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The Flora of Australia As a Measure of the Antiquity of the Angiosperms

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The Flora of Australia As a Measure of the Antiquity of the Angiosperms The Flora of Australia as a Measure of the Antiquity of the Angiosperms KARL SUESSENGUTH 1 INTRODUCTION only in Australia and in New Guinea. These , IF ONE INTENDS to consider as difficult a mammals resemble reptiles more than any of problem as the early history of a large land the other mammals because they lay eggs, area, Australia would seem to be particularly have a cloaca, and still possess the number of suitable for such a study: its long geographic shoulder bones of primitive animals. As fos­ isolation and the great number of scientific sil evidence has shown, their ancestors ap­ investigations to which it has been subjected peared during the Triassic formation , to be­ make it a natural choice. come, in fact, the first of the mammals. They Australia's land connection with the island increased in number during the Jurassic pe­ groups of Malaysia (except for New Guinea) riod, but, to a great extent, they died out was ended in the Upper Cretaceous period. as early as the Eocene epoch. In Australia, According to physicists' calculations, based however, some of these primitive mammals on the rates of disintegration of radioactive have survived to this day, affording us illus­ elements, about 30-40 million years have tration of the concept of "endemism by passed since the Eocene epoch in early Ceno­ conservarion." zoic time. Inasmuch as the Upper Cretaceous The marsupials, too, are notably typical of period occurred before the Eocene epoch, it Australia. They are not limited to Australia, can be concluded that, on the whole, the Aus- for there are opossums in North and South .tralian flora and fauna have remained undis­ America, and Chironectides in South Ameri­ turbed for an extremely long time. Under ca; but the great majority of the marsupials these circumstances of isolation, ancient is found only in Australia. They have devel­ forms of plant and animal life have been oped there, it is interesting to observe, in a preserved, while, during the long periods manner analogous to the development of since the isolation began, new species of . placental mammals of the other continents, plants and animals have developed from notably the carnivores, rodents, insectivores, . them as well. It must not be forgotten, how­ . and ungulates. ever, that immigration also has occurred, in­ The survival of the Monotremata and of troducing new species into Australia's plant the Marsupialia can be attributed to the fact and animal life since the beginning of its that, before man's appearance on the 'Aus­ geographic isolation. tralian continent, no other placental mam­ The many investigations into the animal . mals existed in Australia to prey upon them. life of Australia have given unequivocal evi­ There were only mice, which sometimes dence of the continent's isolation. Today the wandered on driftwood from island to island, most primitive mammals-the Monotremata and bats; but these were not significant .ene­ (Ornithodelphia) and the species of Echidna, mies. (It may be possible, of course,that the Proechidna, and Ornithorhynchus-appear mice and the bats first arrived in Australia in times subsequent to the Upper Cretaceous 1 Botanische Staatssamrnlung, Munich 38, Ger­ many. Manuscript received November 20, 1948. period and, therefore, subsequent to the be- [ 287] 288 PACIFIC SCIENCE, Vol. IV, October, 1950 ginning of Australia's geographic isolation.) tive, while others think that the Polycarpicae As far as the dingo is concerned, it is sup­ among the Choripetales are the most ancient. posed that aboriginal Austral inhabitants en­ Once, even certain of the Monocotyledones tering Australia from Malaysia were the (the Pand anales ) were considered for the first to bring this animal to the -southern distinction, although this claim, of course, continent. could not be proved by any significant argu­ It is also pertinent to our thesis to note ments. Inasmuch as this is hardly the place that zoologists believe that some of the opos­ for a discussion of the phylogenetic criteria sums (Caenolestes) migrated to North Amer­ by which plants are judged, the more perti­ ica from South America. Fossils of Marsu­ nent portions of my book, Neue Z iele der pialia have been found in Europe and in Botanik (1938), are suggested for reference. North and South America, an indication that If, in our search for the oldest angio­ they must have been distributed over vast sperms, and in our analysis of the Australian regions of the earth. The Marsupialia have flora, we hold the opinion that those families survived chiefly in Australia, and for this which are put at the head of the Monochla­ reason Australia today has the oldest and mydeae in the Engler and Prantl system of most primitive mammal types in the world. .classification are the ones which show the Now because, geologically speaking, mam­ most prim itive characteristics, we should be mals and angiosperms are of about the same supported in this assumption by the Casuari­ age, it is natural to ask if the oldest and most naceae found in Australia. In their original primitive of the flowering plants are also to distribution they extended from Sumatra to be found in Australia today. The answer to the Philippines, New Caledonia, and the Fiji this question would help us to determine the archipelago (Diels, 1926 ), and to Tahiti, the antiquity of the angiosperms. Austral Islands, and the Marquesas ( Brown, This is a question that is difficult to an­ 1935 ). As yet, however, there is no reliable swer,if only . because botanists are not in basis for the hypothesis that the Casuarina­ agreement on the most primitive species of ceae are more nearly related to the Gymno­ angiosperms. This one question asks other spermae than is any other family of the An­ questions : If the earliest species of angio­ giospermae. Neither can this supposition be sperms could be defined, would it be found proved for the Proteaceae and the Balanop­ that they exist preeminently or even exclu­ sidaceae, which are also placed at the begin­ sively in Australia? Or, if the earliest species ningof the Monochlamydeae in the Engler c annot be defined, is it possible to determine , and Prantl taxonomic system. In Australia from the Australian flora of today, which are more than half of all ofthe species of Mono­ the most primitive species of angiosperms chlamydeae are Proteaceae (a bout 600 .spe­ that have succeeded in persisting until this cies) , although the family has extended to time ? These are the problems to be investi- southern Africa, southern Asia, and South gated in this paper. America (Vester, 1940) . The species of Proteaceae are almost exclusively ligneous WHAT ARE THE MOST PRIMITIVE FAMILIES plants, which would indicate that, phyloge­ OF THE FLOWERING PLANTS? netically, they are rather an old group, but in the absence of paleontological evidence As every botanist knows, the question of we cannot be certain that these species are primitiveness in flowering plants is a contro­ really older than many others we might con­ versial one. Some think that certain species sider, so we have to be content only with of the Monochlamydeaeare the most primi- supposing thatthey might be. Antiquity of the Angiosperms-SUESSENGUTH 289 On the other hand, if we hold the opinion, ing plants? Or may those various plant spe­ as many botanists do, that the Polycarpicae cies now found in Australia have migrated are the most primitive of the Angiospermae, to the continent since its separation because we should find that only two very small fami­ they-or rather, their seeds-could cross the lies of Polycarpicae are endemic in Australia: ocean gap while the animals were not able the Eupomatiaceae and the Himantandraceae, to do so? which are related .to the 'Eupomatiaceae, but To obtain a general view of the whole which possess neither calyx nor corolla flora of Australia, let us consider thecata­ (Diels, 1919: 126 et seq.). The other fami­ logue of F. von Mueller, his Census of Aus­ lies of Polycarpicae are not well represented tralian Plants (1 889). Because of the recent in Australia : the Magnoliaceae afford only advances in our knowledge, Mueller's list is 4 species, the Annonaceae 18, the Nymphaea­ neither complete nor infallible, yet it is not ceae 5, the Ranunculaceae 17, and the Myris­ likely that the proportions of the numbers ticaceae 1. In view of these facts, it cannot of species within large groups and of the . be claimed that the Polycarpicae are the most endemic species have changed significantly primitive of angiosperms in Australia and since that time. Therefore we may use the that a comparison with the preservation of Census without hesitation , all the more neces­ the earliest mammals could be made. sarily because there does not exist a later cat­ This brief appraisal is enough to show that alogue for the whole Australian territory the angiosperms which most botanists con­ (including Tasmania but not Ne w Zealand ). sider to be the oldest of flowering plants Mueller's catalogue lists 8,842 species, (Polycarpicae) do not exist exclusively or pre­ and, because it does not mention those spe­ eminently in Australia. But in the informa­ cies introduced in recent times (since about tion we have learned about the Casuarinaceae 1800), it is well fitted for our purpose. Of and' the' Proteaceae we may have found cer­ these 8,842 species, 7,734 (that is, 87.5 per tain clues which will be of value later when cent) are endemic in the larger sense of the we investigate their degree of primitiveness. word- that is to say, they are found in Aus- - tralia itself but may also extend to New W H ICH SPECI ES OF ANG IOSPERM S Zealand and to parts of Polynesia as well.
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