Genetic variability in French populations of the Corsican mouflon (Ovis ammon musimon): analysis of 2 blood proteins and red-cell blood groups

HAL Id: hal-00894035

https://hal.archives-ouvertes.fr/hal-00894035

Submitted on 1 Jan 1994

HAL is a multi-disciplinary open access

archive for the deposit and dissemination of

sci-entific research documents, whether they are

pub-lished or not. The documents may come from

teaching and research institutions in France or

abroad, or from public or private research centers.

L’archive ouverte pluridisciplinaire HAL, est

destinée au dépôt et à la diffusion de documents

scientifiques de niveau recherche, publiés ou non,

émanant des établissements d’enseignement et de

recherche français ou étrangers, des laboratoires

publics ou privés.

Genetic variability in French populations of the Corsican

mouflon (Ovis ammon musimon): analysis of 2 blood

proteins and red-cell blood groups

C Montgelard, Tc Nguyen, D Dubray

To cite this version:

Original

article

Genetic

_variability

in French

populations

of the Corsican mouflon

(Ovis

ammon

musimon):

analysis

of

2

blood

_proteins

and

red-cell

blood

_groups

C

_Montgelard

TC

_Nguyen

₂

D

_Dubray

1

Institut des Sciences de

l’Évolution

_(URA

827

_CNRS),

Laboratoire de

_{Paléontologie}

des Vertébrés

_(EPHE),

UM

_II,

CC 64, 3!095

Montpellier

Cedex 5;

2

INRA,

Unité

_{Polymorphisme Sanguin}

Ovin et

_Caprin,

Laboratoire des

_{Groupes Sanguins,}

78350

_{Jouy-en-Josas;}

3

Office

National de la

Chasse, CNERA,

Faune de

_Montagne,

BP _607,!,

84030

Montpellier

Cedex 1, France

(Received

8

_April

1993;

accepted

11

_February

₁₉₉₄₎

Summary - Genetic variation in 7 French

_populations

of the Corsican mouflon

_(Ovis

ammon

musimon)

was

_{investigated by}

red-cell

_{blood-group typing}

and

electrophoresis

of

hemoglobin

and transferrin. The 7

_populations

included 3 that were _captive, 3 established

by introducing

animals into the wild and 1 native

_population

on Corsica. Raw data were treated

_by

classical monofactorial indexes and multivariate

_analyses.

These

_analyses

revealed no

_significant

reduction of

genetic

variability

in either the captive or the

introduced feral

_{populations, although}

the number of founder individuals was _very low in both cases. The maintenance of _genetic variation is

_{explained by}

the diverse

_geographical

origins

and

_by

the

_{genetic divergence}

of the founder individuals. Moreover, multivariate

analyses

revealed a

_genetic

structure related to the

_origin

of the founder

populations.

Long-term resilience of gene combinations of blood factors and absence of gene flow between

populations

are

_suggested

as the factors

_explaining

conservation of the

_{genetic make-up}

of the founder individuals over several

generations. Finally,

the presence of the

hemoglobin

A allele in 4 of the

_populations

allows discussion of the emergence of the mouflon

by

feralization from archaic

_sheep.

Ovis ammon musimon

_/

red-cell blood groups

/

genetic variability

/

origin of

populations

/

multivariate analysis

Résumé - Variabilité _génétique chez les populations françaises du mouflon de

Corse

_(Ovis

ammon

musimon) :

analyse de 2 _protéines du sang et des groupes

sanguins érythrocytaires. La variabilité

_génétique

de 7

_{populations françaises}

du

mouflon

de

l’hémoglobine,

de la

_transferrine

et des

_facteurs

antigéniques

des groupes sanguins

érythrocytaires.

Trois

_populations

élevées en

captivité,

3 populations

introduites sur le

continent et la

_population

autochtone de Corse ont été

_analysées.

Les données sont

traitées par des indices

monofactoriels

et des

analyses

multivariées. Ces

analyses

ne révèlent pas de réduction de la variabilité

génétique,

ni pour les

populations

en

_captivité

ni _pour les

populations

sauvages

introduites,

bien que ces

_populations

aient été

fondées

à partir d’un petit nombre d’individus. Le maintien de la variabilité est

expliqué

par la

diversité de

_{l’origine géographique}

et la

_{divergence génétique}

des animaux

_fondateurs.

La

différenciation génétique

observée entre les

_populations

est en outre corrélée avec

_l’origine

des individus introduits. Ce résultat est

_expliqué

en _invoquant l’absence de

flux génique

entre les

populations

et par la conservation de combinaisons de

gènes

(phénogroupes

des

groupes

sanguins)

sur

plusieurs générations.

La

présence

de l’allèle A de

l’hémoglobine

permet aussi d’aborder la question de

l’origine

du

_mouflon

par marronnage.

Ovis ammon musimon

_/

groupes sanguins érythrocytaires

/

variabilité génétique

/

origine des populations

/

analyses multivariées

INTRODUCTION

The

present

distribution of the Corsican mouflon

_{( Ovis}

ammon

_mvsimon)

is

essen-tially

the result of mainland introductions from native

_populations

on Corsican and

Sardinian islands

_(Tomiczek,

_1985;

Uloth and

Prien,

1985).

Numerous

European

countries

_(from

_Spain

to

_ex-USSR)

and other

_parts

of the world

_(USA,

_Kerguelen

Islands)

also harbor introduced mouflon

_populations

_(Nadler

et

_al,

_1971;

ONC,

1985).

In

France,

introductions took

_place

in the years 1950-1965 and

wild-living

populations

are now

_present

in the mountains of the

Alps,

the

Pyr6n6es,

the

C6vennes and the Massif Central.

The

_genetic

_variability

of the Corsican mouflon has been assessed

mainly

by

means of

_{electrophoretic}

analyses,

which revealed

_large

differences between

populations.

Stratil and Boback

₍₁₉₈₈₎

_analyzed

90 mouflons from Czechoslovakia for 10 blood

_proteins,

5 of which were

polymorphic

(transferrin,

hemopexin,

esterase-A,

X-protein

and

_catalase).

A similar result was obtained

_by

Randi

et al

₍₁₉₉₁₎

who found 4

_polymorphic

loci

_(hexokinase,

isocitrate

_{deshydrogenase,}

glucose-6-phosphate dehydrogenase

and

_{malico-enzyme)}

among the 33 loci scored in 10 Italian individuals. On the other

_hand,

the

_study

of Hartl

₍₁₉₉₀₎

revealed a

very low biochemical

variability

in 4

populations

from the Austrian

Alps: only

one

locus

_(esterase-2)

of the 31

_protein

_systems

studied was

_polymorphic.

The difference

between the latter results is worth

_{noting considering}

that 22

_loci,

_including

the 5

polymorphic

loci

mentioned,

were common to both studies.

In French mouflon

_populations,

genetic

diversity

was determined for 20

captive

individuals

_(Jardin

des Plantes from Paris and INRA from

_{Jouy-en-Josas).}

Elec-trophoresis

of transferrin and

_hemoglobin

_(Nadler

et

_al,

_1971;

Bunch et

_al,

₁₉₇₈₎

showed that both loci were

_polymorphic

and the

_analysis

of red-cell blood _groups

(Nguyen

and

_{Bunch, 1980;}

Bunch and

_Nguyen,

₁₉₈₂₎

indicated a

_great

_proximity

between domestic

_sheep

and the Corsican mouflon.

In the

_present

_paper, we

_{analyse genetic}

variation in 7 French

populations

on the

_genetic

variability

of introduced mouflon

populations.

Genetic variation and

structure are discussed in relation to the

_history

of the introductions

_(origin

and

number of founder

_{individuals).}

MATERIALS AND METHODS

Animals

Blood

_samples

were collected from 172 mouflons

_belonging

to 3

_captive

_populations

(Paris,

Jouy-en-Josas

and

_Lunaret)

and 4 feral ones

_(Corsica,

_Bauges,

Caroux and

Italy) (table I).

Genetic

_systems

Polymorphism

of transferrin and

_hemoglobin

was detected

by

starch-gel

elec-trophoresis

(Nguyen

and

Bunch,

1980)

and

_by

electrofocalisation in the case of

hemoglobin.

Genetic distances were calculated from allelic

frequencies

using

Gre-gorius’

distance,

which satisfies all the

_properties

of a mathematical distance

(Gre-gorius,

1984).

Relationships

between

_populations

were inferred

using

the

Neighbor

program of the

Phylip package

(Felsenstein, 1990).

Seven

_genetic

_systems

of red-cell blood groups

(OEA:

ovine

erythrocyte

antigens)

were

analysed: OEA-A,

B, C, D, M,

R and F30

(Committee

on Genetic Nomen-clature of

Sheep

and

_Goat;

_Nguyen,

_1986).

_Twenty-five

different

sheep

reagents

were

_tested,

corresponding

to the

factors,

Aa,

Ab and A16 for

OEA-A; Bb, Be, Bd,

Be, Bf,

Bg,

Bh, Bi, BF3, BF35,

BF38 and BF418 for

_OEA-B;

Ca,

Cb and CF5 for

_OEA-C;

Da for

_OEA-D;

Ma and Mb for

OEA-M;

R and 0 for

OEA-R;

F30 and F275.

_Hemolytic

and

_{hemagglutination}

techniques

were used in

_blood-group

typing

(Nguyen,

1972;

Nguyen

and

Ruffet,

1975).

For each

_population,

the level

of

_polymorphism

was

computed by

considered a blood factor as

_polymorphic

if its

frequency

in a

population

was between 5 and

95%.

Multivariate

_analyses

Among

multivariate factor

methods, correspondence

analysis

is

_particularly

suited

for

_qualitative

data. Raw data are transformed into a

_contingency

table

(individuals

x

_genetic

_systems).

For each blood

_factor,

each individual is

assigned

the value 1

(positive reaction)

or 0

(negative reaction).

For the

hemoglobin

and transferrin

loci,

each individual is coded 1 for the

_genotype

it

_presents.

Theoretical and

_practical

aspects

of this method

_applied

to

_genetic

_systems

are described in She et al

(1987).

A hierarchical classification of the coordinates of individuals on the factor axes

was

performed

from the

correspondence analysis according

to the method of Roux

(1985).

Major

groups in the classification were retained to

_produce

a

_dendogram.

Group

compositions

were tested

by

a

_contingency

table

analysis

(populations

x

group

compositions)

and the randomness of the distribution

(null

hypothesis)

was

assessed

by

a

_X

_-squared

test.

Multivariate

_analyses

were

_performed

on Biomeco software

(Lebreton

et

al,

RESULTS

Blood

_proteins

Table II

_provides

the allelic

_frequencies

for the

_hemoglobin

and transferrin loci in the 7

_populations.

Four alleles were detected at the transferrin

_locus,

_{TrfD being}

the most common, as in other mouflon

_populations

_(Nadler

et

_al,

_1971;

Stratil and

the Corsican mouflon. At the

_hemoglobin

locus,

2 alleles HbA and HbB were scored. The

_populations

of

_Jouy,

Caroux and Lunaret were

_monomorphic

for the HbB allele.

This was also the case for the Czechoslovakian

(Stratil

and

Bobak,

1988),

Sardinian

(Naitana

et

_al,

₁₉₉₀₎

and Italian

_(Randi

et

_al,

₁₉₉₁₎

_{populations. Finally,}

HbA was

observed

_only

in the

_populations

of

_{Paris, Corsica, Bauges}

and

_Italy.

Genetic differentiation

_computed

_using

_Gregorius’

_genetic

distance

_yielded

dis-tance values between 0 and 0.302 with a mean of 0.176

(table III).

From the distance

matrix,

the

_{neighbour-joining}

method was used to construct an unrooted network

(no

assumption

of molecular

_clock)

_showing

the

_{genetic relationships}

between popu-lations

_{(fig 1).}

Based on these 2

loci,

the

Bauges

and Paris

populations appeared

to be

_closely

related. Another cluster included the

_populations

of

Corsica,

Jouy,

Lunaret and

Caroux,

whereas the Italian

_{sample occupied}

an intermediate

_position

between the _{2 groups.}

Blood _group

_antigens

22

_polymorphic

_reagents

are

provided

in table IV.

Large frequency

differences were observed between

_populations,

but

_only

the Bh and Ca factors showed an overall low

_frequency

_{(< 5%).}

Our results are

roughly

similar to those

reported by Nguyen

and Bunch

₍₁₉₈₀₎

_concerning

the number and

frequency

of blood _group factors. For each

_population,

the level of

_{polymorphism,}

ie the

percentage

of

_polymorphic

blood

factors,

was calculated

_{(table IV).}

The lowest values were observed in

populations

of

_{Paris, Bauges}

and

_Italy

_(0.26,

0.28 and

_0.4,

_{respectively),}

which were also the smallest

samples

analyzed.

In the other

populations, polymorphism

ranged

between 0.45 and

_0.61,

with the Corsican

_population

_showing

the

_highest

value.

Multivariate

_analysis

performed

on 21

_polymorphic

_genetic

_systems

_(hemoglobin,

transferrin and the 19

remaining

blood

factors,

see table

IV)

yielding

44 variables.

Figure

2 shows the

_projections

_{(’envelopes’}

_{joining points}

of extreme

_{distribution)}

of each

sample

in the

_plane

defined

by

the first 2 factor axes, which account for

24.5%

of the total

_variability.

The factor axis 1

_mainly

characterized the individuals from Paris due to the _presence of the

_Bg

and Mb blood

factors,

the absence of Ma and

heterozygosity

at the

_hemoglobin

locus. The factor axis 2 opposes individuals which do not

_possess

_Bf,

BF3 and Be and those

_presenting

BF3 and Be. These variables

separate

the

_majority

of Corsican animals from the

populations

of

Lunaret, Jouy,

Italy

and

_Bauges.

The individuals from Caroux were distributed all

along

this

axis,

The hierarchical classification was

performed

on coordinates of individuals on the

first 6

_principal

axes which

_represented

50.8%

of the overall

_variability.

Five

_major

groups are

_apparent

_(fig

3)

between which individuals are not

_randomly

distributed.

Groups

A and B which include all individuals from Corsica are the

only

ones between which the

composition

was not

_{statistically}

different

₍

_X

_-squared

=

_9.64,

df

=

_5,

p >

_0.05).

On the other

hand,

the

_composition

of _groups A +

B,

D and E

appeared statistically

different

_(x-squared

=

81.3,

df

=

_18,

p <

_0.001).

Group

C

(the

sample

from

_Paris)

was not included in the test because of its small

_sample

size. Most individuals from

_{Jouy, Bauges}

and

_{Italy belong}

to _group E. The Caroux

DISCUSSION

Maintenance of

_{genetic variability}

Except

for the native Corsican

_population,

all the other

_populations

studied were established

_{by releasing}

a small number of animals either in the wild or in enclosures

_(see

table

_I).

In small founder

_populations,

loss of

_genetic

information is

expected

due to

_genetic

bottlenecks and

_{subsequent inbreeding.}

_{Nevertheless,}

a clear

reduction in

_genetic

_variability

was not observed in our

_study

between introduced and native mouflon

_populations.

_Among

the 25

_reagents

_{tested, only}

2

_(Bh

and

CF5)

present

in the Corsican

_population

were not recovered in the other

_samples.

The

_{only populations showing}

a decrease in

variability

are the 3

(Paris,

Bauges

and

_Italy)

for which the

_sample

sizes are the smallest

(4 individuals).

In the

_larger

samples,

the level of

_polymorphism

of blood factors

_{(table IV)}

is similar _among

captive

(Jouy

and

_Lunaret),

feral

_(Caroux)

and native

_(Corsica)

_populations.

Reduction in

genetic

variability subsequent

to low effective

_population

size is not rare for

large

mammals reintroduced from

refuge

zones. This

_phenomenon

has

been observed for the fallow deer

_(Randi

and

_Apollonio,

₁₉₈₈₎

and for the

_Alpine

Ibex

_(Stuwe

and

Scribner,

1989).

The maintenance of

_genetic

_variability

observed in the mouflon

_populations

may be

explained by

the

geographical diversity

and

genetic

divergence

of the founder individuals. As seen in table

_I,

French

populations

always

originated

from animals

_coming

from localities as diverse _as Corsica and

Czechoslovakia. The

_genetic

studies

_performed

on

Czechoslovakian

(Stratil

and

Bobak,

1988),

Austrian

_{(Hartl, 1990)}

and Italian

_(Randi

et

al,

1991)

mouflon

populations

indicated that these

_populations

were

genetically

differentiated. Such

results

_suggest

that the founder mouflons may have been

sufficiently divergent

genetically

to overcome

_genetic

drift and

_inbreeding

at the initial

_stages

of the

introduction. This

_geographical

_genetic

_diversity

observed in

_European

mouflon

populations

may be related to the

history

of introductions which are known to differ

_according

to the

_regions

considered

_(Tomiczek,

_1985;

Uloth and

Prien,

1985)

including breeding

with domestic

_sheep

or with Middle-Eastern

subspecies

of

mouflon

_{(Pfeffer, 1984).}

Genetic structure and

_origin

of founder animals

Multivariate

_analyses

_{(correspondence}

_analysis

and hierarchical

_{classification)}

re-vealed a

_genetic

classification into 5

_major

_groups

_{(fig 3);}

the native Corsican

_sample

(all

individuals

_belong

to the _groups A

_plus

B in

_figure

₃₎

_being

_relatively

differen-tiated from the introduced

_populations.

The

pattern

of introductions and the

_origin

of founder

_populations

_{(table I)}

may

provide

an

_explanation

for this

_genetic

struc-ture. Most individuals of

_{Italy, Bauges}

and

_Jouy,

which are classified in the same

group

(E

in

fig

3),

originated

from

founders,

part of which were introduced from

Chambord between 1954 and 1962. The

_population

from Paris

_{(group C)}

appears

well

differentiated,

which may be

partly

due to a

sampling

effect but also to its _very

diverse

_pool

of founders.

_Finally,

the individuals from Caroux are distributed into the same _{3 groups}

_(A,

D and

_E)

as the Lunaret

_sample

which was created with

A correlation between the

_genetic

structure and the

_origin

of founder mouflons

implies

the

_persistence

for several years of the

genetic

make-up

of the founder

animals.

Two factors

_{might explain}

this

_phenomenon.

First the absence of _gene

flow due to

_geographical

isolation has

_prevented

_genetic

_{homogenization.}

The

populations

studied are not in

_contact,

either because

_they

are

_captive

or because

they

live in isolated mountains. The second factor

_allowing

a resilience of the

genetic

make-up

may be the

genetic

systems

under

_study

_{(blood factors),}

which are

_composed

of several

_genetic

determinants

_(haplotype).

The

_genetic

structure

observed is

_mostly

based on factors of the B

_system

_(Bg,

_Be,

Bf and BF3 in the

correspondence

analysis)

which is known to be controlled

_by

a set of

_closely

linked genes

(Grosclaude

et

al,

1983).

The

_{quasi-absence}

of recombination events _may have allowed

_particular

_genetic

combinations to

_persist

over several

_generations.

Origin

of the mouflon

_by

feralization

The

_origin

of the Corsican mouflon is not well known.

_According

to

_{paleontological}

and anatomical data

_(Poplin,

_1979;

_Vigne,

_1983),

the mouflon

_originated

from the first domesticated

_sheep

_(Neolithic)

followed

_by

feralization in the Corsico-Sardinian

islands. _{The emergence of the mouflon would have therefore occurred between 6 000} and 4 000 BC

_{(Vigne, 1983).}

_Support

for this

_hypothesis

was

_provided

_by

_’

_Bunch

et al

₍₁₉₇₈₎

on the basis of the

_hemoglobin

_{(,0 chain)}

allele distribution. The Hb

A allele is

_reported

to occur

_only

in

_sheep

and Corsican

_mouflon,

but not in the

closest relatives of the

_European

mouflon

_(Asiatic

and Middle-Eastern

_mouflon).

Bunch et al

₍₁₉₇₈₎

_suggested

that the Corsican mouflon

_originated

from an archaic

strain of domestic

_sheep

that carried the HbA allele. Until now, the HbA allele has been recorded

_only

in some of the

_populations

of our

_study

(Corsica,

_Paris,

Bauges

and

_Italy)

but was not detected in Czechoslovakia

_(Stratil

and

Bobak,

1988),

Italy

(Randi

et

_al,

₁₉₉₁₎

or

Sardinia,

where Naitana et al

₍₁₉₉₀₎

studied 100 wild mouflons. If the

_hypothesis

of Bunch et al is correct the

_{subsequent disappearance}

of HbA in Sardinia needs to be

_postulated.

However,

the

_question

arises as to whether HbA recorded in

_sheep

and mouflon

corresponds

to the same allele. Stratil and Bobak

(1988)

as well as Naitana

et al

₍₁₉₉₀₎

have shown

_by

isoelectric

_focusing

and HPLC

_{(reverse-phase}

_high

performance liquid

chromatography)

that HbB of

_sheep

and mouflon are different

although

they appeared

indistinguishable

by

starch

_{electrophoresis.}

At

_present,

support

for the _emergence of the mouflon

by

feralization

_requires

additional HPLC

data

_establishing

the

_identity

of the HbA alleles in

_sheep

and mouflon.

ACKNOWLEDGMENTS

We would like to thank all our collaborators who have taken part in the collection of

samples:

MP

Battesti,

JC

Franceschetti,

D

Roux,

J Vitti and the ONC

_gamekeepers

from the BMI Corsica; P

_Gibert,

H Houssin and the

_personnel

of the

_Bauges

reserve; ME Cresci

and E Bracco for the Italian

sample;

JM

_Cugnasse

and M Garcia for the Caroux

sample;

and M Gallet and the

personnel

from the Zoo de Lunaret. We thank J Britton-Davidian and L Ellison for

_helpful

comments on the manuscript. Data

analysis

was financed

by

the

Office

National de la Chasse. This is contribution 94-016 of the Institut des Sciences de

REFERENCES

Bunch

_{TD, Nguyen}

TC

₍₁₉₈₂₎

Blood _{group comparisons} between _European mouflon

sheep

and North American desert

_{bighorn sheep.}

J Hered _73, 112-114

Bunch

_{TD, Nguyen TC, Lauvergne}

JJ

₍₁₉₇₈₎

_Hemoglobins

of the Corsico-Sardinian mouflon

_(Ovis

_musimon)

and their

_implications

for the

_origin

of HbA in domestic

sheep

(Ovis

aries).

Ann Genet Sel Anim 10, 503-506

Felsenstein

₍₁₉₉₀₎

Phylip,

version 4.0, University of

_{Washington, Seattle,}

WA

Gregorius

H

₍₁₉₈₄₎

A

_{unique genetic}

distance. Biometrical J 26, 13-18

Grosclaude F, Lefebvre

_J,

No6 G

₍₁₉₈₃₎

Nouvelles

_precisions

sur la carte

_g6n6tique

du

syst6me

de _groupes

_sanguins

B des bovins. Genet Sel

_{Evol 15,}

45-54

Hartl GB

₍₁₉₉₀₎

Genetische Variabilitat beim Muflon

_(Ovis

ammon

_musimon).

Z

_Jagdwiss

36,

95-103

Lebreton

JD,

Roux

M,

Bacou

AM,

Banco G

₍₁₉₉₀₎

_Logiciel

Biomeco 3.93. Unit

Biom6trie,

CEFE, CNRS

Nadler

CF, Woolf A,

Harris KE

₍₁₉₇₁₎

The transferrins and

_hemoglobins

of

_{bighorn sheep}

(Ovis canadensis),

dall

_sheep

_{(Ovis dalli)}

and mouflon

_{(Ovis mv,simon).}

Comp

Biochem

Physiol

40, 567-570

Naitana

_S,

Ledda

_S,

Cocco

_E,

Manca L, Masala B

₍₁₉₉₀₎

_{Hemoglobin phenotypes}

of the wild European mouflon

_{sheep living}

on the island of Sardinia. Arcim Genet 21, 65-75

Nguyen

TC

₍₁₉₇₂₎

_{Les groupes}

_sanguins

des ovins. I. Relations entre les groupes

sanguins

des ovins et des bovins. Ann Genet Sel Anim 4, 363-374

Nguyen

TC

₍₁₉₈₆₎

Loci for

_{serologic polymorphisms}

in the

_Ovicaprinae.

In: Standardized Genetic Nomenclature

for

Sheep and Goats 1986

_(Proc

_{COGNOSAG Workshop}

₁₉₈₆₎

Technique

et _{Documentation, Lavoisier, Paris,}

53-56

Nguyen TC,

Ruffet G

₍₁₉₇₅₎

Les groupes

sanguins

des ovins. II. Facteurs

_antig6niques

suppl6mentaires

dans les

_{syst6mes A, B,}

C et M; estimation des

fr6quences ’all6liques’

aux

_{syst6mes A,}

_B,

_C,

_D, M et R dans les races

_{franqaises : Berriclaon-dv.-Cher,}

Ile-de-France et Texel. Ann Genet Sel Anim _7, 145-157

Nguyen TC,

Bunch TD

₍₁₉₈₀₎

Blood _groups and

_{evolutionary relationships}

among

sheep

(Ovis aries),

domestic goat

(Capra hircvs),

aoudad

_{(Ammotragvs lervia)}

and

European

mouflon

_{(Ovis musimon).}

Ann G!n6t Sel Anim 12, 169-180

ONC

₍₁₉₈₅₎

Le mouflon de Corse.

_Supplement

au Bulletin mensuel de 1’ONC 88

P f

effer P

₍₁₉₈₄₎

Le mouflon de Corse

_(Ovis

ammon

_musimon).

In: Atlas des

_mammif!res

sauvages de France, Societe

Fran!aise

pour

l’Étude

et la Protection des

Mammif6res,

Paris, p 226

Poplin

F

₍₁₉₇₉₎

_Origine

du mouflon de Corse dans une nouvelle

_perspective

paleontologi-que, par marronnage. Ann Genet Sel Anim 11, 133-143

Randi

_{E, Apollonio}

M

₍₁₉₈₈₎

Low biochemical

variability

in

_European

fallow deer

_(Dama

dama

_L):

natural bottlenecks and the effects of domestication.

Heredity

61, 405-410

Randi

_E,

Fusco G, Lorenzini _R, Toso

_S,

Tosi G

₍₁₉₉₁₎

Allozyme divergence

and

phylo-genetic relationships

among

Capra,

Ovis and

_Rupicapra

_{(Artiodactyla, Bovidae)}

Hered-ity 67,

281-286

Roux M

₍₁₉₈₅₎

_Algorithmes

de

classification.

Masson, Paris

She JX, Autem M, Kotulas G, Pasteur N, Bonhomme F

₍₁₉₈₇₎

Multivariate

_analysis

of

genetic exchanges

between Solea _aegyptiaca and Solea

_senegalensis

_{(Teleosts, Soleidae).}

Biol J Lin Soc 32, 357-371

Stratil

A,

Bobak P

₍₁₉₈₈₎

_Comparison

of biochemical

_{polymorphisms}

in mouflon and

sheep:

isoelectric differences in

_haemoglobins

and quantitative variation of mouflon

Stuwe

_M,

Scribner KT

₍₁₉₈₉₎

Low _genetic

_variability

in reintroduced

_alpine

ibex

_(Capra

ibex

_ibex)

_populations.

J Mamm 70, 370-373

Tomiczek H

₍₁₉₈₅₎

The muflon

_(Ovis

ammon musimon

_Schreber,

₁₇₈₂₎

in the southern and western countries of

_Europe.

In: Northern Wild

_Sheep

and Goat _Council,

_Special

Report,

127-132

Uloth W, Prien S

₍₁₉₈₅₎

The

_history

of introductions of mouflon sheep

(Ovis

ammon

musimon

Schreber,

1782)

in central and eastern Europe, and the

_development

and

management of these wild

_{sheep populations.}

In: Northern Wild

_Sheep

and Goat

Council, Special Report,

133-137

Vigne

JD

₍₁₉₈₃₎

Les mammifères

_{post-glaciaires}

de Corse.

Etude

arch6ozoologique.

Gallia