|
The Ecology, Evolution, and Behavior of Keas and Kakas
Keas
(Nestor notabilis) are omnivorous olive-green parrots
endemic to alpine scrub and mountain beech forests on the
South Island of New Zealand. They are compulsively neophilic, fearless,
persistent, and ingeniously destructive. They congregate around
rich food resources, displaying a complex, stratified social system;
juveniles associate with both related and unrelated adults for several
years after fledging, following them around and apparently imitating
their foraging behavior. And they show more elaborate and ritualized
social play than any other species of bird. But what primarily attracted
us to the species was the suggestion that they were "open program"
animals, species that were specialized for learning and that displayed
an unusual ability to adapt to changing conditions and circumstances.
Social
Foraging and Ontogeny
We
conducted three seasons of field observations on a banded population
of keas at a rubbish dump outside of Arthur’s Pass National
Park, obtaining quantitative recordings of foraging and social behavior
from 38 keas of both sexes, including substantial numbers of all
four distinguishable age classes – fledglings, juveniles,
subadults, and adults. We used hypergeometric probabilities to define
dissimilarities between behavioral events and compared activity patterns
across sex and age classes with principal coordinates analysis. Keas displayed
characteristic differences according to sex and age in foraging
ability and in the social behavior used to obtain access to resources.
Adult males performed most of the excavation that uncovered new
food resources. Fledglings explored and manipulated objects almost
continuously, but they discovered little food on their own and were
commonly fed directly by adults. Juveniles obtained the highest
foraging yields for the amount of time spent searching of any age
class, aided by appeasement behavior that gave them preferential
access to foods discovered by adults. Kleptoparasitism served as
a primary foraging strategy for subadults, who were otherwise commonly
displaced from any food resources they discovered. Females were
subordinate to males of any age and primarily fed by gleaning small
items from around the edge of the aggregation. We concluded that
social factors influenced the acquisition and display of foraging
expertise in this species in different ways at different stages
of development, and that social transmission during group foraging
was a far more complex phenomenon than had been hypothesized
in the literature. Our initial field study
(Diamond & Bond 1991)
was
broadened into a book-length monograph on kea behavior, biology, and
evolution, published by the University of California Press
(Diamond & Bond 1999).
Comparative
Studies of Social Play
In
our monograph, we argued that many of the more unusual features of
kea behavior could be considered as specific adaptations to their
harsh and unreliable habitat in the high alpine regions, which suggested
a fertile ground for subsequent comparative research. The species
most closely related to the kea is the kaka (Nestor meridionalis),
a parrot from the more equable, lowland temperate rainforests of
New Zealand. Because of their close phylogenetic relationship, one might expect
keas and kakas to display similar behavior patterns, but kakas sociality
is strikingly different. Kakas show more ritualized behavior patterns, particularly
in the context of aggression, and juvenile kakas are independent of adults
at an earlier age. And although kakas display very complex and sophisticated
foraging techniques, at least as adults, they show much less interest in novel
food items. These observations provided the basis for several additional years
of field work, in which we made extensive recordings of social play from both
keas and kakas, examining the two species differed in
behavioral flexibility. A quantitative comparison of play behavior
in keas and kakas was published in Behaviour. We also
published a larger review article on the incidence of social play
in birds, testing for differences between avian taxa, as well as
for correlations between play complexity, brain size, and age of
first reproduction. We found that several bird families with large
numbers of playful species -- corvids, parrots, and hornbills --
had larger relative brain sizes than would be predicted from a class-level
allometric regression, but brain size was not associated with the
complexity of social play among genera within taxa. Play complexity
within parrots and corvids was, however, significantly associated
with the age of first reproduction. The likelihood of complex social
play appeared to increase when delayed reproduction is accompanied
by persisting relationships between adults and post-fledging juveniles,
an intriguing parallel to the incidence of social play in mammals. This line
of research was published in Diamond and Bond
(2003 and
2004) and
Diamond et al.
(2006).
Vocal
Dialects in Keas and Kakas
We
have recorded vocalizations, both spontaneous calls and responses
to playback, at a number of locations throughout the range of both
keas and kakas, documenting the vocal repertoires of the species
and quantifying their geographic variation. The initial results
have been exceedingly interesting, suggesting that keas and kakas
exemplify two distinctive patterns of parrot sociality, patterns
that are echoed in the behavior of a broad range of other parrot
species. The initial work from this research program, on vocal dialects
in keas, was published in
Bond & Diamond 2005,
and a detailed description of vocal repertoires in these species is
included in our book on cognition and behavior in wild parrots
(Bond & Diamond 2019),
placing such divergent strategies in the broader context of the entire group.
|