Generic interrelationships of theCardueae (Compositae): A cladistic analysis of morphological data

A cladistic analysis of 45 genera ofCardueae for 75 characters is presented, taking into account characters of bracts, corolla, stamens, styles and cypselas. Forty-six binary characters completeBremer's (1994) work. Some of the characters considered byBremer are criticised. The traditionalCarlininae is a paraphyletic group, the ten genera studied sharing no common derived character. A new classification of theCardueae is proposed and the distribution of the tribe is presented. Several new combinations are proposed. The nomenclature ofCentaurea is discussed and the status of the genusSerratula is analysed.     

Two new species of the genusMenziesia from Japan

Two species ofMenziesia, M. katsumatae andM. yakushimensis, are described as new to science. The former is characterized by having a long axis of the inflorescence with 15–20 bracts while the other species has a short axis with about ten bracts.M. katsumatae is restricted to the Hakusan range extending northward to Mt. Hōdatsuzan in the Noto Peninsula.M. yakushimensis has been considered asM. purpurea Maxim. in Kyūshū, but it is clearly distinguishable from the latter by the color and shape of the corolla and the presence of fasciculate glandular-hairs at the base of the corolla lobes. It is endemic to the higher altitudes of Yaku-shima Island, Kagoshima Prefecture.     

Phylogeny ofRubiaceae-Rubieae inferred from the sequence of a cpDNA intergene region

A phylogenetic analysis of 25 species, representing eight genera of theRubieae tribe (Rubiaceae), has been made using the DNA sequence of the chloroplastatp B-rbc L intergene region. Six tropical genera from other tribes ofRubiaceae have been used as outgroups. Whatever the method of analysis (distance, parsimony or maximum likelihood), five groups are clearly separated and described as informal clades. Their relative relationships are not clearly resolved by the parsimony analysis, resulting in eight equally parsimonious trees, 327 steps long, with a consistency index (CI) of 0.749 (excluding uninformative sites). TheRubieae tribe appears monophyletic from the data available. Some new and partly unexpected phylogenetic relationships are suggested. The genusRubia forms a separate clade and appears to be the relatively advanced sister group of the remaining taxa. TheSherardia clade also includes the generaCrucianella andPhuopsis. Galium sect.Aparinoides appears closely attached to theAsperula sect.Glabella clade. The remaining taxa ofGalium are paraphyletic:Galium sect.Platygalium (in theCruciata clade) is linked to the advanced generaCruciata andValantia; the more apomorphic groups ofGalium form theGalium sect.Galium clade, including the perennial sectionsGalium, Leiogalium, andLeptogalium as well as the annual (and possibly polyphyletic) sect.Kolgyda.     

Theligonum cynocrambe: Developmental morphology of a peculiar rubiaceous herb

The annual Mediterranean herbTheligonum cynocrambe shows a peculiar combination of morphological characters, e.g., switch from decussate to spiral phyllotaxis with 90–100° divergence, combined with a change from interpetiolar to lateral stipules, anemophily, lack of calyx, flowers often dimerous to trimerous, corolla fused in both male and female flowers, male flowers extra-axillary, with 2–19 stamens per flower, female flowers axillary, with inferior uniovulate ovary, basilateral style and perianth, nut-like fruits with elaiosome. In male flowers the androecium emerges as an (uneven) elliptical rim with a central depression. This common girdling primordium is divided up into several stamen primordia. In male flowers with low stamen number the stamen primordia may occupy the corners alternating with the corolla lobes. There are no epipetalous androecial primordia that secondarily divide into stamens. Male flowers occasionally show a hemispherical base that may be interpreted as remnant of the inferior ovary. In female flowers a ring primordium grows into a tube on which the petal lobes arise. The perianth and style become displaced adaxially by uneven growth of the inferior ovary. The ovary is basically bilocular. The lower region of the ovary is provided with a septum that is overtopped and hidden by the single curved ovule.Theligonum is referred to theRubiaceae-Rubioideae, with theAnthospermeae andPaederieae as most closely related tribes.     

The systematic relationships ofAegialitis (Plumbaginaceae) as revealed by pollen morphology

Light and first SEM observations on the pollen ofAegialitis indicate that the two species of the genus are clearly distinguishable from all other genera of the tribeStaticeae to whichAegialitis presently is relegated. Intraspecific exine or aperture dimorphism which appears frequently in this tribe, is not recorded in this genus.A. annulata andA. rotundifolia have distinct monomorphic pollen and show a great resemblance in exine features with the tribePlumbagineae, particularly species ofPlumbago. Considering these and other features, separate subtribal status forAegialitis within thePlumbagineae is proposed.     

Molecular phylogenetic analysis ofChrysosplenium (Saxifragaceae) in Japan

The phylogeny of Japanese species ofChrysosplenium (Saxifragaceae) was examined using variation in DNA sequences. Sequences ofrbcL andmatK genes were compared for their feasibility for reconstructing the phylogeny ofChrysosplenium, and thematK sequences was found to give greater resolution. All but one of the 17 Japanese species have been examined formatK gene sequences and phylogenetic analysis of these data resulted in eight most parsimonious trees of 390 steps and a consistency index (Cl) of 0.823. The molecular phylogeny obtained was generally in agreement with Hara's (1957) classification based upon phenotypic similarity, although a conclusion needs extensive examination of the genus on a world-wide level. Using the phylogenetic data, character evolution was examined, especially in the characters traditionally used for grouping infrageneric taxa. Differentiation of opposite and alternate phyllotaxis appears to have occurred only once in the course of evolution ofChrysosplenium.     

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.     

The phylogeny ofActaea (Ranunculaceae): A biogeographical approach

Phylogenetic analyses of the genusActaea were performed using morphological, ecological and biogeographical characters. Using solely morphological characters, the relationships of the three identified species-groups remain uncertain. Close biogeographical examination and comparison of the areas with ecological peculiarities as well as climate data gave important insight into the phylogeny ofActaea and the whole tribe. Consequently, the obtained biogeographical data were used for phylogenetic reconstructions. Both, from the point of view of morphological and biogeographical data,A. pachypoda andA. asiatica are the most ancestral species. They grow on the east sides of the continents, mainly in broad-leaved forests. In West Eurasia the apomorphicA. spicata andA. acuminata occur under similar climatic and ecological conditions, but these species are adapted to another climate rhythm. The most advanced species (A. erythrocarpa, A. rubra) are to be found in the boreal forests where they are widely distributed. This biogeographical approach revealed that the evolution of the species led to a gradual widening and shifting of their ecological constitutions.     

Androecial development and systematics inFlacourtiaceae s. l.

Androecial development of 13 species belonging to six tribes ofFlacourtiaceae has been investigated. While inScolopieae andFlacourtieae the stamens develop centrifugally, inErythrospermeae, Oncobeae andPangieae they are initiated in a centripetal sequence or a sequence that is neither distinctly centripetal nor centrifugal. The distribution of these developmental patterns coincides with the distribution of other characters (e.g. cyanogenic compounds, salicoid leaf teeth) and therefore supports a split of the family intoFlacourtiaceae s. str. (containing theScolopieae, Homalieae, Prockieae, Flacourtieae, Casearieae andBembicieae) andKiggelariaceae (withErythrospermeae, Oncobeae andPangieae) and is in accordance with results of recentrbcL studies.     

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.     

Floral evolution and phylogeny in the tribeThelymitreae (Orchidaceae: Neottioideae)

The Australian orchid tribeThelymitreae, composed ofCalochilus, Epiblema, andThelymitra, is unique in theOrchidaceae because of the presence of a mitra or staminodal complex. Evidence from floral structure suggests thatEpiblema andThelymitra are sister genera and thatCalochilus is derived from aThelymitra ancestor. A Gene Pool Vortex model and a hypothetical phylogeny illustrate that introgressive hybridization, allopatry, and long distance dispersal have played a major role in the evolution of the tribe. Pollination and hybridization in the tribe are discussed with major emphasis on floral mimicry.     

Highly repeated DNA analysis and systematics of the Lemuridae,a family of Malagasy prosimians

Hybridization of highly repeated DNA sequences ofEulemur fulvus mayottensis, Lemur catta, andVarecia has been performed on blots of different species of Lemuridae (L. catta, Hapalemur griseus, Varecia variegata variegata, V. v. rubra, E. macaco macaco, E. coronatus, E. mongoz, andE. rubriventer). The probe ofE. fulvus only hybridized with the differentEulemur species, whereas that ofVarecia hybridized with the two subspecies ofVarecia and that ofL. catta with bothL. catta andHapalemur. These results were used to confirm the classification ofVarecia in a separate genus and to review the separation of theL. catta/Hapalemur group from the other species ofEulemur. Comparison of the migration patterns from DNA fragments of these different species has been used to propose a cladogram of the differentEulemur species.     

Hybridization,transgressive segregation and evolution of new genetic systems in<Emphasis Type="Italic">Drosophila</Emphasis>

Introgressive hybridization facilitates incorporation of genes from one species into the gene pool of another. Studies on long-term effects of introgressive hybridization in animal systems are sparse.Drosophila nasuta (2n = 8) andD. albomicans (2n = 6)—a pair of allopatric, morphologically almost identical, cross-fertile members of thenasuta subgroup of theimmigrans species group-constitute an excellent system to analyse the impact of hybridization followed by transgressive segregation of parental characters in the hybrid progeny. Hybrid populations ofD. nasuta andD. albomicans maintained for over 500 generations in the laboratory constitute new recombinant hybrid genomes, here termed cytoraces. The impact of hybridization, followed by introgression and transgressive segregation, on chromosomal constitution and karyotypes, some fitness parameters, isozymes, components of mating behaviour and mating preference reveals a complex pattern of interracial divergence among parental species and cytoraces. This assemblage of characters in different combinations in a laboratory hybrid zone allows us to study the emergence of new genetic systems. Here, we summarize results from our ongoing studies comparing these hybrid cytoraces with the parental species, and discuss the implications of these findings for our understanding of the evolution of new genetic systems. This paper is dedicated to the memory of our teacher, Prof. N. B. Krishnamurthy.     

A cladistic analysis of the tribeAstereae (Asteraceae) with notes on their evolution and subtribal classification

TheAstereae were surveyed and the genera arranged in 23 informal groups. The generic groups were used to sample representative genera for a cladistic analysis based on morphological characters. The resulting cladogram was used for discussion of evolution and subtribal classification within the tribe. The lower basic chromosome numbers x = 4, 5, 6, and 8 are interpreted as reductions from a primitive x = 9. The subtribeGrangeinae occupies a phylogenetically basal position as sister group to the rest of the tribe. This may be divided into two large groups, largely corresponding to the homochromousSolidagininae and to the heterochromousAsterinae sensu lato, i.e. including theBellidinae, Hinterhuberinae, Conyzinae, andBaccharidinae. The latter four subtribes are derived within theAsterinae, and hence reduced to synonymy. Several intercontinental relationships indicate that a geographical subdivision of the tribe should be avoided, although in our analysis most of the groups proved to be restricted to one of five major regions.     

Evolution of thedec-1 eggshell locus inDrosophila. I. Restriction site mapping and limited sequence comparison in themelanogaster species subgroup

Summary We have analyzed 18 kb of DNA in and upstream of thedefective chorion-1 (dec-1) locus of the eight known species of themelanogaster species subgroup ofDrosophila. The restriction maps ofD. simulans, D. mauritiana, D. sechellia, D. erecta, andD. orena are shown to have basically the restriction map ofD. melanogaster, whereas the maps ofD. teissieri andD. yakuba were more difficult to align. However, the basic amount of DNA and sequence arrangement appear to have been conserved in these species. A small deletion of varying length (65–200 bp) is found in a repeated sequence of the central transcribed region ofD. melanogaster, D. simulans, andD. erecta. Restriction site mapping indicated that thedec-1 gene is highly conserved in themelanogaster species subgroup. However, sequence comparison revealed that the amount of nucleotide and amino acid substitution in the repeated region is much larger than in the 5 translated region. The 5 flanking region showed noticeable restriction site polymorphisms between species. Based on calculations from the restriction maps a dendrogram was derived that supports earlier published phylogenetic relationships within themelanogaster species subgroup except that theerecta-orena pair is placed closer to themelanogaster complex than toD. teissieri andD. yakuba.     

Phylogenetic analysis ofUlmaceae

A phylogenetic analysis of theUlmaceae, Cannabaceae, Barbeyaceae, andBroussonetia of theMoraceae produced nine equally parsimonious trees with 127 steps. TheUlmoideae (Ulmaceae, sensuGrudzinskaya) are a monophyletic group and distinct from theCeltidoideae. The genusAmpelocera occupies an isolated taxonomic position among the celtidoids. The similarity ofAmpelocera to the fossil celtidoid flowerEoceltis of North America suggests thatAmpelocera posesses an archaic suite of characters, and occupies a primitive position among the celtidoids, theCannabaceae and theMoraceae. The relationships among the other celtidoid taxa,Cannabaceae, andBroussonetia are problematic. TheCannabaceae andBroussonetia of theMoraceae are nested within the celtidoids suggesting that this is a paraphyletic group. The close, but unresolved, relationship of the celtidoids to theMoraceae andCannabaceae observed in this analysis, and the appearance of the celtidoids in the fossil record prior to the ulmoids suggests that the evolutionary relationship of the ulmoids and celtidoids may be more distant than current taxonomic treatments reflect.     

The natural delimitation ofCentaurea (Asteraceae: Cardueae): ITS sequence analysis of theCentaurea jacea group

Taxonomic complexity has hindered partitioning the genusCentaurea into natural subdivisions, even though it has long been recognized as an unwieldy, artificial assemblage. Most of the remaining difficulties center in theCentaurea jacea group, whose taxa share a common advanced type of pollen. Because it comprises half the species of the genusCentaurea, as well as five other disputed genera previously segregated fromCentaurea (Chartolepis, Cheirolepis, Cnicus, Grossheimia andTomanthea), theCentaurea jacea group is a significant taxonomic challenge. Newer molecular approaches are useful for resolving complex relationships because they provide more precise inferences of evolutionary relationships than traditional morphological characters. Sequences of the Internal Transcribed Spacers (ITS) of nuclear ribosomal DNA were analyzed for a comprehensive sample of this group. Results indicated that theCentaurea jacea group is monophyletic and includes the segregated genera, but not two other genera (Oligochaeta andZoegea), whose inclusion in theCentaurea jacea group was doubtful. In addition to pollen morphology, the ITS phylogeny is also supported by karyological evidence and by good correlation with biogeographic distribution of the species. The monophyly of theCentaurea jacea group suggests that a natural delimitation ofCentaurea that minimizes nomenclatural changes is possible, but only if a new type of the genus is designated.     

狮耳草属——中国唇形科植物一新归化属

该文报道了中国唇形科植物一新归化种——荆芥叶狮耳草[Leonotis nepetifolia(L.)R.Br.],该种的发现代表狮耳草属[Leonotis(Pers.)R.Br.]在中国的新归化。狮耳草属的主要特征为轮伞花序腋生,苞片和萼齿顶端针刺状,花冠桔红色或黄色,冠唇上唇较大,下唇较小;其与绣球防风属(Leucas R.Br)较为接近,但后者的萼齿等大,花冠通常白色,花冠筒不超出萼外,冠檐下唇长于上唇,二者区别较为显著。荆芥叶狮耳草为一年生草本。轮伞花序球形,多花密集,其下承以多数密集苞片;苞片细长,向下微弯曲,先端针刺状。花萼管状,先端膨大而略折曲;萼齿针刺状,最上面一枚较大。花冠桔红色,密被桔红色绒毛;花冠筒细长,伸出萼筒之外;冠檐二唇形,上唇较大,下唇较小。荆芥叶狮耳草与同属植物Leonotis leonurus R.Br.最为相似,但前者叶为卵圆形或心形,后者的叶片多为披针形。根据该研究的标本和相关文献资料,该文还提供了狮耳草属和荆芥叶狮耳草的形态描述、地理分布的讨论,以及荆芥叶狮耳草的资源价值、归化途经、繁殖和危害的讨论和野外生态照片。     

Comparison of the Giemsa C-banded and N-banded karyotypes of twoElymus species,E. dentatus andE. glaucescens (Poaceae: Triticeae)

The karyotypes ofElymus dentatus from Kashmir andE. glaucescens from Tierra del Fuego, both carrying genomesS andH, were investigated by C- and N-banding. Both taxa had 2n = 4x = 28. The karyotype ofE. dentatus was symmetrical with large chromosomes. It had 18 metacentric, four submetacentric and six satellited chromosomes. The karyotype ofE. glaucescens resembled that ofE. dentatus, but a satellited chromosome pair was replaced by a morphologically similar, non-satellited pair. The C-banding patterns of both species had from one to five conspicuous and a few inconspicuous bands per chromosome. N-banding differentiated the chromosomes of the constituent genomes by producing bands in theH genome only. TheS genomes of both species were similar with five metacentric and two satellited chromosomes having most conspicuous C-bands at telomeric and distal positions. They resembled theS genome of the genusPseudoroegneria. TheH genomes had four similar metacentric and two submetacentric chromosomes. The seventhH genome chromosome ofE. dentatus was satellited, that ofE. glaucescens nonsatellited, but otherwise morphologically similar. The C-bands were distributed at no preferential positions. TheH genome ofE. dentatus resembles theH genomes of some diploidHordeum taxa.     

Relationships, circumscription, and biogeography of Arcytophyllum (Rubiaceae) based on evidence from cpDNA

A cladistic analysis was performed using nucleotide sequence variation in therps16 intron and thetrmL-F region (plastid DNA).Arcytophyllum belongs in a subclade of the tribe Spermacoceae (s.l.) together with the American species presently classified in the generaHedyotis andHoustonia. This subclade is morphologically characterized by cymbiform seeds.Arcytophyllum is the sister group of all AmericanHedyotis andHoustonia and it is suggeste that these latter would be most conveniently treated as a single genus, the correct name of which would beHoustonia.Arcytophyllum should be circumscribed such that it excludesA. serpyllaceum, which is not a member of theArcytophyllum-Houstonia clade but more closely related toBouvardia. The phylogeny that was reconstructed suggests that the ancestral area of theArcytophyllum-Houstonia clade is the South American tectonic plate.