THE DISTRIBUTION OF THE PROTEACEi®: PAST AND PRESENT

B y s . B. K a u sik , D.Sc. Department o f Botany, University o f Mysore Central College, Bangalore (With two maps and appendix) (Communicated by B. Sahni)

Received for publication on January 8, 1943

I ntroduction

T h e distribution of the Froteaceas at the present day in certain ■widely separated localities in the southern hemisphere is not only one of the most fascinating problems in geography, but is also such an intricate one that it has always evaded au explana­ tion which approaches finality. Similarly, much controversy also existed at one time with regard to the fossils ascribed to this family, and the opinions expressed by botanists were, therefore*, invariably conflicting. The fossil forms belonging to the Proteacese have been dealt with in the past by Unger, V. Bttingshausen, Hecr, Saporta and others, and also by several recent workers. Many of the earlier determinations, based on winged seeds and fruits, used to bo generally regarded as useless, for the view was often expressed that they prob­ ably belonged to entirely different families. For the same reason, several of the leaf impressions stated to belong to the Proteacea^ were also often viewed with a certain amount of scepticism, and very recently Gothari (1932) has also expressed doubts regarding the validity of the forms ascribed to this family. He remarks : “Es ist klar, dasa die Anwesenheit dieser Familie iiii europaischen Tertiar ein sehr wichtiges pflanzengeographisches Problem bietet; nach dem, was man Bonst von der Tertiarpflanzengemeinschaft weiss, ist dies nicht gerade wahrscheinlich.” However, Berry (1916), one of the leading American palseobotanists who has devoted a good deal of careful attention to this subject, has expressed that ‘‘though excep­ tion may justly be taken to some determinations of Unger, Ettings- liausen and Heer, these doubtful determinations in nowise affect the main body of facts ”, and has also further remarked that the “fossil forms are sufficient, after doubtful determinations are excluded, to show that the originated in the northern hemisphere, first appearing at the close of the Lower Cretaceous epoch and becoming practically cosmopolitan in the Upper Cretaceous time, when they reached the Australian region from south-eastern Asia.” Berry (1916) has reviewed the innumerable fossil forms of this family in a detailed account of the geological history of the family. These and also some additional forms which have since been described by several recent workers are mentioned in the following paper, and some interesting features in the distribution of the modern members are also pointed out. The paper, however, does not deal with any fresh or additional forms ; on the other hand, it is intended to give only a summary of our present knowledge of this subject.

The F ossil H istory of the PaoTEACEiE In Europe the fossil Proteaceje are represented by a large mimber of Cretaceous and Tertiary forms found abundantly in various localities (Map 1 and appendix). These forma belong to the following genera, of which some are also known by living types in the southern hemisphere at the present time, while the others are strictly extinct types:— and the allied generalized types Proteoides, Proteophyllum and Proteolepis, Genarrhencs, , , Eakea, Knightites, and the allied genus Loma- tites, Stenooarpus, , Petrophiloidfis,^ Leucodcndrites, Orevillea, and the closely related genus Emhothrites, Dryandra and the allied genus Bryandroides, and the related form Banksites, and lastly Boupala with the related genera Bho- palospermitee and Ehopalophyllum. In addition to these forms, some fresh ones have also been reported recently. Keid and Eeid (1915) have described the half of a hard fruit from the Plio(jene of Holland and state th at it is “so like that of , and so unlike anything else ” that it must be regarded as a new species of this genus. They further observe that this is perhaps the “ last representative of this dying out family in Europe ”. Principi (1919) has reported a species of Lomatia from the Oligocene of Italy, and very recently Velenovsky and Viniklar (1926-31) have recorded from the Cretaceous of Bohemia one species under each of the genera Dryandra, Conospermitea and Banksicarpns, and also a number of species under the generalized type Proteophyllum. They think that it is only probable that the forms referred to the genus Pro­ teophyllum belong to the Proteace®, but remark in the case of Banksicarpm, the determinations of wMch are based on inflores­ cences bearing round sessile fruits, that there are strong affinities with the well-known modem Australian genus Banksia. The fossil forms found in North America and which belong to the Cretaceous and Tertiary are represented by the genera

* Bowerbank, in his classic study of the fossils from the Isle of Sbeppey, established the genus PetrophUoides from its supposed resemblance to the modern ProCeaceous genus Petrophila. Several species were referred to this, but one of these was subsequently shown by Starkie Gardner (1879) to be an Alnus, while some of the other species were referred to Sequoia by Ettingshauscn. However, Ettings- hausen (1879) retained Onally three English Eocene forms after a careful study of these fossils (Berry, 1916). Even these must now be regarded as useless, for Reid and Chandler (1933) have recently shown that, in ^eir fruit characters, they belong unmistakably to the Juglandace®, except one s ^ ie s, P. oviformis, which they place in the Lauracea. Palceodendron, two species of which are also fouud in the Atane beds of Greenland ; Proteophyllum, established by Fontaine, but regarded by Berry (1.916) as not being a trustworthy type; Gonospermites, deter­ mined by Fontaine and again regarded as useless ; Lomatia, of which the Miocene forms preserved in the volcanic ash at Florissant, Colo­ rado, are stated to he extremely convincing (Knowlton, 1926); Fer- soonia, Ernbothriopsis, Embothrites, recorded from the Dakota sand­ stone, but regarded as doubtful ; Dryandroides, Bankaia, includ­ ing an Alaskan (f) Eocene species and three well-marked species in the Wilcox flora; Enightiophyllum, and a probable Banksia fruit referred to the form genus Oarpolithus. Eecently, a new Eocene (middle or upper) species of Banksia from south-eastern North America (Berry, 1924) and a Cretaceous species of Dryandroides from Colorado (Brown, 1933) have also been reported. In Berry’s (1916) original account there is no mention of any fossil form from South America, but three forms have lately been reported by him from this continent also. These are a pre-Miocene species of Embothrites from Argentina (Berry, 1932), a Miocene species of Lomatia from Patagonia (Berry, 1934), and a Miocene species oi Guevina from Northern Patagonia (Berry, 1934). I t may be pointed out here that Lomatia and Oucvina are found in South America at the present time also, and further it seems to be signi­ ficant that Guevina is peculiar only to this continent even to-day and is not found anywhere else in the other Protead centres. lu the continent of Australia, where we have the largest number of the Proteaceai living at present, there are, by contrast, only a very few fossil forms. Tlie Cretaceous forms of these fossils belong to the genera Proteoides, Gonosper miles, and Bhopalophyllum (V), and the Tertiary forms are an Oligocene species of , a few Oligocene (but more probably Miocene) species of Lomatia, one of which is also found in Tasmania, a Tertiary species of Per- soonia in New South Wales, an Oligocene species of Eoupala, and Oligocene or Miocene species of Dryandra, Dryandroides (in Tasmania) and Banksia. In the Antarctic continent three Oligocene species referred to the genera Knightia, Lomatia and Embothriophyllum are recorded from Grahamland. • Berry (1916) does not refer to any fossil Proteaceae in the continent of Asia, but a few forms are known from this region recently. Two of these are Cretaceous forms referred to the genera Proiecephyllum from Siberia (Kryshtofovieh, 1929) and Dryandra from south-eastern Trans-Caucasia (Palibin, 1930). With regard to the former, however, it has already been mentioned that it cannot be regarded as a trustworthy type. In addition, Kryshtofovieh (1929) has also referred some Tertiary fossils from Oligocene beds in Turkestan to the genus Dr^artdra; however, this determination has now been shown to be incorrect by Korovin (1932) on the evidence of additional material from the same locality showing axillary inflor- escenoes and thus bearing a strong affinity with the tribe GrevilleaB. 4o

Korovin has, therefore, established a new genus Palibinia for these forms. We may now refer here to certain interesting facts in the dis­ tribution of the fossils belonging to this family. We find that several of these fossil genera are represented as common types to two or more regions, a fact which is in sharp contrast with what is seen with regard to the modern members. These fossil types are Lomatia in Europe. N orth and South America, Australia and Antarctica; Emboihriie^ in Europe and North arid South America ; Dryandra in Europe, Australia and Trans-Caucasia; Dryandroides in Europe, North America and Australia; Banksia in Europe, North America and Australia; Knightia in Australia, and the related forms Enightitea and Enightiophyllum in Europe and North America respectively ; Gonospennites in Europe, North America and Austra­ lia ; Qrevillea in Europe and Australia; Proteoides in Europe and Australia ; and Roupala and Rhopalophyllum in Europe and Austra­ lia. On the other hand, the living forms to which these fossils are most closely related are found to-day in distinct and far-sepa­ rated regions where they occur as purely endemic types, all in the southern hemisphere. There is also another interesting fact regarding the past and present distribution of certain members of this family. It will be seen from the appendix at the end of this paper that there are thirty-six genera represented in the fossil record of this family, and of these, fifteen genera are also found to-day as living types. l?hese a re: Protea, Genarrhenes, Eelicia, Lambertia, Hakea, Lomatia, Steno. carpus, Persoonia, Qrevillea, Embothrium, Dryandra, Banksia, Roupala, Guevina and Knightia. The living types of these genera are found in isolated regions. Protea in Africa, Eclicia in south-eastern Asia and Australia, Lomatia, Embothriium and Roupala in South America and Australia, Guevina in South America, and the rest in Australia. In contrast with this distribution of the living types, the fossil forms referred to these genera are found mostly in Europe and a few in South America and Australia. Further, in the above summary of the Cretaceous and Tertiary fossils of the Proteacese, it is also clear that the majority of these forms belong to Europe, although ^everal are represented in North America and some in the other parts of the world (Map 1 and appendix). It is also apparent that it is in the northern hemsiphere that we have an abundance of these forms, while in the southern hemisphere the fossil record of this family is still very meagre ; probably this part of the earth’s surface has not yet furnish­ ed all its fossil material. Our present knowledge of the southern fossil forms is far from being complete, and the few that have come to our attention here are from such isolated regions as Australia, South America and the Antarctic continent. This wide occurrence of these fossils is in striking contrast with the peculiar distiibution of the members of this family at the present day in only certain localities in the sonthem hemisphere where they are mostly endemic (Map 2). Scott (1924), therefore, thinks that probably the few geuuine fossil ProteacesB are limited to Australia. On the other hand, Berry (1916) observes that the present distribution of certain genera, as Boupala with thirty-six species in tropical America, two in New Caledonia and one in Queensland, Bmboihrium with one species in Australia and four in South America ranging from Chile to the Strait of Magellan, and Lomaiia with three species in Chile, four in Australia and two in Tasmania, shows that “these genera must have ranged over intervening areas ”, “ unless one is prepared to subscribe to the doctrine of special creation for each continent or to the independent evolution on separate continents of different species of the same genus He, therefore, concludes that the Proteaceoe must have been practically cosmopolitan during the Upper Cretaceous time. Similarly, Beward (1933), another distinguished pateobotanist, remarks.: “The subject is too controversial. ... ; it must sufiBce to express the opinion that the family was represented in the novthern hemisphere during the Tertiary period.” From such opinions it is clear that the family must be of great antiquity. Probably it is already on the wane at the present time and some of its members even fail to set seed freely. It may be of interest to note here the following remarks by Bolus (1905)“ Few botanists who .... have spent many years in South Africa, and especially in the south-western districts, have not been ponetratod by a gloomy impression that the soi;th-west flora is dying and is doomed to extinction. In general, species of the Bruniace®, Penroace® and Proteace®, so peculiar to this region, have become .so much more rare.” Again, Brough (1933) observes that the “ infertility of the ProteaceiB in general has long been known, and is widely commented upon ”. He ramarks on Lawson’s (1930) observa­ tions that a high percentage of sterile pollen prevails iu numerous Protead forms, but concludes from a study of that, while pollen sterility is only a minor factor, the chief factors controlling seed setting are the visit of the pollinating agents and the weather conditions prevailing at the time of pollen exposure.

• T h e P r e s e n t D istribution o f t h e PROTEACEiE Wo may turn our attention now' to the distribution of the modern members belonging to tj^e Proteacese. Some of the facts here are interesting and they are dealt with briefly in the following pages. According to Erigler and Gilg (1912), and Engler and Diels (1936), the family includes 50 genera and 1,100 species ; of these, 720 are in Australia, 262 in Africa, 27 in New Caledonia, 25 in Bast Asia and 36 in tropical South America. Thus the family belongs character­ istically to the southern hemisphere, although some of the members do extend slightly north of the equator (Map 2). The latter are represented by some species belongii^ to the genera Protao and Leuoospermum in Africa, Helieia in Asia, and Soupdla and Ewplatsa in South America. The accompanying sketch map of the world (Map 2) shows the approximate areas of distribution of these a>nd some of the oth«r important genera of the Proteacese.

In the distribution of the modem members it is seen that Australia forms the largest centre at the present time. It includes nearly three-quarter the total number of species which belong to about thirty-five genera, and these are “ massed chiefly in Western Australia ”, where, it is stated, there are nearly twice as many species as in all the other states of the continent (Brough, 1933). A number of these Australian forms are also found in Tasmania and in the more distant islands of New Caledonia and New Zealand, while there is only a single member of the Proteaoese, namely, a species belong­ ing to Banksia in New Guinea. Africa ranks next in the number of forms which it includes. There are altogether 15 genera and a little less than a quarter the total number of the species in this family. Probably the proportion of the African species to the total number is slightly greater than this, for according to the Flora Gapcnsis (Thisclton-Dyer, 1912), there are here in all about 330 species, including some imperfectly known ones, while it is only 262 as given by Engler and Biels (1936). There seems also to be some discrepancy even with regard to the number of genera represented in Afric.a; only 12 are mentioned by Engler and Prantl (1894), whereas in the Flora Gapensis the number is 14, even after excluding the genus which is confined entirely to Madagascar and is not mentioned in the latter work. The majority of the African Proteacese are found in the Cape district where they are said to form a characteristic component of the vegetation. Two of those forms, namely Protea and Leuaospermum, have rather a wide range, some of their species being found above the equator in tropical Africa and on the mountains of Abyssinia. A genus which is not represented in the Cape Province, but which shows a discontinuous distribution elsewhere in Angola, Natal, the Zambesiland and Madagascar is . Its peculiar distribution is stated to be due to the dispersal of its fruits by birds (Engler, 1894). Lastly, Dilobeia is another interesting African type and this is found, as already mentioned, only in Madagascar. In South America the distribution of certain genera appears again to bo noteworthy. There are 7 genera here and their species are scattered over a wide area throughout the length of the continent. Two of these genera extend slightly north of the equator, being represented in Guiana, and Roupala even beyond in southern Central America. Further, the genus Roupala, the majority of the species of -which are found in various parts of South America, is also represented in the Australian region, with one species in Queensland and two in New Caledonia. Similarly, the genera , Emhothrium and Lomatia are represented both in South America and Australia. Berry (1916) remarks that “ during their geologic history these genera must have ranged over intervening areas ” ; the evidence from the fossils seems to favour this conclusion. Of the remaining genera which are all confined to South America, Quevina is an interesting one, for it is now known that a fossil species which is referred to this genus has been found in Patagonia (Berry, 1934). In south-eastern Asia the Proteacese are represented by a number of species belonging to the genus Helicia. This is characteristically an Indo-Malayan genus and its several species are found scattered over a wide area including various parts of India, Ceylon, Cochin- China, Sumatrg,, Java, Malacca, the Philippines, and also a few in Australia.* A point of especial interest h(;re is that one of the species, H. lancijolia, occurs in southern Japan which is to-day the most northern limit of the Proteacese.

General Conclusions After having known the fossil history and development of the Proteaceaj, it would be useful to consider here if any light is thrown by the study of this group of on the distribution of continents and oceans in former geological times. At present there are two views regarding the formation of our modern continents, and these are (a) the older theory of permanence of continents and oceans, and (h) Wegener’s hypothesis of continental drift. According to the former, it is understood that the present continental areas liave always remained as«such, but that vast portions of land masses which at various times in the past formed connecting bridges between them, have now disappeared beneath the oceans. On the other hand, accord­ ing to the hypothesis of drift it is supposed that there existed during pre-Cretaceous times an immense universal land, the Pangwa, from which later the present continents were formed by fragmentation and drifting away. It is not our purpose here to enter into a detailed discussion of this subject; it is too very controversial and is such an intricate and many-sided problem that it cannot be adequately dealt with within the scope of the present study, which refers to b u t. one group of plants. However, we may briefly note hc'.re ct^rtain points which appear to be of some interest. EeoentJy, in a comprehensive account of the distribution of several families and genera of the flowering plants, Irmscher (1922) has pointed out that the Proteacese show a two-phase development and a great discontinuity in the distribution of the modern membera. He finds that there are in this distribution four well-defined meridional groups, the so-called areas, which, he concludes, must be frag­ ments of an original single unit which later separated away and became vastly isolated consequent on the drifting of large continental areas. On the other hand, Diels (1928) strongly objects to this treatment of vegetation according to meridional arrangement, and emphasizes the fact that everywhere on the earth’s surface the units of vegetation succeed one another in strictly climatic zones extending from the polar to the equatorial regions. Diels makes a brief survey of this characteristic feature in plant distribution in general and finds that the theory of continental drift fails to explain some of the peculiarities in distribution. • In the sketch-map of the world given by Berry (1916) to show the distribution of the Proteaccae, the range of Helicia does not include southern Japan, Ceylon and parts of South India (Travancore and the Nilgiris); however, according to Hooker, Engler, Gamble and others, some species of this genus occur in these regions also. Similarly, Eeid and Chandler (1933) have also been forced to regard that Wegener’s hypothesis leads to the following anomalous conclusions with regard to the Tertiary floras of Europe : “(1) that although America was, by hypothesis, the nearest neighbour to West Europe, and could have conserved the European Tertiary relies, yet it is in the Far East, 120° of latitude away, that these are best preserved, and (2) that as, by hypothesis again, the separation between America and Europe became greater, so, in actuality, did the alliance between their floras become closer. The liypothesis does not agree with the facts.” Again, Berry (1928) objects strongly to the hypothesis of conti­ nental drift and even questions altogether the validity of the argu­ ments. Ho remarks that tlie hypothesis “ rests ou an unscientific principle, the facts of geophysics do not support it, it fails in expla­ nation of geological climates and paJffiontologically it raises more distributional problems than it solves.” In his o])iniou (in the words of van der Gracht, 1928), “ the geographical pattern could not possibly be preserved in comparatively thin masses of continental dimensions drifting along over such distanses.” We may also refer here to the opinion held by Sahni (1930) re­ garding the question of continental drift in general. He discusses the peculiar relationships of the two contemporaneous, but climato- logically very unlike floras, namely the 6lo8soptcris-&or», of Indo- Australia and the GigantopUris-^or& of sTno-Sumatra and remarks : “We have not as yet enough palteobotanical data to prove the drifting apart of the different portions of Gondwanaland. B ut we at least seem compelled to agree that drift movements of large • magnitude elsewhere have brought into juxtaposition continents once separated by wide oceans. It therefore seems impossible to get away altogether from the idea of continental drift. But the details of Wegener’s theory must stand on their individual merits.” From bhe point of view of the past and present distribution of the Proteaceee, it appears that there are considerable diflSculties in fitting this liypothesis with the known facts. If, as held by Wegener and others, the continents had lain in juxtaposition for a long time until the Cretaceous, it is reasonable to assume that tliere should have been to-day a much more uniform distribution of the members of this family, perhaps also with some at least of the genera occurring as common types between two or more of the Protead centres. On the other Jiand, many of the forms are so very endemic. If North America had long been in close contact with Europe, then there seems to be no reason why many of the European Cretaceous and Tertiary forms could not have migrated to that continent and a larger proportion of the Proteacese conserved to-day in South America, as the migration of forms could not have been difBcult here; the north-south range of the mountain chains in the American continents could have hardly been an obstacle for such migration. Actually, we have in South America only a very small percentage of the Proteacese, whereas it is in Australia that we see to-day the majority of the forma, and for the nearest fossil representatives of these we have to turn to the numerous European forms. Further, there are to-day some genera in South America, and these are also represented ill Australia ; it appears that those must have had a common origin and tho.t their species spread subsequently to the two continents, probably by way of land bridges which liave now disappeared. Again, in the case of the African Proteace», at. least some of tlie forms must have made their way to this region from somewhere in the northern hemisphere, probably from Europe, at a time when the east-wefit mountain barriers that we see to-dajiarere not liigli enough to prevent their southward migration. At the present time the African forms are mostly coulined to South Africa wJiere they form a higlily insular flora with not even a single genua occurring in the other Protead centres, for no more opportunities were available to them subsequently to migrate to other regions and spread rapidly over wider areas. Lastly, the distribution of the different species of Hclicia seems also to be rather significant. As already pointed out, these species are scattered over a very wide area, and there appear to be two possibilities for this ; either the species have acquired this wide distribution only in recent times, perhaps even since late Tertiary times, or the different species are persisting outliers of an old stock and are to-day like sigii-posts that serve to mark the southward route that the ancestors of the modern Proteaceas took when they migrated from some loc,ality in the northern hemisphere. It appears that the latter is the more probable, especially as wo know that this family is of great antiquity, not only as revealed from the fossil record, but also from tlie fact tliat many members seem to be already on the wane at the present time {cf. Bolus, 1905 ; Brough, 1933). Tn this connection it may be noted here that Brough (1933) observes that, since Australia and Africa have remained separate since the Carboniferous period, and “ Australia is considered to have been joined with the Asiatic mainland at least during the Upper Cretaceous period, but probably a complete Bepa.ration has existed since the beginning oC the Eocene,” the ancestors of the ProteacesB must have “ migrated southwards from an Asiatic centre of distribution into Australasia nnd Africa, as then constituted, during later Cretaceous times - It appears, therefore, that the hypothesis of continental drift does not help us any more than l;he older, but less fascinating theory of the permanence of continents and oceans in explaining some of the I)eculiarities of the past and present distribution of the Proteaceee. The opinions on this subject are so very conflicting that, with our present knowledge of this subject, the facts that we now have seem to be inadequate to discuss further Wegener’s hypothesis in relation to the present problem. We do not as yet have a full record of the fossils in the southern hemispehre, and what we know of them is so pre­ cious little, that it seems that we should wait until more orideace is available. Probably, for a really satisfactory explanation, we have eventually to turn our attention to regions in the southern hemi­ sphere, at present unexplored, or only incompletely studied, but which must yield their fossil record. Per the present, therefore, it must Bufflce to say that the drifting apart of the continents does not appear to bear well with all the known facts. In conclusion, the present discussion may be summed up here in the following words of Berry (1916): “ The Proteacea? originated in the Northern Hemisphere, first appearing at the close of the Lower Cretaceous time and becoming practically cosmopolitan in the Upper Cretaceous time, when they reached the Australian region from south-eastern Asia. New Zealand must have been already segregated but not the land mass now represented by New Caledonia. During the early half of the Tertiary period Africa and southern Europe were essentially a single floral province, and in the Western Hemisphere the Proteaceffi ranged from the United States through South Amcrica and an unknown distance across tlie Antarctic Continent. Concomitant with the continent building and the consequent climatic c’ anges of the Miocene epoch the area of distribution commenced t hat shrinking which culminated during the Pleistocene epoch, leaving the stranded remnants in the present widely separated localities of the Southern Hemisphere. Not all the modern genera tooJc part in these migra­ tions, since the local peculiarities of poor soil and rigorous climate, combined with relative freedom from outside competition, were the factors that stimulated the evolution of forms in Australia during the Tertiary period...... ’.

S u m m a r y This paper contains an account of the numerous Cretaceous and Tertiary fossils belonging to the Proteacece, and also deals about the geographical distribution of the modern members belonging to this family. The bearing of the facts of distribution on Wegener’s hypothesis of continental drift is discussed in the concluding paragraphs. The hypothesis does not appear to help us much in explaining some of the peculiarities noted in this distribution. Finally, it is pointed out that we have to wait until more is known of the fossils of this family, especially in the southern hemi­ sphere which, perhaps, holds the key J!or a satisiactory explanation of the problem in question. ^ A cknowledgments It is my pleasant duty to acknowledge here the many kind courte­ sies and valuable advice extended to me by Prof. E. W. Berry, of the Johns Hopkins University, Baltimore, U.S.A., and Prof. B. Sahni, of the University of Lucknow, India, in the preparation of this paper. I wish to express my gratitude and sincere thanks to Prof. Berry for kindly reading through the account of the past and present distri­ bution of the Proteace®, and to Prof. Sahni for discussing with me the several points relating to the hypothesis of continental drift and for helping me with some relevant literature. Further, I wish also to mention appreciation of the kindness of Prof. M. A. Sampathkumaran, University of Mysore, for the constant encouragement and many useful suggestions which he gave me in the course of this work.

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R R LITERATURE CITED

Berry, E. W. (1916) “ Tlie Lower Eocene Floras of South-eastern North America,” U. S. GeoL Siirv., Prof. Paper, 91, 1 -353. ------(1924) “ The Middle and Upper Eocene Floras of South­ eastern North America,” Ibid., 92, 1-206 {Biol. Abst., 1925, 14, 1, entry No. 555). ------(1928) Theory o f Coiiliiicntal D rift; A Symposium, Amcr. Assocn. Petrol. Geol. ------(1932) “ Fossil Plants from Chubut Territory collected by the Scarritt Patagonian Expedition,” Amer. Mui. Novitates, 536, 1-10. ------(1934) “ Miocene Patagonia,” Proc. Nat. Acad. Sc., 20, 5, 280-82. Bolus, H. (1905) “ Floral Regions of South Africa,” Science in South Africa, p. 159. Brough, P. (1933) “ The Life-history of Grevillea rohusta Cunn.,” Proc. Unit. Soc. N.S.fV., 58, 33-73. Brown, R. W. (1933) “ Fossil Plants from the Aspen Shale of South­ western Wyoming,” Proc. U.S. Nat. Miis., 82, 12, 1-10 {Biol. Abut., 1934, 8, 1, entry No. 1874). Diels, L. (1928) “ Kontinentalverschiebung und Pflanzengeographie,” Ber. d. Deutsch. Bot. Cesellsch., 46, 49-58. Engler, A., and Syllabus der Pflaiizenfamilien, Berlin. Diels, L. (1936) ■ and Gilg, R. (1912) Ibid. and Prantl, K. (1894) Die naiiirlichen Pflanzenfamilien, 3, 1 Hiilftc, Berlin. Gotahn, W. (1932) PalUobotanik ; Handwiirterhuch der Natur- wissenschaften, Zweite Auflage, Bd. 7, 562-608 (Proteaceae, 601-02). Irmscher, E. (1922) “ Pflanzenverbreitung und Entwicklung der IConti- nente,” Mitt. Instit. attg. Bot. Hamburg, 5, 17-235. Knowlton, F, H. (1927) Plants o f the Past, Princeton Univ. Press. Korovin, E. P. (1932) “ Noryi tretichnyi tip semeistva Proteacese is Srednei Azii tOenus novum fossile palso- genum families Proteacearum),” Journ. Bot. U.K. S. S., 17, 5/6, 506-22. Kryshtofovich, A. (1929) “ The oldest angiosperms in the Cretaceous of Asia and other contributions to the Mesozxjic flora of the maritime province of Siberia,” Amer. Journ. Sc., 18, 519-25. Lawson, A. A. (1930) “ Endemism in the Angiosperm Flora of Australia,” Proc. Linn. Soc. N.S.1V., 55, 4 (cited from Brough, P.. 1933). Palihin, !. V. (1930) “ Verklme-melovaia flora mgovostoka Zakav- kazia (The Upper Cretaceous Flora of South­ East Trans-Caucasia),” V .S .S .K . Bull. Geol. ami Prosp. Surv., 49, 7, 125-34 (Biol. Abst., 1935, 9, 7, entry No. 14664). Principi, P. 1916, (1919) “ Le Dicotiledoni fossil) del giacemento oligo- cenico di Santa Giustina e Sassello in Liguria,” Mem. Desc. Carta. Ceol. d'ltal., 6, 294 (Ibid., 1920, 5, 1, entry No. 717), Reid, C., and Reid, E. M. “ The Pliocene Floras of the Dutch-Prussian (1915) Border,” Meded. Rijks. van Deljstojj., 6, 1-178. Reid, E. M., and The London Clay Flora, British Mus. Nat. Hist., Chandler, M. E. J. (1933) london. Sahni, B. (1936) “ Wegener’s Theory of Continental Drift in the Light of Pala:obotanical Evidence,” Journ. Ind. Bot. Soc., 15, 319-32. Scott, D. H. (1924) Extinct Plants and Problems o f Evolution, London. Seward, A. C. (1933) Plant Life through the Ages, Cambridge. Thiselton-Dycr, W. (I9I2) Flora Capensis, 5, 1. van dcr Gracht, W. A. J. M. Theory o f Continental Drift : A Symposium," van Waterschoot (1928) Amcr. Assocn. Petrol. Gcol. Vclenovsky, J., and ‘‘ Flora Crctacea Bohemia;,” Rorpr. Stat. Geol. Viniklar, L. (1926-31) V.itava Ceskoil. Repttb., Parts 1-4.