Karyomorphology and relationships in some genera of Saururaceae and piperaceae

Karyomorphological observations were carried out on three genera belonging to the Saururaceae and four genera of the Piperacea. All of the genera of Saururaceae show the same karyomorphological characteristics from interphase to metaphase in the somatic cell divisions. However there are two types of the karyomorphology in Piperaceae, i) the first type observed inPiper, Pothomorphe andZippelia, and ii) the second type inPeperomia. Each group corresponds to Thorne's two subfamilies (1974, 1976), Piperoideae and Peperomioideae. The basic chromosome numbers of the genera are confirmed or newly proposed as follows:Saururus x=11,Houttuynia x=12,Anemopsis x=22 (Saururaceae),Peperomia x=11,Piper andPothomorphe (=Heckeria) x=13,Zippelia x=19 (Piperaceae). The relationships of these basic chromosome numbers are presumed to be as shown schematically in Fig. 4. The original basic chromosome number of the common ancestral stock of Saururaceae and Piperaceae is presumed to be x=11.     

Polyploidy and aneuploidy inOphrys,Orchis, andAnacamptis (Orchidaceae)

Studies on chromosome numbers and karyotypes in Orchid taxa from Apulia (Italy) revealed triploid complements inOphrys tenthredinifera andOrchis italica. InO. tenthredinifera there is no significant difference between the diploid and the triploid karyotypes. The tetraploid cytotype ofAnacamptis pyramidalis forms 36 bivalents during metaphase I in embryo sac mother cells. Aneuploidy was noticed inOphrys bertolonii ×O. tarentina with chromosome numbers n = 19 and 2n = 38. There were diploid (2n = 2x = 36), tetraploid (2n = 4x = 72), hexaploid (2n = 6x = 108) and octoploid (2n = 8x = 144) cells in the ovary wall of the diploid hybridOphrys apulica ×O. bombyliflora. Evolutionary trends inOphrys andOrchis chromosomes are discussed.     

Evolution ofPeperomia (Piperaceae) in the Juan Fernandez Islands,Chile

Four species ofPeperomia (Piperaceae) occur in the Juan Fernandez Islands, Chile:P. berteroana, P. margaritifera, P. skottsbergii, andP. fernandeziana. The last species is found also in continental Chile, whereas the other three are endemic to the archipelago.Peperomia margaritifera is found only on the older island of Masatierra, whereasP. skottsbergii is confined to the younger island of Masafuera, andP. berteroana occurs on both islands. Phenetic analyses of mainland taxa suggest thatP. fernandeziana belongs to subg.Sphaerocarpidium whereas the endemic taxa form their own subg.Tildenidium connecting to subg.Tildenia. Cladistic analyses indicate thatP. margaritifera is the most primitive species in the archipelago and thatP. berteroana is the most derived, especially patristically. Chromosomally, the four species are all n = 22, which may be tetraploid on a base of x = 11. Sulfated flavones occur only inP. berteroana andP. skottsbergii, which are otherwise unknown for the family. Dispersal of propagules to the islands from the continent and between islands is believed to have been accomplished by birds.     

The neotropical angiosperm familiesBrunelliaceae andCaryocaraceae: First karyosystematical data and affinities

Chromosome numbers are polyploid, 2n = 28 inBrunellia comocladiifolia andB. mexicana, and 2n = 46 inCaryocar brasiliense, C. microcarpum andC. villosum. The chromosome are small in both genera, with a length of ca. 1,6-0,4µm. Interphase nuclei correspond to the prochromosomal and the chromocentric type, respectively. This is in conformance with the systematic placement ofBrunelliaceae intoCunoniales, and ofCaryocaraceae intoTheales. Brunellia exhibits affinities to various other orders ofRosidae (andHamamelididae), and is suggested to be primarily apetalous. On a comparative basis, the chromosome numbers found in both families are interpreted as paleopolyploid (4 x and 6 x). This apparently is in correspondence with their rather primitive features, systematic isolation, relatively depauperate status, and evidently great age.     

Karyosystematic studies on threeCrepis species (Asteraceae) endemic to Greece

This work examines the cytogeographical distribution, the morphological characters, and the karyotypes of threeCrepis species endemic to Greece (C. sibthorpiana, C. incana, andC. heldreichiana). C. sibthorpiana is diploid (2n = 2x = 8),C. incana is diploid (2n = 2x = 8) and tetraploid (2n = 4x = 16, 17), andC. heldreichiana is always dekaploid (2n = 10x = 40). The Giemsa positive bands, usually pairs of dots, are mainly centromeric inC. incana, while they are terminal inC. sibthorpiana (on the short arm of all chromosomes) and inC. heldreichiana (on both arms of all chromosomes). Intercalary C-bands are scarce and usually variable within karyotypes, individuals, and species. The most variable karyotype both in Feulgen and Giemsa preparations is that ofC. incana, in which also supernumerary chromosomes were observed, which are polysomic to standard set members. On the basis of morphological and karyological data the evolutionary relationships among the threeCrepis taxa are discussed.     

The karyology of some neotropicalStyracaceae

Chromosome numbers and karyomorphological characters have been investigated inPamphilia andStyrax (Styracaceae). Counted for the first time in the genus, two species ofPamphilia were found to have 2n = 16. The twoStyrax spp. investigated share withPamphilia the same chromosome number, a peculiar condensation behaviour of the chromosomes (Fig. 1a–c) and the same type of semi-reticulate interphase nucleus, results which indicate a close relationship of the two genera. The base number inStyracaceae is probably x = 8 (2n = 2x = 16) with stabilized triploids inHalesia andPterostyrax (2n = 3x = 24). A preliminary comparison withSapotaceae andEbenaceae does not allow a general karyological characterisation of the orderEbenales.     

Comparative karyomorphology and interrelationships ofSelaginella in Japan

Karyomorphological comparisons were made of 16 native and cultivated species ofSelaginella in Japan. The somatic chromosome numbers are 2n=16 inS. boninensis; 2n=18 inS. doederleinii, S. helvetica, S. limbata, S. lutchuensis, S. nipponica, S. selaginoides, S. tama-montana, andS. uncinata; 2n=20 inS. biformis, S. involvens, S. moellendorffii, S. remotifolia, andS. tamariscina; 2n=30 inS. rossii; and 2n=32 inS. heterostachys. The interphase nuclei of all species examined are uniformly assigned to the simple chromocenter type. The metaphase karyotype of 2n=16 (x=8) is 8 m (=median centromeric chromosomes)+8(st+t)(=subterminal and terminal). The group of the species having 2n=18 (x=9) is heterogeneous karyomorphologically: The karyotype ofS. nipponica is 2n=18=6 m+12(st+t),S. tama-montana 10 m+2 sm(=submedian)+6(st+t), andS. uncinata 6 m+7 sm+5(st+t). Although the remaining five species have the common karyotype 8 m+4 sm+6(st+t), the values of mean chromosome length are variable. Another group of the specles having 2n=20 (x=10) is homogeneous, since all species have the same karyotypes 8 m+4 sm+8(st+t) and have similar chromosome size. The karyotype of 2n=30 is 12 m+6 sm+12(st+t) and is suggested to be a triploid of x=10, and 2n=32=16m+16(st+t), a tetraploid of x=8. Thus, three kinds of basic chromosome numbers, x=8, 9, 10 are present in JapaneseSelaginella examined, and their karyomorphological relationships are discussed.     

Variation of Giemsa C-band and fluorochrome banded karyotypes,and relationships inAllium subgen.Molium

The somatic karyotypes of 10 taxa belonging toAllium subgen.Molium (Liliaceae) from the Mediterranean area have been investigated using Giemsa C-band and fluorochrome (Hoechst, Quinacrine) banding techniques. A wide range of banding patterns has been revealed. InAllium moly (2n = 14),A. oreophilum (2n = 16) andA. paradoxum (2n = 16) C-banding is restricted to a region on each side of the nucleolar organisers and the satellites show reduced fluorescence with fluorochromes. The satellites are also C-banded and with reduced fluorescence inA. triquetrum (2n = 18), but two other chromosome pairs also have telomeric bands which are not distinguished by fluorochrome treatment. InA. erdelii (2n = 16) 4 pairs of metacentric chromosomes have telomeric C-bands while 2 pairs of telocentric chromosomes have centromeric C-banding. InA. subhirsutum (2n = 14),A. neapolitanum (2n = 28),A. trifoliatum subsp.hirsutum (2n = 14) andA. trifoliatum subsp.trifoliatum (2n = 21) chromosomes with long centromeres, consisting of a centromere and nucleolar organiser are positively C-banded on each side of the constriction. InA. subhirsutum banding is confined to the pair of chromosomes with this feature, whereas inA. neapolitanum one additional chromosome pair has telomeric bands and inA. trifoliatum there are varying numbers of chromosomes with centromeric and telomeric bands, depending on the subspecies.A. zebdanense (2n = 18) shows no C-bands. The banding patterns in this subgenus are compared with those recorded for otherAllium species and with the sectional divisions in the genus. Evidence from the banding patterns for allopolyploidy inA. trifoliatum subsp.trifoliatum andA. neapolitanum is discussed.     

Karyotaxonomical analysis in the genusAngelica (Umbelliferae)

Morphometric karyotype characters were studied in 25Angelica spp. (Umbelliferae, Apioideae) and in one species of the related genusTommasinia. For three species the chromosome numbers are new. In our study the majority of the species investigated are diploids with 2n = 22, some are tetraploids with 2n = 44 (for these tetraploids also diploid cytotypes are reported in the literature). Among the diploid species,A. miqueliana has a distinct karyotype consisting of submetacentric and acrocentric chromosomes only, the remaining diploids with 2n = 22 as well as tetraploids with 2n = 44 have rather symmetrical karyotypes, consisting of metacentric and submetacentric chromosomes. The very different chromosome number 2n = 28 has been found inA. gmelinii. Its karyotype includes two distinct groups of chromosomes: 8 pairs of rather large metacentrics and submetacentrics and 6 pairs of very short and asymmetrical chromosomes. Chromosome numbers and structures appear to be useful in the taxonomy of some intrageneric taxa inAngelica.     

Cytotaxonomic analysis of the Himalayan species of the genusTorilis (Apiaceae)

Chromosome studies of four HimalayanTorilis species reveal a remarkable interand intraspecific differentiation of chromosome numbers and karyotypes:T. arvensis (2n = 12),T. leptophylla (2n = 12),T. Stocksiana (2n = 36) andT. japonica (2n = 16). Base numbers inTorilis are x = 6, 8, 9 and 11.     

Zur systematischen Stellung der GattungProckia: Karyologie und Epidermisskulptur im Vergleich zuFlacourtia (Flacourtiaceae),Grewia (Tiliaceae) und verwandten Gattungen

Detailed analyses of karyology and leaf morphology do not support relationships betweenFlacourtiaceae andTiliaceae. In spite of different chromosome numbers,Prockia (2n = 18),Flacourtia (2n = 22) andRawsonia (2n = 22) are very similar in karyomorphology, indicating a certain karyological uniformity withinFlacourtiaceae. Lacistema (2n = ca. 62) appears more isolated. On the other hand, theTiliaceae Grewia (2n = 18) andLuhea (2n = 36) have much in common and differ remarkably from the Flacourtiaceous genera. The salicoid leaf-teeth ofProckia are also found inIdesia, but never inTiliaceae. Epidermis ultrastructure reveals certain relationships betweenProckia andFlacourtia in contrast to the strongly differingGrewia. Idesia has a rare und unique epidermis sculpture. — Basic chromosome numbers and chromosomal evolution within theFlacourtiaceae are discussed.

     

Karyotype evolution by centromeric fission inZamia (Cycadales)

The chromosome numbers of several species ofZamia from Mexico are reported.Z. paucijuga, distributed from central Oaxaca to Nayarit, has been found to have 2n = 23, 25, 26, 27 and 28. 2n = 28 is the highest chromosome number yet found in the cycads. Karyotypes of this species differ principally in the number of telocentric and metacentric chromosomes present in each; 2n = 23, 25, 26, 27 and 28 were found to have 5, 3, 2, 1 and 0 metacentric and 8, 12, 14, 16 and 18 telocentric chromosomes, respectively.Z. fischeri has been found to be 2n = 16,Z. furfuracea andZ. loddigesii 2n = 18.Zamia paucijuga on the basis of morphological and ecological characteristics, is considered to be an advanced member of this genus. Chromosome and karyotype evolution inZ. paucijuga may have occurred by centromeric fission of metacentric chromosomes; the karyotypes ofZ. paucijuga are strongly asymmetrical, suggesting that they evolved recently.     

Cytotaxonomical studies on AsianAnnonaceae

Chromosome numbers of 42 species and 3 varieties from 24 genera of theAnnonaceae have been determined (Table 1); reports for 15 of the genera are new. Among Asian genera 2n = 14 occurs only in the specializedMezzettia, while 2n = 16 is wide-spread and also has been found inAnaxagorea with some primitive characters. 2n = 18 is reported for 11 genera, and tetraploidy (2n = 36) has been observed inPolyalthia. Therefore, an original basic number of x = 8 or x = 9 is suggested at least for the Asian genera of theAnnonaceae.—Cytotaxonomical notes on the critical speciesPolyalthia rumphii andP. affinis are given, and the new combinationNeouvaria parallelivenia (Boerl.)Okada & Ueda is proposed.     

A karyomorphological study of the genusHypericum (Hypericaceae) in Japan

Chromosome numbers were determined in 226 collections ofHypericum of Japan, representing nine species and one interspecific hybrid. These included the first cytological records forH. erectum var.caespitosum, H. samaniense, H. hakonense, H. sikokumontanum, H. kamtschaticum var.kamtschaticum, H. kamtschaticum var.hondoense, H. pseudopetiolatum, H. yojiroanum andH. tosaense. Counts of 2n=16 were made throughout for collections of six species, and those of 2n=18 forH. ascyron. Intraspecific polyploidy was found inH. samaniense (2x and 3x, x=8) andH. pseudopetiolatum (2x and 4x, x=8). Results of the karyotype analysis showed that three different karyotypes could be recognized, and they were parallel to the subdivision ofHypericum by Kimura (1951). The chromosomes were very small and mostly median centromeric. It was suggested that the role of polyploidy in the evolutionary differentiation ofHypericum in Japan might have been rather limited.     

Chromosome banding and pairing behaviour inFestuca andVulpia (Poaceae,Pooideae)

Giemsa C-banding patterns of the grassesFestuca rubra (2n = 6x = 42),Vulpia fasciculata (2n = 4x = 28) and their wild F1 hybrid ×Festulpia hubbardii (2n = 5x = 35) show marked differences between the chromosomes of the two parents that enable unequivocal identification ofFestuca andVulpia chromosomes in the hybrid. Moreover, meiotic banding patterns reveal that both homogenetic (Festuca-Festuca andVulpia-Vulpia) and heterogenetic (Festuca-Vulpia) pairing occurs in the F1. The significance of this in relation to the formation of backcrosses to theFestuca parent and to the evolution of theFestuca polyploid complex in general is discussed.     

Cytotaxonomic study of genusHymenasplenium (Aspleniaceae) in Xishuangbanna, Southwestern China

The gametic chromosome numbers of sevenHymenasplenium (Aspleniaceae) species from Xishuangbanna, Yunnan Prov., China, were investigated. All the examined individuals ofH. obscurum, H. cheilosorum andH. latipinnum were sexual diploids with n=39 chromosomes. Intraspecific cytological variation was found inH. excisum, which has a sexual diploid (n=39) and a tetraploid (n=78). Only a triploid apogamous cytotype (n=ca.117) was found inH. laterepens. Hymenasplenium apogamum showed the most complicated intraspecific variation and included a sexual diploid (n=39), a sexual tetraploid (n=78) and an apogamous triploid (n=ca.117). This work reports for the first time the sexual diploids ofH. cheilosorum andH. apogamum, which are only apogamous elsewhere in east Asia, Himalayas and Indochina. These results may indicate that this area is one of the diversity centers ofHymenasplenium. Most of the above species have chromosome numbers based on x=39. In contrast,H. costarisorum contains a sexual diploid (n=36) and a sexual tetraploid (n=72), indicating that its basic number is x=36.     

Cytology and systematics in JapaneseArisaema (Araceae)

Chromosome numbers are determined from 37 populations attributed to 22 taxa of JapaneseArisaema. Of them, chromosome numbers ofA. limbatum var.conspicuum (2n=26),A. minus (2n=26),A. nambae (2n=28) andA. seppikoense (2n=26) are determined for the first time. New chromosome numbers, 2n=26, are reported forA. aequinoctiale, A. limbatum, A. stenophyllum, A. undulatifolium andA. yoshinagae. Three modes of basic chromosome numbers,x=14,x=13 andx=12, occur in JapaneseArisaema. Precise karyotypic comparisons of 20 taxa reveal that taxa withx=14 andx=13 share 26 major chromosome arms and have an obvious chromosomal relationship. One of two submeta-centric chromosomes inx=13 corresponds to two telo-centric chromosomes inx=14. InA. ternatipartitum with 2n=6x=72, ten out of 12 basic chromosomes are the most similar in size and arm ratio with larger ten chromosomes ofA. ringens among JapaneseArisaema examined. A basic chromosome number ofx=14 is the commonest in the genusArisaema and the remaining basic chromosome numbers,x=13 andx=12, seem to be derived through dysploidal reduction by translocating large segments of major arm of telo-centric chromosome onto other minor arm of telo-centric followed by loss of the remainings including a centromere, and by loss of two telo-centrics fromx=14, respectively. Some systematic problems of JapaneseArisaema are discussed based on new cytological data.Arisaema hatizyoense, A. minus andA. nambae are accepted as independent species.     

Pathways of karyological differentiation in palms (Arecaceae)

Karyological data are given for 56 palm taxa coming from all 6 palm subfamilies. In 11 genera and 17 species, chromosome numbers are reported for the first time. Most chromosome numbers in palms range between 2n = 36 and 2n = 26 in dysploid series. Species of the same genus usually exhibit identical chromosome numbers which additionally may be constant in larger groups of closely related genera (Coryphoideae trib.Corypheae with nearly always 2n = 36,Arecoideae subtribesEuterpeinae andRoystoneinae with 2n = 36,Arecoideae subtrib.Butiinae with mostly 2n = 32). Polyploidy among palms is of minor significance but the endemic Madagascan genusVoanioala (2n = 606 ± 3) is the most striking exception. — With respect to structure of interphase nuclei and longitudinal differentiation of prophase and metaphase chromosomes, the palm family is highly differentiated. Euchromatin types with different prophase condensation properties and fluorochrome and C-banding patterns of heterochromatin permit a discrimination of several subfamilies on the nuclear level (Arecoideae, Ceroxyloideae, Nypoideae, Phytelephantoideae, Calamoideae).Arecoideae andCeroxyloideae, andNypoideae andPhytelephantoideae have some features in common. Subfam.Coryphoideae s. l. is a non-uniform group. — Nuclear characters among palms exclusively found in recentCoryphoideae subtrib.Thrinacinae link palms with other monocotyledons. Most probably, such a nuclear condition represents an ancestral state in the evolution of palm genomes within subfam.Coryphoideae s. l., but also the conspicuous nuclear characters of the other modern palm subfamilies appear to be derived from a similar starting point, since transitional character states are still present in subfam.Calamoideae and some taxa of subfam.Arecoideae. Early karyoevolution in palms obviously did not involve numerical change of the ancient chromosome number of 2n = 36 which started subsequently, as a dysploid reduction in numerous parallel series, independent in subfam.Coryphoideae (2n = 36 to 2n = 28),Calamoideae (2n = 36 to 2n = 26),Ceroxyloideae (2n = 34 to 2n = 26), andArecoideae (2n = 36 to 2n = 28). Possible mechanisms of karyological change are discussed. — Karyological characters are compared to morphological, ecological, taxonomical, and chorological features, and give some new insight into older and more recent phases of palm evolution. (1) Strong deviations in vegetative or floral morphology are often accompanied by major karyological differences, and sometimes the direction of advancement can be traced through intermediate stages. (2) Apart fromCoryphoideae subtrib.Thrinacinae, the strongest concentration of apparently original karyological traits is found in the more basal members of each subfamily. (3) The most successful and actively radiating colonizers of the forest floors in evergreen tropical forests which belong to completely different subfamilies (Old WorldLicuala, New WorldChamaedorea andGeonoma), appear to be very advanced karyologically.     

Taxonomic studies ofAsarum sensu lato

The karyotype and the C-banding pattern in two species ofHexastylis andAsarum epigynum were analysed in detail, and the results obtained were compared with those of the other species ofAsarum, Asiasarum andHeterotropa previously reported. The present results were partially different from the previous reports related to the karyotypes of these species. The karyotype observed in two species ofHexastylis (2n=26) was represented by ten pairs of metacentric chromosomes and three pairs of small subtelocentric chromosomes, which is very similar to that ofAsiasarum in eastern Asia. The C-banding patterns ofHexastylis andAsiasarum, however, were clearly different from each other. A striking difference was found in one of the three pairs of small subtelocentric chromosomes. A Formosan speciesAsarum epigynum had the somatic chromosome number 2n=12 and a highly asymmetrical karyotype composed of mainly subtelocentric chromosomes. These karyological features were remarkably different from those of the other groups inAsarum s.l.     

Karyological studies onGentiana sect.Chondrophyllae (Gentianaceae) from China

Nineteen populations of fifteen species ofGentiana sect.Chondrophyllae from China were observed cytologically.Gentiana alsinoides, G. anisostemon, G. asterocalyx, G. exigua, G. heterostemon, G. intricata, G. praticola, G. pseudoaquatica, G. spathulifolia, andG. subintricata all had the same chromosome number of 2n = 20 (or n = 10), whereasG. piasezkii had 2n = 36,G. squarrosa 2n = 38,G. prattii 2n = 18,G. aristata 2n = 14 (n = 7), andG. heleonastes 2n = 12. All these chromosome numbers are documented here for the first time, except forG. squarrosa, where it is a new number report. The basic numbers of x = 6, x = 7 and x = 19 are new for the section. Karyotype analyses of some species revealed that, except for a few cases, the species examined mainly had metacentric chromosomes. 2n = 20 = 2m(SAT) + 18m was found to be the main type of karyotype for the species with 2n = 20. Chromosomal evolution and its mechanism in this section are also discussed.