Citation
Ecology and behavior of the Florida Scrub Jay

Material Information

Title:
Ecology and behavior of the Florida Scrub Jay
Creator:
Westcott, Peter Walter, 1938-
Publication Date:
Language:
English
Physical Description:
viii, 85 leaves. : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Birds -- Florida ( lcsh )
Birds -- Behavior ( lcsh )
Dissertations, Academic -- Zoology -- UF
Scrub Jay ( lcsh )
Zoology thesis Ph. D
Greater Orlando ( local )
Birds ( jstor )
Thickets ( jstor )
Bird nesting ( jstor )
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis--University of Florida, 1970.
Bibliography:
Bibliography: leaves 83-84.
General Note:
Manuscript copy.
General Note:
Vita.

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University of Florida
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University of Florida
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This item is presumed in the public domain according to the terms of the Retrospective Dissertation Scanning (RDS) policy, which may be viewed at http://ufdc.ufl.edu/AA00007596/00001. The University of Florida George A. Smathers Libraries respect the intellectual property rights of others and do not claim any copyright interest in this item. Users of this work have responsibility for determining copyright status prior to reusing, publishing or reproducing this item for purposes other than what is allowed by fair use or other copyright exemptions. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder. The Smathers Libraries would like to learn more about this item and invite individuals or organizations to contact the RDS coordinator(ufdissertations@uflib.ufl.edu) with any additional information they can provide.
Resource Identifier:
021880081 ( ALEPH )
13385442 ( OCLC )

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Full Text










ECOLOGY AND BEHAVIOR OF THE

FLORIDA SCRUB JAY
















By

PETER WALTER WESTCOTI


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












UNIVERSITY OF FLORIDA


1970

















I wish to extend Thanks to the fOllowing persons and organizations who aided me in various ,:ays in carryira out the present work. Professors Pierce 3rcdkorb, Archie F. Carr, Jr., Albert M. Laessle, David W. Johnston, and Ernest S. Ford of the University of Florida acted aZ my supervisory committee while I was a student at the University of Florida. Professor 3rodkorb was my principal advisor, and I especially wish to acknowledge his tirel.ess efforts in the preparation of this. manuscript. Dr, Oliver L. Austin, Jr., Curator, Florida State M.XLseum, -rovidsd

both banding material and suggestions during the course of this study. Dr. James N. Layne of the Archbo.d Biological Station and Dr. Glen Woolfend, nt o:: South Florida, provided much advice and encouragement. Mr. Donald E. Goodman greatly aided in the tiresome enterprise of ca-curing birds. am mindful of the invaluable grants-in-aid received from the Frank M. Chapman "I7oriDl Fund of the kmerican Museum o: Natur-al History and from Sigma li.





















ACKNG L D IENTS . . . . . . . ,
LI-ST OF FIL'ES
L S T 0 F G..- -'' . I . . . . .

ABSTRACT . . . . . . . . .

INTRRODUJCUTT. . . . . . . . . .

HICORIA STUDY AREA. . . . . . . .

METHODS . . . . . . . . . .

RECOGNITION, SIGNALS . . . . . . . .

Auditory Signa.l.s . . . . . . .

Visual Signals . . . . . . .

,. r..S Ip.. . . . . . . . . . .

Pair Formatio .. . . . . . . .

Cot ulator e v r . . . . TERRITORIALITY . . . . . . .

Nest Placi2e ..t . . . . . . .

Territorial Zefense . . . .

Site Tenacitv and Mate Consta'.....

"del ers" . . . . . . . .

I: te.ra.:t'n s -,,7;,7

Rol of. "eCerS"..... . . . .



REACTTOU TO OTH spc .


ilL


Page Vi VL .vi









S10 10



. 14 14 16 18 18





28



33

. 39

-1








Page
ReactiDn to Predtors . . . . . 42

Reaction to Man . . . . . . 45

DISPERSAL . . . . . . . . 47

HABITAT AND DISTR3 -CT- . . . . . . 52

Scrub Jay Haitat in Ficrida . . . . 52

Hicoria . . . . . . . . 55

CaPe Kennedy Air Force station.......

Jonathan Dickinson State Park . . 59 Mt. Plvmouth Area . . . . . 59

Lake Jackson . . . . . . .63

Ridge Scrub . . . . . . . 0

Scrubby Flatwoods West of iicoria 61

Atlantic Coast Dune Thickets .... 64 ij Scrub" . . . . . . . 7

Ocala National Fores.. . . . . 6 7

Habitat Summar7 . . . . . . 68

Szrub H-abitat Without EreEcin scrub Ja.s. 69

Kissi.ee Saint Cloud . . . . 69 Weeki Wachee . . . . . . 72

Lake Wales . . . . . . . 72

Florida Panhandle Scrub . . . . 73 FACTORS CONTRLLING DI STRIBTil i7....... ,.76 LITEPATUE CTD........... . . . . .' 3

BIOGRAPHICAL SKETCH . . . . . . . 35


iv

















Figure Page

1. ,'ap of study area at H-Lcoria 1968 . . 6 2. M4ap of study area at Hicoria 1969 . . 20 3. Mar of study area at Hicoria 1.970 ..... 22 4. View of Hicoria study area . . . . 57

5. View of scrub thicket tus.o miles southeast of Hicoria . . . . . . 63

6. View of Atlantic coast thicket at
Marineland . . . . . . . 66

7. Map of central Florida peninsuLa with
Scrub Jay habitat . . . . . . 71

8. View of scrub thicke-, at 'ort Pickens
State Park . . . . . . . 75


*1







A- h ra
of the University cf Florida in :art-ial Fulfillment of the Requirements for the Degree of Dccor of Philosoph'


ECOLOGY AYD B2IAVIOR OF TE Z FLORDA SCRUB JAY



Peter alter ,estcott

August, 1370

Chairman: Prcfessor Pierce ?rodkorb Major Department: Department of Zoology


This study gives an outline of the social structuring of breeding Florida Scrub Jays (Aphelocoma C. c.erulesces: and discusses the restrictions replace. on distribution by the type of habitat selected for nesting. The sccial structure was studied at Hicoria, Highlands County, yloida, where 230 individuals were banded.

Certain calls and postures used in social interactions are described, and pair formation and coculatory behavior are discussed.

In Florida the Scrub Jay is territorial and pairs for life. The territory: is d endeI fcm' intrusion b> o hsr Scrut Jays, but firsZt-year birds (young of that pair -7ro, the year before) are retained in the territcry. They occasionally act as "helCers" caring for and :requently aid in territorial defense. Although they restrict their movements to the territory, usually these extra Tr.cr renaina-:a: fr-. The breeZng I-r. 2-. bond, similar to a Dair o "nd, may be present betw.;-en the hreed








male and his helperss" since courtship f ding occurs between them. Territories -ithout "helpers" usually are those in which the nest or fledglings were destroyed the year before. Parely were first-year birds found away from parenrza t-rritories.

Dispersal by Florida Scrub Jays is rearkedly restricted. Movement of first-vear birds is d.2laved until the second year. The retention of young birds irn territories may allow for a form of site tenacity, so thaL when these "helpers" disperse it is only for short dictanczes, The farthest any banded bird moved to establish a territory was 1.8 miles. No birds have moved between Hicoria and the Archbold Biological Station, 2..0 miles north, where several hundred Scrub Jays have been banded by another worker.

The reaction of Scrub Jays to other species and to

predators is described. Whenever Scrub Ja-s and Blue Jays (Cvanocitta cristata) interact, the Scrub Ja%; acminates.

Florida Scrub Jays are restricted to a plant community that combine' 7cl dense thickets witi numerous oren sany spaces. Such habitat occurs primarily at the edge of sand pine scrub, in recently burned scrub, as scrubby flatwoods, and as shore dune thickets. 7o:, dense scrub thickets frequently do not include open sandy ipaces and sc are a : favored by iestni. ays, cu na:-maie 0cenings in S habitat have increased the amount of suitable nesting habitar, i.e., alonc roa&'ay2 and 7eg- ; o:'velc:sens.


vii









Even so, the total ars: available to Scrub Jays for nesting is small. This .bitat frequently exists as small areas of thickets isolated from similar suitable habitat.

The distribution of breeding Scrub Jays in Florida is outlined. Only a few large exoanses of suitable habitat suoport dense populations of Scrub Jays. Although dispersal distances are not great, most small isolated patches of scrub habitat are occupied by Scrub Jays when in the vicinity of these centers of population, but seemingly suitable habitat isolated by 20 or 30 miles from the nearest Scrub Jay population is occasionally unoccupied. The Florida Panhandle has a large stretch of sand pine scrub community and much of it appears ideal as Scrub Jay nesting habitat. No Scrub Jays inhabit this extensive habitat which is separated by 100 miles from the closest peninsular Scrub Jay population.

The retention of first-year birds in parental territories and the reduced dispersal abilities .of nonbreeding adults are discussed as a controlling factor in the distribution of the Florida Scrub Jay. The sociality of the breeding jays may be a behaviorial adaptation to a limited breeding habitat similar to the case of the :exican Jay ultrararina) in Arizona.


viii

















The Scrub Jay, Aphalocoma coerulescens (Boso), occurs in two se'zarate areas of North America. Throughout the western United States and the mountains of Mexico this species lives in a variety of habitats, often in large numbers (Pitelka, 1951; Wesrcctt, in press). Although the western population is composed of sixteen subspecies, all appear to have similar social organizations. -The single eastern subspecies, A. c. coerulescens, inha.its central Florida. Not only is i isolated by over a thousand miles from the closest .e.tern population but it also exhibits completely anomalous patterns of socialit., Unlike its western representatives, the Florida Scrub Jay is restricted in habitat and is often quite scarce. Western Scr-ub Jays are normally secretive, yet Florida Scrub Jays are notably tame.

Reports or extra individuals acting as "helcers' about the nest (Grimes, 1940) also suggest a degree of sociality unlike that of western Scrub Jays, zut little information n-a2 1:eared co::cerning this example of

sociality in the :2crida Scrub .Jay. Amadon (1944) examined the act ii s oZ treading pairs about :>r nest and presents details of courshin, nest buildings, and care


1.




2


of the young. Altiroc:h '- occasionally observed nonbreeding birds around active nests, Amadon concIluded that they do little to aid in the zare of young.

Although Florida Scrub Jays are found scattered over much of central :lorida, most local populations appear small, and some areas, such as in the Florida Panhandle, lock ideal for nesting Scrub Jays but none have been reported. The study reported here gives a detailed outline of the social structuring of a breeding group of Florida Scrub Jays and analyzes the social behavior of these birds in relation to dispersal and habitat distribution.

















The stud. area at Hicoria is located two miles due south of the Archbold Biological Station, Lake Placid, Highlands County, Florida, and is at the extreme southern end of the sand ridge region of central Florida. At an elevation of 140 feet above sea level, this lccale contains a plant community that combines sand pine scrub and Pine flatw-:ood elements; its special vegetational colponents are termed scrubby flatwoods by Laessle (1942). The vegetation is discussed in detail in a laer section.

A dominant feature of this particular site is a

multitude of large and small open sandy spces. As discussed later, this con-bination of iense scrub thickets and open space represents the optimum habitat for 'lorida Scrub Jays and gives a density of breeding birds at Hicoria that facilitate the studies of social behavior.

The oven a f this site are not typical of -che

scrubby cover .hich elsewhere is extremely dense and extensive. uring the early years of this century H-icoria was the site of a to.:n of one thcuc;and 7erSons, a glass factor, and a lu.-.-ber i1. Todey the only remains are the scattered c en s aes )here houses cr other buildings once stood. The outline of the study area in Fia. 1 dtn f t tn and atr

don:ment of 4the to--wn and frartcrzi__s.


3









To the eat. thns site is separated froa. the main

body of the sa:. ridge scrub by extensive orange groves. The western edge of the tudy area is a1 the border of the sand ridge and drops several feet to pine flatwoods, although many scrub thickets continue to the west. Scrubby vegetation extends south for a mile and then merges into pine flatwoods with widely scattered scrub thickets. The scrub at Hicoria continues north along a railroad in an unbroken, although narrow, band to the Archbold Biological Station.

Little human interference occurs at Hicoria. Occasionally a hunter stops to shoot at a rabbit, someone cuts some hickory for barbecue wood, or checks on the cattle run on the extreme north edge of this area. Otherwise 1 saw few people during my three years of field work at Hicoria. Although Florida Scrub Jays are notoriously tame and in many areas readily follow people to beg for food, at Hicoria the birds usually ignored my presence. I regularly camped in the middle of the study area, but took pains not to leave my food about.













Figure I Lap of sLudy area at Hicoria showing territory boundaries and Scrub Jays residing in each territory during the breeding season
1968. Note: in territory 12 SY-Y died and
S-BY moved to territory 4.

KEY

1 inch = 300 feet


Shaded area = scrubby thickets Light area = open sandy spaces


paved road dirt road


railroad


fence


territory boundary


Letters designate a speci fic Scrub Jay, i.e.,
RS-R


= breeding male


2 = breeding female


= "Helpers"

All birds without a special sign following their letter designation are young of the year, i.e., lG-YS in territory 2.










sj -Os


/rsf









i '




rnn



4- t''* I
K --N'S."""

1s '7"'6 c
"''N .1119


/ H'7jfl tI
f-----t--t--s-- ssI
ii I I
II

ji Wt 4"' "'4"

















The social ehavicr is based upon information gathered from the banc.ina of 230 Scrub Jays at Hicoria and at scattered sites to the west and south. To simplify the report discussion is limited tD mov;ents of banded birds in tne 200 acre study plot outlined in Fig. 1, the center of the Hicoria region.

Jays were captured in mist nests and baited Potter trans. They quickly learned to avoid the mist nets, and

various methods had to be devised to lure them in. ::zt effective ..as the use of a hand-held Scrub Jay which, when shaken slightly, gave loud alarit calls. This attracted many local jays and Ithen chased the_: toward the nets.

Each bird received three bands, two colored :stic coil bands and one aluminum =ilver colored Unit-d States Fish and ildlife Service band. Each individual was identified bv a series o letters connoting ,te color seausnce of bands: RS-R was a bird with red elastic and silver aluminum bands on the right lea and red plastic band of the lefL leg, l1G- had light blue a-d 'ight green bands on riht leg and silver band on left leg. All birds captured in trirs or ets had :he silver Lanc olaced on the right 1. winless definiteIv known to b3 a


'i




a


when adult had silver bands on t e ri ht Ieg. The juveniles banded during 1963 had one colored band on the right and one colorsd and tbe silver bard on the left, and juveniles banded in 1969 had two colored bands on the right and one slver on the left le. Duco cement placed on nlaStic colored bands effected a durable saal, but occasionally bands were lost and the bird had to be recaptured.

To follow the movements of banded birds I used a

30 power telescope mounted on a gun stock. In this wav I could read band sequences from. as far as 100 yards and so did not have to disturb the bird. Occasionally a single-sided blind was used but normally it was not needed.

To study the local distribution of Florida Scrub Jays I attempted to examine all contemporary Scrub Jay breeding sites in Florida. This appears a prohibitively difficult task, and several small areas were undoubtedly overlooked. Yet I believe I was able to find nearly all of the brecding regions. The many letters from Florida Audubon members in rel to solicitations for information greatly facilitated this search.

An invaluable aid .ien hun ing for Scrub Jays is a

portable tape-record-r with recording- of Scrub Jay alarm calls. These calls quikly attract any local Scrub Jays.

As a measure of referred habiat I used tha :uho of active neco located in one hundred acres. Difficul-~~~~~~~~~~ a---- a .--~eo. --C' sr o




9


when nesting areas consisted of many scattered catches. Even so, centers of ovulation numbers were located and

their comn physicgraphic features assessed.

Ten areas t1at included a variety of types of plant cover, yet harbcred nesting Scrub Jays, were examined in detail. A nest census was made. Plant cover was analyzed and the amount of ground in the thickets shaded by the dominant woody plants was estimated. Unless I am discussing a specific plant cc=,unity such as the Florida

sand pine scrub, the word scrub connotes a thicket of evergreen woody plants that is low and dense.

Field work began in June, 1967, and was completed in May, 1970.














RECOG .TICN S IG NALS


The calls, pcstures, and movements that are used in social interacti.cns are treated; other variations have been described by Amadon (1944). In this section the author describes individual calls and postures and sometimes implies their motivation. Later sections discuss how these calls and postures are combined in complex behavior.


Auditorv Sianals


K -err or ker-.--The Florida Scrub Jay utters a

variety of calls, yet most are variations of a rat-her harsh and grating note. Amadon phoneticizes ohis basic call as ka. To me the basic call contains more consonant sounds (kwerr). This call, as Amadon notes, is used for several functions, and certain variations appear to be used cosi ently in similar situations or to express particular motivations.

An ur;D ard irnletion of the kwerr call normally

indicates a heightened motivational level. Such calls are given when some alarming object is present such as a predator.

An increase in intensity or loudness of thiis call a Is :.............





11


a confrontation with another Scrub Jay. As Amadon states, this extra loud call is citen lengthened into a screech.

Finally, the ra krr- :er -kuerD calls are heard. Again this is a sign of increased excitement. It is normally used in antagonistic bouts with other jays, especially after a series of loud kwerrs.

"Hiccuo'.--This peculiar sound apparently is produced only by the female. Amadon's pronunciation of it as klok is quite exact. It is a hollow sound repeated rapidly so that it sounds like a hiccun. It is often used in antagonistic contexts yet it frequently is given when birds are quietly perched alone or during courtship. The loudness of presentation expresses a more excited state.

Beggina call.--This cry is always used with a wing flutter and is the characteristic sound of the youna in the nest as a parent arrives. The same call is used by a female toward her mate. I have never heard a paired male use it, although I have heard it used by an adult male toward a female in a small cage.


Visua.1 Sigrnals


As with the auditory sLnals most of these visua postures are those used by the Scrub Jay where reacting in some agonistic cc' xt zicrtoarc chtir con s -ecies

or toward a predator.









Upic'ht zosture.--This is difficult to classify as a special behavior because Florida Scrub Jays appear to perch natural with the body upright and tail hanging straight domn. Yet the male of a breeding pair habitually perches in this posture on a tall object from which the bird has a clear view )%er ipuch terrain. It is often possible to identify a Scrub Jay from a distance by this posture. This position can be classed as neutral in motivation.

DiD.---In this posture the head and body are bent downward and the tail is raised above the horizontal. This posture always expresses increased excitement and usually is an alar, reaction.

Bobbing.--Again when a bird is excited one reaction is a rapid bobbing of the whole body without moving from the perch. The degree of excitement can be roughly measured by the rapidity of the bob.

Undulating flight.--Normally a Scrub Jay flies from one place to another with a steady, slightly undulating flight. When highly mczivated chis flight is more deeply undulating. This pattern is normally accompanied by a series of loud kearp calls and is often exaggerated by flying slowly. Undulating flight often follows an encounter with another jay.

Bill up.--The bird tins the bill up and remains

silent, and the body is often slightly crouched. This is a 4ling y: i b e

bird flying aw:ay.









Wing flutter and groh.--This is a begging posture and is almost aL-:s usec -ith a begging cry. The body is pressed close to the perch or the ground and the wings are rapidly fluttered.

Bill wiing.--The bill is rapidly rubbed on both sides of the *oerch as though attempting to clean something off. This movement is used at the conclusion of a wide variety of activities: after fighting, after swallowing, after courtship feeding, or after drinking. It is so frequently used that perhaps it can be viewed as the final consummating portion of a sequence of actions.

















Unmated birds frecqunntlv travel and feea in flocks, but no special group activity relating to courtship has been found. This is unlike the Blue Jay, Cyanocitta cristaca (Linnaeus), which performs a highly ritualized group activity (Hardy, 1961).


Pair Formati)n


The initial stage of forming a new pair bond begmns with the male restricting his movements to a particular undefended area of scrub. After the male is first found restricting his movements, a female soon accompanies him.

On 10 January 1970 M-dBS, a male, had restricted his movements to a region just west of territory 4. He cantinued feeding in territory 4, and five days later S-BY, a female, was first seen following the male. After this

date Ltev rarely returned to the birds in territory 4 and the male began to defend actively their new region. I first saw M-dBS chase another jay from the new region on 15 January, and S-BY followed him. The female invariably followed the male during the next few weeks as he wandered through ihe territory and especially when he chased intruders. Such constant accompaniment undoubtedly helps reinforce th ar acnc.


1 4









Althcug> more tan 70 juvenile Scrub Jays have been banded within the 200 acre study area at Hicoria, M-dBS was the only male ba-ndd here as a juvenile to establish a new pair bond and territory in this same area. One other bird banded as a juvenile settled to the west of the study area, and I followed him as he established a pair bond and territory. O-SW, the male, settled in a small patch of scrubby live oaks surrounded by wet flatwoods approximately 1200 yards west of territory 3, in which he was banded as a juvenile in 1968. On

8 December 1969 he was first seen in the scrubby patch and had another bird in attendance. This bird was not banded but gave the female "hiccup" calls as soon as I approached. She was subsequently captured and banded RS-dB.

On 8, 9 and 10 January 1970 0-SW continually approached the female who at once crouched and gave begging calls and wing flutters. The male immediately passed a morsel of food to the female. After passing the food no cther special calls or movementE occurred except that usuaLly they both bill-wiped. Occasionally a female gave "hiccup" calls after courtship feeding, but usually both began some completely unrelated activity such as feeding or preening.

As I slowly approached C-SW on 10 February 1970 he

was perched or- the tallest branch available in this scrub patch. There -,,eac, r I cculd anproac' u .ctioed and at 70 r e gave urwaril inflected kwern calls. At




16


once RS-dB jurped ap ircm the ground. 0-SW bobbed and continued calling, and RS-dB "hiccupped." Then the male flew off giving rapid V :zrcs and the female immediately followed, giving the same calls.

Whenever a di sturbance occurs it is the male that

first flies and the female follows; this pattern is followed by all mated pairs. Pair formation then is a matter of the two birds remaining in close association, one bird, the male, passing food to the female and the female following closely after the male in all encounters.

Although the above discussion deals with the formation of a new bond, it also explains the interaction of paired birds, frequent courtship feeding even during the fall and winter, and persistent following of the rale by the female.


Copulatorv Behavior


The complete copulatory pattern was observed only

once, although I spent many hours behind a blind hopeful of following ffis sequence.

On 10 2.ril 1969 GS-G, the male, presented a caterpillar to his mare S-RB at 1030 and brought her food ei:ht tires ring the next 50 minutes, even though she continued to feed on her own. Frequently, after courtship feeding, the female gave rather suidue6 "hiccup" calls and a series of melodic notes and calls as though

s~~~e~ aer a ~U2JY o'e ovezr to her an-'









as usual she at once gave begging cries, and crouched down. He fed her but remained standing before her. She continued to beg and GS-G began to hop around her. His circling movement was a curious stiff-legged affair with his neck extendcad and b:1l pointed slightly down. His tail was sTread and slcwly'. swung from side to side. The wings were half spread and lowered, almost dragging on the ground. He circled her three times while she remained crouched with her bill raised, a submissive posture. The male was silent, and just before mounting, the female also became silent. The male mounted the female for approximately six seconds and then hopped to a nearby perch where he vigorously bill-wiped and shook his plumage. The female also began bill-wiping, ruffled her feathers, and bill-wiped again. She then "hiccupped" rather loudly and followed the male when he flew off. This pattern is essentially the same as that reported by Brown (1963) for the Scrub Jay in California.














TER:iJTORIALITY


The Florida Scrub Jay is a highly territorial bird. At Hicoria territorial boundaries frecuently made broad contact and border conflicts were colmon. The boundaries were fairly easy to map and Figs. 1, 2 and 3 outline the territories present at hicoria during the three years of my study. Some daily shifting in the location of the defended border occurred but it was minimal. Whether it shifted or not, at all times a definite boundary wa recognized by other breeding jays, and if a forei-gn jay crossed, some sort of conflict ensued.

The area defended, among the 25 territories measured over three years, ranged from 3.8 to 7.5 acres, but most were close to the average of 6.0 acres. Each territory included a large amount of scrub edge and open soace with which Hicoria abounds. The western side of the study area included a large stretch of very dense scrub. The only jays nesting in it were at the edges. I never found pairs defending ground in the center of this dense area.


Nest Placem'ent


The birds invariably placed their nests t.O or tree feet in f: the ed-e of an o*en soace. Usually myrtle caks were slectcd d nest building but alic used :ere wax myrtle, Chaman oak, and in one case a scrub oalmetto
























Figure 2 of stLy i e at 7i,-ori. shwing trrritorial bounctaries and Scrub Jays residing
in each territory during the breeding season
1969. (See key for Fig. 1).



















tit,



R x
I'










\- i !





Ir U) I 2



a tr Z s

tt C~,
ZrIL
t Al
a C'""'


'-I xi

sa ., ~ .
























Figure 3 Map of study area at Hicoria showning territorial boundaries and Scrub Jays residing in
each territory during the breeding season
L970. Note: no young were banded this year
(See key for Fig. 1).













7 '--1 N




z' .j... ...
6 n CL -z





















I c.M o
I ..
,,.









Zn
". ~s

r *. I .. t

__ _ __I J .,k'?I7

( __ _-_




i.*--~1
..\.,
____ ___ ___ ___ ___ ____ ___ ___ ___ __ ____ ___ ___ ___ ___









that was intert-:ined -it. a myrtle oak. Scrub Jay nests were well caz-,ouflaged en were difficult to see. In order to locate all the nests I had to search though practically every tree in the area.

Nest building began as earl- as 20 February and extended to 2 June at Hicoria. There was no definite time when most pairs were nest building or incubating; it was a Drotracted period. One pair had young out of the nest while another was nest building. Unlike Blue Jays (Hardy, 1961) the Scrub Jay pair defended its territory from all Scrub Jay intruders no matter what. the staae of the breeding cycle of the defender or the intruder. GS-G and S-RD did not cowmence nesl building until 18 May 1968, yek they actively defended a rather large territory from all Scrub Jay intruders from .Iarch onwards. The nest was completed on 21 May, but no eggs were laid until 4 June.


Territorial Defense


The male of a pair spends much time perched in an upright position with tail held straight down. This behavior is most comnion during breeding season and occurs wrile the female is both on and off the nest. Trom the perch the bird can readily see all of his territory. This same uostcre is used by Scrub Jays in the fall and winter when in a f1ock Consistently one sees a single ezrd, not a.zessarilv a male, oarchaz i aboy:e the othe' feeding below. If this bird drops to the ground, witin




24


a short time another flies up and replaces him. A bird may perch there for as lcng as 30 minutes. The high bird invariably is first to sight an approaching intruder and begins bobbing an4 c-lLing, usually with upwardly inflected k:err. At cnce the other birds fly or hop into the branches -.earest to them. This position of the bird is undoubtedly a form of sentinel behavior both during and after the breedi ng season. Yet on such a perch not only can he see over his whole territory, but others outside can readily see him. Possibly, then, this posture may function as a means of proclamation of territory.

Rarely does an outside bird intrude far into a

territory during the nesting period, but boundary disputes are frequent. Chasing is the normal means of territorial defense. When another bird approaches or crosses the boundary the male of the invaded territory flies directly toward the intruder. Physical contact is almost never

necessary. The encounter quickly terminates with the intruder rapidly retreating. The attacking male veers off abructlv at his territory boundary and almost never ex-ends his flight beyond it. His return flight often includes a de-plv undulating Flight and a series O loud raid kwerp calls. Ar times the initial attack by the defender is silent- when a call is used it is a loud kwer .

The mate often follows the male but rarely takes

)art Ln 3.-i:.e.se cerches nearby in the territory,









actively oozrxc and iving loud "hiccup calls 'hen the male flies back, the fea. usually -ollows in an undulating pattern and c-l11'ng She "hiccups" in flight.

These bcr>-: rebuffs are not cne-sided affairs. If a male chases a zird from his territory the roles are reversed when th: latter's territory is invaded. Each breeding male then has a definite boundary over which he will allow no intrusion during the breeding season.

These border disputes occur most frequently In the

mor:ling. Later in the day when the heat and humidity are high Scrub Jays become quiet and little activity is evident. At sunrise the jays are very active and territorial pairs perform a series of encounters that are repeated at the same place and time, day after day during tha breeding season.

The following description of the movements of GS-G and mate in territory 3 will describe these encounter.. On 21 May 1969 at 0630 just before sunrise GS-G came swooping northward toward the boundary between his and BS-dB's res-:ccive territory (territories 3 and 5). GS-G was flying an exaggeratedly slow and very undulating

pattern, calling loudly and rapidly keru-k-wa r At once BS-dB flew toward GS-G using the same pattern of flight and alsc calling loudly. When approximately 20 yards apart t7ey si!ataeously circled and fle-.- back to:ard the center of their res-sective territories, still calling









similar calls or ln-e arby and bobbed and "hiccuoped' furiously. Fesm.les never initiated nor passed ahead of the males during such activities. The "helpers," to be discussed later, ofter joined in but remained behind the female. GS--G was then attracted to the southern border of the territcry by a similar pattern of flight and calls from SR-0. Fifteen minutes later RS-R and PiS-0 had initiated a bout at their common border (territories 1 and 7).

Only early in the morning did I see these attempts

of one territorial bird to draw another. These seemingly ritualized patterns of border conflict were common during the breeding season; during the rest of the year they were less frequent although now and then they were repeated.


Site Tenacity and Mate Constancy


During the three years that banded hi ds wer followed at Hicoria, the majority of pairs remained at the same place (Figs. 1, 2 and 3). At any time of year each pair usually could be found in or close to the territory defended during the breeding season, and from year to year the same sites were generally defended.

Even after the young successfully leeve the nest, territorial defense remains at a high intensity! with freqcuent early morning bouts and continual reaction whenever a strange bird wanders too closB. Eut after midsu -er such tia'-t border co-tro 7 relaxes. Until the r.ext





22


breeding sot cse r spend core time wanderIng out of their a:. mixing with other Scrub Jays.

Yet whenever the pair returns to their territory they chase intruders. -n this fashion the boundaries vary little frcm one season to the next.

Apparently the Florida Scrub Jay mates for life. Two of the pairs at Hicoria are known to have been paired for at least four consecutive breeding seasons, RS-P with WS-Pi and GS-S with S-RB. WS-Pi, a female, is of special interest. I banded her as an adult in the summer of 1960 at Hicoria within 200 yards of the territory now occupied. She is thus at least 12 years old and still breeding.

Mate constancy was the rule with the majority of the

other mated pairs. The following pairs were together during 1968 and continued as pairs through 1970: SR-B with S-BO, SB-N with SB-dB, RS-Pi with MS-B, and Sr-B with SW-R.

Only three pairs broke up in this 200 acre site. NS-N, S-WN and young disappeared in late October 1968. I have no specific knowle.dge as to the cause- of their disappearance, except that Scrub Jay feathers were found in the territory. I surmise that some predator killed one or more of this group, probably including one of the

parents, although two of the young were later found associated with a group of non-territorial birds 500 yards to the -orth-est. The parents an,; remaining young were not located during the next two years, alihcugh I roa:ed over









the region as far as three mile! fr.L Hicorla looking for banded birds.

SG-B and "-R nested :or two years in territory 9,

but each year the ycung vers destroyed while in the nest. During 1970 ti-1 pir disap~neared and the territory rerrained unoccupied. ?erhacs continual nest destruction ir.duced their departure. Yet other pairs continued territorial occupancy arer nzst destruction.

On 20 May 1968, a feral cat located the nest of SY-Y and S-BY. All the nestlings were killed and acparentlv so was the male, SY-Y, for he was never seen again. Dislocation of a breeding pair probably occurs only after some major catastrophe. Otherwise a mated pair remains at its original terriLry year after year.


"He lcers"


I have stressed the territorial n trc of t e social organization of breeding Florida Scrub Jays. They are territorial, but each territory often contains more than a mated pair. For instance on 12 June 1969 territory 8 contained IS-B (female), RS-Pi (male), O-SG (female), O-SW (male), dBR-S, PiY-S, and PPi-S. The first two were the mated pair, the last three juveniles. The remaining two looked like adults (juveniles have brown heads until their first molt in July) yEt they were definitely tolerated in Lhe territory and hed b-eU. ar spring. They were, in fact, the young -roduce b% thei




29


pair in that territory the preceding season of 1968, for I banded them in the nest.

This pattern o rate.tion. of young of the year into and through the next breeding season as members of the errotOry is prevalent at Hicoria and at other areas in Florida from which I have banding information. The presence of helpers at the nest was first reported for the Corvidan by Skutch (1D35) Grimes (0.940) first suggested. that Florida Scrub Jays had extra birds active about the nest, and Amadcn (1944) in his study of nesting Scrub Jays at the Archbold Station occasionally found extra birds tolerated at cartain nest sites. He tentativelv concluded that many first year birds may not breed and are sometimes tolerated about an active nest.

During the breeding season of 1968 certain birds

banCded earlier were found associated with breeding pairs. As outlined in Fig. 1 four extra birds were found, one in each of four out of nine territories. Using the flight feather technique of Pitelka (1945, 1951), 1 tentatively ident fied three of the four as first-year birds. Plumage characters were difficult to analyze becaaL all the jays had worn plumage at this time of year.

In 1969 seven "helpers" were found in four of ten territ ries, one in territory 2, three in territory 4, two in territory S, and one in territory 10. The other six territories hac only a :nitec pair in occupancv. Si.-





30


of the seven "helters .are first-year individuIls that had been banded the previous year in the nest of the pair with which theY -.r : associating. The other w:as S-BY,

an adult, and zjic discussed later.

Finally in 1970 six of nine territories and nests in the same -icoria site had "helpers," one in territory I, one in territory 2, one in territory 3, one in territory 5, three in territory 4, and one in territory S. All of these "helpers" ha' been ban.ed the year before as juveniles and in the same territory.

Approximately half of the territories at Hicoria had

extra members. Banding data show that they were invariably juveniles of that pair from the year before, with the exception of S-BY ncted previously. Although reminiscent of the social organization of the Mexican Jay (Aphelocoma ultramarina) in Arizona, in which nonbreeding birds frequently functicn about an active nest (Brown, 19E3), this case is special in its combination of territoriality and sociality (c.f. Skutch, 1961).

The territories that had no "helpers" during 1969 and 1970 were those in with the fledglings either were destroyed by predators or disappeared from the study area and surrounding habitat.


Interactions of Adults and Juveniles


Afterth or e-ote'- >"-''





31


.Even when a ditu:'ban2: al rms the parents, the young rema-in 1ell hidden a S --lent. As newly fledged youngsters are vulnerable to edaaion, this stage of extreme timidity

stv I h arants quickly recognize a Dredator and

may draw its a.tenzo_ -ith their loud alarm call, upwardly in Iected Z7ern; tche silent yoing may remain unseen as the predn.tor is distracted by the parents. In fact. many young are lost during these few weeks, yet possibly many more would be killedd if they were not secretive. About two weeks after leaving the nest the juveniles first become active.

Juvenile Florida Scrub Jas have light brown plumage on head and back and lack the light blue forehead of the adult unti their first mcolt in late July. It is easy to recognize juveniles by this trait. I saw no special reaction by adults to birds with brownish heads except when young from other territories inadvertentl-y wancred over another adult's territory. The resident male quickly chased these wandering youngsters. The ensuing chase was especially intense and afforded the only occasions that I viewed physical contact during an attack. The male's cortege of mate, helpers, and young often join the attack by following :ae male and calling raccoS1-. AsT border disputes, normally only the male initiates these attacks.

in (1961) suc-ests that "the juvenile usually

laCL e.asa Ca:Gces ;hii: a'IleiaCz 7L42 ign:iicant in





32


aggressive behavior in adults." By implication juveniles with distinctly different clumnage should trigger a less aggressive response by an adult. Within the territories at Hicoria this <-as not so. Apparently territorial aggressiveness cverridCes any recognition of juvenile status. The brownish pattern undoubtedly helps as a concealing coloration while the birds are feeding in thickets

although adults are, tiumselves, extremely difficult to see in thickes.

The juvenile birds in their proper territories give begging cries and wing flutters whenever either parent comes close. Such begging continues through the year although less frequently. if the parent has food it is often passed to the begging juvenile. An adult-like plumage is obtained in. July, but even so the parent still responds to a juvenile when it begs.

The parents frequently travel out of the territory,

usually to a food source. Male and female normally travel together even during the nonbreeding season and often their offsping -rocp along. In these cases.thev join with other Scrub Jays. As many as 30 Scrub Jays may congregate at a baited site. not in any territory. Juveniles beg to zheir oj%7 parents alCho:-gh they are usually ignored. The young may even beg to a strange adult and again are igncrec.. 3reedincr :ales hrescci here are not motivated to attack juva-n-ils w4ith brown plumrage.



exact v :Lmila- to courtshi begging o a emale to her









mate, and the reszcmse is usually thie same, food being passed to the beggar. OZ-jously the continued occurrence ol courtshic feed ing, 'reen mated birds is a means of pair bond reinfcrce-ent. It is seen most frequently during the breeding season but occurs all year round. PossLbly as juvenile< 1iirds continue to beg and the parents respond and feed them, an attachment something like a cair bond is 2stablihed. As the next breeding season approaches the oaren : ar icrcasiznjl attentive to each Qther, vet the young from the previous season are still present and tolerated by the male. Occasionally they beg and he responds b-7 passing a morsel of food. I suggest thal- these one year old birds can remain in their parents' territory

because a loose bond is present between the helperss' the breedincT male similar to a pair bond. Role of the "Heloers"


The role of these "helpers" in the organization of

territorial members is often vague. I call them "helpers" but this term is, a- cies, mi sleacin. It is a term that has been used in the JasC and I shali. continue its use, but "auxiliary territorial member" is perhaps more accur e .

I never sa-. these "helpers" take oart in nest building. usually :he first-yerc-: birds igno:-e thr- a nesting procedures and quietly feed sc-where in the terrizC:27, Anen ;y>tng are in the nest and the z.ents




34


actively feed thi. irsct-yer birds occasionally follow one parent toward the nest. At such times the breeding male often splan'ts -n first-year bird by landing next to it and rmi:es poking. The "helper" either gives a submissive ostue (slightly crouched and bill upraised) and then flies off, or else flies a short distance at once. Rarely does the male chase after the "helper."

On 12 May 1969 in territory 2, SR-0, male, was not in sight as PiS-W, female, flew to the nest and fed a youngster. She flew off and immediately lG-YS, a female first-year bird, flew to the nest and peered in. The hestlings began begging and lG-YS moved her bill toward them as though to pass food but none was passed. Again two days later, just after the female left the nest and the male was not in sight, lG-YS flew up to the nest and this time fed one of the youngsters.

Then the young leave the nest the parents continue

feeding them and now the first-year birds more frequently bring food to the fledglings. In territory 1 in 1968, thz younv left the nest on 13 :'arch and for the next three days S1-W, female first-year bird, brought food to the fle-ci5,s ai last tw ice each day. Ln 1969 M-PS in territory 10 fed young after they were out of the nest. Thus, the first--Year birds present in the territory occasionaily function as "hlners" with the young. }'iost of te time, howev-r, they remain somewhere in the territory

-2- O








During a2ress 'ncounters ;iLh neighboring birds these first-'ear birdh most readily take an active role or act as "Iheloers." Whenever the male gives chase to an intruder, the first-,'ear members of the territory at once flv towar' him and follow behind, frequently calling as loudly a- the male. A border encounter often entails three or four birds, loudly calling and chasing up to the invisible ter-itory boundary. The male then wheels back and the 'ther3 follow him back into the territory, all still calling loudly. Certainly a mob of jays, all seemingly bent on attack or chase, is much more effective in instilling a sense of invulnerability about a territory border than would only a single male.

In the same way the seemingly ritualized early morning border encounters include the "helpers," if present in that territory. In these cases two mobs of jays may

approach each other and suddenly, wheel and fly apart. If a territory contains more than one "helper" all do not necessarily join in these border contests.

In 1969 territory 4 contained twc adult. breeding birds and three "helpers." On 10 May an unbanded bird flew through the territory. At once SR-B, the male, took after the intruder and the other four birds feeding with the male followed. The unbanded bird sailed rapidly across the zerr'tory. The five aitac--s oamr back to the feeding site. but all remained in the hiahli agitated sa 1 ai least 10 mic- s. S-f',t-e emale, saz on




36


a tall perch anD bobbed and "hiccuuped." This enticed S-BY, a female helper, o fly up whei- she commenced "hiccu7-Lng." As ';e- fcll silent Sl-B (male) flew off 50 feet, qivinq loud. rapid kwerls. Again the other four members of the te rritory' followed, and another series of bobbing and "hiccupping" ensued.

Not only did SR-B and S-BO in territory 4 have more helpers than did other breeding airs, they also had one helper that was an adult female. This bird, S-BY, was the mate of SY-Y in 1968. On 20 May a feral cat attacked anc, killed their young in the nest and apparently also killed the male, SY-Y. For the next week S-BY remained in the region of the destroyed nest but more and more regularly she fed with the members of territory 4 at a feeding site that I had been supplying between their territories. By 25 May she no longer spent any time in

her old territory and followed along with SR-B, his mate, and helners into territory 4. At this time SR-B and rate were attending young in their nest, and S-BY began to visit the nest with fooi. Whenever SR-B saw her at the nest he gave chase, but she never flew far. She always moved off when he was aggressive. On 26 May I first saw her beg toward SR-2. Ho ignored her but did not chase. Thereafter she frequentlv begged toward him, and he usually responded with a zorsal of food. She remaIned in that territory as the ory adult "helner" that T have found. S-BO, female,




3 -)7


S-BY remained ti d to this pair that it appeared that SR-B had two s.ates. Although he regularly fed her when S-BY bezc.ed, --30 ren.ec i.s dominant mate and was the breedi 1%emale t>7 --.t -,ear.

In 1.96 S-EY was still in attendance. She did not help with nest building, but regularly brought food to the young in the nest. During this breeding season the male did not attempt to chase S-BY from the nest. She was then truly a "heloer' at the nest. The other two first-vear birds in attendance did not attempt any such close association about the nest; they acted similarly to first-vear birds in other territories.

Interestingly, in February, 1970, S-BY and M-dBS

paired and established a territory just west and adjacent to territory 4. :7-dSP was one of the first-year birds in attendarce in territory 4 while S-BY was also a helperr."

Thz Tlorid_ race is unique ai:i-cng Scrub Jays in that

family flocks defend their territories with all individuals helping in various degrees. As far as known it is found in western races (ardy, 1961).

Te social organization of these flocks is as would

be ex-ecLej. ?he Dreeding male is dominant over the firstyear 'hlcs: ." alThough normally they appear to ignore each other. Only occasionally did an adult female supplant any oZ her irs--year offspring in the territory and I never saw a Female chase or attack a "helcer." In a territory









e sta bli sh mon the "he7erss" at first. During the first winter, at cime a rale first-year bird would supplant a female si'LKLr. by displacing her from a perch or by pecking at her, but acain the usual pattern was of no interference.

















About 60 ,-r n of the terrain at the Hicoria study area was tied u;p in defended territories. The remaining area was inhabited by small loose groups of nonbresding birds and bi territorial birds which often wander out of their home area. In 1962 two feeding stations were maintained, one in a non-defended area and one immediately between two territories. At feed site 2, placed between territories 5 and 4, only the birds inhabiting those two territories visited this food site., accompanied by S-BY after her mate dis~ascnarZ. Althcugh border conflicts between BS-dB and RS-B occurred frequently, encounters were infrequent at this food site. Birds from the two territories hardly ever fed here at the same tir7e, and this obviously reduced the necessity for territorial defense at this site. During the few times both groups were together AS-B once chased S-2N and occasionally supplanted BS-dB.

Feed site I was visited not only by territorial birds frcm adjacent i:rritories but also by ten other birds that frequented this general area and apparently had no territorial affiliations. By pliumaage c aracters e'tr.c 7ere first-year birds and two were adults.


39




40


I expected to fird a definite dominance hierarchy among the Scrub Jays at feed site 1, but did not. The only bird to exert his -ominance frequently was RS-R, whose territorial boundary and nest was the closest. The other breeding males, NS-N from territory 6 and RS-O from territory 2, occasionally supplanted the nonbreeding birds but most frequently coexisted, often side by side with both non-territorial jays and with most of the territorial birds visiting this site. Although many birds often frequented the site at the same general time of day, usually only two or three dropped down for food at once. As they flew off a few more dashed in. There was little opportunity, then, for much conflict, or supplanting. many hours were spent watching the movements of jays at this site, and no definite hierarchical ordering could be assessed except that breeding males were generally dominant over most other jays.

In many bird species, the more closely two individuals approach, the more likely displays or attacks become. Each individual has a specific individual distance within which the bird will not tolerate others, excepting sexual encounters. -mong the Florida Scru_ Jay;s at Hicoria this trait was not sen. Birds frequently fed side by side or spent considrable time close to each other.

















Alth g territorial Florida Scrub Jays react vigorously toward Inltruders of their own species, they usually ignore other bird species. The Brown Thrasher (Toxostoma rufumn) is similar in size and ecological requirements. Pairs are scattered over the Hicoria area and they frequently feed in the same places as Scrub Jays, yet conflicts rarely occur. Only when a thrasher or 2ockinabird

(Mimus 2o.lyglottos) perches especially close to an active nest dces the male or female jay displace the intruder. Elsewhere in the territory other species are tolerated.


Reaction to Blue Jays


No Blue Jays nest in the Hicoria area, yet numbers

are always present, either feeding or travelling through, often in groups of four end five. Blue Jays frequently feed with Scrub Jays. Usually the Blue Jays.remain in the background and only after the Scrub Jays leave the feeding si 0 tu e Blue Jays dash in and grab some food.

Frequ ntly t f7edi ng sitec crub Jays and Blue Jays corne ctcse tof:thor and invariably -he Scrub Jays dominate the Blua Jays. ccn cunberg rarely include an attack

or chase; merely a neck or Jab toward the Blue Jay is


41









Blue Jays oftan forage thrc-.gh Scrub Jay territories. Only when the B ae Jay cporoaches the Scrub Jay or blunders

near a nest does the Scrub Jay react. In such cases the offending Blue Ja-y is often chased. As with intras-ecific chases, the other -erritoriel members quickly join in and follow to the edgre of the territory. Occasionally the breeding female Scrub Jay begins such a chase, but the "helpers" join, ard the al usually takes over and completes the -h1se to the border. These interspecific encounters are infrequent, for thE Blue Jays normally stay away from Scrub Jays in their territories.

Scrub Jays recognize something in the Blue Jay a necessitating an aggressive reaction. Yet the Blue Ja: is not treated as a distinct threat, for it is frequently left alone in Scrub Jay territories and at feeding sites.


Reaction to PredaCctrseAlthough the male or any otner territorial member in

a sentinel position frequently warns the others of predators, during 1.938 and 1969 five of 20 nests with young were destroyed. Probably the common predator of young Scrub Jays are snakes. At Hicoria I frequently found Corn Snakes (Ela 7 t z--ca), Yellc: Rat Sn:es (Ela!he obsolete Pine Snakes (PituoThis melanoleucus ngsnakes (Lamuro' agi cgetulu.s'. Black Racers (Coluber constrictor), and esreciail' the eastern Coach hin CMasticohis flacellum), The Ccachwhio a. :rlc zso cu









On 22 April 1959 I banded the nestlings in territory 9 and replaced them in che nest. The next day the young were still in the nest. but on 24 A1ril the nest was empty and two adults were tre:.ly agitated. Dead on the paved road 200 feet from the nest was a six-foot Coachwhip that had been hit by a car. Upon opening the stomach, I found the two young from territory 9 still wearing my bands.

Once out of .he nest the young are still highly vulnerable for at least a :eek and many disappear. Feral cats are frequent at Hicoria and prey heavily on fledglings. Gray Foxes (Urocyon cinereoargentous) and :aas (Felis rufus) also hunt through this area.

Whenever a predator is noticed, the Jays in the territory mob about it, using all alarm and excitement behavior patterns. Some give loud upwardly inflected kwerp calls and bob vigorously. Females frequently give loud "hiccup" calls and bob. A jay often dips its head toward the predator with its body hanging straight dow.n. Sc:e, expecially the breeding pair, hop about the ground, calling loudly and nipping the tai2 of the attacker.

The output of noise easily carries over many other territories, but other Scrub Jays normally do not cross into the territory ;here the trouble occurs. Birds in other territories are aware of the attack for they become extremely excited. Only when the predator is found close to the territorial boandary or when the encounter continues

-or an Csreciall leng tim:. do chr t-ritori s




44


enter and join in the melee. As tha new jays approach, the local male ofton dishes suddenly at one of them. These are shcrt chases, and ">e zimale quickly returns to the predator. These short qoick attacks at a foreign jay rarely continue to the territorial border. During nonbreeding seasons when mated p-irs are less insistent about territorial defense, alarm calls more frequently attract other Scrub Jays.

These observations concerning the attraction of other Scrub Jays to distressed breeding birds are csriistent -.ith Hardy's (1961) comments concerning degrees of territoriality among species of jays. The IMexican Jay with no territoriality and Blue Jay with a loose territoriality both give loud assembly calls that readily attract others, whereas the highly territorial western Scrub Jays do not. Even though possessing a degree of sociality above that of western races, the Florida Scrub Jay is highly territorial and does not have a distinct assembly call. Its alarm calls carry far, but as with western races, outside birds :req ue:-tzly d not assemble when a pair is in distress.

Nearly every other resident species at Hicoria does

assemble.' The alarm calls of the Scrub Jay quickly attract Brown Thrashers, Blue Jays, Mockinabirds, Towhees (Dipilo erhthroohthallmus), White-eyed Vireos (Vireo griseus), and especially wood eckers. Red-bellied C7cdDeckers (Csnturus carolinus), Red-headed Woodoeckers (Melanerres ervthrocecaus I,- a rs I cl e _I ag re eas il e xcit S








by the loud u:;ardly inflcte. kwerp calls of the Scrub Jay and remain wich the distressed pair as long as they continue calling. These other species probably do little to aid the distressed Scrub Jays, for they remain well away from the action. Possibly Scrub Jays aid other nesting species by :redator recognition.


Reaction to %an


One feature which makes the Florida Scrub Jay especially intriguing to work with is its extreme tameness toward man. On 17 June 1967 when I first started banding at Hicoria I walked down the dirt road by territory 3. A Scrub Jay landed ahead of me and a juvenile near my feet. I was surprised at their seeming unawareness of my presence. Then the juvenile hopped between my feet, and the adult landed on my head. I held out my hand and the adult jumped to my arm and then to a nearby branch. I fumbled in my pockets as though to bring out some food and again held my hand out, but they did not fly to it. I walked up to the adult, which remained calm until I was three-feet way, and then it flew 10 feet down the trail.

This encounter surprised me for, to the best of my kno...leJ no one has spent any time at Hicoria feeding Scrub Jays. In fact they did not appear to be attracted to me for Jcxx, mrely inquisitive about my presence. At the Archbold St-:i Scrub Jays readil.V arproach a person

a:d 1: to z::.s ani. These birds have been conditioned




46


to expect a haniouz of food, yet they have become so tame that one can frequen-l1/ pick up a brooding female from the nest, something that co-ld ne7er be done at Hicoria.

An interestin;; example of this tameness occurs at a small scrub thicket north of Englewood, Sarasota County. For several y,:ars people have been feeding Scrub Jays there. As one stops the car several jays fly out of the thicket and often land on the car or an outstretched hand. If no birds are in sight, all one has to do is sound the car horn and the jays quickly appear.

It is difficult to explain this tameness. Although most jay species are easily enticed to feeding stations, only the Canada Jay (Perisoreus canadensis) is as overtly tame as the Florida Scrub Jay (Bent, 1946).

















The striking feature cf the Iovement of Scrub Jays about the Hicoria area is that there is so little of it. Apparently, once a pair establishes a breeding territory, it remains as long as conditions are suitable. Of nine pairs nestir.g at iicoria in 1966 six were still there nesting in 1970. Onii of these was a female WS-Pi banded as a breeding adult in 1960 within 200 yards of the territory she now occupies.

Dispersal of certain members does occur, for in late

summer and fall the first-year "helpers" begin to disappear from their parental territories. During this period Scrub Jays are prone to make long flights. A single bird wil fly as much as 0.3 mile from the territory. Such long flights are usually silent, unaccompanied and at such a height (100 to 200 feet) that other Scrub Jays ignore them. Normally these long-fliers return later the same day. All birds that made long flights and that I could identify were "helpers"; I never saw a breeding adult make such flights.

This is not a phenomenon restricted to a certain time of year. Even during the breeding season, occasional long flights occur. They are much more frequent, however, in thn fI'I. Terha-s it is s of unrest by narti'ular ir5C-, J _relufe to a final mov. Bv late fall most


47




48


old "khepers" have disperse, leaving the new set of young to take on he role "h.

Dispersal i apparently celaved until the second year.

I have not found any first-year birds far from their parental territories. As rezported above, in 1963 several first-year birds were members of a loose nonbreeding flock. As they had just been banded I know nothing of their origin. In 1969 a group of non-territorial individuals roamed about the large open area west of territories 7, 3 and 5. A few were members of the nonbreeding group of 1968 and others were unbanded. Two, P-OS and P-dBS, were first-year birds and were banded as juveniles in territory 6 during 1968. P-OS and P-dBS had moved approximately 500 yards from their parental territory. These were the only birds banded as juveniles and found away from the parental territory during their next year. The breeding pair of territory 6 also deserted the territory, and this event undoubtedly influenced the movement of their young.

The remaining birds banded as juveniles and not present as helperss" probably succumbed to predation. I searched as far as three miles from Hicoria for banded birds, but no first-year individuals were found. It was difficult to check all the many small scrubby thickets to the west ard south; some birds from H1-icoria might have been overlooked.

Tuo new territories were established at Hicoria during 1969 (7 and 10) and one during 1970 (11). Territory 10 was

.tle. by unrked hirds, an& 7IJ? no knowledge of their




49


origins. Territorv 7 ;a3 settled by PiS-O and S-GB who were first-year birds in this same general area during 1968. Their parental territories were also unknown. Territory 11 was established by S-BY and M4-dBS who were both "helpers" during 1969 in territory 4. As these two areas were adjacent, this new pair was filling in unoccupied habitat at Hicoria and only a minimal dispersal occurred.

Three first-year "helpers" moved completely away from the Hicoria region during their second year. S-RY moved 1.2 miles due south from territory 6 and nested in 1969, O-SG moved 0.9 mile westnorthwest from territory 8 and nested in 1970, and C-SW moved 0.4 mile westsouthwest from territory 8 and nested in 1970. These last tw.'o settled in scattered scrubby thickets in the wet flatwoods region, while S-RY remained on the sand ridge but well to the south of Hicoria.

One other bird banded at Hicoria moved to a new locale. SO-B was banded as an adult during the summer of 1967 at a banding site just south of territory 6. Several birds that

wera banded there subsequently disacceared. SO-B was found

1.8 miles to the northwest in February 1969 and nested during that year on a narrow band o, scrub. This strip of scrub he. been censused during 19%?, :h2re SO-B was not present; its movements during 1968 are unknown. This was the farthest a;;a/ from the banding site that I located any of my banded birds.




50


During the Lr dIng seasons of 196) and 1970

Glen oolfenden color banded the local Scrub Jays at the Archbold BiologicFl Staticn and censused for other birds. We attempted to use ncn-overlaing band series so that hirds from th: Station and icoria could be distinguished. The Station is 2.0 miles due north of Hicoria, and to date no birds from Hicoria have been found at the Archbold Station and no Station birds have appeared in Hicoria.

As doc- '-nted at Hicoria, the movement of Scrub Jays from hatching sites is delayed for an entire year by the retention cf offspring as "helpers." Breeding maies may induce this retention by' continuing to feed juveniles throughou-Ithe year and so form with the offspring a loose bond, similar to a pair bond. When birds do disTrerse it is apparently for short distances. Yet in certain cases family groups are found far from scrubby habitat. in on-case a fire destrce. a large, secti on of scrub in late summer, and a qrou, of five Scrub Jays was found two days later approximately three miles from the fire, feeding at the edge of aae watermelon field, with no scrubbV habitat about. The next day the birds had disappeared.

Four miles northeast of Hicoria the scrub on several

thousand acres was destro'-d during the surm-mr of 1959 and orange trees wanteded. For a week thereafter I found small groups o: crb Ja7,s in situation wre I -vr -,: t

before and where they were not seen subsezuentl. As suasted ir t> .CVement of br ig pairs a- .icoria, long




r:1



distance overnents by Florida Scrun Jays apparently are rare and norm-.:-y !Iv e ind2Lced by some major catastrophe such as desIt,.ctio C the habitat by fire or clearing.


















--,E: -S elect a nesting ha.it .- of vjer i mitet a aT nd are absent even thcu. +the

~rc~cre~nz a~t-~4s ;=aabje.






'=tj-,-:c ~s are 4narahl lccaz 7

Of a.n. scr_... vegetation. The ve,tat nal co:.-n~n...

.. ....... .. xer.c .ants tla: a

reS trizd to tihe '71~se.and sor ted" Sz.izous ds

of modern andanojent shore 1_ re Duri r 1 late Terti-c.-:
an th- F en







andtheintril: a ~oo~os c he Pleistozen,- t :ie Jdrast1 r is e Z i, I lI S tr n C i t r a a s s oF deep sa n

-r .5 S : ..t ( .

The z present fa-: snd pe tron.b Tveaveita tis clzt nJenon these stret-ch-es o- S_=._--:Ls ol t-he Sain L-cie and Lk o5

-'-ez-n ..ne ..cur.r..ns. wave ao

and irds ae s.ie 19h8). Since -DheSre t sands ere


C~p~ as loon/ anao


an thes a inet:r .. -ia rti j or te leis-: are the rca."Ta res -i ianin Cru v t and on itslved .onZ

the -i sctrnltrsal -iihlands C ount. et Laksete T1953 and: wins (Lati 1 t8)e. Si. ef ort:d Sad we extre i sscr lie rsn wLarc today~* Jan eies









di s t fr qe ntl Ih CA D Ln at 1 ln' ts T)lpne

clauga), whiz. oe for:as a cormnpact zor st. tce lfeS d~en .se stands of se;f. c ir~a the understor- is lo.' ticket oZ va rious .oody --na:s, and the around is ofte, one: -hit sand, raps with a thin lyer -of Cl aonia 6. Scve herbacecuc .::rirz pLants cccur, and cccaicnaly fe? grras.s -'ndrczogon floridanus and Paniu a. pa-enti folium) mav occur.

WThen irasent as an understzr- to sand ine-, toe

wood; plants may be well spaced, but when sand aines are lacking, these shrubs frecaentlv make an extenive, d "s scrub thicket, in form strikingly siffil'rto communities of Arizona and California. The dominant Kan'ts of these low scrub thickets are three species of xeronorehio oaks (Quercus virainiana maritima, Q. mvrtifolia, and Q. chaumanii); these three frequently comuose ovar 60 oar cent of the ground cover in the thickets. The remaini-ng plans ay consist of a variety of species, including Serenoa reoen-is, Sabal etonia, 1ex cumulicola., Rhus curallina, Lyonia ferrcinea. Erica cerif ra, and occasionall Ceratiola ericoides. As Laessle (1958) reports, Florida sand oine scrub normally includes 7,any other shrubs and her-a.os rc'e, but thilt ab.ve incluoez the wcx:; spscie! witr. wh:ch Scrub Jays are :recuantly associated. Alt':.... :n c ten bare of grass, .ccd ;1:nc









in ra hr san-d -ine is desBu asnEsz and no Scrub Ja. ccur :;&- eing birds. -s te orida

Scrub a a.-ai-s 20inTh t:e Biu Jay e, ,he t;o are together, I doubt if 3'.Ie lue Jays nave di1ace the scruh Javs fro te urs sand rine scrub. Lt is rore

probable that th- Foria Scrub Jay finds this habitat



Sanj p-in treec: e bae asent from t"e rU 44--it

for s e 2... rC&aC. Sand nires ars escpec:ill suscep-tibie to fire and once f1aMs reach th- canopv, large regions quick y burn, Sut Webber (19351 states that .atur sa:pina forest may act as a fir inhibiting co m'uity because the ground is oftcn bare of grasses and :ire 'as IFoicult: in sprsading. Thus ma;; scrub cozzunitiis ma e~i covered with pine forest a-id are uravailable to Scrub Jays. Today the State Forest Service fire protection program is aicinci the p.r-eservation of sand pines.

F reIuent the washe an U1 sorted sa ns are th ir'n sa do not supTort tvpica'l scrub ccununities. At such sites u~nderszory wcc p .anzs of tie scrub comunitv are mixed with srecies normally present in DJne fjat.ocdS of nearby areas.

The 7S- ogrcwth in these xeriz c .ranities ani rejects as unsuita0:ef ts true sanc cin- scru. So ilus. 71 e

special habitaK requirenonts rf *-he t'lrida Scrub Ja'r. a









Scr Jy is presence below. Thes- sitcs were ch.san to demcnstratZe the Vaie:- of ulant crmunities that Florida Scrub Jays select for nesting. Hicoria, 'iihlands Count,


Elevation of Hicoria is 140 feet. As described earlier. this site has artificially open areas, and after effect of a now deserted and razed town (Fi. 4) ; at least 60 per cent of this 200 acre study area is com-posed of open sandy or grassy patches. The scrub thickets consist primarily of the three scrub oaks (C0 per cent of the ground cover in thickets) with a scattering of the other planr's, principally Ser noa repons, Sabal etonia, _iTcoria floridana, and M-yrica cerie0ra. The thickets averaa 10 feet in height and a though each is small the woody plants are dense. Scrub Jay nests occur oredominantly in oaks, almost always in a dens thicket but adjacent to an open soace.

The degree o crass cover (Aristida stricta) and the presence of scattered trees of QuErcus laevi s and Pinus elliotti indicate that this site is a comaination of plants from several communities, i.e. scrubby flatwoods. From t:.e nunaner of Scrub Jays breeding at Hicoria it obviously contains ontimal features that Scrub Jays select for red~ing areas. The population density was 10 nests per 100 acres.


























Figure 4 View of Hicoria study area, center of territory
6.






57


1!
I

r ~*. ~ eq
I-.



~ ,r &



$








4 '~
-~

A






4~ .4 :jIA
I



4.


'W


4;





:3


Ca-e Kennedv Air Ezrcc Ste.ion, 3:.7ra d County


Elevation is 6 feet. T hs large area of approxiraely 1200 acres of scrub sustains a large population of Scrub Jays. This community is on recent shore front sands and is unlike most interior scrubs because wax mvrtie (30 per cent of :rcund cover in thickets) and saw palmetto (25 per cent of grcund cover in thickets) are much commoner. Here the three species of oaks are prevalent (35 per cent of thicket ground cover) The thickets are 8-12 feet high and dcnse, especially near the shore line where onshore winds and salt spray have stunted the upward growth of plants and produced an axLrearely tangled, Lo7 thicket.

The outstanding feature of this site is a network of interconnecting dirt and paved roads crisscrossing the area. All these roads have cleared shoulders 30-50 fec: wide, often of open white sand. Of course, there are large open spaces with rocket launching comrnlexes, but in additi to these industrialize_ parts, open -a:-6y places co7rise onout 40 ner cent of thi3 study area. D uing 1969 censi s. t.;o -plots of scrub thicket-s in the center of the 1_ -:iere there wore y oen soaces but away from t!,e rcketry =xplexes; St each I found 10-.1 active nests per 130 acres. This cape region with many artificial open spaces rciobles Hi ar and the density f bre'3in bird-3 is sioular. Most nests are located in sc;ub'y oacs aLthc 1 seCAl were found in the ;:.





~i:I


Jon at-_a.n c'-_ Z-~ St~~~k X= n n-~ Coun, 'r Elevation is 12 faet. This park preserves a large sretch cF sanc Pi-e scre u A. Aithouah sand cines cover much of this regicn, there are extensive scrub thickets without sanz cine cc'er, and Scrub Jays nest in much of this area. ::rtle oaks dominate this habitat comprising 30 cer cent cc around cover in thickets although the other two cak- species are -resent and make 10 per cent of the thicket ground cover.

The low scrub thickets without sand pines are rather dense and J-r: larce stretches of habitat with few ocen areas except along roadwavs. A census in this extensive cover shc';ed six nests per 100 acres. But in developed areas of the cark where camping facilities are scattered through the low dense sorub, I Sfou'd 9-10 nests per 100 acres.


Mlt. Pl-mo'tn Area, Crance Count'


elevation is 130 feet. Just west of this small town is an extznsive region o_ low scrub thickets without any sand pines. The wooc7 Ulants here are relatively low, 4-7 feet hich. Numerous ev een cdv species occur, although again. the oaks predominate. This is one of the few areas of scrub thickets where numerous olen spaces ocu--int------sd among the denser thickets and aparently notcas Some fom of haman interfc.;rence. Approximately









40 per cent 'vf -e crt :s open sandy s-aces, which are

Practically ceviz cc i:-- ncrbaceou cor:er. A nesting census catec. r-1- ests -er 100 acres and all were found in oak trees


Lake Jac7:scn, ietrinz, Hiihlands Co1unty


Elevazin is 11G, feet. Undoubtedly a fire destroyed an extensive sar.d -ine forest at this tract on the west shore of te k at least ten years ago to judge by scars on a few lae sand tines. Today a young scattered sand of sand nine is present, but most of the region has dense thickets of woody plants 6-10 feet high. As usua2, oaks dominate, but Labal etonia and !"ex cumulicola are cozron. There are -a-ural open sandy spaces, but a new housing development ?as a series of parallel rcads through this scr-, and an unaltered strin of thicket 100-130 feet wide between roa.s has been left. The developer, thus, has produced a series of open spaces, often of white sand, throughout the dense scrub thicket. No nesting census was made, 3ut 1., diiv of Scrub a s .s a vcvs high ana reminded :L s the densities of jars at Cane Kennedy and Hicoria.


Ridre Scr-b icnhlands Co ur.tv


E evaticn is 1J feat. ';ide oxcanses o1 very lo dse

scrub exists along U.S. Highway 27. two miles east of Hicoria. r7 >Cz 71 ar e C. Z e 5J:~ ~~~




61


this terrain. J'dfing b- large sani pineE scattered along the highway: ruch o- s area -a nave been covered with extensive sand in for-st. Scraz thickets extend several miles in a s as a sl dense round cover 3-6 feet h gh. The three scrub cak species dominate (60 per cent of ground cover in thicket) and Sa':a -tonia is comron (20 -er cent of ground over in thicket) (ig. 5). Although grasses are infrequent and the surface is usually bare white sand, very few large oen spaces accear. It is a difficult area to census fcr nes-in- Scrub Jays, because cf the imoene-rable nature of the low woody plants. Nesting jays occur over much of this terrain but congregate along road edaes or about the few ocen sandv spaces. During 1968 and 1969 two areas were censured. Approximatesl- 3-1 cairs nest ner 100 acres. but in the densest areas away from o:-en spaces jays rare>, nested.


Scru~bybv F' oods 7est of Hicoia, -chlands Count:7


Elevation is 130 fee:. Just west of the scrub covered

tillsc ninis Euzzyan extensiverei o l ca

exists. Within three miles of the ridge, numerous small nar- -O-7 srr :~' ~ Poal n

were snaIl sand bars in a Pleistocene sea. The thickets on these strips are extremely dense. The eicT> these

l thickets are usual> b= ---S -slonali' ;;a.>; spacef scrubs of rosemary.

c

es-ecia"Lly h closc- -o -t-e s-x-z7nsi-ve scrub ridge a- tho

























Figure 5 View of low, dense scrub thicket two miles
southeast of Hicoria, Hichlands County.
The only open spaces are along the dirt
road. In the background is a patch of sand
pine without nesting scrub jays.

















ato






*+

a-


&


'Ma -~ w




'0 '


Arc.- bld Station and 'icoria. Although large numbers of breeding Scrur Jays aro suwported in these scattered islands of scrub thickets, the density per unit area is rather low (2-3 pairs per 100 acres).


Atlantic Coast Dune Thickets


An extre nely long yet narrow band of very low and dense thickets exists on shore-front sand dunes of the Florida Atlantic coast from Jac>3sonville to Fort Pierce. Sand pines are not present and saw Palmetto often is the dominant plant. Live and myrtle oaks comprise from 20-40 per cent of around cover in thickets. Onshore winds and salt spray have led to an extremely dense, often impenetrable; ;rowth (Fig. 6). Open areas exist only along roaC;;ays or where houses have been built.

The vegetation 4s similar all along the coast but

the distribution of Scrub Jays varies. The snore thickets about i-elbourne Beach, Brevard County, 20 miles south cf the large Cace Kennedy scrub area contain many scattered breeui2g coLonies. The beach thicket north of Cape Kennedv toward NEaw Smvrna Beach, Volusia County, also contains numerous breeding Scrub Jays. These birds nest only along the ed
North of Daytona Beach, Volusia County, in the same type 'f vegetation, Scrub Jays are much less au:. nn. Se'ZYa1 mils o1 seemi-1y suitable thickets exist before a ot 4--






















Figure 6 View of low and extremely dense Atlantic coast
thickets at Marineland, Flagler County. Salt
spray has retarded the upward growth o the
plants producing an extremely tangled thicket.
The only open spaces are along the road edge
at the extreme foreground of the picture.




















- -








"BigScra: -::


levation~ is 1.e:. -his is ar exzanse C1 scrub

1.7 nies iie an I. niles 1cng, locate Li -s west of

Bellevieu:. DEnSe stands of sand pine cover much of this area withno rreei J .'-at large stretches of

low cense thickets wichrut saId cines are Oresent. As usual the :round is usually open sani with few hers, but larve open spas r lac:ir. The three oak species occuy 30-40 per cent of the ground cover in thickets, and Ilex :viu1icola and Lyon f~vzaiea are cornmon (30 per cent Of grcun' cover in thicket) This vecetation varies from 2--5 fe-t in he -t

The on"-- -art that contains breeding Scrun Jays is the narrow zone alono State Road 464. Here jays ar= fre:uent-ly seen :eecing alcnc tn c'en road;:av, but onehalf mile into- the scrub rcm the roa no Scrub Jays are zound. Eight miles ncrth of this road, Interstate 72 crosses a Oeco:n of scru.. Pars of the scrub bordering .Int~erstate % are simi~lar to that alcng faate Road 434, yet no 7cru Javs nest Here. Interstat-e 73 is a relatively, new road, and awparently Scrub Jayz h ve not located this ne;\l coe n-td h a.i


Ccala Tationa1 --orest on CoutElevation 4- 13C fozt. Clt ^ __ 4: the are

s -- Lorida, fZwoparts -F ->is extensive

-ecv..c s2 rcr a .::s:. h of this area is









covered ;,ith relative dense stands of sand Line with associated Blue Jays. B3t many low dense scrub thickets exist and as at t2e "i i S-rub" to the west, most of th e have f&w onen spaces eind. few nesting Scrub Jays. Breeding birds congregate aloncT r-oacdways, whose edges are frc-iE: ---used as feeding sites.

Only at cts southern edge of Ocala National Forest are Scrub Jays numerous and here the scrub thickets away from the roadsides are cften inters-cersed with open spaces and sand oine are not abundant. Here some mingling of scrub and tu'ke- oak comunit-. occurs, and apparently; the resulting ecotone frecuentlv contains.-- oen sac2s. E-Tn here most breeding birds nest along roadways. Habi tat Summary


The Florida Scrub Jay requires low, dense, largely everarean o0. Thickets for nesting and extensive open spacc for feeding. At the 7rchbold Station, Amadon (1944) fund Scrub X&'s concentrated alcng the edges of man-made clearinas. This tendency is especially obvious throughout its breedin range in Florida. The largest concentrations of nesting birds are found in scrubby thickets that have been oi-neJ p :y man, such as Cape _Kennedy and Hicoria.

Although nesting Scrub Jays almost always select xeric habitats, most Florida sand pine scrub is unsuitable because



so e' f ai te 21ee.n.sfr .ense thioko:j intcerscerseL









with open sandS -ac-s. csricZ them to certain bcrd:r zones such as scrummv :lawoods, co recently burned regions, and es-ecial now tc road-ays and other disrupted scrub areas produce b' hu:an :nter:erence.

Usually whereverr the zrcper ocxbination of dense evergresn oak th'ickats and open sandy spaces exists, sore nes-ing Srub Jays are found. Thus small populations occur at Cedmar Ke, Levy County; at Camp 2ianding, Clay County; and scattered alcnq the Atlantic coast often miles from other breeding Scrub Jays. Although such habitat is scattered over much of central Florida, it is normally in small acreages oZ open thickets. The distribuzion of Scrub Ja breeding habitat indicated in -ag. 7 illustrates the restric-ez a-ount of land available to nesting Flcrida Scrub Jays. This map also shows certain areas that acpear to have suit le Scrub Jay habitat but lack resident ulins. Sor.e of these areas are disS 7-7 4 71- ,fJnc Q--:1 7 p,-*
Scrb EiatTitou Sredi ___ru Jays


A fe"'t raher sizea< thickets with aparently suitable habitat have nc resident Scrub Jays. Several of these scrub chick ameas ace indicated in Fig. 7. Xis simee-Sai:t Cloud, Oscla County


Many so.L scrubby th~ cKets exist about this region

t ':a r h- ca et a c n ad o- 3. d Orance cfroves. The small
























Figure 7 '.a, of co:tral region of Florida ceniis"la
illustrating areas apparently suitable for breeding Scrub Jays and areas where breeding Scrub Jays were located. J.D.S.P. is
the Jonathan Dickinson State Park.







71


PEACHI
K EY


O BrAT SUa BLE FCP CAMP BL! D NG NE STING SCRUB JAYS


OR N ESTING SCRUB JAYS PRESENT


I IZCH IS MILES MAR.NELA4D
N








NOCA.A CAfTONA

I EY NAT. S KACH




SCRUB FOREST




11T
PLYMOUTH





nAYPoTty W EE
j'NECAPE KENNEDY KISSIMMEE










L LAKE WALES










0 .~ SEERINS
S E N F O R T P IE R C E










OD VE MUS

-. ~I.TS





72


s c r u by thic s, _Len eerely 200-300 acres, appear to be excel1 lent Sr Jay cover and resermbI the scrub at Mt. Plyv -..h. Uc Scrub Jays exist in this region, but 20 miles to tLe -or similar habitat supports numerous scattered con actions of Scrub Jays. Week V e a? -do Ccunty


An extensive scr: Cxi sts about this natural spring. Much of it is covered with sand cines and so cold not be elected to succort Scrub Jays. Small sections have reCentl- burned, ha;ever and low dense thickets are -rowing back, with nu r- us onen sandyx sc:aces. >:oreova:, a1o g the southern border of this scrub much openness exis-c in a broad ecotone between the scrub and turkey oak vegetation. I estimate approximately 1000 acres of low ooen scrub thickets exist here, but no Scrub Jays are present. Scrub Jays reside in small nu-bers 20 'ile -o the south and 28 .iles to -e east.


Lake Kaes Polk CountA few islands of scrub thickets, similar to the ones west of iL-ccria, are present to the west of Lake Wales, but no Saruv Jays reside on them. The plant ccn:Gunity on the sand ri close to these scrub islands at Lake Wales

is turkey oa vegetation without any Scrub Jay population.





73


A ver lng strE::':. of sand nine scrub exists along

the Gulf Cc&st frc-i 2.eresa, Frankland County, westward into Alahaua. ::ucn f thcis scrub supports sand pine forest, but man7 extensive acrea-es of low dense scrub oak thickets occur, with onumerus open sandy stretches. An especially larre exransa of tis type of habitat exists on Santa Rosa Island south of Pensacola, Santa Rosa County, and especially at Fort Pickens State Park (Fig. 8), but nowhere in these several hundred miles 'of scrub habitat are t-here any Scrub Jays. The nearest S-rub Ja = Dopulati= is near Cedar 7Ke-, Levy Ccunt, 100 mi les to the southeast of the eastern border of the Panhandle Scrub.


























Figure 8 View of low scrub thicket with open sandy
spaces at Fort Pickens State Park, Santa Rosa Island, Escambia County. This type
of scrub thicket supports large numbers of
Scrub Jays when present in the Florida
Peninsula, but here no jays occur.






















































4 V IA7


41

















The Florida Srub Jay territorial croclivities are as intense as tnese escribe- by Nice (1937) for the SonSparrow (elosniza :elcdia). These jays defend a territo rv that normally includes eating, nesting, and feeding grounds, and thus belongs in territor-y category "A" of Nice (1941). The Florida Scrub Jay forms a pair bond that is retained throughout the year, and frequently> a rair defends the saMe ttory for more than three years.

This pattern of territoriit is similar to that

described for most western races of A. o er l o s bcn Hardy (1961) and Westcott (in press). Eut the Florida race exhibits a degree of sociality not found amog- its western relatives, as first-year birds are retained as "helpers" in the terri Discussing the social structure of the Arizona roculations of the :exican Jar in

itcra-il--- is ackino "n o floc.s are tolerated

at the nest, Eardy (1961) suggests that a species that has a restriott. incunt of habitat available for nesting may adapt by allowing a social breeding structure. Thus the aexican which nests mainly in the narrow riparian

woodland of southeaster- Arizona, talerates flocks of nonbreeding birds about the nest, but the western subspecies of the -a - _-aer anc-e of habitats

availc ._ _rio rerriteriality.


76









Ccntr a to (1961) -+ t. e Frida

Scrub Jay Is terir L a s m uch so as any of the ;estern

races. Yet in Ylorida Cc cerulescens inhabits as narrcw an ecologic zone as .. lraia i Arizona. The

habitat selected b- these two have nc features in common. A. u1tramarina selects the arborescent parts of the woodland that ara strioeito narro. riparian situations, vwhiie the -lzrida Scrub Jaw typicallv selects the xeric, .ow scrub tikets with open sandy s-aces and ignores the tall sand cine -orest and all the mesic forests of Florida.

Even thcach breedi ng Scrub Ja1s are scattered throu-h out much cf t-e n1o -id ieni nl a (,ia. 7), close =7aminai reveals that the amount or accentabla neszing nab]tat is restricter. often r 7 a border of open scrub tnickets along road:ays, the eg'es of the taller san. tine forest, or rece:-.tl L ed regions. On> limited areas consist of extensive scw t2:c:aeI ana nere 2t Is usua-y nan-maae interference that has produced the open spaces that attract dense breedino ojulations. Suitable 4ab tat to induce

Scruc Jays to :est Is re-t-r-icted both in type of cover and in to'al acreaze of this special vegetation. The tolerance shown v r-ng tairs of Florida Scrub Jays to certain nencers cc mhe co ation conceivably is a reaction to a restricted range o suitable habitat, caralleling the situation of t> :


miniu7?_=U4tOn size is crobabl-: necessary for survival,









and species with limited breeding range nlay be unable to maintain such a population level unless special social structures are inclucec. With the retention of first-year birds as territorial helcers," the Florida Scrub Jay has developed a degree of sociality as an adjunct to an essentially territorial nature.

This combination of sociality and territoriality is possible, for the breeding male may courtship-feed the "helpers" and so form a loose bond with these birds. All members of the territory frequently react together in territorial defense. Thus the family group remains territorial, unlike :exican Jay flocks that show no territoriality. Interestingly, compared with Scrub-Jays, the breeding male Mexican Jay utilizes courtship feeding infrequently and only with his mate (Brown, 1963; Hardy, 1961).

In Mexico and western United States many different

habitats are selected by breeding Scrub Jays, but riost are of a xeric nature and, as Pitelka (1951) points out, the Scrub Jay appears best adapted to feeding in and about the low dense thickets of chaparral or open oak woods. Even though isolated by over 100 miles from these western races, the Florida subspecies continues to utilize habitats of similar ecologic structure. Presumably' at one time a ontinuous scrub habitat connected the Florida peninsula with western habitats and Scrub Jays invaded Florida through this connecting habita-t (Pitelka, 1951).

Whether : not large expanses ci suit : habi ta e,,:er existed in Florida is difficult to ascertain, but









Laessle (1953i stones t8)t sand pine scrub areas have Probably existed os presently situated since the formation of -Ecause this xeric community

exists onl on ;efi----ad dune sands, and Florida Scrub Javs require ic: thickets with open spaces that occur mainl.y at. s:Cub borders, I doubt if Scrub Jay habitat was ever wide3sread In Florida.

Hardy (1961) speculates that A. c. coerulescens has survived in Florida as a relict population that may have preserved certain traits indicative of a pr tive condition, i.e., sociality and supposed lack of territoriality. Evidences from the present study suggest the opposite, i.e., territoriality as a basic and possibly primitive trait and social nesting as behavioral adaptation to the special restrictive characters of the xeric scrub thickets in

Florida.

Although western Scrub Jays, especially first-.ear birds, frequently travel great distances away from the nesting region (Westcott, 1969), the Florida Scrub Jays are sedentary. As fi:st-vear birds normally remain in the parental -erritories, dispersal of juveniles is infrequenc. lceria monezen: of local birds to new breeding areas was :oestried. SCe settled to nnst close to parent rri in unoccupied habitat, and a few

inaividuals, usually adults, roamed as far as 1.5 miles Ve.'-e But few long-ranve novenents have been

observed. >:n.oo:' (19 ) C~eseciOJ oone td on the




so


sedentary habits o f c rida Scrub Ja. Only following some majcr ca-a.stro:e, as a ibull-dzzing of large areas ci szaub, -ere Slcrica Scrub Jays found r from scrub bimat.

Only a fe7 areas ccur Jr. Florida with broad expanses of owen zcruzzy cover and suowrt relatively larce and dense op ions of Scrub Jays such as the Lake Placid

areas (Archbold Station, Hicoria, etc.), Jonathan Dickinson State Park7 Cape Kennedy, and the ',t. Plymouth region (Fig. 7). Not only do Scrub Jays abound in these areas, but nearly all similar habitats nearby have resid-nt Scrub Jays. Presum-ably eno-, r ros i-'-bt these rc-hl-tion centers so that most small scrub thic-et5 ar5 locates and colonized by the jays. Similar small sites further away (10-2; miles) have only a few scatt-ered airs of birds or rnzr: at all. The density of nesing Scrub Jays gradually dacre=ses alonz the coastal thickets north :rcr Cawe Kennedy. North of Daytona 3each nesting Scrub --as are noticeably less frequent, and from Marinland to St. Augusti'e nestin7 ai r s-arat- -a

miles of low dense thickets even t:ouah the habitat is similar to the e1Sel i nhabit hcts just cureh

of Cace 7 an:nedy.

Similar patterns of densit.- of this species occur aboit the other large centers o i- *zooula i:n. Close to densely inhabited thickets most small scrub habitats nave nest-2r birds, but harthe: awavfeer birds are









fon. ialysm ites, usa Lly well isol at hve

no Scruz :av5. .a:TzCn t-.se scrubs without

j. -...-. of-.g c ~-.. -. e-;.... appear to meet the

Florida. s szeco ecoloa:cal recTuireents.

The s. of luve:.iles n the generally

short distYnce o: 3isersal b< no-breedirg afu.lts mus eXlai2 the XFence of nesting ScruB Jas from suitable habita.. s ca s-rucur allc::inZ for a degree of
1' -7
si du~c iErsal ruverile birds which

account fcr =ncst of the dis-ersing iniviua s in western races (Westcott. 1969). As "helcers" they remain in the parental te:rt:.ory fr e ee periods, which .-y perrit an increase. failiait7 of that specific size or the surrouncino habitat. Thus a forc o site tenacity ma be present i the younger nondroeding birds and may reduce the distance of dispersal 7,en they do move out of the



iLng t '.e:ets -u- ocu., as itM'

with nesting jays are isolated b many miles from th nearest SCrub f7 copulazicn. Ucaoub- L-,, a -o-. inc: vidals dis-crse long distances, but freuen-li 'x>n Scrub Jays ar found in unlikely habitats some major

catstczh~z ocuredto a J ccr-'- : rcz tne

hirds to move. ?robah> man- distartlv isolated sites ae C y s ch irdf. erhnzs such ca'.aszrophes

SC-Iu. Ja-. 3 fr::. isolated yet suitable"+:g~r: Afir ::a hav dete- Zd le l10 scr7




82


frcin resent -i rd away; an-z. -nerin birds may t have located the nM-C-- aailah habitatafteritrercws.

The extensive 'an>andi: :crub is 100 miles from the nearest small Scrub Jay cdllny. If straying birds cannot occupy I the suit:Ilo sites in central Florida, then they should take many years longer to invade the panhandle region.

The adactaticn of a type of sociality by the FIorida Scrub Jay has produced certain restraints on the distri.bution of these birds. Because the- have become exceticnallv

sedentary, and dispersal abilities have been reduced, some areas cf sui able a'it at in Canzral :Lcri:a and a Iar e expanse o7 habitat in the Pan>a le are unccupied today.


















Amadon, D.
1944. A preliminary life history study of the Florida
Jay, Cvanocitta c. coerulescens. Results Archbold Exped. No. 50. Am. Ilus. Novitates, No.
1252:1-22.

Bent, A. C.
1946. Life histories of North American jays, crow,
and titmice. Bull. U. S. Nat. 1us., No. 1'1.

Brown, J. L.
1963. Social organization and behavior of the Mexican
Jay. Condor, 65:126-153.

Hardy, J. '.
1961. Studies in behavior and ohylo-eny of certain
New World jays (Garrulinae). Univ. Kansas
Sci. Bull., 42(2):13-149.

11inde, R. A.
1961. Behavior. In A. J. Marshall, ed., Biology and
Comparative Physio logy of Birds. i. New York.
Academic Press.

Grimes, S. A.
1940. Scrub Jay reminiscences. Bird Lore, 42:431-436.

Laessle, A. :.
1942. The olant communities of the Welaka area. Univ.
of Florida Press. Biol. Sci. Series, vol. 4,
No. 1:1-143.

1958. The origin and successional relationships of
sandhill vegetaticn and sand -ine scrub. Ecci.
Monog., 28:361-387.


1950.



.L037.


Q .
Pleistocene shore lines in Florida and Georaia. U. S. Geol. Surv., Prof. Paper 221-F, pp. 95-107.


studies in the life history% of the Song S'arrov. I. Trans. Linn. Soc. N. Y., 247--.


N. oa o t ror in 1r2 i r .


83




84



Pitelka, A.
1945. Pterylograohy, molt, and age determination of
American 7a-'s Of the genus A-helocoma. Condor,
47:229-2')".

1951. Speciat*cn ndq ecoo0gic distribution in Americ:n
jays of the genus Aohelocma. Univ. California
U -ZOO ., 70:195-464.


193:. H'p1cers at the nest. Auk, 52:257-273.

1961. Helpers among birds. Condor, 63:198-226.

Webber, H.
1935. Tne Florida scrub, a fire fighting association.
Amer. Jour. Bot., 22:344-361.

Westcott, P. W.
1969. Relationships among three species of jays wi-atering in southeastern Arizona. Condor, 71:353-359.

Westcott, P.W.
In cress. Territorialit'v of the Scrub Jay in Arizona.
Condor.














BITJ:PhICAL KBTCH


Peter WValter .estcott was born 3 November 1938 at Mt. Vernon, Ue. York. In June, 1956, he was graduated from Madison High School, aisn New Jersey. In June, 1960, he received the degree of Bachelor of Arts with a major in Zooioc; from ,- ers, Colege. In 1962 he enrolled in the Graduate School of the University of Arizona. He worked as a graduate teaching assistant in the Department of Zoology and as a graduate research assistant in the Department of -.irolav until June 1962, when he received the degree of _aster of Science with a major in Eoology. From 1963 to 1966 he served in the United States Navy and was staticned in Japan and Ncrfolk, Virginin. From January, 19r7. ntil the -resent time he has pursued his work toward the degree of Doctor of Philosoohy at the University of Ftorida, ;ich a major in the Department of Zoojocv.

Peter alter estcott is married to the former ar7Le : Yin:. He is a member of Sigma Xi, American OrnithlcIist3' Union, and the CooPer Ornithological Socie t.














This discc.3tio wus -rf arI nd r4 the direction of the chairman of the candidate's supervisory committee and has been appro;Lvedy all members of that committee. It was submitted to the Dean of the College of Arts and Sciences and to the Graduate Council-, and was approved as partial ful.fillment of the requirements for tihe degree of Doctor of Philosophy. August, 1970.




Dean, College ofArts and Sziencen




ealr,~cGraduate Sclhool Supervisory Commi ttee:



Chairman










I -




Full Text
68
covered with relatively dense stands of sand pine with
associated Slue Jays. Bat many low dense scrub thickets
exist, and as at the "Big Scrub" to the west, most of
these have few open spaces and few nesting Scrub Jays.
Breeding birds congregate along roadways, whose edges
are frequently used as feeding sites.
Only at the southern edge of Ocala National Forest
are Scrub Jays numerous and here the scrub thickets away
from the roadsides are often interspersed with open spaces
and sand pine are not abundant. Here some mingling of
scrub and turkey oak community occurs, and apparently the
resulting ecotone frequently contains open spaces. Even
here most breeding birds nest along roadways.
Habitat Summary
The Florida Scrub Jay requires low, dense, largely
evergreen oak thickets for nesting and extensive open
spaces for feeding. At the Archbold Station, Amadcn (1944)
found Scrub Jays concentrated along the edges of man-made
clearings. This tendency is especially obvious throughout
its breeding range in Florida. The largest concentrations
of nesting birds are found in scrubby thickets that have
been opened up by man, such as Cape Kennedy and Hicoria.
Although nesting Scrub Jays almost always select xeric
habitats,
most Florida
sand pine scrub
is unsuitable because
the pines
are in dense
stands. The FI
orida Scrub Jay's
specific
habitat prefe
rencas for dense
thickets inierspersed


BIOGRAPHICAL SKETCH
Peter Walter Westcott was born 5 November 1938 at
Mt. Vernon, New York. In June, 1956, he was graduated
from Madison High School, Madison, New Jersey. In June,
1960, he received the degree of Bachelor of Arts with a
major in Zoology from Amherst College. In 1962 he enrolled
in the Graduate School of the University of Arizona. He
worked as a graduate teaching assistant in the Department
of Zoology and as a graduate research assistant in the
Department of Virology until June 1962, when he received
the degree of Master of Science with a major in Zoology.
From 1963 to 1966 he served in the United States Navy
and was stationed in Japan and Norfolk, Virginia. From
January, 1967, until the present time he has pursued his
work toward the degree of Doctor of Philosophy at the
University of Florida, with a major in the Department of
Zoology.
Peter Walter Westcott is married to the former
Marybeth King. He is a member of Sigma Xi, American
Ornithologists' Union, and the Cooper Ornithological
Society.
85


LIST OF FIGURES
Figure Page
1. Map of study area at Hicoria 1968 6
2. Map of study area at Hicoria 1969 20
3. Map of study area at Hicoria 1970 22
4. View of Hicoria study area 57
5. View of scrub thicket two miles south
east of Hicoria 63
6. View of Atlantic coast thicket at
Marineland 66
7. Map of central Florida peninsula with
Scrub Jay habitat 71
8. View of scrub thicket at Fort Pickens
State Park 75
v


73
Florida Panhandle Scrub
A very long stretch of sand pine scrub exists along
the Gulf Coast frcm St. Teresa, Frankland County, westward
into Alabama. Much of this scrub supports sand pine forest,
but many extensive acreages of low dense scrub oak thickets
occur, with numerous open sandy stretches. An especially
large expanse of this type of habitat exists on Santa Rosa
Island south of Pensacola, Santa Rosa County, and especially
at Fort Pickens State Park (Fig. 8), but nowhere in these
several hundred miles of scrub habitat are there any Scrub
Jays. The nearest Scrub Jay population is near Cedar Key,
Levy County, 100 miles to the southeast of the eastern bor
der of the Panhandle Scrub.


51
distance
rare and
such as
movements by Florida Scrub Jays apparently are
normally must be induced by some major catastrophe
destruction of the habitat by fire or clearing.


METHODS
The social behavior is based upon information gathered
from the banding of 230 Scrub Jays at Kicoria and at scat
tered sites to the west and south- To simplify the report
discussion is limited to movements of banded birds in the
200 acre study plot outlined in Fig. 1, the center of the
Hicoria region.
Jays were captured in mist nests and baited Potter
traps. They quickly learned to avoid the mist nets, and
various methods had to be devised to lure them in. Most
effective was the use of a hand-held Scrub Jay which, when
shaken slightly, gave loud alarm calls. This attracted
many local jays and I rhen chased them toward the nets.
Each bird received three bands, two colored plastic
coil bands and or.e aluminum silver colored United States
Fish and Wildlife Service band. Each individual was
identified by a series of letters connoting .the color
sequence of bands: RS-R was a bird with red plastic and
silver aluminum bands on the right leg and red plastic
band of the left leg, 1B1G-S had light blue and light
green bands on right leg and silver band on left leg.
All birds captured in traps or nets had the silver band
placed on the right leg unless definitely known to be a


4
To the east, this site is separated from the main
body of the sand ridge scrub by extensive orange groves.
The western edge of the study area is at the border of
the sand ridge and drops several feet to pine flatwoods,
although many scrub thickets continue to the west. Scrubby
vegetation extends south for a mile and then merges into
pine flatwoods with widely scattered scrub thickets. The
scrub at Hicoria continues north along a railroad in an
unbroken, although narrow, band to the Archbold Eiological
Station.
Little human interference occurs at Hicoria. Occa
sionally a hunter stops to shoot at a rabbit, someone cuts
some hickory for barbecue wood, or checks on the cattle
run on the extreme north edge of this area. Otherwise I
saw few people during my three years of field work at
Hicoria. Although Florida Scrub Jays are notoriously tame
and in many areas readily follow people to beg for food,
at Hicoria the birds usually ignored my presence. I
regularly camped in the middle of the study area, but
took pains not to leave my food about.


75


27
breeding season the pairs spend more time wandering out
of their territories and mixing with other Scrub Jays.
Yet whenever the pair returns to their territory they
chase intruders. In this fashion the boundaries vary
little from one season to the next.
Apparently the Florida Scrub Jay mates for life. Two
of the pairs at Hicoria are known to have been paired for
at least four consecutive breeding seasons, RS-R with v'7S-Pi
and GS-G with S-RB. WS-Pi, a female, is of special interest.
I banded her as an adult in the summer of 1960 at Hicoria
within 200 yards of the territory now occupied. She is
thus at least 12 years old and still breading.
Mate constancy was the rule with the majority of the
other mated pairs. The following pairs were together during
1968 and continued as pairs through 1970: SR-B with
S-BO, SB-N with SB-dB, RS-Pi with MS-B, and SG-B with
SW-R.
Only three pairs broke up in this 200 acre site.
NS-N, S-WN and young disappeared in late October 1968.
I have no specific knowledge as to the cause- of their
disappearance, except that Scrub Jay feathers were found
in the territory. I surmise that some predator killed
one or more of this group, probably including one of the
parents, although two of the young were later found asso
ciated with a group of non-territorial birds 500 yards to
the northwest. The parents and remaining young were not
located during the next two years, although I roamed over


COURTSHIP
Unmated birds frequently travel and feed in flocks,
but no special group activity relating to courtship has
been found. This is unlike the Blue Jay, Cyanocitta
cristate (Linnaeus), which performs a highly ritualized
group activity (Hardy, 1961).
Pair Formation
The initial stage of forming a new pair bond begins
with the male restricting his movements to a particular
undefended area of scrub. After the male is first found
restricting his movements, a female soon accompanies him.
On 10 January 1970 M-dBS, a male, had restricted his
movements to a region just west of territory 4. He con
tinued feeding in territory 4, and five days later S-BY,
a female, was first seen following the male. After this
date they rarely returned to the birds in territory 4 and
the male began to defend actively their new region. I
first saw M-dBS chase another jay from the new region
on 15 January, and S-BY followed him. The female invariably
followed the male during the next few weeks as he wan
dered through the territory and especially when he chased
intruders. Such constant accompaniment undoubtedly helos
reinforce the pair bond.
14


79
Laessle (1953) states that sand pine scrub areas have
probably existed as presently situated since the forma
tion of the terrace ridges. Because this xeric community
exists only on well-washed dune sands, and Florida Scrub
Jays require low thickets with open spaces that occur
mainly at scrub borders, I doubt if Scrub Jay habitat was
ever widespread in Florida.
Hardy (1961) speculates that A^_ cu_ cosrulescens has
survived in Florida as a relict population that may have
preserved certain traits indicative of a primitive con
dition, i.e., sociality and supposed lack of territoriality
Evidences from the present study suggest the opposite, i.e.
territoriality as a basic and possibly primitive trait
and social nesting as behavioral adaptation to the special
restrictive characters of the xeric scrub thickets in
Florida.
Although western Scrub Jays, especially first-year
birds, frequently travel great distances away from the
nesting region (Westcott, 1969), the Florida Scrub Jays
are sedentary. As first-year birds normally remain in
the parental territories, dispersal of juveniles is infre
quent. At Kicoria movement of local birds to new breeding
areas was restricted. Some settled to nest close to
parental territories in unoccupied habitat, and a few
individuals, usually adults, roamed as far as 1.5 miles
before settling. But few long-range movements have been
observed. Amador. (1944) especially commented on the


30

of the seven "helpers were first-year individuals that
had been banded the previous year in the nest of the pair
with which they were new associating. The other was S-EY,
an adult, and will be discussed later.
Finally in 1570 six of nine territories and nests in
the sane Hicoria site had "helpers," one in territory 1,
one in territory 2, one in territory 3, one in territory 5,
three in territory 4, and one in territory S. All of these
"helpers" had been banded the year before as juveniles and
in the sane territory.
Approximately half of the territories at Hicoria had
extra members. Banding data show that they were invariably
juveniles of that pair from the year before, with the excep
tion of S--BY noted previously. Although reminiscent of
the social organization of the Mexican Jay (Aphelocoma
ultramarina) in Arizona, in which nonbreeding birds
frequently function about an active nest (Brown, 1963),
this case is special in its combination of territoriality
and sociality (c.f. Skutch, 1961).
The territories that had no "helpers" during 1969
and 1970 were those in with the fledglings either were
destroyed by predarors or disappeared from the study area
and surrounding habitat.
Interactions of Adults and Juveniles
re
After the young leave the nest thev become
ring, remaining near the nest site and very
extremely
cruiet.


Figure 7
- Map of central region of Florida peninsula
illustrating areas apparently suitable for
breeding Scrub Jays and areas where breed
ing Scrub Jays were located. J.D.S.P. is
the Jonathan Dickinson State Park.


55
Scrub Jays is presented
demonstrate the variety
below. These sites were chosen to
of plant communities that Florida
Scrub Jays select for nesting.
Hicoria, Highlands Countv
Elevation of Hicoria is 140 feet. As described earlier,
this site has artificially open areas, and after effect of
a now deserted and razed town (Fig. 4); at least 60 per
cent of this 200 acre study area is composed of open sandy
or grassy patches. The scrub thickets consist primarily of
the three scrub oaks (80 per cent of the ground cover in
thickets) with a scattering of the other plants, principally
Serenoa repsns, Sabai etonia, Hicoria floridana, and Myrica
cerfera. The thickets average 10 feet in height and al
though each is small the woody plants are dense. Scrub
Jay nests occur predominantly in oaks, almost always in a
dense thicket but adjacent to an open space.
The degree of grass cover (Aristida stricta) and the
presence of scattered trees of Quercus laevis and Pinus
elliottii indicate that this site is a combination of plants
from several communities, i.e., scrubby flatwoods. From
the number of Scrub Jays breeding at Hicoria it obviously
contains optimal features that Scrub Jays select for breed
ing areas. The population density was 10 nests per 100
3.C2TSS


24
a short time another flies up and replaces him. A bird
may perch there for as long as 30 minutes. The high
bird invariably is first to sight an approaching intruder
and begins bobbing and calling, usually with upwardly
inflected kwerr. At once the other birds fly or hop into
the branches nearest to them. This position of the bird
is undoubtedly a form of sentinel behavior both during
and after the breeding season. Yet on such a perch not
only can he see over his whole territory, but others
outside can readily see him. Possibly, then, this posture
may function as a means of proclamation of territory.
Rarely does an outside bird intrude far into a
territory during the nesting period, but boundary disputes
are frequent. Chasing is the normal means of territorial
defense. When another bird approaches or crosses the
boundary the male of the invaded territory flies directly
toward the intruder. Physical contact is almost never
necessary. The encounter quickly terminates with the
intruder rapidly retreating. The attacking male veers
off abruptly at his territory boundary and almost never
extends his flight beyond it. His return flight often
includes a deeply undulating flight and a series of loud
rapid kwerp calls. At times the initial attack by the
defender is silent; when a call is used it is a loud
kwerp.
The mate often follows the male but rarely takes
part in any dispute. She perches nearby in the territory,


77
Contrary to Hardy's (1961) statement, the Florida
Scrub Jay is territorial, as much so as any of the western
races. Yet in Florida A. c coerulescens inhabits as nar
row an ecologic zone as A. ultramarina in Arizona. The
habitat selected by these two have no features in common.
A. ultramarina selects the arborescent parts of the wood
land that are restricted to narrow riparian situations,
while the Florida Scrub Jay typically selects the xeric,
low scrub thickets with open sandy spaces and ignores the
tall sand pine forest and ail the mesic forests of Florida.
Even though breeding Scrub Jays are scattered through
out much of the Florida peninsula (Fig. 7), close examination
reveals that the amount of acceptable nesting habitat is
restricted, often merely a border of open scrub thickets
along roadways, the edges of the taller sand pine forest,
or recently burned regions. Only limited areas consist of
extensive low thickets and here it is usually man-made
interference that has produced the open spaces that attract
dense breeding populations. Suitable habitat to induce
Scrub Jays to nest is restricted both in type of cover and
in total acreage of this special vegetation. The tolerance
shown by breeding pairs of Florida Scrub Jays to certain
members of the population conceivably is a reaction to a
restricted range of suitable habitat, paralleling the situa
tion of the Mexican Jay in Arizona.
As suggested by Hardy (1961) for the Mexican Jay, a
minimum population size is probably necessary for survival,


63


40
I expected to find a definite dominance hierarchy
among the Scrub Jays at feed site 1, but did not. The
only bird to exert his dominance frequently was RS-R,
whose territorial boundary and nest was the closest.
The other breeding males, NS-N from territory 6 and
RS-0 from territory 2, occasionally supplanted the non
breeding birds but most frequently coexisted, often side
by side with both non-territorial jays and with most of
the territorial birds visiting this site. Although many
birds often frequented the site at the same general time
of day, usually only two or three dropped down for food
at once. As they flew off a few more dashed in. There
was little opportunity, then, for much conflict, or
supplanting. Many hours were spent watching the move
ments of jays at this site, and no definite hierarchical
ordering could be assessed except that breeding males
were generally dominant over most other jays.
In many bird species, the more closely two individuals
approach, the more likely displays or attacks become. Each
individual has a specific individual distance within which
the bird will not tolerate others, excepting sexual encoun
ters. Among the Florida Scrub Jays at Kicoria this trait
was not seen. Birds frequently fed side by side or spent
considerable time close to each other.


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59
Jonathan Dickinson State Park, Martin County
Elevation is 12 feet. This park preserves a large
stretch of sand pine scrub. Although sand pines cover
much of this region, there are extensive scrub thickets
without sand pine cover, and Scrub Jays nest in much of
this area. Myrtle oaks dominate this habitat comprising
30 per cent of ground cover in thickets although the other
two oak species are present and make 10 per cent of the
thicket ground cover.
The low scrub thickets without sand pines are rather
dense and form large stretches of habitat with few open
areas except along roadways. A census in this extensive
cover showed six nests per 100 acres. But in developed
areas of the park where camping facilities are scattered
through the low dense scrub, I found 9-10 nests per 100
acres.
Mt. Plymouth Area,, Orange County
Elevation is 130 feet. Just west of this small town
is an extensive region of low scrub thickets without any
sand pines. The woody plants here are relatively low,
4-7 feet high. Numerous evergreen woody soecies occur,
although again, the oaks predominate. This is one of the
few areas of scrub thickets where numerous open spaces
occur interspersed among the denser thickets and apparently
not caused by some form of human interference. Approximately


FACTORS CONTROLLING DISTRIBUTION
The Florida Scrub Jay territorial proclivities are as
intense as those described by Nice (1937) for the Sor.g
Sparrow (Melospiza Ilslodia) These jays defend a territory
that normally includes mating, nesting, and feeding grounds,
and thus belongs in territory category "A" of Nice (1941).
The Florida Scrub Jay forms a pair bond that is retained
throughout the year, and frequently a pair defends the
same territory for more than three years.
This pattern of territoriality is similar to that
described for most western races of A. coerulescens by
Hardy (1961) and Westcott (in press). But the Florida
race exhibits a degree of sociality not found among its
western relatives, as first-year birds are retained as
"helpers" in the territory. Discussing the social struc
ture of the Arizona populations of the Mexican Jay in
which territoriality is lacking and flocks are tolerated
at the nest, Hardy (1961) suggests that a species that has a
restricted amount of habitat available for nesting may
adapt by allowing a social breeding structure. Thus
the Mexican Jay, which nests mainly in the narrow rioarian
woodland of southeastern Arizona, tele
breeding birds about the nest, but the
of the Scrub Jay, which have a much wi
available, can utilize strict territor
rates flocks of non
western subspecies
der range of habitats
iality.
76


29
pair in that territory the preceding season of 1968, for
I banded them in the nest.
This pattern of retention of young of the year into
and through the next breeding season as members of the
territory is prevalent at Hicoria and at other areas in
Florida from which I have banding information. The
presence of helpers at the nest was first reported for
the Corvidae by Skutch (1335). Grimes (1940) first
suggested that Florida Scrub Jays had extra birds active
about the nest, and Am.adcn (1944) in his study of nest
ing Scrub Jays at the Archbold Station occasionally found
extra birds tolerated at certain nest sites. He tenta
tively concluded that many first year birds may not breed
and are sometimes tolerated about an active nest.
During the breeding season of 1968 certain birds
banded earlier were found associated with breeding pairs.
As outlined in Fig. 1 four extra birds were found, one
in each of four out of nine territories. Using the
flight feather technique of Pitelka (1945, 1951), I
tentatively identified three of the four as first-year
birds. Plumage characters were difficult to analyze
because all the jays had worn plumage at this time of
year.
In 1969 seven "helpers" were found in four of ten
territories, one in territory 2, three in territory 4,
two in territory 8, and one in territory 10. The other
six territories had only a mated pair in occupancy. Six


This dissertation was prepared under the direction of
the chairman of the candidate's supervisory committee and
has been approved by all members of that committee. It
was submitted to the Dean of the College of Arts and
Sciences and to the Graduate Council, and was approved
as partial fulfillment of the requirements for the degree
of Doctor of Philosophy.
August, 1970.
College"0
sCt£
Dean
f Arts and Sciences
Dean, Graduate School
Supervisory Committee:
Chairman


when adult had silver bands on the right leg. The juve
niles banded during 1963 had one colored band on the
right and one colored and the silver band on the lefr,
and juveniles banded m 19S9 had two colored bands on
the right and one silver on the left leg. Duco cement
placed on plastic colored bands effected a durable seal,
but occasionally bands were lost and the bird had to be
recaptured.
To follow the movements of banded birds I used a
30 power telescope mounted on a gun stock. In this way
I could read band sequences from as far as 100 yards and
sc did not have to disturb the bird. Occasionally a
single-sided blind was used but normally it was not needed
To study the local distribution of Florida Scrub Jays
I attempted to examine all contemporary Scrub Jay breeding
sites in Florida. This appears a prohibitively difficult
task, and several small areas were undoubtedly overlooked.
Yet I believe I was able to find nearly all of the breed
ing regions. The many letters from Florida Audubon member
in reply to my solicitations for information, greatly fcil
itated this search.
An invaluable aid when hunting for Scrub Jays is a
portable tape-recorder with recordings of Scrub Jay alarm
calls. These calls quickly attract any local Scrub Jays.
As a measure of preferred habitat I used the number
of active nests located in one hundred acres.
Difficul-


fl


28
the region as far as three miles from Hicoria looking for
banded birds.
SG-B and SW-R nested for two years in territory 9,
but each year the young vera destroyed while in the nest.
During 1970 the pair disappeared and the territory remained
unoccupied. Perhaps continual nest destruction induced
their departure. Yet other pairs continued territorial
occupancy after nest destruction.
On 20 May 1568, a feral cat located the nest of SY-Y
and S-BY. All the nestlings were killed and apparently
so was the male, SY-Y, for he was never seen again. Dis
location of a breeding pair probably occurs only after
some major catastrophe. Otherwise a mated pair remains
at its original territory year after year.
"Helpers"
I have stressed the territorial nature of the social
organization of breeding Florida Scrub Jays. They are
territorial, but each territory often contains more than
a mated pair. For instance on 12 June 1969 territory 8
contained MS-B (female), RS-Pi (male), O-SG (female),
O-SW (male), dBR-S, PiY-S, and PPi-S. The first two
were the mated pair, the last three juveniles. The
remaining two looked like adults (juveniles have brown
heads until their first molt in July), vet they were
definitely tolerated in the territory and had been all
spring. They were, in fact, the young produced by that


2
of the young. Although he occasionally observed non
breeding birds around active nests, Amadon concluded
that they do little to aid in the care of young.
Although Florida Scrub Jays are found scattered
over much of central Florida, most local populations
appear small, and some areas, such as in the Florida
Panhandle, lock ideal for nesting Scrub Jays but none
have been reported. The study reported here gives a
detailed outline of the social structuring of a breed
ing group of Florida Scrub Jays and analyzes the social
behavior of these birds in relation to dispersal and
habitat distribution.


69
with open sandy spaces restrict them to certain border
zones such as scrubby flacwoods, to recently burned regions,
and especially now to roadways and other disrupted scrub
areas produced by human interference.
Usually wherever the proper combination of dense
evergreen oak thickets and open sandy spaces exists,
some nesting Scrub Jays are found. Thus small populations
occur at Cedar Key, Levy County; at Camp Blanding, Clay
County; and scattered along the Atlantic coast often
miles from other breeding Scrub Jays. Although such
habitat is scattered over much of central Florida, it is
normally in small acreages of open thickets. The distri
bucin of Scrub Jay breeding habitat indicated in Fig. 7
illustrates the restricted amount of land available to
nesting Florida Scrub Jays. This map also shows certain
areas that appear to have suitable Scrub Jay habitat but
lack resident populations. Some of these areas are dis
cussed below.
Scrub Habitat Hitbout Breeding Scrub Jays
A few rather sizeable thickets with apparently suit
able habitat have no resident Scrub Jays. Several of these
scrub chicket areas are indicated in Fig. 7.
Kissimmee-Saint Cloud, Osceola County
Msnv sczrubfov thi ckGins exist
although most cf the sand ridge was on
tat ion anc. is now cultivated in orange
about this region
e turkey oak vege-
groves. The small


Even so, the total area available to Scrub Jays for
nesting is small. This habitat frequently exists as
small areas of thickets isolated from similar suitable
habitat.
The distribution of breeding Scrub Jays in Florida
is outlined. Only a few large expanses of suitable
habitat support dense populations of Scrub Jays. Al
though dispersal distances are not great, most small
isolated patches of scrub habitat are occupied by
Scrub Jays when in the vicinity of these centers of
population, but seemingly suitable habitat isolated by
20 or 30 miles from the nearest Scrub Jay population
is occasionally unoccupied. The Florida Panhandle -
has a large stretch of sand pine scrub community and
much of it appears ideal as Scrub Jay nesting habitat.
No Scrub Jays inhabit this extensive habitat which is
separated by 100 miles from the closest peninsular Scrub
Jay population.
The retention of first-year birds in parental ter
ritories and the reduced dispersal abilities -of non
breeding adults are discussed as a controlling factor in
the distribution of the Florida Scrub Jay. The sociality
of the breeding jays may be a behaviorial adaptation to
a limited breeding habitat similar to the case of the
Mexican Jay (A. ultramarina) in Arizona.
Vlli


TERRITORIALITY
The Florida Scrub Jay is a highly territorial bird.
At Hicoria territorial boundaries frequently made broad
contact and border conflicts were common. The boundaries
were fairly easy to map and Figs. 1, 2 and 3 outline the
territories present at Hicoria during the three years of
my study. Some daily shifting in the location of the
defended border occurred but it was minimal. Whether it
shifted or not, at all times a definite boundary was
recognized by other breeding jays,- and if a foreign jay
crossed, some sort of conflict ensued.
The area defended, among the 25 territories measured
over three years, ranged from 3.8 to 7.5 acres, but most
were close to the average of 6.0 acres. Each territory
included a large amount of scrub edge and open space with
which Hicoria abounds. The western side of the study
area included a large stretch of very dense scrub. The
only jays nesting in it were at the edges. I never found
pairs defending ground in the center of this dense area.
Nest Placement
The birds invariably placed their nests two or three
feet in from the edge cf an open space. Usually myrtle
oaks wars selected for nest building but also used were
wax myrtle, Chapman oak, and in one case a scrub palmetto
18


17
as usual she at once gave begging cries, and crouched
down. He fed her but remained standing before her. She
continued to beg and GS-G began to hop around her. His
circling movement was a curious stiff-legged affair with
his neck extended and bill pointed slightly down. His
tail was spread and slowly swung from side to side. The
wings were half spread and lowered, almost dragging on
the ground. He circled her three times while she remained
crouched with her bill raised, a submissive posture. The
male was silent, and just before mounting, the female also
became silent. The male mounted the female for approxi
mately six seconds and then hopped to a nearby perch
where he vigorously bill-wiped and shook his plumage.
The female also began bill-wiping, ruffled her feathers,
and bill-wiped again. She then "hiccupped" rather
loudly and followed the male when he flew off. This
pattern is essentially the same as that reported by Brown
(1963) for the Scrub Jay in California.


23
that was intertwined with a myrtle oak. Scrub Jay nests
were well camouflaged and were difficult to see. In order
to locate all the nests I had to search through practi
cally every tree in the area.
Nest building began as early as 20 February and
extended to 2 June at Hicoria. There was no definite
time when most pairs were nest building or incubating;
it was a protracted period. One pair had young out of
the nest while another was nest building. Unlike Blue
Jays (Hardy, 1961) the Scrub Jay pair defended its
territory from all Scrub Jay intruders no matter what
the stage of the breeding cycle of the defender or the
intruder. GS-G and S-RB did not commence nest building
until 18 May 1968, yet they actively defended a rather
large territory from all Scrub Jay intruders from March
onwards. The nest was completed on 21 May, but no eggs
were laid until 4 June.
Territorial Defense
The male of a pair spends much time perched in an
upright position with tail held straight down. This
behavior is most common during breeding season and occurs
while the female is both on and off the nest. From the
perch the bird can readily see all of his territory. This
same posture is used by Scrub Jays in the fall and winter
when in a flock. Consistently one sees a single bird,
not necessarily a male, perched high above the others
feeding below. If this bird drops to the ground, within


TABLE CF CONTENTS
ACKNOWLEDGMENTS
LIST OF FIGURES
ABSTRACT
INTRODUCTION
HICORIA STUDY AREA
METHODS
RECOGNITION SIGNALS
Auditory Signals
Visual Signals
COURTSHIP
Pair Formation
Copulatorv Behavior .
TERRITORIALITY
Nest Placement
Territorial Defense
Site Tenacity and Mate Constancy
"Helpers11
Interactions of Adults and Juveniles
Role of "Helpers11
EXTRA-TERRITORIAL MOVEMENTS ....
REACTION TO OTHER SPECIES
Reaction to Blue Jays .
Fage
ii
v
vi
1
3
10
10
11
14
14
16
18
18
23
26
28
30
33
39
41
A 1
.11
I


Even when a disturbance alarms the parents, the young
remain well hidden and silent. As newly iledged youngsters
are vulnerable to predation, this stage of extreme timidity
is adaptive. The parents quickly recognize a predator and
may draw its attention with their loud alarm call, upwardly
inflected kwerp; the silent young may remain unseen as
the predator is distracted by the parents. In fact, many
young are lost during these few weeks, yet possibly many
more would be killed if they were not secretive. About
two weeks after leaving the nest the juveniles first
become active.
Juvenile Florida Scrub Jays have light brown plumage
on head and back and lack the light blue forehead of the
adult until their first molt in late July. It is easy to
recognize juveniles by this trait. I saw no special
reaction by adults to birds with brownish heads except
when young from other territories inadvertently wandered
over another adult's territory. The resident male quickly
chased these wandering youngsters. The ensuing chase was
especially intense and afforded the only occasions that I
viewed physical contact during an attack. The male's
cortege of mate, helpers, and young often join the attack
by following the male and calling raucously. As with
border disputes, normally only the male initiates these
attacks.
Hir.de (1961) suggests that "the juvenile usually


Page
Reaction to Predators 42
Reaction to Man . 45
DISPERSAL 47
HABITAT AMD DISTRIBUTION 52
Scrub Jay Habitat in Florida 52
Hicoria 55
Cape Kennedy Air Force Station 5 8
Jonathan Dickinson State Park 59
Ht. Plymouth Area 5 9
Lake Jackson 60
Ridge Scrub 60
Scrubby Flatwoods West of Hicoria ... 61
Atlantic Coast Dune Thickets 64
"Big Scrub" 67
Ocala National Forest 6 7
Habitat Summary 68
Scrub Habitat Without Breeding Scrub Jays. 69
Kissimmee Saint Cloud 6 9
Weeki Wachee 72
Lake Wales 72
Florida Panhandle Scrub 73
FACTORS CONTROLLING DISTRIBUTION 76
LITERATURE CITED 83
BIOGRAPHICAL SKETCH 35


72
scrubby thickets, often merely 200-300 acres, appear to
be excellent Scrub Jay cover and resemble the scrub at
Mt. Plymouth. No Scrub Jays exist in this region, but 20
miles to the north similar habitat supports numerous scat
tered populations of Scrub Jays.
Weeki Nachae, Hernando County
An extensive scrub exists about this natural spring.
Much of if is covered with sand pines and so could not
be expected to support Scrub Jays. Small sections have
recently burned, however, and low dense thickets are grow
ing back, with numerous open sandy spaces. Moreover, along
the southern border of this scrub much openness exists in
a broad ecotcne between the scrub and turkey oak vegetation.
I estimate approximately 1000 acres of low open scrub thickets
exist here, but no Scrub Jays are present. Scrub Jays reside
in small numbers 20 miles to the south and 28 miles to the
east.
Lake Nales, Polk County
A few islands of scrub thickets, similar to the ones
west of Hiccria, are present to the west of Lake Wales,
but no Scrub Jays reside on them. The plant community on
the sand ridge close to these scrub islands at Lake Wales
is turkey oak vegetation without any Scrub Jay population.


7c
and species with limited breeding range may be unable to
maintain such a population level unless special social
structures are included. With the retention of first-year
birds as territorial "helpers," the Florida Scrub Jay has
developed a degree of sociality as an adjunct to an essen
tially territorial nature.
This combination of sociality and territoriality is
possible, for the breeding male may courtship-feed the
"helpers" and so form a loose bond with these birds. All
members of the territory frequently react together in
territorial defense. Thus the family group remains terri
torial, unlike Mexican Jay flocks that show no territoriality.
Interestingly, compared with Scrub-Jays, the breeding male
Mexican Jay utilizes courtship feeding infrequently and
only with his mate (Brown, 1963; Hardy, 1961).
In Mexico and western United States many different
habitats are selected by breeding Scrub Jays, but most are
of a xeric nature and, as Pitelka (1951) points out, the
Scrub Jay appears bast adapted to feeding in and about the
low dense thickets of chaparral or open oak woods. Even
though isolated by over 1000 miles from these western races,
the Florida subspecies continues to utilize habitats of
similar ecologic structure. Presumably at one time a con
tinuous scrub habitat connected the Florida peninsula with
western habitats and Scrub Jays invaded Florida through this
connecting habitat (Pitelka, 1951).
Whether or not large expanses of suitable habitat
ever existed an Florida is difficult to ascertain, but


33
reate, and the response is usually the sane, food being
passed to the beggar. Obviously the continued occurrence
of courtship feeding between mated birds is a means of
pair bend reinforcement. It is seen most frequently
during the breeding season but occurs all year round.
Possibly as juvenile birds continue to beg and the parents
respond and feed them, an attachment something like a pair
bond is established. As the next breeding season approaches
the parents are increasingly attentive to each other, yet
the young from the previous season are still present and
tolerated by the male. Occasionally they beg and he
responds by passing a morsel of food. I suggest that these
one year old birds can remain in their parents' territory
because a loose bond is present between the "helpers and
the breeding male similar to a pair bond.
Role of the "Helpers"
The role of these "helpers" in the organization of
territorial members is often vague. I call them "helpers"
but this term is, at times, misleading. It is a term
that has been used in the past and I shall continue its
use, but "auxiliary territorial member" is perhaps more
accurate.
I never saw these "helpers" take part in nest build
ing. Usually the first-year birds ignore the adults'
nesting procedures and quietly feed somewhere in the
territory. When voung are in the nest and the aars-ts


igure 3 Map of study area at Hicoria showning terri
torial boundaries and Scrub Jays residing in
each territory during the breeding season
1970. Note: no young were banded this year
(See key for Fig. 1).


In areas where
no Scrub Jays occur
Scrub Jay always do
together, I doubr i
sand pine is dense, Blue Jays nest and
as breeding birds. As the Florida
minabas the Blue Jay when the two are
f the Blue Jays have displaced the
Scrub Jays from the mature sand pine scrub. It is more
probable that the Florida Scrub Jay finds this habitat
unsuitable for nesting.
Sand pine trees may be absent from the scrub community
for several reasons. Sand pines are especially susceptible
to fire and once flames reach the canopy, large regions
quickly burn. But Webber (1935) states that mature sand
pine forest may act as a fire inhibiting community because
the ground is often bare of grasses and fire has difficulty
in spreading. Thus many scrub communities may remain
covered with pine forest and are unavailable to Scrub Jays.
Today the State Forest Service fire protection program is
aiding the preservation of sand pines.
Frequently the washed and sorted sands are thin and
do not support typical scrub communities. At such sites
understory woody planes of che scrub community are mixed
with species normally present in pine flatwoods of nearby
areas.
The Florida Scrub Jay selects a specific pattern of
growth in these xeric communities and rejects as unsuit
able much or ens true sand pme
special habitat requirements of
detailed examination of several
scrub. !o illustrate
the Florida Scrub Jay
different sices sueno
the
a


16
once RS-dB jumped up from the ground. O-SW bobbed and
continued calling, and RS-dB "hiccupped." Then the male
flew off giving rapid kwsrps and the female immediately
followed, giving the same calls.
Whenever a disturbance occurs it is the male that
first flies and the female follows; this pattern is fol
lowed by all mated pairs. Pair formation then is a
matter of the two birds remaining in close association,
one bird, the male, passing food to the female and the
female following closely after the male in all encounters.
Although the above discussion deals with the forma
tion of a new bond, it also explains the interaction of
paired birds, frequent courtship feeding even during the
fall and winter, and persistent following of the male by
the female.
Copulatory Behavior
The complete copulatory pattern was observed only
once, although I spent many hours behind a blind hopeful
of following this sequence.
On 10 April 1969 GS-G, the male, presented a cater
pillar to his mate S-RB at 1030 and brought her food
eight times during the next 50 minutes, even though she
continued to feed on her own. Frequently, after court
ship feeding, the female gave rather subdued "hiccup"
calls and a series of melodic notes and calls as though
she were whispering. At 1120 he nopoed over to her and


82
forcing resident bi
have located the ne
The extensive
nearest small Scrub
occupy all the suit
they should take r.ia
rds away; and wandering birds may not
wly available habitat after it regrows,
panhandle scrub is 100 miles from the
Jay colony. If straying birds cannot
able sites in central Florida, then
ny years longer to invade the pan
handle region.
The adaptation of a type of sociality by the Florida
Scrub Jay has produced certain restraints on the distribu
tion of these birds. Because they have become exceptionally
sedentary, and dispersal abilities have been reduced, some
areas of suitable habitat in Central Florida and a large
expanse of habitat in the Panhandle are unoccupied today.


61
this terrain. Judging by large sand pines scattered along
the highway much of -this area may have been covered with
extensive sand pine forest. Scrub thickens extend several
miles in places as a solid dense ground cover 3-6 feet high.
The three scrub oak species dominate (60 per cent of ground
cover in thicket) and Sabal etonia is common (20 per cent
of ground cover in thicket)(Fig. 5). Although grasses are
infrequent and the surface is usually bare white sand, very
few large open spaces appear. It is a difficult area to
census for nesting Scrub Jays, because cf the impenetrable
nature of the low woody plants. Nesting jays occur over
much of this terrain but congregate along read edges or
about the few open sandy spaces. During 1968 and 1969
two areas vrere censused. Approximately 3-4 pairs nest per
100 acres, but in the densest areas away from open spaces
jays rarely nested.
Scrubby Flatwoods West of Hicoria, Highlands County
Elevation is 130 feet. Just west of the scrub covered
hills of Highlands County an extensive region of flatwoods
exists. Within three miles cf the ridge, numerous small
narrow strips of white sand are scattered. Probably these
were small sand bars in a Pleistocene sea. The thickets
on these strips are extremely dense. The edges of these
small islands of thickets are usually bare sand or occa
sionally widely spaced scrubs of rosemary.
Nesting Scrub Jays occur in the scattered thickets,
especially '.'hen close to the extensive scrub ridge at the


Abstract of Dissertation Presented to the Graduare Council
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
ECOLOGY AND BEHAVIOR OF
THE FLORIDA SCRUB JAY
By
Peter Walter Westcott
August, 1970
Chairman: Professor Pierce Brodkcrb
Major Department: Department, of Zoology
This study gives an outline of the social structuring
of breeding Florida Scrub Jays (Apheiocoma c. coerulescens)
and discusses the restrictions placed on distribution by
the type of habitat selected for nesting. The social struc
ture was studied at Hicoria, Highlands County, Florida,
where 230 individuals were banded.
Certain calls and postures used in social interactions
are described, and pair formation and copulatory behavior
are discussed.
In Florida the Scrub Jay is territorial and pairs for
life. The territory is defended from intrusion by other
Scrub Jays, but first-year birds (young of that pair from
the year before) are retained in the territory. They
occasionally act as helpers" by caring for the young,
and frequently aid in territorial defense. Although they
restrict their movements to the territory, usually these
extra members remain away from the breeding pair. A bond,
similar to a pair bond, may be present between the breeding
vi


36
a tall perch and bobbed and "hiccupped." This enticed
S-BY, a female helper, to fly up where she commenced
"hiccupping. As they fell silent SR-B (male) flew off
50 feet, giving loud rapid kwerps. Again the other four
members of the territory followed, and another series of
bobbing and "hiccupping" ensued.
Not only did SR-B and S-BO in territory 4 have more
helpers than did other breeding pairs, they also had one
helper that was an adult female. This bird, S-BY, was
the mate of SY-Y in 196S. On 20 May a feral cat attacked
and killed their young in the nest and apparently also
killed the male, SY-Y. For the next week S-BY remained
in the region of the destroyed nest but more and more
regularly she fed with the members of territory 4 at a
feeding site that I had been supplying between their
territories. By 25 May she no longer spent any time in
her old territory and followed along with SR-B, his mate,
and helpers into territory 4. At this time SR-B and mate
were attending young in their nest, and S-BY began to visit
the nest with food. Whenever SR-B saw her at the nest he
gave chase, but she never flew far. She always rrtoved off
when he was aggressive. On 26 May I first saw her beg
toward SR-B. He ignored her but did not chase. Thereafter
she frequently bagged toward him, and he usually responded
with a morsel of food. She remained in that territory as
the only adult "helper" that I have found. S-BO, female,
never became upset by the presence of S-2Y and for 13 months


60
4G per cent of the habitat is open sandy spaces, which are
practically devoid of any herbaceous cover. A nesting
census located 910 nests per 100 acres and all were found
in oak trees.
Lake Jackson, Jeering, Highlands County
Elevation is 110 feet. Undoubtedly a fire destroyed
an extensive sand pine forest at this tract on the west
shore of the lake at least ten years ago to judge by scars
on a few large sand pines. Today a young scattered stand
of sand pine is present, but most of the region has dense
thickets of woody plants 6-10 feet high. As usual, oaks
dominate, but Sabal etonia and Ilex cumulicola are common.
There are few natural open sandy spaces, but a new housing
development has a series of parallel roads through this
scrub, and an unaltered strip of thicket 100-150 feet wide
between roads has been left. The developer, thus, has
produced a series of open spaces, often of ^^7hite sand,
throughout the dense scrub thicket. No nesting census was
made, but the density of Scrub Jays was always high and
reminded me of the densities of jays at Cape Kennedy and
Eicoria.
Ridge Scrub, Highlands County
Elevation is 150 feet. Wide expanses of very low dense
scrub exists along U.S. Highway 27, two miles east of Hicoria.
In recent years orange groves have usurped the majority of


35
During aggressive encounters with neighboring birds
these first-year birds m.cst readily take an active role
or act as "helpers." Whenever the male gives chase to
an intruder, the first-year members of the territory at
once fly toward him and follow behind, frequently calling
as loudly as the male. A border encounter often entails
three or four birds, loudly calling and chasing up to the
invisible territory boundary. The male then wheels back
and the others follow him back into the territory, all
still calling loudly. Certainly a mob of jays, all
seemingly bent on attack or chase, is much more effective
in instilling a sense of invulnerability about a territory
border than would only a single male.
In the same way the seemingly ritualized early morn
ing border encounters include the "helpers," if present,
in that territory. In these cases two mobs of jays may
approach each other and suddenly wheel and fly apart.
If a territory contains more than one "helper" all do
not necessarily join in these border contests.
In 1969 territory 4 contained two adult.breeding
birds and three "helpers." On 10 May an unbanded bird
flew through the territory. At cnce SR-3, the male, took
after the intruder and the other four birds feeding with
the male followed. The unbanded bird sailed rapidly
across the territory. The five attackers came back to
the feeding site, but all remained in the highly agitated
state for at least 10 minutes. S-30, the female, sat on


43
On 22 April 19-39 I banded the nestlings in territory
9 and replaced them in che nest. The next day the young
were still in the nest, but on 24 April the nest was empty
and two adults were extremely agitated. Dead on the paved
road 200 feet from the nest was a six-foot Coachwhip that
had been hit by a car. Upon opening the stomach, I found
the two young from territory 9 still wearing my bands.
Once out of the nest the young are still highly vul
nerable for at least a week and many disappear. Feral cats
are frequent at Hicoria and prey heavily on fledglings.
Gray Foxes (UrOcyon cinereoargenteus) and Bobcats (Felis
rufus) also hunt through this area.
Whenever a predator is noticed, the Jays in the ter
ritory mob about it, using all alarm and excitement behavio
patterns. Some give loud upwardly inflected kwerp calls
and bob vigorously. Females frequently give loud "hiccup"
calls and bob. A jay often dips its head toward the
predator with its body hanging straight down. Some, expe-
daily the breeding pair, hop about the ground, calling
.loudly and nipping the tail of the attacker.
The output of noise easily carries over many other
territories, but other Scrub Jays normally do not cross
into the territory where the trouble occurs. Birds in
ocher territories are aware of the attack for they become
extremely excited. Only when the predator is found close
to the territorial boundary' or when the encounter continues
for an especially long time, do other territorial birds


so
sedentary habits of Florida Scrub Jays. Only following
some major catastrophe, as a fire or bulldozing of large
areas cf scrub, were Florida Scrub Jays found far from.
scrub habitat.
Only a few areas occur in Florida with broad expanses
of open scrubby cover and support relatively large and
dense populations of Scrub Jays such as the Lake Placid
areas (Archbold Station, Eicoria, etc.), Jonathan Dickinson
State Park, Cape Kennedy, and the Mt. Plymouth region
(Fig. 7). Not only do Scrub Jays abound in these areas,
but nearly all similar habitats nearby have resident Scrub
Jays. Presumably enough extra birds inhabit these popula
tion centers so that most small scrub thickets are located
and colonized by the jays. Similar small sites further
away (10-20 miles) have only a few scattered pairs of
birds or none at all. The density of nesting Scrub Jays
gradually decreases along the coastal thickets north from
Cape Kennedy. North of Daytona Beach nesting Scrub Jays
are noticeably less frequent, and from Marineland to
St. Augustine nesting pairs are often separated by several
miles of low dense thickets even though the habitat is
similar to the densely inhabited shore thickets just north
of Cape Kennedy.
Similar patterns of density of this species occur
about the other large centers of its population. Close
to densely inhabited thickets most small scrub habitats
have nesting birds, but farther awav fewer birds are


44
enter and join in the melee. As the new jays approach,
the local male often dashes suddenly at one of them. These
are short chases, and the male quickly returns to the
predator. These short quick attacks at a foreign jay
rarely continue to the territorial border. During non
breeding seasons when mated pairs are less insistent about
territorial defense, alarm calls more frequently attract
other Scrub Jays.
These observations concerning the attraction of other
Scrub Jays to distressed breeding birds are consistent with
Hardy's (1961) comments concerning degrees of territoriality
among species of jays. The Mexican Jay with no territori
ality and Blue Jay with a loose territoriality both give
loud assembly calls that readily attract others, whereas
the highly territorial western Scrub Jays do not. Even
though possessing a degree of sociality above that of
western races, the Florida Scrub Jay is highly territorial
and does not have a distinct assembly call. Its alarm
calls carry far, but as with western races, outside birds
frequently do not assemble when a pair is in distress.
Nearly every other resident species at Hicoria does
assemble. The alarm calls of the Scrub Jay quickly attract
Brown Thrashers, Blue Jays, Mockingbirds, Towhees (Pipilo
erythrophthalmus), White-eyed Vireos (Vireo griseus), and
especially woodpeckers. Red-bellied Woodpeckers (Centurus
carolinus), Red-headed Woodpeckers (Melanerpes ervthrcce-
phalus), and Flickers (Celantes auratus? are easily excited


REACTION TO OTHER SPECIES
Although territorial Florida Scrub Jays react vigor
ously toward intruders of their own species, they usually
ignore other bird species. The Brown Thrasher (Toxostorna
rufum) is similar in size and ecological requirements.
Pairs are scattered over the Hicoria area and they fre
quently feed in the same places as Scrub Jays, yet con
flicts rarely occur. Only when a thrasher or Mockingbird
(Mimus polyglottos) perches especially close to an active
nest dees the male or female jay displace the intruder.
Elsewhere in the territory other species are tolerated.
Reaction to Blue Jays
No Blue Jays nest in the Hicoria area, yet numbers
are alv/ays present, either feeding or travelling through,
often in groups of four and five. Blue Jays frequently
feed with Scrub Jays. Usually the Blue Jays.remain in
the background and only after the Scrub Jays leave the
feeding site do the Blue Jays dash in and grab some food.
Frequently at feeding sites Scrub Jays and Blue Jays
come close together and invariably the Scrub Jays dominate
the Blue Jays. Such encounters rarely include an attack
or chase; merely a peck or jab toward the Blue Jay is
sufficient to cause it to back off and fly.
41


INTRODUCTION
The Scrub Jay, Aphelocoma coerulescens (Bose), occurs
in two separate areas of North America. Throughout the
western United States and the mountains of Mexico this
species lives in a variety cf habitats, often in large
numbers (Pitelka, 1951; Westcott, in press). Although
the western population is composed of sixteen subspecies,
all appear to have similar social organizations. sin
gle eastern subspecies, A. c. coerulescens, inhabits
central Florida. Not only is it isolated by over a
thousand miles from the closest western population but it
also exhibits completely anomalous patterns of sociality.
Unlike its western representatives, the Florida Scrub Jay
is restricted in habitat and is often quite scarce. Western
Scrub Jays are normally secretive, yet Florida Scrub Jays
are notably tame.
Reports of extra individuals acting as "helpers"
about the nest (Grimes, 1940) also suggest a degree of
sociality unlike that of western Scrub Jays, but little
information has appeared concerning this example of
sociality in the Florida Scrub Jay. Amadon (1944) exam
ined the activities of breading pairs about the nest and
presents details of courtship, nest building, and care
1


AC KNOWLEDGMENTS
I wish to extend thanks to the following persons and
organisations who aided me in various ways in carrying
out the present work. Frofessors Pierce Brodkorb, Archie
F. Carr, Jr., Albert M. Laesse, David W, Johnston, and
Ernest S. Ford of the University of Florida acted as my
supervisory committee while I was a student at the Uni
versity of Florida. Professor Brodkorb was my principal
advisor, and I especially wish to acknowledge his tireless
efforts in the preparation of this-manuscript. Dr. Oliver
L. Austin, Jr., Curator, Florida State Museum, provided
both banding material and suggestions during the course
of this study. Dr. James N. Layne of the Archbold Biologi
cal Station and Dr. Glen E. Woolfenden, University of
South Florida, provided much advice and encouragement.
Mr. Donald E. Goodman greatly aided in the tiresome enter
prise of capturing birds. I am mindful of the invaluable
grants-in-aid received from the Frank M. Chapman Memorial
Fund of the American Museum of Natural History and from
Sigma Xi.


84
Pitelka, F.
1945.
A.
Pterylography, molt, and age determination of
American jays of the genus Aphelocoma. Condor,
47:229-250.
1951. Speciation and ecologic distribution in American
jays of the genus Aphelocoma. Univ. California
Pubi. Sool., 50:195-484.
Skutch. A. F.
1935. Helpers at the nest. Auk,
52:257-273.
1961. Helpers among birds. Condor, 63:198-226
Webber, H. J.
1935. The Florida scrub, a fire fighting association*
Amer. Jour. Bot., 22:344-361.
Westcott, P. W.
1969. Relationships among three species of jays winter
ing in southeastern Arizona. Condor, 71:353-359.
Westcott, P.W.
In press. Territoriality of the Scrub Jay in Arizona.
Condor.


Figure 8 View of low scrub thicket with open sandy
spaces at Fort Pickens State Park, Santa
Rosa Island, Escambia County. This type
of scrub thicket supports large numbers of
Scrub Jays when present in the Florida
Peninsula, but here no jays occur.


HABITAT AND DISTRIBUTION
Florida Scrub Jays select a nesting habitat of very
limited range and at times are absent even though the
proper breeding habitat is available.
Scrub Jav Habitat in Florida
Nesting Scrub Jays are invariably located in types
of sand pine scrub vegetation. The vegetational components
of sand pine scrub are primarily xeric plants that are
restricted to the veil washed and sorted siliceous sands
of modern and ancient shore lines. During the late Tertiary
and the interglacial portions of the Pleistocene the drastic
rises in sea level left four distinct ridges of deep sand
ranging in elevation from 150 feet to S feet (MacKeii, 1950)
The present day sand pine scrub vegetation is located on
these stretches cf sands of the Saint Lucie and Lakewood
types of soils, deposited by marine currents, wave action,-
and winds (Laessie, 1958). Since these sorted sands were
laid down as shore lines or sand bars, today sand pine
scrub communities are found primarily as long narrow
strips of vegetation. Two major exceptions are the Ocala
National Forest in Zlarion County and the extensive
O L. Ulv
communities cf central Highlands County. Yet Laessie (1953)
concludes that these, too, are of dune origin, and the
extreme width is the result of winds blowing dune series
inland and new dunes forming at the sea front.
sea


DISPERSAL
The striking feature cf the movement of Scrub Jays
about the Hicoria area is that there is so little of it.
Apparently, once a pair establishes a breeding territory,
it remains as long as conditions are suitable. Of nine
pairs nesting at Hicoria in 1968 six were still there
nesting in 1970. One of these was a female WS-Pi banded as
a breeding adult in 1960 within 200 yards of the territory
she now occupies.
Dispersal of certain members does occur, for in late
summer and fall the first-year "helpers" begin to disappear
from their parental territories. During this period Scrub
Jays are prone to make long flights. A single bird will
fly as much as 0.3 mile from the territory. Such long
flights are usually silent, unaccompanied and at such a
height (100 to 200 feet) that other Scrub Jays ignore them.
Normally these long-fliers return later the same day. All
birds that made long flights and that I could identify were
"helpers"; I never saw a breeding adult make such flights.
This is not a phenomenon restricted to a certain time
of year. Even during the breeding season, occasional long
flights occur. They are much more frequent, however, in
the fall. Perhaps it is an expression of unrest by particu
lar birds, a prelude to a final move. 3y late fall most
47


HICORIA STUDY AREA
The study area at Hicoria is located two miles due
south of the Archbold Biological Station, Lake Placid,
Highlands County, Florida, and is at the extreme southern
end of the sand ridge region of central Florida. At an
elevation of 140 feet above sea level, this locale con
tains a plant community that combines sand pine scrub
and pine flatwood elements; its special vegetational com
ponents are termed scrubby flatwoods by Laessle (1942).
The vegetation is discussed in detail in a later section.
A dominant feature of this particular site is a
multitude of large and small open sandy spaces. As dis
cussed later, this combination of dense scrub thickets
and open space represents the optimum habitat for Florida
Scrub Jays and gives a density of breeding birds at
Hicoria that facilitates the studies of social behavior.
The open areas of this site are not typical of the
scrubby cover which elsewhere is extremely dense and
extensive. During the early years of this century Hicoria
was the site of a town of one thousand persons, a glass
factory, and a lumber mill. Today the only remains are
the scattered open spaces where houses or other build
ings once stood. The outline of the study area in Fig. 1
shows the pattern of open spaces resulting from the aban
donment of the town and factories.
3


34
actively feed them, first-year birds occasionally follow
one parent toward the nest. At such times the breeding
male often supplants the first-year bird by landing next
to it and sometimes pecking. The "helper" either gives
a submissive posture (slightly crouched and bill upraised)
and then flies off, or else flies a short distance at
once. Rarely does the male chase after the "helper."
On 12 May 1969 in territory 2, SR-O, male, was not
in sight as PiS-W, female, flew to the nest and fed a
youngster. She flew off and immediately 1G-YS, a female
first-year bird, flew to the nest and peered in. The
hestlings began begging and 1G-YS moved her bill toward
them as though to pass food but none was passed. Again
two days later, just after the female left the nest and
the male was not in sight, 1G-YS flew up to the nest and
this time fed one of the youngsters.
When the young leave the nest the parents continue
feeding them and now the first-year birds more frequently
bring food to the fledglings. In territory 1 in 1968,
the young left the nest on 18 March and for the next three
days SR-W, female first-year bird, brought food to the
fledglings at least twice each day. In 1969 M-PS in
territory 10 fed young after they were out of the nest.
Thus, the first-year birds present in the territory occa
sionally function as "helpers" with the young. Most of
the time, however, they remain somewhere in the territory
but away from the nest or new set of youncr.


Wing flutter and crouch.This is a begging posture
and is almost always used with a begging cry. The body
is pressed close to the perch or the ground and the wings
are rapidly fluttered.
Bill wiping.The bill is rapidly rubbed on both
sides of the perch as though attempting to clean some
thing off. This movement is used at the conclusion of
a wide variety of activities: after fighting, after
swallowing, after courtship feeding, or after drinking.
It is so frequently used that perhaps it can be viewed
as the final consummating portion of a sequence of actions.




42
Blue Jays often forage through Scrub Jay territories.
Only when the Blue Jay approaches the Scrub Jay or blunders
near a nest does the Scrub Jay react. In such cases the
offending Blue Jay is often chased. As with intraspecific
chases, the other territorial members quickly join in and
follow to the edge of the territory. Occasionally the
breeding feraele Scrub Jay begins such a chase, but the
"helpers" join, and the male usually takes over and com
pletes the chase to the border. These interspecific
encounters are infrequent, for the Blue Jays normally stay
away from Scrub Jays in their territories.
Scrub Jays recognize something in the Blue Jay as
necessitating an aggressive reaction. Yet the Blue Jay
is not treated as a distinct threat, for it is frequently
left alone in Scrub Jay territories and at feeding sites.
Reaction to
Predators
Although the male or any other territorial member in
a sentinel position frequently warns the others of predators,
during 1SS8 and 1969 five of 20 nests with young were des
troyed. Probably the common predator of young Scrub Jays
are snakes. At Hicoria I frequently found Corn Snakes
(Elaphe guttata}, Yellow Rat Snakes (Elaohe obsoleta), Pine
Snakes (Pituophis melar.oleucns) Kingsnakes (LampropeItis
getulus), Black Racers (Coluber constrictor), and especially
the Eastern Coacnwhip (Masticoohis flagellum), The Ccachwhip
as especia.
coursing through the
ms or scrim
thickets.


Figure 4 Viev; of Hicoria study area, center of territory
6.


22


5
Although more than 50 juvenile Scrub Jays have been
banded within the 200 acre study area at Hicoria, M-dBS
was the only male banded here as a juvenile to establish
a new pair bond and territory in this same area. One
other bird banded as a juvenile settled to the west of
the study area, and I followed him as he established a
pair bond and territory. O-SW, the male, settled in
a small patch of scrubby live oaks surrounded by wet
flatwoods approximately 1200 yards west of territory 8,
in which he was banded as a juvenile in 1968. On
8 December 1969 he was first seen in the scrubby patch
and had another bird in attendance. This bird was not
banded but gave the female "hiccup" calls as soon as I
approached. She was subsequently captured and banded
RS-dB.
On 8, 9 and 10 January 1970 O-SW continually approached
the female who at once crouched and gave begging calls and
wing flutters. The male immediately passed a morsel of
food to the female. After passing the food no ether
special calls cr movements occurred except that usually
they both bill-wiped. Occasionally a female gave "hiccup"
calls after courtship feeding, but usually both began some
completely unrelated activity such as feeding or preening.
As I slowly approached O-SW on 10 February 1970 he
was perched on the tallest branch available in this scrub
patch. There was no way I could approach unnoticed and
at 70 yards he gave upwardly inflected kwerp calls. At


37
S-EY remained so tied to this pair that it appeared that
SR-3 had two mates. Although he regularly fed her when
S-BY begged, 3-BO remained his dominant mate and was the
breeding female the next year.
In. 1963 S-EY was still in attendance. She did not
help with nest building, but regularly brought food to
the young in the nest. During this breeding season the
male .id not attempt to chase S-BY from the nest. She
was then truly a "helper" at the nest. The other two
first-year birds in attendance did not attempt any such
close association about the nest; they acted similarly to
first-year birds in other territories.
Interestingly, in February, 1970, S-BY and M-dBS
paired and established a territory just west and adjacent
to territory 4. M-dBS was one of the first-year birds in
attendance in territory 4 while S-BY was also a "helper."
The Florida race is unique among Scrub Jays in that
family flocks defend their territories with all individuals
helping in various degrees. As far as known it is found
in none of the western races {Hardy, 1961). .
The social organization of these flocks is as would
be expected. The breeding male is dominant over the first-
year "helpers," although normally they appear to ignore
each other. Only occasionally did an adult female supplant
any of her first-year offspring in the territory and I never
saw a female chase or attack a "helper." In a territory
that had more than one "helper" no special hierarchy was


11
a confrontation with another Scrub Jay. As Amadon states,
this extra loud call is often lengthened into a screech.
Finally, the rapidity of presentation of the call
represents another variation of the kwerr. Especially
during the early morning a rapid series of kwerr or more
frequently kwerp-kwerp-kwer? calls are heard. Again
this is a sign of increased excitement. It is normally
used in antagonistic bouts with other jays, especially
after a series of loud kwerrs.
"Hiccuo'1This peculiar sound apparently is produced
only by the female. Amadon's pronunciation of it as klok
is quite exact. It is a hollow sound repeated rapidly so
that it sounds like a hiccup. It is often used in antag
onistic contexts yet it frequently is given when birds
are quietly perched alone or during courtship. The loud
ness of presentation expresses a more excited state.
Begging call.This cry is always used with a wing
flutter and is the characteristic sound of the young in
the nest as a parent arrives. The same call is used by
a female toward her mate. I have never heard a paired
male use it, although I have heard it used by an adult
male toward a female in a small cage.
Visual Siar.als
As with the auditory signals most of these visual
postures are those used by the Scrub Jav when reacting
in some agonistic context either toward their own soecies
or toward a predator.


LITERATURE CITED
Amadon, D.
1944. A preliminary life history study of the Florida
Jay, Cvanocitta c. coerulescens. Results Arch
bold Exped. No. 50. Am. Mus. Novitates, No.
1252:1-22.
Bent, A. C.
1946. Life histories of North American jays, crows,
and titmice. Bull. U. S. Nat. Mus., No. 191.
Brown, J. I..
1963. Social organization and behavior of the Mexican
Jay. Condor, 65:126-153.
Hardy, J. W.
1961. Studies in behavior and phylogeny of certain
New World jays (Garrulinae). Univ. Kansas
Sci. Bull., 42 (2):13-149.
Hinde, R. A.
1961. Behavior. In A. J. Marshall, ed., Biology and
Comparative Physiology of Birds. II. New York:
Academic Press.
Grimes, S. A.
1940. Scrub Jay reminiscences. Bird Lore, 42:431-436.
Laessle, A. M.
1942. The plant communities of the Welaka area. Univ.
of Florida Press. Bio.1. Sci. Series, vol. 4,
No. 1:1-143.
1958. The origin and successional relationships of
sandhill vegetation and sand pine scrub. Eccl.
Monog., 28:361-387.
MacNeil, F.
1950.
Pleistocene shore lines in Florida and Georgia.
U. S. Geol. Surv., Prof. Paper 221-F, pp. 95-107.
Nice, M. M.
19 37.
Studies in the life history of the Song Sparrow.
I. Trans. Linn. Soc. N. Y., 247pp.
941* The role: of herritCa
Nat., 26:441-487.
in bird life,
Ame
Midi,
83


49
origins. Territory 7 was settled by Pis-O and S-GB
who were first-year birds in this same general area during
1963. Their parental territories were also unknown. Terri
tory 11 was established by S-BY and M-dBS who were both
"helpers" during 1969 in territory 4. As these two areas
were adjacent, this new pair was filling in unoccupied habi
tat at Hicoria and only a minimal dispersal occurred.
Three first-year "helpers" moved completely away from
the Hicoria region during their second year. S-RY moved
1.2 miles due south from territory 6 and nested in 1969,
O-SG moved 0.9 mile westnorthwest from territory 8 and
nested in 1970, and O-SW moved 0.4 mile westsouthwest
from territory 8 and nested in 1970. These last two
settled in scattered scrubby thickets in the wet flat-
woods region, while S-RY remained on the sand ridge but
well to the south of Hicoria.
One other bird banded at Hicoria moved to a new locale
SO-B was banded as an adult during the summer of 1967 at a
banding site just south of territory 6. Several birds that
were banded there subsequently disappeared. SO-B was found
1.8 miles to the northwest in February 1969 and nested
during that year on a narrow band of scrub. This strip of
scrub had been censused during 1968, where SO-B was not
present; its movements during 1968 are unknown. This was
the farthest away from the banding site that I located any
of my banded birds.


26
similar calls or landed nearby and bobbed and "hiccupped''
furiously. Females never initiated nor passed ahead of
the males during such activities. The "helpers," to be
discussed later, often joined in but remained behind the
female. GS-G was then attracted to the southern border
of the territory by a similar pattern of flight and calls
from SR-O. Fifteen minutes later RS-R and PiS-0 had
initiated a bout at their common border (territories 1
and 7).
Only early in the morning did I see these attempts
of one territorial bird to draw another. These seemingly
ritualized patterns of border conflict were common during
the breeding season; during the rest of the year they were
less frequent although now and then they were repeated.
Site Tenacity and Mate Constancy
During the three years that banded birds were fol
lowed at Kicoria, the majority of pairs remained at the
same place (Figs. 1, 2 and 3). At any time of year each
pair usually could be found in or close to the territory
defended during the breeding season, and from year to
year the same sites were generally defended.
Even after the young successfully leave the nest,
territorial defense remains at a high intensity with
frequent early morning bouts and continual reaction when
ever a strange bird wanders too close. But after mid
summer such tight border control relaxes. Until the next


45
by the loud upwardly inflected kwerp calls of the Scrub
Jay and remain with the distressed pair as long as they
continue calling. These other species probably do little
to aid the distressed Scrub Jays, for they remain well
away from the action. Possibly Scrub Jays aid other nest
ing species by predator recognition.
Reaction to Man
One feature which makes the Florida Scrub Jay especially
intriguing to work with is its extreme tameness toward man.
On 17 June 1967 when I first started banding at Hicoria
I walked down the dirt road by territory 3. A Scrub Jay
landed ahead of me and a juvenile near my feet. I was
surprised at their seeming unawareness of my presence.
Then the juvenile hopped between my feet, and the adult
landed on my head. I held out my hand and the adult jumped
to my arm and then to a nearby branch. I fumbled in my
pockets as though to bring out some food and again held my
hand out, but they did not fly to it. I walked up to the
adult, which remained calm until I was three-feet way,
and then it flew 10 feet down the trail.
This encounter surprised me for, to the best of my
knowledge, no one has spent any time at Hicoria feeding
Scrub Jays. In fact they did not appear to be attracted
to me for food, merely inquisitive about my presence. At
the Archbold Station Scrub Jays readily approach a person
and fly to the hand. These birds have been conditioned


53
Cape Kennedy Air Force Station, Brevard County
Elevation is 6 feet. This large area of approximately
1200 acres of scrub sustains a large population of Scrub
Jays. This community is on recent shore front sands and
is unlike most interior scrubs because wax myrtle (30 per
cent of ground cover in thickets) and saw palmetto (25 per
cent of ground cover in thickets) are much commoner. Here
the three species of oaks are prevalent (35 per cent
of thicket ground cover). The thickets are 8-12 feet
high and dense, especially near the shore line where
onshore winds and salt spray have stunted the upward
growth of plants and produced an extremely tangled, low
thicket.
The outstanding feature of this site is a network of
interconnecting dirt and paved roads crisscrossing the
area. All these roads have cleared shoulders 30-50 feet
wide, often of open white sand. Of course, there are
large open spaces with rocket launching complexes, but in
addition to these industrialized parts, open sandy places
comprise about 40 per cent of this study area. During
1969 I censused two plots of scrub thickets in the center
of the Cape where there were many open spaces but away
from the rocketry complexes; at each I found 10-11 active
nests per 100 acres. This cape region with many artificial
open, spaces resembles Hicoria and the density cf breedinc
birds is similar. Most nests are located in scrubby oaks
although several were found in the wax myrtle.


64
Archbold Station and Hicoria. Although large numbers of
breeding Scrub Jays are supported in these scattered islands
of scrub thickets, the density per unit area is rather low
(2-3 pairs per 100 acres).
Atlantic Coast Dune Thickets
An extremely long yet narrow band of very low and
dense, thickets exists on shore-front sand dunes of the
Florida Atlantic coast from Jacksonville to Fort Pierce.
Sand pines are not present and saw palmetto often is the
dominant plant. Live and myrtle oaks comprise from 20-40
per cent of ground cover in thickets. Onshore winds and
salt spray have led to an extremely dense, often impene
trable, growth (Fig. 6). Open areas exist only along
roadways or where houses have been built.
The vegetation is similar all along the coast but
the distribution of Scrub Jays varies. The snore thickets
about. Melbourne Beach, Brevard County, 20 miles south of
the iai'ge Cape Kennedy scrub area contain many scattered
breeding colonies. The beach thicket north of Cape Kennedy
toward New Smyrna Beach, Volusia County, also contains
numerous breeding Scrub Jays. These birds nest only along
the edges of the danse thickets, normally along the highway.
North of Daytona Beach, Volusia County, in the same
type of vegetation, Scrub Jays are much less abundant.
Several miles of seemingly suitable thickets exist before
another nest is located.


male and his "helpers" since courtship feeding occurs
between them. Territories without "helpers" usually
are those in which the nest or fledglings were destroyed
the year before. Rarely were first-year birds found away
from parental territories.
Dispersal by Florida Scrub Jays is remarkedly restricted.
Movement of first-year birds is delayed until the second
year. The retention of young birds in territories may
allow for a form of site tenacity, so that when these
"helpers" disperse it is only for short distances. The
farthest any banded bird moved to establish a territory
was 1.8 miles. No birds have moved between Hicoria and
the Archbold Biological Station, 2.0 miles north, where
several hundred Scrub Jays have been banded by another
worker.
The reaction of Scrub Jays to other species and to
predators is described. Whenever Scrub Jays and Blue. Jays
(Cyanocitta cristate) interact, the Scrub Jay dominates.
Florida Scrub Jays are restricted to a plant community
that combines low dense thickets with numerous open sandy
spaces. Such habitat occurs primarily at the edge of sand
pine scrub, in recently burned scrub, as scrubby flatwoods,
and as shore dune thickets. Low, dense scrub thickets
frequently do not include open sandy spaces and sc are not
favored by nesting jays, but man-made openings in scrub
habitat have increased the amount of suitable nesting habi
tat, i.a., along roadways and at edges of developments.
vii


32
aggressive behavior in adults." Ey implication juveniles
with distinctly different plumage should trigger a less
aggressive response by an adult. Within the territories
at Hicoria this was not so. Apparently territorial aggres
siveness overrides any recognition of juvenile status.
The brownish pattern undoubtedly helps as a concealing
coloration while the birds are feeding in thickets
although adults are, themselves, extremely difficult to
see in thickets.
The juvenile birds in their proper territories give
begging cries and wing flutters whenever either parent
comes close. Such begging continues through the year
although less frequently. If the parent has food it is
often passed to the begging juvenile. An adult-like
plumage is obtained in July, but even so the parent still
responds to a juvenile when it begs.
The parents frequently travel out of the territory,
usually to a food source. Male and female normally travel
together even during the nonbreeding season and often
their offspring troop along. In these cases.they join
with other Scrub Jays. As many as 30 Scrub Jays may con
gregate at a baited site not in any territory. Juveniles
beg to their own parents although they are usually ignored
The young may even beg to a strange adult and again are
ignored. Breeding males present here are not motivated
to attack juveniles with brown plumage.
The begging sequence of juveniles to their parents is
exactly similar to courtship begging of a female to her


ECOLOGY AND BEHAVIOR OF THE
FLORIDA SCRUB JAY
b7
PETER WALTER WESTCOTT
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THB
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1970

AC KNOWLEDGMENTS
I wish to extend thanks to the following persons and
organisations who aided me in various ways in carrying
out the present work. Frofessors Pierce Brodkorb, Archie
F. Carr, Jr., Albert M. Laesse, David W, Johnston, and
Ernest S. Ford of the University of Florida acted as my
supervisory committee while I was a student at the Uni
versity of Florida. Professor Brodkorb was my principal
advisor, and I especially wish to acknowledge his tireless
efforts in the preparation of this-manuscript. Dr. Oliver
L. Austin, Jr., Curator, Florida State Museum, provided
both banding material and suggestions during the course
of this study. Dr. James N. Layne of the Archbold Biologi
cal Station and Dr. Glen E. Woolfenden, University of
South Florida, provided much advice and encouragement.
Mr. Donald E. Goodman greatly aided in the tiresome enter
prise of capturing birds. I am mindful of the invaluable
grants-in-aid received from the Frank M. Chapman Memorial
Fund of the American Museum of Natural History and from
Sigma Xi.

TABLE CF CONTENTS
ACKNOWLEDGMENTS
LIST OF FIGURES
ABSTRACT
INTRODUCTION
HICORIA STUDY AREA
METHODS
RECOGNITION SIGNALS
Auditory Signals
Visual Signals
COURTSHIP
Pair Formation
Copulatorv Behavior .
TERRITORIALITY
Nest Placement
Territorial Defense
Site Tenacity and Mate Constancy
"Helpers11
Interactions of Adults and Juveniles
Role of "Helpers11
EXTRA-TERRITORIAL MOVEMENTS ....
REACTION TO OTHER SPECIES
Reaction to Blue Jays .
Fage
ii
v
vi
1
3
10
10
11
14
14
16
18
18
23
26
28
30
33
39
41
A 1
.11
I

Page
Reaction to Predators 42
Reaction to Man . 45
DISPERSAL 47
HABITAT AMD DISTRIBUTION 52
Scrub Jay Habitat in Florida 52
Hicoria 55
Cape Kennedy Air Force Station 5 8
Jonathan Dickinson State Park 59
Ht. Plymouth Area 5 9
Lake Jackson 60
Ridge Scrub 60
Scrubby Flatwoods West of Hicoria ... 61
Atlantic Coast Dune Thickets 64
"Big Scrub" 67
Ocala National Forest 6 7
Habitat Summary 68
Scrub Habitat Without Breeding Scrub Jays. 69
Kissimmee Saint Cloud 6 9
Weeki Wachee 72
Lake Wales 72
Florida Panhandle Scrub 73
FACTORS CONTROLLING DISTRIBUTION 76
LITERATURE CITED 83
BIOGRAPHICAL SKETCH 35

LIST OF FIGURES
Figure Page
1. Map of study area at Hicoria 1968 6
2. Map of study area at Hicoria 1969 20
3. Map of study area at Hicoria 1970 22
4. View of Hicoria study area 57
5. View of scrub thicket two miles south
east of Hicoria 63
6. View of Atlantic coast thicket at
Marineland 66
7. Map of central Florida peninsula with
Scrub Jay habitat 71
8. View of scrub thicket at Fort Pickens
State Park 75
v

Abstract of Dissertation Presented to the Graduare Council
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
ECOLOGY AND BEHAVIOR OF
THE FLORIDA SCRUB JAY
By
Peter Walter Westcott
August, 1970
Chairman: Professor Pierce Brodkcrb
Major Department: Department, of Zoology
This study gives an outline of the social structuring
of breeding Florida Scrub Jays (Apheiocoma c. coerulescens)
and discusses the restrictions placed on distribution by
the type of habitat selected for nesting. The social struc
ture was studied at Hicoria, Highlands County, Florida,
where 230 individuals were banded.
Certain calls and postures used in social interactions
are described, and pair formation and copulatory behavior
are discussed.
In Florida the Scrub Jay is territorial and pairs for
life. The territory is defended from intrusion by other
Scrub Jays, but first-year birds (young of that pair from
the year before) are retained in the territory. They
occasionally act as helpers" by caring for the young,
and frequently aid in territorial defense. Although they
restrict their movements to the territory, usually these
extra members remain away from the breeding pair. A bond,
similar to a pair bond, may be present between the breeding
vi

male and his "helpers" since courtship feeding occurs
between them. Territories without "helpers" usually
are those in which the nest or fledglings were destroyed
the year before. Rarely were first-year birds found away
from parental territories.
Dispersal by Florida Scrub Jays is remarkedly restricted.
Movement of first-year birds is delayed until the second
year. The retention of young birds in territories may
allow for a form of site tenacity, so that when these
"helpers" disperse it is only for short distances. The
farthest any banded bird moved to establish a territory
was 1.8 miles. No birds have moved between Hicoria and
the Archbold Biological Station, 2.0 miles north, where
several hundred Scrub Jays have been banded by another
worker.
The reaction of Scrub Jays to other species and to
predators is described. Whenever Scrub Jays and Blue. Jays
(Cyanocitta cristate) interact, the Scrub Jay dominates.
Florida Scrub Jays are restricted to a plant community
that combines low dense thickets with numerous open sandy
spaces. Such habitat occurs primarily at the edge of sand
pine scrub, in recently burned scrub, as scrubby flatwoods,
and as shore dune thickets. Low, dense scrub thickets
frequently do not include open sandy spaces and sc are not
favored by nesting jays, but man-made openings in scrub
habitat have increased the amount of suitable nesting habi
tat, i.a., along roadways and at edges of developments.
vii

Even so, the total area available to Scrub Jays for
nesting is small. This habitat frequently exists as
small areas of thickets isolated from similar suitable
habitat.
The distribution of breeding Scrub Jays in Florida
is outlined. Only a few large expanses of suitable
habitat support dense populations of Scrub Jays. Al
though dispersal distances are not great, most small
isolated patches of scrub habitat are occupied by
Scrub Jays when in the vicinity of these centers of
population, but seemingly suitable habitat isolated by
20 or 30 miles from the nearest Scrub Jay population
is occasionally unoccupied. The Florida Panhandle -
has a large stretch of sand pine scrub community and
much of it appears ideal as Scrub Jay nesting habitat.
No Scrub Jays inhabit this extensive habitat which is
separated by 100 miles from the closest peninsular Scrub
Jay population.
The retention of first-year birds in parental ter
ritories and the reduced dispersal abilities -of non
breeding adults are discussed as a controlling factor in
the distribution of the Florida Scrub Jay. The sociality
of the breeding jays may be a behaviorial adaptation to
a limited breeding habitat similar to the case of the
Mexican Jay (A. ultramarina) in Arizona.
Vlli

INTRODUCTION
The Scrub Jay, Aphelocoma coerulescens (Bose), occurs
in two separate areas of North America. Throughout the
western United States and the mountains of Mexico this
species lives in a variety cf habitats, often in large
numbers (Pitelka, 1951; Westcott, in press). Although
the western population is composed of sixteen subspecies,
all appear to have similar social organizations. sin
gle eastern subspecies, A. c. coerulescens, inhabits
central Florida. Not only is it isolated by over a
thousand miles from the closest western population but it
also exhibits completely anomalous patterns of sociality.
Unlike its western representatives, the Florida Scrub Jay
is restricted in habitat and is often quite scarce. Western
Scrub Jays are normally secretive, yet Florida Scrub Jays
are notably tame.
Reports of extra individuals acting as "helpers"
about the nest (Grimes, 1940) also suggest a degree of
sociality unlike that of western Scrub Jays, but little
information has appeared concerning this example of
sociality in the Florida Scrub Jay. Amadon (1944) exam
ined the activities of breading pairs about the nest and
presents details of courtship, nest building, and care
1

2
of the young. Although he occasionally observed non
breeding birds around active nests, Amadon concluded
that they do little to aid in the care of young.
Although Florida Scrub Jays are found scattered
over much of central Florida, most local populations
appear small, and some areas, such as in the Florida
Panhandle, lock ideal for nesting Scrub Jays but none
have been reported. The study reported here gives a
detailed outline of the social structuring of a breed
ing group of Florida Scrub Jays and analyzes the social
behavior of these birds in relation to dispersal and
habitat distribution.

HICORIA STUDY AREA
The study area at Hicoria is located two miles due
south of the Archbold Biological Station, Lake Placid,
Highlands County, Florida, and is at the extreme southern
end of the sand ridge region of central Florida. At an
elevation of 140 feet above sea level, this locale con
tains a plant community that combines sand pine scrub
and pine flatwood elements; its special vegetational com
ponents are termed scrubby flatwoods by Laessle (1942).
The vegetation is discussed in detail in a later section.
A dominant feature of this particular site is a
multitude of large and small open sandy spaces. As dis
cussed later, this combination of dense scrub thickets
and open space represents the optimum habitat for Florida
Scrub Jays and gives a density of breeding birds at
Hicoria that facilitates the studies of social behavior.
The open areas of this site are not typical of the
scrubby cover which elsewhere is extremely dense and
extensive. During the early years of this century Hicoria
was the site of a town of one thousand persons, a glass
factory, and a lumber mill. Today the only remains are
the scattered open spaces where houses or other build
ings once stood. The outline of the study area in Fig. 1
shows the pattern of open spaces resulting from the aban
donment of the town and factories.
3

4
To the east, this site is separated from the main
body of the sand ridge scrub by extensive orange groves.
The western edge of the study area is at the border of
the sand ridge and drops several feet to pine flatwoods,
although many scrub thickets continue to the west. Scrubby
vegetation extends south for a mile and then merges into
pine flatwoods with widely scattered scrub thickets. The
scrub at Hicoria continues north along a railroad in an
unbroken, although narrow, band to the Archbold Eiological
Station.
Little human interference occurs at Hicoria. Occa
sionally a hunter stops to shoot at a rabbit, someone cuts
some hickory for barbecue wood, or checks on the cattle
run on the extreme north edge of this area. Otherwise I
saw few people during my three years of field work at
Hicoria. Although Florida Scrub Jays are notoriously tame
and in many areas readily follow people to beg for food,
at Hicoria the birds usually ignored my presence. I
regularly camped in the middle of the study area, but
took pains not to leave my food about.

Figure 1
- Map of study area at Hicoria showing terri
tory boundaries and Scrub Jays residing in
each territory during the breeding season
1968. Note: in territory 12 SY-Y died and
S-EY moved to territory 4.
KEY
1 inch = 300 feet
Shaded area = scrubby thickets
Light area = open sandy spaces
Jj = paved road
j = dirt road
railroad
fence
territory boundary
Letters designate a specific Scrub Jay, i.e
RS-R
o* = breeding male
$ = breeding female
* = "Helpers"
All birds without a special sign following
their letter designation are young of the
year, i.e., 1G-YS in territory 2.

6

METHODS
The social behavior is based upon information gathered
from the banding of 230 Scrub Jays at Kicoria and at scat
tered sites to the west and south- To simplify the report
discussion is limited to movements of banded birds in the
200 acre study plot outlined in Fig. 1, the center of the
Hicoria region.
Jays were captured in mist nests and baited Potter
traps. They quickly learned to avoid the mist nets, and
various methods had to be devised to lure them in. Most
effective was the use of a hand-held Scrub Jay which, when
shaken slightly, gave loud alarm calls. This attracted
many local jays and I rhen chased them toward the nets.
Each bird received three bands, two colored plastic
coil bands and or.e aluminum silver colored United States
Fish and Wildlife Service band. Each individual was
identified by a series of letters connoting .the color
sequence of bands: RS-R was a bird with red plastic and
silver aluminum bands on the right leg and red plastic
band of the left leg, 1B1G-S had light blue and light
green bands on right leg and silver band on left leg.
All birds captured in traps or nets had the silver band
placed on the right leg unless definitely known to be a

when adult had silver bands on the right leg. The juve
niles banded during 1963 had one colored band on the
right and one colored and the silver band on the lefr,
and juveniles banded m 19S9 had two colored bands on
the right and one silver on the left leg. Duco cement
placed on plastic colored bands effected a durable seal,
but occasionally bands were lost and the bird had to be
recaptured.
To follow the movements of banded birds I used a
30 power telescope mounted on a gun stock. In this way
I could read band sequences from as far as 100 yards and
sc did not have to disturb the bird. Occasionally a
single-sided blind was used but normally it was not needed
To study the local distribution of Florida Scrub Jays
I attempted to examine all contemporary Scrub Jay breeding
sites in Florida. This appears a prohibitively difficult
task, and several small areas were undoubtedly overlooked.
Yet I believe I was able to find nearly all of the breed
ing regions. The many letters from Florida Audubon member
in reply to my solicitations for information, greatly fcil
itated this search.
An invaluable aid when hunting for Scrub Jays is a
portable tape-recorder with recordings of Scrub Jay alarm
calls. These calls quickly attract any local Scrub Jays.
As a measure of preferred habitat I used the number
of active nests located in one hundred acres.
Difficul-

9
when nesting areas consisted of many scattered patches.
Even so, centers of population numbers were located and
their common physiographic features assessed.
Ten areas that included a variety of types of plant
cover, yet harbored nesting Scrub Jays, were examined in
detail. A nest census was made. Plant cover was analyzed
and the amount of ground in the thickets shaded by the
dominant woody plants was estimated. Unless I am dis
cussing a specific plant community such as the Florida
sand pine scrub, the word scrub connotes a thicket of
evergreen woody plants that is low and dense.
Field work began in June, 1967, and was completed in
May, 1970.

RECOGNITION SIGNALS
The calls, postures, and movements that are used in
social interactions are treated; other variations have
been described by Amadon (1944). In this section the
author describes individual calls and postures and some
times implies their motivation. Later sections discuss
how these calls and postures are combined in complex
behavior.
Auditory Signals
Kwerr or kwerp.--The Florida Scrub Jay utters a
variety of calls, yet most are variations of a rather
harsh and grating note. Amadon phoneticizes this basic
call as ka. To me the basic call contains more consonant
sounds (kwerr). This call, as Amadon notes, is used for
several functions, and certain variations appear to be
used consistently in similar situations or to express
particular motivations.
An upward inflection of the kwerr call normally
indicates a heightened motivational level. Such calls
are given when some alarming object is present such as
a predator.
An increase in intensity or loudness of this call
also indicates excitement but is normally used when in
10

11
a confrontation with another Scrub Jay. As Amadon states,
this extra loud call is often lengthened into a screech.
Finally, the rapidity of presentation of the call
represents another variation of the kwerr. Especially
during the early morning a rapid series of kwerr or more
frequently kwerp-kwerp-kwer? calls are heard. Again
this is a sign of increased excitement. It is normally
used in antagonistic bouts with other jays, especially
after a series of loud kwerrs.
"Hiccuo'1This peculiar sound apparently is produced
only by the female. Amadon's pronunciation of it as klok
is quite exact. It is a hollow sound repeated rapidly so
that it sounds like a hiccup. It is often used in antag
onistic contexts yet it frequently is given when birds
are quietly perched alone or during courtship. The loud
ness of presentation expresses a more excited state.
Begging call.This cry is always used with a wing
flutter and is the characteristic sound of the young in
the nest as a parent arrives. The same call is used by
a female toward her mate. I have never heard a paired
male use it, although I have heard it used by an adult
male toward a female in a small cage.
Visual Siar.als
As with the auditory signals most of these visual
postures are those used by the Scrub Jav when reacting
in some agonistic context either toward their own soecies
or toward a predator.

12
Upright posture.This is difficult to classify as
a special behavior because Florida Scrub Jays appear to
perch naturally with the body upright and tail hanging
straight down. Yet the male of a breeding pair habit
ually perches in this posture on a. tall object from which
the bird has a clear view over much terrain. It is often
possible to identify a Scrub Jay from a distance by this
posture. This position can be. classed as neutral in
motivation.
Dip.--In this posture the head and body are bent
downward and the tail is raised above the horizontal.
This posture always expresses increased excitement and
usually is an alarm, reaction.
Bobbing.Again when a bird is excited one reaction
is a rapid bobbing of the whole body without moving from
the perch. The degree of excitement can be roughly
measured by the rapidity of the bob.
Undulating flight.--Normally a Scrub Jay flies from
one place to another with a steady, slightly undulating
flight. When highly motivated this flight is more deeply
undulating. This pattern is normally accompanied by a
series of loud kwarp calls and is often exaggerated by
flying slowly. Undulating flight often follows an encoun
ter with another jay.
Bill up.The bird tips the. bill up and remains
silent, and the body is often slightly crouched. This
is a submissive posture and is usually followed by the
bird flying away.

Wing flutter and crouch.This is a begging posture
and is almost always used with a begging cry. The body
is pressed close to the perch or the ground and the wings
are rapidly fluttered.
Bill wiping.The bill is rapidly rubbed on both
sides of the perch as though attempting to clean some
thing off. This movement is used at the conclusion of
a wide variety of activities: after fighting, after
swallowing, after courtship feeding, or after drinking.
It is so frequently used that perhaps it can be viewed
as the final consummating portion of a sequence of actions.

COURTSHIP
Unmated birds frequently travel and feed in flocks,
but no special group activity relating to courtship has
been found. This is unlike the Blue Jay, Cyanocitta
cristate (Linnaeus), which performs a highly ritualized
group activity (Hardy, 1961).
Pair Formation
The initial stage of forming a new pair bond begins
with the male restricting his movements to a particular
undefended area of scrub. After the male is first found
restricting his movements, a female soon accompanies him.
On 10 January 1970 M-dBS, a male, had restricted his
movements to a region just west of territory 4. He con
tinued feeding in territory 4, and five days later S-BY,
a female, was first seen following the male. After this
date they rarely returned to the birds in territory 4 and
the male began to defend actively their new region. I
first saw M-dBS chase another jay from the new region
on 15 January, and S-BY followed him. The female invariably
followed the male during the next few weeks as he wan
dered through the territory and especially when he chased
intruders. Such constant accompaniment undoubtedly helos
reinforce the pair bond.
14

5
Although more than 50 juvenile Scrub Jays have been
banded within the 200 acre study area at Hicoria, M-dBS
was the only male banded here as a juvenile to establish
a new pair bond and territory in this same area. One
other bird banded as a juvenile settled to the west of
the study area, and I followed him as he established a
pair bond and territory. O-SW, the male, settled in
a small patch of scrubby live oaks surrounded by wet
flatwoods approximately 1200 yards west of territory 8,
in which he was banded as a juvenile in 1968. On
8 December 1969 he was first seen in the scrubby patch
and had another bird in attendance. This bird was not
banded but gave the female "hiccup" calls as soon as I
approached. She was subsequently captured and banded
RS-dB.
On 8, 9 and 10 January 1970 O-SW continually approached
the female who at once crouched and gave begging calls and
wing flutters. The male immediately passed a morsel of
food to the female. After passing the food no ether
special calls cr movements occurred except that usually
they both bill-wiped. Occasionally a female gave "hiccup"
calls after courtship feeding, but usually both began some
completely unrelated activity such as feeding or preening.
As I slowly approached O-SW on 10 February 1970 he
was perched on the tallest branch available in this scrub
patch. There was no way I could approach unnoticed and
at 70 yards he gave upwardly inflected kwerp calls. At

16
once RS-dB jumped up from the ground. O-SW bobbed and
continued calling, and RS-dB "hiccupped." Then the male
flew off giving rapid kwsrps and the female immediately
followed, giving the same calls.
Whenever a disturbance occurs it is the male that
first flies and the female follows; this pattern is fol
lowed by all mated pairs. Pair formation then is a
matter of the two birds remaining in close association,
one bird, the male, passing food to the female and the
female following closely after the male in all encounters.
Although the above discussion deals with the forma
tion of a new bond, it also explains the interaction of
paired birds, frequent courtship feeding even during the
fall and winter, and persistent following of the male by
the female.
Copulatory Behavior
The complete copulatory pattern was observed only
once, although I spent many hours behind a blind hopeful
of following this sequence.
On 10 April 1969 GS-G, the male, presented a cater
pillar to his mate S-RB at 1030 and brought her food
eight times during the next 50 minutes, even though she
continued to feed on her own. Frequently, after court
ship feeding, the female gave rather subdued "hiccup"
calls and a series of melodic notes and calls as though
she were whispering. At 1120 he nopoed over to her and

17
as usual she at once gave begging cries, and crouched
down. He fed her but remained standing before her. She
continued to beg and GS-G began to hop around her. His
circling movement was a curious stiff-legged affair with
his neck extended and bill pointed slightly down. His
tail was spread and slowly swung from side to side. The
wings were half spread and lowered, almost dragging on
the ground. He circled her three times while she remained
crouched with her bill raised, a submissive posture. The
male was silent, and just before mounting, the female also
became silent. The male mounted the female for approxi
mately six seconds and then hopped to a nearby perch
where he vigorously bill-wiped and shook his plumage.
The female also began bill-wiping, ruffled her feathers,
and bill-wiped again. She then "hiccupped" rather
loudly and followed the male when he flew off. This
pattern is essentially the same as that reported by Brown
(1963) for the Scrub Jay in California.

TERRITORIALITY
The Florida Scrub Jay is a highly territorial bird.
At Hicoria territorial boundaries frequently made broad
contact and border conflicts were common. The boundaries
were fairly easy to map and Figs. 1, 2 and 3 outline the
territories present at Hicoria during the three years of
my study. Some daily shifting in the location of the
defended border occurred but it was minimal. Whether it
shifted or not, at all times a definite boundary was
recognized by other breeding jays,- and if a foreign jay
crossed, some sort of conflict ensued.
The area defended, among the 25 territories measured
over three years, ranged from 3.8 to 7.5 acres, but most
were close to the average of 6.0 acres. Each territory
included a large amount of scrub edge and open space with
which Hicoria abounds. The western side of the study
area included a large stretch of very dense scrub. The
only jays nesting in it were at the edges. I never found
pairs defending ground in the center of this dense area.
Nest Placement
The birds invariably placed their nests two or three
feet in from the edge cf an open space. Usually myrtle
oaks wars selected for nest building but also used were
wax myrtle, Chapman oak, and in one case a scrub palmetto
18

Figure 2
- Map of study area at Hicoria showing terri
torial boundaries and Scrub Jays residing
in each territory during the breeding season
1969. (See key for Fig. 1).

fl

igure 3 Map of study area at Hicoria showning terri
torial boundaries and Scrub Jays residing in
each territory during the breeding season
1970. Note: no young were banded this year
(See key for Fig. 1).

22

23
that was intertwined with a myrtle oak. Scrub Jay nests
were well camouflaged and were difficult to see. In order
to locate all the nests I had to search through practi
cally every tree in the area.
Nest building began as early as 20 February and
extended to 2 June at Hicoria. There was no definite
time when most pairs were nest building or incubating;
it was a protracted period. One pair had young out of
the nest while another was nest building. Unlike Blue
Jays (Hardy, 1961) the Scrub Jay pair defended its
territory from all Scrub Jay intruders no matter what
the stage of the breeding cycle of the defender or the
intruder. GS-G and S-RB did not commence nest building
until 18 May 1968, yet they actively defended a rather
large territory from all Scrub Jay intruders from March
onwards. The nest was completed on 21 May, but no eggs
were laid until 4 June.
Territorial Defense
The male of a pair spends much time perched in an
upright position with tail held straight down. This
behavior is most common during breeding season and occurs
while the female is both on and off the nest. From the
perch the bird can readily see all of his territory. This
same posture is used by Scrub Jays in the fall and winter
when in a flock. Consistently one sees a single bird,
not necessarily a male, perched high above the others
feeding below. If this bird drops to the ground, within

24
a short time another flies up and replaces him. A bird
may perch there for as long as 30 minutes. The high
bird invariably is first to sight an approaching intruder
and begins bobbing and calling, usually with upwardly
inflected kwerr. At once the other birds fly or hop into
the branches nearest to them. This position of the bird
is undoubtedly a form of sentinel behavior both during
and after the breeding season. Yet on such a perch not
only can he see over his whole territory, but others
outside can readily see him. Possibly, then, this posture
may function as a means of proclamation of territory.
Rarely does an outside bird intrude far into a
territory during the nesting period, but boundary disputes
are frequent. Chasing is the normal means of territorial
defense. When another bird approaches or crosses the
boundary the male of the invaded territory flies directly
toward the intruder. Physical contact is almost never
necessary. The encounter quickly terminates with the
intruder rapidly retreating. The attacking male veers
off abruptly at his territory boundary and almost never
extends his flight beyond it. His return flight often
includes a deeply undulating flight and a series of loud
rapid kwerp calls. At times the initial attack by the
defender is silent; when a call is used it is a loud
kwerp.
The mate often follows the male but rarely takes
part in any dispute. She perches nearby in the territory,

25
actively bobbing and giving loud "hiccup" calls. When the
male flies back, the female usually follows in an undulating
pattern and calling. She sometimes "hiccups" in flight.
These border rebuffs are not cne-sided affairs. If
a male chases a bird from his territory the roles are
reversed when the latter's territory is invaded. Each
breeding male then has a definite boundary over which he
will allow no intrusion during the breeding season.
These border disputes occur most frequently in the
morning. Later in the day when the heat and humidity are
high Scrub Jays become quiet and little activity is evi
dent. At sunrise the jays are very active and territorial
pairs perform a series of encounters that are repeated
at the same place and time, day after day during the
breeding season.
The following description of the movements of GS-G
and mate in territory 3 will describe these encounters.
On 21 May 1969 at 0650 just before sunrise GS-G came
swooping northward toward the boundary between his and
BS-d3's respective territory (territories 3 and 5).
GS-G was flying an exaggeratedly slow and very undulating
pattern, calling loudly and rapidly kwerp-kwsrp. At once
BS-dB flew toward GS-G using the same pattern of flight
and else calling loudly. When approximately 20 yards
apart they simultaneously circled and flew back toward
the center of their respective territories, still calling
rapidly. The female sometimes followed behind and gave

26
similar calls or landed nearby and bobbed and "hiccupped''
furiously. Females never initiated nor passed ahead of
the males during such activities. The "helpers," to be
discussed later, often joined in but remained behind the
female. GS-G was then attracted to the southern border
of the territory by a similar pattern of flight and calls
from SR-O. Fifteen minutes later RS-R and PiS-0 had
initiated a bout at their common border (territories 1
and 7).
Only early in the morning did I see these attempts
of one territorial bird to draw another. These seemingly
ritualized patterns of border conflict were common during
the breeding season; during the rest of the year they were
less frequent although now and then they were repeated.
Site Tenacity and Mate Constancy
During the three years that banded birds were fol
lowed at Kicoria, the majority of pairs remained at the
same place (Figs. 1, 2 and 3). At any time of year each
pair usually could be found in or close to the territory
defended during the breeding season, and from year to
year the same sites were generally defended.
Even after the young successfully leave the nest,
territorial defense remains at a high intensity with
frequent early morning bouts and continual reaction when
ever a strange bird wanders too close. But after mid
summer such tight border control relaxes. Until the next

27
breeding season the pairs spend more time wandering out
of their territories and mixing with other Scrub Jays.
Yet whenever the pair returns to their territory they
chase intruders. In this fashion the boundaries vary
little from one season to the next.
Apparently the Florida Scrub Jay mates for life. Two
of the pairs at Hicoria are known to have been paired for
at least four consecutive breeding seasons, RS-R with v'7S-Pi
and GS-G with S-RB. WS-Pi, a female, is of special interest.
I banded her as an adult in the summer of 1960 at Hicoria
within 200 yards of the territory now occupied. She is
thus at least 12 years old and still breading.
Mate constancy was the rule with the majority of the
other mated pairs. The following pairs were together during
1968 and continued as pairs through 1970: SR-B with
S-BO, SB-N with SB-dB, RS-Pi with MS-B, and SG-B with
SW-R.
Only three pairs broke up in this 200 acre site.
NS-N, S-WN and young disappeared in late October 1968.
I have no specific knowledge as to the cause- of their
disappearance, except that Scrub Jay feathers were found
in the territory. I surmise that some predator killed
one or more of this group, probably including one of the
parents, although two of the young were later found asso
ciated with a group of non-territorial birds 500 yards to
the northwest. The parents and remaining young were not
located during the next two years, although I roamed over

28
the region as far as three miles from Hicoria looking for
banded birds.
SG-B and SW-R nested for two years in territory 9,
but each year the young vera destroyed while in the nest.
During 1970 the pair disappeared and the territory remained
unoccupied. Perhaps continual nest destruction induced
their departure. Yet other pairs continued territorial
occupancy after nest destruction.
On 20 May 1568, a feral cat located the nest of SY-Y
and S-BY. All the nestlings were killed and apparently
so was the male, SY-Y, for he was never seen again. Dis
location of a breeding pair probably occurs only after
some major catastrophe. Otherwise a mated pair remains
at its original territory year after year.
"Helpers"
I have stressed the territorial nature of the social
organization of breeding Florida Scrub Jays. They are
territorial, but each territory often contains more than
a mated pair. For instance on 12 June 1969 territory 8
contained MS-B (female), RS-Pi (male), O-SG (female),
O-SW (male), dBR-S, PiY-S, and PPi-S. The first two
were the mated pair, the last three juveniles. The
remaining two looked like adults (juveniles have brown
heads until their first molt in July), vet they were
definitely tolerated in the territory and had been all
spring. They were, in fact, the young produced by that

29
pair in that territory the preceding season of 1968, for
I banded them in the nest.
This pattern of retention of young of the year into
and through the next breeding season as members of the
territory is prevalent at Hicoria and at other areas in
Florida from which I have banding information. The
presence of helpers at the nest was first reported for
the Corvidae by Skutch (1335). Grimes (1940) first
suggested that Florida Scrub Jays had extra birds active
about the nest, and Am.adcn (1944) in his study of nest
ing Scrub Jays at the Archbold Station occasionally found
extra birds tolerated at certain nest sites. He tenta
tively concluded that many first year birds may not breed
and are sometimes tolerated about an active nest.
During the breeding season of 1968 certain birds
banded earlier were found associated with breeding pairs.
As outlined in Fig. 1 four extra birds were found, one
in each of four out of nine territories. Using the
flight feather technique of Pitelka (1945, 1951), I
tentatively identified three of the four as first-year
birds. Plumage characters were difficult to analyze
because all the jays had worn plumage at this time of
year.
In 1969 seven "helpers" were found in four of ten
territories, one in territory 2, three in territory 4,
two in territory 8, and one in territory 10. The other
six territories had only a mated pair in occupancy. Six

30

of the seven "helpers were first-year individuals that
had been banded the previous year in the nest of the pair
with which they were new associating. The other was S-EY,
an adult, and will be discussed later.
Finally in 1570 six of nine territories and nests in
the sane Hicoria site had "helpers," one in territory 1,
one in territory 2, one in territory 3, one in territory 5,
three in territory 4, and one in territory S. All of these
"helpers" had been banded the year before as juveniles and
in the sane territory.
Approximately half of the territories at Hicoria had
extra members. Banding data show that they were invariably
juveniles of that pair from the year before, with the excep
tion of S--BY noted previously. Although reminiscent of
the social organization of the Mexican Jay (Aphelocoma
ultramarina) in Arizona, in which nonbreeding birds
frequently function about an active nest (Brown, 1963),
this case is special in its combination of territoriality
and sociality (c.f. Skutch, 1961).
The territories that had no "helpers" during 1969
and 1970 were those in with the fledglings either were
destroyed by predarors or disappeared from the study area
and surrounding habitat.
Interactions of Adults and Juveniles
re
After the young leave the nest thev become
ring, remaining near the nest site and very
extremely
cruiet.

Even when a disturbance alarms the parents, the young
remain well hidden and silent. As newly iledged youngsters
are vulnerable to predation, this stage of extreme timidity
is adaptive. The parents quickly recognize a predator and
may draw its attention with their loud alarm call, upwardly
inflected kwerp; the silent young may remain unseen as
the predator is distracted by the parents. In fact, many
young are lost during these few weeks, yet possibly many
more would be killed if they were not secretive. About
two weeks after leaving the nest the juveniles first
become active.
Juvenile Florida Scrub Jays have light brown plumage
on head and back and lack the light blue forehead of the
adult until their first molt in late July. It is easy to
recognize juveniles by this trait. I saw no special
reaction by adults to birds with brownish heads except
when young from other territories inadvertently wandered
over another adult's territory. The resident male quickly
chased these wandering youngsters. The ensuing chase was
especially intense and afforded the only occasions that I
viewed physical contact during an attack. The male's
cortege of mate, helpers, and young often join the attack
by following the male and calling raucously. As with
border disputes, normally only the male initiates these
attacks.
Hir.de (1961) suggests that "the juvenile usually

32
aggressive behavior in adults." Ey implication juveniles
with distinctly different plumage should trigger a less
aggressive response by an adult. Within the territories
at Hicoria this was not so. Apparently territorial aggres
siveness overrides any recognition of juvenile status.
The brownish pattern undoubtedly helps as a concealing
coloration while the birds are feeding in thickets
although adults are, themselves, extremely difficult to
see in thickets.
The juvenile birds in their proper territories give
begging cries and wing flutters whenever either parent
comes close. Such begging continues through the year
although less frequently. If the parent has food it is
often passed to the begging juvenile. An adult-like
plumage is obtained in July, but even so the parent still
responds to a juvenile when it begs.
The parents frequently travel out of the territory,
usually to a food source. Male and female normally travel
together even during the nonbreeding season and often
their offspring troop along. In these cases.they join
with other Scrub Jays. As many as 30 Scrub Jays may con
gregate at a baited site not in any territory. Juveniles
beg to their own parents although they are usually ignored
The young may even beg to a strange adult and again are
ignored. Breeding males present here are not motivated
to attack juveniles with brown plumage.
The begging sequence of juveniles to their parents is
exactly similar to courtship begging of a female to her

33
reate, and the response is usually the sane, food being
passed to the beggar. Obviously the continued occurrence
of courtship feeding between mated birds is a means of
pair bend reinforcement. It is seen most frequently
during the breeding season but occurs all year round.
Possibly as juvenile birds continue to beg and the parents
respond and feed them, an attachment something like a pair
bond is established. As the next breeding season approaches
the parents are increasingly attentive to each other, yet
the young from the previous season are still present and
tolerated by the male. Occasionally they beg and he
responds by passing a morsel of food. I suggest that these
one year old birds can remain in their parents' territory
because a loose bond is present between the "helpers and
the breeding male similar to a pair bond.
Role of the "Helpers"
The role of these "helpers" in the organization of
territorial members is often vague. I call them "helpers"
but this term is, at times, misleading. It is a term
that has been used in the past and I shall continue its
use, but "auxiliary territorial member" is perhaps more
accurate.
I never saw these "helpers" take part in nest build
ing. Usually the first-year birds ignore the adults'
nesting procedures and quietly feed somewhere in the
territory. When voung are in the nest and the aars-ts

34
actively feed them, first-year birds occasionally follow
one parent toward the nest. At such times the breeding
male often supplants the first-year bird by landing next
to it and sometimes pecking. The "helper" either gives
a submissive posture (slightly crouched and bill upraised)
and then flies off, or else flies a short distance at
once. Rarely does the male chase after the "helper."
On 12 May 1969 in territory 2, SR-O, male, was not
in sight as PiS-W, female, flew to the nest and fed a
youngster. She flew off and immediately 1G-YS, a female
first-year bird, flew to the nest and peered in. The
hestlings began begging and 1G-YS moved her bill toward
them as though to pass food but none was passed. Again
two days later, just after the female left the nest and
the male was not in sight, 1G-YS flew up to the nest and
this time fed one of the youngsters.
When the young leave the nest the parents continue
feeding them and now the first-year birds more frequently
bring food to the fledglings. In territory 1 in 1968,
the young left the nest on 18 March and for the next three
days SR-W, female first-year bird, brought food to the
fledglings at least twice each day. In 1969 M-PS in
territory 10 fed young after they were out of the nest.
Thus, the first-year birds present in the territory occa
sionally function as "helpers" with the young. Most of
the time, however, they remain somewhere in the territory
but away from the nest or new set of youncr.

35
During aggressive encounters with neighboring birds
these first-year birds m.cst readily take an active role
or act as "helpers." Whenever the male gives chase to
an intruder, the first-year members of the territory at
once fly toward him and follow behind, frequently calling
as loudly as the male. A border encounter often entails
three or four birds, loudly calling and chasing up to the
invisible territory boundary. The male then wheels back
and the others follow him back into the territory, all
still calling loudly. Certainly a mob of jays, all
seemingly bent on attack or chase, is much more effective
in instilling a sense of invulnerability about a territory
border than would only a single male.
In the same way the seemingly ritualized early morn
ing border encounters include the "helpers," if present,
in that territory. In these cases two mobs of jays may
approach each other and suddenly wheel and fly apart.
If a territory contains more than one "helper" all do
not necessarily join in these border contests.
In 1969 territory 4 contained two adult.breeding
birds and three "helpers." On 10 May an unbanded bird
flew through the territory. At cnce SR-3, the male, took
after the intruder and the other four birds feeding with
the male followed. The unbanded bird sailed rapidly
across the territory. The five attackers came back to
the feeding site, but all remained in the highly agitated
state for at least 10 minutes. S-30, the female, sat on

36
a tall perch and bobbed and "hiccupped." This enticed
S-BY, a female helper, to fly up where she commenced
"hiccupping. As they fell silent SR-B (male) flew off
50 feet, giving loud rapid kwerps. Again the other four
members of the territory followed, and another series of
bobbing and "hiccupping" ensued.
Not only did SR-B and S-BO in territory 4 have more
helpers than did other breeding pairs, they also had one
helper that was an adult female. This bird, S-BY, was
the mate of SY-Y in 196S. On 20 May a feral cat attacked
and killed their young in the nest and apparently also
killed the male, SY-Y. For the next week S-BY remained
in the region of the destroyed nest but more and more
regularly she fed with the members of territory 4 at a
feeding site that I had been supplying between their
territories. By 25 May she no longer spent any time in
her old territory and followed along with SR-B, his mate,
and helpers into territory 4. At this time SR-B and mate
were attending young in their nest, and S-BY began to visit
the nest with food. Whenever SR-B saw her at the nest he
gave chase, but she never flew far. She always rrtoved off
when he was aggressive. On 26 May I first saw her beg
toward SR-B. He ignored her but did not chase. Thereafter
she frequently bagged toward him, and he usually responded
with a morsel of food. She remained in that territory as
the only adult "helper" that I have found. S-BO, female,
never became upset by the presence of S-2Y and for 13 months

37
S-EY remained so tied to this pair that it appeared that
SR-3 had two mates. Although he regularly fed her when
S-BY begged, 3-BO remained his dominant mate and was the
breeding female the next year.
In. 1963 S-EY was still in attendance. She did not
help with nest building, but regularly brought food to
the young in the nest. During this breeding season the
male .id not attempt to chase S-BY from the nest. She
was then truly a "helper" at the nest. The other two
first-year birds in attendance did not attempt any such
close association about the nest; they acted similarly to
first-year birds in other territories.
Interestingly, in February, 1970, S-BY and M-dBS
paired and established a territory just west and adjacent
to territory 4. M-dBS was one of the first-year birds in
attendance in territory 4 while S-BY was also a "helper."
The Florida race is unique among Scrub Jays in that
family flocks defend their territories with all individuals
helping in various degrees. As far as known it is found
in none of the western races {Hardy, 1961). .
The social organization of these flocks is as would
be expected. The breeding male is dominant over the first-
year "helpers," although normally they appear to ignore
each other. Only occasionally did an adult female supplant
any of her first-year offspring in the territory and I never
saw a female chase or attack a "helper." In a territory
that had more than one "helper" no special hierarchy was

33
established among the "helpers" at first. During the
first winter, at times a male first-year bird would sup
plant a female sibling by displacing her from a perch or
by pecking at her, but again the usual pattern was of no
interference.

EXTRA-TERRITORIAL MOVEMENTS
About 60 per cant of the terrain at the Hicoria study
area was tied up in defended territories. The remaining
area was inhabited by small loose groups of nonbreeding
birds and by territorial birds which often wander out of
their home area. In 1968 two feeding stations were main
tained, one in a non-defended area and one immediately
between two territories. At feed site 2, placed between
territories 5 and 4, only the birds inhabiting those two
territories visited this food site-, accompanied by S-BY
after her mate disappeared. Although border conflicts
between BS-d3 and RS-B occurred frequently, encounters
were infrequent at this food site. Birds from the two
territories hardly ever fed here at the same time, and
this obviously reduced the necessity for territorial de
fense at this site. During the few times both groups
were together RS-B once chased S-BN and occasionally
supplanted BS-dB.
Feed site I was visited not only by territorial birds
from adjacent territories but also by ten other birds that
frequented this general area and apparently had no terri
torial affiliations. By plumage characters eight were
first-year birds and two were adults.
39

40
I expected to find a definite dominance hierarchy
among the Scrub Jays at feed site 1, but did not. The
only bird to exert his dominance frequently was RS-R,
whose territorial boundary and nest was the closest.
The other breeding males, NS-N from territory 6 and
RS-0 from territory 2, occasionally supplanted the non
breeding birds but most frequently coexisted, often side
by side with both non-territorial jays and with most of
the territorial birds visiting this site. Although many
birds often frequented the site at the same general time
of day, usually only two or three dropped down for food
at once. As they flew off a few more dashed in. There
was little opportunity, then, for much conflict, or
supplanting. Many hours were spent watching the move
ments of jays at this site, and no definite hierarchical
ordering could be assessed except that breeding males
were generally dominant over most other jays.
In many bird species, the more closely two individuals
approach, the more likely displays or attacks become. Each
individual has a specific individual distance within which
the bird will not tolerate others, excepting sexual encoun
ters. Among the Florida Scrub Jays at Kicoria this trait
was not seen. Birds frequently fed side by side or spent
considerable time close to each other.

REACTION TO OTHER SPECIES
Although territorial Florida Scrub Jays react vigor
ously toward intruders of their own species, they usually
ignore other bird species. The Brown Thrasher (Toxostorna
rufum) is similar in size and ecological requirements.
Pairs are scattered over the Hicoria area and they fre
quently feed in the same places as Scrub Jays, yet con
flicts rarely occur. Only when a thrasher or Mockingbird
(Mimus polyglottos) perches especially close to an active
nest dees the male or female jay displace the intruder.
Elsewhere in the territory other species are tolerated.
Reaction to Blue Jays
No Blue Jays nest in the Hicoria area, yet numbers
are alv/ays present, either feeding or travelling through,
often in groups of four and five. Blue Jays frequently
feed with Scrub Jays. Usually the Blue Jays.remain in
the background and only after the Scrub Jays leave the
feeding site do the Blue Jays dash in and grab some food.
Frequently at feeding sites Scrub Jays and Blue Jays
come close together and invariably the Scrub Jays dominate
the Blue Jays. Such encounters rarely include an attack
or chase; merely a peck or jab toward the Blue Jay is
sufficient to cause it to back off and fly.
41

42
Blue Jays often forage through Scrub Jay territories.
Only when the Blue Jay approaches the Scrub Jay or blunders
near a nest does the Scrub Jay react. In such cases the
offending Blue Jay is often chased. As with intraspecific
chases, the other territorial members quickly join in and
follow to the edge of the territory. Occasionally the
breeding feraele Scrub Jay begins such a chase, but the
"helpers" join, and the male usually takes over and com
pletes the chase to the border. These interspecific
encounters are infrequent, for the Blue Jays normally stay
away from Scrub Jays in their territories.
Scrub Jays recognize something in the Blue Jay as
necessitating an aggressive reaction. Yet the Blue Jay
is not treated as a distinct threat, for it is frequently
left alone in Scrub Jay territories and at feeding sites.
Reaction to
Predators
Although the male or any other territorial member in
a sentinel position frequently warns the others of predators,
during 1SS8 and 1969 five of 20 nests with young were des
troyed. Probably the common predator of young Scrub Jays
are snakes. At Hicoria I frequently found Corn Snakes
(Elaphe guttata}, Yellow Rat Snakes (Elaohe obsoleta), Pine
Snakes (Pituophis melar.oleucns) Kingsnakes (LampropeItis
getulus), Black Racers (Coluber constrictor), and especially
the Eastern Coacnwhip (Masticoohis flagellum), The Ccachwhip
as especia.
coursing through the
ms or scrim
thickets.

43
On 22 April 19-39 I banded the nestlings in territory
9 and replaced them in che nest. The next day the young
were still in the nest, but on 24 April the nest was empty
and two adults were extremely agitated. Dead on the paved
road 200 feet from the nest was a six-foot Coachwhip that
had been hit by a car. Upon opening the stomach, I found
the two young from territory 9 still wearing my bands.
Once out of the nest the young are still highly vul
nerable for at least a week and many disappear. Feral cats
are frequent at Hicoria and prey heavily on fledglings.
Gray Foxes (UrOcyon cinereoargenteus) and Bobcats (Felis
rufus) also hunt through this area.
Whenever a predator is noticed, the Jays in the ter
ritory mob about it, using all alarm and excitement behavio
patterns. Some give loud upwardly inflected kwerp calls
and bob vigorously. Females frequently give loud "hiccup"
calls and bob. A jay often dips its head toward the
predator with its body hanging straight down. Some, expe-
daily the breeding pair, hop about the ground, calling
.loudly and nipping the tail of the attacker.
The output of noise easily carries over many other
territories, but other Scrub Jays normally do not cross
into the territory where the trouble occurs. Birds in
ocher territories are aware of the attack for they become
extremely excited. Only when the predator is found close
to the territorial boundary' or when the encounter continues
for an especially long time, do other territorial birds

44
enter and join in the melee. As the new jays approach,
the local male often dashes suddenly at one of them. These
are short chases, and the male quickly returns to the
predator. These short quick attacks at a foreign jay
rarely continue to the territorial border. During non
breeding seasons when mated pairs are less insistent about
territorial defense, alarm calls more frequently attract
other Scrub Jays.
These observations concerning the attraction of other
Scrub Jays to distressed breeding birds are consistent with
Hardy's (1961) comments concerning degrees of territoriality
among species of jays. The Mexican Jay with no territori
ality and Blue Jay with a loose territoriality both give
loud assembly calls that readily attract others, whereas
the highly territorial western Scrub Jays do not. Even
though possessing a degree of sociality above that of
western races, the Florida Scrub Jay is highly territorial
and does not have a distinct assembly call. Its alarm
calls carry far, but as with western races, outside birds
frequently do not assemble when a pair is in distress.
Nearly every other resident species at Hicoria does
assemble. The alarm calls of the Scrub Jay quickly attract
Brown Thrashers, Blue Jays, Mockingbirds, Towhees (Pipilo
erythrophthalmus), White-eyed Vireos (Vireo griseus), and
especially woodpeckers. Red-bellied Woodpeckers (Centurus
carolinus), Red-headed Woodpeckers (Melanerpes ervthrcce-
phalus), and Flickers (Celantes auratus? are easily excited

45
by the loud upwardly inflected kwerp calls of the Scrub
Jay and remain with the distressed pair as long as they
continue calling. These other species probably do little
to aid the distressed Scrub Jays, for they remain well
away from the action. Possibly Scrub Jays aid other nest
ing species by predator recognition.
Reaction to Man
One feature which makes the Florida Scrub Jay especially
intriguing to work with is its extreme tameness toward man.
On 17 June 1967 when I first started banding at Hicoria
I walked down the dirt road by territory 3. A Scrub Jay
landed ahead of me and a juvenile near my feet. I was
surprised at their seeming unawareness of my presence.
Then the juvenile hopped between my feet, and the adult
landed on my head. I held out my hand and the adult jumped
to my arm and then to a nearby branch. I fumbled in my
pockets as though to bring out some food and again held my
hand out, but they did not fly to it. I walked up to the
adult, which remained calm until I was three-feet way,
and then it flew 10 feet down the trail.
This encounter surprised me for, to the best of my
knowledge, no one has spent any time at Hicoria feeding
Scrub Jays. In fact they did not appear to be attracted
to me for food, merely inquisitive about my presence. At
the Archbold Station Scrub Jays readily approach a person
and fly to the hand. These birds have been conditioned

46
to expect a handout of food, yet they have become so tame
that one can frequently pick up a brooding female from
the nest, something that could never be done at Hicoria.
An interesting example of this tameness occurs at a
small scrub thicket north of Englewood, Sarasota County.
For several years people have been feeding Scrub Jays there.
As one stops the car several jays fly out of the thicket
and often land on the car or an outstretched hand. If no
birds are in sight, all one has to do is sound the car
horn and the jays quickly appear.
It is difficult to explain this tameness. Although
most jay species are easily enticed to feeding stations,
only the Canada Jay (Perisoreus canadensis) is as overtly
tame as the Florida Scrub Jay (Bent, 1946).

DISPERSAL
The striking feature cf the movement of Scrub Jays
about the Hicoria area is that there is so little of it.
Apparently, once a pair establishes a breeding territory,
it remains as long as conditions are suitable. Of nine
pairs nesting at Hicoria in 1968 six were still there
nesting in 1970. One of these was a female WS-Pi banded as
a breeding adult in 1960 within 200 yards of the territory
she now occupies.
Dispersal of certain members does occur, for in late
summer and fall the first-year "helpers" begin to disappear
from their parental territories. During this period Scrub
Jays are prone to make long flights. A single bird will
fly as much as 0.3 mile from the territory. Such long
flights are usually silent, unaccompanied and at such a
height (100 to 200 feet) that other Scrub Jays ignore them.
Normally these long-fliers return later the same day. All
birds that made long flights and that I could identify were
"helpers"; I never saw a breeding adult make such flights.
This is not a phenomenon restricted to a certain time
of year. Even during the breeding season, occasional long
flights occur. They are much more frequent, however, in
the fall. Perhaps it is an expression of unrest by particu
lar birds, a prelude to a final move. 3y late fall most
47

4S
old "helpers" have dispersed, leaving the new set of young
to take on the role of "helpers."
Dispersal is apparently delayed until the second year.
I have not found any first-year birds far from their parental
territories. As reported above, in 1963 several first-year
birds were members of a loose nonbreeding flock. As they
had just been banded I know nothing of their origin. In
1969 a group of non-territorial individuals roamed about
the large open area west of territories 7, 3 and 5. A few
were members of the nonbreeding group of 1968 and others
were unbanded. Two, P-OS and P-dBS, were first-year birds
and were banded as juveniles in territory 6 during 1968.
P-OS and P-dBS had moved approximately 500 yards from their
parental territory. These were the only birds banded as
juveniles and found away from the parental territory during
their next year. The breeding pair of territory 6 also
deserted the territory, and this event undoubtedly influenced
the movement of their young.
The remaining birds banded as juveniles and not present
as "helpers" probably succumbed to predation. I searched
as far as three miles from Hicoria for banded birds, but
no first-year individuals were found. It was difficult to
check all the many small scrubby thickets to the west and
south; some birds from Hicoria might have been overlooked.
Two new territories were established at Hicoria during
1969 (7 and 10) and one during 1970 (11). Territory 10 was
settled by unmarked bi.x'ds, and. I have no knowledge of their

49
origins. Territory 7 was settled by Pis-O and S-GB
who were first-year birds in this same general area during
1963. Their parental territories were also unknown. Terri
tory 11 was established by S-BY and M-dBS who were both
"helpers" during 1969 in territory 4. As these two areas
were adjacent, this new pair was filling in unoccupied habi
tat at Hicoria and only a minimal dispersal occurred.
Three first-year "helpers" moved completely away from
the Hicoria region during their second year. S-RY moved
1.2 miles due south from territory 6 and nested in 1969,
O-SG moved 0.9 mile westnorthwest from territory 8 and
nested in 1970, and O-SW moved 0.4 mile westsouthwest
from territory 8 and nested in 1970. These last two
settled in scattered scrubby thickets in the wet flat-
woods region, while S-RY remained on the sand ridge but
well to the south of Hicoria.
One other bird banded at Hicoria moved to a new locale
SO-B was banded as an adult during the summer of 1967 at a
banding site just south of territory 6. Several birds that
were banded there subsequently disappeared. SO-B was found
1.8 miles to the northwest in February 1969 and nested
during that year on a narrow band of scrub. This strip of
scrub had been censused during 1968, where SO-B was not
present; its movements during 1968 are unknown. This was
the farthest away from the banding site that I located any
of my banded birds.

50
During the breeding seasons of 1969 and 1970
Glen Woolfenden color banded the local Scrub Jays at the
Archbold Biological Station and censused for other birds.
We attempted to use non-overlapping band series so that
birds from t.hs Station and Hicoria could be. distinguished.
The Station is 2.0 miles due north of Hicoria, and to date
no birds from Hicoria have been found at the Archbold
Station and no Station birds have appeared in Hicoria.
As documented at Hicoria, the movement of Scrub Jays
from hatching sites is delayed for an entire year by the
retention of offspring as "helpers.,! Breeding males may
induce this retention by continuing to feed juveniles
throughout the year and so form with the offspring a loose
bond, similar to a pair bond. When birds do disperse it
is apparently for short distances. Yet in certain cases
family groups are found far from scrubby habitat. In one
case a fire destroyed a large section of scrub in late
summer, and a group of five Scrub Jays was found two days
later approximately three miles from the fire, feeding at
the edge of a large watermelon field, with no scrubby
habitat about. The next day the birds had disappeared.
Four miles northeast of Hicoria the scrub on several
thousand acres was destroyed during the summer of 1959 and
orange trees planted. For a week thereafter I found small
groups of Scrub Jays in situations where I never saw them
before and where they were not seen subsequentlv. As sug
gested for the movement of brsedincr oairs at Hicoria,
long

51
distance
rare and
such as
movements by Florida Scrub Jays apparently are
normally must be induced by some major catastrophe
destruction of the habitat by fire or clearing.

HABITAT AND DISTRIBUTION
Florida Scrub Jays select a nesting habitat of very
limited range and at times are absent even though the
proper breeding habitat is available.
Scrub Jav Habitat in Florida
Nesting Scrub Jays are invariably located in types
of sand pine scrub vegetation. The vegetational components
of sand pine scrub are primarily xeric plants that are
restricted to the veil washed and sorted siliceous sands
of modern and ancient shore lines. During the late Tertiary
and the interglacial portions of the Pleistocene the drastic
rises in sea level left four distinct ridges of deep sand
ranging in elevation from 150 feet to S feet (MacKeii, 1950)
The present day sand pine scrub vegetation is located on
these stretches cf sands of the Saint Lucie and Lakewood
types of soils, deposited by marine currents, wave action,-
and winds (Laessie, 1958). Since these sorted sands were
laid down as shore lines or sand bars, today sand pine
scrub communities are found primarily as long narrow
strips of vegetation. Two major exceptions are the Ocala
National Forest in Zlarion County and the extensive
O L. Ulv
communities cf central Highlands County. Yet Laessie (1953)
concludes that these, too, are of dune origin, and the
extreme width is the result of winds blowing dune series
inland and new dunes forming at the sea front.
sea

53
Most frequently the dominant plant is sand pine (Pints
clausa), which often forms a compact forest. In the less
dense stands of sand pine the understory is a low thicket
of various woody plants, and the ground is often open white
sand, perhaps with a thin layer of Cladonla spp. Seme
herbaceous flowering plants occur, and occasionally a
few grasses (Andropcgon f loridanus and ? an i cura patent!-
folium) nav occur.
\viren present as an understory to sand pines, the
woody plants may be well spaced, but when sand pines are
lacking, these shrubs frequently make an extensive, dense
scrub thicket, in form strikingly similar to the chaparral
communities of Arizona and California. The dominant plants
of these low scrub thickets are three species of xeromorphic
oaks (Quereus virginiana martima, Q. myrtifolia, and Q._
chapmanii); these three frequently compose over 50 per cent
of the ground cover in the thickets. The remaining planus
may consist of a variety of species, including Serenoa
repens, Sabai etonia, Ilex cumulicola, Rhus coralline,
Lyonia ferrv.gir.ea, Myrica cerfera, and occasionally
Ceratiola ericoides. As Laessle (1958) reports, Florida
sand pine scrub normally includes many other shrubs and
herbaceous species, but the list above includes the woody
species with which Scrub Jays are frequently associated.
Although the ground is often bare of grass, the woody plants
form dense thickets with few oren areas.

In areas where
no Scrub Jays occur
Scrub Jay always do
together, I doubr i
sand pine is dense, Blue Jays nest and
as breeding birds. As the Florida
minabas the Blue Jay when the two are
f the Blue Jays have displaced the
Scrub Jays from the mature sand pine scrub. It is more
probable that the Florida Scrub Jay finds this habitat
unsuitable for nesting.
Sand pine trees may be absent from the scrub community
for several reasons. Sand pines are especially susceptible
to fire and once flames reach the canopy, large regions
quickly burn. But Webber (1935) states that mature sand
pine forest may act as a fire inhibiting community because
the ground is often bare of grasses and fire has difficulty
in spreading. Thus many scrub communities may remain
covered with pine forest and are unavailable to Scrub Jays.
Today the State Forest Service fire protection program is
aiding the preservation of sand pines.
Frequently the washed and sorted sands are thin and
do not support typical scrub communities. At such sites
understory woody planes of che scrub community are mixed
with species normally present in pine flatwoods of nearby
areas.
The Florida Scrub Jay selects a specific pattern of
growth in these xeric communities and rejects as unsuit
able much or ens true sand pme
special habitat requirements of
detailed examination of several
scrub. !o illustrate
the Florida Scrub Jay
different sices sueno
the
a

55
Scrub Jays is presented
demonstrate the variety
below. These sites were chosen to
of plant communities that Florida
Scrub Jays select for nesting.
Hicoria, Highlands Countv
Elevation of Hicoria is 140 feet. As described earlier,
this site has artificially open areas, and after effect of
a now deserted and razed town (Fig. 4); at least 60 per
cent of this 200 acre study area is composed of open sandy
or grassy patches. The scrub thickets consist primarily of
the three scrub oaks (80 per cent of the ground cover in
thickets) with a scattering of the other plants, principally
Serenoa repsns, Sabai etonia, Hicoria floridana, and Myrica
cerfera. The thickets average 10 feet in height and al
though each is small the woody plants are dense. Scrub
Jay nests occur predominantly in oaks, almost always in a
dense thicket but adjacent to an open space.
The degree of grass cover (Aristida stricta) and the
presence of scattered trees of Quercus laevis and Pinus
elliottii indicate that this site is a combination of plants
from several communities, i.e., scrubby flatwoods. From
the number of Scrub Jays breeding at Hicoria it obviously
contains optimal features that Scrub Jays select for breed
ing areas. The population density was 10 nests per 100
3.C2TSS

Figure 4 Viev; of Hicoria study area, center of territory
6.


53
Cape Kennedy Air Force Station, Brevard County
Elevation is 6 feet. This large area of approximately
1200 acres of scrub sustains a large population of Scrub
Jays. This community is on recent shore front sands and
is unlike most interior scrubs because wax myrtle (30 per
cent of ground cover in thickets) and saw palmetto (25 per
cent of ground cover in thickets) are much commoner. Here
the three species of oaks are prevalent (35 per cent
of thicket ground cover). The thickets are 8-12 feet
high and dense, especially near the shore line where
onshore winds and salt spray have stunted the upward
growth of plants and produced an extremely tangled, low
thicket.
The outstanding feature of this site is a network of
interconnecting dirt and paved roads crisscrossing the
area. All these roads have cleared shoulders 30-50 feet
wide, often of open white sand. Of course, there are
large open spaces with rocket launching complexes, but in
addition to these industrialized parts, open sandy places
comprise about 40 per cent of this study area. During
1969 I censused two plots of scrub thickets in the center
of the Cape where there were many open spaces but away
from the rocketry complexes; at each I found 10-11 active
nests per 100 acres. This cape region with many artificial
open, spaces resembles Hicoria and the density cf breedinc
birds is similar. Most nests are located in scrubby oaks
although several were found in the wax myrtle.

59
Jonathan Dickinson State Park, Martin County
Elevation is 12 feet. This park preserves a large
stretch of sand pine scrub. Although sand pines cover
much of this region, there are extensive scrub thickets
without sand pine cover, and Scrub Jays nest in much of
this area. Myrtle oaks dominate this habitat comprising
30 per cent of ground cover in thickets although the other
two oak species are present and make 10 per cent of the
thicket ground cover.
The low scrub thickets without sand pines are rather
dense and form large stretches of habitat with few open
areas except along roadways. A census in this extensive
cover showed six nests per 100 acres. But in developed
areas of the park where camping facilities are scattered
through the low dense scrub, I found 9-10 nests per 100
acres.
Mt. Plymouth Area,, Orange County
Elevation is 130 feet. Just west of this small town
is an extensive region of low scrub thickets without any
sand pines. The woody plants here are relatively low,
4-7 feet high. Numerous evergreen woody soecies occur,
although again, the oaks predominate. This is one of the
few areas of scrub thickets where numerous open spaces
occur interspersed among the denser thickets and apparently
not caused by some form of human interference. Approximately

60
4G per cent of the habitat is open sandy spaces, which are
practically devoid of any herbaceous cover. A nesting
census located 910 nests per 100 acres and all were found
in oak trees.
Lake Jackson, Jeering, Highlands County
Elevation is 110 feet. Undoubtedly a fire destroyed
an extensive sand pine forest at this tract on the west
shore of the lake at least ten years ago to judge by scars
on a few large sand pines. Today a young scattered stand
of sand pine is present, but most of the region has dense
thickets of woody plants 6-10 feet high. As usual, oaks
dominate, but Sabal etonia and Ilex cumulicola are common.
There are few natural open sandy spaces, but a new housing
development has a series of parallel roads through this
scrub, and an unaltered strip of thicket 100-150 feet wide
between roads has been left. The developer, thus, has
produced a series of open spaces, often of ^^7hite sand,
throughout the dense scrub thicket. No nesting census was
made, but the density of Scrub Jays was always high and
reminded me of the densities of jays at Cape Kennedy and
Eicoria.
Ridge Scrub, Highlands County
Elevation is 150 feet. Wide expanses of very low dense
scrub exists along U.S. Highway 27, two miles east of Hicoria.
In recent years orange groves have usurped the majority of

61
this terrain. Judging by large sand pines scattered along
the highway much of -this area may have been covered with
extensive sand pine forest. Scrub thickens extend several
miles in places as a solid dense ground cover 3-6 feet high.
The three scrub oak species dominate (60 per cent of ground
cover in thicket) and Sabal etonia is common (20 per cent
of ground cover in thicket)(Fig. 5). Although grasses are
infrequent and the surface is usually bare white sand, very
few large open spaces appear. It is a difficult area to
census for nesting Scrub Jays, because cf the impenetrable
nature of the low woody plants. Nesting jays occur over
much of this terrain but congregate along read edges or
about the few open sandy spaces. During 1968 and 1969
two areas vrere censused. Approximately 3-4 pairs nest per
100 acres, but in the densest areas away from open spaces
jays rarely nested.
Scrubby Flatwoods West of Hicoria, Highlands County
Elevation is 130 feet. Just west of the scrub covered
hills of Highlands County an extensive region of flatwoods
exists. Within three miles cf the ridge, numerous small
narrow strips of white sand are scattered. Probably these
were small sand bars in a Pleistocene sea. The thickets
on these strips are extremely dense. The edges of these
small islands of thickets are usually bare sand or occa
sionally widely spaced scrubs of rosemary.
Nesting Scrub Jays occur in the scattered thickets,
especially '.'hen close to the extensive scrub ridge at the

Figure 5
- View of low, dense scrub thicket two miles
southeast of Hicoria, Highlands County.
The only open spaces are along the dirt
road. In the background is a patch of sand
pine without nesting scrub jays.

63

64
Archbold Station and Hicoria. Although large numbers of
breeding Scrub Jays are supported in these scattered islands
of scrub thickets, the density per unit area is rather low
(2-3 pairs per 100 acres).
Atlantic Coast Dune Thickets
An extremely long yet narrow band of very low and
dense, thickets exists on shore-front sand dunes of the
Florida Atlantic coast from Jacksonville to Fort Pierce.
Sand pines are not present and saw palmetto often is the
dominant plant. Live and myrtle oaks comprise from 20-40
per cent of ground cover in thickets. Onshore winds and
salt spray have led to an extremely dense, often impene
trable, growth (Fig. 6). Open areas exist only along
roadways or where houses have been built.
The vegetation is similar all along the coast but
the distribution of Scrub Jays varies. The snore thickets
about. Melbourne Beach, Brevard County, 20 miles south of
the iai'ge Cape Kennedy scrub area contain many scattered
breeding colonies. The beach thicket north of Cape Kennedy
toward New Smyrna Beach, Volusia County, also contains
numerous breeding Scrub Jays. These birds nest only along
the edges of the danse thickets, normally along the highway.
North of Daytona Beach, Volusia County, in the same
type of vegetation, Scrub Jays are much less abundant.
Several miles of seemingly suitable thickets exist before
another nest is located.

Figure 6 View of low and extremely dense Atlantic coast
thickets at Marineland, Flagler County. Salt
spray has retarded the upward growth of the
plants producing an extremely tangled thicket.
The only open spaces are along the road edge
at the extreme foreground of the picture.

66

67
"3iq Scrub, Marion C:~:ntv
Elevation is 140 feet. This is an expanse cf scrub
1.7 miles wide and 12 miles long, located 12 miles west of
Belleview. Dense stands of sand pine cover much of this
area with no breeding Scrub Jays. Yet large stretches of
low dense thickets without sand pines are present. As
usual the ground is usually open sand with few hems,- but
large open spaces are lacking. The three oak species
occupy 30-40 per cent of the ground cover in thickets, and
Ilex cunulicola and Lyonia ferrnginea are common (30 per
cent of ground cover in thicket). This vegetation varies
from 8-15 feet in height.
The only part that contains breeding Scrub Jays is
the narrow zone along State Road 484. Here jays are
frequently seen feeding along the open roadway, but one-
half mile into the scrub from the road no Scrub Jays are
found. Eight miles north of this road, Interstate 75
crosses a section of scrub. Parts of the scrub bordering
Interstate 75 are similar to that along State Road 434,
yet no Scrub Jays nest here. Interstate 75 is a relatively
new road, and apparently Scrub Jays have not located this
newly opened habitat.
Ocala National forest, Marion County
Elevation is 130 feet. Although this is the largest
sand pine scrub in Florida, few parts of chis extensive
regc.cn suppers nesting Scrub Jays. Much of this area is

68
covered with relatively dense stands of sand pine with
associated Slue Jays. Bat many low dense scrub thickets
exist, and as at the "Big Scrub" to the west, most of
these have few open spaces and few nesting Scrub Jays.
Breeding birds congregate along roadways, whose edges
are frequently used as feeding sites.
Only at the southern edge of Ocala National Forest
are Scrub Jays numerous and here the scrub thickets away
from the roadsides are often interspersed with open spaces
and sand pine are not abundant. Here some mingling of
scrub and turkey oak community occurs, and apparently the
resulting ecotone frequently contains open spaces. Even
here most breeding birds nest along roadways.
Habitat Summary
The Florida Scrub Jay requires low, dense, largely
evergreen oak thickets for nesting and extensive open
spaces for feeding. At the Archbold Station, Amadcn (1944)
found Scrub Jays concentrated along the edges of man-made
clearings. This tendency is especially obvious throughout
its breeding range in Florida. The largest concentrations
of nesting birds are found in scrubby thickets that have
been opened up by man, such as Cape Kennedy and Hicoria.
Although nesting Scrub Jays almost always select xeric
habitats,
most Florida
sand pine scrub
is unsuitable because
the pines
are in dense
stands. The FI
orida Scrub Jay's
specific
habitat prefe
rencas for dense
thickets inierspersed

69
with open sandy spaces restrict them to certain border
zones such as scrubby flacwoods, to recently burned regions,
and especially now to roadways and other disrupted scrub
areas produced by human interference.
Usually wherever the proper combination of dense
evergreen oak thickets and open sandy spaces exists,
some nesting Scrub Jays are found. Thus small populations
occur at Cedar Key, Levy County; at Camp Blanding, Clay
County; and scattered along the Atlantic coast often
miles from other breeding Scrub Jays. Although such
habitat is scattered over much of central Florida, it is
normally in small acreages of open thickets. The distri
bucin of Scrub Jay breeding habitat indicated in Fig. 7
illustrates the restricted amount of land available to
nesting Florida Scrub Jays. This map also shows certain
areas that appear to have suitable Scrub Jay habitat but
lack resident populations. Some of these areas are dis
cussed below.
Scrub Habitat Hitbout Breeding Scrub Jays
A few rather sizeable thickets with apparently suit
able habitat have no resident Scrub Jays. Several of these
scrub chicket areas are indicated in Fig. 7.
Kissimmee-Saint Cloud, Osceola County
Msnv sczrubfov thi ckGins exist
although most cf the sand ridge was on
tat ion anc. is now cultivated in orange
about this region
e turkey oak vege-
groves. The small

Figure 7
- Map of central region of Florida peninsula
illustrating areas apparently suitable for
breeding Scrub Jays and areas where breed
ing Scrub Jays were located. J.D.S.P. is
the Jonathan Dickinson State Park.

71

72
scrubby thickets, often merely 200-300 acres, appear to
be excellent Scrub Jay cover and resemble the scrub at
Mt. Plymouth. No Scrub Jays exist in this region, but 20
miles to the north similar habitat supports numerous scat
tered populations of Scrub Jays.
Weeki Nachae, Hernando County
An extensive scrub exists about this natural spring.
Much of if is covered with sand pines and so could not
be expected to support Scrub Jays. Small sections have
recently burned, however, and low dense thickets are grow
ing back, with numerous open sandy spaces. Moreover, along
the southern border of this scrub much openness exists in
a broad ecotcne between the scrub and turkey oak vegetation.
I estimate approximately 1000 acres of low open scrub thickets
exist here, but no Scrub Jays are present. Scrub Jays reside
in small numbers 20 miles to the south and 28 miles to the
east.
Lake Nales, Polk County
A few islands of scrub thickets, similar to the ones
west of Hiccria, are present to the west of Lake Wales,
but no Scrub Jays reside on them. The plant community on
the sand ridge close to these scrub islands at Lake Wales
is turkey oak vegetation without any Scrub Jay population.

73
Florida Panhandle Scrub
A very long stretch of sand pine scrub exists along
the Gulf Coast frcm St. Teresa, Frankland County, westward
into Alabama. Much of this scrub supports sand pine forest,
but many extensive acreages of low dense scrub oak thickets
occur, with numerous open sandy stretches. An especially
large expanse of this type of habitat exists on Santa Rosa
Island south of Pensacola, Santa Rosa County, and especially
at Fort Pickens State Park (Fig. 8), but nowhere in these
several hundred miles of scrub habitat are there any Scrub
Jays. The nearest Scrub Jay population is near Cedar Key,
Levy County, 100 miles to the southeast of the eastern bor
der of the Panhandle Scrub.

Figure 8 View of low scrub thicket with open sandy
spaces at Fort Pickens State Park, Santa
Rosa Island, Escambia County. This type
of scrub thicket supports large numbers of
Scrub Jays when present in the Florida
Peninsula, but here no jays occur.

75

FACTORS CONTROLLING DISTRIBUTION
The Florida Scrub Jay territorial proclivities are as
intense as those described by Nice (1937) for the Sor.g
Sparrow (Melospiza Ilslodia) These jays defend a territory
that normally includes mating, nesting, and feeding grounds,
and thus belongs in territory category "A" of Nice (1941).
The Florida Scrub Jay forms a pair bond that is retained
throughout the year, and frequently a pair defends the
same territory for more than three years.
This pattern of territoriality is similar to that
described for most western races of A. coerulescens by
Hardy (1961) and Westcott (in press). But the Florida
race exhibits a degree of sociality not found among its
western relatives, as first-year birds are retained as
"helpers" in the territory. Discussing the social struc
ture of the Arizona populations of the Mexican Jay in
which territoriality is lacking and flocks are tolerated
at the nest, Hardy (1961) suggests that a species that has a
restricted amount of habitat available for nesting may
adapt by allowing a social breeding structure. Thus
the Mexican Jay, which nests mainly in the narrow rioarian
woodland of southeastern Arizona, tele
breeding birds about the nest, but the
of the Scrub Jay, which have a much wi
available, can utilize strict territor
rates flocks of non
western subspecies
der range of habitats
iality.
76

77
Contrary to Hardy's (1961) statement, the Florida
Scrub Jay is territorial, as much so as any of the western
races. Yet in Florida A. c coerulescens inhabits as nar
row an ecologic zone as A. ultramarina in Arizona. The
habitat selected by these two have no features in common.
A. ultramarina selects the arborescent parts of the wood
land that are restricted to narrow riparian situations,
while the Florida Scrub Jay typically selects the xeric,
low scrub thickets with open sandy spaces and ignores the
tall sand pine forest and ail the mesic forests of Florida.
Even though breeding Scrub Jays are scattered through
out much of the Florida peninsula (Fig. 7), close examination
reveals that the amount of acceptable nesting habitat is
restricted, often merely a border of open scrub thickets
along roadways, the edges of the taller sand pine forest,
or recently burned regions. Only limited areas consist of
extensive low thickets and here it is usually man-made
interference that has produced the open spaces that attract
dense breeding populations. Suitable habitat to induce
Scrub Jays to nest is restricted both in type of cover and
in total acreage of this special vegetation. The tolerance
shown by breeding pairs of Florida Scrub Jays to certain
members of the population conceivably is a reaction to a
restricted range of suitable habitat, paralleling the situa
tion of the Mexican Jay in Arizona.
As suggested by Hardy (1961) for the Mexican Jay, a
minimum population size is probably necessary for survival,

7c
and species with limited breeding range may be unable to
maintain such a population level unless special social
structures are included. With the retention of first-year
birds as territorial "helpers," the Florida Scrub Jay has
developed a degree of sociality as an adjunct to an essen
tially territorial nature.
This combination of sociality and territoriality is
possible, for the breeding male may courtship-feed the
"helpers" and so form a loose bond with these birds. All
members of the territory frequently react together in
territorial defense. Thus the family group remains terri
torial, unlike Mexican Jay flocks that show no territoriality.
Interestingly, compared with Scrub-Jays, the breeding male
Mexican Jay utilizes courtship feeding infrequently and
only with his mate (Brown, 1963; Hardy, 1961).
In Mexico and western United States many different
habitats are selected by breeding Scrub Jays, but most are
of a xeric nature and, as Pitelka (1951) points out, the
Scrub Jay appears bast adapted to feeding in and about the
low dense thickets of chaparral or open oak woods. Even
though isolated by over 1000 miles from these western races,
the Florida subspecies continues to utilize habitats of
similar ecologic structure. Presumably at one time a con
tinuous scrub habitat connected the Florida peninsula with
western habitats and Scrub Jays invaded Florida through this
connecting habitat (Pitelka, 1951).
Whether or not large expanses of suitable habitat
ever existed an Florida is difficult to ascertain, but

79
Laessle (1953) states that sand pine scrub areas have
probably existed as presently situated since the forma
tion of the terrace ridges. Because this xeric community
exists only on well-washed dune sands, and Florida Scrub
Jays require low thickets with open spaces that occur
mainly at scrub borders, I doubt if Scrub Jay habitat was
ever widespread in Florida.
Hardy (1961) speculates that A^_ cu_ cosrulescens has
survived in Florida as a relict population that may have
preserved certain traits indicative of a primitive con
dition, i.e., sociality and supposed lack of territoriality
Evidences from the present study suggest the opposite, i.e.
territoriality as a basic and possibly primitive trait
and social nesting as behavioral adaptation to the special
restrictive characters of the xeric scrub thickets in
Florida.
Although western Scrub Jays, especially first-year
birds, frequently travel great distances away from the
nesting region (Westcott, 1969), the Florida Scrub Jays
are sedentary. As first-year birds normally remain in
the parental territories, dispersal of juveniles is infre
quent. At Kicoria movement of local birds to new breeding
areas was restricted. Some settled to nest close to
parental territories in unoccupied habitat, and a few
individuals, usually adults, roamed as far as 1.5 miles
before settling. But few long-range movements have been
observed. Amador. (1944) especially commented on the

so
sedentary habits of Florida Scrub Jays. Only following
some major catastrophe, as a fire or bulldozing of large
areas cf scrub, were Florida Scrub Jays found far from.
scrub habitat.
Only a few areas occur in Florida with broad expanses
of open scrubby cover and support relatively large and
dense populations of Scrub Jays such as the Lake Placid
areas (Archbold Station, Eicoria, etc.), Jonathan Dickinson
State Park, Cape Kennedy, and the Mt. Plymouth region
(Fig. 7). Not only do Scrub Jays abound in these areas,
but nearly all similar habitats nearby have resident Scrub
Jays. Presumably enough extra birds inhabit these popula
tion centers so that most small scrub thickets are located
and colonized by the jays. Similar small sites further
away (10-20 miles) have only a few scattered pairs of
birds or none at all. The density of nesting Scrub Jays
gradually decreases along the coastal thickets north from
Cape Kennedy. North of Daytona Beach nesting Scrub Jays
are noticeably less frequent, and from Marineland to
St. Augustine nesting pairs are often separated by several
miles of low dense thickets even though the habitat is
similar to the densely inhabited shore thickets just north
of Cape Kennedy.
Similar patterns of density of this species occur
about the other large centers of its population. Close
to densely inhabited thickets most small scrub habitats
have nesting birds, but farther awav fewer birds are

81
found.. Finally soma sites, usually well isolated, have
no Scrub Jays. As an Waeki Washes these scrubs without
jays often contain stretches that appear to meet the
Florida Scrub Jay:s special ecological requirements.
The lack of dispersal of juveniles and the generally
short distance of dispersal by nonbreeding adults must
explain the absence of nesting Scrub Jays from suitable
habitan. A social structure allowing for a degree of
sociality has reduced dispersal cf juvenile birds which
account for most of the dispersing individuals in western
races (Westcott, 1959). As "helpers" they remain in the
parental territory for extended periods, which may permit
an increased familiarity of that specific site cr the
surrounding habitat. Thus a form of site tenacity may
be present in the younger nonbreeding birds and may reduce
the distance of dispersal when they do move out of the
parental territory.
Long distance movements must occur, as some sites
with nesting jays are isolated by many miles from the
nearest Scrub Jay population. Undoubtedly, a few indi
viduals disperse long distances, but frequently when
Scrub Jays are found in unlikely habitaos seme major
catastrophe has occurred to a nearby scrub, forcing the
birds to move. Probably many distantly isolated sites
are first located bv such birds. Perhaps such catastrophes
explain the absence of Scrub Jays from isolated yet suit
able vegetation. A fire may have destroyed the low scrub

82
forcing resident bi
have located the ne
The extensive
nearest small Scrub
occupy all the suit
they should take r.ia
rds away; and wandering birds may not
wly available habitat after it regrows,
panhandle scrub is 100 miles from the
Jay colony. If straying birds cannot
able sites in central Florida, then
ny years longer to invade the pan
handle region.
The adaptation of a type of sociality by the Florida
Scrub Jay has produced certain restraints on the distribu
tion of these birds. Because they have become exceptionally
sedentary, and dispersal abilities have been reduced, some
areas of suitable habitat in Central Florida and a large
expanse of habitat in the Panhandle are unoccupied today.

LITERATURE CITED
Amadon, D.
1944. A preliminary life history study of the Florida
Jay, Cvanocitta c. coerulescens. Results Arch
bold Exped. No. 50. Am. Mus. Novitates, No.
1252:1-22.
Bent, A. C.
1946. Life histories of North American jays, crows,
and titmice. Bull. U. S. Nat. Mus., No. 191.
Brown, J. I..
1963. Social organization and behavior of the Mexican
Jay. Condor, 65:126-153.
Hardy, J. W.
1961. Studies in behavior and phylogeny of certain
New World jays (Garrulinae). Univ. Kansas
Sci. Bull., 42 (2):13-149.
Hinde, R. A.
1961. Behavior. In A. J. Marshall, ed., Biology and
Comparative Physiology of Birds. II. New York:
Academic Press.
Grimes, S. A.
1940. Scrub Jay reminiscences. Bird Lore, 42:431-436.
Laessle, A. M.
1942. The plant communities of the Welaka area. Univ.
of Florida Press. Bio.1. Sci. Series, vol. 4,
No. 1:1-143.
1958. The origin and successional relationships of
sandhill vegetation and sand pine scrub. Eccl.
Monog., 28:361-387.
MacNeil, F.
1950.
Pleistocene shore lines in Florida and Georgia.
U. S. Geol. Surv., Prof. Paper 221-F, pp. 95-107.
Nice, M. M.
19 37.
Studies in the life history of the Song Sparrow.
I. Trans. Linn. Soc. N. Y., 247pp.
941* The role: of herritCa
Nat., 26:441-487.
in bird life,
Ame
Midi,
83

84
Pitelka, F.
1945.
A.
Pterylography, molt, and age determination of
American jays of the genus Aphelocoma. Condor,
47:229-250.
1951. Speciation and ecologic distribution in American
jays of the genus Aphelocoma. Univ. California
Pubi. Sool., 50:195-484.
Skutch. A. F.
1935. Helpers at the nest. Auk,
52:257-273.
1961. Helpers among birds. Condor, 63:198-226
Webber, H. J.
1935. The Florida scrub, a fire fighting association*
Amer. Jour. Bot., 22:344-361.
Westcott, P. W.
1969. Relationships among three species of jays winter
ing in southeastern Arizona. Condor, 71:353-359.
Westcott, P.W.
In press. Territoriality of the Scrub Jay in Arizona.
Condor.

BIOGRAPHICAL SKETCH
Peter Walter Westcott was born 5 November 1938 at
Mt. Vernon, New York. In June, 1956, he was graduated
from Madison High School, Madison, New Jersey. In June,
1960, he received the degree of Bachelor of Arts with a
major in Zoology from Amherst College. In 1962 he enrolled
in the Graduate School of the University of Arizona. He
worked as a graduate teaching assistant in the Department
of Zoology and as a graduate research assistant in the
Department of Virology until June 1962, when he received
the degree of Master of Science with a major in Zoology.
From 1963 to 1966 he served in the United States Navy
and was stationed in Japan and Norfolk, Virginia. From
January, 1967, until the present time he has pursued his
work toward the degree of Doctor of Philosophy at the
University of Florida, with a major in the Department of
Zoology.
Peter Walter Westcott is married to the former
Marybeth King. He is a member of Sigma Xi, American
Ornithologists' Union, and the Cooper Ornithological
Society.
85

This dissertation was prepared under the direction of
the chairman of the candidate's supervisory committee and
has been approved by all members of that committee. It
was submitted to the Dean of the College of Arts and
Sciences and to the Graduate Council, and was approved
as partial fulfillment of the requirements for the degree
of Doctor of Philosophy.
August, 1970.
College"0
sCt£
Dean
f Arts and Sciences
Dean, Graduate School
Supervisory Committee:
Chairman



4S
old "helpers" have dispersed, leaving the new set of young
to take on the role of "helpers."
Dispersal is apparently delayed until the second year.
I have not found any first-year birds far from their parental
territories. As reported above, in 1963 several first-year
birds were members of a loose nonbreeding flock. As they
had just been banded I know nothing of their origin. In
1969 a group of non-territorial individuals roamed about
the large open area west of territories 7, 3 and 5. A few
were members of the nonbreeding group of 1968 and others
were unbanded. Two, P-OS and P-dBS, were first-year birds
and were banded as juveniles in territory 6 during 1968.
P-OS and P-dBS had moved approximately 500 yards from their
parental territory. These were the only birds banded as
juveniles and found away from the parental territory during
their next year. The breeding pair of territory 6 also
deserted the territory, and this event undoubtedly influenced
the movement of their young.
The remaining birds banded as juveniles and not present
as "helpers" probably succumbed to predation. I searched
as far as three miles from Hicoria for banded birds, but
no first-year individuals were found. It was difficult to
check all the many small scrubby thickets to the west and
south; some birds from Hicoria might have been overlooked.
Two new territories were established at Hicoria during
1969 (7 and 10) and one during 1970 (11). Territory 10 was
settled by unmarked bi.x'ds, and. I have no knowledge of their


81
found.. Finally soma sites, usually well isolated, have
no Scrub Jays. As an Waeki Washes these scrubs without
jays often contain stretches that appear to meet the
Florida Scrub Jay:s special ecological requirements.
The lack of dispersal of juveniles and the generally
short distance of dispersal by nonbreeding adults must
explain the absence of nesting Scrub Jays from suitable
habitan. A social structure allowing for a degree of
sociality has reduced dispersal cf juvenile birds which
account for most of the dispersing individuals in western
races (Westcott, 1959). As "helpers" they remain in the
parental territory for extended periods, which may permit
an increased familiarity of that specific site cr the
surrounding habitat. Thus a form of site tenacity may
be present in the younger nonbreeding birds and may reduce
the distance of dispersal when they do move out of the
parental territory.
Long distance movements must occur, as some sites
with nesting jays are isolated by many miles from the
nearest Scrub Jay population. Undoubtedly, a few indi
viduals disperse long distances, but frequently when
Scrub Jays are found in unlikely habitaos seme major
catastrophe has occurred to a nearby scrub, forcing the
birds to move. Probably many distantly isolated sites
are first located bv such birds. Perhaps such catastrophes
explain the absence of Scrub Jays from isolated yet suit
able vegetation. A fire may have destroyed the low scrub


50
During the breeding seasons of 1969 and 1970
Glen Woolfenden color banded the local Scrub Jays at the
Archbold Biological Station and censused for other birds.
We attempted to use non-overlapping band series so that
birds from t.hs Station and Hicoria could be. distinguished.
The Station is 2.0 miles due north of Hicoria, and to date
no birds from Hicoria have been found at the Archbold
Station and no Station birds have appeared in Hicoria.
As documented at Hicoria, the movement of Scrub Jays
from hatching sites is delayed for an entire year by the
retention of offspring as "helpers.,! Breeding males may
induce this retention by continuing to feed juveniles
throughout the year and so form with the offspring a loose
bond, similar to a pair bond. When birds do disperse it
is apparently for short distances. Yet in certain cases
family groups are found far from scrubby habitat. In one
case a fire destroyed a large section of scrub in late
summer, and a group of five Scrub Jays was found two days
later approximately three miles from the fire, feeding at
the edge of a large watermelon field, with no scrubby
habitat about. The next day the birds had disappeared.
Four miles northeast of Hicoria the scrub on several
thousand acres was destroyed during the summer of 1959 and
orange trees planted. For a week thereafter I found small
groups of Scrub Jays in situations where I never saw them
before and where they were not seen subsequentlv. As sug
gested for the movement of brsedincr oairs at Hicoria,
long


Figure 2
- Map of study area at Hicoria showing terri
torial boundaries and Scrub Jays residing
in each territory during the breeding season
1969. (See key for Fig. 1).


EXTRA-TERRITORIAL MOVEMENTS
About 60 per cant of the terrain at the Hicoria study
area was tied up in defended territories. The remaining
area was inhabited by small loose groups of nonbreeding
birds and by territorial birds which often wander out of
their home area. In 1968 two feeding stations were main
tained, one in a non-defended area and one immediately
between two territories. At feed site 2, placed between
territories 5 and 4, only the birds inhabiting those two
territories visited this food site-, accompanied by S-BY
after her mate disappeared. Although border conflicts
between BS-d3 and RS-B occurred frequently, encounters
were infrequent at this food site. Birds from the two
territories hardly ever fed here at the same time, and
this obviously reduced the necessity for territorial de
fense at this site. During the few times both groups
were together RS-B once chased S-BN and occasionally
supplanted BS-dB.
Feed site I was visited not only by territorial birds
from adjacent territories but also by ten other birds that
frequented this general area and apparently had no terri
torial affiliations. By plumage characters eight were
first-year birds and two were adults.
39


33
established among the "helpers" at first. During the
first winter, at times a male first-year bird would sup
plant a female sibling by displacing her from a perch or
by pecking at her, but again the usual pattern was of no
interference.


6


66


71


67
"3iq Scrub, Marion C:~:ntv
Elevation is 140 feet. This is an expanse cf scrub
1.7 miles wide and 12 miles long, located 12 miles west of
Belleview. Dense stands of sand pine cover much of this
area with no breeding Scrub Jays. Yet large stretches of
low dense thickets without sand pines are present. As
usual the ground is usually open sand with few hems,- but
large open spaces are lacking. The three oak species
occupy 30-40 per cent of the ground cover in thickets, and
Ilex cunulicola and Lyonia ferrnginea are common (30 per
cent of ground cover in thicket). This vegetation varies
from 8-15 feet in height.
The only part that contains breeding Scrub Jays is
the narrow zone along State Road 484. Here jays are
frequently seen feeding along the open roadway, but one-
half mile into the scrub from the road no Scrub Jays are
found. Eight miles north of this road, Interstate 75
crosses a section of scrub. Parts of the scrub bordering
Interstate 75 are similar to that along State Road 434,
yet no Scrub Jays nest here. Interstate 75 is a relatively
new road, and apparently Scrub Jays have not located this
newly opened habitat.
Ocala National forest, Marion County
Elevation is 130 feet. Although this is the largest
sand pine scrub in Florida, few parts of chis extensive
regc.cn suppers nesting Scrub Jays. Much of this area is


Figure 6 View of low and extremely dense Atlantic coast
thickets at Marineland, Flagler County. Salt
spray has retarded the upward growth of the
plants producing an extremely tangled thicket.
The only open spaces are along the road edge
at the extreme foreground of the picture.


53
Most frequently the dominant plant is sand pine (Pints
clausa), which often forms a compact forest. In the less
dense stands of sand pine the understory is a low thicket
of various woody plants, and the ground is often open white
sand, perhaps with a thin layer of Cladonla spp. Seme
herbaceous flowering plants occur, and occasionally a
few grasses (Andropcgon f loridanus and ? an i cura patent!-
folium) nav occur.
\viren present as an understory to sand pines, the
woody plants may be well spaced, but when sand pines are
lacking, these shrubs frequently make an extensive, dense
scrub thicket, in form strikingly similar to the chaparral
communities of Arizona and California. The dominant plants
of these low scrub thickets are three species of xeromorphic
oaks (Quereus virginiana martima, Q. myrtifolia, and Q._
chapmanii); these three frequently compose over 50 per cent
of the ground cover in the thickets. The remaining planus
may consist of a variety of species, including Serenoa
repens, Sabai etonia, Ilex cumulicola, Rhus coralline,
Lyonia ferrv.gir.ea, Myrica cerfera, and occasionally
Ceratiola ericoides. As Laessle (1958) reports, Florida
sand pine scrub normally includes many other shrubs and
herbaceous species, but the list above includes the woody
species with which Scrub Jays are frequently associated.
Although the ground is often bare of grass, the woody plants
form dense thickets with few oren areas.


25
actively bobbing and giving loud "hiccup" calls. When the
male flies back, the female usually follows in an undulating
pattern and calling. She sometimes "hiccups" in flight.
These border rebuffs are not cne-sided affairs. If
a male chases a bird from his territory the roles are
reversed when the latter's territory is invaded. Each
breeding male then has a definite boundary over which he
will allow no intrusion during the breeding season.
These border disputes occur most frequently in the
morning. Later in the day when the heat and humidity are
high Scrub Jays become quiet and little activity is evi
dent. At sunrise the jays are very active and territorial
pairs perform a series of encounters that are repeated
at the same place and time, day after day during the
breeding season.
The following description of the movements of GS-G
and mate in territory 3 will describe these encounters.
On 21 May 1969 at 0650 just before sunrise GS-G came
swooping northward toward the boundary between his and
BS-d3's respective territory (territories 3 and 5).
GS-G was flying an exaggeratedly slow and very undulating
pattern, calling loudly and rapidly kwerp-kwsrp. At once
BS-dB flew toward GS-G using the same pattern of flight
and else calling loudly. When approximately 20 yards
apart they simultaneously circled and flew back toward
the center of their respective territories, still calling
rapidly. The female sometimes followed behind and gave


Figure 1
- Map of study area at Hicoria showing terri
tory boundaries and Scrub Jays residing in
each territory during the breeding season
1968. Note: in territory 12 SY-Y died and
S-EY moved to territory 4.
KEY
1 inch = 300 feet
Shaded area = scrubby thickets
Light area = open sandy spaces
Jj = paved road
j = dirt road
railroad
fence
territory boundary
Letters designate a specific Scrub Jay, i.e
RS-R
o* = breeding male
$ = breeding female
* = "Helpers"
All birds without a special sign following
their letter designation are young of the
year, i.e., 1G-YS in territory 2.


12
Upright posture.This is difficult to classify as
a special behavior because Florida Scrub Jays appear to
perch naturally with the body upright and tail hanging
straight down. Yet the male of a breeding pair habit
ually perches in this posture on a. tall object from which
the bird has a clear view over much terrain. It is often
possible to identify a Scrub Jay from a distance by this
posture. This position can be. classed as neutral in
motivation.
Dip.--In this posture the head and body are bent
downward and the tail is raised above the horizontal.
This posture always expresses increased excitement and
usually is an alarm, reaction.
Bobbing.Again when a bird is excited one reaction
is a rapid bobbing of the whole body without moving from
the perch. The degree of excitement can be roughly
measured by the rapidity of the bob.
Undulating flight.--Normally a Scrub Jay flies from
one place to another with a steady, slightly undulating
flight. When highly motivated this flight is more deeply
undulating. This pattern is normally accompanied by a
series of loud kwarp calls and is often exaggerated by
flying slowly. Undulating flight often follows an encoun
ter with another jay.
Bill up.The bird tips the. bill up and remains
silent, and the body is often slightly crouched. This
is a submissive posture and is usually followed by the
bird flying away.


9
when nesting areas consisted of many scattered patches.
Even so, centers of population numbers were located and
their common physiographic features assessed.
Ten areas that included a variety of types of plant
cover, yet harbored nesting Scrub Jays, were examined in
detail. A nest census was made. Plant cover was analyzed
and the amount of ground in the thickets shaded by the
dominant woody plants was estimated. Unless I am dis
cussing a specific plant community such as the Florida
sand pine scrub, the word scrub connotes a thicket of
evergreen woody plants that is low and dense.
Field work began in June, 1967, and was completed in
May, 1970.


Figure 5
- View of low, dense scrub thicket two miles
southeast of Hicoria, Highlands County.
The only open spaces are along the dirt
road. In the background is a patch of sand
pine without nesting scrub jays.


ECOLOGY AND BEHAVIOR OF THE
FLORIDA SCRUB JAY
b7
PETER WALTER WESTCOTT
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THB
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1970


RECOGNITION SIGNALS
The calls, postures, and movements that are used in
social interactions are treated; other variations have
been described by Amadon (1944). In this section the
author describes individual calls and postures and some
times implies their motivation. Later sections discuss
how these calls and postures are combined in complex
behavior.
Auditory Signals
Kwerr or kwerp.--The Florida Scrub Jay utters a
variety of calls, yet most are variations of a rather
harsh and grating note. Amadon phoneticizes this basic
call as ka. To me the basic call contains more consonant
sounds (kwerr). This call, as Amadon notes, is used for
several functions, and certain variations appear to be
used consistently in similar situations or to express
particular motivations.
An upward inflection of the kwerr call normally
indicates a heightened motivational level. Such calls
are given when some alarming object is present such as
a predator.
An increase in intensity or loudness of this call
also indicates excitement but is normally used when in
10


46
to expect a handout of food, yet they have become so tame
that one can frequently pick up a brooding female from
the nest, something that could never be done at Hicoria.
An interesting example of this tameness occurs at a
small scrub thicket north of Englewood, Sarasota County.
For several years people have been feeding Scrub Jays there.
As one stops the car several jays fly out of the thicket
and often land on the car or an outstretched hand. If no
birds are in sight, all one has to do is sound the car
horn and the jays quickly appear.
It is difficult to explain this tameness. Although
most jay species are easily enticed to feeding stations,
only the Canada Jay (Perisoreus canadensis) is as overtly
tame as the Florida Scrub Jay (Bent, 1946).