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Chromosomal Diversity in Scorpaenidae ( Teleostei

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Chromosomal Diversity in Scorpaenidae ( Teleostei C 1997 The Japan Mendel Society Cytologia 62 : 397 - 404, 1997 Chromosomal Diversity in Scorpaenidae ( Teleostei, Scorpaeniformes ) : Cytogenetic Studies in Scorpaena brasiliensis and Scorpaena isthmensis from the Coast of Rio de Janeiro, Brazil Margaret Mariade OliveiraCorre1 and Pedro ManoelGaletti Junior2 1Universidade Federal doRio de Janeiro , DepartamentodeZoologia, IB-21941 -590, Rio de Janeiro, RJ,Brazil 2Universidade Federal deSao Carlos , DepartamentodeGenetica e Evolucao -13565 -905 , Sao Carlos, SP,Brazil Accepted October 15, 1997 The Scorpaeniformes are morphologically quite diverse representing the fifth largest order of fish, in which most of the species are marine (Nelson 1994). A bony suborbital stay extending pos- teriorly from the third infraorbital to the preopercle, has been reported as the unique morphological character defining this fish group (Washington et al. 1984a ). This order shows a confuse taxonomy and their phylogenetic relationships are poorly known. Among these fishes, the family Scor - paenidae has approximately 400 species widely distributed in shallow waters of tropical and tem- perate seas, living on rockies, corals, sandys and algae's bottoms (Nelson 1994). Although most of the cytogenetic works in this family describe only the gross karyotype structure, some others have described the heterochromatin distribution pattern (C-banding), the nucleolar organizer regions (Ag-NOR) (Thode et al. 1985, Yokoyama et al. 1992) and replication bands (Giles et al. 1988) in few species. The scorpaenids show a significative chromosome diversity, specially the subfamily Scorpaeninae, whose diploid numbers vary 34 to 48 (Nogusa 1960, Cataudella et al. 1973 , among others). In the present study the Giemsa staining karyotypes, the nucleolar organizer regions (Ag and mithramycin staining) and the C-banded heterochromatin of Scorpaena brasiliensis and S. isthmen - sis are described, and also showed a significant chromosome divergence observed between them which may be related to centric fusion and pericentric inversions. Materials and methods Two individuals (one female and one male) of Scorpaena brasiliensis and 21 of Scorpaena isthmensis (8 females, 12 males and one individual of sex undetermined) were studied. The speci- mens were collected in Guanabara Bay, in the coast of Rio de Janeiro State (Brazil), by scuba div- ing in Botafogo Creek (22•‹56'34"S-43•‹9'42"W) and with a trawlnet in Central Channel (22•‹49'24"S; 43•‹08'36"W-22•‹47'48"S; 43•‹08'36"W). The specimens were deposited in the Ichthyological Collection of the Museu Nacional, of Uni - versidade Federal do Rio de Janeiro, and registered with numbers MNRJ 13566—MNRJ 13567 (Scorpaena brasiliensis) and MNRJ 13568 to MNRJ13580—MNRJ 14263 to MNRJ 14266 (Scor - paena isthmensis). The chromosome preparations were made by direct method from the kidney cells as described by Brum et al. (1992) or sometimes a short term culture method for solid tissue was applied (Fenocchio et al. 1991). In order to improve the quality and quantity of metaphase cells, an yeast suspension was administered before the direct preparations (Lee and Elder 1980). The nucleolar or- ganizer regions were visualized by silver nitrate (Howell and Black 1980) and mithramycin A stain- ing (MM), which was carried out as described by Schmid (1980 ). The C-banding was performed 398 Margaret Maria de Oliveira Correa and Pedro Manoel Galetti Junior Cytologia 62 according to the method by Sumner (1972). Results Scorpaena brasiliensis showed 2n=46, consisting of 14 metacentric-submetacentrics (M-SM) and 32 subtelocentric and acrocentric (ST-A) chromosomes, fundamental number (FN)=60, in males and females (Fig. 1). S. isthmensis showed 2n=40 and FN=54, consisting of 14 M-SM and 26 ST-A in both sexes (Fig. 2). C-banding revealed the presence of small centromeric heterochomatin blocks in almost all chromosomes of both species (Figs. 3, 4). In S. brasiliensis, a conspicuous heterochromatic block was also observed on the short arm of a medium-sized submetacentric pair. The silver nitrate and Fig . 1 . Karyotype of Scorpaena brasiliensis. The bar equals 10ƒÊm. 1997 Chromosomal Diversity in Scorpaenids from Brazil 399 Fig . 2 . Karyotype of Scorpaena isthmensis. The bar equals 10ƒÊm. mithramycin A staining revealed two submetacentric chromosomes (pair 2) bearing telomeric NOR sites on the short arms in S. brasilensis, apparently coincident with the conspicuous telomeric hete- rochromatic blocks (Fig. 3A, B). In S. isthmensis these sites were identified in a comparable chro- mosome region, but in the chromosome pair 5 (Fig. 4A, B). Moreover, size heteromorphisms be- tween the homologous NOR were detected in S. brasiliensis (Fig. 3A, B). Discussion The results presented in this study, along with data from the literature, show the existence of a great karyotypical diversity in the family Scorpaenidae. This is specially true for the subfamily 400 Margaret Maria de Oliveira Correa and Pedro Manoel Galetti Junior Cytologia 62 A B Fig. 3. C-banding in Scorpaena brasiliensis; A) NOR-bearing chromosomes (Pair 2) through Ag stain- ing; B) NOR-bearing chromosomes through MM staining. Scorpaeninae. This subfamily has probably undergone a great number of chromosomal rearrange- ments during its evolution. However, some of its species still show a diploid number of 48 chromo- somes. On the other hand, in the subfamily Sebastinae, the chromosomal evolution has been appar- ently more conservative. In this subfamily, 90% of the species have 48 chromosomes, with FN of 50 in 80% of these species (Table 1). Ohno (1974), in a classic study on fish cytogenetics, suggested that the ancestral karyotype of the Teleostei has 48 acrocentric chromosomes. Recently, Brum (1995) observed that this karyotype could be an evolutionary novelty of the Clupeocephala shared by the Euteleostei. Among the latter, the Scorpaeniformes (Nogusa 1960, Arai and Fujiki 1978, Yokoyama et al. 1992, present study, among others) and the Perciformes (Denton 1973, Ohno 1974, Sola et al. 1981, Cano et al. 1982, Oliveira et al. 1988, Brum 1995, among others) have apparently maintained that basic structure in their karyotypes. According to Regan (1913), the family Scorpaenidae, which is the most generalized group in the Scorpaeniformes, is strictly related to the more generalized Perciformes (e.g. Serranidae), and it has been suggested that the families Scorpaenidae and Serranidae share a common ancestor (Frei- hofer 1963). Moreover, it has been suggested that the primitive, karyotype of the Serranidae has 48 acrocentric chromosomes (FN=48), with some species presenting 2n=48 and FN=50 (Aguilar and Galetti 1997, among others). Thus, up to now, the hypothesis of a 2n=48, FN=50 karyotype as the ancestral state in the Scorpaeniformes seems consistent. This karyotype has remained almost un- changed in the Sebastinae, undergoing more remarkable modifications in the Scorpaeninae. This difference corroborates the idea that Sebastinae is a more generalized group, while the Scorpaeni- nae is a more derived one (Matsubara apud Washington et al. 1984a). Chromosomal markers, other 1997 Chromosomal Diversity in Scorpaenids from Brazil 401 A B Fig. 4. C-banding in Scorpaena isthmensis; A) NOR-bearing chromosomes (pair 5) through Ag stain- ing; B) NOR-bearing chromosomes through MM staining. than 2n and FN, should be investigated in order to exclude thoroughly the possibility of convergent evolution in the karyotypes of Scorpaenidae and Serranidae. Taking into account the presumable primitive karyotye of the Scorpaeniformes (2n=48 and FN=50), the two species of Scorpaena here studied have diminished their karyotype structure through several chromosome rearrangements. In S. brasiliensis at least one centric fusion and six pericentric inversions have apparently taken place (Fig. 1). In S. isthmensis three centric fusions and at least three or four pericentric inversions may have occurred (Fig. 2). In the remaining subfamilies of Scorpaenidae which show wide divergence in the karyotype structure and diploid number almost invariable (Table 1), pericentric inversions must have mainly occurred. These three subfamilies are believed to be more specialized than the Sebastinae (Wash- ington et al. 1984b), and the occurrence of sexual chromosomes in Hypodytes rubripinnis (Ueno and Kong 1992) corroborates the hypothesis of derivation in their FN values. Silver nitrate staining in 10 scorpaenid species have demonstrated up to two NOR-bearing chromosomes per complement, although the NOR sites can be found in different chromosomes among species (Thode et al. 1985, Yokoyama et al. 1992). In both Scorpaena species here studied only two chromosomes were detected bearing NORs, and due to their morphological similarity, it is quite probable that they are homologous between both species. The size heteromorphism between homologous NORs detected in S. brasiliensis, have also been reported in S. porcus and S. notata (Thode et al. 1985) and is a widespread phenomenon among fishes (Foresti et al. 1981). Except to the Ag-NOR sites, no other chromosome regions showed positive bands after 402 Margaret Maria de Oliveira Correa and Pedro Manoel Galetti Junior Cytologia 62 Table 1 . Summary of the karyotypic data of the family Scorpaenidae mithramycin staining in both Scorpaena species. Similar results have been several times described in fishes (e.g. Schmid and Guttenbach 1988). Few heterochromatin segments were detected in both species and reductions of these segments also appear to have an important
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