Genetic relationships in Spanish dog breeds. I. The analysis of morphological characters

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Genetic relationships in Spanish dog breeds. I. The

analysis of morphological characters

J Jordana, J Piedrafita, A Sanchez

To cite this version:

Original

article

Genetic

_{relationships}

in

_Spanish

dog

breeds.

I. The

_analysis

of

_{morphological}

characters

J Jordana J

_Piedrafita,

A

Sanchez

Universitat Aut6noma de

Barcelona,

Unitat de Genètica i Millora

_Animal,

Departament

de

Patologia

i de Producci6

Animals,

Facultat de

_Veterindria,

08193-Bellaterra, Barcelona, Spain

(Received

27

_{July 1990; accepted}

19 December

₁₉₉₂₎

Summary - The

_{relationships}

between 10

_{Spanish dog}

breeds have been studied using

qualitative

and

_{quantitative analyses}

of data from 32

_{morphological}

characters. The

average distance between

breeds,

measured as a

_{morphological index,}

has a value of _4.228

(! 0.681),

with extreme values of 1.732 between Mastin del Pirineo and Mastin

_Espanol,

and of 5.099 for the Gos d’Atura - Sabueso

Espanol pair.

The

_{morphological phylogeny}

obtained in this

_study

confirms the classifications made

previously by

means of

_dental,

cranial,

historical and behavioral

_comparative

criteria. The results suggest the formation of 2

_{large clusters;}

one formed

_by

the breeds

belonging

to the ancestral trunks Canis

fa7rciliaris

intermedius and Canis

familiaris inostranzewi,

and the other which includes the members of the Canis

familiaris

leineri and Canis

_{familiaris metris-optirrtae}

trunks.

Spanish dog breeds

_/

genetic distance

_/

morphological character

_/

_dendrogram

_/

morphological analysis

Résumé - Relations _génétiques entre des races canines espagnoles. I. Analyse des

caractères morphologiques.

À

partir de

l’analyse qualitative

et quantitative des données

provenant de 3! caractères

morphologiques,

on a étudié les relations existant entre 10 races

canines

_espagnoles.

La distance moyenne entre’races, mesurée par un indice de distance

morphologique, prend

une valeur de _4,228

(::1:: 0,681),

avec des valeurs extrêmes de 1,7.i2

entre Mastin del Pirineo et Mastin

_Espanol,

et

_5,099

pour le

couple

Gos d’Atura - Sabueso

Espanol.

La

_{phylogénie morphologique}

obtenue dans ce

travail, confirme

les

classifications

précédentes,

réalisées à

_partir

de critères

_{comparatifs dentaires, crâniens, historiques}

et _{comportementaux.} Les résultats

_suggèrent

la

_formation

de deux

grands

groupes. L’un

comprend

les races _{qui appartiennent} aux troncs ancestraux du Canis familiaris intermedius

et du Canis familiaris

_{inostranzewi,}

et l’autre serait

_formé

_par les composants des troncs

du Canis familiaris leineri et du Canis familiaris

metris-optimae.

’

INTRODUCTION

Archaeological

studies show the existence of differences within

_populations

of

prehistoric dogs

in the same area. These studies also show that there were

already

distinguishable

and

_separated

classes of

_dogs

about 5 000 years ago

(Villemont

et

al,

1970).

Two main factors have determined the differentiation of canine breeds: natural selection in the environment and conscious selection

_by

man. The

_length

of time from

_prehistoric

times to the

present

and the number of

_generations

_{elapsed explain}

the

_{proliferation}

of canine breeds. Added to this has been the modern

_tendency

of selective

_breeding

to

_{produce specialist}

and

_{distinguishable}

breeds,

with strict definitions of desirable and undesirable traits for each breed.

Man first

_began

to influence the classes of canines when he

_began

to

_adapt

them

to his needs.

Sheep farming,

extensive

_throughout

Eurasia,

created the need for

gentle, intelligent

animals which would

_respond

to orders from the

_shepherd

and

help

manage the flock.

Dogs

were

adapted

for defence: here the desired traits were

fierceness,

toughness

and

_suspicion

of

_strangers.

_Dogs

were also used for

hunting:

some would have to be very fast to catch their prey, others would track and flush the prey and others would retrieve the dead prey. Each had a

_specialist

task.

Finally,

a

general

category

of

_dogs

served for

_defence,

for company or

_merely

for decoration.

The first known classification of

_dogs

dates from 1486 and is found in the St Albar!s’

Book,

attributed to Juliana

Barnes, prioress

of the convent of

_Sopwell,

England

(Peters, 1969).

But the

_systematic

classification of different

_dog

breeds

began

to have

_greater

_importance

at the end of the 19th

_century

with the creation

of the Kennel Clubs in

_England

and North America.

Despite

the

_huge

difficulties involved in the reconstruction of the

_phylogenies

of the more than 400

_dog

breeds

currently recognized,

the

systematic

classification

into groups, as

closely

related as

_possible,

as well as the search for their

phylogenic

relationships

has been an

uninterrupted

task. There have been studies based on

archaeological findings

(Olsen

and

Olsen,

1977;

Clutton-Brock,

1984),

historical studies

_{(Gomez-Toldra, 1985),}

cranial,

dental and skeletal

_morphology

(Clutton-Brock et

al, 1976;

Wayne,

1986),

comparative

studies of behaviour

_{(Scott, 1968),}

and

_{immunological}

and

_{electrophoretic}

studies of

proteins

and blood enzymes

(Leone

and

_Anthony,

1966;

Tanabe et

_al,

_1974).

Although

part

of the variation observed among

morphological

traits may have

an environmental

_component,

in

general,

the

heritability

values for

morphological

traits are

relatively high.

The differences observed _among breeds therefore should

be

_good

indicators of the

_genetic

_{relationships}

among them.

So

_far,

_however,

no studies have been

_published

on the

_genetic

relationships

between

_{Spanish dog}

breeds from the

_analyses

of

_{morphological}

characters. Since statistical methods and

_computing

_packages

are available to

_perform

such

_analyses

(Felsenstein,

1986; Swofford,

1991),

the

_present

paper is a contribution to the

study

of the

_{genetic relationships}

between

_Spanish

canids from

qualitative

and

MATERIALS AND METHODS Breeds studied

We have studied 9

_{Spanish dog}

breeds

_{recognized by}

the Federation

_Cynologique

Internationale

_(FCI):

Gos

_d’Atura,

Mastin del

_Pirineo,

Mastin

_{Espanol, Perdiguero}

de

_{Burgos, Galgo}

_Espanol,

Sabueso

_Espanol,

Ca de

_Bestiar,

Podenco Ibicenco and

Podenco

Canario,

and a tenth breed not

_yet

_recognized,

Podenco Ib6rico. The

geographical

distribution of the

_original

breeds is shown in

_figure

1. There are several

existing

hypotheses

about their

_origin

_(Jordana

et

_al,

_1990),

which we summarize in the

_following

way:

Gos d’Atura

_{(Catalonian Sheepdog)}

or Perro de Pastor Catalin

Andreu

₍₁₉₈₄₎

_points

out that the Romans took and ancient

_{Shepherd dog}

on their

campaigns,

which could have been the

_Bergamasco.

This

_dog

was

_adapted

to the

different climatic environments and

_types

of

_shepherding,

and was the basis of a

large

number of breeds

_existing

_today

in Central

_Europe.

Gomez-Toldra

₍₁₉₈₅₎

and Delalix

₍₁₉₈₆₎

_agree with the

_opinion

of the Roman

_origin

of the Gos d’Atura

_breed,

Mastin

_Espanol

and,-Mastin

del Pirinea.

_(Spanish

Mastiff and

_Pyrenean

Mastiff)

These are breeds included in the

&dquo;ortognated

moloses&dquo; which seem to descend from the

_legendary

Mastiff of Tibet

_(in

central

_Asia).

These

_dogs

are

_supposed

to have reached

_{Spain by}

2 routes: the Central

_European

route and via the Mediterranean

(Esquir6, 1982).

Perdiguero

de

_Burgos

_{(Burgos Pointer)}

This breed

_{probably originated}

from

_matings

between the Sabueso

_Espafiol

and

the short-coated Pachones from Navarra

_(Sanz

Timón,

1982;

Rousselet-Blanc,

1983;

Gomez-Toldra,

1985;

Delalix,

1986).

These Pachones from

Navarra,

also

called Perros de Punta

_Ib4ricos,

are the ancestors of the current

_English

Pointer

(Rousselet-Blanc,

1983;

Sotillo and

Serrano,

1985).

Sabueso

_Espanol

_(Spanish

_Bloodhound)

Several authors

_(Villemont

et

_al,

_1970;

Gondrexon and

_Browne,

_1982;

Rousselet-Blanc, 1983;

Gomez-Toldra,

1985)

have attributed a Celtic

_origin

to the

Blood-hounds. Most of the

_European

Bloodhound breeds seem to descend from the Saint

Hubert,

a

_{modern-day Belgian}

breed,

the direct descendant of the

Segusius

of the

Celts and the

Gauls,

which the Greek historian Arrian of Nicomedia talks about in

his

_Cinegetics

_(Villemont

et

_al,

_1970;

_{Rousselet-Blanc,}

_1983).

Ca de Bestiar

_(Balearic

_Sheepdog):

also called Perro de Pastor

Mallorquin

and Ca

_Garriguer

The FCI includes this breed in the second group, within the molosoid

breeds,

together

with the Boxer and the

_Dogo

among others. Several authors

(Guasp,

1982;

Sotillo and

_{Serrano, 1985; Delalix,}

₁₉₈₆₎

_{agree that the}

_origin

of this breed seems

to be the result of

_crossing

between Podencos

_Ibicencos,

_Perdigueros

_{(Ca NIe)}

and Mastiffs.

Galgo Espanol

(Spanish

Greyhound)

For some authors

(Villemont

et

al,

1970;

Sotillo and

Serrano,

1985)

the

_English

Greyhound

and the

_{Galgo Espafiol}

are descendants of the Arabian

_{Sloughi, brought}

to

_Europe

via

_{Spain during}

the Moslem invasion. Another

_hypothesis

(Rousselet-Blanc,

1983)

supports

the idea that the

_Galgo

was

_brought

to Western

_Europe

_by

the ancient Celts when

_they

settled down in Gaul.

_{Nevertheless,}

the same author

points

out a second contribution of blood from the

_Sloughi.

Podenco Ibicenco

_{(Ibizan Hound):}

also known as Ca

_Eivissenc,

Xarnelo,

Lebrel de

_{Mallorca, Mallorqui}

or

_Charneque

It is

_{generally accepted}

that the Podenco Ibincenco breed descends from the

_Dog

Rousselet-Blanc,

1983; G6mez-Toldrh,

1985)

and that it was

_brought

to Ibiza

_by

the Phoenicians

_(Pugnetti,

_1981;

_Maza,

_1982;

_Delalix,

_1986),

even

_though

other

hypotheses

state that it arrived much

_later,

with the

_Moslems,

at the same time as the

_Galgo

_(Villemont

et

_al,

_1970;

_{Rousselet-Blanc,}

_1983).

Podenco Canario

_{(Canary Hound)}

Certain

_hypotheses

_{(Delalix, 1986)}

_suppose that this hunter came from

_Egypt

and that it was taken to the

_{Canary Islands, probably by}

the

_Phoenicians,

Greeks, Carthaginians

or even

_by

the

_Egyptians,

but it is

possible

that

Majorcan

monks,

forced to

_emigrate

to these islands

_by

the

_Vatican,

introduced these

_dogs

(Anonymous, 1982).

Podenco Ib6rico

_{(Iberian Hound):}

also known as Podenco

_Espanol,

Podenco

_Andaluz,

Podenco Ib6rico Andaluz

_Malagueno

and

_Campanero

The Podenco Ib6rico is a recent

_product

obtained

_by

_crossing

the Podenco Rondeno

from Andalusia with the Podenco Ibicenco

_(Garcia

et

_al,

_1982).

Qualitative

and

_{quantitative analyses}

In an ideal

_specimen

of each of 10

_{Spanish dog}

_breeds,

a total of 32 characters have

been studied. Some of the characters were established

by

the official standards of the breed while the other characters came from data of a review

(Avila,

1982;

I

Symposium

Nacional de las Razas Caninas

_Espanolas,

_1982;

Gomez-Toldra, 1985 ;

Sotillo and

_{Serrano, 1985;}

_Delalix,

_1986).

The numbers were

_assigned

to each state

of the different characters in an

_arbitrary

manner. These numbers did not

_represent

any

specific weighting

of the state. The number of states for each character was established

_depending

upon the number of

distinguishable phenotypic

classes. The

characters used and their states are shown in table I.

Qualitative analysis

For the

qualitative analysis,

discrete characters were recoded into a series of

(0, 1)

2-state

_characters,

_denoting

absence or _presence of the

character,

respectively.

Continuous

_quantitative

characters

_(D

and E characters in table

_I)

may be divided into a small number of

classes,

each

_representing

one of the states of the character

in the data matrix. For

_recoding

a character with several states we have used the

and so on. The

_original

and recoded matrices of

_{morphological}

resemblances are

shown in tables II and III

_{respectively.}

The MIX _{program of the}

_phylogeny

inference

_package

_{(PHYLIP) (Felsenstein,}

1986)

was used to construct the

_dendogram

of

_Spanish

breeds of

_dogs

from

qualitative

data of

_{morphological}

characters. This

_analysis

is based _{upon the}

&dquo;parsimony&dquo; principle,

and the criterion is to find the tree

_requiring

the minimum

number of

_changes.

Two

_dendrograms

can be obtained: the

_{first, using}

_Wagner

parsimony

(Farris, 1970),

is used when the ancestral state of the character is

unknown;

the

_second,

_using

Camin and Sokal’s method

_(1965),

_presupposes the

knowledge

of the

_{ancestrality.}

Several

_possible

criteria have been

_proposed

to infer the ancestral state of the character: the fossil

record,

the

_frequency

criterion and

outgroup

analysis

(Avise, 1983).

Each of these criteria has been

_seriously

and

justifiably

criticized

_{(Stevens, 1980),}

_although

it has been

_recognized

that the

outgroup

analysis provides

a

_{particulary compelling}

rationale for

estimating

the

character state

_polarity

_(in

our case, for

_example,

the

wolf,

Canis

_lupus).

We have

chosen, however,

the

_frequency

criterion

_(the

state of the character

_appearing

most

frequently

in the _group

_being

_examined)

in order to make

_comparisons

between these

_dendrograms

and those obtained in a second

study

(Jordana

et

_al,

₁₉₉₂₎

on

the

_{phylogenetic relationships}

_among

_{Spanish dog}

breeds derived from the

_analysis

of biochemical

_{polymorphisms.}

The reason for

_choosing

the

_frequency

criterion was

the lack of

_adequate

literature on

electrophoretic

results of _any

_species

of wolf

candidate to be used as an

_outgroup.

The tree

_generated

_by

_Wagner

_parsimony

is

unrooted,

so we chose

_arbitrarily

the

_Galgo

_Espanol

breed as an

_outgroup

in order

to make

_comparisons

with other

_dendrograms.

An

_evolutionary

tree

_{generated by}

a

_parsimony

criterion was also

_computed

using

the

_{phylogenetic analysis}

_{using parsimony}

_computer

_package

_{(PAUP) (Swof}

ford,

1991).

The

_resulting

tree was rooted and the

_midpoint

_rooting

method

_(Farris,

1972)

was chosen to

_give

the tree an

_evolutionary

direction. The PAUP

_package

allows us also to

_compute

the confidence limits of the

_{topology by}

means of a

boot-strap

analysis

(Efron, 1979),

adapted

to the inference of

_phylogenies

_{(Felsenstein,}

1985).

One hundred

_{bootstrap replicates}

were

made,

and a consensus tree was

ob-tained based _upon the

_{majority-rule}

method

_(Margush

and

_McMorris,

_1981).

The minimum

_frequency

of the

_{bootstrap replicates-}

in which a _{group- is-}

_supported

in

Quantitative analysis

For the

_quantitative

_analysis

of

_{morphological characters,}

_qualitative

data were

transformed and introduced in the form of a matrix of distances. An Euclidean

dis-tance

_(Sneath

and

_Sokal,

₁₉₇₃₎

was used to estimate distances between

_populations,

under the

_assumption

of

_independence

between characters.

where:

d!!,!!

= value of the distance between

the j and

the breed k. The distance

(J!,j —

Xi

k

)

= alternative values

(0, 1)

for the differences

between j

and k breeds

within the character i.

The mean character difference

(MCD)

_{proposed by}

Cain and Harrison

(1958)

was also calculated as a measure of taxonomic resemblance. MCD varies between

0 and 1.

Fitch and

_{Margoliash’s}

method

₍₁₉₆₇₎

was used to find the unrooted tree that

would best

_adapt

to the matrix

_(FITCH

_{program in} PHYLIP

_package).

The tree that minimizes the sum of _squares SS was searched for

_by

means of the

_following

expression:

&dquo;

where:

D

jk

= observed distance between

populations j

and

_k;

d

jk

=

expected

distance between

_{populations j}

and

_k,

_computed

as the addition of

tree

_segment

_lengths,

from

_{population j}

_to

_population

k

_{(patristic distance).}

Alternatively,

a rooted tree was

_{computed by applying}

the KITSCH _program

(PHYLIP package).

In this

_method,

a tree similar to that

_{generated by}

the cluster

analysis

was

computed

and

subsequently

the

topology

of the tree was altered in

order to

_improve

its

_{goodness-of-fit. By}

_assuming:

_a),

that the

_expected

rates

of

_change

are constant

_through

all

_lines;

_b),

that all the

_{subpopulations}

are

contemporary;

and

_c),

that the

_phenotypes

behave as an

evolutionary

clock,

this method can be

_regarded

as an estimator of the

_phylogeny

_{(Felsenstein,}

_1984,

_198G).

RESULTS

Qualitative

analysis

The

_{dendrograms resulting}

from the

_application

of

_Wagner

_parsimony

and Camin

and Sokal’s methods are shown in

_figures

2 and 3

_{respectively.}

Two

_large

_groups

can be observed in each tree. One of the _{groups is} formed

_by

4 breeds: Mastin del

Pirineo,

Mastin

Espanol,

Sabueso

Espanol

and

_Perdiguero

de

_Burgos;

the other

group includes Podenco

Ibicenco,

Podenco

Canario,

Podenco Ib6rico and

_Galgo

Espanol.

In the

_{dendrogram resulting}

from

_Wagner

_parsimony,

the breeds Ca de

Bestiar and Gos d’Atura are

halfway

between the 2

large

_groups, even

though

Gos d’Atura is nearer the

greyhound

_group

(Podencos

and

Galgo)

and Ca de Bestiar is

nearer the other _group.

by

Sabueso

_Espafiol

and

_Perdiguero

de

_Burgos.

Both

_topologies

are

possible,

even

though

the tree obtained

_by

_{applying Wagner}

_parsimony

needed

_only

96

_steps

to rearrange the characters and to obtain the most

parsimonious

tree,

while for the tree

_{generated by}

Camin and Sokal’s

_method,

101

steps

were needed. This difference

in the number of

_steps,

_however,

_probably

reflects the differences in the

_assumptions

of the kinds of

_changes

used in both methods

_{(Felsenstein, 1986),}

and

_consequently

cannot be considered as a definitive criterion to infer the true

_{relationships.}

Figure

4 shows a

_dendrogram

of the

_{Spanish dog}

breeds estimated

_according

to the

_parsimony

and

_midpoint

_rooting

criteria

_(PAUP

_package).

This

_dendrogram

again

shows the _{2 groups} described above. Branch and internodal distances are

proportional

to the number of

_{character-stage changes required.}

The total

_length

measure of the

_homoplasy)

was 0.671. Included within

_parentheses

are the values of the number of

_replicates

from the

_{bootstrap analysis}

_(loosely,

the width of the confidence

_interval).

Quantitative analysis

The results of the

_{morphological}

distance indexes between

_{Spanish dog}

breeds are shown in table IV. The _average distance between breeds has a value of 4.228

(f

0.681),

with extreme values of 1.732 between Nlastin del Pirineo and Mastin

Espanol,

and 5.099 for the Gos d’Atura - Sabueso

_Espanol

_pair.

The values of the distances within the Podenco _group

_(Ibicenco,

Canario and

_Ib6rico)

are

_small,

as are the distances between Mastiff breeds

_(Mastin

del Pirineo and Mastin

_Espanol),

and between

_Perdiguero

de

_Burgos

and Sabueso

Espanol.

The values for the mean character differences

_(NICD)

between

_{Spanish dog}

breeds are shown table V. In the same _{way, the average MCD} between breeds has a value of 0.5645

(t

0.1552),

with

extreme values of 0.0937 between Nlastin del Pirineo and Mastfn

_Espanol,

and of

0.8125 for the Gos d’Atura - Sabueso

_Espanol

_pair.

The trees obtained

_using

FITCH and KITSCH programs are shown in

figures

5 and 6. The

_dendrograms

obtained

_by

the FITCH program are

unrooted,

so we

arbitrarily

used the

_{Galgo Espafiol}

breed as an

_outgroup.

Two hundred and

twenty-six

_possible

trees were examined.

_Figure

5 shows the tree that best

_adjusts

to the

matrix of data. The sum of _squares had a value of

_0.183,

whereas the _average

_percent

standard deviation was

4.5G%.

In the tree in

_figure

_5,

the 2 groups

previously

described are

_again

observed. The

_Greyhound

cluster

_(Podenco

Ibicenco,

Podenco

Canario,

Podenco lb6rico and

_Galgo

_{Espanol) additionally}

contains the Gos d’Atura

breed. The Ca de Bestiar breed remains in an intermediate

_position,

_slightly

closer

to the

_Greyhound

_group.

In the

_resulting

_{tree from}

the

_application

of the KITSCH program, the 2

large

the

_greyhound

_group, even

though

an unresolved

trichotomy

is

presented

between

Galgo Espauol,

Gos d’Atura and Ca de Bestiar breeds. The sum of _squares had a

value of 0.248 _{and the average}

_percent

standard deviation was

5.31%.

DISCUSSION

In

_examining

all the

_topologies

of the trees

_resulting

from the

_analysis

of

morpho-logical

characters,

it is

_possible

to

_verify

some stable

_{relationships}

_among different

groups of breeds. Sabueso

Espanol

and

_Perdiguero

de

_Burgos

form a

_separate

clus-ter from Mastin

_Espanol

and Mastin del Pirineo breeds. The last 2

_clusters,

in

their

_turn,

are related and form a new cluster. The

bootstrap analysis

(figure

4)

confirms this

_grouping

_(79%

of the

_bootstrap

_replicates).

Podenco

_Ibicenco,

Po-denco

_Canario,

Podenco Ib6rico and

_{Galgo Espanol}

breeds are related in all trees.

Espafiol

and Podenco breeds

_(PE,

PC and

_PI),

as the value from the

_bootstrap

analysis

was below

50%.

These breeds

_correspond

to the

_Greyhound

group.

The

_relationship

described above is consistent with

_assigning

the

_Spanish

breeds

to the known ancestral trunks. The

_{Spanish dog}

breeds have been

_assigned

to

their

_hypothetical

ancestral trunks

_by

_comparing

their

_morphology

with some

European

breeds whose

_phylogeny

was taken as known

(table VI).

The

phylogeny

of the

_European

breeds was inferred

_by

_comparative

studies of dental and cranial

morphology,

as well as

_{archaeological,}

historical and behavioral studies

(Studer,

1901;

Antonius, 1922;

Villemont et

_al,

_1970;

_{Rousselet-Blanc,}

_1983).

The

_phylogeny

resulting

from the

_qualitative

and

quantitative

analysis

of

_{morphological}

data seems

to confirm these classifications. There is a

disagreement,

however,

with Villemont

et al

_(1970),

who

_assigned

the

_Bloodhound,

_Beagle

and Grand Bleu de

_Gascogne

It can be observed from the tree in

_figure

6 that the cluster formed

_by

the Mastin del Pirineo and Mastin

Espaiiol

breeds would fit in with the ancestral trunk of

_Cf

_{inostranzewi,}

and the cluster that Sabueso

_Espafiol

and

_Perdiguero

de

_Burgos

form will fit with

_Cf

_intermedius,

both

_being

_{related groups and}

_forming

in their

turn a new cluster. On the other

hand,

a close

relationship

is observed between

the 3 breeds of Podencos that would form the trunk of

Cf

leineri.

Galgo Espanol

remains a little farther _away,

_{although taking}

as a basis the trees

_resulting

from the

qualitative analysis

(MIX

program and PAUP

program)

and

quantitative

analysis

(FITCH program),

the breed

might

be included in the trunk of

_Cf

leineri. Gos

d’Atura would be the

only

representative

of

Cf metris-optimae,

and Ca de Bestiar would remain isolated. Due to the

_{particular origin}

of the Ca de Bestiar breed

_(it

is believed that it comes from

_crossings

between

_Podencos,

_Perdigueros

and

Mlastines)

the breed has not been

_assigned

to _any

_specific

ancestral trunk.

According

to Felsenstein

_(1986),

the resultant tree could be considered as an

estimation of the

_phylogeny

of the

_breeds,

which would

_suggest

that the Sabueso

Espafiol, Perdiguero

de

_Burgos,

Mastfn del Pirineo and Mastin

_Espauol

breeds would be related and would

descend

fiom a

hypothetical

common ancestor. On the

other

hand,

Gos d’Atura

_{( Cf}

_{metris-optimae)}

and Ca de Bestiar would be more

related to the members of the

_Cf

leineri. A common ancestor

_might

be

_postulated

for

the

_Cf

_{m.etris-optim.ae}

and

_Cf

leineri trunks.

_Figure

7 summarizes the

_hypothetical

relationships

between ancestral trunks described above.

The methods

_applied

in this

study

were devised

_mainly

to

_analyze

natural popu-lations. This _paper deals with

_populations

of domestic animals whose

characteristics

were fixed

by

man in a _process of artificial

_selection,

assumed to be very intensive

been _very

_{complex, including}

both characters related to some

specific ability

and other traits derived from the

_caprice

of the breeders.

Nevertheless,

we think that selection is the

_{evolutionary strength}

that could have had the

_greatest

_weight

in the process of breed differentiation. In most

_species

of domestic

_animals,

and in a

spe-cial manner in the canine

_species,

the characteristics that

_usually

define a breed are

basically morphological.

The

breed, consciously

or

unconsciously,

has been created

by

man, even

though

the contribution of the environment has

operated through

nat-ural selection. Orozco’s words

₍₁₉₈₅₎

about the breed

_concept

in domestic animals

are illustrative:

&dquo;Nobody

can

_stop

a

breeder,

a

technician,

or anyone who has access to a group

of

_animals,

from

_establishing

a

particular population

as a

breed,

if he bases this on

_fixed,

_objective,

uniform and different characteristics from other breeds. He

can

_speak,

if he wants to, about a new breed. The breed has

_simply

to _{agree with}

definitive and _very strict characteristics:

_perfection

of

_colour,

_type,

_appearance, well determined measures of different

_parts

of the

_body,

etc. If the breed is

established in this manner, there is no

_objection

to make&dquo;.

This assertion

_acquires

_{great importance}

in the case of

dog

breeds.

Here,

the

patterns

or

_prototypes

for the inclusion of an animal in a

_particular

breed are

very

strict,

resting

on

multiple morphological

_assessments,

both

_qualitative

and

quantitative,

that should be within certain limits. If the

_qualifiers

consider that an

animal does not achieve the proper

requirements,

nobody

doubts that this animal

does not

_belong

to the breed. This consideration should

_preclude

for most breeds

inter-racial

_crossings

that would have resulted in a less tree-like

genealogy.

In an

_ecological

_context,

_Crouau-Roy

₍₁₉₉₀₎

affirms that

_{morphological}

data _may reflect historical _processes but are much more under the influence of differential selective pressures

(micro-

and macro-environmental

influences)

than biochemical data. This affirmation

_might

be also

_applicable

to the case of the evolution of

canine breeds. In this sense, it has also been

argued

that the

study

of the values of

enzymes, without any relation with

fitness,

ie assumed as neutral _genes, would be a

good

indicator of the

_genetic

_similarity

or

_divergence

between

_populations

_(Kimura,

1983).

This kind of

_analysis

_may allow us to

_study

whether there is an

evolutionary

parallelism

between both

_types

of characters -

_{morphological}

and blood substances

- with would be of

great

interest in

_establishing

more

_accurately

the

_{relationships}

between the canine breeds under

_study.

ACKNOWLEDGMENTS

The suggestions of two referees are

_{greatly appreciated.}

We thank J Torrent and W Kelson for assistance with the

_preparation

of this

_{manuscript. Finally,}

the authors thank Gallina Blanca

_Purina,

for its initial contribution to the

_funding

of this

study.

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