Nordic Society Oikos

Territory Quality and Feather Growth in the White-Backed Woodpecker Dendrocopos leucotos
Author(s): Allan Carlson
Source: Journal of Avian Biology, Vol. 29, No. 2 (Jun., 1998), pp. 205-207
Published by: Wiley on behalf of Nordic Society Oikos
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JOURNALOF AVIANBIOLOGY29: 205-207. Copenhagen1998

Territoryquality and feather growth in the White-backed

WoodpeckerDendrocoposleucotos
Allan Carlson, Department of Conservation Biology, Swedish University of Agricultural Sciences, Box 7002,
S-750 07 Uppsala, Sweden. E-mail: Allan.Carlson@nvb.slu.se
During the last decades the White-backed Woodpecker has
shown a precipitous decline in many regions of the western
Palearctic. In this study I ask whether detoriation of the forest
breeding habitat might have contributed to this population decline and contraction of the species' range. By using the technique of ptilochronology it is shown that the bird's condition
reflects the quality of the breeding territory as estimated by the
density of dead and deciduous stems. Feather growth bars were
wider on old museum specimens, suggesting that birds living
55-150 years ago experienced a forest landscape of better
quality than birds do today.

Once widespread in the boreal forests of Fennoscandia, the White-backed Woodpecker Dendrocopos leucotos has declined dramatically during the second
half of this century (Tiainen 1990, Carlson and
Aulen 1992). Several reasons for the decline have
been suggested, but attention has mainly focused on
habitat loss (Aulen 1988, Haland and Ugelvik 1990,
Virkkala et al. 1993). The White-backed Woodpecker
has a highly specialized diet, consisting mainly of
wood-boring and bark-living insects, collected in
dead and decaying trees (Aulen 1988, 1991). Despite
a great interest in the species on the population level
(Virkkala et al. 1993), little is known about possible
effects of habitat deterioration and fragmentation on
the condition of individual birds. Here, I use the
technique of ptilochronology (Grubb 1989) to explore whether estimates of territory quality are correlated with the nutritional condition of the birds
living in these territories. Furthermore, this data set
is contrasted to feather growth measured on museum
skins from the period prior to the introduction of
modern forestry methods.

The technique of ptilochronology uses the width of

feather growth bars to assess the nutritional status of
a bird (Grubb 1989). Experimental evidence indicates
that growth bar width reflects the bird's nutritional
condition at the time of moult and feather growth
(Grubb 1995). Assumptions and utility of the method
are discussed in detail in Murphy and King (1991),
Murphy (1992) and Grubb (1992). During field work
in spring (1990-1992), the right fourth rectrix was
sampled from nine birds (3 females and 6 males) captured at their breeding sites (nine different territories).
This was done in two of the three Swedish subpopulations. The method adopted was that outlined in
Grubb (1989). Because rectrices were worn and dirty
it was often possible to discern only five growth bars.
These five growth bars were measured, and their
means were used in the analysis.
Adult White-backed Woodpeckers show a high degree of territory fidelity (I. Stenberg and A. Carlson
unpubl.). Therefore, I assume that sampled feathers
had been grown at the breeding site during the previous moult. White-backed Woodpeckers moult their
tail in summer (Cramp 1988).
The Museum of Natural History (Stockholm) has a
fairly large collection of White-backed Woodpecker
skins covering several of the species' races. Eight
skins were of the nominate form leucotos leucotos and
these were used in this study. Five of the skins were
from Swedish birds, and one skin each from Norway,
Lithuania and NW Russia. These skins were collected
between 1832 and 1942, thus being from 55 to 150
years old.

JOURNAL OF AVIAN BIOLOGY 29:2 (1998)

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205

3.0

E 2.52.5

-T

21.0

Ta

1.5
.

cS0
) 0.5

r-

0.0

58

117

175

233

292

350

Density (stems /ha)

Fig. 1. Relationshipbetweenestimatesof territoryqualityand
feather growth. Regression analysis; growth= 1.48+ 3.8 x
10-3 deciduousstems/ha (brokenline); F,7= 9.4, r2 =0.57,
p < 0.02; growth= 1.48+ 7.56 x 10-3 dead stems/ha (solid
line); F1,7= 9.01, r2= 0.56, p < 0.05.
Two components of territory quality, density of dead
stems and density of deciduous trees were quantified as
follows. At intervals of 100 m, areas of 20 x 50 m were
censused for growing and dead stems (standing or
fallen) with a breast height diameter (bhd) > 10 cm.
This was done at 10 points along a north-south transect
and at 10 points in an east-west transect centered on the
previous year's cavity tree. Thus, for each territory 20
plots of 0.1 ha were censused.
Due to the limited sample size it was not possible to
control for the effect of sex.
Feather growth bar width was measured on eight
55-150 years old White-backed Woodpecker and were
on average broader (3.71 + 0.41 mm) than on birds
living today (2.20 + 0.36 mm; Mann-Whitney U-test,
U = 9, p < 0.05).
A linear regression analysis revealed a positive relationship between the densities of dead and live deciduous stems found in the White-backed Woodpecker
territories and feather growth (density of dead trees
r2 = 0.56, density of deciduous trees r2 = 0.58) (Fig. 1).
A multiple step-wise regression analysis indicated that
68% of the variation in feather growth bar width was
explained by these two variables (F2,6 = 6.2, r2 = 0.68,
p = 0.03).
The important result of this study is that the bird's
nutritional condition, measured as feather growth bar
width, was associated with two estimates of territory
quality, namely the density of dead stems and the
density of deciduous trees. This result corroborates the
findings of another study in which the density of hunting perches and habitat type influenced feather growth
in the Loggerhead Shrike Lanius ludovicianus(Josef and
Grubb 1992, Grubb and Josef 1994). Due to the Whitebacked Woodpecker's specialization on larvae of wood206

boring insects (Aul6n 1988), it was not surprising that

dead stems affected the birds' nutritional condition.
That also the density of deciduous trees can influence
the birds' nutritional condition could likewise be expected. Deciduous trees, especially old stems, can carry
many dead and decaying branches (Carlson unpubl.),
and these are frequently used by foraging White-backed
Woodpeckers (Aul6n 1988).
An alternative explanation for the results is that high
quality birds are found in high quality territories while
low quality ones reside in territories of poor quality.
Thus, the observed relationship between feather growth
and territory quality may simply depend on the quality
of the individual bird rather than habitat. However, I
have not assessed the quality of individual birds. On the
museum skins, wing lengths were measured as an index
of bird size. There was no indication that long-winged
birds grew feathers with wider bars (r = -0.43, p =
0.28).
Interestingly, the historical comparison suggests that
birds were living under better nutritional conditions in
the past. It is a well-known fact that in today's managed forest landscape the density of both deciduous
trees and dead stems has been severely reduced. In the
late 19th century, the volume of dead stems in Swedish
forests was approximately 20% while the corresponding
figure today is as low as 1% (Linder and Ostlund 1990).
The proportion of deciduous trees was probably
around 30% by the end of the last century (Olsson
1992). Within the early 20th century distribution range
of the species, today the proportion of deciduous trees
in the landscape is 2-4% (Olsson 1992).
To conclude, suitable habitat for the White-backed
Woodpecker, old-growth deciduous forest (Carlson and
Stenberg 1995), is an extremely scarce resource in today's rationally managed forest landscape. The results
presented here support the view that the observed decline of White-backed Woodpeckers in Sweden is
caused by deterioration of the forest landscape.
- I thank Dr. T. Part for commentsand
Acknowledgements
discussionof the manuscript.The help of Dr. G. Friskat the
Museumof NaturalHistorywas invaluable.This projectwas
supportedby grants from WWF-Swedenand the Swedish
ResearchCouncilfor Agricultureand Forestry.The paperwas
writtenwhile I was financedby the latterorganization.

References
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trees:a methodof improvingthe food situationfor endangeredwoodpeckers.- Holarct.Ecol. 14: 68-80.
Carlson,A. and Aul6n, G. 1992. Territorialdynamicsin an
isolated White-backedWoodpeckerDendrocoposleucotos
population.- Conserv.Biol. 6: 450-454.
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All use subject to JSTOR Terms and Conditions

- and Stenberg,I. 1995. VitryggighackspettDendrocopos

leucotos: Biotopval och sdrbarhetsanalys.- Swedish
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WildlifeEcology, Report No. 27. (In Swedish.)
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Linder, P. and Ostlund, L. 1992. F6rindringar i Norra
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Olsson, R. 1992. Levandeskog - skogsbruketoch den biologiskamdngfalden.- SNF, Stockholm.(In Swedish.)
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JOURNAL OF AVIAN BIOLOGY 29: 207-208. Copenhagen 1998

Genetic confirmationof non-identicalembryonictwins in the

House Sparrow Passer domesticus
Simon C. Griffithand Reuven Stewart, Department of Zoology, University of Leicester, Leicester, UK
LE1 7RH. E-mail: sg4Oje.ac.uk
During routine ornithological monitoring of a freeliving population of House Sparrows Passer domesticus on Lundy Island, England, an unhatched egg
from an otherwise successful clutch of four was
found to contain two intact embryos. The embryos
were comparable in size, and were at an equally advanced stage of development (estimated as being
within two days of hatching, assuming an incubation
period of 11 days (Seel 1968)).
PCR amplification of extracted DNA (Primmer et
al. 1995) at three microsatellite loci (Neumann and
Wetton 1996) identified the twin embryos as full siblings, but resulting from separate maternal gametes
(Fig. 1, Table 1). In addition, SSCP analysis of a
sex-specific PCR product (Griffiths 1995) distinguished the embryos as of different sexes.
Twinning in birds is rare (Berger 1953, Batt et al.

1975) and is typically attributed to cleavage during

the early stages of development resulting in identical,
monozygotic twins (Sturkie 1946). Clearly, the presence of dizygotic House Sparrow twins of different
sexes cannot be due to cleavage. Two scenarios

Table 1. Allotypes of parents and twin offspringat three

microsatelliteloci. Alleleshave been assignedarbitraryletters
for illustrativepurposes.

Male
Female
Twin 2

Locus 1

Locus 2

Locus 3

(Pdop3)

(Pdo L4)

(Pdo gS5)

ab
cd
ad

ef
gh
eh

ij
kj
jj

JOURNAL OF AVIAN BIOLOGY 29:2 (1998)

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207