:. . 00M POSITAE 6 * NEWSLETTER Number 48 4 June 2010 Scientific Editor: BERTIL NORDENSTAM Technical Editor: GUNNEL WirENIUS NOHLIN : Published and distributed by The Swedish Museum of Natural History, Department of Phanerogamic Botany, P.O. Box 50007, SE-104 05 Stockholm, Sweden ISSN 0284-8422 CONTENTS Bremer, K.: Personal reflections on a new Compositae book 1 Funk, V. A.: Ten Things I learned on the way to ve Mother Tree (ite: . Mother Ship) 6 Marry, D. & G. G. Marr: Taxonomic delimitation of the genus Tibetoseris SENNIKOv and the new genus Pseudoyoungia of the _Compositae-Cichorieae from Eastern Himalaya 22 J. Noroozi, Y. Asant & B. Norpenstam: A new annual species of Senecio (Compositae-Senecioneae) from subnival zone of southern Iran with comments on phytogeographical aspects of the area 43 MukuerJee, S. K. & B. Norpenstam: Distribution of calcium oxalate crystals in the cypselar walls in some members of the Compositae and their taxonomic significance 63 Avecsite, A. E.: Pollen studies on some populations of Aspilia (Asteraceae) in Nigeria 89 New taxa and combinations published in this issue 102 ENT ONIA NV . JUL 232010 LIBRARIES. Comp. Newsl. 48, 20010 1 Personal reflections on a new Compositae book KARE BREMER Stockholm University, SE-106 91 Stockholm, Sweden The editor of this newsletter has asked me to write a review of Systematics, Evolution, and Biogeography of Compositae by V. A. FuNK, A. SUSANNA, T. F. Stuessy and R. J. Bayer (eds.) published by IAPT in Vienna 2009. It is an impressive volume and I will not try to comment on all chapters and groups covered by the book. Rather than trying to write a regular review I will here with the editor’s kind permission instead take my own experience with the family as a setting for some personal reflections on FuNnK et al.’s book. Above all, FuNK et al.’s Compositae provides an invaluable reference to the current status of our knowledge of phylogeny above the genus level in the family, together with a new tribal and subfamilial classification. For many other aspects, it is necessary to consult earlier standard works, notably KApDEREIT & JEFFREY 2007 (generic classification), H1np 1996 (many general subjects), and even HEywoop et al. 1977 (e.g. secondary chemistry). My own book on the family (BREMER 1994), with contributions from several colleagues, also focused on phylogeny and classification, and is consequently now more or less outdated, although it may provide an interesting picture of what could be understood — and misunderstood — by cladistic analysis of morphological data compared to the current analyses based mainly on molecular data. The contents of the new book follow a familiar outline with introductory chapters on history and morphology followed by an extensive systematic part dealing mainly with phylogeny, classification, and biogeography. I enjoyed reading the first chapter with biographies of earlier synantherologists. Six introductory chapters cover character evolution in the family. Some of the authors, of chapters on chromosomes, secondary chemistry, and pollen, have taken the opportunity to map character evolution on the family cladogram, illustrating classical questions such as the ancestral chromosome base number for the family (x=9). I was a little disappointed to see that other authors of chapters on character evolution have missed this opportunity. The new more resolved family phylogeny is fascinating to see, especially for myself since I struggled with this problem for many years using morphological data. About 25 years ago I decided to explore tribal interrelationships in the family by cladistic analysis of morphological characters. I had together with ? Comp. Newsl. 48, 2010 my wife received an invitation by PeTer RAVEN to spend a year at the Missouri Botanical Garden and chose to devote my time there to this project. At that time | was heavily influenced by ArTHUR CRONQUIST’S heliocentric view of the family, implying that the Heliantheae s. lat. represent the ancestral complex of the family. After about two months of unsuccessful studies of Heliantheae genera in the Missouri Botanical Garden herbarium in 1985, | realized that I was on the wrong track and that the Mutisieae s. lat. are the key to understanding early evolution of the family. Monophyly of the Barnadesioideae turned out to be well supported by morphological data but the sister group relationship between Barnadesioideae and the rest of the family was not possible to retrieve by morphological data alone and was discovered solely by RoBerT JANSEN from his early studies of chloroplast DNA together with Jerr PALMER (JANSEN & PALMER 1987). I heard about their results in 1986 and incorporated the cpDNA inversion uniting all Compositae except Barnadesioideae in my data matrix. It resolved a basal polytomy of various Mutisieae groups into the basal dichotomy we know today. My analysis was presented at the Berlin International Botanical Congress 1987, where it received sour comments from Cronauist. It was published later the same year (BREMER 1987) and formed a starting point for my 1994 book. i In Funk et al.’s Compositae Mutisieae s. lat. are reclassified into several different tribes and subfamilies. The basal phylogeny of the family, excluding the first branch of the Barnadesioideae, is however still not fully understood. The new tree appearing throughout the new book is fully resolved at the base, however only with low support, and this is indicated by dotted lines in some of the illustrations. If these poorly supported dotted branches are collapsed, the family tree excluding Barnadesioideae actually comprises a basal polytomy of six branches, five of them representing former Mutisieae groups, the sixth the rest of the family. Hence, there is still much interesting work to do on early evolution of the Compositae. The overall phylogeny assembled by Funk et al. provides many new insights and well supported relationships. Some former Mutisieae groups are reclassified in subfamily Carduoideae, for example the African Dicoma, Oldenburgia, Tarchonanthus, and their relatives, as tribes Dicomeae, Oldenburgieae, and Tarchonantheae, respectively. These groups were formerly particularly difficult to associate with the mainly South American Mutisieae and their reclassification in the essentially Old World subfamily Carduoideae seems very natural now that the sister group relationships have been established. Other odd genera, treated as orphans or provisionally included in a variety of tribes in traditional classifications of the family have been identified as sister groups of huge subgroups of the family involving one, two or three of the large subfamilies Comp. Newsl. 48, 20010 3 Carduoideae, Cichorioideae, and Asteroideae. Examples are the North American Hecastocleis, the Mediterranean Gymnarrhena and the South African Corymbium. ‘Consequently, these genera are elevated to subfamilies of their own. Because of their phylogenetic position, they are particularly valuable representatives of biodiversity. Suprageneric classification of the family now comprises 12 subfamilies and 43 tribes, a substantial increase, not only in names but also in information, compared to the 19" century classification of 2—3 subfamilies and 12-13 tribes that essentially survived throughout the 20" century. Within the large tribes there is also much new and interesting information to read about. Because of my own experience with the Anthemideae, I will take this tribe as an example. The South African genus Osmitopsis was the subject of my B.Sc. thesis at Stockholm University. I attended the Reading symposium on ’ the Compositae 1975 (HEYwoop et al. 1977), when there was much discussion about isolated genera and their correct classification in the 13 tribes of GEORGE BeENTHAM’s 19" century classification. Osmitopsis was one such genus that was discussed. It certainly looks like other Anthemideae but it was not possible to point out its close relatives within the tribe. This problem now receives its explanation: in the Anthemideae phylogeny in Funk et al.’s Compositae the genus Osmitopsis appears as the sister group of all other Anthemideae. At the Reading symposium I met Curis Humpnrries (1947-2009), who studied other genera of the Anthemideae, Argyranthemum and Anacyclus. We later decided to try cladistic analysis and subtribal reclassification of the whole tribe. We worked on this, using morphological data, for much of the 1980s and after about 15 years the analysis was finally published (BREMER & Humpuries 1993). This study provides another example of what can be understood and misunderstood by cladistic analysis of morphological data alone compared to the current analyses based also on molecular data, as seen in FUNK et al.’s Compositae. Fruit characters provided much information to HumpHrigs’s and my Anthemideae data matrix, and they also misled us to establish a number of artificial subtribes. One part of our phylogenetic hypothesis that has turned out to be correct is that the South African genera represent the ancestral complex of the Anthemideae. Biogeographical hypotheses play a prominent part in FUNK et al.’s Compositae. It is generally assumed that the family originated in southern South America. The family phylogeny in the book is accompanied by general distribution areas optimized on the tree, illustrated with different colours on the branches. Based on this optimization it is hypothesized that the Compositae migrated northwards via North America to the Old World and Africa where they underwent substantial evolution with subsequent cosmopolitan expansion. FUNK et al. even speculate that the phylogenetic and geographical position of the North American genus 4 Comp. Newsl. 48, 2010 Hecastocleis, the sister group to the supposedly originally African subfamilies Carduoideae+Cichorioideae+Asteroideae, represent an ancient track from the expansion of the family from South America to the rest of the World. I must admit I find the various biogeographical hypotheses put forward by Funk et al. fascinating, although I suspect Curis HUMPHRIES, who was always very skeptical about dispersal scenarios, would have regarded much of them as speculation. What strikes me when I look at all new findings of generic interrelationships is that close relatives are often to be found in the same geographic region. This also holds for isolated genera. Oldenburgia and Tarchonanthus (with their immediate relatives), both from Africa, are two examples mentioned above. Within the large tribes, e.g. Anthemideae (cf. above), Astereae, and Lactuceae, new subtribal delimitations to a large extent follow geographical distribution. If morphology is indecisive for identifying close relatives of a particular genus, it seems a useful recommendation would be: search nearby. Osmitopsis, the subject of my own first studies in Compositae, was nomenclaturally confused with another South African genus, Rel/hania of the Gnaphalieae — Linnaeus had got the types mixed up — and the latter genus thus became the subject of my Ph.D. thesis. As circumscribed in my thesis, Re/hania unfortunately turned out to be paraphyletic. I had simply missed that another Anthemideae genus Oedera was an ingroup in Relhania. The species were later reclassified in several different genera based on morphological data. This classification still remains to be evaluated by a cladistic analysis based on molecular data. The sampling in the Gnaphalieae treatment in FUNK et al.’s Compositae lacks many genera I am interested in, including a few I have described myself. There is no criticism of the Gnaphalieae treatment in Funk et al.’s Compositae on this point — it is simply an illustration of the fact that there is still a lot of interesting work to do in this and many other large tribes of the family. I look forward to future even larger trees with all genera well sampled. Many persons contributed to this book. I would like to mention Vicki FUNK especially. With never failing enthusiasm she organized The. International Compositae Alliance and orchestrated the whole book project to its successful completion. Comp. Newsl. 48, 20010 5 References BREMER, K. 1987. Tribal interrelationships of the Asteraceae. Cladistics 3: 210-253. Bremer, K. 1994. Asteraceae — Cladistics and Classification. Timber Press, Portland. Bremer, K. & C. J. Humpuries 1993. Generic Monograph of the Asteraceae- Anthemideae. Bulletin of the Natural History Museum, London, Botany Series 23:71—177. Heywoop, V. H., HARBORNE, J. B. & B. L. TURNER (eds.) 1977. The Biology and Chemistry of Compositae, 2 vols. Academic Press, London. _ Hinp, D.J.N. (ed.) 1996. Proceedings of the International Compositae Conference, Kew, 1994, 2 vols. Royal Botanic Gardens, Kew. JANSEN, R. K. & J. D. PALMER 1987. A chloroplast DNA inversion marks an ancient evolutionary split in the sunflower family (Asteraceae). Proceedings of the National Academy of Sciences USA 84:5818—5822. KKADERETT, J. W. & C. JEFFREY (eds.) 2007. The Families and Genera of Vascular Plants, Vol. 8, Flowering Plants, Eudicots, Asterales. Springer, Berlin. 6 Comp. Newsl. 48,2010 Ten Things I learned on the way to the Mother Tree (i. e., Mother Ship)* V. A. FUNK US National Herbarium, Department of Botany Smithsonian Institution, Washington D.C. 20560, USA funkv@si.edu * Adapted from a talk and two posters presented at the Botany 2009 meeting at Snowbird Utah, USA Abstract The completion of the recent book on the Compositae has answered some questions about evolution and biogeography of the family but it has also produced new and interesting areas for research. Ten such topics are discussed and some suggestions about future topics are made. Introduction The Compositae (Asteraceae) family is nested high in the Angiosperm phylogeny in the Asterideae/Asterales. The family contains the largest number of described, accepted, species of any plant family, ca. 24,000, with estimates of the total number reaching 30,000. There are 1600—1700 known genera distributed around the globe except for Antarctica. That the family is monophyletic has never been in question. Every early worker in plant classification recognized the Compositae as a group at some level (e.g., TOURNEFORT 1700, BERKHEY 1760,VAILLANT 1719- 1723) and in every type of analysis the family is monophyletic (e.g., SMALL 1919. BREMER 1987, JANSEN & PALMER 1987, HANSEN 1991a & b, MICHAELS et al. 1993, LUNDBERG & BREMER 2003). Possibly this is because morphologically the family is well characterized: flowers (florets) arranged on a receptacle in centripetal heads and surrounded by bracts, anther thecae fused at the margins to form a ring but the filaments are free, pollen pushed out by the style, calyx (when present) developed into a pappus, and the fruit is an achene (cypsela). Family wide treatments are few, BREMER’s 1994 cladistic analysis was the first revision of the whole family based on morphology since those of BENTHAM (1873a) and HoFrrMANN (1890) and although he recognized many of the problem areas in the cladograms of the family, the morphology did not generate enough data to resolve many of the issues. Just over ten years later KADEREIT & JEFFREY (2007) Comp. Newsl. 48, 2010 7 and their numerous co-workers reordered the genera, tribes, and subfamilies within the family based on recent morphology and molecular results and this work is now the standard reference for descriptions of the tribes and genera of the family. From the beginning those who studied this family thought the ray and disk pattern represented the basic head structure. Cassini in his famous 1816 diagram (FUNK et al. 2009a: Chapters 2, 6, 41) placed the Heliantheae at the center, the Vernonieae and Eupatorieae at one end, and the Mutisieae and Cichorieae (Lactuceae) at the other. Cronquist (1977) agreed and pointed out that BENTHAM thought the Heliantheae was most primitive (BENTHAM 1873b). CroNnquist (1955, 1977) and TURNER (1977) also thought that the Heliantheae was the most primitive tribe of the family, and accordingly assumed that the ancestor was a perennial herb (or shrub) with opposite leaves and yellow-flowered, radiate capitula. BREMER and JANSEN and their colleagues (BREMER 1987, JANSEN & PALMER 1987, 1988, JANSEN et al. 1991a, 1991b, BrREMER & JANSEN 1992, MicHaELs et al. 1993, JANSEN & KIM 1996, LUNDBERG & BREMER 2003) have used morphological and molecular data to change that view to one with a basal grade made up of members of the former Mutisieae (sensu CABRERA). The Book and the ‘Mother Tree’ The recent publication Systematics, Evolution, and Biogeography of Compositae (FUNK et al. 2009a) links the most recent molecular trees together in a metatree framework (FUNK & SpEecHT 2007) and uses that tree to provide a basis for understanding the evolution and biogeography of the family. The basic structure of the tree was taken from PANERO & Funk (2002, 2008) and BALDwmn (et al. 2002, 2009). The trees in PANERO & FuNK-(2008) contained extensive sampling from the base of the tree, the Mutisieae (sensu CABRERA), 3—10 genera representing all other tribes, and many taxa that had been “hard to place” in previous studies. The phylogeny was based on data from 10 chloroplast gene regions (ndhF, trnL-trnF, matK, ndhD, rbcL, rpoB, rpoC1, exon1, 23S-trnl, and ndhl). The Heliantheae Alliance portion of the base tree was taken from work by BALpwin and his collaborators (BALDWIN et al. 2002, BALDwin 2009) and is the result of nuclear rDNA of the ITS region. The phylogenies of the individual tribes are the work of many individuals (see the Acknowledgements) and a variety of types of molecular data, the most frequent being ITS and ndhF or matK. These molecular data have given us the opportunity to examine the history and morphology of the family in a way never before possible. The book (Funk et al. 2009a, c) includes overview chapters and a chapter on each tribe; for more information on the book see the The International Compositae Alliance (TICA) website (www.compositae.org), Comp. Newsl. 48, 2010 visit www. YouTube.com (search Compositae), or email compositaebook@gmail. com. All proceeds from the sale of the book go to the /nternational Association for Plant Taxonomy. Figure | is a summary tree of the metatree (taken from FUNK et al. 2009c) where each tribe or clade has been reduced to a branch (or visit www.compositae.org to see the ca. 900 taxon tree). Some of the biogeographic conclusions from the study include: I) 2) 3) 4) 5) 6) 7) Extant members of the basal radiation are found in southern South America; the basal clades inhabit the oldest rocks in South America (Southern Andes, Guiana Shield, Brazilian Shield). The members of the basal radiation comprise ca. 5 % of the extant genera in the Compositae. The central part of the area cladogram is a grade dominated by African based radiations with numerous movements into Eurasia and beyond. The members of these clades make up 66 % of the extant genera in the Compositae. The origin of the extant core Heliantheae Alliance is North America and Mexico with numerous links to South America and back to Mexico and North America. This clade contains 29 % of the extant genera. The Northern A frica-Mediterranean area and the Asia-Eurasia- European area played important roles in the history of the family especially in the radiation of the Cardueae, Lactuceae, Anthemideae, and Senecioneae. Major radiations in Hawaii, Australia, and New Zealand are in more highly nested clades. The north and northwest Andes, long thought to be the cradle of Compositae evolution, hold large numbers of taxa but all are highly nested. Western North America and Mexico hold many clades from a variety of tribes but all are highly nested. What we don’t know! I) 2) 3) What happened between the time of the South American radiation and the base of the African radiation? What happened among the clades in the ambiguous areas of the phylogeny? What happened between the Inuleae & Athroismeae clades, extant members of both originate in Africa, and the origin of the Heliantheae Alliance in SW United States -NW Mexico? Comp. Newsl. 48, 2010 9 During the construction of the Compositae metatree we began to call it the ‘Mother Tree’ (i.e., the Mother Ship) because we liked the analogy of a large facility where smaller ships could dock. These smaller ships (trees of individual tribes) could leave and others could dock as information about individual tribes change and when the Mother Ship becomes outdated, a new one can take its place and the smaller ships can dock at the new facility. In the process of producing the metatree several things became apparent that seemed interesting to relate to others. I have organized them into ten things that I thought were worth mentioning. Certainly there are many other interesting things related to the book and I take full responsibility for selecting those I discuss. The Ten Things 1. Unrooted trees are important. Trees can be rotated at the nodes and there is always a chance that personal bias will be reflected in the final shape of the tree. It is always a good idea to examine the results as an unrooted tree periodically during a project so that such bias can be removed. Figure 2 is an unrooted version of the summary tree (Figure 1). In The Book (Funk et al. 2009a) the unrooted tree is printed on the back of the bookmark so it can be turned in any direction. 2. All tribes except one were monophyletic or easily modified to be so(=GOOD JOB!). Most of the traditional 13 tribes were found to be monophyletic or could be made monophyletic with a few rearrangements (FUNK et al. 2009b). Tribes such as Cardueae, Vernonieae, Cichorieae, Senecioneae, Asteraeae, Anthemideae, etc. are all pretty much the same as they were from Cassini to the 1977 volumes (HEYwoop et al.1977). A few adjustments were necessary, for instance the tribe Inuleae ended up being divided into two tribes that are not closely related (Gnaphalieae and Inuleae) and it is still unclear exactly what goes in the Arctotideae, but these rearrangements were easily accomplished or the subtribes were found to be good. Even the Heliantheae s. lat., which has been divided into 12 or 13 tribes (depending on acceptance of Feddeeae), was a grade and monophyletic if one includes the Eupatorieae. In fact, it was the finding that the Eupatorieae were nested inside the rest of the Heliantheae s. lat. that resulted in the breakup of the group. However, it should be noted that only two of the newly recognized tribes in the Heliantheae Alliance had to be described as new, all of the others had been used previously. Cassini, in his famous 1816 diagram (reproduced in BALDWIN 2009), showed the Calyceraceae and Campanulaceae to be closely related to the Compositae and even though he did not have it in the diagram the text indicates that he also thought 10 Comp. Newsl. 48, 2010 the Goodeniaceae was close. He has turned out to be correct about this as well. So, if the previous tribes were so good, why do we have 42-43 tribes now instead of the traditional 13? The breakup of the Heliantheae s. lat. increased the number as did the breakup of the Mutisieae (sensu CABRERA; see #3 below) and several anomalous taxa did not fall into any group and were recognized as separate tribes. As we continue to examine the family the total number of tribes will likely continue to be in flux. However, the original 13 tribes described by our predecessors were, for the most part, good, and it is important that we acknowledge the latter for their insight. 3.The big exception is (as is usual) the basal grade. Frequently the base of a phylogeny is occupied by a basal grade. At the level of the Compositae family, the basal grade involves members of the Mutisieae (sensu CABRERA 1977). The group is largely paraphyletic but does have disjunct clades that make it polyphyletic. Unlike the Heliantheae there was no easy solution concerning the taxonomy and most of the clades had not been recognized at the tribal level before. The Mutisieae of CABRERA are now placed into 14 tribes (18 clades; Figure 1). In the Compositae Book (FUNK et al. 2009a) and here the use of the taxon “Mutisieae (sensu CABRERA)” 1s meant to represent the historic circumscription of the tribe as defined by CABRERA (1977). This is inno way meant as a negative reflection on the many contributions of CABRERA (see BONIFACINO et al. 2009). In fact, his work is the foundation for all modern work in the tribe (OrTIz et al. 2009) and within his 1977 paper he mentions groups of taxa that have a direct correlation to the results of the molecular analyses. Also, unlike the Heliantheae Alliance, the relationships between the morphological and molecular treatments are not always clear (Ortiz et al. 2009) and there is no universal agreement on the taxonomic solutions. The creation of the metatree has been the impetus for most of the taxonomists studying these groups to get together to find out more about the morphology of the various taxa. Another contrast between the Heliantheae Alliance and the Mutisieae (sensu CaBrerRA) is that until Katinas et al. (2008), CaBRERA’S treatment (1977) was the only one for the tribe in modern times while members of the Heliantheae Alliance had been studied by many individuals. As a result most of the clades within the Heliantheae Alliance already had tribal names that had been proposed while the Mutisieae had very few. 4. Chloroplast and nuclear data don’t usually agree. Chloroplast data appear to be more conservative and provide estimates of relationship for bigger picture questions such as the base tree for the family and relationships among tribes. Nuclear data such as ITS and ETS appear to Comp. Newsl. 48, 2010 11 change more rapidly and are commonly used to provide species and generic level relationships; sometimes they can also be used at the tribal level. When examining the relationships among clades at the ‘generic’ and ‘groups of genera’ level both types of molecular data can be used. It seems to be the case that at this level one often finds different arrangements of the groups in question. In addition, the attachment of outgroups can be quite different in the two types of data. For instance, repeated analyses of the tribes within the subfamily Cichorioideae (FUNK & CHAN 2009) show big differences in the placement of the tribes and unplaced taxa in the subfamily. BarKER et al. (2008) have indicated that there is a paleopolyploidization event at the base of the Compositae. This could help explain the discrepancy between the types of data and we may be in a position of always having different trees for different types of molecular data. We will have to be careful to select the type of data that will best answer the questions we are asking. In addition PENNisI (2008) mentions that data sets will have to be reanalyzed with different methods in order to determine the best tree and that the latter is not necessarily guaranteed by more data. 5. What about morphology; a iot of it is missing! It is imperative that we include morphology in our studies. Otherwise we will never understand evolution in the family. Unfortunately, much of the necessary work has yet to be done. Often characters are discussed only if they are informative within a certain group and that means we are missing information across the family. It is now incumbent on the community to organize and try to fill in the missing information. Two TICA projects have been started, a preliminary checklist of the species in the family including distribution (CHRISTINA FLANN is in charge of this project - christinaflann@gmail.com) and a virtual key to the species in the family starting with the USA (J. Mauricio BonIFAcINo is in charge of this project - mbonifa@gmail.com). It is hoped that these two projects will help provide some of the missing data. In addition, many of the tribes now have working groups that are making great strides in our understanding of the morphology. 6. Details of the distribution and morphology don’t matter to the big questions; but they are interesting. One of the most difficult aspects of the metatree project is trying to maintain the correct scale. Close enough that we would maintain the information we needed for the analyses but distant enough that we do not get lost in the details. An example from the distribution is the radiations within the Gnaphalieae in New Zealand and Australia. These are large and important radiations and we must examine them to achieve an understanding of the evolution within the tribe, however, that does not change the fact that the base of the tribe is in Africa and for family #2 Comp. Newsl. 48, 2010 wide biogeographic purposes we use that rooting. In morphology there are many examples, one is the ligulate corolla that is characteristic of the tribe Cichorieae but similar corollas are also found in several other taxa, Stokesia (Vernonieae), Dinoseris and Hyaloseris (Hyaloseris Clade; Stifftieae) and Catamixis (an anomalous taxon now tentatively placed in the Pertyeae). The evolution of a ligulate corolla in several isolated taxa does not diminish the utility of the character in the Cichorieae. A common expression is “Don’t throw out the baby with the bath water.” 7. Extinction is important & fossils would be nice or dates for the Compositae are difficult. Considering the size and importance of the family little has been published about its possible age. One reason may be the lack of any reliable macrofossils from the early diversification of the family. Without such fossils it is impossible to discount ideas of radiation followed by extinction that would drastically change the pattern produced by the extant taxa. There have been various estimates but several recent ones center on an origin of 41—50 Ma (see discussion in FUNK et al. 2009c) based on largely circumstantial evidence. Within the family, most authorities agree that, based on pollen data (GERMERAAD et al. 1968, MULLER 1970), most of the current tribes were in existence by the end of the Oligocene (25—22 Ma; MuLter 1981). Other dates of tribes or small clades have been proposed; many of these are just speculation. What is needed is a detailed analysis of pollen cores from the Antarctic, Australia, and southern South America to determine when the Calyceraceae + Compositae clade separated from the ancestors of the Goodeniaceae. It is likely that this will involve the ultra-structure of the grains since the external appearance of the grains of the base of the Compositae and its related families is similar. 8. The Biogeography makes a fairly complete story. I have been asked repeatedly why the biogeographic pattern is relatively clear in the Compositae but not so obvious in other families. One possible reason is that the family is relatively young and extinction has not wreaked havoc on its distributions. There are exceptions, of course, but in general it is pretty good. Of course, one can turn this around and say that this pattern may be the result of extinction! 9. Some things just don’t make sense. What in the world is Hecastocleis doing between the basal radiation and the jump to the old world? How did the huge radiation of the Heliantheae Alliance get from Africa to western North America and is it possible that such an event happened only once? Why in the world are Stifftia, the Gongylolepis clade, and the Hyaloseris clade grouping together? Why is the Asian Leucomeris clade Comp. Newsl. 48, 2010 3 grouping with the Brazilian and tepui Wunderlichieae? There are many such questions which illustrate that a good research project generates more questions than it answers. 10. The Real fun starts now! Now that we have the metatree what do we do with it? There are many things that can be investigated: the history of family through time, vicariance versus dispersal hypotheses, origin of floras in natural areas, character evolution, adaptive radiations on islands, convergent evolution, evolution of pollination systems, coevolution, characteristics of invasive species, and the evolution of unusual structure such as secondary heads, are just a few of the topics that come to mind. Each of these could be a separate paper(s). Truly, there are many interesting things to investigate. This project is a good example of the power of phylogenies in our quest to understand the history and evolution of life. Without such an overall organizing principle we will continue to look at small pieces of the puzzle and have no way to put them together. Acknowledgements The Compositae book, and hence this talk, would not have been possible without the work of many colleagues. Therefore it is with pleasure that I thank the many authors of the chapters in the book: ARNE A. ANDERBERG, GREGORY J. ANDERSON, BRUCE G. BALDWIN, NIGEL P. BARKER, RANDALL J. BAYER, GABRIEL BERNARDELLO, STEPHEN BLACKMORE, MAuRICIO BONIFACINO, ILSE BREITWIESER, Luc BROUILLET, RODRIGO CARBAJAL, RAYMUND CHAN, ANTONIO X. P. CouTINHO, DANIEL J. CRAWFORD; LALITA M. CALABRIA, JORGE V. Crisci, MICHAEL O. DILLON, VICENTE P. EMERENCIANO, CHRISTIAN FEUILLET, ORI FRAGMAN-SAPIR, SUSANA E. FREIRE, MercE GALBANY-CASALS, NURIA GARCIA-JACAS, BIRGIT GEMEINHOLZER, MICHAEL GRUENSTAEUDL, HANS V. HANSEN, VERNON H. Heywoop, Sven HImMMELREICH, D. J. NicHoLas HInp, CHARLES JEFFREY, JOACHIM W. KADEREIT, MARI KALLERSJO, VESNA KARAMAN-CASTRO, PER OLA Karis, LILIANA KATINAS, STERLING C. KEELEY, DaAvID J. Kem, NorsBert KILiAn, REBECCA T. KIMBALL, MARINDA KOEKEMOER, TIMOTHY K. Lowrey, JOHANNES LUNDBERG, MESFIN TADESSE, RoBERT J. McKenzie, Tom J. Masry, Mark E. Mort, Berti. NoRDENSTAM, CHRISTOPH OBERPRIELER, SANTIAGO Ortiz, PIETER B. PELSER, CHRISTOPHER P. RANDLE, PETER RAVEN, HAROLD ROBINSON, NApIA RoguE, GISELA SANCHO, ARNOLDO SANTOS-GUERRA, EDWARD SCHILLING, Marcus T. Scorti, JOHN C. SEMPLE, MIGUEL SERRANO, BERYL B. Simpson, Ros SMISSEN, FRANZ STADLER, Top F. Stuessy, ALFONSO SUSANNA, MAriA CRISTINA TELLERIA, MatTHEW Unwin, LoweLt Urpatscu, ESTRELLA UrTUBEY, JOAN VALLES, 14 Comp. Newsl. 48, 2010 RoBert VoGT, GERHARD WAGENITZ, STEVE WAGSTAFF, JOSEPHINE M. WARD, KUNIAKI WATANABE, LINDA E. Watson, & ALEXANDRA H. WortLey. They are however not to held responsible for the opinions I have expressed in this paper. I also thank the International Association for Plant Taxonomy (IAPT) and the Smithsonian Institution for sponsoring the publication of the book so that prices are reasonable. All proceeds from the book go to IAPT. References BaLpwin, B. G., Wessa, B. L. & J. L. PANERO 2002. Nuclear rDNA evidence for major lineages of Helenioid Heliantheae (Compositae). Syst. Bot. 27: 161-198. BaLpwin, B. G. 2009. Heliantheae Alliance. Pp. 689-711 Jn: Funk, V. A., SUSANNA, A., STUESSY, T.F. & R.J. BAYER (eds.), Systematics, Evolution, and Biogeography of Compositae. [APT, Vienna. Barker, M. S., KANng, N. C., MATVIENKO, M., Kozik, A., MICHELMORE, R. W., Knapp, S. J. & L. H. RieseserG 2008. Multiple Paleopolyploidizations during the Evolution of the Compositae Reveal Parallel Patterns of Duplicate Gene Retention after Millions of Years. Molecular Biology and Evolution 25(11): 2445-2455. ' BENTHAM, G. 1873a. 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VoctT, R., WAGSTAFF, S., WARD, J.M. & L. E. Watson 2009c. Compositae metatrees: the next generation. Pp. 747-777 In: Funk, V. A., SUSANNA, A., STUESSY, T.F. & R.J. BAYER (eds.), Systematics, Evolution, and Biogeography of Compositae. 1APT, Vienna. GERMERAAD, J. H., Hoppinc, C. A. & J. MULLER 1968. Palynology of Tertiary sediments from tropical areas. Review of Palaeobotany and Palynology 6: 189-348. Hansen, H. V. 1991a. Phylogenetic studies in Compositae tribe Mutisieae. Opera Bot. 109: 1-50. 16 Comp. Newsl. 48, 2010 Hansen, H. V. 1991b. SEM-studies and general comments on pollen in tribe Mutisieae (Compositae) sensu CABRERA. Nord. J. Bot. 10: 607-623. Heywoop, V. H., HARBORNE, J. B. & B. L. TURNER (eds.) 1977. The Biology and Chemistry of the Compositae. Academic Press, London. HOFFMANN, O. 1890-1894. Compositae. Pp. 87-391 Jn: ENGLER, A. & K. PRANTL (eds.), Die Natiirlichen Pflanzenfamilien \V(5). Leipzig: Engelmann. JANSEN, R. K. & J. D. PALMER 1987. A chloroplast DNA inversion marks an ancient evolutionary split in the sunflower family (Asteraceae). Proc. National Acad., U.S.A. 84: 5818-5822. JANSEN, R. K. & J. D. PALMER 1988. Phylogenetic implications of chloroplast DNA restriction site variation in the Mutisieae (Asteraceae). Amer. J. Bot. 75: 753-766. JANSEN, R. K., MICHAELS, H. J. & J. D. PALMER 1991a. Phylogeny and character evolution in Asteraceae based on chloroplast DNA restriction site mapping. Syst. Bot. 16: 98-115. JANSEN, R. K., MicHaexs, H. J., WALLACE, R., Kim, K.-J., KEELEY, S. C., Watson, L. E. & J. D. PALMER 1991b. Chloroplast DNA variation in the Asteraceae: phylogenetic and evolutionary implications. Pp. 252-279 In: Sottis, D. E., Sottis, P. S. & J. J. DoyLe (eds.), Molecular Systematics of Plants. Chapman Hall, New York. JANSEN, R. K. & K.-J. Kim 1996. Implications of chloroplast DNA for the classification and phylogeny of the Asteraceae. Pp 317-339 Jn: Hinp, D. J. N. & H. J. BEENTJE (eds.), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew, 1994. Vol 1. Royal Botanic Gardens, Kew. KADEREIT, J. W. & C. JEFFREY (eds.) 2006 [2007]. Asteraceae. The Families and Genera of Vascular Plants (Series Editor: K. Kupirzkt), Vol. VIII. Asterales. Springer, Heidelberg. Katinas, L., Pruski, J. F., SancHo, G. & M. C. TELLERIA 2008. The subfamily Mutisioideae (Asteraceae). Bot. Review 74: 469-716. LunpBerG, J. & K. BREMER 2003. A phylogenetic study of the order Asterales using one morphological and three molecular data sets. Intern. J. Plant Sciences 164: 553-578. Micnae.s, H. J., Scott, K. M., OLMsTEAD, R. G., SzAro, T., JANSEN, R. K. & J.D. Patmer 1993. Interfamilial relationships of the Asteraceae: insights from rbcL sequence variation. Ann. Missouri Bot. Gard. 80: 742-751. MULLER, J. 1970. Palynological evidence on early differentiation of angiosperms. Comp. Newsl. 48, 2010 7 Biol. Review 45: 417-450. Mutter, J. 1981. Fossil pollen records of extant angiosperms. Bot. Review 47: 1-142. Ortiz, S., Bontracino, J. M., Crisci, J. V., Funk, V. A., HANSEN, H. V., HIND, D. J.N., Katinas, L., RoQuE, N., SANCHO, G., SUSANNA, A. & M. C. TELLERIA 2009. The basal grade of the Compositae: Mutisieae (sensu CABRERA) and Carduoideae. Pp. 193-213 In: Funk, V. A., SUSANNA, A., Stussy, T. F. & R. J. Bayer (eds.), Systematics, Evolution, and Biogeography of Compositae. IAPT, Vienna. Panero, J. L. & V.A. Funk 2002. Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae). Proc. Biol. Soc. Washington 115: 909-922. Panero, J. L. & V. A. Funk 2008. The value of sampling anomalous taxa in phylogenetic studies: Major clades of the Asteraceae revealed. Molecular Phylogenetics and Evolution 47: 757-782. Pennisi, E. 2008. Building the tree of life, genome by genome. Science (Wash.) S208 ASNT SMALL, J. 1919. The origin and development of the Compositae. New Phytologist Reprint No. 11. Wesley & Son, London. Tournerort, J.P. DE 1700. Institutiones Rei Herbariae, 3 vols. Typographia Regia, Paris. Turner, B. L. 1977. Fossil history and geography. Pp. 19-39 In: Heywoon, V. H., Harporne, J. B. & B.L. Turner (eds.), The Biology and Chemistry of the Compositae. Vol. 1. Academic Press, London. VAILLANT, S. 1719-1725. Etablissement de nouveaux caractéres de trois Familles ou Classes de Plantes a Fleurs composées; s¢avoir, des Cynarocéphales, des Corymbiféres, et des Cichoracées. Histoire de 1’Academie Royale des Sciences avec les Memoires de Mathematique & de Physique 1718: 143— IO, th Dp Os LWMG2 NTs BUTS otis AVS ZS AOE RSS), ia Be IAs 1721: 174-224, t. 7, 8. 1725. Comp. Newsl. 48, 2010 CICHORIOIDEAE Cichorieae WUNDER. CARDU. es MUT. (+00S|) 9ealuOUJa/\ (z) eeaiuinboyy (Ov) snueydeajsig (p21) eeeqery (¢) aeaydueohye|q (+L€1) eeulayO9-y (¢) sidajosajayH (+92) aeuIpnojoiy-v (¢) eeeuweyjowaly (+00Z) SO eeaoYsiD (+006) 7O eeaOYoID (ELE) €-LO ejuoueM (L) eeeueyueUWAS (QZ) eeekued (7) seaiBinquap|o (EL) eeeyjueuoyoe (00SZ) eeenpieg (00L-gZ) eeawosig (|) eeapiajooj}seoay (02) eeayeuysos (0¢) Pan snpedoua}s 3 SG) elyoiapuny SI( ae suawoone7 €) apejo syeAY (62) Sas sidajojABuo5 (2) apejo suasojesy (8) EAWNS (pSz) seaisiny (€L¢) eealanessen (ZG) eeapliasoud (16) eaisepeweg (09) eeaoesa0/je9 (Ov) aea0e1uapooy (09) aeeceyjuehuayy (9z) eeaoe||Aydobuy (LL) eeeoeul|eud (OL) eee0e1wWsonas|y (Spz) seaceIpiiA}s Outgroups Al Leia | jo e'! ——S-. i a | 1 | SeSoa SSeS Sses seo SeSeSsSeoossee sesso anal =4 1 r u = | Fig. 1 Comp. Newsl. 48, 2010 ASTEROIDEAE 19 Senecioneae (3500) S-Tussilagininae Grade S-Othonninae Calenduleae (120) Gnaphalieae (1240) Astereae (3080) Anthemideae (1800) Inuleae (687) Athroismeae (55) Coreopsideae (550) Neurolaeneae (153) Tageteae (267) Feddeeae (1) Helenieae (120) Corymbieae. (9) Doronicum (40) Abrotanella (20) S-Senecioninae Bahieae (83) Chaenactideae (29) Polymnieae (3) Heliantheae (1461) & South America a 22 HE cuiana shieta Eee North & central Andes Southern Andes, southern South America ees General South America North America Sas North America, Mexico Central America, Caribbean Fig. 1 (contd.) S iS) S =_N = 2285 aVoeo —— oN o%o— o C= ® oVOS8 Beas OB e7s) = O05 8 Saws Eurasia Eurasia, Europe Eastern & central Asia Southern Africa _ Madagascar, tropical Africa Northern Africa, Mediterranean, southern Europi General Africa Australia and the Pacific Australia, N Guinea, N Caledonia, New Zealand Widespread or ambiguous 20 Comp. Newsl. 48, 2010 Figure Legends Fig. |. A summary tree based on the most recent supertree of the Compositae (Chapter 44), branches and internodes are colored according to the distribution of the terminal taxa or the optimization of those distributions. Some major radiations that are highly nested within the clade are show on the branches. [Hya. = Hyalideae; Wun. = Wunderlichieae; A-Arctotidinae = Arctotideae-Arctotidinae; A-Gorteriinae = Arctotideae-Gorteriinae]. Reproduced from Funk et al. 2009b with permission from IAPT. Fig. 2. An unrooted representation of the summary tree. The size of the circle indicates the number of species found in that clade. Colors are the same as in Fig. 1. Comp. Newsl. 48, 2010 1. Cichorieae Eupatonese 2. Eremothamneae 3. Moquineae 4 Calenduleae - Perityleae Madieae Millerieae Heliantheae Polymnieae Neurolaeneae Bahieae - Chaenactideae 3 Coreopsideae Tageteae Helenieae Feddeeae s e f © Athroismeae Inuleae = Astereae Senecioneae Abrotanella Anthemidtae Doronicum S Gnaphalieae A-Gorteriinae Heterolepis Corymbieae Platycarpheae Liabeae Gymnarrheneae Distephanus On @ Pertyeae Vernonieae ¢ Dicomeae Oldenburgieae & Tarchonantheae Hecastocleideae Cardueae Gochnatieae ~ Stenopadus clade . e Wundertichia 4 : ea s ® Leucomenis clade . Hyalis clade 7 Ly Gongylolepis clade 5 Onoserideae e Dinoseris clade Mutisieae Stifftia Nassauvieae Bamadesieae Calyceraceae Fig. 2 2). 22 Comp. Newsl. 48, 2010 Taxonomic delimitation of the genus 7ibetoseris SENNIKOV and the new genus Pseudoyoungia of the Compositae-Cichorieae from Eastern Himalaya D. Mairy* & G. G. Mairi** *Department of Botany, Taxonomy and Biosystematics Lab. University of Calcutta 35, Ballygunge Circular Road, Kolkata - 700 019, West Bengal, India debmaity@yahoo.com **Department of Botany, University of Kalyani Kalyani-741235, Nadia, West Bengal, India Abstract The genus TJibetoseris SENNIKOV is circumscribed here as monotypic with the single species 7. depressa (Hook. f. & THOMSON) SENNIKOV, which was known as Crepis depressa Hook. f. & THOMSON or Youngia depressa (Hook. f. & THOMSON) Basc. & STEBBINS. Its independent taxonomic status is explained and a new variety is recognized. The new genus Pseudoyoungia D. Mairy & Maiti (Compositae- Cichorieae) is proposed based on the remaining nine species of Tibetoseris SENNIKOV kept under two sections including the typical one. New combinations are made for these nine species. Introduction Cassini (1831) had established the genus Youngia Cass. for some ‘diversified tropical weeds mainly distributed in East Asia (SENNIKOV & ILLARIONOVA 2008). After that several studies including monographic work had been done to highlight the generic delimitation of the genus till the 20" century (LEDEBouR 1843-46, Bascock & STEBBINS 1937, KAMELIN & KovALEvsKAyYA 1993). About 25-30 species had also been added to the genus Youngia by different workers in the form of scattered papers. Presently the genus Youngia comprises 30 species (LACK 2007) or about 40 species (BREMER 1994, MaBBERLEY 2005). Recently SENNIKOV (in TZvELEV 2007) and SENNIKOV & ILLARIONOVA (2008) have tried to resolve the generic delimitation of the genus Youngia and have segregated three new genera based on the sections recognized by BaBcock & STEBBINS (1937), viz., Tibetoseris Comp. Newsl. 48, 2010 73 SENNIKov, Crepidifolium SENNiKov, and Sonchella Sennikov. The genus Youngia together with these three segregates and Ixeris (Cass.) Cass., Crepidiastrum Nakal, Lxeridium (A. Gray) TzveLey and Askellia W.A. WEBER were united ina new subtribe Iveridinae SENNIKOV. Tibetoseris SENNIKOV is currently considered to have 10 species of which 5 were previously kept under Youngia sect. Desiphylum by Bascock & STEBBINS (1937). Two further species were described by Sui (1995) and Sun & Cal (in SHiH 1995) in Youngia, another two new species and a new combination based on Crepis tianschanica Sui were published by TzveLev (2007). At present the 10 species of Tibetoseris are classified in three sections: Tibetoseris sect. Tibetoseris with only one species (7. depressa); sect. Parvae SENNIKOV with two species (7 parva and T. conjunctiva) and sect. Simulatrices SENNIKOV with the remaining 7 species (TZ. simulatrix, T. gracilipes, T. cristata, T: sericea, T. angustifolia, T. ladyginii and T. tianschanica). In the last six years the present authors have came across a large number of Compositae species in the Sikkim Himalaya including-many members of the tribe Cichorieae (Marty & CHAUHAN 2002, Marty & Marti 2001, 2007a, b, Malty 2005). Among these are Dubyaea hispida (D. Don) DC. (a member of a primitive genus of 14 species, cf. BABcock & STEBBINS 1937, Bremer 1994, Lack 2007), many species of Lactuca L., Youngia (s.1.), Stebbinsia umbrella (FRANCH.) LIPSCH. [syn. Crepis umbrella FRANCH. or Soroseris umbrella (FRANCH.) STEBBINS] as well as many collections of Tibetoseris depressa (Hoox. f. & THOMSON) SENNIKOV (=Crepis depressa Hook. f. & THOMsoN). The latter taxon, known to us from several collections from different localities in Sikkim, appeared odd in several respects, and this triggered a critical study on its taxonomic position. It has been thoroughly studied both in herbaria (CAL and BSHC) and in the field, along with anatomical features of cypselas, and it is finally concluded that Crepis depressa is a unique taxon which requires independent generic status, although it shares some characters with the related genera like Youngia (s.1.), and Stebbinsia Lirscu. (a genus often included in Soroseris STEBBINS).A fter our critical study we would like to recognize the genus Tibetoseris as monotypic with the species 7. depressa alone, excluding the remaining 9 species, which are placed under the presently proposed new genus Pseudoyoungia with two sections, Pseudoyoungia and Simulatrices. This proposal and its justification are explained in the following. Materials and Methods A total of 19 herbarium specimens of Crepis depressa, 12 specimens of Youngia 24 Comp. Newsl. 48, 2010 gracilipes, 10 specimens of ¥. simulatrix and 35 specimens of Youngia japonica at Central National Herbarium (CAL) as well as the recent collections at the herbarium of Botanical Survey of India, Sikkim Himalayan Circle, Gangtok (BSHC) were studied. The relevant literature was also consulted. Based on the studied specimens the morphological features of leaves, receptacle, relative length of corolla tube and ligule, stigmatic surface, cypselas and also the transverse section of cypselas, cellular view of pericarp were studied and the illustrations, camera lucida drawings and the photographs taken from Leica DME image analyzer and Magnus binocular microscope are provided to analyse the characteristic features of Crepis (Youngia or Tibetoseris) depressa and compared with genera Youngia (s.1.), and Stebbinsia. Results Crepis depressa is strikingly distinct by the orbicular-ovate to deltoid-triangular leaf blades with denticulate to entire margin, or if lyrate, then the terminal lobe is alike a typical blade, and the numerous, congested and relatively large capitula surrounded by the crowded leaves like a crown. In these features the taxon differs significantly from all species of Youngia. Furthermore, the receptacle is areolate and fimbrillate instead of areolate and glabrous (naked) as in other species of Youngia. In the genus Youngia the corolla tube is much shorter than the ligule, while in Crepis depressa the relative length of tube and ligule is fairly constantly 1:1 (studied in the specimens deposited at CAL and BSHC and also reported by Bascock & StepBins 1937). The corolla tube is glabrous, whereas there is a general tendency of hairiness on the outer surface of the corolla tube in most species of Youngia (s.1.). Anther tails are mostly free in Crepis depressa, but they are united in the other species of sect. Desiphylum as well as in most species of Youngia (s.1.). The pollen grains have an echinate exine as found in many Asteraceous species including Stebbinsia umbrella. The stylar surface and the stigmatic branches are densely barbellate. The barbs on stylar surface below the forking part are few and initially they are in groups below the forking part. The colour of stigmatic branches is dark-brown to black both in fresh material and after drying. The cypsela has a distinct coarse beak and is straw-coloured with numerous randomly scattered brown patches (Fig.1H; Pl.1C). The coloration is depending on the deposition of pigment in the epidermal cells of the pericarp as visible under light microscopic view. In other species of Youngia (s.1.) the cypselas are usually not beaked and have a uniform colour According to the opinion of BABcock & STEsBBINs (1937, pp.8—11) in Youngia Comp. Newsl. 48, 2010 25 (s.1.), “as a general rule, at least the outer achenes (cypselas) are consistently flattened” and on the basis of this character only these authors placed Crepis depressa under Youngia. A study of the recent collections (RAJU 4421, Rai 7356, SINHA & SHUKLA 20455, SHUKLA & Marti 18900, Marry 26880, Marri & SINHA 22485, Marty & PRADHAN 26880 — all at BSHC) does not show much difference between the inner and outer cypselas. Outer cypselas are slightly flattened and possess alternating broad and narrow ribs (Fig. IJ, PI.1C,D). Obviously the slight compression or flattening of outer cypselas does not constitute a basis for generic delimitation. In other respects, the cypsela of C. depressa is very different from those of Youngia (s.1.) including the species of sect. Desiphylum. In C. depressa the cypselas are significantly larger (more than (S—)7 mm long) with 10 ribs, while in Youngia (s.1.) cypselas are smaller, always less than 5 mm long, and with 12-15 ribs (cf. Lack 2007). The ribs are free and not fused in triplets at base. But in Youngia (s. str.) the ribs are fused in triplets at base (SENNIKov & ILLARIONOVA 2008). Bascock & Steppins (1937) had reported a 10—12-ribbed cypsela in C. depressa. However, in our specimens as well as specimens at CAL the cypselas always have 10 ribs (5 major and 5 minor) and very rarely 9 ribs, which may be due to immaturity. Similarly 11 ribs are found due to separation of one large rib at middle part of cypsela, but the basal and apical regions are again with 10 ribs (P1.1C,). Moreover, the figure provided by Bascock & STEBBINS (1937, Fig. 1g) shows a typical 10-ribbed cypsela with 5 major ribs alternating with 5 minor ribs, and not 12 ribs as mentioned in the text (op.cit., p. 34). The cypselas of C. depressa possess two stronger and broader/flattened lateral ribs compared to the narrowly wing-like ribs in Youngia. Interestingly, these large strong lateral ribs are not separable from other large ribs and are equal to them. This observation was also supported by Bascock & STEBBINS (1937). The anatomical sections of cypselas, further, show that the nature of the ridges and furrows of cypselas of Crepis depressa demand special attention. It is noteworthy that in the cross sections of the pericarp the ridges are widely rounded but furrows are acute on outer surface. The outline of inner surface of pericarp, in transverse section also shows similar characteristic features of both ridges and furrows forming a distinct strongly undulating line due to presence of large vallecular canals (Fig.1J,K, Pl.1D,E,F). However, this feature does not occur in other species of sect. Desiphylum nor in Youngia (s.1.), where the inner surface of pericarp is always straight or entire, and does not show distinct ridges or furrows except in Y. scaposa. The anatomy of the cypsela is quite unique. Epidermal cells contain brownish substances (tanniferous?) and are mostly invisible, covered with a thick cuticle. Papillate outgrowths or projections have been seen. Pericarp is entirely made up of sclerenchymatous cells except epidermis. Interestingly, in other species of 26 Comp. Newsl. 48, 2010 Youngia including sect. Desiphylum the pericarp is made up of parenchymatous cells with small patches of sclerenchymatous cells only in ridges (SENNIKOV & ILLARIONOVA 2008, Fig. 2. no. 1-16). In the acute furrows there are a few disorganized (parenchymatous?) cells. Very large vallecular canals are present below the ridges. There are 10 ribs, alternately large and small, 5 vascular strands and the destroyed testa. Endosperm cells are elongated (Fig.1J, K, Pl.1D-—F). Comparative morpho-anatomical studies of the closely related species of Youngia (s.l.), Stebbinsia umbrella and the new genus Pseudoyoungia described below emphasized the distinct taxonomic position of C. depressa (Table 1). Discussion and Conclusions Our current study shows that the outer cypselas are only slightly more flattened than the inner ones, and this feature is not of much use in the delimitation of taxa. The much smaller outer involucral bracts in comparison to the inner ones is a character also present in the genus Stebbinsia (Matty & Marti 2007a) as well as in the whole subtribe Crepidinae (LAck 2007). Therefore, these characters cannot be used for the delimitation of the genera within Crepidinae. Moreover, these features are rightly considered rather as subtribe characters by SENNIKOV & ILLARIONOVA (2008). Our study nevertheless shows that Crepis depressa is a unique taxon in morphological and reproductive characters. The related genera are Youngia (s.1.), and Stebbinsia (S. umbrella). The affinities of this taxon within Youngia (s.1.) may be with species like Y¥. pratti (BABc.) BABc. & STEBBINS having areolate or subfimbrillate receptacle (fimbrillae low, naked), ¥. stenoma (Turcz.) LEDEB. where corolla tube is slightly shorter than ligule, and Y. mairei (H. Lev.) BABc. & Srepsins and Y. henryi (Diets) Basc. & STEBBINS both having free anther tails. The alternating larger and smaller ribs of cypsela constitute another important character linking C. depressa with Youngia (s.1.), but the different wall structure of inner surface of pericarp, presence of large vallecular canals as well as entire sclerenchymatous cellular view of pericarp (mesocarp) in Crepis depressa immediately separate it from Youngia (s.l.). Moreover, the wider, flattened lateral ribs (not wing-like) which are inseparable from other major ribs found in C. depressa are very uncommon in this group. The much smaller outer involucral bracts are already considered a subtribe character by SENNIKOv & ILLARIONOVA (2008). Stebbinsia umbrella is the taxon most similar to Crepis depressa in vegetative features, and also in pollen exine ornamentation and size ratio of ligule and corolla tube in such a way that they may be mistaken for the same species, although Comp. Newsl. 48, 2010 7 there is major difference with respect to cypselar morphology (Marty & Marti 2007a). Stebbinsia (Crepis) umbrella is very different from all other species of Soroseris. Distinctive generic features in Soroseris are the only two outer involucral bracts, shorter than or exceeding inner ones; inner bracts 4, herbaceous with broad scarious margins and 4—6 florets in each capitulum. On the other hand in S. umbrella the involucral bracts are many, biseriate with outer ones (3—)5 in number, much shorter than the 13-19 strongly coriaceous inner ones, and there are 1543 florets per capitulum (GriERSON & SpRINGATE 2001, Marry & Marti 2007a). So, Liescuitz (1956) justified these differences by creating a new genus Stebbinsia Liescu. based on this unique taxon. Lack (2007) had returned back the species Stebbinsia umbrella to Soroseris as Soroseris umbrella (FRANCH.) STEBBINS and thus, unfortunately creating a heterogeneity to this generic character as was done by Stepsins (1940). We strongly support the independent status of the distinct genus Stebbinsia leaving the remaining species of Soroseris in that genus. This opinion is also supported by GrIERSON & SpRINGATE (2001). Crepis depressa is related only to this taxon and not to other species of Soroseris by having similar morphological appearance, biseriate involucres, and equal length of ligule and corolla tube. In fact Crepis depressa is a unique taxon having its own distinctive characters, though related to Stebbinsia or Youngia (s.1.), by some similar features, but these relationships are largely indirect as mentioned by Bascock & STEBBINS (1937). It is not a species of Youngia or Crepis, not even a close relative at all (Bascock & STEBBINS 1937, SENNIKOV & ILLARIONOVA 2008). Lactuca cooperi J. ANTHONY shares vegetative as well as reproductive features with C. depressa and is treated as a synonym (as also done by GRIERSON & SPRINGATE 2001). Also the specimens from Sikkim described by us (Matty & Marri 2001) as Lactuca pseudoumbrella D. Mairy & Maiti along with its var. chauhani D. Marry & Mairiare truly C. depressa. Thus the same species has been variously considered under Youngia (BABcOocK & STEBBINS 1937), as a new species of Lactuca by ANTHONY (1934) and again by Marry & Maiti (2001). Obviously it is a taxon subject to repeated misidentification, and its deceptive similarity to Stebbinsia has also been referred to above. SENNIKOV (in TZVELEV 2007) established the new genus Tibetoseris SENNIKOV based on Youngia sect. Desiphylum Basc. & STEBBINS. However, unfortunately the problem regarding the taxonomic status of Crepis depressa has not been solved, and the genus Jibetoseris remains a heterogeneous group as it was as sect. Desiphylum in BABcock & STEBBINS (1937). However, the latter authors indicated that this species /i.e. C. depressa/ is not closely related to the other tufted species of this genus” (BaBcock & STEBBINS 1937, p. 35). SENNIKOV & ILLARIONOVA 28 Comp. Newsl. 48, 2010 (2008) also mentioned that the first section of Youngia s.l. in BABCOCK’s system, Desiphylum, is most problematic: “For the time being we keep this group separate in /xeridinae until further evidences show the other way to classify it” (p. 77). The many diagnostic features of Crepis depressa, viz., suborbicular, entire to remotely denticulate or lyrate leaves, unwinged petiole, absence of old petiole bases on poorly developed stem, presence of cataphylls, areolate and fimbrillate receptacle, 14—21 florets per capitulum, equal length of ligule and corolla tube, free anther tails, dark brown to black style branches, exceptionally large cypsela with a coarse strong beak and the unique colour of cypsela and the nature of pericarp in transverse section, cellular view of pericarp and very long pappus readily refute the inclusion of this taxon in the genus Youngia (s.1.) or the genus Tibetoseris as presently circumscribed and demand its independent generic position. In conclusion the genus Zibefoseris should be monotypic with its single species T. depressa (Hook. f. & THOMSON) SENNIKOv and for the remaining nine species the new genus Pseudoyoungia is here proposed. A new variety of 7: depressa 1s also included here. Amplified diagnosis of the genus Tibetoseris Tibeotoseris SENNIKOV in TzvELEV, Bot. Zhurn. (Moscow & Leningrad) 92 (11): 1749. 2007; SENNIKOv, Komarovia 5(2): 90. 2008. (Fig. 1; Pl. 1). Tufted, perennial, laticiferous herb with strong vertical taproot. Caudex thick, strong, without withered old leaf bases. Stem absent. Leaves radical, rosulate, few to many, orbicular to broadly ovate or deltoid, entire or remotely denticulate or if lyrate, then terminal lobe alike the typical blade, glabrous or sparsely hairy along veins towards base; petioles long, unwinged; cataphylls often present. Capitula ligulate, few or many (>30), large, congested, surrounded by crown of leaves, with 14—21 florets. Involucre biseriate; outer phyllaries much shorter ('/,) than inner, setose outside along midrib, ciliate at apex; inner phyllaries (14—)15—17 mm long, setose outside along midrib, ciliate and crested at apex. Receptacle areolate-fimbrillate. Florets 14-17 mm long; corolla tube glabrous, equal to ligule in length. Anther tails free. Pollen echinate. Style branches black. Cypselas slightly compressed, more than (5—)7 mm long, with a strong coarse beak, 10-ribbed, hispid towards apex, yellow with dense blackish-brown patches; pericarp sclerenchymatous throughout; inner surface of pericarp ridged and furrowed with strong undulate line. Vallecular canals very large. Pappus biseriate, 11-13 mm long, white or stramineous, persistent. Chromosome number n=8. TYPE: Tibetoseris depressa (Hook. f. & THOMSON) SENNIKOV. Comp. Newsl. 48, 2010 29 Tibetoseris depressa (Hook. f. & THomson) SENNIKov var. depressa TYPE: INDIA: Sikkim, Kupup, 4500--5000 m alt., 9.X.1849, J. D. Hooker (K, lectotype, selected by Bascock & StTepsins 1937; B, G-DL, isolectotypes). Crepis depressa Hook. f. & THomson [C. B. CLARKE, Comp. Indicae 255. 1878, nom. nud., pro syn.] in Hook. f., Fl. Brit. India 3: 397. 1881. Youngia depressa (Hook. f. & THoMson) Basc. & STEBBINS, Publ. Carnegie Inst. Wash. 484: 33. 1937. Lactuca cooperi J. ANTHONY in Notes Royal Bot. Gard. Edinburgh 18: 198. 1934. Lactuca pseudoumbrella D. Marty & Mairi var. chauhani D. Matty & Matt! in J. Econ. Taxon. Bot. 25(3): 750. 2001, syn. nov. The specimens described as Lactuca pseudoumbrella vat. pseudoumbrella have a larger growth-form than Tibetoseris depressa var. depressa and more numerous capitula, but are here regarded as conspecific, although worthy of distinction on an infraspecific level. The leaves are also variable in shape from orbicular to deltoid-triangular. Tibetoseris depressa (Hook. f. & THOMSON) SENNIKOV var. pseudoumbrella (D. Marry & Marri) D. Marty & Marri stat. et comb. nov. Basionym: Lactuca pseudoumbrella D. Matty & Maiti var. pseudoumbrella in J. Econ. Taxon. Bot. 25(3): 750. 2001. - TYPE: Muguthang, North Sikkim, Muguthang (Lhonak valley), 31.VII.1999, Marri & Sinna 22485 (CAL!, holotype; BSHC!, isotype). Key to the varieties of Tibetoseris depressa: 1. Plants 6—7 cm in diam., capitula few to several (up to 20)........... var depressa 1. Plants larger, 10-15 cm in diam., capitula numerous (more than 30) ................ I esti 24 Voltas desi atss sgacmanth odeisac obi oeishttsidnicevndeyestuases eect var. pseudoumbrella Description of the new genus Pseudoyoungia Pseudoyoungia D. Matty & Mairi, gen. nov. — [ Youngia Cass. sect. Desiphylum Basc. & STEBBINS, Publ. Carnegie Inst. Wash. 484: 25. 1937, nom. inval., p.p.; Tibetoseris SENNIKOV in TzveELEV, Bot. Zhurn. (Moscow & Leningrad) 92(11): 1749. 2007, p.p., excl. typo; SENNIKov, Komarovia 5(2): 90. 2008, p.p.]. Plantae perennes, radice lignoso verticali et rhizomate tenui repenti; acaules vel subacaules; folia petiolata laminis lyratis vel pinnatifidis; petioli alati foliorum 30 Comp. Newsl. 48, 2010 veterum basibus plerumque persistentibus; calathidia (5—)9—20(—30) flora; involucrum biseriale; phyllis internis 8-12, 8—11(—16) mm lg., glabris vel nervo centrali setosis, ad apicem excrescentiis adnatis vel nullis; phyllis externis glabris, internis quadruplo (-*/,) brevioribus; corolla (13—)14-17 mm lg., tubo medio, 4-5 mm lg.; cypsela cylindrica, leviter compressa, costis 10-15 distinctis, valde inaequalibus vel subaequalibus; pappus biseriatus, albus vel stramineus, fragilis. Numerus chromosomatum n = 8. Plants tufted, perennial, with vertical strong taproot or slender creeping rhizome; stem absent or very short; leaves with sinuate-dentate, lyrate or pinnatifid blades, distinctly petiolate; petioles winged; old petiolar base often persistent; capitula with (S—)9—20(—30) florets; involucres biseriate; inner phyllaries 8-12 in number, 8—11(—16) mm long, glabrous or setose along the central nerve, sometimes conspicuously crested at the apex; outer phyllaries glabrous, '4 (— 7/,) as long as the inner ones; corolla (13—)14—17 mm long, with a medium sized tube (4-5 mm); cypsela cylindric, slightly compressed, with 10—15 prominently unequal (alternately wide and narrow) or almost equal ribs; pappus biseriate, white or straw-yellow, caducous. Chromosome no. n= 8. TYPE: Pseudoyoungia parva (BABc. & STEBBINS) D. Marry & Mairi. Pseudoyoungia sect. 1. Pseudoyoungia | Tibetoseris SENNIKOv sect. Parvae SENNIKOV | 1. Pseudoyoungia parva (Basc. & STEBBINS) D. Marry & Maiti, comb. nov. Basionym: Youngia parva BABc. & STEBBINS, Publ. Carnegie Inst. Wash. 484: 35. 1937. Syn.: Tibetoseris parva (BABC. & STEBBINS) SENNIKOV, Komarovia 5(2): 91. 2008. - TYPE: CHINA. Northern Szechwan, Sancha-trii, precipice, 4300-4500 m alt., 10. VII.1922, Harry SmitH 3218 (UPS, holotype). Plants with strong taproot; leaves sinuately or runcinately dentate to pinnatifid; involucres 10-11 mm long; phyllaries ventrally densely pilose; corolla tube 4—5 mm long, pilose. Distribution: CHINA. 2. Pseudoyoungia conjunctiva (Basc. & Stessins) D. Mairy & Maiti, comb. nov. Basionym: Youngia conjunctiva Basc. & STEBBINS, Publ. Carnegie Inst. Wash. 484: 37. 1937. Comp. Newsl. 48, 2010 3] Syn.: Tibetoseris conjunctiva (BAsBc. & STEBBINS) SENNIKOV, Komarovia 5 (2): 91. 2008. - TYPE: CHINA. Southwestern Kansu, upper Tebbu region, grassy slopes at foot of Shimen, 12000 feet, 7.VHI.1925, J. F. Rock 13062 (UC 489434, holotype; B, GH, isotypes). Distribution: CHINA. Pseudoyoungia sect. 2. Simulatrices (SENNIKOV) D. Marry & Marr, comb. nov. Basionym: TJibetoseris sect. Simulatrices SENNIKOV, Komarovia 5(2): 91. 2008. Plants with a slender creeping rhizome; leaves sinuately dentate to pinnately lobed; involucre 9-16 mm long; phyllaries ventrally glabrous; corolla tube 4—5 mm long, glabrous. TYPE: Pseudoyoungia simulatrix (BaBc.) D. Matry & Maiti. 3. Pseudoyoungia simulatrix (Basc.) D. Marry & Maiti, comb. nov. Basionym: Crepis simulatrix Basc., Univ. Calif. Publ. Bot. 14: 329. 1928. Syn.: Youngia simulatrix (BABC.) BABC. & STEBBINS, Publ. Carnegie Inst. Wash. 484: 39. 1937; Tibetoseris simulatrix (BABC.) SENNIKOV, Komarovia 5(2): 91. 2008. (Fig.2A—D). - TYPE: CHINA. Xizang: Southern Tibet, Nalamla, sandy place, 4200 m, 1882, Gyatsko (Dr. K1NnG’s collector) (G, holotype; B, CAL, GH, P, isotypes). Crepis smithiana Hanp.-Mazz., Acta Horti Gothob. 12: 357. 1938. - TYPE: CHINA. Sichuan: Taofu (Dawo), Taining (Ngata); in ripa glareosa fluminis, 3600 m, 04.1X.1934, Harry SmitH 11746 (UPS, holotype; A, isotype). Taraxacum altune D. T. ZHAI & C. H. An, J. Aug. 1‘ Agric. College 18(3): 1. 1995 (n. v.). - TYPE: CHINA. Xinjiang: Qiemo, Y. H. Wu 2644 (HNWP, holotype). Distribution: INDIA: Himalaya: Sikkim; NEPAL; CHINA. 4. Pseudoyoungia gracilipes (Hook. f.) D. Marry & Maiti, comb. nov. Basionym: Crepis gracilipes Hook. f., Fl. Brit. India 3: 396. 1882. Syn.: Youngia gracilipes (Hook. f.) BABc. & STEBBINS, Publ. Carnegie Inst. Wash. 484: 40. 1937; Tibetoseris gracilipes (Hook. f.) SENNIKov, Komarovia 5(2): 92. 2008. (Fig. 2 E-G). - TYPE: INDIA. Sikkim, alpine region, 1849, J. D. Hooker (K, lectotype, selected by Bascock & SteBBiNs 1937). Distribution: INDIA. Himalaya: Uttaranchal, Sikkim; NEPAL; BHUTAN; 32 Comp. Newsl. 48, 2010 CHINA. 5. Pseudoyoungia cristata (C. Summ & C. Q. Cat) D. Marry & Mairi, comb. nov. Basionym: Youngia cristata C. Sum & C. Q. Cai, Acta Phytotax. Sin. 33(2): 186. 1995. Syn.: Tibetoseris cristata (C. SHtq & C. Q. Cat) SENNIKOV, Komarovia 5(2): 92. 2008. - TYPE: CHINA. Xizang: Zayii, alt. 3900 m, [X.1935, C. W. Wanc 66121 (PE, holotype). Distribution: CHINA. 6. Pseudoyoungia sericea (C. Suma) D. Matty & Mairi, comb. nov. Basionym: Youngia sericea C. Sut, Acta Phytotax. Sin. 33(2): 186. 1995, nom. inval. (2 types cited); C. SHmH in SENNIKOv, Komarovia 5(1): 48. 2007. Syn.: Tibetoseris sericeus (C. SHIH) SENNIKOV, Komarovia 5(2): 92. 2008. - TYPE: CHINA. Xizang: Zayii, Mt. Karwar-kar-boo, Tsa-wa-rung, 3400 m, IX.1935, C. W. WANG 66254 (P, holotype; A, isotype). Distribution: CHINA. 7. Pseudoyoungia angustifolia (TZVELEV) D. Mairy & Maiti, comb. nov. Basionym: Tibetoseris angustifolia TzvELEvV, Bot. Zhurn. (Moscow & Leningrad) 92(11): 1750. 2007. - TYPE: CHINA. Kam (Tibet), systema fl. Jan-tzy-tzsjan (Golubaja), in cursu superiore fl. J-czju, 13000 ft., in fissuris rupium, 29. VII.1900, V. F. Lapyain 432 (LE, holotype). Distribution: CHINA. This species may be conspecific with P. gracilipes. 8. Pseudoyoungia ladyginii (TzvELEv) D. Maity & Mairi, comb. nov. Basionym: Tibetoseris ladyginii TzvELEV, Bot. Zhurn. (Moscow & Leningrad) 92(11): 1750. 2007. - TYPE: CHINA. Kam (Tibet), systema fl. Jan-Czy-Czjan (Golubaja), locus Nru-czju ad fl. Golubaja, 11700 ft., 25.VII.1900, V. F. LapyGin 380 (LE, holotype). Distribution: CHINA. Comp. Newsl. 48, 2010 33 9. Pseudoyoungia tianshanica (C. Sutin) D. Matry & Mairi, comb. nov. Basionym: Crepis tianshanica C. Suin (as “tianschanica”), Acta Phytotax. Sin. S890: 1995: Syn.: Tibetoseris tianshanica (C. Sut) TzveLeEv, Bot. Zhurn. (Moscow & Leningrad) 92(11): 1751. 2007.—TYPE: CHINA. Xinjiang: Thianshan Mt., Daniu He, in declivitate, alt. 2600 m, 23VII.1947, K. C. Kuan 2212 (PE, holotype). Key to the species of Pseudoyoungia: 1. Plants with taproot; phyllaries ventrally pilose; corolla tube pilose .............. 2 - Plants with creeping rhizome; phyllaries ventrally glabrous; corolla tube 5) RUSTROUIS 5 isd Se seecres reer ecclsar necro e Reertan REMC nc ROLE RA chi OE Rt Mm ABER a ed 3 2, [Penis (ey oro rN) Me ae ee ane me ea oe ties ep ee P. parva melalants amore thant 7 cra tall. o....c ces eehg Gia stcaccccoe setberteanaceokeces P. conjunctiva 3. Inner phyllaries dorsally appendaged .....................s.scesssseeseeneeseeneene P. cristata une pliyllanes dorsally novappendaged)..2)2.)....5.5. meee ee 4 soappus more than VOhmm Jong 7k 2.qy.8en.--seecddle « Aten celsecs hsste sce 2 Se aD OUSHLESS arn, /omnMAlOMe 4... se. .eec oP estes MeN eee ie eee tae ce 6 5. Peduncles pilose; inner phyllaries dorsally not spongy-thickened. P ladyginii - Peduncles glabrous; inner phyllaries dorsally spongy -thickened at base......... 20s 300d dd8SNaD SOR OCA BRASH ESA eS RDUIBNN = a ans be Ua JEL AOE ORI En Rae P. simulatrix eeinverepliyvilanes: dorsally Waitys. \.... Il "s]jao o1dreoid s]esAID ONVWISL (So See) crema | Ne ea ae ead wmnnuJsad unisadivy Ol ‘quo0z o1diesida ou} (TE “LZ ‘SBI4) oVoTNUyT OPISUL poyenyls “s[TeySAID o}e[eXO WUNTOTLS a]qISIA s]esAIO OIVBWUISIIg wnunssisoleds 6 Aq poausoy sinjydjnos Arepuoses ‘s]jo9 ordieoidgq wnuyoylydng (TE “8 ‘Sdiq) evoynuy | SIOAR] [¥}SO} pure Piles oidievside yog s]eisA1O ONeUWISIIg winnjofioyos unuoyydng g ‘Quoz oIdieorda apisurl poyenyis ‘s~e}sArO (O€ “SIq) 9vaIEISV eS sjeysAIo Osniq pJuoposajay a) ayl-esnip Aq pouwlloy oimjdjnos Arepuosas ‘IeA DJUOPOJAJaYy “F 70 Comp. Newsl. 48, 2010 “s]]90 o1divs0sou ‘payjeM-yory) ‘snoyewAyouosed “uoise1 MOLIN UT "s]]90 o1duevsosaw snoyewAyouored JO 19K] JSOULIOINO “UOISII MOLING UT ‘(paraAey] -1UN) JdAv] [2}S9} JO S]]9O BUUAYOUSJO[OS UI puNno.y s[eysAIo OeUIsIg ‘sioAvy Aue ul sjja0 eUAYouared o1dies0sow pur oidsesidg (pZ “B1q) SBOIUOUIOA, pIOofijs19 DIUOUAaY (€Z “‘SI4) SBTUOUIOA, “QUOZ [PISO ‘s]]90 o1drevs0sowl payjeM-Yory L, ‘UOISOI s]eSAIO OIVVUUISLI pJDsUuapUuod DANYJUBUOUA, ue (ZZ ‘SI4) svoplmoyjuYy seqsAi0 oNeWISLIg partie a a | ot wnusadsosina] aid], (17 ‘SIq) svaprmoyyuy | s]eisAIo asniq winjjdydossvu wnjaovuL] ae (OZ SIA) 9vaUOTDIUSS ; syeysAI9 ONeusiid SaJasv] SNISIPOAlA]S ie (6 SIA) SvoTUOUIOA, sjeish19 oNeUISIg 0¢ puvaysod psopajog (S] “SIq) evoTUOUIAA | qli Jo soviq eWAYoUaII]NS SUIpUNONS “s]]99 s]eSAIO DIVBUISIIg ee 6l veurAyouoied payjem-yoryy d1diwo0soul Ul punoy aD. eee ‘quoz o1diesida ayy Drapes’ = oie ky (Se “LT Sid) evoqnuy | PISUI PoyeNyIs ‘s]e}SAIO aye[eXO WUNIO[LS d|[QISIA s]eISAIO OIVBUISII Pio ise 81 Aq powiioy oinjdjnos Arepuosas ‘s][ao ordaesidgq oe ie "S][99 94} JO [[BM 1O]NO JeaU payenqts sjeysA10 ‘AjUo divsosaul JO sjjao euUTAYoUared (Q| “SI4) avolus}oH opesiyjed ‘payjem-ury} “poraAey-au0 “19jnO s]eiSAIO ONBUISLIg DIDISIAD DIPAD]]IDH) ‘LI Ti Comp. Newsl. 48, 2010 1 Gers o pug (9 “Brg) avauoura, ‘anssi} snoyewAyouaiapos poyfem-yoiyy “oiduvoosay S[eqSAIO O1BUISI Ig sisuajDSauas DIUOULa{ Lic - onss ae é s[eqsAIo OBUISTIg (SZ “SIq) SBOTUOLLIOA, oF SNOJBUWIAYOUSIOIOS poy]eM-YOIy} ‘o1idrwo0sajy msdajad DIUOULA, 72 Comp. Newsl. 48, 2010 Table 2. No. of studied taxa of Compositae, scrutinized for the presence or absence of CaQOx crystals. SI | Number of genera Number of genera No Tribe and species with and species without ae CaOx crystals CaOx crystals ie Eupatorieae - 5 genera, 7 species 2 Vernonieae 7 genera, 9 species 5 genera, 9 species a | Astereae 2 genera, 3 species 4 genera, 10 species 4. Gnaphalieae | 1 genus, 1 species - > Inuleae 3 genera, 5 species 12 genera, 20 species 6. Madieae - 1 genus, | species - i. Coreopsideae - 5 genera, 7 species 8. Heliantheae - 4 genera, 5 species Helenieae | 1 genus, | species - 10. | Tageteae - 1 genus, | species Me Senecioneae 2 genera, 2 species 6 genera, 16 species WE Anthemideae 3 genera, 3 species 4 genera, 5 species 13: Arctotideae - 3 genera, 3 species | 14. Calenduleae - | 2 genera, 2 species [Ss Cardueae 2 genera, 2 species 6 genera, 10 species 16. Pertyeae - | genus, 2 species 7 | Dicomeae ae 1 genus, | species 18. | Mutisieae - 1 genus, | species no: Cichorieae 1 genus, | species 10 genera, 14 species 22 genera, 27 species | 71 genera, 114 species es studied 93 genera 141 species Comp. Newsl. 48, 2010 73 Discussion and Conclusion The present study shows that 22 genera out of 93 genera investigated, and 27 species out of 141 species have calcium oxalate crystals; i.e. about 24 % of the genera and 19 % of the species possess calcium oxalate crystals. There are no correlations among the tribes of Compositae on the basis of distributional and data of calcium oxalate crystals, but this study indicates that the formation of calcium oxalate crystals may be antagonistic to the formation of phytomelanin pigment, since all taxa with a phytomelanin layer (belonging to the tribes Eupatorieae, Heliantheae, Coreopsideae and Tageteae) do not possess crystals. Similar type of statement had been advocated by Ropinson & Kinc (1977) on the achene wall of the tribe Eupatorieae. Although Gaillardia used to belong to the Heliantheae, it is now included in the tribe Helenieae, subtribe Gaillardiinae, which has no phytomelanin layer, and this genus possesses crystalliferous tissues in pericarp. In the mature cypselar wall there are two basic types of crystals. Predominant is the prismatic crystal type. Seldom druses are also available from the pericarpic zone (Brachycome campylocarpa, Brachycome heterodonta var. heterodonta, Tanacetum macrophyllum). Calcium oxalate crystals have been noted from cross sectional configuration of cypselar wall, i.e., pericarp associated with seed coat of cypselas. Sometimes structure of crystals is also observable in dry condition from the SEM photographs of cypselas, where crystals are distributed in the epicarpic zone of cypselas as in Aster thomsonii, Brachycome ‘heterodonta var. heterodonta, Buphthaimum salicifolium, Buphthalmum speciosissimum, Carpesium cernuum, Carpesium nepalense, Inula ensifolia. In these taxa, secondary sculpture is formed by the visible calcium oxalate crystals, which are situated inside the cell as cell inclusion. On the basis of present observation, the distribution of calcium oxalate crystals and druses in the pericarp and/or testa is particularly important for the tribes of Vernonieae, Astereae, Inuleae, Senecioneae, Anthemideae and Cardueae and is less important in Arctotideae, Calenduleae, Pertyeae, Dicomeae, Mutiseae and Cichorieae. On the basis of distribution of calcium oxalate crystals in different tissue region of cypselas, the studied taxa could be grouped into the following five categories: Crystals are distributed in different tissue regions of mesocarpic zone of pericarp; viz., Anthemis tinctoria, Arctium lappa, Baccharoides calvoana ssp. meridionalis, Bothriocline CategoryI Jaxa ssp. laxa, Brachycome campylocarpa, Catananche caerulea, Elephantopus scaber, Gaillardia aristata, Linzia melleri, Polydora poskeana, Vernonanthura condensata and 3 Vernonia spp. 74 Comp. Newsl. 48, 2010 Crystals are distributed only in epicarpic zone of pericarp; viz., Aster thomsonii, Brachycome heterodonta var. heterodonta, Carpesium cernuum, Carpesium nepalense, Inula ensifolia and Buphthalmum speciosissimum. Category Il Crystals are distributed in testal zone; viz., Cirsium vulgare, Category III = Craspedia uniflora, Emilia flammea, Tripleurospermum maritimum and Steirodiscus tagetes. Crystals are distributed in epicarpic zone of pericarp and testal Category IV zone; viz., Buphthalmum salicifolium. Crystals are distributed in epicarpic and mesocarpic zone of Category V pericarp; viz., Zanacetum macrophyllum. Among the five categories, most prevalent is first category and least common are forth and fifth categories, whereas second and third categories are intermediate in frequency. In first category, crystals are distributed either in one row (Anthemis, Arctium, Bothriocline, Gaillardia and Vernonanthura) or in many rows in other taxa. Sometimes crystals are restricted to a specific region of mesocarpic zone. For example, in Brachycome campylocarpa, crystals occur in anterio-posterior lobes region, in other taxa in furrow region (Catananche, Elephantopus, Vernonia cistifolia) or rib region (Linzia and Vernonia petersii). In Baccharoides calvoana ssp. meridionalis, the crystals are found only on the basal arm of inverted *T’- shaped sclerenchyma bundle of mesocarp. Regarding distribution of crystals, one diacritical thing is the position or location of crystals. In Anthemis, these are always found towards inner wall of cells, whereas in Bothriocline and Gaillardia, they are towards outer wall of cells. In the second category, crystals are arranged in one row of cells. Secondary sculpture is formed by calcium oxalate crystals, which exist inside the cell, and are visible in SEM pictures as well as cross sections of mature cypselas. In the third category, crystals are found in definitely organized testal cells as in Cirsium, Emilia and Tripleurospermum, or in crusted disorganized zone (Craspedia, Steirodiscus). According to Dormer (1961), calcium oxalate crystals occur in the inner part of ovary wall. Our study indicates that this statement is true for the tribe Cardueae, but is not valid for all other taxa of Compositae. The observations by KALLERSIO (1985, 1990) on presence or absence of druses in Comp. Newsl. 48, 2010 75 the pericarp in some members of Compositae has a significant value for the tribe Anthemideae. Prycuip & RUDALL (1999) mentioned that druses are frequently occurring in dicotyledons, whereas raphides are commonly distributed in monocotyledons. The present study does not fit well with this statement, since the majority of the studied species have prismatic crystals and in the Compositae druses or raphides are less common, (reported e.g. from the Gynura-Solanecio Clade, JEFFREY 1986). Other than ovary wall, crystals have been noted from floral parts of two species of Helianthus (H. annuus, H. tuberosus) by Meric & DANE ( 2004). We have noted the absence of calcium oxalate crystals in the mature cypselar wall in Helianthus annuus (recorded as a taxon without CaOx crystals; Table 2). Observation on calcium oxalate crystals in various tissues of cypselas in the tribe Cardueae (ZAREMBo & BoyKo 2008) shows that inner region of mesocarp, adjacent to testa in Serratula, possesses crystals. Similarly, the border between pericarp and testa in Synurus contains crystals. Identical type of distribution of oxalate crystals is seen here in Anthemis, Arctium and Baccharoides. Presence of calcium oxalate crystals in the vegetative parts of Aster squamatus was recorded by Meric (2009). Although morphology and distribution of calcium oxalate crystals are now somewhat known, their functional aspects are not clearly known or understood. There are various opinions regarding the possible functions of oxalate crystals as follows: 1) to regulate the calcium levels in plant cells and tissues (FRANCESCHI & Nakata 2005). ii) to enhance the strength of the plant tissue or tissues (FRANCESCHI & Horner 1980). lii) to protect against animals especially herbivores (MoOLANO-FLORES 2001). Iv) to actas storage tissue of calcium and oxalic acid (FRANCESCHI & HORNER 1980, Prycuip & Rupa. 1999). v) to detoxify heavy metals (Nakata 2003). vi) to precipitate calcium salt in same specific environmental condition (Waite & Broapiey 2003). Vii) to remove toxic substances (BorcHERT 1984). Vill) to overcome salt stress and homeostasis (HURKMAN & TANAKA 1996). ix) to regulate light intensity in plant tissues (FRANCESCHI & HORNER 1980). 76 Comp. Newsl. 48, 2010 x) to provide mechanical support to the plant tissues (NAKaTA 2003). Xi) to participate in transformation of light energy to the chloroplast of the parenchyma cells of leaves during photosynthetic process (KUO-HUANG et al. 2007) and xii) to facilitate pollination by providing a visual signal or a scent interesting to insect (CHASE & PEACOR 1987, D’Arcy et al. 1996). Some of these proposed functions are closely related or near synonymous, and some may seem more plausible than others. One recent study (JAUREGUI-ZUNIGA et al. 2005) suggests that CaOx crystals are important in bulk calcium regulation but not in metal detoxification. Whatever their functions may be, the distribution, shape and structure of calcium oxalate crystals are very specific and obviously taxonomically useful at least in certain plant families (MoLANO-FLoREs 2001). From the taxonomic viewpoint, it is worth noting that they are not universally present in all plant organs, but instead they are confined to specific crystalliferous tissue only. Their morphology and distribution have been indicated as genetically controlled by the cell (ILARSLAN et al. 2001, Nakata & McConn 2000). Usually a specific taxon possesses a specific type of crystals (PRycHID & RUDALL 1999). The present study shows similar results. Although crystal formation is gene controlled, various external factors such as temperature, air pressure, light intensity, pH of soil and other environmental factors may affect the formation of calcium oxalate crystals (FRANCESCHI & HorNER 1980, Garry et al. 2002, Kuo-Huanc et al. 2007, Meric 2008, 2009). ILARSLAN et al. (1997, 2001) have also suggested that deposition of calcium oxalate crystals serves as a storage source for calcium and it is assumed that both the mitosis and cytokinesis are regulated by calcium ions (HEPLER & WayNE 1985). The climatic condition of an area associated with phenology controls the formation of calcium oxalate crystals in the cambial zone of Citharexylum (Verbenaceae) as has been observed by Marcati & VERONICA (2005). They have indicated that an abundance of crystals was observed during water deficit condition, whereas during flowering time associated with rainy season, crystals were rarely observed. Absence of calcium oxalate crystals in the mature cypsela of non-phytomelanin containing taxa is probably the primitive condition. Presence of calcium oxalate crystals without presence of phytomelanin may be one step evolved condition, and absence of calcium oxalate crystals in phytomelanin bearing taxa may be highest evolved state, since these taxa have been considered as belonging to the most advanced lines in the family (BREMER 1994, 1996, KADEREIT & JEFFREY 2007). Evolutionary relationships of crystals within the cypselar wall are diagrammatically shown in Table 3. From this observation it is obvious that formation and distribution of calcium oxalate crystals have an evolutionary role. Comp. Newsl. 48, 2010 77 Therefore, further detailed critical studies regarding the distribution of calcium oxalate crystals in mature cypselar walls in other taxa of Compositae will help developing an improved classification of the Compositae. Table 3. Probable evolution of cypselar structure of Compositae in relation to calcium oxalate crystals. Presence of phytomelanin in pericarp: Absence of CaOx crystals in cypselar wall Absence of phytomelanin in pericarp: Presence of CaOx crystals in cypselar wall Absence of phytomelanin in pericarp: Absence of CaOx crystals in cypselar wall Acknowledgements The authors are highly grateful to the Directors and Curators of the following herbaria: AD (Adelaide, South Australia); BRI (Queensland, Australia); LISC (Lisboa, Portugal); NSW (Sydney, Australia); RB (Rio de Janeiro, Brazil); SRGH (Harare, Zimbabwe);:and Z (Zurich, Switzerland) who kindly have provided mature cypselas for this study. 78 Comp. Newsl. 48, 2010 References AHMED, K.A., KHAFAGI, A.F. & A. EL-GAzzAr 1986. Taxonomic studies on Aster L. (Compositae). Phytologia 61 (5): 299-310. ANDERBERG, A. A. 2009. 39, Inuleae. /n: Funk, V. A., SUSANNA, A., STUESSY, T. F. & R. J. 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Herbivory and calcium concentrations affect calctum oxalate crystal formation in leaves of Sida (Malvaceae). Ann. Bot. 88 (3): 387-391. Nakata, P. A. & M. M. McConn 2000. Isolation of Medicago truncatula mutants defective in calcium oxalate crystal formation. Plant Physiology 124: 1097-1104. Nakata, P. A. 2003. Advances in our understanding of calcium oxalate crystal formation and function in plants. Plant Science 164 (6): 901-909. NorDENSTAM, B. 1977. Senecioneae and Liabeae—systematic review. /n: HEYwoop, V. H., HARBorNE, J. B., & B. L. TuRNEr (eds.), The Biology and Chemistry Comp. Newsl. 48, 2010 | 81 of the Compositae 2, pp. 799-830. Academic Press, London. NorpenstaM, B. 1978. Taxonomic studies in the tribe Senecioneae (Compositae). Opera Bot. 44: 1-83. NorpenstaM, B. & G. Ex-Guazaty 1977. Floral micromorphology and pollen ultrastructure in some Centaureinae (Compositae) mainly from Egypt. Publ. Cairo Univ. Herb. 7 & 8: 143-155. Panpey, A. K., Sincu, R. P. & S. Cuopra 1978. Development and structure of seeds and fruits in Compositae: Cichorieae. Phytomorph. 28 (2): 198-206. Prycup, C. J. & P. J. RuDALL 1999. Calcium oxalate crystals in monocotyledons: a review of their structure and systematics. Amn. Bot. 84 : 725-739. Reese, H. 1989. Development of pericarp and testa in Calenduleae and Arctotideae (Asteraceae). Bot. Jahrb. Syst. 110 (3) : 325-419. Rosinson, H. & R. M. Kin 1977. Eupatorieae —systematic review. Jn: HEywoop, V. H., HarBorne J. B. & B. L. Turner (eds.), The Biology and Chemistry of the Compositae 1, pp. 437-485. Academic Press, London. Sincu, R. P. & A. K. PANDEY 1984. Development and structure of seeds and fruits in Compositae-Cardueae. Phytomorph. 34 (1-4): 1-10. Waiter, P. J. & M. R. BroapLey 2003. Calcium in plants. Ann. Bot. 92 (A): 487-511. ZAREMBO, E. V. & E. V. Boyko 2008. Carpologia de algunas Cardueae (Asteraceae) de Asia oriental. Anales del Jardin Botanico de Madrid 65 (1): 123-134. Figs. 1-27. Distribution of calcium oxalate crystals in different taxa. 1,2: Anthemis tinctoria. 3: Arctium lappa. 4: Aster thomsonii. 5: Bothriocline laxa. 6: Brachycome campylocarpa. 7: Buphthalmum salicifolium. 8: Carpesium cernuum. 9: C. nepalense. 10: Catananche caerulea. 11: Cirsium vulgare. 12: Craspedia uniflora. 13: Elephantopus scaber. 14: Emilia flammea. 15: Gaillardia aristata. 16: Inula ensifolia. 17: Linzia melleri. 18: Tripleurospermum maritimum. 19: Polydora poskeana. 20: Steirodiscus tagetes. 21: Tanacetum macrophyllum. 22: Vernonanthura condensata. 23: Vernonia cistifolia. 24: Baccharoides calvoana ssp. meridionalis. 25: Vernonia petersii. 26: V. senegalensis. 27: Buphthalmum speciosissimum. (C - Cuticle. CR - Crystal. D - Druse. E - Endosperm. EP — Epicarp. HO- Hollow. ME - Mesocarp. NCP- Non Cellular Pellicle. P- Pericarp. PA - Parenchyma. PH — Papillate Hair. RCV—Resin cavity or Resin duct. SC — Sclerosed cells. SCC — Secretory cells. SCL - Sclerenchyma. SCLB — Sclerenchyma brace or bundle. SCV — Secretory cavity. T - Testa. TE — Testa epidermis. TI — Testal inner zone. TM — Testal middle zone. TPA - Thin-walled parenchyma. TWPA- Thick-walled parenchyma. VB — Vesicular body. VT - Vascular trace or vascular tissue). 82 Comp. Newsl. 48, 2010 2 o = 2 2 iS i) ‘= Arctium lappa [ = bi S Se See Os. ORI SSSS 6% ~ 0g): oe! 2) ‘eo Or, U) are" Baccharoides calvoan 4 Aster thomsonii i de! } ines Ho-@® o aN 0 (On O} Aieh\ 83 Comp. Newsl. 48, 2010 asuajedeu winisedied wniyjoyoyes wnujeyjydng exe] ‘dss exe; aulja0uYyjog ARDY eee ay fe ESS Comp. Newsl. 48, 2010 84 Jaqeos sndojueydejz (apie eae Cee Soren D> E> ested > Dan BEG 4 eioyiun eipedses 2 Comp. Newsl. 48, 2010 85 NOG eo KK erlaG~s 10; COC OSEI Es 7 1 ~ {eC % B6) CO . \\, SSBCOo Ose PN Bs 530256 @} ORS J) \ EF Linzia meller Inula ensifolia Ko =. % s, IG ES, } S Tams £ IA} b= aC SSK yy aS EVES . D, a Comp. Newsl. 48, 2010 86 PJESUBPUOD BINYJUBUOUII/ mi Cy - A l MOYO A wnuwiuew wnuuedsoin aah . ys ad] TZ 300 Comp. Newsl. 48, 2010 AG RY Sh ER > DN Ne SCLB Buphthalmum speciosissimum Vernonia petersii —~ Vernonia senegalensis Comp. Newsl. 48, 2010 88 ‘008 x “p1j0fisua DjnuT :¢¢ “OOS x ‘asuajodou UINISadUDD -hE “YOR x “wnnusad unsadiny :¢¢ ‘wnuussisoisads unupoysydng :7¢ “OOP x ‘winofinyvs wnujoyjydng :1¢ ‘O09T x “Piuoposajay “Tea DJUOpodajay aulordyooig :Q¢ ‘Bdupz0jAdiup9 aulorAYyoDAY 267 ‘OR x ‘MUOSMOY) 121s 187 ‘s]eISATO OJe[eXO WINTOT[eO JO UOINGISIp oy) SurMoys sydesZojoyd was "SE-8Z “SSI ae ») g ‘te Comp. Newsl. 48, 2010 89 Pollen studies on some populations of Aspilia (Asteraceae) in Nigeria A. E. ADEGBITE Department of Biological Sciences, University of Agriculture Abeokuta, Nigeria eadegb@unaab.edu.ng Abstract Pollen studies were carried out on twenty populations of three species of Aspilia namely A. africana, A.kotschyi and A. helianthoides and two hybrids obtained from them. All the populations produced tricolporate and tetracolporate pollen grains that showed high pollen fertility and viable seeds except the interspecific hybrid and one population of A. helianthoides which showed low pollen fertility and seed sterility. The occurrence of tetracolporate pollen grains in the Aspilia populations and the mode of flower colour segregation in the interspecific hybrid which implicated polygenic inheritance provide evidence for the previously hypothesised polyploidy state of the species in the genus Aspilia. Low pollen fertility and high seed sterility were suggested to be responsible for the absence or rarity of natural hybrids among species in the genus Aspilia. Introduction The importance of pollen grain size and morphology in the characterization of plant species have been stressed by many authors. CLAUSEN (1962) reported the use of pollen size as a reliable character for distinguishing two species of Betula. OLoropE & Torres (1970) used pollen characters to characterize species in the genera Zinnia (Sect. Mendezia) and Tragoceras of the family Asteraceae and artificial hybrids obtained from them. The use of pollen characters as reliable indicators of ploidy level in related species has been reported (STEBBINS 1950, OLORODE & BAQUAR 1976). The genus Aspilia THouars contains highly variable species with considerable overlap in their morphological and cytological characters but without evidence or report of natural hybridization, even in locations where their populations overlap. There are controversies over the mode of evolution and genomic status of the species of the genus (TURNER & Lewis 1965, PoweLL & Cuarrecasas 1970, 90 Comp. Newsl.48, 2010 SoLsriG et al. 1972, OLoRODE 1974, Gitt & Omoicui 1988, ADEGBITE & OLORODE 2002a). While some workers suggested the species to be polyploids as a result of the high chromosome numbers reported for the species, others suggested the species to be diploid in view of the high incidence of regular bivalent formation in the microsporogenesis of the species (ADEGBITE & OLORODE 2002b). This study therefore attempts a pollen characterization of the Nigerian species of the genus Aspilia with a view to unravelling the controversies about the genomic status and mode of evolution of the species in the genus and provide explanations for the absence or rarity of natural hybridization among species of the genus Aspilia. Materials and Methods Pollen grains were obtained from flower heads collected from field and screen- house populations of the species under study. Table 1 shows the sources and distinguishing characteristics of the populations studied. Slides for pollen studies were prepared by shaking pollen grains from open capitula into a drop of stain (cotton blue in lactophenol) on the slide or by squashing mature anthers in the stain and allowed to stay for 24 hours. Five such slides from five different capitula were prepared for each population and pollen counts were made from ten fields on each slide at x 100 magnification. . Full pollen grains with the cytoplasmic content stained blue were considered fertile, while those that were not stained or only partially stained and with collapsed outline were counted as sterile (JACKSON 1962, OLORODE & BAQUAR 1976). Measurement of the diameter of full and deeply stained pollen grains (excluding the walls and the spine) were taken at <400 using an ocular micrometer. The measurements were later converted to microns using a stage micrometer. Estimate of mean pollen size was based on measurements of 30 pollen grains from five different slides for each population. Photomicrographs of pollen grains stained with FLP-orcein were taken at x400 magnification for pollen structural. studies. Results Pollen data for the populations studied are recorded in Table 2, while the photomicrographs of the fertile pollen grains of the species and sterile and abnormal pollen rains of the hybrid are shown in Plate 1. All the populations possessed tricolporate and fenestrate pollen grains. These are single spheroid pollen grains with a cover of coarse network of pointed projecting Comp. Newsl. 48, 2010 9] spinous elements that are interrupted by three (or sometimes four) protuberances (germ pores) in a fixed geometrical pattern. This type of pollen is characteristic of the subdivision Tubuliflorae (= subfam. Asteroideae) of the family Asteraceae (Moore & WesB 1978). The mean pollen size was found to range between 22,53y1m and 32.51um in A. africana (Prrs.) C.D. Apams with the Northern Nigeria accessions having relatively higher values. Pollen fertility in A. africana ranged from 51,83 % to 98.55 % being lowest in population a,f and highest in a,,s (cf. Tables 1 & 2). The mean pollen sizes and percentage fertility values were comparable in the different populations and in the screen house and field populations of the same accession except in accessions a, and a,, where the field populations showed relatively low pollen fertility percentages of 51.83 and 58.58 respectively. The pollen fertility can be said to be high in A. africana except in the field populations of a, and a,, which coincidentally have high coefficient of variation (CV) values of 17.26 and 12.34 respectively. Populations of A. kotschyi (Scu. Bir.) Outv. were found to have mean pollen sizes that ranged between 23.5lum and 27.14um and pollen fertility percentage ranging from 62.67 to 99.19. Relatively low pollen fertility percentages of 62.67 and 65.97 were recorded in the field and screen house populations of accession k, respectively which were also found to have relatively high CV values of 8.91 and 9.44 respectively. The screen house and field populations are comparable in their pollen size and pollen fertility percentage. Aspilia helianthoides (ScuuM. & THONN.) Otiv. & HIERN has mean pollen sizes ranging from 23,00um to 25.77pm and pollen fertility percentages ranging from 63.99 to 98.92. The field and screen house populations are comparable in their pollen size. and pollen fertility percentage. Some unusually small and infertile pollen grains with mean pollen size of 9.98 +1.66um and high CV value of 16.63 were observed in population h, (Table 2, Plate Ic). The A. kotschyi and A. helianthoides(k,x h,) F, hybrid has a mean pollen size of 26.62 + 4.66um, a relatively low pollen fertility of 54.38 % and high CV value of 17.51. Small infertile pollen grains with a mean size of 11.83 + 1.73 um and a high CV value of 14.62 were observed in this hybrid (Table 2, Plate 1b). The pollen size interval exhibited by this hybrid extends beyond the size limits of the two parent plants (Table 2). The A. africana * A. africana intraspecific hybrid (a,, x a,) F, has a mean pollen size of 24.83 + 1.19um and a pollen fertility of 82.84 %. The pollen, ligule and stigma colours are yellow in A. africana and white in A. helianthoides. In A. kotschyi, while the pollen colour is yellow, the ligule is deep purple. The colours of pollen, ligule and stigma in the interspecific hybrid 92 Comp. Newsl.48, 2010 are intermediate between those of the parent plants. The pollen is light yellow or cream, the ligule is light purple or mauve and the stigma is near white with purple tinge. Discussion The use of pollen grain size and morphology in the characterization of plant species cannot be over-emphasised. The three species and the hybrids showed considerable overlap in their pollen size range and fertility. Pollen fertility is generally high in all the three species except in the field populations of accessions a, and a,, of A. africana, field and screen house populations of accession k, of A. kotschyi and screen house population of accession h, of A. helianthoides which also showed high CV value (Table 2). The observation in this study of low pollen fertility in populations having high coefficient variation (with few exceptions) has been reported earlier for A. africana and Melanthera scandens (Scuum. & THONN.) BRENAN by ADEGBITE et al. (1984). This suggests a correlation of high pollen size variance to a low pollen percentage fertility, which was attributed to the level of meiotic irregularities observed in those populations concerned (ADEGBITE & OLORODE 2002b). Meiotic irregularities in the interspecific hybrid reported by ADEGBITE & OLORODE (2002b) can be said to be responsible for the low pollen fertility and total seed sterility (even with backcrossing) of the hybrid. The high pollen fertility recorded for the populations of the three species was not unexpected in view of the regularity of meiosis observed in them (ADEGBITE & OLORODE 2002b). The low pollen fertility and high seed sterility observed in the interspecific hybrid provide explanations for why natural hybridization or any evidence of it was not observed and has not been observed or reported in the field populations, even where the species are sympatric. The low pollen fertility and seed sterility in the hybrid are expected to have chromosomal and/or genic origins. Some situations are known where directed processes produce unexpectedly high frequency of fertile megaspore in angiosperms (SwANson 1957, BURNHAM 1962). The failure of backcrosses with pollen from the parents to produce seeds contrary to expectation indicates that no preferential or directed segregational processes were operative in the formation of megaspores in the hybrids. The unusually small infertile pollen grains observed in the interspecific hybrid (k, x h,) F, and population h, of 4. helianthoides are microspores formed from micronuclei which apparently resulted from meiotic irregularities such as laggards, uneven distribution of chromosomes and chromosome clusters from multipolar second telophase during microsporogenesis in the hybrid and population h, Comp. Newsl. 48, 2010 ; 93 (ADEGBITE & OLORODE 2002b). Tetracolporate pollen grains in addition to the tricolporate ones were observed in all three species of Aspiia studied. The presence of pollen grains with four germ pores corroborates the proposition that the genomes of West African species of Aspilia are polyploids (STEBBINS 1950, OLoRODE & BAQuaR 1976). The pollen colour of the species corresponds to the colours of the ligule and stigma except in A. kotschyi and the interspecific hybrid. A comparison of the flower and pollen colours of A. kotschyi and A. helianthoides and their F, hybrid (Table 2) indicates that flower colour is polygenically determined with additive gene effect i.e. colour expression depends on gene dosage. The number of gene pairs involved will be ascertained through flower colour segregation ratio in the F.,. While it can be deduced from parts of the observations in this study that the same gene(s) may condition pollen, ligule and stigma colours in Aspilia as suggested by OLorobE (1976), the situations in A. kotschyi and the interspecific hybrid suggest that the characters are influenced by other gene interactions which modified their expressions. The sterility of the F, interspecific hybrid and the low number of the amphiploids (polyploidised interspecific hybrid plant) that survived to flowering stage made it difficult to draw precise conclusions in respect of the inheritance of flower colour in the F, of the interspecific hybrid. References ADEGBITE, A. E., Oso, G. M. & K. APENSILE 1984. Morphological and Cytological Studies of some species in the genus Aspilia THouars (Compositae). B. Sc. Botany dissertation, University of Ife, Ife-Ife (unpublished). ApveGsitE, A. E. & O. CLoRODE 2002a. Karyotype studies of three species of Aspilia THouars (Heliantheae—Asteraceae) in Nigeria. Plant Science Research Communications | (3): 11-26. Apecsitr, A E. & O. OLoropE 2002b. Meiotic studies of some populations of three species of Aspilia THouars (Heliantheae — Asteraceae) in Nigeria. Nig. J. Bot. 15: 74-83. BurnuaM, C. R. 1962. Discussions in Cytogenetics. Burgess Publ. Co., Minneapolis, Minnesota. CLAUSEN, K. E. 1962. Size variations in pollen of three taxa of Betula (1). Pollen et spores 4 (1): 169-174. 94 Comp. Newsl.48, 2010 Git, L. S. & I. D. OmMorcur 1988. Cytomorphology of the tribe Heliantheae (Asteraceae) from Southern Nigeria. Feddes Repertorium 99 (1-2): 1-13. Jackson, R. C. 1962. Interspecific hybridization in Haplopappus and its bearing on chromosome evolution in the Blepharodon section. Amer. J. Bot. 49 (2): 119-132. Mook, P. D. & J. A. Wess 1978. An Illustrated Guide to Pollen Analysis. Hodder & Stoughton, London. OvoropE, O. 1974. Chromosome numbers in Nigerian Compositae. Bot. J. Linn. Soc. 68: 329-335. OLoropE, O. 1976. Genetic studies on Nigerian weeds. Chromosome strategies in evolution. Paper presented at the 7" Annual Conference of the Science Association of Nigeria, Zaria. OLoRODE, O. & S. R. BAQuar 1976. The Hyparrhenia involucrata—H. subplumosa (Gramineae) Complex in Nigeria: Morphological and Cytological Characterization. Bot. J. Linn. Soc. 72: 212-222. O.oropE, O. & A. E. OLORUNFEMI 1973. The hybrid origin of Emilia praetermissa (Senecioneae: Compositae). Ann. Bot. 37: 185-191. OLoRoDE, O. & A. M. Torres 1970. Artificial hybridization of the genera Zinnia (Sect. Mendezia) and Tragoceras (Compositae-Zinninae). Brittonia 22: 359-369. . PoweELL, A. M. & J. Cuatrecasas 1970. Chromosome numbers in Compositae: Colombian and Venezuelan species. Ann. Missouri Bot. Gard. 57: 3/4379) SousriG, O. T., KyHos, D. W., PowELL, M. & P. H. Raven 1972. Chromosome numbers in the Compositae. VIII: Heliantheae. Amer J. Bot. 59 (8): 869-878. STEBBINS, G. L. 1950. Variation and Evolution in Plants. Columbia University Press, New York. Swanson, C. P. 1957. Cytology and Cytogenetics. Prentice-Hall Inc., Englewood Cliffs, New Jersey. Turner, B. L. & W. H. Lewis 1965. Chromosome numbers in the Compositae, IX. African Species. Journ. S. Afr. Bot. 31: 207-217. 95 Comp. 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Newsl. 48, 2010 SHUM Suu SHUM ve | LI OI-v0'S LL’S7-00'€Z Saploujunijay “V ajding g{ding nee é ope ee YOSIOY 131] dooq IISA Ve VV 6-69'°S VRE GeSaSG IAYISION “V DUDILAD “VW MOTOR MOTIOX MOTIOA me | wellieiery WS CEES CE SS°86-E8 TS Axevuiuing | (‘4) vuvoiifo yp MOTTOX. MOTTOX. MOTTOX € 6L V 61 IFES ET | 878 CSCI 'A(ex"'®) 8C < te y oAney] he sae S| 10 Molin = 9b] EL IFES IT | 00°0 oss | ‘aC ux) | (Q) zz ysyding | ajding Ms é Nisha met Cp) saployjupyay “y eee wieaig x 1MYOS]OY “py oyyM 10 (0) MOTTO . . . 5 I (' I ) (e) WE ystjding aiding ; os ie IS LI 99° VFC9'9T | BE PS LLvl AC Ux ¥ 1317 Yor] 102 Comp. Newsl. 48, 2010 New taxa and combinations published in this issue Pseudoyoungia D. Maity & Maiti, gen. nov.: p. 29 Pseudoyoungia sect. Simulatrices (SENNIKOV) D. Matry & Maiti, comb. nov.: p. 31 Pseudoyoungia angustifolia (TzvELEV) D. Mairy & Maiti, comb. nov.: p. 32 Pseudoyoungia conjunctiva (Basc. & StepBins) D. Mairy & Maiti, comb. nov.: p. 30 Pseudoyoungia cristata (C. Sum & C. Q. Cat) D. Mairy & Maiti, comb. nov.: p. 32 Pseudoyoungia gracilipes (Hook. f.) D. Mairy & Maiti, comb. nov.: p. 31 Pseudoyoungia ladyginii (TzvELEV) D. Marry & Mairi, comb. nov.: p. 32 Pseudoyoungia parva (Basc. & Stepsins) D. Mairy & Maiti, comb. nov.: p. 30 Pseudoyoungia sericea (C. Summ) D. Mairy & Maiti, comb. nov.: p. 32 Pseudoyoungia simulatrix (Basc.) D. Maitry & Mairi, comb. nov.: p. 31 Pseudoyoungia tianshanica (C. Sum) D. Maitry & Maiti, comb. nov.: p. 33 Senecio subnivalis Y. AJANI, J. Noroozi & B. NorD., sp. nov.: p. 47 Tibetoseris depressa (Hook. f. & THOMSON) SENNIKOV var. pseudoumbrella (D. Marry & Marti) D. Marry & Mairi, stat. et comb. nov.: p. 29 a ‘ 7 » a0 y k vo : a ‘oe + ? Z a ‘i - 7 i ia ; ett n - os : arn ‘ r ae He q 7 7 ; _ . CORTES in tte mie i * = » ‘i A. hal Wacre TX ent by WV i . “+ ; ‘ i vas ’ ‘ ‘ ¥ ke. t a a ‘ t ' { j RENT i i af \ } ra a >i ; ; 4\ : Ae Ea ITHSONIAN INSTITUTION Pcp wil 88 01551 1538