*Corresponding author: gilberto.lemus@uan.edu.mx

Keywords:

Genetics

PCR

Local breeds

Haplotypes

Linage

Phylogenetic relationships of Yucatan hairless pig with Asian and European breeds by

mitochondrial DNA D-loop

Relaciones logenéticas del cerdo pelón de Yucatán con razas asiáticas y europeas mediante el ADN

mitocondrial

Relações logenéticas do porco sem pelo de Yucatán com raças asiáticas e europeias através do

DNA mitocondrial

Clemente Lemus Flores

Gilberto Lemus Avalos

Job Oswaldo Bugarín Prado

Rogelio A. Alonso Morales

Raúl Ulloa Arvizu

José Candelario Segura Correa

Miguel Ángel Ayala Valdovinos

Raúl Sansor Nah

Rev. Fac. Agron. (LUZ). 2025, 42(4): e254251

ISSN 2477-9407

DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v42.n4.VIII

Animal production

Associate editor: Dra. Rosa Razz

University of Zulia, Faculty of Agronomy

Bolivarian Republic of Venezuela

Postgraduate Program in Biological and Agricultural

Sciences. Academic Unit of Veterinary Medicine and

Zootechnics and Academic Unit of Agriculture, Autonomous

University of Nayarit, Mexico.

Molecular Genetics Laboratory, Department of Genetics

and Biostatistics. Faculty of Veterinary Medicine and

Zootechnics, National Autonomous University of Mexico.

Campus of Biological and Agricultural Sciences,

Autonomous University of Yucatan, Yucatan, Mexico.

Department of Animal Production, University of

Guadalajara, Mexico.

Mexican Association of Iberian Pig Breeders, Yucatan,

A.C., Mexico.

Received: 08-08-2025

Accepted: 22-10-2025

Published: 04-11-2025

Abstract

In Mexico native pig genotypes exist whose populations face

serious threats to survival. One of them is the Yucatan hairless pig

(YUCMEX), for which limited information is available regarding

its current conservation status. This study aimed to determine

the phylogenetic relationship of YUCMEX with Iberian, wild

pigs (WB), European, Asian, and commercial pigs using the

mtDNA D-loop region. A total of 31 YUCMEX sequences and 77

mitochondrial haplotypes from GenBank were analyzed, aligned to

reference sequence AJ002189. The study fragment, trimmed between

positions 15435 and 15977, resulted in 543 base pairs. Genetic

distances were calculated to compare YUCMEX with the other pig

groups. Phylogenetic trees were constructed using the Neighbor-

Joining method with Kimura’s two-parameter distance and 1000

bootstrap replicates. Additionally, a principal component analysis

(PCA) was performed based on evolutionary distances. Among the

108 sequences analyzed, 41 variable sites and 44 haplotypes were

identied. YUCMEX individuals grouped into four haplogroups (HA,

HB, HC, HD), showing lower D-loop diversity and genetic distance

from the Duroc breed. European and Asian haplotypes formed seven

phylogenetic groups, clearly separating both regions. The YUCMEX

haplogroups clustered into three lineages close to WB from Portugal

and Spain but were distinct from Asian pigs and Eastern European

WB haplotypes. These ndings conrm the European—specically

Iberian—origin of the Yucatan hairless pig.

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Rev. Fac. Agron. (LUZ). 2025, 42(4): e254251 October-December. ISSN 2477-9409.

2-6 |

Resumen

En México existen genotipos de cerdos nativos cuyas poblaciones

enfrentan serias amenazas para su supervivencia. Uno de ellos es

el cerdo pelón de Yucatán (YUCMEX), para el cual se dispone de

información limitada sobre su estado de conservación actual. Este

estudio tuvo como objetivo determinar la relación logenética de

YUCMEX con cerdos ibéricos, jabalíes (WB), europeos, asiáticos y

comerciales utilizando la región D-loop del ADNmt. Se analizaron un

total de 31 secuencias de YUCMEX y 77 haplotipos mitocondriales

de GenBank, alineados a la secuencia de referencia AJ002189. El

fragmento de estudio, recortado entre las posiciones 15435 y 15977,

resultó en 543 pares de bases. Se calcularon distancias genéticas para

comparar YUCMEX con los otros grupos de cerdos. Los árboles

logenéticos se construyeron utilizando el método Neighbor-Joining

con la distancia de dos parámetros de Kimura y 1000 réplicas

bootstrap. Además, se realizó un análisis de componentes principales

(ACP) basado en distancias evolutivas. Entre las 108 secuencias

analizadas, se identicaron 41 sitios variables y 44 haplotipos. Los

individuos YUCMEX se agruparon en cuatro haplogrupos (HA,

HB, HC, HD), mostrando menor diversidad de bucle D y distancia

genética con respecto a la raza Duroc. Los haplotipos europeos y

asiáticos formaron siete grupos logenéticos, separando claramente

ambas regiones. Los haplogrupos YUCMEX se agruparon en tres

linajes cercanos al WB de Portugal y España, pero distintos de los

haplotipos WB de los cerdos asiáticos y de Europa del Este. Estos

hallazgos conrman el origen europeo, especícamente ibérico, del

cerdo pelón de Yucatán.

Palabras clave: genética, PCR, razas locales, haplotipos, linaje.

Resumo

No México existem genótipos nativos de suínos cujas populações

enfrentam sérias ameaças à sua sobrevivência. Um deles é o porco

sem pelo de Yucatán (YUCMEX), para o qual há informações

limitadas disponíveis sobre seu status de conservação atual. Este

estudo teve como objetivo determinar a relação logenética de

YUCMEX com suínos ibéricos, javalis (WB), europeus, asiáticos

e comerciais usando a região D-loop do mtDNA. Um total de 31

sequências de YUCMEX e 77 haplótipos mitocondriais do GenBank

foram analisados, alinhados à sequência de referência AJ002189.

O fragmento de estudo, aparado entre as posições 15435 e 15977,

resultou em 543 pares de bases. As distâncias genéticas foram

calculadas para comparar YUCMEX com outros grupos de suínos.

Árvores logenéticas foram construídas usando o método Neighbor-

Joining com a distância de dois parâmetros de Kimura e 1000 réplicas

bootstrap. Além disso, uma análise de componentes principais (ACP)

baseada em distâncias evolutivas foi realizada. Entre as 108 sequências

analisadas, 41 sítios variáveis e 44 haplótipos foram identicados.

Indivíduos YUCMEX agruparam-se em quatro haplótipos (HA,

HB, HC, HD), mostrando menor diversidade de D-loop e distância

genética em relação à raça Duroc. Haplótipos europeus e asiáticos

formaram sete grupos logenéticos, separando claramente ambas

as regiões. Haplótipos YUCMEX agruparam-se em três linhagens

próximas aos haplótipos WB de Portugal e Espanha, mas distintas

dos haplótipos WB de suínos asiáticos e do Leste Europeu. Essas

descobertas conrmam a origem europeia, especicamente ibérica,

do porco sem pelo de Yucatán.

Palavras-chave: genética, PCR, raças locais, haplótipos, linhagem.

Introduction

There is wide evidence, using molecular studies, that the wild

pigs (Sus scrofa) distributed throughout Europe and Asia separated

phylogeographically, indicating that Central Europe was a center of its

early domestication (Giura et al., 2000; van Asch et al., 2012). The

Iberian haplotypes are close to the European (other than Iberian pigs),

but there is a second European group between Asians and Iberians

from the region of Italy, considering that the divergence between

Europeans and Asians has 600,000 years (Alves et al., 2003). Similar

studies with mitochondrial DNA analysis in wild boars introduced in

the Ural region, indicate that the contribution of lineages originating

in Eastern Europe was greater than expected than the proportions of

European and Asian animals (Markov et al., 2022). In native pigs,

their ancestral origin is presumed and hybridization is assumed, so

knowledge of their phylogeneia is necessary to know their identity.

The study by Banayo et al. (2023) showed that the Philippine native

pigs have originated from at least three Sus scrofa lineage and that they

were not domesticated from the endemic wild pigs of the Philippines,

indicating that the Philippine native pigs had other genetic origins;

further analysis revealed its multiple ancestral origins, from East and

Southeast Asia. The Iberian pigs exported during the colonization of

America were ancestors of several breeds, which contributed to the

origin of the Duroc in the United States and the hairless pig in Mexico

(Jones, 1998; Burgos-Paz et al., 2013; Lemus-Flores et al., 2023).

According to Alves et al. (2003), the characterization of genetic

resources is needed to make clear their relationship and origin with

other pig populations. It is documented that the hairless pig arrived

in America on the second voyage of Christopher Columbus (Ogata,

2019). The entry of the Iberian pig into Mexico was along the Atlantic

coast, reaching the Yucatan Peninsula (Ortega et al., 2019), where it

has remained under rural breeding and management, and local people

have developed a gastronomy based on this pig (Hernández et al.,

2020; Ramos-Canché et al., 2020). Other populations have been

derived from the hairless pig populations, such as the one developed

in Colorado, United States, patented as Yucatan minipig (Burgos-Paz

et al., 2013). There is no information on the ow of hairless pigs to

the Yucatan Peninsula; however, the local culture, family exploitation

system, and demography have allowed the conservation of the

hairless pig (Hernández et al., 2020; Ramos-Canché et al., 2020).

It is known that its phenotype and genetics are similar to Iberian

hairless pigs (Lemus-Flores et al., 2023), and that it shows a wide

genetic diversity (Lemus-Flores et al., 2020). For the conservation

of the hairless pig in Yucatan, it is necessary to know its origin and

genetic place concerning its ancestors. The objective of this study

was to determine the phylogenetic relationship of the Yucatan hairless

pig with the Iberian, wild European, Asian, and commercial pigs.

The information here generated will be useful for the conservation

programs carried out in Mexico, and for the identication of pigs that

will be used for breeding. Another benet is to give added value to

their genetic identity in the promotion of their products in demand

within the tourist gastronomy in the Yucatan Peninsula.

Materials and methods

Sample collection and isolation of mitochondrial DNA

(mtDNA)

This project was registered at the Autonomous University of

Nayarit-México, under number SIP18-076. Following animal

management protocols NOM-051-ZOO-1995 (Secretary of

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Lemus et al. Rev. Fac. Agron. (LUZ). 2025, 42(4): e254251

3-6 |

commercial, European and Asian pigs. Phylogenetic trees were

constructed with neighbor-joining (Saitou and Nei, 1987) incorporated

in the MEGA 11 software (Tamura et al., 2021). The dierences in the

composition bias among sequences were considered in evolutionary

comparisons (Tamura and Kumar, 2002). All positions containing

gaps and missing data were eliminated. Standard errors were

computed with the bootstrap method using 1000 replicates (Tamura

et al., 2021). With the evolutionary distances, principal component

analysis (PCA) was carried out with the Darwin program (Perrier et

al., 2003) and graphed with Minitab (2021).

Results and discussion

The analysis of 108 sequences identied 41 change sites grouped

into 44 haplotypes (table 1). It should be noted that a greater number

of haplotypes than change sites suggests high genetic diversity

(Niedziałkowska et al., 2021).

Agriculture, Livestock and Rural Development, 1998) and NOM-

062-ZOO-1999 (Secretary of Agriculture, Livestock, Rural

Development, Fisheries and Food, 2001), blood samples were

collected from the jugular vein in Vacutainer EDTA K2 tubes (Becton

Dickinson, Franklin Lakes, NJ, USA), from 31 unrelated hairless

females from a group of hairless pig from dierent farms in Yucatan

Mexico (YUCMEX). Total DNA was puried following the protocol

established by Miller et al. (1989) in the Molecular Biotechnology

laboratories of the Faculty of Veterinary Medicine at the National

Autonomous University of Mexico.

PCR amplication and sequencing of mitochondrial DNA

(mtDNA)

Considering the methodology used by van Asch et al. (2012),

a 662 bp mtDNA fragment from the D-loop region was amplied.

The whole D-loop was amplied using a package of PCR reagents,

adding the amounts recommended by the manufacturer (Biogenica,

S.A of C.V), and under the following conditions: 1 cycle at 94 °C (5

min); 30 cycles at 94 °C (1 min), 56 °C (30 sec) and 72 °C (1 min);

and a nal cycle of 72 °C (5 min). mtDNA fragment was amplied

by PCR using the primers SCMtF-CTAACTCCGCCATCAGCAC

y SCMtR-CTGTGTTAGGGCCTTTGACG. Then, the amplied

fragments were puried by the sodium iodide (NAI) plus silica

pearls method and used in sequence reactions carried out with the

commercial sequence kit ABIPRISM BigDye Terminator Cycle

Sequencing v3.1 (Applied Biosystems Division, Perkin-Elmer,

Foster City, CA, USA). The manufacturer’s protocol was followed

under the following conditions: 25 cycles at 96 °C (10 sec), 50 °C

(5 sec), 60 °C (4 min). Then, the samples were ltered in columns

of Sephadex G50 (SIGMA, St. Louis, MO, USA), and reading was

performed with an automatic ABI prism 310 DNA Genetic Analyzer,

(Applied Biosystems Division, Perkin-Elmer, Foster City, CA, USA).

Two readings were obtained for all samples: one for each chain. The

consensus sequences from both readings were obtained with the

CHROMAS v.1.62 software (Technelyuim Pty. Ltd., Queensland,

Australia). The amplied fragment was aligned to the GenBank

AJ002189 sequence as a reference, trimming the studied fragment

from position 15435 to 15977 with 543 bp (Alves et al., 2003).

Mitochondrial DNA analysis

To identify the haplotypes, present in the 31 YUCMEX pigs

sequences, 77 mitochondrial haplotype sequences (Alves et al.,

2003; Alves et al., 2010; van Asch et al., 2012; Wu et al., 2007),

representative of European, Asian and commercial populations were

used. The amplied fragments of YUCMEX pigs were aligned to

the GenBank AJ002189 sequence as a reference, trimming the study

fragment from position 15435 to 15977 with 543 bp (Alves et al.,

2003) using the MUSCLE tool in the MEGA software version 11

(Tamura et al., 2021).

Phylogenetic analyses

Using haplotypes from the dierent pig populations, genetic

distance analyses were carried out, comparing YUCMEX pigs with

Table 1. Change sites according to the variable position of the DNAmt D-loop, AJ002189 sequence as a reference.

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5

4 4 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 9

5 5 1 2 4 4 5 6 6 7 8 9 1 1 4 7 7 8 9 1 1 1 1 2 2 3 4 5 7 2 2 4 4 7 8 8 8 9 9 3 4

0 8 1 4 3 4 8 2 5 9 7 2 5 6 8 4 5 2 4 1 4 5 7 3 9 6 1 8 8 2 5 0 8 8 3 4 7 5 7 6 0

T G C G T G A A G C C A T T T T T G C T T T C A A C C T A A C T C A C T C C C A C

In the mitochondrial study within the YUCMEX population,

four subsets (Haplotypes) were observed. The greatest amount

corresponded to the HB and HD haplotypes (table 2) suggesting that

the maternal line of descent associated with the HB haplotype is the

most common haplotype in the YUCMEX population.

Table 2. Haplotypes identied in YUCMEX pigs according to the

variable position of the mtDNA D-loop.

Haplotypes 15544 15558 15615 15714 15758 15878 n

HA A A T C C G 1

HB G A T C C G 22

HC G A T C T A 2

HD G T C C C A 6

n: Sample size, HA: haplogroup A, HB: haplogroup B, HC: haplogroup C, HD: haplogroup

Mitochondrial study between Yucatan hairless pigs and

commercial breeds

When the mitochondrial sequences of YUCMEX pigs with

commercial populations were analyzed to measure the average

distances in each population, it was observed that the genetic distance

of the YUCMEX population concerning the Duroc (0.06 ± 0.03)

and Landraces (0.03 ± 0.02) breeds was the smallest. These small

distances indicate lower diversity according to the haplotypes used

in each population and also suggests that they are genetically more

similar to these breeds than to other populations. It appears that the

YUCMEX population and the Duroc and Landrace breeds have either

a shared genetic origin or a closely related recent evolutionary history.

With other populations was higher: Hampshire 0.48 ± 0.25,

Pietrain 0.44 ± 0.19, Large White 0.21 ± 0.09 and among YUCMEX

0.03 ± 0.02.

The short genetic distance in the YUCMEX pig reects the

conservation of the hairless variant in that region of the Yucatán

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Rev. Fac. Agron. (LUZ). 2025, 42(4): e254251 October-December. ISSN 2477-9409.

4-6 |

or locally adapted pigs, such as YUCMEX, is more common than

in commercial pigs, as pointed out by Zhang et al. (2016). In this

work, introgression of commercial breeds cannot be attributed, to the

distance observed because the phenotypic appearance of YUCMEX

pigs is distant from the white and spotted colors, retaining the black

and hairless color (Lemus-Flores et al., 2023). However, evidence of

interspecic hybridization is common in local pigs, such as that found

in Philippine pigs (Banayo et al., 2023).

Mitochondrial study among hairless pigs from Yucatan and

European and Asian racial groups.

The grouping of the haplotypes of European and Asian populations

identied seven groups in the phylogenetic analysis (table 3).

Table 3. Average distance within the populations of pigs.

Group Population Distance

Standard

Error

A Asian 0.1525 0.0489

B French WB, Italian WB 0.2744 0.0896

C Portuguese WB 0.1154 0.0582

Spanish WB, Iberian with hair, Bísa-

ro, Malhado

0.0499 0.0214

Spanish WB, Portuguese WB, Hair-

less Iberian

0.0227 0.0103

Red Iberian, French WB, Italian

WB, Portuguese WB

0.0075 0.0077

Mangalica, Croatian WB, Austrian

0.0488 0.0272

YUCMEX México 0.0300 0.0200

The greatest distance within each group is identied in groups

B (French WB Italian WB), A (Asians) and C (Portuguese WB);

and the smallest distance in the YUCMEX and Iberian haplotypes

indicates strong genetic homogeneity and close kinship conrming

the historical theory that American Creole pigs, like the Yucatán pig,

descend from pigs that arrived from the Iberian Peninsula during

colonization.

In the phylogenetic analysis and principal component analysis

(gure 3), the YUCMEX HA and HB haplotypes are located close

to group C (Portuguese WB), the HC haplotype with D (WB from

Spain, Iberian with hair, Bísaro, Malhado) and E (Spanish WB,

Portuguese WB, Iberian hairless), the HD haplotype near group F

(Iberian red, French WB, Italian WB, and Portuguese). Groups A, B

and G (Mangalica, WB from Croatia, Austrian WB) are more distant

from the YUCMEX haplotypes.

Figure 3. Principle component analysis (PCA) of haplotypes of

European, Asian wild and YUCMEX pigs.

Peninsula, supported by local breeding programs (Hernández et al.,

2020), without using crossbreeding with commercial pigs (Lemus

-Flores et al., 2020).

In the grouping with commercial pigs, a greater closeness of the

YUCMEX pigs to the Duroc breed is identied in the four haplotypes

(gure 1). The close relationship to the Duroc breed in all four

haplotypes indicates that the maternal lines of descent of YUCMEX

pigs are genetically very similar to those of the Duroc breed. This

reinforces the idea of a common origin or signicant crossbreeding

between the two populations. Likewise, it is seen through the principal

component analyses (gure 2) that the YUCMEX haplotypes are

grouped closely meaning that the pigs are genetically very similar

to each other in their maternal ancestry, and corroborates a lower

diversity in the D-loop mtDNA, but close to the Duroc population

revealing evidence of genetic conservation rather than genetic erosion.

Figure 1. Haplotypes groups in the DLoop del DNAm region.

Figure 2. Principal component analysis of commercial pig

populations and YUCMEX haplotypes.

The phylogenetic and principal component analysis reveals a

smaller distance with commercial pigs of the Duroc breed, because

they have a similar ancestral origin with Iberian pigs (Jones, 1998)

since there is no evidence of interbreeding of hairless pigs with

this commercial breed in Yucatan. The loss of diversity in local

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Lemus et al. Rev. Fac. Agron. (LUZ). 2025, 42(4): e254251

5-6 |

A separation is observed between Asian (A) and European groups,

where group B (French and Italian WB) was closer to the Asian pigs.

The four YUCMEX haplotypes (HA, HB, HC and HD) clustered

within clusters from Portugal and Spain pigs, indicating a European

origin.

The presence of only four haplogroups in YUCMEX pigs suggest

little genetic variability, compared to that of Iberian pigs, in which 14

haplogroups were identied (Alves et al., 2003), six to 11 haplotypes

in Indian pigs (Laxmivandana et al., 2022). Asian groups retain a

distinct lineage from Europeans, which has been mentioned in several

papers using mtDNA D-loop analysis (Giura et al., 2000; van Asch

et al., 2012; Alves et al., 2010) and with SNP analysis (Ramírez et al.,

2015). Ishihara et al. (2023) detected 50 haplotypes from Vietnamese

native pigs, with 27 novel haplotypes and no European haplotypes

found.

Concerning YUCMEX pigs the four haplogroups were grouped

into three lineages, within the Portuguese WB and Iberian pigs, away

from the Asian pigs, and groups formed by haplotypes from Eastern

Europe with French WB, Italian WB, Mangalica, WB from Croatia,

and Austrian WB; conrming their European origin from the Iberian

Peninsula from where they were originally left for America. Alves et

al. (2010) suggest that the Iberian Peninsula was a refuge for the pig

(Sus scrofa) during the ice age and that there is no evidence of Asian

mtDNA introgression. Analysis by Vergara et al. (2021) indicates

that Ecuadorian Creole pigs probably diverged from the Asian pig

population and that, like the YUCMEX pigs, they appear to be

genetically inuenced by European and Iberian populations raised in

Spain.

In the Yucatan hairless pig, little diversity was quantied with four

haplotypes that distance them from white commercial pigs (Landrace

and Large White) and spotted pigs (Pietrain), but close to Duroc.

There was a greater phylogenetic relationship of the Yucatan hairless

pig with Iberian and European wild pigs from the Iberian Peninsula,

far from Asian and wild pigs from Eastern Europe. Like the Iberian

pigs, the hairless pigs of Yucatan are distanced from the Asian pigs.

Conclusions

Four YUCMEX haplogroups were grouped into three lineages

close to the WB from Portugal and Spain, distant from the Asian pigs

and the groups formed by WB haplotypes from Eastern Europe. The

European origin, from the Iberian Peninsula of the YUCMEX pig is

conrmed.

This information could contribute to a better understanding of

the origin of the Yucatan hairless pig and to assess its conservation.

Further exploration is needed through complete mitochondrial DNA

sequencing and analysis.

Acknowledgments

This research was funded by the Secretariat for Research,

Innovation and Higher Education, Mérida, Yucatán, Mexico.

Literature cited

Alves, E., Ovilo, C., Rodríguez, M.C., & Silió, L. (2003). Mitochondrial DNA

sequence variation and phylogenetic relationships among Iberian pigs and

other domestic and wild pig populations. Animal Genetics, 34(5), 319-24.

https://doi.org/10.1046/j.1365-2052.2003.01010.x

Alves, P. C., Pinheiro, I., Godinho, R., Vicente, J., Gortáazar, C., & Scandura,

M. (2010). Genetic diversity of wild boar populations and domestic

pig breeds (Sus scrofa) in South-western Europe. Biological Journal of

the Linnean Society, 101(4), 797-822. https://doi.org/10.1111/j.1095-

8312.2010.01530.x

Banayo, J.B., Manese, K.L.V., & Salces, A.J. (2023). Phylogeny and Genetic

Diversity of Philippine Native Pigs (Sus scrofa) as Revealed by

Mitochondrial DNA Analysis. Biochemical Genetics, 61, 1401–1417.

https://doi.org/10.1007/s10528-022-10318-0

Burgos-Paz W., Souza, C.A., Megens, H.J., Ramayo-Caldas, Y., Melo, M.,

LemúsFlores, C., Caal, E., Soto, H. W., Martínez, R., Álvarez, L.

A., Aguirre, L., Iñiguez, V., Revidatti, M. A., MartínezLópez, O. R.,

Llambi, S., EsteveCodina, A., Rodríguez, M. C., Crooijmans, R. P. M.

A., Groenen, M. A. M., & Pérez-Enciso, M. (2013). Porcine colonization

of the Americas: a 60k SNP story. Heredity, 110(4), 321-330. https://doi.

org/10.1038/hdy.2012.109

Giura, E., Kijas, J.M., Amarger, V., Carlborg, O., Jeon, J.T., & Andersson, L.

(2000). The origin of the domestic pig: Independent domestication

and subsequent introgression. Genetics, 154(4),1785–91. https://doi.

org/10.1093/genetics/154.4.1785

Hernández, A.A., García-Munguía, C.A., García-Munguía, A.M., Ortiz-Ortiz,

J.R., Sierra-Vásquez, A.C., & Morales-Flores, S. (2020). Sistema de

producción del cerdo pelón mexicano en la Península de Yucatán. Nova

Scientia, 12(24), 1-22. https://doi.org/10.21640/ns.v12i24.2234

Ishihara S., Arakawa A., Ba N.V., Dinh N.C., Ninh P.H., Okamura, T., DangNguyen,

T. Q., Kikuchi, K., Pham, L. D., & Taniguchi, M. (2023). Population

structure of Vietnamese pigs using mitochondrial DNA. Animal Science

Journal, 94(1), e13875. https://doi.org/10.1111/asj.13875

Jones, G.F. (1998) Genetic Aspects of Domestication, Common Breeds and Their

Origin. In: Rothschild, M.F., & Ruvinsky, A. (Eds). The Genetics of the

Pig. CAB International, Wallingford, 17-50.

Laxmivandana, R., Vashi, Y., Kalita, D., Banik, S., Sahoo, N.R., & Naskar

S. (2022). Genetic diversity in mitochondrial DNA D-loop region of

indigenous pig breeds of India. Journal of Genetic,101, 5. https://doi.

org/10.1007/s12041-021-01353-8

Lemus-Flores, C., Alonso-Morales, R., Toledo-Alvarado, H., Sansor-Nah,

R., Burgos-Paz, W., & DzibCauich, D. (2020). Genetic diversity and

population structure of Yucatan black hairless pig using SNP50K chip.

Abanico Veterinario, 10, 1-12. https://doi.org/10.21929/abavet2020.10

Lemus-Flores, C., Prado, J., Bernal, R.V., Segura-Correa, J.C., & Sansor-

Nah, R. (2023). Genetic relationships of the Yucatan black hairless

pig with Iberian breeds using single nucleotide polymorphism.

Brazilian Journal of Veterinary Research and Animal Science, 60,

e195697. https://doi.org/10.11606/issn.1678-4456.bjvras.2023.195697

Markov, N.I., Ranyuk, M.N., Babaev, E.A., Seryodkin, I.V., Senchik, A.V. Bykova,

E. A., Esipov, A. V., Nurtazin, S. T., Pavlova, O. S., & Matrosova,

V. A, (2022). Introduced, Mixed, and Peripheral: Conservation of

Mitochondrial-DNA Lineages in the Wild Boar (Sus scrofa L.) Population

in the Urals. Diversity, 14(11), 916. https://doi.org/10.3390/d14110916

Miller, S.A., Dykes, D., & Polesky, H.F. (1989). A simple procedure for extracting

DNA from human nucleated cells. Nucleic Acids Research, 16(3),1216.

https://doi.org/10.1093/nar/16.3.1215

Minitab, LLC. (2021). Minitab 15 Statistical Software. State College, PA: Minitab,

Inc. Available from: www.minitab.com.

Niedziałkowska, M., Tarnowska, E., Ligmanowska, J., Jędrzejewska, B.,

Podgórski, T., Radziszewska, A., Ratajczyk, I., Kusza, S., Bunevich, A.

N., Danila, G., Shkvyria, M., Grzybowski, T., & Woźniak, M. (2021).

Clear phylogeographic pattern and genetic structure of wild boar Sus

scrofa population in Central and Eastern Europe. Scientic Reports, 11(1),

9680. https://doi.org/10.1038/s41598-021-88991-1

Ogata, N. (2019). 1519: Hernán Cortés y la llegada del Cerdo a la diversicación

productiva Mesoamericana. Diversidad Biológica y Cultural Trópico

Americano, Universidad Veracruzana. https://tropico-americano.uv.mx/

cerdo-pelon-mexicano/

Ortega-S, J. A., Delgado-Acevedo, J., Villarreal-González, J. G., Borroto-Páez, R.,

& Tamez-González, R. (2019). Wild Pigs in Mexico and the Caribbean.

Invasive Wild Pigs in North America, Edit. CRC Press, 1

edition, p. 423-

438. https://doi.org/10.1201/b22014-18

Perrier, X., Flori, A., & Bonnot, F. (2003). Genetic diversity of cultivated

tropical plants. Data analysis methods. By Perla Hamon, P. 1

. Edit.,

Eneld, Science Publishers. Montpellier. p 43 - 76. https://doi.

org/10.1201/9781482280043

Ramírez, O., Burgos-Paz, W., Casas, E., Ballester, M., Bianco, E., Olalde,

I. Santpere, G., Novella, V., Gut, M., Lalueza-Fox, C., Saña, M., &

Pérez-Enciso, M. (2015). Genome data from a sixteenth century pig

illuminate modern breed relationships. Heredity 114, 175–184. https://

doi.org/10.1038/hdy.2014.81

Ramos-Canché, M. E., Magaña-Magaña, M. A., Aguilar-Urquizo, E., Pech-Zapata,

A., Piñeiro-Vázquez, A. T., Toledo-López, V.M., & Sanginés-García, J.R.

et al. (2020). Óptimos económicos en la cría del cerdo pelón mexicano:

propuesta de integración para cadena productiva. Ecosistemas y Recursos

Agropecuarios, 7(1), e2302. https://doi.org/10.19136/era.a7nl.2302

Saitou, N., Nei, M. (1987). The Neighbor-joining method: a new method for

reconstructing hylogenetic trees. Molecular Biology and Evolution, 4,

406–425. http://doi.org/10.1093/oxfordjournals.molbev.a040454

Secretaría de Agricultura y Desarrollo Rural. (1998). NOM-051-ZOO-1995.

Trato humanitario en la movilización de animales. Diario Ocial de

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Rev. Fac. Agron. (LUZ). 2025, 42(4): e254251 October-December. ISSN 2477-9409.

6-6 |

la Federación. Vigente desde el 24 de marzo de 1998. http://publico.

senasica.gob.mx/?doc=531

Secretaría de Agricultura y Desarrollo Rural. (2001). NOM-062-ZOO-1999.

Especicaciones técnicas para la producción, cuidado y uso de los

animales de laboratorio. Diario Ocial de la Federación. Vigente desde

el 23 de agosto de 2001. http://publico.senasica.gob.mx/?doc=743

Tamura, K., & Kumar, S. (2002). Evolutionary distance estimation under

heterogeneous substitution pattern among lineages. Molecular

Biology and Evolution, 19(10), 1727–1736. https://doi.org/10.1093/

oxfordjournals.molbev.a003995

Tamura, K., Stecher, G., & Kumar, S. (2021). MEGA 11: Molecular Evolutionary

Genetics Analysis Version 11. Molecular Biology and Evolution, 38, 7,

3022–3027. https://doi.org/10.1093/molbev/msab120.

van Asch, B., Pereira, F., Santos, L.S., Carneiro, J., Santos, N., & Amorim, A.

(2012). Mitochondrial lineages reveal intense gene ow between Iberian

wild boars and South Iberian pig breed. Animal Genetics, 43, 35–41.

https://doi.org/10.1111/j.1365-2052.2011.02222.x

Vergara, A.M.C., Martínez, A.M., Bermejo, J.V.D., Macri, M., Nájera, P.R.A.,

Duchi, N.A.D., &. Vargas, P.A.T. (2021). A Matrilineal Study on the

Origin and Genetic Relations of the Ecuadorian Pillareño Creole Pig

Population through D-Loop Mitochondrial DNA Analysis. Animals, 11,

3322. https://doi.org/10.3390/ani11113322

Wu, G.S., Yao, Y.G., Qu, K.X., Ding, Z.L., Li, H., Palanichamy, M.G., Duan, Z.Y.,

Li, N., Chen, Y.S., &. Zhang, Y.P. (2007). Population phylogenomic

analysis of mitochondrial DNA in wild boars and domestic pigs revealed

multiple domestication events in East Asia. Genome Biology, 8, R245.

https://doi.org/10.1186/gb-2007-8-11-r245

Zhang, J., Jiao, T., & Zhao, S. (2016). Genetic diversity in the mitochondrial DNA

D-loop region of global swine (Sus scrofa) populations. Biochemical and

Biophysical Research Communications, 473(4), 814-820. https://doi.

org/10.1016/j.bbrc.2016.03.125