The Forehead Patch of Tufted Titmice:
A Possible Status Signal
Shawn P. Moses and Gary Ritchison
Department of Biological Sciences
Eastern Kentucky University
Richmond, KY 40475 USA
Abstract.-- Among birds, morphological features, called status
signals or badges, may provide information about relative fighting ability
and, as such, permit birds to avoid costly fights. Tufted Titmice (Baeolophus
bicolor bicolor) have black forehead patches that vary in size, and
our objective was to determine the possible relationship between patch
size and dominance status in winter flocks in central Kentucky. Patch width
was significantly correlated with dominance rank, with higher ranking birds
having wider patches. Such results suggest that the forehead patch of titmice
might serve as a status signal, particularly during encounters between
unfamiliar individuals. Variation in patch size may also be an important
intersexual signal.
Status signals are morphological features that convey
information about relative fighting ability. These signals may allow assessment
of potential competitors and permit birds to avoid costly fights with dominant
individuals (Rohwer 1975). In birds, prominent plumage patterns often function
as status signals and such signals have been reported in several species,
including the House Sparrow (Passer domesticus; Møller 1987)
and Dark-eyed Junco (Junco hyemalis; Yasukawa and Bick 1983). Status
signals have also been reported among species in the family Paridae, including
Great Tits (Parus major; Jarvi and Bakken 1984, Wilson 1992) and
Willow Tits (P. montanus; Hogstad and Kroglund 1993). Among North
American parids, Otter and Ratcliffe (1999) found that bib morphology varies
among Black-capped Chickadees (Poecile atricapillus), but the signal
value of such variation remains unclear. Similarly, Tufted Titmice (Baeolophus
bicolor bicolor) have black forehead patches that vary in size among
individuals. The objective of our study was to determine if the size
of these forehead patches is correlated with dominance status in winter
flocks of titmice in Kentucky.
During the non-breeding season, Tufted Titmice occur
in small flocks of 2 - 5 birds that typically consist of a pair of adults
plus a variable number of juveniles (offspring and juveniles from unknown
natal sites) (Grubb and Pravosudov 1994, Pravosudova et al. 1999). Dominance
hierarchies develop within these flocks and flock members may also interact
with titmice from other flocks (Brawn and Samson 1983, Grubb and Pravosudov
1994). Thus, as with other flocking species, Tufted Titmice might benefit
from the presence of a status signal.
METHODS
We studied titmice at the Central Kentucky Wildlife
Management Area (CKWMA), located 17 km southeast of Richmond, Madison County,
Kentucky. Beginning in September 1996, six feeding stations were stocked
with sunflower seeds. From October 1996 through February 1997, titmice
visiting these stations were captured in mist nets and banded with numbered
aluminum bands plus a unique combination of colored bands. The size
of the forehead patch (top width, bottom width, and height) was also determined.
From December 1996 - March 1997, titmice were observed at feeding stations
to determine the dominance relationships among flock members. During interactions
at feeders, an individual was considered dominant if it either chased or
supplanted another titmouse or if another titmouse waited nearby and did
not visit the feeder until the dominant bird left.
RESULTS
We captured 29 titmice, and were able to determine
the age of 13 (4 hatching year and 9 after-hatching year) (Pyle et al.
1987). AHY birds had longer (Mann-Whitney U test, P = 0.02) wing chords
(mean length = 81.3 mm) than HY birds (mean length = 76.8 mm), but AHY
and HY birds did not differ (P > 0.15) in tarsus length, tail length,
mass, or any measure of patch size (top width, bottom width, height, or
total area).
Fifteen titmice were observed throughout the study
and formed five flocks, with flock size ranging from 2 to 5 birds. Dominance
hierarchies in titmouse flocks are linear peck-right, with each member
of a flock consistently either dominant or submissive to other flock members
(Brawn and Samson 1983). For all flocks combined, we observed an average
of 2.9 interactions per dyad and, based on these interaction, assigned
each flock member a dominance rank. We found a significant correlation
between rank and patch width (measured along the top of the patch), with
higher ranking birds having wider patches (rs = 0.55,
n = 15, P = 0.032). Correlations between the width of the bottom of the
patch and rank and between patch area and rank, respectively, approached
significance, with higher ranking birds having both wider (rs
= 0.44, n = 15, P = 0.099) and larger patches (rs
= 0.47, n = 15, P = 0.07). The height of the forehead patch was not
correlated with rank (rs =0.002, n = 17, P =
0.99).
DISCUSSION
The significant correlation between the width of
the forehead patch of Tufted Titmice and dominance status suggests the
patch may serve as a status signal. However, the results of previous
studies of titmice also suggest that variation in patch morphology may
be important only during intrasexual interactions with unfamiliar individuals
and as an intersexual signal. Several factors appear to support this hypothesis.
First, dominance status within flocks of titmice is typically age- and
sex-related, with males dominant to females and, within a sex, adults dominant
to juveniles (Brawn and Samson 1983). If so, and assuming that male titmice
were the top-ranked individuals in the flocks observed during our study,
the observed correlation between patch size and status in our study may
have due, in part, to sex-related differences in plumage. In addition,
flocks of titmice are stable (Samson and Lewis 1979) and often composed
of parents and offspring (Pravosudova et al. 1999). As a result, flock
members typically know, and know the status of, other flock members, making
status signals unnecessary.
However, even badges that reflect age or sex may
serve as a signal of potential fighting ability among unfamiliar individuals
(Lemel and Wallin 1993). Thus, status signaling may be important during
encounters between members of different flocks because they might not remember
the fighting ability of birds with which they interact only rarely (Jarvi
and Bakken 1984). Because they interact occasionally with conspecifics
from other flocks and with satellite birds with no flock membership (Brawn
and Samson 1983, Grubb and Pravosudov 1994), titmice might benefit from
the presence of a status signal.
Morphological variation in the forehead patch of
Tufted Titmice may also be important as an intersexual signal. For example,
Norris (1990a, b) found that female Great Tits may use the breast stripe
of males as a criterion for mate choice, and may benefit from such discrimination
because large-striped males are more attentive to the nest and more likely
to defend the brood. Similarly, female House Sparrows prefer large-bibbed
males as mates, and such males tend to obtain better territories and nest
sites (Møller 1988). Additional study is needed to determine the
possible role, if any, of the forehead patch in the mate choice decisions
of male and female Tufted Titmice.
ACKNOWLEDGMENTS
We thank Joel Beverly for help with the field work,
and the Eastern Kentucky University Research Committee for financial support
.
LITERATURE CITED
Brawn, J. D. and F. B. Samson. 1983. Winter behavior of Tufted Titmice. Wilson Bull. 95:222-232.
Grubb, T. C., Jr. and V. V. Pravosudov. 1994. Tufted Titmouse (Parus bicolor). In The Birds of North America, No. 86 (A. Poole and F. Gill, eds.). The Academy of Natural Sciences, Philadelphia, PA; The American Ornithologists’ Union, Washington, DC.
Hogstad, O. and R. T. Kroglund. 1987. The throat badge as a status signal in juvenile male Willow Tits, Parus montanus. J. Ornithol. 134:413-423.
Jarvi, T. and M. Bakken. 1984. The function of the variation in the breast stripe of the Great Tit (Parus major). Anim. Behav. 32:590-596.
Lemel, J. and K. Wallin. 1993. Status signalling, motivational condition and dominance: an experimental study in the Great Tit, Parus major. Anim. Behav. 45:549-558.
Møller, A. P. 1987. Variation in badge size in male House Sparrows Passer domesticus: evidence for status signalling. Anim. Behav. 35:1637-1644.
Møller, A. P. 1988. Badge size in the House Sparrow, Passer domesticus. Effects of intra- and intersexual selection. Behav. Ecol. Sociobiol. 22:373-378.
Norris, K. J. 1990a. Female choice and the evolution of conspicuous coloration of monogamous male Great Tits. Behav. Ecol. Sociobiol. 26:129-138.
Norris, K. J. 1990b. Female choice and the quality of parental care in the Great Tit Parus major. Behav. Ecol. Sociobiol. 26:275-281.
Otter, K. and L. Ratcliffe. 1999. Relationship of bib size to age and sex in the Black-capped Chickadee. J. Field Ornithol. 70:567-577.
Pravosudova, E. V., T. C. Grubb, Jr., P. G. Parker, and P. F. Doherty, Jr. 1999. Patch size and composition of social groups in wintering Tufted Titmice. Auk 116:1152-1155.
Pyle, P., S. N. Howell, R. P. Yunick, and D. F. DeSante. 1987. Identification
guide to North American passerines. Slate Creek Press, Bolinas, CA.
Samson, F. B. and S. J. Lewis. 1979. Experiments on population regulation
in two North American parids. Wilson Bull. 91:222-233.
Watt, D. J. 1986. Relationship of plumage variability, size and sex to social dominance in Harris’ Sparrows. Anim. Behav. 34:16-27.
Wilson, J. D. 1992. A re-assessment of the significance of status signalling
in populations of wild Great Tits, Parus major. Anim. Behav. 43:999-1009.
Yasukawa, K. and E. I. Bick. 1983. Dominance hierarchies in Dark-eyed
Juncos (Junco hyemalis): a test of a game-theory model. Anim. Behav.
31:439-448.
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