Chromosomal evolution withinPiperaceae

Chromosome numbers for 26 different species of the generaPiper, Peperomia andPothomorphe (Piperaceae) are reported. The basic chromosome numbers are 2n = 26, x = 13 (Piper, Pothomorphe) and 2n = 22, x = 11 (Peperomia), polyploid series are characteristic forPiper andPeperomia. Piper has the smallest chromosomes and prochromosomal interphase nuclei,Peperomia the largest ones and mostly reticulate to euchromatic nuclei.Pothomorphe is intermediate in both characters. The karyomorphological differences betweenPothomorphe andPiper underline their generic separation. Interspecific size variation of chromosomes occurs inPiper andPeperomia. Infraspecific polyploidy was observed inPiper betle. C-banding reveals different patterns of heterochromatin (hc) distribution between the genera investigated. The genome evolution is discussed.     

Karyology and systematics of the genusAmbavia and otherAnnonaceae from Madagascar

New chromosome counts and nuclei structures are reported forAnnonaceae species from Madagascar:Ambavia (2n = 14),Xylopia (2n = 16) andPolyalthia (2n = 18). This first generic count ofAmbavia and its nucleotypic parameters underline its long term isolation from the African continent and it should be regarded as an extremely distinct member of the basic stock of AfricanAnnonaceae. Some karyological similarities are found with the African generaCleistopholis (2n = 14) andUvariopsis (2n = 16) and the AmericanTetrameranthus (2n = 14, 28). The karyology ofXylopia is completely in line with previous results from the Palaeo- and Neotropics.Polyalthia has 2n = 18, and x = 9, probably the only base number within the whole genus. Fluorochrome and Giemsa-C-band patterns are identical with different congeneric species.Dedicated to Prof. DrElisabeth Tschermak-Woess on the occasion of her 70th birthday.     

Allozyme diversity and systematics inAnnonaceae—a pilot project

Five species ofAnnona and one species fromArtabotrys, Cananga, Polyalthia, andRollinia were investigated in regard to 11 different allozyme loci. Preliminary studies on small population samples ofAnnona suggest genetic uniformity in three species and variability within and between populations in two other species. The allotetraploid origin ofA. glabra is clearly shown by its hybrid enzyme bands. The genetic distance between fiveAnnona species partly corresponds with their morphological relationships; onlyA. muricata appears more separated than is suggested by morphology. A comparison of the five genera demonstrates close relationship betweenAnnona andRollinia. Two enzyme loci are identical within all taxa investigated and possibly may serve as a genetic marker for the family.     

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.     

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.     

Karyological Differentiation in Sapindaceae with Special Reference to Serjania and Cardiospermum

New chromosome counts for 9 species and 2 genera of Sapindaceae are presented and compared with a review of all available chromosome numbers of the family. In 4 species diploid numbers differing from previous reports are found. In 4 species of the tribe Paullinieae (S. diversiflora, S. subdentata, C. grandiflorum and C. halicacabum) detailed studies on interphase nucleus structure, condensing behaviour and chromosome banding patterns are presented. The karyological differentiation of Paullinieae is generally characterized by dysploid reduction of chromosome numbers and the increase of chromosome size. Sequential staining of nuclei with CMA/DAPI and Giemsa-C-banding demonstrates diversification of constitutive heterochromatin (= hc) and different types of chromatin organization in Serjania and Cardiospermum. The obvious lack of polyploid series and the karyological evolution within the family is discussed. The outstanding small genome size found in Cardiospermum halicacabum is considered to be due to a secondary loss of DNA in the course of the change to herbaceous growth.     

Systematics and karyoevolution inMagnoliidae:Tetrameranthus as compared with otherAnnonaceae genera of the same chromosome number

First generic chromosome counts reveal the base number x=7 for the generaTetrameranthus andRollinia. T. umbellatus from the Peruvian Amazon is diploid (2n=14),T. duckei from Brazil (Manaus) is tetraploid (2n=28). In the NeotropicsRollinia (7 species counted) has developed diploid to octoploid taxa (2n=14, 28, 42, 56). Counts of 7 South AmericanAnnona species are presented for comparison (2n=14, 28). The West AfricanCleistopholis patens has 2n=14. The Asian genusMezettia: 2n=14 and the neotropicalGuatteria tribe: 2n=28 are also revised. A detailed karyomorphological comparison, including karyotypes, banding patterns, condensing behaviour of chromosomes and structure of interphase nuclei reveals that the closely related generaAnnona andRollinia are almost identical in their diploid genomes, whereas the polyploid ones differ in their heterochromatin (=hc) composition and number of NO-chromosomes.Cleistopholis, Mezettia and theGuatteria tribe are karyologically and systematically distinct from each other and fromAnnona/Rollinia. Tetrameranthus as compared with the karyomorphology of about 60 other Annonaceous genera has a very peculiar and unusual karyomorphology which underlines its isolated position. Nuclear structures are almost identical in the African genusUvariopsis (2n = 16) and partly similar in theGuatteria tribe; both also share some morphological similarities and possibly are related. From a comparison ofTetrameranthus with several nuclear types within theMagnoliidae, a new model of chromosome evolution in primitive Angiosperms is suggested. In respect to their eco-morphological differentiation the genera investigated differ strongly from each other.Dedicated to Prof. Dr. K.-H.Rechinger on the occasion of His 80th birthday.     

Karyosystematics and evolution of AustralianAnnonaceae as compared withEupomatiaceae,Himantandraceae, andAustrobaileyaceae

Chromosome counts are presented for 12 genera and 20 species of AustralianAnnonaceae (all diploid with 2n = 16 or 18; Table 1) and two species ofEupomatiaceae (2n = 20, partly from Papua New Guinea). Detailed studies on interphase nuclear structure, condensing behaviour of chromosomes, and fluorochrome and Giemsa C-banding patterns also includeHimantandraceae (Galbulimima) andAustrobaileyaceae. — Eupomatiaceae completely correspond withAnnonaceae karyologically, their base number 2n = 20 is interpreted to have evolved from 2n = 18 by ascending dysploidy from common ancestors.Eupomatia laurina andE. benettii differ in DNA and constitutive heterochromatin (hc) quantity; their evolution from high to low DNA content probably corresponds to general progressions inMagnoliidae. Austrobaileya has nuclei of the presumably primitive Tetrameranthus type which is closely related to that ofGalbulimima and several other primitive taxa inMagnoliidae. Karyomorphology and other characters support the maintainance of two main branches within theMagnoliidae, Laurales andMagnoliales, withAustrobaileya probably intermediate; theWinteraceae appear more remote.—InAnnonaceae the reestablishment ofAncana is underlined by its chromosome number (2n = 18) the unexpected and specialized disulcate pollen, and various morphological characters which point to a close alliance with the Australian endemic generaFitzalania andHaplostichanthus (also disulcate) and the American genus pairSapranthus/Desmopsis; they are united in the provisionalSapranthus tribe, with a more distant position toFissistigma s. str. (2n = 16). AustralianAnnonaceae exhibit a high generic and a low species diversity; they can be considered as an ± old and partly impoverished outpost of the family with phytogeographical relationships to Asia, Africa and America.—On the base of field observations three main types of floral development inAnnonaceae are proposed, the most elaborated one found in the fly pollinated genusPseuduvaria. The growth form change from shrubs to lianas during the ontogeny ofDesmos andMelodorum, the vegetative propagation of anAncana species and the ecological and evolutionary patterns of the taxa investigated are discussed.     

C-banding inLiriodendron tulipifera (Magnoliaceae): Some karyological and systematic implications

InLiriodendron tulipifera the Giemsa C-banding pattern differs remarkably from the distribution of condensed chromatin in prophase as revealed by conventional methods. The nature of proximal bands in metaphase chromosomes is interpreted to be different from the terminal ones. A comparison ofLiriodendron with some C-banded karyotypes of ± related taxa indicates a relatively high degree of karyomorphological differentiation in some woodyPolycarpicae. This has to be considered in karyosystematic comparisons.     

Interspecific differences in differentiation and dedifferentiation patterns in the genusNicotiana

Differentiation on hormoneless media, habituation ability and crown gall induction inNicotiana tissue cultures have been used as physiological parameters of evolutionary differentiation between species. Some of them on hormone free media differentiated whole plantlets, others produced only shoots or roots or showed undifferentiated growth (habituation), some eventually died. Moreover, the same genotypes showed a differential behaviour as far as tumor formation byAgrobacterium tumefaciens was concerned. Particularly, the competence for crown gall transformation inNicotiana species seems negatively correlated with differentiation capacity and may be ascribed to differences in the plants capacity to synthesize growth regulators. The correlation between the results obtained and the phylogenetic position of the genotypes tested is finally discussed.     

Chromosome numbers and differentiation of centrospermous families

The patterns of variation of chromosome numbers in theCentrospermae suggest a common ancestry of centrospermous anthocyanin and betalain families. Phylogenetic divergence in the order may have originated with progenitors similar to extantMolluginaceae, Aizoaceae orPhytolaccaceae taxa with x = 9. Evolutionary radiation and advancement in several lines then seems to have been paralleled by trends for increasing chromosome base numbers through dysploidy and polyploidy, e.g. towardsCaryophyllaceae, Portulacaceae-Basellaceae, Hectorellaceae, Cactaceae, Didieraceae, Nyctaginaceae andChenopodiaceae-Amaranthaceae. Presented in the Symposium Evolution of Centrospermous Families, during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

The cytotaxonomy ofEmilia spp. (Asteraceae: Senecioneae) occurring in Brazil

Analysis of several populations in a large part of the distribution area of the genusEmilia in Brazil has revealed only two species: the diploidE. sonchifolia and the tetraploidE. fosbergii. The more widely reportedE. coccinea was not found. They show a karyotype constancy in morphology and chromosome number (2n = 10 and 2n = 20, respectively), C-banding pattern and number of secondary constrictions. Some indications were found thatE. fosbergii may be an allopolyploid and that its ancestors had different genome sizes.     

The genusBuxus (Buxaceae): Aspects of its differentiation in space and time

Trends of pollen grain aperture evolution and exine characters as well as characters of leaf venation, petiole and axial vascularization are briefly described and related to geographical distribution and classification ofBuxus. A review of fossil records is given. Three major taxonomic groups can be delimitated within the genus, and aspects of their relationships and chorogenesis are presented. The level of differentiation, the pattern of distribution and the fossil record speak in favour of an ancient origin of the genus.     

Gamete composition and chromosome variation in pollen-derived plants from octoploid triticale x common wheat hybrids

Summary Anther culture of secondary octoploid triticale (AABBDDRR) and F₁ hybrids (AABBDDR) of octoploid triticale x common wheat crosses was carried out, and 96 pollen-derived plants were developed and studied cytologically. In addition to the 8 types of pollen-derived plants with the theoretically predicted chromosome number, plants with the chromosome constitutions of 2n = 38, 43, 45, 47, 74, and mixoploids were obtained. The haploids and the diploids had different distributions. The frequencies of plants with one and two (pairs of) rye chromosomes were extremely high, and anther culture may be an expeditious route for creating alien addition lines of distant hybrid F₁s. Chromosome aberrations, including deletions, inversions, translocations, as well as isochromosomes and ring chromosomes, were observed in some plants. Abnormal meioses, such as chromosome non-disjunction, were also found. The reasons for the chromosome aberrations are discussed.     

Heterochromatic banding patterns in the chromosomes ofBrimeura (Liliaceae)

The two species of the genusBrimeura (Liliaceae),B. amethystina andB. fastigiata, show a strikingly disjunct geographical distribution. Both species have a bimodal chromosome complement, 2n = 28, which includes 10 large (L) and 18 small (S) chromosomes. Most of the L-chromosomes possess heterochromatic segments which show enhanced quinacrine fluorescence. The resulting banding patterns are species-specific. Two triploid plants have been found inB. fastigiata.     

A natural hybrid ofMonophyllaea (Gesneriaceae) in the tropical rain forests of West Sumatra

A natural hybrid (2n = 21) between the parapatric rain forest speciesMonophyllaea hirtella (2n = 20) andM. horsfieldii (2n = 22) (Gesneriaceae) has been observed at Sg. Lubuk Paraku, Padang, W. Sumatra. The hybrids showed intermediary characters between the parental species in the inflorescence structure, flower size and colour, indumentum, chromosome numbers, and habitat. The 29% pollen fertility of a single hybrid specimen suggests that the genetical isolation between the parental species may break down and reproduction over subsequent generations may occur.     

Natural abundance of 15N in tropical plants with emphasis on tree legumes

Natural abundance of ¹⁵N ( ¹⁵N) of leaves harvested from tropical plants in Brazil and Thailand was analyzed. The ¹⁵N values of non-N₂-fixing trees in Brazil were +4.5±1.9, which is lower than those of soil nitrogen (+8.0±2.2). In contrast, mimosa and kudzu had very low ¹⁵N values (–1.4+0.5). The ¹⁵N values of Panicum maximum and leguminous trees, except Leucaena leucocephala, were similar to those of non-N₂-fixing trees, suggesting that the contribution of fixed N in these plants is negligible. The ¹⁵N values of non-N₂-fixing trees in Thailand were +4.9±2.0. Leucaena leucocephala, Sesbania grandiflora, Casuarina spp. and Cycas spp. had low ¹⁵N values, close to the value of atmospheric N₂ (0), pointing to a major contribution of N₂ fixation in these plants. Cassia spp. and Tamarindus indica had high ¹⁵N values, which confirms that these species are non-nodulating legumes. The ¹⁵N values of Acacia spp. and Gliricidia sepium and other potentially nodulating tree legumes were, on average, slightly lower than those of non-N₂-fixing trees, indicating a small contribution of N₂ fixation in these legumes.     

How stable are genomes of tropical woody plants? Heterozygosity in C-banded karyotypes ofPorcelia as compared withAnnona (Annonaceae) andDrimys (Winteraceae)

InPorcelia goyazensis (2n = 18) Giemsa C-banding patterns differ from those ofAnnona reticulata (2n = 14) and reveal structural heterozygosity. The amplitude of karyological variation in theAnnonaceae is greater than expected for a primitive woody family. In a comparison with other tropical angiosperm groups, the highly differentiated karyotype ofDrimys brasiliensis (2n = 86) is interpreted as being the end-point of numerous karyological changes.     

Karyologie und ökologisch-morphologische Differenzierung vonPeumus boldus (Monimiaceae,Laurales)

The karyotype analysis ofPeumus boldus reveals a remarkable differentiation of chromosomes in size and form. C-banding pattern and interphase nuclei are discribed and compared with other genera. The great variation in leaf-shape and growth-form is continuous and correlated with the very diverse habitats of the species. The recognition of infraspecific taxa is therefore not recommended. The lower leaf surface is very uniform within the monotypic genus and ± similar with that ofSiparuna. The different evolutionary strategies ofPeumus and related genera are discussed and show a different eco-systematic situation than inMollinedia andSiparuna.

     

C-banding patterns in the AustralianBulbine (Liliaceae): The annual group,B. semibarbata s. lato

Chromosome C-band patterns have been studied in 34 populations of the Australian annualBulbine group, which comprises 4x (2n = 26, 28), 8x (2n = 52, 54) and 12x (2n = 78) populations. The 2n = 26B. semibarbata populations have a simple, low heterochromatin pattern with very minor polytypic variation. The 2n = 28 populations, corresponding morphologically to a group given separate status asB. alata, are similar in pattern but exhibit pronounced enhancement of telomeric and, more particularly, centromeric dot bands. NOR heterochromatin and satellites are difficult to identify inB. alata but appear to occur in different positions from the 26-chromosome karyotype. Eastern Australian 8 x patterns are consistent with a proposed hybrid ancestry,B. semibarbata ×B. alata. Annual and perennial C-band profiles in the AustralianBulbine are discussed briefly in relation to the additive and transformation models of heterochromatin evolution and to the possible adaptive significance of variation in heterochromatin content.Cytoevolution in the AustralianBulbine 2; for part 1 see Pl. Syst. Evol.157, 201–217.