Biosystematic studies on the genusIsoetes in Japan I. Variations of the somatic chromosome numbers

The somatic chromosome number of three Japanese species ofIsoetes, I. asiatica, I. japonica andI. sinensis, was determined in 199 individuals from 49 populations. The chromosome number ofI. asiatica was 2n=22, confirming previous reports. However,I. japonica andI. sinensis displayed a diversity in chromosome number. Six cytotypes, 2n=66, 67, 77, 87, 88 and 89, were found inI. japonica; 2n=67, 87, 88 and 89 are new counts in the genusIsoetes. The plants with 2n=66 were the most frequent (72% of total individuals examined) and were distributed throughout Honshu and Shikoku. The plants with 2n=88 occurred in western Honshu and a limited region in northeastern Honshu where the plants with 2n=77 were also found. In contrast, four cytotypes, 2n=44, 65, 66 and 68, were found inI. sinensis. The chromosome numbers ofI. sinensis were reported here for the first time. The plants with 2n=44 occurred only in Kyushu, while the plants with 2n=66 were found throughout a large area of western Japan.     

Caryological analysis of South American species of Vernonia (Vernonieae,Asteraceae)

Abstract

In the present study 16 populations belonging to 13 species of the genus Vernonia Schreb. were examined cytologically. In total, six different chromosome numbers, which represent three basic numbers: x = 10, x = 16 and x = 17, were found. These results include the first chromosome number reports for the following four species: V. lanifera Cristóbal & Dematt. (2n = 2x = 32), V. membranacea Gardner (2n = 2x = 34), V. salzmannii DC. (2n = 2x = 20) and V. scabrifoliata Hieron. (2n = 2x = 128). Besides, a new chromosome number was found in V. saltensis Hieron. (2n = 2x = 32), for which only tetraploid populations have been previously recorded. The data obtained in this work, along with the information available from the literature, show that the genus Vernonia in South America is heterogeneous with basic chromosome numbers that range between x = 9 and x = 19. These numbers suggest that a combination of polyploidy and aneuploidy has played an important role in the evolution of the genus.     

Karyotypic analysis of <Emphasis Type="Italic">Skimmia japonica</Emphasis> (Rutaceae) and related species

A karyotypic analysis of three species of Skimmia (Rutaceae) in East Asia was performed that examined 88 individuals from 53 localities. Chromosome numbers of S. japonica, S. reevesiana and S. arisanensis were 2n=30, 31, 32 (=30+0−2B), 2n=60 and 2n=60, respectively. The chromosome number of S. arisanensis was reported for the first time. All species had a large chromosome pair or quartet (the first pair or quartet) with a median–submedian centromere in the karyotype. In S. japonica the arm ratio of this first pair was considerably variable and showed a geographical pattern. In the northern half of the distribution range, Sakhalin, Hokkaido, Honshu, Shikoku and part of Kyushu, the arm ratio was 1–1.2, while in the southern half, part of Kyushu, Ryukyu and Taiwan, the arm ratio was very variable and ranged from 1.2 to 2.4. In S. japonica the first pair was sometimes rather heteromorphic; however, the heteromorphism was not related to sex of the plant.     

Cytogeography and meiotic chromosome configurations of six intraspecific aneuploids ofCarex conica Boott (Cyperaceae) in Japan

Chromosome numbers were determined for 340 plants ofCarex conica from 83 populations in Japan. Six aneuploids, 2n=32, 33, 34, 36, 37 and 38, were found. Plants with even diploid chromosome numbers 2n=32, 34, and 36 were the most common and had different geographical distributions. Individuals with 2n=32 were from islands in the Seto Inland Sea and nearby coastal areas of the Chugoku District of Honshu; those with 2n=34 were from the Kanto, Chubu and Kinki Districts of Honshu; those with 2n=36 were from the mountainous areas of Chugoku, Shikoku and Kyushu Districts. Canonical discriminant analysis of 17 morphological characters demonstrated that the plants with 2n=32 were clearly distinct from those with 2n=34 or 36. All four aneuploids with even chromosome numbers showed normal bivalent pairing at meiotic metaphase I and probably represent cytogenetically stable cytodemes. Plants with 2n=33 had one heteromorphic trivalent and 15 bivalents, indicating a structural mutation. At mitotic metaphase I, one chromosome was markedly larger than the others, suggesting that the 2n=33 plants arose from 2n=34 plants by fusion of two chromsomes. The plant with 2n=37 was intermediate in morphology betweenCarex conica (2n=36) andC. morrowii (2n=38) and probably originated as an interspecific hybrid between these species.     

CHROMOSOME NUMBERS IN CUPHEA (LYTHRACEAE): NEW COUNTS AND A SUMMARY

The American genus Cuphea with ca. 260 species is extremely diverse with respect to chromosome number. Counts are now available for 78 species and/or varieties, or 29% of the genus. Included in this study are first reports for 15 taxa from Brazil, Cuba, Dominican Republic, Mexico, and Venezuela. Twenty-two different numbers are known for the genus, ranging from n = 6 to n = 54. The most common number in the primary center of species diversity in Brazil is n = 8, which is regarded as the base number of the genus. Two numbers are most common in the secondary center in Mexico, n = 10 and n = 12. Species with n = 14 or higher are considered to be of polyploid origin. Polyploids comprise 46% of the total species counted and appear in 9 of the 11 sections for which chromosome numbers have been reported. Aneuploid species comprise ca. 25% of the genus and are known from 7 of the 11 sections. The two subgenera are not characterized by different chromosome numbers or sequences of numbers. None of the 14 sections are circumscribed by a single chromosome number. Morphological and ecological variability in widespread, weedy species is correlated with differing chromosome numbers in some species whereas in others the chromosome number is stable. Summary of chromosome numbers by taxonomic section is presented. Section Euandra, centered in eastern Brazil, and the largest section of the genus, appears to be chromosomally most diverse. In section Trispermum, characterized by difficult, variable species with intermediate forms, two of the four species studied have polyploid races. Section Heterodon, endemic to Mexico and Central America and comprising most of the annual species of the genus, is best known chromosomally. Chromosome numbers have been counted for 25 of 28 species, and 12 different numbers are reported. The most advanced sections, Melvilla and Diploptychia, with numerous species occurring at higher altitudes, are characterized by high polyploids. Apomictic species occur in sect. Diploptycia. The cytoevolution of Cuphea is complex with frequent polyploid and aneuploid events apparently playing a significant role in speciation in both centers of diversity.     

Chromosome numbers of miscellaneous Malagasy Acanthaceae

Meiotic chromosome numbers are reported for 12 species in eight genera of Acanthaceae from Madagascar. Chromosome numbers of 11 species are reported for the first time. Counts inMendoncia (n=19) andNeuracanthus (n=20) are the first for these genera. A new chromosome number (n=30) is reported inJusticia. Systematic implications of the chromosome counts are addressed and basic chromosome numbers for these eight genera of Malagasy Acanthaceae are discussed.     

Comparison of genomes of eight species of sections <Emphasis Type="Italic">Linum</Emphasis> and <Emphasis Type="Italic">Adenolinum</Emphasis> from the genus<Emphasis Type="Italic"> Linum</Emphasis> based on chromosome banding,molecular markers and RAPD analysis

Karyotypes of species sects. Linum and Adenolinum have been studied using C/DAPI-banding, Ag-NOR staining, FISH with 5S and 26S rDNA and RAPD analysis. C/DAPI-banding patterns enabled identification of all homologous chromosome pairs in the studied karyotypes. The revealed high similarity between species L. grandiflorum (2n = 16) and L. decumbens by chromosome and molecular markers proved their close genome relationship and identified the chromosome number in L. decumbens as 2n = 16. The similarity found for C/DAPI-banding patterns between species with the same chromosome numbers corresponds with the results obtained by RAPD-analysis, showing clusterization of 16-, 18- and 30-chromosome species into three separate groups. 5S rDNA and 26S rDNA were co-localized in NOR-chromosome 1 in the genomes of all species investigated. In 30-chromosome species, there were three separate 5S rDNA sites in chromosomes 3, 8 and 13. In 16-chromosome species, a separate 5S rDNA site was also located in chromosome 3, whereas in 18-chromosome species it was found in the long arm of NOR-chromosome 1. Thus, the difference in localization of rDNA sites in species with 2n = 16, 2n = 30 and 2n = 18 confirms taxonomists opinion, who attributed these species to different sects. Linum and Adenolinum, respectively. The obtained results suggest that species with 2n = 16, 2n = 18 and 2n = 30 originated from a 16-chromosome ancestor.     

New chromosome counts and evidence of polyploidy in Haageocereus and related genera in tribe Trichocereeae and other tribes of Cactaceae

Chromosome numbers for a total of 54 individuals representing 13 genera and 40 species of Cactaceae, mostly in tribe Trichocereeae, are reported. Five additional taxa examined belong to subfamily Opuntioideae and other tribes of Cactoideae (Browningieae, Pachycereeae, Notocacteae, and Cereeae). Among Trichocereeae, counts for 35 taxa in eight genera are reported, with half of these (17 species) for the genus Haageocereus. These are the first chromosome numbers reported for 36 of the 40 taxa examined, as well as the first counts for the genus Haageocereus. Both diploid and polyploid counts were obtained. Twenty nine species were diploid with 2n=2x=22. Polyploid counts were obtained from the genera Espostoa, Cleistocactus, Haageocereus, and Weberbauerocereus; we detected one triploid (2n=3x=33), nine tetraploids (2n=4x=44), one hexaploid (2n=6x=66), and three octoploids (2n=8x=88). In two cases, different counts were recorded for different individuals of the same species (Espostoa lanata, with 2n=22, 44, and 66; and Weberbauerocereus rauhii, with 2n=44 and 88). These are the first reported polyploid counts for Haageocereus, Cleistocactus, and Espostoa. Our counts support the hypothesis that polyploidy and hybridization have played prominent roles in the evolution of Haageocereus, Weberbauerocereus, and other Trichocereeae.     

CHROMOSOMES IN TWENTY-FIVE SPECIES OF CHINESE MEM-BRACIDS (HOMOPTERA: MEMBRACIDAE)*

Abstract Chromosomes in 25 species of Membracidae are recorded. Numbers, size and behaviors of the chromosomes during meiosis and mitosis are used as specific or generic features for the taxonomy of this group. Chromosome numbers vary from n=5 to 12, and sex mechanism are of XO type except two species with neo-XY system. The histogram indicates that 2n=11 with XO sex mechanism is the modal chromosome number, and another type n=10 is also commonly found in this family.     

The chromosomes of Salmo salar

A new chromosome number, 2n=58, is reported for Salmo solar, the Atlantic salmon. The complement comprises 8 pairs of metacentrics, 21 pairs of acrocentrics. The question of variation in chromosome numbers for different salmon stocks is very briefly discussed.     

Cytogenetic variations in eight species of Saussurea DC. (Asteraceae,Cardueae) from Northwest Himalaya

The genus Saussurea comprises many species with highly medicinal, religious or other economical values. The chromosome number in the genus ranges from 2n = 24 to 108, based on x = 12, 13, 16, or 17, but the primary base number is still not clear. The present meiotic study cover 8 species (14 populations) of Saussurea from the Northwest Himalayas, and add new or varied cytotypes, as an attempt to solve the enigmatic cytogenetic variations in the genus. The chromosome numbers in Saussurea auriculata (n = 16) is new to the world, while S. costus (n = 17), S. jacea (n = 17) and S. roylei (n = 17) reveal the varied cytotypes at world level. Besides, S. taraxicifolia (n = 16) is a first ever report for Indian populations. The meiotic studies on different populations reveal a substantial amount of meiotic abnormalities in the form of chromosome stickiness, un-oriented bivalents, cytomixis, laggards, etc. leading to the meiocytes with less (aneuploidy) or more (polyploidy) chromosome numbers. These abnormalities may produce unreduced pollen grains and adversely affect pollen viability. The high number of these meiotic abnormalities may be responsible for the chromosome number variation in the genus.     

Morphological and chromosomal variation in Fallopia section Reynoutria (Polygonaceae) in Korea

We have examined morphological and chromosomal variation inFallopia sect.Reynoutria in Korea to clarify their taxonomic identities and to determine whether their morphological variability is associated with ploidy levels. Principal components analysis (PCA) of individuals from 21 populations, using major distinguishing characters, revealed the presence of four major entiries of sect.Reynoutria in Korea; these includeF. sachalinensis, F. japonica var.japonica, F. forbesii, and the Nonsan population consisting of presumed hybrids. Based on morphology, it is hypothesized that the Nonsan population was probably derived from multiple hybridization events involving the three named taxa. The results also indicate thatF. forbesii is distinct fromF. japonica var.japonica. Polyploidy is more prevalent in sect.Reynoutria than has been previously recognized.Fallopia sachalinensis in Korea occurs as dodecaploids with 2n=132; our count is the first dodecaploid count for the species, and represents the highest chromosome number known in the genus.Fallopia japonica var.japonica occurs as tetraploids (2n=44), hexaploids (2n=66), and octoploids (2n=88), whileF. forbesii occurs as hexaploids (2n=66) and octoploids (2n=88); our counts appear to be the first reported chromosome numbers forF. forbesii. Morphological analysis indicates that there is no apparent correlation between the ploidy levels in these taxa and the morphological characters that we have considered in this study except that the tetraploids ofF. japonica var.japonica tend to have somewhat thicker leaves.     

A study on sex-determination and karyotypic evolution inTetranychidae

The chromosome complements of 45 species of spider mites (Tetranychidae) were studied, making the total number of species now examined in this family 57, approximately 10% of all species known. The chromosome numbers range fromn=2 ton=7. The modal number of the family is 3 (found in 44% of the species). It is argued that the ancestral number isn=2 (21% of the species).In the more primitive subfamily of theBryobiinae both thelytokous and arrhenotokous species occur, whereas the subfamily of theTetranychinae exclusively exhibits arrhenotoky. The karyotype evolution is discussed in connection with arrhenotoky. It is stated that karyotype information is a useful tool for the spider mite taxonomist.This study was financially supported by the Netherlands Foundation for the Advancement of Tropical Research (W.O.T.R.O.) and the Uyttenboogaart-Eliasen Foundation.     

Karyotype and meiosis studies in three South African Pyrgomorpha species (Orthoptera: Pyrgomorphidae)

Two South African Pyrgomorpha species have reduced chromosome numbers, due to centric fusions between the largest autosomes and the medium and small autosomes. P. rugosa has 2n=11(XO) (4 pairs of submetacentric and 1 pair of acrocentric autosomes) and P. granulata has 2n=13(XO) (3 pairs of submetacentric and 3 pairs of acrocentric autosomes). A third South African species has a typical Pyrgomorphidae number of 2n=19(XO) (acrocentrics). The mean chiasma frequency of the 2n=19 species is higher than that of the other two, although the frequencies of distal chiasmata in all three are similar. The recombination potential of the two species with lower chromosome numbers has been reduced, due to fewer crossovers in comparison to the 2n=19 species, as well as to independent assortment.     

Interspecific crosses inHordeum (Poaceae)

A crossing programme including 30 species and 40 cytotypes within the genusHordeum was undertaken. Viable hybrids were obtained in 302 combinations, 15 of which were intraspecific. Differences in seed set and in germination were observed in crosses between different groups of species. Obtaining crosses between different taxonomic groups was generally more difficult when diploid material was used. Some species, e.g.,H. lechleri, H. jubatum, andH. brachyantherum showed a higher crossability than others. The chromosome numbers of the hybrids were usually those expected from the parental numbers but aneuploid series around the expected numbers were rather frequent. Three cases of unreduced gametes were found. Selective chromosome elimination was restricted to combinations including eitherH. vulgare orH. bulbosum.—Despite a very diverse morphology, all South American diploid species together with the two North American diploidsH. intercedens andH. pusillum appear to be closely related. The hexaploid American speciesH. procerum, H. lechleri, andH. arizonicum are also related. The two North American tetraploid speciesH. jubatum andH. brachyantherum sometimes form semifertile hybrids. The Asiatic speciesH. roshevitzii appears to be related to both North and South American taxa.     

Cytotaxonomy of some species of the South American genus Lessingianthus (Asteraceae, Vernonieae)

Mitotic chromosome numbers of 32 populations belonging to 23 species of the genus Lessingianthus H.Rob. (Vernonieae, Asteraceae) were determined. The chromosome number of all examined plants was found to be based on x = 16. The numbers observed varied from 2n = 32 to 2n = 176. The results include the first report of the chromosome number for 11 species: L. lanatus (2n = 32), L. varroniifolius (2n = 32), L. cataractarum (2n = 64), L. intermedius (2n = 64), L. argenteus (2n = 96), L. centauropsideus (2n = 96), L. profusus (2n = 96), Lessingianthus sp. nov. 1 (2n = 96), Lessingianthus sp. nov. 2 (2n = 128), L. robustus (2n = 160), and L. macrocephalus (2n = 176). New chromosome numbers were found in the four other species: L. rubricaulis, L. laniferus, and L. sellowii were tetraploid with 2n = 64, while L. oxyodontus was hexaploid with 2n = 96. B chromosomes were observed in L. coriaceus and L. varroniifolius. Lessingianthus macrocephalus (2n = 11x = 176) is reported as the first case of an odd polyploid and the higher chromosome number of Lessingianthus. The significance of the results is discussed in relation to chromosomal data available for the genus.     

Karyological features and cytotaxonomy of the tribe Vernonieae (Asteraceae)

Chromosome numbers are reported for 29 populations of 19 Vernonieae taxa collected mainly in the northeastern region of Brazil. Among them, data for five genera (Blanchetia, Rolandra, Pithecoseris, Stilpnopappus and Vanillosmopsis) are here reported for the first time, and the first chromosome counts are presented for 12 species. Chromosome numbers are quite diverse among and sometimes within genera, especially in the controversial and large subtribe Vernoniinae. The numbers varied from 2n = 18 to 2n = ~72. The main karyoevolutionary mechanism seems to be dysploidy, while polyploidy is probably associated with ancient hybridization processes generating most paleotetraploid genera. All studied species presented semi-reticulated interphase nuclei and proximal-early condensing behavior in prophase to prometaphase. In one species (Vernonia condensata with 2n = 40) fluorochrome staining with CMA/DAPI revealed five chromosome pairs bearing subterminal CMA⁺/DAPI^? heterochromatin, probably NOR-associated, revealing the existence of low amounts of satellite DNA. The role of these features in the evolution of the tribe is discussed, revealing some interesting aspects for understanding of the Vernonieae karyoevolution, especially regarding neotropical members.     

Geographical distribution of Phagocata vivida in the Far East

Phagocata vivida (Ijima et Kaburaki, 1916) is common in cold-water habitats in mountainous and hilly regions in Japan; in Northern Japan it occurs in lowland areas. Comparative studies of the material from South Korea and Primorskiy in Northeast Siberia, Russia, show that Ph. vivida is distributed widely in these geographic areas. Phagocata miyadii Okugawa, 1939, reported from North Korea and Northeastern China is a synonym of Ph vivida. Geographical distribution of this species in the Japanese Islands now becomes very clear. Judging by its geographical and vertical distributions, the species probably is a preglacial faunal element that entered Japan by the northern route to Old Honshû Island along the coast of the Old Sea of Japan.     

Chromosome multiformity in the genus Ctenomys (Rodentia,Octodontidae)

The karotypes of eleven species of the South American burrowing rodents, genus Ctenomys are described, and information on the somatic number of two other species of the same genus is given. The studied species are: C. torquatus (2n =68), C. tuconax (2n =61), C. minutus (2n=50), C. talarum (2n= 48), C. porteousi (2n=48), C. cf. minutus (2n=48), C. australis (2n=46), C. azarae (2n=48), C. latro (2n=42), C. magellanicus fueguinus (2n=36), C. tucumanus (2n=28), C. opimus luteolus (2n=26), and C. occultus (2n =22). This extreme intrageneric variation in somatic number is also reflected by a great amount of diversity in chromosome structure. Karyotypes seem to be rather constant at the species level. Autosomal polymorphism has been found in two of the species, namely C. talarum and C. latro. The hypothesis of the superimposition of Robertsoman rearrangements, pericentric inversions and translocations in the evolution of the karyotype of Ctenomys is advanced. The direction of chromosome change, either toward increase or decrease in chromosome number, is discussed. It is emphasized that high chromosome multiformity is correlated in Ctenomys with a rapid and explosive pattern of species diversification; the meaning of the small size of populations in enhancing the role of chromosome rearrangements in the evolution of Ctenomys is discussed.