Dwight (1900) says of the Tree Swallow (Tachycineta bicolor): "The
female has corresponding plumages and moults [to those of the male],
but is usually duller with less iridescence and browner wings and tail
until adult [=second] winter plumage is assumed which is usually indis-
tinguishable from that of the male." The same sequence of plumages
is indicated by Low (1933): "The dull, brown birds and those with a
faint tinge of green appear to be one-year old females. The male obtains
its brilliant plumage the first year, but the female apparently does not
acquire it until the second year. The female may be slightly duller, but
frequently she is indistinguishable from the male except when both are
held together in the hand."
Although the relationship between age and plumage color appears
from the above quotations to have been worked out more than 50 years
ago, some doubt remains about the accuracy of ageing females as year-
lings or older by these methods, as was done by DeSteven (1978). For
example, Kuerzi (1941) reported 5 one-year-old females (banded as
nestlings) in the green-tinged brown plumage, but one brown female
also returned the following year in the green-tinged brown plumage
indicating that fully iridescent upper parts are not always acquired by
the second breeding season. On the other hand Chapman's (1955) de-
scription of 1 of 5 one-year-old females as "dull blue" indicates that a
predominantly blue iridescent plumage may be acquired by the first
breeding season.
These uncertainties are partially reflected in the 1980 revision to the
key to age and sex of Tree Swallows in the North American bird banding
manual (BBM) (Canadian Wildlife Service and U.S. Fish and Wildlife
Service 1977) in which breeding adults with brown upper parts are
classified as yearling females (SY-F in banding terminology--see Meth-
ods) but no provision is made for ageing predominantly blue-plumaged
females as older than yearling (ASY).
Can female Tree Swallows be aged accurately by plumage color? Here
I present evidence that most breeding females can be classified as year-
lings or as older than yearlings with better than 95% accuracy, but a
small percentage must be assigned to unknown age. A key for ageing
and sexing adult Tree Swallows in the breeding season is also provided.
METHODS
Nest boxes for Tree Swallows have been maintained by Long Point
Bird Observatory at Long Point, Ontario (42ø33'N, 80ø04'W) since be-
fore 1970 and at 2 nearby mainland sites since 1975 and 1977. Nearly
all young were banded each year. Adults were captured in the nest
boxes, usually during the nestling stage, and were sexed by presence of
a brood patch (females) or cloacal protuberance (males). Data are pre-
sented here on plumage color of 171 adults banded as nestlings in 1970-
1981 which were recaptured as breeding females in 1971-198:2 and
generated :288 plumage-age records. Data were collected by the author,
other researchers, and several experienced and inexperienced assistants.
Color of the surface of the body plumage of the upper parts of females
was classified as Brown, Intermediate, or Blue (capitalized plumage de-
scriptions below represent these categories). In the early years of the
study females with predominantly brown or greenish-tinged brown upper
parts were called Brown, those with completely or almost completely blue
or blue-green iridescent upper parts were classified as Blue, and others
which did not fit these categories were fully described. The latter birds
were later assigned to the appropriate categories, which by 1978 were
defined as: Brown = up to 50c of the surface of body plumage of the
upper parts consists of iridescent blue or blue-green feathers (i.e., 50c/c
or more is made up of brown or greenish-tinged brown feathers); In-
termediate = more than 50c and up to 90c/c blue or blue-green irides-
cent feathers; and Blue = more than 90% blue or blue-green iridescent
feathers.
Age designations follow bird-banding terminology: SY = second cal-
endar year of the bird's life, TY = third calendar year, AHY = after
hatch year (i.e., SY or older), ASY = after second year (i.e., older than
SY), ATY = after third year. Note that an SY bird is 1 year old and is
in its first breeding season, a TY bird is :2 years old, and that AHY,
ASY, and ATY birds are at least 1, :2, and :3 years old, respectively. M =
Male, F = Female, U = Unknown (age or sex). "Adults" are birds oœ
breeding age (SY or older).
RESULTS
Variation in plumage color with age in female Tree Swallows is shown
in Table 1. Of :288 age-color records 67 (:23.3) were Brown, 18 (6.:2)
were Intermediate, and :203 (70.5c/r) were Blue. Although this distri-
bution depends partly on the age structure in our colonies, which may
not be representative of the overall population, it is clear that Inter-
mediates are rare.
Among Brown females 64 of 67 records (95.5%) were SY birds and
the remaining 3 were TY. The 18 Intermediates were made up of 9
SY, 8 TY, and 1 fourth year bird. Only 3 of :203 Blues (1.5%) were SY,
and the remainder were ASY with ages ranging from :2 to 7 years. Of
113 ATY birds (3 to 7 years old), 11:2 were Blue and 1 (a fourth year
bird) was Intermediate.
These results suggest that Brown and Blue females can be aged SY
and ASY with better than 95c confidence. Intermediates are usually
either 1 or :2 years old and in banding terminology must be assigned to
the AHY category.
Plumages of known-age individuals caught in successive years (Table
:2) show that one-year-old (SY) Browns and Intermediates usually change
TABLE 1. Breeding season plumage color of female Tree Swallows banded as nestlings.
Color of Number of birds of age (years)
upper parts 2 1 2 3 4-7
Brown 64 3
Intermediate 9 8 1
Blue 3 88 57 553
' Total individuals = 171.
2 See text for definitions.
55 plumage-age records representing 39 individuals: 4 years = 31, 5 years = 18, 6
years = 5, 7 years = 1.
to Blue by the following breeding season. Two-year-old Browns and
Intermediates are nearly always Blue at 3 years, but 1 Intermediate did
not change between 2 and 3 years. Although I have not analyzed all of
our data on females banded as breeding adults, it is worth noting that
we have a record of 1 individual which was banded as an Intermediate
(about 80% blue upperparts), in 1976 and was still in Intermediate
plumage in 1982 when it was at least 7 years old. This emphasizes that
AHY is the appropriate age designation for Intermediates.
DISCUSSION
The data presented here show that about 94% of breeding female
Tree Swallows can be aged SY or ASY with better than 95% accuracy
by the color of the upper parts alone. This conclusion is supported by
detailed plumage color data on known-age females in Colorado (Cohen
1980 and pers. comm.).
Separation of females into the categories Brown, Intermediate, and
Blue, for ageing purposes, is somewhat arbitrary as there appears to be
a more or less continuous series of plumages from dull brown to bright
iridescent blue-green, although some plumages are rarer than others.
The following additional comments on plumages may aid assignment
of females to the correct plumage category.
Brown females often have faint green iridescent tips to the brown
feathers and many have scattered feathers or patches of feathers which
are bright iridescent blue-green, commonly on the crown, upper back,
wing coverts, or upper tail coverts. Intermediates have at least 50% and
up to 90% of the surface of the upper parts of the body plumage con-
sisting of the bright iridescent blue-green feathers. Typical Interme-
diates have patches of brown feathers (lacking the bright blue-green
iridescence) on the forehead, nape, and rump. Many Blue females are
easily categorized as they are in bright blue-green plumage which is
indistinguishable from that of males; but some have a few feathers
lacking blue-green iridescence, usually on the forehead and sometimes
on the nape or elsewhere. If there is doubt about the percentage of
blue-green iridescent feathers, it is safest to call females Intermediate,
as this leads to age AHY (see key below).
TABLE 2. Plumage changes between breeding seasons in known-age female Tree
Swallows.
Plumage color at age No. of
1 year 2 years 3 years birds
Brown Intermediate -- 1
Brown Blue -- 24
Intermediate Intermediate -- 1
Intermediate Blue -- 2
Blue Blue -- 3
Brown Blue 1
Intermediate Intermediate 1
Intermediate Blue 3
Blue Blue 33
The BBM (1980 revision of Tree Swallow age-sex key) uses 2 addi-
tional characters to age and sex Tree Swallows: (1) degree of skull
pneumatization (birds with small "windows" are designated SY in the
breeding season), and (2) color of forehead and feathers at base of
nostrils (birds with brown feathers in these areas are called female, others
unknown sex).
We have not used skull pneumatization for ageing breeding Tree
Swallows at Long Point, but presumably this character would improve
age discrimination of males, Intermediate females, and those Blue fe-
males which are actually SY. The BBM key does not indicate what
percentage discrimination of SYs may be expected from the skull pneu-
matization criterion, but in the Bank Swallow (Riparia riparia) Freer and
Belanger (1981) found that 17% of 18 SY birds retained small "win-
dows" in their first breeding season. They advocate use of paired win-
dows at least 2 mm in diameter as the criterion for ageing birds as SY,
as single or paired "pinholes" were sometimes retained by older birds.
A similarly cautious approach for ageing adult Tree Swallows by this
method will be adopted here in view of the lack of data on the rate of
skull pneumatization in this species. Although it is not usually necessary
to use plumage color to sex breeding Tree Swallows, our experience
agrees with the BBM in that birds with brown feathers on the forehead
and at the base of the nostrils are females.
Based on the results presented here and the 2 additional characters
in the BBM, the following key has been constructed for ageing and
sexing adult Tree Swallows in the hand during the breeding season
(May-July).
First check for presence of brood patch (F) or cloacal protuberance
(M). If present, these should be consistent with sex designations derived
from the key.
la. Brown upper parts (0-50% of surface of upper parts of body plumage iridescent
blue-green, remainder of upper parts dull brown or brown lightly tinged with
green iridescence) ............................................... SY-F
lb. Intermediate upper parts (>50-90cA of surface of upper parts of body plumage
iridescent blue-green; remainder of upper parts brown or brown lightly tinged
with green. Brown areas most often on forehead, nape and rump) ...... F (see 4)
lc. Blue upper parts (>90-100% surface of upper parts of body plumage blue-green;
remainder of upper parts brown or brown lightly tinged with green. Brown areas
most often on forehead and nape) .................................. see 2
2a. Forehead ad feathers at base of nostrils brown ................. F (see 4)
2b. Forehead and feathers at base of nostrils blue, green, or black ........ see 3
Brood patch present .......................................... F (see 4)
Cloacal protuberance present ................................... M (see 4)
Not as 3a or 3b .......................................... Sex U (see 4)
4a. Small paired unpneumatized "windows" (>-2 mm diameter) at rear sides of
skull ....................................................... SY
4b. Skull pneumatization more advanced than 4a or not determined ....... see 5
5a. Blue female (from 2a or 3a) ..................................... ASY-F
5b. Not as 5a (from lb, 3b or 3c) ..................................... AHY
Because Tree Swallows in juvenal plumage are dull sooty-brown above
(Dwight 1900), care must be taken late in the breeding season to dis-
tinguish SY-F from HY birds. With experience the sooty-brown upper
parts (lacking iridescence) and dusky-white under parts of HY birds are
easily recognizable in comparison with the worn and faded plumage of
Brown SY females. By late July at Long Point, however, even some HY
birds have quite worn and faded plumage.
The key given above has certain implications for ageing and sexing
Tree Swallows during and after the postjuvenal and postnuptial molts.
Both molts take place before fall migration and are complete; moreover
the birds acquire plumages which are essentially identical to those of
the following breeding season except that the new tertials are broadly
tipped with white (Dwight 1900). Thus after the molts, birds in fresh
plumage can be aged and sexed as follows: Brown birds are HY-F;
Intermediates are either HY-F or SY-F and must be aged by skull pneu-
matization; Blue birds with brown foreheads and brown feathers at the
base of nostrils are AHY-F, but those which retain small "windows"
in the skull can be called SY-F; other Blue birds are either U-M or
AHY-F so must be called U-U unknown, except that age can be designated
HY, SY, or AHY by skull pneumatization and HY birds are males.
During the molt in late summer and early fall, birds changing from
sooty-brown (juvenal) plumage to new Brown plumage and new Blue
plumage are HY-F and HY-M, respectively. Birds changing from Brown
to Blue are SY-F. A bird changing from old Blue to new Blue plumage
is either AHY-M or ASY-F and must be called AHY-U unless "windows"
in the skull indicate that it is SY-M or traces of brown plumage on the
forehead and above the nostrils allow it to be sexed F (but it should be
aged AHY--see below). Intermediate plumages are unlikely to be ad-
equately distinguishable from Blue plumages during molt, but 10-15cA
of all females are likely to be Intermediate in either the old or new
plumage. Birds entering Intermediate plumage from sooty-brown and
Brown plumage are HY-F and SY-F, respectively. As Intermediates in
3a.
molt may be misidentified as Blue and are likely to be identified as female
by presence of brown plumage on the forehead, I would expect them
to make up more than 15c/c of this group of females. Therefore females
which appear to be changing from old Blue to new Blue plumage should
be aged AHY unless the presence of small "windows" in the skull in-
dicates that they are SY. These age-sex designations based on plumages
during molt should not be attempted unless one or both plumages can
be clearly distinguished. Ageing by skull pneumatization will usually
help in determining sex by narrowing the possibilities, but is often dif-
ficult if the head feathers are in molt.
SUMMARY
Based on 288 plumage-age records from 171 breeding female Tree Swallows, most females can be classified by plumage color as SY or ASY with better than 95% accuracy, but about 6% have to be called AHY. A key is provided for ageing and sexing adult Tree Swallows in the breeding season and criteria for ageing and sexing Tree Swallows during and after the postjuvenal and postnuptial molts are discussed.
ACKNOWLEDGMENTS
This paper is a contribution of the Long Point Bird Observatory.
Thanks to D. DeSteven, G. L. Holroyd, T. E. Quinney, and numerous
assistants who took part in the fieldwork, and to Co. A. Curtis who com-
piled much of the data from their records. The Long Point Region
Conservation Authority, the Ontario Ministry of the Environment, and
the Canadian Ministry of Transport permitted use of the colony sites.
R. R. Cohen, K. C. Parkes, R. B. H. Smith, and two anonymous reviewers
commented on the manuscript. My research in 1971-1979 was sup-
ported by a Frank M. Chapman Fellowship at the American Museum
of Natural History, New York. Long Point Bird Observatory provided
other support.
LITERATURE CITED
CANADIAN WILLDIFE SERVICE AND U.S. FISH AND WILDLIFE SERVICE. 1977. North Amer-
ican bird banding techniques, Vol. II. Fisheries and Environment Canada, Ottawa.
CHAPMAN, m. B. 1955. Studies of a Tree Swallow colony (Third paper). Bird-Banding
26:45-70.
COHEN, R. R. 1980. Color versus age in female Tree Swallows. J. Colorado-Wyoming
Acad. Sci. 12:44-45.
DESTEVEN, D. 1978. The influence of age on the breeding biology of the Tree Swallow
Iridoprocne bicolor. Ibis 120:516-523.
DWIGHr, J., Jr. 1900. The sequence of plumages and moults of the passerine birds of
New York. Ann. N.Y. Acad. Sci. 13:73-360.
FREER, V., AND B. BELANGER. 1981. A technique for distinguishing age classes of adult
Bank Swallows. J. Field Ornithol. 52:341-343.
Kt;ERZl, R. G. 1941. Life-history studies of the Tree Swallow. Proc. Linn. Soc. New York
52-53:1-52.
Low, S. H. 1933. Further notes on the nesting of the Tree Swallows. Bird-Banding 4:
76-87.
Long Point Bird Observatory, P.O. Box 160, Port Rowan, Ontario NOE IMO
(Present address: Wildlife Research Section, Otario Mitistry of Natural
Resources, P.O. Box 50, Vlaple, Ottario LOJ IEO). Received 31 Aug. 1982;
accepted 7 Feb. 198:3.