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Meetings of the Royal Botanical Society Of Acta Bot. Neerl. 497-505 41(4), December 1992, p. Meetings of the Royal Botanical Society of The Netherlands MEETING OF THE SECTION FOR PLANT SYSTEMATICS AND GEOGRAPHY ON 24 APRIL 1992 the variation these is Significance of Cytological Characters in cytological among higher taxa not yet clear. Systematic and Evolutionary Studies of Crassulaceae Chloroplast DNA Variationand Henk ’t Hart. Departmentof Plant Ecology and in the Evolutionary Biology, University of Utrecht, Evolutionary Relationships Padualaan 8, 3584 CH Utrecht, The Netherlands Crassulaceae R.C.H.J. van Ham. Department of Plant Ecology Cytologically the family Crassulaceae is probably the and Evolutionary Biology, Padualaan 8, 3584 CH most diverse group of angiosperms. Chromosome Utrecht, The Netherlands numbers range from 2n = 8 to In = 640, and basic numbers from = = and The x 4 to x 37 more. genus Evolutionary relationships within the Crassulaceae Sedum especially is extremely variable. The 54 were investigated at various taxonomic levels. To European species for instance, comprise about 140 investigate characters independentfrom those thus far different cytotypes. used ('t Hart, H. (1991): Flora Mediterranea 1: 31-61) Chromosomes of Crassulaceae are generally very we studied variation in the conservatively evolving less than 2 but variation in chromo- restriction-site small, usually pm, chloroplast genome by comparative some size is nevertheless considerable. Chromosome mapping. As a basis for this study we generated a 10-fold well of length may vary among species as as complete clone bank the chloroplast DNA among chromosomes of a single karyotype. The (cpDNA) of the European Sedum album L. from of DNA nucleus is which clones used in amount per proportional to were as homologous probes chromosome number and chromosome length. species comparisons (van Ham, R.C.H.J. et al. 1992. 2C values low in Biochem. Ecol. 20: Although are extremely many species Syst. 243-253), (0-5 pg and less), 40-fold differences have been CpDNA restriction-site variation in Sedum series observed among species. Rupestria revealed new insights in the evolution of the to Cytological variation in Sedum is due autopoly- seven species of this comparium, i.e. a different basal trans- ploidy, descending dysploidy (Robertsonian branching and a more complicatedreticulate terminal locations), and amphiploidy, and in most cases the branchingpattern includingone taxon (S. montanum direction of the transformations of the of A cytological ssp. montanum) putative triple hybrid origin. characters can be determined unequivocally. Further- preliminary analysis of restriction-site variation at more, variation in 2C values is often correlated with the familial level provided evidence for a basal split in life-form or adaptations to more extreme habitats. In the family between a Crassula-lineage and a Sedum- small variation is Within the infrageneric groups, cytological lineage. Crassula-lineage, the divergence often a powerful tool for reconstructing phylogenies. between Crassula and the disputed genus Tillaea must For instance, in Sedum series Rupestria the phylogeny, have taken place relatively early in the evolutionary based on cytological variation, fully agrees with a history of the family. The evolution of the Sedum- states cladogrambased on 240 different characters of lineage is much more diverse and complicated. The 104 morphologicalcharacters. analyses suggest that (a) the classical subfamilies Among the European infrageneric taxa of Sedum, Kalanchoideae, Cotyledonoideae, Sempervivoideae different levels of chromosomal evolution can be and Echeverioideae are terminal taxa of the Sedum- distinguished. Groups either have a single basic lineage; (b) there is a close, possibly monophyletic, chromosome number or their basic number varies. relationship between the Kalanchoideae and Cotyle- Some groups have low primary basic numbers ranging donoideae; (c) the genus Sedum, is paraphyletic and from x = 5 to x = 13, whereas others comprise possibly comprises three major lineages, each ofwhich primary as well as secondary basic numbers (from has members on more than onecontinent. Finally, a = 6 to = have 14 of characters x x 37), or only secondary (x= or parsimony analysis morphological basic numbers. similar of showed little with the higher) A pattern cyto- congruence cpDNA phylogeny, contributed the logical variation can be observed among genera and indicating that parallelism greatly to subfamilies. However, the evolutionary significance of phenotypic diversification in the Crassulaceae. 497 498 MEETINGS Of the Crassulaceae for the of Rosularia (Crassulaceae) —A Systematic investigated presence alkaloids, Sedum acre has received most attention. Study Some 20 piperidine alkaloids have been reported Urs Eggli. Stadtische Sukkulenten-Sammlung, for this species alone (Hegnauer, R. (1989): Chemo- Mythenquai88, CH-8002 Zurich, Switzerland taxononie der Planzen, 8, Birkhauser, Basel). S. acre has retained combination of charac- Rosularia (De Candolle) Stapf 1923, based on a morphological Umbilicus sect. Rosularia De Candolle 1828, was mono- ters, which are considered to be primitive within the graphed by the author (Eggli, U. (1988): Bradleya 6: Crassulaceae (’t Hart, H. & Koek-Noorman, J. Taxon (Suppl.) 1-120) and counts 36 taxa (26 species (1989): 38:535). Because a similar combination S. including one described after the completion of the of primitive characters occurrs in S. aetnense, monographand 10 infraspecific taxa). anglicum, S. brissemoreti, S. farinosum. S. fusiforme, to in most The genus is closely related Sedum S. lancerottense,S. melanantherum, and S. nudum.we characters and is accordingly placed in subfamily investigated these species for the presence of alkaloids. Sedoideae, at variance with Bergers’ treatment. It A number of pyrrolidine and piperidine alkaloids, differs from Sedum. in that its leaves are always rosu- some of which have not been reported as natural late, its petals are sympetaly (petals united for one- constituents, were detected in the leafy parts. tenth to three-quarters of their length), and its carpels Thedistribution ofthe alkaloids agrees perfectly well remain erect even at fruiting time. The pollen is tri- with the infragenericclassification ofthe Europeanand colporate and may be without ornamentation, or Macaronesian Sedum species (’t Hart, H. (1991): Flora rugulate-striate; the seeds are of the bipapillate type Mediterranea 1:31),except for S. farinosum ofS. series and are longitudinally striate to costate. Cytologi- Macaronesia. S. farinosum should be separated from the falls into dissimilar the other of S. series i.e. S. cally, genus two widely groups, species Macaronesia, 7 either with x= or x = 9. brissemoreti, S. fusiforme. S. lancerottense. and S. The is classified into four sections the base based its alkaloids genus on nudum, on composition, ecological of comparative morphology and cytology. Section preference,polyploidy level, colour ofthe flowers, and Rosularia is widespread in the Near and Middle East its hybridizationpattern. (from Crete to Afghanistan), has x=9, and the only Primitive as well as advanced morphological known which character- characters in the which polyploids are tetraploids are are present acre-group, com- istic for three Turkish subspecies of the widespread prises S. ursi, S. alpestre, S. grisebachii, S. laconicum, R. sempervivum. S. annuum. S. borissovae. S. tuberiferum, S. urvillei, S. Section Sempervivella ( = Setnpervivella as separate apoleipon. S. multiceps. S. sexangulare, S. samium, genus; but also including Sedum hirsutum)) is widely and S. litoreum in addition to S. acre itself. Except for to distributed in the western Mediterranean,North Africa, S. acre, these species were found contain only Ethiopia, and the Hindukush-Himalaya region; its pyrrolidines and 2-monosubstituted piperidines ofthe largely disjunct area is interpreted as being relictic. pelletierine-type. A good concordance was observed The section is cytologically heterogeneous,but at least between the composition of alkaloids and the some taxa also have x = 9. hybridizationpattern. Section is distributed in the In the of alkaloids in the Ornithogalopsis contrast to presence more Inner Himalaya-Hindukush area and in South Asia. primitive Sedum species, pyrrolidine and piperidine in of series This is the least well-known section; the basic chromo- alkaloids were absent species Sedum to be = and all and some number seems x 7, specimens Rupestria, Aeonium, Greenovia, Jovibarba, investigatedweretetraploid. Sempervivum, which all share the same combi- Section Chrysanthae has a relatively narrow distri- nation ofadvanced characters. These results indicate a bution in the mountains of Turkey and immediately correlation between the distribution of alkaloids and adjacent regions of Iran, Iraq and Soviet Armenia. the major evolutionary trends in the European = of alkaloids The base number is x 7, with predominant poly- Crassulaceae. Whether the occurrence to 16 taxa taxa ploidy (up x ) in all investigated; all are may be considered a primitive character in other neoendemics. of Sedoideae well is clear. interpreted as groups as not yet quite Alkaloids of some European Sedoideaeand Systematics of Monanthes (Crassulaceae) Sempervivoideae Reto Nyffeler. Institut fur Systematische Botanik, J.F, Stevens, H. t’ Hart,* H. Hendriks and Th. M. Universitat Zurich, Zollikerstrasse 107, CH-8008 Malingre. Department ofPharmacognosy, University Zurich, Switzerland. ofGroningen.Ant. Deusinglaan2,9713
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