TERRITORIALITY

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The Bell Vireo exhibits "classic" passerine territoriality. Within a specific area, a pair of this species carries out pair-formation, courtship activities, copulation, nesting, rearing the young, and foraging. With the cessation of reproductive activities, a pair continues to restrict its other daily activities to the same general area.

Establishment of Territory

In early May the segment of the total suitable habitat within which a Bell Vireo restricts its activities is not rigidly defined and the first male of the season ranges over an area too large to be maintained permanently—one that seems greatly to exceed the needs of breeding. Male 1 (1960), for instance, was first seen foraging over an area of approximately seven acres. With the influx of other males, portions of this large tract were usurped and the territory of the original male was gradually reduced to an area of little more than an acre.

In this initial period, a male becomes identified with a large area but is restricted to an area of nearly typical size by the encroachment of other males. Territorial disputes in this period often involve physical contact, as well as protracted sessions of high-intensity singing at rates exceeding three hundred song-phrases per hour.

Eventually the carrying capacity of the habitat is reached and no further partitioning occurs. The beginning of nestbuilding coincides with this relative stabilization of the territorial boundaries. Through the remainder of the cycle of behavior associated with any one nest, all activity is that of the occupant pair within its territory.

Size of Territories

The nine original territories established in 1960 varied in size from 0.26 acre to 3.1 acres (Table 2). Fitch (1958:270) found the territories of several pairs of Bell Vireos at the University of Kansas Natural History Reservation to vary from 0.4 to 1 acre. Hensley (1950:243) estimated the size of the territory of a pair of Bell Vireos observed in Piatt County, Illinois, at 3.1 acres. Nolan (1960:227) records home ranges of 2 to 3 acres. The pairs that he studied were sole occupants of fields several acres larger than the portions actually utilized. His description of the vegetation indicates that most of the second growth was not much taller than 7 feet. As indicated elsewhere, the second-growth in my tract averaged 15 feet tall. The smaller average size of territory (1.25 acres) that I found is probably a function both of this greater vertical range of available foraging area and the much higher gross density of birds (40 pairs per 100 acres).

Permanence of Territories

Most pairs remain in their original territories throughout the summer, although some shift certain territorial boundaries. In 1960 pairs 2 and 6, in the course of selecting a site for a replacement nest, annexed adjacent areas previously occupied by other pairs. Pair 2 relocated in a space that originally included territories 1 and 4, and pair 6 built a nest in an area formerly occupied by pair 7. Males 1 and 4 were sacrificed for specimens and pair 7 probably was destroyed by a predator. Owing to the presence of a nest, the annexed area becomes the focal point of the activities of a pair, but the original area is regularly visited and may be returned to in a later renesting.

Table 2. Size of the Nine Original Territories Occupied in 1960.

Territory Date first occupied Dimensions
1. May 3, 1960 1.6 acres
2. May 5, 1960 0.6 acre
3. May 7, 1960 0.26 acre
4. May 11, 1960 1.03 acres
5. May 12, 1960 2.07 acres
6. May 14, 1960 3.1 acres
7. May 13, 1960 1.7 acres
8. May 14, 1960 0.46 acre
9. May 14, 1960 0.4 acre
Average 1.25 acres

Maintenance of Territory

Except in the early stages of nesting, territory is maintained primarily by song. In the period of incubation a male regularly patrols his territory between sessions of sitting on the eggs. He sings several songs from each of several perches. A male follows a predictable path, rarely traveling more than 150 feet from the nest. Incipient patrolling is seen early in the breeding season when territorial boundaries are in a state of flux.

The male White-eyed Vireo travels a semi-predictable route, as does the Solitary Vireo (R. F. Johnston, MS). According to Lawrence (1953:50), the male Red-eyed Vireo has a distinct singing area completely divorced from the nest area dominated by the female. Southern (1958:109), working with this same species in Michigan, did not recognize separate areas, but found that the male wandered randomly over the territory. In a species so highly active as the Bell Vireo, the degrees of hostile action associated with an encounter overlap in such a fashion that no clearcut distinction can be drawn among the various displays. Nevertheless, certain generalized patterns are characteristic of all situations in which members of this species are in a state of anxiety. The threat displays described in the succeeding paragraphs may all be utilized within as little as two minutes; mutual agonism may be terminated at any stage by concerted attack of the dominant bird.

1. Vocal threat. When an intruder is discovered the resident male markedly increases his rate of singing. The alarm note, eh-eH-EH, is the first call uttered during the nestbuilding and egglaying periods.

2. Head-forward threat. If the intruder does not flee, the resident male adopts a specific threat posture. The head and neck are extended. The feathers of the crown are erected, but those of the body are sleeked. The bird crouches slightly and the tail is flicked laterally, but not fanned. The intensity of the singing increases and is supplemented by scolding, also delivered at a rapid rate. The intruder normally retreats at this juncture.

3. Wing-flicking and submaximal tail-fanning. If the interloper remains, the anxiety of the resident male increases. He slightly depresses the tail and, at the same time, rapidly fans and closes it. The tail is only partially fanned. The wings are held slightly away from the body and rapidly flicked above the back. This flicking should not be confused with quivering of the wings associated with begging and other solicitory postures. Song is now almost completely replaced by high-intensity scolding. Associated with this high degree of anxiety are displacement behaviorisms, including bill-wiping, reversal of direction on a single perch, and a nervous hopping from one perch to another.

4. Ruffling and maximum tail-fanning. This display is most often seen in conjunction with the harassment of predators, but occasionally it is observed in territorial disputes occurring at the boundary of adjacent territories where neither male is strictly dominant and in which there is much vacillation prior to attack. The feathers of the abdomen are ruffled. The term "ruffled" pertains to a full erection of the feathers, giving a ragged appearance to the body outline (Morris, 1956:80). Ruffling of the abdominal feathers emphasizes their yellow color and seemingly heightens the intimidatory effect. The tail is fully fanned, and so maintained, for a few seconds at a time; it is held at a 45° angle to the body. The scold becomes an extremely intense, stacatto buzz, ZZ-ZZ-ZZ-ZZ.

5. Supplanting attack. The attack directed against a trespassing male is initiated as a lunge that results in a collision with the opponent in mid-air or on his perch. The bird attacked is struck by his adversary's open beak or body.

Hinde (1952:71-72) indicates four courses of action followed by a Great Tit (Parus major) when attacked under similar circumstances. "(a) It flies away: The attacker usually flies after it and a chase ensues. (b) It shifts its perch a few inches: the attacker lands in its place, and both usually show head-up postures. (c) It remains where it is, but adopts a head-up posture: the attacker usually then shows upright flight. (d) It may fly up and meet the attacker in mid-air: in that case an actual combat may result, or both combatants may show upright flight."

Head-up posturing and upright flight are not presently recognized components of the behavior of the Bell Vireo. The behavior of the attacked Bell Vireo is similar to that described in (a), (b), and (d) above, and is clearly dictated by the proximity of his own "home base."

Eleven disputes among occupants of adjacent territories were witnessed between May 6 and June 3, 1960, in which some or all of the described threat displays were manifest (Table 3). In each instance, patrolling males were gradually attracted to each other. As they approached, their rates of song increased from an average of six repetitions per minute to 15 per minute. Eight of the disputes involved physical combat.

On May 6, 1960, when male 2 (1960) was in the process of usurping an eastern segment of the original territory of male 1 (1960), a violent, protracted dispute was observed. By this date male 1 (1960) had obtained a mate and had begun construction of nest 1-a (1960); male 2 (1960) had not yet acquired a mate. At first the two males were singing vigorously, from one to 10 feet apart. Female 1 (1960) followed her mate closely and scolded, at the same time partially fanning her tail. In the course of vocal dueling the males had traveled to within 50 feet of nest 1-a (1960), when male 1 (1960) suddenly lunged at 2 (1960). The males plunged to the ground, locking bills and clutching at each other with their feet as they fell. As soon as they touched the ground they separated. Male 2 flew east with male 1 in pursuit. This conflict lasted three minutes.

Additional physical combat was witnessed several minutes later. This again involved striking with the bill, wings and feet. A high pitched squeaky chee was uttered by both combatants. The female scolded from a nearby perch. Upon separating, the males engaged in a wild, looping flight. At about 350 feet from nest 1-a (1960), the chase abruptly ended. For ten minutes thereafter, both males sang at a high rate from perches about 10 feet apart. This terminated the physical combat, but three additional protracted, vocal duels occurred in the remainder of the morning.

Table 3. Intraspecific Disputes in Maintenance of Territory.

Behavior

Number of conflicts Vocal dueling Combat Average length of disputes
Prenesting 3 3 2 6 min. 40 sec.
Building 8 8 6 3 min. 8 sec.
Incubation 1[B] 1 ... 20 min.
Totals 12 12 8 5 min. 30 sec.

[B] Directed against a stuffed Bell Vireo.

Probably as a direct result of these conflicts, a neutral zone approximately 300 feet wide developed between the two territories. By May 14 this intervening area was occupied by male 4 (1960). By this date both 1 (1960) and 2 (1960) were involved in nestbuilding and 4 (1960) was not challenged for several days.

Maximum tail-fanning prior to attack also appears as an element of aggressive behavior in White-eyed Vireos. A brief skirmish between a male of this species and a small, greenish passerine was observed at the Natural History Reservation on May 25, 1960. The White-eyed Vireo was singing from a perch 30 feet high in a dead elm, when the unidentified passerine landed 10 feet distant. The white-eye ceased regular song and uttered several catbirdlike calls, and at the same time slightly depressed and fully fanned the tail. After 10 seconds, the white-eye lunged at the intruder, striking it in mid-air. A brief looping flight ensued through the branches of the elm before the intruder was able effectively to retreat.

Aggressive Behavior of the Female

The female Bell Vireo is concerned primarily with the defense of the nest and the young and she rarely assists the male in defense of distant parts of the territory. She employs the same threat displays as the male.

Interspecific Relationships

A number of meetings between Bell Vireos and other species were observed in the course of the study (Table 4). Resident pairs of this species exhibited different degrees of tolerance toward other species. Many birds, including Cardinals, Field Sparrows, Painted Buntings and Mourning Doves were ignored completely. Chickadees evoked responses characterized by slight increase in song and some anxiety; this was perhaps owing to similarity in size, motion and call notes. Warblers, when met with, were invariably chased. They may be momentarily mistaken for rival vireos.

Table 4. Interspecific Conflict Observed in 1959 and 1960.

Species Number of conflicts Phase of breeding cycle Behavior of Bell Vireos
HFT[C] S TF A
Coccyzus americanus 1 Nestling period x
Cyanocitta cristata 3[D] Nestling and incubation period x x x x
Parus atricapillus 1 Prenesting x
Molothrus ater 1 Nestling period x x
Dendroica petechia 1 Prenesting x x
Geothlypis trichas 1 Nestbuilding x x
Pituophis catenifer[E] 1 Post-fledging x x

[C] HFT = head-forward threat; S = scolding; TF = tail-fanning; A = attack.

[D] Includes attack against a dummy Blue Jay.

[E] The Bull Snake is here included because the vireos directed typical aggressive displays towards it.

Blue Jays were vigorously attacked, especially late in incubation and throughout the nestling period of the Bell Vireo. I did not see a jay struck, but a vireo would circle one closely as it perched and pursue it when it flew, following as far as 100 yards beyond territorial bounds. The buzz, ZZ-ZZ-ZZ-ZZ, was uttered in conjunction with this harassment.

A stuffed jay placed eight feet from a nest elicited threat display and displacement behavior from the owners of the nest, but no attack. Incubation had just begun at this nest. A dummy Bell Vireo placed close to another nest only momentarily disturbed the male, and the female completely ignored it. Incubation had also recently begun at this nest. At this same general stage, moreover, nesting pairs showed little inclination to harass me.

Discussion

Hinde (1956:341-342) indicates that territory has been defined in a number of ways by many workers. All of the definitions involve modification of Howard's classic "defended area." Pitelka (1959:253) has reacted against this behaviorally-oriented concept. He thinks that the concept of territory should be based on exclusive use of an area by its occupants, and not so much the defense by which they maintain it.

Methods of treating territoriality in the Bell Vireo seemingly incorporate features of both schools of thought. The area used exclusively for all biological needs by a single pair of Bell Vireos is vigorously defended both physically and vocally early in the breeding season and vocally as the season progresses.

In the period of territorial establishment a relatively large area is actively defended. The building of a nest establishes a focal point of activity in a somewhat more restricted area than that originally occupied. After the success or failure of a nest, a new site is selected to which the focal point of activity is shifted. If suitable habitat adjacent to the extant territory is unoccupied by other Bell Vireos the unoccupied area may be annexed in the course of searching for a new site. Such annexation occurs only when pairs formerly occupying adjacent suitable habitat disappear from this territory; possibly the size of the territory of any one pair is dictated by the density of population of the species as well as by the presence of suitable habitat. This may not always be true as indicated by Kliujver (1951:40), who in studying the Great Tit, found no appreciable difference in the size of territory in two different habitats even though there was a marked difference in population density of the birds.

Fluctuation of territorial boundaries is not uncommon in passerines, especially when no rivals exist to contest movement. Hinde (1956:351) indicates that fluctuations in size of territory are to be expected although the territories of different species of birds have different mean sizes.

Once nesting activities commence there is a marked reduction in the amount of territory utilized and a distinct decrease in the aggressive tendencies of the male; it would seem that energy previously utilized in regular fighting is rechanneled for nestbuilding, incubation and care of the young. Further, contraction of the area of activity obviates high-intensity territorial defense, as adjacent males, even in regions of high population density, are isolated from one another by an area no longer regularly traversed.

With cessation of breeding activities physiological mechanisms governing maintenance of territory seemingly are no longer active and yet the pairs of Bell Vireos remain within a restricted area which they alone use. Earlier definitions of territory as a "defended area" do not adequately cover such situations and yet from the standpoint of Pitelka the area still retains the characteristics of true territory. In fact, territory as defined by Pitelka is clearly manifest at this time. Whether the birds remain in an area through "force of habit" is of little consequence.

I have retained the term "territory" in preference to the term "home range" used by Nolan (1960:227). His failure to observe territorial defense is responsible for his terminology, although it is readily understandable that such defense would be lacking in a population of relatively low density in which pairs were isolated from one another by areas of unfavorable habitat. This isolation in itself would tend to preclude territorial conflict but territories were, in fact, maintained.

The marked similarity in the essential features of aggressive behavior in North American vireos attests to their close relationship. Flicking and fanning of the tail are distinct components of the hostile behavior of the Bell Vireo, White-eyed Vireo, Red-eyed Vireo (Lawrence, 1953:69), and the Black-whiskered Vireo (Vireo altiloquus; Bent, 1950:319), and, presumably, of the remaining species of the genus. The occurrence of these same displays as intrinsic behavioral elements of interspecific hostility suggests a common derivation. Moynihan (1955:256) indicates that all intraspecific hostile displays, and probably most interspecific hostile displays, evolved originally as social signals having the same general function. Further, Hinde (1956:344) points out that there is a fundamental similarity in the motor patterns used in fighting in different contexts, including both interspecific and intraspecific fighting.


                                                                                                                                                                                                                                                                                                           

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