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The Nesting of The Long Tailed Tit
The Nesting of The Long Tailed Tit
The Nesting of The Long Tailed Tit
To cite this article: David Lack & Elizabeth Lack (1958) The Nesting of the Long-Tailed Tit, Bird
Study, 5:1, 1-19, DOI: 10.1080/00063655809475897
SUMMARY
1. Winter flocks of Long-tailed Tits roosted communally.
2. In each of five years, 5-6 pairs nested on 8o acres, the nests being well
spaced, but the species is not territorial.
3. In woodland, many nests were in thorns, usually Bramble or Hawthorn,
usually 3-5 feet up, many others were in a bare fork, usually of Oak or Ash,
usually 30-70 feet up, and some were in thornless twigs, such as Clematis or
Spruce, usually at an intermediate height.
4. The birds often changed their type of nesting site for repeat nests, which
were usually 50-80 yards away, occasionally 550-700 yards away from the first
nest.
5. Other British observers have found nearly all nests in low thorns, while
in Sweden most nests are in Birch or Juniper, and hardly any in thorns.
6. In first attempts the first egg usually follows three weeks after the start
of building, but the interval is shorter in repeats, in late repeats being,8-10 days.
7. The first eggs were usually laid at the end of March, but up to three
weeks later in a cold spring. Repeat nests follow until the end of April, but
rarely later.
8. Clutches were unusually small in the late season of 1955.
g. The frequency with which three or four parents feed one brood is
attributed to the high rate of nest destruction, the restricted breeding season
and the absence of territorial behaviour.
to. Young were raised from only 16% of the Wytham nests. All in bare forks
and thornless twigs were destroyed, nearly all before laying. Of those in low
thorns, 12% were successful inside and 47% outside the woodland, and losses
were highest during incubation.
INTRODUCTION
These observations on the Long-tailed Tit (Aegithalos caudatus)
were made by E.L. in 194.9-5I and D.L. in 1955-57, in mixed
broad-leaved woodland on the Wytham Estate near Oxford,
especially in Marley Wood. The latter is but moderately suitable,
since most of the canopy trees of Oak and Ash (a favourite feeding
place) were felled in 1914-18, and much of the wood now consists
of overgrown Hazel coppice:, there are also some small marshes
and extensive Elder scrub, where Long-tailed Tits rarely occur.
Many further nests were found in the Great Wood and two planta-
tions, in which canopy trees are at normal density. In 195o and
1951, most of the individuals in Marley Wood were trapped during
the winter and colour-ringed.
Personal observations were supplemented by nesting data from
the diaries of A. Whitaker, F. C. R. Jourdain and J. H. Owen
(partly published in Owen 1945), deposited at the Edward Grey
2 D. AND E. LACK
PRE-NESTING BEHAVIOUR
In at least two winters, two flocks of Long-tailed Tits, each ten
or a dozen strong, frequented Marley Wood. Each flock kept to its
own part of the wood and they were not seen to meet. When an
individual got separated from its flock, it became extremely agi-
tated, calling loudly and flying rapidly about until it again made
contact.
Tits of the genus Parus often feed for a time with Long-tailed
Tits, as is well known in winter, but the attraction continues in
March and April when, should a pair of Long-tailed Tits stop build-
ing and move into an Oak tree to feed, they are usually joined after
a very short time by Parus tits, and sometimes by a Nuthatch (Sitta
europaea). Our impression is that the Long-tailed Tits attract the
Parus species rather than the reverse. The different species tend
to feed on different parts of the trees, and the survival value of the
association is probably that the chance of a predator approaching
undetected is smaller for a party than a solitary individual. If the
trilling hawk-alarm call is given by one of the group, tits of all
species respond by immediately diving from the upper parts of the
trees to cover below.
In different winters we have found five communal roosting
places of the Long-tailed Tit in Marley Wood, each involving 9-12
individuals, which in some cases certainly, and in the rest presum-
ably, were the same as the flock which fed together during the
day. One roost, used regularly for at least two months, was nine
feet above the ground on the bare branch of a Hawthorn. Another,
used for six weeks until the birds deserted it through human dis-
turbance, was six feet up in the middle of a dense thicket of Black-
thorn ; it was in use again two years later, though not in the inter-
vening winter so far as we know. Another roost, fifteen feet up on
a bare Hawthorn branch, was not used after the evening on which
it was found, though the birds were not disturbed there by the
observer. Two others, found by D. F. Owen following hard after a
flock at dusk, were used only once each, probably because of the
unavoidable disturbance that his actions entailed. One of them was
five feet up in a Blackthorn thicket and the other nearly thirty feet
up on a bare Hawthorn branch, the latter seemingly being a last-
minute choice of a flock previously disturbed from their normal
place by the chasing. When roosting on a branch, the birds usually
huddled together in a line, but in a thicket they sometimes appeared
to form into a ball, with tails sticking out at diverse angles. Such
roosts were found between November and March. One used regu-
larly in March was deserted by each pair successively when their
nest was roofed, after which they roosted in their own nest. Else-
where, E.L. once found a communal roost in June, presumably a
family party.
1 95 8 NESTING OF LONG-TAILED TIT 3
At least some colour-ringed individuals nested within the area
over which they fed in their winter flock. In March, each pair
separates from the flock in order to build, and each also feeds
together, but later in the day they may rejoin the flock to feed and
for roosting.
NUMBERS
Wytham Great Wood, with many canopy trees and good under-
growth, is fairly typical of English woodland, and here in 1 957
9 pairs of Long-tailed Tits nested on about 65 acres of what is per-
haps an optimum habitat. Marley Wood, with few canopy trees
and some marshes and scrub, in all about 8o acres (which includes
some bordering ground beyond the wood proper), held 6 pairs in
1950, 1951, 1955 and 1957, and 5 pairs in 1956. None of these years
was preceded by a hard winter.
The nearly constant density each year in Marley Wood, coupled
with the fact that each pair nested away from its neighbours, might
suggest that the Long-tailed Tit is territorial, but it is not. The
male does not sing, nor does either of the pair otherwise proclaim
itself, or patrol the ground to drive out other Long-tailed Tits.
Further, when collecting feathers or feeding, the pair wanders
freely over ground which, had the species been territorial, should
have belonged to other pairs, and the birds do this without any
change from conspicuous to inconspicuous behaviour, such as
characterises, for instance, a trespassing Robin (Erithacus rube-
cula). Both near their nest and on these longer excursions, the pair
stays near together, keeping in touch by calls and getting highly
agitated should they lose each other; similar behaviour is found in
other species which form pairs in flocks, but not in territorial
species, which, having a common meeting ground in the territory,
do not seem to need contact with - their mates in this way.
Moreover the occasional fights between neighbours do not occur
at fixed places, such as might suggest territorial boundaries. They
happen when one individual comes extremely close to a pair, where-
upon one of the pair drives it off, after which it quickly returns to
its mate. The impression given is that the bird is defending its
mate and certainly not an area. In one instance two building pairs
met when each was some 6o yards from their nest, and then all
four fought, each of one pair tackling a member of the other. We
have also seen strange Long-tailed Tits driven away from the nest,
but such chases, like the others, have been discontinued after a
short distance. We did not identify the sexes in these encounters.
As discussed later, while many repeat nests are built near the
destroyed one, three were 550-700 yards away, and two of these in-
volved the pair crossing over ground used mainly by other pairs,
which again suggests the absence of territorial behaviour. So does
the unusual frequency with which one and sometimes two extra
adults join with a breeding pair, as mentioned later. Indeed, every-
thing suggests that the Long-tailed Tit is not territorial. Because
the pairs make rather long excursions for food and feathers, they
may well be aware of the position of neighbouring pairs ; and the
D. AND E. LACK
4 5(I )
TABLE II
NESTING SITES OF LONG-TAILED TIT
B. THORNLESS TWIGS
Broom (Cytisus) 3
Honeysuckle (Lonicera) 3
Clematis (Clematis) 4
Privet (Ligustrum) 1 1 3
Ivy (Hedera) .. 1 1
Box (Buxus) .. .. 1 3
Laurel (Laurus) .. 1
Elder (Sambucus) 1
Ash (Fraxinus) .. 1
Scots Pine (Pinus) 1
Spruce (Picea) 4 1 2 1 2 4 5
Cypress (Cupressus) 1
Yew (Taxus) . . 4 1
Larch (Larix) . 1
Juniper (Juniperus) 29
Dead brushwood 1
C. BARE FORK
Oak (Quercus) .. .. 7 5 3 1 2 18
Ash (Fraxinus) 6 1 1 1
Elm (Ubnus) . . .. 1
Alder (Alnus) .. .. 1 2 29
Willow (Salix) .. 1 1 5
Birch (Betula) . . 2 112
Beech (Fagus) . .. 4
Aspen (Populus) .. 4
Rowan (Sorbus) 1
Elder (Sambucus) .. 1 2
Larch (Larix) . .. 1
Fruit Tree (Pyrus) 2
NOTE: The Swedish records were given only for the species of tree, not as to whether
the nest was in a fork or among twigs. Two further Swedish records were in nesting
boxes.
Percentage found in
Total found A
Sweden 212 1 99
kind of plant). Birch was much the commonest tree and Juniper
(Juniperus communis) the commonest shrub. Some nests were
alongside nests of the Wood Ant (Formica rufa) and others amid
colonies of the Fieldfare (Turdus pilaris), which presumably give
protection from predators (Durango 1 949).
The marked differences in nesting between Swedish A.c. caud-
atus and English A.c. rosaceus may well be due wholly to differ-
ences in the opportunities provided by Swedish and English habi-
tats respectively, and there is no need on present evidence to
attribute them to hereditary differences. In support of this view,
Scottish A.c. rosaceus often use forks in Birch like A.c. caudatus
in Sweden, and English A.c. rosaceus nest in the introduced
Spruce, although it provides a very different nesting site from
those in natural English woodland.
Summarising, the Long-tailed Tit has more varied nesting sites
than any other British passerine species, and it would be hard to
match it anywhere in the world. The nest may be from 2 to 70 feet
above the ground, and deep in a thorny bush, or among thornless
twigs, usually evergreen but sometimes bare, or in a bare open
fork, large or small. Finally, protection may be given by inaccessi-
bility on a terminal branch, or by a lichen covering which matches
the lichens on a trunk, or by a screen of creepers or dense conifer
needles, or by numerous thorns, or by proximity to the nest of a
fiercer bird or insect. As an abnormality the species may even breed
in a nesting box (per S. Durango) or in a hole in the ground (Pfeifer
1938, Thielemann 1956).
NOTES:
(i) All these were in Marley Wood except for the last three, which were in Great Wood
or adjoining plantations.
(ii) A. Whitaker (unpublished diary) recorded a pair which first nested 5 ft. up in a
Wild Rose, then in the fork of an Oak at 25 ft.
1958 NESTING OF LONG-TAILED TIT 9
29 March, 9 weeks after building started. In the Great Tit, like the
Long-tailed "Tit, building takes a much smaller number of days in
late than first nests (J. A. Gibb, in Litt.).
UNUSUAL CONSTRUCTIONS
In one second repeat in 195o, the start of which we missed, the
birds hurried so much that the nest was still undomed on I I May,
although by then it held its fifth egg; it was robbed soon after-
wards. In another nest, of the normal pattern when built and until
the young were several days old, the roof later collapsed and the
young thereafter sat in an open cup ; they were reared successfully.
Another nest had an extra hole in the top, through which the brood-
ing bird's tail projected (cf. Robertson & Porter 1952). Another
became tipped completely sideways, so that the long axis was hori-
zontal, but the young were successfully raised. A few of those in
high forks were blown loose during gales. Owen (1945) recorded
other cases of structural damage to nests.
Record Cards for the Blackbird (Turdus merula) and Song Thrush
(T. ericetorum) in May as compared with April (Myres 1955), and
this does not reflect what happens in nature. But the drop in these
thrushes is much smaller than in the Long-tailed Tit. Further, there
is no drop in the number of Long-tailed Tit records in late as com-
pared with early April. We conclude that in southern England only
a few Long-tailed Tits re-nest after April, though the proportion
doing so may be rather higher than indicated by the Nest Record
Cards. It may be added that the speed with which repeat nests are
built in April suggests that breeding as late as May is disadvanta-
geous.
A. Whitaker's records for the Derby—Yorks. border, and the
rather few B.T.O. Nest Record Cards for England north of the
Wash, show that here, as was to be expected, Long-tailed Tits
tend to start and continue breeding rather later than further south,
and nests up to the middle of May are not uncommon, with a very
few towards the end of the month.
Jourdain (1938), apparently based on Brown (1924), stated that
there are occasional very late nests. As already mentioned, one nest
in Marley Wood was nearing completion on 3 June, while A.
Whitaker recorded one with 5 fresh eggs in Shropshire on 16 June
(but, as he did not mention whether he saw the parent birds, this
was conceivably a deserted nest begun earlier). We have no further
records of June nests (one B.T.O. record was clearly erroneous).
CLUTCH
Feeling in with one finger, we found it impossible to be sure of
the number of eggs present when it exceeded 7, which it did in
most years. But of 5 clutches in the late season of 1955, four con-
sisted of seven eggs and one of six. The Great Tit likewise laid
smaller clutches in a late year (Lack 1 955).
F. C. R. Jourdain's diaries record only 7 clutches of the Long-
tailed Tits in southern England (1/9, 2/1o, 4/11). A. Whitaker's
records, set out in Table VI, show that in Derby—Yorks. the usual
clutch is 9-11, with an average near 10. But the average for the
B.T.O. Nest Records, also set out in Table VI, is probably too
low, since some observers felt in with one finger, which is likely to
result in an underestimate. Brown (1924) considered the normal
clutch in Cumberland to be 8-12, occasionally 5 or 6, but gave no
dated records. The figures in Table VI for Sweden show that the
average here is rather higher than in northern England.
In England, it is generally believed that clutches of more than
1 2 eggs are due to two hens. The dates of laying in a B.T.O. nest
with 14 eggs showed that on at least two consecutive days, 2 eggs
were laid each day, which almost certainly means that two hens were
involved. A more definite instance of two hens laying in the same
nest was observed by A. Whitaker. Laying started on zo April ;
on 3o April 4 adults were put out of the nest at dusk, which then
contained 6 eggs, which were taken ; on 13 May it contained 9
more eggs, 4 of which were larger than the others, so presumably
two hens had laid there. In some of the instances discussed later of
more than two adults at a nest, the clutch or brood exceeded 12.
1958 NESTING OF LONG-TAILED TIT 13
TABLE VI
CLUTCH SIZE
Number of records
NOTE: All these nests were found during building and followed through to their fate.
NESTING LOSSES
Our figures for nesting losses on the Wytham Estate are set out
in Table VII, for all those nests which we found during building
and followed through to failure or the successful raising of ypung.
We took care not to disturb the nests and think it most unlikely
that our visits increased their liability to predation, while no losses
were due to small boys or other human causes.
In all, 56 out of 67 nests were destroyed before the young left,
a loss of 84%, which was higher than that recorded for any other
species of bird (cf. Lack 1954). However, D. W. Snow (1958)
has found that 86% of the Blackbird nests in Wytham Woods were
destroyed by predation. In the Long-tailed Tit, there were no clear
differences between different years as such, or between first and re-
peat nests as such, but marked differences between different nest-
ing sites. None of the 15 nests in bare forks, and none of the 9 in
thornless twigs, were successful, whereas of those in low thorns
II out of 43, or 26%, were successful. Actually, of the nests in
thorns in the woodland proper, only 3 out of 26, or 12% were suc-
cessful, as compared with 8 out of 17, or 47%, in thorns at the
wood-edge (the latter including 8 in isolated Hawthorns and
Brambles in the Park, 5 in an area of Hawthorn and Bramble scrub
with hardly any trees, 3 in irregularly clipped hedges between wood
and field, and one in an isolated clump of Gorse). If these latter
nests are excluded, only 3 out of the 5o nests in the woodland
proper, or 6%, were successful, but their exclusion is not justi-
fiable since the birds in question fed in the woods, though nesting
at its edge.
Table VIII shows the stage at which each nest was destroyed.
Omitting the few nests visited too infrequently for precise dating,
all I I nests in bare forks were destroyed before laying (if we assume
16 D. AND E. LACK 5(1)
TABLE VIII
STAGE AT WHICH NEST WAS DESTROYED
SUCCESSFUL 0 0 3 8
NOTE: The percentage destroyed of those at risk is calculated with reference to the total
available, including any successful nests but omitting those in which the stage at which
destruction occurred was not recorded. Thus for nests in thorns, 42 were available
initially, of which 8, or 19%, were destroyed before laying; this leaves 34 of which 3,
or 9%, were destroyed during laying; this in turn leaves 31, of which 16, or 52%, were
destroyed during incubation, and so on.
REFERENCES
BROWN, R. H. 1924. The nest-building and other breeding habits of
the Long-tailed Tit. Brit. Birds, 17:206-7.
DURANGO, s. 1949. The nesting associations of birds with social in-
sects and with birds of different species. Ibis, 91 : 1 4o-3•
JOURDAIN, F. C. R. 1938. In The Handbook of British Birds, ed.
H. F. Witherby et al., 1 :272. London.
LACK, D. 1 954. The Natural Regulation of Animal Numbers. Ox-
ford. Pp. 74-81.
LACK, D. 1955. British Tits (Parus spp.) in nesting boxes. Ardea,
43:5 0-8 4.
MANSE, V. 1942. Four Long-tailed Tits feeding young in nest. Brit.
Birds, 3 6 : 55
MvRES, M. T. 1955. The breeding of Blackbird, Song Thrush and
Mistle Thrush in Great Britain. Pt. 1. Bird Study, 2:2-24.
OWEN, J. H. 1945. The nesting of the Long-tailed Tit. Brit. Birds,
38 : 271-3.
1958 NESTING OF LONG-TAILED TIT 19
SUMMARY
The Collared Turtle Dove usually nests on horizontal branches of trees at
various heights, though occasional nests are situated on buildings. It is excep-
tional for it to take over the old nest of another species, e.g. Woodpigeon.
The birds take several days to build their nests, but they work only in the
morning. The intensity of their work is greater early in the morning than later.
Nest building occurs from March to May.
INTRODUCTION
The Collared Turtle Dove (Streptopelia decaocto) is a bird about
which many authors have recently written in European ornitho-
logical journals. This is because the boundary line of its geographi-
cal distribution is moving northwards very quickly (Miczynski,
195o and 1951 ; Fisher, 1953 ; Stresemann, 195o). The Collared
Turtle Dove is a common bird throughout Poland, except the north-
east districts (Dyrcz, 1957). Since 195o I have observed many
Collared Turtle Doves in Krakow where they live both in the town
and in its outskirts, wherever they find trees, which are their
typical nest habitat.