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Chromosome Polymorphism in the Brazilian Dwarf Brocket Deer, Mazama Nana (Mammalia, Cervidae)

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Chromosome Polymorphism in the Brazilian Dwarf Brocket Deer, Mazama Nana (Mammalia, Cervidae) Genetics and Molecular Biology, 31, 1, 53-57 (2008) Copyright by the Brazilian Society of Genetics. Printed in Brazil www.sbg.org.br Research Article Chromosome polymorphism in the Brazilian dwarf brocket deer, Mazama nana (Mammalia, Cervidae) Vanessa Veltrini Abril1,2 and José Maurício Barbanti Duarte2 1Programa de Pós-Graduação em Genética e Melhoramento Animal, Faculdade de Ciências Agrárias e Veterinárias, Universidade Estadual Paulista, Campus de Jaboticabal, Jaboticabal, SP, Brazil. 2Núcleo de Pesquisa e Conservação de Cervídeos, Departamento de Zootecnia, Faculdade de Ciências Agrárias e Veterinárias, Universidade Estadual Paulista, Campus de Jaboticabal, Jaboticabal, SP, Brazil. Abstract The Brazilian dwarf brocket deer (Mazama nana) is the smallest deer species in Brazil and is considered threatened due to the reduction and alteration of its habitat, the Atlantic Rainforest. Moreover, previous work suggested the pres- ence of intraspecific chromosome polymorphisms which may contribute to further population instability because of the reduced fertility arising from the deleterious effects of chromosome rearrangements during meiosis. We used G- and C-banding, and nucleolus organizer regions localization by silver-nitrate staining (Ag-NOR) to investigate the causes of this variation. Mazama nana exhibited eight different karyotypes (2n = 36 through 39 and FN = 58) result- ing from centric fusions and from inter and intraindividual variation in the number of B chromosomes (one to six). Most of the animals were heterozygous for a single fusion, suggesting one or several of the following: a) genetic in- stability in a species that has not reached its optimal karyotypic evolutionary state yet; b) negative selective pressure acting on accumulated rearrangements; and c) probable positive selection pressure for heterozygous individuals which maintains the polymorphism in the population (in contrast with the negative selection for many rearrangements within a single individual). Key words: Mazama nana, Brazilian dwarf brocket deer, cytogenetics, chromosome polymorphism, centric fusions. Received: June 28, 2007; Accepted: October 16, 2007. Introduction the tail. Its anterior limbs are shorter and darker than the The genus Mazama (Artiodactyla, Cervidae) is com- posterior ones (Duarte, 1996; Mikich & Bérnils, 2004). posed of eight species: M. rufina, M. pandora, M. chunyi, Mazama nana is found from northern Paraná State to cen- M. americana, M. gouazoubira, M. nana, M. nemorivaga tral Rio Grande do Sul State, encompassing parts of Para- and M. bororo (Wemmer, 1998; Duarte, 2007). It is distrib- guay and Argentina. These areas are presently undergoing uted from Mexico to central Argentina and is classified as a significant fragmentation and anthropic alteration (Duarte, “small solitary forest deer” because of its ecological and 1996). The Brazilian dwarf brocket deer was identified as behavioral characteristics (Barrette, 1987; Eisenberg & vulnerable in the official list of Brazilian threatened species Redford, 1999). In Brazil, five species are recognized based in 2003 following the recommendation of the Deer Special- on morphology and cytogenetics: M. gouazoubira, M. ist Group and was classified as DD (deficient data) in “The nemorivaga, M. americana, M. nana and M. bororo (Duar- 2006 IUCN Red List” (IUCN, 2006; IBAMA, 2007). te & Merino, 1997; Duarte & Jorge, 2003; Duarte, 2007). The M. nana karyotype was initially described from The Brazilian dwarf brocket (Mazama nana)isthe analyses of three animals from the Paraná River basin in the smallest deer species in Brazil, reaching a weight of 15 kg Foz do Iguaçu region. These individuals possessed 2n = 36 and a height of 45 cm. This species has small, slightly and FN = 58 with four or five B chromosomes. Chromo- pointed and almost hairless ears and displays little grada- somes 1, 2, 6, 7, 8, 9 and 10 were metacentric; chromo- tion in its reddish coloration, other than a few white hairs on somes 3, 4 and 5 were submetacentric; chromosomes 11 through 17 were acrocentric; the X chromosome was a midium-sized metacentric and the Y was probably a small- Send correspondence to José Maurício Barbanti Duarte. Departa- mento de Zootecnia, Faculdade de Ciências Agrárias e Veteriná- sized metacentric. A karyotype with 2n = 39+5B and rias, Universidade Estadual Paulista, 14884-900 Jaboticabal, SP, FN = 58, heterozygous for a centric fusion involving pair 5 Brazil. E-mail: [email protected]. and lacking the metacentric pair 10 was also described phisms described in somes and theC-banding rearrangements involved and in Ag-NOR the to polymor identify individualmosome chromo fusions/fissions. clearly ascertained and it was suggested topatterns result from were chro obtained, the causecias, of Universidade Estadual this Paulista, variation 1998). was AsArtiodactyla) no not banding no Brasil. PhD thesis, Instituto de Biociên Material andMethods genética e taxonômicawere up seven do B chromosomes (Duarte gênero varied JMB, from Análise 36 to cito 40, the FNdo ranged Sul from 56 States) to 60 and andlocalities Paraguay there of in Brazil (Paraná, which Santa the Catarinapolymorphisms and diploid Rio were number Grande described indade Estadual animals Paulista, from 1992). More different recently,Faculdade chromosome de Ciênciasalgumas Agrárias espécies e de cervídeos Veterinárias/Universi brasileiros.(Duarte Master’s thesis. JMB, Aspectos taxonômicos e citogenéticos de 54 Results andDiscussion Duarte (PhD thesis, 1992) and the differencesautosome between both pairs. This karyotype(pairs is 2, 4, the 5 same andmetacentric presented 9) (pairs by 1, and 3, seven 6, acrocentric 7,tively (pairs 8 11 and (Figure 10), to four 17) 1). submetacentric and This seven chromosome karyotype pairs in was groupsand A, one C composed to and six E, of B respec chromosomes six and FN = 58, withpossibly six, a four small-sized metacentric. mosome was a medium-sizedseveral metacentric rearrangements (V01 and to V07; the Table YFN 1). The = was X 58 chro and seven distinct karyotypes were found due to variation. damental numberssomes were because not considered to there calculateextranumerary the diploid chromosomes and with was fun RL > 1.0%).somes intraindividual B chromo with RLchromosomes > with 3.0%) RL and <C (biarmed B 3.0%), chromosomes with E (microchromosomes RL > (acrocentriclengths or 2.5%), (RL): D chromo A (acrocentric (biarmed chromosomesand with were RL organized > into 2.5%), groupssubmetacentric according or to acrocentric according their to relative their arms ratio Sumner (1972) and Howell &Ag-NOR Black (1980), were respectively. obtainedcultures according (Verma to & Seabrightchromosome Babu, preparations after 1995). (1971), lymphocyte and G- fibroblasts Mazama and nana C-banding and blood and skin samples were collected from 24 captive The aim of the present study was to use G- and The most frequent karyotype (V01) showed 2n = 36 The diploid number varied from 2n = 36 to 39 with The chromosomes were classified as metacentric, After physical and chemical restraint of the animals, (13 males and 11 females) and used to obtain Mazama nana . Mazama (Cervidae; - - - - - - - - - - - Table 1 - Cytogenetic data of Mazama nana. V 2n FN Number of Figure Autosome morfology Variation Number of analysed autosomes reference of B chro- animals 1234567891011121314151617mosome male female V01 36 58 34 1(a-d) MSMS SMMMSMAAAAAAA 1to2 01 0 2 01 0 2to4 0 02 2to5 03 0 4to6 0 01 V02 375835- MSHS SMMMSMAAAAAAA 2to40 01 V03 3758352aMSMS SMHMSMAAAAAAA 1to3 01 0 1to4 0 01 2 01 0 3to6 0 01 V04 3858362bMSMS SMMHHMAAAAAAA 2to5010 V05 3858362cMSMS SMAMSMAAAAAAA 1to2 0 01 2to3 0 01 2to4 0 01 Abril and Duarte 4to6 01 0 V06 395837- HSMS SMAMSMAAAAAAA 2to3010 V07 3958372dMSHS SMAMSMAAAAAAA 2to4 01 01 2to5 02 01 V - karyotype variants; 2n - diploid number; FN - fundamental number; M - metacentric; S - submetacentric; H - heterozygous pair; A - homozygous acrocentric pair. Karyotypic variation in Mazama nana 55 Figure 1 - Karyotype V01 of Mazama nana (2n = 36; FN = 58) after (a) conventional Giemsa staining, (b) C-banding, (c) Ag-NOR staining and (d) G-banding. classifications are due to chromosome ordering. All chro- V04(2n=38+2to5B;FN=58)wastheonly mosome pairs, except pairs 2 and 10, presented pericentro- karyotypic variant with two heterozygous rearrangements meric constitutive heterochromatin after C-banding. Pairs 2 involving pairs 8 and 9. V05 also presented 2n = 38 and and 10 showed a faint positive C-band in the centromere. FN = 58, but had an acrocentric pair 7 (Figures 2c, 2d) and Interstitial hetrochromatin was detected in both chromo- one to six B chromosomes. some arms of pairs 1 and 4 and on the long arm of pairs 2, 3 V06 and V07 shared the same diploid and fundamen- and 5 (Figure 1b). The Ag-NORs were terminally located tal numbers (2n = 39; FN = 58) and the absence of the on the short arms of pair 2 and on the long arms of pair 6 metacentric pair 7, represented by identifiable acrocentric (Figure 1c). G-banding allowed the identification of all chromosomes. These two karyotypes differed in the chro- chromosomes (Figure 1d). mosome pairs involved in heterozygous centric fusions, pair 1 in V06 and pair 3 in V07 (Figure 2d). The number of V01(2n=36+1to6B;FN=58)waschosen as the B chromosomes varied from two to three in V06 and from standard karyotype for the species based on its higher fre- two to five in V07. quency among the animals analyzed (8 out of 24) and on the The hypothetical ancestral karyotype of Cervidae
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