Title White feathers in black birds

Authors van Grouw, Hein

Date Submitted 2021-02-09

 White feathers in
black birds
Hein van Grouw
Richard Allen

Blackbird Turdus merula with progressive greying

Abstract The most common plumage abnormalities in birds involve some form
of white feathering, ranging from birds with just a few white feathers to individuals
that are completely white. The causes of aberrant white feathers are diverse and,
in many cases, unknown. Some are heritable, based on simple, genetically
determined changes in the pigmentation process. More commonly, the causes are
less clear-cut and can include environmental conditions (particularly in relation to
food availability), and the physical condition and/or age of the bird. In this paper,
white feathering is explored in three common species: Carrion Crow Corvus
corone, Hooded Crow C. cornix and Blackbird Turdus merula. Results from the BTO
Abnormal Plumage Survey are summarised, and data from a museum-based study
of Blackbirds with plumage abnormalities are reported. In all three species, partly
white plumage is recorded regularly and is often referred to incorrectly as albinism
or leucism.
Introduction partial albino or leucistic are used for almost
Aberrant white feathers in birds have always all different forms of aberrant white feath-
intrigued people, resulting in many pub- ering, yet in only a tiny proportion of cases
lished records in the ornithological literature. are they used correctly (Mahabal et al. 2016).
In these publications, a variety of names are A ‘partial albino’ is simply impossible, since
used to identify and classify the pigment albinos cannot produce melanin pigment at
abnormalities. The terminology used is, all. The true albino aberration is encountered
however, often conflicting, confusing and far less often than is generally supposed. In
incorrect. Most commonly, the terms albino, fact, aberrant white feathers are hardly ever

250 © British Birds 111 • May 2018 • 250 – 263

 White feathers in black birds

caused by albinism; typically, they are either a Jackdaw Coloeus monedula and Carrion
form of leucism or, more commonly, ‘pro- Crow. More than 30% of respondents have
gressive greying’. Non-heritable causes, such been watching birds in their garden for over
as dietary imbalance, can also be responsible. 20 years, showing that many records are
This paper explores two of these abnormali- coming from areas that are routinely
ties – progressive greying and dietary imbal- observed, and over 90% of gardens were
ance – on the basis of their occurrence in classed as suburban or rural.
three common species: Carrion Crow Corvus For Carrion Crow, the survey received 198
corone, Hooded Crow C. cornix and Black- records, but ten of these lacked further
bird Turdus merula. description so the data from 188 records
As a source of data on the occurrence of were used. For Blackbird, the data from 1,516
different plumage abnormalities, the results records (1,588 in total minus 72 without
of the BTO’s Abnormal Plumage Survey are further description) were used. For most
reported. This survey began in December records, a more accurate definition for each
2011, in conjunction with the familiar aberration could be assigned, since many
Garden BirdWatch (GBW) survey, in which descriptions gave sufficient clues to make a
several thousand volunteers across the UK positive identification possible. Many records
record weekly lists of the birds in their were accompanied by a photograph, which
garden, throughout the year; www.bto. made identification easier and more accurate.
org/volunteer-sur veys/gbw). The
Abnormal Plumage Survey aimed to
find out more about the plumage
abnormalities being seen by GBW par-
ticipants.
For recording birds with aberrant
white feathers, participants could
select from three different types of
aberration: melanism, leucism and
albinism. Since many aberrations can
be difficult to distinguish from each
other without some experience, the
BTO used ‘Leucism’ as an umbrella
term to encompass a range of plumage
irregularities involving lighter, white,
or partially white plumage. Impor-
tantly, however, the ‘Description’
section of each record – a free text
field in which the participant is asked
to describe the bird’s appearance in
detail – turned out to be extremely
Hans-Martin Berg, NMW

helpful and it was by analysing these

descriptions that (most of) the aberra-
tions could be distinguished and cate-
gorised more accurately.
The records used for this
paper spanned a period of five years,
from December 2011 to November
2016. By November 2016, about 4,350
records, involving over 70 species, had 171. Juvenile Blackbird Turdus merula showing leucism
been received. Easily the most com- in its typical form: the face, primaries, belly and feet lack
pigment (specimen at Natural History Museum Vienna,
monly reported species was Blackbird, NMW 13424). Note that the white feathering is already
with over 1,550 records, followed by present in the bird’s first (juvenile) plumage, and this
House Sparrow Passer domesticus, pattern will not change during later life.

British Birds 111 • May 2018 • 250 – 263 251

 van Grouw

White feathers
Leucism, from the
Greek leukos (for
white), can be defined
as the lack of melanin
from all or parts of
the plumage and skin.
Leucism thus occurs
in different forms:
all-white and pied. In
cases of all-white
plumage, no pigment
cells at all are present
in the skin to provide
the growing feathers
with melanin
pigment. In cases of

Anne Riley
Partial leucism, the
pigment cells are
absent from only
parts of the skin, 172. Adult male Blackbird, Yorkshire, April 2011. Progressive greying is a
leaving only these progressive loss of melanin pigment with each successive moult. In the
early stages the affected feathers are often spread randomly over any part
areas without pig- of the plumage, but finally the bird can become entirely white (plate 173).
mentation. Owing to
the way the early pigment cells (melano - 2013), the white pattern caused by Partial
blasts) migrate from their embryonic origin leucism is normally patchy and bilaterally
into the rest of the body (see van Grouw symmetrical – typical examples include a few
white outer flight feathers on
both wings and/or some white
feathers in the face. In more
extreme cases, the entire face, all
the primaries, the belly and the
feet lack pigment (plate 171).
Fully white birds may cause con-
fusion with an albino, but eye
colour is the most obvious dif-
ference: in albino the eyes are
red, in leucism eye colour is
unaffected. The underlying
causes of the two aberrations are
completely different. An albino
has pigment cells but lacks the
necessary enzyme (tyrosinase)
to start melanin synthesis,
Tim Heathfield

whereas a leucistic bird lacks

pigment cells altogether and is
thus unable to provide its
feathers with melanin (see van
173. Blackbirds that become (almost) entirely white often also Grouw 2014). Leucism is con-
lose the melanin from their bare parts (bill and feet). One or genital (present from birth) and
two remaining pigmented feathers in these birds are evidence
that their white plumage is the result of progressive greying in heritable.
its final stages. Owing to the loss of melanin pigment in the A far more common cause of
skin, the bill colour in both sexes is yellow. white feathers is ‘progressive

252 British Birds 111 • May 2018 • 250 –263

 White feathers in black birds

greying’ (van Grouw 2012b,

2013), which is the progressive
loss of melanin pigment with
each successive moult. The loss
of pigment can be the result of
a gradual reduction of tyrosi-
nase activity in the pigment
cells, but also of the pigment
cells dying off. In the early
stages of progressive greying,
the affected feathers are
usually spread randomly all
over the bird (plate 172), but
finally the entire plumage can
become white (plate 173). Dif-
ferent forms of progressive
greying occur and, for many,
the causes are still unknown.
In some cases, the cause may
be simple Mendelian (straight-
forward) inheritance (van
Grouw & Hume 2016), but
these cases seem to be rare.
More commonly, the condi-
tion may be linked with
factors such as environmental

Austin Brady
conditions or age, while the
progressive loss of pigment
cells may also be due to heri-
table disorders such as vitiligo
(pigment disease). 174. Carrion Crow Corvus corone, after its partial post-juvenile
External, non-heritable moult, UK, March 2016. The tail feathers and larger wing feathers
factors such as illness or are still juvenile feathers, and these clearly show a lack of
pigment, as a result of dietary imbalance.
dietary imbalance can also
result in pigment loss (plate 174). In such that are habitual scavengers, including wild-
cases, the bird is unable to extract the neces- fowl, gulls, some raptors (such as kites
sary nutrients from its food, which affects Milvus) and passerines (such as sparrows).
melanin synthesis. The bird’s pigmentation Plumage aberrations seem markedly less
will return to normal as soon as the external common in many of these groups than in the
causes are removed. Progressive greying was crows. Perhaps one reason for this is that
initially thought to be related to diet (Rollin these other species are in reality more omniv-
1964), but whereas a dietary deficiency orous than the crows, which may require
causes wide white bars in the feathers and much more animal protein to grow healthy
also often poor feather structure, in progres- feathers. This distinction may operate even
sive greying the feathers are completely white within the corvid family. From personal
and their structure is unaffected. Dietary experience, I know that hand-reared juvenile
imbalance is generally rare in wild birds, but Carrion Crows do not thrive on a diet that
one exception seems to be Carrion and juvenile Jackdaws and Rooks C. frugilegus
Hooded Crows, since individuals with develop very well on. Carrion and Hooded
pigment loss resulting from dietary defi- Crows are perhaps far less omnivorous than
ciency are not uncommon. often assumed and thus more prone to devel-
Why are these crows particularly affected? opmental problems (including loss of
Many other groups of birds include species pigment) owing to diet imbalances.

British Birds 111 • May 2018 • 250 – 263 253

 van Grouw
Harry Taylor, NHM

175. The degree of white feathering depends on the stage of progressive greying and can be
divided into four categories: clockwise from top left, <25% of the total plumage, 25–50%, 50–75%
and 75–100% (specimens at the Natural History Museum, Tring, respectively NHMUK 1987.24.315,
NHMUK 1987.24.318, NHMUK 1996.42.2330, NHMUK 2014.73.139).

Blackbirds and progressive greying table leucism found in domesticated birds

with age like pigeons, budgerigars, canaries and zebra
Although we know about feather pigmenta- finches. Almost all of the aberrant museum
tion and how mutations can change a bird’s specimens had white feathers randomly scat-
colour, there are still aberrations for which tered throughout their plumage. Some had
the cause remains unclear. One of these is only a few white feathers, others had more,
progressive greying: the occurrence of a while others were almost completely white
seemingly random spattering of white (plate 175). None of them seemed to follow
feathers, giving the bird a pied or variegated the normally patchy and bilaterally symmet-
appearance. At least 1,371 of the 1,516 aber- rical pattern of leucism. Furthermore, I
rant Blackbirds (90%) recorded in the BTO found no juvenile pied birds, even though the
survey were birds with some form of pro- white pattern caused by partial leucism is
gressive greying. The sex ratio within this already apparent in juvenile plumage, and
subset is unknown as birds that are more does not change with age. However, the
white than coloured are often difficult to sex diversity in the degree of white plumage
reliably in the field. found in the specimens, from hardly any to
During many years of personal research to almost totally white, seemed to suggest that
identify and record the many aberrantly in this aberration the numbers of white
plumaged bird specimens in museums all feathers increases with age.
over Europe (and in America), I noticed that Given that aberrantly coloured birds were
most specimens with white feathers were at often targeted by collectors in the past, it seems
odds with any of the known forms of heri- likely that at least some young birds would

254 British Birds 111 • May 2018 • 250 –263

 White feathers in black birds

have become study skins before

reaching adulthood. So, the absence
of juvenile birds with white feathers
among museum samples supports
the theory that this loss of pigment
occurs in later life. However,
museum specimens obviously show
only one plumage, and do not
provide data on how the condition
alters through successive moults in
individual birds. The BTO survey
provides some useful evidence in

Bill Barnett
this regard, charting the changes in
resident individual Blackbirds. A
typical comment reported was: ‘the
resident Blackbird with white
feathers becomes whiter every year’,
proving that a progressive loss of
pigment over time is more than just
conjecture (plates 176 & 177).
Further evidence was already avail-
able from ringers. In the late 1950s,
Band (1956) reported that a
‘normal’ adult male Blackbird
trapped in Lancashire on 24th
November 1950 was retrapped on
4th December 1955 when ‘there

Bill Barnett
were considerable patches of white
over most of its plumage’. In
response to Band’s observation,
several other ringers also reported 176 & 177. Male Blackbird photographed in September
cases of ‘albinism related to age’ in 2012 (176) and September 2013 (177) showing a significant
the Blackbird (Foott 1956; Spencer increase of white feathers during the intervening year due to
1956; Wigzell 1956). Spencer (1956) progressive greying. Note, however, that the bare parts (the
had noticed that the birds became legs and feet) are not (yet?) affected.
whiter over successive years and
called it ‘progressive albinism’. Rankin (1954) Progressive greying was the most common
had also reported an increase of white feathers colour aberration found in the sample (147
over time. specimens, or 66.5%; table 1). The other 74
specimens were assigned as follows: brown
Museum data on Blackbirds (8.6%), ino (6.8%), dilution (6.3%), grizzle
The information regarding museum speci- (3.6%), melanism (3.2%), albinism (2.7%)
mens presented below is based on my exami- and leucism (2.3%). Given that most forms
nation of 221 aberrant Blackbird specimens of progressive greying are probably not heri-
in several museum collections (see Acknowl- table, Brown was therefore the most common
edgments). Although aberrant birds were heritable aberration found in this sample of
often targeted by collectors, and the number Blackbirds.
of such specimens compared with normally Since the degree of white feathering
plumaged birds in collections does not repre- depends on the stage of progressive greying,
sent the ratio in the wild, we can assume that the 147 specimens were categorised into four
the ratio between the different aberrations is groups: up to 25% of the total plumage, up
quite representative; any unusual bird tended to 50%, up to 75% and up to 100% (plate
to be collected, regardless of its colour. 175). The majority, 89 specimens (61%) were

British Birds 111 • May 2018 • 250 – 263 255

van Grouw

Table 1. Colour aberrations found in the Blackbird Turdus merula, based on a sample of 221
museum specimens.

MUTATION (definitions based on van Grouw 2013) EFFECT ON COLOUR NO. %

ALBINISM Complete lack of melanin in feathers, All-white plumage, red eyes, 6 2.7
eyes and skin due to the heritable absence of the yellowish bill and pink feet.
enzyme tyrosinase in the pigment cells.

LEUCISM 5 2.3
Complete leucism Lack of melanin from all All-white plumage, yellow bill, pink 0 0
parts of the plumage and skin owing to the feet, normally coloured eyes.
heritable absence of pigment cells from all
of the skin areas.
Partial leucism Lack of both melanins from parts All-white feathers adjacent to normally 5 0.8
of the plumage and skin owing to the heritable coloured ones. White pattern
absence of pigment cells from some areas of skin. bilaterally symmetrical. Yellow bill
and pink feet or normally coloured
bill and feet; normally coloured eyes.

PROGRESSIVE GREYING Lack of melanin in All-white plumage or all-white 147 66.5

parts of the plumage owing to progressive loss feathers mixed randomly with
of pigment with age. Most forms seem to be normally coloured ones. Yellow bill
non-heritable. and pink feet or normal coloured bill
and feet; normally coloured eyes.
<25% white feathers. 89 40.3
25–50% white feathers. 26 11.8
50–75% white feathers. 12 5.4
75–100% white feathers. 20 9.0

BROWN Qualitative reduction of eumelanin owing Black becomes brown and brown 19 8.6
to incomplete synthesis (oxidation) of eumelanin. becomes light-brown; reddish-/
Phaeomelanin unaffected. Mutation is sex-linked yellowish-brown is unaffected.
and therefore only female birds in the sample.

DILUTION Quantitative reduction of one or both Black and brown (eumelanin) 14 6.3
melanins. Since male Blackbirds have eumelanin becomes silvery grey; reddish-/
only, certain forms of dilution, e.g. pastel and yellowish-brown (phaeomelanin)
isabel, cannot be separated without breeding may or may not be affected.
tests. However, at least three different forms are
present in the sample.

INO 15 6.8
Ino – light Strong qualitative reduction of both Black and brown becomes pale cream; 10 4.5
melanins due to incomplete synthesis (oxidation) reddish-/yellowish-brown becomes
of both melanins. Mutation is sex-linked and hardly visible. Yellow feet
therefore only female birds in the sample. and bill; eyes pinkish.
Ino – dark Qualitative reduction of both melanins Black and brown becomes pale brown; 5 2.3
due to incomplete synthesis (oxidation) of both reddish-/yellowish-brown becomes
melanins. Mutation is sex-linked and therefore hardly visible. Yellowish feet and bill;
only female birds in the sample. eyes dark pinkish.

MELANISM Abnormal deposit of melanin. Increase of black and/or reddish-brown. 7 3.2

GRIZZLE Lack of both melanins in part of Grizzled-white plumage. Pink feet, 8 3.6
the feather barbs in each feather. yellow bill and normally coloured eyes.

Total number aberrant specimens 221 100

256 British Birds 111 • May 2018 • 250 –263

 White feathers in black birds

in the first category, up to 25%; 26 birds feathers are also affected. The first records of
(17.7%) had between 25% and 50% white ‘white wing-barring’ in the Carrion Crow in
feathers; 12 (8.2 %) were more than half Britain are from the early 1950s (Sage 1954,
white and 20 (13.6%) had 75% or more of 1956a,b; Harrison 1957a,b). Both authors
their plumage white. assumed the white wing-barring to be genetic
The large percentage of birds in the early in origin and J. M. Harrison believed it was a
stage is easy to explain. A bird affected by symptom of an ‘ancestral’ plumage in the
progressive greying has to moult several Corvidae. However, C. J. O. Harrison (1963)
times before becoming entirely white, but considered the loss of pigment in Carrion
any aberrant Blackbird with a few odd white Crows to be diet-related, stating that the
feathers would have been desirable for collec- species ‘seems particularly prone to such
tors. Furthermore, it can be assumed that rel- defect, especially in urban or suburban areas
atively few birds become old enough to reach where the principal feeding places are often
the final stages. Annual mortality of adult rubbish tips, and the diet is likely to be
Blackbirds varies between 34% and 69%, abnormal or deficient.’ Sage (1964) disagreed,
depending on habitat and year (Simms but the evidence gathered subsequently sug-
1978), but it may be higher in urban than gests that C. J. O. Harrison was correct.
rural areas (Snow 1988). All specimens exam- Slagsvold et al. (1988) investigated the
ined were in adult plumage, and about three- causes of white wing-barring in Hooded
quarters were males (109 males, 35 females Crows in Norway. They concluded that the
and three unknown). For the fourth category, condition (which they misleadingly labelled
up to 100% white feathers, the sex ratio is less ‘partial albinism’) was primarily related to
clear as the three unidentified specimens food availability during the nestling stage
belonged in this category (15 males, 2 rather than to any genetic factors. Dietary
females and 3 unknown) but it can be imbalance was also found to be the cause of
assumed to be similar to other categories. white wing-barring in Carrion Crows in the
Netherlands, Germany and France (Terluin
Crows and loss of pigment as a 1996, 1998, 2009; Malher 2003; Bosch 2004).
result of diet
The occurrence of abnormal white feathering The effects of dietary imbalance
in the crows typically shows as whitish trans- The white wing-bars in crows are often com-
verse bars in the flight feathers (plate 178); in pared with fault-bars, but in fact fault-bars
are a subset of the wider category of white
more extreme cases, the wing-coverts and tail
wing-barring. Fault-bars are defined as inter-
ruptions in feather growth caused by a tem-
porary lack of the required nutrients. As a
result, the structure of the feather is incom-
pletely developed in the
corresponding areas,
appearing as an almost
transparent line or lines
right across the feather.
Fault-bars weaken the
Harry Taylor, NHM

feather considerably,
owing to the incomplete
development of the shaft
and barbs. Although
fault-bars can be accom-
panied by narrow lines
178. Carrion Crow wing showing the effects of moderate dietary where pigment is
imbalance in juvenile plumage (specimen at Natural History Museum,
Tring, NHMUK 2016.28.1). Since the juvenile feathers grow lacking, the feathers of
simultaneously, the loss of pigment will show in the same part of each the crows being dis-
feather and form the continuous white bars in the open wing. cussed here often lack

British Birds 111 • May 2018 • 250 – 263 257

 van Grouw

(Slagsvold et al. 1988;

Terluin 2009). The most
severely affected birds
rarely survive their first
year because of stunted
growth and poor overall
physical condition, while
the poor quality of their
feathers often reduces
Harry Taylor, NHM

their capability to fly.

Research has shown that
affected birds also have
smaller thyroid glands
and that those which die
early are nearly all
179. Carrion Crow wing, showing a more severe effect of dietary
imbalance in its juvenile feathers adjacent to newly moulted and females (Terluin 2009).
unaffected adult-type inner primaries (specimen at Natural History Crows that survive their
Museum, Tring, NHMUK 2016.28.4). first year usually grow
healthy and fully pig-
pigment over more than half of their length. mented feathers during their first complete
Close examination reveals the presence of moult (plate 179).
fault-bars, but also that the loss of pigment Since the feathers of a nestling all grow at
extends over a much larger part of the feather. the same time, the loss of pigment will show
The amino acids necessary for growth in in the same part of each feather to form the
general, and melanin development in partic- continuous white bars in the open wing (plate
ular, are tyrosine and lysine, and these need 178). Consequently, ‘white wing-barring’ as a
to be processed directly or indirectly from result of dietary deficiency is found mainly in
food. A lack of essential nutrients over an juvenile crows. Crows undergo only a partial
extended period during feather growth not post-juvenile moult in their first year of life,
only causes a loss of pigment, but the affected replacing their body plumage and some wing-
birds are also smaller than unaffected birds coverts only and retaining their juvenile flight
feathers (plate 174).
Most records of ‘adult’
crows with white wing-
barring are in fact birds
early in their second cal-
endar-year, before the
first moult of the flight
feathers in May or June.
Although pigment
loss resulting from
dietary imbalance
Olivier Poncin

appears to be largely
restricted to juveniles fed
on inappropriate food
by the parents during
180. Adult Carrion Crow, Brussels, Belgium, April 2010, showing the the nestling stage, the
result of dietary imbalance during part of the bird’s last wing moult. problem can also occur
The innermost primaries (P1–P3) were unaffected and show normal in older birds, even if
pigmentation. P4 shows the first signs of dietary deficiency and P5–P7
they were not previously
are strongly affected, being extensively white but also slightly shorter,
as are the two outermost secondaries (S1–S2), which grew at the same affected (plate 180).
time. The bird’s feeding habits must subsequently have improved, since Slagsvold et al. (1988)
P8–P10 and the remaining secondaries are only weakly affected. examined almost 3,500

258 British Birds 111 • May 2018 • 250 –263

 White feathers in black birds

birds and found that 5% of

juveniles showed pigment loss
as a result of dietary deficiency
compared with only 1% of
adults. More recent observa-
tions from a study of Carrion
Crows in Belgium showed a
higher proportion of affected
adults (van Grouw 2012a).
Around 7.2% of all crows
sampled (855 individuals) had
white feathers, which is a
remarkably high percentage
(Olivier Poncin pers. comm.).

Eddy Vaes
In juveniles (n=668), 5.5% had
white feathers, while no less
than 8.9% of the adults
(n=180) were affected. Birds 181. Adult Carrion Crow, Belgium, October 2015. Dietary
with white feathering were imbalance in adult plumage results in a more patchy appearance
also smaller (both weight and than in juvenile plumage, since each individual feather has
wing length were on average regrown at a different time.
around 3% less than for normal birds). first, which is the most common, progressive
In adult plumage, the pattern is generally greying begins around the head (plate 182).
different from the more regular ‘white wing- The second form affects mainly the wings and
barring’ in juveniles and is best referred to body first (plate 183), and in the third type all
more simply as a ‘lack of pigment’ (plate 181). of the plumage is affected equally (plate 184).
It seems very likely that the plumage abnor- In the first two types the colour change is pro-
malities of these Belgian crows stem from the gressive but the birds probably never obtain
birds’ diet, considering that all the birds were fully white plumage (plate 185). In the third
caught in urban areas or next to a rubbish type, however, the condition seems to
dump. However, further research is still neces- progress more rapidly and birds with this
sary to examine the precise way(s) in which type of progressive greying appear to be those
poor diet results in lack of pigment in crows. that do become completely white (plate 173).
Many of these fully white Blackbirds in the
BTO survey data BTO survey were recorded in autumn, winter
The BTO survey results suggest that pigment or early spring, which might suggest that they
loss due to dietary imbalance, as described are winter migrants. In the case of many resi-
above, is the most common colour aberra- dent birds, however, survey participants were
tion in Carrion Crows in the UK. Plumage able to observe the transformation of indi-
descriptions indicated that in 168 of the 188 vidual birds, four of which were recorded as
records the white feathering was due to becoming fully white.
dietary deficiency. The other 20 records could In the final stage of progressive greying,
be assigned to brown (6), ino (3), ‘all-white’ when almost the entire plumage is white,
(5), acromelanism (4) and unknown (2); see many Blackbirds also lose the melanin
van Grouw (2013) for more information pigment in the bare parts (bill and feet) as
about other colour mutations. well as the plumage, resulting in pink feet
and a yellow bill in both sexes (plate 173). In
Discussion Blackbirds, both sexes have a yellow bill, but
Based on both the museum specimens exam- in females the yellow pigment (carotenoid) is
ined and data from the BTO Survey, it masked by the melanin; in the absence of
appears that the distribution of white feathers melanin, the bill therefore appears yellow.
in Blackbirds showing progressive greying can Based on pigmentary defects in humans, the
be roughly divided into three types. In the disappearance of melanin from the skin

British Birds 111 • May 2018 • 250 – 263 259

 van Grouw

Rollin (1953a,b, 1959, 1964,

1981) considered that the white
feathering is non-hereditary and
solely diet-related. Sage (1962),
while recognising food defi-
ciency as a possible cause for
pigment loss, believed the white
feathering in Blackbirds to be
heritable and disagreed with
Rollin over its relationship to
diet: ‘I know of no evidence that
wild birds ever encounter condi-
tions which compel them to exist
on an unbalanced diet for any
John Oakley

length of time.’ Later research,

summarised above, indicates
that plumage abnormalities in
wild Carrion and Hooded Crows
182. Female Blackbird, UK, July 2010. It appears that
progressive greying in the Blackbird occurs in broadly three
are affected by diet. As far as
different forms. In the most common form, the progressive Blackbirds are concerned,
whitening starts at the head, and although the extent of white however, both Rollin and Sage
does increase with each successive moult, most birds probably may have been wrong. Rollin’s
never become fully white (see plates 176 & 177). experiments with captive birds
did show that artificial food
could cause loss of pigment, but
the affected feathers were not
solidly white and still showed
remains of pigment, as seen in
crows. No convincing evidence
that the development of white
feathers after a certain age has a
straightforward heritable origin
has been found so far. Rollin
(1964) associated the occurrence
David King

of abnormal white feathers in

wild birds with ‘the haunts of
man’, but certainly for Blackbirds
183. Male Blackbird, UK, November 2013. The rarest form of this is questionable. In the mid
progressive greying affects mainly the wings and body first; while 1800s, when the Blackbird was
the extent of white increases with each successive moult, these still predominantly a rural bird
birds also probably never become fully white (see plate 185). and the bird table had yet to be
seems to suggest a progressive loss of the invented, Blackbirds with white feathers were
pigment-producing cells rather than a already being noted (Thompson 1849; Johns
gradual reduction of tyrosinase activity. Grey 1862), while Watters (1853) stated that ‘…
hair in elderly people, for example, is caused white-marked varieties are nothing unusual.’
by a tyrosinase reduction; only the colour of Furthermore, many of the museum specimens
the hair is affected and not that of the skin. with white feathers from my study sample
In many bird species other than Blackbird, were collected in the 1800s (the earliest in
progressive greying does not appear to affect 1851). In addition, progressive greying is also
the melanin in the skin. rather common in the Fieldfare T. pilaris and
Progressive greying is clearly the most Ring Ouzel T. torquatus, two species not typi-
common colour aberration in the Blackbird, cally closely associated with humans (plates
but the causes are often unknown (see above). 186 & 187).

260 British Birds 111 • May 2018 • 250 –263

 White feathers in black birds

Progressive greying thus

remains something of an
enigma. We know that white
feathers are never present in the
juvenile plumage but that they
can develop any time after the
bird reaches maturity, and in
some individuals the condition
is more progressive than in
others. Since progressive
greying in the Blackbird often
also affects the pigment in the
skin (like the heritable human
pigment disease vitiligo), a heri-
table cause for this form cannot

Ian York
be ruled out. Although in most
cases there is no evidence for a
straightforward genetic basis to
184. Male Blackbird, UK, May 2014. In the third form of
progressive greying, there are no progressive greying, the entire plumage can be affected and
clear external factors either. The the condition probably progresses more rapidly; these appear
lack of a demonstrable external to be the birds that do become fully white (see plate 173).
cause for the condition may
mean that ‘becoming grey’
is, in fact, a natural feature of
the species, with a (complex)
genetic component, just as it is
in humans!

Acknowledgments
First of all, a massive thank you to all
participants in the BTO’s Abnormal
Plumage Survey; their records of odd-
coloured birds are a great help
Mary Payne
in understanding the occurrence
and nature of these aberrations.
Participants also supplied the
photographs used as plates 172–174,
176, 177, and 182–185 in this paper, 185. Female Blackbird, Stoke Mandeville, Buckinghamshire,
which are greatly appreciated. I should April 2011. Probably not all birds affected by progressive
also like to thank Tim Harrison and
greying become fully white, even if they live long enough.
Clare Simm, who organised the
Abnormal Plumage Survey, together Note that although this bird has over 75% white feathers,
with Kate Risely, Mike Toms and the the pigmentation in the feet is not or hardly affected.
other members of the BTO’s Garden
Ecology Team. I wish to thank David Noble-Rollin for Zoological Research Centre and Museum Alexander
his invaluable information and help in finding the initial Koenig in Bonn; Tineke Prins and Kees Roselaar,
analyses of the survey carried out by his father. Olivier Zoological Museum in Amsterdam (now Netherlands
Poncin is thanked for sharing his findings in 2010 on the Centre for Biodiversity Naturalis in Leiden); Sylke
Carrion Crows around Brussels, and the following Frahnert, Zoological Museum in Berlin; Cordula Bracker,
museum curators, in no particular order, are thanked Zoological Museum in Hamburg; Georges Lenglet,
for allowing me to examine the specimens under their Royal Belgian Institute of Natural Sciences in Brussels;
care: Paul Sweet and Peter Capainolo, American Anne Préviato, National Museum of Natural History in
Museum of Natural History in New York; Jiri Mlikovski, Paris; Bob McGowan, National Museums Scotland in
Natural History Museum in Prague; Christane Schilling, Edinburgh; and Malcolm Pearch, Harrison Institute in
Natural History Museum in Hannover; Anita Gamauf Sevenoaks, Kent. Last but by no means least, thanks to
and Hans-Martin Berg, Natural History Museum in Katrina van Grouw for editing, suggestions and
Vienna; Gerald Mayr, Senckenberg Museum in Frankfurt constructive criticism which improved the manuscript
am Main; Michaela Forthuber, Natural History Museum enormously.
in Braunschweig; Renate van den Elzen and Till Töpfer,

British Birds 111 • May 2018 • 250 – 263 261

 van Grouw

–– & Hume, J. P. 2016. The history and

morphology of Lord Howe Gallinule
or Swamphen Porphyrio albus (Rallidae).
Bull. Brit. Orn. Club 136: 172–198.
Harrison, C. J. O. 1963. Mottled plumage
in the genus Corvus, its causation and
relationship to fundamental barring.
Bull. Brit. Orn. Club 83: 41–50.
Harrison, J. M. 1957a. The ‘white wing-
barring’ and other variants in the
Carrion Crow and Rook. Bull. Brit.
Orn. Club 77: 84–85.
— 1957b. Significant pattern variations
in European Corvidae. Bull. Brit. Orn.
Club 77: 131–133.
Johns, C. A. 1862. British Birds in their
Haunts. Society for Promoting
Christian Knowledge, London.
Mahabal, A., van Grouw, H., Sharma,
R. M., & Thakur, S. 2016. How
common is albinism really? Colour
aberrations in Indian birds reviewed.
Dutch Birding 38: 301–309.
Malher, F. 2003. Les Corneilles
‘Bigarrées’ Corvus corone en Europe.
Alauda 71: 13–19.
Rankin, W. T. C. 1954. Variation in partial
albinism of Blackbird. Brit. Birds 47:
406.
Rollin, N. 1953a. Abnormal plumage
Harry Taylor, NHM

study. Dawn Song and All Day 1: 82.

— 1953b. A note on abnormally
marked Song Thrushes and Blackbirds.
Trans. Nat. Hist. Soc. Northumberland
and Durham 10: 183–184.
— 1959. White plumage in Blackbirds.
Bull. Brit. Orn. Club 79: 92–96.
186 & 187. Ring Ouzels Turdus torquatus (specimens at the — 1964. Abnormal Plumage. Bird
Natural History Museum, Tring: left, NHMUK 1987.24.313 and Research 2: 1–44.
right NHMUK 1996.41.2325, both collected between 1882 and — 1981. Breeding tests on albinistic
European Blackbirds. Bird Research
1900) and Fieldfares Turdus pilaris (left, NHMUK 1996.41.2335;
2: 5.
right, NHMUK 1987.24.321 (right), both collected between Sage, B. L. 1954. An unusual example of
1883 and 1890). Progressive Greying is not uncommon in symmetrical albinism in the Carrion-
these two species. Crow Corvus corone corone Linnaeus.
Bull. Brit. Orn. Club 74: 104.
References — 1956a. On the occurrence of ‘mottled’ plumage in
Band, R. M. 1956. Albinism related to age. Brit. Birds 49: the Carrion-Crow Corvus corone corone Linnaeus.
153–154. Bull. Brit. Orn. Club 76: 13–14.
Bosch, S. 2004. Schwarz und weiss: Weisse — 1956b. Notes on an aberrant Carrion-Crow Corvus
Gefiederanteil bei Aaskrahe Corvus corone in corone corone Linnaeus obtained in Hertfordshire.
Deutchland. Orn. Mitt. 56: 396–403. Bull. Brit. Orn. Club 76: 64–65.
Foott, K. G. 1956. Albinism related to age. Brit. Birds 49: — 1962. Albinism and melanism in birds. Brit. Birds 55:
500. 201–225.
van Grouw, H. 2012a. Geen kraai zo bont of er zit wel — 1964. Mottled plumage in the genus Corvus.
een vlekje aan; het fenomeen ‘witte veren’ in Bull. Brit. Orn. Club 84: 25–30.
kraaiachtigen. Het Vogeljaar 60: 3–20. Simms, E. 1978. British Thrushes. New Naturalist No.
— 2012b. What colour is that Sparrow? A case study: 63. Collins, London.
colour aberrations in the House Sparrow, Passer Slagsvold, T., Rofstad, G., & Sandvik, J. 1988. Partial
domesticus. International studies on Sparrows 36: 30– albinism and natural selection in the Hooded Crow
55. Corvus corone cornix. J. Zool. 214: 157–166.
— 2013. What colour is that bird? The causes and Snow, D. W. 1988. A Study of Blackbirds. BMNH, London.
recognition of common colour aberrations in birds. Spencer, K. G. 1956. Albinism related to age.
Brit. Birds 106: 17–29. Brit. Birds 49: 500.
— 2014. Some black-and-white facts about the Terluin, A. B. 1996. Opmerkelijke toename van partieel
Faeroese White-speckled Common Raven Corvus albinistische Zwarte kraaien Corvus corone corone in
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— 1998. Witgevlekte Zwarte kraaien Corvus corone Birds. Vol. 1. Reeve & Benham, London.
corone kampen met een tekort aan goede voeding. Watters, J. J. 1853. The Natural History of the Birds
Het Vogeljaar 46: 107–118. of Ireland, indigenous and migratory. Dublin.
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Thompson, W. M. 1849. The Natural History of Ireland:

Hein van Grouw, Curator, Bird Group, Department of Life Sciences,

The Natural History Museum, Akeman Street, Tring, Hertfordshire HP23 6AP;
e-mail h.van-grouw@nhm.ac.uk

Hein van Grouw has been a Bird Curator since 1997, first at the National Museum of Natural History Naturalis,
Leiden, the Netherlands, and now at the Natural History Museum in Tring. His main interest and ongoing
research is into the occurrence and correct identification of colour aberrations, heritable and non-heritable, in
European birds. This research, conducted over the last 19 years, involves both practical breeding experiments
with domesticated birds and examination of over 4,000 aberrantly coloured bird specimens in museum
collections.

Appendix 1. The abnormal plumage survey by the World Bird Research

Station, Glanton, Northumberland
Calvert Noble Rollin (1906–2004) had a lifetime’s interest in birds and their behaviour. After
training as a metallurgical chemist, he worked for the North Eastern Marine Engine Company in
Northumberland, at Wallsend. In 1930, upon the death of his father, Charles, he inherited Grey-
stones, the family house at Glanton, and started the Bird Research and Educational Station,
which opened to the public in 1931. During this time, he began to formulate his main areas of
study into the daily behaviour of birds and, in particular, the dawn chorus.
The Second World War forced Rollin to return to his former job as a metallurgical chemist on
Tyneside. However, at weekends he cycled to Greystones (a round trip of almost 120 km) and,
during the week, spent all his spare time watching and studying birds on the Tyne, sometimes
getting into trouble with the authorities as a possible German spy. After the war, he again left his
job and returned to Glanton to continue his ornithological work, now also with a growing
interest in abnormal plumage. He began comparative studies on domesticated birds to study the
inheritance of plumage abnormalities, in an attempt to understand what he believed to be non-
inherited plumage in wild birds. In 1952 he was asked to teach ornithology at the Extra-Mural
Department of Newcastle University, which he did up to the 1980s. This work provided a reser-
voir of enthusiastic and willing adult students to help in his research. At this time, he expanded
the scope of the research station at Glanton, which became known as the World Bird Research
Station, and continued his long-term study of abnormal plumage.
The inspiration for that study were three young Song Thrushes T. philomelos and two young
Blackbirds, which were hand-reared in May 1951 and which subsequently developed partly white
plumage. Rollin assumed that the loss of pigment in these birds was due to diet. Continued field
observations on Blackbirds in different areas convinced him that an intensive study of the distri-
bution of abnormally coloured Blackbirds and related species was needed to ascertain whether
the condition had any relationship with rural and urban environments. He began the study in
early April 1962 with a public request for reports of any wild bird with odd white feathers, pub-
lished in newspapers across the country. The final conclusions of that survey were never pub-
lished, though interim results were reported (Rollin 1964).
Based on that 1964 paper and unpublished work, it is clear that Rollin recognised that the vast
majority of cases of aberrant white feathers were not heritable and that the whitening was often
progressive. He also believed that this phenomenon was solely due to diet and occurred mainly in
urban areas. Calvert Rollin was well ahead of his time in his understanding of the nature of the
colour aberrations in birds. Yet despite his efforts, an understanding of aberrant white feathering
in Blackbirds (and other species) was still lacking. He would surely have been pleased to know
that his work eventually encouraged further research into the phenomenon.

British Birds 111 • May 2018 • 250 – 263 263