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The structure and function of nests of Long

Tailed Tits Aegithalos caudatus
ARTICLE in FUNCTIONAL ECOLOGY JULY 2004
Impact Factor: 4.83 DOI: 10.1111/j.0269-8463.2004.00883.x

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Functional
Ecology 2004
18, 578583

The structure and function of nests of Long-Tailed Tits

Aegithalos caudatus
Blackwell Publishing, Ltd.

A. McGOWAN,* S. P. SHARP and B. J. HATCHWELL

Evolution and Behaviour Group, Department of Animal and Plant Sciences, University of Sheffield,
Sheffield S10 2TN, UK

Summary
1. The aim of this study was to investigate the structure and thermoregulatory function
of nests of the Long-Tailed Tit, Aegithalos caudatus.
2. The feather lining of Long-Tailed Tit nests represents a major portion (41%) of the
total nest mass.
3. The mass of feathers varied among nests and declined through the breeding season,
but there was no seasonal loss of nest insulation quality because of increasing ambient
temperatures.
4. In an experiment to investigate the seasonal decline in the feather mass of nests,
feathers were added to nests at an early stage of the lining phase of nest construction. Nest
structure and insulating properties were then examined following nest completion.
5. The total mass of feathers in treatment and control nests did not differ significantly
and there was no significant difference in their nest insulation quality.
6. Our results demonstrate that Long-Tailed Tits adjust their nest-building behaviour
according to the nests thermal environment. Moreover, nest structure appears to be
adjusted to prevailing environmental conditions rather than being a function of feather
availability or time constraints.
Key-words: Feathers, nest construction, nest insulation
Functional Ecology (2004) 18, 578 583

Introduction
Conditions during the incubation and nestling periods
of reproduction by birds may have significant effects
on offspring growth and development (Drent 1975;
Deeming, Rowlett & Simkiss 1987; Webb 1987), but the
regulation of conditions within acceptable limits may be
energetically costly for parents (Pearson 1994; Williams
1996). The structure of nests may mitigate this energetic
demand on parents (Collias & Collias 1984; Hansell
2000). Well-insulated nests slow the rate of egg cooling
when an incubating parent departs, thereby reducing
the energy required for reheating and maintaining eggs
at incubation temperatures (White & Kinney 1974; Drent
1975; Reid, Monaghan & Ruxton 2000a). These energy
savings may be reallocated to other phases of reproduction (Reid, Monaghan & Ruxton 2000b). Poorly
insulated nests may affect reproductive success by prolonging the incubation period and reducing nestling
growth rates (Winkler 1993; Lombardo et al. 1995).

2004 British
Ecological Society

Author to whom correspondence should be addressed.

E-mail: amcgowan@seaturtle.org
*Present address: School of Biological Sciences, University of
Exeter, Hatherley Laboratories, Prince of Wales Road, Exeter,
EX4 4PS, UK.

However, the construction of a thermally optimal nest

must be traded off against building costs, so variation in
nest location, structure and properties may result if parents
adjust nest construction to suit prevailing conditions
(Schaefer 1976, 1980; Kern 1984; Kern & van Riper 1984).
Several studies have investigated spatial variation in
the insulating properties of nests across a geographical
range, but changing climatic conditions through a
breeding season may also result in temporal variation
in nest structure and function (Reid et al. 2002).
In this paper, we investigate the structure and function
of nests of the Long-Tailed Tit, Aegithalos caudatus.
Long-Tailed Tits build elaborate domed nests, lined with
up to 2600 feathers (Lack & Lack 1958; Gaston 1973;
Hansell 1993, 2000). Pairs are single-brooded, but repeat
nests are common (Hatchwell et al. 1999). Repeat nests
are built faster than first nests (Gaston 1973), and
Riehm (1970) found that they contained fewer feathers
and were less compact. The aim of this study was to
investigate the causes and consequences of variation in
the insulating properties of Long-Tailed Tit nests.
First, we quantify seasonal changes in the structure of
nests and determine which variables influence their
insulating properties. We show that the feather lining
affects nest insulation quality and that there is a seasonal decline in the mass of feather lining. Secondly, we
578

579
Nest structure
and function

describe an experiment to test three hypotheses for the

seasonal decline of feather mass: (1) feather mass declines
because breeders match insulating properties to
environmental conditions, so as ambient temperature
increases feather mass drops; (2) feather mass declines
because late in the season feathers are limiting; and
(3) feather mass declines due to time constraints; if collecting feathers is costly (Hansell 2000) it should be traded
off against the benefit of completing breeding early
in the season (MacColl & Hatchwell 2002; Hatchwell
et al. 2004). The experiment involved the addition of
feathers to nests. If feather mass is matched to environmental conditions we predicted that supply of extra
feathers would not increase the total feather mass of nests
relative to controls. However, if feathers are limiting,
feather mass would be higher in nests provided with
feathers compared with control nests. Finally, if feather
mass declines because of time constraints, the interval
between the start of lining and laying would be shorter
for nests given feathers compared with controls.

Materials and methods

A colour-ringed population of 3284 pairs of LongTailed Tits was studied from 2000 to 2002 in Melton
Wood (5331N, 113W), Doncaster, UK. Nests were
found during the building stage and visited every 1
2 days until they were depredated or until broods fledged.
The first nest built by a pair in a breeding season was
termed the first nest and subsequent nests by the same
pair in the same breeding season were classified as
second nests (few pairs built more than two complete
nests in a season). Both sexes build the nest and there are
two construction phases: building of the outer structure
and lining of the interior with feathers, termed feathering hereafter (Brown 1924). The outer structure comprises
various materials including moss, plant fibres, spiders
silk and lichen flakes (Hansell 1993, 2000; A. McGowan,
S. P. Sharp & B. J. Hatchwell, personal observation). The
outer structure of first nests took an average of 227
117 SE days (n = 30 nests found at the very start of
building) to complete, followed by 156 125 SE days
(n = 25 nests) of feathering by both birds. However,
second nests are built much quicker (112 188 days,
n = 11) than first nests (383 155 days, n = 26; Mann
Whitney U-test, z = 487, P < 0001).

2004 British
Ecological Society,
Functional Ecology,
18, 578583

In 2001, we measured the height of the entrance hole

from the ground (nest height) and the direction that
the nest hole faced (nest bearing) of 41 nests. Nest
contents were checked by touch to determine the start
of laying and nests were assumed to be complete when
the first egg appeared. On the day the first egg was laid,
cooling trials were conducted to determine nest insulation quality (the egg was removed during trials).
Two ellipsoid blocks of hardened Fimo modelling clay
(Eberhart, Neumarkt, Germany) were submersed in

water at 90 C, and left until the water temperature had

fallen to 75 C. Both blocks were then removed, dried
and a thermistor (PB-3221-S10, Gemini Data Loggers
UK Ltd, Chichester, West Sussex, UK) connected to a
Tinytag Plus datalogger (TGP-0073, Gemini Dataloggers
UK Ltd), was positioned into the centre of each block.
One block was placed in the nest while the other was
placed just outside the nest at an equivalent height.
Both blocks were left to cool for 90 min, dataloggers
recording the temperature of each block every 20 s.
Trials showed that this protocol allowed both blocks to
reach ambient temperature during cooling. Cooling rates
were obtained for each block by fitting an exponential
equation to the resultant temperature traces:
Temperature of block = ambient temperature + [B
exp (C time)],
eqn 1
where B and C are fitted positive constants. B is the initial gradient between the block and the ambient temperature, and C describes the rate of cooling of a block.
Nest insulation quality was calculated as the value of
C of the nest block minus the value of C of its control
block; a large positive difference reflects a high value of
nest insulation quality. The cooling rates of the two
blocks did not differ significantly when cooled under
the same environmental conditions (paired t-test: t9 =
0199, P = 085), so any differences in cooling rates during
trials was attributable to the insulating effect of nests.
Under controlled environmental conditions this method
of calculating nest insulation quality had a repeatability
of 89% (F32,65 = 1757) (Lessells & Boag 1987).
After breeding, Long-Tailed Tit nests were collected
and measured using a set of callipers with the nest in
an upright position and the nest hole facing forward.
Height was measured from the lowest to the highest
point of nest material while width was measured at the
base of the entrance hole. Depth was measured from
the base of the entrance hole to the back of the nest.
Nest volume was calculated by assuming nests were
ellipsoid in shape:
Nest volume = 4/3 rh rw rd,

eqn 2

where rh = height radius, rw = width radius and rd =

depth radius. After being placed in conditions of 15 C
and 70% humidity for at least 72 h to allow moisture
content to stabilize, nests were dissected and separated
into structural material, feathers and dried faeces.
Finally, the mass of each component part was weighed
electronically.

In 2002, a feather addition experiment was conducted
to investigate the seasonal decline in feather mass. Body
or contour feathers from domestic poultry in the size
range 20 40 mm were used, the preferred size for LongTailed Tits (Riehm 1970). Twenty pairs of first nests

580
A. McGowan,
S. P. Sharp &
B. J. Hatchwell

were matched for the start of the feathering phase

and randomly assigned to a feather addition (treatment)
or non-addition (control) group. When more than two
nests were available to choose from they were paired
for their proximity to one another in order to minimize
any habitat effect (mean distance between nest pairs:
208 365 m, n = 20). All nests were first nests. Feather
addition started at least 48 h after feathering had started.
Treatment nests had 2 g of feathers added to the interior per day for 4 consecutive days, giving a total of 8 g
of extra feathers per experimental nest; this represents
approximately 50% of the expected total feather mass
of a nest built at that time of year (see Results). The
added feathers were smoothed down to fit nest contours.
Nests in the control group were treated in the same
way, except for feather addition, to control for repeated
nest disturbance. No pairs were observed removing
feathers from their nest after additions had taken place
and no nests were deserted. The first egg date, clutch
size, hatch date, number of nestlings at day 11, average
chick weight at day 11, and fledging success was recorded
for all nests.
Experimental nests were collected after breeding and
their volumes calculated. Those nests depredated by
mustelids or small rodents remained structurally intact
and so were also collected to maximize sample sizes;
nests depredated by corvids were torn apart and could
not be used for analysis. Nests were left in controlled
conditions of 15 C and 70% humidity for a minimum
of 72 h before cooling rates were determined, as described
above, in a 5 C constant temperature room. Finally,
nests were dissected and feather and structural masses
measured.

Analyses were conducted using SPSS v.101 and S-Plus
v.61. All tests were two-tailed and means 1 SE are
presented. Observational data were analysed using General
Linear Models (GLM). Non-significant effects were
removed from the GLM by stepwise deletion (Crawley
1993). Insulation quality was used as the response
variable and all variables were included in the starting
model as covariates, except nest order (first or second),
which was included as a categorical variable. The starting model also included higher-order interaction terms
considered biologically meaningful. Model residuals
were checked for normality and homoscedasticity at
each step of removal.

Results

2004 British
Ecological Society,
Functional Ecology,
18, 578583

Thirteen of the initial 41 nests were destroyed by corvids before collection, leaving 28 whole nests for analysis. The mean mass of nests was 286 14 g (n = 28,
range = 154 446 g), feathers representing 41% (range =
2152%) and the outer structure 59% (range = 4879%)

Fig. 1. The relationship between (a) the feather mass of a nest

and the date the first egg was laid (Spearmans rank correlation
rs28 = 060, P < 0001) and (b) the structural mass of a nest
and the date the first egg was laid (Spearmans rank correlation:
rs28 = 037, P = 006).

Table 1. Results of GLM analysis to determine the physical

and biological nest variables that affected nest insulation
quality. An asterisk represents an interaction between two
variables (adjusted R2 = 046). Initially non-significant
variables removed from the model were: first egg date, nest
height, nest bearing and structural mass

Source

Type III sum

of squares
d.f. F

Corrected model
Intercept
Feather mass
Nest volume
Nest order
Nest volume * feather mass
Error
Total
Corrected total

472E 04
877E 06
122E 04
540E 05
963E 05
113E 04
389E 04
175E 02
862E 04

4
1
1
1
1
1
21
26
25

637
047
658
291
519
610

P
0002
0499
0018
0103
0033
0022

of total nest mass. There was a significant negative

relationship between a nests feather mass and the date
the first egg was laid (Fig. 1a), but no significant relationship between structural mass and lay date, although
mass tended to decline through the season (Fig. 1b).
Insulation quality was determined for 26 nests owing
to technical difficulties at two nests. Feather mass varied
from 49 g to 183 g and had a significant positive effect
on nest insulation quality (Table 1; Fig. 2a). There was

 0001 C per 20 s, n = 7) had poorer insulation quality

than first nests (00270 0001 C per 20 s, n = 19;
Table 1). Note that all these effects were independent
of date, which was non-significant and therefore removed
from the final model (Table 1).

581
Nest structure
and function

Fig. 2. The relationship between the nest insulation quality and (a) the mass of feather
lining; and (b) the feather massnest volume interaction (Table 1). In (b), nests were
categorized as high or low volume according to whether nest volume exceeded or was
less than or equal to the mean value for nest volume (785 cm3). Data points show the
models fitted values for nest insulation quality and incorporate all the variables of the
final model.

also a significant interaction between feather mass and

nest volume (Table 1): small nests had better insulating
properties if they had a larger feather mass while there
was no effect of feather mass on the insulation quality
of large nests (Fig. 2b). In addition, second nests (00207

There was no significant effect of feather addition on the

time between the onset of feathering and first egg date
(Table 2a; 8/40 nests were destroyed by predators before
laying, so n = 32). A further 14 nests (7 treatment, 7 control) were destroyed by corvids during the incubation
and nestling periods, so nest structure and insulation
quality were analysed for the remaining 18 nests. There
was no significant difference in the outer structure mass
of control and experimental nests (Table 2a). This is
not surprising because the experimental treatment started
only after nest structures were complete. More importantly, the mass of feathers in treatment and control nests
did not differ significantly (Table 2a). Furthermore,
there was no significant difference in the nest insulation
quality of treatment or control nests (Table 2a).
Of the 18 collected nests, 14 (7 treatment, 7 control)
had contained nestlings, and 13 nests (6 treatment, 7
control) had produced fledglings. No significant differences were found between experimental and control
nests in clutch size or the number and mass of nestlings
(Table 2b). Experimental and control nests did not differ significantly in the duration of the incubation or
nestling periods either (Table 2b).

Discussion
Feather mass was the most important nest component
for insulation quality. The mass of feathers in nests
declined through the breeding season and first nests
had better insulation quality than second nests. In
addition, there was a significant interaction between
feather mass and nest volume, low-volume nests with

Table 2. The effect of experimental addition of feathers to Long-Tailed Tit nests on: (a) nest construction and nest insulation
quality; and (b) measures of parental effort and reproductive success. Experimental nests either received 8 g of feathers
(treatment, n = 20) or received similar levels of disturbance but no feathers (control, n = 20). Means SE are shown

(a) Nest construction

Feathering duration (days)
Structural mass (g)
Feather mass (g)
Nest insulation quality (C/20 s)

2004 British
Ecological Society,
Functional Ecology,
18, 578583

(b) Parental effort and reproductive success

Clutch size
Number of nestlings at day 11
Mean nestling weight per nest at day 11 (g)
Duration of incubation period (days)
Duration of nestling rearing period (days)

Wilcoxon Signed
Ranks test

Experimental condition

Paired t-test

Treatment

Control

147 093
163 191
196 199
0023 00004

153 079
140 108
180 193
0024 00004

16
9
9
9

050
178
024
049

062
011
082
064

100 038
88 040
75 020
159 014
165 020

90 044
85 076
73 020
166 048
162 048

7
6
6
7
6

092
037
073
088
071

036
072
046
038
048

582
A. McGowan,
S. P. Sharp &
B. J. Hatchwell

2004 British
Ecological Society,
Functional Ecology,
18, 578583

more feathers having been better insulated than lowvolume nests with fewer feathers. We had no evidence
that Long-Tailed Tits further enhanced nest thermal
properties by orientating the nest towards the sun,
as reported in some bird species (e.g. Ricklefs &
Hainsworth 1969; Inouye et al. 1981), including LongTailed Tits (Riehm 1970).
The mass of feathers used for lining had a positive
effect on nests insulating properties. This is not surprising because the insulation qualities of feathers are
well established (Collias & Collias 1984), but the effect
of nest volume on the relationship between feather mass
and nest insulation quality would also be expected.
Small nests have a higher surface area to volume ratio
than large nests and therefore lose heat at a higher rate.
Long-Tailed Tits that build small nests could counteract
this effect by lining the nest with a larger number of
feathers. Alternatively, birds could counter this effect
by initially building a larger nest. However, larger nests
did not have as many feathers and were no better insulators than small nests that had a large number of feathers
(Fig. 2b).
The seasonal decline in the mass of feathers used to
line nests, also reported in an earlier study (Riehm 1970),
contrasts with the absence of any significant change in
structural mass with date. These two results suggest
that Long-Tailed Tits selectively adjust the lining component of their nests. Despite the seasonal decline in
feather mass there was no associated reduction in nest
insulation quality (Table 1); this is probably attributable to increasing ambient temperatures counteracting
the effect of a decreased feather mass. This suggests
that Long-Tailed Tits build nests with a microclimate
that allows eggs to cool within acceptable limits that
balance water loss with the risk of chilling, the key threats
to embryo viability (Drent 1975; Deeming et al. 1987;
Webb 1987). When controlling for the effects of feathers
and nest volume, second nests were of poorer insulation quality than first nests. There was no significant
difference in structural mass between first and second
nests, but the composition or construction of the outer
structure might have some influence on nest insulating
properties. We did not measure this directly, but Riehm
(1970) found that later nests were less compact than
early nests and it is certainly the case that the outer
structure of second nests is built far more rapidly than
that of first nests. Similar results have been reported in
other species (Schaefer 1976, 1980; Kern 1984).
Results of the feather provisioning experiment supported the environment-matching hypothesis. Pairs of
Long-Tailed Tits supplied with extra feathers brought
approximately 50% fewer feathers themselves so that
the total mass of feathers and nest insulating properties did not differ between control and treatment nests.
This reduction in feathering effort had no significant
effect on the duration of the feathering period, suggesting that the seasonal decline in feather mass is due
to Long-Tailed Tits tailoring feather mass to environmental conditions. Furthermore, this result shows that

Long-Tailed Tits must be able to accurately gauge the

thermal environment within their nest and adjust their
nest-building behaviour accordingly. There was no
evidence that energetic savings during feathering were
reallocated to other phases of reproduction, but it should
be noted that samples of experimental and control nests
available for these comparisons were small because of
nest predators.
Results of the experiment also suggested that feathers
were not a limiting resource in our study site. This
is consistent with the only previous study that has
investigated the availability of feathers for nest-builders,
which found that supplied feathers were barely used by
a range of passerine species, including Long-Tailed
Tits (Hansell & Ruxton 2002). Finally, the experiment
indicated that the seasonal reduction in the feather lining of nests was not due to time constraints. However,
this hypothesis cannot be ruled out entirely because only
first nests were involved in the experiment. If LongTailed Tits time completion of first nests to the time
when they are physiologically capable of laying eggs,
treatment birds may not have been able to lay eggs any
earlier even though their nests were complete. Given
that productivity declines sharply through the season
(MacColl & Hatchwell 2002), time constraints on the
construction of late nests are likely and it would be
worth repeating this experiment with second nests.
In conclusion, the feather lining of Long-Tailed Tit
nests is important in determining a nests insulating
properties; breeders appear able to gauge the thermal
environment within a nest and adjust their nest-building
behaviour accordingly. This study focused on the thermal
environment of nests with regards to egg cooling rates
and it would be interesting to extend this approach to
consider the effects of nest construction on nest humidity
and the thermal environment for nestlings and roosting adults.

Acknowledgements
We thank Andrew MacColl, Ian Hartley, Tim Birkhead and Dave Hazard for their invaluable input and
Doncaster City Council for allowing us to watch birds
on their land. We thank Robert Ricklefs and an
anonymous referee whose comments helped to improve
the manuscript. We also thank the numerous undergraduates who helped in measuring and dissecting
nests. AMcG was supported by a NERC studentship.

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