ON THE PRIMARY STUDY OF THE SYSTEMATICS AND FLORISTICS OF COMPOSITAE
Ling Yeou-ruenn
1993, 13(2):
151-201.
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The paper would presnt the Phylogeny and Floristics of Compositae(Asteraceae) in the world. According to the reseach from Fossils and Paleopollen grains, thecentre of the original speciation and the earliest divergence of Compositaecould be at the areas around the Ancient Mediterranean and Montana of USA in Eocene or earlier of it in Tertiary era. The Pro-Asterales, especiallyPro-Heliantheae was perhaps, firstly, derived from Pro-Rosales, Pro-Spir-aeoideae, and from Pro-Asterales the primitive species from Trib. Heli-antheae, Trib. Mutisieae and Trib. Cichorieae, were diverged separatelyfrom the ancestors of Pro-Heliantheae, Pro-Mutisieae and Pro-Cichorieae.As the result of the climate became drought and cold, and the strong stim-ulation from insect-pollination in earlier Tertiary era, the tubular corolla, which sepals were adnated regularly or irregularly as bilabiate at the base, as in Pro-Heliantheae and in Pro-Mutisieae, or sepals adnated laterally onone-side, as in the ligulate corolla in Pro-Cichorieaea, syngenesious stamenfrom united free anthers and capitula, which were aggregated the florets onthalamous, were formed. Pro-Vernonieae Pro-Eupatorieae, Pro-Inuleae andSenecioneae, maybe derived earlierly or laterly from the ancestor ofPro-Heliantheae at tropics of C.& S. America, when the ancestorshad migrated and extended the areas into there from the speciation cen-tre. Pro-Astereae and Pro-Anthemideae, perhaps, derived from Pro-Inuleaethere, where the climate became drought or cold in Eurasia, when theancestors had migrated there;Pro-Cynareae and Pro-Arctotideae derivedfrom Pro-Mutisieae, and Pro-Calenduleae from Pro-Senecioneae in droughtand hot areas in Africa and S.-W. Asia, when ancestors of them migratedthere. Trib. Cichorieae solitarily developed from Pro-Cichorieae. The pres-ent species and genera of tribes in the world were separately divergedfrom above pro-tribes. The corolla of florets in capitula from ancestors in the family in ear-lier Tertiary age developed, perhaps, from homogamous capitula with wholebisexual florets and adnated regularly or irregularly tubular at base orbilabiate corolla, into heterogamous, i. e. the marginal florets had beenreduced into pistillate then to asexual with ligulate corolla and it usuallywith teeth 3→2 and 0 at apex of it. Those species are usually found at theareas of insect-pollination. However, in the drought area of wind-pollinationthe marginals were reduced into pistillates with narrow tubular corolla and 4→3→2→0-teeth at the apex. The inner bisexual florets have also the red-uction of pistillate absorption, but absorbed ovaries from present to abs-ent. Then, the unisexual and monoeceous or dioeceous capitula and reducedcorolla in it latererly occured. Those are the evolution in subfam. Tubulif-lorae. With the exception of the capitula, in subfam. Liguliflorae, thosehave whole bisexual ligulate florets, which are accompanied with milkysap in plants, were solitarily developed. Both ligulate corollae with pistill-ate or asexual florets in heterogamous. capitula and with whole bisexualflorets, so-called "concordance", were derived solitarily from differentancestors, but in the same or similar environment. The present pollen grains have more or less shown the differentiationsand relationships among the tribes, for example, in Heliantheae, Vernonie-ae, Eupatorieae, Inuleae, Senecioneae, Astereae and Anthemideae, those arewith 3-zonocolporate, the extine cuneate and the surface spinate-spinulateand with an excrescence system that composed of the column and basalcushion. But in Heliantheae the columellar union with exposed foot layerunlike in others without the layer. In Mutisieae and Cynareae those are byunique extine strata with megacolumellar and microcolumella layers, butin the former it is by a little ellipsoid apertural lacuna and in the latterit is by the ellipsoid and furrowed apertural lacuna. Contrarily, it havecommon lophate extine surface and homogeneous ellipsoid apertural lacunain Cichorieae. It seems to be commended in cp-DNA and Cladistics, especially in cp-DNA., but it still is, now, difficult to clarify naturally the relationship,.among the tribes and a part of not whole, subtribes, which have mentioned data in Cladistics, because by means of equal-parallel. characters of them, it can not show what is the primary or the main from that the secondary or minor of the characters, and it have not shown what is the individual variation of development in evolution for categories of the family. In addition, it do not sufficient to be one only, not more, for the com-parated family, as in Campanulaceae, included Lobeliaceae for the considerable relationship with Compositae. By the mentioned ideas, author'd like to surnmarize Compositae into 2 subfamilies, 5 supertribes and 12 tribes (1990*) according to the reseach from morphology, incl. Pollen grains and Plant Geography, incl. Paleo-eeography. The category is as follows: Compositae——Asteraceae I, Subfarn, Tubuliflorae DC.——Carduoideae Kitam.——Asteroideae Carlq 1) Supertrib Helianthanae Y, R. Ling 1. Trib. Heliantheae Cass. (incl. syn. Trib. Helenieae Cass.) 2. Trib. Inuleae Cass, 3. Trib. Astereae Cass. 4. Trib. Anthemideae Cass. 2) Supertrib. Eupatorianae Y. R. Ling 5. Trib. Eupatorieae Cass. 6. Trib. Senecioneae Casse 7. Trib. Calenduleae Cass. 3) Supertrib, Vernonianae Y. R. Ling 8. Trib. Vernonieae Cass, 9) Supertrib. Mutisianae Y. R, Ling 9. Trib。Mutisieae Cass. 10. Trib. Cynareae Less, 11. Trib. Arctotideae Cass。Ⅱ, Subfam Liguliflorae DC——Cichorioideae Kitam.——Cichoriace〇e Juss. 5) Supertrib. Cichorianae Y. R. Ling 12. Trib. Cichorieae Riechb. Since the alternates with Ice-ages and intermittence of Ice-ages from Tertiary to Quaternary eras in the world, the ancestors of tribes of Com-positae usually migrated towards warm areas, and new groups, including new genera and species, were diverged and arisen from the ancestors and survival species of the family. In C. & Se America it is either the diverging centre or one of the present distributional mass centres, because there, distribute the most accounts of genera and species, especially the genera with monospecies and oligospecies in the western Hemisphere in the:Q world. The area with the second accounts of genera and spe'cies of the family is in Africa, especially in S, S.-E. Africa, included Madacascar Island, S.-W. and Tropical West Africa, where the ancestors of the family moved there by the Equator had shift towards midland from the:north in Africa. So the area, including Se, S.-W., Tropical West and S——E. Africa, is another one of diverging and present distributed mass centres, in Eastern Hemisphere, Nevertheless, in Eurasia most of genera and species, which occured earlierly there, were migrated from 2 routes, one was, through the "route";C. Asia, W. & S.-We Asia from the Ancient Mediterranean;another one is via the ancient adjacent "route" between N.-E, Asia-N-W America from N.-W. America migrated into there. Those are by the evidence from Paileopollen grains, However, the "Hengduang-Himal-ayan Mts."in S.-W. China is a special area by the Floristic point of view. It is either the refuge of Compositae from N# Asia in Ice Age, or-an area inhabited by some ancient Tertiary endemic and present regional endemic species, latter were arisen after Ice Age. Besides, it is also one of the secondary speciation centres after Ice Age and one of present distributional mass centres, Caucasus in Europe, although there are less genera and species of Compositae, it as fact, is an exchange-convergence area by Floristics among the Europe, C.& W. Asia and Africa for the family, and the Alps is a small refuge with a few endemic genera and species of Compositae in C. Europe。In W. & N.-W. America it have more endemic species, which arose before and after Ice Ages, because it is the high mountain areas and also a refuge of Compositae in Ice Age, unlike in E. &C.of N. America, where is instead of hills or plains only, and many genera and species of the family were destroyed in Ice Age, In addition, although it distribute less genera and species of Compositae in Pacific Continent, that wats separated from Gundwana before Tertiary Era, than in other Continents , the genera and species of the family were solitarily diverged while the Continent was in drifting towards the Equator, and more regional endemic genera and species of it are included in the Continent now, and only a few common genera and species were moved into there from C.-S. America, S. Asia and Africa. By the Floristic point of view author divides the areas of Compositae into 4 Kingdoms, 27 subkingdoms. I。Holarctic Kingdom 1) Eurasia Division 1. Subholarctic subkingdom 2. East Sibiria Far East subkingdom 3. Eastern Asi'a forest subkingdom 4. Hengduang-Himalayan Mts。forest subkingdom 5. Eurasia steppe or semi-desert steppe subkingdom 6. C., W & N. Europe forest or forest-steppe subkingdom 2) Ancient Mediterranean Division 7. C. Asia temperate desert subkingdom 8. Qinghai-Xizang plateau;subkingdom 9. W. & S.-W. Asia_Arab_N.-E. Africa tropical desert and savanna subkingdom 10. The Mediterranean tropical, subtropical and winter-green forest subkingdom 3) N。America Division 11. N. & N,-E America lower mountain forest subkingdoni 12. W. & N.-W. America mountain forest subkingdom 13. C. of N. America steppe subkingdom Ⅱ。Paleotropical and subtropical in S. Hemisphere Kingdom 14. Indo-China E. & S.-E. Indian tropical forest and savanna. subkingdoni 15. Iadian-Pakistan tropical desert and savanna subkingdom 16. Malaysian-Philippine tropical Jungle subkingdom 17. N. & N.-W African tropical desert and Xerophyte subkingdom 18. E. & C。African plateau and savanna subkingdom 19. Tropical West Africa——S. Africa——S.-E, Africa forest & steppe subkingdom Ⅲ. Neotropical Kingdom 20. C. America We Indian Islands subkingdom 21. Amazon Tropical Jungle subkingdom 22. Brazil Pampas subkingdom 23. Andes C. of S. American forest subkingdom Ⅳ. Australian Kingdom 24. N. & E. Australian forest & steppe subkingdoni 25. C. & N.-W。Australian tropical desert & semidesert-steppe subkingdom 26. S.-W Australian subtropical winter-green forest subkingdom 27. The tropical forest subkingdom in Pacific Islands.
