Group Title: comparative analysis of the vocal communication systems of the Carolina chickadee and the tufted titmouse /
Title: A comparative analysis of the vocal communication systems of the Carolina chickadee and the tufted titmouse /
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Title: A comparative analysis of the vocal communication systems of the Carolina chickadee and the tufted titmouse /
Physical Description: v, 96 leaves : ill. ; 28 cm.
Language: English
Creator: Gaddis, Philip, 1945-
Publication Date: 1979
Copyright Date: 1979
 Subjects
Subject: Carolina chickadee -- Anatomy   ( lcsh )
Titmice -- Anatomy   ( lcsh )
Animal communication   ( lcsh )
Zoology thesis Ph. D
Dissertations, Academic -- Zoology -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: Thesis--University of Florida.
Bibliography: Bibliography: leaves 91-95.
Statement of Responsibility: by Philip Gaddis.
General Note: Typescript.
General Note: Vita.
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Bibliographic ID: UF00099113
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000097066
oclc - 06496833
notis - AAL2501

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A COMPARATIVE ANALYSIS OF THE VOCAL COMMUNICATION SYSTEMS
OF THE CAROLINA CHICKADEE AND THE TUFTED TITMOUSE













BY

PHILIP GADOIS















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





UNIVERSITY OF FLORIDA

1979


j








ACKNOWLEGEMENTS



Sound recording and analysis equipment were provided by the Bioacoustics

Laboratory of the Florida State Museum, Gainesville, Florida.

Financial assistance was provided by a Sigma-Xi grant-in-aid of

research. Vickie Duncan Tillman helped with the field work.

Critical comments and advice were provided at various stages of the

research by Philip Callahan, Peter Feinsinger, J. W. Hardy, John H. Kaufmann,

and Horst Schwassmann. The figures were prepared by Esta Belcher.
















TABLE OF CONTENTS


ABSTRACT . . . . ... .


INTRODUCTION . . . . . . . . . . . .


BACKGROUND ON THE SPECIES STUDIED .


METHODS . . . . . . . . . . .


Study Area . . . .
Observational Techniques.
Definitions . . . .

RESULTS . . . . . . .

Chickadee Subsystem I . .
Structure . . .
Usage and Contexts .
Chickadee Subsystem Ila .
Structure . . .
Usage . . . .
Chickadee Subsystem lb .
Structure, Usage, and
Chickadee Subsystem Ilc .
Structure . . .
Usage and Contexts. .
Chickadee Subsystem Ilia.
Structure . . .
Usage and Contexts.
Chickadee Subsystem Illb. .
Structure . . .
Usage and Contexts.
Chickadee Subsystem IV.
Structure . . .
Usage and Contexts. .
Titmouse Subsystem I .
Structure . . .
Usaqe and Contexts. .
Titmouse Subsystem Ia. .
Structure . . .
Usage and Contexts.
Titmouse Subsystem Ilb. .
Structure, Usage, and


Contexts.

























Contexts.


. Iv


2


. . 13

. . 13
. . 13
. . 13
. . . . 15
. . 15
19
. . 24
. 24
. 26
. 26
. 26
. . 30
. . 30
. . 31
. . 33
. . 33
. . 33
. . 36
. . 36
. 36
. . 39
. . 39
. . . 40
. . 40

. . 40
. . 46
. . . 48
. . . 48


. .











Titmouse Subsystem III . . . . .
Structure . . . . . . . . . . .
Usage and Contexts .. .... .................
Titmouse Subsystem IVa . . . . . ... . .
Structure ...............................
Usage and Contexts . . . . . . ... ..
Titmouse Subsystem IVb . . . . .
Structure ..... ..........................
Usage and Contexts .. . . . . .......
Titmouse Rasp .................................
Structure, Usage, and Contexts . . . .... .

DISCUSSION . . . . . . . . ............


Functional Interpretations . . . .. . .
Chickadee Subsystem . . . . . . .
Chickadee Subsystem Ila . . . . . .
Chickadee Subsystem Ib . . . . . .
Chickadee Subsystem Ilc . . . . . .
Chickadee Subsystem lIla . . ... .
Chickadee Subsystem Illb. . . . . . .
Chickadee Subsystem IV. . . . . . .
Titmouse Subsystem I . . . . . . .
Titmouse Subsystem Ila. . . . . . .
Titmouse Subsystem lib. . . . . . .
Titmouse Subsystem IIl . . . . ... ..
Titmouse Subsystem IVa . . . . . .
Titmouse Subsystem IVb . . . ... .
Titmouse Rasp . . . . . . . .
Intraspecific Subsystem Comparison .. . ....
Chickadee . . .................
Titmouse . ................
Interspecific Comparison of Subsystems . . .
Review of Communication Systems of the Genus Parns.
Interspecific Effects on Vocal Communication Signals.


LITERATURE CITED . . . .

BIOGRAPHICAL SKETCH . . . .


. 61
. 61
. 61
. 62
. 62
. 63
. 63
63
64
. . 64
. 65
. 65
. 65
. 66
. 66
. 67
. 67
. 69
. . 70
. 73
. 83

. 91

. 96


: : : : :













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


A COMPARATIVE ANALYSIS OF THE VOCAL COMMUNICATION SYSTEMS
OF THE CAROLINA CHICKADEE AND THE TUFTED TITMOUSE

By

Philip Gaddis

December, 1979


Chairman: John H. Kaufmann
Major Department: Zoology


Repertoires of the vocal communication signals of two interspecif-

ically social species, Parus carolinensis and P. bicolor, are described

and compared according to acoustic structure and behavioral function.

Calls of both species were placed in four major subsystems on the basis

of similarities in acoustic structure. The subsystems that corresponded

in function between the two species were compared and their similarities

evaluated. The extent of structural similarities in these subsystems

correlated with the extent of interspecific relevance of their communi-

cative functions. Calls used for close-range contact were very similar

between the two species. Calls used for mid-range contact and contact

under changing conditions differed in details of acoustic structure

between the two species but were basically similar, and call subunits

were arranged by the two species according to similar rules of syntax.

I interpret this as a maintenance of sufficient similarity to facilitate

interspecific flocking while allowing sufficient divergence to enable

intraspecific group cohesion within the mixed flock. Calls used in

aggressive encounters, which were primarily intraspecific, were completely











different between the two species, although both subsystems were derived

from the mid-range contact calls of their respective species. Calls

used for long-range contact and territorial advertisement were also

completely different.













INTRODUCTION


The Carolina chickadee (Parus carolinensis) and the tufted

titmouse (P. bicolor) flock together throughout most of the year where

their ranges overlap. Even in the breeding season, when flocking is

less common, the two species nevertheless spend much of their time

together. When the mixed flocks do form, from late summer to early

spring, many other species are associated with the flocks, but the

vocalizations of the two parid species are unmistakably predominant

in the flocks' acoustic output. Some of the vocalizations of the chicka-

dees are humanly indistinguishable from those of the titmice, some are

similar, some bear vague resemblance in certain aspects, and some are

quite different. This study was undertaken to evaluate more precisely

these similarities in vocalization structure, to study relations between

structural and functional similarities, and ultimately to describe the

extent of interspecific language.

Peter Marler (1957) described the extent of similarities in vocal-

izations of several sympatric, British bird species and formulated

expectations for similarities in communication signals. These

expectations were confirmed in studies on two sympatric and inter-

specifically social species of Cercopithicus monkeys (Marler, 1973). My

study of the two sympatric and interspecifically social species of Parus

closely parallels this work on the monkeys and further confirms Marler's

predictions.














BACKGROUND ON THE SPECIES STUDIED


The genus Pcaus is a relatively distinctive group that comprises

44 of the 46 species of the family, Paridae (Snow, 1967). No pre-

pleistocene fossils are known for the genus (Brodkorb, 1978), and thus

very little can be said with certainty regarding its history. However,

this genus reaches its greatest diversity in Eurasia, especially in the

eastern Himalayas and mountains of western China where 14 species occur

(Lack, 1971), and it seems likely that the genus arose there. The

genus can be subdivided into ten subgenera based on the classification

of Hellmayr (1903). Five or more species may coexist in many areas of

Eurasia, but, with only a few exceptions, coexisting species are from

different subgenera (Lack, 1971). The exceptions are P. palustris and

P. montana, which coexist over much of temperate Eurasia including

England and Japan, and P. atricapillus and P. rufe3oens, which coexist in

the Pacific northwest of the United States (Smith, S. M., 1967). Both

species pairs are from the subgenus Poecile, the largest of the sub-

genera. Coexisting forms are usually well separated by habitat, or by

food and microhabitat preferences (Lack, 1971). Only one subgenus

(Melaniparps) occurs in Africa south of the Sahara, and, although ten

species exist, overlap between them is rare (Lack, 1971).

Two subgenera occur in North America. The subgenus Poecile

consists of six North American and six Eurasian species, one of which,

P. cinctus, occurs in Alaska as well as Eurasia from Scandinavia to

Siberia. All of the species of the subgenus Poecile are very similar









in appearance, with only P. speritiouas of the mountains of western

China, and P. jivnbcli of the mountains of western U.S. deviating from

the general pattern by having a white superciliary line. The other

North American species can be divided further into brown-capped and

black-capped groups (Lack, 1971: superspecies). The brown-capped group

consists of P. cinctun, P. hudsonicus, and P. fuf:;c.cn,, the latter two

of which were considered sibling species by Grinnell (1904). The black-

capped group consists of P. sclateZi, P. atricapillus, and P. carolineni.s,

the latter two of which were considered sibling species by Brewer (1963),

and were thought to have diverged during the Illinoian glaciation in the

eastern U.S. Parus carolinensis presently occupies the southeastern

U.S. from the Atlantic ocean to central Missouri, Kansas, Oklahoma,

and Texas and from southern Pennsylvania, Ohio, Indiana, and Illinois

south to the Gulf of Mexico. Paeis carolinensis is subdivided into four

subspecies (Snow, 1967; five subspecies in A.O.U. Checklist, 1957) based

on subtleties of plumage coloration and on measurements of wings and

tails, with the larger subspecies in the north.

The subgenus Baeolophus is considered to be a uniquely North

American group (Lack, 1971; Dixon, 1961) although P. wollweberi was

considered by Hellmayr (1903) to be more closely related to the subgenus

Lophopoanes of Eurasia. Besides P. mollweber Baeolophls consists of

only two other species. Parus inonatus occurs in southwestern U.S.

north to southern Oregon and east to the Rocky Mountains. I, :aw bicnlor

is distributed from southern New England, extreme southern Ontario, and

southeastern Nebraska south to northern Hidalgo and Veracruz, Mexico.

The subspecies of P. bicolor occupying the eastern United States,











P. b. bicolor, is relatively distinct from the other four subspecies,

and was recognized as a species by the A.O.U. until 1976 (A.O.U. Check-

list, Thirty-third Supplement, 1967). Pawis b. hicolor averages around

30% heavier than its neighboring subspecies, P. b. bcnzncti (Dixon, 1961),

and bears distinctive crest markings, but the two subspecies interbreed

where their ranges meet (Dixon, 1955) and so have been considered one

species.

The range of P. carolinensis is virtually contained within the

range of P. b. bicolor. Parus b. bicolor, however, extends into the

range of P. atricapillus several hundred miles along its (P. b. bicolor's)

northern and northwestern boundaries. The ranges of both P. carolinensis

and P. b. bicolor are advancing northward (Brewer, 1963; Pielou, 1957).

In general, members of the genus Parus are present wherever forest

or savanna occurs in Eurasia, Africa, or North America. They have also

taken well to areas of human habitations, and persist in gardens and

hedgerows. They are broadly omnivorous in their food preferences. They

are partial to caterpillars, but also take many other arthropods as well

as acorns and the seeds of many other trees and plants. Their ability

to exploit plant resources no doubt enables them to spend the winter

in cold temperate climates. The genus is basically non-migratory but

some of the far northern species make regular migrations southward in

the winter (Havlin and Jurlov, 1977; Hilden, 1977).

Except during the breeding season, species of the genus Parus

are nearly always associated with mixed species flocks. Parus luiubris

of southeastern Europe and the northern part of the mideast, and P.

inornatus of southwestern U.S., however, show little flocking behavior






5



(Hinde, 1952; Dixon, 1949). Mixed species Flocks appear to be a nearly

universal phenomenon in forest habitats throughout the world. Wherever

parids occur they are nuclear species in the mixed flocks in that the

flocks tend to form around them. Throughout Eurasia and North America,

the most regular flocking associates of the parids are nuthatches,

Sittidae, treecreepers, Certhiidae, and various species of Sylviidae.

In Eurasia, muscicapid flycatchers are regular associates (Hinde, 1952;

Macdonald and Henderson, 1977; Filonov, 1960). In southern and south-

eastern Asia, many species of babblers, Timaliidae, are associated with

the mixed flocks (Ali, 1977). In North America, various species of

the family Parulidae are regular associates (Morse, 1970). The dis-

tribution ranges of the Paridae, Sittidae, Certhiidae, and Sylviidae

in North America are largely coincident.

Throughout the fall, winter and early spring in central Florida and

probably wherever else they co-occur, P. carolinensis and P. bicolor

spend virtually all of their time in each others company. In my study

area, the two species tended to roost colonially in tree holes and also

in piles of Spanish moss, Tillandsia usneoides, in the case of the

titmouse. The birds went to roost together one half to one hour before

sunset and emerged together at 3/4 to one hour after sunrise, staying

in longer in cold weather. The birds often gathered at traditional

centers following morning emergence, as well as at other times during

the day, with a great deal of calling. The mixed flock would then leave

all together or in smaller mixed groups in separate directions.

The most regular attendant species in my study area were the black

and white warbler, Mniotilta varia, yellow-throated warbler, Dendroica










dominica, and brown creeper, Carthia j'filiardi, all of which occurred

at a density of only one individual per flock and were only rarely

seen away from flocks. The solitary vireo, Vireo solitariu:, downy

woodpecker, Picoides pubeocens, red-bellied woodpecker, 'e lancripcs

c,'olinczsis, pine warbler, P:rndrUoica pinus, and orange-crowned warbler,

Verinivora celata, were also regular attendant species Theyoccurred at

one to three individuals per flock and were frequently seen away from

the flocks. The yellow-rumped warbler, Dendr'oi'a cor.naZta, American

goldfinch, Carduelis tristis, and ruby crowned kinglet, Regul zs calZendla,

occurred irregularly with the flocks in groups of four or more to

more than thirty (more than fifty in the case of the yellow-rumped

warbler) per flock. The brown headed nuthatch, Sitta pusilla, did not

occur on my study area probably due to the small size of the pine

"island" but, in more extensive areas of pine in the southeast, it

regularly occurs with the parids in mixed flocks (Morse, 1970;

pers. obs.).

While the parids were feeding on long leaf pine seeds during the

fall and early winter, the flocks moved slowly (< 2 km/hr), were often

widely dispersed, and often became fragmented. When the pine seeds

ran out in early winter, and especially in cold weather, the flocks

were more cohesive and moved faster through the woods (2-5 km/hr).

The flocks were most active during early morning and late afternoon.

When pine seeds were available, and during warm weather when insects

were apparently more available, the flocks' activity declined during

mid-day and the birds would perch quietly together and preen. During

periods of cold weather in mid-winter, the flocks continued to forage

actively throughout the day.











The flocks were exposed to predator pressure at a rate of around

two attacks or flybys per day by the sharp-shinned hawk, Accipiter

striatus, and occasionally the Cooper's hawk, Aci'ipitea coopeirii, from

November to March.

Supplanting attacks and attempts at food piracy were rare when

pine seeds were available but became frequent later in the winter.

These were usually directed against members of the same species but

titmice were occasionally supplanted by red-bellied woodpeckers, chicka-

dees by titmice, warblers by chickadees and titmice, and yellow-throated

warblers by all other species, especially the pine warblers (cf. Ficken

et al., 1968). The vulnerability of chickadees and titmice to piracy

is due to their habit of taking food items that require manipulation by

holding the item with the feet while tearing it up with the bill.

Although attempts at piracy were at times frequent, no successful

attempts were observed. The food item was occasionally dropped in the

process and lost. Birds were occasionally displaced from a cone or

other foraging site after it had yielded a bit of food.

The flocks defended a winter territory against neighboring flocks.

The chickadees and titmice fought only against their own species.

Occasionally only the titmice fought, and, after the two factions

separated, the chickadees continued on with the opposite group from the

one they had originally been traveling with, indicating that the chicka-

dees' flock range spanned more than one titmouse flock range. The

groups of four to six titmice defended territories of thirty to forty

acres. The groups of 5 to 8 chickadees defended territories that were

probably at least twice the size of the titmouse flock territories.


I










The flocks appeared to disregard flock boundaries and to continue

foraging into neighboring territories until discovered and expelled by

the neighboring flock.

Pair formation presumably takes place during the winter. Display

flights (cf. Tooby, 1948; Smith, S. T., 1972) were observed during

January and February. They were seen to be performed on four occasions

by titmice and once by a chickadee in the areas where the flocks

gathered in the morning and at other times during the day (see above).

When the weather warmed up in March, the dominant pair of the

flock carved out a breeding territory from the flock territory and

excluded others of its species. The remaining area of the flock

territory was then taken over by a second pair from the remaining

flock and the others were excluded. The two species nested in tree-

holes and raised three to six fledglings. The fledglings were led around

the breeding territory for around two weeks for the chickadee and

four or five weeks for the titmouse until the parents gradually

stopped feeding the young and the flocks began mixing. Border con-

frontations during this period consisted of fights between parents

only. Strife between fledglings within family groups became more and

more intense as they got older. Following this period of family

group activity, the flocks began mixing and a period of intense and

complicated fighting followed, while winter flocks and flock territories

were formed. This period lasted from mid-summer to early fall.

This general life history pattern is typical of the genus (cf.

Bent, 1946; Hinde, 1952; Lohrl, 1964; etc.).














METHODS


Study Area


The study area consisted of approximately 50 ha in the center of

the 2200 ha San Felasco Hammock State Preserve. The center of the study

area is 20 km NW of the University of Florida campus and at least 3 km

from the nearest public road.

At the center of the study area is an open stand of long-leaf pine

(Pinus palustris) and turkey oak (Quercus laevis). This stand slopes off

gradually on three of four sides into denser second growth loblolly

pine (Pinus taeda) and turkey oak. Much denser second growth woods

consisting of sweetgum (Liquidamber stylaciftla), southern red oak

(Quercus falcata), mockernut hickory (Carya tomentosa), and laurel oak

(Quercus henisphaerica) occur on the fourth side. This second growth

deciduous woods continues to slope off to a small swamp surrounded by

a relatively mature stand of laurel oak, pignut hickory (Carya glabra),

basket oak (Quercus michauxiii), and magnolia (Magnolia grandiflora and

M. virginiana).

Around my home in Gainesville, tall laurel oaks are abundant.

Many live oaks (Quercus virginiana), long leaf, and loblolly pines are

also scattered throughout the neighborhood.


Observational Techniques

Flocks containing Carolina chickadees (Paruos carolincnsis) and/or

tufted titmice (P. bicolor) were followed in San Felasco Hammock State











Preserve from March, 1977, to December, 1978. The most intensive obser-

vations were made between September, 1977,and June, 1978, during which

time the flocks were followed on a daily basis. Beginning in December,

1977,attempts were made to mark birds for individual identification.

Mist nets were used to catch the birds. The chickadees and titmice were

marked with colored plastic leg bands on both legs and with a U.S.

Fish and Wildlife Service aluminum leg band on one leg. The birds were

extremely difficult to catch; it took nearly four months to catch and

mark 12 titmice and six chickadees. Attempts to bait and then trap

the birds were unsuccessful in San Felasco Hammock. An additional

six titmice and two chickadees were trapped and marked after having

been attracted to feeders at my home in a residential neighborhood of

Gainesville, three blocks north of the U.F. campus. These birds were

not given the systematic attention given the birds at San Felasco

Hammock, but many recordings of their vocalizations were made.

After a few weeks of following the flocks regularly, the birds

habituated to my presence and I was able to stay in the midst of the

flock and observe the birds at close range. Recordings of their vocali-

zations were made on a Superscope C105 cassette recorder with a

Realistic electret condenser microphone and also on a Nagra IIIN open

reel recorder with a Gibson P200 microphone in a 46 cm parabolic

reflector. Approximately 40 hours of tape were recorded and then

analyzed on a Kay Electric 7029A Sona-Graph.

Detailed transcripts of the tapes were made and sonograms of

representative examples of each call from each bout of its occurrence

were made. The resulting sonograms were then sorted and arranged into


I









categories of calls and then the calls into major subsystems. The

calls were given names and then an index of each of these calls was

compiled from the tape transcripts. Contextual information, if avail-

able, was included with each index entry. This contextual information

was summarized for each call and then used in the process of inferring

functions and messages of the calls. Tape recordings of all calls have

been deposited in the Florida State Museum Bioacoustics Laboratory.
Definitions


The process of communication is interpreted in many different ways

by its various investigators. It is a complex subject and I find no

set of definitions of all relevant variables entirely satisfactory.

None of the following definitions are entirely my own but no single

work in the literature contains them all.

I use the term function according to Williams (1966) as a conse-

quence of an act or an effect of an organ's activities that enhances the

fitness of an individual organism. An adaptation is the evolutionary

means of accomplishing a function. Information is anything that stimu-

lates a sense organ of any kind (Smith, W. J., 1977). A signal is a

behavioral, physiological, or morphological characteristic adapted for

conveying information (Otte, 1974). A message is an abstract quality

that defines the packaging of the information made available by a

signal (sensu Smith, W. J., 1977). A message is an effect of a signal.

The function of a signal is the manipulation of the behavior of another

organism (sensu Dawkins and Krebs, 1978). Communication is the process

of transmitting messages.

A call is any vocalization. A call type is a class of calls with

certain specified similarities. A form is a specific call configuration.










A note is a sound with virtually no frequency inflection. A syllable

is a continuous sound with one or more frequency inflections. A phrase

is a more or less stereotyped cluster of notes and/or syllables used as

a subunit in longer calls. A series is a string of notes, syllables,

and/or phrases considered to be one call unit. The notes and phrases

of a series are also referred to as call subunits. A bout is an

episode of calling. Two sounds are considered to be harmonically related

if the frequency difference between them is integral to the absolute

frequencies of the two sounds (Roederer, 1975). One sound is a

harmonic of another if its frequency is an integral multiple of that

other sound. Onset and cutoff arms refer to the vertical components

seen in sonograms at the beginning and ending of certain vocalizations.














RESULTS


Chickadee Subsystem I


Structure


The calls of this subsystem consist of four basic types (Fig. 1).

They all sound like a soft, quick "tseet." Types A and B represent

extremes in a continuum in which intermediate forms are expressed

with descending onset portions of variable length. Type B consists

of those forms without descending onset. The portion of the call that

was horizontal or nearly so (as seen in sonograms) varied in length in

those cases where unambiguously measurable from 12-30 msec (x = 20,

SD = 6.12) for type A and 25-45 msec for types B and C (B x = 34.7

msec; SD = 5.3; C x = 32.5 msec, SD = 6). However, most cases were not

safely measurable due to differential reverberation at various call

amplitudes and in acoustic media of variable structure and density (for

a discussion of reverberation, see Wiley and Richards, 1978). Calls of

all four types when given at sufficiently high amplitudes, were followed

by an "echo" at much lower amplitude. This "echo" was invariably

identical to the original call and followed at a latency of approximately

214-218 msec.


Usage and Contexts

Calls of this subsystem were given during virtually all kinds of

activities. The calls appeared to be correlated with short hops or


















I I
8
dI

1I
O


i'k OrJ'0


f

12


4


5'.


SECOrM'




Figure 1. a-e show calls of P. carolinensis subsystem I. a shows
type A'; b, type A; c, type B; d, type C; e, type D.
f-j show calls of P. bicolor subsystem I. f shows type
A'; g, type A; h, type B; i, type C; j, type D.











other foraging, food manipulating, or preening movements. Those calls

for which unambiguous contextual data were available indicated that type

A calls were given during active foraging (23 out of 25 cases) whereas

type B calls were given rmre frequently during blood manipulation and

periods of inactivity (10 out of 18 cases). This difference in usage
2
between types A and B is statistically significant (2 = 11.88; df = I,

p < .001). Those cases in which type B calls were given during active

foraging tended to be later in the day when flock movement rates and

individual foraging rates were more relaxed than earlier in the day.

Type C calls were most frequently given during active foraging

(10 out of 18 cases) but much less conclusively so than type A calls.

Calls of type D were always given during long flights and usually by

the first of a group to fly. Calls that appeared to be intermediate

between types C and D occurred, and those of type C were associated with

a high likelihood of a long flight following their occurrence (.6).

During aggressive confrontations, an extremely shortened form of

type A referred to as A' was frequently given. The contexts in which

these calls were given constitute the highest level of excitement a.id

activity rate that chickadees experienced. The shortness of A' calls

and the prominence of the descending onset fits the pattern of type A

and B in that these variables appeared to be correlated with activity

rate.


Chickadee Subsystem I I

Structure

The calls of this subsystem are made up of two to several subunits

of three basic types which differ primarily in the magnitude of emphasized










frequency and in the extent of harmonic content (Fig. 2). It is for the

calls of this subsystem that the chickadee is named. The subunits of

type A occurred in a graded series varying in duration, length of onset

and cutoff arms, and maximum frequency. These variables were related

in such a way that the high frequency extreme was the longest in duration

and had no onset or cutoff arms. Those subunits at lower frequencies had

correspondingly shorter durations and greater onset and cutoff arms.

The subunits toward the low frequency extreme often carried sound energy

at a second frequency associated with the onset arm but these subunits

did not carry sound energy at more than two frequencies simultaneously.

I recognize six forms that appear to represent modes in a graded series

of type A subunits.

Subunits of type B also occurred in a graded series which varied in

harmonic complexity. The increase in complexity was associated with an

increase in duration. The emphasized frequency remained constant through-

out the series. Those subunits at the simplest extreme of the series

resembled the shortest, lowest frequency subunits of type A; those at

the most complex extreme resembled the subunits of type C but still

carried the peaked ending. Thus, the type B series represented a tran-

sition from type A to type C. I recognize three forms that appear to

represent modes in a graded series of type B subunits.

Subunits of type C were relatively flat stacks (as seen in sono-

grams) of what appeared to be harmonics of a fundamental at approximately

450 Hz (cf. Greenewalt, 1968). These were the "dee" notes of the "chick-

ka-dee" series. Those harmonics between 3 1/2 and 8 kHz were typically

represented. The subunits varied in the flatness of these harmonics

in a very irregular way and I was unable to recognize a pattern in the


























Figure 2. a-k calls of P. carolinensis subsystem Ila. a, c, f, h, and
i are series containing subunits of types A and C. h shows
a call with fused type A and C subunits. i shows a call
introduced by a call from subsystem I type A. b, d, e,
and j show series containing subunits of types A, B, and
C. g shows a series of short type C subunits. This
series was given during an interflock border confrontation
and is also typical of the calls given during mobbing.
I shows a call from titmouse subsystem Ila given in exchange
with a chickadee whose call is shown in k. The type A
subunits given by the chickadee in k are unlike those
given in exchange with other chickadees but are very
similar to those of the titmouse. The type C subunits
of the titmouse call in I are unusually short and are
comparable in length to those of the chickadee in k.









b

,- r, - ^ .s


8]


0

c d


-I






blCONlDS

f 9



4-


0


h







--------~----v
I-

0
SECOND$

j k


8a


SiCONLD)


erir













~rdDNPU8 l


~X ~



---T~-










variability. The subunits also varied in duration and I have recognized

3 forms based on this variable.


Usage

Any of the subunit forms could occur alone and nearly all were

combined with other forms in a remarkable variety of combinations, but

only according to certain rigidly upheld rules of syntax. The highest

frequency form of type A (0 or "high zee" alarm call) if in combination

occurred only with the next highest form (Al). Either of the two forms

could occur first, but frequency transitions within a series were

progressive and no more than one frequency reversal within a series was

ever recorded. Any of the other forms could co-occur in the same

series. Type A forms always preceded forms of types B and C; forms of

type B always preceded those of C. Within type A, the forms with

higher frequency and longer duration always preceded those with lower

frequency and shorter duration. Within type B, forms with simplest

harmonic content preceded the more complex forms. The subunits of type

C could occur in any order. They could grade into each other or occur

in distinct runs of a single form but no more than one trend reversal

or more than three runs of a single form per series were recorded. A

series could begin with any subunit form but it was followed only by

those forms that did not violate the above rules, i.e. by those forms

at equal or lower frequency emphasis and equal or greater harmonic

complexity.

A sample of 402 series for which sonograms were made averaged

8.02 subunits/series (Fig. 3A) (SD = 3.3, mode = 7, range = 1.23).

Of the 295 of these 402 that contained type A subunits, the mean number
























Figure 3. A.


Occurrence frequency of P. carolinensis sub-
system Ila series containing different numbers
of subunits per series x = 8.02, N = 402.


B. Percentage occurrence of P. carolinensis Ila
series with different type compositions. N = 402.

C. Percentage occurrence of P. carolinensis Ila
series with different numbers of subunit forms
per series. N = 402.









60-
A
o -
50

O --
L 40
LL

L 30--
10
z



S0


1 2 3 4 5 6 7 +8 9 10 II 12 13 14 15 16 17 I
X
NO. SUBUNITS /SERIES







100- 100-
S B -


O -
D 60- 60
o 0
o 0
040- -- 40-
2 L
LU
0 20- C 20
x n

0 0 -- -F4-H-:---- [----j
LL


c c m ooco I 3 4 5
< mo < < m < NO. SUBUNIT

SUBUNIT TYPES FORMS/SERIES


B 19 23









of type A subunits/series = 3.2 (SD = 2.5, mode = 1, range = 1-I1). Of

the 131 containing type B subunits, the mean number of subunits/series =

3.6 (SD = 2.4, no strong node, range = 1-12). Of the 296 containing type C

subunits, the mean number of subunits/series = 6 (SD = 3.3, range = 1-16,

no strong mode). An average of 2.2 subunit forms was represented in any

call (Fig. 3C) (SD = .86, range = 1-5). Twenty series contained fused sub-

units of types A and C (Fig. 2h). Two other series contained fused subunits

of types B and C. Only one fused pair occurred per series. The distribution

of subunit type combinations is shown in Fig. 3B. From this figure, it is

clear that the predominant combination is of subunits from types A and C

(42.5 The distribution of the individual subunit forms is shown in Fig.4.


Contexts

The calls of subsystem Ila are collectively the most conspicuous

aspect of the chickadees' behavior, and it is quite appropriate that the

bird should be named after this call. As with the calls of subsystem I,

those of Ila were given during virtually all kinds of behavior. Neverthe-

less, two general features appeared to be consistently associated with

their occurrence. One was an exchange of varying versions by two or more

birds over distances from less than 5 to 40 or more meters, and the

other was a change in the movement pattern of one caller or as many as

all birds in a flock. The calls were given when accelerations (velocity

and/or direction changes) or cohesion changes were made by a flock,

especially when a flock left an area, arrived at another, or was

obstructed in its movement by a vegetational disctontinuity. The calls

were given when an individual approached and joined other birds, or when

an individual left others. When making these movements, the calls were

given on landing, or shortly after landing on a flight. The calls were


























>- I
0

LL 100
0
Lt
2
Li-

LLJ
0


rX 40 --

O
0 20



I 2 3 4 5 6 7 8 9 1(
A B

SUBUNIT FORMS


Figure 4. Occurrence frequency of P. carolininsis Ila subunit
forms in a sample of 402 series.










also given by birds that had been prevented from carrying out an intended

activity such as visiting a drinking hole, pine cone, or feeder by the

presence there of a dominant bird.

Calls of this subsystem were given during fights and these tended

to be series that predominated in one type. Series of type A only were

given in the initial stages of fights and whenever a potentially

dangerous situation was at hand. Series of type B only were given in

later stages of fights and as the birds drew apart and left. Similarly,

type A subunits were prominent in combination series when birds arrived

at an area or a group of other birds, and type B subunits were prominent

in combinations when the birds left an area or group. Series consisting

entirely of type C subunits occurred regularly in fights. Series of short

type C subunits were given during mobbing as well as during fights.

IlaO and IlaAl to IlaO combination series were also heard during

fights, but most frequently these calls were given when the birds were

exposed to a threat of predation.


Chickadee Subsystem llb


Structure, Usage, and Contexts

This call was made up of five to eight very gradually descending

subunits bearing a strong resemblance to Ila-C subunits (Fig. 5A). The

calls were given by females from very early in the breeding season until

the young had hatched. They were given by young chickadees from shortly

before fledging until after dispersal from their natal flocks. When

given by females they were given at very low amplitude at close distances

from their mates who usually responded by feeding them. The calls were























a

12
r 4-


0 5 1~g ~
S r0 D


Figure 5. A. Call from P. carolinensiz subsystem Ilb. This
call was given by an adult female.

8. Call from P. biolor subsystem lib. This call
was given by a fledgling.










also given prior to copulation. When given by juveniles in their first

few weeks following fledging, the calls were much louder than when given

by adult females, and became louder on the approach of a parent bird.

After the period of fledgling dispersal began, the juveniles continued

to give the calls, but they were no longer responded to with feeding.

The calls were usually associated with wing quivering when given by

females as well as juveniles.


Chickadee Subsystem Ilc


Structure

The calls of this subsystem were made up of subunits from subsystem

Ila and were arranged in accordance with the same syntax rules with one

major exception: the series could be repeated in rapid succession.

These calls were also set apart by a high degree of stereotypy. Two

basic types of call occurred (Fig. 6). Type I consisted of a four-subunit

series and two variations that differed from this form in lacking one

of the two internal subunits. Type 2 occurred in only one form but it

was occasionally extended beyond the basic five-subunit form to include

a variable number of introductory or terminal subunits. Only the five-

subunit form was repeated in continuous series. The extended, non-repetitive

form frequently preceded the repetitive form or was mixed with it.


Usage and Contexts

The calls of type 1 occurred in series with one to six units per

series. The four-subunit form did not occur more than three times per

series. The three forms of type I were occasionally mixed in the same
































Figure 6. a, b, and c show calls from P. carolinensis subsystem
IIc type 2. c shows an example of the extended form of
this call. d shows a series of llc type 1 subunit
phrases. All three phrase forms are represented in
this unusual call. e-m show T R calls of P. carolinensis
subsystem Ilia. e shows TlaR; f, TlbR; g, T8R; h, T4aR;
i, T4cR; j, T6R; k and I, T3aR; m, T3bR.









b


N -.


c


\ %


d


e


0


h






41 ri
4I1

' j

0


jr


~ JI-


SECOND



Ref


SECONO)4


jtCCir,[~,
f









I,



41


V


* iF L


r









2


SECOND






S2-



0 i
SKONLDS


U
.


2


V

" 1ol, ,


2

0


Figure 6. continued. n-t show calls of P. carolinensis Ilia
containing the X phrase. n shows XTlbR; o, XT7R; p,
XTIcR; q, XT4bR; r, XR; s, T2aXR; t, T2bXR. u shows
an isolated P. cavolinensio rasp. v shows a rasp from
a P. bicolor juvenile.











series. Type 2 occurred at two per series in the majority (70%) of the

cases.

Type I calls occurred at the end of the interflock or interpair

fights as the birds began to leave the area. They were usually exchanged

by two or more birds and the calls continued to be given as the birds

left. They were usually mixed with series from subsystem IIa A, Ila B,

and calls from subsystem Ill. The calls were given usually during fast

retreat, usually under pursuit.

Type 2 calls were also given toward the ends of fights but were

not necessarily followed by exchanges or immediate retreat. They were

usually well mixed with the variable calls of Ila and especially series

of Ila B.



Chickadee Subsystem Ilia


Structure


Calls of this subsystem were stereotyped combinations of two basic

phrase types that appeared with or without one of several rasp forms

(Fig. 6). These calls had a distinctive fast, liquid, sputtery quality.

Fifteen stereotyped combinations were distinguished.

Phrase type X occurred in one basic form only. It was preceded by

a prefix of variable length when it preceded a phrase of the other type,

type T. It was followed by a suffix when preceded by a T phrase. X very

rarely occurred without T, in which case it carried both prefix and

suffix. All four recordings of X without T were made in Gainesville.

I distinguish 12 variations of the type T phrase. Of these, six

never occurred in combination with X. Of the remaining six forms that










did occur with X, three never occurred without X, and three could occur

with or without X. Of the six T forms that occurred with X, four were

always preceded by X; the other two forms were always followed by X.

Eleven rasp forms are distinguished (this includes two forms used

with calls of Illb, below). Seven forms occurred with one call form only.

Of the remaining four rasp forms, one occurred in six different calls,

in each case following a T phrase that appeared to be closely related to

Tl. One occurred in three combinations that all contained X. Two

other rasp forms occurred with two different T phrases, and in each

case the two forms were closely related. Thus where the same rasp

form occurred in different combinations; the combinations'T phrases

bore some structural similarity. Certain T phrases (Tla and T4a) and

certain XT combinations (XTIb, XTIc, XT4b) could occasionally occur

without the terminal rasp. Very rarely, the rasp occurred alone.

I was able to distinguish the three major categories of these

calls (TR, TXR, and XTR) in the field. The specific forms within the

major categories, however, could be safely identified only after sono-

graphic analysis.


Usage and Contexts

Calls of this subsystem occurred during interflock and inter-

individual fights and during territorial boundary advertisement and

exchange. In 89 observed fights, at least one call from Ilia was given

in each case. A mean of 3.6 calls of all kinds were given in these

fights (SD = 1.54, range 1-9). Seventy-one percent of these fights

contained only one representation from Ilia, 23% contained 2, and 6%

contained 3. Most fights contained calls from only one major category










(TR, TXR, XTR) and this category was usually TR. Combinations of these

categories were relatively unusual. Nevertheless, calls ot one category

were more likely to be mixed with calls of another category than with

calls of their own (10% of fights had two different calls from the same

category, 19% had calls from two or three categories).

Of 35 pairwise combinations of specific call forms, 23 (66%)

occurred only once and another 10 (29%) occurred only twice. The

remaining two combinations occurred three and five times, respectively.

The calls occurred during all phases of fights and tended to be

most prominent in the initial stages. The likelihood of attack follow-

ing calls of any major category (TR, XTR, and TXR) was not significantly

different from any of the others and ranged from .25-.275. When T and

XT were considered separately, however, XT did not prove to be different

from the others, but T alone showed a .625 likelihood of being followed

by attack. No significant difference was found in the occurrence of

these categories in interflock vs. interpair fights (X2).

Calls containing X were prominent during territorial border

patrolling and/or displacement of one bird by another at a clumped

resource (i.e., drinking hole, pine cone, feeder, etc.).

I could determine no consistent pattern from the contextual

data on any specific call from subsystem Ilia. Due to the tendency

of the chickadees to fight in dense vegetation, I was usually unable

to determine the identity of the callers, and it may be that some of the

variability in these calls can be explained by individual specificity.

However, most of the calls occurred without variaion at widely separated

localities and many over distances of 20 or more km. I therefore consider

it unlikely that individual variation can account for very much of the

observed variation in the calls of this suhsystem.










Chickadee Subsystem I lb


Structure


This subsystem contained two calls that were both structurally and

functionally distinct from the calls of Ilia although clearly related in

basic features of structure (Fig. 7). Type Illb-l was unique in sub-

system III in that it appeared to contain two T phrases preceding the

rasp. The two T phrases were structurally different from each other but

both appeared to be related to T4. Type Illb-2 appeared to be made up

of the final two notes of T4 followed by a unique, high frequency rasp.

The rasp was often followed immediately by the T4 segment. It was

occasionally followed by T3aR, and occasionally by only the rasp of

T3aR. The call was always preceded and occasionally followed by long

series of high frequency (8-10 kHz) liquid, chattery notes (HSC).

Long series of these notes, interspersed with Illb-2 could continue

for several seconds. These high frequency notes were extremely

irregular in form and often carried sound energy at two harmonically

unrelated frequencies, giving them a distinctive, squeaky quality.


Usage and Contexts

Type II b-l occurred during interflock fights and was always

followed by a departure of one of the flocks. In many cases, it was

given by onlookers rather than those directly involved in fighting.

Type Illb-2 was always given at very close range to another

chickadee of the same flock or to a mate (by female). It was never

followed by attack by either bird. It occasionally occurred at the

beginning of interflock or interpair fights. On three occasions, a

chickadee approached me very closely and gave this call.
































Figure 7. Calls from P. carolinensis subsystem Illb. a shows
type 1; b and c, type 2. c shows both rasp forms
associated with this call. Both b and c show HSC
introductory notes. d and e show long series of these
HSC notes. e shows an unusual series of very irregular
notes in series with the more typical HSC notes. These
may be homologous with the "broken see" series recorded
by Smith (1972) in Pennsylvania.









a








2
C, S



b

12






2
0 5


c




":; ~^ ?'Talwto

j


SIION[)S












5
SECONDS


- -


St CO'NDS


(I


SECOND II4'


,
Ili,


--N

I U


/


*^- ..-r. -r-


III










Chickadee Subsystem IV


Structure


Calls of subsystem IV were made up of two and occasionally three

repetitions of one of two basic phrases (Fig. 8). The phrases could

be slightly modified within one call. They could be transcribed as

"feebee-feebee."

Type I consisted of two and rarely three repetitions of a phrase

with two long notes and a short note at an intermediate frequency. The

phrase could be repeated without variation or with the first note of the

second phrase at the frequency of the short intermediate note. The

terminal note was frequently deleted. This variation did not occur in

a three-phrase form.

Type 2 consisted of a two note phrase whose first note had a

long descending cutoff arm. This phrase was repeated at a lower frequency.

A second variation of this call occurred with the first note of the

phrase reduced to only its descending cutoff arm. The phrase was

repeated a second time at a lower frequency, and the phrase could be

repeated a third time at still a lower frequency than the second.


Usage and Contexts

Calls of this subsystem were most frequently given during patrolling

of territory boundaries by males. The calls were given at regular

"song posts" in series of up to more than 60 calls that could last

many minutes. Three territorial males whose repertoires were thoroughly

surveyed expressed both types and both two-phrase variations of types 1

and 2. The singing bird typically sang one type many times, switched

































Figure 8. Calls of P. carolinensis subsystem IV.
a, b, and c are type 1 calls. d and e show calls
of type 2. f and g show irregular versions with
rearranged subunits.






















'Ii IlLr


SECONDS





4








SiCONDS

d e
8



S2-



1-1
0I
SECONDS


9


f


4

22




C0

SECONDS


2










to another type and repeated it many times. It then switched back to the

first type, or moved on to another "song post." If answered, the

chickadee exchanged for only a few times before approaching the source of

the answer with a variable medley of calls from all three other major

subsystems. Calls of subsystem Illa were also included in territory

border calling series in the absence of exchange. These usually

contained the X phrase.

The calls were also given from the nest and at any other place in

the chickadees' terriLory while the bird foraged. The calls from the nest

were usually of type 2, and were often incomplete and occasionally mixed

with call subunits from subsystem Ila A. The calls occasionally contained

rearranged phrases and occasionally were totally garbled and unrecog-

nizable. The calls were also given by chickadees that were presumed

to be wandering, landless birds that were invading new territory. The

calls were also given during the non-breeding season, especially follow-

ing interflock fights.


Titmouse Subsystem I


Structure


As with chickadee subsystem I, titmouse subsystem I consisted of

four basic types (Fig. 1). Calls of type A were variable in the length

of the descending onset and appear to grade into the calls of type B.

The portion of the call that was flat or nearly so varied in length in

type A from 4 to 38 msec (x = 20.4, SD = 7.9 msec, N = 30), in type B

from 17 to 58 msec (x = 37.1 msec, SD = 16.3 msec, N = 10), and in type

C from 2.5 to 25msec (x = 15.9 msec, SD = 5.5 msec, N = 6). Calls of all










four types were occasionally followed by an "echo" at much lower

amplitude and at a latency of approximately 214-218 msec.


Usage and Contexts


The titmice tended to give type A calls while actively foraging

(35 of 43 cases) and type B when foraging more slowly, while manipulating

food or when otherwise inactive (11 of 20 cases). The distinction is

less conclusive for type B, but the difference in usage between types A

and B is statistically significant (X = 9.06, p < .01). A very short

version of type A, called type A', was given during fights. The cases

in which type B calls were given during active foraging tended to occur

later in the day or when flock movement was slower. Type C was given

rarely by titmice. In three of the six cases, it was given just before

flying. In the remaining three cases, the birds were foraging actively.

Type D was given during long flights by the first of a group to cross

an open space. It was occasionally given by successive birds but not

by the last. On one occasion, the call was chorused by four perched

and inactive titmice just before they left the area in long flights.



Titmouse Subsystem Ila


Structure


Calls of this subsystem were made up of subunits from two basic

types. Those of the first, type A, were clear, whistle-like tones with

sound energy at only one frequency at a time. Those of the second,

type C, were broad frequency-spectrum harmonic series with a harsh

scolding quality (Fig. 9). These calls sounded somewhat like "seejert,"

"seejawer," etc.

































Figure 9. Calls of P. bicolor subsystem IIa. a shows
AI-Cl; b, A3-C2; c, A4-C2; d, fused A4-C1 with
low frequency emphasis; e, A5-CI; f, A6-C2.










a

12
S....










121

i-




21
'I_ _







8j^ *


52


0




12J












21
4
u





^:


,lI tO ',


AICON Ds


I~1~


SECONDS

e







-r N

SiCON [I

f


2ONI
st ONI S


t-
i;~a~o!
































Figure 9. continued. g shows A7-C2; h, A8-C3; i, A9-C3;
j, A9-C3; k, C3; 1, C4; m, C5.












12








8




02
StCONUS


h




>7U %'\


I2 I 1V




0 5
SECOND)


k
12




2-
O

5 -_____________


fl


SECONDS




,$..U i,: FP
(af *,,

,i


SECONDS


- ------~-
I


i L'1









Type A subunits were variable in frequency and ranged from seven to

near 10 kHz. Those over 9 kHz were constant in frequency (i.e., flat)

and were rarely followed by notes of type C. Those between eight and

nine kHz could be constant in frequency or carried a slightly descending

onset. They also carried a cutoff arm of variable length. In those

subunits that were followed by others of type A, the cutoff arm, if

present,was very short. In those that were followed by a subunit from

type C, the cutoff arm was always present and covered a frequency range

of five + kHz. The cutoff arm could be sharp (Al), broadly rounded

(A3-6), or more highly modified (A7-9).

Type C subunits were harmonic series of a fundamental at about

500 Hz. Cl was the most commonly occurring form. It covered a frequency

range from 3 to 9 kHz. It could vary in the emphasized frequencies: a

high frequency emphasis (> 4 1/2 kHz) and a low frequency emphasis

(< 4 1/2 kHz) form occurred. C2 carried only two harmonics with a faint

third occasionally present. The harmonics present in this form were

between 4 and 6 kHz. Other forms could show frequency inflection

patterns: C3 rose evenly in frequency, C4 had a strongly peaked ending,

C5 descended in frequency.

Subunits of both types could occur alone but combinations of the

two types were more frequently expressed. Whereas the simplest forms

of type A, Al through A6, were most likely to occur alone, it was the

more complex forms of type C, C3 through C5, that were most frequently

given without introduction from type A. When subunits from the two

types were combined, the transition was always from type A to type C.

In a sample of 268 calls, the overall number of subunits/call averaged










3.9 (SD = 1.77, range = 1-14, mode = 3 [36%]) (Fig. 10). Type A

subunits, when present, averaged two subunits/call (SD = .58, range =

1-4). Type C subunits, when present, averaged 2.4 subunits/call (mode =

1 [46%], range = 1-14). The most frequently occurring combination was

two subunits of type A and one of type C. The subunit transition

involved was usually A3 to Cl. An Introductory subunit of type A was

occasionally fused to the first subunit of type C (Fig. 9d). These

combinations involved A3 and a Cl form with low frequency emphasis.


Usage and Contexts

Calls of this subsystem were extremely common and were the most

conspicuous aspect of the titmouse's vocal communication system. The

calls were given in a great variety of circumstances, but, in general,

they appeared to be correlated with changes in flock movement patterns

and with essentially hostile encounters (fights, mobbings, etc.).

Al and A2 with or without C3 accompanied major changes in flock

movement patterns such as when a flock left an area or arrived at a new

one. The calls were given from a perch or on landing from a flight,

especially when others were following. The calls were given when the

previous path of movement was obstructed by vegetational discontinuities

or by the presence of danger. In the latter case, high frequency type A

calls (9 to 10+ kHz) were given. When this was caused by a human

observer, A3-Cl calls were mixed with the high frequency calls. When

the flock approached a possible border confrontation with another flock,

Al and AS through AS in association with Cl and C2 were given. C3

through CS also occurred in this context but usually without introduction.

C3 was the most common form in these encounters and, when introduced, it

























>-
00
Z 100-

O
J 80
LL-

Lii 60


J 40-


20
0
Q: ---



I 2 3 4 5 6 7 8 9 JO1 11 12 13



NO. SUBUNITS/ SERIES




Figure 10. Occurrence frequency of P. hicoZor Ila
series with different numbers of subunits
per series. x = 3.9; N = 268.










was usually by Al. During mobbing, C3 was the predominant call and

occurred in series of 7-11 shortened subunits. C4 occurred at flock

borders when no opponent was apparent and also following border fights

after the opponent had withdrawn. C5 was a rare call that occurred

following border exchanges that did not escalate to actual combat.

The call was occasionally chorused by several of the birds.

A5 and A6 were frequently used in interflock and interpair border

exchanges. During fights, they were given by onlookers. A7 through A9

were given during and following actual combat. Al was also given during

close confrontation, and the vertical portion was occasionally given

alone during intense fighting.

The high frequency forms of type 1 (9 to 10+ kHz) were given

during close chases, probably by the fleeing bird. The most frequent

context of these forms, however, was during alarm responses, especially

when an accipiter appeared. The very high frequency forms (10+ kHz)

were then repeated in long series as all birds stopped activity.


Titmouse Subsystem Ilb


Structure, Usage, and Contexts

This call consisted of one descending harmonically simple subunit

followed by one or two ascending harmonically rich subunits (Fig. 5b).

The introductory subunit had one or two harmonics, one below and one

above the predominant frequency. The terminal subunits were much shorter.

The two variations (two- and three-subunit series) were alternated

irregularly. They were given by females from early in the breeding

season until after the eggs had hatched, and by young titmice from

shortly before fledging to the period of dispersal. The calls were










accompanied by wing quivering and responded to by mates or parents with

feeding. When given by the female, it was at a low amplitude and close

to her mate. When given by the juveniles, it was much louder and became

still more so on the approach of a parent. When being fed, the juveniles

gave many more of the terminal subunits, which were also extended in

duration. Parent titmice gradually stopped responding with feeding

after four weeks from fledging, at which time juveniles were beginning

to disperse.


Titmouse Subsystem Ill


Structure

This subsystemwas made up of six basic call forms (Fig. 11), all

given at high frequencies (7 1/2 to 9 kHz). All forms had a strong

dip in frequency, and all except III-1 had a sharp vertical component

at the onset. This latter feature gave the calls a distinctive quality.

These calls could be transcribed as "kwee' or "kchee." Several

distinctive variations of 111-6 occurred in isolated cases and could

have been individual variations.


Usage and Contexts

The calls all occurred during fights and hostile border exchanges

at close range. They were given following exchanges of other calls

from subsystem IIa as the birds prepared for attack, and during attack

itself; shortly thereafter, the calls were given by attacking and

victorious birds. The calls could occur in series of one to four.

More than one form often occurred in the same bout of calling. Forms




















b C









S ICO N[)S
e



Vl


'ECON S'


Figure 11. Calls of P. bicolor subsystem III. a shows III-1;
b, 111-2; c, 111-4; d, 111-6; e, 111-5; (d and e
were given by the same bird in the relationship
shown) f, I11-8.


a

B

24


2
C)










111-4, 5, 6, and 8 were followed immediately by short series of IVb-I or

6 in 68% of their (calls of subsystem I I) occurrence. The IVb calls

were always given at a much lower amplitude than the Ill call, but

appeared to be given by the same bird. When combined with the IVb

calls, they were given at greater distances from opponent birds than

when uncombined, and were associated with much lower likelihood of

being followed immediately by attack. The likelihood of attack follow-

ing a call of Ill was .67. When the calls were combined with calls

of IVb, the likelihood of immediate attack was .06. All forms of III

were given in interflock, intraflock, and interpair fights.

The simpler forms of ll, namely 111-1, 2, 4, and 9 were

occasionally followed by and combined with type C calls of Ila. This

interchangeability suggests a probable derivation of III from laA.


Titmouse Subsystem IVa


Structure

Calls of this subsystem were made up of repetitive series of

phrases from three basic types (Fig. 12). All forms sounded somewhat

like "peter" or "peto." The phrase types varied in duration, but overall

series length for all types was a relatively constant .9 sec (SD = .19,

N = 106). Thus phrase length correlated well with number of phrases/
2
series (Fig. 13) (r = .912). I have subdivided these phrases into

three types based on phrase length and number of phrases/series:

IVaS 1-3/series, 200-325 msec/phrase

IVaM 3-6/series, 100-200 msec/phrase

IVaF 5-10/series, 80-100 msec/phrase
































Figure 12. Calls from P. bicolor subsystem IVa and IVb.
a-h show calls from IVaS. a shows IVaS-1; b,
IVaS-2; c, IVaS-3; d, e, and f show IVaS-5 as
given by three different males, all of which
had common territory borders. g shows IVaS-4;
h, IVaS-8. i-l shows calls from IVaM. i shows
IVaM-I; j, IVaM-4; k, IVaM-5; 1, IVaM-7.










a b c



4 -


|d


41















2-









8


SI






SECONrLS






2 2
k








0I

SECCNr[CS


ms%,~~ ~*\~


I------
































Figure 12. continued. m shows IVaM-6; n, IVaF.
o shows a series of the female's version of
IVa. r shows IVaM-8. p, q, and s-w show calls
of IVb. p shows IVb-6; q, IVb-4; s, IVb-I as
given in the San Felasco population; t, IVb-I as
given in the Gainesville (home) population; u,
IVb-3; v, IVb-2; w, IVb-5.









m










4
a2


8
u








0
4











8








0


NIV4 %N*-*"


"H Or[W,


-


, CON S,





ItoNOS
5c Ot's


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8-








0 2














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$it OrDS





























) 300- *S3

E S4
250- *55
I
OS8
09 200 M7
Z **M4
LJ MIO
_1 150
*LJ OM6
cn 100 -
Q
I 50

110

0 2 4 6 8

NO. PHRASES/SERIES


Figure 13. Phrase length of P. bicoZor IVa calls as a function
of number of phrases/series. Only the forms with a
sample size greater than five and given by the four
most thoroughly surveyed titmice of the San Felasco
population were included in this graph. The
correlation coefficient (r2) of a linear regression
analysis of these points is .912.










The first phrase of a series was usually shorter than the others. This

tendency was most pronounced in the IVaM calls. A very irregular form

of IVa was given by females during the breeding season. Individual

variations occurred in the IVaS calls. For example, three neighboring

males gave and often exchanged slightly different versions of IVaS-5

(Fig. 12, d-f). IVaM and IVaF calls were matched by exchanging birds

and were virtually indistinguishable.


Usage and Contexts

Calls of this subsystem were given by males primarily during

territory boundary calling and exchanging. During the winter, the calls

were given in border exchanges by the dominant bird in the flock. They

were also used by flockmates that had become separated from the flock.

IVaM-6 was usually used in this context. IVaM-7 was used by adults in

calling to their young when leading them to a new foraging area.

During the breeding season, territorial males gave their calls

from prominent and consistent song posts along territory borders. The

males usually began calling with IVaM. They frequently continued with

the same call for 100 or more repetitions before switching to a new

form or leaving the area. Switches in call form were usually to a

different call type (i.e., from M to S or F). If these calls were

answered, the two birds exchanged the same call for 10-20 repetitions

before switching to a different type and approaching each other more

closely. Among four adjacent territorial males that were thoroughly

surveyed, 12 call forms were used with a mean of eight/bird. Five

forms were used by all birds, and four were used by one bird only.

Eighteen forms in all were recorded from the San Felasco Hammock










Preserve in general, and still others were recorded elsewhere. Each

male tended to have preferred forms from IVaS and M call types, but

call forms were always matched in exchanges. The IVaF call type was

given predominantly during escalated exchanges after the birds had

exchanged IVaM and approached each other more closely. IVaS calls were

most frequently given after the birds had appeared to lose interest in

the exchange and response latency had increased. The IVaS calls were

also given following intraflock squabbles. The female version of IVaM

was given as the female left the nest and established contact with the

male. The call was given until an answer was received from her mate in

a typical IVaM. She then exchanged with her mate using a combination

call from Ila as she approached further. The female version was also

given as she approached her mate more closely if he was engaged in a

border exchange.


Titmouse Subsystem IVb


Structure

Calls of this subsystem were clearly derived from calls of IVa.

IVb calls resembled those of IVa both in phrase structure and, except

for IVb-2, in their repetitive nature (Fig. 12). IVb-1 and 4 resembled

IVaM forms in structure but both were given at a lower number of

repetitions/call and both tended to vary in structure within a given

series. IVb-l occurred in a strongly modified form by the birds at my

home (Fig. 12c). IVb-5 appeared to contain two IVb-4 subunits with a

third introductory subunit. The call was always repeated without

variation in this form. IVb-3 appeared to be made up of one and a










half extended IVb-4 subunits. It occurred as two, two and a half, or

three and a half of these subunits. IVb-2 bore no strong resemblance

to any other subunit form. It had the general character of a IVaS

form when heard. It was never repeated in a continuous series.

All of the calls of IVb could easily be distinguished in the

field.


Usage and Contexts


IVb-1 was most frequently given in combination with a call from

III during fights between flocks or between individual birds. A

typical sequence sounded like "kwee-kwee-wadee-wadee." The call was

also given by itself usually in series of two or three and as many

as six but in the same circumstances and usually in the same bout of

calling as the combined form. They were given during all phases of

the fights.

IVb-2 was always given in association with III calls but not

directly combined as were the IVb-l calls. The call was rarely used,

and no pattern was evident in its usage.

IVb-3 was given following long interflock fights, following

interruptions in escalated exchanges between individuals, or temporary

separation of opponents. It is perhaps best described as a drawn out

"weee-ba-weee...." The call was used by one male in response to IVaM

playback after he had approached with answers of the same IVaM call.

He then gave IVb-3 as he excitedly circled the area apparently search-

ing for the missing opponent. In other cases, it was given by the

dominant bird in flock fights.










IVb-4 was given during interflock and interpair fights by sub-

ordinate birds and females that were onlookers to fights. This call

was usually associated with calls from subsystem III. In the population

at my home, a call very similar to this was given as a IVaM form during

territory advertisement (Fig. 12r).

IVb-5 was also given during fights. To my ear, it sounded like

"pyuweetaweet." The call was usually given by subordinates that were

isolated from their flockmates. Other flockmates responded by approach-

ing the calling bird. On one occasion, a titmouse that was foraging at

the periphery of the flock discovered another small group of trespass-

ing titmice. It retreated toward its flock, gave IVb-5 calls, and was

joined immediately by its flockmatesand together they attacked the

intruders.



Titmouse Rasp


Structure, Usage, and Contexts

This section contains only one call that strongly resembles the

rasp of the chickadee (Fig. 6v). It was one of the first calls given

by developing fledglings and was given in association with overt

aggression. The rate of occurrence of this call increased as the

fledglings got older and reached its highest occurrence rate during

the period of fledgling dispersal. They occasionally gave it during

the fall and it became rare during winter. No adult was observed

giving this call. It was occasionally associated with IVb-I and 6 as

were the calls of Ill.














DISCUSSION


Functional Interpretations


Chickadee Subsystem I

In general, it appeared that the information made available by

the calls of subsystem I dealt with the close range maintenance of

contact during steady-state flock movements. In addition to indicating

position, the type A calls indicated a high activity rate, type B calls

indicated a low activity rate, those of type C indicated a disposition

to leave, and those of type D indicated leaving. These and all sub-

sequent functional interpretations are summarized in Table 1.


Chickadee Subsystem Ila

In general, the calls of subsystem Ila carried information dealing

with movement changes and establishment and maintenance of mid-range

contact under rapidly changing circumstances. IlaO carried the message

of alarm or of a strong disposition to stop and escape danger. This

indication of escape tendency was probably the relevant message when

the call was used during fights. Al appeared to be a milder form of

alarm indicating surprise or suspicion of danger, and it appeared to

be used as a general alert. The other type A forms, when uncombined

with other forms, appeared to indicate the presence of danger. The

higher frequency, slower forms indicated greater escape tendency, and

the lower frequency, faster forms indicated greater aggressiveness

61



















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and perhaps "indignation" toward the cause of alert. Type A subunits in

combination indicated that the caller was arriving in an area or a

group.

Type B subunits, used alone and in combination, appeared to

indicate that the caller was leaving an area or group.

Type C subunits alone, as used in scolding and mobbing responses,

appeared to be requests or perhaps demands for increased distance between

the caller and the target animal. (This message can be paraphrased as

"go away."). To a predator stalking the birds, the call could indicate

that the predator's presence was known and that it would be useless for

it to remain in the area. To other flockmates, the call could specify

the location of the predator and call for their help in mobbing it.

To young and inexperienced birds, mobbing could have educational value,

and, in this context, the mobbing calls could carry the message "this

is your enemy."


Chickadee Subsystem Ilb

This call was very distinctive structurally, highly stereotyped,

and unambiguous in its meaning. It was clearly a demand for food. Only

one variation in this pattern was observed: its use for copulation

solicitation.


Chickadee Subsystem lie

Both types of llc calls appeared to function in synchronizing

retreat from some sort of hostile confrontation. This interpretation

is well supported in the case of type 1 but tentative for type 2. Type

2 appeared to indicate more of a tendency to retreat than an urgent need.










Chickadee Subsystem llla

The calls of Illa in general appeared to indicate an aggressive

disposition. They appeared to indicate not only a tendency to attack as

in T and TR, but also readiness to defend as in those containing X. In

TX combinations, the T could indicate readiness to fight and the X could

indicate that it is priority of access to a specific location that is to

be fought for. This location could be a territory boundary, a roost

hole, a pine cone, or perhaps even status position.


Chickadee Subsystem Illb

The two calls of this subsystem had very little in common apart

from their probable common derivation from calls of Ila. In keeping

with their structural distinctiveness, their messages appeared to be

well defined: Illb-I proposed flock retreat; Illb-2 indicated a lack

of hostile intentions. In the context of an impending interflock or

interpair fight, lllb-2 may say in effect "don't attack me, I'm on

your side." (The message of Illb-1 is not completely distinguished

from Ilc-2. It appeared, however, that llc-2 had the function of

synchronizing rapid flock retreat.)


Chickadee Subsystem IV

Calls of this subsystem appeared to express a claim to territory

ownership when given by resident males. When given by wandering or

invading birds, the calls probably conveyed as much of an inquiry into

tenancy as a claim to ownership. I was unable to distinguish specific

messages for the different call forms.










Titmouse Subsystem I


In general, it appeared that the calls of this subsystem dealt with

close range maintenance of contact and steady-state flock movement.

Calls of types A and B indicated position and activity rate: type A

indicated high activity, type B indicated lower activity. Type C

also indicated high activity rate but also probably indicated a dis-

position to leave in long flight. Type D indicated that a long flight

was in progress.


Titmouse Subsystem Ila

The calls of subsystem Ila appeared to deal with the establishment

and maintenance of mid-range contact during activities in which the

problems of behavioral synchronization were much more complex than in

those activities controlled by calls of subsystem I, namely during

changes in flock movement patterns, during fights, or over greater

distances.

The very high frequency forms indicated alarm and escape tendencies,

and the incorporation of the higher frequency forms of type 1 into

combination calls probably indicated the extent to which the caller

perceived that a dangerous situation was at hand. When type 2 calls

were used alone in mobbing and aggressive scolding during fights, they

probably expressed a demand for increased inter-bird distance along with

a threat of aggressive consequences for noncompliance. Incidental

messages in mobbing, as with chickadees, could also be a request for

participation in the mobbing to other flockmates, indication of identity

of enemies to juveniles, and indication to the predator that its presence










is well known to the flock. When type 2 calls were mixed with type 1

calls, the type 2 calls could have somehow specified the desired inter-

bird distance for use in regulation of flock cohesion. These specifi-

cations of inter-bird distance could have been the primary messages

when used in border exchanges as well as in exchanges by temporarily

separated flockmates.


Titmouse Subsystem llb


As with the chickadee, this call was structurally distinct, highly

stereotyped, and unambiguous in its meaning. It was clearly a demand

for food. No copulations were observed, so the possible use of ilb in

sexual solicitation is unknown.


Titmouse Subsystem III

When used without the calls from IVb, the calls of III clearly

carry a threat of impending attack. The combination of III with IVb

(calls derived from distance decreasers) at greater distances could

indicate the desire to come closer to the prospective opponent. Perhaps

this was a dare to the other bird to come close and risk attack. The

high frequency nature of the calls could indicate a high degree of

escape tendency in the caller (cf. Morton, 1977). This could also be

interpreted as indicating that a very dangerous situation was at hand.


Titmouse Subsystem IVa

In general, the calls of IVa functioned in establishing contact

over long distances. The ultimate objective of this association varied

with context and call form. In boundary calling and exchanging, claim










to territory ownership was clearly indicated, and association with

neighbors was sought for the purpose of confirming those claims and

establishing borders. Threat of aggressive retaliation for noncompliance

with those claims was also explicit, especially in IVaF. In an intra-

flock context IVaM-6 was used for re-establishing contact with separated

birds for the purpose of reuniting the flock. IVaM-7 clearly called for

decreasing inter-bird distance when used by parents in leading their

fledglings. Dispersing young could use these calls for inquiring into

tenancy of unfamiliar areas.


Titmouse Subsystem IVb

In general, the calls of this subsystem appeared to be specialized

applications of the general function of coordinating approach (except

IVb-4). IVb-I appeared to call for closer approach during escalated

fights for the purpose of combat. This was especially apparent when

combined with III. IVb-3 called for reestablishing contact after the

disappearance of an opponent or other interruption. Data on IVb-4

was inconclusive but its use by subordinates and onlookers while others

were escalating exchanges or actually fighting suggests that it may

have been used to reaffirm support in battle. IVb-5 clearly called for

flockmates to come and help fight. IVb-6 appeared to be similar in

meaning to IVb-1.


Titmouse Rasp

The titmouse rasp was apparently a juvenile version of the calls of

Ill, although the juveniles did use calls of Ill in addition to the rasp.

Its association with IVb-I indicates an interchangeability with calls of

Ill. Its message was clearly a threat of aggression.









The structural similarity of this call to the rasp of the chickadee,

along with its usage in similar contexts, suggests a possible homology.

I suspect that the titmouse rasp may be a nearly vestigal remnant of a

homologue to the chickadee rasp. This situation is analogous to that

found by Dilger (1960) working with lovebirds of the genus Anaornis,

by Moynihan (1962) with gulls and terns, and by Smith (1977) with

tyrranid flycatchers, in which common displays of some species were

rarely expressed in other closely related species.


Intraspecific Subsystem Coiparison


Chickadee

Subsystem I was distinct from other chickadee subsystems in call

structure and usage patterns. The calls were given during virtually

all kinds of behavior and were thus heard in association with calls from

other subsystems. In certain cases, the association appeared to be

more than coincidental. In these cases, calls of subsystem IIc and

Illa ( lc-l, T8R, T2aXR, T3aR) were prefixed with short call notes

that could have been calls of subsystem I. These combinations were

repeated without variation in the same bout of calling or, as with

lc-I from one bout to the next. In all of these combinations, the

call from subsystem 1 that was involved was type A, the type that most

clearly correlated with the excited activity that characterized the

contexts of these calls. The closeness and regularity of the calls in

these combinations could represent compound messages, but the specific

nature of these messages was unclear.










Subsystems II and III were made up of relatively distinctive sub-

units, but the patterns of subunit arrangement bore strong resemblance

to each other. Calls of both subsystems were made up of series of

several subunits that began with high frequency, harmonically simpler

subunits. Except for calls in which X precedes T, successive subunits

were at the same or lower emphasized frequency and increased in frequency

range and/or harmonic complexity. The calls ended in subunits with

very broad frequency range. T1 and T8 of subsystem III were particularly

reminiscent of subsystem II. TlbR was prefixed by a subunit that strongly

resembled subunit A3 from subsystem Ila.

The calls of subsystem Ila and Ilc differed primarily in the extent

of their variability. Whereas the calls of Ila were extremely variable,

those of ilc, especially Ilc-I showed little variability. The calls of

Ilc were further distinguished by their repetitive nature, a character-

istic that was not observed in lla but was in Ilia. The calls of Ilc

thus appeared to represent a transitional stage between Ila and III in

that their subunits were structurally similar to lla but were repeated

in stereotyped phrases as in subsystem I1l. It is conceivable that calls

of subsystem III were derived from Ila by the development of stereo-

typy in certain combinations and their further modification as they

became associated with a more specialized message and function. The

general function of establishing and/or maintaining contact during

rapidly changing conditions that characterized calls of subsystem IIa

was extended in the calls of subsystem Ilia to facilitating approach

during aggressive encounters or for expressing the intent or willing-

ness to approach aggressively.










Calls of subsystem IV were distinct from the other subsystems both

in structure and syntax. When calls of subsystem IV were given near the

nest, they were occasionally introduced by type A subunits from subsystem

Ila. In their function as long distance contact calls, IV calls shared

with calls of subsystem II the general function of contact, but at a

greater distance. A combination of the two subsystems in this context

was not repeated and may represent a momentary confusion of purposes--the

bird was highly motivated to assert territory ownership but at the same

time needed to establish contact with his mate and indicate his arrival

at the nest.


Titmouse


Calls of subsystem I are distinct from the calls of all other

subsystems. A call of type A occasionally occurred in introduction to

an otherwise unintroduced series of type 2 subunits of subsystem II.

Calls of subsystem III were distinguished from the type A calls

of subsystem I 1 by their possession of frequency inflection patterns.

However, they shared with type A calls of subsystem Ila a common

frequency range and harmonic simplicity. They were occasionally intro-

duced by type A subunits of subsystem Ila, and, in other cases, they

were followed by Subsystem Ila type C subunits in close association.

The relatedness of the two subsystems was further indicated by the

occurrence of intergradations between them. Calls of subsystem III

thus appeared to be derived from type A calls of subsystem Ila. In

terms of information, the two subsystems had in common the message of

establishing and/or maintaining contact under rapidly changing conditions.

Calls of subsystem III represented application of this message to the


I











specific function of indicating intent or willingness to approach

aggressively.

Calls of subsystem IV were distinct from those of other subsystems.

Certain calls of IVb regularly occurred in combination with those of III

and probably represented compound messages rather than relatedness.



Interspecific Comparison of Subsystems

The calls of subsystem I of the chickadee were very similar to

those of the titmouse subsystem I in acoustic structure and usage patterns.

Type A calls were nearly identical in duration (Ax=.4 msec; t = .16;

df = 42; p > .5). In terms of the caller's activity rate in two categories

(1, active foraging; 2, relaxed foraging, manipulation food, and perching

quietly), the two species showed no difference in the contexts

of usage of type A calls (x2 = .698; df = 1; .5 > P > .3). Type B calls

had similar mean duration (34.7: 37.1 msec) but were very different

in range and variation. Nevertheless, in terms of activity rate, analysed

as before with type A, no significant difference was shown in the usage

pattern of type B (x2 = .085; df = 1; .8 > p >.7). Type C calls showed

the greatest disparity in acoustic structure with a mean duration in the

chickadee version nearly twice that of the titmouse version. However,

the two species' versions were essentially similar in overall inflection

pattern and showed no difference in usage in terms of activity rate.

Small sample size for this call prohibited statistical testing. The calls

of both species also had high likelihood of being followed by long flight.

Type D was also very similar in the two species, both structurally

and functionally.









There was thus a high degree of correspondence between the two

species both structurally and functionally in subsystem I. This corres-

pondence can probably be accounted for by the high degree of inter-

specific relevance of the information made available by these calls.

The calls of subsystem 11 of both species were similar in many

features of acoustic structure, rules of syntax, and functions. Type

A subunits of both species were characterized by their high frequency

and harmonic simplicity. Subunits of titmouse Ila-C were very similar

to chickadee Ila-C in that bothwere harmonic series covering a broad

range of frequency. The titmouse apparently had no counterpart to

the intermediate subunits of chickadee Ila-B. The rules of subunit

combination were the same for both species. The acoustic structure of

the very high frequency subunits (> 9 kHz) were indistinguishable in

the two species. Otherwise, the structure of the chickadee subunits

was distinct from those of the titmouse in duration and in patterns

of frequency inflection within the subunits. The chickadees appeared

to modulate the high frequency and intermediate subunits to a much

greater extent that the broad frequency range forms. The opposite

appeared to be the case with the titmouse. On a few occasions, however,

the calls of the two species converged to an intermediate form during

exchanges between the two speices (Fig. 2 k and 1).

In terms of the functions of the calls of subsystem II, there is

also a high degree of correspondence in that both are involved in the

regulation of flock movements under changing conditions and in facili-

tating mid-range contact and approach. These functions are not always

of equal interspecific relevance. Regulating flock movements










certainly was of interest to both species, but the calls were only

rarely exchanged by the two species, and the messages carried by the

calls appeared to be primarily intraspecific. The two species maintained

flock cohesion intraspecifically since they rarely appeared to be evenly

mixed throughout the floxk, but overall flock velocity and direction

must have been synchronized if the two species were to remain together.

The calls must therefore have had some components that were sufficiently

species-distinctive and others that were held in common to accomplish

this duality of purpose.

The very high frequency subunits of both species carried the message

that extreme danger was at hand and, in the context of fights, strong

escape tendency. The essential ingredient was the same in both cases,

i.e. avoiding danger. This message was of such immediacy that selection

has apparently allowed little ambiguity and the call was strikingly

different from all other calls.

Subsystems lib were clearly derived from calls of Ila. However, they

were both structurally and syntactically distinct. Although fledglings of

both species were observed making obvious attempts to beg from adults of

other species, none of these attempts were rewarded. It appeared that the

adult birds have effectively selected for distinctive begging calls.

Subsystem III of the two species appeared to share a common origin

in subsystem II. Chickadee III calls appeared to use all subunit types

of subsystem II, whereas titmouse II calls used only the type A sub-

units. Chickadee Ill calls were much more complex in structure and

occurred in more distinctively different forms that titmouse Ill calls.

Although specific messages or even consistent contextual features could









be reliably associated with only a few of the chickadee III calls, it was

clear that the range of contexts in which the calls occurred was greater

than for the calls of titmouse IIl. These calls were very rarely

directed to the other species and thus had little, if any, interspecific

relevance. It is not surprising therefore that the calls of subsystem

III should bear no structural similarity between the two species.

Subsystems IV had very little in common outside of their basically

repetitive nature. The calls of chickadee IV showed little variety

compared to those of titmouse IV and were used in a correspondingly

smaller variety of contexts. The messages carried by calls of chickadee

IV appeared to be carried by the calls of titmouse IV, which also appeared

to cover those messages carried by chickadee III but not by titmouse III.

The lack of correspondence between subsystems IV in acoustic structure

was consistent with their lack of messages with interspecific relevance.

The relatedness of the major subsystems of the two species is

summarized in Table 1.

To summarize this section, I have argued that each of the four call

subsystems of the tufted titmouse and Carolina chickadee bore an extent

of similarity in acoustic structure and/or syntax that corresponded to

the extent of interspecific relevance of their messages and functions.

This correspondence is summarized in Table 2.


Review of Communication Systems of the Genus l'iis


Relatively complete surveys of the vocal communication systems of

six species in the genus Parus have been prepared by various authors.

Only three of these, however, were done with the aid of the sound
















Table 2. Summary of correspondence between structural
similarity and interspecific relevance in the
vocal communication subsystems of Pczuus
crw'olinunoic and P. bicolor.


Structural


Interspecific


Subsystems Similarity Relevance

I high high

IlaO high high

Ila intermediate intermediate

lib low low

Ill low low

IV low low










spectrograph (P. carolinensis: Smith, 1972; P. atricapillus: Ficken

et al., 1977; and P. hniconicus: McLaren, 1976). Surveys based on

phonetic renderings were made of the vocal repertoires of P. carolinensis

by Brewer (1961), P. at'fi' apilZul by Odum (1942), P. pil7?intria by

Morley (1953), P. intormlau/: by Dixon (1949) and i'. mciajo by Gompertz

(1961). Discussions of several selected vocalizations have also been

made by various authors (e.g., Dixon et al., 1970) and two authors have

attempted broad surveys of a few vocalizations primarily among European

members of the genus Parus (Latimer, 1977; Thielcke, 1968).

None of these authors organized the vocalizations as I did, but

using their sonograms and descriptions, I have reclassified the vocali-

zations according to the subsystems that I used in the analysis of P.

carolinensis and P. bicolor. This reclassification is summarized in

Table 3.

S. T. Smith (1972) prepared a survey of the vocalizations of P.

carolinensis in a residential area near Philadelphia, Pennsylvania.

Although all subsystems appear to be represented in the vocalizations

of the population she studied and most calls appear to be used in

basically the same way as the birds I studied in Florida, there are

some notable differences. I suspect that these differences are due to

three factors. First, her recordings were apparently made primarily

at feeders and nest boxes in a residential neighborhood. Although she

is confident that no "major categories of behavior were overlooked,"

the tendency for aggressive reactions to be intensified at feeders, and

the limited opportunities for flock movements in gardens and hedgerows

may have biased the interpretation of the patterns of usage of the vocali-

zations. Second, her study population was nearly a thousand miles from











Table 3. Comparison of the vocal communication subsystem
of seven species of the genus Parus.


Close-Range Contact

Flight

Mid-Range Contact

Surprise, Alert

Alarm, Escape


Mobbing

Begging

Aggressive Conflict


Supplication


Long-Range Contact,
Territorial
Advertisement


I*

?







I IaA*

Ilaa0*
IlaA*

I aC

7

I laA'*
III
I a "


I

I-D

I la

I laA


I laO

I laC

I lb

IlaA
III

HSC
III


0




I l


I aC
(?)
I laA

(7)
I laA

?7

I lb

I laA
I I I

(7)
HSC


I la







IlaC*
Ib*


1



I la





I laO

I laC

I Ib

I laB
IIl

HSC
III


1*



I la*

1 laA*


I laO

I laC

I Ib*


HSC* HSC*


" I* III I1I I la
IV* IV III IV IVa IV IV


Designation based on verbal transcription, no sonograms available.










mine and very close to the border of the range of P. atricapillus (Ward

and Ward, 1974). Hybrids are known to occur in some areas along this

border (Rising, 1968; Johnston, 1971), and they are suspected in others

(Brewer, 1963). Third, a very different approach to the problem of

organizing the vocal repertoire was taken by Smith. Classification was

based on assumed message structure of the calls rather than on an

analysis of their physical structure. Sonograms of all call classifi-

cations were presented, but these apparently played little part in

analyzing the relationships among the calls.

Calls of subsystem I (Smith: "Tseet") were present but they were

given a very different interpretation from my own. Smith considered

them to indicate a conflict between escape and "general set" (Smith,

W. J., 1969), i.e. just about anything, possibly encoding "social

interaction hesitance," and to "function to alert other chickadees of

aversive conditions." As discussed earlier, calls of subsystem I occur

during all activities and are associated with nearly all other vocali-

zations. It must also be true that all activities of the chickadees

are accompanied by escape tendencies and "social interaction hesitance"

in such a treacherous world as they live in. But to assert that these

calls are adaptations for making available information concerning

aversivee conditions" is unwarranted by the evidence. Smith's emphasis

on aggressive contexts may have resulted from her over-abundance of

observations at feeders, where aggression is concentrated, and from an

apparent under-represenatation of observations of more typical flock

foraging movements in her sample. Her conclusions regarding this call

differ not only from my own but from all other descriptions of this









call's homologues in other congeners (e.g., Odum, 1942; Ficken et at.,

1977; etc.).

Calls of subsystem IIaA of P. carolinensis are broken down by

Smith into several functional categories without providing sufficient

physical description to distinguish between them (e.g., lisping tee,

loud tee, chip, soft dee). Their relatedness in the continuum of sub-

system Ila is obscured by Smith's classification. Nevertheless, the

contextual information provided with these calls indicates that their

usage patterns are comparable if not identical to those of the Florida

population. This is also true of the begging calls (subsystem Ilb) and

High See alarm calls (IlaO).

Subsystem III was clearly represented in the Pennsylvania popu-

lation but the calls were very different from those of the Florida

population in details of structure. Again Smith's classification system

proved unworkable and homologies to the calls of my population are un-

clear. On first mention of these calls, Smith described them as

variable. She later discussed "variants" of these calls apparently

treating them as stereotyped. She counted 23 "variants" in the nine most

thoroughly studied individuals (I counted 17 in six to ten individuals)

with each individual having at least some variants unique to itself

within its flock. Her recordings at various distances from the study

population revealed distinctly different variants. Smith reported

three different calls of subsystem III in a brief period of observation

16 km from her main study population. I recorded only one minor

variation, an X-R with no T unit, in many hours of observation of the

Gainesville (home) population 20 km from the San Felasco population.










Nearly all of the other III calls of the San Felasco population were

represented in the Gainesville (home) population.

One "variant" in Smith's population (not illustrated) was frequently

associated with "variable see" (HSC) in situations that, in at least some

instances, appeared comparable to situations in which I observed HSC-lllb-

2 in my Florida population. These calls may be homologous.

Smith reported a greater likelihood of attack following calls with

rasps than following calls without rasps. This pattern fits well to

Morton's M-S rules (1977), but is contradicted by my results. As dis-

cussed earlier (see Results), some T forms occasionally were given

without the rasp and were associated with a much higher likelihood of

being followed by an attack than were the complete T-R forms.

Calls of IV were similar in structure and usage between the two

populations. The only song form for which a sonogram was published by

Smith was very similar to one in the Florida population. Others,

designated "song variants" were not necessarily represented in the Florida

population but comparable vocalizations with rearranged subunits did

occur there.

Calls of P. atricapillus appear remarkably similar to those of

P. carolinensis (Odum, 1942; Brewer, 1961; Ficken et al., 1977). All

subunits were represented in P. atricapillus and usage patterns appear

to be very similar. The most notable differences occur in the details

of structure of subsystem III and IV, but not enough information was

made available on III to generalize about the nature of the differences.

A "special type" of III (Dixon et al., 1970; not illustrated) was given

in sexual contexts. This call may be homologous with Illb-2 of










P. carolinensis. Subsystem IV of P. atricapillus differed from IV of

P. carolinensis in being restricted to two notes instead of four.

Parus hudsonicus differed from P. carolinensis and P. atricapillus

in its lack of subsystem IV. McLaren (1976) makes no mention of how

the function of territory advertisement was accomplished in her study

population in Ontario, but Townsend and Allen (1907) in Labrador and

Allen (1910) in New Hampshire reported a "song" that may have been

homologous to calls of subsystem III (Townsend and Allen: "low, bubbly,

warbling'). Calls of III were regularly associated with border patrol-

ling in P. carolinensis and apparently P. atricapillus (Odum, 1942),

and it is easy to imagine their promotion to the role of song in

P. hudsonicus. Patrs rufescens, thought to be a sibling species of

P. hudsonious (Grinnell, 1904), also lacked subsystem IV, but compensated

with a "repetitive trill" (Dawson, 1923).

McLaren (1976) noted the tendency of P. hudsonicus to fuse subunits

of Ila type A to type C subunits. This tendency was expressed by P.

carolinensis and P. bicolor.

Parus palustris, a Eurasian member of the subgenus Poecile,

appears to have communication signals similar to those of its North

American relatives, the chickadees. Sonograms for only subsystems IV

and Ila type C are available in Thielcke (1968) and Latimer (1977).

Other calls have been described by Morley (1953). From these descrip-

tions it appears that P. palustris uses calls homologous to subsystem

I, Ila-0, Ila-A, a combination call of la-A and C, and Ill. These

calls appear to have uses similar to those of the North American

chickadees.









Sonograms of subsystems IV and Ila in part of P. inornaitus

are available in Dixon (1969). Other calls can be inferred from trans-

criptions in Dixon (1949). No mention is made in either of these papers

of any calls resembling subsystem III of P. bicolor, and it is unclear

what calls are associated with hostile interactions in P, incrnatus. A

call that appears to be homologous with HSC was used by females early

in the breeding season in place of the more normal begging calls of

subsystem Ilb. Begging calls of Ilb were used later in the season.

This usage of HSC is consistent with their function of supplication

in other congeners. The calls of Ila were considered by Dixon (1949)

to be more similar to those of P. atricapillus than to those of the

more closely related P. biuolor.

Sonograms are available for subsystem IV of P. major in Gompertz

(1961), Thielcke (1968), Krebs (1976), and Latimer (1977) and for Ila

in part in Thielcke (1968) and Latimer (1977). Other calls can be

inferred from transcriptions in Gompertz (1961) and Hinde (1952).

Begging calls that appear to be homologous with lib, close-range contact

calls of subsystem I, and HSC all were represented, but nothing com-

parable to III of the species discussed above appears to be present in

the repertoire of P. major. They appear to compensate for this deficiency

with calls of la and several other calls of uncertain affinities

(Gompertz: "tsee, pee, tink;" possibly isolated IV notes).

Thielcke (1968) and Latimer (1977) both made surveys of subystems

IV and Ila (termed alarm calls, but this assumed function appears to be

too broadly applied) of many species, primarily European, of the genus

Parus. The majority of the species surveyed in these works had "songs"

similar to subsystem IV of P. carolinensis or P. bicolor in that they








were repetitive series of a simple phrase. Others appear more similar

to P. 'jlincfzil u subsystem I Ill, for example I'. in': { ',il'f'l'' 'l in

Thielcke (1968). Still others appear more similar to subsystem I Ia,

for example, P. cia 'ul ua in Thielcke (1968), or P. 'iT: tice in Latimer

(1977). Subsystem liaC, or at least notes comparable to IlaC of

P. carolinensis, appear to be a constant feature of the communication

systems in the genus Przit,:.



Interspecific Effects on Vocal Communication Signals



Vocal communication signals of any species can be expected to be

influenced by those of other sympatric species in several ways. In a

community of many species, competition for the acoustic medium can be

intense. Species may avoid each other by finely partitioning the

frequency spectrum ( Rand and Drury, ms), by distinctively patterning

their calls (Bremond, 1978), by adjusting their schedules to call at

different times of the day (Smith, 1. J., 1977, pg. 372) or by alter-

nating calls with competitors (Cody and Brown, 1969; Ficken, et al.,

1974; Marler and Tenaza, 1977). Distinctive patterning in long distance

calls is especially critical for closely related species that are in

danger of hybridizing (Brown and Wilson, 1956; Marler, 1957; Sibley,

1961) and sibling species can be expected to diverge in these calls.

The opposite trend, character convergence, has been suggested

to account for many cases of similarity in appearance of song between birds

that could possibly be interspecifically territorial (Cody, 1969; 1973).









Although none of the cases used by Cody to argue for the idea were

adequately supported by evidence, and many have been discredited (Murray,

1976; Brown, 1977; Hardy and Murray, ms), the idea may yet have some

validity under certain circumstances. The hypothesis suggested by

Cody (1969, 1973) is that birds with considerable resource overlap

should be interspecifically territorial and should develop common

signals for mutual recognition that will facilitate avoidance. As

pointed out by Murray (1971, 1976), such interspecific territoriality

is unlikely to be in the mutual interest of both species since one

species will tend to dominate the other and exclude it from optimal

habitat in keeping with the competitive exclusion principle. Thus, it

would hardly be in the interest of the subordinate species to retain

the signals that provoke the dominant species to react aggressively

to it. Sibling species, however, may be very closely matched in

aggressive prowess as well as ecological requirements. In areas of

secondary contact, it may serve these species to exclude each other

from their territories and to develop the communicative adaptations to

facilitate avoidance. Emlen et al. (1975) reported that indigo and

lazuli buntings (Passerina cyanea and P. aznoena) maintained interspecific

territories in their zone of sympatry. "Figures" (syllables) of the

songs of the other species were incorporated into each species' songs

in sympatry but not in allopatry, and playback response was elicited

to the other species song in sympatry but not in allopatry. Similarly,

Ward and Ward (1974) found that Paris atricapillus and P. car'olinensis

sang both their own songs and songs of the other species in a zone of

sympatry in SE Pennsylvania but not outside that zone. Response to










playback was elicited to the other species' song in sympatry but not

in allopatry. Interspecific territoriality was not investigated. Abnormal

songs have been reported in several areas of overlap (Rising, 1968;

Brewer, 1963; Johnston, 1971; Ward and Ward, 1974). "Bivalent" and

abnormal song in these areas may be the result of hybridization but

they may also be adaptations for facilitating interspecific territoriality.

Convergence, or lack of divergence, has been demonstrated in

alarm and mobbing calls among sympatric bird species (Marler, 1957) and

monkeys (Marler, 1973). That the calls for alarm should need to be very

high frequency, whistle-like tones has been suggested by Marler (1955)

since this type of tone is presumably minimally locatable. Shalter

(1978) has suggested that a better explanation for the high frequency

of alarm calls may be in accordance with Darwin's (1872) principle of

antithesis in that the higher the sound, the more unlike it is to

aggressive growls and scolds and thus indicative of escape tendency.

Both arguments suggest that there is an ideal acoustic form for an

alarm call, and, by their arguments, it is reasonable to expect that

the alarm calls of titmice and chickadees should converge on this

common form.

In the case of mobbing calls, the advantages of recruiting intra-

specific participation are clear. Marler (1955) suggests that here

too the physical structure of the calls suits them to their function

in that their broad frequency-spectrum makes them locatable. Their

harsh quality, along with their low frequency components, gives them a

universally recognizable aggressive and aggravated connotation. The

convergence of mobbing calls to this common form among many species is









to be expected. In my study, chickadees and titmice participated in

mobbings with unintroduced series of IlaC subunits. The calls of the

two species were distinguishable but similar. Carolina wrens, ruby-

crowned kinglets, and solitary vireos also participated in mobbings with

the characteristically broad spectrum, harmonically rich calls. The

warblers also participated, but with emphatic chips that also covered

a broad frequency range but were harmonically simple.

The extent of divergence in the vocal communication systensof the

Carolina chickadee and tufted titmouse is in agreement with the con-

clusions and predictions of Marler (1957) and also with his conclusions

on the extent of divergence in the calls of two sympatric and inter-

specifically social Cercopithecus monkeys (Marler, 1973). In his

studies, as with mine, the long distance signals (subsystems IV of the

parids) were the most distinctive. Unlike the monkeys of Marler's

study, however, the subjects of my study were probably in very little

danger of hybridizing, and the divergence in their long distance calls

was probably due as much to the different strategies of territory

defense and aggression by the two species as to the need for species-

distinctiveness. Once contact was established with calls of subsystem

IV, the chickadees approached each other and exchanged calls of sub-

systems II and Ill. The titmice, however, continued to exchange

with calls of sybsystem IV. The greater communication burden on

subsystem IV of the titmouse results in a more elaborate and thus

distinctively different complex of calls from subsystem IV of the chicka-

dee. Specific distinctiveness may, however, be important to these two

species, not in reference to each other, but to the more closely related

and geographically adjacent species of their respective subgenera.










In addition to the high frequency alarm calls and mobbing calls of

subsystem II, the calls made up of combinations of other subunits also

showed similarities not only in the two parids of my study but in all

parid species for which data are available. The extent of divergence in

the two species of my study probably represented a compromise in that

a certain degree of specificity was necessary for the two species to

maintain group cohesiveness independently of each other and at the same

time regulate overall mixed flock movements. The problem has apparently

been resolved by allowing call subunits to diverge in details of acoustic

structure while retaining at least the basic features of a common syntax.

This case is similar to that found by Marler (1973) in the Cercopithecus

monkeys. The calls known as "phrased grunts," were given by both species

in similar contexts and appeared to function in coordinating group

movements. The calls were very similar in basic acoustic structure

between the two species and were considered to be clearly homologous.

However, the calls of the two species differed in "phrase" duration,

"phrases" per call, variability of these two parameters, and in the

extent of high frequency emphasis. Marler interpreted these differences

as having arisen in response to the need for maintaining intraspecific

cohesiveness within the interspecific group.

That birds should be responsive to patterns of syntax is not with-

out precedent. The songs of many birds are made up of repetitions of

various song types, the precise arrangement of which are species

specific. Falls (1963) demonstrated that the exact timing of the

subunits in the song of the ovenbird, Seiurus aurocapillus, is necessary

for species recognition. Boughey and Thompson (1976) found that brown











thrashers (Texostoma rufun) responded to songs of catbirds (LDuatella

carolinensis) and mockingbirds (Miras po;ly'lottos) when altered to

match the phrase repetition pattern of normal brown thrasher song.

Although data are incomplete, the syntax rules of subsystem IIa

of P. carolinensis appear also to be used by P. atricapillus (Ficken,

et. al., 1978) and P. h;dsonicus (McLaren, 1976).

The intra and interspecific relationships of the various subsystems

are summarized in figure 14. Subsystems IIa are considered to be

central in each species vocal communication systems. Subsystems I

and IV are shown as possibly having been derived from IIaA, but

these derivations are highly tentative and based only on their

acoustic simplicity and common basic functions, i.e. contact at various

distances. Subsystems I showed strong correspondence in structure

and function, and it is very likely that this was the result of con-

vergence due to an interspecific influence. IIaO, the high whistle

alarm calls, also appear to be convergent. This convergence could be

due as much to selection for physical properties of the calls as to

interspecific influence. Subsystems IIa were convergent primarily

in syntax. The precise structural configuration of the subunits

showed less correspondence. Subsystems IIb appear to be divergent due

to selection for specific distinctiveness. The divergence of subsystems

IV could be due as much to differences in strategies of territory

defense as to the need for specific distinctiveness.



































Figure 14. Summary of intra- and interspecific relationships
of vocal communication subsystem of Parus
caro'inensis and P. bicolor.












CHICKADEE


N.


--------- possible derivation
-probable derivation
Sintersp. influence resulting in convergence
intersp. influence resulting in divergence


TITMOUSE













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