Zo Wreggitt

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ADAPTATIONS OF WINTER ACCLIMATIZED BLACK-CAPPED CHICKADEES (Poecile

atricapilllus)

The black-capped chickadee (Poecile atricapilllus) is a common passerine bird found

throughout much of North America (Smith, 1991). They are non-migratory and thrive in a
diverse range of habitats (Smith, 1991). This broad range is due to the ability for chickadees to
withstand temperatures as low as -50C (Sharbaugh, 2001), decreased food supply, and reduced
foraging time (Fig. 1) (Kessel 1976). P. atricapillus have numerous strategies that allow them to
overwinter in extreme conditions including physical, behavioural and physiological adaptations.
Physical Adaptations
To survive the winter birds will change various physical features including fat scores and
plumage. Chickadees are small birds, weighing between 10-14g (Kessel, 1976). This small size
indicates that they experience large energy expenses (Kleiber in Sharbaugh, 2001). Some bird
species may increase body mass and abdominal fat scores to survive cold winter temperatures.
However, a study conducted by Cooper and Swanson (1994) found that there was no significant
difference in mass and fat scores between summer and winter chickadees. The summer birds
weighed in at 13.1 0.7g (n = 50) and the winter birds at 13.0 .09g (n = 60). There was no
significant difference in furcular visible fat scores and abdominal visible fat scores. The most
effective physical adjustment that chickadees experience to survive winter appears to be from the
increase in feather insulation after molting (Rising and Hudson, 1974). The efficiency of
insulation from plumage is quite outstanding as there is at least a 40C temperature difference
between the skin of P. atricapillus and the environment (Hill et al. 1980).
Behavioural Adaptations
Chickadees have many behavioural adaptations to winter including flocking, length of
foraging period, roosting location/posture, and food caching. Throughout the year, chickadees

remain with their breeding mate, however, once the breeding season is over, chickadees form
flocks (Odum, 1942). These flocks range in size and vary in numbers from day to day (Odum,
1942). Kessel (1976) found that on cold days the feeding range of flocks are reduced. Chickadee
flocks experience a reduced feeding time if: temperatures are extremely low; it is snowing; there
is high cloud cover; it is windy (Kessel, 1976).
During the breeding season chickadees roost in nest cavities, however, in winter they
roost in the upper dense foliage of conifers more often than cavities (Odum, 1942). Smith (1991)
suggests that the cavities that chickadees roost in over winter are smaller than the ones that they
breed in. Along with finding suitable roosting sites, Smith (1991) suggests that the posture of
chickadees when roosting can reduce heat loss by having the head inclined forward with the beak
covered by shoulder feathers.
P. atricapillus also store food throughout the fall and winter (Kessel, 1976). A study by
Petit and Petit (1989) found that for food caching to be beneficial, chickadees must be able to
locate hidden food before cache robbers steal it, environmental conditions prevent chickadees
from retrieving it, and before chickadees forget location of cached food. Chickadees are scatterhoarders, meaning that they store food in a variety of different places within 40m of the food
source. They do not protect their caches from other birds, instead they depend on the reduced
chance of cache robbers discovering all of their cache sites. Petit and Petit (1989) also found
that chickadees strategically placed 58% of their forages on the underside of branches and twigs
to reduce snow and ice cover. A food cache that proved to be unique to chickadees, in this study,
was the storage of seeds (8%) in dead leaves that remained on deciduous trees. Chickadees also
have to remember where their cache sites are located. Relocation requires memory of spatial

locations which relies on the hippocampus and neuron recruitment that occurs in the fall
(discussed below) (Barnea and Nottebohm, 1994).
Physiological Adaptations
Chickadees have two fascinating physiological adaptations to winter acclimatization:
nocturnal hypothermia and hippocampal neuron recruitment. Chaplin (1974) showed that
chickadees are able to survive harsh winter conditions without significant fat storage due to their
ability to reduce their metabolism at night. The low amount of energy that P. atricapillus expend
at night occurs because chickadees experience nocturnal hypothermia. As a result, chickadees
keep their body at temperatures 10-12C lower than their temperature of 41C throughout the
day (Sharbaugh, 2001).
Another study conducted by Chaplin (1976) showed that once hypothermia is achieved in
P. atricapillus, it occurs in short bursts throughout the night. These short bursts consist of
shivering actions that are controlled by the flight muscles. Shivering throughout the night
provides chickadees with heat, but it is costly as it consumes large amounts of the fat store.
Many bird species go into moderate hypothermia when their energy sources are running low due
to a small forage. However, P. atricapillus intentionally induce hypothermia on cold winter
nights as a defence mechanism opposed to a last-chance survival mechanism. Another cost of
hypothermia is the effect that it has on alertness. Small birds need to be ready for flight at all
times to escape predators. P. atricapillus are one of the few winter acclimatized species that can
fly while in hypothermia. (Chaplin, 1976)
As mentioned earlier, the hippocampus is important in retrieving food caches. The study
by Barnea and Nottebohm (1994) showed that new neurons are constantly being created in the

hippocampus of adult chickadees. However, they also noted a significant increase in the
recruitment of neurons in the fall. This suggested that the increase in neurons was due to the need
for chickadees to generate precise spatial memories for the upcoming lifestyle change as a result
of colder temperatures. In fact, the more severe the weather conditions, the higher the number of
new hippocampal neurons recruited (Freas et al. 2013).
In summary, chickadees have numerous physical, behavioural, and physiological
adaptations that allow them to survive winter. The most outstanding of these adaptations is likely
the new recruitment of neurons in the hippocampus. Numerous studies are being executed to
further understand how the hippocampus affects the spatial memory of P. atricapillus.

References
Barnea, A. and Nottebohm, F. 1994. Seasonal recruitment of hippocampal neurons in adult freeranging black-capped chickadees. Proc. Natl. Acad. Sci. USA. 91: 11217-11221.
Chaplin, S. B. 1974. Daily Energetics of the Black-capped Chickadee, Parus atricapillus, in
Winter. Journal of Comparative Physiology. 89: 321-330.
Chaplin, S.B. 1976. The Physiology of Hypothermia in the Black-capped Chickadee, Parus
atricapillus. Journal of Comparative Physiology. 112: 335-44.
Cooper, S. J., Swanson, D. L. 1994. Seasonal acclimatization of thermoregulation in the blackcapped chickadee. The Condor. 96: 638-646.
Freas, C. A., Roth, T. C., LaDage, L. D., Pravosudov, V. V. 2013. Hippocampal neuron soma size
is associated with population differences in winter climate severity in food-caching
chickadees. Functional Ecology. 27: 1341-1349. doi: 10.1111/1365-2435.12125
Hill, R. W., Beaver, D. L., and Veghte J. H. 1980. Body Surface Temperatures and
Thermoregulation in the Black-Capped Chickadee (Parus atricapillus). Physiological
Zoology. 53(3): 305-321.
Kessel, B. 1976. Winter activity patterns of Black-capped Chickadees in interior Alaska. The
Wilson Bulletin. 88(1): 36-61.
Odum, E. P. 1942. Annual Cycle of the Black-Capped Chickadee: 3. The Auk. 59(4): 499-531.
Petit, D. R. and Petit L. J. 1989. Winter caching ecology of deciduous woodland birds and
adaptations for protection of stored food. The Condor. 91: 766-776.
Rising, J. D., and Hudson, J. W. 1974. Seasonal variation in the metabolism and thyroid activity
of the Black-capped Chickadee (Paurs atricapillus). The Condor. 76: 198-203.

Sharbaugh, S. M. 2001. Seasonal Acclimatization to Extreme Climatic Conditions by BlackCapped Chickadees (Poecile atricapilla) in Interior Alaska (64N). Physiological and
Biochemical Zoology. 74(4): 568-575.
Smith, S. M. 1991. The Black-capped Chickadee. Cornell University Press. Ithaca, NY.

Figures
40

THE

WILSON

BULLETIN

Vol. 88, No. 1, March 1976

FIG. 1. Times of first and last activity of chickadees recorded from 1960 through 1967.
Times have been adjusted for longitude and are true solar times. The dotted lines
represent the times of the beginning of morning Civil Twilight and ending of evening
Civil Twilight, respectively; the solid lines are best-fit polynomial curves, adjusted
somewhat by eye. (from Kessel 1976)

autumn, morning activity began after the beginning of Civil Twilight.

Within

a few days of 1 November, first activity began to occur prior to the beginning
of Civil Twilight;

and, except during occasional periods of cold stress (see

below), chickadees generally continued to become active before the beginning

of Civil Twilight

until sometime during the first few days of January, after

which they again usually remained at roost until after the beginning of Civil
Twilight.

(During

the unusual winter of 1960-61,

when temperatures were

relatively warm and cloud cover relatively low, chickadees continued to become
active before the beginning of Civil Twilight until the third week of January.)
Activity ceased prior to the end of Civil Twilight throughout the year, and it
ceased even before sunset from sometime during the first half of February
until mid-September.
Regression analyses showed that the daily light cycle (= time of beginning
and ending of Civil Twilight)
affecting the chickadees

is by far the dominant environmental

activity cycle (P < 0.001).

factor

The time of 249 first

activity observations made between September and May showed a 93.1% correlation with the time of the beginning of morning Civil Twilight and a 96.0%
correlation of 193 last activity observations over the same period with the end-