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Poneromorfas Do Brasil Miolo.Indd 10 - Citogenética e evolução do cariótipo em formigas poneromorfas Cléa S. F. Mariano Igor S. Santos Janisete Gomes da Silva Marco Antonio Costa Silvia das Graças Pompolo SciELO Books / SciELO Livros / SciELO Libros MARIANO, CSF., et al. Citogenética e evolução do cariótipo em formigas poneromorfas. In: DELABIE, JHC., et al., orgs. As formigas poneromorfas do Brasil [online]. Ilhéus, BA: Editus, 2015, pp. 103-125. ISBN 978-85-7455-441-9. Available from SciELO Books <http://books.scielo.org>. All the contents of this work, except where otherwise noted, is licensed under a Creative Commons Attribution 4.0 International license. Todo o conteúdo deste trabalho, exceto quando houver ressalva, é publicado sob a licença Creative Commons Atribição 4.0. Todo el contenido de esta obra, excepto donde se indique lo contrario, está bajo licencia de la licencia Creative Commons Reconocimento 4.0. 10 Citogenética e evolução do cariótipo em formigas poneromorfas Cléa S.F. Mariano, Igor S. Santos, Janisete Gomes da Silva, Marco Antonio Costa, Silvia das Graças Pompolo Resumo A expansão dos estudos citogenéticos a cromossomos de todas as subfamílias e aquela partir do século XIX permitiu que informações que apresenta mais informações a respeito de ca- acerca do número e composição dos cromosso- riótipos é também a mais diversa em número de mos fossem aplicadas em estudos evolutivos, ta- espécies: Ponerinae Lepeletier de Saint Fargeau, xonômicos e na medicina humana. Em insetos, 1835. Apenas nessa subfamília observamos carió- são conhecidos os cariótipos em diversas ordens tipos com número cromossômico variando entre onde diversos padrões cariotípicos podem ser ob- 2n=8 a 120, gêneros com cariótipos estáveis, pa- servados. Dentre os himenópteros, são conheci- drões de cromossomos exclusivos para um gênero dos os números cromossômicos de cerca de 800 como Dinoponera Roger, 1861, variação intraes- morfoespécies de formigas de diversas regiões pecífi ca em espécies de Neoponera Emery, 1901 e biogeográfi cas, embora nem todos esses estudos a presença de cromossomos com potencial para defi nam o cariótipo dos organismos estudados. serem marcadores de uma tribo. As hipóteses aqui Essas informações correspondem a cerca de 6% lançadas têm por base a análise de mais de 100 do número de espécies descritas para a família e, cariótipos de poneromorfas. São sucessivamente mesmo escassas, apontam uma grande variação abordados i) Situação atual do conhecimento ci- nos cariótipos conhecidos. Em diferentes subfa- togenético em Formicidae; ii) Conhecimento de mílias, padrões distintos de cariótipos são encon- citogenética em poneromorfas; iii) Estudos de trados e, dentre essas, as espécies agrupadas no caso: Tribo Ponerini, possíveis agrupamentos ge- conjunto artifi cial denominado “poneromorfas” néricos/cariótipos; iv) Análises dos cariótipos co- apresentam interessantes modelos para elabora- nhecidos para esse agrupamento por meio de mo- ção de hipóteses acerca da evolução cariotípica delos como o cariógrafo e uma comparação com que serão apresentados neste capítulo. Dentre as outras subfamílias. Esses dados oferecem subsí- seis subfamílias agregadas sob a denominação dios para formular hipóteses sobre a evolução dos poneromorfas, existem informações acerca dos cariótipos em formigas poneromorfas. MARIANO, Cléa S.F.; SANTOS, Igor S.; SILVA, Janisete Gomes da; COSTA, Marco Antonio; POMPOLO, Silvia das Graças. Citogenética e evolução do cariótipo em formigas poneromorfas. In: DELABIE, Jacques H. C. et al. As formigas poneromorfas do Brasil. Ilhéus: Editus, 2015. p. 103-125. Citogenética e evolução do cariótipo em formigas poneromorfas | 103 PPoneromorfasoneromorfas ddoo BBrasil_miolo.inddrasil_miolo.indd 110303 113/01/20163/01/2016 111:41:49::0404 Abstract Cytogenetics and karyotype evolution for all six poneromorph subfamilies. Th e subfamily in poneromorph ants - From the 19th century for which most karyotype information is available, onwards, the continuing development of the Ponerinae Lepeletier de Saint Fargeau, 1835, cytogenetic procedures has allowed information is also the most diverse in species numbers. Only to be gathered that can be applied to studies on within this family has the following been reported: evolution and taxonomy. In insects, karyotypes are karyotypes with chromosome numbers ranging known for several orders and various karyotype from 2n=8 to 120; genera with stable karyotypes; patterns can be observed. Among hymenopterans, chromosome patterns exclusive to a genus such as chromosome numbers are known for about 800 Dinoponera Roger, 1861; intraspecifi c variation in ant morphospecies from distinct biogeographical species of Neoponera Emery, 1901; and the presence regions, although not all of the studies have of chromosomes with the potential for being a defi ned the karyotype of the studied organisms. marker for the tribe Ponerini. Th e hypotheses Th is karyotype information corresponds to about advanced in this chapter are based on the analysis 6% of described Formicidae species and, although of more than 100 karyotypes of poneromorphs. We scarce, has demonstrated that there is large here discuss: i) the state of the art of cytogenetics variation among the known karyotypes. Among of Formicidae; ii) cytogenetics of poneromorphs; the diff erent subfamilies, distinct karyotype iii) case studies concerning the tribe Ponerini, patterns are found, and of these subfamilies, the with possible groupings of genera/karyotypes; iv) species included in the artifi cial group named analysis of known karyotypes for this grouping poneromorphs are interesting models for the using methods such as the karyograph; and v) construction of hypotheses regarding karyotype a comparison with other subfamilies. Th ese data evolution, as discussed in this chapter. For this provide background for formulating hypotheses group, information on chromosomes is available on karyotype evolution in poneromorph ants. Citogenética Na ordem Hymenoptera, os trabalhos de Crozier (1975) e Imai e colaboradores (1977, 1984a, b) fo- Em comparação com outras disciplinas da ram importantes contribuições ao conhecimento biologia, a citogenética pode ser considerada recen- citogenético, os quais determinaram o cariótipo de te, pois somente apareceu no fi nal do século XIX. Os centenas de espécies de diferentes regiões geográ- primeiros estudos descreviam os processos de mi- fi cas cujas informações forneceram a base para a tose e a presença de cromossomos denominados de elaboração de uma teoria de evolução cariotípica. “material cromático nuclear” (SUMNER, 2003). Ra- A análise de cromossomos mitóticos é utili- pidamente foi possível descrever cromossomos mi- zada com frequência em estudos citotaxonômicos tóticos em animais (FLEMMING, 1882) e estes es- em alguns grupos de eucariotos (IMAI, 1983; BAI- tudos são considerados os pioneiros da citogenética. MAI, 1998; MENEZES et al., 2014). Através do estu- A expansão dos estudos citogenéticos per- do de cariótipos, além do número e morfologia dos mitiu que esta disciplina fosse aplicada em estu- cromossomos, é possível inferir sobre diferenciação dos evolutivos, taxonômicos e na medicina. Em de espécies, grau de parentesco entre estas, proces- insetos, diversos exemplos podem ser apontados sos evolutivos tais como especiação simpátrica e/ou em ordens como Lepidoptera (KANDUL et al., alopátrica, e posição fi logenética dos taxa estudados, 2007; LUKHTANOV et al., 2011), Diptera (SBI- assim como elaborar hipóteses sobre a evolução dos LORDO et al, 2010) e Hymenoptera, entre outras cariótipos e seu papel na especiação (LUKHTA- (GOKHMAN, 2006). Nessa classe de Hexapoda, NOV et al., 2006; LORITE; PALOMEQUE, 2010). diversos modelos cromossômicos foram defi ni- A princípio, cada espécie tem um cariótipo dos como, por exemplo, os cromossomos politê- (que é o conjunto cromossômico defi nido por nú- nicos em dípteros quironomídeos (METZ, 1916). mero + morfologia cromossômica) conservado e 104 | Cléa S.F. Mariano, Igor S. Santos, Janisete Gomes da Silva, Marco Antonio Costa, Silvia das Graças Pompolo PPoneromorfasoneromorfas ddoo BBrasil_miolo.inddrasil_miolo.indd 110404 113/01/20163/01/2016 111:41:49::0505 compartilhado pelos indivíduos, independente da de braços cromossômicos: este número aumenta sua distribuição geográfi ca. Porém, existem me- quando a inversão é do tipo AM (um cromosso- canismos que causam mudanças nos cariótipos. mo do tipo A sofre quebras e transforma-se em um Estas mudanças podem ser numéricas e/ou estru- cromossomo do tipo M) (Figura 10.2); ou diminui turais e podem ser observadas em diversos níveis: o número de braços quando a inversão é do tipo intrapopulacional (quando mais de um cariótipo MA (um cromossomo do tipo M sofre uma quebra é observado em uma única população) (LORITE; na região centromérica e transforma-se em um cro- PALOMEQUE, 2010); entre populações e entre mossomo do tipo A) (IMAI; MARUYAMA, 1978). espécies irmãs ou espécies crípticas (defi nição de Inversões do tipo MA são consideradas raras em SEIFERT, 2009). Essas alterações são geradas por formicídeos porque tenderiam a aumentar a ocor- eventos conhecidos como rearranjos cromossômi- rência de interações entre os cromossomos, prová- cos, os quais alteram o número e/ou a morfologia vel causa de efeitos deletérios (IMAI et al., 1988). dos cromossomos (Figuras 10.1, 10.2). As inversões do tipo paracêntrica (fora da região As alterações numéricas referem-se à varia- do centrômero) são imperceptíveis e não são visu- ção no número de cromossomos e são causadas alizadas por
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