44
NEW DEVELOPMENTS IN VERTEBRATE CYTOTAXONOMY” III. KARYOLOGY OF BONY FISHES: A REVIEW L.SOLA, S.CATAUDELLA & E. CAF’ANNA 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 cyto- logical 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 karyo- type 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 chro- mosomes 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 evolu- tion (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 inter- esting . * 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 ‘Cytotaxo- nomy and Vertebrate Evolution’ (A.B. Chiarelli & E. Capan- no, eds, Academic Press, London & New York, 1973) - Eds. The presence of diversified karyotypes in restrict- ed 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 in- dicative 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 out- lined by recent studies. For example, new evidence suggests that the karyotype can play a primary role in speciation (cf. White, 1978) contrary to Mayr’s original hypothesis (1963) which stated that only polypIoidy and geographical isolation lead to the for- mation 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% 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. 285

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Page 1: New Developments in Vertebrate Cytotaxonomy

NEW DEVELOPMENTS IN VERTEBRATE CYTOTAXONOMY” III. KARYOLOGY OF BONY FISHES: A REVIEW

L.SOLA, S.CATAUDELLA & E. CAF’ANNA

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 cyto- logical 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 karyo- type 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 species or 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 chro- mosomes 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 evolu- tion (Fig. 1) and it is therefore often impossible to establish a definite correlation between karyotype and karyotype transformations and to identify the processes involved.

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 inter- esting .

* 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 ‘Cytotaxo- nomy and Vertebrate Evolution’ (A.B. Chiarelli & E. Capan- no, eds, Academic Press, London & New York, 1973) - Eds.

The presence of diversified karyotypes in restrict- ed 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 in- dicative 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 out- lined by recent studies. For example, new evidence suggests that the karyotype can play a primary role in speciation (cf. White, 1978) contrary to Mayr’s original hypothesis (1963) which stated that only polypIoidy and geographical isolation lead to the for- mation 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%

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. 285

Page 2: New Developments in Vertebrate Cytotaxonomy

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 Engrauli- dae [Ohno et al. 19691). In some groups currently considered to be in expansion, such as the Salmonidae (Fukuoka 1972, Thorgaard 1976, 77, Ohno et al. 1965), cases of chromosomal polymorphism are known and they sliould be followed closely.

Fig.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.

In the Poecilidae speciation may occur by inter- specific hybridization. This apparently applies to Poecilia formosa whose hybrid origin is widely ac- cepted on the basis of morphological and karyologi- cal 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 mo- Aacha-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 prob- lems. They display almost all possible forms of sexu- ality, ranging from simultaneous or successive herma- phroditism 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 allo- patric populations of the same species may display either female or male heterogamety (e.g. in Xipho- phorus maculatus [Kallman, 19731, in which the sex chromosomes are identified by means of pigment gene

286

Page 3: New Developments in Vertebrate Cytotaxonomy

markers showing a sex-linked hereditary pattern). Uyeno & Miller (1971, 1972) described several Mexi- can 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 (Cypri- nidae, Centrarchidae , etc .) evidence is frequently found of interspecific and even intergeneric hybrids where sy’mpatric 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 success in 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 con- clusions about phyletic relationships from the karyo- logical comparison of different species, karyotype analyses may supplement other taxonomic evalua- tions. The systematic definition of a species depends on a thorough morphological analysis, and the karyo- type is only one characteristic among several. How- ever, 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 develop- mental stages can be followed, which is usually dif- ficult. In such cases a knowledge of the karyotype and the existence of diversified ktiological situa- tions, if any, can provide a precise diagnostic criterion suitable for solving identification problems (Fig. 3).

However, this point also illustrates the most im- portant 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 re- arrangements, except for such ‘dramatic’ events as translocations and inversions or fusions between whole chromosomes.

In the fishes, an accurate morphological character- isation of the chromosomes is hindered by the large

Fig. 2. Karyotypes of fish hybrids: (a) Salmo trutta x Sal- velinus fontinalis. 2n = 82 and marker chromosomes in single copies (arrows) indicating the combination of the two parent- ial 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

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a

b

C Fig. 3. Karyotype diversity in the genus Scorpaena: (a) S. scrofa L., 2n = 46; - (b) S. porcus L., 2n = 42; - (c) S. notata Raf., 2n = 34.

number and the small size of the chromosomes. The banding techniques commonly used in the karyo- logical 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 tech- niques (G-, Q-, R-, etc.) must be encouraged and ex- panded, 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 differ- entiations between the karyotypes of-related species.

Furthermore, the karyological evidence must be integrated with quantitative and qualitative evalua- tions of DNA contents, autoradiographic and in situ hybridization evidence, and electrophoresis data on isoenzymes, especially when populations with chro- mosome 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 im- prove 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 in- cludes 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.

Page 5: New Developments in Vertebrate Cytotaxonomy

References K.R. Lewis pp. 139-147.

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.

Ohno, S. (1974). <Animal cytogenetics’. Chord&a I. Proto- chordata, Cyclostomata and Pisces. Borntraeger, Berlin- Stuttgart 1974.

Ojima, Y., K. Ueno & M. Hayashi (1976). A review of the chromosome numbers in fishes. La Kromosomo II, 1: 1947.

Greenwood, P.H., D.E. Rosen, S.H. Weitzman & G.S. Myers (1966). Phyletic studies of teleostean fishes, with a provi- sional 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.

Park, E.H. (1974). A list of the chromosome numbers of fishes. Coil. Rev. Coil. Lib. Arts & Sciences, Seoul Nut. Univ. 20 (l-2): 346-372. 1

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 vivi- parous fish Poeciliopsis. Science 157: 1564-1567.

Schultz, R.J. (1969). Hybridization, unisexuality and poly- ploidy 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 chromo- somes of the rainbow trout (Salmo gairdneri). otogenet. Cell Genet. 17 (4): 174-186.

Nevo, E. & C.R. Shaw (1972). Genetic variation in a sub- terranean 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). Post- zygotic 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 &

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). Poly- ploidization in the fish family Cyprinidae, order Cyprini- formes. I. DNA content and chromosome sets in various species of Cyprinidae. Humangenetik 7: 240-244.

Table 1

Species

OSTEICHTHYES

n 2n Karyotype Number Hetero- Reference of gametic arms* sex

Actinopterygii

SUPERORDER CHONDROSTEI

Order A CIPENSER IFORMES Family Acipenseridae

Acipenser naccurii 239 i 7 Fontana et al. 1974 Acipenser sturio 116+4 Fontana et al. 1974 Huso huso 116+4 Fontana et al. 1974 Scaphirhvnchus pkztyrhynchus 112 * Ohno et al. 1969b

* When in brackets, number of arms calculated by the present authors as following: m,sm = arms; st, a = 1 arm.

289

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Table 1 (Continued)

Species n 2n Kwotype Number Hetero- Reference of gametic arms sex

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

120 32m+8sm+8a+ 72michrochromosomes

68 28m + 14a + 26minute Ohno et al. 1969b

46 20m + 26a

38 24m + 8st + 6a 38 12m+lOsm+16a 38 22m, sm + 16a 38 20m, sm + 18a 38 22m, sm + 16a 38 20m, sm + 18a 38 12m+8sm+18a 38 14m + 6st + 18a 38 22m, sm + 1 &a

50 Subrahmanyah et al. 1971

38 12m+8sm+l8a 38 14m + 4sm + 20a

38 20m + 14sm + 4st 16 Nishikawa et al 1977b 38 lOm+16sm+4st+8a 68 Nishikawa et al. 1977b

48 52 54 52 52

48a (48)

8m + 44a

66-70

(Z)

(66) Ohno et al. 1969b

(6’3 Q? Chiarelli et al. 1969a (60) Q Passakas 1978 (60) - Sola et al. 1980 (58) - Nishikawa et al. 197 1 (60) - Sola et al. 1980 (58) - Nishikawa et al. 1971 (58) Q Park et al. 1979 (52) Q? Ohno et al. 1973 (601 - Sola et al. 1980

(58) Q Park et al. 1979 56 Nishikawa et al. 197713

Dingerkus et al. 1976

Mayers et al. 1969 Roberts 1966 Skvortsova 1975 Skvortsova 1975 Ohno et al. 1968

290

Page 7: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero Reference of gametic arms sex

Family Engraulidae Engraulis mordax

SUPERORDER OSTEOGLOSSOMORPHA

Order OSTEOGLOSSIFORMES Suborder Osteoglossoidei

Family Osteoglossidae Arapaima gigas Osteoglossum bicirrhosum Scleropages formosus

Suborder Notopteroidei Family Hiodontidae

Hiodon alosoides

Family Notopteridae Notoptenrs chitala Notopterus chitata Notopterus notopterus Notopterus notopterus Notoptenrs notopterus Papyrocranus afer Xenomystus nigri

SUPERORDER PROTACANTHOPTERYGII

Order SALMONIFORMES Suborder Salmonoidei

Family Salmonidae Brachymystax lenok Coregonus albula Coregonus artedii Coregonus clupeaformis Coregonus hoyi Coregonus lavaretus Coregonus lavaretus Coregonus lavaretus Coregonus lavaretus baeri Coregonus nasus Coregonus nasus Coregonus oxyrhynchus Coregonus peled Coregonus peled Coregonus pidschian Coregonus pidschian Coregonus reighardi Coregonus ussuriensis Coregonus zenithicus Oncorhynchus gorbuscha

48 48a (48) Ohno et al. 1968

56 4m + 12sm, st + 40t 56 lsm+ lst+54a 50 4sm + lost + 36t

50 40m, sm + I Ost 100

48 12m+36a 60 Nayyar 1965 42 42a 42 Uyeno 1973 42 42a (42) Srivastava et al. 1964 42 42a (42) Urushido et al. 1975 42 Prasad et al. 1976 34 4sm + 30a 38 Uyeno 1973 42 42a 42 Uyeno 1973

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 Kang et al. 1973a 96 Nygren et al. 1971a

106 Booke 1968 108 Booke 1968

98 Booke 1968 96 Svardson 1945

118 Karbe 1964 92(98) Nygren et al. 1971c

(96) Prokofieva 1934 96 Karbe 1964 92 Nygren et al. 1971b 96 Karbe 1964 92 Viktoroski 1964 92(98) Nygren et al. 1971b 96 Karbe 1964 98(92) Nygren et al. 1971b

104 Booke 1968 100 Viktoroski et al. 1978

98 Booke 1968 104 Simon 1963

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

Uyeno 1973

291

Page 8: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero Reference of gametic arms sex

Oncorhynchus gorbuscha Oncorhynchus ketu Oncorhynchus kisutch Oncorhynchus kisutch Oneorhynchus masou masou Oncorhynchus masou ishikawue Oncorhynchus nerka Oncorhynchus nerka Oncorhynchus nerka

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.

292

52 14 60 60 66 66 58 58 58(56)

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

100 Muramoto et al. 1974 106 Sasaki et al. 1968 112 Simon 1963 106 Uyeno 1972 124 Muramoto et al. 1974 124 Muramoto et al. 1974 104 Sasaki et al. 1968 104 Muramoto et al. 1974 104(102) Gorshkova 1978

58(57) 44(45)m + 2st + 12(10)a 104 Fukuokal972b 56 46m, sm + 10a 102 Simon 1963 66 34m, sm + 24st + 8a 124 Muramoto et al. 1974

66 68 68

ii: 78 64 14 18 58 80 80 58 80 68 64 64 60 60(58)

64 90 60 58-60 60 59 58 60 58-65 60 60 60 56 58 58 51 56 54-56 58

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

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

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

104 104

12 12 12 14 74 12 12 72 14

d? d? d

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 Vasil’ev 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

Page 9: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero Reference of gametic arms sex

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

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

Family Plecoglossidae Plecoglossus altivelis

Family Osmeridae Osmerus eperlanus Osmerus eperlanus

56 80 77-82 84 80 80 80 80 80 82

49m, sm + 7a 20m + 60a

16m + 2st : 62a 14m + 6sm + 2st + 58a

16m + 2st + 62a 16m + 2st + 62a 12m, sm + 70a

78 84 84 84 84 84 82 80 82 84 74

102 102

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

80

66

66

60

72

70 70 69 82

92-106

35m, sm + 4st + 27a 105

35m, sm + 4st + 27a 105

38m + 8sm + 14a 106

14m +‘56a

15m + 3sm + 4st + 60a

56 12m, st + 44a

58 10m + 48a 54 16m + 38a

105 100 102

(96) (100)

(96) (96) (94)

96 (100) 100 106

(100) (100)

(98) 98

102 100 108

170

(84) 91

(100)

133-159

68

(68) 70

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

Sasaki et al. 1968

Sasaki et al. 1968

Uyeno 1972

Uyeno 1972

Prokofieva 1934 Svardson 1945 Nygren et al. 1975b Capanna et al. 1973b

Cataudella et al. 1975b

Yamazaki 1971

Svardson 1945 Nygren et al. 1971d

293

Page 10: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

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 Stomiatoidei Family Gonostomatidae

Bonapartia pedaliota Gonostoma bathyphilum Gonostoma elongatum

44 17m + 27a (d) 61 Ebeling et al. 1971a

60 lm + 16m + 43a (d) 77 2m + 16m + 42a (0) 78

54 64

18 36 19m + 14sm + 2sat + la (d)

71

Ebeling et al. 1971a, 1971b

Chen 1969 Chen 1969 Chen et al. 1966

64 82 Ebeling et al. 1971a

38 22 22

64 42

(2) (50)

d?

d

d?

6

Campos 1972 Campos 1972 Merrilees 1975

30

10m + 16sm + 12a 8m+12sm+2a 8m + 1Osm + 4st lm + 18sm + lla (d) 2m + 18sm + 10a (Q) Campos 1972

50 50 50 46 48 50 50 50 50 50 50 50 50 50

50a 50a 50a

50a 50a 50a 50a 50a 50a 50a 50a 50a

(Z, (50)

(2:)

(Z) 50

(50) 50

(50) 50

(44) (44) (44)

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

78 48 22 22 22

22m 22m, sm 22m

Beamish et al. 1971a Beamish et al. 1971a Beamish et al. 1971a Kligerman et al. 1975 Beamish et al. 1971a

24 Post 1974 6 Post 1974

24 Post 1974

294

Page 11: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Suborder Myctophoidei Family Synodontidae

Saurida elongata Saurida undosquamis Synodus lucioceps

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

Lepidophanes guentheri Lepidophanes photothorax Lepidophanes supralateralis Lobianchia gemelari Myctophum nitidulum Notoscopelus resplendens Protomyctophum crockeri Stenobranchius leucopsarus Symbolophorus californiensis Triphoturus mexicanus

Family Neoscopelidae Scopetengys tristis

Order GONOR YNCHLFORMES Suborder Chanoidei

Family Chanidae Chanos chanos

SUPERORDER OSTARIOPHYSI

Order CYPRINIFORMES Suborder Characoidei

Family Characidae Anostomus anostomus Chnlceus maerolepidotus Chaleeus macrolepidotus Hemigrammus caudovittatus Lepolinus fasciatus

48 48a (48) Q Nishikawa et al. 1978 48 48a (48) Q Nishikawa et al. 1978 48 28m, sm + 20a (76) Chen et al. 1974

48 24 24 24 24 24 24

48 24 24 24

48 49 48 41 48

24 24 24 24

48 24 48

48 48 48 48

48a (48)

48a (48) lm + 48a (6) (50) 48a (48) lm + 46a (d) (48) 2m + 46a (0) (50)

d?

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

48a (48)

48a (48) 48a (48) 2m, sm + 46a (50) 48a (48)

d Chen et al. 1974

Post 1912 Post 1972 Post 1972 Post 1972 Chen et al. 1974 Post 1972 Chen et al. 1974 Chen et al. 1974 Chen et al. 1974 Chen et al. 1974

48 2m, sm + 46a (50) Chen et al. 1974

32 14m + 4sm + 14a 50 Arai et al. 1976~

54 54m, sm (108) Ojima et al. 1976 52 44m, sm + 8st, a (96) Ojima et al. 1976 54 32m, sm + 22st 636) Muramoto et al. 1968 50 2m + 32sm + 16a (84) Fontana et al. 1970a 54 54m, sm (108) Ojima et al. 1976

295

Page 12: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Methinis schreiinuelleri 62 60m, sm + 2a Myloplus arnoldi 58 34m, sm + 24st, a Mylossoma duriventris 54 50m, sm + 4a Serrasalmus hollandi 64 30m, sm + 16st + 18a

Suborder Gymnotoidei Family Apteronotidae

Apteronotus albifrons 24 14m+2sm+2st+6a l (40) Howell 1972

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

25 50 52

26 52 50 52

22 44 25 50

48 48

22 44 48 44 44 44 44 44 44 44 50 50 52 48 52

100 100 100

52 52

18m

30m 82

8m + 20sm, st + 20a 76 10m + 18sm + 20st, a (76)

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

26m + 18sm + 18st + 38a

30m, sm + 4st + 16a + 2min.

Barbus meridionalis 100 Barbus meridionalis petenyi 100 Barbus (Barbodes) pentazona 24 Barbus schuberti 24 Barbus semifasciolatus 26 52 Barbus (Capoetaj tetrazona 24 Barbus tetrazona 50 Barbus (Capoeta) titteya 24 Barbus (Barbodes) viviparus 24 Brachidanio (Danio) frankei 24 Capoeta oligolepis 24 Carassius auratus 41 94

22m + 20sm + 12st + 46a (142)

34m, sm + 6st + 1Oa (84)

296

(122)

(104) (94)

IO

(76) (72) (72)

16 (72)

16 (72) (72) (76) (82) (60) (68)

94 148

(144)

(8’3

Ojima et al. 1976 Ojima et al. 1976 Ojima et al. 1976 Muramoto et al. 1968

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 Vasil’ev 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

Page 13: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Carassius aura&s 100 Carassius aura&s 104 Carassius auratus 100 Carassius aura&s auratus 100 Carassius auratus buergeri 100 Carassius auratus buergeri 156 Carassius auratus cuvieri 100 Carassius auratus gibeho 100 Carassius auratus gibelio 156 Carassius auratus grandoculis 100 Carassius auratus langsdorfii 100 Carassius auratus langsdorfii 156 Carassius auratus langsdorfii 206 Carassius carassius 47 94 Carassius carassius 100 Catla ca tla 25 50 Chela bacaila 52 Chela bacaila 50 Chondrostoma soetta 50 Chondrostoma toxostoma 50 Chrosomus eos 50 Chrosomus erythrogaster 50 Cirrhina reba 48 Crossocheilus iatius punjabensis 48 Ctenopharyngodon idellus 48 Ctenopharyngodon idellus 48 Ctenopharyngodon idellus 48 Cyprinus carpio 100 Cyprinus carpio 104

Cyprinus carpio 98 Cyprinus carpio 100 Danio albolineatus 24 Danio albolineatus 50 Danio devario 50 Danio malabaricus 50 Danio rerio 50 Danio rerio 50 Danio (Brachidanio) rerio 50 Esomus danrica 25 50 Exoglossum maxilingua 48 Garra rufa 22-25 Gila bicolor 50 Cila orcu tti 50 Gnathopogon coerulescens 50 Gnathopogon elongatus 50 tinathopogon elongatus elongatus 50 Gobio albipinnatus vladykovi 50 Gobio bessleri banaticus 50 Gobio gobio 24 Gobio gobio 50 Gobio gobio 50

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

148 (150) (160) (148) (160) (252) (160) (160) (252) (160) (160) (252) (332)

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 (96) 18m+20sm+6st+4a (86) 12m + 36a (60)

16m + 20sm, st + L2a 14m + 20sm + 8st + 6a (2) 12m + 36sm, st + 52a 46m, sm + 18st + 36a + (150) 4min.

24m + 24sm, st + 52a

10m + 39sm, st + la 10m + 40a 10m + 40a

10m + 39sm + la 10m + 12sm, st + 14a

WO (60)

(99)

12m+16sm+lOst+12a (78)

44m, sm + 6a 94 14m t 24st t 12a (64)

12m + 32sm, st + 6a 94 28m + 20sm + 2a 98 24m + 20sm + 4st + 2a 98

22m + 24sm + 2st + 2a 98 38m + 12a (88)

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

Page 14: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Gobio uranoscopus 52 Hemibarbus barbus 50 Hemibarbus labeo 50 Hemibarbus longhostris 25 50 Hemibarbus longirostris 50 Hemigrammocypris rasborella 24 48 Hesperoleucus symmetricus 50 Hesperoleucus symmetricus 50 Hybopsis plumbea 50 Hypophthalmichthys molitrix 48 Hypophthalmichthys molitrix 48 Hypophthalmichthys molitrlx 48 Ischikauia steenackeri 48 Ischikauia steenackeri 48 Labeo bata 25 50 Labeo chrysophekadion 50 Labeo gonius 54 Lavinia exilicauda 50 Lepidomeda albivallis 50 Lepidomeda mollispinis 50 Levidomeda vittata 50 Leuciscus albidus alborella 50 Leuciscus aula 50 Leuciscus cephalus 50 Leuciscus cephalus 50 Leuciscus cephalus 50 Leuciscus cephalus albus 50 Leuciscus cephalus cabeda 50 Leuciscus cephalus cephalus 50 Leuciscus leuciscus 50 Leuciscus idus 50 Leuciscus souffio agassizi 50 Leuciscus souffia muticellus 50 Leuciscus souffia muticellus 50 Leuciscus svallize 50 Leuciscus svallize svallize 50 Leuciscus turskyi 50 Leuciscus turskyi tenellus 50 Meda fulgida 50 Moroco jouyi 50 Moroco lagowskii 50 Moroco oxycephalus 50 Moroco percnurus 27 Moroco steindachneri 27 54 Moroco steindachneri 50 Mylopharodon conocephalus 50 Notemigonus crysoleucas 50 Notemigonus crysoleucas 50 Notropis cornutus 50 No tropis lu trensis 50 Notropus texanus 50 Notropis venustus 50

298

24m+12sm+12st+4a 100 14m + 2&m, st + 8a 94 18m + 22sm, st + 10a 90

14m + 28sm, st + 4a 92

20m + 20st + 10a 42m, sm + 8a

(70) 92

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

88 99

(82) (54) 92

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

70 86

(78) 86 90 88 92

32m, sm + 18a

34m, sm + 18a 18m + 24sm, st + 8a 14m + 22sm + 8st + 6a 12m + 2&m, st + 10a 12m +2&m, st + 10a

(82) 74 82 68 84

92 90 90

14m + 22sm + 8st + 6a 44m, sm + 6a

44m, sm + 6a 6m + 40sm + 4a

2m + 42sm + 6a 2m + 38sm + 10a

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 Vasil’ev 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

Page 15: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Kawotype Number Hetero- Reference of gametic arms sex

Opsariichthys uncirostris 66 Opsariichthys uncirostris 78 Opsopoeodus (Notropis) emiliae 48 Orthodon microlepidotus 50 Oxygaster bacaila 48 Pachychilon pictum 50 Paraphoxinus adspersus 50 Paraphoxinus alepidotus 50 Paraphoxinus alepidotus 50 Paraphoxinus croaticus 50 Paraphoxinus metohiensis 50 Paraphoxinus pstrossi 50 Pelteobagrus nudiceps 56 Phoxinus percnurus 27 Phoxinus phoxinus 50 Phoxinus phoxinus 50 Pimephales promelas 50 Plagopterus argentissimus 50 Pogonichthys macrolepidotus 50 Pseudogobio esocinus 26 52 Pseudogobio esocinus 50 Pseudoperilampus typus 44 Pseudorasbora parva 25 50 Pseudorasbora parva 50 Ptychocheilus grandis 50 Pungtungia herzi 50 Pungtungia herzi 50 Pungtungia hilgendorfi 50 Puntius conchonius 50 Puntius conchonius 24 Puntius conchonius 50 Puntius filamentosus 24 Puntius sophore 50 Puntius stigma 48 Puntius stigma 50 Puntius stigma 48 Puntius ticto 50 Rasbora buchanani 50 Rasbora heteromorpha 24 Rasbora trilineata 24 Rhinichthys atratulus 50 Rhinichthys cataracta 50 Rhodeus atremius 46 Rhodeus ocellatus 44 Rhodeus ocellatus ocellatus 48 Rhodeus ocellatus ocellatus 48 Rhodeus ocellatus smithii 48 Rhodeus sericeus amarus 48 Rhodeus suigensis 46 Richardsonius egregius 50 Rohtee cotio 48 Rut&s rubilio 50

8sm, st + 70a 86 2m + 30sm + 16a (80) 44m, sm + 6a 94 48a (48)

38m, sm + 12a 26m, sm + 24a 28m, sm + 22a 36m, sm + 14a 32m, sm + 18a 30m, sm + 20a

32m, sm + 18a 82 44m + 12sm + 16st + 8a 86

18m + 26sm, st + 6a 14m, sm + 6a 94

16m + 28sm, st + 6a 14m + 14sm + 16a

14m + 36sm, st 42m, sm + 8a

94

(72)

100 92

20m + 30sm, st

50a

100

(50)

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

0’6) 18 86 82 80

(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 Vasil’ev 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

Page 16: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Rutilus rutilus 52 Rutilus rutilus 50 Rutilus rutilus 50 Rutilus rutilus 50 Sarcocheilichthys variegatus 25 50 Sarcocheilichthys variegatus 50 Scardinius erythrophthalmus 48 Scardinius erythrophthalmus 48 Scardinius erythrophthalmus 50 Semotilus atromaculatus 52 Semotilus corporalis 52 Semotilus margarita 50 Tanakia tanago 48 Tinca tinca 48 Tinca tinca 48 Tinca tinca 48 Tinca tinca 48 Tribolodon ezoe 50 Tribolodon ezoe 50 Tribolodon hakonensis 50 Tribolodon hakonensis 50 Tribolodon hakuensis 25 50 Typhlogarra widdowsoni 24 Vimba vimba 52 Zacco platypus 48 Zacco temminckii 48

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

100 98 96-98 98 96-98

100 100

50 100 100

96-98 96-98 96-98

Family Cobitidae Acanthophthalmus khulli Barbatula oreas Barbatula toni Botia macracantha Cobitis biwae Cobitis biwae Cobitis biwae Cobitis biwae Cobitis biwae Cobitis biwae

50 24 48

50 98

27 54 96 48 96 48 48

300

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

10m + 18sm + 20st, a (76)

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 (76) 18m + 22sm. st + 8a 88 18m + 22sm, st + 8a 88

8m + 90sm. a

12m + 86sm, a

14m. sm + 12st + 24a (64)

16m, sm + 34st, a (66) 28m, sm + 70a (126)

58m, sm + 38st, a (154) 18m + 30sm (96) 36m + 52sm + 8st (184) 2m, sm + 46a (50) 20m + 22sm, st + 6a 90

68 76

90 (82)

84 (68) 92 92 92 92

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 1 a 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

Page 17: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero Reference of gametic arms sex

Cobitis biwae 96 Cobitis delicata 50 Co bitis delieata 50 Cobitis taenia 50 Cobitis taenia taenia 86 Cobitis taenia taenia 50 Cobitis taen& taenia 86 Cobitis taenia taenia 94 Cobitis taenia striata 50 Cobitis taenia striata 50 Cobitis taenia striata 98 Lefua echigonia 25 50 Lefua nikkonis 50 Leptobotia curta 50 Misgurnus anguillicaudatus 26 52 Misgurnus anguillicaudatus 50 Misgurnus fossilis 24 Misgurnus fossilis 100

Order SIL URIFORMES Family Ictaluridae

Ictalurus nebulosus Ictalurus punctatus

Family Bagridae Coreobagrus lshikaua Liobagrus reini Mystus gulio Mysius seenghala Mystus tengara

56 38 58 50 54

Mystus tengara 54

Mystus vittatus 50 Pseudobagrus aurantiacus 56 Pseudobagrus aurantiacus 48 Rita chrysea 54 Rita rita 54

Family Siluridae

Callichromis bimaculatus

Ompok bimaculatus Parasilurus asotus Parasilurus asotus Parasilurus biwaensis Parasilurus lithophilus Wallago attu

Family Schilbeidae Clupisoma parua Eutropiichthys vacha

60 20m, sm + 40a 80 Berberovic et al. 1975 56 16m, sm + 22st + 18a (72) Muramoto et al. 1968

41 d 17m + 12sm,mt + 12a 42 9 18m + 12sm, st + 12a 40 4m + 36a 58 38m, sm + 8st + 12a 58 44m, sm + 14st, a 58 44m, sm + 14st, a 58 44m, sm + 14st, a 86 86a

66 58

32m + 54sm, st + 10a 182 18m, sm + 32st, a (68) 18m, sm + 32a (68) 12m + 6sm + 16st + 16a (68) 64m, sm + 22a (150) 12m+4sm+34a 66 32m + 32sm, st + 22a 150 26m + 32sm, st + 36a 152 16m, sm + 34a (66) 12m+4sm+34a 66 20m + 22sm + 56a 140

12m, sm + 38st, a (62) 8m + 1Osm + 16st + 16a (68)

14m, sm + 36a (64)

36m, sm + 64a 136

28m + 24sm + 4st, a (108) 28m + 1Osm 66 30m + 12sm + 2st + 140 102

86 10m + 44a (64) 10m + 38sm + 6a (d) 102 9m + 38sm + 7a (Q) 101

24m + 12sm + 20st, a (92) 20m + 12sm + 16st, a (80) 28m + 20sm + 6st 102 16m + 38a (70)

(44) 104 102 102 102 (86)

66a 10m + 20sm + 12st + 4t + 12T

(66) 100

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

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

d 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

301

Page 18: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Family Clariidae Clarias batrachus Clarias batrachus Clarias batrachus Clarias fuscus

Family Heteropneustidae Heteropneustes fossilis Heteropneustes fossilis Heteropneustes fossilis Heteropneustes fossilis

Family Plotosidae Plo~osus anguillaris

Family Pimelodidae Pimelodella sp. Rhamdia hilarii

Family Loricariidae Hypostomus plecosromus Loricaria macrodon Loricaria parva

Plecostomus ancistroides

Plecostomus macrops Plecosromus paulinus Plecosiomus s09gatieeps

SUPERORDER PARACANTHOPTERYGII

Order GADIFORMES Suborder Gadoidei

Family Gadidae Gadus morrhua Gaidropsarus medirerraneus Micromesistius poutassou Pollachius pollachius Pollachius virens Raniceps raninus Trisopterus minutus

SUPERORDER ATHERINOMORPHA

Order A THERINIFORMES Suborder Exocoetoidei

Family Belonidae Xenentodon cancila

52 52 50 56

52a

16m + 8sm + 14st + 12a

(52) Nayyar 1966 58 Srivastava et al. 1968

(74) Rishi 1976b 88 Arai et al. 1974a

58 56 56 56

58a 18m + 18sm + 20a 14m + 26sm + 16a 18m + 18sm + 20a

(58) Nayyar 1966 (92) Vasudevan et al. 1973 (96) Prasad et al. 1974 (92) Rishi 1976b

48 12m, sm + 36st, a 60 Arai et al. 1974b

46 28m + 1Osm + 8a 84 Toledo et al. 1976b 62 36m+18sm+8a 116 Toledo et al. 1976b

54 58

24

24m, sm + 12st + 18a (78) 18m + 2sm + 38a 78

Muramoto et al. 1968 Michele et al. 1977b Post 1965

68

68 14 74

10m + 27sm + 31a d 105 10m + 28sm + 30a Q 106 10m + 14sm + 44a 92 10m + 20sm + 44a 104 8m + 4sm + 62a 86

d Michele et al. 1977b

Michele et al. 1977b Michele et al. 1977b Michele et al. 1977b

46 28 44 38 40 48 48

24m, sm + 4a

10m + 28a 10m + 30a 30m + 18a

50 Srivastava et al. 1964

52

48 50 78

Nygren et al. 1974 Vasil’ev 1978 Nygren et al. 1974 Nygren et al. 1974 Nygren et al. 1974 Nygren et al. 1974 Nygren et al. 1974

302

Page 19: New Developments in Vertebrate Cytotaxonomy

Table I (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Suborder Cyprinodontoidei Family Oryziatidae

Oryzias latipes Oryzias melastigma

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

48 48

32 24 48 24 24 24 24 24 48 18

38 15 30 20 40 19 38

38 38 36 32

17 34 34 34 34 30 26

20 40 19 38 16 32 17 34

34 32 32 30 28 30 24 26

10 20 9 18

20 40 26 28 30

18 36 18 36

30 32 30 32

64

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

20m, sm + 28a 68 Ojima et al. 1969a 50 Scheel 1972

Scheel 1972b Karbe 196 1 Karbe 1961 Karbe 1961 Karbe 1961 Karbe 196 1 Scheell968a Scheel 1968a Scheel 1972b Scheel 1968a Scheel1968a, 1972b Scheel 1968a, 1972b Scheel 1972b Scheell972b Scheel 1972b Scheel 1972b Scheel 1968a, 1972b Scheel 1972b Scheel1972b Scheel 1972b Scheel 1972b Scheel1972b Scheel 1968a, 1972b Scheel 1968a, 1972b Scheel 1968a Scheel 1968a, 1972b Scheel 1972b Scheel1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1968a, 1972b Scheel1968a, 1972b Scheel1968a, 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel1968a, 1972b Scheel 1968a, 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b

303

Page 20: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetew Reference of gametic arms sex

Aphyosemion franzwerneri Aphyosemion gardneri Aphyosemion gardneri Aphyosemion gardneri Aphyosemion gardneri Aphyosemion gardneri Aphyosemion geryi Aphyosemion guineense Aphyosemion gulare Aphyosemion labarrei Aphyosemion labarrei Aphyosemion liberiense Aphyosemion louessense Aphyosemion melanopteron Aphyosemion mirabile Aphyosemion mirabile Aphyosemion mirabile Aphyosemion mirabile Aphyosemion nidanum Aphyosemion obscurum Aphyosemion occidentale Aphyosemion ogoense Aphyosemion petersii Aphyosemion roloffi Aphyosemion santaisabellae Aphyosemion scheeli Aphyosemion schoutedeni Aphyosemion seymouri Aphyosemion sjoestedti Aphyosemion toddi Aphyosemion walkeri Aplocheilichthys jlavipinnis Aplocheilichthys katangae Aplocheilichthys macrophthalmus Aplocheilichthys normani Aplocheilichthys spilauchen Aplocheilichthys spilauchen Aplocheilus barmoiensis Aplocheilus bifasciatus Aplocheilus blocki Aplocheilus chaperi Aplocheilus dageti Aplocheilus dayi Aplocheilus duboisi Aplocheilus eskanus Aplocheilus fasciolatus Aplocheilus fasciolatus Aplocheilus grahami Aplocheilus grahami Aplocheilus grahami Aplocheilus lineatus Aplocheilus lineatus

11 22 20 40 18 36

38 40 36 40 38

16 32 14 28

26 42

10 20 30 32 34 36 38

20 40 17 34

46 40 40 42

20 40 40 22 28

20 40 46

18 36 24 24

48 48 48

24 17 34 20 40 24 48 25 50 25 50 24 48

48 21 42

40 19 38

48 23 46

48 25 50

48

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

60

Scheel 1968a, 1972b Scheel 1968a, 197213 Scheel1968a, 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1968a, 1972b Scheel1968a, 1972b Scheel 1972b Scheel 1972b Scheel 1968a, 1972b Scheel 1972b Scheel 1972b Scheel 1972b c Scheel 19721, Scheel 1972b Scheel 1968a, 1972b Scheel 1968a Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1972b Scheel 1968a, 1972b Scheel 1972b Scheel 1972b Scheel 1972b . Scheel 1968a, 1972b Scheel 1972b Scheel1968a, 1972b Post 1965 Post 1965 Scheel 1972b Scheel1972b Scheel 1972b Post 1965 Scheel 1972b Scheel 1968a, 1972b Scheel1968a, 1972b Scheel 1968a, 1972b Scheel 1968a, 1972b Scheel 1968a, 1972b Scheel 19721, Scheel 1968a, 1972b Scheel 1972b Scheel 1968a, 1972b Scheel 1972b Scheel 1968a, 1972b Scheel 1972b Scheel1968a, 1972b Scheel 1972b

304

Page 21: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero Reference of gametic arms sex

Aplocheilus panchax 38 Aplocheilus panchax 18 36 Aplocheilus sangmelinensis 24 48 Aplocheilus sexfasciatus 24 48 Aplocheilus spilargyreius 17 34 Aplocheilusquemeri 48 Austrofundulus dolichopterus 22 44 Austrofundulus transilis 44 Cynolebias bellotti 48 Cynolebias bellotti 24 Cynolebias nigripinnis 24 Cynolebias nigripinnis 48 Cynolebias whitei 24 Cynolebias whitei 46 Cynopoecilus ladigesi 24 Cynopoecilus melanotaenia 24 Cynopoecilus melanotaenia 44 Cyprinodon macularis 24 Fundulus chrysotus 34 Fundulus cingolatus 46 Fundulus confluentus 48 Fundulus diaphanus 48 Fundulus diaphanus 48 Fundulus diaphanus 48 Fund&s diaphanus 48 Fundulus grandis 48 Fundulus heteroclitus 24 Fundulus heteroclitus 48 Fundulus heteroclitus 48 Fundulus heteroclitus 48 Fundulus kansae 48 Fundulus lineolatus 46 Fundulus luciae 32 Fundulus majalis 48 Fundulus majalis 48 Fundulus majalis 48 Fundulus notatus 40 Fundulus notatus 48 Fundulus notatus 40 Fundulus notatus 44 Fundulus notti 46 Fundulus olivaceus 48 Fundulus olivaceus 48 Fundulus olivaceus 48 Fundulus olivaceus 48

Fundulus parvipinnis 48

Fundulus parvipinnis 48 Fundulus pulvereus 48 Fundulus rathbuni 48 FunduIus sciadicus 44 Fundulus seminolis 48

14m + 20a (4sat) 48 2m + 44a (2sat) 48 48a (2sat) 48 4sm + 44a (2sat) 52 lm + 3sm + 42a + 2sat 52 4sm + 42a + 2sat 52 4sm + 38a + 6t (52) 2sm + 46a (2sat) 50

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

(ii) 48 48 52 50 50

(48) 52 52 50 50 48 52 52 54 50 49 50 48 48 50 50 48

56

50

88 54 80 52

72

92

52

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 1971 Chen 1971 Chen 1971 Chen 1971 Chen 1971

305

Page 22: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Heterw Reference of gametic arms sex

Fund&us similis 48 Fundulus waccamensis 46 Jordanella floridae 24 Jordanella floridae 48 Kosswigichthys asquamatus 24 Nothobranchius guentheri 19 Nothobranchius kirki 36 Nothobranchius mayeri 19 Nothobranchius orthonotus 18 Nothobranchius palmquisti 19 Nothobranchius palmquisti 18 Nothobranchius rachovii 8 16 Nothobranchius rachovii 18 Nothobranchius thirty 42 Nothobranchius thirry 44 Orestias agassize 24 Orestias luteus 24 Pachypanchax playfairi 24 48 Pachypanchax playfairi 48 Procatopus aberrans 48 Procatopus nigromarginatus 24 Procatopus nototaenia 24 Procatopus roseipinnis 24 Procatopus similis 24 48 Pterolebias longipinnis 10 Pterolebias longipinnis 20 Pterolebias peruensis 34 54 Rivulus cylindraceus 48 Rivulus dorni 48 Rivulus harti 44 Rivulus marmoratus 48 Rivulus milesi 46 Rivulus ocellatus 48 + Rivulus ornatus 40 Rivulus strigatus 46 Rivulus urophthalmus 46 Valencia hispanica 48

Family Goodeidae Ameca splendens Characodon later&s Hubbsina turneri Xenotoca eiseni Xenotoca melanosoma Xenotoca variata

26 22m + 2st + 2a (48) Miller et al. 1971 24 24m (48) Uyeno et al. 1972a 48 48a (48) Uyeno et al. 1972a 48 6st + 42a (48) Fitzsimons 1972 48 8st + 40a (48) Fitzsimons 1972 48 4st + 44a (48) Fitzsimons 1972

Family Poecilidae

Gambusia affinis

Gambusia affnis

Gambusia affinis

48

48

48

306

2smf2sat) + 46a 4sm (2sat) + 42a

50 50

60

58

30 48 48

96 96

96

20 90 56 70 56 52 92 54 66 88 86 92

2sm + 46a d (50) lm+2sm+45a P (51) lm + 47a 9 (4% 48a d (48) Im+47aQ (49)

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,

P Chen et al. 1968

O? Yosida et al. 1971

Q Itahashi et al. 1975

Page 23: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Gambusia affinis holbrooki 48 Gambusia holbrooki 24

Gambusia gaigei 48

Gambusia hurtadoi 48

Gambusia marshi 42

Gambusia nobilis 48

Gambusia regani 48 Gambusia vittata 48 Lebistes reticulatus 46 Limia melanogaster 24 Mollienisia latipinna 24 Mollienisia sphenops 24

Mollienisia sphenops 46

Mollienisia velifera 23 46 Platypoecilus maculatus 48 Poecilia formosa 46 Poecilia formosa 46 Poecilia formosa 69 Poecilia latipinna 24 Poecilia latipinna 23 46 Poecilia latipunctata 23 Poecilia mexicana 23 46 Poecilia reticulatus 46 Poecilia sphenops 24 Poecilia sphenops 23 46 Poecilia velifera 23 46 Poeciliopsis lucida 48 Poeciliopsis lucida 72 Poeciliopsis monacha-lucida 48 Poeciliopsis monacha-2lucida 72 Poeciliopsis 2monacha-lucida 72 Xiphophorus helleri 48 Xiphophorus helleri 48 Xiphophorus helleri 48 Xiphophorus helleri 48 Xiphophorus maculatus 48 Xiphophorus maculatus 48

Family Atherinidae Melanotaenia maccullochi 24

Family Anoplogasteridae

48a d, Q

48a d lm +47aQ 48a d lm+47aQ 42a 48a d lm +47aQ 48a 48a 46a

46a d Im+45aQ

46a 1st + la + 44minute 1st + 2a + 66minute

48a 72a 48a 72a 72a 48a

48a

48a

48

(48) (49) (48) (49) 42

(48) (49) (48) (48) (46)

(46) (47)

46 (46) (6%

48

(ii, (72) (72) (48)

(2) 48

(48)

Seopelogadus microlepis bispinosus

Order GASTEROSTEIFORMES Suborder Gasterosteoidei

Family Gasterosteidae

Ape&es quadracus

46

46 12m+2Osm+l4ad 78 12m + 19sm + 15a Q 77

Cataudella et al. 1977a Post 1965

Q Campos et al. 1971

Q 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

Chen 1969

Q Chen et al. 1970a

307

Page 24: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero Reference of gametic arms sex

Culuea inconstans 46 Gasterosteus acuieatus 42 Gasterosteus aculeatus 42 Gasterosteus wheatlandi 42 Pungitius pungitius 42 Pungitius pungitius 42 Pungitius pungitius 42 Pungitius tymensis 42

Order CHANNIFORMES Family Channidae

Channa marulius Channa punctatus Channa punctatus Channa striatus

40 10m + 30a (50) Nayyar 1966 34 16m + 18a (50) Nayyar 1966 32 10m + 18sm + 4a (58) Rishi 1973 40 10m + 30a (50) Nayyar 1966

Order SYNBRANCHIFORMES Family Synbranchidae

Flu ta alba 24 24a (24) Kitada et al. 1972

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

48 48 48

41 34 42 46 36 46 48 48 4s 48

Family Synanceidae Inimicus japonicus Inimicus japonicus

50 48

Suborder Hexagrammoidei Family Hexagrammidae

Hexagrammus octogrammus Hexagrammus otakii

24 Makino 1937b 48 Nogusa 1960

Suborder Congiopodoidei Family Congiopodidae

Hypodytes rubripinnis 48 52 Arai et al. 1973a

8sm + 38a

6m + 6sm + 30a 4m + 6sm + 32a

54

54 52

16m + 12sm + 14a 70

2sm + 46a 2m + 28st + 18a 6m + 32st + 10a 7m + 32st + 8a 18st + 16a 6m + lost + 26a 20st + 26a

4m + 42a 2m + 46a 2m + 46a 2m + 46a 2m + 46a

lOm, sm + 6st + 32a 64

60) Sola et al. 1978a (78) Nishikawa et al. 1977a

86 Nishikawa et al. 1977a 86 Nishikawa et al. 1977a

(34) Sola et al. 1978a (48) Sola et al. 1978a (46) Sola et al. 1978a

68 Arai et al. 1957a 50 Nishikawa et al. 1977a 50 Nishikawa et al. 1977a 50 Nishikawa et al. 1977a 52 Nishikawa et al. 1977a 50 Nishikawa et al. 1977a

Chen et al. 1970a Post 1965 Chen et al. 1970a Chen et al. 1970a Makino 1934a Post 1965 Chen et al. 1970a Makino 1934a

Nogusa 1960 Nishikawa et al. 1977a

308

Page 25: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

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

Family Centropomidae Chanda ranga

Family Serranidae Dicentrarchus labrax Coreoperca herzi Coreoperca kawamebari Morone saxa tilis

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

24 52 48 40 48 48

24 48 48 48 40

6m, sm + 46a 58 6m, sm + 42st, a 54 18m, sm + 22a 58 lOm, sm + 38st, a 58 8m, sm + 40a 56

IOm, sm + 38a 58 12m, sm + 36a 60 24m, sm + 16a 64

Post 1965 Starmach 1972 Abe 1972 Abe 1976 Abe 1972 Starmach 1972 Nogusa 1957a Abe 1976 Abe 1976 Abe 1976

22 Post 1965

48 2st + 46a (48) Cataudella et al. 1973b 48 2sm + 46st, a (50) Ueno et al. 1977 48 4sm + 44st, a (52) Ueno et al. 1977 48 2sm + 8st + 38a (50) Rachlin et al. 1978

48 2m + 46st, a 50 Arai et al. 1976b

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 48 48a 48 48a 48 48a 48

48a 48a 48a 48a 48a

48a 48a lm + 46a 2m + 44a 48a 48a 48a 48a 48a 48a 48a 2m, sm + 44st, a

48 48 48 48 48 48 48 48 48 48 48 48 48 48 48 48

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

Page 26: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Micropterus salmoides 48 Micropterus salmoides 46 Micropterus treculi 46 Pomoxis annularis 48 Pomoxis nigromaculatus 48

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

Family Branchiostegidae Sillago sihama 48 Nogusa 1960

Family Carangidae Seriola quinqueradiata Trachurus mediterraneus ponticus

48 2sm + 2st + 44a (501 Ida et al. 1978 48 4m + 6sm + 38st. a 58 Vasil’ev 1978

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

Family Sciaenidae Sciaena umbra 48 2m + 4sm + 42st. a 54 Vasil’ev 1978

Family Monodactylidae Monodactylus argenteus 24 Post 1965

Family Kyphosidae Girella melanichthys Microcanthus strigatus

48 48a (48) Nishikawa et al. 1972 48 2sm + 46a 50 Arai et al. 1975a

Family Ephippidae Platax orbicularis 48 48a 48 Arai et al. 1976b

310

48a 2m, sm + 44st, a 2m, sm + 44st, a 48a 48a

48a 2m + 46a 4m + 2sm + 42a 2m + 2sm + 44a 4m + 44a 4m + 44a

48a (48)

48a (48)

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

48 Roberts 1964 (48) Thompson et al. 1978 (48) Thompson et al. 1978 48 Roberts 1964 48 Roberts 1964

(48) (50) (54) (52) (52) (52)

(48) Nishikawa et al. 1972 (54) Sola et al. 1978b

48 Vasil’ev 197 8 (54) Sola et al. 1978b (54) Sola et al. 1978b (52) Sola et al. 1978b (50) Sola et al. 19781, (48) Sola et al. 1978b (54) Sola et al. 1978b (56) Sola et al. 1978b

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

Page 27: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of game tic arms sex

Family Scatophagidae

Scatophagus argus 48

Family Chaetodontidae Centropyge vrolikii Chaetodon auriga Chaetodon collaris Chaetodon lunula Chaetodon trifasciatus Chaetodon vagabundus Megaprotodon plebe&s Megaprotodon strigangulus Pomacanthus semicirculatus

48 48a 48 Arai et al. 1975a 48 48a 48 Arai et al. 1975a 48 48a 48 Arai et al. 1975a 48 48a 48 Arai et al. 1975a 48 48a 48 Arai et al. 1975a 48 48a 48 Arai et al. 1975a 48 2m + 46a 50 Arai et al. 1975a 48 2sm + 46a 50 Arai et al. 1975a 48 48a 48 Arai et al. 1975a

Family Nandidae Nandus nandus Nandus nandus

48 24 48

Family Oplegnathidae Oplegnathus fasciatus 48 2m + 46a (50) Nishikawa et al. 1972

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

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

lm+47ad (49) 48a P (48)

48a 4m + 36sm + 6st + 2a

36sm, st + 12a

3sm + 45a

44m + 16a (104)

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

d? Khuda-Bukhsh et al. 1974

(48) Kaur et al. 1965 (88) Manna et al. 1977~

51

(42) 52

(42)

Post 1965 Post 1965 Nishikawa et al. 1973 Post 1965

d 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

48 Arai et al. 1976a 50 Arai et al. 1976a 52 Arai et al. 1976a 52 Arai et al. 1976a 78 Arai et al. 1976a 84 Arai et al. 1976b 84 Arai et al. 1976b 48 Arai et al. 1976a 48 Arai et al. 1976a

311

Page 28: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

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

312

48 2st + 46a (48) Cataudella et al. 1974 48 2st + 46a (48) Cataudella et al. 1974 48 2st + 46a (48) Cataudella et al. 1974 48 2st + 46a (48) Cataudella et al. 1974 48 48a (48) Cataudella et al. 1974 48 48a (48) Le Grande et al. 1976 48 48a (48) Khuda-Bukhsh et al. 1974 28 20m + 4st + 4a 48 Le Grande et al. 1976 48 48a (48) Chatterjee et al. 1973 48 48a (48) Khuda-Bukhsh et al. 1974 48 2st + 46a (48) Cataudella et al. 1974

46 Nogusa 1960

48 48a 48 8st + 40a 48 2st + 46a 48 8st + 40a 48 8st + 40a 48 2st + 46a 48 2m+6sm+18st+22a 48 6m + 12st + 30a 40 8m + 32st, a 48 2m + 46a 48 48st, a 42 10m + 2sm + 6st + 24a 48 48a 48 48st, a

(48) (48) (48) (48) (56) 54 48 50 48 54 48 48

Arai et al. 1973c Cataudella et al. 1975a Cataudella et al. 1975a Cataudella et al. 1975a Cataudella et al. 1975a Cataudella et al. 1975a Cataudella et al. 1975a Arai et al. 1974e Arai et al. 1974e Arai et al. 1973~ Arai et al. 1974e Arai et al. 1974e Arai et al. 1973~ Arai et al. 1974e

48 12m + 18sm + 6st + 12a 78 Arai et al. 1974e 46 14m+18sm+8st+6t 86 Nishikawa et al. 1978

48

19 38

40 44 44 44

Makino 1937

Nogusa 1960

40st, a 44a

(40)

lOm, sm + 34st, a 54 36st + 8a 44

Arai et al. 1974d Nogusa 1960 Nishikawa et al. 1974 Arai et al. 1975b

Page 29: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of ganietic arms sex

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 fi’umineus 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 98 2m + 42st, a 46 48a 48

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

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

92 78

46 46 52

62 80 56 56 44 60 44 84 46 46 56 64

(46) (46) 44 46 74

$62) 46 66

(62) 44 44

(46) 80 76 58 58 52 42

(44) (44) 44 44 44 44

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

Page 30: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of game tic arms sex

Rhodoniichthys laevis 42 Synechogobius hasta 44 Tridentiger obscurus 44 Tridentiger obscurus brevispinis 44 Tridentiger obscurus obscurus 44 Tridentiger obscurus obscurus 44 Pidentiger trigonocephalus 44 Triden tiger trigonocephalus 44 Trypauchen vagina 46 Tukugobius flumineus 44

Suborder Acan thuroidei Family Acanthuridae

Acanthurus triostegus Prionurus microlepidotus

Suborder Scombroidei Family Scombridae

Scomber tapeinocephalus Thunnus thynnus

Suborder Anabantoidei Family Anabantidae

Anabas testudineus Anabas testudineus Betta splendens Ctenopoma ansorgei Macropodus opercularis Macropodus opercularis

Family Belontiidae Colisa fasciatus Colisa lalia

Colisa lalius

Helostoma temmincki Sphaerichthys osphromonoides

Family Osphronemidae

Trichogaster fasciatus

Trichogaster fasciatus

Trichogaster Iabiosus 24 48

Trichogaster Ieeri 46 Trichogaster sumatranus 24 48 Trichogaster trichopterus 23-24 Trichogaster trichoptetus 46

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

58 46

(44) 54 70 54 72

(6604) (44)

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

48 48a 48 Arai et al. 1976a 48 48a 48 Arai et al. 1976a

48 2st + 46a (48) Ida et al. 1978 48 2m + 2st + 44a (50) Ida et al. 1978

48 46

21 42 24 21

46

48a (48) 4m, sm + 42a (50)

12m, sm + 34a (5%

Kaur et al. 1965 Abe 1975 Post 1965 Post 1965 Post 1965 Abe 1975

48 46 46 d 45 Q 48 16

48a 24m, sm + 22a 14m + 12sm, st + 20a 14m + 12sm, st + 19a 48a

(48) Kaur et al. 1965 (70) Abe 1975

14m + 2a (48) (30)

Q Rishi 1976b

Abe 1975 Calton et al. 1974

24 48 d 48 Q 48

14m + 12sm, st + 22a 15m + 12sm, st + 21a 8m+20sm+12st+ +6t+2T 2M+lOm+6sm+ + 12st + lot + 8T 46a

Q Rishi 1975

Manna et al. 1977~

Manna et al. 1977~

(46) 48

46a (46)

Abe 1975 Calton et al. 1974 Post 1965 Abe 1975

314

Page 31: New Developments in Vertebrate Cytotaxonomy

Table 1 (Continued)

Species n 2n Karyotype Number Hetero- Reference of gametic arms sex

Suborder Mastacembeloidei Family Mastacembelidae

Macrognathus aculeatum Mastacembelus armatus Mastacembelus armatus Mastacembelus pancalus

48 48 48

8m + 2sm + 38T

10m + 4m + 2st + 32T 2M + 14m + 6sm + +8st+18t

64

Manna et al. 1977~ Khuda-Bukhsh et al. 1974 Manna et al. 1978 Manna et al. 1977~

Rhyncobdelta aculeata 48 Rhyncobdella aculeata 48 48a (48)

Nogusa 1960 Kaur et al. 1965

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 48 16m + 20sm, st + 2a 56 48st, a 48

48a (48)

Le Grande 1975 Le Grande 1975 Le Grande 1975 Nogusa 1960 Ohno et al. 1966

Family Pleuronectidae Cleisthenes pinetorum Kareius bicoloratus Limanda herzensteini Limanda punctatissima Limanda schrenki Limanda yolihamae Microstomus achne Platichthys stellatus Pleuronichthys verticalis Pleuronectes platessa

44 4sm + 40a 48 Fukuoka et al. 1970 48 48a (48) Fukuoka et al. 1970 48 48a (48) Fukuoka et al. 1970 48 48a (48) Fukuoka et al. 1970 48 48a (48) Fukuoka et al. 1970 48 48a (48) Fukuoka et al. 1970 48 48a (48) Fukuoka et al. 1970 48 48a (48) Fukuoka et al. 1970 48 48a (48) Ohno et al. 1966 48 48a (48) Barker 1972

Suborder Soleoidei Family Soleidae

Achirus lineatus Solea lascaris nasuta Trinectes maculatus

Family Cynoglossidae

40 42 40

6m + 2sm + 16st + 16a 48 Le Grande 1975 2m + 6sm + 34st, a 50 Vasil’ev 1978 6m+2sm+2st+30a 48 Le Grande 1975

Symphurus plagiusa 45 d 46 9

17m + 6sm + 2st + 20a 18m + 6sm + 2st + 20a

68 70

d Le Grande 1975

Order TETRAODONTIFORMES Suborder Balistoidei

Family Balistidae Amanses sandwichiensis 40 Hemibalistes chtysopterus 46 Novodon modestus 20 40 Oxymonacanthus Iongirostris 36 Pseudobalistes flavimarginatus 44 Rhinecanthus aculeatus 44 Rhinecanthus verrucosus 44 Rudarius ercodes 36 Stephanolepis cirrhifer 34

40st, a 40 46a 46

36a 36 2m + 42st, a 46 44a 44 44a 44 36a 36

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

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Table 1 (Continued)

Species

Family Ostraciodontidae Ostraciodon tuberculatus

n 2n Karyotype Number Hetero- Reference of gametic arms sex

50 4sm + 46st, a 54 Arai et al. 1976a

Suborder Tetraodontoidei Family Tetraodontidae

Canthigaster rivulata Chelonodon patoca Fugu niphobles Sphaeroides chvsops Tetraodon immaculatus Tetraodon nigropunctatus

34 4m+6sm+lOst+14a 44 40 14m + 16sm + lost, a 70 44 64 44 6m + 14sm + 24st, a 64 42 14m + 16sm + 12st 72 38 14m + 20sm + 4st 72

Arai et al. 1976c Arai et al. 1976c Arai et al. 1973a Arai et al. 1976c Arai et al. 1976c Arai et al. 1976c

Family Diodontidae Diodon bleekeri 46 6m + 6sm + 34st, a 58 Arai et al. 1976c

Crossopterygii

ORDER DIPNOI Family Lepidosirenidae

Lepidosiren paradoxa

Family Protopteridae Protopteius annectens

38 38m

34

(76) Ohno et aL 1966

Wickbom 1945

Brachiopterygii

ORDER POL YPTERIFORMES Family Polypteridae

Calamoichthys calabaricus Calamoich thys calabaricus Polypterus delhezi Polypterus endlicheri congicus Polyptetus palmas Polyptencs palmas

36 32m + 4st 36 30m + 6st 36 32m + 4st 36 32m + 4st 36 24m + 12sm 36 32m + 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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