doi:10.1111/j.1365-2052.2008.01837.

Mitochondrial DNA diversity and origins of South and Central

American goats
M. Amills*, O. Ramı́rez*, A. Tomàs*, B. Badaoui*, J. Marmi†, J. Acosta‡, A. Sànchez* and
J. Capote§
*Departament de Ciència Animal i dels Aliments, Universitat Autònoma de Barcelona, Bellaterra 08193, Spain. †Institut Català de
Paleontologia, Universitat Autònoma de Barcelona, Bellaterra 08193, Spain. ‡Fundación Canaria-Instituto de Investigación y Ciencia de
Puerto del Rosario, Puerto del Rosario 35600, Fuerteventura, Spain. §Instituto Canario de Investigaciones Agrarias, La Laguna 38108,
Tenerife, Spain

Summary We have analysed the genetic diversity of South and Central American (SCA) goats
by partially sequencing the mitochondrial control region of 93 individuals with a wide
geographical distribution. Nucleotide and haplotype diversities reached values of
0.020 ± 0.00081 and 0.963 ± 0.0012 respectively. We have also observed a rather weak
phylogeographic structure, with almost 69% of genetic variation included in the within-
breed variance component. The topology of a median-joining network analysis including 286
European, Iberian, Atlantic and SCA mitochondrial sequences was very complex, with most
of the haplotypes forming part of independent small clusters. SCA sequences showed a
scattered distribution throughout the network, and clustering with Spanish and Portuguese
sequences occurred only occasionally, not allowing the distinguishing of a clear Iberian
signature. Conversely, we found a prominent cluster including Canarian, Chilean, Argen-
tinian and Bolivian mitochondrial haplotypes. This result was independently confirmed by
constructing a Bayesian phylogenetic tree (posterior probability of 0.97). Sharing of mito-
chondrial haplotypes by SCA and Canarian goats suggests that goat populations from the
Atlantic archipelagos, where Spanish and Portuguese ships en route to the New World used to
stow food and supplies, participated in the foundation of SCA caprine breeds.

Keywords goat, migration, mitochondrial, phylogeography, South America.

Fernández et al. 2006). The high versatility, moderate size

Introduction
A considerable effort has been devoted in the last few years
to characterize the genetic diversity of African, Asian and
European goats, with studies performed at both continental
(Luikart et al. 2001; Cañón et al. 2006; Naderi et al. 2007)
and regional (Joshi et al. 2003; Sultana et al. 2003; Azor
et al. 2005; Chen et al. 2005; Pereira et al. 2005; Sardina
et al. 2006) levels. These surveys have revealed the absence
of clear ties between the genetic make-up of goats and
geography, with most genetic variation occurring at the
within-population level. This weak phylogeographic struc-
ture has been attributed to the extensive intercontinental
transportation of goats in ancient times (Luikart et al. 2001;
Address for correspondence
M. Amills, Departament de Ciència Animal i dels Aliments, Universitat
Autònoma de Barcelona, Bellaterra 08193, Spain.
E-mail: marcel.amills@uab.es
Accepted for publication 7 November 2008
and hardy nature of goats made them ideal as a food
resource in the lengthy commercial and exploratory jour-
neys that took place in the Old World a long time ago.
South and Central America (SCA) is one of the few
geographical areas where the genetic diversity of goats has
not been yet properly analysed. In poorly developed rural
areas of SCA, goats are an important meat and milk resource,
with a current census recording 38 million individuals (FAO-
STAT, http://faostat.fao.org). The absence of well-managed
conservation genetics programmes and the uncontrolled
introgression of highly productive foreign breeds seriously
threaten the future of many SCA populations that, through-
out the centuries, have adapted successfully to the harsh
conditions in which they were bred (Pariacote 2006). The
main objective of the present study was to provide a general
perspective of the phylogeographic structure and genetic
variation of SCA goats at a continental level. This would be a
first milestone before regional-specific analyses are con-
ducted. Moreover, we were interested in understanding the

 2009 The Authors, Journal compilation  2009 International Society for Animal Genetics, Animal Genetics, 40, 315–322 315
316 Amills et al.
migratory processes under which these SCA populations were
formed and evolved. Upon the conquest and colonization of
the New World by the Spanish and the Portuguese, goats and
other livestock were massively transported through the
Atlantic Ocean to be used as a food supply in exploratory and
military expeditions (Rodero et al. 1992). There are abundant
historical records demonstrating beyond any doubt the
essential role that Andalusian and Lusitanic breeds played in
this process, with most of the transatlantic exploratory jour-
neys in the 15th century departing from a coastal area that
goes from the San Vicente Cape to the Bay of Cádiz (Rodero
et al. 1992). In contrast, the magnitude of the contribution of
the Atlantic archipelagos, which were invariably ports of call
for ships and galleys en route to SCA, to the formation of
American creole breeds has been scarcely documented and it
is still a matter of debate.
Materials and methods
Sample collection and partial sequencing of the
goat mitochondrial control region
Genomic DNA was extracted from either hair or blood
samples corresponding to 152 goats from the South of Spain
(Malagueña, Murciano-Granadina breeds, n = 22), Portu-
gal (Bravia breed, n = 9), Cape Verde (n = 28), Cuba
(n = 25), México (n = 22), Brazil (n = 12), Venezuela
(n = 4), Argentina (n = 9), Chile (n = 10), United States
(Spanish goat, a breed of Mexican ancestry, n = 6) and
Bolivia (n = 5). Specimens were collected by local personnel
with expertise in zootechny in order to ensure that they
correspond to indigenous populations without introgression
with foreign commercial breeds. Hair DNA extraction was
performed with the DNeasy Blood & Tissue Kit (Qiagen)
according to the manufacturerÕs instructions, with the
exception of the lysis method, which instead involved an
incubation in 350 ll lysis buffer (100 mM Tris–Cl pH 8,
100 mM NaCl, 3 mM CaCl2, 2% SDS, 40 mM dithiothreitol
and 250 lg/ml proteinase K) at 56 C for 3–4 h. The pro-
tocol for DNA isolation from peripheral blood mononuclear
cells has been previously reported (Amills et al. 2004).
Purified genomic DNA was used as a template to amplify
and sequence 653 bp of the mitochondrial control region
with a previously described technique (Amills et al. 2004).
Sequences corresponding to different mitochondrial haplo-
types were submitted to GenBank (GenBank accession
numbers: EF372411–EF372515). The basic local alignment
search tool (BLASTN, http://blast.ncbi.nlm.nih.gov/Blast.cgi)
with default parameters was employed to perform compar-
ative searches throughout the GenBank nucleotide collec-
tion database. Other mitochondrial control region
sequences from Spanish (Malagueña, Murciano Granadina
and Guadarrama, n = 8) and Canarian (Palmera, Majorera
and Tinerfeña, n = 20) breeds characterized in a previous
study (Amills et al. 2004) were also incorporated to our
 2009 The Authors, Journal compilation  2009 International Society for Animal Genetics, Animal Genetics, 40, 315–322
analysis. The comparison of the genetic diversity of the
American, Iberian and European goat breeds was achieved by
using the aforementioned dataset and a pool of 106 sequences
extracted from the GenBank. This additional dataset included
sequences from Swiss (Chamois, Verzaska, Saanen and
Toggenburg, n = 35), Portuguese (Serpentina, Serrana,
Algarvia and Charnequeira, n = 41), Italian (Girgentana,
Maltese and Derivata di Siria, n = 21), Irish (Irish Island,
n = 4), Canarian (n = 4) and Danish (n = 1) breeds (Luikart
et al. 2001; Pereira et al. 2005; Sardina et al. 2006).
Phylogenetic and genetic diversity analyses
Phylogenetic and genetic diversity analyses were performed
using a 463 bp region shared by mitochondrial control
sequences obtained by us and others. Nucleotide diversity
(p), Fu and Li statistics (F) and haplotype diversity were
computed with the DNASP 4.00 software (Rozas et al. 2003),
whereas the significance of F was calculated with a coa-
lescent algorithm implemented in this program. The spatial
analysis of the molecular variance was performed with the
SAMOVA software (Dupanloup et al. 2002). The median-
joining network was constructed with the NETWORK 4.2
program (Bandelt et al. 2000). Positions were weighted
according to their mutational frequency and the relative
weight of transversions vs. transitions was set to 3
according to Bandelt (2007). Bayesian phylogenetic trees
were calculated with MRBAYES V3.1.2 (Ronquist & Huelsen-
beck 2003). In this Bayesian analysis, we employed a
Hasegawa, Kishino & Yano (1985) evolutionary model
previously selected with MODELTEST (Posada & Crandall
1998). Evolutionary parameters (proportion of invariant
sites, a-value and transition to transversion rates) were
estimated with both MODELTEST and MRBAYES. After 2.5 mil-
lion iterations, the initial 5000 trees were discarded and the
remaining 20 000 ones were employed to construct a
consensus tree that was visualized with TREEVIEW (Page
1996). A neighbour-joining tree and a multidimensional
scaling plot based on Da distances (number of net nucleotide
substitutions per site between populations) amongst popu-
lations were constructed with MEGA 3.1 and XLSTAT softwares
respectively. Da distances were calculated with the DNASP 4.00
software (Rozas et al. 2003) as follows:
Da ¼ DXY  ðDX þ DY Þ=2
where DXY is the average distance between populations X
and Y, and DX and DY are the mean within-population
distances (Nei & Miller 1990).
Results
Genetic diversity of SCA goats
Sequencing of the mitochondrial control region of SCA
goats (n = 93) with a wide geographical distribution
 Atlantic origin of American goats 317

Table 1 Genetic diversity at the mitochondrial control region (463 bp) Table 2 Spatial analysis of the molecular variance of the mitochondrial
of South and Central American (SCA), Atlantic, Iberian and European control region (463 bp) in South and Central American (SCA) goat
(non-Iberian) goats. breeds.

Haplotype No.
Population N1 p2 Nh3 diversity groups Populations r2WB r2AG r2ABWG

SCA 93 20.09 (0.81) 54 0.963 (0.012)
Atlantic 52 (24) 19.53 (1.07) 28 0.965 (0.011)
Iberian 80 (49) 22.23 (0.73) 71 0.996 (0.003)
European4 61 (61) 19.02 (0.92) 53 0.994 (0.005)
1
N: Number of sampled individuals (sequences extracted from GenBank
are shown between parentheses).
2
Nucleotide diversity · 103.
3
Nh: Number of haplotypes.
4
Pool of Italian, French, Swiss, Irish and Danish breeds.
revealed the existence of 54 different haplotypes, all of them
belonging to the A lineage, which is the most frequent all
over the world (Table 1). Measurement of nucleotide and
haplotype diversities in our SCA continental population
yielded values of 0.020 ± 0.00081 and 0.963 ± 0.012
respectively (Table 1). The sample size employed (almost
100 individuals) and the wide geographical distribution of
the populations studied make this analysis fairly represen-
tative of the existing genetic variation at a continental level.
These levels of nucleotide diversity are notably similar to the
ones observed in the Atlantic, Iberian and European popu-
lations (Table 1). However, if we consider the ratio between
the number of different haplotypes and the total number of
analysed sequences (54/93 = 0.58 for SCA; 28/52 = 0.54
for Atlantic; 71/80 = 0.89 for Iberian; 53/61 = 0.87 for
European population), it is clear that SCA and Atlantic
populations have a relatively smaller proportion of different
haplotypes than Iberian and European populations. A
regional analysis of genetic variability is presented in
Table S1. These results should be taken with caution given
the small sample size of several of the analysed populations,
but they suggest that genetic diversity is particularly
reduced in Argentinian and Chilean goats.
Population structure
The Fu and Li F-statistic corresponding to the analysis
of SCA mitochondrial sequences yielded a negative and
non-significant value of )1.578 (P = 0.071). Analysis of
geographical structure using SAMOVA 1.0 (Dupanloup et al.
2002), a software based on a simulated annealing algo-
rithm that maximizes the proportion of total genetic vari-
ance due to differences between groups of populations,
demonstrated that most of genetic variation occurs within
breeds (Table 2). When the number of groups was set to
two, the spatial analysis of the molecular variance (SAMOVA)
defined the existence of two groups, 1 (Argentina and Chile)
and 2 (remaining populations) with almost 69% of the
genetic variation occurring within breeds (variance
2 Argentina, Chile 68.85% 19.90% 11.25%
Remaining SCA populations
3 Argentina, Chile, Bolivia 73.54% 24.62% 1.84%
México
Remaining SCA populations
r2WB, variance within breeds; r2AG, variance amongst groups; r2AGWB,
variance amongst groups and within breeds.
components are shown in Table 2). When the number of
groups was set to three, we observed the following groups:
(i) Chile, Argentina and Bolivia, (ii) México and (iii)
remaining SCA populations.
Relationships between SCA goats and other Iberian,
Atlantic and European populations
A median-joining network-based phylogenetic analysis
involving 286 American, Iberian, Atlantic and European
goat mitochondrial sequences is shown in Fig. 1. In general,
SCA sequences showed a scattered distribution throughout
the network, with a few clusters encompassing individuals
with the same geographic origin. This diffuse topology,
which is rather complex because a significant amount
of haplotypes are unique or are shared by a small number of
individuals, has been observed in previous works dealing
with Portuguese (Pereira et al. 2005) and Chinese (Chen
et al. 2005) goats. Occasional clustering between SCA
sequences and their European, Iberian and Cape Verdean
counterparts was observed, but it was difficult to identify a
clear relationship between these three populations and SCA
goats because of their scattered distribution. The only
exception to this general trend was a cluster of considerable
size that included Canarian, Chilean, Argentinian and
Bolivian mitochondrial haplotypes (Fig. 1 and Fig. S1). The
existence of this cluster, which was also closely connected to
Cuban, Mexican and Cape Verdean haplotypes (Fig. S1),
might suggest that Canarian goats contributed to the
foundation of the genetic pool of SCA breeds.
With the aim of confirming the results obtained with the
median-joining network approach and inferring their
statistical significance, we performed an independent phy-
logenetic analysis. The 286 sequences employed in the
network analysis corresponded to 195 different haplotypes
that were subsequently analysed by means of a Bayesian
tree (Fig. 2). This phylogenetic tree reliably recovered the
aforementioned Canarian cluster with a highly significant
posterior probability (97%). Moreover, and as previously
observed, clustering of SCA sequences with their Iberian,
Cape Verdean and European counterparts was occasional

 2009 The Authors, Journal compilation  2009 International Society for Animal Genetics, Animal Genetics, 40, 315–322
318 Amills et al.

South American
Cape Verde
Canary Islands
Iberian
European pool

Figure 1 Median-joining network analysis of 286 American, Iberian, Atlantic (Canarian and Cape Verdean) and European goat mitochondrial control
region sequences. The size of the circles is proportional to haplotype frequencies. Transversions were weighted three times as high as transitions
according to Bandelt (2007), and sites were weighted according to their mutational frequencies. The MP option was employed to delete all superfluous
median vectors and links that were not contained in the shortest trees of the network. This figure is in colour in the online version of the paper.

and, with a few exceptions, individuals with the same
geographical origin did not group together, confirming the
lack of a well-defined phylogeographic structure. A third
analysis exploring genetic relationships amongst goat pop-
ulations was carried out by constructing a multidimen-
sional scaling plot (Fig. 3a) and a neighbour-joining tree
(Fig. 3b), where the number of net nucleotide substitutions
per site between populations (Da) was used as a distance
measurement. This analysis highlighted the existence
of three main groups: one including Canarian, Bolivian,
Argentinian and Chilean goats; a second involving Euro-
pean, Iberian and Mexican goats; and a third encompassing
the remaining populations (Fig. 3a,b).
Discussion
The current gene pool of SCA goats was founded in the last
five centuries as a result of the introduction of this species to
America by the Spanish and Portuguese colonizers. Our
main goal was to acquire a general perspective of the final
consequences of this process by exploring mitochondrial
genetic variation of caprine breeds with a large geographi-
cal distribution in SCA. We can conclude from this survey
that all the SCA mitochondrial control region sequences
belonged to lineage A. Our inability to detect mitochondrial
lineages B, C, D, F and G in SCA goats is not surprising given
that these lineages are also very rare, or even absent, in
Europe (Luikart et al. 2001; Joshi et al. 2003; Naderi et al.
2007). More specifically, in the Iberian Peninsula, lineage C
has a very low frequency (<5%) and lineages B, D, F and G
seem to be completely absent (Amills et al. 2004; Azor et al.
2005; Pereira et al. 2005; Naderi et al. 2007). According to
our data, in the Atlantic archipelagos, the situation is rather
similar, with an absolute predominance of lineage
A. Nucleotide diversities in Atlantic and SCA goats were
similar to the ones characterized in Iberian and European
breeds, but haplotype diversities were somewhat lower and
the ratios between the number of different haplotypes and
the total number of sequences were much lower. Although
data presented in Table S1 should be interpreted with
caution, because sample size for several populations is quite
limited, this regional analysis of genetic diversity suggests
that nucleotide and haplotype diversities are particularly
reduced in two Andean populations located in Chile and
Argentina. Our interpretation is that these two populations
descended from Canarian goats, which show a decreased

 2009 The Authors, Journal compilation  2009 International Society for Animal Genetics, Animal Genetics, 40, 315–322
 Atlantic origin of American goats 319

FALCO1
100
95

97

92
93

98
91

96

100

100

100
100
96
100

South American 96
100 100
97
Cape Verde
100
96
Figure 2 Bayesian phylogenetic tree displaying Canary Islands
the genetic relationships between 195
American, European, Atlantic (Canarian and Iberian 100

Cape Verdean) and Iberian goat mitochondrial

control region haplotypes. Posterior probabil- European pool 96
99

ities higher than 90% are shown. A sequence 100

corresponding to Capra falconeri (Falco 1, 100

GenBank accession number: AB044306) was

used as an outgroup. A cluster containing 21 100
Canarian, Cape Verdean and SCA mitochon-
drial haplotypes (Canarian cluster, posterior 91

97
probability: 97) is indicated with a red arrow.
98
This figure is in colour in the online version of
100
the paper.

genetic diversity when compared, for instance, to Cape
Verdean goats. The most likely explanation for this low
genetic variability of Canarian goats is that an important
founder effect took place 2200 years ago, when the first
inhabitants of the Canary Islands settled the archipelago,
carrying with them a small number of domestic animals
(Capote et al. 2004).
The SAMOVA analysis of SCA sequences showed that
around 69% of the genetic variation corresponds to the
within-population component, meaning that creole breeds
are poorly differentiated at the genetic level. This low phy-
logeographic structure observed in SCA breeds is a recurring
theme when analysing mitochondrial variation of caprine
populations. The AMOVA analysis of 406 European, African
and Asian goat sequences revealed that 78.7% of
mitochondrial variation corresponded to the within-breed
component (Luikart et al. 2001). Naderi et al. (2007), in an
analysis involving 2430 individuals from all over the world,
also reported that 77% of mitochondrial DNA variation is
distributed within breeds. Similarly, in Indian goats, 83% of
the total molecular variance was included in the within-
breed component (Joshi et al. 2003). As a reference, in cattle,
the within-breed and amongst-group components include
45% and 51% of the total variation respectively (Luikart
et al. 2001). The weak structuring of goat breeds is one of
the main pieces of evidence supporting the utilization of this
domestic species as a portable food resource accompanying
human migratory movements from time immemorial.
There is compelling and abundant historical evidence
supporting the idea that Iberian livestock was extensively
transported from the South of Spain and Portugal to
America (Rodero et al. 1992). In Spain, Sevilla, Cádiz, San-

 2009 The Authors, Journal compilation  2009 International Society for Animal Genetics, Animal Genetics, 40, 315–322
320 Amills et al.

lations had a poor phylogeographic structure at the time of

(a)
the American colonization, and conceivably much earlier
0.003
Argentina (Fernández et al. 2006), and for this reason they did not
Chile leave an identifiable genetic footprint that can be recognized
0.002
Mexico now. This lack of an identifiable Iberian genetic signature
Portugal 0.001 Canary Islands contrasts very markedly with the detectable Canarian
Europe Bolivia
Spain footprint found in the genetic background of American
0
–0.003 –0.002 –0.001 0.001 0.002 0.003 0.004 0.005 goats. In this study, median-joining network and Bayesian
0
CV phylogenetic analyses consistently identified a cluster
Venezuela –0.001
including Canarian, Argentinian, Chilean, Bolivian, Mexi-
USA
–0.002 can, Cuban and Cape Verdean mitochondrial sequences
Brazil
(Figs 1 & 2 and Fig. S1). This cluster was supported when
Cuba –0.003 performing a multidimensional scaling plot and a neigh-
bour-joining tree based on Da distances (Fig. 3a,b). We
believe that sequences included in this cluster have a
Cuba Canarian rather than European origin for several reasons:
(b)
Brazil
(i) They do not cluster with any of the 141 Spanish, Por-
United States of America (Spanish breed)
tuguese and European sequences employed in the current
study; (ii) The BLASTN analysis of these sequences did not
Venezuela
allow the identification of any 100% match with the hun-
Cape Verde
dreds of African, Asian and European goat mitochondrial
Mexico
sequences stored in the GenBank database; (iii) The Canary
Spain
Islands had a goat population of considerable size and with
Europe
a likely North African origin, which remained geographi-
Portugal
cally isolated for 1700 years until Spanish colonization
Bolivia
(Capote et al. 2004); (iv) These sequences correspond to a
Canary Islands mitochondrial haplotype that has a frequency of
Chile approximately >90% in the La Palma and Tenerife islands
Argentina (Amills et al. 2004). Moreover, this relatedness between
American and Canarian goats has also been shown at the
0.0005 morphological level by Capote et al. (2004). In summary,
Figure 3 Multidimensional scaling plot (a) and neighbour-joining tree although we cannot completely rule out the possibility that
(b) displaying the genetic relationships amongst South American, these American mitochondrial haplotypes have an Iberian
Iberian, European and Atlantic (Canarian and Cape Verdean) goat or European origin, the scenario of a Canarian introduction
populations based on the calculation of Da distances (number of net to SCA seems much more plausible from a historical and
nucleotide substitutions per site between populations). CV, Cape
Verde, USA, United States of America (Spanish breed).
genetic perspective. If this is correct, Canarian haplotypes
might have reached a considerable geographical distribu-
tion (México, Cuba, Argentina, Chile and Bolivia) in the
lúcar de Barrameda and other Andalusian ports participated American continent. This would mean that either the
very actively in commerce with the American continent transportation of goats from the Canary Islands to SCA had
(Rodero et al. 1992). Conversely, the magnitude of the been performed at a considerable scale or, conversely, that
contribution of the Atlantic archipelagos to this large-scale Canarian goats, after one or few introduction events, rapidly
process of livestock exportation to America has been less disseminated in SCA. We favour a scenario with multiple
well-studied. Although it is clear that most of the ships and introductions because there is compelling historical evi-
galleys en route to the New World stopped in the Canary dence indicating that all Spanish expeditions en route to the
Islands, Cape Verde and other nearby insular territories to New World transported livestock that generally came either
take food and supplies, the impact of this marine practice on from Andalusia or the Canary Islands (Rodero et al. 1992).
the formation of creole breeds still remains to be evaluated. It is well known, for instance, that Columbus in his second
With the aim of analysing this issue from a genetic per- trip took goats, calves, sheep and pigs from the Canarian
spective, we have dissected the relationships between SCA island of La Gomera and shipped them to the Antilles
goats and caprine breeds from Spain, Portugal, Cape Verde, (Rodero et al. 1992). Goats were also transported in the
Canary Islands and several European countries. third and fourth trips of Columbus as well as in the expe-
It is worth mentioning that we were unable to find a ditions of Pizarro and Almagro (1530) to Perú and of Ñuflo
distinctive Iberian genetic signature in SCA goats. This de Chávez (1548) to Argentina (Rodero et al. 1992). The
finding might be explained by the fact that Iberian popu- relatively small size of the Spanish galleys combined with

 2009 The Authors, Journal compilation  2009 International Society for Animal Genetics, Animal Genetics, 40, 315–322

the fact that Spanish expeditions targeted many different
locations in the vast SCA subcontinent probably made it
necessary to repeatedly transport livestock, either from the
Iberian Peninsula or the Atlantic islands, in order to be able
to sustain the ever-increasing population of Spanish colo-
nizers. According to Rodero et al. (1992), goats had an
excellent adaptation to both hot and cold areas of SCA,
disseminating rapidly throughout the subcontinent. Finally,
it is worth mentioning that the presence of a Canarian
haplotype in Cape Verde is consistent with historical records
indicating that, in the 15th century, Portuguese sailors
brought livestock including goats from Portugal, Africa and
the Canary Islands to populate these uninhabited archipel-
agos (Rodero et al. 1992).
The genetic wealth of caprine SCA breeds has remained
uncharacterized to date, in spite of its potential as a pow-
erful tool in conservation and selection programmes. In
this study, we have analysed the genetic diversity of several
SCA goat populations and we have found evidence sug-
gesting that Atlantic goat populations, localized in the
West Indies route, participated in the foundation of Ar-
gentinian and Chilean breeds. However, there are several
countries such as Colombia and Perú with large popula-
tions of caprines, which have not been sampled in our
study. A pending issue would be to characterize the genetic
diversity of South American creole breeds by analysing
thousands of individuals with a wide geographical distri-
bution with the aid of a combination of nuclear and
mitochondrial markers. Additional efforts to understand
how this American genetic patrimony was built up and
shaped, combined with the genetic characterization of local
breeds, will play a fundamental role in the preservation of
a legacy with deep roots in the Iberian Peninsula and
Atlantic archipelagos.
Acknowledgements
We are grateful to Jean-Marie Luginbuhl, Liliana Allegretti,
Alfredo Teixeira, Vladimir Ribera, Norge Fonseca, José A.
Atto Mendives, Angelika Stemmer, Noemı́ Castro, Afonso
Semedo, Morse B. Solomon, Hector Andrade, Jorge Olmos,
Emilio Cabrera, Felipe Trujillo and René Garcés for provid-
ing goat samples. Thanks to Jorge Frı́as for his helpful
comments on the manuscript.
References
Amills M., Capote J., Tomas A., Kelly L., Obexer-Ruff G., Angiolillo
A. & Sànchez A. (2004) Strong phylogeographic relationships
among three goat breeds from the Canary Islands. Journal of Dairy
Research 71, 257–62.
Azor P.J., Monteagudo L.V., Luque M., Tejedor M.T., Rodero E.,
Sierra I., Herrera M., Rodero A. & Arruga M.V. (2005) Phylo-
genetic relationships among Spanish goats breeds. Animal
Genetics 36, 423–5.
 2009 The Authors, Journal compilation  2009 International Society for Animal Genetics, Animal Genetics, 40, 315–322
Atlantic origin of American goats 321
Bandelt H.J. (2007) Network 4.2.0.1. User Guide, pp. 3–35. Fluxus
Technology Ltd, Suffolk, UK.
Bandelt H.J., Macaulay V. & Richards M. (2000) Median networks,
speedy construction and greedy reduction, one simulation, and
two case studies from human mtDNA. Molecular Phylogenetics and
Evolution 16, 8–28.
Cañón J., Garcı́a D., Garcı́a-Atance M.A., Obexer-Ruff G., Lenstra
J.A., Ajmone-Marsan P., Dunner S. & Econogene Consortium
(2006) Geographical partitioning of goat diversity in Europe and
the Middle East. Animal Genetics 37, 327–34.
Capote J., Tejera A., Amills M., Argüello A., Fresno M. & López J.L.
(2004) Influencia histórica y actual de los genotipos canarios en
la población caprina americana. AGRI 35, 49–60.
Chen S.Y., Su Y.H., Wu S.F., Sha T. & Zhang Y.P. (2005)
Mitochondrial diversity and phylogeographic structure of
Chinese domestic goats. Molecular Phylogenetics and Evolution
37, 804–14.
Dupanloup I., Schneider S. & Excoffier L. (2002) A simulated
annealing approach to define the genetic structure of popula-
tions. Molecular Ecology 11, 2571–81.
Fernández H., Hughes S., Vigne J.D., Helmer D., Hodgins G., Miquel
C., Hanni C., Luikart G. & Taberlet P. (2006) Divergent mtDNA
lineages of goats in an early Neolithic site, far from the initial
domestication areas. Proceedings of the National Academy of
Sciences of the United States of America 103, 15375–9.
Hasegawa M., Kishino H. & Yano T. (1985) Dating of the human-
ape splitting by a molecular clock of mitochondrial DNA. Journal
of Molecular Evolution 22, 160–74.
Joshi M.B., Rout P.K., Mandal A.K., Tyler-Smith C., Singh I. &
Thangaraj K. (2003) Phylogeography and origin of Indian
domestic goats. Molecular Biology and Evolution 21, 454–62.
Luikart G., Gielly I., Excoffier L., Vigne J.D., Bouvet J. & Taberlet P.
(2001) Multiple maternal origins and weak phylogeographic
structure in domestic goats. Proceedings of the National Academy of
Sciences of the United States of America 98, 5927–32.
Naderi S., Rezaei H.R., Taberlet P., Zundel S., Rafat S.A., Naghash
H.R., El-Barody M.A., Ertugrul O., Pompanon F. & Econogene
Consortium (2007) Large-scale mitochondrial DNA analysis of
the domestic goat reveals six haplogroups with high diversity.
PLoS ONE 2, e1012.
Nei M. & Miller J.C. (1990) A simple method for estimating average
number of nucleotide substitutions within and between popula-
tions from restriction data. Genetics 125, 873–9.
Page R.D. (1996) TREEVIEW, an application to display phylogenetic
trees on personal computers. Computer Applications in the
Biosciences 12, 357–8.
Pariacote F. (2006) The goat in South America and development
perspectives. In: International Symposium Goat Farming Central,
Eastern European countries, present and future. IGA-CAPRIROM.
Constanta, Romania.
Pereira F., Pereira I., van Asch B., Bradley D.G. & Amorim A. (2005)
The mtDNA catalogue of all Portuguese autochthonous goat
(Capra hircus) breeds, high diversity of female lineages at the wes-
tern fringe of European distribution. Molecular Ecology 14, 2313–8.
Posada D. & Crandall K.A. (1998) MODELTEST, testing the model of
DNA substitution. Bioinformatics 14, 817–8.
Rodero A., Delgado J.V. & Rodero E. (1992) Primitive andalusian
livestock, their implications in the discovery of America. Archivos
de Zootecnia 41, 383–400.
322 Amills et al.

Ronquist F. & Huelsenbeck J.P. (2003) MRBAYES 3, Bayesian
phylogenetic inference under mixed models. Bioinformatics 19,
1572–4.
Rozas J., Sanchez-Delbarrio J.C., Messeguer X. & Rozas R. (2003)
DNASP, DNA polymorphism analyses by the coalescent and other
methods. Bioinformatics 19, 2496–7.
Sardina M.T., Ballester M., Marmi J., Finocchiaro R., van Kaam
J.B., Portolano B. & Folch J.M. (2006) Phylogenetic analysis of
Sicilian goats reveals a new mtDNA lineage. Animal Genetics
37, 376–8.
Sultana S., Mannen H. & Tsuji S. (2003) Mitochondrial DNA
diversity of Pakistani goats. Animal Genetics 34, 417–21.
Figure S1 Drawing based on the median-joining network
shown in Fig. 1, displaying each one of the populations
included in or connected with the Canarian cluster. Num-
bers indicate mutated positions.
Table S1 Regional analysis of the genetic diversity at the
mitochondrial control region (463 bp) of South and Central
American, Atlantic, Iberian and European (non-Iberian)
goat populations.
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