New Developments in Vertebrate Cytotaxonomy

NEW DEVELOPMENTS IN VERTEBRATE CYTOTAXONOMY III. KARYOLOGY OF BONY FISHES: A REVIEW L.SOLA, S.CATAUDELLA & E.
CAFANNA
Institute of Comparative Anatomy, University of Rome, Via Alfonso Borelli 50, 00161 Rome, Italy
Received andAccepted Octobber 2.1979
Introduction Several years after the introduction of modern cytological techniques in the karyological study of the bony fishes, an attempt may be made to take stock of the results obtained, the problems raised and the difficulties encountered by workers engaged in this type of investigation. The first limitation is set by the value of the karyotype itself. Because it is purely morphological and its immediate adaptive value is unknown, karyotype transformations are difficult to interpret in analyses of populations of the same speciesor of related species. When related species are being compared, similar karyotypes are considered indicative of a relatively recent separation, whereas widely differing karyotypes are believed to indicate an earlier separation.However, this assumption is not always valid. In the bony fishes a karyotype composed of 48 acrocentric chromosomes occurs not only in related species but also in species that are phylogenetically further apart. In some taxa the karyotype therefore appears neutral with respect to both speciation and phyletic evolution (Fig. 1) and it is therefore often impossible to establish a definite correlation between karyotype and karyotype transformations and to identify the processesinvolved. Although this situation diminishes the immediate value of the karyological evidence, it raises a series of questions that, when set in the right problematical context, make karyological analyses extremely interesting .
* This is the third in a series of articles, by different authors, to appear in this journal under the editorship of Professor Chiarelli, supplementing the data published in Cytotaxonomy and Vertebrate Evolution (A.B. Chiarelli & E. Capanno, eds, Academic Press, London & New York, 1973) - Eds.
The presence of diversified karyotypes in restricted systematic groupings poses the following questions. - Do the diversifications indicate that karyotype transformations play a primary role in speciation? - Do they indicate a selective pressure on certain chromosome arrangements, which maintain specific associations? - Or are these arrangements random, statistical events, not subject to control by selection, except perhaps during the initial phase in which they may contribute to the formation of isolating mechanisms; the taxonomist may therefore consider them as indicative of the time elapsing since phyletic separation? - Or can the presence of diversified karyotypes in systematic groups which are considered to be related be interpreted as evidence of convergent evolution? The significance of the homogeneous karyological situations may also become apparent when related to the above questions. Answers to some of these questions are being outlined by recent studies. For example, new evidence suggests that the karyotype can play a primary role in speciation (cf. White, 1978) contrary to Mayrs original hypothesis (1963) which stated that only polypIoidy and geographical isolation lead to the formation of new species. In some species of insects (Neanura, Cassagnau 1974), mollusks (Thais or Nucella, Staiger, 1954; Bantock & Cockayne 1975) and rodents (Thomornys, Berry & Baker, 1971; Spalax, Nevo 1972, 1974) various types of chromosome rearrangements are directly adaptive to peculiar ecological situations. Fish comprise a polyphyletic group which has undergone an enormous expansion - the number of existing species totals about 25,000, i.e. about 60% 285
Genetica 54,285-328 (1981): 0016-6707/8l/0543-0285 $8.80. 0 Dr. W. Junk B.V. Publishers, The Hague. Printed in The Netherlands.
of all Vertebrate species. The capacity of this group for adaptation has led to the colonisation of diverse aquatic habitats and to the occupation of extremely specialized niches. This species explosion cannot be explained wholly in terms of geographical isolation, and chromosome mechanisms probably played a part in speciation. Although only a small number of species have been studied (850-900 out of 25,000), evidence has been found of evolution by polyploidization (inside the Cyprinidae family [Wolf et al. 19691 and the Salmonidae if referred to the Clupeidae and Engraulidae [Ohno et al. 19691). In some groups currently considered to be in expansion, such asthe Salmonidae (Fukuoka 1972, Thorgaard 1976, 77, Ohno et al. 1965), cases of chromosomal polymorphism are known and they sliould be followed closely. In the Poecilidae speciation may occur by interspecific hybridization. This apparently applies to Poecilia formosa whose hybrid origin is widely accepted on the basis of morphological and karyological characteristics (Hubbs & Hubbs, 1932; Prehn & Rasch, 1969). The stabilisation of the original hybrid form was, however, achieved by gynogenesis and does not suggest significant evolutionary possibilities. The Poecilidae include other hybrid forms, for example, the three populations of Poeciliopsis moAacha-Zucida in northern Mexico described by Schultz (1967,1969). The condition of gynogenesis is present in one population, but in a situation of allotriploidy which could represent a step in the direction of an allotetraploid stabilization, a condition which would allow a return to sexual reproduction. The bony fishes provide excellent material for cytogeneticists working on sex determination problems. They display almost all possible forms of sexuality, ranging from simultaneous or successivehermaphroditism to gonochorism, with early or late sexual differentiation, to the condition of gynogenesis with occasional evidence of true parthenogenesis, The karyological picture is equally varied. In most species sex chromosomes are absent or at least not morphologically identifiable. However, some species are known to exhibit male heterogamety , whereas others have female heterogamety. Furthermore alloFig.1. Metaphase plates, with 48 acrocentric chromosomes, of three phylogenetically unrelated species: (a) Gambusia uffinis, Cyprinodontiformes; - (b) Mugil cephalus, Mugilidae; Perciformes; - (c) Pagellus erythrinus, Sparidae, Perciformes. patric either populations of the same species may display female or male heterogamety (e.g. in Xiphomaculatus
phorus
[Kallman, 19731, in which the sex

are identified by means of pigment gene
chromosomes
286
markers showing a sex-linked hereditary pattern). Uyeno & Miller (1971, 1972) described several Mexican populations of unidentified species belonging to the Cyprinodontidae; a multiple sexchromosome system was apparently found. Thorgaard (1977) discovered a primitive differentiation stage of sex chromosomes in Salmo gairdneri. The interest of cytogeneticists and evolutionists is closely connected with that of ichthyologists in the study of hybrids. In the freshwater bony fish (Cyprinidae, Centrarchidae , etc .) evidence is frequently found of interspecific and even intergeneric hybrids where sympatric populations of different species exist. The development of fish culture has resulted in numerous experiments on hybridization to obtain new strains which are more resistant, quicker-growing, etc. Karyological knowledge allows the parent species to be recognised in the case of natural hybrids and gives some indication of the likelihood of successin the case of artificial cross-breeding. Further it may help to make predictions concerning hybrid fertility and the interactions of the two parental sets (Fig. 2). Although it is not possible to draw absolute conclusions about phyletic relationships from the karyological comparison of different species, karyotype analyses may supplement other taxonomic evaluations. The systematic definition of a species depends on a thorough morphological analysis, and the karyotype is only one characteristic among several. However, it is often necessary, and certainly correct to include this. Sometimes karyological analyses are a useful work and study tool for ichthyologists, for example, in the classification of related species. Often there are no obvious morphological characters allowing definite identification and ecotypism occurs frequently in many fish species. Identification of larval forms and early stages is often difficult because these resemble one another and are often different from their adult stage. Therefore, morphological observations are not sufficient to identify the species unless the developmental stages can be followed, which is usually difficult. In such cases a knowledge of the karyotype and the existence of diversified ktiological situations, if any, can provide a precise diagnostic criterion suitable for solving identification problems (Fig. 3). However, this point also illustrates the most important of the present limitations of the karyological studies of bony fishes. If karyotypes are found to be
morphologically diversified then they can be used as a diagnostic criterion to distinguish the species analysed. On the other hand, if they are morphologically similar the cytotaxonomic evidence loses some of its importance, as similarity of karyotypes may be superficial only. Although the genetic material is organised into chromosomes, the latter only represent its form, i.e. the outer shell, and tell us nothing about the DNA rearrangements, except for such dramatic events as translocations and inversions or fusions between whole chromosomes. In the fishes, an accurate morphological characterisation of the chromosomes is hindered by the large
Fig. 2. Karyotypes of fish hybrids: (a) Salmo trutta x Salvelinus fontinalis. 2n = 82 and marker chromosomes in single copies (arrows) indicating the combination of the two parential haploid complements; - (b) Salmo gairdneri x Salvelinus fontinalis, 2n = 96 and N.F. = 141 suggesting aneu-triploidy, possibly resulting from combination of paternal haploid complement with diploid complement of the maternal species known to present Robertsonian variability. 287
C
Fig. 3. Karyotype 2n = 34. diversity
in
the genus
Scorpaena:
(a)
S. scrofa
L., 2n
= 46;
(b)
S. porcus
L., 2n
= 42;
- (c) S. notata
Raf.,
number and the small size of the chromosomes. The banding techniques commonly used in the karyological study of other Vertebrate classes have therefore found little application. Only C-banding was tried on a few species. However, the use of all banding techniques (G-, Q-, R-, etc.) must be encouraged and expanded, so that cytotaxonomic evidence may become a useful work tool for ichthyologists and so that a more correct evaluation may be made of the changes occurring inside the chromosome and of the differentiations between the karyotypes of-related species. Furthermore, the karyological evidence must be integrated with quantitative and qualitative evaluations of DNA contents, autoradiographic and in situ hybridization evidence, and electrophoresis data on isoenzymes, especially when populations with chromosome polymorphism are being studied. Only by such multiple contributions can the complex process of speciation be reconstructed. What, then, is the significance of updating this list following the excellent reviews by Eun Ho Park (1974) and Ojima et al. (1976) and the book on fish cytogenetics by Ohno (1974)? Because few species have been studied and little
288
is known of the details of karyotypes, it may seem of limited value to present a new list of species. However, only by collecting karyological data will the ma+ lines of the karyological evolution of the class of bony fish be worked out, and only in this way can groups be identified whose cytogenetic characteristics are useful in the construction of cytogenetic speciation models. Research workers are therefore invited to study systematically homogeneous groups, to pay careful attention to the geographical distribution, and to improve the level of research by adopting the techniques described above. The bibliography is believed to be complete up to June 1979 but it has not been possible to find the abstracts for all the references given. The list includes 8 10 species arranged systematically after Greenwood et al. (1966).
Acknowledgement
We wish to thank Mr. Pier Luigi Piccioni and Mr. Luciano Veroli for their kindest help in preparing the manuscript.
References
Bantock, C.R. & WC. Cockayne (1975). Chromosomal polymorphism in Nucella lapillus. Heredity 34: 231-245. Berry, D.L. & R.J. Baker (1971). Apparent convergence of karyotypes in two species of pocket gophers of the genus Thomomys (Mammaha, Rodentia). Qtogenetics 10: l-9. Cassagnau, P. (1974). Les chromosomes polytenes de Neanura monticola Cassagnau (Collemboles) I. Polymorphisme Bcologique du chromosome X. Chromosoma 46: 343-363. Fukuoka, H. (1972). Chromosome-number variations in the raimbow trout (Salmo gairdneri irideus Gibbons). Jap. J.
Genet. 47: 455458.
K.R. Lewis pp. 139-147. Ohno, S. (1974). <Animal cytogenetics.

chordata, Cyclostomata and Pisces.
Chord&a
I. Proto-
Greenwood, P.H., D.E. Rosen, S.H. Weitzman & G.S. Myers (1966). Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bull. Am. Museum
nut. Hist. 131 (4): 393403.
Hubbs, C.L. & L.C. Hubbs (1932). Apparent parthenogenesis in nature, in a form of fish of hybrid origin. Science 76: 628630. Kallman, K.D. (1973). The sex-determining mechanism of the platyfish Xiphophorus maculatus. In: Genetics and mutagenesis of fish ed. J.H. Schroder. Springer-Verlag Berlin 1973: 19-28. Mayr, E. (1963). Animal species and evolution. Harvard University Press. Nevo, E. & C.R. Shaw (1972). Genetic variation in a subterranean mammal Spalax ehrenbergi. Biochem. Genet. 7: 235-241. Nevo, E. & A. Shkolnik (1974). Adaptive metabolic variation of chromosome forms in mole rats, Spalax. Experientiu 30: 724-726. Ohno, S., C. Stenius, E. Faisst & M.T. Zenzes (1965). Postzygotic chromosomal rearrangements in rainbow trout (Salmo irideus Gibbons). Cytogenetics 4: 117-129. Ohno, S., J. Muramoto, J. Klein & N.B. Atkin (1969). Diploid-tetraploid relationship in clupeoid and salmonoid fish. In: Chromosomes today II. Eds. C.D. Darlington & Table 1 Species n 2n Karyotype
BerlinStuttgart 1974. Ojima, Y., K. Ueno & M. Hayashi (1976). A review of the chromosome numbers in fishes. La Kromosomo II, 1: 1947. Park, E.H. (1974). A list of the chromosome numbers of fishes. Coil. Rev. Coil. Lib. Arts & Sciences, Seoul Nut. 1 Univ. 20 (l-2): 346-372. Prehn, LM. & E.M. Rasch (1969). Cytogenetic studies of Poecilia (Pisces). I. Chromosome numbers of naturally occurring poeciliid species and their hybrids from eastern Mexico. Can. J. Genet. Cytol. 11: 880-895. Schultz, R.J. (1967). Gynogenesis and triploidy in the viviparous fish Poeciliopsis. Science 157: 1564-1567. Schultz, R.J. (1969). Hybridization, unisexuality and polyploidy in the teleost Poecilopsis (Poecilidae) and other Vertebrates. Am. Nat. 103: 605619. Staiger, H. (1954). Der Chromosomendimorphismus beim Prosobranchier Purpura lapillus in Beziehung zur tikologie der Art. Chromosoma 6: 419478. Thorgaard, G.H. (1976). Robertsonian polymorphism and constitutive heterochromatin distribution in chromosomes of the rainbow trout (Salmo gairdneri). otogenet. Cell Genet. 17 (4): 174-186. Thorgaard, G.H. (1977). Heteromorphic sex chromosomes in male rainbow trout. Science 196: 900-902. Uyeno, T. & R.R. Miller (1971). Multiple sex chromosomes in a Mexican cyprinodontid fish. Nature 231: 452453. Uyeno, T. & R.R. Miller (1972). Second discovery of multiple sex chromosomes among fishes. Experientia 28: 223-225. White, M.J.D. (1978). Modes of speciation. W.H. Freeman, San Francisco. Wolf, Il., H. Ritter, N.B. Atkin & S. Ohno (1969). Polyploidization in the fish family Cyprinidae, order Cypriniformes. I. DNA content and chromosome sets in various species of Cyprinidae. Humangenetik 7: 240-244.
Borntraeger,
Number of arms*
Heterogametic sex
Reference
OSTEICHTHYES Actinopterygii SUPERORDER CHONDROSTEI

Order A CIPENSER IFORMES
Family Acipenseridae
Acipenser naccurii Acipenser sturio Huso huso Scaphirhvnchus pkztyrhynchus
239 i 7 116+4 116+4 112 *
Fontana Fontana Fontana Ohno et m,sm = arms; st, a = 1 arm.
et al. 1974 et al. 1974 et al. 1974 al. 1969b
* When in brackets, number of arms calculated by the present authors as following:
289
Table 1 (Continued) Species n 2n Kwotype Number of arms Heterogametic sex Reference
Family Polyodontidae Polyodon spathula SUPERORDER HOLOSTEI Order LEPISOSTEIFORMES Family Lepisosteidae Lepisosteus productus Order AMIIFORMES Family Amiidae Amia calva Teleostean fishes SUPERORDER ELOPOMORPHA Order ANGUILLIFORMES Suborder Anguilloidei Family Anguillidae Anguilla anguilla Anguilla anguilla A ngu illa anguilla Anguilla australis Anguilla australis A nguilla japonica Anguilla japonica Anguilla rostrata Anguilla rostrata Family Moringuidae Moringua linearis Family Congridae Astroconger myriaster Conger myriuster Family Ophichthidae Ophiswus macrorhynchus Pisoiidonophis zophistius SUPERORDER CLUPEOMORFA Order CL VPEIFORMES Suborder Clupeoidei Family Clupeidae Alosa pseudoharengus Clupea harengus Clupea harengus harengus n. membras Clupea harengus pallasi n. maris albi Clupea pallasi 290
120
32m+8sm+8a+ 72michrochromosomes
Dingerkus et al. 1976
68
28m + 14a + 26minute
Ohno et al. 1969b
46
20m + 26a
(66)
Ohno et al. 1969b
38 38 38 38 38 38 38 38 38
24m + 8st + 6a 12m+lOsm+16a 22m, sm + 16a 20m, sm + 18a 22m, sm + 16a 20m, sm + 18a 12m+8sm+18a 14m + 6st + 18a 22m, sm + 1&a
(63
(60)
(60)
(58)
(60)
(58) (58) (52)
(601
Q? Q Q Q? -
Chiarelli et al. 1969a Passakas 1978 Sola et al. 1980 Nishikawa et al. 197 1 Sola et al. 1980 Nishikawa et al. 1971 Park et al. 1979 Ohno et al. 1973 Sola et al. 1980 Subrahmanyah et al. 1971
50
38 38 38 38
12m+8sm+l8a 14m + 4sm + 20a 20m + 14sm + 4st lOm+16sm+4st+8a
(58)
56 16 68
Park et al. 1979 Nishikawa et al. 197713 Nishikawa et al 1977b Nishikawa et al. 1977b
48 52 54 52 52
48a
(48)
66-70
8m + 44a
(Z)
Mayers et al. 1969 Roberts 1966 Skvortsova 1975 Skvortsova 1975 Ohno et al. 1968
Table 1 (Continued) Species n 2n Karyotype Number of arms (48) Hetero gametic sex Reference
Family Engraulidae
Engraulis mordax
48
48a
Ohno et al. 1968
SUPERORDER OSTEOGLOSSOMORPHA
Order OSTEOGLOSSIFORMES Suborder Osteoglossoidei
Family Osteoglossidae
Arapaima gigas Osteoglossum bicirrhosum Scleropages formosus Suborder Hiodon Notopteroidei
56 56 50
4m + 12sm, st + 40t lsm+ lst+54a 4sm + lost + 36t
Urushido et al. 1975 Uyeno 1973 Urushido et al. 1975
Family Hiodontidae
alosoides
50 48 42 42 42 42 34 42
40m, sm + I Ost 12m+36a 42a 42a 42a 4sm + 30a 42a
100 60 42 (42) (42) 38 42
Uyeno 1973 Nayyar 1965 Uyeno 1973 Srivastava et al. 1964 Urushido et al. 1975 Prasad et al. 1976 Uyeno 1973 Uyeno 1973
Family Notopteridae
Notoptenrs Notopterus Notopterus Notopterus Notoptenrs Papyrocranus Xenomystus chitala chitata notopterus notopterus notopterus afer nigri
SUPERORDER PROTACANTHOPTERYGII
Order SALMONIFORMES Suborder Salmonoidei
Family Salmonidae
Brachymystax Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Coregonus Oncorhynchus lenok albula artedii clupeaformis hoyi lavaretus lavaretus lavaretus lavaretus baeri nasus nasus oxyrhynchus peled peled pidschian pidschian reighardi ussuriensis zenithicus gorbuscha
90 80 80 80 80 80 96 80 80 96 80 96 80 80 96 80 80 80(82) 80 52
26m, sm + 64a 16m + 64a 16m + lost + 54a 20m + 8st + 52a 10m + 8st + 62a 16m + 64a 32m, sm + 64a 12(18)m + 68(62)a 12m+&m+54a 96a 12m + 68a 96a 12m, sm + 68a 12(18)m + 68(62)a 96a 18(12)m + 62(68)a 12m + 12st + 56a lO(12)m + 8m + 32(30)a 10m + 8 sm + 62a 52m
116 96 106 108 98 96 118 92(98) (96) 96 92 96 92 92(98) 96 98(92) 104 100 98 104
Kang et al. 1973a Nygren et al. 1971a Booke 1968 Booke 1968 Booke 1968 Svardson 1945 Karbe 1964 Nygren et al. 1971c Prokofieva 1934 Karbe 1964 Nygren et al. 1971b Karbe 1964 Viktoroski 1964 Nygren et al. 1971b Karbe 1964 Nygren et al. 1971b Booke 1968 Viktoroski et al. 1978 Booke 1968 Simon 1963 291
Table 1 (Continued) Species n 2n Karyotype Number of arms 100 106 112 106 124 124 104 104 104(102) 104 102 124 124 128 104 100 100 100 100 100 100 106 W-9 100 106 (98)
104 104 106 104 104 104 156 106 104 104 104 104 104 104 12 12 12 14 74 12 12 72 14
Hetero gametic sex
Reference
Oncorhynchus Oncorhynchus Oncorhynchus Oncorhynchus Oneorhynchus Oncorhynchus Oncorhynchus Oncorhynchus Oncorhynchus
gorbuscha ketu kisutch kisutch masou masou masou ishikawue nerka nerka nerka
52 14 60 60 66 66 58 58 58(56) 58(57) 56 66 66 68 68 ii: 78 64 14 18 58 80 80 58 80 68 64 64 60 60(58)
48m, sm + 4st 26m, sm + 6st + 42a 52m + 8a 38m + 8sm + 14a 34m, sm + 24st + 8a 34m, sm + 24st + 8a 44m, sm + 2st + 12a 44m, sm + 2st + 12a 44(45)m + 2st + 12(10)a 46m, sm + 10a 34m, sm + 24st + 8a 34m, sm + 24st + 8a 32m, sm + 28st + 8a 36m + 32a 12m + 16sm + 44a 10m + 8sm + 64a 12m + 1Osm + 56a 24m + 12sm + 28a 12m + 14sm + 48a 8m + 14sm + 56a 44m + 2sm + 2st + 10a 16m + 64a 20m + 60a 48m, sm + 10a 16m + 2sm + 62a 36m, sm + 32a 40m, sm + 24a 42m + 22a 44m + 16a 44(46)m + 2st + 14(10)a 44m, sm + 2st + 14a 66m, sm + 3st + 21a 44m, sm + 2st + 14a 44m + 4st + 12a 45m + 3st + lla 46m + 2st + 10a 39-46m, sm + l,2-26a 8m + 4sm + 48a 12m + 48a 16m, sm + 40a 16m, sm + 42a 16m, sm + 42a 16m, sm +41a 16m, sm + 40a 18m,sm+36a 14m + 2sm + 42a
Muramoto et al. 1974 Sasaki et al. 1968 Simon 1963 Uyeno 1972 Muramoto et al. 1974 Muramoto et al. 1974 Sasaki et al. 1968 Muramoto et al. 1974 Gorshkova 1978 Fukuokal972b Simon 1963 Muramoto et al. 1974 Muramoto et al. 1974 Muramoto et al. 1974 Simon 1963 Booke 1970 Booke 1970 Booke 1970 Booke 1970 Booke 1970 Booke 1970 Gold et al. 1975 Svardson 1945 Nygren et al. 1971c Miller 1972 Merlo 1957 Gold 1977 Gold 1977 Simon et al. 1963 Simon et al. 1963 Fukuoka 1972a Cuellar et al. 1972 Cuellar et al. 1972 Muramoto et al. 1974 Vasilev 1975 Thorgaard 1976,1977 Thorgaard 1976, 1977 Thorgaard 1976,1977 Bungenberg de Jong 1955 Ohno et al. 1965 Raicu et al. 1977 Prokofieva 1934 Svardson 1945 Boothroyd 1959 Rees 1967 Nygren et al. 1968b Roberts 1968 Roberts 1968 Roberts 1970 Grammeltvedt 1975
spring(summer)embryos
Oncorhynchus nerka nerka Oncorhynchus nerku nerka Oncorhynchus rhodurus macrostomus Oncorhynchus rhodurus rhodurus Oncorhynchus tschawtscha Oncorhynchus tschawtscha Prosopium abyssicolu Prosopium coulteri Prosopium cylindraceum Prosopium gemmiferum Prosopium spilonotus Prosopium wilhamsoni Salmo aguabonita Sulmo alpinus Salmo alpinus Salmo apache Sulmo curpio Salmo clarki clarki Salmo clarki henshawi Salmo clurki lewisi Sulmo guirdneri Salmo gairdneri Salmo gairdneri Sulmo gairdneri Sulmo gairdneri Sulmo gairdneri Salmo gairdneri SQlmO gairdneri Salmo guirdneri SQlmO gairdneri irideus Salmo irideus Salmo irideus Salmo saiar Atlantic salmon Eur. SQlmO solar Atlantic sahnon Eur. Salmo salar Atlantic salmon Amer. Salmo salur Atlantic salmon Eur. Salmo salar Atlantic salmon Eur. Salmo salar landlocked form Salmo salar culls Sulmo salar Atlantic salmon Amer. Salmo salar Atlantic salmon Eur.
64
90 60 58-60 60 59 58 60 58-65 60 60 60 56 58 58 51 56 54-56 58
d? d? d
292
Table 1 (Continued) Species n 2n Karyotype Number of arms 105 100 102 (96) (100) (96) (96) (94) 96 (100) 100 106 (100) (100) (98) 98 102 100 108 170 Hetero gametic sex Reference
Salmo gilae Salmo trutta Salmo trutta Salmo fario Salmo trutta fario Salmo trutta fario Salmo trutta fario Salmo trutta lacustris Salmo trutta trutta Salmothymus obtusirostris oxyrhynchus Salvelinus alpinus Salmo (Salvelinus) fontinalis Salvelinus fontinalis Salvelinus fontinalis Salvelinus fontinalis Salvelinus leucomaenis Salvelinus malma Salvelinus malma Salvelinus malma Salvelinus namaycush Stenodus leucichthys Thymallus thymallus Thymallus thymallus
56 80 77-82 84 80 80 80 80 80 82 78 84 84 84 84 84 82 80 82 84 74 102 102 80 66 66 60 72 70 70 69 82 92-106
49m, sm + 7a 20m + 60a 16m + 2st : 62a 14m + 6sm + 2st + 58a 16m + 2st + 62a 16m + 2st + 62a 12m, sm + 70a 16m + 2sm + 60a 16m + 68a 16m + 68a 16m, sm + 6st + 62a 16m + 6st + 62a 16m, sm + 2st + 66a 16m, sm + 4st + 62a 16m + 2sm+ 62a 16m, sm + 4st + 62a 16m, sm + 68a 20m + 14sm + 40a 68m + 34a
Beamish et al. 1977 Nygren et al. 1971c Zenzes 1975 Prokofieva 19 34 Svardson 1945 Cataudella et al. 1975b Raicu et al. 1977 Svardson 1945 Svardson 1945 Berberovic et al. 1970 Viktorosky 1978 Svardson 1945 Davisson et al. 19 72a Muramoto et al. 1974 Cataudella et al. 1973b Abe et al. 1974 Abe et al. I974 Viktorosky 1978 Muramoto et al. 1974 WahllY60 Booke 1975 Svardson 1945 Nygren et al. 1971a Prokofieva 19 34
Salmonid Hybrids
Coregonus lavaretus baeri x Salvelinus fontinalis Oncorhynchus keta x Oncorhynchus nerka Oncorhynchus nerka x Oncorhynchus keta Oncorhynchus kisutch x Salvelinus fontinalis Salvelinus fontinalis x Oncorhynchus kisutch Salmo salar x Salvelinus fontinalis Salmo salar x Salmo trutta Salmo salar x Salmo trutta Salmo trutta fario x Salvelinus fontinalis Salmo gairdneri x Salvelinus fontinalis
35m, sm + 4st + 27a 35m, sm + 4st + 27a 38m + 8sm + 14a
105 105 106
Sasaki et al. 1968 Sasaki et al. 1968 Uyeno 1972 Uyeno 1972
14m +56a 15m + 3sm + 4st + 60a
(84) 91 (100) 133-159
Prokofieva 1934 Svardson 1945 Nygren et al. 1975b Capanna et al. 1973b Cataudella et al. 1975b
Family Plecoglossidae
Plecoglossus altivelis
56 58 54
12m, st + 44a 10m + 48a 16m + 38a
68
Yamazaki 1971 Svardson 1945 Nygren et al. 1971d
Family Osmeridae
Osmerus Osmerus eperlanus eperlanus
(68)
70
293
Table 1 (Continued) Species n 2n Karyotype Number of arms Heterogametic sex Reference
Suborder Argentinoidei Family Argentinidae Argentina silus Family Bathylagidae Bathylagus milleri Bathylagus ochotensis Bathylagus stilbius Bathylagus wesethy Leuroglossus stilbius Suborder Galaxioidei Family Galaxiidae Brachygalaxias bullocki Galaxias maculatus Galaxias maculatus Galaxias platei Suborder Esocoidei Family Esocidae Esox americanus americanus Esox americanus americanus Esox americanus vermiculatus Esox lucius Esox lucius Esox lucius Esox lucius Esox lucius Esox masquinongy Esox masquinongy Esox niger Esox niger Esox reicherti Esox reicherti Family Umbridae Dallia pectoralis Novumbra hubbsi Umbra limi Umbra limi Urn bra pygmaea Suborder Family Bonapartia Gonostoma Gonostoma Stomiatoidei Gonostomatidae pedaliota bathyphilum elongatum
44
17m + 27a (d) lm + 16m + 43a (d) 2m + 16m + 42a (0) 19m + 14sm + 2sat + la (d)
61 77 78 71 82
d?
Ebeling et al. 1971a
60 54 64 36 64
Ebeling et al. 1971a, 1971b Chen 1969 Chen 1969 Chen et al. 1966 Ebeling et al. 1971a
18
d?
38 22 22 30
10m + 16sm + 12a 8m+12sm+2a 8m + 1Osm + 4st lm + 18sm + lla (d) 2m + 18sm + 10a (Q)
64 42 (2) (50) 6
Campos 1972 Campos 1972 Merrilees 1975 Campos 1972
50 50 50 46 48 50 50 50 50 50 50 50 50 50 78 48 22 22 22
50a 50a 50a 50a 50a 50a 50a 50a 50a 50a 50a 50a
(Z, (50)
(2:) (Z) 50 (50) 50 (50) 50
Davisson 1972 Beamish et al. 1971a Beamish et al. 1971a Lieder 1953 Prakken et al. 1955 Nygren et al. 1968a Beamish et al. 1971a Davisson 1972 Beamish et al 1971a Davisson 1972 Beamish et al. 1971a Davisson 1972 Beamish et al. 1971a Davisson 1972 Beamish et al. 1971a Beamish et al. 1971a Beamish et al. 1971a Kligerman et al. 1975 Beamish et al. 1971a
22m 22m, sm 22m
(44) (44) (44)
24 6 24
Post 1974 Post 1974 Post 1974
294
Suborder Saurida Saurida Synodus
Myctophoidei elongata undosquamis lucioceps
Family Synodontidae 48 48 48 48 24 24 24 24 24 24 48 24 24 24 48 49 48 41 48 24 24 24 24 24 48 48 48 48 48 48 48 48a 48a 48a 2m, sm + 46a 48a 2m, sm + 46a (48) 48a lm + 48a (6) 48a lm + 46a (d) 2m + 46a (0) 48a 48a 28m, sm + 20a 48a (48) (48) (76)
Q
Nishikawa et al. 1978 Nishikawa et al. 1978 Chen et al. 1974 Chen et al. 1974 Post 1972 Post 1972 Post 1972 Post 1972 Post 1972 Post 1972 Chen et al. 1974 Post 1972 Post 1972 Post 1972 Chen et al. 1974 Chen et al. 1974 Chen et al. 1974 Chen et al. 1974 Post 1912 Post 1972 Post 1972 Post 1972 Chen et al. Post 1972 Chen et al. Chen et al. Chen et al. Chen et al.
Family Myctophidae
Ceratoscopelus towsendi Ceratoscopelus warmingi Ctenoscopelus phengodes Diaphus brachycephalus Diaphus dumerili Diaphus fragilis Diaphus rafinesquei Diaphus theta Electrona rissoi Hygophum hygomi Lampadena chavesi Lampadena urophaos Lampanyctus ingens Lampanyctus regalis Lampanyctus ritteri
(48)
(48)
(50) (48) d? d
(48)
(50)
Lepidophanes guentheri Lepidophanes photothorax Lepidophanes supralateralis Lobianchia gemelari Myctophum nitidulum Notoscopelus resplendens Protomyctophum crockeri Stenobranchius leucopsarus Symbolophorus californiensis Triphoturus mexicanus
1974 1974 1974 1974 1974
(48)
(48) (50) (48)
Family Neoscopelidae
Scopetengys tristis
(50)
Chen et al. 1974
Order GONOR YNCHLFORMES Suborder Chanoidei
Family Chanidae
Chanos chanos
32 OSTARIOPHYSI
14m + 4sm + 14a
50
Arai et al. 1976~
SUPERORDER
Order CYPRINIFORMES Suborder Characoidei
Family Characidae
Anostomus anostomus Chnlceus maerolepidotus Chaleeus macrolepidotus Hemigrammus caudovittatus Lepolinus fasciatus
54 52 54 50 54
54m, 44m, 32m, 2m + 54m,
sm sm + 8st, a sm + 22st 32sm + 16a sm
(108)
(96)
636)
(84)
(108)
Ojima et al. 1976 Ojima et al. 1976 Muramoto et al. 1968 Fontana et al. 1970a Ojima et al. 1976
295
Table 1 (Continued) Species n 2n Karyotype Number of arms 2a 24st, a 4a 16st + 18a Heterogametic sex Reference
Methinis schreiinuelleri Myloplus arnoldi Mylossoma duriventris Serrasalmus hollandi Suborder Gymnotoidei Family Apteronotidae Apteronotus albifrons Suborder Cyprinoidei Family Cyprinidae Abbottina rivularis Abramis ballerus Abramis brama Abramis brama Abramis brama Acheilognathus cyanostigma Acheilognathuslanceolatus Acheilognathus Ianceolatus Acheilognathus lanceolatus Acheilognathus limbata Acheilognathus limbata Acheilognathus longipinnis Acheilognathus moriokae Acheilognathus rhombea Acheilognathus rhombea Acheilognathus tabira tabira Acheilognathus tabira tabira Acheilognathus tabira subsp. Alburnus alburnus Amblypharyngodon calbasumola Amblypharyngodon mola Aristichthys nobilis Aspius aspius Aulopyge hugeli Barbus barbus Barbus barbus plebejus Barbus fasciatus Barbus fasciatus Barbus meridionalis Barbus meridionalis petenyi Barbus (Barbodes) pentazona Barbus schuberti Barbus semifasciolatus Barbus (Capoetaj tetrazona Barbus tetrazona Barbus (Capoeta) titteya Barbus (Barbodes) viviparus Brachidanio (Danio) frankei Capoeta oligolepis Carassius auratus 296
62 58 54 64
60m, 34m, 50m, 30m,
sm + sm + sm + sm +
(122)
(104)
(94)
Ojima et al. 1976 Ojima et al. 1976 Ojima et al. 1976 Muramoto et al. 1968
24
14m+2sm+2st+6a
(40)
Howell 1972
25 26
22 25
22
50 52 52 50 52 44 50 48 48 44 48 44 44 44 44 44 44 44 50 50 52 48 52
18m 30m 8m + 20sm, st + 20a 10m + 18sm + 20st, a 10m + 18sm + 20st, a 14m + 14sm + 16st, a 14m + 14sm + 16st, a 12m + 20sm, st + 12a 14m + 14sm + 16st, a 12m + 20sm, st + 12a 14m + 14sm + 16st, a 14m + 14sm + 16st, a 16m + 1Osm + 16st + 8a 12m+20sm+8st+lOa 8m, sm + 44a 20m, sm + 20st + 8a 42m 48m, sm + .52st, a
IO
82
76 (76) (76) (72)
(72)
16 (72) 16 (72) (72) (76)
(82) (60) (68)

94
100 100 100

52 52
148
26m + 18sm + 18st + 38a (144) 30m, sm + 4st + 16a + (83 2min. 22m + 20sm + 12st + 46a (142)
Nogusa 1955a Nygren et al. 197Sa Post 1965 Wolf et al. 1969 Nygren et al. 1975a Nogusa 1960 Makino 1939 Ojima et al. 1972 Ojima et al. 1973 Nogusa 1960 Ojima et al. 1973 Ojima et al. 1973 Ojima et al. 1973 Ojima et al. 1972 Ojima et al. 1973 Ojima et al. 1972 Ojima et al. 1973 Ojima et al. 1973 Cataudella et al. 1977b Manna et al. 1977b Nayyar 1964 Vasilev et al. 1978 Nygren et al. 197 5a Berberovic et al. 1973 Wolf et al. 1969 Cataudella et al. 1977b Nogusa 1960 Ohno et al. 1967 Cataudella et al 1977b Sofradzija et al. 1973 Post 1965 Post 1965 Nogusa 1960 Post 1965 Ohno et al. 1967 Post 1965 Post 1965 Post 1965 Post 1965 Makino 1941b
100 100
24 24 26 24 24 24 24 24 41 52 50
34m, sm + 6st + 1Oa
(84)
94
Table 1 (Continued) Species n 2n Karyotype Number of arms 148

(150)
Heterogametic sex
Reference
Carassius aura&s Carassius aura&s Carassius auratus Carassius aura&s auratus Carassius auratus buergeri Carassius auratus buergeri Carassius auratus cuvieri Carassius auratus gibeho Carassius auratus gibelio Carassius auratus grandoculis Carassius auratus langsdorfii Carassius auratus langsdorfii Carassius auratus langsdorfii Carassius carassius Carassius carassius Catla ca tla Chela bacaila Chela bacaila Chondrostoma soetta Chondrostoma toxostoma Chrosomus eos Chrosomus erythrogaster Cirrhina reba Crossocheilus iatius punjabensis Ctenopharyngodon idellus Ctenopharyngodon idellus Ctenopharyngodon idellus Cyprinus carpio Cyprinus carpio Cyprinus carpio Cyprinus carpio Danio albolineatus Danio albolineatus Danio devario Danio malabaricus Danio rerio Danio rerio Danio (Brachidanio) rerio Esomus danrica Exoglossum maxilingua Garra rufa Gila bicolor Cila orcu tti Gnathopogon coerulescens Gnathopogon elongatus tinathopogon elongatus elongatus Gobio albipinnatus vladykovi Gobio bessleri banaticus Gobio gobio Gobio gobio Gobio gobio
100 104 100 100 100 156 100 100 156 100 100 156
47 25 206 94
12m + 36sm + 52st, a 46m, sm + 16st + 42a 20m + 40sm + 40a 12m + 36sm + 52st, a 20m + 40sm + 40a 34m + 62sm + 60a 20m + 40sm + 40a 20m + 4Osm + 40a 34m + 62sm + 60a 20m + 40sm + 40a 20m + 40sm + 40a 34m + 62sm + 60a 44m + 82sm + 80a
(160)
(148)
(160)
(252) (160) (160) (252) (160) (160) (252) (332)
100
50 52 50 50 50 50 50 48 48 48 48 48
20m + 40sm + 40a (160) 8m + 16sm + 14st + 12a (74) 52a (52) lOm+12sm+1Ost+18a (72) 16m + 14sm + 14st + 6a (80) 16m + 14sm + 14st + 6a (80) 10m + 36sm + 4a 18m+20sm+6st+4a 12m + 36a 16m + 20sm, st + L2a 14m + 20sm + 8st + 6a 12m + 36sm, st + 52a 46m, sm + 18st + 36a + 4min. 24m + 24sm, st + 52a 10m + 39sm, st + la 10m + 40a 10m + 40a 10m + 39sm + la 10m + 12sm, st + 14a 12m+16sm+lOst+12a
(96)
(86)
(60)
(2) (150)
100 104
98 100 24 50 50 50 50 50 50 50 48 50 50 50 50 50 50 50 24 50 50
Ojima et al. 1966,1967 Ohno et al. 1967 Kobayasi et al 1970 Ojima et al. 1966 Kobayasi et al. 1973b Kobayasi et al. 1973b Kobayasi et al. 1973b Kobayasi et al. 1973a Kobayasi et al. 1973a Kobayasi et al. 1973b Kobayasi 1977 Kobayasi 1977 Kobayasi 1977 Makino 1941b Kobayasi et al 1973b Manna et al. 1977b Nayyar 1964 Manna et al. 1977b Cataudella et al. 197713 Cataudella et al. 1977b Legendre et al. 1959 Greenfield et al. 1973 Manna et al. 1977b Nayyar 1964 Nogusa 1960 Ojima et al. 1972 Manna et al. 1977b Ojima et al. 1967 Ohno et al. 1967 Szollar et al. 1971 Raicu et al. 1972b Post 1965 Fontana et al. 1970a Fontana et al. 1970a Fontana et al. 1970a Endo et al. 1968 Fontana et al. 1970a Rishi 1976b Manna et al. 1977b Legendre et al. 1959 Post 1965 Avise et al. 1977b Greenfield et al. 1972 Nogusa 1960 Nogusa 1960 Ojima et al. 1972 Raicu et al. 1973 Raicu et al. 1973 Post 1965 Raicu et al. 1973 Sofradzija et al. 1975a 297
WO
(60) (99)
25 22-25
(78)
44m, sm + 6a 14m t 24st t 12a 12m + 32sm, st + 6a 28m + 20sm + 2a 24m + 20sm + 4st + 2a 22m + 24sm + 2st + 2a 38m + 12a
94 (64)
94 98
98
98
(88)
Table 1 (Continued) Species n 2n Karyotype Number of arms 100 94 90 92 (70) 92 Heterogametic sex Reference
Gobio uranoscopus Hemibarbus barbus Hemibarbus labeo Hemibarbus longhostris Hemibarbus longirostris Hemigrammocypris rasborella Hesperoleucus symmetricus Hesperoleucus symmetricus Hybopsis plumbea Hypophthalmichthys molitrix Hypophthalmichthys molitrix Hypophthalmichthys molitrlx Ischikauia steenackeri Ischikauia steenackeri Labeo bata Labeo chrysophekadion Labeo gonius Lavinia exilicauda Lepidomeda albivallis Lepidomeda mollispinis Levidomeda vittata Leuciscus albidus alborella Leuciscus aula Leuciscus cephalus Leuciscus cephalus Leuciscus cephalus Leuciscus cephalus albus Leuciscus cephalus cabeda Leuciscus cephalus cephalus Leuciscus leuciscus Leuciscus idus Leuciscus souffio agassizi Leuciscus souffia muticellus Leuciscus souffia muticellus Leuciscus svallize Leuciscus svallize svallize Leuciscus turskyi Leuciscus turskyi tenellus Meda fulgida Moroco jouyi Moroco lagowskii Moroco oxycephalus Moroco percnurus Moroco steindachneri Moroco steindachneri Mylopharodon conocephalus Notemigonus crysoleucas Notemigonus crysoleucas Notropis cornutus No tropis lu trensis Notropus texanus Notropis venustus
298
52 50 50 25 50 50 24 48 50 50 50 48 48 48 48 48 25 50 50 54 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 50 27 27 54 50 50 50 50 50 50 50 50
24m+12sm+12st+4a 14m + 2&m, st + 8a 18m + 22sm, st + 10a 14m + 28sm, st + 4a 20m + 20st + 10a 42m, sm + 8a
20m + 12sm + 6st + 10a 20m, sm + 20st + 8a 10m + 30sm, st + 8a 6m+18m+16st+10a 14m, sm + 18st + 18a 2st + 52a 42m, sm + 8a 12m + 32sm, st + 6a 12m + 30sm, st + 8a 14m + 32sm, st + 4a 12m + 25m, st + 13a 15m + 23sm, st + 12a 1 lm + 29sm, st + 10a 36m, sm + 14a 16m + 12sm + 12st + 10a 36m, sm + 14a 40m, sm f 10a 38m, sm + 12a 42m, sm + 8a 15m + 23sm, st + 12a 20m + 12sm + 8st + 10a 32m, sm + 18a 34m, sm + 18a 18m + 24sm, st + 8a 14m + 22sm + 8st + 6a 12m + 2&m, st + 10a 12m +2&m, st + 10a 14m + 22sm + 8st + 6a 44m, sm + 6a 44m, sm + 6a 6m + 40sm + 4a 2m + 42sm + 6a 2m + 38sm + 10a 70 86 (78) 86 90 88 92 88 99
(82)
(54) 92
(82)
74 82 68 84 92 90 90 92 94 90 94 (96) (E) (90)
Raicu et al. 1973 Ojima et al. 1972 Kang et al. 1973b Nogusa 1960 Kang et al. 1973b Nogusa 1960 Greenfield et al. 1972 Gold et al. 1977a Legendre et al. 1959 Khuda-Bukhsh et al. 1974 Manna et al. 1977b Vasilev et al. 1978 Nogusa 1960 Ojima et al. 1972 Manna et al. 1977b Muramoto et al. 1968 Srivastava et al. 1964 Gold et al. 1977a Uyeno et al. 1973 Uyeno et al. 1973 Uyeno et al. 1973 Fontana et al. 1970a Fontana et al. 1970a Wolf et al. 1969 Fontana et al. 1970a Berberovic et al. 1972 Sofradzija 1977 Cataudella et al. 1977b Sofradzija 1977 Sofradzija 1977 Sofradzija 1977 Sofradzija 1977 Fontana et al. 1970a Cataudella et al. 1977b Berberovic et al. 1972 Sofradzija 1977 Berberovic et al. 1972 Sofradzija 1977 Uyeno et al. 1973 Ojima et al. 1976 Kang et al. 19731, Kang et al. 1973b Nogusa 1960 Nogusa 1960 Ojima 1976 Gold et al. 1977a Lieppman et al. 1969 Gold et al. 1977a Greenfield et al. 1973 Lieppman et al. 1969 Campos et al. 1973 Campos et al. 1973
Table 1 (Continued) Species n 2n Kawotype Number of arms Heterogametic sex Reference
Opsariichthys uncirostris Opsariichthys uncirostris Opsopoeodus (Notropis) emiliae Orthodon microlepidotus Oxygaster bacaila Pachychilon pictum Paraphoxinus adspersus Paraphoxinus alepidotus Paraphoxinus alepidotus Paraphoxinus croaticus Paraphoxinus metohiensis Paraphoxinus pstrossi Pelteobagrus nudiceps Phoxinus percnurus Phoxinus phoxinus Phoxinus phoxinus Pimephales promelas Plagopterus argentissimus Pogonichthys macrolepidotus Pseudogobio esocinus Pseudogobio esocinus Pseudoperilampus typus Pseudorasbora parva Pseudorasbora parva Ptychocheilus grandis Pungtungia herzi Pungtungia herzi Pungtungia hilgendorfi Puntius conchonius Puntius conchonius Puntius conchonius Puntius filamentosus Puntius sophore Puntius stigma Puntius stigma Puntius stigma Puntius ticto Rasbora buchanani Rasbora heteromorpha Rasbora trilineata Rhinichthys atratulus Rhinichthys cataracta Rhodeus atremius Rhodeus ocellatus Rhodeus ocellatus ocellatus Rhodeus ocellatus ocellatus Rhodeus ocellatus smithii Rhodeus sericeus amarus Rhodeus suigensis Richardsonius egregius Rohtee cotio Rut&s rubilio
66 78 48 50 48 50 50 50 50 50 50 50 56 27 50 50 50 50 50 52 50 44 50 50 50 50 50 50 50 50 24 50 48 50 48 50 50 24 24 50 50 46 44 48 48 48 48 46 50 48 50
8sm, st + 70a 2m + 30sm + 16a 44m, sm + 6a 48a 38m, 26m, 28m, 36m, 32m, 30m, sm + sm + sm + sm + sm + sm + 12a 24a 22a 14a 18a 20a
86
(80)
94
(48) 06)
18 86 82 80
32m, sm + 18a 44m + 12sm + 16st + 8a 18m + 26sm, st + 6a 14m, sm + 6a 16m + 28sm, st + 6a 14m + 14sm + 16a 14m + 36sm, st 42m, sm + 8a 20m + 30sm, st 50a
82 86
94 94 (72) 100 92
26
25
100
(50)
24
50a 48a 50a 2m + 2sm + 44a 50a 30m + 18sm + 2st 16m + 28sm + 2st + 4a 16m + 28sm + 2st + 4a 4sm + 42st, a 10m + 18sm + 20st, a 8m + 20sm, st + 20a 10m + 18sm + 20st, a 34m, sm + 14a 4sm + 42st, a 36m, sm + 14a 12m + 14sm + 8st + 4a 18m+lOsm+16st+6a
(50) (48) (50)
(52)
(50)
(98)
(94) (94)
16 82 86
(84)
(78)
Nogusa 1960 Ojima et al. 1972 Campos et al. 1973 Gold et al. 1977a Srivastava et al. 1964 Berberovic et al. 1972 Sofradzija et al. 1972 Berberovic et al. 1969b Sofradzija et al. 1972 Sofradzija et al. 1972 Sofradzija et al. 1972 Sofradzija et al. 1972 Nogusa 1960 Nogusa 1960 Berberovic et al. 1972 Cataudella et al. 1977b Gravell et al. 1969 Uyeno et al. 1973 Gold et al. 1977a Nogusa 1960 Ojima et al. 1972 Ojima et al. 1973 Makino 1939 Ojima et al. 1972 Gold et al. 1977a Nogusa 1960 Ojima et al. 1972 Nogusa 1960 Nayyar 1964 Post 1965 Vasilev et al. 1978 Post 1965 Nayyar 1964 Srivastava et al. 1964 Nayyar 1964 Rishi 1973 Srivastava et al. 1964 Manna et al. 1977b Post 1965 Post 1965 Howell et al. 1976 Howell et al. 1976 Ojima et al. 1973 Nogusa 1960 Ojima et al. 1973 Ojima et al. 1972 Ojima et al. 1973 Sofradzija et al. 1975 Ojima et al. 1973 Gold et al. 1917a Manna et al. 1977b Cataudella et al. 1977b 299
Rutilus rutilus Rutilus rutilus Rutilus rutilus Rutilus rutilus Sarcocheilichthys variegatus Sarcocheilichthys variegatus Scardinius erythrophthalmus Scardinius erythrophthalmus Scardinius erythrophthalmus Semotilus atromaculatus Semotilus corporalis Semotilus margarita Tanakia tanago Tinca tinca Tinca tinca Tinca tinca Tinca tinca Tribolodon ezoe Tribolodon ezoe Tribolodon hakonensis Tribolodon hakonensis Tribolodon hakuensis Typhlogarra widdowsoni Vimba vimba Zacco platypus Zacco temminckii Family Catostomidae Carpiodes carpio Catostomus catostomus Catostomus clarki Catostomus commersoni Catostomus commersoni Catostomus discobolus Catostomus latipinnis Erymizon sucetta Hypentelium nigricans Moxostoma duquesnei Moxostoma erythrurum Moxostoma macrolepidotum Family Cobitidae Acanthophthalmus khulli Barbatula oreas Barbatula toni Botia macracantha Cobitis biwae Cobitis biwae Cobitis biwae Cobitis biwae Cobitis biwae Cobitis biwae 300
25
25 24
52 50 50 50 50 50 48 48 50 52 52 50 48 48 48 48 48 50 50 50 50 50 52 48 48 100 98 96-98 98 96-98 100 100 100 100 96-98 96-98 96-98 50 48 50 98 54 96 48 96 48 48
68 76 12m + 30sm, st + 8a 13m + 35a 42m 20m + 12sm + 12st + 6a
90 (82)
10m + 18sm + 20st, a 6m + 28sm, st + 14a 36m 8m + 12sm + lost + 18a 10m + 20sm + 12st + 8a 14m + 22sm + 8st + 6a 10m + 20sm + 12st + 8a 14m + 22sm + 8st + 6a 24m, sm + 28st, a 18m + 22sm. st + 8a 18m + 22sm, st + 8a
(76)
84 (68) 92 92 92 92 (76) 88 88
Lieder 1954a Wolf et al. 1969 Berberovic et al. 1972 Nygren et al. 1975a Nogusa 1960 Ojima et al. 1972 Fontana et al. 1970a Nygren et al. 1975a Cataudella et al. 197713 Legendre et al. 1959 Legendre et al. 1959 Legendre et al. 1959 Ojima et al. 1973 Wolf et al. 1969 Fontana et al. 1970a Nygren et al. 1975 Cataudella et al. 1977b Itoh et al. 1972 Ojima et al. 1976 Itoh et al. 1972 Ojima et al. 1976 Makino 1939 Post 1965 Shinderite et al. 1976 Ojima et al. 1972 Ojima et al. 1972 Uyeno et al. 1972b Beamish et al. 1971b Uyeno et al. 1972b Beamish et al. 1971b Uyeno et al. 1972b Uyeno et al. 1972b Uyeno et al. 1972b Uyeno et al. 1972b Uyeno et al. 19721, Uyeno et al. 1972b Uyeno et al. 1972b Uyeno et al. 1972b Muramoto et al. 1968 Makino 194 1a Hitotsumachi et al. 1969 Muramoto et al. 1968 Post 1965 Hitotsumachi et al. 1969 Kobayasi 1976 Kobayasi 1976 Takahasi et al. 1976 Ueno et al. 1976
8m + 90sm. a 12m + 86sm, a
50
14m. sm + 12st + 24a 16m, sm + 34st, a 28m, sm + 70a 58m, sm + 38st, a 18m + 30sm 36m + 52sm + 8st 2m, sm + 46a 20m + 22sm, st + 6a
24 27
(64)
(66)
(126)
(154) (96)
(184)
(50) 90
Table 1 (Continued) Species n 2n Karyotype Number of arms 182 (68) Hetero gametic sex Reference
Cobitis biwae Cobitis delicata Co bitis delieata Cobitis taenia Cobitis taenia taenia Cobitis taenia taenia Cobitis taen& taenia Cobitis taenia taenia Cobitis taenia striata Cobitis taenia striata Cobitis taenia striata Lefua echigonia Lefua nikkonis Leptobotia curta Misgurnus anguillicaudatus Misgurnus anguillicaudatus Misgurnus fossilis Misgurnus fossilis Order Ictalurus Ictalurus SIL URIFORMES
96
25 26 24
50 50 50 86 50 86 94 50 50 98 50 50 50 52 50 100
32m + 54sm, st + 10a 18m, sm + 32st, a 18m, sm + 32a 12m + 6sm + 16st + 16a 64m, sm + 22a 12m+4sm+34a 32m + 32sm, st + 22a 26m + 32sm, st + 36a 16m, sm + 34a 12m+4sm+34a 20m + 22sm + 56a 12m, sm + 38st, a 8m + 1Osm + 16st + 16a 14m, sm + 36a 36m, sm + 64a
(68) (68)
(150) 66 150 152 (66) 66 140
(62) (68) (64)

136
Ueno et al. 1976 Hitotsumachi et al. 1969 Takahasi et al. 1976 Cataudella et al. 1977b Takahasi et al. 1976 Ueno et al. 1976 Ueno et al. 1976 Ueno et al. 1976 Takahasi et al. 1976 Ueno et al. 1976 Ueno et al. 1976 Nogusa 1960 Hitotsumachi et al. 1969 Sawada 1976 Makino 1941a Hitotsumachi et al. 1969 Post 1965 Raicu et al. 1972a
Family Ictaluridae
nebulosus punctatus
60 56 56 38 58 50 54 54 50 56 48 54 54 41 d 42 9 40 58 58 58 58 86 66 58
20m, sm + 40a 16m, sm + 22st + 18a 28m + 24sm + 4st, a 28m + 1Osm 30m + 12sm + 2st + 140 10m + 44a 10m + 38sm + 6a (d) 9m + 38sm + 7a (Q) 24m 20m 28m 16m + + + + 12sm + 20st, a 12sm + 16st, a 20sm + 6st 38a
80 (72)
Berberovic et al. 1975 Muramoto et al. 1968 Ojima 1976 Ojima 1976 Manna et al. 1978 Srivastava et al. 1969 Nayyar 1966 Q Rishi 1973 Srivastava et al. 1969 Ueno 1974 Ueno 1974 Das et al. 1977 Nayyar 1966
Family Bagridae
Coreobagrus lshikaua Liobagrus reini Mystus gulio Mysius seenghala Mystus tengara Mystus tengara
(108)
66 102 86
(64)
102 101
Mystus vittatus Pseudobagrus aurantiacus Pseudobagrus aurantiacus Rita chrysea Rita rita
(92) (80)
102 (70)
Family Siluridae
Callichromis bimaculatus
Ompok bimaculatus Parasilurus asotus Parasilurus asotus Parasilurus biwaensis Parasilurus lithophilus Wallago attu
17m + 12sm,mt + 12a 18m + 12sm, st + 12a 4m + 36a 38m, sm + 8st + 12a 44m, sm + 14st, a 44m, sm + 14st, a 44m, sm + 14st, a 86a 66a 10m + 20sm + 12st + 4t + 12T
Rishi 1976a Nayyar 1966 Muramoto 1969 Arai et al. 1974b Arai et al. 1974b Arai et al. 1974b Nayyar 1966 Nayyar 1966 Manna et al. 1978
(44) 104 102 102 102
(86) (66)
Family Schilbeidae
Clupisoma parua Eutropiichthys vacha
100
301
Family Clariidae
Clarias Clarias Clarias Clarias batrachus batrachus batrachus fuscus
52 52 50 56 58 56 56 56 48 46 62 54 58 24 68 68 14 74
52a 16m + 8sm + 14st + 12a
(52)
58 (74) 88
Nayyar 1966 Srivastava et al. 1968 Rishi 1976b Arai et al. 1974a Nayyar 1966 Vasudevan et al. 1973 Prasad et al. 1974 Rishi 1976b Arai et al. 1974b Toledo et al. 1976b Toledo et al. 1976b Muramoto et al. 1968 Michele et al. 1977b Post 1965 d Michele et al. 1977b Michele et al. 1977b Michele et al. 1977b Michele et al. 1977b
Family Heteropneustidae
Heteropneustes Heteropneustes Heteropneustes Heteropneustes fossilis fossilis fossilis fossilis
58a 18m + 18sm + 20a 14m + 26sm + 16a 18m + 18sm + 20a 12m, sm + 36st, a 28m + 1Osm + 8a 36m+18sm+8a 24m, sm + 12st + 18a 18m + 2sm + 38a 10m + 27sm + 31a d 10m + 28sm + 30a Q 10m + 14sm + 44a 10m + 20sm + 44a 8m + 4sm + 62a
(58) (92)
(96) (92) 60 84 116
Family Plotosidae Plo~osus anguillaris Family Pimelodidae

Pimelodella sp. Rhamdia hilarii
Family Loricariidae
Hypostomus plecosromus Loricaria macrodon Loricaria parva Plecostomus Plecostomus Plecosromus Plecosiomus ancistroides macrops paulinus s09gatieeps
(78) 78 105 106 92 104 86
SUPERORDER PARACANTHOPTERYGII
Order GADIFORMES Suborder Gadoidei
Family Gadidae
Gadus morrhua Gaidropsarus medirerraneus Micromesistius poutassou Pollachius pollachius Pollachius virens Raniceps raninus Trisopterus minutus
46 28 44 38 40 48 48
24m, sm + 4a 10m + 28a 10m + 30a 30m + 18a
52 48 50 78
Nygren Vasilev Nygren Nygren Nygren Nygren Nygren
et al. 1974 1978 et al. 1974 et al. 1974 et al. 1974 et al. 1974 et al. 1974
SUPERORDER ATHERINOMORPHA
Order A THERINIFORMES Suborder Exocoetoidei
Family Belonidae
Xenentodon cancila 50
Srivastava et al. 1964
302
Table I (Continued) Species n 2n Karyotype Number of arms Heterogametic sex Reference
Suborder Oryzias Oryzias
Cyprinodontoidei latipes melastigma
Family Oryziatidae 48 48 32 48 20m, sm + 28a 68 50 64 Ojima et al. 1969a Scheel 1972 Scheel 1972b Karbe 196 1 Karbe 1961 Karbe 1961 Karbe 1961 Karbe 196 1 Scheell968a Scheel 1968a Scheel 1972b Scheel 1968a Scheel1968a, Scheel 1968a, Scheel 1972b Scheell972b Scheel 1972b Scheel 1972b Scheel 1968a, Scheel 1972b Scheel1972b Scheel 1972b Scheel 1972b Scheel1972b Scheel 1968a, Scheel 1968a, Scheel 1968a Scheel 1968a, Scheel 1972b Scheel1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1968a, Scheel1968a, Scheel1968a, Scheel 1972b Scheel 1972b Scheel 1972b Scheel1968a, Scheel 1968a, Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b
Family Cyprinodontidae
Adinia multifhsciatus Aphanius cypris Aphanius dispar Aphanius fascia&s Aphanius iberus Aphanius sophiae Aphyoplatys duboisi Aphyosemion ahh Aphyosemion arnoldi Aphyosemion australe Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bivittatum Aphyosemion bualanum Aphyosemion bualanum Aphyosemion calliurum Aphyosemion cameronense Aphyosemion cameronense Aphyosemion cameronense Aphyosemion cameronense Aphyosemion cameronense Aphyosemion cameronense Aphyosemion cameronense Aphyosemion cameronense Aphyosemion cameronense Aphyosemion celiae Aphyosemion christyi Aphyosemion cinnamomeum Aphyosemion cognatum Aphyosemion cognatum Aphyosemion cognatum Aphyosemion exiguum Aphyosemion filamentosum Aphyosemion filamentosum Aphyosemion filamentosum Aphyosemion filamentosum Aphyosemion filamentosum
24 24 24 24 24 24 18 15 20 19
48 38 30 40 38 38 38 36 32 34 34 34 34 30 26 40 38 32 34 34 32 32 30 28 30 24 26 20 18 40 26 28 30 36 36 30 32 30 32 72 54 50 40 38 54 54 54 52 50 38 44 44 68 66 44 46 42 46 46 44 44 44 44 40 36 60 36 36 36 72 48 42 42 40 40
1972b 1972b
17
1972b
20 19 16 17
1972b 1972b 1972b
10 9 20
1972b 1972b 1972b
18 18
1972b 1972b
303
Table 1 (Continued) Species n 2n Karyotype Number of arms 44 54 56 56 56 54 42 48 32 50 50 44 40 36 60 60 60 60 88 72 56 46 40 70 36 48 40 86 48 48 48 94 50 48 70 52 92 86 52 50 76 76 50 Hetew gametic sex Reference
Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aphyosemion Aplocheilichthys Aplocheilichthys Aplocheilichthys Aplocheilichthys Aplocheilichthys Aplocheilichthys Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus Aplocheilus
franzwerneri gardneri gardneri gardneri gardneri gardneri geryi guineense gulare labarrei labarrei liberiense louessense melanopteron mirabile mirabile mirabile mirabile nidanum obscurum occidentale ogoense petersii roloffi santaisabellae scheeli schoutedeni seymouri sjoestedti toddi walkeri jlavipinnis katangae macrophthalmus normani spilauchen spilauchen barmoiensis bifasciatus blocki chaperi dageti dayi duboisi eskanus fasciolatus fasciolatus grahami grahami grahami lineatus lineatus
11 20 18
16 14 10
20 17
20
20 18 24 24
22 40 36 38 40 36 40 38 32 28 26 42 20 30 32 34 36 38 40 34 46 40 40 42 40 40 22 28 40 46 36 48 48 48
24 17 20 24 25 25 24 21 19 23 25
34 40 48 50 50 48 48 42 40 38 48 46 48 50 48
60
Scheel 1968a, Scheel 1968a, Scheel1968a, Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1968a, Scheel1968a, Scheel 1972b Scheel 1972b Scheel 1968a, Scheel 1972b Scheel 1972b Scheel 1972b Scheel 19721, Scheel 1972b Scheel 1968a, Scheel 1968a Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1968a, Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1968a, Scheel 1972b Scheel1968a, Post 1965 Post 1965 Scheel 1972b Scheel1972b Scheel 1972b Post 1965 Scheel 1972b Scheel 1968a, Scheel1968a, Scheel 1968a, Scheel 1968a, Scheel 1968a, Scheel 19721, Scheel 1968a, Scheel 1972b Scheel 1968a, Scheel 1972b Scheel 1968a, Scheel 1972b Scheel1968a, Scheel 1972b
1972b 197213 1972b
1972b 1972b 1972b c 1972b
1972b . 1972b 1972b
1972b 1972b 1972b 1972b 1972b 1972b 1972b 1972b 1972b
304
Table 1 (Continued) Species n 2n Karyotype Number of arms 56 50 88 54 80 52 72 92 52 14m + 20a (4sat) 2m + 44a (2sat) 48a (2sat) 4sm + 44a (2sat) lm + 3sm + 42a + 2sat 4sm + 42a + 2sat 4sm + 38a + 6t 2sm + 46a (2sat) 48a (2sat) 46a + 2sat 48a 48a (2sat) 2m + 44a (2sat) 16m + 4sm (2sat) + 12a 2sm (2sat) + 46a 46a + 2sat 48a 10m + 2sm + 28a (2sat) 10m + 12st + 18a 6m+12st+26a 2m + 44a (2sat) 2m + 2sm + 44a (2sat) 2m + 12st * 34a lsm + 47a d 2sm + 46a o 48a (2sat) 48a (2sat) 2sm + 46a (2sat) 4m + 2sm + 38a (2sat) 48a (2sat) 48 48 48 52 52 52 Hetero gametic sex Reference
Aplocheilus panchax Aplocheilus panchax Aplocheilus sangmelinensis Aplocheilus sexfasciatus Aplocheilus spilargyreius Aplocheilusquemeri Austrofundulus dolichopterus Austrofundulus transilis Cynolebias bellotti Cynolebias bellotti Cynolebias nigripinnis Cynolebias nigripinnis Cynolebias whitei Cynolebias whitei Cynopoecilus ladigesi Cynopoecilus melanotaenia Cynopoecilus melanotaenia Cyprinodon macularis Fundulus chrysotus Fundulus cingolatus Fundulus confluentus Fundulus diaphanus Fundulus diaphanus Fundulus diaphanus Fund&s diaphanus Fundulus grandis Fundulus heteroclitus Fundulus heteroclitus Fundulus heteroclitus Fundulus heteroclitus Fundulus kansae Fundulus lineolatus Fundulus luciae Fundulus majalis Fundulus majalis Fundulus majalis Fundulus notatus Fundulus notatus Fundulus notatus Fundulus notatus Fundulus notti Fundulus olivaceus Fundulus olivaceus Fundulus olivaceus Fundulus olivaceus Fundulus Fundulus Fundulus Fundulus FunduIus Fundulus parvipinnis parvipinnis pulvereus rathbuni sciadicus seminolis
18 24 24 17 22 24 24
38 36 48 48 34 48 44 44 48 48
24 46 24 24 44 24 34 46 48 48 48 48 48 48 24 48 48 48 48 46 32 48 48 48 40 48 40 44 46 48 48 48 48 48 48 48 48 44 48 48 (ii) 48 48 52 50 50 (48) 52 52 50 50 48 52 52 54 50 49 50 48 48 50 50 48
(52)
50
Scheel1972b Scheell972b Scheel1968a, 1972b Scheel 1968a, 1972b Scheel1968a, 1972b Scheel 1972b Scheel 1968a, 1972b Scheell972b Scheel 197223 Post 1965 Post 1965 Scheel 1972b Post 1965 Scheel1972b Post 1965 Post 1965 Scheel 1972b Post 1965 Chen 1971 Chen 1971 Chen 1971 Chen 1971 Chen et al. 1970b Chen et al. 1970b Arcement et al. 1976 Chen 1971 Post 1965 Chen 197 1 Chen et al. 1970b Fisher et al. 1972 Chen 1971 Chen 1971 Chen 1971 Chen 1971 Chen et al. 1970b Fisher et al. 1972 Chen 1971 Setzer 1970 Black et al. 1978 Black et al. 1978 Chen 1971 Setzer 1970 Chen 1971 Scheel1972b Black et al. 1978 d Chen et al. 1970b Chen Chen Chen Chen Chen 1971 1971 1971 1971 1971 305
Table 1 (Continued) Species n 2n Karyotype Number of arms 50 50 60 58 Heterw gametic sex Reference
Fund&us similis Fundulus waccamensis Jordanella floridae Jordanella floridae Kosswigichthys asquamatus Nothobranchius guentheri Nothobranchius kirki Nothobranchius mayeri Nothobranchius orthonotus Nothobranchius palmquisti Nothobranchius palmquisti Nothobranchius rachovii Nothobranchius rachovii Nothobranchius thirty Nothobranchius thirry Orestias agassize Orestias luteus Pachypanchax playfairi Pachypanchax playfairi Procatopus aberrans Procatopus nigromarginatus Procatopus nototaenia Procatopus roseipinnis Procatopus similis Pterolebias longipinnis Pterolebias longipinnis Pterolebias peruensis Rivulus cylindraceus Rivulus dorni Rivulus harti Rivulus marmoratus Rivulus milesi Rivulus ocellatus + Rivulus ornatus Rivulus strigatus Rivulus urophthalmus Valencia hispanica
48 46 24 48 24 19 36 19 18 19 18 8
2smf2sat) + 46a 4sm (2sat) + 42a
16 18 42 44 48 48 48
30 48 48
24 24 24 24 24 24 24 10 34
96 96
48 20 54 48 48 44 48 46 48 40 46 46 48
96 20 90 56 70 56 52 92 54 66 88 86 92
Chen 1971 Chen 1971 Post 1965 Scheel 1972b Karbe 1961 Post 1965 Scheel 1972b Post 1965 Scheel1968a Post 1965 Scheel1968a Post 1965 Scheel 1972b Scheel 1972b Scheel1972b Lueken 1962 Lueken 1962 Post 1965 Scheel 1972b Scheel 1972b Scheel 1968a Post 1965 Scheel1968a Scheel 1968a, 1972b Post 1965 Scheel 1972b Scheel1968a, 1972b Scheei 1972b Scheel1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 19721, Scheel 1972b Scheel 1972b Scheel 1972b Scheel 19721,
Family Goodeidae
Ameca splendens Characodon later&s Hubbsina turneri Xenotoca eiseni Xenotoca melanosoma Xenotoca variata
26 24 48 48 48 48
22m + 2st + 2a 24m 48a 6st + 42a 8st + 40a 4st + 44a 2sm + 46a lm+2sm+45a lm + 47a 9 48a d Im+47aQ d P
(48) (48) (48) (48) (48) (48)
Miller et al. 1971 Uyeno et al. 1972a Uyeno et al. 1972a Fitzsimons 1972 Fitzsimons 1972 Fitzsimons 1972
Family Poecilidae
Gambusia Gambusia Gambusia affinis affnis affinis
48 48 48
(50) (51) (4%
P O?
Q
Chen et al. 1968 Yosida et al. 1971 Itahashi et al. 1975
(48) (49)
306
Table 1 (Continued) Species n 2n Karyotype Number of arms 48 (48) (49) (48) (49) 42 (48) (49) (48) (48) (46) Q Q Heterogametic sex Reference
Gambusia Gambusia Gambusia Gambusia Gambusia Gambusia
affinis holbrooki holbrooki gaigei hurtadoi marshi nobilis
48 24 48 48 42 48 48 48 46 24 24 24 46 23 46 48 46 46 69 46 46 46 46 46 48 72 48 72 72 48 48 48 48 48 48
48a d, Q 48a d lm +47aQ 48a d lm+47aQ 42a 48a d lm +47aQ 48a 48a 46a
Cataudella et al. 1977a Post 1965 Campos et al. 1971 Campos et al. 1971 Campos et al. 1971 Campos et al. 1971 Campos et al. 1971 Campos et al. 1971 Yosida et al. 1971 Post 1965 Post 1965 Post 1965 Q Rishi et al. 1976 Post 1965 Freye et al. 1967 Prehn et al. 1969 Rasch et al. 1970 Rasch et al. 1970 Post 1965 Prehn et al. 1969 Prehn et al. 1969 Prehn et al. 1969 Scheel 1972b Post 1965 Prehn et al. 1969 Prehn et al. 1969 Schultz 1967 Schultz 1967 Cimino 1973 Cimino 1973 Cimino 1973 Ohno et al. 1966 Freye et al. 1967 Scheel1972b Itahashi et al. 1975 Scheel 1972b Itahashi et al. 1975 Post 1965
Gambusia regani Gambusia vittata Lebistes reticulatus Limia melanogaster Mollienisia latipinna Mollienisia sphenops Mollienisia sphenops
46a d Im+45aQ 46a 1st + la + 44minute 1st + 2a + 66minute
(46) (47) 46 (46)
Mollienisia velifera Platypoecilus maculatus Poecilia formosa Poecilia formosa Poecilia formosa Poecilia latipinna Poecilia latipinna Poecilia latipunctata Poecilia mexicana Poecilia reticulatus Poecilia sphenops Poecilia sphenops Poecilia velifera Poeciliopsis lucida Poeciliopsis lucida Poeciliopsis monacha-lucida Poeciliopsis monacha-2lucida Poeciliopsis 2monacha-lucida Xiphophorus helleri Xiphophorus helleri Xiphophorus helleri Xiphophorus helleri Xiphophorus maculatus Xiphophorus maculatus
(6%
24 23 23 23 24 23 23
48a 72a 48a 72a 72a 48a 48a 48a
48 (ii, (72) (72) (48) (2) 48 (48)
Family Atherinidae
Melanotaenia maccullochi
24 46
Family Anoplogasteridae
Seopelogadus microlepis bispinosus
Chen 1969
Order GASTEROSTEIFORMES Suborder Gasterosteoidei
Family Gasterosteidae
Ape&es quadracus
46
12m+2Osm+l4ad 12m + 19sm + 15a Q
78 77
Chen et al. 1970a

307
Table 1 (Continued) Species n 2n Karyotype Number of arms

54 54 52
Hetero gametic sex
Reference
Culuea inconstans Gasterosteus acuieatus Gasterosteus aculeatus Gasterosteus wheatlandi Pungitius pungitius Pungitius pungitius Pungitius pungitius Pungitius tymensis Order CHANNIFORMES Family Channidae Channa marulius Channa punctatus Channa punctatus Channa striatus Order SYNBRANCHIFORMES Family Synbranchidae Flu ta alba Order SCORPAENIFORMES Suborder Scorpaenoidei Family Scorpaenidae Helicolenus dactylopterus Pterois lunulata Scorpaena neglecta neglecta Scorpaena neglecta neglecta Scorpaena notata Scorpaena porcus Scorpaena scrofa Scorpaenodes littoralis Sebastes hubbsi Sebastes inermis Sebastes longispinis Sebastes schlegeli Sebasticus marmoratus Family Synanceidae Inimicus japonicus Inimicus japonicus Suborder Hexagrammoidei Family Hexagrammidae Hexagrammus octogrammus Hexagrammus otakii Suborder Congiopodoidei Family Congiopodidae Hypodytes rubripinnis
46 42 42 42 42 42 42 42
8sm + 38a 6m + 6sm + 30a 4m + 6sm + 32a 16m + 12sm + 14a
70
Chen et al. 1970a Post 1965 Chen et al. 1970a Chen et al. 1970a Makino 1934a Post 1965 Chen et al. 1970a Makino 1934a
40 34 32 40
10m 16m 10m 10m
+ + + +
30a 18a 18sm + 4a 30a
(50) (50)
(58)
(50)
Nayyar 1966 Nayyar 1966 Rishi 1973 Nayyar 1966
24
24a
(24)
Kitada et al. 1972
48 48 48 41 34 42 46 36 46 48 48 4s 48
2sm + 46a 2m + 28st + 18a 6m + 32st + 10a 7m + 32st + 8a 18st + 16a 6m + lost + 26a 20st + 26a 4m 2m 2m 2m 2m + 42a + 46a + 46a + 46a + 46a
60)
(78) 86 86 (34) (48) (46) 68
50 50 50
52
50
Sola et al. Nishikawa Nishikawa Nishikawa Sola et al. Sola et al. Sola et al. Arai et al. Nishikawa Nishikawa Nishikawa Nishikawa Nishikawa
1978a et al. 1977a et al. 1977a et al. 1977a 1978a 1978a 1978a 1957a et al. 1977a et al. 1977a et al. 1977a et al. 1977a et al. 1977a
50
48
lOm, sm + 6st + 32a
64
Nogusa 1960 Nishikawa et al. 1977a
24 48
Makino 1937b Nogusa 1960
48
52
Arai et al. 1973a
308
Suborder
Cottoidei
Family Cottidae
Cottus gobio Cot&s gobio Cottus hangiongensis Cottus kazika Cottus nozawae Cottus poecilopus Cottus poNux Cottus pollux Cot&s reinii Trachidermus fasciatus Order PER CYFORMES Suborder Percoidei
24 52 48 40 48 48 48 48 48 40 6m, sm + 46a 6m, sm + 42st, a 18m, sm + 22a lOm, sm + 38st, a 8m, sm + 40a IOm, sm + 38a 12m, sm + 36a 24m, sm + 16a 58 54 58 58 56 58 60 64
24
Post 1965 Starmach 1972 Abe 1972 Abe 1976 Abe 1972 Starmach 1972 Nogusa 1957a Abe 1976 Abe 1976 Abe 1976
Family Centropomidae
Chanda ranga
22 48 48 48 48 48 48 48 48 48 24 24 48 48 48 48 48 46 48 48 41 46 48 48 48 48 48 48 48 46 48a 48a
48a
Post 1965 2st + 46a 2sm + 46st, a 4sm + 44st, a 2sm + 8st + 38a 2m + 46st, a 48a 48a 48a 48a Cataudella et al. 1973b Ueno et al. 1977 Ueno et al. 1977 Rachlin et al. 1978 Arai et al. 1976b Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Post 1965 Post 1965 Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Ohno et al. 1966 Ohno et al. 1966 Ohno et al. 1966 Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Roberts 1964 Thompson et al. 1978
Family Serranidae
Dicentrarchus labrax Coreoperca herzi Coreoperca kawamebari Morone saxa tilis
(48) (50)
(52)
(50)
50
Family Plesiopidae
Plesiops caerulolineatus
Family Centrarchidae
Acantharchus pomotis Ambloplites rupestris Centrarchus macropterus Chaenobryttus gulosus Elassoma evergladei Elassoma zonatum Elassoma zonatum Enneacanthus chaetodon Enneacanthus gloriosus Enneacanthus obesus Lepomis auritus Lepomis cyanellus Lepomis cyanelhis Lepomis cyanellus Lepomis cyanellus Lepomis cyanellus Lepomisgibbosus Lepomis humilis Lepomis machrochirus Lepomis marginatus Lepomis megalotis Lepomis microlophus Micropterus dolomieui Micropterus punctulatus
48 48 48 48 48 48 48 48 48 48 48 48 48 48 48 48 48 48 48 48
48a 48a
48a
48a lm + 46a 2m + 44a 48a 48a 48a 48a 48a 48a 48a 2m, sm + 44st, a
Table 1 (Continued) Species n 2n Karyotype Number of arms

48 (48) (48) 48 48
Heterogametic sex
Reference
Micropterus salmoides Micropterus salmoides Micropterus treculi Pomoxis annularis Pomoxis nigromaculatus
48
46 46 48 48
48a 2m, sm + 44st, a 2m, sm + 44st, a 48a 48a
Roberts 1964 Thompson et al. 1978 Thompson et al. 1978 Roberts 1964 Roberts 1964 Lieder 1963 Nygren et al. 1968 Ross 1973 Ross 1973 Ross 1973 Ross 1973 Ross 1973 Svtidson et al. 1939 Lieder 1963 Nygren et al. 1968a Svlrdson et al. 1939 Nygren et al. 1968a Nogusa 1960
Family Percidae
Acerina cernua Acerina cernua Ethiostoma blennioides Ethiostoma caeruleum Ethiostoma camurum Ethiostoma jlabellare Ethiostoma nigrum Lucioperca Iucioperca Lucioperca lucioperca Lucioperca lucioperca Perca fluviatilis Perca fluvia tilis 24 48 48 48 48 48 48 48 24 46 48 28 48
48a 2m + 46a 4m + 2sm + 42a 2m + 2sm + 44a 4m + 44a 4m + 44a 48a 48a
(48) (50) (54)
(52)
(52)
(52)
(48) (48)
Family Branchiostegidae
Sillago sihama 48
Family Carangidae
Seriola quinqueradiata Trachurus mediterraneus ponticus 48 48
2sm + 2st + 44a 4m + 6sm + 38st. a 2st + 46a 6m, sm + 2st + 40a 6m, sm + 2st + 40a 6m, sm + 16st + 26a 6m, sm + 6st + 34a 2m + 6st + 40a 48a 6m, sm + lost + 32a 8m. sm + lost + 30a 2m + 4sm + 42st. a
(501 58
Ida et al. 1978 Vasilev 1978 Nishikawa et al. 1972 Sola et al. 1978b Vasilev 197 8 Sola et al. 1978b Sola et al. 1978b Sola et al. 1978b Sola et al. 19781, Sola et al. 1978b Sola et al. 1978b Sola et al. 1978b Vasilev 1978 Post 1965
Family Sparidae
Chrysophrys major Diplodus annularis Diplodus annularis Diplodus sargus Lithognathus mormyrus Oblada melanura Page&s acarne Pagellus erythrinus Sarpa salpa Sparus aurata 48 48 48 48 48 46 48 48 48 48 (48) (54) 48 (54) (54)
(52)
(50)
(48)
(54)
(56)
54
Family Sciaenidae
Sciaena umbra 48
Family Monodactylidae
Monodactylus argenteus 24
Family Kyphosidae
Girella melanichthys Microcanthus strigatus 48 48
48a 2sm + 46a 48a
(48) 50
Nishikawa et al. 1972 Arai et al. 1975a Arai et al. 1976b
Family Ephippidae
Platax orbicularis 48 48
310
Table 1 (Continued) Species n 2n Karyotype Number of arms Heterogame tic sex Reference
Family Scatophagidae
Scatophagus argus 48
lm+47ad 48a P 48a 48a 48a 48a 48a 48a 2m + 46a 2sm + 46a 48a 48a 4m + 36sm + 6st + 2a 2m + 46a
(49)
(48)
48 48 48 48 48 48 50 50 48
d?
Khuda-Bukhsh Arai Arai Arai Arai Arai Arai Arai Arai Arai et al. et al. et al. et al. et al. et al. et al. et al. et al.
et al. 1974
Family Chaetodontidae
Centropyge Chaetodon Chaetodon Chaetodon Chaetodon Chaetodon Megaprotodon Megaprotodon Pomacanthus vrolikii auriga collaris lunula trifasciatus vagabundus plebe&s strigangulus semicirculatus 48 48 48 48 48 48 48 48 48
1975a 1975a 1975a 1975a 1975a 1975a 1975a 1975a 1975a
Family Nandidae
Nandus Nandus nandus nandus 24 48 48 (48)
(88)
(50)
Kaur et al. 1965 Manna et al. 1977~ Nishikawa et al. 1972 Post 1965 Post 1965 Nishikawa et al. 1973 Post 1965 Michele et al. 1977a Post 1965 Post 1965 Post 1965 Post 1965 Post 1965 Post 1965 Ohno et al. 1966 Badr et al. 1977a Post 1965 Prasad et al. 1976 Badr et al. 1977a Michele et al. 1977a Badr et al. 1977a Arai Arai Arai Arai Arai Arai Arai Arai Arai et et et et et et et et et al. al. al. al. al. al. al. al. al. 1976a 1976a 1976a 1976a 1976a 1976b 1976b 1976a 1976a
Family Oplegnathidae
Oplegnathus fasciatus 48
Family Cichlidae
Apistogramma pertense Apistogramma ramirezi Cichlasoma citrinella Cichlasoma sever-urn Geophagus brasiliensis Haplochromis multicolor Hemichromis bimaculatus Lamprologus leleupi Nannacara anomala Pelmatochromis kribensis Pterophyllum eimechei Symphysodon aequifasciata Tilapia galileae Tilapia grahami Tilapia mossam bica Tilapia nilo tica Tilapia rendalli Tilapia zilli 24 24 48 24 48 22 22 24 24 24 24 60 44 24 44 40 44 38 3sm
36sm, st + 12a + 45a 51 d
44m + 16a 6st + 38a 2m + 38a 8sm + 36a 2m + 2sm + 34a 4st + 44a 2m + 2st f 44a 2m + 2sm + 2st + 42a 2m + 2sm + 2st + 42a 14m + 16sm + 18st, a 14m + 22sm + lost + 2a 14m + 22sm + 12st, a lm + 46a 48a
(104)
(42) 52 (42)
Family Pomacentridae
Abudefduf leucozonus Abudefduf notatus Abudefduf sordidus Abudefduf vaigiensis Amphiprion clarkii Amphiprion frenatus Amphiprion ocellaris Dascyllus trimaculatum Pomacentrus coelestis 48 48 48 48 48 48 48 41 48 48 50 52 52 78 84 84 48 48
311
Suborder Mugiloidei Family Mugilidae Chelon labrosus Liza aurata L iza ram ada Liza saliens Mugil cephalus Mugil cephalus Mugil corsula Mugil curema Mugil par& Mugil parsia Oedalechilus labeo Suborder Labroidei Family Labridae Pseudolabrus japonicus Suborder Blennioidei Family Blenniidae A trosalarias fuscus Blennius canevae Blennius incognitus Blennius pavo Blennius sanguinolentus Blennius sphinx Blennius trigloides Blennius yatabei Dasson trossulus Istiblennius enosimae Istiblennius lineatus Omobranchus elegans Salarias jasciatus Salarias luctuosus Family Sticheidae Dictyosoma burgeri Dictyosoma burgeri Family Pholididae Pholis pictus Suborder Callionymoidei Family Callionymidae Callionymus richardsonii Suborder Gobioidei Family Gobiidae A boma lactripes Acanthogobiusjlavimanus Acanthogobius j7avimanus Acan thogobius jlavimanus
48 48 48 48 48 48 48 28 48 48 48
2st + 46a 2st + 46a 2st + 46a 2st + 46a 48a 48a 48a 20m + 4st + 4a 48a 48a 2st + 46a
(48) (48) (48) (48) (48) (48) (48) 48 (48) (48) (48)
Cataudella et al. 1974 Cataudella et al. 1974 Cataudella et al. 1974 Cataudella et al. 1974 Cataudella et al. 1974 Le Grande et al. 1976 Khuda-Bukhsh et al. 1974 Le Grande et al. 1976 Chatterjee et al. 1973 Khuda-Bukhsh et al. 1974 Cataudella et al. 1974
46
Nogusa 1960
48 48 48 48 48 48 48 48 40 48 48 42 48 48 48 46 48
48a 8st + 40a 2st + 46a 8st + 40a 8st + 40a 2st + 46a 2m+6sm+18st+22a 6m + 12st + 30a 8m + 32st, a 2m + 46a 48st, a 10m + 2sm + 6st + 24a 48a 48st, a 12m + 18sm + 6st + 12a 14m+18sm+8st+6t
(48) (48) (48) (48)
(56)
54 48 50 48 54 48 48 78 86
Arai et al. Cataudella Cataudella Cataudella Cataudella Cataudella Cataudella Arai et al. Arai et al. Arai et al. Arai et al. Arai et al. Arai et al. Arai et al.
1973c et al. 1975a et al. 1975a et al. 1975a et al. 1975a et al. 1975a et al. 1975a 1974e 1974e 1973~ 1974e 1974e 1973~ 1974e
Arai et al. 1974e Nishikawa et al. 1978 Makino 1937
19
38
Nogusa 1960
40 44 44 44
40st, a 44a lOm, sm + 34st, a 36st + 8a
(40) 54 44
Arai et al. 1974d Nogusa 1960 Nishikawa et al. 1974 Arai et al. 1975b
312
Table 1 (Continued) Species n 2n Karyotype Number of arms 98 46 48 92 78 46 46 52 62 80 56 56 44 60 44 84 46 46 56 64 (46) (46) 44 46 74 Heteroganietic sex Reference
Acentrogobius pflaumi Amblygobius albimaculatus Bathygobius fuscus Brachygobius nunus Boleophthalmus boddaerti Boleoph thalmus glaucus Boleophthalmus pectinirostris Boleophthalmus pectinirostris Boleophthalmus pectinirostris Bostrichthys sinensis Chaenogobius annularis Chaenogobius annularis Chaenogobius castanea Chaenogobius isaza Chaenogobius urotaenia Chasmichthys dolicognathus Chasmichthys gulosus Chasmichthys gulosus Ctenogobius criniger Eleotriodes strigatus Eleotris acanthopomus Gillichthys mirabilis Gillichthys seta Glossogobiusgiuris Glossogobius giuris Gobiodon citrinus Gobioides rubicundus Gobiosoma macrodon Gobiosoma zebrella Gobius abei Luciogobiusguttatus Mogurnda mogurnda Mugilogobius abei Mugilogobius abei Odontamblyopus rubicundus Odontobutis obscura Odontobutis obscura Odontobutis obscura Periophthalmus cantonensis Periophthalmus cantonensis Periophthalmus cantonensis Pterogobius elapoides Pterogobius zonoleucus Quietula guaymasia Quietula y-cauda Rhinogobius brunneus Rhinogobius brunneus Rhinogobius brunneus Rhinogobius brunneus Rhinogobius fiumineus Rhinogobius giurinus
50 44 48 24 46 46 46 46 46 48 44 44 44 44 42 44 44 44 50 44 46 44 44 46 46 43 44 46 38 38 52 44 46 46 46 46 62 44 44 46 46 46 44 44 42 42 44 44 44 44 44 44
48m, sm + 2st, a 2m + 42st, a 48a 12m + 20sm + 14a 46a 46st, a 46st, a 4m, sm + 44st, a 44a 18sm + 26st 36m, sm + 8st, a 12sm + 32st 14sm + 28a 44st, a 16m, sm + 28st, a 44st, a 34m, sm + 6st + 10a 2m + 42st, a 46st, a 12st + 32a 6m + 14st + 24a 46a 46a lm + 42st, a d 2m + 42st, a Q 2m+26sm+18a
Nishikawa et al. 1974 Arai et al. 1974c Arai et al. 1975b Post 1965 Subrahmanyam 1969 Manna et al. 1974b Nogusa 1960 Nishikawa et al. 1974 Arai et al. 1975b Arai et al. 1974c Nogusa 1960 Arai et al. 1975b Nishikawa et al. 1974 Arai et al. 1975b Yamada 1967 Arai et al. 1975b Nishikawa et al. 1974 Arai et al. 1975b Arai et al. 1974d Arai et al. 1974d Arai et al. 1974d Chen et al. 1971 Chen et al. 1971 Kaur et al. 1965 Manna et al. 1974b d Arai et al. 1974d Manna et al. 1974b Ahmed 1974 Ahmed 1974 Lieder 1964a Arai et al. 1973b Arai et al. 1974c Post 1965 Arai et al. 1974c Arai et al. 19751, Nogusa 1960 Arai et al. 1974c Nishikawa et al. 1974 Nogusa 1960 Nishikawa et al. 1974 Arai et al. 1975b Arai et al. 1975b Arai et al. 1975b Cook 1978 Cook 1978 Nogusa 1960 Nadamitsu 1974 Nishikawa et al. 1974 Arai et al. 1975b Arai et al. 1973b Nishikawa et al. 1974 313
6sm + 40st, a 46a 46st, a 4m + 16sm + 26st, a 62a 44st, a 44a 46a 34m, sm + 24st, a 18m + 12sm + 16st, a 14sm + 30st, a 14sm + 3Ost, a 6m + 4sm + 32a 42a 44a 44a 44st, a 44st, a 44a 44st. a
$62) 46 66 44 44 (46) 80 76 58 58 52 42 (44) (44) 44 44 44 44
(62)
Table 1 (Continued) Species n 2n Karyotype Number of arms 58 46 (44) 54 70 54 72 (6604) (44) Heterogame tic sex Reference
Rhodoniichthys Synechogobius Tridentiger Tridentiger Tridentiger Tridentiger Pidentiger Triden tiger Trypauchen Tukugobius Suborder Acanthurus Prionurus Suborder Scomber Thunnus Suborder
laevis hasta obscurus obscurus brevispinis obscurus obscurus obscurus obscurus trigonocephalus trigonocephalus vagina flumineus Acan thuroidei triostegus microlepidotus Scombroidei
42 44 44 44 44 44 44 44 46 44
14m, sm + 28st 2m + 22st, a 44a lOm, sm + 34a 26m, sm + 18st, a lOm, sm + 34st 28m + 16a 16m, sm + 28st, a 12m+6sm+lOst+18a 44a
Arai et al. 1974~ Arai et al. 1975b Nogusa 1960 Arai et al. 1974~ Nishikawa et al. 1974 Arai et al. 1974~ Nishikawa et al. 1974 Arai et al. 1974~ Khuda-Bukhsh 1978 Nadamitsu 1974
Family Acanthuridae 48 48 48a 48a 48 48 Arai et al. 1976a Arai et al. 1976a
Family Scombridae
tapeinocephalus thynnus Anabantoidei
48 48
2st + 46a 2m + 2st + 44a
(48) (50)
Ida et al. 1978 Ida et al. 1978
Family Anabantidae
Anabas testudineus Anabas testudineus Betta splendens Ctenopoma ansorgei Macropodus opercularis Macropodus opercularis
21 24 21
48 46 42 46 48 46 46 d 45 Q 48 16
48a 4m, sm + 42a
(48)
(50)
12m, sm + 34a 48a 24m, sm + 22a 14m + 12sm, st + 20a 14m + 12sm, st + 19a 48a 14m + 2a 14m + 12sm, st + 22a 15m + 12sm, st + 21a 8m+20sm+12st+ +6t+2T 2M+lOm+6sm+ + 12st + lot + 8T 46a 46a
(5%
(48) (70) Q
Kaur Abe Post Post Post Abe
et al. 1965 1975 1965 1965 1965 1975
Family Belontiidae
Colisa Colisa Colisa fasciatus lalia lalius temmincki osphromonoides
Kaur et al. 1965 Abe 1975 Rishi 1976b Abe 1975 Calton et al. 1974
Helostoma Sphaerichthys
(48)
(30)
Family Osphronemidae
Trichogaster Trichogaster Trichogaster Trichogaster Trichogaster Trichogaster Trichogaster fasciatus fasciatus Iabiosus Ieeri sumatranus trichopterus trichoptetus
24
48 d 48 Q 48 48
Rishi 1975 Manna et al. 1977~ Manna et al. 1977~
24
46 24 48 23-24 46
(46) 48 (46)
Abe 1975 Calton et al. 1974 Post 1965 Abe 1975
314
Suborder Macrognathus Mastacembelus Mastacembelus Mastacembelus Rhyncobdelta Rhyncobdella
Mastacembeloidei aculeatum armatus armatus pancalus aculeata aculeata 48 48 48
Family Mastacembelidae 8m + 2sm + 38T 10m + 4m + 2st + 32T 2M + 14m + 6sm + +8st+18t 48a
64
Manna et al. 1977~ Khuda-Bukhsh et al. 1974 Manna et al. 1978 Manna et al. 1977~ Nogusa 1960 Kaur et al. 1965
48 48
(48)
Order PLEURONECTIFORMES Suborder Pleuronectoidei
Family Bothidae
Citarichthys spilopterus Etropus crossotus Paralich thys lethostigma Paralich thys olivaceus Xystreurys liolepis 28 38 48 46 48
14m + 6sm + 2st + 6a 16m + 20sm, st + 2a 48st, a 48a 4sm + 40a 48a 48a 48a 48a 48a 48a 48a 48a 48a
48 56 48 (48)
Le Grande 1975 Le Grande 1975 Le Grande 1975 Nogusa 1960 Ohno et al. 1966 Fukuoka et al. 1970 Fukuoka et al. 1970 Fukuoka et al. 1970 Fukuoka et al. 1970 Fukuoka et al. 1970 Fukuoka et al. 1970 Fukuoka et al. 1970 Fukuoka et al. 1970 Ohno et al. 1966 Barker 1972
Family Pleuronectidae
Cleisthenes pinetorum Kareius bicoloratus Limanda herzensteini Limanda punctatissima Limanda schrenki Limanda yolihamae Microstomus achne Platichthys stellatus Pleuronichthys verticalis Pleuronectes platessa 44 48 48 48 48 48 48 48 48 48 48 (48) (48) (48) (48) (48) (48) (48) (48) (48)
Suborder
Soleoidei
Family Soleidae
Achirus lineatus Solea lascaris nasuta Trinectes maculatus
40 42 40
45 d 46 9
6m + 2sm + 16st + 16a 2m + 6sm + 34st, a 6m+2sm+2st+30a 17m + 6sm + 2st + 20a 18m + 6sm + 2st + 20a
48 50 48 68 70
Le Grande 1975 Vasilev 1978 Le Grande 1975 Le Grande 1975
Family Cynoglossidae
Symphurus plagiusa
d
Order TETRAODONTIFORMES Suborder Balistoidei
Family Balistidae
Amanses sandwichiensis Hemibalistes chtysopterus Novodon modestus Oxymonacanthus Iongirostris Pseudobalistes flavimarginatus Rhinecanthus aculeatus Rhinecanthus verrucosus Rudarius ercodes Stephanolepis cirrhifer 40 46 40 36 44 44 44 36 34
40st, a 46a 36a 2m + 42st, a 44a 44a 36a
40 46 36 46 44 44 36
20
Arai et al. 1976a Arai et al. 1976a Nogusa 1960 Arai et al. 1976a Arai et al. 1976a Arai et al. 1976a Arai et al. 1976a Arai et al. 1976a Nogusa 1960 315
Table 1 (Continued) Species n 2n Karyotype Number of arms 54 Heterogametic sex Reference
Family Ostraciodontidae
Ostraciodon Suborder tuberculatus Tetraodontoidei 50
4sm + 46st, a
Arai et al. 1976a
Family Tetraodontidae
Canthigaster rivulata Chelonodon patoca Fugu niphobles Sphaeroides chvsops Tetraodon immaculatus Tetraodon nigropunctatus
34 40 44 44 42 38 46
4m+6sm+lOst+14a 14m + 16sm + lost, a 6m + 14sm + 24st, a 14m + 16sm + 12st 14m + 20sm + 4st 6m + 6sm + 34st, a
44 70 64 64 72 72 58
Arai Arai Arai Arai Arai Arai
et et et et et et
al. al. al. al. al. al.
1976c 1976c 1973a 1976c 1976c 1976c
Family Diodontidae
Diodon bleekeri
Arai et al. 1976c
Crossopterygii
ORDER Lepidosiren DIPNOI paradoxa
Family Lepidosirenidae 38 34 38m (76) Ohno et aL 1966 Wickbom 1945 Family Protopteridae
Protopteius annectens
Brachiopterygii
ORDER POL YPTERIFORMES calabaricus thys calabaricus delhezi endlicheri congicus palmas palmas
Family Polypteridae
Calamoichthys Calamoich Polypterus Polypterus Polyptetus Polyptencs
36 36 36 36 36 36
32m 30m 32m 32m 24m 32m
+ 4st + 6st + 4st + 4st + 12sm + 4st
(68) 72
(38) (68)
72
Capanna et al. 1973a Denton et al. 1973 Cataudella et al. 1978 Cataudella et al. 1978 Denton et aL 1973 Cataudella et al. 1978
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New Developments in Vertebrate Cytotaxonomy

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NEW DEVELOPMENTS IN VERTEBRATE CYTOTAXONOMY III. KARYOLOGY OF BONY FISHES: A REVIEW L.SOLA, S.CATAUDELLA & E.

[Kallman, 19731, in which the sex

K.R. Lewis pp. 139-147. Ohno, S. (1974). <Animal cytogenetics.

OSTEICHTHYES Actinopterygii SUPERORDER CHONDROSTEI

239 i 7 116+4 116+4 112 *

Fontana Fontana Fontana Ohno et m,sm = arms; st, a = 1 arm.

et al. 1974 et al. 1974 et al. 1974 al. 1969b

* When in brackets, number of arms calculated by the present authors as following:

Table 1 (Continued) Species n 2n Kwotype Number of arms Heterogametic sex Reference

Dingerkus et al. 1976

28m + 14a + 26minute

Ohno et al. 1969b

Ohno et al. 1969b

12m+8sm+l8a 14m + 4sm + 20a 20m + 14sm + 4st lOm+16sm+4st+8a

Ohno et al. 1968

4m + 12sm, st + 40t lsm+ lst+54a 4sm + lost + 36t

Urushido et al. 1975 Uyeno 1973 Urushido et al. 1975

40m, sm + I Ost 12m+36a 42a 42a 42a 4sm + 30a 42a

100 60 42 (42) (42) 38 42

Hetero gametic sex

Oncorhynchus Oncorhynchus Oncorhynchus Oncorhynchus Oneorhynchus Oncorhynchus Oncorhynchus Oncorhynchus Oncorhynchus

52 14 60 60 66 66 58 58 58(56) 58(57) 56 66 66 68 68 ii: 78 64 14 18 58 80 80 58 80 68 64 64 60 60(58)

56 80 77-82 84 80 80 80 80 80 82 78 84 84 84 84 84 82 80 82 84 74 102 102 80 66 66 60 72 70 70 69 82 92-106

35m, sm + 4st + 27a 35m, sm + 4st + 27a 38m + 8sm + 14a

105 105 106

14m +56a 15m + 3sm + 4st + 60a

(84) 91 (100) 133-159

12m, st + 44a 10m + 48a 16m + 38a

Yamazaki 1971 Svardson 1945 Nygren et al. 1971d

Table 1 (Continued) Species n 2n Karyotype Number of arms Heterogametic sex Reference

Ebeling et al. 1971a

Campos 1972 Campos 1972 Merrilees 1975 Campos 1972

(2:) (Z) 50 (50) 50 (50) 50

22m 22m, sm 22m

(44) (44) (44)

Post 1974 Post 1974 Post 1974

Table 1 (Continued) Species n 2n Karyotype Number of arms Heterogametic sex Reference

Suborder Saurida Saurida Synodus

Myctophoidei elongata undosquamis lucioceps

1974 1974 1974 1974 1974

Chen et al. 1974

Order GONOR YNCHLFORMES Suborder Chanoidei

14m + 4sm + 14a

Arai et al. 1976~

Order CYPRINIFORMES Suborder Characoidei

54m, 44m, 32m, 2m + 54m,

sm sm + 8st, a sm + 22st 32sm + 16a sm

60m, 34m, 50m, 30m,

76 (76) (76) (72)

(82) (60) (68)

100 100 100

34m, sm + 6st + 1Oa

Table 1 (Continued) Species n 2n Karyotype Number of arms 148

(252) (160) (160) (252) (160) (160) (252) (332)

Table 1 (Continued) Species n 2n Kawotype Number of arms Heterogametic sex Reference

(50) (48) (50)

Table 1 (Continued) Species n 2n Karyotype Number of arms Heterogametic sex Reference

68 76 12m + 30sm, st + 8a 13m + 35a 42m 20m + 12sm + 12st + 6a

8m + 90sm. a 12m + 86sm, a

(62) (68) (64)

(44) 104 102 102 102

Table 1 (Continued) Species n 2n Karyotype Number of arms Heterogametic sex Reference

52a 16m + 8sm + 14st + 12a