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A Biodiversity Approach in the Neotropical Erythrinidae ¢Sh, Hoplias Malabaricus

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A Biodiversity Approach in the Neotropical Erythrinidae ¢Sh, Hoplias Malabaricus Chromosome Research 8: 603^613, 2000. 603 # 2000 Kluwer Academic Publishers. Printed in the Netherlands A biodiversity approach in the neotropical Erythrinidae ¢sh, Hoplias malabaricus. Karyotypic survey, geographic distribution of cytotypes and cytotaxonomic considerations Luiz A. C. Bertollo1, Guassenir Gonc°alves Born2, Jorge A. Dergam3, Alberto Sergio Fenocchio4 & Orlando Moreira-Filho1 1 Departamento de Gene¨ tica e Evoluc°a¬o, Universidade Federal de Sa¬o Carlos, C.P. 676, 13565-905, Sa¬o Carlos, SP, Brazil; Tel: (016) 260.8309; Fax: 55 16 261.2081; E-mail: [email protected]; 2 Departamento de Cieª ncias Morfobiolo¨ gicas, Universidade do Rio Grande, Rio Grande, RS, Brazil; 3 Departamento de Biologia Animal, Universidade Federal de Vic°osa, Vic°osa, MG, Brazil; 4 Departamento de Gene¨ tica, Universidad Nacional de Misiones, Posadas, Argentina Received 20 May 2000; received in revised form and accepted for publication by M. Schmid 10 July 2000 Key words: cytotaxonomy, geographic distribution, Hoplias malabaricus ¢sh, karyotypic diversity, sympatric cytotypes Abstract Hoplias malabaricus, a widely distributed neotropical freshwater ¢sh, shows a conspicuous karyotypic diversi¢cation. An overview of this diversity is presented here comprising several Brazilian populations, and some others from Argentina, Uruguay and Surinam. Seven general cytotypes are clearly identi¢ed on the basis of their diploid number (2n 39 to 2n 42), chromosomal morphology and sex chromosome systems, which can be clustered into two major karyotypic groups. This clustering suggests that karyotype structure would be more informative than the diploid number regarding cytotype relationships in this ¢sh group. While some cytotypes show a wide geographical distribution, some others appear to be endemic to speci¢c hydrographic basins. Sympatric cytotypes can occur without detection of hybrid forms; this situation points to a lack of gene £ow, a fact that is also reinforced by studies with genomic markers. The karyotypic data support the view that the nominal taxon H. malabaricus corresponds to a species complex comprising distinct evolutionary units, each with well-established chromosomal differences. Introduction dence has pointed to the karyotypic diversity of H. malabaricus, showing interpopulational differ- The Erythrinidae family comprises some ences in the diploid number and chromosome neotropical ¢shes with a wide distribution in South morphology, as well as in sex chromosome systems America (Britski et al. 1986). Within this taxon, (Bertollo et al. 1979, 1983, Ferreira et al. 1989, Hoplias malabaricus is the most widespread Dergam & Bertollo 1990, Scavone et al. 1994, species. Although usually considered as a single Lopes & Fenocchio 1994, Bertollo et al. 1997a, biological species, the taxonomy of this group is 1997b, Lopes et al. 1998, Bertollo & Mestriner poorly understood (Oyakawa 1990). Growing evi- 1998, Born & Bertollo 2000). 604 L. A. C. Bertollo et al. Specimens with a putative hybrid karyotype submetacentrics, subtelocentrics, and acrocentrics have not been found when distinct chromosomal according to their morphology and arm ratios forms (cytotypes) are sympatric. Such is the case (Levan et al. 1964). in the rio Aguapey (northeastern Argentina) where specimens with 2n 40 and 2n 42 chromosomes are found together, without a 2n 41 intermediary Results and discussion form (Lopes et al. 1998). Similar situations are also observed in some Brazilian localities (Scavone Based on their macrostructure, we are able to et al. 1994, Bertollo et al. 1997a). determine seven basic karyotypic con¢gurations In this paper, we provide an overview of the referred to hereafter as cytotypes (Figures 2 & karyological diversity in Hoplias malabaricus, 3). Each of these cytotypes shows unique com- with the description of a new cytotype from the binations of chromosome numbers and/or Amazon basin and the comparative analysis of morphologies; some of their most remarkable the several known cytotypes, their geographic aspects are summarized here. The distribution distributions and sympatric regions, compiled of the cytotypes is based on the available data from our studies with this ¢sh group over the last up till now. two decades. The available karyotypic data for Hoplias malabaricus have led to the hypothesis Cytotype A that this ¢sh represents a species complex (Bertollo et al. 1986, Dergam & Bertollo 1990, Bertollo et al. Cytotype A presents 2n 42 meta- and sub- 1997a, Lopes et al. 1998); this is reinforced by the metacentric chromosomes in both sexes (Figures present study. 2A & 3A), without an apparent sex chromosome system. This cytotype shows a wide distribution, from northern to southern Brazil, Uruguay Materials and methods (Dergan, unpublished), and northern Argentina (Figure 1). Data are available from thirty-six distinct localities, thirty-two in Brazil, two in Argentina, Cytotype B one in Uruguay and one in Surinam (Table 1, Fig- ure 1). Samples sizes are also given in Table 1. Cytotype B also shows 2n 42 chromosomes both Karyological analyses were performed from in males and females, the general karyotypic struc- cephalic kidney cells, with either of the following ture being similar to cytotype A. However, this protocols: short-term culture cells (Fenocchio et cytotype can be differentiated by an exclusive al. 1991), Hank's saline treatment (Foresti et al. XX/XY sex chromosome system: females present 1993) or the conventional air-drying method two subtelocentric X chromosomes (pair 6); in (Bertollo et al. 1978). In the latter case, specimens the male karyotype, only one of this chromosome were previously treated with 0.05% colchicine sol- is identi¢ed, together with the Y chromosome, ution (1 ml/100 g body weight), 50^60 minutes probably the smallest submetacentric in the comp- before sacri¢ce. Some specimens were also ¢rst lement (Figures 2B and 3B). The X chromosome stimulated with a yeast solution as a mitogenic also carries ribosomal cistrons and can be (Lee & Elder 1980). Meiotic preparations were polymorphic in size (Born & Bertollo 2000). This basically obtained by the method of Kligerman cytotype has a geographic distribution restricted & Bloom (1977), according to the description in to a lake system in the Vale do Rio Doce, Minas Bertollo & Mestriner (1998). Gerais State ^ Brazil (Figure 1). The diploid number was determined for each specimen studied. The homologous pairs were Cytotype C arranged in decreasing order of size in the karyotype, and partial idiograms were drawn to Cytotype C is characterized by 2n 40 meta- and depict some relevant aspects of the karyotypes. submetacentric chromosomes, both in males and The chromosomes were classi¢ed as metacentrics, females, without an apparent sex chromosome Chromosomal diversity in Hoplias ¢sh 605 Table 1. Collection sites of Hoplias malabaricus, with the respective cytotypes and sample sizes. Locality Cytotype n References 1. Manaus (AM) ^ igarape¨ Mindu¨A22 2. Pocone¨ (MT) ^ lagoons: rio Bento Gomes A 1 2 3. Araguaiana (MT) ^ co¨ rrego Dois de Agosto A 2 2, 11 4. Treª sMarias(MG)^rioSa¬ o Francisco A 3 2 5. Reserva Ecolo¨ gica do Jata|¨ (SP) ^ lagoons: rio Mogi-Guac°u A 3 9 6. S. J. do Marinheiro (SP) ^ Aè gua Vermelha reservoir: rio Grande A 2 2, 14 7. Conceic°a¬ o das Alagoas (MG) ^ Volta Grande reservoir: rio Grande A 6 10 8. Juquia¨ (SP) ^ rio Juquia¨ A912 9. Itatinga (SP) ^ Jurumirim reservoir: rio Paranapanema A 7 2 10. Poc°o Preto (SC) ^ rio Iguac°u A 1 2 11. Gua|¨ba (RS) ^ rio Gua|¨ba A 12 2 12. Corrientes ^ Argentina ^ r|¨oAguapey A 21 3,4 13. Tacuarembo¨ ^Uruguai^r|¨o Negro A 6 15 14. Parque Florestal do Rio Doce (MG) ^ lagoons: rio Doce B 11 12, 16 15. Manaus (AM) ^ rio Negro; igarape¨ Mindu C 15 1, 2 16. Tucuru|¨ (PA) ^ rio Tocantins C 7 1 17. Porto Velho (RO) ^ rio Madeira C 6 1, 2 18. Aripuana¬ (MT) ^ rio Aripuana¬ C11 19. Cuiaba¨ (MT) ^ lagoons: rio Cuiaba¨ C141,2 20. Aragarc°as (GO) ^ lagoons: rio Araguaia C 7 2 21. Goia¨ s Velho (GO) C 2 1 22. Corumba¨ (MS) ^ rio Paraguai C 8 1, 2 23. Miranda (MS) ^ lagoons: rio Miranda C 10 1, 2 24. Misiones ^ Argentina ^ r|¨oParana¨ C183 25. Corrientes ^ Argentina ^ r|¨o Aguapey, r|¨o Riachuelo C 4 4, 5 26. Itirapina (SP) ^ Lobo reservoir: ribeira¬ o do Lobo D 40 2, 6 27. Sa¬ o Carlos (SP) ^ UFSCar reservoir: ribeira¬ o Monjolinho D 42 2, 7, 8 28. Pirassununga (SP) ^ rio Mogi-Guac°u D 9 2, 8 29. Ipeu¨ na (SP) ^ rio Passa-Cinco D 9 9 30. Piracicacaba (SP) ^ rio Piracicaba D 7 9 31. Novo Horizonte (SP) ^ rio Treª sPontes D 3 9 32. Mirassolaª ndia (SP) ^ ribeira¬ o Barra Grande D 3 2 33. Reserva Ecolo¨ gica Jata|¨ (SP) ^ lagoons: rio Mogi-Guac°u D 8 9 34. Conceic°a¬ o das Alagoas (MG) ^ Volta Grande reservoir: rio Grande D 4 10 35. Londrina (PR) ^ ribeira¬ oTreª sBocas D 7 2 36. Porto Trombetas (PA) ^ rio Trombetas E 1 2 37. Paramaribo ^ Surinam F 6 1 38. Tucurui (PA) ^ rio Tocantins F 2 1 39. Sa¬ oLuiz(MA) F 3 1 40. Natal (RN) ^ lagoa Redonda: N|¨zia Floresta F 4 1, 2 41. Recife (PE) F 4 1 42. Treª s Marias (MG) ^ rio Sa¬ o Francisco F 4 1, 7 43. Porto Trombetas (PA) ^ rio Trombetas G 1 2 44. Porto Velho (RO) ^ rio Madeira G 3 2 45. Aripuana¬ (MT) ^ rio Aripuana¬ G152,6 n Number of specimens studied; Brazilian States in brackets: AM: Amazonas; GO: Goia¨ s; MA: Maranha¬ o; MG: Minas Gerais; MS: Mato Grosso do Sul; MT: Mato Grosso; PA: Para¨ ; PE: Pernambuco; PR: Parana¨ ; RN: Rio Grande do Norte; RO: Rondoª nia; RS: Rio Grande do Sul; SC: Santa Catarina; SP: Sa¬ oPaulo. References: 1. Bertollo et al. (1997a); 2. Present paper; 3. Lopes & Fenocchio (1994); 4. Lopes et al. (1998); 5. Jorge (1995); 6. Bertollo et al. (1983); 7.
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