Part 5

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 | | | Average | of | Vocal | | length of | conflicts | dueling | Combat | disputes -------------+-----------+---------+--------+----------------- Prenesting | 3 | 3 | 2 | 6 min. 40 sec. Building | 8 | 8 | 6 | 3 min. 8 sec. Incubation | 1 | 1 | ... | 20 min. +-----------+---------+--------+----------------- Totals | 12 | 12 | 8 | 5 min. 30 sec. -------------+-----------+---------+--------+-----------------

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.

================================================================== | Number | Phase of | Behavior of | | | Bell Vireos Species | of | breeding +------+---+----+--- |conflicts| cycle |HFT| S | TF | A ---------------------+---------+--------------+------+---+----+--- Coccyzus | 1 | Nestling | | | | americanus | | period | -- | x | | | | | | | | Cyanocitta | 3 | Nestling and | | | | cristata | | 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 | 1 | Post-fledging| -- | x | -- | x ---------------------+---------+--------------+------+---+----+---

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

Includes attack against a dummy Blue Jay.

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.