attempt) time
Whisper song time
Countersong time (matched)
Countersong time (not matched)
Tuck call time
Tuck/tsip call time
Si twitter call time
Tchi/Tyu call time
Zuck call time
Tchi flight frequency
Wing raise time
Chase time
Grass pick time
Sham-preen time
Bob and bill jab time
Latency of reaction time
Within 25 m model time
Within 5 m model time
Within 0.5 m model time
Contact model frequency
3Changes in Muted birds' territory sizes were compared Before-During,
and During-After their songless states and tested with one-tailed
Wilcoxon matched-pairs signed-ranks tests (WSR). Behavioral changes were
compared Before-During (but not During-After) with two-tailed Wilcoxon
matched-pairs signed-ranks tests.

104
Table IV—continued
^Differences between the groups' (Muted, Sham-Operated, and Undisturbed)
abilities to acquire and re,£ain mates, and acquire and retain
territories compared with X~ or Fisher exact probability tests. Because
size change in one treatment group may cause concurrent size change in
another group, territory size comparisons among the 3 groups should not
be made with X~ or Mann-Whitney U_-tests, which assume independence of
groups. Thus one-tailed Wilcoxon matched-pairs signed-ranks tests are
used to compare indirectly size changes for treatment groups (see text).
Behavioral differences between the groups (Muted to Sham-Operated, and
Sham-Operated to Undisturbed) compared with two-tailed Mann-Whitney U_
tests (MWU).
frequency measured as occurrences/hour. Time measured as minutes/hour.

105
behaviors were only rarely seen; thus similar behaviors were grouped for
statistical analyses of results.
I made two types of comparisons for each round of mutings. First I
compared treatment groups: Muted to Sham-Operated and also Sham-Operated
to Undisturbed. Additionally, I was interested in how being muted
affected a bird; therefore I made Before, During, and some After
comparisons for the individuals.
My experimental protocol was as follows. After preliminary
observations on birds as they began to sing in early February, I
designated approximately 30 males each year to be Muted, Sham-Operated,
or Undisturbed birds for the Early Season round, and then re-designated
a treatment for the Mid-Season round. The treatment assignment was
random, except: no bird underwent the same treatment twice; territories
of muted birds were not contiguous; and only mated males were assigned a
treatment and studied in the Mid-Season round.
I chose the dates of the Early Season manipulation each year
according to when heightened male-female pairing behavior began. Birds
were also in the process of establishing territories at this time. They
had preliminary territories, but boundaries were not fixed and changes
in ownership sometimes occurred during those early spring weeks. I
considered "territory acquisition" to be successful defense of any area
on the study site (or the censused adjacent marsh) within 0-10 days
after manipulation.
For both rounds of manipulation, I observed each of the subjects at
least every third day for a total of >20 h (or more) per season.
Behavioral data used for analyses were those collected 0530-0930 and
under moderate weather conditions. Muted and Sham-Operated birds were

106
studied more intensively just before and after their surgery. I drew
territorial maps during every observation period and described
behavioral events in detail on cassette tapes and field data forms. I
later transcribed notes and tabulated and analyzed the data.
To investigate further behavior under more controlled conditions, I
simulated territory intrusion with playback experiments. 1 hid a
playback tape recorder at the approximate center of the bird's territory
and placed a stuffed model seaside sparrow in Juncus 1 m above the
ground. The seaside sparrow playback song had been recorded in 1981, 2
km from the study site. It was recorded on a Sony TC 150 cassette
recorder (Mineroff-modified) with a Bell and Howell "Shotgun"
unidirectional electret condenser microphone with wind screen. The song
was played back with the same recorder at what I estimated to be a
normal song volume. The playback period consisted of 7 min blank tape,
8 min song, and 15 min blank tape. I tested all Muted, Sham-Operated,
and Undisturbed birds on three occasions: Before (1 day prior), During
(3 days following), and After (20 days after) their manipulations.
Reactions listed in Table IV-1 (Playback Experiments) were noted during
the 30 min experiment while the tape ran. For comparative purposes,
playback experiment results were standarized, from min/0.5 h playback to
min/hour by multiplying by two.
Additional recordings of vocalizations were made with the equipment
described above. Audiospectrograms were prepared on a Kay Elemetrics
7029A Sona-graph (wide-band [300 Hz] filter and 80-8000 Hz scale).
Muting Procedure
Birds muted or sham-operated were first caught in mist nets on their
territories. After surgery I carried the bird back to his territory and

107
released him. I used a modification of Smith's technique of
temporarily muting the bird by rupturing the interclavicular air sac
membrane (Smith 1976 and 1979). In pilot work, my first attempts to
mute birds in the laboratory and field were unsuccessful for several
reasons. I had difficulty adjusting the anesthesia dosage to a level
that would relax the bird yet not persist for hours, thus exposing the
bird to predation, inclement weather, and food deprivation. I decided
that the most humane and experimentally straightforward alternative was
to perform the surgery quickly and without anesthesia, which enabled me
to release the bird back on his territory within 15 min after capture.
Another source of early failure was my closing the skin incision with
too many sutures, thus offsetting the desired effect of rupturing the
air sac to direct air away from the syrinx.
I present a stepwise instructional outline of my temporary muting
procedure:
1. Hold the bird in one hand and work with the other. Keep the head and
wings from getting in the way with fingers of your holding hand.
2. Part the feathers in the area of the furculum and neck, then swab
with antiseptic. I used Betadine solution (Purdue Frederick Company).
3. Mat down the parted feathers with water-soluable lubricating jelly.
This step is unnecessary if one works quickly enough so that the wet
antiseptic holds the feathers away from the incision.
4. Make a skin incision from the base of the furculum extending
rostrally about 1.2 cm. I used a new #12 scalpel blade.
5. Push aside connective tissue with blunt forceps and expose the
interclavicular air sac.

108
6a. To mute birds: Grasp the air sac and puncture it. I found this
somewhat difficult—the air sac is thin, slippery, and rapidly rising
and falling in a breathing bird. To make the initial 1 mm puncture, I
used the slightly bent tip point of a #12 scalpel blade, then widened
the hole to about 1 cm by inserting and spreading small forceps. One
must be careful at this step—the small carotid arteries and other
vessels lie just beneath the air sac. Keep a styptic pencil handy for
bleeding emergencies.
6b. To sham-operate birds: Touch the air sac with forceps.
7. Close the skin incision with one suture. I used a sterile 0.5 inch
circular cutting-edge needle attached to 5-0 silk suture.
8. Swab again with antiseptic and comb the feathers back into place.
After practice I was able to perform this procedure in the field in
less than five minutes. Of my total 57 field Muted and Sham-Operated
birds, only four were known or presumed to have died as a direct
consequence of the surgery—two died during surgery and two permanently
disappeared after release.
Statistical Analyses
Non-parametric statistics were used for all analyses (Siegel 1956;
Conover 1980). I compared mate attraction and retention, and territory
2
acquisiton and retention with X~ contingency (or Fisher exact
probability) tests. The individual's Before and During behaviors were
compared with two-tailed Wilcoxon matched-pairs signed-ranks tests.
Wilcoxon tests were also used to compare Before, During, and After
territorial size changes. Because territorial size change in one
treatment group (Muted, Sham-Operated, or Undisturbed) probably caused a
concurrent size change in another group, comparisons of size changes

109
o
among the 3 groups could not be made with >C or Mann-Whitney U_-tests,
because both assume independence of groups. Therefore I indirectly
tested the significance of territory and size changes between groups
using Wilcoxon tests. That is, 1 noted whether there was a significant
size change from Before to During in the Muted group and whether there
was a significant size change in the Sham-Operated group from Before to
During. Territorial loss was included in this analysis by considering it
2
to be a size change from the original area to 0 m area. Likewise, in
the analysis of size change from During to After, permanent territorial
2
loss was considered to be an expansion of 0 m in area. All other group
comparisons (Muted, Sham-Operated, and Undisturbed) were made with
Mann-Whitney U tes ts.
Results
Voice and Post-Operative Recovery
I found Muted and Sham-Operated birds back on their territories a
few hours to two days after their surgery. Sham-Operated birds sang and
apparently behaved normally in every respect. Comparisons between
Sham-Operated and Undisturbed groups revealed only one significant
difference among all of the attributes listed in Table IV-1. Thus,
Undisturbed birds were pooled with Sham-Operated birds for some
statistical tests, as noted.
Two unexpected yet fortuitous outcomes of the muting manipulation
were: 1) Muted birds attempted to sing by going through the motions of
singing; and 2) Muted birds gave all of their normal calls. Thus these
birds were actually "de-songed" rather than completely "muted," although
for convenience I use the latter term throughout this paper.

no
By Day 5 following surgery most of the muted birds began to utter
croaks and squeaks as they attempted to sing. Nearly all Muted birds
fully recovered their reportoire of 2-3 distinct song types by Day 12.
Fig. IV-1 shows the audiospectrogram of Before song, and Fig. IV-2 shows
the During "song'' of a muted bird. Within 10 days after surgery skin
incisions healed completely and the suture dropped out. I never
detected any infection or other detrimental consequence of the surgery
other than the mortality mentioned above.
Mate Attraction and Retention
Results of the Female Attraction and Female Retention tests clearly
indicate the importance of song in obtaining and retaining a mate
(Fig. IV-3). None of the 10 Early Season Muted birds had obtained a
mate by Day 7 after surgery, as contrasted to 11 of the 13 Sham-operated
and 32 of the 37 Undisturbed birds having mates by then. The difference
between Muted and Sham-Operated birds is highly significant (one-tailed
Fisher exact probability test, P<0.0001).
All female mates of the 21 Mid-Season muted birds began ignoring
their mates immediately following manipulation. These females did,
however, continue giving solicitation calls at about the same frequency
and remained on the territory for 1-2 days. By three days post-muting,
most females deserted their mates and paired with intruders or
neighboring males. Only one muted bird retained his female beyond three
days after manipulation. I never saw her associating with her
mate; she probably stayed on his territory because she had a nest with
hatchlings. This mate retention difference between treatment groups is
? 2
also highly significant (one-tailed X_ 2X2 test, =59.6, N=21 Muted and
13 + 30 pooled Sham-Operated + Undisturbed birds, df=2, P_<0.001).

Figure IV-1. Audiospectrogram of bird ABOR recorded 3 days prior to muting.

N
X
O
TIME IN SECONDS
1
112

Figure IV-2. Audiospectrogram of "songs" of muted bird ABOR recorded 5 days after muting.

VII

figure IV 3. Relative abilities of Muted versus Sham-Operated and Undisturbed males to attract (a) and
retain (B) females, 19&3 and 1934 combined* Sample sizes as given.

PROPORTION OF TOTAL MALES (%)
100—1
75
50
25-
0
N=13
N = 1 0
N=37
MUTE SHAM UND
ATTRACTED
A
N=13 N=30
B
116

117
Table IV-2. During songless comparison of territory ownership and size
changes for the Mid-Season mu tings
Treatment
Change
Sample
Size
Lost
Terr.
Smaller
Terr.
Expanded
Terr.
No
Change
Mean Size
Change
Muted
21
6
15
0
0
-79/**
Sham-Opera ted
13
0
1
10
2
+44%**
Undisturbed
30
0
0
17
13
+31%**
**Significant size change from Before to During, one-tailed Wilcoxon
matched-pairs signed-ranks test, P<0.005. Also see text.

118
Table IV-3. "After song regained" comparison of territory ownership and
size changes for the Mid-Season mutings
Change
Sample New
Smaller Expanded
"Shared"
No
Mean
Treatment
Size Terr.
Terr.
Terr.
Terr.
Change
Change
Muted
21 3
0
14
3
1
+76%*
Sham-Opera ted
13 0
8
0
0
5
-13Z*
Undisturbed
30 0
12
0
0
18
-15%*
*Significant
size change from During
to After,
one-tailed Wilcoxon
matched-pairs
signed-ranks
test, P<0.
005. Also
see text.

119
Territory Establishment, Retention, and Size Change
Territory acquisition was delayed for the songless males. None of
the 10 Early Season muted birds had obtained a territory by 10 days
after manipulation, in contrast to 10 of the 13 Sham-Operated and 23 of
the 37 Undisturbed birds. This difference was highly significant
2 2
(one-tailed £ 2X2 test, pooled Sham-Operated + Undisturbed, £ =25.4, 1
df, £<0.001). Eight of the muted birds eventually obtained territories
by 12-29 days (mean = 13.2) after surgery.
The Mid-Season post-muting changes in territorial ownership and
sizes are summarized in Tables IV-2 and IV-3. Six muted birds incurred
territory losses during the time they remained songless compared to no
Sham-Operated or Undisturbed birds losing their territories. This is a
highly significant difference (one-tailed Fisher exact probability test,
N=21 Muted and 13 + 30 pooled Sham-Operated + Undisturbed birds,
£<0.001). Expansion of adjacent Sham-operated and Undisturbed birds'
territories caused significant territory size decrease for the remaining
15 muted birds. This territory size decrease for Muted birds from
Before to During was highly significant (one-tailed Wilcoxon
matched-pairs signed-ranks test, £=0, £=21 paired observations,
£<0.005). There was a Before to During significant size increase for
both the Sham-Operated (T=2, £=13 paired observations, £<0.005) and
Undisturbed groups (£=0, £=30 paired observations, £<0.005).
Territorial changes at 10 days after the Muted birds regained their
singing ability were also evident (Table IV-3). Many ex-muted birds
expanded nearly to their original territorial boundaries. Three
established new territories, and three remained on ("shared") their
original territories, seemingly behaving subdominant to the males that

120
had taken over these territories. One-tailed Wilcoxon tests indicated a
significant size increase for ex-muted birds (T=0, N=17 paired
observations, P_<0.005) and significant size decreases for Sham-Operated
0T=O, N==8 paired observations, P<0.005) and Undisturbed birds OT=0, Nj=18
paired observations, P<0.005).
Behavioral Changes of Muted Birds
I compared Muted birds' behaviors Before and During their songless
states, and also compared the behavior of Muted to Sham-Operated and
Sham-Operated to Undisturbed birds. I did not compare statistically
"After" to "Before" and "During" behaviors because by the time song was
fully recovered some birds of all treatment groups exhibited a marked
natural seasonal decrease in territorial and other observable behavior.
The results of Before and During behavior for individuals (Muted birds)
and a comparison of Muted, Sham-Operated, and Undisturbed groups are
given in Tables IV-4 and IV-5, respectively. Considering all
comparisons, the behaviors of Sham-Operated and Undisturbed groups
differed significantly in only one measurement—primary song time during
intrusion. I have no explanation for this difference, other than the
fact that the cummulative probablility of a Type I error becomes great
when many comparisions are made.

121
Table IV-4. Behavior changes for individuals, from "Before" to "During"
mu ted
Event/Behavior Individual Comparisons
(Muted Birds)
Before
During
P
(A) No Intruder Present
Primary song (or attempt) time'-
11.99
7.37
kkk
Flight song frequency
0.10
0.06
â– kirk
Tuck, Tsip, Si Twitter call time
<0.01
0.03
NS
Tchi-Tyu and Zuck call time
<0.01
0.02
NS
Female solicitation call time
0.45
0.47
NS
Associate with female time
2.03
0.00
'ick'k
(B) Intruder Present
Intrusion occurrence frequency
0.11
3.69
kkk
Repel intruder frequency
0.11
0.00
kkk
Total intrusion time
0.42
33.20
irk'k
Intr. time >5 m between birds
0.29
23.53
kirk
Intr. time 0.5-5 m between birds
0.19
8.06
kkk
Intr. time <0.5 m and contact
<0.01
1.61
k'k'k
Primary sogg (or attempt) timed
Chase timea
1.86
0.28
kkk
2.51
4.71
kkk
"Alarm/Threat" call time
"Attack" call timed
<0.01
1.81
kkk
<0.01
<0.01
NS
Wing raise time^
2.52
4.71
kk
Grass pick time^
0.02
0.20
kkk
Sham-preen time
0.00
2.65
kkk
Bob and bill jab time
0.00
0.14
kkk
(C) Playback Reaction
Primary song (or attempt) time
4.59
3.34
k
Tuck, Tsip, Si Twitter time
11.10
10.77
NS
Tchi-Tyu and Zuck call time
0.10
<0.01
NS
Tchi flight frequency
0.77
0.62
NS
Wing raise time
0.59
0.79
NS
Grass pick time
0.49
0.55
NS
Sham-preen time
3.83
3.41
NS
Bob and bill jab time
0.36
0.35
NS
e
Latency of reaction time
0.88
0.99
NS
Within 5-25 m model time
22.04
21.11
NS
Within 0.5-5 m model time
12.41
11.52
NS
Within 0.5 m model time
2.92
2.71
NS
Contact model frequency
1.01
0.91
NS

122
Table IV-4—continued
q
Similar vocalizations and behaviors are grouped: Tuck, Tsip, and Si
Twitter calls ("alarm/threat calls") are given in reaction to nearby
(0-15 m) predators and avian invaders; Tchi-Tyu, Tchi, and Zuck call
("attack calls") are vocalizations often used by males and females
invading another territory; Wing raises, grass picking, sham-preening,
bobbing and bill jabbing are visual displays given in the presence of
nearby (0-5m) predators and avian invaders.
^Figures are mean values per bird for 21 muted birds. Minimum
observation time per bird was 10 h for (A) and (B). Time results
reported as min/h; playback results standarized to min/h. Frequency
results reported as mean occurrences/h. Individual Before-During
comparisons tested with two-tailed Wilcoxon matched-pairs signed-ranks
tests. Significance of differences between Before and After:
***P<0.01; **£<0.02; *£<0.05; NS=Not Significant.
C
Includes countersinging with another males off territory.
^Standardized to min/h intrusion.
0
Minutes from first song on tape to first observed reaction; not
standarized to min/h.

123
Table IV-5. Behavior differences comparing Muted, Sham, and
groups
Event/Behaviora
Mute
Group Comparisons
Sham Und P_
M/S
Undisturbed
P_
S/U
(A) No Intruder Present
Primary song (or attempt) time0
Flight song frequency
Tuck, Tsip, Si Twitter call time
Tchi-Tyu and Zuck call time
Female solicitation call time
Associate with female time
(B) Intruder Present
Intrusion occurrence frequency
Repel intruder frequency
Total intrusion time
Intr. time >5 m between birds
Intr. time 0.5-5 m between birds
Intr. time <0.5 m and contact
Primary sogg (or attempt) time^
Chase time11
"Alarm/Threat" call time
"Attack" call ti^me0
Wing raise time0
Grass pick time^
Sham-preen time
Bob and bill jab time0
(C) Playback Reaction
Primary song (or attempt) time
Tuck, Tsip, Si Twitter time
Tchi-Tyu and Zuck call time
Tchi flight frequency
Wing raise time
Grass pick time
Sham-preen time
Bob and bill jab time
0
Latency of reaction time
Within 5-25 m model time
Within 0.5-5 m model time
Within 0.5 m model time
Contact model frequency
7.37
11.43
12.51
*
NS
0.06
0.13
0.11
*
NS
0.03
<0.01
<0.01
NS
NS
0.02
<0.01
<0.01
NS
NS
0.47
0.53
0.42
NS
NS
0.00
2.01
1.94
kkk
NS
3.69
0.07
0.16
â– kirk
NS
0.00
0.07
0.16
kick
NS
33.20
0.51
0.39
-kkk
NS
23.53
0.33
0.20
kick
NS
8.06
0.17
0.19
kkk
NS
1.61
<0.01
<0.01
kkk
NS
0.28
2.62
1.56
kkk
k
4.71
2.52
2.86
kkk
NS
1.81
<0.01
<0.01
kkk
NS
<0.01
<0.01
<0.01
NS
NS
4.71
2.52
2.86
kick
NS
0.20
0.00
0.00
kick
NS
2.65
0.00
0.00
kkk
NS
0.14
0.00
0.00
kkk
NS
3.84
4.71
4.44
k
NS
10.77
12.03
11.11
NS
NS
<0.01
0.11
0.08
NS
NS
0.62
0.86
0.31
NS
NS
0.79
0.55
0.66
NS
NS
0.55
0.22
0.36
NS
NS
3.41
2.98
3.71
NS
NS
0.35
0.22
0.41
NS
NS
0.99
0.91
0.88
NS
NS
21.11
25.37
23.78
NS
NS
11.52
10.37
12.21
NS
NS
2.71
2.69
3.01
NS
NS
0.91
1.11
1.32
NS
NS

124
Table IV-5—continued
Similar vocalizations and behaviors are grouped: Tuck, Tsip, and Si
Twitter calls ("alarm/threat calls") are given in reaction to nearby
(0-15 m) predators and avian invaders; Tchi-Tyu, Tchi, and Zuck call
("attack calls") are vocalizations often used by males and females
invading another territory; Wing raises, grass picking, sham-preening,
bobbing and bill jabbing are visual displays given in the presence of
nearby (0-5m) predators and avian invaders.
^Figures are mean values per bird for 21 Muted, 13 Sham-Operated, and 20
Undisturbed birds. Minimum observation time per bird was 10 h for (A)
and (B). Time results reported as min/h; playback results standarized
to min/h. Frequency results reported as mean occurrences/h. Group
comparisons tested with two-tailed Mann-Whitney U_-tests. "M/S" compares
mute to Sham-Operated groups; "S/U" compares Sham-Operated to
Undisturbed groups. Significance of differences between groups:
***p_<0.002; **P<0.02; *P<0.05; NS=Not Significant.
Q
Includes countersinging with another males off territory.
^Standardized to min/h intrusion.
Minutes from first song on tape to first observed reaction; not
standarized to min/h.

125
Behavior with no Intruders present. Singing rate and other
behaviors were measured under two conditions: unprovoked ("no intruder
present”) and intruder-induced ("intruder present"). The unprovoked
"song" (attempted song) rate and flight song frequency of Muted birds
was less during their songless state than before they were muted. They
also spent slightly less time giving unprovoked song than Sham-Operated
and Undisturbed birds.
Muted birds spent less time interacting with females while muted
than before. Also, compared with the Sham-Operated group, Muted birds
spent much less time with females. Other behavioral observations
collected under non-intrusion conditions showed no significant
differences between Before and During muted, and no difference between
the Muted and Sham-Operated groups.
Behavior during intrusions. Songless birds experienced major
problems in their confrontations with other males. Normally territorial
boundaries are well established by April. Thereafter, neighboring males
squabble only occasionally, and rarely do floaters (non-territorial
males) challege territorial birds. Territory owners usually react to
intruders by first increasing their singing activity, then if necessary
with short-distance aggressive calls and displays, and infrequently with
physical attack. Frequency of intrusions and behavioral reactions to
intrusion differed considerably for muted birds both in comparison with
themselves prior to muting and in comparison with Sham-Operated and
Undisturbed birds. Invasions into the Muted birds’ territories by
neighbors and floaters were immediate, and these provoked heightened
aggressive interactions. Often intruders came in and began to assert
themselves by singing and by associating with the resident females

126
within several hours after the muting surgery. Muted territory owners
reacted to intruders with their most intense and aroused calls and
close-range displays. Intruders usually ignored or simply avoided muted
birds, or seemingly treated them as mere annoyances.
Prior to manipulations I measured intrusion frequency, total time
of intrusion, and birds' behavior during intrusions. 1 then made Before
and During comparisons of intrusion rates and reactions. I compared
total time of intrusion, and Before and During changes in four
categories of behaviors measured for the 21 Mid-Season Muted birds (Fig.
IV-4). Tests of behavioral changes were made with two-tailed Wilcoxon
matched-pairs signed-ranks tests (Table IV-4).
Invaders spent significantly more time on Muted birds' territories
and none was successfully repelled. While songless, Muted birds reacted
to intruders with more chases, "alarm/threat" calls, and close displays
(wing raises, grass picking, sham-preening, and bob and bill jabs) than
they did before being muted. There was no significant difference in the
time birds reacted to intruders with "attack" calls. During their
songless state, however, the Muted birds spent less time reacting to
intruders with song attempts, compared to the time they spent singing
during intrusions before they were muted.

Figure IV—4• behavioral changes of 21 muted birds Before and During their songless periods. Changes are
compared as mean time (min/h) birds were involved in designated activities. The two bars on the left (A)
compare the overall time the bird's territory was invaded upon. The eight bars on the right (b) compared
(in min/h intrusion) how birds reacted during the time intruders were present. All changes were
significantly different (one-tailed Wilcoxon matched-pairs signed-ranks tests), 21 paired observations
pel test.

TOTAL
â–¡ BEFORE
DURING
N=21 MUTED BIRDS
n
i
s\S
i
n
X,
1
1
x\
SHORT PRIMARY
DISTANCE SONG
INTRUSION INTRUSION
CHASE
ALARM/
THREAT
CALLS
SHORT
DISTANCE
DISPLAYS
->
DURING INTRUSION

129
Reaction to Playback
Playback experiments simulated intrusions under more controlled
conditions. I compared the reactions to playback of Muted birds before
and During their manipulation using two-tailed Wilcoxon matched-pairs
signed-ranks tests, and I compared the Mute to Sham-Operated groups with
Mann-Whitney _U tests. I did not compare statistically the After to
Before and During playback reactions because of the end-of-season
decline in observable behavior.
Playback experiment results are summarized in Tables IV-4(C) and
IV-5(C). For all treatment groups the pattern of reactions to playback
was generally similar to reaction to natural intrusion (Tables IV-4(B)
and IV-5(B)). Playback stimulated more intensive reaction behavior
overall than did natural intrusions. Muted birds responded to playback
with "songs" less During muted than Before. Other playback reaction
behaviors did not differ significantly between Before and During muted.
Treatment group comparisons (Table IV-5(C)) indicated that Muted
birds attempted song less that Sham-Operated. Other reaction behaviors
did not differ significantly between Muted and Sham-Operated. For all
birds, their reactions to playback both Before and During was generally
similar to their reactions to natural intrusions (Table IV-4(B)). Muted
birds did, however, react to playback with "songs" slightly less During
muted than Before. Other playback reaction behaviors did not differ
significantly between Before and During for both treatment groups.
Sham-Operated and Undisturbed groups did not differ significantly in any
measurement.

130
Voice Recovery and Subsequent Behavior
After regaining their singing ability in 10-15 days, 8 of the 10
birds muted in the Early Season round eventually acquired mates and
territories. Mid-Season Muted birds did not rebound as quickly. Only 2
of these 21 birds re-acquired a mate by the end of the breeding season.
Most of these males did, however, regain their own or a new territory by
21 days after muting (Table IV-3).
One year following surgery 14 of the 21 Mid-Season muted birds
returned and appeared to function normally in every respect. The
numbers of returning Muted (67% of 21), Sham-Operated (62% of 13) and
Undisturbed (73% of 30) birds did not differ significnatly (X2 2X3 test,
X2=0.11, 2 df, P>0.7 ).
Discussion
Singing ability is unquestionably crucial for a male seaside sparrow
to attract a mate and maintain a territory. These two proximate
functions of song are not, however, mutually exclusive. As discussed in
Catchpole (1981), both female attraction and male repulsion are related
to sexual selection—the first intersexual, the latter intrasexual.
Catchpole proposed that song function may be better represented as a
continuum with predicted characteristics of song at each end (Table
IV-6). For some species proximate song function is mostly intrasexual,
used in male-male interactions. For birds at the other end of the
continuum, the primary function is intersexual, used in male-female
breeding behavior. This model is relevant to seaside sparrow song.
Their singing behavior is a mixture of the characteristics listed in
Table IV-6. They sing short, simple songs and sing after pairing. Yet

131
Table IV-6. Sexual selection related to two main song types
Selection Pressure
Attribute Intersexual Selection Intrasexual Selection
Main proximate
function of song
Female attraction
Male repulsion
Female choice
Direct
Indirect
Song structure
Long, complex,
variable
Songs not repeated
Continuous singers
Short, simple,
s tereotyped
Song types repeated
Discontinuous singers
Contextual
correlations
Stops singing after
pairing
Tend to be migratory
Continues singing
after pairing
Tend to be resident
Direct effect
on males
No matched countersing
Little playback effect
in repelling rivals
Matched countersing
Playback repels rival
males from territory
Direct effect
on females
Larger repertoires
attract females first
Larger repertoires
do not attract
females first
Source: Adapted from Gatchpole (1982)

132
they are continuous singers and probably do not match songs
significantly while countersinging. Experimental results from this
study indicate that seaside sparrow singing falls midway along this
continium, functionally as well as diagnostically.
Why did muted birds behave differently? Their responses to intruders
changed after manipulation, both as compared to themselves prior to
muting and as compared to Sham-Operated controls. A normal bird's first
reaction to territorial threat is increased singing, and this is usually
successful in repelling intruders or intrusion attempts. Because the
muted birds' song attempts were ignored by intruders, however, they
resorted to more intensive defense behaviors—aroused calls, close-range
visual displays, chases, and even contact fights.
These results are consistent with several other ethological studies
where removing socially significant signals resulted in animals spending
more time and energy in intrasexual conflicts as a response to the
behavior of surrounding individuals (Peek 1972; Rohwer 1977). The
increased territorial intrusions and behavioral changes of muted seaside
sparrows were similar in some respects to the changes described for
muted red-winged blackbirds (Peek 1972; Smith 1976; 1979). There were,
however, several notable differences. Muting red-winged blackbirds had
no apparent effect on their mate-retention abilities. Furthermore,
although territorial intrusion rates increased, most males were still
able to maintain their territories. Smith (1979) concluded that the
visual component of the song-spread display, combined with other visual
displays, were effective intruder-repulsion behaviors in red-winged
blackbirds.

133
Smith also speculated that female red-winged blackbirds may also
have at least partially chosen males on the basis of their visual
displays and their territory quality. Female choice based on resources,
the most familiar being territories, is also known in other birds
(Selander 1972). Female seaside sparrows, however, deserted the
territories of their muted mates within a few days after muting. Most
solicited copulations from the nearest persistent singer. Thus female
seaside sparrows seemed to be assessing males based on a behavioral
characterisitic—singing ability—rather than their territory quality.
Because females of muted males associated with singing males who
already had mates, I unintentionally induced temporary artificial
polygyny in my study population. I have no data indicating whether the
females thus sharing non-mu ted males were aware of their polygynous
condition. If they were aware, then apparently these females were opting
to cross the "polygyny threshold" (Orians 1969) and share a mate rather
than remain as the exclusive female on a muted males's territory.
Apparently for female seaside sparrows, monogamy and mate fidelity are
not rigid innate behaviors. The female's immediate choice to associate
with a given male seemed instead to be more of a matter of convenience
and her attraction to males with greater song output.
Intruding floater males seemed to appear from nowhere after I muted
birds. Floaters took over 6 of the 21 Mid-Season Muted birds'
territories within hours to several days. Five other floaters wedged
themselves into the areas adjacent to muted birds territories, causing
these territories to shrink appreciably. Surprisingly, three of the six
lost territories were taken over by floaters that had been territory
owners at the exact location the year before. Prior to these

134
experiments, the existence of floater males had not been detected in
this well-studied population. Perhaps the existence of floaters in avian
populations is more common than generally recognized.
The muting technique described herein has great potential in the
field study of bird song. The role of song in duetting species, in
determining the relative contribution of parents and neighbors in song
learning, and in investigating the relative importance of sexual and
territorial functions of song in other species, could all be approached
using judicious muting experiments. Also of interest would be studies on
visually displaying birds designed to test the relative contribution of
acoustic versus visual display in social interactions.
This study has shown that the sound message, the auditory modality
of singing behavior (and not the visual component), is critical for a
male seaside sparrow to attract a female and maintain his territory.
Because "muted" birds did give normal calls and went through the motor
patterns of singing, there was an unexpected yet perfect control for
comparing "song" to "no song" behavior: the design problem of
separating the visual component of the song display from the auditory
message was overcome. The effect of the surgery itself was surprisingly
minimal, as evidenced by the undaunted behavior of the Sham-Operated
birds. This is one of the first well-controlled field studies that
clearly substantiates the dual function of song hypothesis—birds sing
both to attract females and to repel males. The results clearly show
that inability to sing has devastating consequences to a male seaside
sparrow's reproductive success.

CHAPTER V
CONCLUSIONS
In this dissertation I have described the reproductive biology and
vocalizations of a population of seaside sparrows. I have also reported
my experiments on the function of song and have interpreted my findings.
Scott's seaside sparrows at Cedar Key have presumably lived in a
relatively stable salt marsh habitat, isolated from human interference,
for thousands of years. Thus, it is likely that the birds' reproductive
strategies, behavior, and vocalizations are well-adapted to their
environment.
Despite considerable nest destruction due to predation and flooding,
the population maintained its numbers over the years of this study
because of the birds' repeated nesting attempts and their relatively
high survival rate.
The seaside sparrow's repertoire of distinct calls and a primary
song were used differentially throughout the year. Some vocalizations
conveyed general messages, such as alarm; other vocalizations
transmitted more specific information, an example being the female's
solicitation call. I experimentally determined that the male's primary
song had two functions: mate attraction and retention, and male
repulsion. My experiments clearly substantiated the "Dual Hypothesis"
of song function, which before this project had only been inferred from
observational studies.
135

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BIOGRAPHICAL SKETCH
Mary Victoria McDonald was born in Winchester, Virgina, 29 December
1952. Mary Victoria attended public schools in Charles Town, West
Virgina. In 1967 she and and her family moved to York, South Carolina,
where she went to high school. She attended Wake Forest University
1971-1975 (B.A. biology) and Virginia Polytechnic Institute and State
University 1975-1977 (M.S. wildlife sciences). Her master's thesis was
"A Computerized Environmental System for Virginia Counties." From
1977-1979 Mary Victoria taught biology in the Life Sciences Department
of Southwest Missouri State University. In 1979 she enrolled in the
graduate program of the Department of Zoology of the Universtiy of
Florida. As a graduate student from 1979-1936, she not only studied
seaside sparrows at Cedar Key, but also taught biology courses and
became a Florida Master Gardener.
145

I certify that I nave read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degrees, of
Doctor of Philosophy.
Jjzihn William Hardy, Chair:
Professor of Zoology'
I certify that I have read this study and that in my opinioif it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
Pierce Brodkorb
Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
£eterFeinsih§er
Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
Q SÍ**,. (\J ■ /Lq*a/{/v\,
John H. Kaufmannn 7
iVofessor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
Professor om/Entomology and
hematology
This dissertation was submitted to the Graduate Faculty of the
Department of Zoology in the College of Liberal Arts and Sciences and to
the Graduate School and was accepted as partial fulfillment of the
requirements for the degree of Doctor of Philosophy.
December 1986
Dean, Graduate School

UNIVERSITY OF FLORIDA
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