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Hexaploidy in Yellowfish Species (Barbus, Pisces, Cyprinidae) from Southern Africa

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Hexaploidy in Yellowfish Species (Barbus, Pisces, Cyprinidae) from Southern Africa Journal of Fish Biology (1990) 37, 105-1 15 Hexaploidy in yellowfish species (Barbus, Pisces, Cyprinidae) from southern Africa L. K. OELLERMANNAND P. H. SICELTON* J.L.B. Smith Institute of Ichthyology, Private Bag 1015, Grahamstown 6140, Republic of South Africa (Received 18 July 1989, Accepted I February 1990) Five small-scaled yellowfish (large Burbus spp.) from southern Africa are shown to have modal 148 or 150chromosomes. Themajority ofcyprinid species have 2N = 50chromosomes, indicating that the yellowfish karyotype is hexaploid in origin. However, as there is no indication that the species are unisexual or that normal reproduction occurs by any means other than bisexual fertilization, the yellowfish karyotype is considered to have reverted to a diploid condition. Key words: karyology; yellowfish; Burbus; hexaploidy; southern Africa. I. INTRODUCTION The name ' yellowfish ' is given to several large-sized Burbus species in southern Africa (Jubb, 1967). The yellowfish fall into two groups on the basis of scale size. The group with smaller scales consists of five recognized species distributed in the Orange River system and adjacent drainage systems (Skelton, 1986). The second group, with larger scales, consists of two species, Barbus rnarequensis Smith, 1841 and Barbus codringtonii Boulenger, 1908, distributed in east coastal drainages from the Phongola River to the Zambezi River system. This study concerns the five small-scaled yellowfish species, viz. Barbus cupensis Smith, 1841 from the Olifants River system, Burbus aeneus (Burchell, 1822) and Burbus kimberleyensis Gilchrist & Thompson, 1913 from the Orange River system, Barbus polylepis Boulenger, 1907 from the Limpopo, lncomati and Phongola River systems, and Burbus natalensis Castelnau, 1861 from the rivers of Natal. Yellowfish taxonomy generally is complicated by the characteristically large degree of intraspecific variation of the species (e.g. Gilchrist & Thompson, 19 13; Barnard, 1943; Groenewald, 1958; Jubb, 1967). Even at present the close similarity between certain species, e.g. B. aeneus and B. cupensis, and B. polylepsis and B. natalensis, and the fact that they are distributed allopatrically raises doubt on their taxonomic status. Yellowfish are popular angling species and most of the species have been successfully bred artificially and raised in captivity. Both B. aeneus and B. natalensis have been translocated beyond their native range in southern Africa (de Moor & Bruton, 1988). Understanding the karyological and genetic structure of these fishes may help to resolve these taxonomic questions. The karyology of fishes has advanced little beyond the description of gross chromosome morphology. A change in a gene sequence causing a phenotypic variation in the species may not necessarily alter the gross shape of the chromo- some. Gold (1980) encountered this problem in the speciose cyprinid genus *Author to whom correspondence should be addressed. 105 0022-1 I12/90/070105+ 11 $03.00/0 0 1990 The Fisheries Society of the British Isles 106 L. K. OELLERMANN AND P. H. SKELTON TABLEI. RUSI catalogue number, localities and sources of the small-scaled yellowfish specimens used for karyological studies Species No. Cat. no. Locality Collector ~ ~ Burbus aeneus 15 28407/8 Kubusie R., L. K. Oellermann Kei R. system, Ciskei B. cupensis 15 28403/4 Olifants R., CDNEC* S.W.Cape B. kimberleyensis 8 28400/1 Le Roux Dam, L. K. Oellermann and Vaal R. Amalinda hatchery CDNEC B. nutalensis 11 28402 MgeniR. Natal Parks Board B. polylepis 4 28406 DorpsR., L. K. Oellermann Olifants-Limpopo R. system ~ *Cape Department of Nature and Environmental Conservation Notropis: 22 of the species he studied displayed similar gross karyotypes. Although much of fish karyology is aimed at recording karyotypes, a growing number of workers are using karyology as a tool for studying taxonomy, as well as genetics, hybridization, polyploidy, sex and sex reversal. Taxonomic studies on the small-scaled yellowfish, involving morphometric and meristic characters and osteology, are in progress (Oellermann, 1989). Karyology has been used with varied success in fish taxonomy (e.g. Campos & Hubbs, 1973; Gjedrem et al., 1977; Loudenslager & Thorgaard, 1979; Rukhkyan, 1984; Anemiya & Gold, 1988). There is a fast-growing field of data on cyprinid karyology inter- nationally, but so far no information has been published on African cyprinids. This study initiates the karyology of southern African cyprinids in order to provide (a) karyological descriptions for comparative studies, (b) cytotaxonomic data on the species, and (c) complementary data that could test hypothesized phylogenetic relationships based on anatomical or electrophoretic characters. 11. MATERIALS AND METHODS The small-scaled yellowfish specimens used for this study were mostly sexually immature juveniles, ranging from about 6 months to 2 years old. Specimens were collected in the field by means of seine nets and gill nets, or were provided by Provincial Nature Conservation authorities from hatchery-held stocks. The number of specimens used and their collection data is given in Table 1. All specimens used were preserved and are deposited in the fish collection of the J.L.B. Smith Institute of Ichthyology (RUSI). CHROMOSOME ISOLATION AND ANALYSIS Chromosomes were isolated using a modification of the method described by Kligerman & Bloom (1977) as follows. The specimen is injected with 0.01 ml gg ' body weight of 0.1 % colchicine and placed for 4-6 h in well-aerated water which is 2-5" C above the holding temperature. It is then killed by pithing and whole gill arches removed, teased apart and placed in 0.4% KCl hypotonic solution for 40 min. The gill tissue is fixed in Carnoy solution (I part glacial acetic acid : 3 parts absolute methanol) and then macerated in 50% acetic acid. A suspension of tissue is released from a bulb dropper onto a heated (40" C) glass slide from which it is withdrawn after 30 s. After drying, the slides are stained in fresh 5% Giemsa stain for 1&15 min, rinsed, dried and mounted. HEXAPLOIDY IN BARBUS SPP. 107 Preparations were viewed under a Nikon Optiphot compound light microscope. Photomicrographs were taken through an oil immersion x 100 objective lens. The large number and small size of the yellowfish chromosomes made them difficult to count. The small size also restricted our ability to subdivide the chromosomes into any classes other than bi-armed and uni-armed. Several authors have encountered similar problems (e.g. Schwartz & Maddock, 1986) and have concluded that further subdivision is completely arbitrary. The method used to count the chromosomes was as follows. A transparent sheet was placed over the photograph, and each chromosome was traced onto the sheet, using water-insoluble marker pens. The chromosomes were classed and coloured separately as either bi-armed or uni-armed and then counted. Each photographed chromosome spread was recounted at least three times. The ten most well defined chromosome spreads from each species were used to estimate the Fundamental Number (FN) of the karyotype as follows: FN=Z(n,)+n,, where n,is the number of bi-armed and nz is the number of uni-armed chromosomes. The final karyotype for each species was taken from the best defined chromosome spread [e.g. Fig. l(a)]. The chromosomes were cut from the photograph and paired according to their type and size into a photokaryotype. These karyotypes were traced on a light table and drawn for presentation [Fig. I(b)]. 111. RESULTS A representative chromosome spread of B. kimberleyensis is shown in Fig. l(a) and the drawn karyotype for this species in Fig. l(b). The small-scaled yellowfish species can be divided into two groups on the basis of their diploid (2N) chromo- some number. Barbus capensis, B. natalensis and B. polylepis each have 150 chromosomes, while B. aeneus and B. kimberleyensis have 148 chromosomes. Chromosome number varied greatly, mostly due to chromosomal loss from the spreads (Fig. 2). The large number of chromosomes caused great difficulty in spreading the chromosomes sufficiently for counting without losing chromosomes from the spread. Difficulties experienced in obtaining accurate and consistent counts were overcome by applying the counting method described in the ' Materials and Methods ' section. Counts from the slides under the microscope were usually at least 10 chromosomes less than those from the photographs. Mistaken chromo- some identification (e.g. a bi-armed chromosome counted as two uni-armed chromosomes) and chromosome imports from other spreads probably resulted in chromosome counts higher than the modal number. No sexual dimorphism was observed within the small-scaled yellowfish karyo- types. Any major dimorphism in chromosome pairs was probably masked by the multiple chromosomes introduced by polyploidy. The FN for each species was relatively similar, and ranged from 208 in B. cupensis to 196 for B. aeneus (Table 11). The modal chromosome arm numbers (Table 11) differed marginally between the species but must, at this stage, be accepted with caution due to the difficulties in obtaining clear chromosome impressions. The variation in individual counts of bi-armed chromosomes in 10 spreads from which the modal number was derived is given in Table 111. IV. DISCUSSION The extremely high number of chromosomes is a striking feature for the yellow- fishes. An extensive literature survey on the karyotypes of cyprinids (see NOllaWS 'H 'd (INV NNVNXIB71BO ')I '1 80 I HEXAPLOIDY IN BARBUS SPP. 109 30401 30 - 20 - 10 - 0 I I Chromosome number 50--I (e 40 t 10 l---4L.090 100 110 120 130 140 150 160 Chromosome number FIG.2. Percentage spreads with various chromosome numbers for (a) Barbus aeneus (n= 36 spreads), (b) B. capensis (n= 40 spreads), (c) B. kimbedeyensis (n= 33 spreads), (d) B. nafa1ensi.s(n = 28 spreads) and (e) B. po/y/epis (n = 3 I spreads). was duplicated in the cyprinid species which they used as examples of the tetraploid group. Thus, at some stage during their evolution these species experienced a polyploidic event (Mayr et al., 1986) which doubled their chromosome numbers to produce a tetraploid condition.
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