ArticlePDF Available

Abstract and Figures

Several groups of birds have suffered high extinction rates, especially rails, pigeons, parrots and passerines. Some island species that disappeared in the early 19th century, e.g. Lord Howe Gallinule Porphyrio albus, Rodrigues Parakeet Psittacula exsul and Mascarene Parrot Mascarinus mascarinus, are known from only a few skins and illustrations, whereas the Huia Heteralocha acutirostris of New Zealand is known from hundreds of specimens. Furthermore, two North American species-Passenger Pigeon Ectopistes migratorius and Carolina Parakeet Conuropsis carolinensis-which became extinct in the early 20th century, are also represented by hundreds of specimens. Other supposedly extinct bird species are enigmatic. Confusion exists concerning the unique specimens of Sharpe's Rail Gallirallns sharpei and Townsend's Bunting Spiza townsendi, paintings of a parrot from the West Indies and an aberrant white Huia, as well as aberrant specimens of the Critically Endangered Kakapo Strigops habroptilus. Much has been written concerning these birds and why they became extinct, or have become extremely rare, but few data are available concerning colour aberrations in certain specimens; the literature is also riddled with incorrect terminology. This paper addresses this shortfall and describes the various colour aberrations in these extinct and endangered birds and why they have occurred.
Content may be subject to copyright.
Julian P. Hume & Hein van Grouw 168 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Colour aberrations in extinct and endangered birds
by Julian P. Hume & Hein van Grouw
Received 20 December 2013
rails, pigeons, parrots and passerines. Some island species that disappeared in the
early 19th century, e.g. Lord Howe Gallinule Porphyrio albus, Rodrigues Parakeet
Psiacula exsul and Mascarene Parrot Mascarinus mascarinus, are known from
only a few skins and illustrations, whereas the Huia Heteralocha acutirostris of
New Zealand is known from hundreds of specimens. Furthermore, two North
American species—Passenger Pigeon Ectopistes migratorius and Carolina Parakeet
Conuropsis carolinensis—which became extinct in the early 20th century, are also
represented by hundreds of specimens. Other supposedly extinct bird species are
enigmatic. Confusion exists concerning the unique specimens of Sharpe’s Rail
Gallirallus sharpei and Townsend’s Bunting Spiza townsendi, paintings of a parrot
from the West Indies and an aberrant white Huia, as well as aberrant specimens
  Strigops habroptilus  
concerning these birds and why they became extinct, or have become extremely
rare, but few data are available concerning colour aberrations in certain specimens;
the literature is also riddled with incorrect terminology. This paper addresses
this shortfall and describes the various colour aberrations in these extinct and
endangered birds and why they have occurred.
       
and macaws) and Passeriformes (passerines) are all widely distributed, occurring on every
           
Walters 2012), especially those genera endemic to oceanic islands. This is primarily due to
        
exotic species. Many of these extinct taxa are known from very few specimens, particularly
those that disappeared in the 19th century. These include Lord Howe Gallinule Porphyrio
albus (ex. c.1844), Rodrigues Parakeet Psiacula exsul (ex. c.1875) and Mascarene Parrot
Mascarinus mascarinus (ex. c          
represented by just two known skins. In contrast, Huia Heteralocha acutirostris (ex. 1907)
of New Zealand is known from hundreds of specimens (Fuller 2000) and the Critically
  Strigops habroptilus by >1,000; both were particularly sought after
carolinensis (ex. 1918) and Passenger Pigeon Ectopistes migratorius (ex. 1914), of the
continental USA, are known from at least 720 skins (Luther 1996) and 1,532 skins (Hahn
(Rothschild 1907, Greenway 1958, 1967, Fuller 1987, 2000, Butcher 1992, Hume 2007, Hume
      
aberrations. The unique specimens of Sharpe’s Rail Gallirallus sharpei, obtained in 1865 and
of unknown provenance but considered an extinct species (Olson 1986), and Townsend’s
Bunting Spiza townsendi, collected once in 1834 and also considered extinct (Paynter 1970),
      , illustrations of an enigmatic
Amazona parrot from the West Indies, and a white Huia are also described.
Julian P. Hume & Hein van Grouw 169 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
This paper addresses the probable reasons why these colour aberrations occurred, and
elucidates some of the erroneous misconceptions associated with these birds. In particular,
     
describe any avian taxon exhibiting partial white feathering (van Grouw 2006, 2012, 2013).
LORD HOWE GALLINULE (SWAMPHEN) Porphyrio albus (White, 1790)
Endemic to Lord Howe, a small, remote island between Australia and New Zealand in the
Tasman Sea, c.600 km east of Australia, P. albus was considered common when discovered
            
predators, becoming extinct between 1832 and 1844 (Hindwood 1940). The population was
derived from Purple Gallinule P. porphyrio
status. Two specimens, as well as several contemporary illustrations and subfossil remains,
skin has accurate provenance data.
             
First Fleet vessels, the Lady Penrhyn
1991). The drawing depicts one white and two pied gallinules, and Smyth described the
      
demonstrate that the species was variably coloured, with some completely white, others
white with blue speckling, some entirely blue. Phillip (1789) described the adult female
Figure 1. Watercolour of live Lord Howe Gallinules
Porphyrio albus showing the various stages of
progressive greying discussed in the text. Fig.
1a (left) shows a pair exhibiting the same colour
aberrations as the extant specimens; illustration by
George Raper c.1790 (from Hindwood 1940). Fig. 1b
(above) shows the three colour stages described by
White (1790); illustration by Thomas Watling c.1792
(from Fuller 2000).
Julian P. Hume & Hein van Grouw 170 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Figure 2. Comparison of the two extant specimens of
Lord Howe Gallinules Porphyrio albus, both of which
     
Purple Gallinule P. porphyria: Fig. 2a (top) holotype
NMW 50761; Fig. 2b (above left) WML D3213; Fig.
2c (above right) artist’s impression of the Liverpool
specimen (Julian P. Hume); Fig. 2d (left) Eastern Purple
Gallinule P. p. melanotus BMNH 1887.5.2.66 dorsal and
ventral views (Hein van Grouw). Note that the holotype
is completely white.
Julian P. Hume & Hein van Grouw 171 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
as all white, but was informed that the male had some blue on the wings. White (1790),
followed by Latham (1824) and Gray (1862), stated that young were all black, turning bluish
grey, then pure white with maturity (Fig. 1b). White’s (1790) statement that juveniles were
strongly suggests that the species exhibited the aberration progressive greying, which can
onset of this condition, the bird gains an increasing number of white feathers with every
moult. In the early stages, they are usually spread randomly over the bird, but eventually
the entire plumage is white. Progressive greying may or may not be inheritable; some forms
are related to age, while in others the progressive loss of pigment cells is due to inheritable
disorders such as vitiligo (van Grouw 2013). Progressive greying is common in birds,
especially in the Corvidae, Passeridae and Turdidae (van Grouw 2012). The aberration
      
Howe Gallinule’s bill and legs retained normal coloration.
The widespread Australasian subspecies of Purple Gallinule, P. p. melanotus (Fig.
2d), still occurs on Lord Howe and some exhibit blue and white feathering (Mayr 1941,
Ripley 1977), but it appears that these birds had disappeared entirely by 1977. Mayr (1941)
considered the survival of blue individuals was due to them being less conspicuous,
after the original population of white birds had disappeared; however, Hindwood (1965)
thought that blue birds occasionally arrived from Australia and interbred with the resident
           
confusing taxonomic literature (Rowley 1875, Forbes 1901, Rothschild 1907, Mathews 1928),
resident on Lord Howe.
NMW (Naturhistorisches Museum Wien) 50.761, type (Fig. 2a). This specimen was
Fulica albus, Norfolk
Island’ (Pelzeln 1873) in error. It is considered the type, as White (1790) mentioned that the
bird on which his drawing was based was deposited in the Leverian collection (Pelzeln
1873). The skin, which was originally mounted, is in good condition, and the natural red
and thus were probably reddish in life. The specimen is all white, with no trace of yellowish
sheen or any purple feathering (contra Rothschild 1907: 144). The remiges and rectrices are
   
shorter than in the Liverpool specimen (Greenway 1967).
WML (Liverpool World Museum) D3213 (Fig. 2b–c). A mount, this specimen was
originally obtained by Sir Joseph Banks, but the collection date is unknown. It must have
            
went to the Bullock collection, was purchased at auction by Lord Stanley in 1819, then
(Forbes 1901). It is generally in good condition, considering its age, but with extensive
and is more concentrated on the head, especially near the dorsal surface of the shield. There
lateral rectrices are purplish brown. The remiges and rectrices are soft, and the primaries
shorter than in P. porphyrio (Forbes 1901, Greenway 1967). The bill, shield, iris, legs and
Julian P. Hume & Hein van Grouw 172 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
feet were bright red or yellow in life (Philip 1789, White 1790), but the bill and feet in the
Liverpool skin have been painted red.
Rothschild (1907) correctly suggested that P. albus was not albinistic, but that the
population was in the evolutionary process of becoming pure white. Our analysis of
   contra
two extant specimens is caused by a presumably inheritable form of progressive greying.
all white. The individuals that occurred on Lord Howe until 1977 exhibiting white and blue
coloration were probably a local population of P. p. melanotus, which also carried the gene
for progressive greying. They too are now extinct.
SHARPE’S RAIL Gallirallus sharpei
Several rails are known from unique skins or illustrations, and the enigmatic Sharpe’s Rail
Gallirallus sharpei   
      Gallirallus philippensis (Olson 1986), an extremely
            
Guinea, New Caledonia and New Zealand, as well as Australia and Indonesia (Ripley 1977,
of which are not threatened, but Macquarie Island Rail G. p. macquariensis became extinct
c.1870 due to the introduction of feral cats and Wekas G. australis
The type of G. sharpei was received at the Rijksmuseum van Natuurlijke Historie, Leiden
1893, Olson 1986). The original range is unknown, but H. Schlegel thought that the bird was
       Pardirallus maculatus  
error that initiated great confusion over its true provenance (R. B. Sharpe in
   
allocation (Olson 1986), described it as a distinct species from South America in honour
in Peters 1934)
placed the rail in Hypotaenidia and, as it was never recorded again, this prompted Peters to
suggest that it might be an extinct species. Hypotaenidia
Gallirallus (Olson 1986), and Olson concluded that Sharpe’s
Rail, which was similar in many aspects of size and morphology to volant G. philippensis,
should be placed there too.
On many occasions, unique bird skins have been ignored by modern ornithologists,
especially those collected on islands during the 18th and 19th centuries, which has
commonly resulted in their relegation to dubious status. Based on the available evidence,
            
coloration, it probably represented an extinct species, and speculated that the skin was
procured from Java, Sumatra or Borneo. Unusually among island rails, which are often
          
reduction in wing length (Olson 1986).
RMNH (Rijksmuseum van Natuurlijke Historie, Leiden) 87485 (Fig. 3). Sharpe’s Rail
remiges dull brown, strongly barred white; sides of head, chin, throat and neck grey; rest of
Julian P. Hume & Hein van Grouw 173 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
         
     
aberration does not cause a qualitative and / or quantitative loss of pigment.
In general the appearance of a melanistic bird is dark, predominantly blackish, but this
           
their usual boundaries (the rest of the plumage is often somewhat darker too); (2) all
pigment distribution is changed, but plumage is not necessarily darker.
feathers’, which means that melanism is not necessarily an increase of pigment, but may be
(van Grouw 2013).
The possibility that G. sharpei represents a colour aberration was not considered until
mtDNA revealed that the specimen can be referred to G. philippensis (Naturalis unpubl.
data). Therefore the unique specimen of Sharpe’s Rail is not an extinct species (van Grouw
Figure 3. Sharpe’s Rail Gallirallus sharpei RMNH 87485 (left) exhibiting category 3 melanism compared with
G. philippensis RMNH 185133 (right) (© Steven van der Mije / Naturalis)
Julian P. Hume & Hein van Grouw 174 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
PASSENGER PIGEON Ectopistes migratorius (Linnaeus, 1766)
Probably once the most abundant bird species on Earth, its extinction is one of the most
             
abundant could be exterminated so quickly. Its range was inextricably linked to the eastern
deciduous forests of North America and Canada, from the Great Plains east to the Atlantic,
north to southernmost Canada, and south to Virginia and northern Mississippi. In winter
 
Cuba (Schorger 1955, Gibbs et al. 2001). Early accounts describe the immense numbers
of individuals that congregated during the nesting season and especially on migration,
        
in the early 1800s, and coincided with an increase in deforestation and commercial
exploitation, particularly following the development of new railroads and telegraph
systems after the American Civil War. Wherever the pigeon concentrated in numbers, it
was subject to wanton slaughter; they were in huge demand for meat and the feather trade
(Butcher 1992). By the end of the century the species was almost extinct. The last recorded
wild specimen was shot on 24 March 1900 (Fuller 2000). A few were kept in private aviaries,
but they proved poor breeders; the last, a captive female called Martha, outlived the rest by
four years. She died on 1 September 1914 in Cincinnati Zoo (Shufeldt 1915).
Figure 4. Passenger Pigeon Ectopistes migratorius BMNH 1939.12.9.4071 exhibiting the brown aberration (left)
van Grouw / © Natural History Museum, Tring).
Julian P. Hume & Hein van Grouw 175 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Passenger Pigeon is known from at least 1,532 skins and 16 skeletons (Hahn 1963).
It was a large dove with long wings and a long graduated tail, and adults were sexually
dichromatic. Males had the head bluish grey; display area at back of neck iridescent bronze,
          
    
primaries and secondaries darker greyish brown; secondaries edged white; throat and
   
          
and orbital ring greyish blue.
BMNH (Natural History Museum, Tring) 1939.12.9.4071 (Fig. 4). This aberrant
specimen, an adult female, has no provenance other than it formed part of the Rothschild
collection at Tring. Rothschild had a great interest in colour aberrations and this specimen is
tail where grey normally occurs; the primaries, lower breast and belly are white; the black
            
            
of eumelanin’ (van Grouw 2013). This mutation is the most common in wild birds, and
               
females occur in the wild, as females need only one gene to express the recessive brown
mutation. In contrast, to produce a brown male, a heterozygous father and brown mother
are essential; the likelihood of this occurring under natural conditions is extremely remote.
The bill and feet are usually slightly paler in brown (not discernible in this specimen), but
eye colour remains the same (van Grouw 2012). The result of this aberration is that the black
      
whereby old feathers can be much paler than fresh ones (van Grouw 2012, 2013). The
RODRIGUES PARAKEET Psiacula exsul (Newton, 1872)
             
   
also observed other parrots (our translation from the original manuscript):
Necropsiacus rodericanus]
are larger than a pigeon and have a very long tail, the head big as well as the beak;
most are from the southern islets to the south of island where they eat a small black
seed that produces a small shrub whose leaves have the scent of lemon, and come to
        
Julian P. Hume & Hein van Grouw 176 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
 Psiacula exsul] is slightly smaller, and much more
 P. exsul] is small, all green
and the beak black.’
visiting the main island for water. The astronomer Pingré, who was on Rodrigues to observe
  
rare, Necropsiacus
 
             
 
described (Newton 1872). A second transit expedition to the island took place in 1874,
           
1879, Hume et al. 2014). The parakeet was by now extremely rare, and Slater remarked on
       
Palaeornis exsul. I saw one specimen of this bird as I was going from my camp to a
distant cavern: unfortunately I had not my gun with me or I could easily have shot it as
Figure 5. The two extant specimens of Rodrigues Parakeet Psiacula exsul exhibiting the parblue aberration.
Male UMZC18/Psi/67/h/I (right) lateral and dorsal views; holotype female UMZC18/Psi/67/h/I (left) lateral
and dorsal views (Julian P. Hume).
Julian P. Hume & Hein van Grouw 177 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
it came within thirty feet of me: I believed it to be a male at the time, and it is probably
the one procured since by Mr Caldwell; I never had time to hunt for it subsequently.
that he had seen two specimens at the N. end of the Island, but I had never time to visit
that region.
Assistant Colonial Secretary, William James Caldwell (1875), who arrived on Rodrigues
12 May 1875, saw several parakeets during his stay, but was unable to obtain a specimen
himself. However, he did receive a male from a local resident and ship’s pilot, William
              
1876); it was probably that seen by Slater. This was the last time the parakeet was recorded.
CUMZ (University Museum of Zoology, Cambridge) 18/Psi/67/h/I holotype (Fig.
overall dull greyish glaucous, darker above, with a blue or green sheen depending on the
  
blue, becoming greener on vent; black collar from nostrils to sides of head; bill black, iris
black with yellow ring, legs and feet greyish black. Originally preserved in alcohol, the
specimen was removed from solution when it was described (Newton 1872).
CUMZ (University Museum of Zoology, Cambridge) 18/Psi/67/h/I 
in being larger and in having the top of head clearer glaucous, less grey; black stripe
more distinct, extending upwards from nostrils to nape of neck and almost meeting there;
primaries with dull black patch on inner web near tip; central secondaries dusky black;
            
Newton 1876). The feathers exhibit more wear than the female, and thus are less well
The unusual coloration of P. exsul has caused much debate, so it is important to establish
a general overview of parrot coloration. The predominant greens of most parrots are not
true colours as they are not formed by green pigments; instead, the green is produced by
in all birds, melanin, or more precisely, eumelanin, is one of the pigments that determines
colours in parrots. Eumelanin is normally black or dark brown, but depending on the way
the microscopic pigment granules are arranged (i.e. their density) in the feather cells, it
deeper cells of the feather barbs, together with their special structure, distort the light as it
passes through, making the feathers appear blue. Thus, blue is a structural colour and not
   
   
              
barb, and combined with the structural blue coloration, creates the bright shades of green
characteristic of most parrot species (Martin 2002).
            
probably the commonest mutation in parrots and is, for obvious reasons, called blue. In
mutation to appear, a bird must inherit a gene for the mutation, in this case blue, from both
parents. The coloration caused by blue mutations varies in shade, depending on the normal
colours (wild type) in the relevant species. For example, in Psiacula the blue coloration
will be evenly distributed and intense, as is the original green of these species. In contrast,
an uneven distribution and intensity of green can result in an uneven distribution of blue
Julian P. Hume & Hein van Grouw 178 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
    
            
described P. exsul 
the angle of light. This suggests that the cause of the colour aberration was due to a parblue
mutation rather than true blue.
MASCARENE PARROT Mascarinus mascarinus (Linnaeus, 1771)
         
species disappeared before anything was recorded of its ecology. A few captive birds
arrived in Paris during the late 18th century, by which time it was probably already extinct
         
           
Greenway 1958, 1967, Fuller 1987, 2000, Forshaw 1989); however, this is extremely unlikely
and the species was almost certainly extinct by c.1800 (Hume 2007). The specimen described
by Hahn was not preserved, but two exist, the holotype in Paris (MNHN 211; Fig. 6a,c) and
     
      
Mascarene Parrot (Latham 1781), as the skin has some white feathering (Fig. 6c). Latham
        Mascarinus], with the mask in front. The
but irregularly sprinkled with white feathers throughout; the tail, part white, part
brown, but not regular; some of the quills and tail feathers being wholly white, while
the opposites, which should have answered them, were white and brown: it is a
particular and beautiful lusus naturae.’
Both skins are now faded, and in the 1790s, the wings and tail of the Paris specimen
 
faded coloration of the specimens and early descriptions and illustrations, which depict
the bird with a purplish head, black mask, pale brown body with darker wings and tail,
and white bases to the lateral rectrices (see Hume 2007). However, birds described in life
        
             
et al. 1999], a black hood on the head, the beak very large and
feathers white at base; bare skin around eye region, bill, bare skin around upper bill and iris
of the original coloration is discernible in the two extant skins (Fig. 6); only the black mask
and red bill are clearly visible.
MNHN (Muséum National d’Histoire Naturelle, Paris) 211 (Fig. 6a). No details exist
as to this specimen’s provenance, but probably it was one of the live individuals that
resided in Paris in the late 1700s (Hume 2007). Overall coloration uniform pale brown, with
Julian P. Hume & Hein van Grouw 179 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Figure 6. The two extant specimens of Mascarene Parrot Mascarinus mascarinus: Fig. 6a (top left) holotype
MNHN 211 (Julian P. Hume); Fig. 6b (top right) NMW 50.688, dorsal and lateral views (Julian P. Hume);
the Paris holotype (centre left and centre) is not discernible in the Vienna specimen, and the tail has been
reconstructed. The irregular white feathering in the Vienna skin is due to poor diet in captivity.
Julian P. Hume & Hein van Grouw 180 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
paler; head brown with black mask; tail dark brown with white bases to lateral feathers; bill
bright red; legs greyish brown.
NMW (Naturhistorisches Museum Wien) 50.688 (Fig. 6b). The Vienna specimen was
purchased from the sale of the Leverian collection in 1806 (Pelzeln 1873, Schifter 1994).
Nothing else is known about it. Overall uniform brown, greyer on the head and neck, with
individual white feathers on mantle, back, rump and underparts; primaries pure white (one
side has been clipped), secondaries white with dark brown tips; head brown with black
mask; tail has white and dark brown feathers; legs and feet reddish brown.
The pale brown coloration is clearly an artefact of fading due to light exposure, from
grey to brown (Hume 2007, van Grouw 2013), which has occurred in other ancient skins
that were predominantly grey or black in life, but are now brown, e.g. a Captain James Cook
Moho nobilis, and Townsend’s Bunting Spiza townsendi (pers. obs.,
 
white feathering on the body, with white primaries and asymmetrical white tail feathers,
            
         
alive in Paris during the 1770s (Hume 2007). The clipped primaries further suggest that it
was caged. The dietary requirements of parrots were doubtless poorly known in the 18th
century, and almost certainly inadequate for maintaining a healthy bird. This specimen of
tyrosine from its food, which is necessary for normal melanin synthesis. In consequence,
      
present. However, vasa parrots Coracopsis sp. and Mascarene Parrot are the only species
   Mascarinus are obscure and not satisfactorily resolved. Based on
           
           
Coracopsis nigraet al
et al
convergent evolution or supports a close relationship remain unclear.
CAROLINA PARAKEET Conuropsis carolinensis (Linnaeus, 1758)
Formerly present over much of the eastern USA, from Florida and southern Virginia
             
subspecies, Eastern Carolina Parakeet C. c. carolinensis and Western Carolina Parakeet C. c.
when described in the late 1700s, but as early as 1831 the species was rapidly declining as
a result of persecution due to its habit of damaging fruiting crops. The birds disappeared
an increase in collecting for aviculture. By the end of the 19th century, Carolina Parakeet
  
low and mortality high. The last bird died in Cincinnati Zoo on 21 February 1918 (Hume
expired just four years earlier.
Carolina Parakeet is known from at least 720 skins and 16 skeletons (Hahn 1963, Luther
1996) and c            
The nominate subspecies was generally green with a bluish tint, paler on the underparts;
Julian P. Hume & Hein van Grouw 181 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
forehead, lores, bare skin around eye and upper cheeks orange; rest of head and upper
part of neck, bend of wing, carpal edge and thighs yellow; outer webs of primaries yellow
     
   
and being generally paler. As far as we are aware, there has not previously been any
description of colour aberrations in the species.
NMW (Naturhistorisches Museum Wien) 50.795 (Fig. 7a). Purchased from the
Leverian sale in 1806, it appears that it may have been a captive individual (data from label).
    
and back being replaced by brownish orange and the underparts with yellow, with more
  
               
also replaced by orange, but the bill, legs and feet are normal. A second captive specimen
(NHMW 50.682) exhibits similar coloration and was also purchased from the Leverian sale.
The aberrant coloration exhibited by these birds is caused by an increase and abnormal
             
(Martin 2002). In very rare cases, the increase of red pigment is due to inheritable factors
          
persistent reddish feathering, and lack of direct sunlight, presumably resulting in a lack
of vitamin D, may also be a contributory factor. Given that the Vienna specimen was in
         
GEORGE EDWARDS’ PARROT Amazona sp. (Fig. 8a)
          
produced a watercolour of an unknown parrot (see Fuller 2000: 209; Fig. 8a). The painting
Figure 7a (left). Carolina Parakeet Conuropsis carolinensis
Julian P. Hume & Hein van Grouw 182 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
     
parrot from Jamaica. Drawn from Nature
the size of life by G. Edwards, July 1764.’
On the back in Edwards’ own hand is a
more detailed description, repeated here
       
yellow, the colours of the upper sides
casting faintly through them. This bird was
lent to me by Dr. Alexander Russel and is
preserved in his collection. It was shot in
Jamaica and brought Dryed to England. The
people on Jamaica did not remember Ever
to have Seen one of this species of Parrots
Geo. Edwards, July, 1764.
Some of the fethers have their tipps red and
Others have them yellow. The fethers on the
under sides, Back and rump have yellow
Figure 8a (top). Watercolour dated July 1764 by George Edwards of an Amazona parrot, probably A. collaria,
exhibiting the opaline aberration; the skin was taken on Jamaica and brought to England, but its whereabouts
are unknown (© Errol Fuller). Fig. 8b (below). The two extant Jamaican Amazona  
Amazon A. collaria           
Amazon A. agilis BMNH 1845.4.29.3, second from left dorsal, and second from right ventral views, alongside
a Cuban Amazon A. leucocephala BMNH1890.6.1.158, third from left dorsal, and far right ventral views (Hein
van Grouw / © Natural History Museum, Tring).
Julian P. Hume & Hein van Grouw 183 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Because of the uniqueness of this specimen and the fact that it is associated with one
of the West Indian Islands, where so many parrot species have become extinct (Rothschild
             
George Edwards’ parrot has been discussed as a possibly extinct species (Shuker 1999,
Fuller 2000). It bears a strong resemblance to an Amazona parrot, a widespread genus in
Furthermore, two extant AmazonaA. collaria,
with a mean 25.62 cm in total length (nA. agilis, mean 23.83
      c.24.3
cm long (from top of head to tail tip), is intermediate between A. agilis and A. collaria, and
fractionally larger than A. agilis. However, wing length is shorter, while combined head and
bill length is much larger than either species; bill depth is intermediate between the two.
from the illustration must be viewed with great caution.
   
      
secondaries, and green uppertail with outer tail feathers red at base of inner webs. Edwards’
depiction has predominantly red coloration, but more importantly it shows the blue in the
            
      
Although not from Jamaica, if there is any doubt concerning the collection locality, Cuban
Amazon A. leucocephala (Fig. 8b) might also be considered as another strong candidate for
     
A. collaria.
Measurements of Jamaican Amazona parrots compared with Edwards’ Parrot painting. All specimens held
deviation]. Edwards’ Parrot falls within low range of A. agilis in all measurements except head length and
bill depth, which are too large compared with other measurements, suggesting that the painting was not
accurately drawn to scale.
Species    
Amazona collaria    
Amazona agilis    
Edwards’ Parrot    
Julian P. Hume & Hein van Grouw 184 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
KAKAPO Strigops habroptilus Gray, 1845
Formerly occurred on North Island, South Island and Stewart Island, and prior to human
occupation of the islands had one of the widest distributions of any New Zealand bird
        
 
especially rats, mustelids, domestic cats and dogs (Sutherland 2002), rapidly eliminated
it from most areas, and the demand for museum skins in the late 19th century, when
thousands were collected (Butler 1989), reduced the species to a few remote localities. The
last report on North Island was in 1927, three males were recorded on South Island in 1987,
but have not been recorded since, and the last record on Stewart Island was in 1997, when
    et al. 2006). Due to
   
           
           
          
           
individual variation. The bill is pearly grey with an ivory tip, iris dark brown, and legs and
CMNZ (Canterbury Museum, New Zealand) Av2059 (Fig. 9a). This specimen was
  
Bay in Preservation Inlet (Galbreath 1989). Walter Buller, the New Zealand ornithologist
and dealer, kept it alive on a diet of potatoes before it became part of his collection (Butler
was vivid canary yellow, fading to pale yellow on the cheeks and throat; the bill was white
it to be one of his most precious specimens.
The yellow coloration is almost certainly due to the mutation ino, which can be
. Among parrot
breeders, this mutation is known as lutino, because of the predominant yellow colour.
  
AM (Auckland War Memorial Museum) LB8526 (Fig. 9b). A second aberrant specimen,
a relaxed mount, was collected by G. Mueller, a surveyor and engineer, in Jackson Bay,
    
  
wings. This aberration vaguely resembles certain forms of recessive leucism in parrots
(Martin 2002), but the overall appearance strongly suggests opaline (see Edwards’ Parrot),
      
melanin only in the feather tips (especially on head and mantle), the absence of melanin
    
Julian P. Hume & Hein van Grouw 185 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
parrots (Martin 2002), and if our opaline
determination is correct, the AM specimen
is probably female.
BMNH (Natural History Museum,
Tring) 1853.6.9.1 (Fig. 9d). The reduction
of melanin resulting in yellow is not the
Figure 9. Three colour aberrations exhibited by
 Strigops habroptilus: Fig. 9a (top left). Ino
aberration CMNZ Av2059, dorsal, lateral and ventral
     
Zealand); Fig. 9b (top right) opaline aberration AM
LB8526, dorsal, lateral and ventral views (© Jason
  
(centre left) originally unpublished painting of CMNZ
      
parblue aberration BMNH 1853.6.9.1 type of S. greyi,
dorsal, lateral and ventral views (© Natural History
     
in the collection of Errol Fuller (Julian P. Hume); Fig.
     
1927.12.18.1 (Hein van Grouw / © Natural History
Museum, Tring) .
Julian P. Hume & Hein van Grouw 186 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
    
One notable specimen (BMNH 1853.6.9.1) is the holotype of S. greyii (Fig. 9d), which has
an intense blue hue to the upperparts and some on the underparts, and was originally
    
feathers are particularly worn, which strongly suggests that it was kept in captivity. In both
specimens, the upperparts exhibit the more extensive and stronger blue hue. Although the
              
Jones in li. 2013), so it is more likely that the Tring specimen was caught because it was an
unusual coloration and subsequently kept in captivity. We believe, therefore, that both of
Because this mutation can be extremely variable depending on the wild type, the
phenotype can exhibit blue coloration that is unevenly distributed. In the case of a parblue
HUIA Heteralocha acutirostris (Gould, 1837)
being relatively short and slightly decurved, and the female’s much longer and heavily
for chiseling at hard substrates such as decaying wood to extract insect larvae, whereas
females used their long, thin bill for probing crevices (Buller 1873). The fossil record reveals
time Europeans arrived the species was largely restricted to the south of the island. Huia
               
1907 (Myers 1923) despite claims that it survived until at least the 1960s.
Huia is known from hundreds of skins (Fuller 2000); a unique egg and subfossil
remains are at the Museum of New Zealand, Wellington. General coloration was black with
feet black. The sexes were similar; the juvenile duller, with a brownish wash. A specimen
was told that the specimen was for sale, but when he inquired about it, the owner had
already sold the bird (Buller 1905); its whereabouts are now unknown.
and therefore the dark eye is not reliable; it is unlikely that the individual exhibited true
albinism. Albinism, from the Latin Albus  
in feathers, eyes and skin’ (van Grouw 2013). The complete lack of melanin is due to
the hereditary absence of the enzyme tyrosinase in the pigment cells, as this enzyme is
            
Julian P. Hume & Hein van Grouw 187 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Vevers 1960). In almost all cases, this results in a completely colourless bird, whereas the
red or pinkish hue present in the eyes and skin is caused by blood being visible through the
colourless tissue. Albinos rarely occur in the wild, although the mutation is not uncommon
in birds. Their scarcity is due to the absence of melanin in the eye, which makes them
        
poor depth of vision. Therefore, it is not primarily the white plumage that makes albinos
vulnerable in the wild, but their poor eyesight; most succumb to starvation and predation
eliminates albinism as a cause. It is more likely that progressive greying, leucism or ino
was involved. In progressive greying, the bird would have moulted several times before
becoming entirely white, but an aberrant pied Huia would surely have been a target
for collectors; it is unlikely that such a bird went unnoticed, especially as it survived to
adulthood. Leucism, which is a congenital and heritable absence of pigment cells from all
of the skin areas where they would normally provide the growing feather with pigment,
is very rare in wild birds (van Grouw 2012, 2013), and thus probably can be ruled out.
As ino is not uncommon in wild birds, and individuals can be all white, it is the most
likely explanation for the aberration. The fresh plumage of an ino may show some very
      
bird almost pure white. Furthermore, although the eyes of an ino are also reddish due to
    
survive comparatively well in the wild and the mutation is not uncommon, we believe that
Heteralocha acutirostris
Julian P. Hume & Hein van Grouw 188 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Although we cannot be absolutely certain about the colour aberration of this Huia
without a specimen, the fact that the illustrated bird is a female strongly supports the
TOWNSEND’S BUNTING Spiza townsendi (Audubon, 1834)
Chester County, Pennsylvania, and described the following year (Audubon 1834). There
has been much debate as to its status, it being variously considered an extinct species
(e.g. Paynter 1970, AOU 1983), a colour variant of Dickcissel S. americana (Parkes 1985) or
a hybrid female S. americana × male Blue Grosbeak Passerina caerulea (E. Coues in Sharpe
1888). The type (USNM 10282) is in poor condition, with the original coloration now barely
discernible; the specimen has foxed, producing an overall wash of pale brownish beige (Fig.
Audubon (1834; Fig. 11c) described the fresh plumage as having the upperparts, head,
           
streaked black on interscapular area; eyestripe, chin, throat, central line on underparts
      
            
back, wings darker, rusty patch on shoulder, and superciliary stripe and underparts yellow,
      
  
speckled. Juvenile females are duller and browner, and have black or brown speckling on
upper breast.
Parkes (1985) suggested that Townsend’s Bunting was a female colour variant of
Dickcissel that lacked normal carotenoid pigments, resulting in the complete absence of
yellow. Holt (2002–03) questioned this conclusion, as Townsend explicitly stated that the
comparison of adult males, females and juvenile Dickcissels with the unique skin and
original illustration of Townsend’s Bunting (Audubon 1834) suggests that the situation is
far more complex. For example, the lack of yellow due to diet or a single genetic change
            
(Martin 2002).
If Townsend’s Bunting was a female Dickcissel as suggested by Parkes (1985), the
diagnostic brown crown and cheeks should still be present. Townsend stated that it was
a male, so it should have exhibited the black throat patch, unless juvenile. Finally, Holt’s
(2002–03) suggestion that the specimen was a juvenile male is supported by the lack of a
full black throat patch, but juvenile males are brown above, not grey. In the Townsend’s
Bunting specimen, there is black speckling (eumelanin) concentrated on the sides of the
At this stage, we cannot determine if the specimen of Townsend’s Bunting is the result
of hybridisation, a colour aberration, or caused by some other genetic change. That it might
be an extinct species is highly unlikely, but it cannot be ruled out. No molecular work has
been undertaken on the skin, but mtDNA analysis may be the only way to resolve this
Julian P. Hume & Hein van Grouw 189 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Figure 11. Townsend’s Bunting Spiza townsendi: Fig.
11a (top) holotype, USNM 10282, male (Julian P.
Hume); Fig. 11b (above left) illustration showing
the specimen today (above), and how it would
have looked when described by Audubon in 1834
(below) (Julian P. Hume); Fig. 11c (above right)
from Audubon (1834); Fig. 11d (left) Dickcissel
S. americana, from left to right in lateral view,
adult male BMNH 1899.2.1.4108, juvenile male
BMNH 1885.13.14.147, and adult female BMNH
1899.2.1.4126 (Hein van Grouw / © Natural History
Museum, Tring). The cause of this probable colour
aberration is unclear.
Julian P. Hume & Hein van Grouw 190 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Colour aberrations are not uncommon in birds (van Grouw 2006, 2013), so it is
unsurprising that they occur in extinct and threatened species. However, few of these
extinct bird colour aberrations had been analysed, despite the specimens themselves
being apparently well known. The factors involved in producing unusual coloration can
be complex, and it appears that in some species discussed here, illness and poor diet in
captivity resulted in aberrant plumage. There may also be a bias in collecting from wild
populations, e.g. the parblue aberration of Psiacula exsul, white Porphyrio albus or parblue
and yellow specimens of Strigops habroptilus
    
of Passenger Pigeon, >700 skins of Carolina Parakeet, and hundreds of Huias having been
collected, we have only located single colour aberrations of Passenger Pigeon and Huia,
and two Carolina Parakeets, none previously described in the literature. During the 19th
century, the demand for unusual colour varieties of birds was extremely high and examples
caused by physical factors and not by true genetic aberrations.
Colour aberration Cause  Species exhibiting
Greying Partial or total lack of melanins in
feathers due to progressive loss of
pigment cells in some or all of the
skin areas with age.
coloured feathers. Bill and feet
Lord Howe Gallinule
Porphyrio albus
Melanism Abnormal deposit of melanin in
skin and / or feathers. 
distribution is changed, but
plumage is not necessarily darker.
Sharpe’s Rail Gallirallus
Brown 
due to incomplete synthesis
(oxidation) of eumelanin.
Black becomes brown. Passenger Pigeon
Ectopistes migratorius
 Poor physical condition in
captivity. 
spread through the plumage. Carolina Parakeet
Conuropsis carolinensis
 
diet, resulting in disturbed melanin
Random white feathering
Mascarene Parrot
Mascarinus mascarinus
Opaline(1) Reduced melanin distribution,
especially in the body plumage,
and an enhanced spread of
A predominant reddish coloration
with minimal dark (melanin)
plumage markings.
Edwards’ Parrot Amazona
cf. collaria
Opaline(2) Reduced melanin distribution,
especially in the body plumage. Predominant yellow coloration
with minimal dark (melanin)
plumage markings.
Ino(1) Strong qualitative reduction of
melanin. 
visible Strigops
Parblue(1)  Coloration between green and blue. Strigops
Parblue(2) As above. Coloration between green and blue. Rodrigues Parakeet
Psiacula exsul
Ino(2) As above 
that is rapidly bleached to white by
exposure to light. Paler bare parts,
reddish eyes.
Huia Heteralocha
Undetermined Lack of lipochrome pigments?
Hybridisation? Loss of yellow pigmentation Townsend’s Bunting Spiza
Julian P. Hume & Hein van Grouw 191 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Buller was considered to be one of his most prized possessions.
In ornithology in general, colour aberrations are poorly understood, which has been
further exacerbated by incorrect terminology (van Grouw 2006, 2012, 2013). For example,
often applied to extinct birds too, e.g. the Vienna Mascarene Parrot. The inherent condition
 
the individual lacks the enzyme tyrosinase, or white feathering can be explained by other
factors, e.g. leucism, ino, progressive greying and poor diet (van Grouw 2006, 2012, 2013).
Studies of extant birds that exhibit unusual coloration can assist understanding of why
similar aberrations occurred in extinct species, and can sometimes provide new information
about the bird’s ecology or a specimen’s history. In these cases in particular, it is important
to understand the processes involved and to use correct terminology (Table 2). The aim of
this study is to highlight extinct or endangered bird species that exhibit colour aberrations
and we would welcome hearing from curators with additional specimens of interest. It is
probable that many more examples exist than those discussed here.
     
(Naturalis) photographed specimens, and Alison Harding (NHM) made accessible rare books in her care. We
for kind hospitality and access to collections; Ross Galbreath for providing information on the white Huia;
    
Abbé Alexandre-Gui Pingré: Courser Venus, voyage scientique à Rodrigue en 1761.
American Ornithologists’ Union (AOU). 1983. Check-list of North American birds. Sixth edn. AOU, Washington
Audubon, J. J. 1834. Ornithological biography, vol. 2. Robert Havell, London.
Cold Spring
Harbour Symp. Quant. Biol. 74: 345–53.
BirdLife International. 2013. Species factsheet: Strigops habroptila. (accessed 19 May 2013).
Brisson, M. J. 1760. Ornithologia
Buller, W. L. 1873. A history of the birds of New Zealand. John van Voorst, London.
Buller, W. L. 1888. A history of the birds of New Zealand, vol. 1. Second edn. John van Voorst, London.
Buller, W. L. 1905. Supplement to the ‘Birds of New Zealand,’ vol. 2. Privately published, London.
Butcher, E. H. 1992. The causes of extinction of the Passenger Pigeon. Current Orn. 9: 1–36.
Butler, D. 1989. Quest for the Kakapo. Heinemann Reed, Auckland.
Notes Leyden Mus. 15: 274–275.
Caldwell, J. 1875. Notes on the zoology of Rodriguez. Proc. Zool. Soc. Lond. 1875: 644–647.
Lost land of the Dodo.
Les voyages fait par le sieur D.B. aux isles Dauphine ou Madagascar et Bourbon ou Mascarenne
es années 1669, 70, 71 et 72. Claude Barbin, Paris.
Forbes, H. O. 1901. Note on Lord Stanley’s Water Hen. Bull. Liverpool Mus. 3: 62–68.
Forshaw, J. M. 1989. Parrots of the world. Third edn. Landsdowne Editions, Melbourne.
The nature of animal colours
Fuller, E. 1987. Extinct birds. Viking, London.
Fuller, E. 2000. Extinct birds. Revised edn. Oxford Univ. Press.
Galbreath, R. 1989. Walter Buller: the reluctant conservationist. GP Books, Wellington.
Pigeons and doves. Pica Press, Robertsbridge.
Gibbs, G. 2007. Ghosts of Gondwana; the history of life in New Zealand
Gray, G. R. 1862. A list of birds of New Zealand and the adjacent islands. Ibis 4: 214–254.
Julian P. Hume & Hein van Grouw 192 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
Greenway, J. C. 1958. Extinct and vanishing birds of the world
Protection, New York.
Greenway, J. C. 1967. Extinct and vanishing birds of the world. Second edn. Dover, New York.
van Grouw, H. 2006. Not every white bird is an albino: sense and nonsense about colour aberrations in birds.
Dutch Birding 28: 79–89.
van Grouw, H. 2010. How to recognise colour aberrations in birds (in museum collections). J. Afrotrop. Zool.
Spec. Iss. 2010: 53–59.
van Grouw, H. 2012. What colour is that sparrow? A case study: colour aberrations in the House Sparrow
Passer domesticus. Intern. Stud. Sparrows 36: 30–55.
van Grouw, H. 2013. What colour is that bird? The causes and recognition of common colour aberrations in
birds. Brit. Birds 106: 17–29.
Phil. Trans. Roy. Soc., extra vol. 168: 423–437.
Hachisuka, M. 1953. The Dodo and kindred birds, or the extinct birds of the Mascarene Islands
Hahn. C. W. 1834. Ornitologische Atlas oder naturgetreue Abbildung und Beschreibung der aussereuropäischen
Vögel. Erste Abteilung: Papagaien. C. H. Zeh’sche Buchhandlung, Nurnberg.
Hahn, P. 1963. Where is that vanished bird: an index to the known specimens of extinct and near extinct North
American species
Emu 40: 1–86.
 Emu 65: 83–95.
                
preliminary account. Rec. Aust. Mus. Suppl. 27: 85–100.
Holt, J. 2002–03. Notes on Audubon’s “mystery” birds. Cassinia 70: 22–24.
Zootaxa 1513: 1–76.
Extinct birds
   
century cave exploration on Rodrigues Island, Indian Ocean. Hist. Biol. 34: 1–22.
Birds of Lord Howe Island, past and present. Lithocraft Graphics, South Melbourne.
Iredale, T. 1910. Birds of Lord Howe and Norfolk Islands. Proc. Linn. Soc. New South Wales 35: 776–777.
                 
Zootaxa 3205: 26–40.
Parrots: a guide to parrots of the world. Pica Press, Robertsbridge.
  Feathers to brush: the Victorian bird artist, John Gerrard Keulemans, 1842–
1912. C. J. Coldewey, Epse.
Mol. Phyl. & Evol. 62: 296–305.
Latham, J. 1781. General synopsis of birds, vol. 1. Benjamin White, London.
Latham, J. 1824. General history of birds, vol. 9. Jacob and Johnson, Winchester.
Leguat, F. 1708. A new voyage to the East-Indies. R. Bonwicke, W. Freeman, T. Goodwin, F. Walthoe, M.
and ontogenetic perspectives. Orn. Monogr. 53: 1–654.
Luther, D. 1996. Die ausgestorbenen Vögel der Welt. Die neue Brehm-Bücherei 424. Fourth edn. Spektrum,
Martin, T. 2002. A guide to colour mutations & genetics in parrots
Mathews, G. M. 1928. Birds of Norfolk and Lord Howe Islands and the Australasian South Polar Quadrant
G. Witherby, London.
Mayr, E. 1941. Taxonomic notes on the birds of Lord Howe Island. Emu 40: 321–322.
Bird-Banding 48: 25–37.
Florida Orn. Soc. Spec. Publ. 2: 1–64.
 
trouvent représentées dans les collections du Muséum d’Histoire Naturelle. Pp. 190–252 in Centenaire de
la fondation du Muséum d’Histoire Naturelle, Paris. Muséum d’Histoire Naturelle, Paris.
Wild south: saving New Zealand’s endangered birds. Second edn. Random House,
New Zealand.
             
in Olson, S. (ed.) Avian paleontology
at the close of the 20th century. Proc. 4th Intern. Meeting Soc. Avian Paleontology and Evolution,
Washington, D.C., 4–7 June 1996. Smiths. Contrib. Paleobiol.
Myers, J. G. 1923. The present position of the endemic birds of New Zealand. NZ J. Sci. Technol. 6: 65–99.
Newton, A. 1872. On an undescribed bird from the island of Rodrigues. Ibis 2: 31–34.
Ibis 6: 281–289.
Julian P. Hume & Hein van Grouw 193 Bull. B.O.C. 2014 134(3)
© 2014 The Authors; Journal compilation © 2014 British Ornithologists’ Club
The island of Rodrigues. Standard Printing Establishment, Mauritius.
Wilson Bull. 85: 381–416.
Olson, S. L. 1986. Gallirallus sharpei
origin. Le Gerfaut 76: 263–269.
Nat. Hist. 94: 88–93.
Paynter, R. A. 1970. Subfamily Emberizinae. Pp. 3–214 in Paynter, R. A. (ed.) Check-list of birds of the world,
vol. 13. Mus. Comp. Zool., Cambridge, MA.
Pelzeln, A. 1873. On the birds in the collection at Vienna obtained from the Leverian Museum. Ibis 15: 14–54.
Peters, J. L. 1934. Check-list of birds of the world, vol. 2. Harvard Univ. Press, Cambridge, MA.
Phillip, A. 1789. The voyage of Governor Phillip to Botany Bay; with an account of the establishment of the colonies
of Port Jackson & Norfolk Island; compiled from authentic papers, which have been obtained from the several
Departments, to which are added the Journals of Lieuts. Shortland, Was, & Capt. Marshall, with an account of
their new discoveries. John Stockdale, London.
      
(Strigops habroptila), and the context of its conservation management. Notornis 53: 3–26.
Ripley, S. D. 1977. Rails of the world. D. R. Godine, Boston.
Rothschild, L. W. 1907. Extinct birds. Hutchinson, London.
Rowley, G. D. 1875. Ornithological miscellany
The birds of Africa, vol. 8. Christopher Helm, London.
Schifter, H. 1994. Historical specimens of parrots in the bird collection of the Museum of Natural History,
Vienna, Austria. Pp. 34–48 in III Intern. Loro Parque Parrot Convention, Loro Parque 14th to 17th September
1994. Puerto de la Cruz, Tenerife.
Schorger, A. W. 1955. The Passenger Pigeon: its natural history and extinction. Univ. of Wisconsin Press, Madison.
Sharpe, R. B. 1888. Catalogue of the birds in the British Museum, vol. 12. Trustees of the Brit. Mus. (Nat. Hist.),
Shufeldt, R. W. 1915. Anatomical and other notes on the Passenger Pigeon (Ectopistes migratorius) lately living
in the Cincinnati Zoölogical Gardens. Auk 32: 29–41.
Mysteries of planet Earth. An encyclopedia of the inexplicable. Carlton Books, London.
Slater, H. H. 1874. Notes on the birds of Rodrigues
Slater, H. H. 1879. Reports on the proceedings of the naturalists. 2. Report of Henry H. Slater, Esq., B.A. Phil.
Trans. Roy. Soc. Lond. 168: 294–295.
Conuropsis carolinensis). Pp. 1–36 in 
(eds.) The birds of North America no. 667. The Birds of North America Inc., Philadelphia, PA.
Nature 419: 265–266.
c.1725–26. Relation de l’isle Rodrigue. Anon. MS in Archives Nationales, Paris. Reprinted in part by
Newton, A. 1875. Proc. Zool. Soc. Lond. 1875: 39–43.
      Rails: a guide to the rails, crakes, gallinules and coots of the world. Pica Press,
White, J. 1790. Journal of a voyage to New South Wales
Macaw Ara tricolor. Bull. Brit. Orn. Cl. 133: 125–156.
The lost world of the Moa. Indiana Univ. Press, Bloomington.
Addresses: Bird Group, Dept. of Life Sciences, Natural History Museum, Akeman Street, Tring, Herts. HP23
... In fact, no individual factor can be singled out for either the extinction of some species or the great reduction in numbers of others; instead, a series of interconnected causes contributed to the relentless demise of Hawaiian avifauna. There is, however, widespread agreement that most of the blame should be ascribed to anthropogenic factors [7,9,10,13,17,18,21,22,31,[33][34][35][36][37][38][39]. Human-associated factors typically include: 1, modification of the natural habitat and destruction of original vegetation after initial settlement by Polynesians, followed by agricultural expansion; 2, bird catching by native Hawaiians (see below); 3, predation by animals introduced by settlers, such as Polynesian rats (Rattus exulans), pigs, chickens, dogs, and, later, mongoose and feral house cats; 4, imported avian parasites/diseases, such as avian malaria and pox; 5, avian competition with introduced birds; 6, shooting and trapping of rare species by collectors, largely in the 1880s-1920s; 7, a host of abiotic factors that include hurricanes, tsunamis, fires, floods, volcanic eruptions, and various recurrent environmental perturbations. ...
Full-text available
This research focuses on the historical demise of Hawaiian avifauna due to hunting by ancient Polynesians. Numerous documents, published since the early 1800s, were scrutinized and evaluated; these provided information on bird hunting and traditional Hawaiian practices. Hawaiians used birds as sources of feathers and food. Feathers were important symbols of power for Polynesians; in Hawai’i, feathers were more highly prized than other types of property. Feathers used for crafts were obtained from at least 24 bird species, however, the golden feathers of ‘ō‘ō and mamo birds made them primary targets for birdhunters; both birds became extinct by the late 1800s. Feathers were utilized for many items, including ‘ahu‘ula [cloaks], mahiole [war helmets], and kāhili [standards]. Most garments utilized a considerable number of feathers; a cloak for Kamehameha consumed the golden feathers of 80,000 mamo birds. Bird meat was an important food item for native Hawaiians. It is believed that most birds were killed after being plucked; historical sources mention ~30 bird species were consumed. The ‘ua‘u (Pterodroma sandwichensis), a currently endangered seabird, was ruthlessly hunted and avidly eaten. Its current geographical range is just a minute fraction of its former one; now, ‘ua‘us are largely restricted to inaccessible cliffs at Haleakalā Crater (Maui).
... NOTES: The White Gallinule, known from two skins collected in the late 18th century as well as a number of illustrations, was a large rallid with short wings and robust legs (McAllan et al. 2004, van Grouw andHume 2016). All blue normal-colored individuals were present, but a predominance of all white, and white and blue individuals, was a result of the "progressive greying" color aberration, a condition in which the cause can be inheritable and is based on a single mutation or is entirely age-related (Hume andvan Grouw 2014, van Grouw andHume 2016). The two surviving skins represent white, and white and blue individuals. ...
Lord Howe Island, situated 790 km northeast of Sydney in the Tasman Sea, was first encountered in February 1788, and one of the last islands discovered by humans. Apparently, Polynesians were unaware of the island. At the time of discovery, seabirds dominated the island, with a terrestrial avifauna comprising five endemic species and eight subspecies. The island was initially a whaling refurbishment station until establishment of a settlement in 1834, which resulted in the extinction of three endemic birds due to over-hunting. The accidental invasion of black rats in 1918 resulted in another wave of extinctions, with the loss of five passerines, followed by an endemic owl c. 1940. Museum skins represent all but one of these species, but almost no skeletal remains exist. Here, we present the results of a palaeontological survey that explored subfossil depositaries on Lord Howe Island, including analysis of hitherto unstudied Lord Howe bird subfossil collections held in Australian museums. Our discoveries include the first known skeletal elements of six extinct species, five recorded in context, and a particularly fossil-rich sand dune site on Lord Howe Island that contained hundreds of terrestrial bird subfossils.
... La hembra similar, pero gris arriba y mucho más opaca (Phelps & Meyer de Schauensse, 1994;Hilty, 2003 Las aberraciones cromáticas en el plumaje de las aves son fenómenos atribuidos a procesos de hibridación que se dan entre poblaciones, también a déficit nutricionales y mutaciones. Según sea el c a s o , e x i s t e n a l m e n o s o c h o v a r i a n t e s documentadas que alteran la coloración típica del plumaje en diferentes grupos orníticos, a saber: albinismo, encanecimiento progresivo, dilución, esquizocroismo, eumelanismo, feomelanismo y leucismo (Martin, 2002;van Grouw, 2006van Grouw, , 2013Hume & van Grouw, 2014). ...
Full-text available
En la avifauna venezolana se han documentado aberraciones cromáticas en un total de 23 especies incluidas en 18 familias, de ellas solo un caso está referido a la familia Trochilidae, especificado como encanecimiento progresivo en el Colibrí Pecho Canela (Glaucis hirsutus Gmelin 1788). En tal sentido en este trabajo se presenta el primer caso aberración cromática en el plumaje del Colibrí Anteado (Leucippus fallax Bourcier, 1843), especificado como dilución en un individuo observado en la isla de Coche, Estado Nueva Esparta.
... Brisson described a live captive bird in the 1750s, whereas the Cabinet du Roi specimen of unknown provenance that Martinet used a decade or so later may have deteriorated. The two specimens that survive are very faded, and one possesses irregular white feathering (Hume 2007, Hume & van Grouw 2014. Cheke (1983) also commented on possible post-mounting alterations to explain colour inconsistencies in the Zosterops plates compared to live birds. ...
... Atypically coloured birds have intrigued ecologists, ornithologists, birdwatchers and nature enthusiasts for a long time (M'Callum 1885, Sage 1963, Gross 1965, Ellegren et al. 1997, Møller & Mousseau 2001, McCardle 2012, van Grouw 2013, Hume & van Grouw 2014. In the past, aberrantly coloured individuals were sometimes described as new species (Brisson 1760, Vigors 1825, Dybowski 1916, van Grouw 2017). ...
Birds with plumage colour aberrations are of interest to both the general public and scientists. However, due to their rarity in nature, information on the presence of colour aberrations is rarely found in the peer‐reviewed literature. Exploration of public observations using modern information technologies such as Internet‐based search engines could facilitate cost‐effective and rapid broad‐scale collection of data on phenotypic aberrations in animals but may also be prone to the same problems as fieldwork, including systematic collection bias. We used both Google Images and asked birdwatchers and ornithologists, via naturalists’ forums and social media, to collate a unique data set of photographs of 936 aberrantly coloured birds of 74 species from Poland. Phylogenetically informed analyses, which included species both with and without reported colour aberrations, revealed that the number of colour aberrations was higher in species with larger populations in Poland, larger body size, and those associated with human settlements. Colour aberrations were also more often reported for species with a wider habitat breadth and which do not migrate over long distances. Habitat openness and diet type were not related to the number of colour aberrations across species in multivariate models. Our study emphasizes not only the power of novel sources to collect large data sets on relatively rare phenotypic aberrations in animals, but also the importance of vigilance when using data mined from public sources because the observed patterns may reflect collection bias rather than the nature of the studied phenomena.
... Its phylogenetic position is currently under debate, as to whether it is close to Coracopsis Wagler, 1832 (Kundu et al. 2012;Jackson et al. 2015) or close to the Psittacula eupatria clade (Podsiadlowski et al. 2017). Based on morphological and osteological traits, †Mascarinus is considered to be a close relative to the Psittacula parakeets (Hume 2007;Cheke & Hume 2008;Hume & van Grouw 2014). ...
Full-text available
The long-tailed parakeets of the genus Psittacula Cuvier, 1800 have thus far been regarded as a homogeneous and mono-phyletic group of parrots. We used nucleotide sequences of two genetic markers (mitochondrial CYTB, nuclear RAG-1) to reconstruct the phylogenetic relationships of Psittacula and closely related species. We found that the Asian genus Psit-tacula is apparently paraphyletic because two genera of short-tailed parrots, Psittinus Blyth, 1842 and Tanygnathus Wagler, 1832, cluster within Psittacula, as does †Mascarinus Lesson, 1830. To create monophyletic genera, we propose recognition of the following genera: Himalayapsitta Braun, 2016 for P. himalayana, P. finschii, P. roseata, and P. cyanocephala; Nicopsitta Braun, 2016 for P. columboides and P. calthrapae; Belocercus S. Müller, 1847 for P. longicauda; Psittacula Cuvier, 1800 for P. alexandri and P. derbiana; Palaeornis Vigors, 1825 for †P. wardi and P. eupatria; and Alexandrinus Braun, 2016 for P. krameri, †P. exsul, and P. (eques) echo. Additionally, Psittacula krameri and P. alexandri are paraphyletic species, which should be split to form monophyletic species.
... That melanistic birds were, especially in the past, mistaken for 'new species' is therefore understandable. Sometimes 'new species' were erected on the basis of a single specimen simply because it was differently coloured, like Sharpe's Rail (Hume & van Grouw 2014). Mostly, however, the confusion was based on melanistic forms that occurred, or still occur, quite commonly in the relevant species / populations. ...
Full-text available
The melanistic variety of Red-legged Partridge Alectoris rufa was described from a small population in western France around the 1850s. In this region, the Red-legged Partridge population as a whole was hunted, but melanistic individuals were targeted for both private and museum bird collections, and by 1865 the variety was extinct in western France. An extensive search for extant specimens documented 13 melanistic birds in six museums, and their details are presented here. Remarkably, some of these specimens were collected in areas elsewhere in France or even in other countries. After 1915, the allele for melanism appears to have been lost within the Red-legged Partridge population as a whole, and we discuss possible reasons for this.
Plumage color aberrations in wild birds are infrequently reported and often misinterpreted. We report a color aberration in the Gray-headed Swamphen (Porphyrio poliocephalus), a large colorful rail with a red bill and a frontal shield. Our report documents the observation of a diluted plumage in an adult Gray-headed Swamphen for the first time. The bird displayed an almost all-white plumage with normal-colored eyes that we assigned to a dilution process. We also summarize plumage color aberrations reported in 3 additional species of the genus Porphyrio.
Full-text available
Plumage anomalies in birds are more prevalent in India, where numerous studies have lately been published. We show an instance of brown plumage aberration in a Common House Crow (Corvus splendens) in this report. The plumage of the Common House Crow has been documented in a variety of morphs, the majority of which are examples of leucistic and other aberrations. However, the brown aberration observed by us was unusual and has been discussed.
The genus Agapornis consists of nine small African parrot species that are globally well known as pets, but are also found in their native habitat. Illegal trapping, poaching and habitat destruction are the main threats these birds face in the wild. In aviculture, Agapornis breeding is highly popular all across the globe. Birds are mainly selected based on their plumage colour variations but very little molecular research has been conducted on this topic. There are 30 known colour variations amongst the nine species and most of these are inherited as Mendelian traits. However, to date none of the genes or polymorphisms linked to these variations have been identified or verified. Due to unethical breeding practices, the need for the development of molecular tests such as identification verification tests or species identity tests is growing. Future research is paramount to ensure the conservation of wild populations as well as aiding breeders in improving breeding strategies.
Full-text available
In this paper 16 distinct, heritable colour aberrations (mutations) in the House Sparrow are described, based on specimens found in museum collections, records of individuals seen in the wild and from bird breeders keeping aberrant coloured sparrows in captivity. Based on the frequency found in the museum specimens Brown is the most common mutation in the House Sparrow, followed by Ino and Albino. Besides the mutations there is also a, presumably, non-heritable aberration called Progressive Greying described. Progressive Greying is in fact by far the most common colour aberration found in the species but was, in the past, always assigned as ‘Partial Albino’ without its real nature being understood. This paper will give some insight in the nature of Progressive Greying.
Full-text available
Extinction of the Passenger Pigeon (Ectopistes migratorius) is unique not only because of its rapid decline from being one of the world’s most abundant birds, but also because a considerable amount of information on the bird’s natural history is available in the records of early naturalists, travelers, and even the press. This information, a great part of which has been reviewed and summarized in Schorger’s (1955) monumental work, allows a detailed analysis of the factors that may have led to the species’ extinction and helps to reduce the number of plausible explanations. Beyond the intellectual challenge of such an analysis, an explanation of the causes of the Passenger Pigeon extinction may provide insights that are helpful in current efforts to prevent the extinction of many endangered species.
Full-text available
The last 20 years have seen a resurgence in systematic studies of parrots (Aves: Psittaciformes). Principally but not solely molecular in nature, this body of work has addressed the circumscription of higher level groupings within the Psittaciformes and relationships among them. Stability has now emerged on many formerly contentious matters at these levels. Accordingly, we consider it appropriate to underpin further work on parrot biology with a freshly revised classification at the taxonomic ranks spanned by family-group nomenclature, i.e., between superfamily and tribe. In light of the body of recent work, we advocate a framework of three superfamilies among parrots (Strigopoidea, Cacatuoidea and Psittacoidea) within which Linnaean taxonomy can accommodate present phylogenetic understanding by employing groupings at the ranks of family, subfamily and tribe. Just as importantly, we have addressed numerous issues of nomenclature towards stabilising the family-group names of parrots. We erect two new subfamily names, Coracopseinae Joseph, Toon, Schirtzinger, Wright & Schodde, subfam. nov. and Psittacellinae Joseph, Toon, Schirtzinger, Wright & Schodde, subfam. nov. We stress that rankings we have applied reflect the state of understanding of parrot phylogeny and how it can be summarized in a Linnaean system; comparisons with rankings in other groups are likely not appropriate nor relevant.
This volume consist of eight main sections. Initially origins and evolutionary relationships are examined, followed by a brief section on the classification of the parrots. Next a section reviews the natural history of the parrots, briefly covering: general behaviour; distribution; habitat; movements; social behaviour; diet; breeding; and nocturnal species. Conservation status ics covered next. The main threats to parrots are then outlined and discussed: habitat loss; live bird trade; introduced species; persecution and hunting; and storms'climatic change. A brief section then looks at captive breeding. The mian body of the book is taken up with colour plates and a systematic section. The systematic section contains the following information, for each species: identification notes; voice; distribution and status (including distribution maps); ecology; description; sex/age; measurements; geographical variation; and references.
Since the last review of kakapo biology, published 50 years ago, much has been learnt as a result of the transfer of all known individuals to offshore islands, and their intensive management to increase adult survival and productivity. This review summarises information on a diversity of topics, including taxonomy, plumage, moult, mass, anatomy, physiology, reasons for decline in distribution, present numbers and status, sex ratio, habitat, home range, foraging activities, diet, voice, breeding biology, nesting success, sexual maturity, and adult survival. In addition, those kakapo attributes that compromise its long-term survival in present-day New Zealand are discussed, along with management practises developed to overcome these problems.