Behavioral Ecology Vol. 15 No.

1: 158–162
DOI: 10.1093/bheco/arg109

Kin selection and reciprocity in flight

formation?
Malte Andersson and Johan Wallander
Department of Zoology, Göteborg University, Box 463, SE 405 30 Gothenburg, Sweden

The reasons for conspicuous ‘‘V’’ and other flight formations in birds are debated. Theory and recent empirical advances show
that energy saving is one important function of flight formations, but some aspects remain poorly understood. Combining
theories of animal flight and sociality, we suggest that some of the variation in flight formations has its base in kin selection and
reciprocation. The bird leading an acute V formation saves less energy than does the trailing participants. The disadvantage of
leading is reduced in more obtuse formations, and when the longitudinal distance between neighbors is small, the leading bird
can save about as much energy as others. Therefore, acute V formations are predicted to occur mainly in circumstances
conducive to kin selection or reciprocity. These mechanisms seem possible, for example, in small flocks of adults with offspring,
such as in swans, geese, and cranes. Inclusive fitness advantages may then favor an energetically expensive leader role for adults.

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In small groups, reciprocity is also possible among unrelated adults that recognize each other and take turns leading the V
formation. In contrast, obtuse formations are expected in large flocks of unrelated individuals, such as spring flocks of waders
migrating long distances. Possibilities for testing these ideas are discussed. Key words: aerial parental care, bird migration, energy
saving, kin selection, prediction, reciprocity, shape variation, testing, theory, V formation. [Behav Ecol 15:158–162 (2004)]

V formation of migrating swans, geese, cranes, cormor- (Figure 1) (see Badgerow, 1988; Speakman and Banks, 1998).
A ants, pelicans, flamingos, or other large birds is a spec-
tacular sight that gives rise to interesting questions. Why such
The trailing birds, in contrast with the leader, can make
substantial energy savings (Hummel, 1995; Weimerskirch et
regular formations? Most smaller birds that migrate in flocks al., 2001). Other species use more obtuse or bow formations
do so in less ordered groups (Alerstam, 1990). What is the (Hummel, 1995), in which the leading bird is only a little
advantage of V formation (Figure 1), and why do mainly large ahead of its nearest neighbors, the longitudinal distance
birds use them? At least two hypotheses may explain such between neighbors increasing along the arms of the forma-
formations: energy saving and communication (for review, see tion (Figure 3). In such formations, energy savings are
Alerstam, 1990; Speakman and Banks, 1998). probably more equable (Hummel, 1995). Why is there such
Energy saving is well supported both theoretically (see variation in formation shape?
Badgerow and Hainsworth, 1981; Hummel, 1995; Lissaman We suggest that the variation is related to flock kinship
and Shollenberger, 1970) and empirically (see Badgerow, structure and to reciprocity that depends on group size. Acute
1988; Cutts and Speakman, 1994; Hainsworth, 1987, 1988; V formations may often include relatives, in particular parents
Hummel, 1995; Speakman and Banks, 1998). Weimerskirch et and offspring and siblings. Gains in inclusive fitness might
al. (2001) showed that great white pelicans (Pelecanus make parents willing to accept a less favorable lead position, if
onocrotalus) have lower heart and wing beat rate, glide more their relatives will benefit. Reciprocity, with individuals taking
often, and save as much as 11–14% energy when flying in the turns at the lead position, may also be involved.
vortex wake from another bird (Figure 2).
Flight formation may also enhance communication and
orientation of the flock (see Cutts and Speakman, 1994;
Gould and Heppner, 1974; Hainsworth, 1988; Hummel, 1995; Energetics of flight formation
Speakman and Banks, 1998). The evidence for this function is In the wake of an aircraft, two counter-rotating trailing
weaker than for energy saving, but both mechanisms might vortices create air downflow behind the wingtips, and uplift
work together. outside the wake (see Hummel, 1995; Lissaman and Shollen-
Still, several interesting aspects of flight formations remain berger 1970) (Figure 2). The vortices are a grim reality; they
poorly understood, such as their variation in occurrence, size, can approach tangential velocities of 100 m/s and extend
and form. Theory suggests that large birds make greater many kilometers behind a heavy aircraft, horizontal tornadoes
energy savings than do small birds by formation flight (see with sometimes disastrous consequences for other planes that
Alerstam, 1990). Yet, not all large long-distance migrants use happen to fly into them. Under controlled conditions,
formation flight. Groups of raptors often soar high together however, a trailing plane can save much energy by suitable
with cranes or pelicans in thermal uplifts during migration, lateral positioning in the upwash behind the leading plane
but raptors rarely use formation flight. Other species often (Chichka et al., 1999; Hummel, 1973, 1995).
leave the uplift in conspicuous formation (Alerstam, 1990). Photographs of the wake of birds flying in clouds of helium
Why don’t raptors use flight formations? bubbles also show that birds leave two trailing vortices (see
Other questions concern formation shape. Swans, cranes,
Pennycuick, 1989; Spedding, 1987). The birds in a flock can
large geese, and other large birds often use acute V formation
therefore fly more cheaply in V formation, saving energy in
the uplift from the frontal neighbor (see Hummel, 1995;
Address correspondence to M. Andersson. E-mail: malte.andersson@ Lissaman and Shollenberger, 1970; for review, see Alerstam,
zool.gu.se. 1990; Norberg, 1990; Speakman and Banks, 1998). Lateral
Received 22 November 2002; revised 22 February 2003; accepted 12 distance strongly influences energy savings (Chichka et al.,
March 2003. 1999; Hummel, 1995; Lissaman and Shollenberger, 1970).
Behavioral Ecology vol. 15 no. 1
International Society for Behavioral Ecology 2004; all rights reserved.
Andersson and Wallander • Kin selection and reciprocity in flight formation? 159

Figure 2
A vortex is formed in the wake of each wingtip, creating downflow
behind the wing and uplift outside the wake, as indicated at the tip of
the right wing of the right-hand bird. A trailing bird can take
energetic advantage of this uplift by flying at a suitably lateral position
relative to the bird ahead. Theory suggests that the optimal wingtip
overlap for the trailing bird is about one tenth of the wingspan b.
A distance of about 0.78b separates the centres of the two trailing
vortices from a bird or aircraft.

tunnels (see Pennycuick et al. 1997) in species such as red

knot (Calidris canutus) (Kvist et al., 2001; Piersma et al., 1990).

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Theory suggests that if formation members differ in size,
Figure 1
energy savings will be particularly high for small light
V formation flights, the upper one with an obtuse formation angle, individuals, for example, juveniles, or females in species with
a .90 . In the lower figure, the greater spacing between birds in the sexual size dimorphism (Hummel, 1995).
longitudinal (flight) direction leads to an acute formation angle, Not all flock migrants use regular V formations or echelons
a ,90 . (see Alerstam, 1990; Heppner, 1974). In passerines and other
birds with short relative wing length, their rapid, large-
amplitude wing strokes create complex wake patterns that
may prevent energy saving by formation flight (Hummel,
Theory suggests that the lateral distance, d, between the 1995).
centres of the two trailing vortices is less than the wingspan,
b (Figure 2), the approximate relation being d ¼ bp/4 ¼ 0.78b
(Badgerow and Hainsworth, 1981). In theory, a bird gains the Kin selection and reciprocity
greatest benefits if its wingtip overlaps laterally with that of the We suggest that kin selection (Hamilton, 1964) plays a role in
bird in front (Figure 2), as described by the optimal wingtip flight formations of related birds. Although the leading
spacing (negative, as the wings overlap): Sopt ¼ (0.78b
b)/ position in acute V formation is energetically more expensive
2 ¼
0.11b (Badgerow and Hainsworth, 1981). Energy saving than are other positions, the leader may gain inclusive fitness
decreases rapidly with increasing lateral distance (Badgerow if the followers are close relatives such as offspring or siblings.
and Hainsworth, 1981; Hummel, 1973, 1995). Likely candidates are small flocks of geese, swans, cranes, or
The longitudinal distance between individuals that mini- other large birds that migrate in family groups of parents and
mizes the total energy expended by the flock is debated (see young (see Alonso and Alonso, 1993; Ely, 1993; Prevett and
Hainsworth, 1987; Higdon and Corrsin, 1978; Speakman and MacInness, 1980; Scott, 1980). The behavior of the leading
Banks, 1998), but it is probably less critical than lateral bird may then be a form of aerial parental care. The
distance (Chichka et al., 1999; Hummel, 1995; Speakman and advantage for young can be large if they are smaller and
Banks, 1998). For individual birds, however, energy saving can lighter than are adults (see above), as is usually the case (see
vary strongly depending on longitudinal distances (Figure 3). Cramp and Simmons, 1977).
In an acute V with long distances, the leading bird makes little Kin selection may also occur in flocks of several families
or no energy saving (Hummel, 1973, 1995; Weimerskirch et migrating together. In waterfowl with female-biased philopa-
al., 2001). A mathematical model of energy savings in relation try (Anderson et al., 1992), related females often breed near
to lateral and longitudinal position (Hummel, 1973) shows each other, and there is evidence that mothers, daughters,
that energy savings are more equable in bow formation, in and sisters can recognize each other as adults (see Andersson
which trailing neighbors are closer to the leader (Figure 3). As and Åhlund, 2000; van der Jeugd et al., 2002). In some arctic
there is a slight upwash preceding each bird, with short geese, wintering adults and offspring in extended family
longitudinal distances the leader can benefit from lift groups may contain members from more than two gener-
generated by the trailing neighbors. ations (Ely, 1993). Migrating flocks of swans or geese may
Owing to longitudinal variation in the ‘‘concertina-wake’’ consist of several related females and their mates, plus
from wing strokes (see Pennycuick, 1989; Spedding, 1987), offspring from several years (Warren et al., 1993).
and to potential effects of phase relationships between Reciprocation (Trivers, 1971) can occur in flight formations
neighbors, it seems likely that longitudinal position is more if individuals take turns as leader, sharing the energetic
critical in birds than in fixed-wing aircraft (see Hainsworth, disadvantage of the front position. Reciprocity can most easily
1987). This may be why there are often longitudinal distances evolve in small groups in which individuals interact repeatedly
of several bird lengths between neighbors in an acute V over long time (Axelrod and Hamilton, 1981; Boyd and
formation, although such long distances are probably un- Richerson, 1988; Trivers, 1971; for review, see Dugatkin,
favorable for the leader. A follower may fly most cheaply at 1997). Large group size tends to favor defecting individuals
some optimal point up to several bird lengths behind the that avoid costly behavior (Boyd and Richerson, 1988). Acute
neighbor ahead, at least in large species with high flight formations and reciprocity are therefore expected mainly in
speed, slow wing beats, and long distance between successive small stable groups with individual recognition. Recognition
equal-phase wake points. This aspect may be testable in wind facilitates punishment of a free-loader that avoids taking the
160 Behavioral Ecology Vol. 15 No. 1

expensive lead position, for example, by mobbing it in the air

or, perhaps more likely, at stopover sites. Reciprocity and kin
selection can have strong synergistic effects in small groups
(Axelrod and Hamilton, 1981; Boyd and Richerson, 1988)
and may be particularly likely in migrating flocks of extended
families. Such flocks may be teams in the sense of Anderson
and Franks (2001).
In large flocks of unrelated individuals, acute formation
shape that saves little energy for the leader may not be stable.
If the nearest neighbors lag behind at positions that benefit
them but not the leader, it may easily reduce flight speed so
far below the optimum that the trailing neighbors will catch
up. The leader therefore has an easy means of achieving
a more egalitarian obtuse formation. This applies also for
other birds in the formation relative to their nearest followers.
Such adjustments of longitudinal distances may contribute to
the dynamic nature of large formations of migrating eiders
(Somateria mollissima) and Branta geese, which may have
several local and varying leading points. Such points may
perhaps also contain related or familiar birds among which

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kin selection or reciprocity can operate locally within the
larger flock.
The scope for leader adjustment of the distance to followers
may be a major reason why obtuse V, echelon, or bow
formations occur in many birds flying in large flocks, for
instance, arctic geese (Branta and Chen), sea ducks such as
eiders (Alerstam, 1990), and also many waders (Piersma et al.,
1990). During the many 1000 km nonstop migrations over
open ocean by some waders (Piersma and Gill, 1998), energy
saving is probably crucial for each individual. Large flocks of
such long-distance migrants are therefore expected to fly in
energetically equable, obtuse formations (Figure 1) or bows in
which all individuals make similar energy savings (Figure 3).
No raptors use V formation, perhaps because selection for
hunting efficiency prevents dense flocking and therefore also Figure 3
formation flight. But raptors do not migrate in formations, Energy savings are unequally distributed among individuals in an
not even when leaving thermal uplifts at great height on acute V formation (top), the leading bird expending more energy
migration, when hunting is probably no option. Lack of flight than the others. A bow-shaped formation (bottom) is more
formation in raptors may in part depend on their lack of kin- egalitarian, and the frontal birds owing to uplift from their neighbors
related social structure after the breeding season, when they can make similar energy savings as the birds farther back in the
lead solitary lives. formation. The average energy gain is similar for both formations, as
the number of birds and the distance between wingtips are similar
(modified from Hummel, 1995).
Alternative explanations
The angle of a V formation may be related to relative neck Predictions and possibilities for tests
length for reasons of optical contact, long-necked birds such
as cranes and geese using acute formations, and short-necked The previous ideas predict that small flocks containing
species using more obtuse formations (Hummel, 1995). Neck relatives will often use acute formations in which the leader
length and communication, however, do not seem to explain saves little or no energy. Energetically more equable obtuse or
the longitudinal distances between neighbors in, for example, bow-shaped formations are predicted when there is little
geese, which are often much longer than a bird length (see possibility for kin selection or reciprocity among individuals
Speakman and Banks, 1998). that recognize each other.
Small birds will probably save relatively less energy by V These ideas are testable by analysis of the occurrence and
formation than do large birds, in which flight formations are shape of flight formations in relation to flock size and kinship
more common (see Alerstam, 1990). But there is no obvious structure. Formation shape is expected to differ between
reason why small birds might not benefit as much from flocks of small to moderate size, especially those containing
coordination and orientation by formation flight as do larger related individuals (e.g., geese and cranes) (Alonso and
birds. The lack of V formations in small flock-migrating Alonso, 1993; Prevett and MacInnes, 1980), and larger flocks
passerines therefore suggests that communication is not the of mostly unrelated individuals, such as spring-migrating
major function of formation flight. waders and arctic geese (see Alerstam, 1990; Piersma et al.,
Formation shape may depend on body size if the slower 1990). To critically test these ideas, quantitative observations
wing strokes of large birds permit closer approach to optimal of formation shape are needed, which may be possible to
positions in a formation (see Hummel, 1995), perhaps obtain at suitable migration stations or in overwintering areas
explaining why acute V formation may be more common in, (see Speakman and Banks, 1998).
for example, large geese than in small geese (Badgerow and Age and sex distribution are expected to influence
Hainsworth, 1981; Speakman and Banks, 1998). Another formation shape. Homogeneity in age, sex, and body size is
possible reason is that smaller species tend to occur in larger expected to favor more egalitarian formations than do mixed
flocks, in which kin selection or reciprocity seem less likely to flocks of, for example, adults and young. Offspring of the year
favor individuals that accept an expensive leader role. are often smaller than parents and may be less able to
Andersson and Wallander • Kin selection and reciprocity in flight formation? 161

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