Serologische Untersuchungen zur Gliederung derBrassicaceae

The serological investigations support the opinion ofJanchen (1942) to combine the generaBunias, Isatis, andSisymbrium in the tribeSisymbrieae; Cheiranthus, Erysimum, andMatthiola in the tribeHesperideae; andBrassica, Crambe, Sinapis, andSuccowia in the tribeBrassiceae. They further underline the central position of theSisymbrieae and the isolated position of theHeliophileae. In accordance withEigner (1973) theBrassiceae are placed closer to theSisymbrieae than inJanchen; the same holds for thePringleeae. No serological justification could be found to uniteArabis andBarbarea in the tribeArabideae, andAlyssum andLunaria in theAlysseae. From the antigen-systems used among the representatives ofJanchen's Lepidieae the generaLepidium andNeslia show remarkable correspondence both toCamelina andThlaspi, but not toCochlearia which appears distant fromCamelina andThlaspi also.

Teil 1/Part 1.     

rbcL sequences support exclusion ofRetzia,Desfontainia, andNicodemia from theGentianales

The taxonomic positions ofRetzia, Desfontainia, andNicodemia have been much discussed, and all three genera have been included inLoganiaceae (Gentianales). We have made a cladistic analysis ofrbcL gene sequences to determine the relationships of these taxa toGentianales. Four newrbcL sequences are presented; i.e., ofRetzia, Desfontainia, Diervilla (Caprifoliaceae), andEuthystachys (Stilbaceae). Our results show thatRetzia, Desfontainia, andNicodemia are not closely related toLoganiaceae or theGentianales. Retzia is most closely related toEuthystachys and is better included inStilbaceae. The positions ofDesfontainia andNicodemia are not settled, butDesfontainia shows affinity for theDipsacales s.l. andNicodemia for theLamiales s.l.     

Relationships in theAcanthaceae and related families as suggested by cladistic analysis ofrbcL nucleotide sequences

A parsimony analysis of DNA sequences of the chloroplast-encoded generbcL from twelve members of theAcanthaceae s.l., including members of the sometimes segregateThunbergioideae andNelsonioideae, and other families in theBignoniales sensuThorne (1992) is presented. The results largely agree with the classification of theAcanthaceae presented byBremekamp (1965) andThorne (1992) and supportNelsonioideae as a sister group to the rest of theAcanthaceae. Thunbergioideae are placed as a sister toAcanthaceae s.str.Acanthus andAphelandra, both representatives ofAcanthoideae, form a sister group toRuellioideae. An analysis of branch support found that many branches throughout theBignoniales are weakly upheld. This points to the need for further studies in the group using more sequences ofrbcL as well as other data. None of the families ofBignoniales as presently circumscribed (includingAcanthaceae s.l.) were strongly supported, although the larger clade containing the families of theBignoniales was robust.     

A molecular systematic study ofCathaya, a relic genus of thePinaceae in China

The systematic position ofCathaya, a relic genus of thePinaceae, was discussed based on therbcL gene sequence. The sequence data were analysed with PAUP and MEGA programs. The great genetic distance value betweenCathaya and any other genus of thePinaceae showed thatCathaya was a distinct and isolated genus. The most parsimonious Fitch tree and neighbor-joining tree showed thatCathaya was distantly related to the clade comprisingAbies, Keteleeria, Pseudolarix andTsuga, and a sister group relationship betweenCathaya andPinus was weakly supported.Pseudotsuga is closely related toLarix. In theAbies-Keteleeria-Pseudolarix-Tsuga clade,Abies has a close relationship toKeteleeria whilePseudolarix is relatively closely related toTsuga.     

Intergeneric hybridization and C-banding patterns inHordelymus (Triticeae,Poaceae)

Crosses ofHordelymus europaeus (2n = 4x = 28) with four genera in theTriticeae were attempted. Adult hybrids were obtained in combinations withHordeum bogdanii (2x),Hordeum depressum (4x), andSecale cereale (2x). The meiotic pairing was very low in the hybrids withH. bogdanii andSecale cereale (0.12 and 0.30 chiasmata/cell, respectively), whereas high pairing (9.90 chiasmata/cell) was found in hybrids withH. depressum due to autosyndetic pairing ofH. depressum chromosomes. The chromosome complement ofHordelymus europaeus comprised 16 metacentrics, 8 submetacentrics, and 4 SAT-chromosomes. The Giemsa C-banding patterns of the chromosomes were characterized by small to minute bands at no preferential positions. It is hypothesized thatHordelymus europaeus may genomically be closest related toTaeniatherum andPsathyrostachys spp.     

Evolution alpiner Populationen vonEuphrasia (Scrophulariaceae): Chromosomenzählungen an diploiden und polyploiden Sippen aus den Ostalpen

Chromosome counts are reported for several E. Alpine taxa ofEuphrasia sect.Euphrasia. First records of diploidy for small-flowered taxa are 2n = 22 forE. inopinata andE. sinuata, related toE. minima (4 x). Aberrant E. AlpineE. hirtella is 2 x, just as the typical W. Alpine populations of this species. Tetraploidy, 2n = 44, has been found inE. pumila, close toE. stricta (also 4 x). The limitation of ploidy levels within sect.Euphrasia to 2 x and 4 x on the chromosome base number x = 11 is confirmed.

     

The delimitation and taxonomic position of the tropical African generaLeptactina andDictyandra (Rubiaceae)

The closely related African generaDictyandra andLeptactina are redelimited. The two genera are distinguished by long tubular corolla-tubes with included anthers (Leptactina) vs. much shorter tubes with exserted anthers (Dictyandra), rather than—as previously thought—presence or absence of multilocellate anthers. The discovery of a new species (Dictyandra congolana from the Congo and W-Zaire) confirms this. It can be concluded, especially from comparative morphological studies of fruits and seeds, thatDictyandra andLeptactina are related toPavetta, Tarenna, Ixora, etc., i.e. the group of genera in theCoffeeae s.l. with terminal inflorescences. It is proposed that the tribePavetteae be revived to accomodate those genera.Presented at the meeting of May 4, 1983 of the Koninklijke Belgische Botanische Vereniging.     

Molecular evidence for hybridization ofIlex x wandoensis (Aquifoliaceae) by RAPD analysis

Based on the presence of intermediate morphological characters, such as serrated leaf margins and flower structures,Ilex x wandoensis was initially described as a putative natural hybrid betweenI. cornuta andI. Integra, and was formally described as a new hybrid species,I. x wandoensis C. F. Mill., and M. Kim. However, using molecular markers generated via random amplified polymorphic DNA (RAPD), we have now discovered hybridization in populations of theI. x wandoensis complex collected from Wando and Jeju Islands, Korea. Marker bands of the putative parent taxa also were found in some populations ofI. x wandoensis, confirming its hybrid origin. Morphological variability within and among those populations was confirmed by model-based clustering methods, using multilocus genotype data. Phenograms generated from RAPD bands indicated that some accessions ofI. x wandoensis clustered with one of the parental species. This implied the occurrence of hybridization and recurring backcrosses of the hybrid to both parents, resulting in various hybrid derivatives because of the segregation and recombination of traits.Ilex x wandoensis was more closely related toI. cornuta than toI. integra suggesting that it backcrossed more with the former than with the latter.     

Weitere Untersuchungen über Proteinkörper in Zellkernen und ihre taxonomische Bedeutung

In 83 species of the familiesMonotropaceae, Apocynaceae, Oleaceae, Scrophulariaceae, Lentibulariaceae, Bignoniaceae, Martyniaceae, Myoporaceae, Verbenaceae, Lamiaceae, Campanulaceae, andLiliaceae, protein bodies in the cell nuclei have been found, in 68 of these species for the first time. On the basis of their structure in accordance with morphological characters the generaBurgsdorfia, Hesiodia, Olisia, andPhlomoides of theLamiaceae are accepted;Lamium is divided intoLamium, Lamiastrum andOrvala; two new combinations are established:Kickxia campyloceras (Rech. fil. &Esfandiari)Speta andEtornotus papilionaceus (Burm. in L.)Speta. Deviating shape or lack of protein bodies corroborate former taxonomic decisions, e.g. the transfer ofMonotropa toMonotropaceae or the separation ofGaleopsis andLadanum; Veronica schmidtiana should not be included inPseudolysimachion. Systematic affinities are discussed primarily withinScrophulariaceae because nuclear protein bodies have been studied already in many species of this family. ForCampanula patula two 2 x populations are reported.

Herrn Professor Dr. L.Geitler zum 80. Geburtstag gewidmet.     

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.     

Interspecific hybridizations among species of theElymus semicostatus andElymus tibeticus groups (Poaceae)

Interspecific hybridizations were made between species of theE. semicostatus group, viz.,E. semicostatus (Nees exSteud.)Meld.,E. validus (Meld.)B. Salomon,E. abolinii (Drob.)Tzvel., andE. fedtschenkoi Tzvel., and species of theE. tibeticus group, viz.,E. pendulinus (Nevski)Tzvel.,E. tibeticus (Meld.)Singh,E. shandongensis B. Salomon, andE. gmelinii (Ledeb.)Tzvel., as well as among species within theE. tibeticus group. All species are tetraploid (2n = 4x = 28) and possess SY genomes. Meiotic pairing data from 24 hybrids involving 17 interspecific combinations are presented. The average number of chiasmata per cell ranged from 17.91 to 26.20 in hybrids within theE. tibeticus group, compared with 7.26 to 22.04 in hybrids between the two species groups. Despite the extensive collection of cytological data, there was no definite evidence for confirming or disproving the separate existence of the two groups.     

Ultrastructure of nuclear inclusions and the separation ofVerbenaceae andOleaceae (incl.Nyctanthes)

TEM observations were carried out on 40 taxa of the familyVerbenaceae and 35 taxa of the familyOleaceae, in order to ascertain distribution, ultrastructure and morphology of the intranuclear proteinic inclusions in the mesophyll parenchymatic cells. The investigated genera amount to some 25% and 60% respectively of the genera of the two families. Inside theVerbenaceae, lamellar inclusions (L-type) occur in 6 out of 23 investigated genera: they are mostly present inside the tribesCitharexyleae andVerbeneae (both belonging toVerbenoideae), while they are absent in other subfamilies. All of the investigatedOleaceae genera show intranuclear crystalline inclusions (C 1-type) of three different shapes. Among theAsteridae this is a character peculiar toOleaceae. They appear to be a well defined natural group, including the controversial genusNyctanthes.     

Craniometrical variations among eastern Brazilian marmosets and their systematic relationships

The craniometrical variations among marmosets on the eastern coast of Brazil or theCallithrix jacchus group (C. jacchus, C. penicillata, C. kuhli, C. geoffroyi, C. flaviceps, andC. aurita) were analyzed to test various hypotheses for the arrangement of the species and phylogenetic relationships ofC. kuhli andC. flaviceps. Q-mode correlation coefficients and principal component analysis were employed to extract important shape factors from the craniometrical data. On this basis, the shape distances among the eastern Brazilian marmosets were found to be larger than those between the marmosets which are recognized as distinct species. This evidence supports the hypothesis that the members of theC. jacchus group are distinct species.C. kuhli shows the strongest resemblance toC. geoffroyi within theC. jacchus group.C. flaviceps is the most similar toC. aurita in its cranial shape. These findings indicate thatC. kuhli is most closely related toC. geoffroyi, whileC. flaviceps is toC. aurita.     

Systematic relationships of theLactoridaceae,an endemic family of the Juan Fernandez Islands,Chile

The systematic relationships ofLactoridaceae are problematical, with alternative assignments toMagnoliales, Laurales, andPiperales. Phenetic analyses suggest thatLactoridaceae are best accomodated withinMagnoliales and are most closely related toAnnonaceae. Cladistic analyses indicate that the family is a relatively derived member of theMagnoliales, with affinities toAnnonaceae, Eupomatiaceae, Himantandraceae, andMyristicaceae. These analyses together with fossil pollen data are used to interpret the phylogeny ofLactoridaceae and its relatives.     

Phylogenetic relationships in theCampanulales based onrbcL sequences

Phylogenetic relationships within the angiosperm orderCampanulales were investigated by comparative sequencing of the chloroplast generbcL. CompleterbcL sequences were obtained for ten species in six families within the order. These data were analyzed along with previously publishedrbcL sequences from other taxa (for a total of 117 species) within the subclassAsteridae and outgroups, producing 32 equally parsimonious trees. A subset consisting of 44 of these taxa was then chosen and more rigorous analyses performed, resulting in four equally parsimonious trees. Results indicate that two major clades roughly corresponding to traditionally circumscribedAsterales andCampanulales exist as sister taxa. In particular, therbcL trees indicate thatSphenoclea is not a member ofCampanulales orAsterales, thatPentaphragma is more closely allied toAsterales thanCampanulales, that theCyphiaceae are not monophyletic, thatCampanulaceae andLobeliaceae are not sister taxa, and thatStylidiaceae are correctly placed withinCampanulales.     

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.     

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.

     

A phylogenetic analysis of chloroplast DNA restriction site variation inPoaceae subfam.Pooideae

A phylogenetic analysis was conducted on chloroplast DNA restriction site variation in 34 genera of grasses (familyPoaceae), including 28 genera from subfam.Pooideae (representing tribesAveneae, Brachypodieae, Bromeae, Meliceae, Poeae, Stipeae, andTriticeae) and representatives of three other subfamilies,Arundinoideae, Oryzoideae, andPanicoideae. Analyses of all 34 genera always distinguishedPooideae as monophyletic, regardless of which nonpooid genus functioned as outgroup; six separate analyses of all 28 pooid genera, each including one of the six nonpooid genera as outgroup, resolved five identically-constituted clades withinPooideae (in four cases), or (in the other two cases) yielded results that were less well resolved, but not in conflict with those of the other four analyses. The four best-resolved analyses distinguishedMeliceae as the earliest diverging lineage withinPooideae, andStipeae as the next. Above the point of divergence ofStipeae is a dichotomy between supertribeTriticodae (including tribesBrachypodieae, Bromeae, andTriticeae), and a clade comprisingPoeae andAveneae. The analysis supports some tribal realignments, specifically the assignment ofBriza, Chascolytrum, Microbriza, andTorreyochloa toAveneae, andArctagrostis, Catabrosa, andSesleria toPoeae. The analysis also suggests that the pooid spikelet (i.e., glumes shorter than lemmas and florets two or more) is plesiomorphic inPooideae, and that spikelets with one floret, and those with glumes longer than the first lemma, each have evolved more than once withinPooideae. Results also indicate that small chromosomes and chromosome numbers based on x=c. 10–12 are plesiomorphic withinPooideae. Alternative states of these characters (chromosomes large, chromosome numbers based on x=7) are interpreted as synapomorphies or parallelisms of clades that includeTriticodae, Aveneae, andPoeae. Lanceolate lodicule shape may be a synapomorphy of the clade that includesStipeae, Triticodae, Aveneae, andPoeae, and loss of lodicule vascularization a synapomorphy of the entirePooideae.     

TheUlmaceae, one family or two? Evidence from chloroplast DNA restriction site mapping

The Ulmaceae is usually split into two subgroups, referred to as either tribes or more commonly subfamilies (Ulmoideae andCeltidoideae). The two groups are separated, with some exceptions, on the basis of leaf venation, fruit type, seed morphology, wood anatomy, palynology, chemistry, and chromosome number. Propositions to separate the two groups as distinct families have never gained general acceptance. Recent morphological and anatomical data have suggested, however, that not only is family status warranted but thatCeltidaceae are more closely related toMoraceae and otherUrticales than toUlmaceae. In order to test these alternative sets of relationships, restriction site mapping of the entire cpDNA was done with nine rare cutting enzymes using 11 genera ofUlmaceae s. l., three other families of theUrticales, and an outgroup family from theHamamelidae. Cladistic analysis of the data indicates thatUlmaceae s. l. is not monophyletic and that distinct families (Ulmaceae andCeltidaceae) are warranted; thatUlmaceae is the sister group toCeltidaceae plus all other families in the order; and thatCannabaceae might be nested withinCeltidaceae. Familial placements of various problematic genera (e.g.Ampelocera, Aphananthe) are resolved and character evolution of key morphological, anatomical, chemical, and chromosomal features are discussed.     

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.