C.I SOME E C O L O G I C A L IN T H E SONG ASPECTS OF S O C I A L BEHAVIOUR SPARROW, § § l o s p J L z a m e l o d i a by RICHARD B Sc., Lakeheaa A THESIS THE WALTER KNAPTON University, SUBMITTED IN P A R T I A L REQUIREMENTS MASTER in FOR OF 1971 FULFILMENT T H E DEGREE OF OP SCIENCE the Department of Zoology We accept this thesis required THE UNIVERSITY OF as conforming to the standard BRITISH September 1973 COLUMBIA In p r e s e n t i n g an the advanced degree at Library I further for this thesis shall the of this thesis written University of B r i t i s h permission for s c h o l a r l y p u r p o s e s may his f u l f i l m e n t of make i t f r e e l y a v a i l a b l e agree that by in partial representatives. be for f i n a n c i a l gain permission. Department The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, C a n a d a the I t i s understood Columbia shall requirements Columbia, for reference extensive g r a n t e d by the and Head o f my be thesis Department copying or that study. copying of t h i s that not I agree for or publication allowed without my i ABSTRACT The purpose experimentally Song of some Sparrows, this ecological Helosjaiza melodia questions Two A. Does the temporal influence the number of territory size, and breeding B. the the opportunity arises, hierarchies established ft necessary answered is that non-territorial are and the showed area one there and that of the probably Two of of a in for both of were c a r r i e d but were was types surplus of birds were and given area? season? to but by the are which resident fall and on and of 1972 breeding the breeding. juveniles, be investigated. the Sparrows the dominance breeding, in of when in replacements Song hence which experimentally p h y s i o l o g i c a l l y capable replacement local a territories birds territories Subsequent in British questions of experiments there a pre-breeding is 1973. on individuals also of Island, territories, surplus taking behaviour territories, p o t e n t i a l l y capable from social settling density the investigate : obtain during to Reifel asked dominant exists of This spring that on occupied that the holders. Removal are pre-requisite prevented territory , pattern juveniles is aspects Columbia. Are main study study All a l l but were origin. of removal experiments, simultaneous and i i successive, were carried out i n both the Simultaneous Removal nine to Significantly and the Further, ten days. increases the mean significantly boundaries The three the was both were completely replacement on days. number of after the than size that and replacement territories and fall after were the before. Further, nor taken, was territory removals. took upto difference i n the the about removals Finally, significant taken, took 40%. areas no On about S u c c e s s i v e Removal was fall. were the territories removals. more spring rearranged following the There total spring territory smaller to four size, in areas, the mean in territory territorial pattern retained. Breeding same before several these density and unmated unmated territories after males than the experiments. territories proved mated which a l l a r e a s , however, with males Factors results on to remained much Therefore, there after have the the were removals, significantly and smaller ones. could have of the simultaneous experiments are discussed, accounted and and the a f o r the different successive proposed removal explanation i s given. Dominance of juvenile certain hierarchies Song Sparrows localities along were that the determined congregated hedgerows. in the over Each loose the group groups winter tended at to i i i be a discrete (both dominant themselves reversals of unit, although and were some interchange subordinate) stable and and t h e o c c a s i o n a l essentially removal experiments the members of the hierarchies that,dominant successful largest obtained ones presented males in i n establishing hierarchy, Factors the position of the bird the possible outcomes The hierarchies linear, with an o p p o r t u n i t y to obtain the few hierarchies which males, could i n the hierarchy of the hierarchy were Further, 4 from possibly are are f o r some territories. territories. o u t o f 12 j u v e n i l e territories. individuals triangle. The was f o u n d occurred. of I t the i n the the top 5 influence discussed, considered. and iv TABLE OF CONTENTS PAGE ABSTRACT i TABLE OF CONTENTS iv L I S T OF TABLES .. L I S T OF FIGURES > v ACKNOWLEDGEMENTS GENERAL vi i i viii 1 INTRODUCTION 1. . AIMS OF THE STUDY 2. THE STUDY AREA 3. L I F E HISTORY U. METHODS PART A. THE EFFECTS OF TEMPORAL PATTERNS ON TERRITORY S I Z E 1 3 5 11 .. SETTLING INTRODUCTION 1. F o o d s u p p l y and t e r r i t o r y s i z e ..... 2. Other environmental factors and t e r r i t o r y s i z e ..................... 3. Temporal p a t t e r n s o f s e t t l i n g on territories I I PROCEDURE I I I RESULTS 1. R e p l a c e m e n t o f removed p a i r s . . . . . . . . 2. Number o f o c c u p i e d t e r r i t o r i e s ...... 3. Size of t e r r i t o r i e s 1. C h a n g e s i n number of- t e r r i t o r i e s on t h e F a l l Removal A r e a o v e r t h e winter 5. Breeding d e n s i t y i n the s p r i n g of 1973 6. Differences in territory size between mated and unmated m a l e s .... 7. The r e p l a c e m e n t b i r d s 16 16 18 I IV DISCUSSION ... A. The r e p l a c e m e n t Song S p a r r o w s and the s u r p l u s B. The t e m p o r a l p a t t e r n s o f s e t t l i n g on t e r r i t o r i e s . . . . . . . . . . . . . . . . . . . . . 2 2 2 4 2 ? 32 32 33 35 40 42 44 47 51 51 55 V 1. C. Fluctuating environmental resource 2. P r e s s u r e from i n t r u d e r s 3. Inexperience of the j u v e n i l e s and the temporal patterns of s e t t l i n g 1. Disruption of territorial boundaries 5. Genetic basis f o r the size of t e r r i t o r y .................. Proposed explanation f o r the different results of the simultaneous and t h e s u c c e s s i v e removal experiments ................ I INTRODUCTION , I I PROCEDURE I I I RESULTS A. The h i e r a r c h i e s B. Success i n gaining t e r r i t o r i e s i n r e l a t i o n t o h i e r a r c h y p o s i t i o n ..... DISCUSSION a. The functional significance of dominance h i e r a r c h i e s .............. B. Factors determining 1 2 3 M 5 6 7 8 9 67 68 72 75 80 8 0 88 94 94 position i n 95 Outcomes o f t h e h i e r a r c h i e s 102 LITERATURE CITED APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX APPENDIX 63 dominance h i e r a r c h y the C. 59 72 PART B"... DOMINANCE HIERARCHIES IV 55 57 106 .-. ., . 118 120 120 123 124 126 127 128 129 vi LIST OP TABLES TABLE 1. fall PAGE D i s p e r s a l d i s t a n c e s o f j u v e n i l e s from 1972 , site 2. R e s u l t s of removal experiments: numbers t e r r i t o r i e s , o f both mated and unmated males of birth: 8 of occupied 3. R e s u l t s o f removal experiments: d i f f e r e n c e s i n numbers o c c u p i e d t e r r i t o r i e s o f mated and unmated males 4. R e s u l t s o f r e m o v a l experiments: mean t e r r i t o r y s i z e . Removal Area • 5. R e s u l t s o f removal* experiments: mean t e r r i t o r y S p r i n g Removal A r e a ................................... 34 of 36 Fall 38 size. 39 6. R e s u l t s o f r e m o v a l experiments: breeding d e n s i t y .... 43 7. D i f f e r e n c e s in males: s p r i n g 1973. 46 t e r r i t o r y s i z e between mated and unmated Simultaneous Removal areas ........ 8 . V e s t i n g d a t a f o r t h e C o n t r o l s and the E x p e r i m e n t a l 1973 9. Number o f j u v e n i l e s banded, f a l l areas: 1972 t o e a r l y s p r i n g 1973 10. L i n e a g e : t e r r i t o r y s i z e s from one g e n e r a t i o n t o the next 11. Numbers o f e n c o u n t e r s (displacements, avoidances f i g h t s ) i n f o u r dominance h i e r a r c h i e s ................. and 12. Analysis Hierarchy B-C of the avoidance interactions in the 54 ^ 84 13. Rank i n h i e r a r c h i e s and sex o f Song Sparrows i n t e r g r o u p movements performing 87 14. A n a l y s i s o f f i g h t s i n the B-C H i e r a r c h y 15. Change i n s t a t u s of a j u v e n i l e female d i s a p p e a r a n c e o f an a d u l t t e r r i t o r i a l female 49 89 following 16. Feeding o r d e r o f h i e r a r c h y : appearance o f f i r s t f e e d i n g s t a t i o n s o f the B-C H i e r a r c h y the 92 b i r d at 1 0 4 vii L I S T OF FIGURES , FIGURE 1. The s t u d y a r e a : Mestham I s l a n d Reifel Island PAGE and an adjacent part of 4 2. Numbers o f j u v e n i l e Song S p a r r o w s r e s i g h t e d on t h e S p r i n g R e m o v a l A r e a f r o m mid-December t o t h e end o f May . . . . . . 10 3. Singlengths o f male and 12 f e m a l e Song S p a r r o w s . . . . . . . . «». Mean w e i g h t s o f male and months female Song Sparrows for a l l 13 5. Fall Removal A r e a : t h e movement and e x p a n s i o n o f C o n t r o l male C6 a f t e r t h e r e m o v a l e x p e r i m e n t and d u r i n g the subsequent winter 41 6. Spring boundaries 7. The B-C Removal b e f o r e and Area: Simultaneous a f t e r removals Removal area 65 Hierarchy 81 8. Number o f i n t e r a c t i o n s v e r s u s distance f o r a l l j u v e n i l e s i n t h e B-C H i e r a r c h y 9. P o s i t i o n i n h i e r a r c h y and Hierarchy within hierarchy number o f i n t e r a c t i o n s : 89 the B-C 103 1 0 . B u l l d o z e d Area: the arrangement of territory boundaries b e f o r e t h e dyke r e b u i l d i n g o c c u r r e d ................... 11*1 11. Cottonwood Stretch Hierarchy 12. Sedge P a t c h 13. Bulldozed 14. First 15. No Hierarchy Path Hierarchy B i g A l d e r and Marsh N a r r o w s H i e r a r c h i e s Name C l e a r i n g H i e r a r c h y 1*9 1£° 131 132 1£3 . viii aCKNOWLEDGEMEHTS I for am m o s t h i sadvice, work. read Dr. I in are C. J. a t the start should who g a v e nest-finding. assistance and Dr. and a d v i c e Dr. J. R. a t a l l stages N. R. was r e c e i v e d Liley from Krebs, of this critically Dr. J. F. o f the study. t o thank during P. the Taitt removal who a s s i s t e d i n t h e b a n d i n g Their D. help due to with t h e computer Finally, patience Krebs assistance a l l the people supervisor. and guidance further like valuable also t o my criticism the manuscript, Bendell and grateful I should and t o l e r a n c e was truly Lauriente, like S. a n d W. Zales, experiments, operations and appreciated. Borden a n d B. Thanks Webb f o r programming. t o thank t h e study my would wife, n o t have without been whose possible. 1 GENERAL 1. Aims of The purpose ecological HelosjDiza of the first of with density and 1970; are evidence some settling of study, settlement test this Tompa and Sparrows, classical examples and the 1964). and some Song i s divided has been dominance This into the shown territories occupied patterns, hypothesis. perplexing in given territory any I in a of mean area environmental Krebs then, A variability density, (see territories the the supporting on in territory 1943; 1972). influence evidence investigate two territoriality, several conceivably show in several populations been population There the behaviour Klomp has to behaviour Sparrows both, is two study sections, second dealing hierarchies. breeding however, Song aspects dealing study social 1937, Territorial hence of (Nice dominance Moss this interaction: considers with of melodia. hierarachy the Study aspects social INTRODUCTION of them 1971) given Watson problem in birds, territory from year and size, to year. that could the produced successive, the There pattern i n determining In and but temporal area. experimentally simultaneous (see unequivocal. the the animals annually, i s not i s important in a determine resources size that to the first two is of number part of types of i n order to 2 In the second hierarchies in congregated over area. The the hierarchies were hierarchies yearlings of but the on from Song 1964). that Sparrows The provided a a of surplus fall of test and that doing so process to the i t by the spring classical independant for the was study is hierarchies were the outcomes spring In the of the when the particular, opportunity, the spring hierarchies the there I the are the established investigation existed worth birds on of of the of breeding, of descriptive another population (Tompa on, there both behaviour Columbia that of surplus good instigated (and 1962, indeed would be such the study area from that the dominance 1973. mentioning territoriality features in British assumption of capable is operating was a territorial There southwestern for), in that the physiologically non-territorial Finally, be was present 1972 hierarchy in Sparrows study in dominance dominance the the given individuals. this of at season. birds, territorial evidence the Song localities and that, pre-requisite problems looked disintegrate. pre-breeding prevented certain territories in I juvenile during to obtain non-territorial of winter, followed necessary study, composition the individuals these at hypothesis that the A the the groups and began investigated of winter structure over during loose the determined dominant part a bird's are not social considered behaviour to (Brown 3 1963; Davis 1959). major parts, with not be m u t u a l l y 2. The Study The the part north west by Westham to of part Topography f o r most into parts two need 1). Island The of the a n d an area i s in and i s bounded to arm o f F r a s e r River, t o the C. Eeifel George by a g r i c u l t u r a l f a r m l a n d i s flat, t h e water o f the year, and various the with early in fall and i s tidal, hedgerows. vegetation table of i s close and t h e l a n d i s of o f woody Appendix species. i s and herbaceous 1 gives a list The s t r u c t u r e similar along and a l l the identical. t h e dykes, Strait species and i m p o r t a n t but i s not varies sloughs two Columbia, to the south grown t o form Georgia levels the Reifel (Figure British and the dykes, Beyond spring and m o r e common hedgerows, level i s divided dyked. have the on by t h e s o u t h Strait surface composition and east Island. Along of that out Island of Delta, Refuge, necessity carried o f Westham and the s o i l plants the understanding was Georgia Waterfowl thesis Area municipality the this exclusive. study adjacent Therefore, t h e marshes are essentially the season: summer winter. rising bordering river the Fraser freshwater. The l e v e l s are high the to f a i r l y low the marshes and daily. In but decrease The in and f a l l i n g twice water during run-off, water River summer. 4 FRASER RIVER N South Jetty FALL REMOVAL AREA SPRING REMOVAL AREA Figure 1 The Spudy A r e a : R e i f e l I s l a n d and an a d j a c e n t p a r t o f Westham Island 5 low tides The partly occur land as in daylight, enclosed pasture The crops grown to season, the for cattle, on and by the i n winter dykes and on year to study as land year, night. the partly agricultural from at area was used agricultural changed owing to from the land. season rotation of crops. The territorial vegetation marshes at along low fall and water levels fields, 3. the tide. winter were either Life Song Sparrows hedgerows, The foraged low. No pasture or were found occasionally non-territorial extensively Song at any in foraging juveniles i n the Sparrow crops, mainly was marshes seen to the in the in the when the use the time. History Song Sparrows have a wide breeding range in North o America, 1963) and the suggests a environmental Beer et (1937), a l . very variety Nice (1943) attachment of the high (1956), of these lower more The Phillips (1956a,b), habitat and of plasicity conditions. Johnston wide In recognition types Tompa birds mainland of 30 the studies and and the of show a of (Dickerman s p e c i e s to Song Dickerman Marshall site races on for breeding, (1964) to than Sparrows (1957), (1948a b) # and varying the Nice reveal studies highly by a of developed birth. British Columbia, the Song 6 Sparrows Hi I i are ™2££^BS# Lowlands, and east, and study The in by however, by Nice flocking Nice, behaviour Sparrows on during winter i n c e p t i o n and bird; place probably in Weather snowfall biotic August data are occurred The patterns Sound areas to the (Munro north and caught in Jjorphna. has birds i n many therefore a Puget A l l birds Sparrows 1943). the occur nu Song concerned, on to discussed Heifel respects detailed relevant been Island to those description this study (Nice 1941). is are, here. have on in Type Heifel the winter months, described and given lasted September of 40 (cf. in Appendix in late December, Tompa whole the on of their to Reifel the for Island territorial inclement weather post-breeding moult. 50 Nice 2; The (1964) year, moulting to within contrast birds periods during time in by The the during moult remained round, Island. length of and territories Island f o r almost only the A year Mandarte territorially ceasing to Forest Certain aspects behaviour The Coast of and boundaries the in subspecies, behaviour holders territory resident subspecies (1937, Sparrows territory behaved the discussed Song is Other behaviour identical unnecessary. Song of general described subspecies morphologically distinct. were detail showed I s l a n d s , and 1947). are The Oberholser, Gulf Cowan, this non-migratory. varied from bird days, and took 1937; the otherwise Tompa only the v 1964). sizeable winter was 7 relatively mild. Territory and one were pair boundaries linearly would have From observations, males into and each arranged there Island along i n general others becoming Sparrows perform end of the were banded long were well-defined, the hedgerows. only was two little territories; independent Hence, neighbouring were any pairs. t r e s p a s s i n g by boundaries this of their e x p l o r a t o r y movements post-juvenile shortly distances resighted adjacent recognized after were moult. leaving (Torapa In of on most parent's Song until young suggested fact, their young 1964), Observations the nest covered. within the boundaries parents, the that that young no were territory at time. In become the f a l l , after territorially dispersal of the individuals whose for a on discussion Mandarte Island, coincided with females, of the post-breeding active young. by some Reifel respected. On some on rise to neighbouring dispersal evidence to suggest from and the study more, 1 gives birthplace the emigration the emigrate Table longthe once was and the the this adults results distances known in of fall ( s e e Howard territorial many young, 50% On Reifel Island, d i s t a n c e s were most area. of the quite of 1960, For example, On behaviour which large, juveniles in covered short-distance dispersers). islands. that moult, were although there i s did o u t o f 52 not banded 8 TABLE 1 Dispersal distances of juveniles from s i t e of b i r t h ; Numbers of juveniles Distance (m) 4 0-150 2 150 - 500 1 500 - 750 1 750 - 1000 2 1000 - 1500 1 1500 - 2000 f a l l 1972 9 juvenile Area during October within to Song and Sparrows August areas juvenile amongst young individuals in favoured centres of hierarchical In as yearlings spring, to Sparrow Waterfowl groups ( s e e PART Island and only one two areas. Song April. ones from and were banded seemed Sparrows Figure the on bird t o be, Tompa their on 2 shows any fall banded of concentration hedgerows. the groups began hostility 1964:26). began May, the hedgerows at These winter as study then, a sharp area no in t h e numbers of 3) area. was the in the the early The April Reifel the study and area April located decline late between non-territorial i n March (see Appendix up, exclude From adjoining to break Also, to 1973, surveyed, the study i n the non-territorial territories. i n late also activity, B). birds recorded as throughout increasing (see of the study Westham of of visits in present. the persisted territorial was along Removal resighted to reveal centres the h i e r a r c h i c a l result Refuge There formed localities themselves the end Song remained birds non-territorial failed birds fall Fall were occasional area that the of t e r r i t o r i a l Sparrows young a the t h e peak Further, on 19% t h e many S o n g the spring, possibly present September, the study the certain were after area. outside The and November, the study that March juvenile the numbers and Song 1973, in i n the on early Sparrows 10 Each month i s d i v i d e d i n t o two two-weekly time i n t e r v a l s Figure 2 Numbers o f j u v e n i l e Song Sparrows r e s i g h t e d on t h e S p r i n g A r e a from mid-December to the end o f May Removal 11 resighted of Hay, two-weekly showing spring. in in Tompa Song a sharp (1964) Sparrows, and been found in 1963; Dhondt 1971b; 1967; Watson left on the 4. Methods The of the banded area of each a 282 few the Each tarsus, and each was banded Sparrows weighed, culmen. and Males winglengths f o r the Although to 72 males throughout the on 1972 mm to one (Figure Island Carrick Song study May have 1961; Smith Sparrows pairs. area 1973. necessary was Because to have a recognizable. aluminium and During mainly in for length females can winglength banded one the or study mistnets, 1937). from more was of sexed birds; females there be (Nice slightly 4), the traps. measured weighed in territoial the caught, in winglength, average year on (e.g. only the end in spring individually with and 3 67 on i n p o r t a b l e sparrow was the i t was were and from Orians bird differences the a l , February behavioural gives birds apparently, the Mandarte species of 1963; to seen i n numbers population from on i n various combinations. Song bird same r e s u l t experiments, bird caught et numbers i n m i d - A p r i l were Sparrow c o l o u r e d bands, period, with declines population, Therefore, two sharp other mid-December i n the the Thus, continually from found Jenkins 1965). study nature decline several Song observed intervals too males 63 to than much wing, both on Figure ranged 68 mm. females overlap NUMBERS Figure 3 W i n g l e n g t h s o f male and female Song Sparrows 13 M E A N WEIGHTS (gm) 4 Figure 4 Mean w e i g h t s o f male and female Song Sparrows f o r a l l months 14 between be 'light* useful of area Song than and culmen was u s e d Juveniles fall plumage feature i s that retained colour (Nice of yellow in decreased proved identified their the dusty, the 1937). fleshy colour. i n size, have a t t h e base parts In juveniles , plumage males I also assume a which a r e have used rounded was the of the mandibles. the and a r e fleshy colouration c h a r a c t e r i s t i c when in distinguishing are extensive, but the yellow t o be an o b v i o u s which other area. retrices, adult the subspecies and then useful pointed whereas case no s u b s p e c i e s moult, A A criterion parts f o r other finely-streaked adults. the winter, in on t h e s t u d y post-juvenile a l l juveniles these taken earlier, the fleshy fledgelings, were As m e n t i o n e d to through retrices of sex. ground loose similar f o r this characteristic to measurements o n e was after females as a wintering Sparrows. the resident late 'heavy' i n determination Tarsus study and bright parts have i s retained, the In bird was and in fe" the in hand. Certain 1972) r e t a i n e d The fleshy dark grey. response Song of Sparrow's described the yellow parts Territory in a yearlings (possibly colouration a t t h e base were territorial male on a t a p e Dhondt 1966). into of late broods February 1973. o f the mandibles boundaries! song members determined i n adults are by noting to the playback recorder This (Sony method the of a stranger 110A) (method proved very useful. 15 Males the a would tape territory boundary any from various clearly and sing. of of by the usually this song or by had f l y to a song of perch along the particularly as Thus, the relatively needed. cross be neighbours, not recorder could two was to elicit not boundaries time, in tape would only size close I could the Males territory territory (1960) flying playing hedgerow. way, by display. short space hedgerow. Suthers birds would very new sing the but In measuring the either along defined i n a linearity proposed spots pair to territorial boundaries, one method quickly r e c o r d e r , and response at react Song owing to the complicated Sparrows as 16 PART A .THE EFFECT OT TEMPORAL SETTLING TERRITORY I. breeding of Brown 1970a; SIZE Klomp five in population . far fully removed means by follows. and to then The a population in limited and they 1 by feel need show no a study in resident territory replacement, (a) behind and a i f (b) any, in removal and to date holders 8 experiment new be breeding occurs, Appendix list to i f territorial individuals removed, other (1970a) limits which animals, some behaviour that on populations which Moss Moss opinion conditions. area, are to of g. Chitty and a l l five conditions rationale way Watson that lines the e. 1966; breeding the which proceed satisfied given limit been years, Watson main been out. show subsequent which In have 8) experiments, determined. stable order two determining has Rowan 1969; i t i s simply (Appendix animals, 1966; Lucas are in recent Lack i t does themselves They Removal (b) populations space satisfied (a) of in 1965; There behaviour; conditions so Crook F r e t w e l l and v e r t e b r a t e s , or factor, i t s importance populations 1972). whose b r e e d i n g and discussion 1962; 1969; territorial are in much Wynne-Edwards 1967; behaviour, density subject has ON INTRODUCTION Territorial of PATTERNS are can be is as from a individuals I 17 subsequently replace replacements had been in that prior area on by t h e p r e s e n c e for presence that breeding, removal of species; 1964), i n fish Healey 1961; Watson Elliot 1967; H a r r i s i n holders of that these territories individuals, a territory deduced limits breeding been carried i s that density territory between 1968), there one year and 1955; Moore (e. g. particularly and Cope Lockie in birds 1951; Orians and J e n k i n s 1968; B e n d e l l 1970; Young 1970; K r e b s i f of t e r r i t o r i a l minimum which within a between territory This variability size i s however associated species: would be and 1971; not so. with and within a habitat in territory size in are differences territory (c) t h e r e and There a r e variations (a) t h e r e i n t h e mean easier were c o n s t a n t , one p a r t i c u l a r p o p u l a t i o n habitats; size behaviour territory are are differences This Jacobs i n mammals 1951; H e n s l e y 1970; Holmes individuals within (b) mean size (e.g. o u t on a 1972). importance problems have Clarice 1970), for a l l individuals. years. on i t c a n be f u r t h e r 1965, 1967; Watson demonstrate three holding insects and A l d r i c h 1971; Young The the then 1967; Smyth Stewart equal settling of the t e r r i t o r i a l experiments (e.g. (e.g. to from the of the territory variety Peek If i t c a n be d e d u c e d area. Such 1966; then prevented to the removals. necessary the them, and size year; within are differences i n over successive has prompted Lack 18 (1966) to reject the regulates breeding territories have in limiting major, density. He minimum the breeding Wood, that to minimum year size, since year territory territorial pointed population Oxford, c o n s i d e r a b l y from argues year that a fixed i n Barley varied hypothesis this for territorial out that not operated of the Great the breeding t o year. size has However, need n o t be behaviour behaviour Tits, i f Parus density Krebs has (1971) constant from t o determine breeding a possible mechanism density, My causing year study was i n t e n d e d the differences within considering an now to investigate i n mean area territory (problem some f a c t o r s (c) which taken from year stressed that the possible factors Further, mechanisms my differences 1. Food approach between su££ly Territory if size of availability only, t o the study individuals and t e r r i t o r y size could territory of year. not food. above). I t year i s It should was worth however t o be d i s c u s s e d to to influence the size ultimate (a) of be refer to mechanisms. not intended (problem to e x p l a i n above). size fluctuate was from could territory proximate to size annually proximately The evidence with food adjusted here i s supply, to the far from unequivocal. Differences i n mean territory size in various habitats 19 have been related some instances, in three Tinbergen 1960), species and and , Mason and numbers there other of other, a one that over Oyster-Catcher, from taken to defend, Orians size a territory, confines, factor by the a 1969), and the Egglishaw times species biomass of richer Turnstone, 1968). Oncorhynchus Dunlin, Laqppus l - _ three (Smith (Glas the Grouse, biomass was In higher kisutch, where on salmonids food of fish, and (1967), in their suggests which the of years Harris a a the in two than the territory size and general Blackbird, that territory food i s related (1970) compromise proportion mean ostralegus, young, (1961)found the in Haematopus probably feed and number conclusive. Red-winged but Red and coelebs, in in indirectly (Kluyver food) greater differences for needed the 1973), a spp. Moss supply. three i s far of 1966; that supply birds a l . showed food population territory the g. (and, differed. Evidence the , e. Fringilla measuring 1970) quantity species, Tamiasciurus, (1965) found streams, a et genus greater hardly within (by food Parus j u v e n i l e Coho Salmon, hand, Scottish Chaffinch, (Settleship the Chapman was the (Miller of various titmice, (Holmes M § £ § l i l lBf®I£I®s, squirrels in of directly alpina, Scoticus differences in quality) 1953), Calidris to of is size the area A ^ e l a i us not regulated that, of the area the they between can the Eljoe n i c e us, obtained i s probably size the correlation food food suggested between the to the within i t s proximate and that 20 territorial behaviour There i s some area can 1958), evidence determine Ovenbird, Se^urus although experience was Evidence can be increase result this that food increased Salmon, demonstrate differing in Salmo a laboratory availability the territories were non-territorial). that fish receiving reverse whist deprived an to be superabundance determine in of true (1962) of latioes. ways food of in Brook found If food for the caused size (the then species. not 1971) that fry (1962) showed than those (1970) found in do the ineonstans, absence work they the subordinate complete aggression in of S t i c k l e b a c k , Culaea aggression might sizes aggressive Smith the could the Yamamoto whereas in fish, different and that (Symons in the their territory he changes size of deprivation feeding salmon a or causes suggested food more reduced and supply sizes experiment that i f age territory However, were food, food the with young the size. the Keenleyside food territory different food dominant abundance Bagnusson Ory.zias of of known influence salar. (Stenger not {1968) in given by i t is a taken year change size in one of associated food important. within can Symons of more territory territory to fish, i n aggression Atlantic # i f manipulation In an of study supply i s much quantity auroca£illus individuals in the way size influencing territories. some that the in obtained territorial in the or Medaka, together so a to apparently 21 Bustard (1969) gekkonid lizard, vary differing with social (1970a) and not cracks used the territory birds were supply, or i f young, whilst production food, in a of new from young however, 1966), in the not that Watson Moss and environments, a l l the suitable i t is to year, not found to size that which not the et mean that i f the In food number of food Peromyscus supported survival population food. d e c l i n e of the settled. food additional the was influence of excess the to the uneguivocal. found prevent the known in a l l of for improved with not territory year evidence showed Pitelka between of S t e r c p r a r i u s £omarinus, difference (1970) winter. relationship in availability spring a and the the in Jaeger, i s also fluctuations (1971) previous but increased failed did a concluded Krebs was the area to and size inverse (1959) Fordham stump the because related adjusting their Bendell as been lemmings, density but, year-to-year Pomarine mammals, population fiSSicuAatus, an there breeders small population supply i n the breeding on He the territory suggested of limiting, (Lack food limit, a of lizards. not that density be Tit, have size potential the young the (1955) by Great size to could upper food. p h y s i c a l s t r u c t u r e of out, f o r the In of females inhabiting the were the supplies of point experimentally and variegata, number an behaviour, related the set territory al. Gehyjra that behaviour In food showed a a of the size and Excess population 22 of Microtus californicus and o f Mus musculus Finally, appears within i n attributable nutritional Watson 2. Other levels are which of the some (1968), (1967), Zimmerman (1970) felt tadorna, area, was influenced (1961), lag. of heather not This directly has been through and Moss high 1970b; factors size been (1956), other from suggested and Snow e.g. Lin (1958), (1963), than year of territories (1961), food to year. as being i n a given D i n g l e and (1963), Davies and a n d Snow Watson (1964), Snyder and T e s t e r and Watson (1973). Young territory that size invertebrates Calhoun (1971), the shape, Warblers, both determined suggested has t h e numbers Dixon and by t h e l i m i t s (1972) Reed that taken, and t e r r i t o r y territory Alexander e.g. Buechner quality (Watson environmental animals, (1965), year chicks habitat area vertebrates the a g g r e s s i v e n e s s working influence i n influencing Caldwell one 1966), 1970). of t e r r i t o r y factors important in a the several of 1965; K r e b s 1971). might nature Grouse, the size to decreased and Belong 1 9 6 7 ; Newsome Red environmental There The the but with and M i l l e r supply (Delong to influence one y e a r , (Krebs size by in topography Tadorna topography o f t h e owner. but not the size, AcrocePhalus Shelduck, t h e immediate of tolerance the l o c a l the of the Catchpole o f reedbeds greatly of the territories scirp_aceous, and Sedge of Warblers, 23 schoenobagnus. and agonistic spacing for a can Tit (Lack LY.£2.1§]J£I» ( study to nestsites has Sparrow is the in of a of 1971; 1961; Zaret passerine Yeaton a shelter for pomacentrid the fish, of the supply shown in (refs. in habitat or the of type of Great Ficedula 1972). Tompa a 1961) shortage territory of nest the Sjostrand size of and a population Flycatcher, and the factor has and Band comm.) for some larger territories an and There effect of taken habitat hence that Brown on limiting Song territory the density and a Sheppard Song that resource Sparrows size, that of more in a given be (e.g. Ayala 1971). the the another 1966; Sparrows, will the vertebrates Michielson 1971; hypothesised the influence i s some e v i d e n c e 1966; particularly of could invertebrates Greenberg 1971; and which animal, both competing the of the occupy). Inger (pers. Sparrows competition. birds, Pied Song obviously, species and the limit been Enemar nature territorial one has 1956; possible populations Connell of birds, This i n f l u e n c e on will interspecific density that responsible n e s t s i t e s can in on limits, further density and date any the of Haartman V o n that A heads density. 1966), suggests Song suggested factors hole-nesting increase (within (1972) were coral shortage example, boxes No over Sale aruanus. Further, of, behaviour patterns Dascyllus Finally, 8. For I. species area, the ('horizontal' 24 rather than 'vertical*). circumstantial conducted to Krebs in date which each method minimum at and territories can Temporal In this that temporal influence on birds and Van the males introduced theoretically density, some be and methods been possible discusses in which related directly methods settling deals pattern on to the with of the settling on territories densities in degrees limit a there Assem of different settling set to the v a r i a b l e one. of in on supporting years could those years. number For settling i s some (1967) gasterosteus considerably territorial have on of the settled hypothesis territories evidence in can work fish. den stickleback, of be not these patterns density, is built density. upper can some temporal different owners however, experiments considers of the an no influence need breeding of way, territory could He one patterns result and presented influence Fluctuating the has size that hypothesis, test i t . length. territory hypothesis be to territory environment, 3. evidence, (1971:17) ways His with males were the found that aculeatus, pattern accomodated introduced successively. in of a the three-spined territory sizes settling. The tank simultaneously He in was as interpreted nearly when the varied number of double when they were results as 25 suggesting in the male settling males that the relative sticklebacks and b u i l d i n g were Turdus (1965) population number compared the was r e d u c e d of b y 55% 200 the pattern of s e t t l i n g year, Great and, compressibility February more Krebs mean showed February and Sedge conclusive adults a might he o b s e r v e d He that evidence size. with the Tits occurs influences birds in the warm to might settle increase. correlation breeding between density. patterns of f o r the variations in i n a three admits, 244 consistent temporal account to forward year was a p o s i t i v e suggested was 93 a s put density and s u b s e q u e n t The consistently and f i r s t the breeding there Warblers. has settling and stable rose territory least of a as suggestive (1971) at i n but synchrony then (1972) evidence influenced Blackbird, snowstorm by G r e a t both size the i s taken i s territories territory found i f snowstorm. on t e r r i t o r i e s temperature on after territories, that Catchpole mean i f synchronously, settling that of causes a severe T i t , Krebs Settling February, European territories after and t h i s evidence hypothesis. of the i n the previous year, that suggestive for held following the levels i n the process of t e m p o r a r i l y s m a l l e r than that the For a l l o f them number territories t o about i n aggressive successively. relates merula, - - were introduced Simms differences year however, study to to substantiate the hypothesis. of having Reed no 26 Possibly in birds i s from 1971). the The study year caused by that and these the and number of of study. I the the and> t h e from number therefore of settling of territories breeding density? the birds Song catastrophic peak active. for Sparrows, and territorial a the settle, now birds increased. the given had The fourth year investigate does hence amongst After territorial high the birds. mortality experimentally occupied, of to of mortality, the territories in in territorial allowed 1964; years of were territories two substantially established to hypothesis (Tompa first little remained out on at the Island territorially occupied also a March, spring not the in for caused of set question: number in the were territories following pattern in resistance Mandarte following snowstorm that on supports increased non-territorial gone, the but apparently yearlings best territories study occurred behaviour, storm, of the which Sparrows constant, freak storm those Song of a example number was third The the the the temporal area influence territory size, 27 II. PROCEDURE In order temporal number to of t e r r i t o r i e s occurred, there of juveniles holding the taking territory for territorial types •Simultaneous' empty then on necessary The i f came were territories for pairs. o f Song birds holders i t was that, surplus Sparrows from preventing the study i s area. i t c a n be f u r t h e r capable a As necessary of birds deduced certain then of the resident loose i t c a n be breeding, behaviour on t h e the Sparrow were removal replacement Song juveniles that can influence the a of removal and t h e removal females, creating show hypothesis deduced breeding, limited the density. Two involved area, juveniles, territory i f the replacement breeding a non-territorial pre-requisite that, i n a given I f the replacement from a on t e r r i t o r i e s would existed the resident the of the t e r r i t o r i a l themselves area. groups experimentally taken out removals experiments that test patterns of settling carry study to i n as an a r e a of 'Successive'. simultaneous Song Sparrows. a space suitable territorial allow Song settling This A type of birds, time habitat some o f removal carried of both out: Removal males and as possible, thus which Sparrows. of were Simultaneous of the t e r r i t o r i a l short of experiments was completely This was i n t e n d e d t o the non-territorial was i n t e n d e d to simulate 28 a catastrophic mortality but i t void of t e r r i t o r i a l for example same time are could here A when to a well birds migratory non-occupied treated at as regular situation time allow be a p p l i e d i n an a r e a of birds area. to other birds, instances cf a of suitable habitat, return i n the spring These types involved intervals, the periodic of t e r r i t o r i a l successive territorial of as at the occupancy equivalent. S u c c e s s i v e Removal pairs of equally o f non-migratory the removal simulating disappearance individuals. settling of the over This some of territorial more normal a longer period was intended non-territorial to Song Sparrows. Simultaneous the fall o f 1972 a n d i n t h e s p r i n g coincided with Sparrows were Each of s e t was such most were removed successful a s t r a n g e r Song they concerned Both responded the that either Sparrow, to removal Two These behaviour different experiments a Control conducted area by m i s t n e t t i n g and f e m a l e s the of times of Song hedgerows was were areas. o r by tape. females The strung over caught only 1). sandwiched shooting. t o be t h e u s e o f a t a p e d and a m i s t n e t in two (see F i g u r e and t h e S u c c e s s i v e Removal means p r o v e d males o f 1973. 1964). s e t o f removal arranged were of t e r r i t o r i a l Tompa the Simultaneous recorder. as 1943; f o r each Birds The t h e two p e a k s (Nice used between and S u c c e s s i v e removals song t h e tape i n the mistnets major i n the f a l l problem (see below). 29 Appendix 4 gives difficulty The Fall possible i n locating Removal This 1972. some females reasons at this accounting time. Area hedgerow contained 8 territories were used 24 territories i n the f a l l f o r the Successive for the Control, a n d 10 f o r t h e t h e S i m u l t a n e o u s 10 territories on adult males from mates, that intrusion on the Simultaneous t h e summer, had during the with and established the early t h e S u c c e s s i v e Removal mateless, a 2 fall. juvenile Removal. 6 The contained 8 males, without f o r themselves contained established of Removal, area Similarly, area territory Removal territories the 8 1 by territories juvenile by i n t r u s i o n male, early i n fall. 9 o u t o f t h e 10 m a l e s o n t h e were f o r the removed removed i n the period on O c t o b e r October 1 to locating him c o u l d completed the October. Also, possible 8 explanations 12. 11, This Simultaneous October last 4 t o 7; t h e l a s t male was and i t i s p o s s i b l e be due to post-breeding the moult on t h e S i m u l t a n e o u s females, for this only 4 were Removal not that male recorded was from the d i f f i c u l t y fact that until t h e second Removal area he area, removed. had d i s c r e p a n c y a r e g i v e n i n Appendix not week o f out Some in of a possible 4. 30 The from 8 males October 3 alternately. Out of a Appendix The 6 after area day an female and March Two 7. of taken to 2. males and Successive alternately, apparently days locate. removed i n the (see spring a from were One the of Removal, Simultaneous apparently missing territorial territory watching a from paired her mate's male h i s mate; i s this male, i n the f a l l disappeared, d i d not appear from The as removed females over reveal disappeared i n PART was to f o r the Simultaneous experiment. hours length one territories taken were had at only four explanations). 5 females male female difficult removed 6 f o r the S u c c e s s i v e Removal. adult juvenile 6 were had failed were o f t h r e e and proved females, c o n t a i n e d 19 the February females area Area that territory and on of (six female 7 territories yearling 29 Removal males and the at intervals for possible f o r the Control, Removal 27, Once a g a i n , hedgerow 7 7 on to again Spring 1973. t h e S u c c e s s i v e Removal possible 4 This on a hide 1972 t o have in this female). territory of a a male's The second between January thought to have taken a incident i s c o n s i d e r e d more B. 4 females Removal area, between missing were at intervals March on removed 3 the days and i n pairs o f t h r e e and 23. Two from the four days females of the experimental were removals. 31 One female and was mate in The disappeared not seen before early shortly afterwards. the before The experiments, the second and removal female remated with a experiments, deserted her Control male March. Controls Controls undisturbed i n both during the Spring and removal Fall Removal experiments. Areas remained 32 III. RESULTS 1. Replacement The Sparing R e m o v a l On the were Pairs Area Within 9 Thus, On Removed Simultaneous were removed. settled. Of the 17 days, birds had Successive removed. Removal 10 replaced pairs 7 males Removal Replacement area, males and the 12 area, 6 settled 7 females females had removed. males on 5 and and vacant 4 females territories I, during the t h r e e or removals. By Harch replaced the The Removal Fall On were male the 10 they were were until the On experiment, the 7, 14 caught spring removed. during Within :\ pairs of Removal 10 days settled. in after 10 the Thus, males the successive 10 i n the fall of with the of birds 1972, number of 4 had of females the Although i t was replacement the and removal settled. not birds females ninth several known i f or was not. not done 1 female were 1973. the replacement three or area, had S u c c e s s i v e Removal As between males assessment the 5 intervals Area paired Therefore, 27, Simultaneous October females day removed. removed. on four four area, 8 Successive birds day males Removal settled intervals on and in the vacant between the spring territories successive 33 removals. Once but By O c t o b e r again, their the After in a settling both and both the were the was of not 1972, known. deferred territories of i n both until slight and covered Fall expansion area Krebs showed who were the Removal occurred removals, (Watson 1967; met (see there was 1971). the existence prevented by t h e r e s i d e n t t e r r i t o r y of from holders. Territories of the removal Removal Areas. mated and Numbers unmated, a r e Removal a r e a s , occupied spring Successive Some experiments area both Spring the breeders Simultaneous number only was an E x p e r i m e n t a l before the r e s u l t s Fall males, and expansion Occupied Spring Increases On Of 2 gives On that potential Table by occurred, a Control on t h e s t u d y occupied in of Numbers therefore example). the replacement surplus 2. where preventing Thus, had 6 as an suggesting pressure i n the f a l l males o f t h e hedgerows i n both (see F i g u r e below), to the t e r r i t o r i a l resettling instances caught territories. 1973. length Areas were of t e r r i t o r i a l females spring of entire 9 m a l e s had e s t a b l i s h e d s e v e r a l females relation Assessment 30, territories and f a l l Removal changes there were areas in the experiments f o r of territories given. was after an the i n the region and on increase removals. of 40%. the Controls, there numbers of occupied 34 TABLE 2 Results of removal experiments: numbers of occupied of both mated and unmated males Spring Removal Area Before After Percentage change i n number of t e r r i t o r i e s Simultaneous: 7 Successive: 6 -16.7% Control: 6 +16.7% 10 +42% F a l l Removal Area Simultaneous: 10 Successive: 8 Control: 14 +40% +12.5% -16.7% territories 35 territories The after (see differences the removal differences Removal the below). experiments between areas, and Controls, in compared. The Simultaneous than i n t h e numbers the those on the i n Spring in areas and Fall occupied the and 3. The Successive Removal areas and Removal Areas, are territories are s i g n i f i c a n t l y Successive before Table and the Simultaneous differences Removal a r e compared Simultaneous between both of t e r r i t o r i e s Removal on higher areas the (P<0.05) and on the Controls. These more data suggest territories the simultaneous taken than i f settling the results settling in occurs successively. 3 • Size An Of increase necessarily In the before Territories implies present and Removal in after numbers that study, the Area. This boundaries territories adjacent removal measure of experiments t h e mean the territory areas were experiments, territorial accurate of t e r r i t o r i e s to the was due of those the size to not e x a c t l y the the the expansion Control differences fixed males areas. before mean t e r r i t o r y same Spring of that Hence and area decreases. particularly removal i s t o compare in a the had a more after the sizes. 36 TABLE 3 R e s u l t s o f removal experiments: d i f f e r e n c e s i n numbers of occupied t e r r i t o r i e s o f b o t h mated and unmated males Simultaneous Successive Spring: +3 -1 Fall: +4 +1 t = 3.13 P < 0.05 (one-tail) Simultaneous Control Spring: +3 +1 Fall: +4 -1 t = 3.13 P< 0.05 (one-tail) 37 Tables 4 Simuntaneous Controls, and the between t h e mean the Fall Simultaneous territory instance Removal sizes smaller are much There and areas, after and on Area, P<0.05 sizes on t h e after differences the i n mean the removals the C o n t r o l s , b u t i n no on both significant. support results territory the the removals smaller are and differences and a f t e r territory further mean the Area. areas settling areas, T h e mean i n the experiments. Removal are the differences simultaneous removal before sizes areas, f o r the Spring before data territory Removal Removal before. Removal These with P<0.01 sizes mean Successive territory than Successive the s p r i n g and f a l l Removal experiments give Simultaneous significant; for 5 the f o r both On are and the i n more size, than hypothesis territories i f the that occupied, settling i s successive. So f a r , the territories, the removal the doing that to and t h e s i z e s experiments. breeding males be remained so, took analyses density unmated over considered. dealt with of t e r r i t o r i e s , I t i s necessary, before during however, c e r t a i n place have and a f t e r the before however, and spring of on t h e F a l l after a s some 1973. of Removal of t o compare the removals, c h a n g e s i n numbers the winter t h e numbers Before territories Area need 38 TABLE 4 R e s u l t s o f the removal experiments: mean t e r r i t o r y F a l l Removal A r e a . N Simultaneous: B e f o r e After * t = Mean T e r r i t o r y 10 2323 14 1659 2.01 P < Size 2 (m ) 2 S(m ) 972 652 0.05 Successive: Before 8 3238 969 After 9 2878 548 t = o.96 0.2 > p y 0.1 C o n t r o l : Before 6 1745 406 After 5 2094 520 t = 1.25 * Statistically significant 0.2 > P > 0.1 size. 39 TABLE 5 R e s u l t s o f removal e x p e r i m e n t s : mean t e r r i t o r y s i z e . S p r i n g Removal A r e a N Mean T e r r i t o r y S i z e Simultaneous: Before After 10 * t = 2.69 P < 0.01 2 (m ) 2 S(m ) 3638 370 2572 887 (one-tail) Successive: Before 1890 766 After 2143 608 t = 0.60 Control: 0.45 > P > 0.3 (one-tail) Before 3956 611 After 3480 1229 t = 0.86 * Statistically significant 0.25 > P ^ 0.2 (one-tail) 40 \ 4. Changes I n Numbers Of T e r r i t o r i e s Over The the the F a l l the Removal Area Winter In on On The F a l l p r e c e d i n g a n a l y s e s , numbers Removal A r e a were d e t e r m i n e d removal territories experiments. on t h e F a l l Changes of replacement males d u r i n g the f a l l after in Removal A r e a the numbers of however o c c u r r e d o v e r t h e winter. By October territorial was spent the the 14 replacements Only Simultaneous Removal were s t i l l i t was apparent a 'natural' 'originals' from from the f a l l , non-existent, partitioned, by incorporated into the two had the t e r r i t o r y of place. of these and one was a newcomer been males, territories. expanded and of a Control events The p a r t i c u l a r 1973, d e f e n d e d 9 the Of had been o c c u p i e d i n t h e f a l l neighbouring sequence incorporation. of that of had t a k e n a l o n g t h e hedgerow; four territories surveys removal the spring In (See PART B ) . had t a k e n o v e r one o f t h e ' o r i g i n a l ' s ' gives area. a t t h e dominance h i e r a r c h i e s on t h e S p r i n g population that now 10 time that were replaced t o mid-March, I d i d n o t s u r v e y t h i s a r e a a s looking were had From 10 males had t e r r i t o r i e s males males present. mid-March, breeding juvenile on Removal A r e a . In 14 males mid-December, mid-December 17, male into, two had male. leading involved to and been Figure the (C6), apparently successfully and a 5 latter in the territory Figure 5 F a l l Removal A r e a : the Simultaneous Removal A r e a - C o n t r o l . The movement and e x p a n s i o n o f C o n t r o l male (C6) a f t e r the removal experiments and d u r i n g the subsequent w i n t e r 42 whose than size any the 14 in the the an the 10 area that the males actually 1972. In that fact, and on covered supported removal, square territory replacement of 8755 12 8 metres, Reifel their less adult territories area combined replacement males than the territories males in larger Island. occupied much far in the fall males out the before removal. On the Successive replacements were neighbouring males, 5- males. Breeding. number Fall Density 6 of gives 1972 and Area, the Despite after both the Spring pairs after. the In the The remained There 1973 that the on Removal essentially were winter. and in S p r i n g . Of of unmated spring of Fall 2 the Both of 9 of the incorporated, by spring of 1973, 7 territories. breeding experiments and into, number the fact the Therefore, territorial Removal area, over expanded a l l paired, held Table Removal disappeared territories of Sparrows 1973 fall after unprecedented Song spring of covered fall an other Thus, in was ,1973 breeding males, of the territories Simultaneous Areas, the between On the the spring compared. number the and for a l l areas. densities are pairs, the same therefore several (5) Removal number - 14 increased of pairs unmated areas on breeding before, 15 territorial 43 TABLE 6 R e s u l t s o f the removal e x p e r i m e n t s : b r e e d i n g d e n s i t y S p r i n g Removal A r e a Breeding p a i r s Simultaneous: Territorial unmated males Number o f territories Before 6 After 7 3 10 Successive: Before 6 0 6 After 5 0 5 Before 6 0 6 After 7 0 7 Before 8 0 8 After 8 2 10 Successive: Before 7 0 7 After 7 0 7 Before 5 After 5 Control: 1 7 F a l l Removal A r e a Simultaneous: Control: 1 0 6 5 44 males present. It same i s of i n t e r e s t result mortality on Mandarte of of occupied whilst the breeding previous was not females of years until As there the sex unmated males unmated males' 1 The areas, 47 the f a l l with and expansion study the roughly much t h e catastrophic i n March nearly i n 1960 a n d 1962, t h e 401 to a t t h e same 69, level 1 9 6 1 , 44 i n 1 9 6 2 . the s e t t l i n g of males the terminated the either or through found raised It juvenile number t o 69. i n t h e summer breeding through the density o f 1973, achieved the disappearance settling of females a of the on these territories. of on t h e the as explained breeding Controls, the pairs Controls territory on remained removal can the Successive essentially experiments. be accounted boundaries of Removal the The for certain same slight by the Control earlier. D i f f e r e n c e s In T e r r i t o r y Males by o f unmated that ratio, after of rose o f 1962 t h a t pairs present number differences males, pairs 1971) Sparrows d e n s i t y remained - and on t h e before 6. territories i s no e v i d e n c e balanced (1964; Following Song on t h e t e r r i t o r i e s territories Tompa Island. territorial number as that Size Between Mated And Unmated 45 An the interesting Simultaneous attract those a did that choice vacant on shortage with of or smaller from banded in the Song that Nice in March else the but does in did not territories than and not Tompa (1964) discriminate male. For there as females that s e t t l e s wherever either 1973, territories males (1943) unmated Island, of smaller Sparrows an size was several there the an is Song overall males discriminated in did not against males territories. 9 banded or Both with females is territory, Reifel mates, Table 7). female male areas on significantly (Table the of had territory Sparrows obtain Removal mate that state her a sidelight gives the fall this 1972 number to period, of non-territorial early 32 February were 1973. known to speculative and females Of be the alive 57 in mid-February. The evidence Although in the i t appears spring males with that, on females 1964). of that 1973 smaller and there that in spring of through in the even the concluded that spring of in other the i t (25) obtained settling there was circumstantial. females available discriminate i s males 1962 passive several did territories, those the were they Island, Tompa Yet, i s both Mandarte apparently females here against i n t e r e s t i n g to note that not have the fall, did mates in of new females (Tompa an o v e r a l l shortage of 1962. years of his study, there were 46 TABLE 7 Differences i n t e r r i t o r y s i z e between mated and unmated males: spring 1973. Simultaneous Removal areas. Simultaneous Removal Areas 2 Mean T e r r i t o r y Size (m ) N 2 S(m ) T e r r i t o r i a l males: mated 15 2828 1401 unmated 5 1736 867 * t = 1.75 * Statistically significant P < 0.05 (one-tail) 47 unmated 7) males that with unmated territories. males had He also smaller shows (Ibid:18,Table territories than mated males. The situation substantiate 7. The or replacement accurately study was known length in before i t The not and they Sparrows the fall the the 3 g.v.). had has i s about the experiment 20 more t h a n 1000 metres from less than a l l non-territorial, attempt to fall, as establish the parentage more at as nest, and In Song 1943). removed a l l of most local juveniles birthplace, metres. juvenile the on birds their 500 (or, defined (Uice probably the the of parents. territorial during disperse is the days that, majority bird i t s the of banded their left were 3) Some been young from of considered 1 shows not to juveniles hence is A i t replaced Appendix yearlings). lineage after aspects were j u v e n i l e s , and interval removal areas females) independence time some (see spring, of time one removal Discussion probably do but males, juveniles that disperse were in (5 Table that both gains this the origin. on question the investigation speculations. considering A l l dependent during Sparrows, during as (the dependent do birds area preceding i t i s worth birds replacement the further Birds birds. replacement more refute Replacement Finally, requires Further, males Song territories for 48 themselves and after coincidentally behaviour at until which On time the removal the Spring males Successive and It hence i s juveniles of early winter, 1972, were Results The slightly that and breeding between the are given pairs on for Experimental i n Table earlier, the pairs success the hierarchies, ( s e e PART B ) . dispersal non-territorial Introduction), Interestingly the The r e m a i n i n g young Several of the of the some of the spring areas enough, fall or i n t h e summer 6 were the these territorial preceding young the preceding as dependent area. birds. during 5 from considering with 1964), replacement the the winter one as a dependent area. during (see G e n e r a l banded fall, territorial 15 i n dominance that, paired of Removal, banded over a l llocal 5 had been Finally, than the winter on t h e s t u d y compared suggested females only banded 1) a n d t h e b e h a v i o u r males males, out •o f the study followed closely during 12 Area, involved the carried out. the Simultaneous were in 1 9 3 7 , 1 9 4 3 ; Tompa was and 2 had been yearlings (Table replacement (Nice experiment o f 1 9 7 2 , om tentatively distances out winter, were birds moult recrudescence ) , 11 h a d b e e n t h e summer replacement the Removal (10 f r o m Removal or early during during of the resident yearling fall the post-juvenile of a l l unhanded. of 1973 was and the C o n t r o l s . 8. Controls and had a h i g h e r on t h e E x p e r i m e n t a l s . laid rate slightly more of successful The d i f f e r e n c e s eggs nests, however 49 TABLE 8 N e s t i n g d a t a f o r the C o n t r o l s and the E x p e r i m e n t a l N Nest success Experimental: 10 40% Control: 11 46% N Experimental: a r e a s : 1973 Clutch size 3.50 t = 0.683 Control: P y 0.5 3.66 N Mean l a y i n g date Experimental: April 19 Control: April 17 (start of f i r s t brood) 50 are not s t a t i s t i c a l l y Tompa (1964) unbalanced, Song in however their first Experimentals unmated In were of territorial summary, a l l t h e new then, (except non-territorial replacement of local remaining replacement suggestive that species 1957; the Island could Yearlings and , t o lay of birds (e.g. Perrins Controls than of interference date 1965). and the of laying i n be a t t r i b u t e d rather highly mortality males. a n d i n t h e mean i t was f o u n d that juveniles, that origin. they some between birds, before obviously much was to interference the from males. one) juveniles was clutches, Richdale size on R e i f e l there in differences Sparrows the catastrophic smaller 1958; i n clutch the sex r a t i o by t h e u n m a t e d later, Snow the inexperience when Island, t o have clutch 1966; Song that, activities are reported Therefore, the on Mandarte breeding Coulson the found a s i n 1962 f o l l o w i n g Sparrows the significant. birds the were the replacement and removal banded were experiments. of l o c a l stock. a l l Those young f o r the origin i s circumstantial, t o o were probably as dependant The evidence birds were of the but i s strongly 51 IV DISCUSSION &. The Replacement It was necessary patterns stated on so The on on show the the replacements the study the replacement removals had The not been breeders have been carried Krebs 1971) since removal summer Hensley of of and and Cope the do and not carried by a first 1950 (1951). a that have prevented the s u r p l u s of Song breeding took capable out the bred breeding subsequent experiments and Song shows that of breeding. the subsequent Song Sparrows possibility elsewhere that i f the out. experiments i s not new. variety large birds by of a settlement but surplus of rule removal exists on a of the existed the that s u r p l u s of behaviour experiments not a breeding fact after removal would territorial 1949 and e x i s t e n c e of out the of physiologically demonstration potential Introduction existence of indeed areas Surplus investigation territorial area, they birds the there the the the area, were show on of that although for The experiments removal replacements However, the study General capable removal replacements the by And the was physiologically birds. place in territories doing Sparrows Sparrows pre-requisite Sparrows, from Song scale was of Removal animals the and a surplus experiments (references experiments conducted Stewart Many of that i n the Aldrich removal on spring in the and (1951) and experiments have 52 shown conclusively exists. The Island that present further a surplus findings add to the on of potential Song material Sparrows on the breeders on Reifel existence of such surpluses. The age determined Sparrow be of the in a few on 2§£dracjapus In (1967) and small predominated (1971) Micrptus replacements (Bendell Boss (Orians e.g. and 1970b), the 1961), and for j?eromv.scus onto an not have been found the Blue the support this Song the Tit clear; (Krebs Healey maniculatus, from but data Red Blackbird, Great plot, to Grouse, 1967) , situation i s less empty been the Red-winged that, could also Like Elliot the in dispersal removal replacements exists 1970), the experiments. mammals, experiments replacement (Orians removal has young Myers on and dispersing spp. Few which individuals i n some b i r d s , obscurus, claimed Krebs the phgeniceus, 1971). the yearlings (Watson Ag;elaius of R e i f e l Island, primarily Grouse replacement 1961; and a descriptive are individuals. drawn at Sparrows Watson similar the case in on from the studies Song have Some r e m o v a l s nonbreeding periphery and of Jenkins (e.g. Great Delius Island the 1968; s i t u a t i o n has Mandarte the a established by been the origin have shown population breeding population Young shown to 1970; exist Carrick 1963; Tompa 1964). This Wood that surplus 1965; T i t i n Bean of (Krebs Harris in some and i s 1971) , in not where 53 vacant woodland already had Sparrows local territories on on from which by the patterns study area that were was should to that be m o r t a l i t y and/or of birds, out Area, Table birds Simultaneous spent on the one Spring areas on the Removal or the known dominance areas, the Song is a of surplus are apparently derived, week o f to and April the surplus Table over which (Figure 2; 9 the further accounted on Area) on Controls. the problem Finally, area the of to appear on both necessarily either more birds tended time so was Successive moved to individuals another be (particularly the tended juveniles some Spring There areas A l s o , some majority for settle not Removal of mid-February. to may number in a l l areas. Proportionately than territorial explanation. this Sparrows shows t h e in Simultaneous The Song winter available but of banded alive investigation those emigration be h i e r a r c h y i n one i n P A R T B. for juveniles reasons. although localities. considered 82 potentially following hedgerows, certain were Removal trapping Removal the 63 of banded 9 r e q u i r e s some more the In the removed. winter for hedgerows. n u m e r i c a l l y exceed Although be that pre-reguisite juveniles to birds second non-territorial Removal mainly 1964). settlement these by replacements necessary birds occupied Island, there further the were i n adjacent disappears Tompa A of Reifel birds, apparently also territories around remain moving elsewhere to in from is congregate 54 TABLE 9 Number of juveniles banded, f a l l 1972 to early spring 1973 Spring Removal Area Males Females Undetermined 1 Total Simultaneous: 29 20 Successive: 7 1 Control: 11 10 1 22 Totals: 47 31 4 82 2 50 10 Fall Removal Area Males Females Undetermined Total Simultaneous: 22 18 1 41 Successive: 11 7 1 19 Control: 4 1 0 5 Totals: 37 26 2 65 55 at certain Spring favoured Removal Area, the junction of Out of than 82 34 Thus, there on establish is not The Temporal to show that the area Spring this be a and the On the was at Control. Area, no less locality. large number The s e v e r a l of fate locality Removal areas. The hedgerows. favoured Removal at removal Patterns removal that a removal size, i s now resulted number can be of not Simultaneous the of potential results these in birds rest of of the were able the surplus enguire areas, Removal smaller mean Territories subsequent in a more into and with given show a smaller area that size, in f a c t o r s which mean territory in significant no replacements a a mean after a successive territories, territory or a given area. could have size on the differences areas. F l u c t u a t i n g Environmental The the also smaller Removal On territories, mean to the and They smaller Successive of result necessary in Settling accomodated removal. does Of experiments greater significantly 1. appeared both The the most the banded territories. simultaneous on on along known. territory It and experiments to show single Simultaneous caught replacements B the j u v e n i l e s banded were removal the localities Resource territory size on the Simultaneous 56 Removal area could environmental be resource, related to some fluctuating on the argument t h a t t e r r i t o r y are a d j u s t e d a n n u a l l y to some environmental variable. were the case, then the t e r r i t o r y s i z e s on the Successive mean Controls and This d i d not happen. t e r r i t o r y s i z e s on the S u c c e s s i v e Removal areas and on the C o n t r o l s s p r i n g and f a l l in If this Removal areas should have f l u c t u a t e d i n accordance with the Simultaneous Removal a r e a s . The sizes territory fluctuated removals, size in opposite although before and directions i n the i n no case was a d i f f e r e n c e after a removal experiment significant. I t i s p o s s i b l e t h a t some r e s o u r c e changed along only p a r t of t h e hedgerow, i n both hedgerows are t h e r e are substantial Simultaneous Removal areas. The not homogeneous i n v e g e t a t i o n a l s t r u c t u r e , and differences in mean territory between, and w i t h i n , the S p r i n g and F a l l Removal Areas size (Tables 4 and 5 ) . A s u b j e c t i v e assessment o f the v e g e t a t i o n along the hedgerows suggested that smaller mean territory r e l a t e d to a higher i n c i d e n c e of brambles, although a n a l y s i s of reasonable in the vegetation was carried out. s i z e was no f u r t h e r , However, assumption would be t h a t a resource would parallel along the whole j u s t i f i a b l e t o r e j e c t the argument hedgerow. that a Thus, resource a fluctuate i t seems changed along only p a r t o f t h e hedgerow from 1972 to 1973. The observed r e s u l t s on mean t e r r i t o r y s i z e , t h e r e f o r e , 57 / cannot e a s i l y be environmental nature the of study Sparrows in resource, the along ) on such habitat. period numbers bewickii explained on the as the basis of food supply or Further, species of hedgerows birds did those species (e.g. that might compete for interspecific a fluctuating change in observations co-existing not in Therefore, casual a reveal any Bewick's with the was on that Song changes Thryomanes some e n v i r o n m e n t a l competition during major Wren, the resource. also apparently unimportant. 2- Pressure In his territory might be the suggestion Krebs pressure are is given (Perrins of 1965; in in Wheatear, found survived the the spring next to and Oenanthe winter on A did a not Mandarte a the relationship density (Conder number of influence Island. the time when behind this Tit between breeding density number has been also 1956) Song and Tompa of found the Tree (1964) Sparrows breeding in winter between 1965). young factor Great relationship (Weeden the rationale For determine possible subsequent oenanthe, the The 7. breeding arborea, that be and 1966). Spizella however Appendix that might s e t t l e r s at established. appears spring suggests potential juveniles birds Sparrow, the from in Lack factors (1971) being Wood, t h e r e population the Intruders discussion size, territories Marley From density that of 58 For the population it was 9). For of example, 82 Area, birds If areas an but areas conclude that, the then would the any of areas (see one might been these were the that on Table on the there settle were on both this may given. the Simultaneous of a suppose exerted No and Spring banded explained earlier, latter areas, on to reasons Controls. for these banded 50 pressure have and Removal i t appears as winter large, available f o r the by settlers, in on large number that a both the similar significant areas, pressure Removal of sort Successive differences t h e r e f o r e one from can intruders was important. It formed and Thus, the was were these, areas, so, caused occurred not of area. Removal pressure Removal juveniles Island, juveniles Simultaneous potentially be Reifel increase in territories was potential the and, necessarily on non-territorial Removal Simultaneous of banded on Simultaneous not Sparrows largest Removal more Song may be worth dominance retained consequence individual has subordinates apparently territory hierarchies them of through the the the collect to in a that the (see PART winter. dominance precedence in holder mentioning a spring. s h r i n k the B) in the vacant The and try late I t i s concluded hierarchy i s that to group non-territorial boundaries force of males an fall, that a dominant territory yearling to the birds over do not existing his territory, to 59 accomodate however, the one that birds dominant (PAST Song B), mean i t or cannot just of from to see a l l that the Successive the i t males i s not most territories competed for competed. pressure a areas, that Although resulting from reduction in Simultaneous alone the 7). stressed, imply obtained caused Removal be not individuals settlers on does Appendix i f dominant taken (It should hierarchies known potential Inexperience Of The 3- - i n the the explain group. intruders' groups eliminated, sufficient the hypothesis size the be of not the number not in i s territory (but up Sparrows Although large more 'pressure gang territories, a or Removal or the the areas Controls), considered nearly results. Juveniles And The Temporal Patterns Of Settling If changes potential then of be the settlers, cannot factor. (Southern than and Dhondt l§M£2£llH2* Chickadee, fully the (simultaneous territories 1968; environmental inexperience of settling a i n an adults Morley explain have i n the 1950), been Marsh the White-crowned (Ralph and Pearson atricapillus, on shown Tit, Great the or T i t (Stefanski and to the take Parus pattern 1967). could smaller palustris, (Dhondt the from results, territories Sparrow, 1971) , a n d pressure observed birds successive) 1971a), Parus the replacement and Yearlings resource, and Huble Zonotrichia Black-capped' Nice (1937) 60 and Tompa that there territory and (1964) was a sizes weeden , however, could significant of f i r s t (1965) found year n o t show f o r S o n g difference birds, similar between and t h o s e results Sparrows the of older i n the Tree mean birds, Sparrow i n Alaska. It was i s playing possible a part Simultaneous on inexperience sizes were although one; territorial the was section again system Successive i n this one unimportant, 3). The respect, explanation could a tendency to take than adult, but this vacancy appears vacancy may is ready to defend, in two would with initially between be m a s k e d i n however, the were territory removals, a l l yearlings areas be l a r g e r up of the i s probably than i n general, smaller territorial territory when enough This of occurred only a pattern. the replacement birds. disappearance replacements, a that, i s not apparent the two r e p l a c e m e n t subsequent were boundaries. be but i t i s not large i f the birds retention of t e r r i t o r i a l that the and i s c o n s i d e r e d a t l e n g t h i n do have single birds birds on a s mean b e f o r e and a f t e r juveniles an taken areas, t h e S u c c e s s i v e Removal on d i s r u p t i o n However, Removal the replacement Appendix on territories as a l lthe replacement apparently see important areas, t h e same b o t h once (except very the much inexperience of the s e t t l i n g i n the smaller Removal yearlings. that to be initial territory The juvenile divided tendency holders, periodically, as 61 happened i n the (catastrophic studies mortality experiments i n the masked a empty i f of From after a between large results of each neighbours, present) tendency with less to then levels of the a take territory. If and a l l the then a set of risen three to four days the ( removal) , the that hedgerow the the Tompa removal not be completely as The in the the or point four days where he as be they take have this removal could result. a tendency of can the The might be are on smaller initial low experiment, If successive his level claim with connected juveniles to in a juvenile disappeared. because simultaneous i s involved, interval(s) initial might replacement however, boundaries had replacements in a time original possibly areas, replacement perhaps smaller territories juvenile might present, therefore inclined a g g r e s s i o n , as have was a smaller territory of after suitable and successive tendency smaller territory level then or initially, ground, Nice S u c c e s s i v e Removal aggression. unfamiliar of established take to i n the The Sparrows indicating aggressive replacement study. successive his and of experiments. three a p p a r e n t l y had (if Song on male Sparrows apart), stretch removal period Song present territorial simultaneous on just removal, aggression whole one of might the 62 territory he Van den explained into did has a acquired . 1 Assent (1967) his observations, tank resulted relative in a differences sticklebacks - settling building males and were Mandarte large of resulted the 1971) that more likely considered in a larger in i s animal greater and no familiar will detail position an levels in smaller of animal be i n PART i f Sparrows on there 1965; i s with an subordinate B under in non-territorial territories Dixon of found settling Further, 1963; male process Song previously the than the than probably, of the In number Brown territories that mortality. 1949; less that experience There subordinate, Brian the showed introductions manner; aggressive temporarily (1964) of similar successively. catastrophic (e.g. a number simultaneously were introduced numbers evidence - in three-spined stickleback simultaneous larger in them I s l a n d , Tompa juveniles after a l l of the that successive introduction, the and for prior taken i s some Phillips area, (this the is occupancy i n the h i e r a r c h y ) . conclusive evidence from the study to 1. An i n d e p e n d a n t m e a s u r e o f ' l e v e l o f a g g r e s s i o n ' s h o u l d be taken i n order to v e r i f y t h i s hypothesis. One p o s s i b l e method w o u l d be t o m o n i t o r a b i r d ' s r e s p o n s e t o a t a p e p l a y b a c k a method w h i c h m i g h t be p a r t i c u l a r l y u s e f u l i n t h e c a s e o f S o n g S p a r r o w s , w h i c h p r o v e d t o be v e r y responsive to a stranger Song Sparrow's song. 63 substantiate the hypothesis initially less smaller territory. hypothesis a g g r e s s i v e , and i s ; non-territorial in the the retention, boundaries, section, time of On to to the days and inclined most be were (PART of discussed i t may evidence were A further Before considering young, adults i s territory leaving the a the factor traditional next. worth take subordinate to B). the to are supporting replacements (b) disruption, i s suggested replacement with neighbours with the the guidelines to territory to longer to f i x . days needed for total Of in This and may time the this problem interval length very of thus partly present quickly Removal that of three needed territory there were might have establishing areas, there boundaries explain of time the fact boundaries bird follow, a re-establish The Simultaneous much Disruption being neighbours. fixed On areas, as birds settling boundaries. 4. more the S u c c e s s i v e Removal boundaries such hence birds replacement. the aided settling hierarchies be however, four for a l l birds, or the Suggestive (a) dominance that may the his were have nine no taken or ten that the replacement. Territorial Boundaries i Both increase Simms in catastrophic (1965) number of mortality and Tompa occupied was (1964) noted territories accompanied following by the a total 64 rearrangement territorial broke down of system hedgerows The Simultaneous in of completely the given territory were the divided 6, Song The up i n t o area with on traditional S p a r r o w s on t h e s t u d y on t h e S i m u l t a n e o u s Removal Figure boundaries. Removal territories the Spring t h e o l d a n d new area areas, entirely Removal and anew. area territory i s boundaries shown. It were i s apparent not removals. occur The only remained new territory pattern i s i s much birds settled that pattern to such account boundaries according territories from year to a n d Snow the the Removal Removal traditional to the the the birds Song the some of of the areas. of landmarks. The experiments. considering features; i n hedgerow. f o r the retention territorial year boundaries contrast stability to certain before the Successive in birds i s preserved. and a f t e r I t i s worth the the the the removals, topographic that of retained, that system territory the i n the Successive possible (Davies along might replacement arrangement before areas. structured philgmelgs, i n the after was the territorial section t h e same Removal due the boundaries explanations territorial 6 that of the territory pattern Simultaneous It by narrowest territory Although the the Figure similarities 1 linearity The areas influenced along general from territory habitat f i x i s their The s t a b i l i t y i n Thrush, 1965) a n d t h e E u r o p e a n Turdus Blackbird, Figure 6 S p r i n g Removal A r e a : b o u n d a r i e s b e f o r e and a f t e r the removal . e x p e r i m e n t s on the Simultaneous Removal a r e a 66 T_«. m e r u l a , (Snow 1958) habitat. Song Sparrow presumably the birds themselves and boundaries, however, interactions directly to occurs, to birds replacement territory area. and after schedule, been the retain one fact i n an some s e d e n t a r y their of i s not traditional of the related how how into i t with boundaries deduce that interval of passerines on throughout would of the for the males the Removal and wait vacated d i d not areas, four boundaries of four the existing boundaries a three a maximum area. along takes into territory that, males the the Successive conducted before Sparrow the adjacent the t e r r i t o r y on settle linearity long When a the vacated Song the long their were can male to territory individuals. bird familiar that territories product neighbouring owing known t o expand can two ,become the removals then the to encroach only or between recognition and and habitat. the experiments determined In to boundaries, attempting that not a themselves, a replacement areas, i s bird From change likely replacement with The the t e r r i t o r i a l begin study, It by new Removal hedgerow. before a contend Successive the b i r d s respected the present have most defined, boundaries neighbours. i s to features of the are c l e a r l y the explanation i s that neighbouring In attributed recognize features of the i s vacancy been territories their between A second pattern have day had days. like t h e Song the year, as Sparrow, i n the 67 European Robin, Erithacus Wren-tit, Chamaea fasciata, activities of newly territory boundaries, former boundaries behaviour has M££2£fii£Ki# the newly show such areas, been (Young This could further to It in the same explain one of to explore Similar on On and Reifel the first the of the exploratory Skua, to Catharacta argue Island on that would Simultaneous male, the the re-establishment replacement would is Mccormick's Sparrows one 194 3) seems r e a s o n a b l e patterns. the birds neighbours. Song any at 1938), leading 1972). boundaries, same t h i n g (Erickson described settled however, (Lack settled with behavioural territory the the rubecula, also Removal exploring find other time, with no guidelines to follow. length of time to fix the replacement males the taken doing the boundaries. 5. Genetic It Klomp Basis i s not known (1972) that evidence that size i s (1962). probable lends Of Territory territories the problem support individuals of fixed Klomp territory that genetic far genetically indications their how Size i s genetically fixed, However, that The discusses territory that For the the make-up. to from states size has a same s p e c i e s of the be an gives Schoener of (1968) basis, of this are based size of circumstantial although genetic compressed. optimum existence that, variations Most of and the can an and i t is there attribute on differences optimum Gibb highly are no among differences in seem of to be 68 a modificatory a territory one generation of lineage particular tested the of a are the C. large to a i s not the after territorial The on immediately the pursued a have but, territory into For the could Different a be i s that the from appear one to brambles, and involved. The to be drawn, Results Of The Experiments not show a or mean in significant territory replacement and Each the occurred traditional replacement defend because area, obtained here. basis, holders a to on conclusions did to of of few territories a l s o be because perhaps vacant small territories, prepared too from examples male passed on parents impression Removal areas only the are i s The retained. been which data passed the particular firm one-to-one was in further removals be incidence Successive of may gives general high Removal pattern territory, neighbouring The f o r any number the initially vacant that However, Explanation And 10 territory i s a sufficient successively may next. Successive in but size area, and small there matter The size, known, or the Proposed change study vegetational factor not Simultaneous Table f o l l o w i n g year. where therefore and the are particular next. statistically, taken data a the from generation be of to male territory taking nature. of only the a juvenile of the reluctance of their part to expand boundaries and p o s s i b l y because of the 69 TABLE 10 Generation 1 L i n e a g e : t e r r i t o r y s i z e s from one g e n e r a t i o n t o the n e x t Territory size 2 (m ) Generation 2 Territory size W/0 2708 R/BL 1824 W/0 2708 LP/BL 1444 W/DP 1625 Y/LP 2166 DP/M 1264 M/W G/R 5054 B/B 722 3249 2 (m ) 70 proposed exploratory boundaries the of the replacement behaviour territory bird, of of the probably the new removed in a bird, bird matter the were of size and taken by three to four days. The Simultaneous significant changes Removal i n number territory sizes. The completely broken down, entirely with anew, the the replacements argue that were possible It levels immediately the were factors that of claim removed a size adults. attempting to settle, territories the summary, study area, physiologically then, of with a the capable size the have of a birds surplus breeding. days). A l l reasonable the less to low a l l to area taken. had likely to that of level of simultaneously in a larger territory experiments the initially were resulted formed territory temporary s m a l l e r mean removal was ten was f i x with of mean comparison to equivalent birds, in were In i t seems hence territory could there the and replacement taken, and to showed system taken (nine replacement Thus, the time unfamiliarity aggression, and boundaries the birds, hand, territorial longer affecting the of In much other guidelines. areas, young aggression of apparent took the territories, territory i n e x p e r i e n c e and/or i s postulated lower and without boundaries of on traditional S u c c e s s i v e Removal territory areas, number size. showed that, on of birds that were The replacement birds 71 were almost a l l j u v e n i l e s , and were probably a l l of local origin. The show successive significant territory smaller account changes sizes. territories explanation The resulted mean for pattern i n numbers results i s given. numbers sizes. are on t e r r i t o r i e s of territories, simultaneous i n higher territory these of settling pattern of Possible d i dnot o r i n mean of settling territories, f a c t o r s which discussed, and a on and might proposed 72 PART I B DOMINANCE HIERARCHIES INTRODUCTION Schelder'up-Ebbe (1922) biologist to recognize the in of animals. groups peck-order He i n domestic or subordinate situation was essentially then, species Wood-Gush 1965) been and vertebrates, Witter 1969); Cynomys in laboratory mice, primates, e.g. spp., Leopard Krebs 1973). (e.g. Ito 1970) be Lemur Devore Indeed, to both wild of a linear was that animals, the one and Frog, either the dyadic Rang of 1965); have the the Mantis and most Cockroach, 1969); the the and and (Jolly hierarchy the hierarchies e.g. e.g. in 1953; pipiens, (McBride (Levick observed (e.g. Guhl Caldwell 1955), spp., been domestic and (Anthony lemurs. first hierarchies bird have vertebrates, musculus, and each o t h e r , and 1970), truncatus, Mus (Hall For higher Tursiops existence hierarchies (Dingle ludoyicianus, Dolphin, the the irreversable. (Ewing the being dominance invertebrates, bredini, as of that any animals, in e.g. such to wild. cinerea, Gongdactylus Papio of documented Nauphoeta discovered dominance numerous credited importance fowl, dominant Since i s Shrimp, in lower (Boice and Prairie Dog, Bottle-nosed Hebb 1948), and Beilharz 1973); in 1966), i n man, and baboons, (Sommer 1967; i s c o n s i d e r e d by important concepts some in 73 animal social There i s hierarchical been found herons behaviour. wealth i n such diverse Worm 1971; B r u l l , (Bennet 1939), cited parrots i n corvids in (Brian parids fringillids Wessell under half possibly Brown sentiments imposing when social Several winter area flocks from although subject Gartlan the of birds in the function of Wynne-Edwards much 1962), 1952; L o c k i e 1956), 1942), and in 1964; S a b i n e 1939). which the that however, animal's also care should conducted outcome in a (e.g. the consequences taken Hence, hierarchy were o f geographic there hierarchy. Lack i n units. different the o f t h e dominance be (see similar i n the wild winter. of behaviour expresses are not s o c i a l they discussion various i n laboratories, a factor that.breed on t h e f i n a l and done studies which 1934b), pigeons were (1968) o n what (Watts and 1967), Odum studies, influenced models that information bird have tree-nesting 1 9 3 3 ; Tompa 1941; Shoemaker he s t r e s s e s of 1966; 1954; Tompkins conditions, 1963:460). a n d Hogan 1963; L o r e n z on relationships lek species and A l l e e Dixon material colonial 1938), (Masure o f these have These as by K r u i j t 1949; and Leigh unnatural could observational groups (Brown (Tordoff Almost of and Schmidt passerines; 1959; v relationships i n birds. (Noble, Stokes a ^ i s no Indeed, has been t h e 1954, are not well 1966; understood. 74 ( A few studies hierarchy through spring. Odum flock the the status of eight Mandarte alpha, Parus four beta, two alpha birds territories, of birds I therefore opportunity the dominant during the on Reifel to at the fall, area had been present territories. of Finally, by l a t e chance f o r question; obtain in neither established of and associations Song experiment their i n the along the Sparrows enabling Song opportunity localities fall, when t h e territories population loose several o n how spring, and an e x c e l l e n t of the o f two difficulty hierarchies favoured t h e removal the hierarchies, formed proportion In on territories. The the at special no d a t a no that presented in Sparrows two o f t h e b e t a season? colour-banded a had in a i n Carolina was c o m p o s e d the following The j u v e n i l e s A large recognition. would nature o f Song i n obtaining males determined the although i n dominance Island and c o n g r e g a t e d hedgerows. but only pre-breeding (1964) are given. are the juveniles individuals consequences. birds, i n the territories results t h e group apparently investigated Sparrows look that succeeded arises, Tompa determined establishing t h e gamma similar i n a group a n d t w o gamma were the top three established found males t h e dominance i t s breakdown that carolinensis. and found relationships the (1965) followed until reported juvenile Island, have Chickadees and Dixon Chickadees, birds winter (1942) of Black-capped spring, the in i n the individual i n the spring yearlings to obtain 75 II The PROCEDURE Spring Removal hierarchies were Feeding male's set millet, each stations they measuring by barley the (about three stations, principally left early flat which the adult tables were of of yellow worst a after December, was put out a t a s t a t i o n of other amount Sparrows, to of on the seed-eating spp., eaten by close to a feeding one feeding passerines, Pipilo Passerella iliaca. day and bait any Towhees, a t t h e end o f t h e at (see monthly Only attracted white intervals weather small plywood, and 2). Fox presumably i n each pieces mainly Juncg and up put out a t i r r e g u l a r a number at the station morning, along Nineteen (a m i x t u r e readily Juncos, erythrophthalmus, food were were of the winter's Appendix set 1972. out was handfuls) as hedgerow were the winter, of Sparrows of Bait oats) period Song made throughout summary. day. (tables) 40cm. and weather the only i n the f a l l were 60cm station after was determined. territory up; Area had Any gone by rodents. i A hide station effected was by recorder activities given was under the (Sony erected observation. The presence the 110a) and at the station; of a birds i n this to t h e i n t e r a c t i o n s between A were way, when d i d not appear hide. microphone station used full the b i r d s . that t o be cassette to record attention tape the was 76 All interactions Interactions feeding stations encounters between and individual were two a causing retreating displacements, below. be Appendix clearly noted at the 5). A l l took place and i n a l l e n c o u n t e r s to retreat was recorded. birds invariably a time, as divided avoidances, A i s feeding Bird flight B at Bird 1943:156) determined. was noted as a The dominant; subordinate. into three categories: and f i g h t s . beak directly either at i s made. dominant bird On A, being i t would Distances ft. driven or t o the ground displaced. be d r i v e n between Should o f f by B i r d the birds threat and Bird determined B below, (Nice has wings lowered, flies would another usually f l y depending a subordinate and o f f before providing A ground posture o u t , head off, the a rapid invariably present. on Bird fluffed who or makes in Bird i s not already been hops feathers Bird to the table had j u s t at body table and e i t h e r A, o r q u i c k l y pointing contact at the bird appears, •out-of-pocket*, then at were other (see Sparrows the other were and noted could individual Sparrows Displacements Bird it also Song loser Song Sparrows birds Encounters (a) Song involving only winner the between between be on where present, A. varied from one-half to one 77 metre, (b) and appeared to increase A is feeding the table appears and hops below. Bird area, whilst adopting •out-of-pocket*. 1943:225), down, and Bird A hops raised away one from once mild A approached. assumes out the an up. feeding more o f t e n , towards body As the posture, alarm feathers, and again or, threatening crest stretched Distances roughly a Bird with neck B at ground on feeding with wings posture feathers Bird B the (Nice sleeked approaches, area. between birds varied, directly facing each but were metre. Fights Two have birds tails 'square out, and quite close to Two generally close bill each remain seconds. and body frequently often off*, fanned •out-of-pocket', and spring Avoidances Bird (c) as open. other, motionless possible to in i t s bill, turns the other in pursuit. fight. through They the shrubbery, spring into 20 are i t s head, There and the (2) air Both depressed, ground, two these outcomes closes chase The about bird close the other. feathers birds to 30 wings generally centimetres postures for then then the flies follows birds together, a are apart, several distinguishable: and puffed (1) off one with prolonged physically pecking, and 78 attempting The two to birds ground, through the Each fight) the the of that Sparrows in gives by to twelve and the surmount feet flies victor the and above the with the off, a other. prolonged chase encounter equal (displacement, weight were at the combined. did analysis, This not and that the encounter was not in interactions seemed involve time, the a in avoidance, and more the juvenile than as doubt. some o t h e r a l l three justifiable decision in and to two who This on was is fringillids in (e.g. 1959). mentioned congregated at earlier, special favoured These localities frequently male's territory, and feeding stations observations of would inevitably and the female. group involving certain involved individuals of the feeding in Interactions juveniles adult stations group were more more than one than females and only at as the the one one adult were adult different Further, different combined, hedgerow. juveniles. than involving attracted of analysis. over with more Sparrows the observations same the Song along encompassed ranging interact different localities therefore were combined group and as way interactions an to As male trying much pursuit given grounds Sabine as one encounters contrast each underbrush. type victor claws, rise close was of Song can before accompanying types clench These the territory territorial adult males interactions juveniles. adult same male Also, and his 1 79 female was during the observation and hence no hierarchy resolved. Matrices (see period, were Results). during the particularly constructed The outcomes spring they for each hierarchical o f t h e h i e r a r c h i e s were a s t h e h i e r a r c h i e s began were followed experiments presented an o p p o r t u n i t y members of the h i e r a r c h i e s to obtain followed t o break-up, and i n the removal removal group areas f o r some territories. where t h e of the 80 III RESULTS (A) The Hierarchies Seven and is matrices included The ranged time bias due i n the text i n the number from amount to the fact over an feeding area down group involving one. the The 7 ) , and t h e involved to five hedgerows, B-C Hierarchy remaining the group i n the six hierarchies individuals. of birds spent incorporating i s given. The amount The heavy on t h e B-C Hierarchy i s partly of birds concerned wandered three territories, As n o t e d earlier, t h e same group a l l hierarchies, Table (displacements, an 1153 example, and observations of birds fights, t o be males) were 11 t h e B-C and shows hence three at different were combined birds only at the in numbers was 9% in based were encounters top more of of in four hierarchies. on a grand On than the involved encounters avoidances, displacements. i n more involved the Hierarchy the rest of i n t e r a c t i o n s and f i g h t s ) o f which involved Hierarchy, the majority avoidances, interactions, tended B-C each that displacements. ^% (Figure along analysis. In As f o r each Sparrows of time stations. stations the o f Song with determined Appendices. twenty-seven spent in were were c o n s t r u c t e d represented are of hierarchies total less t h e whole, feaales. hierarchy interactions than birds of than males In the (mostly lower 81 Figure 7 B-C HIERARCHY LOSER 9 tf tf tf LP G Y w® 0 • G R Y 4 5 7 22 13 7 3 6 2 7 7 6 5 8 1 <f tf B 8 G B8I R YR W R ROB 8 B \ G B Bl W R \ R Y R R DB 6 LP 2. Y £ OG R T w BG 8 BS tt 3 Y W ® Y 1 tf w 9 w© R® B RW LP 6 11 1 4 11 2 5 5 16 11 5 '6 8 2 8 1 2 2 1 5 1 3 7 2 5 5 3 1 11 2 22 2 3 24 1 1 5 17 3 1 15 6 5 5 9 3 10 11 9 9 16 6 6 \ 10 2 16 6 5 2 6 1 2 1 \ 1 7 \ 2 BG B BG BY BY 6 11 3 4 4 1 4 13 3 4 5 1 1 34 3 14 9 17 \ 5 \ 1 2 4 BY BY c 1 1 2 2 10 tf \ \ W © R ® 5 1 1 B RW LP 1 BO 6 Y® 3 9 tf tf tf 5 1 1 9 80 tf B y<B BY \ 1 10 9 9 G \ 1 Q© 1 G OB DB 1 6 13 2 13 4 6 5 3 4 10 6 2 8 4 6 2 9 6 18 1 12 2 1 5 3 18 4 20 1 3 1 2 6 6 3 B 08 IS. R 8© 7 2 1 3 10 12 5 1 1 3 4 2 6 1 14 2 20 1 2 1 3 1 1 3 2 3 1 3 8 9 1 4 2 1 2 4 16 7 \ 2 1 4 \ 7 1 2 2 6 \| 1 7 1 AVOIDANCES FIGHTS 3 1 1 2 1 1 1 4 1 1 0 B© B© 2 , 9 9 G© B® 1153 I N T E R A C T I O N S 80 M A N H O U R S 9 JL. LO DB R 1 B BY 8© 9 B® 3 7 9 \ \ \ 82 TABLE 11 Numbers o f e n c o u n t e r s ( d i s p l a c e m e n t s , avoidances and f i g h t s ) i n f o u r dominance h i e r a r c h i e s Hierarchies B-C Sedge P a t c h Bulldozed Path Cottonwood S t r e t c h Displacements Avoidances Fights Total 1043 101 7 1153 142 41 3 186 160 11 3 174 153 17 0 170 83 down (mostly fights and in the very the B-C seventh slightly (Table females) more 9 below). H i e r a r c h y , s i x were male-female. involved the male-male However, i n avoidance Of seven encounters, males were interactions than only females 12). All seven hierarchies few reversals involved corresponds with the not the (1922), and (1934a). Nice she Sparrows pass clear-cut peck-order Reifel was this should of by period early after I t seems been to As December, likely that young Song Song to three old. months Song linear amongst one more might a Sparrows 'peck-right* observations then allee peck-dominance established winter. and juvenile juvenile stable B-C arrangement on that are the the Masure through they are Schelderup-Ebbe observations time There in of of of concluded then, have type * peck-right* from the time Island, many r e v e r s a l s . This supercedence by linear. a l l interactions •peck-dominance* raising, started of classical relationships hierarchies strikingly reversals. from juveniles 4% (1943), Sparrows Extrapolating are ; Only 1 Hierarchy on (see F i g u r e the on groups the not expect reversals would 1. In encounters between a n y two b i r d s (a a n d B ) , a w o u l d p r e d o m i n a n t l y b e t h e w i n n e r o v e r B. O c c a s i o n a l l y , B would win s u c h an e n c o u n t e r , a n d t h i s w o u l d be c l a s s e d as a reversal. In the matrices, numbers of reversals appear below the diagonal lines. 84 TABLE 12 Analysis of the avoidance interactions i n the B-C Hierarchy Won Lost Total Males: 83 26 109 Females: 18 75 93 Total: 101 101 -% 2 = 64.74 P < 0.001 85 have been recorded b i r d was being Reversals often. e a r l i e r i n the winter, decided. d i d not i n v o l v e i n d i v i d u a l s t h a t were seen l e s s In the B-C Hierarchy, involved in members of reversals as the rank of each those i n d i v i d u a l s (9) that were more than one r e v e r s a l were a l l prominent, d a i l y the hierarchy, when under observation. Also, d i d not tend to occur l a t e r than e a r l i e r during the observation period. 40 man hours before Out of 43 r e v e r s a l s on the B-C February 12 showed 24 Hierarchy, reversals, 40 man hours a f t e r February 12 showed 19 r e v e r s a l s . •Triangles', dominates A, frequently the where and dominates B, B dominates C, and C relationship observed e. g. i n domestic fowl Juncos A phenomena (Masure and remaining of stable, dominance Allee 1934b); stelleri, itself. (Odum 1942); Jay, In Song Sparrows, t r i a n g l e s occurred and remained s t a b l e during Brown Cyanositta the h i e r a r c h y only, period. middle of the a d u l t male and h i s female were the dominant b i r d s a t the f e e d i n g The the B-C with both sexes i n v o l v e d . a l l hierarchies, territory. only i n the the o b s e r v a t i o n was found t h a t t r i a n g l e s i n v o l v e d b i r d s i n In Oregon were t r a n s i e n t , the l i n e a r s i t u a t i o n soon a s s e r t i n g Hierarchy, It in (Kikkawa 1961). (1963) found t h a t t r i a n g l e s i n the S t e l l e r ' s a hierarchies, (Sabine 1956); i n Black-capped Chickadees i n White-eyes, Zosterops l a t e r a l i s , are only time station that located pairs from in their adjoining 86 territories f e d a t t h e same s t a t i o n placed on, pairs, as or very The the the Cottonwood respected ranging might of each table was s e p a r a t i n g t h e two Stretch territory hierarchy i n numbers from determine Hierarchy. boundaries, and was composed two t o t w e n t y - t w o . t h e dominant or subordinate of Factors status i n hierarchy are included i n the discussion. Although intact, did the some (Sabine Some maintained of juveniles of individuals of Chickadees et i n v o l v e d b i r d s o f low rank Tompa (1964) were involved low mentioned This i n position that only was n o t t h e c a s e Song same s t a t u s i n t h e new old one. These Song groups of low which joined. Sparrows on Mandarte on Reifel rank of middle wandering group as t h a t they Sparrows movements, b u t a l s o i n d i v i d u a l s 13). Wood birds intergroup (Table and o n l y , and t h e s e between individuals Juncos group Not ranks a l . the 1966), Island. upper did 1965), subordinate for i n I n Oregon i n t h e new g r o u p interchanges only involved (Dixon (fiurton relatively between h i e r a r c h i e s a l l juveniles palumbus, their remained i n t e r g r o u p movements. Carolina Columba interchanges Island. 14% performed 1959), Pigeons, groups interchange occur. hierarchies the the d i d not occur. remainder juveniles, which i n adults trespassing when c l o s e t o , the boundary happened Otherwise, was perform and even b i r d s assumed t h e they Most examples a r e o f low ranked had held birds, in and o f 87 TABLE 13 Rank i n h i e r a r c h y and sex o f Song Sparrows p e r f o r m i n g intergroup movements Sex Hierarchy position Top t h i r d Male Female Total 2 0 2 Middle t h i r d 2 1 3 Bottom t h i r d 0 5 5 Total 4 6 10 Each h i e r a r c h y was d i v i d e d i n t o t h r e e s e c t i o n s i n o r d e r to e l i m i n a t e the d i f f e r e n c e s i n numbers o f j u v e n i l e s i n each h i e r a r c h y . The a d u l t males and females were n o t used i n the a n a l y s i s ; hence o n l y the j u v e n i l e s were n u m e r i c a l l y m i d d l e , and bottom. d i v i d e d i n t o the t h r e e c a t e g o r i e s , t o p , 88 females the in particular, Bulldozed early February mid-February Stretch male, male Path high was in a An shows of that fights the analysis hierarchy than of and the gives the results There i s no of to was are run more of the in the interact than hierarchy Success in birds January, B-C was the found in and in Hierarchy, male in the 1000 only a was Cottonwood metres. dominant territory in This juvenile (see (B) The (Table 14) ). the B-C Hierarchy are closer together more likely to be apart. B-C Hierarchy birds those farther the involved in Therefore, a (displacements, to closer in investigate together apart. in Figure the 8 analysis. correlation are about that interactions. together of {Red/Yellow) a l l interactions the than the farther on in in in that that late obtain fights possibility interact (B) the fights) v hierarchy "numbers enough, not in male dominant significantly analysis further the Hierarchies those avoidances, scale as individuals are complete the hierarchies Consequences dominant distance interestingly the the Hierarchy found Hierarchy, in but between Thus, distance birds hierarchy which are not necessarily which are farther Gaining in are more and closer likely to apart. T e r r i t o r i e s in r e l a t i o n to Hierarchy Position Of the seven hierarchies (Figure 7 and Appendix 8) NUMBER OF INTERACTIONS 10 CO o o —I— _1_ J>0 Oo o o • >• • MOOo DO D X 4^J>t> >&•• • ipt> O D D C J O o • J>o oo o• > " • •a a o• oo-Jpaa (tOOO • a* $00 "D a ro co b 00 x + o CD CO o o o o•o o o o -< 11 1 p p > oa J>a oo < > J Oa o o d> • o o JI o o o <po oo o o o o > X o • DO N)JP O mm> > 2 s r r > > m rn mm .< .< < 5: "n 2 • m> 5 >>m ? "m > m r~ m r r Figure 8 Number o f i n t e r a c t i o n s v e r s u s I a l l j u v e n i l e s i n t h e B-C H i e r a r c h y distance within hierarchy fo 89 TABLE 14 Analysis of f i g h t s i n the B-C Hierarchy Number of juveniles i n the B-C Hierarchy = 22 Number of f i g h t s observed = 7 Random sample s i z e ('pairs' from Random D i g i t s Tables) = 70 Distance i n hierarchy* 1 2 3 Numbers of 'pairs' of b i r d s : Observed 1 0 1 2 Expected D = 0.76 P .3 .5 .2 4 .3 5 6 2 1 .7 .4 7-22 0 4.6 0.01 Kolmogorov-Smirnov one-sample test * Distance i n the hierarchy refers to how f a r apart i n numbers one member was from the other member of a 'pair'. Thus, i f the distance i n the hierarchy i s 1, then the two birds involved i n a f i g h t were next to each other i n the hierarchy. I f the distance was 5, then there were 4 other birds i n the hierarchy between the 'pair'. 90 determined, Sedge four Path) Narrows) were was (Big Alder, i n the Simultaneous i n the Stretch) was the Simultaneous and area (1) incorporated Simultaneous Removal explained i n PART the Simultaneous holders the had been area, The area i n the one (Marsh and seventh one (B-C) and t h e C o n t r o l Simultaneous met, Removal Area A, t e n m a l e s Removal removed. area established after Of t h e s e territories the resident t e n , seven territory had f i g u r e d in hierarchies. The dominant territories: second males obtained Hierarchy The Hierarchy males i n obtained a dominant male stretch hedgerow. was c o n s p i c u o u s member this male was when the removal a n d B-C the f i f t h and B-C. The Hierarchies also male i n the B-C territory. the only prominent Big Alder, Path and f i n a l l y Removal of Patch, the Bulldozed territories, was i n the following hierarchies obtained No-Name, S e d g e Simultaneous male Removal two t e r r i t o r i e s area, Removal was o n t h e C o n t r o l . No-Name, a n d and one on t h e C o n t r o l . As in Path, Removal Successive (Cottonwood where Bulldozed top bird area i n not t o obtain by i t s wandering not present in the As e x p l a i n e d of three took (Red/Yellow) hierarchies. along place. the Bulldozed hierarchies a territory earlier, this behaviour, on the along this particular having been I t i s conceivable the Simultaneous Path Removal a that area 91 Three part on males that established i n the hierarchies. the Control, and One was simultaneous removal occasionally throughout Removal area, b u t was during the time unbanded male hedgerows quite the study males position of Island this join made. of the seen Simultaneous feeding The station last was i n the region the study individual December was the any territory that paired the removals, only one in the February Table interactions in seen on January the adult 15 This Sedge 3. spent shows area, and the winter 29, female's male and and date, an where i t i s outside after February during replacement (2) S u c c e s s i v e Removal i n early the removals, member had a low Hierarchies moved difference that territorial prominent she r a p i d l y i t i s thought position a B u l l d o z e d Path of the Bulldozed Path the one lost t h e new (Black/Blue) a significant and area this with had ^ e e n female Patch After won the the adult Only along x t h e week second at females female of recorded a until The winter established hierarchies. scale. in banded take area. after before was o b s e r v a t i o n s were that the seven the the not flestham possible Of in from d i d not not r e s i g h t e d experiment. that that of thes territories up the i n the number The adult 3. H i e r a r c h y was Black/Blue assumed February. On she with paired last the death one of males. Areas hierarchy was determined along the Successive 92 TABLE 15 Change i n s t a t u s o f a j u v e n i l e female f o l l o w i n g d i s a p p e a r a n c e o f an a d u l t territorial B e f o r e Feb.3 Interactions: X The a d u l t 2 the female A f t e r Feb.3 Total Win 2 23 25 Lo*se 11 5 16 Totals 13 28 41 = 17.09 female was P < 0.005 l a s t seen on January 29. J u v e n i l e ( B l a c k / B l u e ) presumably female assumes the a d u l t female's r o l e i n e a r l y F e b r u a r y . The j u v e n i l e female wins s i g n i f i c a n t l y more i n t e r a c t i o n s , not o n l y w i t h b i r d s below h e r , b u t p r i m a r i l y w i t h b i r d s above h e r ; hence h e r change i n s t a t u s . 93 Removal were area. juveniles, succeeded other female winter across with which Ewen birds, females. a new male One after It i s possible were Slough of which unbanded, i n the F a l l of these the that two females removal, the could Removal only the replacement have Area spent the (Figure 1). Control 11 Of Hierarchy, In corpse this juveniles that took none were seen after May, however, early was found on territory, replacement The B-C the only amongst five have and male had 4). one males obtained given territories hierarchies the are one 5, been third took bird more chance part established than This (the taken ever female. This Hierarchy. of the hierarchies, one territorial hierarchy, i n the h i e r a r c h y , i s highly four and male out of 12 from the unlikely to 0.01). strongly opportunity, the had the adult Analysing this (P = then, a male males d i e d i n t h e B-C the largest Stretch experiments. of the adult May territories. evidence, i n the Cottonwood the removal On supplying that part apparently also H i e r a r c h y was o c c u r r e d by The that, May the replacements. juvenile top of of five both disappeared. most was and in pairing birds, (3) I t consisted the supports juveniles dominant individuals during the pre-breeding the hypothesis that in obtain dominance season. 94 IV DISCUSSION (a) The F u n c t i o n a l S i g n i f i c a n c e of Dominance H i e r a r c h i e s Gartlan social (1968) suggests dominance available can subordinate although food function when l i m i t , regarded the (1962) suggests a social guillotine times of as whilst food that for shortage, few starve or numbers of the s p e c i e s are up a g a i n s t the bottom of concurring to the the the f u n c t i o n of to r e l a t e p o p u l a t i o n s i z e during (1966), i n d i v i d u a l s near is Wynne-Edwards individuals Lack emigrate hierarchy resources. hierarchies that the primary the that peck-order t h i s as a consequence, not a function. Lack argues t h a t the h i e r a r c h y a r i s e s as a response to s c a r c e food resources, and Wynne-Edwards, pre-adaptive not however, however, differentiate the food that the is cause shortage of hierarchy frequently and effect females! food shortage, than males. The is guillotining both difficult of I t i s even more hypothesis supporting the to dominance difficult to with the h i e r a r c h i e s i n Song Sparrows, where a l l the bottom, hence subordinate are them. privation. t h a t can be put forward it between anticipation maintains h i e r a r c h i e s i n most i n s t a n c e s . relate an f o r c o n d i t i o n s of There i s evidence arguments; as of subordinates birds, i n times then, would presumably e f f e c t females f a r of more 95 A further reducing aggressive conserving and aggressive Calhoun states that the and food because i t saves lose with anyway, elsewhere. between the are and animals. the identify that a might any one of (b) Factors wasting energies are on can Sparrows i s as the to not determining fights various are for he both they those lower they would for that searching aggression because of rare, reversals like ways functional reduce aggressive determine territory for reduced behaviour, Sparrows, to been when stable. in be has Guhl because which argue 1955; g. lines conserved to reducing (e. value fights be reasonable s u r p l u s , or them survival time function could a similar and hierarchy obtain birds fighting, little hierarchical hierarchy along has particularly Song domestic in is Anthony very Song could (e.g. i s a There important mammals a l s o argued in thus this hierarchy. are on that higher seems For and the argued f o r those It the on be function individuals, individuals, their Dominance behaviour, 1971), and hierarchies, few, i n work them individual between i t can peck-order weaker obtain again hierarchies f u n c t i o n , of (1954) that that hierarchies Kinsey Lack stronger a encounters 1971; 1956). but not evidence is encounters energy, consequence, good argument given the territorial for different significance of interactions, to dominant individuals the opportunity. the Dominance Data unequivocal. position in Hierarchy for 96 It has Sparrows already been i s essentially considering dominance some of shown linear the that the hierarchy and s t a b l e . factors or subordinance of the which I t i s might individuals i n Song worth now determine the comprising the hierarchy. Sex and p o s i t i o n In Song juvenile 22 is Sparrows, females. juveniles, arrangement (e. i n hierarchy g. from c a n be o r i n Steller»s takes the rank M£ii£2ia2[» are graded of i n some the male males. Black-crowned pairs this species 1942; B r i a n where the with. female In c e r t a i n Nycticorax 1939) , m a l e s sex, of 1955)), i t Night-Herons, and Schmidt to passerine (Tordoff over out Although 1 9 6 6 ; Odum 1964), she dominant f o r example, other (Dixon (Brown regard were curyi.rostra, species Jay ( N o b l e , Worm without i n Loxia i n Parus 1949), as t h e t o p 14 w e r e found Red C r o s s b i l l , instances, males I n t h e B-C H i e r a r c h y , 12 not the case juvenile and females suggesting an asexual fowl that cocks situation., Guhl normally two (1956) peck hens, hierarchies, Sparrows involved An found cannot for domestic and t h e r e f o r e one f o r each be d i v i d e d a breeding flock sex. i n this The usually hierarchies manner, do n o t has i n Song sexes were by p a i r i n g with as both i n the interactions. individual can climb the s o c i a l ladder 97 a dominant instance pairing of of a with juvenile adult bird a male female had mate, moving scale the female. disappeared paired this with after the the up the the in adult spring the replacement young Size and position hierarchy (1963) of the relates dominance Song hierarchy period (see male on removal the an after Sparrows, after January interactions period with in In in success after and, up adult Her significantly his Brown ( B l a c k / B l u e ) moved 3. apparently opposite sex. dominant female February the a the 29 to increased Table 14). She disappearance experiment, she of paired male. f Size has hierarchical Hebb been found position 1948), and Forest (1969) found that, i n the wings than subordinates. weight (Figure and relationship between hierarchy was found. largest or heaviest be a factor in Bottle-nosed i n New 4) to or i t birds 1972). vary were the be longer considerably 3) , and ones but position argued and Fretwell had (Figure winglength cannot (McBride dominants Sparrows winglength weight Thus, (Tyler Junco, Song in Dolphins Ponies Oregon in determining that that in no in the obtained territories. Age and position Age domestic passerines and fowl (R. in hierarchy dominance have (Shoemaker I. been 1939), Stewart pers found but not coram). to correlate in A in short-lived more subtle 98 relationship Grouse, is a Centrocercus centripetal mating centre In of between Song a t most o f t h e ^Lek^)as Age juveniles the first prior movement two a d u l t the birds were of only dominant over could evidence, hierarchy. T h e a g e o f t h e two B-C Hierarchy determined (see second was was bird the calculated of he 6 was those the banded when respect to brood, a second younger or third the ideas of that, position males i n in the i n t h e summer age be of age). The juvenile top brood, the of could but the or third than from next. calculations dominant rest to suggest their a comprised second with meagre, were of a f i r s t 60 d a y s from i n determining for either t o be a b o u t Nevertheless, as they Appendix product i s with top juvenile development a product towards i f the individuals albeit the of there the years. group t o be d i s c u s s e d age i s n o t i m p o r t a n t at a stage over be c o n f u s e d juveniles, a r e known, where males be i m p o r t a n t for 1972, arise i n t h e Sage i n any one t e r r i t o r y , and t h e and e x p e r i e n c e , some 1970), territorial f o r example R e s u l t s , however, i s occurs (Wiley vacancies could themselves, occupancy dominance any one h i e r a r c h i c a l then broods There and urophasianus, Sparrows, juveniles. broods. age bird and was second bird. i n the largest hierarchy. Prior occupancy Experience familiarity and e x p e r i e n c e and/or with prior territory) and p o s i t i o n occupancy and the i n the hierarchy (prior residency, relationship with 99 hierarchy These rank has relationships (1971); Krebs in birds, e. have g. much a t t e n t i o n been the 1965), Great Blue Jay dominance order the Brown (Brian Parus (Brown place; with i t s the distance Territorial adults could even food seems juncos their The at was u n l i k e l y that food, not recent fish, and e. even feeding territories recognition juveniles they years. g. in Phillips man, e. (Colguhoun p o s i t i o n of the distance of the g. its enticed in were by not the of their of the feeding The incentive was apparently not strong of, and and the rest therefore the of enough the not in territory varied be In Song apply. territories territory. presence of behaviour of then territorial would bird not neighbour's aware for, and its does noisy respect 1942), territory. from their the (Dixon individual probably be Chickadee stations. comprising non-territorial, Carolina from supplied particularly the the rank phenomenon when 1949), with this the (1964); 1963), Sparrows, the in caeruleus, corresponds feeding inversely Tit Tit, Steller's It found in (1973). In from received to to leave overcome boundaries. hierarchy involved in were this idea. Sabine first on groups that thought (1959) the by suggested wintering arrived Kikkawa that grounds later. (1961) Prior to those were juncos in,general residency, be a that dominant however, factor in arrived was to not determining 100 hierarchy position advantage area was conferred masked The situation position in juveniles their Song can own formed birds with the large number regarding same during the of the case, an area, to Introduction), would have supporting an or It seems, a factor in however, cannot factors The lower top rank. the first, one during of idea that hierarchy. and ruled the birds out Further, be the are as a in birds in likely along fall territorial the suggestion to that is arrange have those of the familiarity dispersal that juveniles phase (see concentration arrivals. prior position possible No data of occupancy a juvenile prior is Song occupancy, factor. hierarchy genetically social top most only' available. long-term the that would centres later fact previous Short-term position may the over this argue an areas appear for with hierarchical are this individual determining be groups If regard possibly therefore, Sparrow Other no advantage in adults. and were of any present, the certain juveniles at refuting by recrudescence could General birds adults that familiarity occupancy in without arrive of juvenile time because One happened all The the positions suggested complicated territorial then familiarity. not is he previous prior considered. the hierarchical that on Sparrows be about behaviour with White-eyes; territories. hedgerow, the by in rank different could be from birds of genetically 101 inherited. not an by hypothesis juvenile based aggressive shows t h a t each an at Sparrow's the an argued that individual, evidence features position dominance in that the dominance but supporting a or might is 'role* refuting determine dominance is a bird was and is a the the involved. behaviour, the bird top the a a hierarchies not number show because incidence of end of the then, there based difficult dominant of as to on determine bird. Figure interactions in Krebs the his i n Song are 9 p o s i t i v e c o r r e l a t i o n between However, hierarchies higher is significant i n d i v i d u a l need that hierarchies i t i s frequently aggressive there aggressive there no genetic in hierarchy alpha fact on have encounters, most position I of has Island. Although the however feature him. Song Reifel if (1967) intrinsic acquired on Sade (1973) which points out, highest freguency position i s stable. Sparrows are interactions stable, hierarchy, requires The and concerning of yet birds further investigation. In summary, evidence in this determine the rank hierarchy. prior Males occupancy short-term prior study that an dominated were not occupancy appears supporting to be any little f a c t o r (s) i n d i v i d u a l Song females. thought and Sparrow Size, to possibly conclusive be age, that held and or in the long-term important, genetic might but aggression 102 features were not ruled out. Finally, i t i s worth hierarchies, individuals obtaining (c) and territories Dixon subordinates position were at idea that the of the t h e dominant successful ones in arose. as often from as bird was Guhl first, followed appeared the B-C just 1956; of the not related Table that i s a 1955; The appearance station there 16 by first to i t s gives the on t h e t e n d a y s that Hierarchy. as o f t e n In fact, as dominates, and males. found Serinus that, serinus, success the feeds hierarchy. first (1939) breeding order. feeding of animals, Anthony individual observing arrived canaries, evidence the groups (e. g. dominance i n Shoemaker higher the of the f i r s t subordinates of order i n descending i n the spent females outcomes the opportunity The dominant Sparrow position when hierarchical feeding 1965). Song the of the Hierarchies certain pronounced particular at i n t h e h i e r a r c h i e s were Outcomes In in looking than present in a laboratory the dominant males subordinates. study population had a There to support much i s or refute no this suggestion. In not only certain lek species, i s i t important (e. g. f o r males Sage Grouse, to obtain a Wiley 1970), territory to 103 POSITION Figure 9 IN HIERARCHY P o s i t i o n i n h i e r a r c h y and number o f i n t e r a c t i o n s : t h e B-C Hierarchy The l i m i t a t i o n s o f t h i s a n a l y s i s a r e acknowledged. Each b i r d d i d n o t spend an e q u i v a l e n t amount o f time a t t h e f e e d i n g s t a t i o n . Some b i r d s f e d w i t h o u t an i n t e r a c t i o n o c c u r i n g . The m a n i f e s t a t i o n s d i s c u s s e d i n the t e x t . o f aggressive behaviour are 104 mate, but i t i s territories to species birds of pre-requisite many in of the in for successful which species, in v obtaining winter Odum flock a the of results correlation, in the that the dominant ones pre-breeding summary, then, in the at Hierarchies presence stable, with In the few long-term in determining Dixon did a are several territory is breeding. dominance in the been that a In hierarchy hierarchy shown the top in (1965) found (1964) study in obtain hierarchies a few males in established similar results Song further that and only three Chickadees Tompa dominance a favoured determined. has individuals in There hence of dominance reported as groups of juvenile favoured formed, Sparrows strengthen this territories established . are during Song localities which by Sparrows along congregated the mid-winter hedgerow. were linear reversals. hierarchies, and been present certain were and certain season. In fall mate, has spring. obtain a c q u i s i t i o n of Black-capped Chickadees, Carolina the to matings. territory (1942) in the a c o r r e l a t i o n between in the the season territories The important obtaining pre-breeding populations. a ensure these The success also prior the males occupancy rank dominated were that an not females. thought individual to Size, be held age, important in the 104 TABLE 16 F e e d i n g o r d e r o f h i e r a r c h y : appearance o f f i r s t a t f e e d i n g s t a t i o n s o f the B-C bird Hierarchy Sex Position of b i r d i n hierarchy Totals: Number o f days Male Female F i r s t seven 3 3 0 Middle eight 4 2 2 L a s t seven 3 0 3 22 b i r d s 10 105 hierarchy. aggression The Short-term prior occupancy features were removal experiments showed that those spring on the not ruled yearlings Spring individuals in the pre-breeding season. that possibly genetic or out. and subsequent obtained Removal hierarchies and Area replacements territories were established the in the dominant during the 106 LITERATURE CITED Alexander, R. 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Lorenz, K. Z. 1966. Lond. 1952. Territory i n small carnivores. 18: 143-65. King Solomon's Ring. London. of 3: Symp. 112 M c B r i d e , A. F . a n d D. 0. Hebb, 1948. Behaviour of the captive Bottle-nosed Dolphin, Tursiops truncatus. J. Comp. Physiol. Psychol. 4 1 : 11T-2 3. ~ ~ Magnuson, J . J . 1 9 6 2 . An a n a l y s i s o f a g g r e s s i v e growth, and competition f o r food and space Oryzias latip.es (Pisces:Cyprinodontidae). Z o o l . ~407 3"l3-63. behaviour, i n medaka, Can. J. M a r s h a l l , J . T. 1948a. E c o l o g i c a l r a c e s o f Song the San Francisco Bay region. Part I . abundance. Condor 50: 193-215. Sparrows i n H a b i t a t and M a r s h a l l , J . T. 1948b. E c o l o g i c a l r a c e s o f Song the San Francisco Bay region. Part I I . variation. Condor 50: 233-56. Sparrows i n Geographic Mason, J. C. a n d D. W. Chapman. 1965. S i g n i f i c a n c e o f early emergence, environmental rearing capacity, and behavioural ecology of juvenile coho salmon i n stream channels. J. Fish. Res. Bd. Can. 22: 173-90. M a s u r e , R. H. a n d w. C. Allee. f l o c k s o f t h e common c h i c k e n 306-27. Masure, R. H. a n d W. C. of the S h e l l Parakeet, 15: 3 8 8 - 9 8 . 1934a. The s o c i a l and the pigeon. order i n Auk 51: Allee. 1934b. Flock organization Melopsittacus undulatus. Ecology Michielson, A. 1966. Intraspecific and interspecific c o m p e t i t i o n i n t h e shrews Sorex araneus and S. m i n u t u s . Arch. Neerl. Zool. 1 7 : 73-1947 Miller, G. R., D. J e n k i n s a n d A. Watson. 1966. Heather performance and Red Grouse populations. I Visual estimates of heather performance. J . Appl. Ecol. 3: 313-26. M o o r e , N. among Moss, W. 1964. Intraand dragonflies. J. Anim. interspecific competition Ecol. 33: 49-71. R. 1969. A c o m p a r i s o n o f Red G r o u s e s t o c k s p r o d u c t i o n and n u t r i t i v e v a l u e of the heather vulgaris. J . Anim. Ecol. 3 8 : 103 - 2 2 . with the Calluna Murton, R. K., A. J . I s a a c s o n a n d N. J. Westwood. 1966. T h e r e l a t i o n s h i p s b e t w e e n Wood P i g e o n s a n d their clover food s u p p l y and t h e mechanism o f p o p u l a t i o n c o n t r o l . J. Appl. Ecol. 3: 5 5 - 9 6 . " Myres, J. H. a n d C. J. Krebs. 1971. Genetic, behavioral 113 and reproductive attributes Microtus pennsvj.anicus and Monogr. 4 1 : 53-787 Nettleship, D. N. 1973. Arenaria interpres, at I b i s Tl5: 202-7. Newsome, A. E. mouse p l a g u e . of fl. dispersing Field pchro_gaster. Breeding ecology of Turnstones, Hazen Camp, Ellesmere Island. 1970. An e x p e r i m e n t a l a t t e m p t t o p r o d u c e J. Anim. Ecol. 39: 299-311. Nice, M. M. Sparrow. 247pp. 1937. Part Nice, M. M. 1941. The role of territory Amer. Midi. Nat. 26: 441-87. N i c e , , M. M. Sparrow. 329pp. Studies i n the l i f e history of the I. Trans. Linn. Soc. N, Y. E. P. Chickadee. O r i a n s , G. social Peek, 1942. P t . 3. bird ecology Monogr. the Phillips, R. behaviour Chasmodes 389-408™ life. of blackbird 31: 285-312. Black-capped (Agelaius) 1971. S e a s o n a l c h a n g e i n t h e b r e e d i n g b e h a v i o u r male Red-winged Blackbird. Wilson B u l l . 83: Perrins, C. M. 1965. Population fluctuations clutch-size i n the Great T i t , Parus major. J. Ecol. 34: 601-47. P h i l l i p s , A. Sparrows Song IV. Schmidt. 1938. Social Night-Heron. Auk 55: Annual cycle of Auk 5 9 : 4 9 9 - 5 3 1 . H. 1961. The systems. Ecol. F. W. of the 383-95. in a 1943. S t u d i e s i n t h e l i f e h i s t o r y o f t h e Song Part II. Trans. Linn. Soc. N. Y. VI. Noble, G. K., M. Worm and A. behaviour of the Black-crowned 7-40. Odum, Voles Ecol. R. a n d R. W. o f t h e Mexican Dickerman. 1957. N o t e s o n Plateau. Auk 7 4 : 3 7 6 - 8 2 . and Anim. Song R. 1971. The relationship between s o c i a l and the use of space i n the Benthic fish bosauianus Lacepede. Z. Tierpsychol. 29: P i t e l k a , F. A . , P. Q. T o m i c h a n d G. W. Treichel. 1955. Ecological relations of jaegers and owls a s lemming p r e d a t o r s near Barrow, Alaska. Ecol, Monogr . 25: 85-117. 1 14 R a l p h , C. J . a n d C. A. Pearson. 1971. C o r r e l a t i o n o f age, size of territory, plumage and breeding success in White-crowned Sparrows. C o n d o r 73: 77-80. R i c h d a l e , L. Oxford. E. 1957. A population Rowan, M. K. 1966. Territory m e c h a n i s m i n some S o u t h A f r i c a n 6: 3 9 7 - 4 0 8 . study of penguins. as a density-regulating birds. Ostrich Suppl. S a b i n e , W. S. 1959. The w i n t e r s o c i e t y o f t h e O r e g o n J u n c o : i n t o l e r a n c e , dominance and t h e p e c k i n g o r d e r . Condor 61: 110-35. Sade, D. S. 1967. Determinants of dominance i n a group of f r e e - r a n g i n g r h e s u s monkeys. IN: Social communications among p r i m a t e s . S. A. A l t m a n n (ed) Chicago. Sale, P. F. 1972. the pomacentrid 741-44. Schjelderup-Ebbe, des Haushuhns S c h o e n e r , T. birds. T. . I n f l u e n c e of c o r a l s i n the fish, DascyjLlus aruanus. Z. W. 1968. E c o l o g y 49 1922. B e i t r a g e zur S o c i a l p s y c h o l o g i e Psychol. 88: 226-52. : Sizes of 123-41. Sheppard, D. H. 1971. s p e c i e s (Eutamias) . S h o e m a k e r , H. canary. H. Auk d i s p e r s i o n of Ecology 53: feeding territories Competition between E c o l o g y 52: 320-29. 1939. Social 56: 381-406. hierarchy in two among chipmunk flocks of the Simms, E. 1965. E f f e c t s o f t h e c o l d w e a t h e r o f 1 9 6 2 - 6 3 on the B l a c k b i r d p o p u l a t i o n of D o l l i s H i l l , London. Brit. B i r d s 58: 33-43. Smith, C. C. organization E c o l monogr. 1968. The i n the genus of 38: 31-63. adaptive nature of social tree squirrels Tamiasciurus. S m i t h , R. F. J. 1970. Effects of food availability on aggression and n e s t b u i l d i n g i n Brook S t i c k l e b a c k , Culaea inconstans. J. Fish. Res. Bd. Can. 27: 2 3 5 0 - 5 5 . S m i t h , S. M. 1967. Seasonal changes i n the s u r v i v a l Black-capped Chickadee. C o n d o r 69: 344-59. Smyth, M. 1968. The effects of removal of individuals of the from a 115 population of Bank Voles, Anim. Ecol. 37: 167-83. Snow, D. W. S n y d e r , W. scaled 19. 1958. A study Clethrionymus of blackbirds. glareolus. London. D. 1967. Experimental habitat improvement quail. Tech. Publ. Colo. Game F i s h P k s . Sommer, R. 1967. 145-52. Small Stefanski, R. territory in 259-67. A. the Stenger, J. Ovenbirds group ecology. J. Psychcl. 1967. Utilization of Black-capped Chickadee. Bull. for Dep. 67: the breeding Condor 69: 1958. Food habits and available food of i n relation to territory size. Auk 7 5 : 3 3 5 - 4 6 . S t e w a r t , R. E. and J. W. Aldrich. 1951. repopulation of breeding birds i n a spruce community. Auk 6 8 : 4 7 1 - 8 2 . Suthers, R. A. territories 232-37. of 1960. the Removal and - f i r forest Measurement of some Song Sparrow. Wilson lakeshore Bull. 72: S y m o n s , P. E. K. 1968. Increase in aggression and in strength o f t h e s o c i a l h i e r a r c h y among j u v e n i l e A t l a n t i c salmon d e p r i v e d o f food. J. Fish. Res. Bd, Can. 25: 2387-401. S y m o n s , P. E. K. 1971. density to available J. Fish. Res. Bd. B e h a v i o r a l adjustment of population f o o d by juvenile Atlantic salmon. Can. 28: 569-87. ° Tester, J. R. a n d A. Watson. t e r r i t o r i a l i t y o f Woodcock, S c o l o p a x roding behaviour. I b i s 115: 135-7. Tompa, F. S. 1962. Territorial controlling factor of a local Sparrows. Auk 7 9 : 6 8 7 - 9 7 . 1973. Spacing rusticola based behaviour: population the of and on main Song Tompa, F. S. 1964. F a c t o r s d e t e r m i n i n g t h e numbers o f S p a r r o w s , M e l o s p i z a m e l o d i a , o n M a n d a r t e I s l a n d , B. Canada. Acta Zool. Fenn. 1 0 9 : 1-73. Song C , Tompa, F. S. 1971. Catastrophic mortality population consequences. Auk 8 8 : 7 5 3 - 5 9 . i t s Tompkins, G. 1933. Individuality and and territoriality as 116 displayed by three passerine species. C o n d o r 35: 98-106, T o r d o f f , H. B, 1954. S o c i a l o r g a n i z a t i o n and b e h a v i o u r in a flock of captive, n o n - b r e e d i n g Red C r o s s b i l l s . Condor 56: 346-58. T y l e r , S. J. the New 187-94. 1972. Forest The b e h a v i o u r and Ponies. Anim. W a t s o n , A. 1964. A g g r e s s i o n and Grouse. N a t u r e 202: 506-7. social organization Behav. Monogr. population of 5: r e g u l a t i o n i n Red Watson, A. 1965. A p o p u l a t i o n study of Ptarmigan, Laggpus mutus , i n S c o t l a n d . J. Anim. Ecol. 34: 135-72. W a t s o n , A. 1967. Red G r o u s e . T e r r i t o r y and p o p u l a t i o n N a t u r e 215: 1274-75. regulation i n Watson, A. a n d D. Jenkins. 1968. Experiments c o n t r o l by t e r r i t o r i a l b e h a v i o u r i n t h e Red Anim. Ecol. 3: 5 9 5 - 6 1 4 . Watson, A. a n d G. R. Miller. a g g r e s s i o n i n a f l u c t u a t i n g Red Anim. Ecol. 40: 367-83. the on p o p u l a t i o n Grouse. J. 1971. Territory size Grouse population. and J. W a t s o n , A. a n d R. Moss. 1970a. Dominance, s p a c i n g behaviour and aggression i n relation to population limitations i n vertebrates. IN: Animal p o p u l a t i o n s i n r e l a t i o n t o t h e i r food resources. A. Watson (ed) Blackwell, Oxford: 167-218. Watson, A. a n d R. Moss. t e r r i t o r i a l behaviour, Grouse. IN: Behaviour b y a n i m a l s a n d men. A. York - London. W a t t s , C. R. a n d A. W. of T u r k e y s . Scient. Weeden, J. Sparrow. S. 1965. Condor 67: 1970b. Spacing as a f f e c t e d by habitat, and nutrition in Red and environment: t h e use o f space H. E s s e r (ed) P l e n u m P r e s s , New Stokes. 1971, The social Amer. 224 ( 6 ) : 112-8. Territorial 193-209. behaviour of the Wessell, J. P. a n d H. Leigh. 1941. Studies of the organization of the White-throated Sparrow. Wilson 53: 222-30. Wood-Gush, D. G. M. 1965. domestic b i r d communities. 14: 2 1 9 - 3 1 . The Symp. order Tree flock Bull. s o c i a l organization of Zool. Soc. Lond. 1 17 W i l e y , R. H. Jr. mating in the Onpubl. Ph. D. York. W y n n e - E d w a r d s , V. C. s o c i a l behaviour. Y o u n g , C. W. 1970. Tadorna tadorna. 1970. Territoriality and non-random Sage Gronse, C e n t r o c e r c u s urophasianus. T h e s i s , The R o c k e f e l l e r U n i v e r s i t y , New 1962. Animal d i s p e r s i o n i n r e l a t i o n O l i v e r and Boyd. Edinburgh. Territoriality in I b i s 112: 3 3 0 - 5 . the Common to Shelduck Young, E. C. 1972. Territory establishment Mccormick's Skua. I b i s 114: 2 3 4 - 4 4 . and Zaret, T. M. a n d A. , S. Rand. 1971. tropical stream fishes: support for exclusion theory. E c o l o g y 52: 336-42. Competition in the competitive Zimmerman, J. L. dependant e f f e c t 1971. The territory and i n S p i z a americana. Auk 8 8 : stability in i t sdensity 590-612. 118 APPENDIX plants 1. List along The occasional J*i Finally, Populus Pseudotsuga herbaceous plants important ones, Vicia spp., speciosa, of : SE®ricanus, Care x of small racemgsa, consisted mainly R.. l a c i n i a t u s , albus, trees the and were A In us Lonicera principally rubra, and . and detailed was n o t however; vegetational attempted. are listed lanceolatum, margaritacea, species adjacent spp., o f mature P.. t r i c h o c a r p a , and s e v e r a l Salix species spectabilis, stands with Sambucus Shrubs the Cirsium marsh hookeriana, Rubus frequently shrubs, T h e main spp. important thick, and spp., Symphoricarpos Anapholis aguilinum, The Salix a Rosa menziesii complete trees. and most marsh. of trees and Prunus tremuloides, A small stolonifera, macrophyllus, common and a d j o i n i n g o f mature douglasii, Cornus o f most comprised of AInus r u b r a , were spp. was tangle stand Crataegus of t h e hedgerows hedgerow impenetrable trees of species The Urtica lyalii, Eguisetum lyngbyei, Typha I^ysichiton the and lanatum, Petasitgs aryense, i n cleared of common : Heraclgum of grass gale, most here t o t h e hedgerow Hyrica survey Pteridiura areas. was c o m p r i s e d 1atifolia, k a rots h a t c e n s e mainly Scirpus . 119 APPENDIX 2 Weather data from the M e t e o r o l o g i c a l O f f i c e , Vancouver I n t e r n a t i o n a l A i r p o r t Mean monthly temperatures Jan. Feb. Mar. Apr. May June July Aug.- Sep. Oct. Nov. Dec. — 39.0 44.6 44.6 54.7 58.3 62.3 64.0 54.3 46.1 43.0 35.2 1973: 36.0 40.5 43.2 47.4 53.7 Normal (37 y r ) :36.3 40.0 42.5 48.1 54.3 59.5 63.4 62.7 57.6 50.1 42.9 38.9 Apr. May June July Aug. Sep.- Oct. Nov. Dec. 1972: Monthly temperature :ranges 1972 Max. Jan. Feb. Mar. — 56.0 62.0 ft 60.9 75.0 73.4 81.4 80.7 77.9 62.1 56.6 51.8 45.9 47.9 48.0 35.0 30.2 25.4 12.0 17.4 30.0 30.8 41.1 1973 Max. 52.9 58.1 53.2 60.6 76.6 Min. 13.1 24.3 28.1 33.9 36.8 Jan. Feb. Mar. Apr. May_ June July Aug. Sep. Oct. Nov. Dec. — 5.25 6.98 45.1 1.17 1.84 3.20 0.92 3.14 2.09 4.61 11.34 1973: 5.43 2.69 2.69 0.51 1.91 Normal: A.95 4.29 3.51 2.39 1.87 1.78 1.17 1.46 2.41 4.81 5.47 5.96 — 0.3 0.4 1.9 1973: 4.7 TR 0.1 TR Normal: 8.8 3.1 2.1 TR 5.28 7.05 4.70 1.27 1973: 5.92 2.69 2.70 0.51 1.91 Normal: 5.80 4.57 3.69 2.40 1.87 Min. ~ Precipitation R a i n f a l l : 1972: Snowfall: 1972: Totals: 1972: — TR 6.4 0.9 5.7 1.84 3.20 0.92 3.14 2.09 4.61 11.82 1.78 1.17 1.46 2.41 4.81 5.56 6.51 120 APPENDIX 3 Destruction of a feet of stretch of dyke ( Bulldozed , Area'). In June, Removal of the This stretch 10 of gives bulldozing 3 4 was males birds rebuilt. the Area and no in less than must bird along when the resident the fall 1973 as about 2000 The Control a spring. the with a on total gravel number of that the Fall detruction and Song rock. Sparrows; existed before a of have the this were the hedgerow stretch. A l l relocated. summer close adjacent was to to removed as the the on a One cryptic Bulldozed Bulldozed October 3/1972 early March experiments. territorial south third bird, the female male about this 4 spent male the after along these (Green/Green) strongly metres 3 territory removal male known June. to apparently i t boundaries were late over A second entailed supported 1 female took during he hedgerow started. non-territorial as This territory (Black/Orange) Area, dyked replacing hedgerow disappeared However, male 800 v e g e t a t i o n , and Figure the Area 1972, 300 bird on metres male's was located in in a stretch Westham (LP/White) from female her of shrubbery Island. , paired destroyed disappeared with a territory, in early F i g u r e 10 Bulldozed before A r e a : the arrangement o f t e r r i t o r y the dyke r e b u i l d i n g o c c u r r e d boundaries 122 These non-breeding, data suggest that adults can survive as birds through the summer, in the surplus of birds non-territorial spite of the fact seemed t o be that composed only of juveniles. \ on the study area 7 123~ APPENDIX locate 4 Possible i n the Some locate 1. fall fall October, found are that September, Island were which 4 partial 2. and females why are not given removals 1972, and were to to of to seemed Removal by Moulting Nice (1937) i n August finishing moulting However, moult place. not September Simultaneous out moulted bird occurred. difficult find. i n Ohio occasional were took difficult Sparrows most the difficult here. Casual observations of from were females a l l females the the made i n until birds on to the be and Beifel month one male removed on area was s t i l l in moult. Females are generally apparently spring that Song moulting October 1972 extremely with mid-October. reasons when the why 1972. of I t i s possible individuals in of possible i n the early reasons (Nice 3. Females and early Tompa desert their Thus, territorial Nice 100-700 i t males mates to d i d not most showed females adjacent metres seems fall than males, than in the 1964). did 3056 w e n t to locate responsive i n the (1943) instances territories. territories less 1943; fall. possible are more d i f f i c u l t often that i n the in only return to late 31% out their territories, and summer of 55 former 33^ to distant. quite likely that have females i n the several of fall 1972. of the 124 APPENDIX and 5 other Interspecific seed-eating Song of Sparrows birds that common frequented most the stations, at of the observations, inetrspecific dominance most which and tsong stations. There Sparrow, the Sparrows of the other stations. J u n c o , Fox mexicanus, occurred From feeding between the feeding Occasionally, other Carjgodacus lincolnii, at i n t e r a c t e d with Oregon abundant.. Finch, passerines fed at the species, interactions junco were and three Towhee, was by seed-eaters species that far such L i n c o l n ' s Sparrow, as the House Helgspiza stations. a very could general be c o m p i l e d list thus in order of : Towhee Fox Sparrow Song Sparrow Lincoln's however, certain Song Fox Sparrows. On Fox Sparrow, but precedence over indifferent and would to feed Juncos House Finch Oregon Junco Sparrows would occasion, this Song was a Song Sparrows. were leave threaten Sparrow r a r e , and the t h r e a t e n i n g and Sparrow would Towhees behaviour driven to o f t h e Song by and off Sparrows tended o f such off Towhees drive u s u a l l y Fox independently readily both a took remain Sparrow, behaviour. a l l the other 125 / seed-eaters, relatively but close retaliation. often to sheer potential numbers allowed antagonists them without to feed inducing 126. APPENDIX the 6 B-C age Hierarchy of the a r e known. banded and on J u l y i t had p r o b a b l y (see Nice 35-40 i n hierarchy: t h e t o p two b i r d s i n Hierarchy The was Age a n d p o s i t i o n top The f i r s t 25, w i t h just 19^3). two dominant no s i g n s gained Its.age juvenile males juvenile t h e B-C (Blue/Blue) of post-juvenile independence on J u l y in 25 was from moult, i t s parents put t e n t a t i v e l y a t days. The banded (Stage second on May 4 of Nice male on 25, the dominance at the 1943).. scale fluttering Therefore i t was (Yellow/LP) stage of between was development 10 and 16 days o l d . Thus, were i n determined, Yellow/LP January, Blue/Blue would be a b o u t product o f most product of a first of the largest when likely a brood. hierarchy. the positions would be about 260 d a y s second old. brood, Yet, Blue/Blue of these 200 days Blue/Blue and was two birds old, would Yellow/ the top and be a LP a juvenile 127 APPENDIX If to (N 7. Pressure each potential a resident's = nos. o f proportional T i s also T assumption to increase makes x i n t r u s i a n s potential settlers). The the resident spends the time ,when T. settler Explanation. the total to being intruders: territory, proportional constant, from Nx that to territory goes number o f i n t r u s i o n s area, up, t e r r i t o r y the bird does p e r d a y on value of n o t have Nx i s i n defence (T). so i n order size Nx = goes much to keep down - t h e spare time 128 8 APPENDIX The Watson conditions a five and which c o n d i t i o n s of Moss need (1970a) t o be Watson put met to and forward show Moss. the that following behaviour five limits breeding population. A. A either substantial because breeding even part animals though of the die; or they p o p u l a t i o n does because survive, they and are may not breed, inhibited breed in from later years. B. if Such the are C non-breeders more dominant The breeding such the resource D. The as, If change both as territorial itself other (B) , the are space, capable ( i . e. of breeding breeding) animals or not completely nest sites. using If up they some are, limiting. depressed in causes following limiting i s or changes (A) , animals food, mortality factor(s) E. or physiologically removed. resource, rate are an of (C) , a n d changes r e c r u i t m e n t due opposite mortality (D) are i n food, sense or to, to and depressed fulfilled, then food the and and at limiting the same recruitment. the numbers behaviour are breeding population. 129 COTTONWOOD STRETCH LOSER \ 6 \LP G LP LP LP M LpUnb R Y Y © LP LP jjBUnt) \ R Y 14 \ o" 6" Y © « G G 9 3 7 10 5 2 5 \ 2 2 1 4 1 2 3 1 LP RW © Black <D RW RW G 1 1 G G w 2 d* d LP Black RW w © © 2 11 11 7 - 11 2 2 3 1 9 ? ? 2 G ® LP L-D B 0 0 B BO B 5 3 5 1 1 3 4 5 2 1 4 1 1 1 1 2 2 1 1 2 5 2 1 \ 2 1 1 1 ® LP L-D 1 BO OB 1 BO B 170 I N T E R A C T I O N S 13 MAN HOURS F i g u r e 12 SEDGE \ GB GB G BY R Y cf cf GB G GB BY \ 2 \ cf R Y 15 ? 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UBC Theses and Dissertations
Some ecological aspects of social behaviour in the song sparrow, Melospiza melodia Knapton, Richard Walter 1973
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