Hybrid origin of the imperial pheasant Lophura imperialis ...viduals (Delacour, 1977), but these died en route to France and their identity was never veriﬁed. In 1938, C. Cordier,

Biological Journal of the Linnean Society, 2003, 80, 573–600. With 3 figures

© 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 573–600 573

Blackwell Science, LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066The Linnean Society of London, 2003? 200380?573600Original Article

IMPERIAL PHEASANT ORIGINA. HENNACHE

ET AL.

*Corresponding author. E-mail: [email protected]

Hybrid origin of the imperial pheasant Lophura imperialis (Delacour and Jabouille, 1924) demonstrated by morphology, hybrid experiments, and DNA analyses

ALAIN HENNACHE1*, PAMELA RASMUSSEN2, VITTORIO LUCCHINI3, SILVIA RIMONDI3 and ETTORE RANDI3

1Muséum National d’Histoire Naturelle, FRE 2632 Conservation des espèces et suivi des populations, Parc Zoologique de Clères, 76690 Clères, France2 Michigan State University Museum and Department of Zoology, East Lansing, MI 48824–1045 USA3Istituto Nazionale per la Fauna Selvatica, Via Cà Fornacetta 9, 40064 Ozzano dell’Emilia (BO), Italy

Received 20 September 2002; accepted for publication 22 May 2003

The imperial pheasant Lophura imperialis was described in 1924 from a captive pair that was obtained in Viet-nam, and that became the sole founders of a captive line in France. Always considered a highly endangered andmysterious species, and despite concerted searches, L. imperialis was not found again in the wild until one wastrapped in 1990, and the captive population gradually died out. Its status as a distinct species was unquestioneduntil the late 1990s when the possibility of a hybrid origin was raised. To elucidate the taxonomic status ofL. imperialis, we studied all the existing museum specimens, carried out captive hybridization experiments, andanalysed mitochondrial DNA and microsatellites. All these lines of evidence demonstrate congruently and conclu-sively that L. imperialis is an occasional hybrid between silver pheasant L. nycthemera and Edwards’s pheasantL. edwardsi, with the 1990 bird probably being a hybrid between L. nycthemera and Vietnamese pheasantL. hatinhensis. Thus L. imperialis has no taxonomic standing and should be removed from lists of species of con-servation concern. However, hybridization with L. nycthemera may pose a further threat to the survival in the wildof the endangered L. edwardsi and L. hatinhensis. © 2003 The Linnean Society of London, Biological Journal ofthe Linnean Society, 2003, 80, 573–600.

ADDITIONAL KEYWORDS: conservation – Edwards’s pheasant – fragmented habitat – hybridization –microsatellites – morphological analyses – mtDNA sequencing – silver pheasant.

INTRODUCTION

The discovery of the imperial pheasant Lophura impe-rialis was perhaps the high point of J. Delacour’s expe-ditions to central Annam (Delacour & Jabouille, 1925).In 1923, he had obtained a single live pair of a here-tofore unknown captive pheasant from missionaries,who said the birds had been obtained on the southernboundary of the Quang Binh province, northern QuangTri, Vietnam (Delacour & Jabouille, 1925). The specieswas named in honour of Khaï Dinh, Emperor of Annam(Delacour & Jabouille, 1924). The pair was sent to

Delacour’s estate in France in 1924 and the followingyear began producing young (Delacour, 1925). How-ever, further attempts to locate L. imperialis in thewild were unproductive. Father Meaunier, a Frenchmissionary, was said to have obtained a few more indi-viduals (Delacour, 1977), but these died en route toFrance and their identity was never verified. In 1938,C. Cordier, a famous collector of live birds, attemptedand failed to procure L. imperialis (Delacour, 1977). Inthe late 1930s B. Björkegren spent two weeks search-ing for it without result (Eames & Ericson, 1996). Formany years after that time, until recently, much ofVietnam was inaccessible to ornithologists, and thelack of further reports of L. imperialis did not seem acause for great concern.

574 A. HENNACHE ET AL.

© 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 573–600

Although L. imperialis has been repeatedly said tooccur in central Laos (Delacour & Jabouille, 1931;David-Beaulieu, 1949; King, Dickinson & Woodcock,1975), no specimens exist from that country, and itsputative occurrence was based only on vague descrip-tions by local hunters. Recent field surveys in Laoshave resulted in no further records (McGowan & Gar-son, 1995), and the past or current occurrence ofL. imperialis in Laos is highly doubtful. Lophuraimperialis has also been included on the Chinese list(Xu et al., 1996), presumably in error. The species istherefore definitely known only from Vietnam, whereit must be an extremely rare endemic, and as such hasbeen the subject of great conservation concern (Collar,Crosby & Stattersfield, 1994; McGowan & Garson,1995; Fuller & Garson, 2000). Despite the recent con-servation concern and field efforts to find a wild pop-ulation, the only other records are of two liveimmature males, one in 1990 (Robson et al., 1993;Eames, Lambert & Nguyen, 1994) and the second in2000 (BirdLife International, 2000). However, both ofthese recently trapped individuals differ distinctlyfrom other specimens in some features (Appendix; VoQuy, unpubl. data), and no populations have beenlocated in the areas where these individuals werefound.

Despite the fact that Delacour’s original pair hadfounded a captive population of L. imperialis, andalthough some of their progeny had been distributedto other aviaries, they failed to thrive and nearly diedout. By 1959 the only known birds remained in Antw-erp Zoo, where these few were hybridized with thesilver pheasant L. nycthemera and with Edwards’spheasant L. edwardsi in an attempt to recreate newstock (Carpentier et al., 1975). Although some result-ant hybrid individuals closely resembled the descrip-tion of the original pair, others looked more like theparental species, and the stock was highly variable.The population’s genetic purity had obviously beenseriously compromised, and attempts to sustain the‘species’ in captivity failed, later pairings resultingonly in male progeny (Carpentier et al., 1975), a seem-ing catastrophe for the survival of the species.

Delacour wrote that L. imperialis occurs only inthe dense forests of the mountains of the interior ofthe Province of Dong Hoi, Vietnam (Delacour &Jabouille, 1931), but the source of his information isuncertain. In fact, this conflicts with both recentrecords, which were located in lowland secondary for-est. On 28 February 1990, a wild-trapped bird wasobtained in lowland forest 12 km west of Cat Bin,approximately 200 km north of Dong Hoi in the samesnare line as two adult male Vietnamese pheasantsL. hatinhensis (Robson et al., 1993), an endemicspecies closely related to L. imperialis. On 27February 2000, another individual considered to be

L. imperialis was obtained in Da Krong district,Quang Tri province, within the historical range ofL. edwardsi, yet another endemic and closely relatedspecies of secondary lowland forest (Truong Van La,pers. comm.; Dang Gia Tung, pers. comm.; HuynhVan Keo, 1999). Thus, L. imperialis is evidently sym-patric with two other congeneric pheasant species,all endemic to a small area in Annam and sharingthe same habitat, a remarkable situation, especiallyamong large-bodied birds. Although wild populationsare known for both L. hatinhensis and L. edwardsi,all attempts to find more L. imperialis other thanthose detailed above have failed. Thus, L. imperialiswas considered a critically endangered Vietnameseendemic species with no known wild population andnone in captivity (McGowan & Garson, 1995; Garson,1998; BirdLife International, 2000; Fuller & Garson,2000). More recently it was treated as ‘data defi-cient’ (BirdLife, 2001) on the basis of evidence pre-sented herein.

In 1997, K. Wood (pers. comm.) suggested that,rather than being an individual species, L. imperialismight actually be a hybrid between L. edwardsi andL. diardi, an idea that would explain its extremerarity and the inability of field workers to locate awild population. This led one of the authors (P.R.)to examine the possibility of a hybrid origin ofL. imperialis. She found that female L. imperialisare intermediate between female L. edwardsi andL. nycthemera of Vietnamese races, and lackcharacters indicative of L. diardi, while the maleL. imperialis phenotype is typical of hybrids betweena glossy black and a more elaborately plumaged spe-cies of Lophura (Rasmussen, 1998, 1999). She alsofound that plumage features and anomalies ofL. imperialis specimens are all present in the hybrid-ized Antwerp Zoo birds, and that the L. imperialisphenotype shows no novel features or characters notexpected from a cross between L. edwardsi andL. nycthemera (Rasmussen, 1998, 1999). After thesemuseum studies indicated the strong likelihood thatL. imperialis was a hybrid between L. nycthemeraand L. edwardsi, another of the authors (A.H.)carried out controlled hybridization experimentsin captivity for the purposes of confirmation.Simultaneously, a third author of this study (E.R.)and colleagues were exploring the molecular phyloge-netics of the three endemic Vietnamese lowlandLophura species with the objective of contributinginformation vital to the process of defining a pro-gramme for their global conservation (Hennache,Randi & Lucchini, 1999). The purpose of this paperis to present the results of three different lines ofinquiry into the origins and status of L. imperialis:morphological studies, hybridization experiments,and genetic analyses.

IMPERIAL PHEASANT ORIGIN 575


MATERIAL AND METHODS

The original description of L. imperialis was basedsolely on the living captive pair that Delacourbrought to France from Vietnam (Delacour &Jabouille, 1924). The female holotype died in Sep-tember 1927 and was preserved as Museum Nationald’Histoire Naturelle (MNHN) no. 1928–1117. Themale holotype, which was still living at Clères in1939, was lost along with its progeny during WorldWar II and thus no male holotype specimen exists,nor do there appear to be any photographs of thisindividual. Thus we were able to make direct com-parisons with the female holotype, but for males wewere limited to comparisons with the rather vaguetype description and the first-generation maleprogeny.

Several of the progeny of the original pair have beenpreserved at the MNHN and at the Natural HistoryMuseum (BMNH), Tring, UK (see Appendix for a com-plete listing). Delacour also sent live descendants toAmerican breeders, and some of these skins are pre-served in US museums; however, as the lineage ofmost of these is unclear and interspecific hybridiza-tion was taking place from early on, these have beenused advisedly. Only those progeny that are labelledas definitely resulting from the original pairing havebeen considered true L. imperialis.

Photographs of the 1990 wild-trapped immaturemale, both before and after its death, were sent by J.Eames, and the specimen was examined by A.H. in1997 and 1999. A description and photographs of theliving immature male wild-trapped in 2000 were sentto us by Prof. Vo Quy.

MORPHOLOGICAL ANALYSES

For all known museum specimens of L. imperialis(see Appendix), numerous close-up photographs weretaken by P.R. to document external morphology. Mea-surements (to the nearest mm) were taken by P.R.when possible for each fully grown specimen of manycharacters, of which the following were measurableon most specimens and so were used for this study:bill length from nares; bill height from nares; uppermandible width from nares; central length of lowermandible; proximal width of lower mandible fromfeathers; crest length; flattened wing length; outerprimary length; outer primary width; tarsus length;tarsus proximal depth; tarsus proximal width; tarsusminimum width; tarsus minimum depth; tarsus dis-tal width; tail length; outer rectrix width; inner rec-trix width; tail graduation (distance between tip ofinner rectrix and tip of outer rectrix); greatest widthof longest uppertail coverts. Many of the L. imperialisspecimens were damaged in some way related to

their captivity, often with overgrown bills, very wornor otherwise abnormal claws, and broken tails andwingtips, making it impossible to measure thesecharacters. Photographs and measurements werealso taken of several frozen and one mounted speci-men(s) of the hybrids produced at the Antwerp Zoo.For comparison, photographs and the same measure-ments were taken of a series of Vietnamese races ofL. nycthemera and of L. edwardsi. In addition, photo-graphs were taken of a series of well-documentedhybrid Lophura specimens in the BMNH. No speci-mens of L. hatinhensis were available for measure-ment, but many photographs were taken of a captivepair in the Antwerp Zoo, and of their immature off-spring. Photographs and measurements of the cap-tive hybrids at Clères (see below) were taken by P.R.when the birds were c. 1 year old (full-sized). Mea-surements taken by J. Eames of the 1990 specimenand considered to be comparable with those taken byP.R. were included. Univariate statistics and princi-pal components analyses (PCA) were done using Sys-tat 5.0. Characters used in PCA were: culmen length,crest length, wing length, tarsus length, and taillength. These characters were chosen to allow theinclusion of the maximum number of specimens inthe analysis. Because specimens at different muse-ums could not readily be directly compared with oneanother, P.R. used her photographs of each individualto construct the Appendix. In several cases, someindividual characters were not visible or discerniblein the photographs, so these cells were labelled ‘notrecorded’ in the Appendix.

CAPTIVE-BREEDING EXPERIMENTS

In 1998, A.H. set up a hybridization experiment atthe Zoological Park of Clères, France (Delacour’sformer estate) with the aim of assessing the initialresults from museum studies (summarized in Ras-mussen, 1998). A captive-reared male L. nycthemeraberliozi was mated using artificial insemination witha captive-reared female L. edwardsi, and five chickswere produced from this cross, of which four weremales and one was a female. One of the males diedat 1 year of age when it was killed by two of itssiblings.

Each of the resultant hybrids was studied fromchick stage to adulthood, and photographs were takenat different stages of growth. Their phenotypes at2 years of age are described herein by A.H. and arecompared with those of Delacour’s description andwith the existing specimens of L. imperialis from theoriginal pair and their progeny. The hybrids’ pheno-types at about 1 year of age were also described fromphotographs taken by P.R., and these data areincluded in the Appendix.



DNA ANALYSES

DNA extraction, amplification and sequencingTotal DNA was extracted from 95% ethanol-preservedtissue or feather root samples, using proceduresdescribed by Randi & Lucchini (1998). The 5¢ domainof the mitochondrial DNA control region (mtDNA CR)was PCR-amplified and sequenced as previouslydescribed (Randi & Lucchini, 1998; Randi et al., 2001).CR sequences were obtained from the L. imperialissamples listed in Table 1, and from L. edwardsi(N = 6), L. hatinhensis (N = 15), and two closelyrelated outgroups L. nycthemera (N = 4), andL. swinhoei (Swinhoe’s pheasant, N = 2). The CRsequences were aligned using CLUSTAL X with thedefault options (Thompson et al., 1997). Phylogeneticanalyses were performed using the software PAUP*(Swofford, 1998), by: (1) a maximum-parsimony pro-cedure (Swofford, 1998), excluding all uninformativenucleotide positions, with unordered and equallyweighted characters; (2) the neighbour-joining algo-rithm (Saitou & Nei, 1987), with Tamura & Nei’s(1993) DNA distances (TN93). Robustness of the phy-logenies was assessed by bootstrap percentages (BP;Felsenstein, 1985), with 1000 random resamplings

with replacement. Details on phylogenetic analysesare given in Randi et al. (2001).

Microsatellite analysesAll samples were genotyped by PCR amplification ofnine microsatellites that were originally isolated atWageningen University from the chicken (Gallus gal-lus) genome. The primers used were: M1 = MCW160;M2 = MCW236; M3 = MCW292; M4 = MCW239;M5 = MCW297; M6 = MCW254; M7 = MCW152;M8 = MCW262; M9 = MCW199. Primer sequencesand information on MCW markers can be retrievedat http://www.zod.wau.nl/abg/index.html. Allelic vari-ability was determined in DNA extracted from theL. imperialis samples listed in Table 1, and fromL. edwardsi (N = 10), L. hatinhensis (N = 5),L. leucomelana (N = 2), L. nycthemera (N = 10) andL. swinhoei (N = 3). Each PCR amplification was per-formed in a total volume of 9 mL using 30–50 ngDNA, in a Perkin-Elmer 9600 Gene-Amp cyclerwith the following cycle: 94∞C ¥ 2 min; 30 cycles at94∞C ¥ 30 s, 45–65∞C ¥ 30 s (depending on theoptimal primer annealing temperature), and72∞C ¥ 1 min; 72∞C ¥ 10 min. Genotypes were deter-

Table 1. Lophura imperialis sampled for the genetic analyses

SampleID Sample Origin

Captive/wild Sex M/F Reference Comments

LIM8 Feather Quang TriProvince

W M Sample fromJ. Eames

Trapped on 27 February 2000

LIM9 Feather Quang TriProvince

W M Sample fromVo Quy

Trapped on 27 February 2000(same individual as LIM8)

LIM11 Skin and toepad fragments

MNHN C (F1) M 1936–1548 Died at Clères 15/04/1936


MNHN C (F1) M 1931–144 Born 1930; died at Clères 15 March 1931


MNHN C M 1934–1388 Died at Clères


MNHN –Dong Hoi

W F Syntype1928–1117

Died at Clères September 1927


MNHN C F 1934–1389 Died at Clères


MNHN C Young F 1927–2376 Donation from Delacour 19 July 1927

LIM17 Toe padfragment

BMNH C M 1931.5.11.5 Died at Clères 01 May 1931


BMNH C F 1948.70.1 Died at Clères


BMNH C Juvenile 1926.9.9.3 Died at Clères 25 August 1926


BMNH C Fledgling 1930.6.12.2. 5 weeks old; died in captivity 20 June 1930

http://www.zod.wau.nl/abg/index.html



Tab

le 2

a. U

niv

aria

te s

tati

stic

s fo

r al

l kn

own

fu

lly

grow

n L

oph

ura

im

peri

alis

mal

e sp

ecim

ens

and

hyb

rids

, an

d co

mpa

rati

ve s

erie

s of

L. e

dw

ard

si a

nd

Vie

tnam

ese

race

s of

L. n

ycth

emer

a

Var

iabl

e

L. i

mpe

rial

is

L. e

dw

ard

siL

. nyc

them

era

All

Ori

gin

al s

erie

sA

ntw

erp

hyb

rids

Cat

Bin

Clè

res

hyb

rids

Bil

l le

ngt

h18

.2 ±

1.5

(15

)17

.1 ±

1.9

(4)

17.9

± 1

.2 (

7)19

.0 (

1)19

.5 ±

0.6

(4)

17.7

± 1

.8 (

12)

20.7

± 1

.5 (

14)

Bil

l h

eigh

t11

.2 ±

0.9

(13

)10

.4 ±

0.5

(5)

11.7

± 0

.5 (

9)–

–9.

9 ±

1.0

(16)

12.4

± 0

.5 (

11)

Upp

er m

andi

ble

wid

th10

.1 ±

0.9

(15

)9.

9 ±

0.7

(6)

10.4

± 0

.9 (

10)

––

9.3

± 0.

6 (2

4)11

.1 ±

0.6

(14

)L

ower

man

dibl

e ce

ntr

al l

engt

h11

.7 ±

1.3

(12

)11

.4 ±

1.1

(5)

11.9

± 1

.4 (

8)–

–11

.7 ±

0.7

(24

)13

.4 ±

0.5

(13

)L

ower

man

dibl

e pr

oxim

al w

idth

12.3

± 0

.9 (

10)

12.5

± 0

.9 (

5)12

.2 ±

0.8

(6)

––

11.8

± 0

.7 (

24)

14.1

± 0

.6 (

14)

Cre

st l

engt

h41

.1 ±

12.

6 (1

9)39

.6 ±

8.2

(6)

42.5

± 1

3.0

(9)

70.0

(1)

33.5

± 4

.3 (

4)28

.6 ±

2.9

(23

)63

.3 ±

6.7

(14

)W

ing

len

gth

(fl

atte

ned

)23

2.9

± 12

.7 (

19)

232.

8 ±

7.9

(5)

229.

5 ±

14.5

(10

)25

2.0

(1)

237.

7 ±

8.3

(4)

227.

9 ±

8.1

(22)

251.

7 ±

1.0

(15)

Ou

ter

prim

ary

len

gth

120.

1 ±

12.0

(8)

125.

7 ±

4.3

(4)

114.

5 ±

15.3

(4)

––

117.

2 ±

9.1

(16)

134.

3 ±

8.4

(9)

Ou

ter

prim

ary

wid

th20

.0 ±

0.9

(4)

20.0

± 0

.9 (

4)–

––

17.1

± 1

.5 (

12)

19.0

± 2

.5 (

12)

Tar

sus

len

gth

81.7

± 1

7.9

(19)

81.5

± 6

.1 (

6)78

.4 ±

6.6

(9)

103.

0 (1

)83

.2 ±

3.5

(4)

75.7

± 4

.8 (

24)

87.2

± 5

.2 (

14)

Tar

sus

prox

imal

dep

th12

.6 ±

0.9

(7)

12.7

± 1

.0 (

6)12

.2 (

1)–

–11

.5 ±

0.8

(24

)13

.2 ±

1.2

(13

)T

arsu

s pr

oxim

al w

idth

13.2

± 1

.0 (

10)

12.5

± 0

.8 (

5)12

.5 (

1)–

14.1

± 0

.3 (

4)11

.6 ±

0.7

(22

)13

.7 ±

0.9

(14

)T

arsu

s m

inim

um

wid

th6.

0 ±

0.3

(10)

5.8

± 0.

4 (6

)–

–6.

2 ±

0.2

(4)

5.5

± 0.

5 (2

4)6.

3 ±

0.5

(14)

Tar

sus

min

imu

m d

epth

9.8

± 0.

5 (6

)9.

8 ±

0.5

(5)

9.7

(1)

––

8.8

± 0.

6 (2

2)10

.0 ±

0.8

(13

)T

arsu

s di

stal

wid

th12

.8 ±

1.1

(10

)12

.3 ±

1.0

(5)

13.0

(1)

–13

.5 ±

1.0

(4)

11.7

± 0

.6 (

23)

13.4

± 1

.1 (

13)

Tai

l le

ngt

h25

8.2

± 38

.3 (

14)

256.

0 ±

29.5

(6)

254.

3 ±

50.1

(6)

303.

0 (1

)25

0.0

(1)

207.

5 ±

17.6

(18

)30

5.1

± 56

.6 (

14)

Ou

ter

rect

rix

wid

th28

.4 ±

5.2

(14

)30

.0 ±

6.5

(5)

26.7

± 4

.3 (

7)–

27.5

± 3

.5 (

2)27

.2 ±

2.1

(20

)31

.1 ±

3.3

(14

)In

ner

rec

trix

wid

th43

.0 ±

5.5

(14

)46

.5 ±

2.8

(5)

40.8

± 6

.6 (

7)–

42.0

± 1

.4 (

2)38

.9 ±

3.8

(21

)54

.0 ±

7.3

(14

)T

ail

grad

uat

ion

143.

3 ±

35.5

(11

)15

4.0

± 20

.5 (

5)14

2.6

± 44

.2 (

5)–

–11

2.0

± 11

.2 (

19)

206.

8 ±

54.8

(14

)U

pper

tail

cov

erts

gre

ates

t w

idth

40.6

± 5

.7 (

10)

42.7

± 6

.0 (

3)39

.2 ±

5.5

(8)

––

34.4

± 3

.4 (

15)

44.0

± 4

.1 (

9)



Tab

le 2

b.

Un

ivar

iate

sta

tist

ics

for

all k

now

n fu

lly

grow

n L

oph

ura

im

peri

alis

fem

ale

spec

imen

s an

d h

ybri

ds, a

nd

com

para

tive

ser

ies

of L

. ed

war

dsi

an

d V

ietn

ames

era

ces

of L

. nyc

them

era

Var

iabl

e

L. i

mpe

rial

is

L. e

dw

ard

siL

. nyc

them

era

All

Ori

gin

al s

erie

sA

ntw

erp

hyb

rids

Clè

res

hyb

rids

Bil

l le

ngt

h16

.8 ±

0.9

(6)

16.9

± 0

.9 (

4)15

.9 (

1)18

.0 (

1)17

.9 ±

1.3

(7)

19.0

± 0

.7 (

10)

Bil

l h

eigh

t9.

7 ±

0.7

(6)

9.5

± 0.

5 (5

)10

.8 (

1)–

9.5

± 0.

6 (4

)11

.2 ±

0.6

(7)

Upp

er m

andi

ble

wid

th9.

3 ±

0.4

(6)

9.2

± 0.

2 (5

)10

.9 (

1)–

9.1

± 0.

8 (7

)10

.1 ±

0.4

(10

)L

ower

man

dibl

e ce

ntr

al l

engt

h11

.3 ±

0.5

(6)

11.3

± 0

.6 (

5)11

.6 (

1)–

11.0

± 0

.1 (

5)12

.2 ±

0.8

(9)

Low

er m

andi

ble

prox

imal

wid

th11

.2 ±

1.1

(6)

11.2

± 1

.2 (

5)11

.3 (

1)–

10.7

± 0

.5 (

6)12

.9 ±

1.1

(11

)C

rest

len

gth

23.1

± 4

.7 (

7)21

.7 ±

4.0

(5)

30.7

(1)

23.0

(1)

17.2

± 2

.9 (

8)44

.6 ±

11.

8 (1

1)W

ing

len

gth

(fl

atte

ned

)22

4 ±

8.3

(4)

229.

5 ±

0.7

(2)

212.

0 (1

)22

5.0

(1)

213.

9 ±

10.7

(8)

233.

1 ±

9.6

(10)

Ou

ter

prim

ary

len

gth

121.

7 ±

7.6

(3)

126.

0 ±

1.4

(2)

113.

0 (1

)–

105.

5 ±

8.3

(4)

123.

3 ±

4.8

(9)

Ou

ter

prim

ary

wid

th18

.3 ±

0.3

(3)

18.3

± 0

.3 (

3)–

–16

.1 ±

1.5

(7)

18.9

± 1

.4 (

10)

Tar

sus

len

gth

72.7

± 4

.5 (

6)71

.2 ±

3.3

(4)

80.3

(1)

71.0

(1)

66.2

± 4

.1 (

9)77

.3 ±

3.5

(11

)T

arsu

s pr

oxim

al d

epth

11.3

± 0

.4 (

4)11

.3 ±

0.4

(4)

––

10.2

± 0

.6 (

8)11

.4 ±

0.8

(10

)T

arsu

s pr

oxim

al w

idth

11.4

± 0

.8 (

5)11

.0 ±

0.3

(4)

–12

.7 (

1)10

.7 ±

0.8

(8)

12.1

± 0

.6 (

11)

Tar

sus

min

imu

m w

idth

5.4

± 0.

3 (5

)5.

3 ±

0.2

(4)

–5.

7 (1

)4.

9 ±

0.3

(9)

5.6

± 0.

3 (1

1)T

arsu

s m

inim

um

dep

th7.

9 ±

0.5

(4)

7.9

± 0.

5 (4

)–

–7.

7 ±

0.5

(7)

8.8

± 0.

6 (1

1)T

arsu

s di

stal

wid

th11

.3 ±

0.5

(5)

11.1

± 0

.3 (

4)–

12.0

(1)

10.8

± 1

.0 (

8)12

.3 ±

0.4

(11

)T

ail

len

gth

194.

2 ±

14.9

(4)

190.

7 ±

16.0

(3)

205.

0 (1

)–

182.

5 ±

7.6

(8)

226.

1 ±

23.7

(10

)O

ute

r re

ctri

x w

idth

24.2

± 2

.2 (

4)24

.2 ±

2.2

(4)

––

24.3

± 2

.8 (

7)28

.1 ±

4.6

(9)

Inn

er r

ectr

ix w

idth

38.5

± 1

.9 (

4)38

.0 ±

2.0

(3)

40.0

(1)

–35

.1 ±

3.9

(8)

41.8

± 3

.1 (

10)

Tai

l gr

adu

atio

n97

.3 ±

4.7

(3)

95.5

± 4

.9 (

2)10

1.0

(1)

–90

.7 ±

7.2

(5)

132.

4 ±

26.1

(8)

Upp

erta

il c

over

ts g

reat

est

wid

th32

.5 ±

3.3

(4)

31.3

± 2

.9 (

3)36

.0 (

1)-

32.0

± 2

.8 (

4)37

.2 ±

3.8

(10

)



mined by analysing the PCR products with an ABI373 A automated sequencer, and the software Gene-can 3.7 and Genotyper 2.1.

RESULTS AND DISCUSSION

MENSURAL ANALYSES

For most measurements, adult L. imperialis speci-mens (including known hybrids and the 1990 birdfrom near Cat Bin) were intermediate in size betweenadult L. nycthemera and L. edwardsi specimens of thesame sex (Table 2). Crest length was highly variableamong adult L. imperialis specimens, including thoseof the original series and even more so among knownhybrids. Outer primary length did not conform well tothe pattern of intermediacy among L. imperialis spec-imens, but the sample size for this character was verysmall. Except for the Cat Bin bird (see following), nospecimen of L. imperialis showed mensural charactersthat would not be expected of hybrids betweenL. nycthemera and L. edwardsi.

The Cat Bin bird had an exceptionally long crest(Table 2), longer even than the mean for adult maleL. nycthemera, and much longer than any otherL. imperialis; this feature was clearly shown in thephotographs. The Cat Bin bird also had long wingsand tail, both similar to the mean for adult maleL. nycthemera and much longer than any otherL. imperialis. Its tarsus length was much longer thanthat of any specimen of L. nycthemera; differences inmeasurement technique probably explain this lastanomaly (the Cat Bin specimen was measured by J.Eames). We were unable to measure any specimens ofL. hatinhensis, but this species appeared (to P.R.) to besomewhat lankier and less chicken-like in shape com-pared with L. edwardsi. If this impression is borne outby measurements, it might explain why the Cat BinL. imperialis was closer to L. nycthemera in some of itsmeasurements, although the possibility of its being abackcross to L. nycthemera might provide an alterna-tive or additional explanation.

PCA (Table 3, Fig. 1) confirmed the intermediacy ofL. imperialis (except for the Cat Bin bird). In the PCAmodel used, by far the greatest amount of variancewas explained on PC-I, a very strong size axis onwhich all characters had strongly positive loadings.PC-II showed a relatively weak contrast between cul-men length and crest length. Other axes were evenweaker and clearly not significant. Thus, for charac-ters included in the PCA, overall size was the mainquantitative difference between L. edwardsi andL. nycthemera, and loadings for L. imperialis speci-mens (including known hybrids) were largely interme-diate between these two species, with scores for a fewindividuals overlapping with L. edwardsi, and the CatBin bird overlapping with L. nycthemera.

PLUMAGE ANALYSES

All specimens of L. imperialis exhibited plumage char-acters that were to some degree intermediate betweenL. edwardsi and Vietnamese races of L. nycthemera(Appendix). Adult male L. imperialis of the originalseries (and most of the known hybrids) were generallyglossy blue-black overall with moderately scallopedrear upperparts and greenish-tinged wing coverts;their crests varied from entirely blue-black to slightlyor extensively mixed or barred with white, and whilecrest length varied greatly among adult maleL. imperialis, most specimens had a wispy, thin crest.They were always much less glossy and less blue com-pared with adult male L. edwardsi, with much lessprominently scalloped rear upperparts, and much lessgreen-tinged wing coverts. Their crests never con-tained as much white as adult male L. edwardsi, norwere they as short and bushy. Compared with adultmale L. nycthemera, adult male L. imperialis of theoriginal series totally lacked any trace of the black-and-white patterning of the upperparts, highly vari-able in but clearly present in all races of the formerspecies. Below, adult male L. imperialis showedglossy black underparts comparable to those ofL. nycthemera, but their side feathers were much lesslanceolate than in adult male L. nycthemera, andmore so than in L. edwardsi (Appendix). The crest ofadult male L. imperialis was much shorter and muchless full than that of L. nycthemera, and unlike the lat-ter species was not always entirely black. The tail ofadult male L. imperialis was intermediate in lengthand degree of curvature between L. edwardsi andL. nycthemera, and while it was usually solid blacklike L. edwardsi, one specimen (CAS 60752) had afinely black-and-white barred tail, and the descriptionof the original male (not preserved) indicates that ithad faint chestnut spotting on the back and centralrectrices (see below).

Table 3. Results from principal components analysis(graphed in Fig. 1) of adult Lophura imperialis,L. edwardsi, and Vietnamese races of L. nycthemera

Character PC-I PC-II

Culmen l from nares 0.70 -0.68Crest l 0.79 0.37Wing l (flattened) 0.89 0.01Tarsus l 0.84 0.05Tail l 0.87 0.16Eigenvalues 3.37 0.63Percent total variance explained 67.46 12.62

The sexes were included in the same analysis but weregraphed separately on Fig. 1 for clarity.



Adult female L. imperialis of the original series (andthe few known hybrids) showed variability and inter-mediacy in each of the several plumage charactersthat distinguished adult female L. edwardsi fromadult females of Vietnamese races of L. nycthemera(Appendix). These included the colour and degree ofshaft-streaking of head and body plumage (darkerand warmer-toned, with very weak shaft-streakingin L. edwardsi; paler and greyer, with more shaft-streaking in L. nycthemera), the intermediate length

of the crest, and the shape and distribution of blackishand chestnut in the tail. No plumage characters wereobserved among adult female L. imperialis that werenot intermediate between the two putative parentalspecies.

Among other age/sex classes of L. imperialis, thesame situation was apparent (Appendix): each age/sexclass of L. imperialis showed only plumage charactersintermediate between those of L. edwardsi andL. nycthemera, or in a few cases they were shared withone of the latter two. Briefly, subadult maleL. imperialis showed a muted L. nycthemera-type pat-tern in the central rectrices, whether the groundcolour was white, buff, or chestnut. Subadult femaleand juveniles of both sexes of L. imperialis typicallyshowed a more chestnut colour on the upperparts com-pared with the adults (as in subadult L. edwardsi) andsome dark, pale-tipped markings on wing coverts.Downy young and birds in first contour plumage (stillwith downy heads) showed generally intermediatehead markings, although further study is needed ofthe variation within downy L. nycthemera.

Thus, the strong and repeating pattern both in men-sural and plumage characters was that L. imperialisof all ages and sexes were intermediate betweenthe two putative parental species (without rulingout L. hatinhensis which was extremely similar toL. edwardsi except for the white central rectrices), andthey exhibited no unique external morphological char-acters. This is clearly consistent with the hypothesis ofa hybrid origin, and provides no support for the statusof L. imperialis as a distinct species.

The fact that adult male L. imperialis showed noneof the obvious species-specific plumage characters ofL. nycthemera probably explains why its hybrid originhad not been suspected previously. Also, femaleL. imperialis did not show distinctive characteristicsof the more familiar Chinese races of L. nycthemera,further contributing to the long-unquestioned statusof L. imperialis as a full species. Instead, in maleL. imperialis these obvious L. nycthemera characterswere masked by the black overall colour of one of theparental species, and the tail and crest ornamentationof one parental species was reduced. This patternis typical of numerous other crosses among Lophuraand other pheasant species in which adult malesof one parental species are generally black (P.R.unpubl. data). In contrast to males, femaleL. imperialis showed complete intermediacy, andspecies-specific characters were not masked by blackcoloration.

EXPERIMENTAL CROSS-BREEDING

Delacour described L. imperialis as follows (Delacour& Jabouille, 1924).

Figure 1. Graphs of results of principal components anal-ysis of Lophura imperialis, L. edwardsi, and Vietnameseraces of L. nycthemera. The sexes were included in thesame analysis (see Table 3) but are graphed separatelyhere for clarity.

PC-I

PC

-II

-2 -1 0 1 2 3

3

2

1

0

-1

-2

-3

size (67.5% variance)

bil

l le

ng

th v

s. c

rest

len

gth

(1

2.6

% v

aria

nce

)

PC-I

PC

-II

-2 -1 0 1 2 3

3

2

1

0

-1

-2

-3

size (67.5% variance)

bil

l le

ng

th v

s. c

rest

len

gth

(1

2.6

% v

aria

nce

)

Males

Females

edwardsi

edwardsi

imperialis original series

imperialis original series

nycthemera

nycthemera

imperialis Antwerp hybrids

imperialis Cleres hybrid

imperialis Cat Bin

imperialis Antwerp hybrid



Adult male‘General colour of the male dark shining blue, withbright blue markings on the wing-coverts, back, rump,and upper tail-coverts; crest moderate and black; tailrather long and slightly curved. The feathers on themiddle back and the two central tail-feathers faintlyspotted with reddish-brown. Iris yellowish-brown;skin of the face bright scarlet; legs and feet crimson,spurs white; bill pale green, darker on the base of theupper mandible.

Wing 252 mm; tail 300 mm; culmen 30 mm; tarsus86 mm.’

Adult female‘The adult female is bright chestnut on the back,wings, and upper and lower tail-coverts; head andneck pale brown, with cheeks and throat greyish-buff;primaries black, suffused with pale grey; secondarieschestnut, spotted and bordered with black; two centraltail feathers chestnut-brown, marked with fine blackspots and streaks, the others being black. All thebrown and chestnut feathers are faintly spotted andstreaked with black.

Wing 214 mm; tarsus 67 mm; tail 214 mm; culmen28 mm.’

The phenotypes of the hybrids described below havebeen referenced according to their ring number. Theydiffered from L. imperialis types in the followingfeatures.

Male N∞317Feathers of the back dark blue with reddish-brownfringes. Upper back and wing coverts black spottedwith brown and white. Lower neck inconspicuouslyspotted with whitish feathers. Central pair of rec-trices chestnut spotted with blackish. Second pair ofrectrices black spotted with brown and white. Wingdark blue vermiculated with black and reddishbrown.

Wing 255 mm; tarsus 86 mm; tail broken; culmen31 mm.

Male N∞334Crest dark blue with the base black, some feathersfaintly spotted with white. Back and wings blackinconspicuously spotted with brown. The inner borderof the second pair of tail feathers faintly spotted brownand white.

Wing 254 mm; tarsus 86 mm; tail 314 mm; culmen30 mm.

Male N∞321Feathers of the wing coverts black with a dark metal-lic turquoise blue border. Lower part of the neck witha few shafts vermiculated white. Crest dark blue with

the lower part black and a few feathers vermiculatedwith white on the base of the shaft.


The bill of these three males was pale yellowishgreen and darker on the base of the upper mandible.

When 1 year old, the hybrid males differed fromsubadult L. imperialis in having the two central tailfeathers dark blue, more strongly vermiculated withwhite and chestnut, the second pair of rectrices chest-nut brown vermiculated with black and the third pairblue faintly vermiculated with chestnut on the innerborder.

Female N∞329Underparts more olive-brown than chestnut. Centralpair of rectrices dark brown marked with chestnutspots and streaks. The other rectrices dark brown withchestnut vermiculations becoming obsolete on theexternal tail feathers. Wing feathers dark brown ver-miculated and streaked with chestnut.


With regard to comparative measurements, thethree males came close to L. imperialis with the excep-tion of the tail of male N∞321, which was slightlylonger. The only female differed slightly in being of alarger size.

The tail of the males was long, pointed, and slightlycurved with the central pair longer than the other rec-trices, like L. imperialis. The crests and the generalcolour of these hybrids were excellent despite aslightly longer crest for male N∞321. The mantle ofmales was dark blue, the body feathers being blackwith a blue fringe and the green gloss being absentfrom the wing coverts even though male N∞321 exhib-ited a dark turquoise blue sheen. The colour of thefacial skin, legs, feet, spurs and bills were like that ofL. imperialis. The general colour of the female wasmore olive brown than the female syntype but such acolour was noted amongst the female progeny of theoriginal pair.

Male hybrid N∞317 differed from its brothers in thered brown fringe of the back feathers, the upper backand the wing coverts spotted with reddish brown andwhite, the wing feathers spotted chestnut and the twocentral tail feathers chestnut. In these features itresembled the specimen trapped on 27th February2000 in Da Krong district.

Males N∞317 and N∞321 showed scarce white marksat the base of the neck. Such a character was shown bysome descendants from the original pair and also bythe 1990 Cat Bin specimen. The inconspicuouslybarred crest of males N∞334 and N∞321 was alsoobserved on museum skins and noted by Delacourhimself (Delacour & Jabouille, 1931).



The tail of the female differed likewise from the syn-type in the mixed black and chestnut feathers but oneF1 female kept in BMNH also exhibited this plumageanomaly (N∞1948.70.1).

The tail pattern, with strong whitish or pale vermic-ulations on the two central feathers and even on thesecond pair for males N∞317 and N∞334, representsthe main difference between the F1 male hybrids andthe type description of male L. imperialis. Althoughnot observed on the adult male museum skins, thischaracter was also noted on the two central tail feath-ers of the 1990 specimen.

The more prominent whitish or pale spots on themales’ central tail feathers may have resulted fromthe L. nycthemera subspecies which we selected forthis experimental cross-breeding. Six subspecies ofL. nycthemera occur in Vietnam, of which three over-lap the range of L. edwardsi or L. hatinhensis:L. n. beaulieui, in Ha Tinh Province; Berlioz’s silverpheasant L. n. berliozi in the western slopes of theAnnamitic chain in Quang Tri and Quang Binh Prov-inces; Bel’s silver pheasant L. n. beli on the ridges ofthe eastern slopes of the Annamitic chain in centralAnnam. There is a great deal of individual variationbut the amount of black in the plumage increases fromthe north to the south (Delacour, 1948; McGowan &Panchen, 1994) resulting in much darker southernforms and two ‘black’ L. nycthemera, as can beobserved in L. n. annamensis in Vietnam andL. n. lewisi in Cambodia (Delacour, 1948).Lophura nycthemera beaulieui is a lighter form closelyrelated to the true silver pheasant L. n. nycthemera,the whitest subspecies; L. n. berliozi is intermediatebetween beaulieui and beli and is a rather unstableform showing a great deal of individual variation(Delacour, 1948); L. n. beli is the darkest of the threesubspecies. A crossing between one of these subspeciesand L. edwardsi may result in a variable amount ofwhite in the hybrid’s plumage, the darker phenotypesbeing obtained with L. n. beli. Furthermore the maleof L. n. berliozi which we used for this experiment wasa whitish individual that may have been itself theresult of a past cross-breeding with L. n. nycthemera(S. Moulin, unpubl. data ), which therefore inducedwhitish spots on the tail not typical of L. imperialis.Unfortunately there is no captive population ofthe rare L. n. beli and only this one male L. n. berlioziwas available to us for the experimental cross-breeding.

Alternatively, the pale vermiculations on the centralfeathers may be characteristic of a first hybrid gener-ation, which would be erased by a backcross toL. edwardsi or breeding between a pairing of siblings.No attempt was made to backcross our hybrids toL. edwardsi but we paired male N∞321 with its sisterand obtained one F2 male that looked quite like

L. imperialis, differing only in having a few smallwhite marks at the base of the neck when 1 year old.Furthermore such experiments were conducted by K.Wood using L. edwardsi and L. diardi (pers. comm.).The resultant offspring exhibited very little or nophenotypic variation. Males were inky violet blueand females closely resembled skins of AMNHL. imperialis, although a bit more russet. The prob-lematic characters in the F1 generation disappearedin the backcross with L. edwardsi. Conversely, pairingof siblings from L. diardi ¥ L. edwardsi producedmore variable progeny, some of which resembleL. imperialis, despite purple or clear violet sheens,and others were similar to ‘black’ L. nycthemera.Therefore it seems that a backcross with L. edwardsior the pairing of siblings may produce L. imperialismorphs exhibiting very little phenotypic variation(K. Wood, pers. comm.).

GENETIC STUDIES

The relationships of the L. imperialis mtDNA CRsequences with other species of Lophura are de-scribed by the neighbour-joining tree (Fig. 2). Therewere two kinds of sequences among L. imperialis: thefirst one was identical to homologue sequences from asample of captive-reared and wild L. edwardsi andL. hatinhensis; the other one was related to theL. nycthemera sequences. These findings suggestthat L. imperialis could have received mtDNA fromL. edwardsi/L. hatinhensis females or from popula-tions closely related to L. nycthemera. The neighbour-joining tree represented in Figure 2 is based on c. 184nucleotides only, because the museum samples ofL. imperialis did not allow sequencing of longer partsof the mtDNA CR. Analyses of microsatellite allelesize in species of Lophura are reported in Table 4.It is noteworthy that samples of L. imperialis hadno unique alleles at any locus. Samples attributedto L. imperialis shared alleles with L. edwardsi,L. hatinhensis and L. nycthemera at loci M1, M2 andM4, and did not show unique alleles or alleles sharedwith L. leucomelana and L. swinhoei at loci M1 andM2. Therefore, it is probable that L. imperialis origi-nated from the crossing of L. edwardsi, L. hatinhensisand L. nycthemera.

CONCLUSION

Lophura imperialis showed no unique plumage orshape features. Indeed, there was no consistentphenotype, and each individual showed plumageanomalies also exhibited by known hybrids. Mor-phological data thus strongly support the hybridorigin of L. imperialis, involving L. edwardsi andL. nycthemera as the parents (Fig. 3).



The experimental cross-breeding of L. n. berliozi¥ L. edwardsi strongly support the museum studiesand the hybrid origin of L. imperialis. Two of threehybrid males were very like the type description ofL. imperialis, and the third closely resembled the birdtrapped in Da Krong Province in early 2000. Most ofthe plumage anomalies shown by the Clères hybridswere also exhibited by museum specimens, even thepale spots on the central tail feathers that occurredon the 1990 specimen. Nevertheless the maleL. imperialis described as the type may not be ahybrid of first generation but the result of a backcrossto L. edwardsi or of a pairing of hybrids of the firstgeneration.

Gene analyses confirmed the morphological dataand the hybrid origin of L. imperialis. All mtDNAsequences from museum skins were identical toL. edwardsi sequences, whilst that of the 2000trapped bird grouped with the L. nycthemera clade.Microsatellites were identical to those of L. edwardsior a combination of L. edwardsi and L. nycthemera.

Thus there is overwhelming evidence to concludethat L. imperialis is a natural hybrid and as such itshould be removed from taxonomic lists, as well aslists of species of conservation concern.

On the basis of the geography as well as morphologyand genetic analyses, the putative parental taxa forthe original pair would be L. n. beli ¥ L. edwardsi,whilst that of the 2000 bird would be L. edwardsi¥ L. n. beli. The putative parental taxa of the 1990bird are more enigmatic as L. edwardsi does not occurin Ha Tinh province, where it is replaced byL. hatinhensis. This taxon differs from L. edwardsionly by having several variable and asymmetricalwhite tail feathers. Recent morphological and genetic

Table 4. Molecular size of alleles at studied microsatellite loci in samples of Lophura

Species M1 M2 M3 M4 M5 M6 M7 M8 M9

L. edwardsi 206, 208, 294, 295, 214, 216, 146, 148 284 113, 114, 171, 173 258, 274, 255, 257214, 216, 297, 299 218, 220 115, 116 276218, 228

L. hatinhensis 206, 214 278, 295,296, 297

214, 216,218, 220

146 284 111, 113,114

169, 171,173

258, 276 255, 257,259

L. leucomelana 210, 220, 297 214, 218 148 276, 284 – – – –228

L. nycthemera 210, 214,218, 220,

297, 299 216, 218 146, 148 276, 284 113, 114,115

171 276 255, 257,263

222, 224,228, 232

L. swinhoei 200, 204 294 218 – 276, 283 – 171 – –

L. imperialis 214, 218 295, 299 214, 218 146 276, 284 – – – –

Figure 2. Neighbour-joining tree (based on c. 184 nucle-otides only; see text) showing phylogenetic relationships ofthe sequenced mitochondrial DNA control regions of thethree endemic Lophura species, L. n. nycthemera andL. swinhoei. (LED, L. edwardsi; LHA, L. hatinhensis; LIM,L. imperialis; LNY, L. n. nycthemera; LSW, L. swinhoei).

LED58LED4LED59LED55LED56

LHA2LHA3LHA13LHA8LHA9LHA10LHA11

LIM12LIM13LIM11LIM15LIM14LIM16LIM17LIM18LIM19LIM20

LED57

LHA7LHA 12

LHA15LHA4LHA1LHA14LHA6

LHA5

LIM9LIM8

LNY5LNY3LNY4LNY6

LSW1LSW2

0.005substitutions/site

Quang Tri



studies suggest it may just be an inbred form ofL. edwardsi (A. Hennache & E. Randi, unpubl. data).Therefore its crossing with L. nycthemera wouldresult in phenotypes also resembling L. imperialis,and the putative parental taxa for the 1990 specimenwould be L. n. beaulieui ¥ L. hatinhensis.

CONSERVATION IMPLICATIONS

Hybridization usually occurs in contact zones or incontinuous zones where characters are graded clinallyfrom one taxon to another. Anthropogenic modifica-tions of natural habitats, such as fragmentation, can

also result in hybridization. In these isolated pocketsspecies may hybridize more than they do undermore natural conditions, due to the rarity of matesof their own species (Dowling & Secor, 1997).Lophura edwardsi is a very rare species, so much sothat it was even for a time thought to be extinct in thewild; it is now restricted to small patches of primary orsecondary evergreen forest, while L. nycthemera is ageneralist species that may tolerate a much widerhabitat range. Lophura hatinhensis is also very rareand was only recently brought to the attention of sci-ence. Today, massive deforestation has resulted in for-est fragments where these taxa may cohabit and

Figure 3. Lophura imperialis (B) between its parental species, L. nycthemera (A) and L. edwardsi (C). Painting by JohnSchmitt.

A

B

C



sometimes mate. Thus the fragmented habitatmight induce introgressive hybridization fromL. nycthemera into the wild L. edwardsi andL. hatinhensis populations, of which the birds trappedin 1990 and 2000 are manifestations. Alternatively, itis noteworthy that L. nycthemera and L. edwardsi aresympatric, and it is possible that low levels of hybrid-ization may be natural between these species, thoughthey do not usually share the same habitat.

Hybridization and introgression could introducegenetic variation and reduce inbreeding depression inthe small population size (Grant & Grant, 1994). Pos-itive consequences include a genetic enrichment of theendangered form when the hybrids backcross (Arnoldet al., 1999) and greater fitness in a fluctuating envi-ronment (Grant & Grant, 1994). Negative conse-quences concern the conservation of the rarest taxaL. edwardsi and L. hatinhensis. In the smallest iso-lated forest fragments, populations may evolve combi-nations of characters inherited from the two parentaltaxa and attain evolutionary independence (Dowling& Secor, 1997) of which L. imperialis might be onemanifestation.

ACKNOWLEDGEMENTS

We are grateful to Kermit Wood (Pleistocene Zoologi-cal Trust, USA) for sharing information about thebackcross of F1 hybrids with L. edwardsi, ProfessorVo Quy (CRES, Hanoi) and Jonathan Eames (BirdLifeVietnam) for providing us with descriptions, samplesand pictures of the 2000 trapped bird, Truong Van La(IEBR, Hanoi) and Dang Gia Tung (Hanoi Zoo) fortheir assistance at Hanoi in helping us to check the1990 trapped bird, Robert Prys-Jones (BMNH) andClaire and Jean-François Voisin (MNHN) who pro-vided us with museum samples, Nigel Collar for sug-gesting that P.R. study this problem and for checkinglabel data at the MNHN, and Michel Saint Jalme(MNHN) who kindly set up the artificial inseminationof an L. edwardsi female with semen fromL. n. berliozi. We thank the staff of the following insti-tutions for providing access to specimen material:American Museum of Natural History, New York(AMNH), California Academy of Sciences, San Fran-cisco (CAS), Museum of Comparative Zoology, Har-vard University, Cambridge, MA (MCZ), MuseumNational d’Histoire Naturelle, Paris, France (MNHN),Royal Ontario Museum, Toronto (ROM), University ofMichigan Museum of Zoology, Ann Arbor (UMMZ),National Museum of Natural History, SmithsonianInstitution, Washington, DC (USNM), PeabodyMuseum, Yale University, New Haven (YPM), Zoolo-gisches Museum, Berlin (ZMB). Staff of the AntwerpZoo allowed P.R. to examine specimens of the cap-tive hybrid population kept frozen there. Funding was

provided by the Museum National d’HistoireNaturelle, the Smithsonian Institution, and theIstituto Nazionale per la Fauna Selvatica (INFS),Bologna, Italy.

REFERENCES

Arnold ML, Bulger MR, Burke JM, Hempel AL, WilliamsJH. 1999. Natural hybridization: How low can you go andstill be important? Ecology 80: 371–381.

BirdLife International. 2000. Threatened birds of the world.Barcelona and Cambridge, UK: Lynx Edicions and BirdLifeInternational.

BirdLife International. 2001. Threatened birds of Asia: theBirdLife International red data book. Cambridge, UK:BirdLife International.

Carpentier J, Yealland JJ, Van Bocxstaele R, Van denBerg W. 1975. Imperial Pheasant hybridization at AntwerpZoo. International Zoo Yearbook 15: 100–105.

Collar NJ, Crosby MJ, Stattersfield AJ. 1994. Birds towatch 2. The world list of threatened birds. Cambridge, UK:BirdLife International.

David-Beaulieu A. 1949. Les oiseaux de la province deSavannakhet (Bas-Laos). L’Oiseau Revue Francaise Orni-thologie 19: 41–84, 153–194.

Delacour J. 1925. La reproduction en captivité du Faisand’Edwards et du Faisan Impérial. L’Oiseau. (Nouvelle Série)1: 68–69.

Delacour J. 1948. The subspecies of Lophura nycthemera.American Museum Novitates. 1377: 1–12.

Delacour J. 1977. The pheasants of the world Hindhead, UK:Spur Publications and WPA.

Delacour J, Jabouille P. 1924. Description of twelve newspecies and subspecies from French Indo-China. Bulletin ofthe British Ornithologists Club 45: 28–35.

Delacour J, Jabouille P. 1925. On the birds of Quangtri,Central Annam; with notes on others from other parts ofFrench Indo-China. Ibis Series 12: 209–260.

Delacour J, Jabouille P. 1931. Les oiseaux de l’IndochineFrançaise Volume 1. Exposition coloniale internationale,Paris, France.

Dowling TE, Secor CL. 1997. The role of hybridization andintrogression in the diversification of animals. AnnualReview of Ecology and Systematics 28: 593–619.

Eames JC, Ericson P. 1996. The Björkegren expeditions toFrench Indochina: a collection of birds from Vietnam andCambodia. Natural History Bulletin of the Siam Society 44:75–111.

Eames JC, Lambert FR, Nguyen Cu. 1994. A survey of theAnnamese lowlands, Vietnam, and its implications for theconservation of Vietnamese and Imperial PheasantsLophura hatinhensis and Lophura imperialis. Bird Conser-vation International 4: 343–382.

Felsenstein J. 1985. Confidence limits on phylogenies: anapproach using the bootstrap. Evolution 39: 83–91.

Fuller RA, Garson PJ. 2000. Pheasants status survey andconservation action plan 2000–04 WPA/Birdlife/SSC pheas-



ant specialist group. Gland, Switzerland and Cambridge,UK: IUCN and World Pheasant Association, Reading.

Garson PJ. 1998. Seminar on threatened Vietnamese lowlandpheasants: Hanoi, 18th September 1997. Oriental Bird ClubBulletin 27: 19–20.

Grant PR, Grant BR. 1994. Phenotypic and genetic effects ofhybridization in Darwin’s finches. Evolution 48: 297–316.

Hennache A, Randi E, Lucchini V. 1999. Genetic diversity,phylogenetic relationships and conservation of Edwards’sPheasant Lophura edwardsi. Bird Conservation Interna-tional 9: 395–410.

Huynh Van Keo. 1999. Existence of Edward’s Pheasant atBach Ma National Park. Eighth Annual Conference of SouthEast Asian Zoos Association. 1–3 November 1999, Ho ChiMinh City, Vietnam, Saigon Zoo and Botanical Garden.

King BF, Dickinson EC, Woodcock MW. 1975. A field guideto the birds of South-East Asia. London: Collins.

McGowan PJK, Garson PJ. 1995. Status and conservationaction plan 1995–99: pheasants. Gland, Switzerland: IUCN.

McGowan PJK, Panchen AL. 1994. Plumage variationand geographical distribution in the Kalij and Silverpheasants. Bulletin of the British Ornithologists Club 114:113–123.

Randi E, Lucchini V. 1998. Organization and evolution ofthe mitochondrial DNA control-region in the avian genusAlectoris. Journal of Molecular Evolution 47: 449–462.

Randi E, Lucchini V, Hennache A, Kimball RT, BraunEL, Ligon JD. 2001. Evolution of the mitochondrial DNAcontrol-region and cytochrome b genes, and the inference

of phylogenetic relationships in the avian genus Lophura(Galliformes). Molecular and Phylogenetic Evolution 19:187–201.

Rasmussen P. 1998. Is the Imperial Pheasant Lophura impe-rialis a hybrid? Work in progress and a call for information.Tragopan 9: 8–10.

Rasmussen P. 1999. On the taxonomic status of the ImperialPheasant Lophura imperialis. WPA International Conven-tion. Clères, France, September 1999. [Abstract]

Robson CR, Eames JC, Nguyen Cu, Truong Van La. 1993.Further recent records of birds from Vietnam. Forktail 8: 25–32.

Saitou M, Nei M. 1987. The neighbor-joining method: a newmethod for reconstructing phylogenetic trees. MolecularBiology and Evolution 4: 406–425.

Swofford DL. 1998. PAUP*: phylogenetic analysis using par-simony (and other methods). Version 4 0b2a. Sunderland,MA: Sinauer Associates.

Tamura K, Nei M. 1993. Estimation of the number of nucle-otide substitutions in the control region of mitochondrialDNA in humans and chimpanzees. Molecular Biology andEvolution 10: 512–526.

Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F,Higgins DG. 1997. The CLUSTAL–windows interface: Flex-ible strategies for multiple sequence alignment aided byquality analysis tools. Nucleic Acids Research 24: 4876–4882.

Xu WS, Zhao ZK, Zheng GM, Yang CW, Tan YK. 1996. Afield guide to the birds of China. Taiwan: Kingfisher Press.



AP

PE

ND

IX

Sel

ecte

d pl

um

age

char

acte

rs f

or e

ach

spe

cim

en o

f L

oph

ura

im

peri

alis

an

d th

e pu

tati

ve p

aren

tal s

peci

es (

L. e

dw

ard

si, L

. hat

inh

ensi

s, a

nd

Vie

tnam

ese

race

s of

L. n

ycth

emer

a). L

abel

dat

a on

ori

gin

of

spec

imen

is in

qu

otes

wh

ere

repr

odu

ced

verb

atim

. See

Ack

now

ledg

emen

ts f

or e

xpla

nat

ion

of

mu

seu

mac

ron

yms.

Tab

le A

.A

dult

mal

es (

1)

Spe

cies

Spe

cim

enbi

rd n

oO

rigi

n

Ch

arac

ter

Cre

st c

olou

rC

rest

len

gth

Nec

k/m

antl

eco

lou

rS

capu

lars

/ru

mp

feat

her

sW

ing

cove

rts

Bre

ast

colo

ur

edw

ard

siw

hit

esh

ort

smoo

th,

very

glos

sy b

lue

stro

ngl

y sc

allo

ped,

very

glo

ssy

blu

est

ron

gly

scal

lope

d,ve

ry g

loss

y gr

een

smoo

th,

glos

sybl

ue

hat

inh

ensi

sw

hit

esh

ort

smoo

th,

very

glos

sy b

lue

stro

ngl

y sc

allo

ped,

very

glo

ssy

blu

e st

ron

gly

scal

lope

d,

very

glo

ssy

gree

nsm

ooth

, gl

ossy

blu

e

impe

rial

isM

NH

N19

34.1

388

‘né

et m

ort

enca

ptiv

ité

à C

lère

s’bl

ack,

wh

itis

hba

ses

mid

-len

gth

sm

ooth

, gl

ossy

blu

em

oder

atel

y sc

allo

ped,

glos

sy b

lue

mod

erat

ely

scal

lope

d,gr

een

ish

-blu

e n

ot r

ecor

ded

impe

rial

isM

NH

N19

36.1

548

‘mor

t à

Clè

res,

iss

ude

par

ents

orig

inai

res

d’A

nn

am. .

. , 1

5 av

ril

1936

’

blac

k, w

hit

ish

ba

ses

mid

-len

gth

smoo

th,

glos

sybl

ue

mod

erat

ely

scal

lope

d,gl

ossy

blu

em

oder

atel

y sc

allo

ped,

gree

nis

h-b

lue

not

rec

orde

d

impe

rial

isB

MN

H19

31.5

.11.

5‘is

sued

fro

m w

ild

bird

s . .

.’ba

rred

bla

ckan

d w

hit

e m

id-l

engt

hsm

ooth

, gl

ossy

blu

e-bl

ack

mod

erat

ely

scal

lope

d,gl

ossy

blu

e m

oder

atel

y sc

allo

ped,

gree

nis

h-b

lue

smoo

th,

glos

sybl

ue-

blac

k

impe

rial

isC

AS

607

52di

ed 1

947

blac

km

id-l

engt

hsm

ooth

, gl

ossy

blu

e-bl

ack

mod

erat

ely

scal

lope

d,gl

ossy

blu

e n

o gr

een

tin

gegl

ossy

blu

e-bl

ack,

wh

ite-

shaf

ted

impe

rial

isA

MN

H64

8214

‘Rec

’d 1

956’

‘rec

eive

dfr

om W

m. J

. M

acK

enso

n b

y Je

anD

elac

our.

..’

gree

nis

h-b

lack

rath

er l

ong

smoo

th,

glos

sybl

ue-

blac

k m

oder

atel

y sc

allo

ped,

glos

sy b

lue

mod

erat

ely

scal

lope

d,gr

een

ish

-blu

e sm

ooth

, glo

ssy

blu

e-bl

ack

impe

rial

ism

oun

tA

ntw

erp

hyb

rid

blu

e-bl

ack

rath

er l

ong

smoo

th,

glos

sybl

ue-

blac

k m

oder

atel

y sc

allo

ped,

glos

sy b

lue

mod

erat

ely

scal

lope

d,gr

een

ish

-blu

e sm

ooth

, glo

ssy

blu

e-bl

ack

impe

rial

is35

2/75

An

twer

p h

ybri

d;‘k

eise

rfas

ant’;

16-6

-75

blu

e-bl

ack

rath

er l

ong

smoo

th,

glos

sybl

ue-

blac

k m

oder

atel

y sc

allo

ped,

glos

sy b

lue

mod

erat

ely

scal

lope

d,gr

een

ish

-blu

e sm

ooth

, glo

ssy

blu

e-bl

ack

impe

rial

is59

0/76

An

twer

p h

ybri

d;‘k

eise

r x

zilv

er’;

29-9

-76

blu

e-bl

ack

mid

-len

gth

smoo

th,

glos

sybl

ue-

blac

k m

oder

atel

y sc

allo

ped,

very

glo

ssy

blu

e m

oder

atel

y sc

allo

ped,

very

glo

ssy

gree

nis

h-

blu

e

smoo

th, g

loss

y bl

ue-

blac

k

impe

rial

is65

1/74

An

twer

p h

ybri

d;‘z

ilve

r x

keis

er’

blu

e-bl

ack

mod

erat

esm

ooth

, gl

ossy

blu

e-bl

ack

mod

erat

ely

scal

lope

d,gl

ossy

blu

e m

oder

atel

y sc

allo

ped,

gree

nis

h-b

lue

smoo

th, b

lue-

blac

k



impe

rial

is34

2/82

An

twer

p h

ybri

d;‘k

eise

rfas

ant’;

27-7

-82

blac

k w

ith

mu

ch w

hit

eat

bas

e

mod

erat

esm

ooth

, gl

ossy

blu

e-bl

ack

stro

ngl

y sc

allo

ped,

very

glo

ssy

blu

e st

ron

gly

scal

lope

d,ve

ry g

loss

y gr

een

not

rec

orde

d

impe

rial

is72

0/75

An

twer

p h

ybri

d;‘k

eise

rfas

ant’;

24-1

2-75

blac

klo

ng

smoo

th,

glos

sybl

ue-

blac

k m

oder

atel

y sc

allo

ped,

glos

sy b

lue

mod

erat

ely

scal

lope

d,gr

een

ish

-blu

e sm

ooth

, gl

ossy

blu

e-bl

ack

impe

rial

is64

An

twer

p h

ybri

d;‘b

asta

ard

keis

erfa

san

t’;26

/2/8

8

blu

e-bl

ack

rath

er l

ong

smoo

th,

glos

sybl

ue-

blac

k;w

hit

e sp

eckl

eson

sid

es o

f h

ead

and

nec

k

mod

erat

ely

scal

lope

d,bl

ue

and

copp

er-t

inge

d bl

acki

sh, c

oppe

r-ti

nge

dgl

ossy

blu

e-bl

ack,

stro

ng

wh

ite

shaf

tst

reak

s on

sid

es

impe

rial

is70

1/76

An

twer

p h

ybri

d;‘k

eise

r x

zilv

er’;

2-2-

76

blu

e-bl

ack

very

lon

gsm

ooth

,m

oder

atel

y sc

allo

ped,

glos

sy b

lue-

blac

k;

wh

ite

spec

kles

on

si

des

of h

ead

and

nec

k

mod

erat

ely

scal

lope

d,gl

ossy

blu

e, r

um

pco

pper

-tin

ged

glos

sy b

lue

gree

nis

h-b

lue

nyc

them

era

(an

nam

ensi

s,be

li,

berl

iozi

)

blac

klo

ng

fin

ely

toco

arse

lyve

rmic

ula

ted

blac

k-w

hit

e

fin

ely

to c

oars

ely

verm

icu

late

dbl

ack-

wh

ite

fin

ely

to c

oars

ely

verm

icu

late

dbl

ack-

wh

ite

slig

htl

y gl

osse

dbl

ue-

blac

k

Spe

cies

Spe

cim

enbi

rd n

oO

rigi

n

Ch

arac

ter

Cre

st c

olou

rC

rest

len

gth

Nec

k/m

antl

eco

lou

rS

capu

lars

/ru

mp

feat

her

sW

ing

cove

rts

Bre

ast

colo

ur

Tab

le B

.A

dult

mal

es (

2)

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Sid

e fe

ath

ersh

ape

Un

dert

ail

cove

rts

colo

ur

Mid

-bel

lyco

lou

rC

entr

alre

ctri

ces

Ou

ter

rect

rice

sT

ail

shap

e

edw

ard

siro

un

ded,

not

elon

gate

scal

lope

d, g

loss

ybl

ue

un

glos

sed

blac

kish

glos

sy b

lue-

blac

kgl

ossy

blu

e-bl

ack

vau

lted

, u

ncu

rved

,ra

ther

sh

ort

hat

inh

ensi

sro

un

ded,

not

elon

gate

sc

allo

ped,

glo

ssy

blu

e u

ngl

osse

dbl

acki

shw

hit

e gl

ossy

blu

e-bl

ack

vau

lted

, u

ncu

rved

,ra

ther

sh

ort

impe

rial

isM

NH

N19

34.1

388

‘né

et m

ort

enca

ptiv

ité

à C

lère

s’tr

ian

gula

r sl

igh

tly

scal

lope

d bl

ue-

blac

k sl

aty

blac

kish

blu

ish

-bla

ckbl

ue-

blac

k va

ult

ed, s

ligh

tly

curv

ed, m

id-l

engt

h

Tab

le A

.C

onti

nu

ed



impe

rial

isM

NH

N19

36.1

548

‘mor

t à

Clè

res,

iss

u d

epa

ren

ts o

rigi

nai

res

d’A

nn

am .

. .,

15av

ril

1936

’

tria

ngu

lar

slig

htl

y sc

allo

ped

blu

e-bl

ack

slat

y bl

acki

shbl

uis

h-b

lack

blu

e-bl

ack

vau

lted

, sli

ghtl

y cu

rved

, mid

-len

gth

impe

rial

isB

MN

H19

31.5

.11.

5‘is

sued

fro

m w

ild

bird

s . .

.’tr

ian

gula

rsl

igh

tly

scal

lope

dbl

ack

blu

e-bl

ack

blu

ish

-bla

ckbl

ue-

blac

kva

ult

ed,

slig

htl

ycu

rved

, mid

-len

gth

impe

rial

isC

AS

607

52di

ed 1

947

not

rec

orde

dn

ot r

ecor

ded

not

rec

orde

dfi

nel

y ba

rred

wh

ite

and

blac

kn

ot r

ecor

ded

not

rec

orde

d

impe

rial

isA

MN

H64

8214

‘Rec

’d 1

956’

‘rec

eive

dfr

om W

m. J

. Mac

Ken

son

by J

ean

Del

acou

r . .

.’

slig

htl

y el

onga

te,

tria

ngu

lar-

tipp

ed

slig

htl

y sc

allo

ped

blu

e-bl

ack

un

glos

sed

blac

kish

blu

ish

-bla

ckbl

ue-

blac

kva

ult

ed, s

ligh

tly

curv

ed, m

id-l

engt

h

impe

rial

ism

oun

tA

ntw

erp

hyb

rid

slig

htl

y el

onga

te,

tria

ngu

lar-

tipp

edn

ot r

ecor

ded

un

glos

sed

blac

kish

blu

ish

-bla

ckn

ot r

ecor

ded

vau

lted

, sli

ghtl

y cu

rved

, mid

-len

gth

impe

rial

is35

2/75

An

twer

p h

ybri

d;‘k

eise

rfas

ant’;

16-

6-75

slig

htl

y el

onga

te,

tria

ngu

lar-

tipp

edn

ot r

ecor

ded

un

glos

sed

blac

kish

blu

ish

-bla

ck

blu

e-bl

ack

vau

lted

, mid

-len

gth

(b

ent

in s

tora

ge)

impe

rial

is59

0/76

An

twer

p h

ybri

d;‘k

eise

r ¥

zilv

er’;

29-9

-76

elon

gate

,la

nce

olat

e sl

igh

tly

scal

lope

d bl

ue-

blac

ku

ngl

osse

dbl

acki

shbl

uis

h-b

lack

blu

e-bl

ack

vau

lted

, m

id-l

engt

h(b

ent

in s

tora

ge)

impe

rial

is65

1/74

An

twer

p h

ybri

d;‘z

ilve

r ¥

keis

er’

slig

htl

y el

onga

te,

tria

ngu

lar-

tipp

edsc

allo

ped

blu

e-bl

ack

un

glos

sed

blac

kish

blu

ish

-bla

ckbl

ue-

blac

kpo

orly

pre

serv

ed

impe

rial

is34

2/82

An

twer

p h

ybri

d;‘k

eise

rfas

ant’;

27-7

-82

not

rec

orde

dsl

igh

tly

scal

lope

dbl

ue-

blac

kn

ot r

ecor

ded

blu

ish

-bla

ckbl

ue-

blac

kva

ult

ed,

mid

-len

gth

(ben

t in

sto

rage

)

impe

rial

is72

0/75

An

twer

p h

ybri

d;‘k

eise

rfas

ant’;

24-1

2-75

slig

htl

y el

onga

te,

tria

ngu

lar-

tipp

edn

ot r

ecor

ded

un

glos

sed

blac

kish

blu

ish

-bla

ckbl

ue-

blac

kpo

orly

pre

serv

ed

impe

rial

is64

An

twer

p h

ybri

d;‘b

asta

ard

keis

erfa

san

t’;26

/2/8

8

slig

htl

y el

onga

te,

tria

ngu

lar-

tipp

edn

ot r

ecor

ded

not

rec

orde

dn

ot r

ecor

ded

not

rec

orde

dn

ot r

ecor

ded

impe

rial

is70

1/76

An

twer

p h

ybri

d;‘k

eise

r ¥

zilv

er’;

2-2-

76el

onga

te,

lan

ceol

ate

not

rec

orde

du

ngl

osse

dbl

acki

shbl

uis

h-b

lack

blu

e-bl

ack

vau

lted

, m

id-l

engt

h(b

ent

in s

tora

ge)

nyc

them

era

(an

nam

ensi

s,be

li,

berl

iozi

)

lan

ceol

ate

slig

htl

y gl

osse

dbl

ue-

blac

ku

ngl

osse

dbl

acki

shal

mos

t so

lid

wh

ite

to v

erm

icu

late

dbl

ack-

wh

ite

fin

ely

toco

arse

lyve

rmic

ula

ted

blac

k-w

hit

e

vau

lted

, cu

rved

,ra

ther

lon

g

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Sid

e fe

ath

ersh

ape

Un

dert

ail

cove

rts

colo

ur

Mid

-bel

lyco

lou

rC

entr

alre

ctri

ces

Ou

ter

rect

rice

sT

ail

shap

e



Tab

le C

.A

dult

fem

ales

(1)

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Cre

st c

olou

rC

rest

len

gth

Nec

k/m

antl

eco

lou

rS

capu

lars

/ru

mp

feat

her

sW

ing

cove

rts

Bre

ast

colo

ur

edw

ard

sigr

ey-b

row

nve

ry s

hor

tu

nif

orm

dar

kgr

ey-b

row

nu

nif

orm

ru

fou

s-br

own

un

ifor

m r

ufo

us-

brow

nu

nif

orm

gre

y-br

own

hat

inh

ensi

sgr

ey-b

row

nve

ry s

hor

tu

nif

orm

dar

kgr

ey-b

row

nu

nif

orm

ru

fou

s-br

own

un

ifor

m r

ufo

us-

brow

nu

nif

orm

gre

y-br

own

impe

rial

isM

NH

N19

34.1

389

‘né

et m

ort

enca

ptiv

ité

à C

lère

s’da

rkgr

ey-b

row

nsh

ort

dark

neu

tral

brow

n d

ark

(fre

sh)

rufo

us-

brow

n(f

resh

)da

rk r

ufo

us-

brow

n (

fres

h)

wea

kly

shaf

t-st

reak

ed g

rey-

br

own

impe

rial

isM

NH

N19

28.1

117

‘mor

t à

Clè

res,

le 1

8 S

epte

mbe

r 27

’‘c

otyp

e’

pale

bro

wn

(wor

n)

shor

tbl

otch

y n

eutr

albr

own

(w

orn

) bl

otch

y w

arm

brow

n (

wor

n)

blot

chy

pale

an

dru

fou

s-br

own

(wor

n)

blot

chy,

pal

en

eutr

al b

row

n

impe

rial

isB

MN

H19

48.7

0.1

‘de

Clè

res’

neu

tral

bro

wn

rath

er s

hor

tsl

igh

tly

shaf

t-st

reak

ed n

eutr

albr

own

slig

htl

y sh

aft-

stre

aked

war

mbr

own

slig

htl

y sh

aft-

stre

aked

war

mbr

own

shaf

t-st

reak

edol

ive-

brow

n

impe

rial

isA

MN

H64

8215

‘Rec

’d 1

956’

‘rec

eive

d fr

omW

m.

J. M

acK

enso

nby

Jea

n D

elac

our

. . .’

neu

tral

bro

wn

mid

-len

gth

very

fin

ely

verm

icu

late

d,sh

aft-

stre

aked

grey

-bro

wn

very

fin

ely

verm

icu

late

d,

shaf

t-st

reak

edru

fou

s-br

own

very

fin

ely

verm

icu

late

d,sh

aft-

stre

aked

rufo

us-

brow

n

very

fin

ely

verm

icu

late

d,sh

aft-

stre

aked

rufo

us-

brow

n

impe

rial

is73

9/75

An

twer

p h

ybri

d;‘k

aise

r fa

san

t’,‘3

0–12

–75’

neu

tral

bro

wn

m

id-l

engt

hsl

igh

tly

shaf

t-st

reak

ed n

eutr

albr

own

slig

htl

y sh

aft-

stre

aked

ru

fou

sbr

own

slig

htl

y sh

aft-

stre

aked

ru

fou

s br

own

shaf

t-st

reak

edol

ive-

brow

n

nyc

them

era

(an

nam

ensi

s,be

li, b

erli

ozi)

dark

grey

-bro

wn

rath

er l

ong

shaf

t-st

reak

edpa

le g

rey-

brow

nto

oli

ve-b

row

n

shaf

t-st

reak

edol

ive-

brow

n

shaf

t-st

reak

edol

ive-

brow

n

shaf

t-st

reak

edol

ive-

brow

n



Tab

le D

.A

dult

fem

ales

(2)

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Sid

e fe

ath

ersh

ape

Un

dert

ail

cove

rts

colo

ur

Mid

-bel

ly c

olou

rC

entr

alre

ctri

ces

Ou

ter

rect

rice

sT

ail

shap

e

edw

ard

siro

un

ded,

not

elo

nga

tebr

oad

rufo

us-

brow

n t

ips,

blac

kish

bas

es

un

ifor

mgr

ey-b

row

nve

ry d

ark

ches

tnu

t to

slig

htl

y gl

ossy

bl

acki

sh

slig

htl

y gl

osse

dbl

ack

vau

lted

,u

ncu

rved

,ra

ther

sh

ort

hat

inh

ensi

sro

un

ded,

not

elo

nga

te

prob

ably

as

edw

ard

siu

nif

orm

gre

y-br

own

wh

ite

slig

htl

y gl

osse

dbl

ack

vau

lted

,u

ncu

rved

,ra

ther

sh

ort

impe

rial

isM

NH

N19

34.1

389

‘né

et m

ort

en c

apti

vité

à C

lère

s’

rou

nde

dtr

ian

gles

dark

ru

fou

s,da

rker

bas

es

grey

-bro

wn

very

fra

yed,

grey

-bro

wn

fin

ely

blac

k-sp

eckl

ed c

hes

tnu

tve

ry f

raye

d

impe

rial

isM

NH

N19

28.1

117

‘mor

t à

Clè

res,

le 1

8 S

epte

mbe

r27

’ ‘co

type

’

rou

nde

dtr

ian

gles

dark

ru

fou

s,da

rker

bas

es

grey

-bro

wn

blac

kish

-ch

estn

ut

blac

kish

-ch

estn

ut

vau

lted

, ve

ryli

ttle

cu

rved

,m

id-l

engt

h

impe

rial

isB

MN

H19

48.7

0.1

‘de

Clè

res’

rou

nde

dtr

ian

gles

blac

k-sp

eckl

edru

fou

s-br

own

gr

ey-b

row

nve

ry d

ark

ches

tnu

t-br

own

very

dar

kch

estn

ut

wit

hbr

oad

blac

k ti

ps

vau

lted

, ve

ryli

ttle

cu

rved

,m

id-l

engt

h

impe

rial

isA

MN

H64

8215

‘Rec

’d 1

956’

,‘r

ecei

ved

from

Wm

. J.

Mac

Ken

son

by J

ean

Del

acou

r . .

.’

rou

nde

dtr

ian

gles

ru

fou

s-br

own

-ti

pped

bla

ckis

hdu

ll b

row

nw

eakl

y ru

fou

s-ve

rmic

ula

ted

glos

sy b

lack

ish

glos

sy b

lack

ish

vau

lted

, ve

ryli

ttle

cu

rved

,m

id-l

engt

h

impe

rial

is73

9/75

An

twer

p h

ybri

d;‘k

aise

r fa

san

t’,‘3

0–12

–75’

rou

nde

dtr

ian

gles

bl

ack-

spec

kled

brow

nn

eutr

al b

row

nw

ith

bla

ckis

hbl

otch

es

fin

ely

ches

tnu

t-

verm

icu

late

dbl

ack

rufo

us-

tin

ged

blac

kish

va

ult

ed,

mid

-le

ngt

h (

ben

t in

stor

age)

nyc

them

era

(an

nam

ensi

s,be

li, b

erli

ozi)

rath

ertr

ian

gula

r br

igh

t ru

fou

s-br

own

pale

gre

y-br

own

dark

ch

estn

ut-

brow

nda

rk c

hes

tnu

t-br

own

vau

lted

, sl

igh

tly

curv

ed,

rath

erlo

ng



Tab

le E

.S

uba

dult

mal

es (

1)

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Cre

st c

olou

rC

rest

len

gth

Nec

k/m

antl

e co

lou

rS

capu

lars

/ru

mp

feat

her

sW

ing

cove

rts

Bre

ast

colo

ur

edw

ard

siba

rred

bla

ck-

and-

wh

ite

shor

tso

oty

(old

) an

dve

ry g

loss

y bl

ue

(new

)

dusk

y, r

ufo

us-

mar

ked

(old

) an

dve

ry g

loss

y bl

ue

(new

)

fin

ely

mar

ked

rufo

us

(old

) an

dve

ry g

loss

y,sc

allo

ped

(new

)

soot

y (o

ld)

and

glos

sy b

lue

(new

)

hat

inh

ensi

sw

hit

ish

(det

ails

not

reco

rded

)

shor

tso

oty

(old

) an

dve

ry g

loss

y bl

ue

(new

)

dusk

y (o

ld)

and

glos

sy b

lue

(new

) du

sky

(old

)du

sky

(old

) an

dgl

ossy

blu

e (n

ew)

impe

rial

isM

NH

N19

31.1

44

‘fils

du

cou

ple

orig

inal

prov

enan

t . .

.’ ‘C

lère

s,n

é en

cap

tivi

té m

ai 1

930,

[d

ied]

15

mar

s 19

31’

blu

e-bl

ack

shor

tsm

ooth

, gl

ossy

blu

e-bl

ack

mix

ed r

ufo

us-

brow

n (

old)

an

dsc

allo

ped

blu

e-bl

ack

(new

)

mix

ed r

ufo

us-

brow

n (

old)

an

dbl

acki

sh (

new

)

blu

e-bl

ack

impe

rial

isC

at B

inbl

ue-

blac

klo

ng

glos

sy b

lue-

blac

kw

ith

som

e w

hit

esp

eckl

ing

and

shaf

t st

reak

s

fin

ely

verm

icu

late

dan

d ru

fou

s-br

own

(old

) bl

ue-

blac

k(n

ew)

fin

ely

verm

icu

late

dru

fou

s-br

own

slig

htl

y gl

ossy

blu

e-bl

ack

wit

hw

hit

ish

sh

aft

stre

aks

impe

rial

is31

2C

lère

s h

ybri

dn

eutr

al b

row

n(o

ld)

and

blu

e-bl

ack

(new

)

shor

tn

eutr

al b

row

n (

old)

and

blu

e-bl

ack

wit

hw

hit

e sp

eckl

ing

on n

eck

side

s

fin

ely

verm

icu

late

dan

d bl

ue-

blac

k(n

ew)

fin

ely

barr

edru

fou

s-br

own

(old

)

slig

htl

y gl

ossy

blu

e-bl

ack

rufo

us-

brow

nan

d bl

ack

impe

rial

is31

7C

lère

s h

ybri

dn

eutr

al b

row

n(o

ld)

and

blu

e-bl

ack

(new

)

rath

er s

hor

tn

eutr

al b

row

n (

old)

and

blu

e-bl

ack

wit

hw

hit

e sp

eckl

ing

on n

eck

side

s

fin

ely

verm

icu

late

dan

d bl

ue-

blac

k(n

ew)

fin

ely

barr

edru

fou

s-br

own

(old

)

slig

htl

y gl

ossy

blu

e-bl

ack

rufo

us-

brow

nan

d bl

ack

impe

rial

is32

1C

lère

s h

ybri

dbl

ue-

blac

km

id-l

engt

hn

eutr

al b

row

n (

old)

and

blu

e-bl

ack

wit

hw

hit

e sp

eckl

ing

onn

eck

side

s

fin

ely

verm

icu

late

dan

d bl

ue-

blac

k(n

ew)

rufo

us-

brow

n(o

ld)

rufo

us-

brow

n a

nd

blac

k

fin

ely

barr

edgl

ossy

blu

e-bl

ack

impe

rial

is33

4C

lère

s h

ybri

dn

eutr

al b

row

n(o

ld)

and

blu

e-bl

ack

(new

)

shor

tn

eutr

al b

row

n (

old)

and

blu

e-bl

ack

wit

hw

hit

e

fin

ely

verm

icu

late

d ru

fou

s-br

own

(ol

d)

spec

klin

g on

nec

ksi

des

fin

ely

barr

ed

rufo

us-

brow

nan

d bl

ack

and

blu

e-bl

ack

(new

)

slig

htl

y gl

ossy

blu

e-bl

ack

nyc

them

era

berl

iozi

Clè

res

blac

k-ti

pped

ra

ther

sh

ort

oliv

e br

own

oliv

e br

own

oliv

e br

own

stro

ngl

y bl

ack-

and-

wh

ite

barr

ed, a

nd

blac

k-bl

otch

edce

ntr

ally



Tab

le F

.S

uba

dult

mal

es (

2)

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Sid

e fe

ath

ersh

ape

Un

dert

ail

cove

rts

colo

ur

Mid

-bel

lyco

lou

rC

entr

alre

ctri

ces

Ou

ter

rect

rice

sT

ail

shap

e

edw

ard

siro

un

ded

blac

kish

, w

eakl

ybl

ue-

scal

lope

d so

oty

blac

kish

blac

kish

vau

lted

, u

ncu

rved

,ra

ther

sh

ort

hat

inh

ensi

sn

ot r

ecor

ded

not

rec

orde

dn

ot r

ecor

ded

blac

kish

blac

kish

vau

lted

, u

ncu

rved

,ra

ther

sh

ort

impe

rial

isM

NH

N19

31.1

44‘fi

ls d

u c

oupl

e or

igin

alpr

oven

ant

. . .’

‘Clè

res,

né

en c

apti

vité

mai

1930

, [di

ed]

15 m

ars

1931

’

rou

nde

dtr

ian

gles

not

rec

orde

ddu

ll g

reyi

sh-

blac

kdu

sky

blu

ish

-bl

ack

not

rec

orde

dva

ult

ed,

slig

htl

ycu

rved

,m

id-l

engt

h

impe

rial

isC

at B

inac

ute

tri

angl

es(s

ligh

tly

lan

ceol

ate)

dusk

y bl

ack

wit

h b

road

wh

itis

h s

haf

tst

reak

s

dull

bro

wn

ish

-bl

ack

blac

kish

wit

hst

ron

g ru

fou

s-bu

ff v

erm

icu

lati

on

dusk

y bl

ack

vau

lted

, sl

igh

tly

curv

ed, r

ath

er l

ong

impe

rial

is31

2C

lère

s h

ybri

dac

ute

tri

angl

es(s

ligh

tly

lan

ceol

ate)

blu

e-bl

ack

un

glos

sed

blac

kfi

nel

y bl

ack-

and-

wh

ite

barr

ed,

tips

ru

fesc

ent

blac

k w

ith

wea

kru

fou

s ba

rrin

gon

edg

ing

vau

lted

, sl

igh

tly

curv

ed, m

id-l

engt

h

impe

rial

is31

7C

lère

s h

ybri

dn

ot r

ecor

ded

not

rec

orde

dn

ot r

ecor

ded

mis

sin

gbl

acki

shm

ost

feat

her

sm

issi

ng

impe

rial

is32

1C

lère

s h

ybri

dac

ute

tri

angl

es(s

ligh

tly

lan

ceol

ate)

dusk

y bl

ack

un

glos

sed

blac

kfi

nel

y bl

ack-

and-

wh

ite

barr

ed, t

ips

rufe

scen

t

dusk

y bl

ack

vau

lted

, sl

igh

tly

curv

ed, m

id-l

engt

h

impe

rial

is33

4C

lère

s h

ybri

dn

ot r

ecor

ded

not

rec

orde

du

ngl

osse

dbl

ack

stro

ngl

y bl

ack-

and-

wh

ite

blac

kish

dam

aged

nyc

them

era

berl

iozi

Clè

res

stro

ngl

y bl

ack-

and-

wh

ite

barr

ed

not

rec

orde

dn

ot r

ecor

ded

barr

ed s

tron

gly

blac

k-an

d-w

hit

eba

rred

stro

ngl

y bl

ack-

and-

wh

ite

barr

edva

ult

ed,

un

curv

ed,

mid

-len

gth



Tab

le G

.S

uba

dult

fem

ales

(1)

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Cre

st c

olou

rC

rest

len

gth

Nec

k/m

antl

eco

lou

rS

capu

lars

/ru

mp

feat

her

sW

ing

cove

rts

Bre

ast

colo

ur

edw

ard

sigr

ey-b

row

nve

ry s

hor

tn

eck

grey

-bro

wn

,m

antl

e ru

fou

s-br

own

rufo

us

rufo

us

wit

h b

uff

and

blac

k sp

eckl

es

shaf

t-st

reak

edru

fesc

ent

brow

n

hat

inh

ensi

s*gr

ey-b

row

nve

ry s

hor

tda

rk b

row

nda

rk w

arm

bro

wn

dark

war

m b

row

nn

ot r

ecor

ded

impe

rial

isA

MN

H38

8264

‘bor

n i

n c

apti

vity

at

Yard

ley,

Pa.

May

195

1;di

ed O

ctob

er 1

2, 1

951’

; ‘5

th g

ener

atio

n f

rom

th

e or

igin

al p

air

. . .’

neu

tral

bro

wn

very

sh

ort

fin

ely

verm

icu

late

dw

arm

bro

wn

fin

ely

verm

icu

late

dw

arm

bro

wn

fi

nel

y ve

rmic

ula

ted

war

m b

row

n

shaf

t-st

reak

ed,

fin

ely

verm

icu

late

dw

arm

bro

wn

impe

rial

is48

8/81

An

twer

p h

ybri

d;‘k

eise

rfas

ant’;

sex

and

age

clas

s n

ot c

erta

in

brow

nis

h

rath

er s

hor

tfi

nel

yve

rmic

ula

ted

war

m b

row

n

fin

ely

verm

icu

late

dw

arm

bro

wn

;ru

mp

feat

her

sbl

uis

h-t

ippe

d

fin

ely

verm

icu

late

dw

arm

bro

wn

wit

hbl

ack

blot

ches

an

dw

hit

ish

tip

s

shaf

t-st

reak

edw

arm

bro

wn

impe

rial

is32

9C

lère

s h

ybri

dda

rk g

rey-

brow

nra

ther

sh

ort

dark

war

mbr

own

dark

war

m b

row

nda

rk w

arm

bro

wn

shaf

t-st

reak

edda

rk w

arm

bro

wn

nyc

them

era

(an

nam

ensi

s,be

li, b

erli

ozi)

dark

bro

wn

rath

er s

hor

tn

eutr

al t

o w

arm

br

own

neu

tral

to

war

m

brow

nn

eutr

al t

o w

arm

brow

n w

ith

bu

ff a

nd

blac

k sp

eckl

es

shaf

t-st

reak

ed

pale

neu

tral

to

rufe

scen

t br

own

*Not

dir

ectl

y co

mpa

red

wit

h e

dw

ard

si; f

urt

her

com

pari

son

s n

eede

d.



Tab

le H

.S

uba

dult

fem

ales

(2)

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Sid

e fe

ath

ersh

ape

Un

dert

ail

cove

rts

colo

ur

Mid

-bel

lyco

lou

rC

entr

al r

ectr

ices

Ou

ter

rect

rice

sT

ail

shap

e

edw

ard

siro

un

ded

rufo

us-

tipp

edbl

acki

shbr

own

ish

-gre

yve

ry d

ark

ches

tnu

tbl

acki

sh w

ith

ches

tnu

tm

arki

ngs

vau

lted

, u

ncu

rved

,ra

ther

sh

ort

hat

inh

ensi

s*n

ot r

ecor

ded

not

rec

orde

dn

ot r

ecor

ded

blac

kish

blac

kish

vau

lted

, u

ncu

rved

,ra

ther

sh

ort

impe

rial

isA

MN

H38

8264

‘b

orn

in

cap

tivi

tyat

Yar

dley

, Pa.

May

195

1; d

ied

Oct

ober

12,

195

1’;

‘5th

gen

erat

ion

from

th

e or

igin

alpa

ir’

rou

nde

dtr

ian

gles

rufo

us-

tipp

edbl

acki

shbr

own

ish

-gre

yfi

nel

y ve

rmic

ula

ted

buff

, ru

fou

s, a

nd

blac

kish

blac

kish

vau

lted

, u

ncu

rved

,ra

ther

sh

ort

impe

rial

is48

8/81

An

twer

p h

ybri

d;‘k

eise

rfas

ant’;

sex

and

age

clas

sn

ot c

erta

in

not

rec

orde

dn

ot r

ecor

ded

not

rec

orde

dfi

nel

y ve

rmic

ula

ted

buff

, ru

fou

s, a

nd

blac

kish

blu

e-bl

ack

vau

lted

, sl

igh

tly

curv

ed, m

id-l

engt

h

impe

rial

is32

9C

lère

s h

ybri

dro

un

ded

tria

ngl

esn

ot r

ecor

ded

not

rec

orde

dm

issi

ng

coar

sely

bar

red

rufo

us-

brow

nan

d bl

acki

sh

mos

t fe

ath

ers

mis

sin

g

nyc

them

era

(an

nam

ensi

s,be

li, b

erli

ozi)

not

rec

orde

dru

fesc

ent

pale

gre

y-br

own

fin

ely

mar

ked

ches

tnu

tfi

nel

y m

arke

dch

estn

ut

vau

lted

, sli

ghtl

y cu

rved

, mid

-len

gth

*Not

dir

ectl

y co

mpa

red

wit

h e

dw

ard

si; f

urt

her

com

pari

son

s n

eede

d.



Tab

le J

.Ju

ven

ile

mal

es

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Cro

wn

colo

ur

Man

tle

colo

ur

Sca

pula

rsW

ing

cove

rts

Bre

ast

colo

ur

Sid

e fe

ath

ersh

ape

Mid

-bel

lyco

lou

rC

entr

alre

ctri

ces

Ou

ter

rect

rice

s

edw

ard

sida

rk g

rey-

brow

nli

ghtl

y bl

ack-

spec

kled

ric

hch

estn

ut

(old

),ve

ry g

loss

ybl

ue

(new

)

dark

-edg

edbr

igh

tch

estn

ut

brig

ht

ches

tnu

tw

ith

dar

ksu

bter

min

alba

nds

, bu

ff t

ips

dark

-spe

ckle

dch

estn

ut

(old

),gl

ossy

blu

eof

ten

wit

hch

estn

ut

shaf

tst

reak

s (n

ew)

dark

-spe

ckle

dch

estn

ut

(old

),gl

ossy

blu

eof

ten

wit

hch

estn

ut

shaf

tst

reak

s (n

ew)

dusk

y gr

eyve

ry d

ark

ches

tnu

t to

blu

e-bl

ack

blac

kish

impe

rial

isY

PM

2466

0 ‘S

tate

Gam

eFa

rm,

Ith

aca,

N

.Y.’;

not

yet

full

-siz

ed

dark

brow

nsc

allo

ped

blac

k an

dch

estn

ut

scap

ula

rsch

estn

ut,

tipp

edbl

ack;

ru

mp

ches

tnu

t

ches

tnu

t w

ith

blu

e-bl

ack

subt

erm

inal

ban

ds, b

uff

tip

s

blac

kish

wit

hch

estn

ut

shaf

tst

reak

s

rou

nde

ddi

ngy

wh

itis

hbl

acki

shbr

own

ish

-bl

ack

impe

rial

isB

MN

H19

26.9

.9.3

‘né

à C

lère

sen

Mai

192

6’;

abou

th

alf-

grow

n

neu

tral

brow

nch

estn

ut

ches

tnu

tch

estn

ut

wit

hbl

ue-

blac

ksu

bter

min

alba

nds

, bu

ff t

ips

war

m b

row

nw

ith

pal

e sh

aft

stre

aks

(old

),bl

ue-

blac

k (n

ew)

not

rec

orde

dbu

ffy

and

grey

very

dar

kch

estn

ut

verm

icu

late

dbl

ack

blac

kish

and

ches

tnu

t

impe

rial

isM

NH

N19

27.2

375

‘4 a

oût

1927

’; n

ot f

ull

-siz

edda

rkbr

own

buff

sh

aft-

stre

aked

dar

kw

arm

bro

wn

buff

sh

aft-

stre

aked

dark

war

mbr

own

buff

sh

aft-

stre

aked

dar

kw

arm

bro

wn

,bl

ack

subt

erm

inal

spot

s an

d bu

ffti

ps

buff

sh

aft-

stre

aked

blac

kish

buff

-tip

ped

blac

kish

bl

acki

sh-

grey

bron

zy-b

lack

ches

tnu

t-an

d-bl

ack

barr

ed



Tab

le K

.Ju

ven

ile

fem

ales

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Cro

wn

colo

ur

Man

tle

colo

ur

Sca

pula

rsW

ing

cove

rts

Bre

ast

colo

ur

Sid

e fe

ath

ersh

ape

Mid

-bel

lyco

lou

rC

entr

alre

ctri

ces

Ou

ter

rect

rice

s

edw

ard

sin

ot f

ull

-siz

edda

rkbr

own

shaf

t-st

reak

ed,

wea

kly

verm

icu

late

dda

rk w

arm

brow

n

shaf

t-st

reak

ed,

wea

kly

verm

icu

late

dda

rk w

arm

brow

n

wea

kly

verm

icu

late

dda

rk w

arm

brow

n,

blac

kish

subt

erm

inal

spot

s, b

uff

tip

s

shaf

t-st

reak

edda

rkw

arm

brow

n

shaf

t-st

reak

edda

rk w

arm

brow

n

dark

gre

yve

ry d

ark

ches

tnu

tn

otre

cord

ed

impe

rial

isM

NH

N19

27.2

376

‘jeu

ne

?[m

ale

sym

bol]

,D

on d

eM

Del

acou

r,19

ju

ille

t 19

27’;

not

fu

ll-s

ized

dark

brow

nsh

aft-

stre

aked

,w

eakl

yve

rmic

ula

ted

neu

tral

bro

wn

shaf

t-st

reak

ed,

wea

kly

verm

icu

late

dn

eutr

al b

row

n

wea

kly

verm

icu

late

dn

eutr

al b

row

nw

ith

a f

ewbl

acki

shsu

bter

min

alsp

ots

shaf

t-st

reak

edn

eutr

albr

own

shaf

t-st

reak

edn

eutr

albr

own

wea

kly,

fin

ely

barr

eddu

ll b

row

n

blac

k-fl

ecke

dda

rkch

estn

ut

blac

k-fl

ecke

dda

rk

ches

tnu

t

nyc

them

era

(an

nam

ensi

s,be

li, b

erli

ozi)

not

fu

ll-s

ized

dark

brow

n

fin

ely

verm

icu

late

d n

eutr

al b

row

n

fin

ely

verm

icu

late

dn

eutr

al b

row

n,

som

e fe

ath

ers

wit

h d

ark

subt

erm

inal

spot

s

fin

ely

verm

icu

late

dn

eutr

al b

row

nw

ith

dar

ksu

bter

min

alsp

ots

and

buff

tips

shaf

t-st

reak

edn

eutr

albr

own

shaf

t-st

reak

edn

eutr

albr

own

slig

htl

yba

rred

grey

ish

dark

ches

tnu

tda

rkch

estn

ut



Tab

le M

.F

irst

con

tou

r pl

um

age

(hea

d st

ill

dow

ny)

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Cro

wn

patt

ern

Cro

wn

colo

ur

Su

perc

iliu

mE

ye-s

trip

eM

antl

e co

lou

rS

capu

lars

Win

g co

vert

sB

reas

t co

lou

r

Mid

-be

lly

colo

ur

edw

ard

siC

lère

s 36

day

sn

ot r

ecor

ded

dark

bro

wn

yell

ow-b

uff

stro

ng

blac

kish

post

ocu

lar

stri

pe

not

rec

orde

dch

estn

ut

wit

h l

arge

blac

ksu

bter

min

alsp

ots

ches

tnu

tw

ith

lar

gebl

ack

subt

erm

inal

sp

ots

and

buff

ter

min

alst

reak

s

bold

ly,

irre

gula

rly

barr

ed a

nd

spot

ted

blac

kan

d ch

estn

ut

wh

itis

h

impe

rial

isB

MN

H19

30.6

.12.

2st

ron

g bl

ack

fore

hea

dst

reak

pale

taw

ny

rufo

us-

buff

post

ocu

lar

stri

pe

stro

ng

blac

kish

fin

ely

verm

icu

late

d

pale

sh

aft-

stre

aked

, an

dpa

le-t

ippe

d,

war

m b

row

n

pale

sh

aft-

stre

aked

and

blac

kish

-ti

pped

, fi

nel

yve

rmic

ula

ted

war

m b

row

n

pale

sh

aft-

stre

aked

war

mbr

own

fin

ely

verm

icu

late

dw

arm

bro

wn

pale

sh

aft-

stre

aked

pa

legr

ey-

brow

n

impe

rial

isM

NH

N19

26.2

39‘p

ouss

in,

Hau

t-A

nn

am,

né

et m

ort

enca

ptiv

ité

à C

lère

s’

not

reco

rded

dark

bro

wn

rufo

us-

buff

fair

ly s

tron

gbl

acki

shpo

stoc

ula

rst

reak

dark

ch

estn

ut

wit

hpr

omin

ent

buff

str

eaks

dark

ch

estn

ut

wit

h p

rom

inen

tbu

ff s

trea

ks a

nd

blac

k sp

ots

dark

ch

estn

ut

wit

h p

rom

inen

tbu

ff s

trea

ksan

d da

rk t

ips

blac

kish

wit

hch

estn

ut

bars

and

rufo

us

shaf

t st

reak

s

pale

grey

-br

own

impe

rial

isM

NH

N19

17.2

374

‘pou

ssin

né

et m

ort

enca

ptiv

ité

àC

lère

s’

not

rec

orde

dda

rk b

row

npa

le b

uff

wea

k da

rkpo

stoc

ula

rst

reak

dark

ch

estn

ut

wit

h w

eak

buff

str

eaks

dark

ch

estn

ut

wit

h w

eak

buff

str

eaks

dark

ch

estn

ut

wit

h d

ark

subt

erm

inal

spot

s an

dn

arro

w p

ale

tips

dark

war

mbr

own

wit

hpa

le s

haf

tst

reak

s

pale

grey

-br

own



impe

rial

isC

lère

sh

ybri

d H

129

an

d36

day

s n

ot r

ecor

ded

dark

bro

wn

yell

ow-b

uff

stro

ng

blac

kish

post

ocu

lar

stri

pe

fin

ely

verm

icu

late

dbr

own

wit

hst

ron

g bu

ffsh

aft

stre

akan

d bl

ack

subt

erm

inal

spot

s

fin

ely

verm

icu

late

dbr

own

wit

hst

ron

g bu

ffsh

aft

stre

akan

d bl

ack

subt

erm

inal

spot

s

fin

ely

verm

icu

late

dbr

own

wit

hsm

all

blac

ksu

bter

min

alsp

ots

and

buff

tips

dark

bro

wn

wit

h s

tron

gbu

ff s

haf

tst

reak

s an

dti

ps

not

reco

rded

impe

rial

isC

lère

sh

ybri

d H

231

day

sn

ot r

ecor

ded

pale

taw

ny

yell

ow-b

uff

stro

ng

blac

kish

post

ocu

lar

stri

pe

not

rec

orde

dn

ot r

ecor

ded

fin

ely

verm

icu

late

dbr

own

wit

hsm

all

blac

ksu

bter

min

alsp

ots

and

buff

tips

rufo

us-

and-

blac

k-ba

rred

wit

h s

tron

gbu

ff s

haf

tst

reak

s an

dti

ps

not

reco

rded

impe

rial

isC

lère

sh

ybri

d H

3 35

day

sst

ron

g bl

ack

fore

hea

dst

reak

dark

bro

wn

ye

llow

-bu

ffst

ron

gbl

acki

shpo

stoc

ula

rst

ripe

not

rec

orde

dn

ot r

ecor

ded

fin

ely

verm

icu

late

dbr

own

wit

hsm

all

blac

ksu

bter

min

alsp

ots

and

buff

tips

dark

bro

wn

wit

h s

tron

gbu

ff s

haf

tst

reak

s an

dti

ps

not

reco

rded

impe

rial

isC

lère

sh

ybri

d H

435

day

sn

ot r

ecor

ded

dark

bro

wn

yell

ow-b

uff

stro

ng

blac

kish

post

ocu

lar

stri

pe

not

rec

orde

dn

ot r

ecor

ded

fin

ely

verm

icu

late

dbr

own

wit

hsm

all

blac

ksu

bter

min

alsp

ots

and

buff

tips

fin

ely

barr

edbr

own

an

dbl

ack

wit

hst

ron

g bu

ffsh

aft

stre

aks

and

tips

not

reco

rded

nyc

them

era

berl

iozi

Clè

res

33 d

ays

not

dow

ny

inin

divi

dual

ph

oto-

grap

hed

not

dow

ny

inin

divi

dual

ph

oto-

grap

hed

rufo

us-

buff

stro

ng

blac

kish

post

ocu

lar

stri

pe

fin

ely

dark

-ba

rred

pal

e br

own

wit

h

stro

ng

pale

sh

aft

stre

aks

fin

ely

dark

-ba

rred

pal

e br

own

wit

h

stro

ng

pale

sh

aft

stre

aks

fin

ely

dark

-ba

rred

pal

e br

own

wit

h

broa

d ye

llow

bu

ff t

ips

fin

ely

barr

ed,

blac

k, w

hit

ean

d bu

ff

pale

grey

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Cro

wn

patt

ern

Cro

wn

colo

ur

Su

perc

iliu

mE

ye-s

trip

eM

antl

e co

lou

rS

capu

lars

Win

g co

vert

sB

reas

t co

lou

r

Mid

-be

lly

colo

ur



Tab

le N

.D

own

y ch

icks

Spe

cies

S

peci

men

bird

no

Ori

gin

Ch

arac

ter

Cro

wn

pat

tern

Cro

wn

col

our

Su

perc

iliu

mE

ye-s

trip

eB

reas

t co

lou

r

edw

ard

sih

atin

hen

sis

evid

entl

yve

ry s

imil

ar

stro

ng

blac

kfo

reh

ead

stre

akta

wn

y br

own

wh

itis

h t

obu

ffy

stro

ng

blac

kish

post

ocu

lar

stri

pest

ron

gly

rufe

scen

t

impe

rial

isA

MN

H74

8400

‘on

e da

y ol

d’w

eak

dark

fore

hea

d st

reak

pa

le r

ufo

us

very

pal

eru

fou

sn

arro

w d

ark

brow

n p

osto

cula

rw

ash

edru

fesc

ent

impe

rial

isB

MN

H19

30.5

.8.1

‘h

atch

ed 1

4th

May

& d

ied

16th

May

193

0’

stro

ng

blac

kfo

reh

ead

stri

pe

dark

bro

wn

wh

itis

hst

ron

g bl

acki

shpo

stoc

ula

r st

ripe

taw

ny

impe

rial

isB

MN

H19

39.6

.2.1

‘5 d

ays

old’

wea

k da

rkfo

reh

ead

stre

ak

pale

ru

fesc

ent

wh

itis

hn

arro

w d

ark

brow

n p

osto

cula

rpa

le t

awn

y

impe

rial

isH

1C

lère

s h

ybri

d,5

days

not

rec

orde

d ru

fou

sru

fou

s-bu

ffst

ron

g bl

acki

shpo

stoc

ula

r st

ripe

was

hed

rufe

scen

t

nyc

them

era

un

patt

ern

ed

rufo

us

rufo

us-

buff

wea

k, r

ufe

scen

tpo

stoc

ula

rli

ttle

to

no

rufe

scen

t w

ash

Documents

Hybrid origin of the imperial pheasant Lophura imperialis ...viduals (Delacour, 1977), but these died en route to France and their identity was never veriﬁed. In 1938, C. Cordier,