Chromosomal studies on Coscoroba coscoroba (Aves: Anseriformes) reinforce the Coscoroba–Cereopsis clade

The Coscoroba (Coscoroba coscoroba), endemic to southern South America, is traditionally considered as an early branch from the common ancestor leading to true geese and swans. Recently, an interesting association between the Coscoroba and Cape Barren goose (Cereopsis novaehollandiae) as sister groups has been proposed. We present here the characterization of the karyotype of C. coscoroba using whole chromosome probes derived from Gallus gallus macrochromosomes. Our data showed that C. coscoroba has the highest diploid number among Anseriformes (2n = 98), and the conservation of macrochromosome pairs 1–10 indicates that the increase in diploid number has occurred by fission events involving only the microchromosomes. Moreover, the similarity between the diploid numbers of C. coscoroba (2n = 98) and Cereopsis novaehollandiae (2n = 92) reinforces the phylogenetic position of these two species as sister groups, considering that other species of geese and swans have diploid numbers close to 2n = 80. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 274–279.     

Karyology and phylogeny of some Mesoamerican species of Zamia (Zamiaceae)

Chromosome numbers and karyotypes of four species of Zamia L. (Zamiaceae) are described. Plants of Z. manicata from Colombia are 2n = 18 with eight metacentric (M), four submetacentric (S), two acrocentric (A), and four telocentric (T) chromosomes. Plants of Z. ipetiensis from Panama are 2n = 23 with 3M + 4S + 2A + 14T. Plants of Z. cunaria from Panama have two different chromosome numbers, 2n = 23 with 3M + 4S + 2A + 14T and 2n = 24 with 2M + 4S + 2A + 16T. Plants of Z. acuminata from Costa Rica and Panama are 2n = 24 with 2M + 4S + 2A + 16T. On the basis of the occurrence of a one-to-two-ratio in the variation of M- and T-chromosome numbers in the karyotypes, centric fission or fusion are considered for their potential involvement in the chromosome variation of these plants. Data deriving from morphology and karyology, interpreted in a cladistic framework, suggest that centric fission rather than centric fusion is involved in the karyotype diversification of the four species and their closest Mesoamerican allies.     

Evidence for eight tandem and five centric fusions in the evolution of the karyotype ofAethomys namaquensis A. Smith (Rodentia: Muridae)

G- and C-banded chromosomes ofAethomys namaquensis (2n=24),A. chrysophilus (2n=44), andPraomys coucha (2n=36) are compared and contrasted with publised material on Australian Muridae and North American Sigmodontidae. Direction and types of chromosomal rearrangements are established using cladistic methodology. An acrocentric morphology for chromosomes 5, 14, 15 and 20 (numbering system fromPeromyscus) are proposed as primitive for the common ancestor of the Muridae and Sigmodontidae rodent lineages. Reduced diploid number ofAethomys namaquensis is derived by eight tandem and five centric fusions since divergence from the common ancestor withA. chrysophilus. The two species ofAethomys share one derived metacentric chromosome that distinguishes them fromPraomys. Praomys has unique chromosomes which can be derived from the proposed primitive condition by five centric fusions and five pericentric inversions. It is concluded that karyotypic orthoselection for tandem and centric fusions is best explained by cellular or biochemical mechanisms rather than variation in population characteristics.     

Heteromorphic sex chromosomes of lizardfish (Synodontidae): focus on the ZZ-ZW1W2 system in Trachinocephalus myops

The karyotype and DNA content of four lizardfish species (family Synodontidae), that is, Saurida elongata, Synodus ulae, Synodus hoshinonis and Trachinocephalus myops, were analyzed. The karyotype of T. myops significantly differed from that of the other three species having diploid chromosome number of 48 with mainly acrocentric chromosomes and the ZZ-ZW sex chromosome system. The chromosome number of male T. myops was 2n=26, while that of female T. myops was 2n=27. The karyotype consisted of 11 pairs of metacentrics, one pair of acrocentrics and, in addition, two large metacentrics in the male and a single large metacentric, a distinctly small subtelocentric and a microchromosome in the female. C-banding demonstrated that in the female the subtelocentric chromosome and the microchromosome were heterochromatic. The karyotype of T. myops was thought to be derived from a 48 chromosome type synodontid fish through the involvement of Robertsonian rearrangement; the rearrangement of the sex chromosomes proceeded during karyotype evolution. Among the chromosomes, the large metacentrics were determined to be neo-Z (a fusion of the original Z and an autosome), the microchromosomes the W₁ (originally W), and the subtelocentric chromosomes the W₂ (derived from an autosome pair). The miniaturization of W₁ and W₂ chromosomes and their heterochromatinization suggested that sex chromosomes in this species have been already highly differentiated. The findings on DNA content implied that the karyotype of T. myops evolved by centric fusion events without loss in DNA amount.     

Karyotypes of parasitic Hymenoptera: Diversity, evolution and taxonomic significance

Haploid chromosome numbers (n) of parasitic Hymenoptera (= traditional Parasitica + Chrysidoidea) vary from 2 to 23. However, this range can be subdivided into three intervals with n= 14–23 (less derived parasitic wasps, e.g., some Ichneumonidae and Braconidae as well as Gasteruptiidae), 8–13 (many other parasitic Hymenoptera) and 2–7 (Dryinidae, the majority of Chalcidoidea and some advanced Braconidae, e.g. Aphidiinae). The symmetric karyotype with a relatively high chromosome number (n= 14–17) and the prevalence of biarmed chromosomes must be considered as a groundplan feature of parasitic Hymenoptera. Independent reductions of chromosome numbers (n≤ 10–11) occurred in some groups of the superfamily Ichneumonoidea as well as in the common ancestor of the Proctotrupoidea sensu lato, Ceraphronoidea, Cynipoidea and Chalcidoidea. Further multiple decreases in chromosome numbers (n≤ 4–6) took place in some Braconidae, various lineages of the superfamily Chalcidoidea as well as in the family Dryinidae. Two main trends prevailed in the karyotype evolution of parasitic wasps: the reduction of chromosome numbers (mainly due to tandem fusions and less frequently due to centric ones) and karyotypic dissymmetrization (through an increase in size differentiation of chromosomes and/or in the share of acrocentrics in a chromosome set). Although karyotypic features of parasitic Hymenoptera can be used for solving taxonomic problems at various levels, this method is the most effective at the species level.     

Chromosomal evolution in serpentes; a comparison of G and C chromosome banding patterns of some colubrid and boid genera

G and C-chromosome banding techniques have been used to compare the structure of the karyotype in a variety of colubrid and boid snakes. The comparison of G-band patterns indicates that while some band sequences have been conserved, either as whole chromosomes or entire arms, there is also evidence of considerable rearrangement especially in the smaller chromosomes. In the colubrid Elaphe subocularis there is also evidence that there has been a relocation of the centromere on chromosome 2 without any accompanying inversion in the sequence of G-bands. Finally, G-banding has facilitated the demonstration of a simple pericentric inversion distinguishing the Z and W chromosomes in Acrantophis dumereli. This represents the first report of differentiated sex chromosomes in a boid snake. The combined banding data thus indicates that snake chromosomes are certainly not lacking in variability. The use of C-banding to detect constitutive heterochromatin has confirmed that in some boids and colubrids macrochromosomes have been derived from microchromosomes by the additions of heterochromatin.     

Karyology of the genus Epidendrum (Orchidaceae: Laeliinae) with emphasis on subgenus Amphiglottium and chromosome number variability in Epidendrum secundum

Epidendrum is one of the largest Neotropical genera of Orchidaceae and comprises approximately 1500 species. Only 2.8% of these species have been studied cytologically, demonstrating chromosome numbers ranging from n = 12 in E. fulgens to n = 120 in E. cinnabarinum. The present work evaluated the evolution of the karyotypes of Epidendrum spp. based on data gathered from the literature and from analyses of the karyotypes of 16 Brazilian species (nine previously unpublished). The appearance of one karyotype with n = 12 with one larger chromosome pair in subgenus Amphiglottium appears to have occurred at the beginning of the divergence of this lineage, and x = 12 probably represents the basic number of this subgenus. Epidendrum secundum exhibits wide variation in chromosome numbers, with ten different cytotypes found in 22 Brazilian populations, seven of which were new counts: 2n = 30, 42, 50, 54, 56, 58 and 84. Most lineages of Epidendrum seem to have been secondarily derived from one ancestral stock with x = 20, as is seen in the majority of the present‐day representatives of the genus. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 329–344.     

Cladistical analysis of primitive G-band sequences for the karyotype of the ancestor of the Cricetidae complex of rodents

Homologous segments identified by G-banding sequences of chromosomes of Peromyscus boylii, Neotoma micropus, Oryzomys capito, (Family Cricetidae) Rattus norvegicus, Melomys burtoni, and Apodemus sylvaticus (Family Muridae) were used to hypothesize a chromosomal condition for the cricetid ancestor. A critical assumption in proposing the primitive G-banding sequences for a given chromosome is that if the outgroup and ingroup taxa have a specific sequence, then the ancestor of the ingroup taxa also had that same sequence. Using this methodology, (chromosome numbers refer to proposed homology to the standardized karyotype for Peromyscus), we propose that: (1) the primitive banding pattern of chromosome 1 was identical to that of Neotoma; (2) the primitive patterns of chromosomes 2, 3, 4, 6, 7, 8, 9, 10, 11, and 12 were primitive banding patterns of 5 and 13 were undetermined; (4) a major portion of the banding patterns of 14 and X were present in the ancestral karyotype. Only the largest 14 autosomes and X were examined because the smaller elements had insufficient G-band definition to ensure reasonable accuracy. The karyotype ancestral to that of Peromyscus, Neotoma, and Oryzomys may be as above and the banding patterns of 5, 13, and 14 were acrocentric and identical to those shown for Peromyscus, Neotoma, and Oryzomys (Fig. 1). In the primitive karyotype, heterochromatin (C-band material) was probably limited to the centromeric regions. If the primitive karyotype is as described above, then it is possible to determine the direction, type, and magnitude of chromosomal evolution evident in the various cricetid lineages. Based on the available data, radiation from the ancestral cytotype is characterized by a nonrandom distribution of types of chromosomal changes. Within many genera, more rearrangements occur in the 14 largest autosomal chromosomes of some congeneric species than distinguish the proposed primitive conditions for the genera Peromyscus, Neotoma, and Oryzomys. It would appear that the extensive morphological radiation from the primitive cricetid ancestor as indicated by the presence of over 100 surviving genera within the family, was not accompanied by extensive karyotypic changes. The magnitude of chromosomal variation that accompanies speciation in these genera appears to range from no detectable chromosomal evolution to a radical reorganization of the genome.     

Chromosome and genome size variation in Luzula (Juncaceae), a genus with holocentric chromosomes

The present study examines chromosome and genome size evolution in Luzula (woodrush; Juncaceae), a monocot genus with holocentric chromosomes. Detailed karyotypes and genome size estimates were obtained for seven Luzula spp., and these were combined with additional data from the literature to enable a comprehensive cytological analysis of the genus. So that the direction of karyotype and genome size changes could be determined, the cytological data were superimposed onto a phylogenetic tree based on the trnL‐F and internal transcribed spacer (ITS) DNA regions. Overall, Luzula shows considerable cytological variation both in terms of chromosome number (2n = 6–66) and genome size (15‐fold variation; 2C = 0.56–8.51 pg; 547.7–8322.8 Mb). In addition, there is considerable diversity in the genomic mechanisms responsible, with the range of karyotypes arising via agmatoploidy (chromosome fission), symploidy (chromosome fusion) and/or polyploidy accompanied, in some cases, by the amplification or elimination of DNA. Viewed in an evolutionary framework, no broad trend in karyotype or genome evolution was apparent across the genus; instead, different mechanisms of karyotype evolution appear to be operating in different clades. It is clear that Luzula exhibits considerable genomic flexibility and tolerance to large, genome‐scale changes. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 529–541.     

The so‐called primitive genera of Genisteae (Fabaceae): systematic and phyletic considerations based on karyological data

A karyological analysis of the so‐called primitive genera of Genisteae has shown that they have a relatively homogeneous chromosome complement: all species tend to have a somatic chromosome number 2n = 48, which can increase to 2n = 52, presumably as a result of hyperaneuploidy. Karyological data suggest that Argyrocytisus, Cytisophyllum and Petteria may be considered as distinct genera rather than being assigned to Cytisus, with 2n = 52 for the first of these and 2n = 50 for the other two genera. They may be interpreted as relict monotypic genera as a result of the presence of a stabilized aneuploidy. Karyological characters exclude a recent origin of Genisteae from Thermopsideae. On the contrary, they are consistent with the hypothesis that Genisteae and Thermopsideae are independently derived from a basic papilionoid stock, of which present day Sophoreae are the remainder. At least two lines would lead from Sophoreae to the taxa of the ‘genistoid alliance’, one to Thermopsideae and the other ‘podalyrioid alliances’ (Podalyrieae and Mirbelieae), with the prevailing basic number of x = 9, and the other to Genisteae, with a basic number of x = 12 persisting in some present day genera, including Cytisus s.l. From this lineage, a wide range of secondary basic numbers has been formed, mostly by descending aneuploidy. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 232–248.     

Chromosome studies in Orchidaceae: karyotype divergence in Neotropical genera in subtribe Maxillariinae

Here, we study karyotype divergence in the closely related genera Brasiliorchis, Christensonella and Trigonidium belonging to subtribe Maxillariinae of subfamily Epidendroideae (Orchidaceae). We compare karyotypes in 15 species by (1) measuring 1C genome sizes, (2) mapping the distribution of 4′,6‐diamidino‐2‐phenylindole and chromomycin A3 chromosome bands and (3) localizing 5S and 45S nuclear ribosomal DNA (rDNA) sequences using fluorescent in situ hybridization. Recently, phylogenetic studies have been conducted to resolve species and genera relationships in subtribe Maxillariinae. We used these phylogenetic trees to map the cytogenetic characters in an evolutionary framework. This has enabled a better understanding of the patterns of genomic divergence in the group. Genome sizes range from 1C = 1.85 to 4.1 pg. The largest, B. schunkeana, shows evidence of genome upsizing, probably through the acquisition of tandem repeats that now form large 4′,6‐diamidino‐2‐phenylindole‐positive blocks of heterochromatin. Our cytogenetic data are consistent with a base chromosome number of 2n = 40, although Christensonella is characterized by a dysploid reduction in chromosome number to 2n = 36. The number of 5S and 45S rDNA sites is variable between species, consistent with high rates of karyotype divergence. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 29–39.     

Population differentiation and speciation in the genus Characidium (Characiformes: Crenuchidae): effects of reproductive and chromosomal barriers

Both time and low gene flow are the key factors by which different biological species arise. The divergence process among lineages and the development of pre‐ or postzygotic isolation occur when gene flow events are lacking. The separation among species of the genus Characidium was analysed in relation to the geomorphological mechanisms in river courses, events of captured adjacent upland drainages in south‐eastern Brazil, and sex chromosome differences. The ZZ/ZW sex chromosomes of Characidium vary in size, morphology, degree of heterochromatinization, and presence/absence of ribosomal DNA. The goal of this study was to understand the mechanism of sex chromosome differentiation, its close association with the geological history of cladogenetic events among drainages, and reproductive isolation leading to Characidium speciation. The W‐specific probe from Characidium gomesi generated a highlighted signal on the entire W chromosome of C. gomesi, Characidium heirmostigmata, Characidium pterostictum, and Characidium sp., instead of karyotypes of three Characidium aff. zebra populations, which showed scattered signals. An evolutionary and biogeographic landscape arose by analysis of ribosomal DNA site location and differentiation of the sex chromosomes, which established mechanisms of reproductive isolation leading to meiotic barriers, keeping the biological unit distinct even if the contact among species was restored. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 541–553.     

A phylogenetic study of bird karyotypes

Karyotypes were compared in 48 species, including 6 subspecies, of birds from 12 orders: Casuariiformes, Rheiformes, Sphenisciformes, Pelecaniformes, Ciconiiformes, Anseriformes, Phoenicopteriformes, Gruiformes, Galliformes, Columbiformes, Falconiformes and Strigiformes. — With the exception of the family Accipitridae, all the species studied are characterized by typical bird karyotypes with several pairs of macrochromosomes and a number of microchromosomes, though the boundary between the two is not necessarily sharp. The comparative study of complements revealed that a karyotype with 3 morphologically distinct pairs of chromosomes is frequently encountered in all orders except the Strigiformes. Those 3 pairs, submetacentric nos. 1 and 2, and a subtelocentric or telocentric no. 3, are not only morphologically alike but also have conspicuous homology revealed by the G-banding patterns. Furthermore, G-banding analysis provided evidence for the derivation of the owl karyotype from a typical bird karyotype.—The above cytogenetic features led to the assumption that the 3 pairs of marker chromosomes had been incorporated into an ancestral bird karyotype. It seems probable that those chromosomes have been transmitted without much structural changes from a common ancestor of birds and turtles, since the presence of the same marker chromosomes in the fresh water turtle Geoclemys reevesii is ascertained by G-banding patterns. — A profile of a primitive bird karyotype emerged through the present findings. Hence, it has become possible to elucidate mechanisms involved in certain structural changes of macrochromosomes observed in birds. It was concluded that a major role had been played by centric fission as well as fusion, translocation, and pericentric inversion.     

New chromosome numbers, meiotic behaviour and pollen fertility in American taxa of Lupinus (Leguminosae): contributions to taxonomic and evolutionary studies

Original chromosome determinations are presented for 20 American Lupinus taxa, including, for the first time, unifoliolate species, together with first data on meiotic behaviour and pollen fertility for some South American species. Most of the Brazilian multifoliolate L. lanatus, L. rubriflorus, L. multiflorus, L. paranensis, L. bracteolaris and L. reitzii and unifoliolate L. crotalarioides, L. guaraniticus and L. velutinus accessions analysed presented regular chromosome pairing. Meiotic indexes and estimations of pollen viability were higher than 90% for all species and accessions analysed, reflecting the generally regular meiotic behaviour of these plants. Chromosome numbers were determined for the first time for the eastern South‐American species L. guaraniticus, L. crotalarioides, L. paranensis, L. paraguariensis and L. velutinus (n = 18 or 2n = 36) and for the Andean L. ballianus, L. eanophyllus, L. huaronensis, L. semperflorens, plus another eight taxa (2n = 48) from Peru and Bolivia, and L. bandelierae (2n = 36) from Bolivia. Chromosome numbers were confirmed for L. lanatus, L. rubriflorus (2n = 36), L. bracteolaris (2n = 34) and L. microphyllus (2n = 48). In the three accessions of the North American unifoliolate species, L. cumulicola and L. villosus, a chromosome number (2n = 52) previously unknown among American taxa was found. The results of the study, plus published data, support the suggestions that south‐eastern South American species are a group cytologically differentiated from the Andean as well as from most other American ones, and that the Brazilian and the North American unifoliolate Lupinus had independent origins. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 150 , 229–240.     

Phylogenomics of several deer species revealed by comparative chromosome painting with Chinese muntjac paints

A set of Chinese muntjac (Muntiacus reevesi) chromosome-specific paints has been hybridized onto the metaphases of sika deer (Cervus nippon, CNI, 2n = 66), red deer (Cervus elaphus, CEL, 2n = 62) and tufted deer (Elaphodus cephalophus, ECE, 2n = 47). Thirty-three homologous autosomal segments were detected in genomes of sika deer and red deer, while 31 autosomal homologous segments were delineated in genome of tufted deer. The Chinese muntjac chromosome X probe painted to the whole X chromosome, and the chromosome Y probe gave signals on the Y chromosome as well as distal region of the X chromosome of each species. Our results confirmed that exclusive Robertsonian translocations have contributed to the karyotypic evolution of sika deer and red deer. In addition to Robertsonian translocation, tandem fusions have played a more important role in the karyotypic evolution of tufted deer. Different types of chromosomal rearrangements have led to great differences in the genome organization between cervinae and muntiacinae species. Our analysis testified that six chromosomal fissions in the proposed 2n = 58 ancestral pecoran karyotype led to the formation of 2n = 70 ancestral cervid karyotype and the deer karyotypes is more derived compare with those of bovid species. Combining previous cytogenetic and molecular systematic studies, we analyzed the genome phylogeny for 11 cervid species.     

Molecular phylogenetics of Paphiopedilum (Cypripedioideae; Orchidaceae) based on nuclear ribosomal ITS and plastid sequences

Phylogenetic relationships in the genus Paphiopedilum were studied using nuclear ribosomal internal transcribed spacer (ITS) and plastid sequence data. The results confirm that the genus Paphiopedilum is monophyletic, and the division of the genus into three subgenera Parvisepalum, Brachypetalum and Paphiopedilum is well supported. Four sections of subgenus Paphiopedilum (Pardalopetalum, Cochlopetalum, Paphiopedilum and Barbata) are recovered as in a recent infrageneric treatment, with strong support. Section Coryopedilum is also recovered, with low bootstrap but high posterior probability values for support of monophyly. Relationships in section Barbata remain unresolved, and short branch lengths and the narrow geographical distribution of many species in the section suggest that it possibly underwent rapid radiation. Mapping chromosome and genome size data (including some new genome size measurements) onto the phylogenetic framework shows that there is no clear trend in increase in chromosome number in the genus. However, the diploid chromosome number of 2n = 26 in subgenera Parvisepalum and Brachypetalum suggests that this is the ancestral condition, and higher chromosome numbers in sections Cochlopetalum and Barbata suggest that centric fission has possibly occurred in parallel in these sections. The trend for genome size evolution is also unclear, although species in section Barbata have larger genome sizes than those in other sections. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 176–196.     

The founder effect theory: quantitative variation and mdg-1 mobile element polymorphism in experimental populations of Drosophila melanogaster

The chromosomes of 14 specimens of the genus Reithrodon from three different localities of Argentina and two localities of Uruguay were studied using G-and C-banding techniques. Specimens of Uruguay showed a karyotype of 2n=28 chromosomes having a large metacentric X, and a telocentric Y chromosome. This karyotype is very similar to that recently described in a sample from southern Brazil, differing only in the nature of the Y chromosome, which is metacentric in the Brazilian form. All specimens from Argentina showed a 2n=34 karyotype, differing from the Brazilian karyotype by two centric fusions, an acquisition of chromosome material, and at least one pericentric inversion, and by the telocentric nature of both the X and the Y chromosomes. G-and C-banding suggest that the metacentric gonosomes in the Brazilian form resulted from a double autosomal-X-Y Robertsonian translocation. The Uruguayan cytotype is interpreted as derived from a hypothetical neo-X/Y₁Y₂ ancestral form by the secondary loss of the Y₁ chromosome. The karyotypic differences between the Brazilian-Uruguayan and the Argentinian forms afford evidence of species differentiation. It is proposed to assign the former to Reithrodon typicus, and the later to R. auritus.     

Genome size evolution in Sapindaceae at subfamily level: a case study of independence in relation to karyological and palynological traits

Sapindaceae s.l. is a moderately large family of trees, shrubs and lianas. Current knowledge on genome size and how it varies in this family is scarce. This research aims to characterize the DNA content in 39 species of Sapindaceae, mainly in tribe Paullinieae s.s., by the analysis of the variation in genome size relative to karyotypic and palynological features. Nuclear DNA amount was measured by flow cytometry, and linear regression analyses were conducted to analyse the relationship between genome size variation and various karyotypic and palynological features. Genome size varied nine‐fold among species, ranging from 1C = 0.305 pg (Lophostigma plumosum) to 2.710 pg (Cardiospermum heringeri). The low regression coefficients obtained suggest that genome size mainly varies independently of karyotypic and palynological features. With regard to karyotype evolution, the constant chromosome number but variable genome size in Houssayanthus, Paullinia and Serjania suggest that structural changes mainly caused by changes in the amounts of repetitive DNA are more important than numerical change. In contrast, in Cardiospermum and Urvillea, variation in chromosome number and genome size supports the suggestion that numerical and structural changes are important in the karyotype evolution of these genera. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 589–600.     

Enzyme Electrophoretic Evidence for Diploidy in Japanese Woodwardia japonica (L.f.) Sm. (Blechnaceae, Pteridophyte) with 2n= 68

Abstract Enzyme-electrophoretic variation was examined in sporophytes of Woodwardia japonica (2n= 68) and segregation was examined in gametophytes. Despite the high chromosome number, W. japonica displayed disomic segregation of isoenzyme patterns. This and karyotype analyses indicate that W. japonica is a diploid derived from a base number of either x= 17 or x= 34.