A. LIFE-HISTORY STUDY OF THE TUFTED TITMOUSE..
PARUS BICOLOR LINNAEUS

' ;

Thesis {or ”no Degree of pk. D.
MICHEGAN STATE UNIVERSITY
Wiiiiam P. Pielou

1957

 

 

 

 

 

 

 

 

 

 

 

121 11111111111111111

29300 3014

This is to certify that the

thesis entitled
A Life-History Study of the Tufted

Titmouse, Pams tricolor Linneaus.

presented by

William P. Pielou

has been accepted towards fulfillment

of the requirements for

M degree in M

in a... - u;
ajor rolessor

DateNovember 15, 1957.

 

0-169

 

 

_‘
‘- _--_._A—_ ——_..

w -

LIBRARY

Michigan State
University

 

 

 

PLACE IN RETURN BOX to remove this checkout from your record.
TO AVOID FINES return on or before date due.

 

DATE DUE/ DATE DUE DATE DUE

 

 

 

I‘W‘.)

r s .
1 § ‘ l
. .,

 

 

 

 

 

 

 

to“ £5.

11 3;
J ”‘9

 

 

 

III/{Riki 2505"

 

 

 

 

 

 

 

 

 

MSU Is An Affirmative ActiorVEqual Opportunity Institution
cm

m3

A LIFE-HISTORY STUDY OF THE TUFTED TITNCUSE,
Parus bicolor Linnaeus

BY

William Percival Pielou

AN ABSTRACT

Submitted to the School of Graduate Studies of Michigan
State University of Agriculture and Applied Science
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Zoology

1957

Approved aged”. I “Pang“

William Percival Pielou

A life-history study of the Tufted Titmouse, Earns
bicolor, was the object of investigation from September 1953
to September 1955. The study is presented in two parts:

(1) classification, nomenclature, and distribution; (2) observ-
ations on the annual cycle. The information in Part I was
based on a review of the literature and extensive correspond-
ence. The mapping of the distribution of the Tufted Titmouse
in Michigan is based in part on the author's own observations
but mainly on correspondence with ornithologists throughout

the state.

The data presented in Part II were gained largely from
field observations in three woodlots on the property of
Michigan State University, East Lansing, Michigan. This was
the principal objective of the research and included a study
of: (l) the nesting cycle, from the time of pairing and
separation of birds from the winter flocks to the dispersal
of juveniles and formation of the fall flocks; (2) ecological
relationships; (3) flock composition and behavior; (A) voice;
(5) roosting; (6) food and feeding habits, based on stomach
analyses and field observations.

Eggting Cycle. Pair formation, and separation of birds
from the winter flocks, occurred in late March and early April.

Nest construction, which was the duty of the female, began

William Percival Pielou
several days after pairing. Territory establishnent seemed
to coincide with nest construction. The male defended the
territory until the young left the nest.

All nests were located in abandoned woodpecker holes
or natural cavities in living deciduous trees. The average
height of twenty nests was 38.5 feet. The loosely constructed
nests all contained dried grass, moss, and hair, and often
other.miscellaneous items.

First eggs were laid from May A to Nay 11, and the eggs
were deposited at the rate of one a day until the full comple-
ment of five or six was reached. COpulation was observed only
during this period. Incubation was performed solely by the
female and she was fed during both her attentive and inatten-
tive periods by the male. Incubation lasted approximately
thirteen days.

Hatching brought about a shift of the male's feeding
attention from his mate to the young. The rate of feeding
the young by both parents gradually increased until the young
left the nest, fifteen days after hatching.

The young remained with their'parents at least several
days and in some cases the family group formed the nucleus
of fall flocks.

Flocking. Flocks of titmice and associated species began

forming as early as August. The size of the flock was variable

William Percival Pielou
but averaged seven birds. During their feeding excursions
in the fall the titmouse groups were enlarged with nigrart
birds, especially warblers.

Xgigg. Of its variable repetoire the Tufted Titmouse's
most commonly heard and characteristic song is the "peto,
peto" expression. In addition it has a location note, alarm
note, reCOgnition note, hissing note, invitational note,
begging note, and distress note.

Roosting. Roosting occurred in evergreen and deciduous
trees and the birds roosted wherever they happened to be at
sundown during the fall and winter. During the nesting cycle
the female slept in the nest cavity while the male roosted
nearby.

Eggd Egg Feeding Habits. Tufted Titmice consume two-
thirds animal matter and one-third vegetable matter annually.
Food was secured from the ground litter to the uppermost
twigs of the tallest trees. Insects were gleaned mostly
from the bark and leaves but hawking of insects was noted

on one occasion.

A LIFE-HISTORY STUDY OF THE TUFTED TITNOUSE,
Parus bicolor Linnaeus

By

William Percival Pielou

Submitted to the School of Graduate Studies of Michigan
State University of Agriculture and Applied Science
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Zoology

Year 1957

TABLE OF CONTENTS

Page

INTRODUCTION . . . . . . . . . . . . . . . 1
ACKNOWLEDGMENTS . . . . . . . . . . . . . . 2
PART I: CLASS FICAI‘ION, NC CEITCL’ITIBE, AETD DISTE‘LIBUTIMJ . 3
Classification . . . . . . . . . . . . . 3
Nomenclature . . . . . . . . . . . . . 3
Nomenclatural History . . . . . . . . . 3

Common Names . . . . . . . . . . . . 5
Distribution in North America . . . . . . . . 6
Geographical Distribution . . . . . . . . 6
Ecological Distribution . . . . . . . . 6
Distribution and Status in Michigan . . . . . 10
Historical Background . . . . . . . . . 10

Current Distribution . . . . . . . 11
Extension of Range . . . . . . . . . . . 14
PART II: OBSERVATIONS ON THE ANNUAL CYCLE . . . . . 18
Introduction . . . . . . . . . . . . 18
Materials and Methods . . . . . . . . . . 18
Banding . . . . . . . . . . . . . . 18
Painting . . . . . . . . . . . . . l9
Trapping . . . . . . . . . . . . . 19

The Study Area . . . . . . . . . . . . . 21
Sanford Woodlot . . . . . . . . . . . 21

Red Cedar WOodlot . . . . . . . . . . 23

Baker Woodlot . . . . . . . . . . . . 23

Climate and Weather . . . . . . . . . . 2h

The Population at Beginning of Study . . . . . 25
Pair Formation . . . . . . . . . . . . . 25
Time of Occurrence . . . . . . . . . . 25
Courtship and Mating . . . . . . . . . 26
Behavior of Newly Paired Birds . . . . . . 27
Permanence of Mating . . . . . . . . . 28
Territory . . . . . . . . . . . . . . 29
Establishment . . . . . . . . . . . . 29

Defense . . . . . . . . . . . 30

Size of Territory . . . . . . . . . . 32

Habitat . . . . . . . . . . . . . 3h
Population Density . . . . . . . . . . 3h

The Nest . . . . .
Nest-Site Selection
Nest Construction
Nesting Associates

Egg Laying . . . .
Deposition of Eggs
Description of Eggs
Behavior during Egg

Incubation . . . .
Role of Sexes . .
Behavior during Incu
Incubation Period
Hatching . . .

Care of the Young . .
Brooding and Feeding
Nest Sanitation . .
Length of Nest Life .

Development of Nestlings
External Appearance .
Behavior . . . .
Number of Broods . .

Postnest Life . . . .
Nest Leaving . . .
Dispersal of Young .

Nesting Mortality and Lon
Nesting Mortality
Longevity . . .
Predators . . .

Flocking . . . . .
Formation . . .
Size and Composition

Associates . . .

Le

C7

ee<eeeeee

Roasting Habits .
Fall and Winter
Breeding Season
Time of Roasting

Voice . . . . .
"Peto" Song .
Location Note .
Alarm Note . -.
Rec0gnition Note
Hissing Note .
Invitational Note
Begging Note . .
Distress Note . .

at

eeoo'eetdeeeeeee
m
‘1'

oeeeeoeHeeH'oeeeeeo

p.
('1’

oeoeeeoeeeoefieeeeeeeoeeeoe

e e 0 e e e e e 0 0 e O

Food and Feeding Habits

Food . . . .
Feeding Habit

SUMMARY . . . . . .
APPENDIX 0 O O O O 0
Tables
Lists
Plates

LITERATURE CITED

Page
. oo

Fig.
Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.
Fig.

LIST OF FIGURES

Distribution of the Tufted Titmouse . . . . 7

Major Biotic Communities of Eastern United
States . . . . . . . . . . . . . 8

The Distribution of the Tufted Titmouse in .
Michigan . . . . . . . . . . . . . 13

The Biotic Communities of the Lower Peninsula
of Michigan . . . . . . . . . . . . 13

Campus and Associated Woodlots, Michigan State
University, E. Lansing, Michigan . . . . . 22

Territories of the Tufted Titmouse in Sanford
Woodlot . . . . . . . . . . . . . 32

Territories of the Tufted Titmouse in Red Cedar
WOOdIOt e e e e e e e e e e e e e 32

Average Hourly Rate of Feeding Nestlings . . L6
Boosting Time in Relation to Sunset, l95h-‘55. 61

Introduction

The Tufted Titmouse, Parus bicolor Linnaeus, was
selected as the subject for this investigation because of
its permanent-resident status locally; because of its
comparative tameness and availability for study; and because,
despite the fact that it is a common bird, many aspects of
its life history have not been recorded.

The chief purposes of the research were to learn more
about the life history of the Tufted Titmouse in Michigan
and to assemble all available data, both new and old, about
its habits. The new facts about its habits were gained
through a field study of its annual cycle in lower Michigan.

The material is presented in two parts: (1) classif-
ication, nomenclature, and distribution; and (2) the annual
cycle in Michigan. The material included in Part I is a
summary of the information obtained by a review of the
literature and by correspondence with selected members of
various state ornithological societies and other individuals.
The events in the annual cycle are presented in the follow-
ing order: pair formation and the coincident dispersal of
the winter flocks, the nesting cycle, dispersal of Juveniles,
mortality, voice, flocking, roosting, and feeding.

Field observations were mainly limited to a two-year

period, from September 1953 to September 1955. Additional

2
information on nesting habits is included from observations
made during the spring of 1956. The study was conducted in
three East Lansing woodlots belonging to Michigan State
University. Titmice were banded from September to April
during the study period. As part of the food-habits study,

I examined the stomach contents of five adults.
Acknowledgments

The completion of this study was partially dependent
upon information obtained from other individuals. Correspond-
ence regarding observations in other hichigan localities was
very helpful, and I am grateful to everyone who took time to
answer letters and to offer information and suggestions.

Acknowledgments are due first to Dr. George J. Wallace,
chairman of my guidance committee, for his advice during the
study and for his constructive criticism in the preparation
of the manuscript. The other members of the guidance
committee, Dr. Walter F. Morofsky, Dr. William B. Drew, and
Dr. Henrik J. Stafseth, have been helpful in reading and
criticizing the manuscript.

Acknowledgments would be remiss without due credit to
my wife, for her constant help and encouragement throughout

this investigation.

PART I
CLASSIFICATION, NCthCLATURE, and DISTRIBUTICK
Classification

The Tufted Titmouse is a member of the Family Paridae,
which consists of titmice, verdins, and bushtits. Each of
these groups constitutes a separate subfamily: Farinae, tit-
mice; Remizinae, verdins; and Psaltriparinae, bushtits. The
Subfamily rarinae now contains only the one genus, Egrgg,
which includes all chickadees and titmice. The 7 species of
chickadees in North America currently include 30 subspecies
(Fifth A.O.U. Check-list of North American Birds, 1957:382-
393). Although three of the titmouse species are polytypic,
Parus bicolor is monotypic, containing only one described

form.
Nomenclature

Nomenclatural History. The genus Parus was established
by Linnaeus (Systems Naturae, ed. X, 1, 1758:189). In the
12th edition, vol. I, of his Systems Naturae, 1766, he refers

to specimen 340 as Parus bicolor. Its designation was based

 

on neither a specimen nor a description, but on a painting
by Mark Catesby (1731) of the Crested Titmouse, Parus
cristatus (Allen, 1951zh66).

A
In 1829 Kaup suggested the generic name Lophophanes

 

for the crested titmice, type Parus cristatus. Thus it is
that Bonaparte's List of the Birds of Europe, 18h2, gives
Lophophanes and Eaggs as the generic names of the crested
titmice and chickadees respectively (Baird, 1875288). In

1850, Cabanis suggested the new genus Baeolophus for the

 

American crested titmice (type, Parus bicolor, L.). In this

 

same year Reichenbach suggested the genus Penthestes for the

 

chickadees (type, Parus lugubris, Temminck). Baird (1858),
disagreeing with Cabanis's suggestion that our genus of
tufted titmice was distinct from the European crested tit-
mice, continued to use Lophophanes for the crested titmice
and Egggg for the chickadees. The First and Second A.O.U.
Check-lists, 1886 and 1895, list the genera Egggg and
Lophophanes. In the 12th A.O.U. Supplement (1903:357-359)

the genus Lophophanes, Kaup, was changed to Baeolophus,

 

Cabanis.
Ridgway (1904) recommended the splitting of the orig-

inal genus Parus into the following genera: Lgphophanes for

 

the crested titmice of Europe, Baeolophus for the crested

species of the Nearctic region, and another genus (Penthestes)

 

for the chickadees. In 1908 the A.O.U. Check-list Committee
acted on Ridgway's recommendation and re-established Reich-
enbach's generic name of Penthestes for the chickadees

(14th Supplement, 1908:386-387).

5
The Third A.O.U. Check-list (1910:3h7-352) lists the genus

Baeolophus, Cabanis, for our crested titmice, and Penthestes,

 

Reichenbach, for our chickadees. This nomenclature was
continued in the Fourth Edition in 1931. Hellmayr suggested
uniting all the Parinae into the single genus Egggg and does
so in his Catalogue of Birds of the Americas (1934:70).
The A.O.U. Check-list Committee--in the 19th Supplement,

19hh--acted on Hellmayr's suggestion and united Penthestes

 

and Baeolophus into the single genus Egggg. Thus the orig-
inal name of Parus bicolor, Linnaeus, for the Tufted Titmouse,
is reinstated.

Common Names. The English name for the Latin generic
name Egrgg is "titmouse.” The specific name, bicolor, means
"of two colors." The most widely used common name for this
species is Tufted Titmouse, often shortened to Tufted Tit.
Other names, sometimes still encountered in older works,
are Crested Titmouse, Top-knot bird, Black-fronted Titmouse,
Tomtit, Peter-bird, Sugar bird, Tufted Chickadee, Peto-bird,

Crested Tomtit, and Tip-top.

Distribution in North America

Geographical Distribution. Since the Tufted Titmouse
is essentially non-migratory, its winter range and breeding
range are practically identical. In distribution it is
limited to the eastern half of the United States (See Fig.1).
The political limits to the north are the southeastern corner
of South Dakota, the southern part of Minnesota and Wisconsin,
the lower half of the lower peninsula of Michigan, the lower
half of New York State, and the western part of Connecticut.
The eastern boundary is the Atlantic coast, from Connecticut
to Florida. The southern limit extends west from Florida
along the Gulf coast to the eastern half of Texas. The
western extension is to the eastern part of Nebraska and
Kansas, and the eastern half of Oklahoma and Texas. The
usual breeding range is slightly less extensive and within
these limits it is a common resident except along the Appal-
achian ridge, where its numbers decrease with altitude.

The occasional nesting records that have been reported out-
side of the established breeding range are indicated on the
accompanying map (Fig. l).

Ecological Distribution. It is well established that
the distribution of any organism depends on the interaction
of many factors-~climatic, physiographic, and biotic-~and

the Tufted Titmouse is no exception. Its range is directly

 

'fil‘g’l‘é
"W
1% . .mi,,,%

- Breeding record outside
regular range

 

§

Legend

[253 Breeding Range
0 Winter Visitor

DISTRIBUTION OF THE TUFTED TITMOUSE
Compiled from data supplied by the Patuxent
Research Refuge, Laurel, Maryland.

Figure l

 

 

\ \ Legend
Deciduous Forest

m Oak-Pine Subclimax C f D id

oni erous- ec uous
Pine Subclimax ’Illl Forests Ecotone
m Deciduous Forest- [:1 Grassland

Grassland Ecotone

 

.‘.:'.°. Coniferous Forest

MAJOR BIOTIC COMMUNITIES OF EASTERN UNITED STATES
Compiled from F. A. Pitelka (19hlzllh)

Figure 2

9
affected by climate, which in turn influences distribution
through its effect on plant life.

Vegetation is of prime importance in supplying such
essentials as nest material, nest sites, food, and cover.
There is a definite correlation between the distribution of
the deciduous forests in North America and the range of the
Tufted Titmouse. Fitelka (1941) states that it occurs in
deciduous forest growths ranking from mid-seral to climax
stages, occurring most frequently, however, in mature flood-
plain forests. A comparison of Figure 1, showing the dis-
tribution of the Tufted Titmouse, and Figure 2, showing the
distribution of the principal vegetation types of North
America, reveals this close correlation. The apparent ex-
tension of the range of this species into the grasslands of
Oklahoma, kansas, Nebraska, and southern South Dakota, is
explained by the presence of bottomland woodlands along
waterways and plantings about farms.

Another biotic factor which is probably of some influ-
ence in the distribution of the Tufted Titmouse is its
association with other hole-nesting species that excavate
their own nest cavities. Because the titmouse does not
have a bill adapted for digging into wood it must rely on
old woodpecker holes or decayed knot-holes for nest sites.
Since these are usually plentiful in fairly mature stands of
deciduous trees, titmice do not ordinarily have difficulty

locating suitable nest cavities.

Distribution and Status in Michigan

Historical Background. The first records of the

 

 

Tufted Titmouse in hichigan were reported toward the end of
the last century. Wood (1921:59h) reported a personal sight
record on October 7, 1878, in the vicinity of Ann Arbor,
Washtenaw County. The first record of a specimen was from
Detroit, Wayne County, collected by E. E. Campbell on Decem-
ber 7, 1878. Other specimens were collected by B. F. Sykes
at Almana, Van Buren County, on hay 12, 1880, and by J. E.
Nichols at Lansing, Ingham County, in September, 1881. On
October 5, 1886, N. A. Wood collected three specimens at
Ann Arbor. Two specimens were taken by S. E. White in June,
1891, in Kent County--the northeranSt station up to that
time. A. J. Cook (1893:126) lists the Tufted Titmouse as a
common winter bird in the more southern parts of the state
and comments that it is a summer resident in the northern
counties of Indiana. At the time of his writing there were
apparently no nesting reports for the state. The first
breeding record was reported by F. M. Kidd on hay 13, 1898,
at which time he collected a set of eggs on Belle Isle,
Wayne County (Wood, 1951). By the turn of the century this
species was observed frequently as a winter bird in the
lower tier of counties but had not yet become a permanent

resident in the state.

11

During the first 25 years of this century the Tufted
Titmouse gradually established itself as a permanent resident
in the southern counties. In 1908, A. D. Tinker reported a
breeding record a few miles west of Ann Arbor. In 1916
another nesting record was reported by F. A. Stuart from
Calhoun County and A. D. Tinker reported another Washtenaw
County breeding record in June, 1918.

Current Distribution. During the last 30 years the
Tufted Titmouse has become firmly established as a breeding
bird in the southern counties of the lower peninsula, has
increased in numbers, and is gradually extending its range
northward. The northernmost nest records are from Brook
Township, Newago County; Alma, Gratiot County; and Imlay
City, Lapeer County. The species has been observed through-
out most of the year at kt. Pleasant, Isabella County, and
at midland, Kidland County, but no actual nest records have
been reported up to 1957. Winter records of this bird in
areas north of its breeding range have been reported from
Stony Lake, Oceans 00.; Manistee, hanistee Co.; Gladwin,
Gladwin Co.; Mio, Oscoda Co.; Loon Lake, Iosco Co.; Prescott,
Ogemaw 00.; Bay City, Bay Co.; Sebewaing, Tuscola Co.; and
Charlevoix, Charlevoix Co. The latter record--the most
northern one yet reported--was of a bird photographed by

J. J. Fessenden on January 7, 1937. This is certainly an

l2

unusual record because the Tufted Titmouse has not been
reported from Charlevoix in any year since.

Nest records indicate that the present breeding range
of the Tufted Titmouse extends northward into lichigan as
far as an imaginary line running from huskegon across the
state through Alma, Saginaw, Imlay City, and Port Huron.
Since it has been reported in almost every month of the year
in ht. Pleasant, Isabella Co., and in Midland, Midland 00.,
it is reasonable to assume that it breeds in those areas,
even though no nests have been reported. Forth of the breed-
ing range its status is that of an occasional winter visit-
ant. The limits of the range of the Tufted Titmouse in
hichigan are indicated on the accompanying map (Fig. 3).

A comparison of Figure 3 and Figure 4 reveals the
close correlation between the range of the Tufted Titmouse
and the distribution of the deciduous forest in Michigan.
The deciduous forest is found in the lower half of the lower
peninsula. The remainder of the state is somewhat ecotonal,
being bounded by the Canadian coniferous forest to the north,
and by the deciduous forest to the south. The dependency
of this species on deciduous trees is explained by its need
for nest sites. Old woodpecker holes and decayed knot-holes

are usually not found in conifers.

 

   
       

IS. “I! “(I‘d (LAD!

  

The Distribution of the
Tufted Titmouse in Niohigan
Figure 3

«no
“'1‘ "mm

   

n "on

The Biotic Communities of the
Lower Peninsula of wichigan
Figure A

  
      

Legend
Perl-neat Reeident

0 Winter Vieitor

Legend

§!§§ Deoiduoue toreet

Coniferous-Deciduoue
rorente lootone

:13

Extension of Range

The Tufted Titmouse has been gradually extending its
range northward in other states as well as in hichigan.
There are many reports of the first appearance and the subse-
.quent establishment of these birds in regions beyond their
regular breeding range. The following examples of this
extension are typical of the situation all along its northern
limits.

In hinnesota the first specimen was taken in 1920,
and by 1932 titmice were breeding in the southeastern part
of the state. Numerous sight records have been reported
recently from several localities north of the hinneapolis-
St. Paul area. A titmouse was banded at St. Cloud in
November, l953--the northernmost record at that time.

In Connecticut there has been a gradual increase in
sight records since 1934. The first breeding was noted at
Westport in 19L9. L. Griscom (195A), reporting for the New
England area, states that the Tufted Titmouse is slowly
spreading northward into western Massachusetts and Vermont
but that there are no nesting records north of Connecticut.

Although it is easy to cite examples of the northward
extension of the range of the Tufted Titmouse, identifying
the causes is more difficult. The environmental changes

that have resulted in conditions favorable for this expansion

15
have been occurring over a long period of time. Apparently
these changes have been unusually favorable during the past
few decades. Such factors as the availability of suitable
nest sites, food, predators, and competition with other
species of similar habits influence extension of the range.
Climatic factors, acting directly on the species, or indi-
rectly through their influence on plant succession, must
be considered also.

In the last few decades the climate has been gradually
changing in the area marking the northern limit of the
range of the Tufted Titmouse. 1n Lichigan, for example,
the average temperature has increased slightly since the
turn of the century. Baten and Eichmeier (1956) state that
the average monthly temperatures for the last 27 years of
this century were higher than those for the first 27 years.
The most pronounced change was recorded for the months of
January and February. The mean temperatures of these months
increased one and three degrees respectively. The fact that
these two months are often the most difficult for bird life
makes these slight changes significant.

This gradual temperature change, especially during
the winter months, could affect the expansion of the range
of the Tufted Titmouse in two ways. First, the warmer
winter temperatures would increase the chances for survival,

causing an increase in the breeding population. This

16

subsequent increase in the population beyond the nest-
carrying capacity in a given region could induce movement
into new localities. Second, if the birds found suitable
habitat in the new area, they might become established as
permanent residents, though only if the winter temperatures
were favorable.

The records indicate that the Tufted Titmouse gener-
ally appears first in areas north of its breeding range
during the winter months. After its initial appearance it
gradually becomes established as a breeding bird. This
behavioral pattern lends support to the possibility that
winter temperatures are an important ecological factor in
the spread of this species.

Vegetation should also be considered as a possible
environmental factor. The range of the Tufted Titmouse
parallels the distribution of the deciduous forests in
eastern North America and experimental evidence indicates
that these forests are spreading northward. In Michigan,
Parmalee (1947) indicates a possible series of floral
changes that have occurred in the southern part of the state
since the period of glaciation. The most recent stage
cited by him is characterized by a return to mesophytic
conditions. The increase in mesic sites and the drying up
of hygric sites has resulted in an invasion of mesophytic

plants, primarily beech and maple. This change in

l7
vegetation has been due mainly to the result of combined
climatic and edaphic changes. These deciduous woodlands,
that have invaded previous xeric and hygric sites, are now
providing suitable habitat for a whole new association of
animals, including the Tufted Titmouse. Its establishment
in these deciduous woodlands, however, did not occur until
there was a sufficient number of old trees in which other
hole-nesting species could excavate cavities. This nest-
site requirement is probably one reason why the northward
spread of the Tufted Titmouse has lagged behind the similar
movement of other cavity-nesting birds.

Several other Species of birds parallel the Tufted
Titmouse in extending their northern limits. Included in
this movement are three warblers--lrothonotary, Blue-winged,
and Cerulean--as well as the Bewick's Wren, the Carolina
Wren, the Blue-gray Gnatcatcher, the Red-bellied Woodpecker,
the Red-shouldered Hawk, and the Cardinal. The range
extension of the latter bird, as well as that of the Tufted
Titmouse, has especially come to the attention of individuals
who maintain feeding stations. Consequently, the literature

contains more references on these species than on the others.

Part II

Observations on the Annual Cycle

Introduction

A field study of the annual cycle of the Tufted
Titmouse at East Lansing was the main objective of this
part of the research. As mentioned previously, the obser-
vations were largely limited to a two-year period-~September,
1953, to September, 1955. Some information on nesting habits
during 1956 is also included. The investigation was carried
on with equal intensity throughout the study period. The
observations on the various aspects of the life of the

titmouse included a survey of associated plants and animals.

Materials and Methods

Banding. In a study of this nature it was necessary
to be able to identify individual birds. For this purpose
numbered aluminum bands and painted aluminum bands (size 1B),
furnished by the Fish and Wildlife Service, were used. The
red, yellow, and green bands, together with the numbered
band, were arranged in various combinations on either one or
both tarsi. For example, individual R-YGA was banded on the

right tarsus with yellow, green, and aluminum bands.

19
No difficulty was experienced in seeing the colors at a
distance of 75 feet. Distances greater than this made
accurate identification questionable.

Twenty-nine birds were marked: ten adults during the
fall and spring of l953-‘5h, nine nestlings in the summer
of l95h, and ten individuals during the fall and spring of
l95h-‘55.

Painting. An adult male (Al9h151) from Fest RC1 was
marked with white "Duco" enamel so that his relationship to
his mate, nestlings, and other birds could be followed.
Approximately five millimeters at the tip of the tail was
painted and allowed to dry before releasing the bird.
Although his mate and other individuals regarded him with
curiosity at first, they soon seemed to accept this alter-
ation of his appearance and no adverse reactions were noted.

Tra in . The method of trapping was modified several
times, until one was finally found which produced the best
results. Conventional government sparrow traps placed on
the ground were used at first--with very poor results.

Even when placed on tables, which were constructed and
arranged to prevent mammalian predation, these traps were
not adequate for trapping titmice, as they soon found their
way out of these funnel traps. As a possible remedy I
constructed Potter treadle traps and Chardonneret swinging

treadle traps. These also were relatively unsuccessful

20
.because many other woodland birds were trapped. A finely
woven bird trapping net, suspended between two trees so
that the birds would fly into it on their way to or from
the feeding table also proved fruitless.

I finally resorted to the use of hand-operated spring
traps, a method which was very time consuming but which
yielded the best results. The construction of the traps was
based on the pattern of the Higgins model, as described by
Lincoln (1947:35). A picture of one of them is presented
on Plate II, Fig. 18. They were made of one-quarter inch
square wire mesh netting and the dimensions were A6X56X18
centimeters. The large entrance doors were placed opposite
each other and a long elastic band, connected to the inside
of the doors, supplied the spring action. The doors were
released by pulling a string attached to a heavy piece of
wire, which extended over the top of the trap and held down
the trigger wires from each door. An observation blind was
unnecessary because I was able to conceal myself behind trees
and shrubs, some 50 to 75 feet away from the trap.

Feeding stations were maintained for two weeks pre-
ceding banding operations so that the birds would get
accustomed to coming to the trapping site. The feeding
stations were covered platforms suspended from trees.

Banding was carried out in each of the three campus

21

woodlots. As much consistency as possible was maintained

in the time of bending, usually early morning and late after-
noon. Birds were trapped on alternate days and were given
free access to the baited traps at other times. The traps
were visited just before dark to make sure no bird had

accidentally become entangled in the wire mesh.
The Study Area

The woodlots involved in this study are located at
Michigan State University, East Lansing, Township AN, Ingham
County, Michigan. The three areas are Known as the Sanford,
Red Cedar, and Baker woodlots. Their location and approxi—
mate size is indicated on the accompanying map (Fig. 5).

Sanford Woodlot. This is a SA-acre stand of relatively
mature deciduous trees, with Fagus ggandifolie and 5&3;
seccharum being the primary dominants. The secondary domi-

nants include Fraxinus americana, Tilia americana, guercus

 

rubra var. borealis, Ulmus americana, and Ulmus fulva.

 

The ground floor is gently rolling, with a gradual
drop in relief from the south side toward the river. Some
of the small depressions contain shallow water part of the
year.

The woodlot is the site of numerous activities,
including the harvesting of maple sugar. Many departments
of the university use it for class field trips and for

individual research.

m ohswfih
:mmflnOfla .m_wncmq .m .mpfimho>wab mumpm dmmHnOflS
.mooauooz wepmaoomm< can mamSmo

15$: 3 . o<om mac: P2302 Ham J v v ‘ :1H
\“s\\‘\\ 94m.m 2W8
\\\ . .
v ........a..... hm“. R: .52. _ SSW
3r). I
,\ if. \s ‘\
H .. \s\ Q
. n \2 to
\.. w \: \\:s¢sv~ft
. m \e w
OtCJEK< \ . ss [AL
u . #01800; .\ :Cttmsowt
«320 \x s‘s‘s‘ss—e
x m . wo._m.u. 2w¢0 zlts‘ww‘wq
FkK)? AICSK _ ss
02 :
fl. afimfiufiwfdflshfiztmub 1.1:...:Qrixxe.:Z:::::..211
.132:1..CZZZ.3. s::s\
~.4.-....a.v... .3 H \‘\\
4....d-.p....*...: W V \\‘\s .
4: 7o W U \‘fisx
H m it],
V m c. .,
w o
O 3 N
0
V U U
N W W
U oz<JSa<u ZwAO O a
m. _ EL, I-.- one“. 280

            

 

 

...()/-/. .o,..4

#93002,
. oxomz<m

32.0.:«5

353 Sow

 

milieu 1
._.---.i...!.-. . .mmzuyq zeeEnEPIllluaL

mumjol

0252/3 Fm<w muwzoI

. , 62.92.. hm<m

 

23

Red Cedar Woodlot. This 35.9 acre stand of relatively
mature deciduous trees is bisected by Kalamazoo Street.
The land is generally low, rising a little over ten feet
above water level. Several depressions contain shallow
water most of the year and during the spring intermittent
streams flow through the area.

The dominant trees are Ulmus americana and the soft
maples, Acer rubrum and Acer saccharinum. Secondary domi-
nants include Fagus grandifolia, Fraxinus americana, Quercus

rubra var. borealis, and the hard maples, Acer seccharum

 

and éggg nigrum.

This woodlot is maintained chiefly for class use and
research. Because selective cutting has not been practiced
in recent years there is a pronounced absence of small trees
and shrubs except in the openings.

Baker Woodlot. This 72-acre mixed deciduous associ-
ation contains such dominants as Fagus grandifolia, Acer
seccharum, Acer rubrum, guercus rubra var. borealis, and
Prunus serotina. Associates occurring less frequently are

Tilia americana, Ulmus americana, Fraxinus americana, and

 

Carya cordiformis.
The area is generally level but does contain three
shallow depressions which are filled with water most of the

year.

2h

Since this woodlot likewise had not been subjected
to selective cutting at the time of this study, there was
a general absence of shrubs and small trees. Although one
large concentration of Acer seccharum is used for maple
sugar harvesting the main use for this woodland is for
class field trips and research.

Climate and Weather. The records of the East Lansing
Weather Bureau (1941) supplied the following information on
climate and weather. I

The average0 temperature for the coldest month,
January, is 22. 9° F., while the average temperature
for the warmest month, July, is 71.1 F., with a
maximum of 1020 F. and a minimum of -26°F. The
average data of the last killing frost in spring is
may 5 and the average date of the first killing frost
in fall is October 10. These average frost dates
encompass a growing season of 158 days. The average
annual precipitation is 30.52 inches. Rainfall is
fairly evenly distributed throughout the growing
season with occasional thunderstorms in the summer.
Snowfall is moderate, rarely exceeding ten inches,
and the ground is seldom continously covered for
long periods. The prevailing winds are moderate
and westerly. The winter months are characterized
by much cloudiness, while spring has a variation of
cloudy and clear days, and the summer often has
many long periods of uninterrupted sunshine.

The first signs of spring may appear as early as late
February or early march, but spring does not usually become
established until the middle of March. This season is
characterized by high winds, cloudiness, and temperatures
ranging close to freezing. Both day and night temperatures
rise during spring, so that by the nesting period in Nay

and June the night readings range from hOO to 50°F., while

25
those during the day range from 60° to 85°F. A summary of
the weather conditions during the winter months and during
the months of nesting is given in Table III. Temperatures
and other data in the study area were assumed to be
comparable to those at the Lansing weather station.

The lopulation
at Beginning of Study

The Tufted Titmouse occupies the woodlots throughout
the year, but there is a seasonal as well as an annual
fluctuation in the population. Numbers are greatest and
most variable in late summer and fall. The fall population
in the three woodlots in 1953 was approximately 3h, as
compared with 18 during the following nesting season. Most
of the breeding birds were present in the fall flocks, but
many of the winter residents left the area in the spring,
one of them moving as far as three-quarters of a mile.
Similar seasonal shifts have been reported for the Black-
capped Chickadee, by Odum {19h1) at Rensselaerville, New
York; by Wallace (1941) at Lenox, Massachusetts; and by

Butts (1931) at Ithaca, New York.

Pair Formation

 

 

Time 33 Occurrence. Pair formation in most passerine
birds is usually preceded by the establishment of a breeding

territory by the male. My observations indicate that it

26

usually occurs earlier in the Tufted Titmouse. Only one
of the seven pairs, in which I was reasonably certain of
the time of pairing and of territory establishment, was
formed after territorialism was evident.

Pairing and the dispersal of the newly formed pairs
from the winter flocks occurred gradually during the period
from Larch 21 to April 12. In 1954 the first indication of
pair formation was evident on march 25 and the last semblance
of a flock (three apparently unmated birds) was seen on
April 10. During the spring of 1955 pairing first became
evident on March 21 and continued until April 12. After
this date the only birds seen were mated pairs or single
unnated wanderers. Prior to the dispersal of the winter
flocks there was movement of individuals from one group to
another and new birds occasionally appeared. Similar early
spring wandering has been reported for the Black—capped
Chickadee (Butts, 1931; Bowdish, 1938; Odum, 19hl).
Gillespie (1930) recorded the same behavior for the Tufted
Titmouse at Glenolden, Pennsylvania.

Courtship and Mating. During the period of pair

 

formation there was no evidence of courtship. The feeding
of the begging female by the male, which was the only
observed indication of courtship, did not occur until the
birds were examining possible nest sites. The male also fed

the female during nest construction, egg laying, and

27
incubation. Approximately two weeks elapsed between the
first Sign of definite pairing and courtship feeding.
Sixteen days elapsed between the formation of Fair 31 and
the first record of the begging-feeding behavior. It was
noted on two different occasions, occurring first while
the pair was examining possible nest sites.

Copulation was witnessed three times during egg laying.
Whether copulation occurred during pair formation as well
as after nest construction was not determined.

Behavior 9; Newly Paired Birds. In one case (Pair Sh)
the newly paired birds remained within the area where mating
occurred. Other birds were found nesting in areas quite
distant from the one in which pair formation took place.

In a few cases the birds were already constructing nests
or incubating before the pair was located so that the
relationship between the place of pairing and the place of
nesting could not be determined.

The newly paired birds remained closely associated
throughout the day. Their daily activities consisted of
feeding, resting, preening, and exploring for prospective
nest sites. During their movements through the woods the
birds usually maintained close contact with each other.

They frequently uttered a soft "tseep" note, which seemed to
act as a means of communication between them. When they

became separated the "tsicka-dee-dee" call was used to

28

locate the mate. On April 15, l95h, when a Red—shouldered
Hawk (Buteo lineatus) flew over Sanford Woodlot, male ln6205
emitted a loud, explosive "tsicka-dee-dee." This note
apparently acted as a warning to the other birds because
they suddenly became quiet and remained that way until the
hawk left. Vocal display and chasing resulted whenever
another titmouse approached a pair of titmice. Both of the
mated birds uttered the "peto, peto" notes, repeated in
rapid succession, and on three occasions when chasing ensued,
they both took part in the pursuit. After several minutes
of calling and chasing, the intruder departed and the pair
resumed its activities. This defense of a mate prior to the
establishment of a nesting territory has been referred to

by Noble (1939) as defense of "sexual territory."

Permanence g; Mating. In several species of Paridae
the mates maintain an attachment for each other beyond the
breeding season. Steinfatt (1938) found that harsh Tits
(Parus palustris) in East Prussia wander about in winter in
pairs and that the Crested Tit (Parug gristatus) and the
Willow Tit (Parus atricapillus) also are permanent residents
and mate permanently. In In years of banding Great Tits
(Parus major) at Budapest Werga (1939) did not record any
change of mates from season to season, and Kenrick (1940)
found that Blue Tits (Parus caeruleus) in England usually

retained the same mates.

29
In the North American Paridae, Price (1936) and

Dixon (l9h9) state that the Plain Titmouse (Parus inornatus)

 

in California is usually seen in pairs throughout the year
and keeps the same mate from year to year. Nice (1933)
reports a case of a pair of Carolina Chickadees (Pagus
carolihonsis which remained associated three winters and
two summers in Ohio. Baldwin (193h) also noted several
pairs of Carolina Chickadees which remained mated for two
or more seasons in Massachusetts. In the Tufted Titmouse
Middleton (l9h9) reports a male that was mated to one female
for three successive years and then to another female for
three more consecutive years. His observations were made
at a nest box in Norristown, Pennsylvania.

In a two-year study little can be determined regarding
permanency of mating, but male 1&6205 and female 146207 were
known to be mated for two successive years. In three other
banded pairs either the male or female was replaced by

another mate the following season.
Territory

Establishment. The Tufted Titmouse territory serves

 

as a nesting and feeding area for the adults and later as
a feeding place for their young. The newly meted birds
wander about for several days before claiming a territory.

Even though Pair 81 was established by April 8, there was

30
no indication of territorialty until April 25, after the
pair had been investigating possible nest sites. Territori-
alism was most evident at the beginning of nest construction
for other pairs as well. Perhaps the selection of a suita-
ble nest cavity induces the establishment of a territory.

It may or may not coincide with the area where pair formation
occurred or with the winter range. One member (146214) of
a flock moved three-quarters of a mile from the winter
range to nest.

Defense. Territorial defense was observed several times
during the nesting season. The male assumed the leading
role but he was occasionally assisted by the female.
Defensive behavior did not seem to be as elaborate as Nice
(1937) describes for the Song Sparrow. Of the five types
of behavior she attributes to the Song Sparrow, I noted
only the following two for the Tufted Titmouse. (l) The
challenge: When the territory was invaded the male challenged
the invader with loud "peto, peto" calls. The intruder
responded with similar notes but left the area upon the
approach of the defender. (2) The chase: In a few instances
the intruder did not retire immediately and a chase ensued.
On two occasions (Pairs 83 and R01) the defending male was
joined by his mate. While chasing the intruder from their

territory the mated birds scolded continuously. Actual

31
fighting was not observed and the only other belligerent
behavior that was noted was the increased erection of the
crest feathers during scolding.

The vocal activity of the mated birds dininished
noticeably after establishment of the territory. It is
this reduction in vocalization that makes the detection of
nesting birds difficult. The literature contains numerous
reports that the birds either "disappear" during nesting or
that they are extremely "shy." host of the singing during
the nesting period was done either by unmated birds or by
males having territorial disputes.

Defense of the territorial boundary against another
species was not recorded, and I witnessed a dispute with

another species only once. When a female Cowbird (Molothrus

 

agar) ventured clOse to the titmouse nest hole, she was
immediately chased by the male titmouse.

The territory was maintained until the young left the
nest, but it was defended most frequently during the early
part of the nesting period, when territories were being
established and the birds were still quite unsettled.
Defense was recorded once after hatching, when the young
were ten days old. The decline in aggressiveness after
hatching may be attributed to the fact that the males were

busy securing food for the young. Protection of the terri-

tory evidently becomes very weak or stops entirely during
the fledging period, as other titmouse families wandered
through occupied territories without challenge.

32
Size 2; Territory. The territories established by 1h
pairs of Tufted Titmice in the Sanford and Red Cedar Wood-

lots are indicated in Figures 6 and 7.

 

 

 

      

 

Scale: 1 inch=10 chains

 

 

 

/
Territories of the Tufted Titmouse in Red Cedar Woodlot
Figure 7
Legend
-- 195h
—-- 1955

o Nest, l95h
o Nest, 1955

33

The solid and broken lines, for 1955 and 1955 re-
spectively, outline the maximum area used and defended by
each pair during the early part of nesting. The numbers
indicate the pair which occupied the territory and the dots
and circles mark the location of the nest. Although the use
of lines gives the impression of definite limitations, rigid
boundaries were apparent only where territories came into
contact. At other points the territorial boundary seemed
to fluctuate, especially as the nesting season progressed.
Overlapping of territories was not apparent. The territories
were determined early in the breeding season, during the
last week of April and the first week of May. The singing
birds were heard from the same localities and pairs were
seen in the same area each day.

The size of the territories, calculated with a planime-
ter, ranged from 7.8 acres (3.2 hectares) to 13.9 acres
(5.6 hectares), with an average of 10.h acres (4.2 hectares).
These dimensions are comparable to those reported for other
Paridae. Dixon (l9h9) reported an average of 6.3 acres
(2.6 hectares) per pair for the Plain Titmouse at Berkeley,
California. Odum (1941) calculated an average of 13.2 acres
(5.3 hectares) per pair for the Black-capped Chickadee at
Rensselaerville, New York.

Although the diagrams indicate the size of the terri-

tories at the beginning of the nesting period, the defended

3h

area diminished as the nesting period progressed. During
nest building, egg laying, and incubation, the birds ranged
widely over their territory, but they limited themselves to
an area closer to the nest after the young had hatched.
The distance traveled for gathering food gradually decreased
with the development of the young. Butts (1931) and Odum
(19hl) noted this same behavior in the Black-capped Chickadee.

Habitat. The titmouse territories included two types
of habitat.“ The territory of lair RCA consisted of a stand
of young aspen, willow, ash, and maple. The territories of
all other pairs were located within mature (climax) stands
of beach, maple, elm, oak, and ash. Abandoned fields,
farmlands, and pine plantings adjacent to the woodlots were
utilized in winter but were not used appreciably during the
nesting season so that territories were essentially restricted
to deciduous woodlands. Titmouse territories extended verti-
cally into all of the various nesting societies--ground
nesting society, low shrub society, high shrub society, and
the upper canopy society.

Population Density. The number of pairs of titmice
in the Sanford and Red Cedar Woodlots averaged 7 pairs per
89 acres--adjusted to include areas immediately adjacent to
the woodlots that were parts of territories. This density

would mean one pair per 12.7 acres (5.1 hectares) of

35
suitable habitat—-excluding roads and open water. Comparing
the territorial boundaries with the vegetation in the wood-
lots indicated that little suitable habitat was left unoccu-
pied.

There is a close correlation between the density found
in this study and the densities reported by others. Twomey
(l9h5) found an average of 13.3 acres (5.4. hectares) per
pair in a two—year study of an elm-maple forest in east-
central Illinois. Williams (1936) calculated an average of
9.3 acres (3.7 hectares) per pair over a four-year period

in a beech-maple forest near Cleveland, Ohio.
The Nest

Nest-Site Selection. Nest-site selection began several

 

days after pair formation. In 1954 the birds were seen
examining cavities from April 1 to April 13, while in 1955,
due to a delayed spring, this activity extended from April 9
to April 20. A pair of titmice searched for possible nest
sites together but the female did most of the inspection of
the cavities while the male waited on a perch a few feet
away. The nests were located by watching either the move-
ments of both birds together or an individual carrying
building material. They were made either in old abandoned

woodpecker holes or in natural cavities.

36

Of the 20 nests in the study area, 9 were in American
elms; 10 in red maples; and one in a beech. Although the
nest sites were limited to these three trees, titmice were
seen inspecting prospective locations in other trees. The
height of the nest hole above ground ranged from 18 to 66
feet, with an average of 38.5 feet. The reported extremes
in nest height range from less than one foot (nest in a fence
post that extended one foot below ground level) to 97 feet
(Bent, 19h6). Cairns (1889) reported an average height of
35 feet, based on data from over one hundred nests.

In addition to the trees mentioned above, Tufted
Titmice have been reported nesting in the following: Pine;
cypress; white, red, blue, and scrub oaks; dogwood; birch;
apple; tupelo; sycamore; hemlock; mulberr‘; sugar maple;
yellow and black locust; white ash; and chestnut. Wayne
(1910) reported an unsuccessful attempt to build a nest in

Spanish moss (Tillandsia usheoides). In addition to such

 

natural sites the Tufted Titmouse sometimes uses bird houses
and abandoned woodpecker cavities in fence posts.

None of my banded titmice reused the nest site that
they occupied the previous year. McLaughlin (1888) reported
a similar lack of reuse, but Bent (l9uéz396) states that
"they will continue to use the same cavity for years, if

unmolested."

37

Nest Construction. The female did the actual nest
building, although both birds carried material. Even in
this latter activity the female was the more active of the
pair. Often the male simply accompanied his mate to and
from the nest hole. When the female entered the nest cavity
the male remained outside, calling softly or waiting in
silence. During the period of nest construction the female
often begged for food and was fed by the male.

Once started, nest building progressed rapidly until
the nest was completed. The birds were just as active in
the afternoon as in the morning. During the morning of the
second day of construction at Nest 81, the female, accompa-
nied by the male, made 8 trips to the nest in a period of
L0 minutes. This same pair made 20 trips in a two-hour
period during the afternoon of the third day of building.
Sometimes they flew directly to the nest while at other
times they lingered along the way.

The average length of time required for the construction
of seven nests was four days. All nests under study (with
the exception of a second nest cOmpleted on June 7) were
completed by the middle of May, the latest on May 16.

After the nesting season four nests were collected
for examination. All contained dried grass, moss, and hair;
two contained pieces of string, dried leaves, and strips of

bark; and one contained four white chicken feathers. While

38
nest composition presumably varied with the availability of
materials, there was a definite preference for dried grass,
green moss, hair and other soft materials. The hair was
from raccoon, opossum, dog, fox squirrel, and red squirrel.
Other sources of hair that have been reported are rabbit,
horse, cattle, cat, mice, woodchuck, and man.

The sizes and shapes of the nests were quite variable,
depending on the character of the cavity, which in one case
was so deep it necessitated a large amount of loose material
to fill the extra space. The depth of the cavities ranged
from 21.2 cm. to 27.5 cm. The depth of the nests ranged
from 13.7 cm. to 20.0 cm., while the outside diameter
ranged from 10.0 cm. to 12.5 cm. The inside diameter of the
hair-lined cup was difficult to measure because of its
irregularity but was approximately 7.5 cm. The depth of
the depression in which the eggs were laid averaged 3.9 cm.
The distance from the top of the nest to the entrance aver-
aged 8.a2 cm., and the diameter of the opening ranged from
h.4 cm. to 5.6 cm. The specific dimensions for each nest
are presented in Table V.

Nesting Associates. The 29 species of birds found
nesting in the study area are given in List II of the
Appendix. None of the titmouse nests contained Cowbird eggs,

even though Cowbirds were present in the area. Friedmann

(1938) reports five records of titmouse nests being
parasitized by Cowbirds.

39
Egg Laying

Deposition 9; Eggs. In this latitude the eggs of the

 

Tufted Titmouse are laid in hay. The dates of first eggs
ranged from May h to Lay 11 in this study. Information

from four nests indicated that egg laying occurred early in
the morning and that the eggs were laid at the rate of one

a day. The female partially or completely covered the first
few eggs with nest material Sets of 5 eggs were found in
Nests RC1, R02b, and 810, whereas Nest RC6 contained 6 eggs.
Bent (19h6) mentions that the number may vary from h-8, with
5 or 6 being the most common.

Description Q; Egg . The eggs examined were generally

 

ovate, with some having an elongate-ovate shape. The ground
color was pure white or creamy white and the surface was
dull and speckled. I noticed no variation in the color of
the spots on the eggs in a given set but there was variation
between sets. The specks on the eggs were hazel in Nest SlO;
dark hazel in Nest RCl; chestnut in Nest RC2b; and light
hazel in Nest R06. The distribution of the spots varied in
each set of eggs, being concentrated at the large end on 5
eggs and evenly distributed on 16 others. Egg measurements
were not attempted because of the difficulty in removing

the eggs. Bent (19A6) lists the average dimensions of 50

eggs as 18.4 by 14.1 mm. The extremes in length were

40
20.7 mm. and 16.8 mm., while the extremes in diameter were
Ibo? mm. and 1207 m.

Behavior During Egg Laying. The sexes remained closely

 

associated during the egg-laying period. A pair spent most
of its time ranging over the territory, carrying on the
usual daily activities of feeding, preening, and resting.
The female occasionally begged successfully but this
behavior was more evident during incubation. During the day
the birds spent little time near the nest, sometimes not
returning for three hours. The female roosted at night in
the nest hole, as she had done during nest building.
Incubation apparently did not begin until all the eggs in a
set were laid because hatching was more or less simultaneous.
Copulation was observed three times during the egg-
laying period, once for each of the following: Pair Sl -
9:30 A.M., May 3, 195A; lair R01 - 10:20 A.M., Way 5, 1954;
and Pair SlO - 2:10 P.M., May 8, 1956. The copulatory
behavior was similar on all three occasions, but the post-
copulatory actions of the male were most unusual in two
cases. The following is a summarized account of the
behavior recorded for fair R01 on May 5, 1954.
(10:15 A.M.) Female (146203) perched 15 feet up in
elm tree 10 feet away, twittering and vibrating wings
rapidly, head extended forward, crest depressed.
(10:17) Male (A194151) flew into same tree and
almost immediately flew to perch next to female.

Female continued to twitter and flutter wings.
Male attempted to mount female. Male resumed perch

41

next to mate, while she continued displaying. First
attempt apparently unsuccessful.

(10:20 A.N.) Male attempted mounting again. Female
flew away immediately after the act to a nearby tree
where she began preening and arranging her feathers,
no longer displaying. Apparently coyulation was
successful this time. hale remained perched on limb
where copulation had occurred. His body was yressed
close to the limb so that his legs were concealed.
Head and tail were extended in line with the body,
eyes gradually closed. He remained in this horizontal
sposition on the branch for a few seconds and then
began to tip slowly backward on the limb. His body
continued to rotate slowly backward and downward
until it was suspended beneath the limb. With eyes
still closed in this stuporous state the male then
fell toward the ground. After falling about eight
feet he suddenly revived and flew off into same
nearby bushes. He remained there resting ard greening
for a few seconds before flyinr into the woods out
of sight.

(10:23:30) Eenale flew off in same direction.

This same behavior was noted for the male (146205) of
fair 81 on may 3, 1954, but no such actions were displayed
during the c0pu1ation of lair 810 on Fay 8, 1956. Immedi-
ately after copulation the two birds flew into a nearby

tree and began greening.
Incubation

5912 g; §gx§§. During the incubation period a total
of 42 hours was spent at four nests. Incubation was
undoubtedly the duty of the female, for I never saw a male
stay in the nest cavity for more than a few seconds.
Incubation by the male at night was not observed and the

males examined did not reveal a brood patch. The time of

t2
the appearance of the fenale's brood patch was not deternined
but it was still present when the young left the nest.

Bent (1946) mentions that incubating and broodinr are
performed mainly by the fenale. Price (1936) and Dixon
(1949) state that incubation is the duty of the female in
the Plain Titmouse. Odum (l9tl) found no evidence that the
male Black—capped Chickadee takes part in incubation at any
time.

Behavior During Incubation. The period of incubation
was characterized by alternating periods of attentiveness
and inattentiveness. During the attentive periods the
female was fed by the male, who announced his approach to
the nest with a low whistled "tuee, tuee." On some occasions
the female was fed on the nest, while at other times her
mate perched near the nest hole and called the fenale off
the nest. Regardless of the place of feeding the male
announced his arrival with a call, corresponding to the
"signal song" used by the male Song Sparrow (Nice, 1937),
Hooded Warbler (Odum, 1931), Black-throated Green Warbler
(Pitelka, l9t0), Black-capped Chickadee (Odum, 19t1) and
others. Whether the female titmouse always responded vocally
to her mate's call was questionable, but on three occasions
she emitted a soft twitter before receiving food from him.

When fed in the nest cavity the female continued incubating

43
and did not come up to the opening. The male perched on
the edge of the hole and thrust his head down into the
cavity, leaving only the lower surface of his tail visible.
Usually the female remained on the nest if she had been
there only a few minutes, but if she had been incubatint
for several minutes, she left the nest just after beinp fed.

Two incubating females nade hissing sounds when
agproached. At other times they either renained quiet or
flushed immediately. This hissing defense also has been
reported for the Blue Titnouse and Great Titmouse
(Armstrong, 1947), and for the Black-capped Chickadee
(Odum, l9tl).

During the inattentive periods the ferale usually
begged for food in the presence of the male. He fed his
mate at intervals and she also fed herself. Cccasionally
the male accompanied the female to within a few feet of the
nest but usually she returned alone. During the return to
the nest, which was quite direct, neither bird emitted any
vocal sounds. Between his feeding visits to the nest the
male moved leisurely about the territory, feeding and
resting. He sang occasionally but generally singing
diminished during incubation.

Table IV summarizes data gained from 2t hours of
observation during incubation at four nests. The observ-

ations at a given nest were not made continuously but in

Ah

one to three-hour periods at different tines of the day
during the latter half of the incubation period. The
average attentive geriod was 25.8 minutes 3rd the average
inattentive Leriod was 8.9 minutes. The number of tines
the male fed the female on the nest during each attentive
period averaged 1.5. In on percent of the cases when the
female left the nest she did so in response to the call of
her mate. There was no pronounced difference in the
behavior of the birds at different times of the day.

These findings are similar to those recorded for other
Farinae. In the Plain Titmouse Dixon (19A9) recorded an
average of 8.5 minutes for the inattentive period and 28.5
minutes for the attentive period; Steinfatt (1938) recorded
a female harsh Tit averaging 37.09 minutes on and 7.9 minutes
off the nest during an all day observation; and Odum (19Ll)
recorded an average of 7.8 minutes for the inattentive
period and 24 minutes for the attentive period in the Black-
capped Chickadee.

Incubation Period. The incubation period was estab-

 

lished with reasonable accuracy in four nests. The time
between the laying of the last egg and its hatching averaged
13 days and 3 hours, as determined by marked eggs.

The incubation periods for other larinae are somewhat
longer than that determined for the Tufted Titmouse. Dixon

(l9h9) recorded 16 days for the Plain Titmouse; Odum (19hl)

A5

gives the incubation period of the Black—capped Chickadee
as 13 days; and Witherby, et al (1938) state that incubation

in the Coal-tit (Parus ater) requires from 14-16 days, while

 

it lasts 13 to 1h days in most other Eurogean Parinae.
Hatching. All 5 eggs in East RC1 hatched within 10
hours; a of the 5 eggs (one infertile) in Vest RC2b hatched
within 18 hours; and 5 of the 6 eggs (one infertile) in
Nest RC6 hatched within 20 hours. The last two records
included hours of darkness. The egg shells were carried a
considerable distance away from the nest and I could not see

whether they were eaten or singly discarded.
Care of the Young

Brooding and Feeding. The behavior of the adults

 

during the first week after hatching was quite sinilar to
that during incubation, but now the male directed his

feeding to the young instead of to his nate, and the female
fed the young when returning from her inattentive periods.
During the first few days after hatching the brooding periods
of the female were approximately of the same length as her
attentive periods during incubation but gradually decreased
as the development of the young Lrogressed. The male trans-

ferred food to the brooding female but whether she swallowed

46
it or fed the young could not be observed. As development
of the nestlings progressed the begging behavior of the

female subsided until both parents were directing their

feeding to the young.

Data on the feeding rate were recorded at three nests,
when the young were one, four, eight, and twelve days old

(Fig. 8 and Table VI).

 

 

 

 

 

 

 

 

 

 

12 . _“.Mi-ii_._iwr_ L._
‘2 i i l l
i t I
10 L l ”L- ..--. i. ,
a) 8 5 ‘ i
p . - a» --— —-‘ ¢ __~
'23 a
H
> 6 i- -
E" s
"C 1+ .. ‘ r t
g 0—“ %
fi‘ 2 «rn- ( f
I l
i i 1' i A h-
0 I? L 6 8 10 12 1h

Age (Days)

Average Hourly Rate of Feeding Nestlings
(based on data from three nests)

Figure 8
During the first four days the male fed the young
approximately twice as often as the female, who spent about
two-thirds of the time brooding. At 8 days the male still
fed more often than the female, but at 12 days the sexes

were feeding at about the same rate and the female was not

A7

brooding during the day. The number of feedings per hour
per pair increased from 8 when the young were one day old
to 17.7 when the nestlings were 12 days old.

The behavior pattern changed during the last few days
of nest life. The male no longer announced his arrival at
the nest and he discontinued feeding the female entirely,
both at the nest and away from it. Vocal activity diminished
and brooding by the female ceased almost entirely, excegt
during cold and wet periods and at night.

Nest Sanitation. Nest sanitation apreared to be the
duty of the male at first, but as nest life progressed he
was assisted by the female. The focal sacs were removed
after feeding the young and were carried a considerable
distance from the nest so that I was not able to determine
whether they were discarded or eaten. Dixon (19h9) mentions
that the male Plain Titmouse sometimes swallows the fecal
sacs.

Length of Nest Life. The young left the nest 15 days
and 17 hours after hatching (average of four nests). This
figure compares favorably with the 15 to 16-day Period
given by Bent (l9ho). The llain Titmouse remains for a
somewhat longer period, given as three weeks by Dixon (l9h9).

Odum (l9hl) recorded 16 days for the Black-capped Chickadee.

h8

Development of Nestlings

hestlings were removed from Nests RC2b and R06 at
regular intervals for examination. To facilitate removing
the young the entrance into the nest cavity was enlarged
with a small Keyhole saw and the removable yiece of wood then
held in place with a leather strap. Examination of the
nestlings was done as quickly as possible to minimize
disturbance. Consequently, information on the develoyment
of nestlings is limited to notes on external arpearance and
behavior.

External Appearance. The first day after hatching the
young were almost entirely naked, Link, blind, and helyless.
The mouth and abdomen were large and the limbs were small in
comparison to the rest of the body. Natal down ayyeared in
six small tufts, two each on the cagital, humeral, and spinal
tracts. The number of feathers per tuft varied but was less
than lb. On the fourth day the eyes were {artly open and the
feather tracts were more distinct. At eight days the eyes
were completely Open, the gray feathers of the juvenal
plumage were developing on the back, and the flanks were
tinged with rusty brown. The remiges were in the pin-feather
stage. By ten days the body was nearly covered with contour

feathers although apteria still showed and the remiges were

1+9
gartly sheathed. At 1h days the nestlings were well
feathered and closely resembled the adults, except for the
shorter grimaries and rectrices, and the yellow rictus of
the mouth.

Behavior. The behavior of the nestlings yaralleled
morphological and physiological changes. On the f‘rst dav
they responded to stimuli either by opening their mouths
for food or by squirming. Sense of fear was absent and did
not appear until the fourth day. From the age of eight days
the nestlings made hissing sounds, sonewhat similar to the
reaction of an incubating female vhen the nest was disturted.
The young were quite active at all ages, the amount of
activity increasing with age.

Number of Broods. One brood is usual for the Tufted
Titmouse. There were no second broods during this study
and the literature revealed only two records of double
broods, one from Ohio and one From Virginia. The other“
resident Paridae, the Black-capped Chickadee, occasionally
raises two broods. Steinfatt (1938) reforts that among
European Paridae the migratory species raise two broods and

the resident species one.

50

Postnest Life

Nest Leaving. Of the seven nests in which the tine of

 

departure was recorded, the young titmice left ftur of them
in the morning and three of them in the afternoon.
Apparently they do not leave until they are capable of

7
O

climbing up to the exit, at which time they are ready to fl;
Fortunately I was present when the brood left Nest

RCl--one of the two nests in which the young were handed.

From my vantage point I could see that the young were

becoming restless, repeatedly com'ny u; to the entrance hole.

At 9:15 A.h. on June 5, l95n, three of the young left the

nest in rapid succession. They flew into some bushes 50

feet from the nest. The other two departed at 10:35 an

joined the rest of the family. While the young were leaving

the nest both Larents flew about excitedly, frequently

emitting the soft "tsick-adee~dee" call note. Soon after

all the fledglings had left the nest the family began to

move through the trees away from the nesting site, with the

parent birds in the lead. While the young were being fed

by one parent the other parent flew to a new location about

50 feet away, and began calling softly, using the "peto"

sonc. he rest of the group then joined him. his procession

continued until 11:30 A.K., at which time the family was

75 yards from the nest cavity. During their movements

through the trees and while being fed, the young voiced an

51
anxious "seep, seep." The parents usually foraged a few I
feet away, although they occasionally traveled 50 yards
from. the young. At the end of their first day out of the
nest the fledglings were found with their perer'tts in a
bladdernut (Staphylea trifolia) thicket, close to the sgot
where I had last seen them that morning. I presumed that
the y roosted in the thicket, as they remained in it and
were quiet when darkness ended observations at 8:15 P.1Vf.
Dispersal of Young. On their second day out of Nest
RC]. the young titmice were quite inactive, remaining almost
Ste tionary in the foliage while we iting for their parents
to feed them. Feeding was occasionally interrupted by a
short flight to a new location. Five days after leaving the
nest the young were first seen feeding themselves. By June
26 the calls of the young closely resembled those of the
adults and by July 3, at six weeks of age, the fledglings
were definitely foraging for themselves and no beg-ging was
noted. At this time there were only four young, individual
1146212 having disappeared. This family of six was still
intact on July 10, but on August 7 two more of the young
(146211 and 11.6211.) had disappeared. One of them (ltéle)

was discovered almost a quarter of a mile away in a flock

of unbended titmice.

52
The postnest history of another family (Nest RCZb) is
summarized in the following paragraph.

July lO--Four young fledged at l P.M.

'July lh--Family seen at north end of territory.

July l7--Attempt to locate birds failed.

July 24--Parents and three young feeding in
northeastern part of territory, fourth
young (1&6218) not ;resent in study area
and was never seen again.

August lh-—Fenale parent and two of her young (lh62lé
and 1A6219) in flock with three unbanded
birds. This was the first indication of
a flock consisting of more than just a
family group.

October 9--Female parent and her two remaining young
came to the feeding shelf with two unbanded
birds.

Of the nine young banded from these two nests, only
two were present in the breeding area the next year. One
of these (1&6213) had replaced the lost mate of a banded
resident, while the other (lh6217) was mated to an unbanded
individual of unknown origin. One of the younr (1&6216)
from Nest RCZb nested three—quarters of a mile fron the
study area.

Van Tyne (1948) has suggested that family ties remain
Strong and that the family group forms the basis for fall
and winter flocks. The data gathered in this study support
this view since two of the fall flocks in 1954 contained

Parent birds and some of their young.

53

Nesting Mortality and Longevity

Nesting mortality. Because of the inaccessibility of
many of the nests, the information on nesting success is
based on four accessible ones. All of these were successful
in that at least one young left the nest. Nineteen fledglinhs
from 21 eggs was a nesting success of 90 percent.

Nice (1937), summarizing data on nesting success, gives
the average for hole-nesting passerines as 65 percent. In
nesting boxes in Holland 61 to 76 percent of the titmice eggs
produced young that left the nest. The higher :ercentage of
nesting success in this study may be due to the limited
number of nests.

Thus the 12 birds which were present at the beginning
of the breeding season (195h) in the Sanford and Red Cedar
Woodlots produced 28 young, a percentage increase of 233.
Since two adults were lost the fall population was not more
than 38 birds. This total, compared with the nunber present
at the beginning of the season, represents a three-fold
increase from April to August.

This summer increase is comparable to the average
found in a lh-year study of the House Wren (Kendeigh, 1937),
where the average number of adults nesting on 15 acres was
19.7 and the number of Young produced was 56, or a percentage

increase of 28A.

5h

Longevity. The longest age record revealed in this

 

study was at least 3 years and 11 months. This male hird
(Al9h151), banded by J. V. Lund on October 23, 1951, was
last seen on May 20, 1955. The oldest bird reported in the
literature was a male of at least 7 years and 4 months
(Middleton, l9h9). During his long life this unusual male
had three mates and raised A7 young in the same nest box.
The oldest Tufted Titmouse that nested in natural cavities
was one of 5 years and 11 months (Laskey, 19A3).

Predators. Mammalian predators {resent in the study

 

area were the Raccoon, Red Squirrel, and Fox Squirrel.
Avian predators were the Red-shouldered Hawk, Red-tailed
Hawk, Sharp-shinned Hawk, and Great Horned Owl. These,
however, are potential predators only, as I had no actual

records of loss from these animals.

Flocking

After the post-nuptial molt in July or August the
titmice assembled in small flocks and remained together
until pair formation separated them in late larch or early
April.

Formation. On August 14, l95h, my first flock that

 

was not strictly a family group was observed. It consisted
of the female from Fair RC2b, two of her young, and three

unbanded juveniles. On September A, 195A, another flock

55
in the same woodlot consisted of both parents from Fair RC ,
two of their young, and two unbanded birds. This eviderce
indicates that the family group ray form the nucleus of the
flock. These findings agree with those of Van Tyne (lQnP),
who mentions that young from differert nests continued to
accompany their parents to his feeding station from the time

of fledging until as late as January 10.

Size and Composition. The determination of the size

 

of a flock, as well as the recognition of its individuals,

was especially difficult in the woods. A flock crossing an

opening afforded the best opportunity for recording numbers

and composition. Two or more birds seen together were

considered a flock. The occasional single birds encountered

during the fall and wintcr are not included in the summariz-

ation of the data on the size of flocks presented in Table XI.
The large postnesting population in September consisted

of a greater number of flocks than during the following winter

months. The decrease in the population by January resulted

in fewer flocks but the average number of birds per flock

was only slightly smaller than it was during the fall months.

For example, in separate groups were recorded on September

25, l95h, while 5 were noted on January 8, 1955. The

average number of birds per flock was 9 and 6, respectively.

56

The behavior of SF} (flock number 3, Sanford Woodlot)
was typical of the instability of the titrice flocks. On
November 17, 1951., SFB consisted of six birds. Two days
later one of the individuals (11.6207) was noticed in
another flock, while the size of SF}. had increased to eight
birds with the addition of three new members. On l’ovember
22 the composition of the flock was age in different.
Individual 1L6207 had returned to it while two other members
(11.6205 and 11.6208) had joined other flocks, thereby reducing
the size of SF3 to seven birds. Similar shifts in other
groups indicated that the titmouse flocks were loose agpreg-
ations, varying, in both size and composition.

Associates. Other species were often seen accompanying

 

the titmouse flocks. If these other birds remained with the
titmice as the group moved from glace to {lace they were
considered members of the flock but if they did not renza in
with the titmice they were considered independent. Various
associations with the Tufted Titmouse were noted, although
they we re somewhat temporary.

In the fall, transient warblers and Ruby-crowned
Kinfilets formed temporary flocks with the titmice. In
Sei‘teznber the loose aggregations ‘of warblers and titmice
were comaonly seen feeding; in the tree toys. The Tufted
Titmice were the noisiest of the group ani were undoubtedly

the leeaders since the warblers followed them in their

57

movements. Liayfield (1937) relorts a similar observation

in Tennessee where Carolina Chickadees and titnice share in
the leadership of mixed autumnal flocks.

warblers most conm.only associated with the Tufted

‘.' urn lax: ,

were the leegnolia, Black-throated Green, Flue}:
Gil, “."Jt'l

Canada, Bleok-and-White, Bay-breasted, Chestnut-sir

the Redstart. Myrtle Warblers, which passed through after
the first wave of warblers, did not nix with the
~ The Ruby—

L).

flocks but remained in their own small groug
ember,

crowned Kinglets, passing through in October and Nov

replaced the warblers in the flocks of Tufted Titgice
their behavior was similar to that of the warblers.

During the winter months Black-capped Chickadees were

the most common and most permanent associates in the flocks

of'hnflmd Titmice but Hairy Woodgeckers, Downy Woodpeckers,

and White-breasted Nuthatches were seen in temporary

association with the titmice.

r H
.4
C0
0‘
Ho
cf
0:

Boosting

Since the roosting habits of the Tufted Titmouse were
slightly different in the winter than in the breeding; season,
they will be described sejarately.

Fall and Winter. During; the fall and winter months

most of the titmice roosted in tall evergreens or in cavities.

Three were found in cavities, two in old Woodpecker holes

58
and one in a decayed Knot-hole. lions of these cavities
was used for nesting the following year. By using a ladder
1 was able to look into the knot-hole without alarming the

sleeping, bird. It was in a squatting position on the floor
of the cavity and appeared to be a ball of gray fluff, with
its head thrust under the scapulars. The tail was raised
against the side of the cavity.

Bent (191.6) mentions that Dickey reported titmice
roosting in Woodpecker holes and natural cavities in
liennsylvania and West Virginia. Huey (1927) reported a
case of a titmouse roosting on a twig surrounded by the
leaves of a broadleaf tree. It is possible that such a

roost surrounded by dense foliage may have afforded conceal-

ment and shelter similar to a cavity.

The places of roosting reported for other Parinae

are about the same as those of the Tufted Titmouse. Odum
(191.2) mentions the use of natural cavities, old woodpecker
holes, and hemlock shelters by the Black-capped Chickadee.
Williams (191.1) found Chestnut-backed Chickadees (£293;
rufescens) roosting singly under caves of houses and in
vines, and quoted Bassett as reporting a Itountain Chickadee

\Parus gambeli) roosting in an old Robin's nest. Mogall

 

(1939) found that Great Tits and Blue Tits commonly roosted
singly in nesting boxes during the winter, and Dixon (191.9)

reported the use of cavities by the Plain Titmouse.

59

Breeding Season. With the breakup of the winter
flock and the pairing of birds in harch and April the
winter roosting sites were abandoned. Until the nesting
site was selected a pair seemed to sleep wherever it was
convenient. In one instance 1 noted that the birds of a
pair (SS) did not roost close together. The male (lt6223)
sought shelter in a spruce tree while his mate remained
active. She later went to roost in a spruce tree in the
same area but about 75 feet from the male.

Not until the nest was completed did the female begin
sleeping on it, but from that time until the young left the
nest she used it regularly. The male roosted in trees in
the general vicinity of the nest. After the young had
fledged the family group roosted wherever they happened to
be at sunset.

Tips 9; Boosting. During the fall and winter of
l954-‘55 the roosting time of a flock of titmice in Sanford
Woodlot was recorded. This was assumed to be the time when
the last call was heard, by which time most of the birds
were settled.

The fall and winter roosting times in relation to
sunset are summarized in Table I, page 60, while Table II
presents the roosting times recorded for Pairs SS and 39

during the nesting season of 1955. The "+" before a number

means minutes after sunset that the birds went to roost,
while the "-" indicates minutes before sunset.

Table I 60

Time of Roosting during Fall and Winter

Months
_ __ _,,_i___,,__ October' November ‘ December: January - February+
- J98£h9r__-
Clear
No. observ. 3 r l O 1 f 1 ‘fi
Average i+13.3 +10 --- -15 -19
Partly cloudy
go. observ. O 2 2 _ 1 1 _
Average --- +h +3 -l8 -25
Overcast or
snowing
.139: abserV- 2 1 2 1 _ 2 ,__
Average +3 -2 -8 -29 ; -36
Total No.
observ. . 5 A h l 3 r A a,
gverage ; -+9.2 - +8 -2.5 . -20.7 -29

—

Variation .+2 to -16 ;-2 to +10 .-10 to +aé-29 to 454-40 to -16

Table II
Time of Roasting during Nesting

9

____quate” ‘VMale E Female ; Condition _

,“mgpril l5 l -56 i :43_ . Pair S5, both in evergreens

_M_April 28 . -34 i -38 . Pair S9, bothin evergreens
May 7 ' ~30 ‘ -32 Pair SS, both in evergreens,

_i- -MA. a r L , nest construction

3119139.--. ; -2_5 . -35 , . Pair 85. easier-ins-

jmay 15 - ~42 ‘ -56 , Pair #59, incubation

_f_May r7 -20 -32 . Pair SS, incubation

1:3?! 20 ? -30 : Pair S9, incubation

Egg; -13 ‘ -3o , Pair as, incubation

:§_¥ay 30 _ 716 . ~22 7 ,Pair S9, feeding young

.fiJun9_5 .1717 n -20 Pair Siimfeeding youngwa“

HLLune 9. ‘? -l7 . Pair S9. feeding young .

‘f;June 15 .j!16 . -l8 __-__Pair_35, feeding young

* Female roosting in cavity, male elsewhere.

61

Although the number of observations was limited in
both sets of data, certain behavioral patterns are evident.
The birds usually went to roost earlier in January and
February than in October and Kovember. Birds roosted earlier
on overcast days than on partly cloudy or clear days (Fig. 9).
Individual birds tended to retire later before nesting began
than after. Once the female began using the nest cavity for

roosting she usually retired before her mate.

 

 

A 15 __.. , _. , ___. 1 . _ a it

p i {
m 10'
o n 3
2 8 i

1
o :3 i
p f
m Sunset
G .
'8
m 5
0)
3 10.
s
3
a f‘ 15
g I

‘J 20 Clear days
H o
o H
n o
5 ‘3 25” Partly cloudy
:3 on days
a 30- *
Q
:4
3. 35 Overcast or
¢ snowing
1+0 _._-._. . H ____ _, ______ 5.
Oct. Nov. Dec. Jan. Feb.

Boosting Time in Relation
to Sunset, l954-‘55.

Figure 9

62

The literature reports similar findings for several
other species. The seasonal variation in roosting time has

been rewarded for the European Tree Creeper (Certhia familiaris

 

britannica)(Rankin and Rankin, I940); Chestnut-backed
Chickadee and Bewick's Wren (Thryomanes bewicki)(Williams,
1941); and the Black-capped Chickadee (Odum, 1942).
Williams and Odum found a similar sex difference in the
roosting of Chestnut-backed and Black-capped Chickadees,
respectively--the female roosting earlier and rising later
than the male.

During the winter of l954-‘55 observations were made
on the time of leaving the roost in relation to sunrise.
The average of three records in November, l95h, was 6 minutes
before sunrise, while the average of three observations in
January, 1955, was 14 minutes before sunrise. My observ-
ations indicate that the Tufted Titmouse goes to roost early
and rises late. The continued activity of other woodland
birds after titmice had retired, and first singing in the
morning, before they had left their shelters, was also noted.
In November, 1954, they spent approximately 1h hours on the
roost; in January, 1955, about 15 hours. Long roosting
times have also been reported for other species of the

genus Parus.

63

Voice

The vocabulary of the Tufted Titmouse consists of a
great number of notes used in various combinations. The
following list includes only those which I have been able
to relate to a specific function, i.e., they were given in

a particular situation and frequently produced definite

observable responses.

 

"Peto" Song. This is a high-pitched, clearly whistled
phrase, usually repeated two to four times. It has various
translations, the most common of which are "peto, peto, peto"
and "peter, peter, peter." The accent falls on the first
syllable when the song is uttered slowly and on the last
when singing is rapid, but in either case the first syllable
of each phrase is higher in pitch. This song was heard in
all months of the year, becoming almost monotonous in the
spring. It was uttered by both sexes but more frequently
and more loudly by the male, especially during courtship
and defense of territory.

Location Note. This note was given by both sexes
throughout the year but most frequently during the fall and
winter flocking periods. It sounded like a dry, scratchy
"tsicka-dee-dee." This note seemed to be used for’re-estab-
lishing contact with other.members of the flock or for

re-establishing contact with the other member of a feeding

6h

pair. A similar note has been recorded for the Plain
Titmouse (Dixon, l9h9) and is similar to the "chick-a-dee-
dee-dee" of the Black-capped and Carolina Chickadees. When
this note is given by the Tufted Titmouse it is louder,
hoarser, and more slurred than it is in these other species.

Alarm Note. When alarmed at close quarters, such as
while feeding on the ground or during banding operations,
the birds gave a short, explosive "chit, chit" or "tsicka-
dee-dee." After dashing to cover or after being released
from the trap, this alarm note was often followed by the
harsh scolding note--a loud, rapidly given "tsicka-dee-dee-
dee," with the emphasis on the "dee" part of the note. This
expression was often given when my presence was detected or
when a predator was sighted. It was also given by titmice
in nearby trees during banding operations.

Recognition Note. While foraging together the members
of a pair frequently gave soft lisping notes which may be
transcribed as "tsip, tsip" or ”tseep, tseep." This note
was heard throughout the year from both sexes but it was
heard most frequently when a pair was feeding. They rarely
used this note during brooding or during exceptionally cold
weather when they were more stationary.

Hissing $233. This sound, sometimes accompanied by
puffing, is probably more the result of a forced expiration

than a true vocal utterance. It was given by incubating

65

and brooding females, and by the young in the nest. According

to Bent (l9h6) it is a characteristic defensive behavior in
several species of the genus Parus.

Invitational Note. On several occasions females gave

an invitational or begging display which consisted of the
soft, high-pitched, rapidly uttered notes "tsee, tsee, tees"
and the simultaneous quivering of her wings. On two occasions
this behavior preceded and accompanied copulation.

Begging Eggg. This note was quite similar to the
invitational note except that it was much louder. It was

given by the female when begging food from the male, and

also by the young from the time they left the nest until

they were independent. The begging note was accompanied

by wing quivering in both the adult female and the young.

Distregg Note. The shrill, high-pitched "see, see,
see” was given by adults and nestlings while they were being
banded. The distress notes usually elicited a scolding
response both from other titmice in the vicinity and from
the parents of the nestlings being banded.

any of the above notes of the Tufted Titmouse are not
described in the literature, but similar notes have been
reported for other members of the genus nggg. Odum (1942)
describes 16 different notes for the Black-capped Chickadee,

while Dixon (1949) describes 13 functionallyndifferent notes

for the southwestern Plain Titmouse. This great variation

66

in vocalization is characteristic of other strongly social
species. Davis (l9h0) lists 13 notes for the highly gregari-
ous Smooth-billed Ani (Crotophaga ani), and Erickson (1938)
lists more than a dozen notes for the Wren-tit (Chamaea
fascists). On the other hand, Hahn (1937) lists but five

call notes for the Ovenbird (Seiurus eurocapillus), a

 

generally non—social species.
Food and Feeding Habits

Feeding is one of the principal diurnal activities of
titmice, especially during the cold fall and winter months,
when nearly all their time is spent in seeking food.

EQQQ, According to Beal (1941) the food of the Tufted
Titmouse, as revealed by the analysis of 186 stomachs
obtained at all season, consists of 66.57 percent animal
matter and 33.43 percent vegetable matter. Table X summarizes
the results or the stomach analyses. One item, caterpillars,
formed more than half the animal food,and together with
wasps, formed more than half the total food. These were
torn into small pieces before being swallowed, thus making
indentification difficult. Coleoptera made up 7.06 percent
of the titmouse diet and more than two-thirds of these were

snout beetles or weevils. The cotton boll weevil was found

in four stomachs.

67

Beal mentions that ants were found only occasionally
but other hymenOpterous insects-—bees, wasps, and sawfly
larvae-~were eaten extensively (12.5 percent). Hemiptera,
principally pentatomids, and Homoptera (membracida and
coccids) were eaten to a moderate extent (b.03 percent) in
7 of the 12 months. A single fly was found in a stomach
collected in January. The only traces of orthopterous
insects (0.42 percent) were eggs of katydids, egg cases of
cockroaches, and a Jew and ovipositor thought to be those
of a grasshopper. Spiders and a few snails made up the
remainder of the animal food.

Beal's analysis of the vegetable matter consumed
indicated that fruit was eaten to a moderate extent (5.15
percent), mostly in midsummer, and included raspberries,
blackberries, strawberries, elderberries, hackberries,
blueberries, huckleberries, and mulberries. Seeds of
various kinds-~sumac, poison ivy, and bayberry--totaled
h.07 percent. Broken seeds and mast formed more than two-
thirds of the vegetable matter. Mast amounted to 23.h per—
cent of the total year-round food, comprising 95 percent
of the food eaten in November, SO-LZ percent in January,
and 55.97 percent in February. Mast was also the principal

form of vegetable matter consumed from August to March.

68

In this study specimens were not deliberately collected
for stomach analyses, but five adult Tufted Titmice, which
had been killed accidentally, were examined. The results
of these examinations are presented in Table IX.

In no instance was food found in any part of the
digestive tract except the gizzard. Because the food was
so finely divided a stomach was often recorded as containing
only one insect when it doubtless contained parts of more
than one.

Feeding Habits. Tufted Titmice were very versatile
in seeking food, foraging from the ground to the tops of
trees in a variety of habitats, although they generally fed
at low or at intermediate levels in trees and shrubs.
Several distinctly different habits associated with feeding
were recorded.

Most of the insect food eaten during the spring,
summer, and autumn months was taken from broadleaf and
coniferous foliage, mainly from the former. The birds often
tore open the cases of leaf-rolling larvae and on two

occasions the forest tent caterpillar (Malacosoma americana)

 

was taken. During the spring months opening buds were

frequently examined, probably for insects. Inspection of
the bark of limbs and twigs occurred throughout the year
but it was especially prevalent during the winter months.

Tufted Titmice were often seen hanging upside down at the

69

ends of twigs in their search for food. In addition to
gleaning small insects from the bark of twigs and limbs,

the birds captured moths from the tree trunks. The wings

of moths were removed and only the soft body eaten. Bent
(l9h6) reported titmice eating small white grubs contained
in an oak apple, and Trautman (l9h0) noted a titmouse eating
the pupal contents of a cecropia moth (Samia cecropia)
cocoon.

Seeds and fruit were eaten mostly in the fall and
winter months. On one occasion a small winter flock was
noted feeding in a hemlock on the campus. The birds hung
upside down on the cones while carefully removing their
seeds. ,Beechnuts were also consumed during the winter
months. No storing of naturally occuring seeds was observed,
but on November 20, l95h, one particular individual (146203)
stored at least 20 sunflower seeds in the bark of a black
cherry tree, located a short distance from a feeding station.
This storage of items from feeding stands is a commonly
observed practice. Titmice were seen eating the fruit of
the following trees, shrubs, and vines: choke-cherry, stag-
horn sumac, wild crab-apple, Virginia creeper, and flowering
dogwood.

Tufted Titmice spent considerable time searching for
food in the ground litter. After locating an edible item

they would carry it to a nearby perch before consuming it.

70
This habit prevailed also at the feeding tray where the

most common practice was to hold the seed against the perch
with their feet, while they deftly and quickly removed the
seed coat.

Hawking insects is apparently not a regular habit in
titmice. In the literature I found only one observation,
that of Dickey (Bent, 1946), of hawking a moth or butterfly.
I witnessed this behavior once. At hz30 P.M. on August 27,
1955, I saw a titmouse that was perched 15 feet from the
ground in the edge of a red maple tree suddenly fly straight
out about 8 feet to capture a cabbage butterfly (Pieris
gapgg). The bird returned immediately to its perch where
it removed the wings before consuming the soft body of the
insect. Although I waited for a possible repeat performance
the bird soon flew out of sight.

Tufted Titmice bathed and drank in the small woodland
pools. They also used artificial bird baths for drinking
and bathing. On five different occasions I watched titmice
drink water from melting snow on trees. While clinging to
the underside of a limb they reached up and drank drops of

water as they formed.

SUMMARY

A life-history study of the Tufted Titmouse, Egggg
bicolor, was the object of this investigation from September
1953 to September 1955. The study included a review of the
classification and nomenclature of this species, an analysis
of its past and present distribution, and its annual cycle
in.Michigan. The latter was the principal objective of the
research.

The Tufted Titmouse is a monotypic species of the
family Paridae and is indigenous to North America. The
scientific name, Parus bicolor, was established by Linnaeus
in 1758. The generic name was changed several times but
was re-established in 19Ah. The English name for the Latin
generic name, nggg, is "titmouse," and the specific name,
bicolor, means "of two colors."

The Tufted Titmouse is a non-migratory species whose
range covers most of the eastern half of the United States.
The political limits to the north are the southern portions

of the northern tier of states from.Minnesota to New York,

and southern New England. The southern limit extends along

the Gulf coast from Florida to the eastern half of Texas.
The western extension closely parallels the 99th meridian.
This distribution coincides very closely with the distrib-

ution of the eastern deciduous forests.

72

In Nichigan the northern limit of the breeding range
extends to a line running roughly from Muskegon across the
state through Mt. Pleasant and Midland to Port Huron.

During the past 25 years the Tufted Titmouse has been
gradually spreading northward in all the states along the
northern limits of its range. This extension may be due to
a number of factors. The gradual rise in temperature,
especially during the critical winter months, is suggested
as a possible contributing factor. Evidence of vegetational
succession is also presented, especially as it relates to
the availability of suitable nesting sites.

Studies on the annual cycle were conducted in three
woodlots at Michigan State University, East Lansing. The
woodlots consist of a mixed deciduous tree association,
tending toward the beech-maple climax stage of succession.

Numbered aluminum bands and colored aluminum bands were
used in various conbinations for the identification of adults
and young.

The hand-operated Higgins trap proved to be the most
successful of the several trapping methods attempted.

At the beginning of the study, in the fall of 1953,
there was a population of approximately 3h titmice in the
three study areas. This number dwindled to 18 by the spring
of 1954.

73
Pairing and the gradual break-up of the winter flocks

occurred during the period from March 25 to April 12. The
newly paired birds wandered about for several days before
investigating possible nest sites.

The establishment of territory, which seemed to coincide
with neat building, was principally for feeding and nesting.
Territories varied from 7.8 to 13.9 acres, averaging 10.A
acres (4.2 hectares). Defense was mainly the duty of the
male although the female sometimes assisted in banishing an
intruder. The size of territory and the vigor of its defense
decreased as nesting progressed.

The titmice did not excavate their nest sites but used
natural cavities or abandoned woodpecker holes. Nests were
found in elm, maple, and beach trees, but of course a much
greater variety of neat sites has been reported in the
literature, including bird boxes. The height of the 20 nests
ranged from 18 to 66 feet, with an average of 38.5 feet.

Nest construction was the duty of the female, although the
male accompanied his mate and occasionally carried building
material. Building of the nest averaged four days. All nests
were loosely constructed and all contained dried grass, moss,
and hair. Additional items in some nests were strips of bark,
dead leaves, string, and paper. The hair formed the shallow

cup in which the eggs were laid.

7A
The dates of first eggs ranged from May A to key 11 and

they were laid at the rate of one a day until the full comple-
ment of five or six was deposited. The eggs were white,
unevenly marked with brown specks. Copulation was observed
only during the egg-laying period.

Incubation was by the female. The male fed his mate
both on the nest and away from it. Attentive periods averaged
25.8 minutes, inattentive periods 8.9 minutes. The incubation
period in four nests averaged 13 days and 3 hours.

After hatching the young were fed by both parents but
mainly by the male during early nestling life. The male now
ignored the begging of his mate and concentrated on feeding
the young. The total number of feedings per hour by both
parents increased from 8, when the young were one day old,
to 17.7, when they were 12 days old. By the time the young
were ready to leave the nest they closely resembled the adults,
except for their fluffier plumage, shorter primaries and
rectrices, and the yellow rictus of the mouth. The young
left the nest 15 days and 17 hours after hatching (average
of four nests).

After fledging, the young followed their parents for
several days or longer. In some cases at least, the young
remained with their parents through the remainder of the
summer, so that the family formed the nucleus of fall flocks.
This was possible in this species since only one brood was

raised.

75

Nesting success in four nests was 90 percent and there
was a three-fold increase in the summer population. The
longest age record in this study was at least 3 years and
11 months.

The flocks of titmice and associated species began to
form in August. They assembled gradually and changed consider-
ably during the fall and winter months. Some of the flocks
were started by family groups, while others were mixed assoc-
iations including young from several nests. The size of the
flocks was quite variable but averaged seven birds. During
the fall and spring the size of the titmouse flocks increased
with the addition of migrants, especially warblers.

During the fall and winter the birds roosted wherever
they happened to be at sundown. Roosting was in conifers as
well as in deciduous trees and shrubs. The birds generally
went to roost much earlier, in relation to sunset, during
January and February than in October and November. They also
retired much earlier during the nesting season than they did
at other times of the year. Data on the time of leaving the
roost, in relation to sunrise, indicate that Tufted Titmice,
compared to most other species, go to roost early and rise
late. In November, 195L, approximately in hours was spent

on the roost, while in January, 1955. the time was extended

76
to 15 hours. During the breeding season the female slept
in the nest cavity, the male nearby. On days of inclement
weather and general cloudiness the birds went to roost earlier
than on days of fair weather.

The songs and calls of the Tufted Titmouse were many and

4 ."fl

varied. The most common song of "peto, peto" was given by

‘ .J‘ .1”.

both sexes throughout the year, but less frequently during
the nesting season. In addition to this song I discerned a
location note, recognition note, alarm note, hissing note,
invitational note, begging note, and distress note. Some of
these are basically similar but have distinct variations
associated with various types of behavior.

The food of the Tufted Titmouse, based on the analysis
of 186 stomachs by Beal (1941), consists of two-thirds animal
matter and one-third vegetable matter. The territory for
securing food extends vertically from the ground litter to
the uppermost twigs of the tallest trees. Most insect food
is gleaned from bark and leaves, but the hawking of a butterfly

was noted on one occasion.

APPENDIX

Table III

Average Weather Conditions at East Lansing, Michigan *
for the months from December to June, 1953- '5h and l95h-‘55

—_ __-. i

Wind

E Sunshine
(M. P. H. ) E(Per cent)

ITemperature uracipitation
(Degrees F.) (Total Inches)

Max. mMin. Av. WaterE Snow i Max.EAv.
1953 '77 ‘ j i E . E
December) 38 25. 231. 6 11.18 7.5 E 45 15.1: 7 29
1954 E . ' E
 January 3o 1 15.923.2Q 1.61 i 12.1 67 113.2} 30
_ February 38. 5 2h.331.h Eh.21 . 13.3 35 (14.2. #3
fmharchm 38 9 22.53o.7 '3.25 ' 13.8 50 '15.27 55
April 59 6 37.368.5 E2.75 i 0.1 62 313.8 LA
I May 364. 5 h1.553.0 :1.16 f 0.3 to 111.5 60
fIJune 80.3 59.269.8 Lu.o7 : o 51 E 8.51 g 66
December 33. 6 22.A28.0 2.71 10.9 17 5.3 26
19557777 If '
E—January 3O 6 18.52A.6 1.h7 ; 5.2 18 6.3. . hl__
February 36.1 19.026.6 1.89 13.A 16 5.8 51
:j16166 63. 5 25. 136.3 8.5 29 :_7.2“ 67
April E65. 7 63. 356. 5 11.56 T 23 6.7 78
- May E73. 0 19. 061. o '1.53“ o 18 . 5.2' ' 60
"86;;- E78 9 56. 667. 7 33.81 I" -0. E 22 1 6.0: _"59_.

 

Data from the U.S.Department of Commerce, Weather Bureau,
Local Climatological Data

 

 

cameos

m.¢~-m.~ .m.w
«wmwumwm,ta.m
m.4Nnm.N n.0H
m.mHnm.m b.m
H.oa-s.4 “.6

.um>713»uow¢
! - Amopzcwev

 

04

NH
0H

m
CH

.>nmmno

Ho
ponasz

vowpmm o>wpcmumeH

 

.zofipm9502H ho ham: Hoppoa mafipso mums mdowumpnomno *
1 .. Eam. W. E ;,,

 

5H
+~
b
m
m
whooom

G30
to: we

pmo: mo>mma m
mmawp mo

A-.., 1-- l

maamo

.m con:

 

l.u).nv! lllr.alll'|!-l.'1 IIIOI

umwwmsmm cmwmmmdocH waw

5.4-

 

III-II. il'll

 

:- 1|!!qu .

 

 

 

 

 

>H magma

[1"]: III".

.ng

 

 

 

_ euo m.H 1.~wrw1w.muia on- nausea

w . E . - :E E..- E: -H . -wEE..i. H 51.1“
m w muo 4.H 041m m.>~ 0H mm .
a W 5.0 o.H mm-o o.n~ 4H mm
m . muo a.H H6-0¢5.6~ a nmom
u mno H.H amam o.¢m m Hum

- -E-nmwwiz Eummm«zs..was1.uop< -
weapon oco mcwhuo A v mpnmmpo .02
% mfimohb mmuH—GHE HO
npwcoa uwmm
mesa» mo .02 nonadz
:-:1- ,, : - 1; aEztiail.

.oz coupon o>wpsopp<

a 1:1; _ s -E.;:.

 

Table V

Dimensions of Nest Cavities and Nests

 

‘erth

_Qiameter (smallest)

_._mleNe§t
Depth
__929er7
__1229rwcup

Edge of nest to
outer edge of

_,39Pran¢e
Diameter
mflQutside

H Inner cup

 

flNesthavity ,
9peging (diameter)

 

 

* Figures in centimeters.

h‘”sé£"
:Péir N0-
_fiHRCl
,, RCZb
, RC6
ATQtals
_Average

_jEELPaér i

40

QWNW

 

Table VI

V’Ager(Days)

a
l

o,

\lO‘fl

20
6.7
10

 

N-?'$‘
o,

N
mF'm-QO

 

$15.6 +16.1 ¥2o.o
3.9 ; n.a

8.a2i10.o

Hourly Rate of Feeding Nestlings

l
2 ! 6‘
6 j 11
6. i 8
5 ‘ 10
17 29
5.7 9.7

13.7

12

40

N
one-wow

17-7fi

.umom mo Aeneas on gamma mmhdmflm *

N 4- -1 _ o e 4 .4 .o -.i. : o o 4.5; -N. -.m--.....- mimw: :Mwmwfiaofla Bag
04 m we 4... m an m w m w w mm m m m a... «a 04 on 0 £8.82 .8...“ Sagas
is m ,m A... if; . 3a. 3. :3 m...m1a4miwzm -55.. z; 3:88 wodfl.»%§
. . ; menu on wraumwmml

 

 

 

 

 

 

zmmmsmz mmmzzzzmzzmmzmmzmm_ 323038333
33 «mm. . 3 233.8 33 :84 .33 83.2% «853% .8. *3:on 86% _omwwmm
...... .8 a: M x .m a x a x x N am a x aural : 32.8..mmwg
84 N x x a 38 3x888: Bo
hpa>ao mo ouspuz
, . _ saga mean new
..MNM. M N N . N M 55.3 same no "Savanna
2.3 xxxxxx x NMNMN x xx {9.3133
cons no chum
m ga. . N . . w maaomamcmpm msmwa
ofoflx . .x; AMEN WM.” ..;Mm - .1 ..MH NM” 4% ”NH- 5?. Tied—Bang-
.36 x N N. . .M N N N N . . _NW iwfimowpmam 2:5:
i W _ m omua mo mouoomm
mmwaewwwwwmammmmemwwxmggmw
at». e i4_n. 01,5 h.,t so 7V.L nu w W
m.mwflm_flw m._ m . “Mme“ i n
me u s _ . _ _ _ _ _
lm-mr@ «H _ w r W ~ -_; fl . w . ._ _ . l _ i P bi...
a m m z m

 

mmpfim pmmz no aofipmooq
HH> magma

Table VIII

Materials Used in Nest Building

 

 

 

N E S T
RClj R02a !R061 810
Plant ? E
Grass 60* 70 65 55
:f:fi§sg_ “‘23 7 16 '17 '20
Bark (Pogulus 3p.) -"l- : 2 j l
-_ Leaves 'Lw
Deciduous 3 2 3
:::§§y§d if“; i '?
Tot§l_u __ _83 . 88 .85 .80
Animal
__§§ir____ H ,, 7W7 , g ‘__16 10 14‘ H20__
_. Feathers _m____ . 1 M. _ .
Totelh_n%mw __.ml6 11 _1h _20_ ‘
Miscellaneous” _VY‘ _fi ‘ __W___ ' _; __
___§§zips~m _‘ . 1 l - _
Paper ‘ l

* Figures indicate percentage by volume relationships.

Table IX

Stomach Analyses of Five Tufted Titmice

p——_——'———l __. _._. ._. _.

 

 

Lf\
4
m
\\
H
\\
r-i
Animal

Insects

unwwwésgéaqatsie-fllarvae’ . 5
ColeOptera

___ Adul§§--_ 2

__. _Larvae ‘ 3

SP§§§{§_ ..

Miscellaneousg ‘
___H39Fe} . , a 10
Vegetable
__§9§9§_h_m_ _ . a .10 .
_WMast(incl. broken seeds) 60 ‘
_J@¥_muw_ue,_ .5.

Miscellaneous (leaves, bud 5

aqalaa. fruit 9919).
__m Total f f '80
Grit __IM.MIWIM_L}9-

 

w

Date of collection

b.)

10
10

3'87 ,
imm8_.

59'

3/10/'su

w

10

_30.

#3

#2

@12/30/'su

O\

1/8/'55

N

* Figures indicate percentage by volume relationships.

Table X

Summary of Food Items from Stomachs
of 186 Tufted Titmice (Baal, 1961)

. _.._.1..__,-‘_

 

Animal

 

. ._ Au”.- .7, 7.. 1/ .

lPercent

 

,_._____.-.W--_— ._ , .-

 

LepidOptera (caterpillars) ! 38.31

 

Hymenoptera (bees, wasps, 1 l2. 50

sawfly larvae)

ColeOptera (beetles)

Hemiptera

 

OrthOptera (eggs)

 

Misc. (snails, spiders)

 

Total (Animal)

 

 

       

 

Size of Flocks of Tufted Titmice

Numb

 

 

 

 

 

 

 

55

er of
Observations
_Xear .,'53<151 '54-'
Month ‘
Sept. 3h 30
Nov. 20 18
Jag: 4L_..16.6 151

'53-'54

lO
8

 

 

Average number of

birds per flock
V'““"‘”1 7"

-H_..._. _._., ... ..

'5h-‘55

) .__ __-_-....-.._. _

1

 

 

 

Vegetable Percent
11‘ 1 Fruit (Q11311111253E? ‘1
,and_cultiv.l1
. Seeds ' 6.07
1 7. O6 flMast ' 24.211 1
; b.03 i 1 1 1
.42” i
E _4駧_§ m - __ 1 . _-
2-66:4211_ _Vegeteble 31.11 _
Table II

Variation

in size

.w._ __r

'53 '511'54-'55_

 

2 12 2- 13
2-11 2-15
2,1§__)2-1h

“M"

List I

Non-Avian Vertebrate Associates of the Tufted Titmouse
(observed in the woodlots)

Mammals

Opossum, Didelphis virginiana
Masked Shrew,Sorex cinereus
Short-tailed Shrew, Blarina brevicauda

Raccoon, Proczon lotor

Long-taile easel, Mustela frenata

Striped Ground Squirrel, Citellus tridecemlineatus
Eastern Chipmunk, Tamias striatus

Red Squirrel, Tamiasciurus hudsonicus

Fox Squirrel, Sciurus niger

Prairie Deer Mouse, Peromyscus maniculatus bairdii
White-footed Mouse, Peromyscug leucogus

Meadow Vole, Microtus pennsxlvanicus

House Rat, Rattus norve icus
Cottontail, levilagus gloridanus
Amphibians

Spotted Salamander, Ambvstoma maculatum
Jefferson's Salamander, Ambggtoma jeTTersonianum
Red-backed Salamander, Plethodon cinereus

Tree Toad, Kyla versicolor

Swamp Cricket Frog, Pseudacris triseriata
Leopard FrOg, Rana pipiens

Eastern Wood FrOg, Rana svlvatica

Reptiles

Garter Snake, Thamnophis sirtalis
Painted Turtle, Chrvsemxs picta

 

 

 

 

 

 

.amoa ..em can .meaam emanated apnea “6 pmaauaoono .s.o.« *.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

,artsiiuv; _ .

M 1 . . . . . .msancnhw msqaonww

N 1 . . . . maoomoamm msmooonuqum

N . . . . .msmoaam> msqooonuqoa

H 1 1 . . . . . msfinm> msoamonhnmw

N _ 1 . . .msHmnqeoonnpmno momnosmaoa

. N 1 1 . . . . . msofiaommo mongoose

M 1 . . . . . . mspaeda modeHoo

_N 1 . . . . . .mmXon oamuoomwoz

H 1 1 . . . . .mfinpsaoo msnooHHnond

N 1 1 1 . . . . . .amwwsaom suspense

N 1 . 1 1 . . . . . . nocme mafiaoonono

N 1 1 1 _ . . . . . . msamficflmnfl> onsm

N 1 1 _ _ . . . . . monooHHoSw msuhoooo

1M 1 1 . . . . . ousonowa mpscquoN

N 1 , 1 . . . . . .QHAmHsoms MHpHpo¢

N 1 1 1 . . . . .msuouMoo> msmncmuono

N w 1 . . . . . msownoHoo mscmmmwmm

N 1 . . . . . mdpmfinpm nouamwoo<

M 1 . . . . . .mfimaooawamfi oopsm

x . . . . . . .mSpmocaa oopsm

M . . . . . . . ..mmmmmm.mm¢
”w w. N. no we “w
939939
MQPQQD.
T..T_.D.I.T..
mamas m

m a *oe6z caeapuoaom
pamcfimom pcmwmcmee

 

 

 

-me hunaupqom.qw mm.;wmnaopmmw.aonmoou< acupm

HH pmHA

 

 

map as eopnuuno menam.

) (( Ir ..1\ 1))1'():\ 11.4(IIII‘IIIIIII

enanwasm newsman
noxoomcooz mason
noxooqooog hawmm
Hoxosmmom codaaonnsoaaow
Hoxoomuoos coomonucom
nmxoomuooz uoaaaonucom
noMoaah cupumnmuaoaaow
“mamaquSM condom
unfinmqfiaasm espoopnpuhnam
passe mmqaano
samnpawaz aoaaoo

H30 voqaom wooed
ooxoso coaaanuSoaao»
upon mafiandoz
Headunwm ooppomm
Hoouaaam

pammmonm onooalwdam
xzom uoadfinmaqnmnm
xSom ooaampnuom

Mann wonocadonmnuom
Mona coo;

dn‘livl lrflllblli) (IOIIII‘) (Pll

oawz nosedo

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

1 1N 1 1 . . . . .ooasnowo mafipmomwoa

1 N 1 1 1 . . . . . . .maawfim wwammm

1 M _ _ 1 . . . . movemoomsm mHnomooawm

1 1H 1 1 1 . . . . . wpmppsm anomooamm

1 N 1 1 1 _ . . . . QGMHopmsE oaaowooamm

1 1M 1 1 . . . . . mSHnopmanm msvnde

_ 1M 1 1 . . . . . . Bayou oSOpmosoB

1 1H 1 1 . . . . .mmmcoamHonmo nHHepoadD

1 . 1 1 N 1 . . . mopvaHmopp mopw‘ won

1 _N 1 1 _ . . . . . cocoa moprOHmnMB

1 1 1 M . . . . . mwmaHHHamw manages

1 1 1M 1 1 1 . . . . . mamaoawHonoo mpvmm

1 1 N 1 1 . . . . . . .nowooan msnmm

1 1 1M 1 . . . . . msaawflmomnpo manna

1 1M 1 . . . . mommchnhnomun ms>noo

N 1 1 . . . . .opopmflno appaooqowm

N1 1 1 . . . . . . . mmnsm oqmowm

M1 1 1 1 . . . . .noHooHp oaoonocHHH

_ N 1 . . . . manpwomwm uawnmoaonm

1N 1 . . . . . . mcoufi> mmmopdoo

N 1 . . . . . . oncomm mmanommm

__ .1 N 1 11.. . . . 4332.11.12.13
m1 w. m1 mw mm M11
a S 9 Mn as a 1
M m“ m” “y ”H mu1

m w .w m. am pm 1 osoz 63333.6

3 1s 1
,1 1
poocfimwm pcoflmamne1

 

1eesaapqoo1 HH swag

 

 

Honopwopodc msuwuosam
unfinosam unopmsm
hnmo>

madame paauom

manage coo;

danom

nonmnnna asoam

unwnpwo

don; Henna:

don; omsom

Homoono QSonm
nopcnpsz woummwnncoawna
omsoaufis wounds
cocoxowno uoquonxooam
3on0 doaaoo

huh msam

navama magnum

Soaaesm cone

mama coanom

oozom woo; savanna
onmonm unwpmom
Hmnopmomam uopwmno nacho

lull llutll ll

 

mama dosaoo

 

 

 

 

 

 

 

 

 

 

 

 

1 N
N
N
1 N
1 N
_ _ 1 1 N
1 1 1 N .
11 11M
_ 1 1 1 N
1 1 1M 1 1
11111un
1 1 1 1 1 N
11111”
1 1 1 N
1 1.1"
1 N 1
N _
M 1
1 x
N
N
mmammw
939989
M 1. “v a n? D
TrTrpI.I
o u nu .} "u u
A A: B u my 9
a 9
a
pcowfimmm pcowmcmne
-q‘,-: III::1{1:I:::J:1;

 

 

ll ’3...

 

 

 

. . mocmpmmo

 

 

 

 

 

 

 

 

 

 

 

 

 

nomonuamn
.moacm>ammqmm wowoncaoa
. . . moms“ momonccma
. . ostnoo woaonnaoa
. . .mdonm> moaoaccmm
. . oumaopoo uofionvmon
.mcoommasnmmo moaonoqma
. . as aw u no cannon
. . meocmma mowoanoQ
. . manomnmm mafionuqoa
. . .wawOHaomm mwsnmm
. oaawmoomhsn wno>fiano>
. .ocwnmonum who>Hauu>
. unopmommnno mHo>HBnob

 

ofinmp apafipo as
. mspawm mmwww

 

. QSOUOG>HHO OOHH>

 

.maouMH>mHH oonwb
.mwhwwas> muchspm

 

. .adnonwmo «Hamomnaom

 

madeono msaswom

 

osaz owgapqoflom

. . ammupwm mdasmmm

‘al‘ lin<

Admscfipaoo1 HH pmfiq

noanumz copmmonnumwm
noanuwa covam-pnqpmmno
Reagan; amadndnxomam
Amannmz qmmasuoo
unannoz coonc gouachnpuaonam
“manna: oapnma

noannag osam vawonapnxomam
Meagan; hmE ammo

moannmz wadedwnfi

uoannaz Soaaow

Hoannws magnum

Meagan; oaaa>nmwz

noanuws ommmoanua

Hoannma commasadocaoo
Reagan: unansanamaxomam
00HH> mafiannog

oouw> cohonuom

oona> dopuonnpusoaao»
mafianopm

maazkma Hmooo

poawuHM cmasonoumnsm
noawnam gonzononQovaoo

 

 

oamz Goaaoo

 

, iii-1

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

1 1 N . . . . . . mHHmSQNm ooqsn ocean couoaoonopmam
x1 . . . . msocwscwm mopooooom Sonnmqm nommo>
1 _ N . . . msaamnpnmonnpmno 0H m 002309 covfimumsomsm
1 1 N . . . . . . mspmfinp manawm nocfiuvaoo nooanoad
1 N1 . . . . msonsawmw macawomuoo , :oafih oamadm
1 1 N . . . . . canowm mcmnommmm wuapadm owfiuaH
1x 1 1 . . . mscmmom>oosa msoaposonm xmonmono copmuonnnomom
1 _ _ N . . . maawufibnmo mqowcoano«m amuwcumo
1H 1 1 1 . . . . . coom>wao mwuaawm Amwmdma uoauoom
_ 1 N 1 1 . . lb . . . nope mannpoao: unansoo deaconuazonm
_ N1 1 . uoaooamnm> oasomwmd msawomfisd, oaxomno wounonm
. _ 1 x 1 . . . . . .mHanmm msuouoH cacauo ohoaapaum
M 1 _ 1 . . . . msoo amomm mswowumm ouwnxowam vmmaaascox
N 1 . . . . . .mcmua aaaoanspm mamasovooz.anopmmm
1 N _ . . . . . msompmmaov nommmm 1 zonnmqm omnom
1 x . . . . oHHMpran umwamapmm 1 uawpmoom amoflnma<
1 H _ . . . . mHmcommqmo wmcomawa “manna; «cameo
1 N . . . cfinmwowwwwmm mmqnonomb Reagan; waaansos
1 N . . . . .mwnomhp mfimNanoob puonnpsoaaow
1 N 1 . . . . mSHHHWdoomsm mandaum cnwnaopo
1 M _ . . . . .SSQmBHmm moHoubaoQ Meagan; Edam
1 M 1 . . . . . mpmappm moaoncqoa Amannmz HHOQMomam
J1 "a .a n1 nu .a
T. O O I O 9
A. s a 9 s o
M n... p. 9 1 D.
I. I. p I. I.
m ow cm m. cm mm 382 3:23 om 382 «38800
n a
pcovfimom pcmamcmne

 

 

 

 

1cmscfipcoov HH pmfiq

 

N . . . . . a cave oufimmOHoa sonnnmm muom

 

 

 

 

 

 

x . . . . . momma“ waaoammmmm zonnoam Nam
N . . . .mWHHmmmem mascahpoaow soanamm copuonnpnopdgz
N . . . .m a: Oode mfinownpoaom Bonnwam nongononopwnz
N . . . . . .mHHHmnm wHHoNHmW Sonammm caufih

 

. . . . . wcfinommmm maaouamw zonnamm maaquno

 

VI .‘ II. V ..II I . . II - . rltfil\l.l| I- ll.iil|l El rill-I-» .1 l.

r

Burnsaai

5

.LT Silluliir

A
.
I

Butqsag’
Euxpaag:

oawz owmfipcofiom camz noaaoo

ISAC MOIg!
'Sugpeeg
Bu:paaag§>¢

 

 

 

 

 

pcoufimmm ucmwmcmne

 

 

 

Awosuapdoov HH pmaq

PLATE I

Fig. A. Sanford Woodlot as it appears from the south,
beyond an Open field. June 18, 1955.

Fig. B. Sanford Woodlot, viewed from within. June 18,
1955.

PLATE I

.1
1

> «.- .'p‘.<~z.a;¢n-~ ..p

 

Fig. A

 

Fig. B

' PLATE II

Fig. A. Sanford Woodlot as it appears during the
winter. February 19, 1955.

Fig. B. Trapping Station. Note the large doors on the
sides of this Higgins trap. The written authority for the
operation of the banding station in the campus woodlots is
posted on the tree to the left. February 19, 1955.

PLATE II

 

Fig. A

 

Fig. B

PLATE III

Fig. A. Nest cavity of Pair $6 in Sanford Woodlot.
The nest site is located 57 feet from the ground in an
American elm tree. February 19, 1955.

Fig. B. Nest cavity of Pair RCZa in Red Cedar Woodlot.
Close-up view was taken through a 60mm. spotting scepe.
The cavity is 45 feet from the ground. February 19, 1955.

PLATE III

 

Fig. A

 

Fig. B

PLATE IV

Nest cavity of Pair Sh in Sanford Woodlot. The nest
site is located in a limb extending over Hagadorn Road.
Beal Pinetum is evident in the background. February 19,
1955-

LITERATURE CITED

Allen, Elsa G.
1951. The history of American ornitholOgy before
Audubon. Trans. Ameri. lhil. Soc., New Series,

41 (3)3387-591-

A.O.U. Check-list of North American Birds.

1886. First ed., Amer. Ornith. Union, Lancaster, Pa.
1895. Second ed., Amer. Ornith. Union, Lancaster, Pa.
1903. Second ed., twelfth supp. Auk, 20:357-359.
1908. Second ed., fourteenth supp. Auk, 25:386-387.
1910. Third ed., Amer. Ornith. Union, Lancaster, Pa.
1931. Fourth ed., Amer. Ornith. Union, Lancaster, Pa.
l9hh. Fourth ed., nineteenth supp. Auk, olzhhl-hoh.
1957. Fifth ed., Amer. Ornith. Union, Lancaster, Pa.

Armstrong, Edward A.
l9h7. Bird Display and Behavior. Oxford University
Press, New York.

Baird, Spencer F.
1858. Catalogue of North American birds. Smithonian
Misc. Collections.

Baird, S.F., T.M. Brewer and R. Ridgway.
1875. A history of North American birds. Little Brown
and Co., Boston, 1:88.

Baldwin, Dorothy A.
193h. Three returning mated pairs of chickadees.
Bird-Banding, 5:47.

Baten, W.D. and A.H. Eichmeier
1956. Is Michigan getting warmer? Mich. Agr. Exp. Sta.,
Quart. Bull. 39:97-101.

Beal, F.E.L.
l9hl. Common birds of southeastern United States in
relation to agriculture. U. S. Fish and Wildlife
Service, Washington, D. 0., Conserv. Bull. 15.

Bent. A.C.
l9h6. Life histories of North American Jays, Crows,
and Titmice. U. S. Natl. nus. Bull. 191:393-406.

Bowdish, B.S.
1938. Chickadee movement at Demarest, Y. C. Bird—Banding,

93159.

Butts, Wilbur K.
1931. A study of the Chickadee and White-breasted
Nuthatch by means of marked individuals. Bird-Banding,
2(1):l-26, 59-75.

Cairns, John S.
1889. The summer birds of Buncombe County, N. Carolina.
Ornith. and 001., lt(2):22.

Cook, Albert J.
1893. Birds of Lichigan. Lich. State Agric. College,
Bull. 94:126.

Dixon, Keith L.
l9h9. Behavior of the Plain Titmouse. Condor, 51(3):

Davis, D.E.
1940. Social nesting habits of the Smooth-billed Ani.

Erickson, Mary M.
1938. Territory, annual cycle, and numbers in a
population of Wren-tits (Chamaea fasciata). Univ.
of California lubl. in 3001., h3:257-333.

Friedmann, Herbert
1938. Additional Hosts of the larasitic Cowbirds.
Auk, 55(l)=h7.

Gillespie, Mabel
1930. Behavior and local distribution of the Tufted

Titmouse in winter and spring. Bird-Banding,
1:113-126.

Griscom, Ludlow
195A. Audubon Field Notes, 8(5):333.

Hann, Barry W.
1937. Life history of the Oven-bird in southern
Michigan. Wilson Bull., 49:1h5-2h0.

. Hellmayr, C.E.

1934. Catalogue of birds of the Americas and the
adjacent islands. Zoological Series, Field Yuseum of
Natural History, Publ. 330, Part VII, 13:70.

Huey, L.M.
1927. Where do birds spend the night? Wilson Bull.,

Kendeigh, 8.0. and S.P. Baldwin.
1937. Factors effecting yearly abundance of passerine
birds. Ecol. Monogr., 7:91-124.

Kenrick, H.
1940. A study of Blue Tits by colour ringing. British
Birds, 33:307-310.

Laskey, Amelia R.
1943. Some age records for banded birds. The Migrant,

14(1):6.

Lincoln, Frederick C.
1947. Manual for bird banders. U. 8. Fish and Wildlife
Service, Washington, D. C.

Mayfield, George R.
1937. Observations on mixed flocks in autumn. The
Migrant, 8:47-49.

McLaughlin, R.B.
1888. Nestin of the Tufted Titmouse. Ornith. and
001., 3(4 :61.

Middleton, Raymond J.

1949. Tufted Titmouse nesting seven years. Bird-Banding,
20(3):151-152.

Mogall, Karl
1939. Beobachtungen an dem.Meisenbestand der Revier-
forsterei Breach. Vogelring, 11:10-40 (review, Bird-
Banding, 10:133).

Nice, Margaret M.
1933. Robins and Carolina Chickadees remating. Bird-
Banding, 4:157.

1937. Studies in the life history of the Song Sparrow.
1. Trans. Linn. Soc. New York, 4:1-247.

Noble, G.K.
1939. The role of dominance in the social life of birds.
Auk, 56:263-273.

Odum, Eugene P.
1931. Notes on the nesting habits of the Hooded Warbler.
Wilson Bull., 43:316-317.

1941. Annual cycle of the Black-capped Chickadee.
Auk. 58(3)=314-333; 58(4)=518-535.

1942. Annual cycle of the Black-capped Chickadee.
AUX, 59(4):499-531-

Parmalee, George W.

1947. Postglacial forest succession in the Lansing area
of Michigan: a study of pollen spectra. Unpublished
haster's thesis, Dept. of Botany, vichigan State
University, 1-70.

Pitelka, Frank A.
1940. Breeding behavior of the Black-throated Green
Warbler. Wilson Bull., 52:3-18.

1941. Distribution of birds in relation to major biotic
communities. Am. kid. Nat., 25:113-137.

Price, J.B.
1936. The family relations of the Plain Titmouse.
Condor, 38:23-28.

Rankin, M.N. and D.H. Rankin
1940. Additional notes on the roosting habits of Tree
Creepers. British Birds, 34:56-60.

Ridgway, Robert
1904. The birds of North and Middle America. U. S.
National Museum, Bull. 50, Part III:308.

Steinfatt, Otto
1938. Das Brutleben der Sumpfmeise und einige Vergleiche
mit dem Brutleben der anderen einheimischen Heisen.
Beitrage z. d. Fortplanzungsbiol. d. Vogel, 14:84-89,
137-144.

Tinker, A.D.
1908. Breeding of the Tufted Titmouse in Nashtenaw
County, Michigan. Auk, 25:322-323.

Trautman, Milton B.
1940. The birds of Buckeye Lake, Ohio. Univ. of Mich.
Mus. Z001. Misc. Publ. 44:1-466.

Twomey, Arthur C.
1945. The bird population of an elm-maple forest with
special reference to aspection, territorialism, and
coactions. Ecol. Monogr., 15(2):172-205.

Van Tyne, Josselyn
1948. Home range and furation of family ties in the
Tufted Titmouse. Wilson Bull., 60(2):121.

Wallace, George J.
1941. Winter studies of color-banded chickadees.
Bird-Banding, 12:49-67.

Warga, Koloman
1939. Dauer-Ehen bei Parus mt mayor and Jahres-Ehen
bei Phoenicurus p. phoenicurus. Journ. f. Ornith.,
87:54-60.

Wayne, Arthur T.
1910. Birds of South Carolina. Charleston Museum,
Charleston, S. C.

Wil1iams, A.B.
1936. The composition and dynamics of a beech-maple
climax community. Ecol. Monogr., 6:317-408.

Williams, Laidlaw
1941. Roosting habits of the Chestnut-backed Chickadee
and the Bewick Wren. Condor, 43:274-285.

Witherby,H.F.,F.C.R.Jourdain,N.F.Ticehurst and B.W.Tucker
1938. The handbook of British birds. London. 1:245.

WOOd, NOAO
1921. Some southern Michigan bird records. Auk, 38(4):594.

1951. The birds of Michigan. Univ. of Mich. Mus. of
2001., Misc. PUbl. l7;l-5590

"Illlllllllll’lllr