SOME REMAINS FROM THE TERTIARY OF HUNGARY

KLARA RASKY Budapest, 1., Lovas ut 23, Hungary

ABSTRACT DESCRIPTION

The plant remains from the Upper Eocene Agrostistachyophyllmn tomharrisi gen. et deposits of Budapest-6buda consist of the following sp. nov. new genera of : Agrostistachyophyl­ lum tOtrlharrisi, A lchorneaephytlum chandteri, PI. 1, Fig. 1 Baliosperrnaphytlum kraeuseti and Codiaeophytlum palaeovariegatum. A new combination - Alchorneae­ Diagnosis gen. et sp.- Leaf oblong-lan­ phyllum grambasli (Rasky) - is proposed for the ceolate. Apex missing. Measurable length leaf remains described previously under the 12·0 cm., maximum width 2·7 cm. Base name Sloaneaephyllum grambasti Rasky (1962). narrowly decurrent. Margin finely ser­ Carpolithus alchorneaefarmis sp. nov. is also des­ cribed from the same locality. From the younger rate except at the base, with apically Tertiary tuff deposits at Ipolytarn6c, North directed glandular teeth. Preserved petiole Hungary, is identified Omphateaephytlum weylandi 3·0 mm. .Midrib prominent, secondaries gen. et sp. nov. 14-15 pairs, subparallel, opposite below, These fossil remains, genera and species show close resemblance with the Jiving members of becoming subopposite and alternate above, group of the Euphorbiaceae (emend. curved 51ightly upward. Subsecondaries Pax & Hoffmann, 1931). branched on the under sides rarely. Inter­ secondaries diverging from the midrib not frequently. Tertiary venation connecting secondaries subvertically. Texture coria­ INTRQDUCTION ceous. Remarks and comparison - The fossil leaf HE genera and species described in remains can be compared with those of the the present paper from the Upper modern Agrostistachys massoana Vidal T Eocene marl formation had been (=Agrostistachys indica Dalz. var. massoana found in the locality of the former j agyba­ (VieJ.) Pax et Hoffm.) The mentioned living tony-Ujlak brickyard at Budapest-Obuda. species furnishes the most satisfactory match From the same locality the author had with the fossil leaves. The most similar published already (RASI

~6! RASKY - SOlliE PLANT RElI1AINS FRONI THE TERTIARY OF HUNGARY 265-

8·5 cm., broader above the l)1iddle. The partment of Botany, Hungarian Natural preserved petiole very stout, length 3·0 History Museum, Budapest. mm. Margin crenate-dentate, except near the base. Midrib stout, secondaries about Alchorneaephyllum grambasti (Risky) 6-7 pairs, curved upward; two basal pairs comb. nov. -of secondaries opposite, curved upward PI. 1. Fig. 2 parallel to the lower lateral margin; other secondaries diverging from the midrib, 1962-S1oaneaephyllum grambasti Risky­ alternate or subopposite, and curved towards Ann. Hist. Nat. Hung. 54: 36, PI. 3, Fig. 1. leaf margin. Secondaries subcamptodrome Description - Leaves broadly ovate in and craspedodrome. From the two basal general outline. Maximum width above at secondaries subsecondaries arising from the the middle. 9-10 cm, length about 13 cm. under side and curving upward in broad Shortly acuminate at the apex, more or less and regular subcamptodrome arches. Ter­ truncate at the base. Petiole not preserved. tiaries percurrent and approximately trans­ Margin serrate, with remote and irregularly verse between the midrib and the secon­ spaced larger and smaller teeth. Midrib daries. Irregular quadrate meshes between stouter than the laterals. The basal secon­ the tertiaries. Texture coriaceous. daries, one on each side, diverging from the Remarks and comparison - The leaf of midrib, ascending about one half or one Alchorneaephyllum chandleri can be asso­ third length of the lamina, and camptod­ ciated with the leaves of the recent Alchornea romely joining with the next secondaries. jioribunda Mull. Arg. species. Alchornea Subsecondaries arching from the other side jioribunda Mull. Arg. ranges nov,' in tropical of the basal secondaries, thinner and sub­ West Africa (Cameroon, Gaboon, Congo), camptodrome. Further secondaries diverg­ sometimes semi-climbing shrub or small tree. ing somewhat il regular spaced from the The fossil leaf remains are somewhat midrib and along them arching subsecond­ similar to the leaves of the living Mallotus aries in subcamptodrome or craspedodrome dispar (BI.) Mull. Arg. from Java, or to manner. Secondary veins on upper half cf Mallotus auriculatus Merr. from Mindanao, the leaf might also be craspeoodrome. The but differ morphologically in the leaf base tertiary venation comprising a series of ami in the secondary venation. These approximately transverse veins between the fossil leaves are somewhat similar to the midrib and secondarie" Texture coria­ leaves of the living Glyphostylus laoticus ceous. Gagnep. from Laos, to Grossera major Pax Remarks and comparison - After a de­ from Cameroon, to A ntidesma laciniatum tailed study and comparison with recent .Mull. Arg. val'. genuinuln Pax et Hoffm. herbal' material I can see no reason for from the Phillippines and also to the leaves doubting that the presen t fossil specimens {)f Hugonia holtzii (tropical Hugon iaceae), represent the leaf forms of Alchornea l:ricu­ but in the configuration of the secondary ra na Casar and its var. genuina Pax et venation, den tation or en tire margin, espe­ Hoffm. Therefore I have changed the name cially the different leaf bases all differ from Sloaneaephyllum gramhasti Risky to Alchnr­ the fossil leaf remains' of the Alchorneae­ neaephyltum grambasti (Risky) comb. nov. phyllum chandleri. Alchornea iricurana Casar. has polymorphic The fossil leaf remains of Mallotus riparius leaves, the juvenile-, normal- and cld-leaf l\IacGini tie, described and figured by Mac­ forms are different. The fossil leaf form Ginitie (1941) and by Becker (1960), com­ from Hungary, figured in the present paper, pared with the living Mallotus japonicus is a transitional form between the young Mull. Arg. from southeastern Asia, cannot and adult leaf forms. Alchornea iricurana be related to the leaf form of Alchorneae<­ is a tree at present. living in tropical South phyllum chandleri. The fossil leaf form of America: in Brasil, Paraguay and Bolivia. Parrotl:a cuneata (Newberry) Berry (1930) Occasionally the large genus of Alchornea from the Eocene Wilcox groups is not iden­ has several species which exhibit similarity tical with our specimens. of leaf characters and are something like Locality - Budapest-Obuda, marl depo­ our fossil leaves, a~ for example the Cen tral sits, Upper Eocene. American (Portorico) species Alchornea lati­ Generotype - PI. 1, Fig. 3.- CoIl. No. folia Swartz or the tropical African (Congo, -65.28.1.- Palaeobotanical Collection, De- Cameroon) species Alchomea laxiflora 266 THE PALAEOBOTANIST

(Benth.) Pax et Hoffm., but the resem­ Locality - Budapest-Obuda, marl depo- blance is not at all close. sits, Upper Eccene. T Locality - Budapest-Obuda, marl depo­ Generotvpe - PI. 2, Fig. 4.- Coil. No. sits, Upper Eocene. 65.30. I.--=- and the counterpart PI. 2, Fig. 5. Paratype - PI. 1, Fig. 2.- Col.!. No. 62. - ColI. No. 65.31.1.- Palaeobotanical Col­ 914.1.- Palaeobotanical collectIOn, De­ leC[ion, Department of Botany, Hungarian. partment of Botany, Hungarian Natural Natural History Museum, Budapest. HistOlY Museum, Budapest. Cf)diaeophyllum palaeovariegatum gen. et Baliospermophyllum kraeuseli gen. et sp. nov. sp. nov. PI. 2. Figs. 4-5 PI. 3. Figs. 9-11 Diagnosis gen. et sp.- Leaves oblong­ Diagnosis gen. et sp.- Leaves oblong­ ovate and linear-Ianceolate. Exceedmgly ellipsoidal to broadly-oblong.. About .13·5 variable in size. Imperfectly preserved. cm. in lenath by 7·0 cm. m maximum width. Measurable length of the lanceolate leaf 9S Narrowed below and terminating in a curved em., width 2·5-3·0 em. Measurable.length truncate or sClltate base. Tip of the apical of the ovate leaf is 10 em. and the maXlUmum portion of the leaf missing, but on the coun­ width 4·0 em. Petiole not presnved. The terpart apiculate. Margm coarsely crenate­ apex of the oblo~g-?vate leaf acuminate. dentate except near the base. VenatIOn Margin entire. Mldnb very. stout, secon­ pinnate, but 3-nerved at the scutate base. daries !>preading from the mldnb more or l\Iidrib stout espeCIally below. Secondanes less at right angles. parallel, co~sPlcuollsly irregularly spaced and dispose?, subparallel, anastomosing before the margm. Subse­ ascending, 6 to 7 on each sIde, alternate. condaries also spreading at right angles from The lowest pair of secondanes thmner and the midrib. Midrib and secondaries con­ opposite, curved upward parallel to t.he nected by short veinlets producing a s:y~tem lower lateral margins. Subsecondanes of elongate or irregularly areolate reticula­ branching from the outer side of the secon­ tion. Texture coriaceous. daries, becoming subcamptodrome and cras­ Remarks and comparison - The leaf re­ pedodrome and terminating. in one of ~he mains found in the Upper Eocene marl for­ marginal dentations. TertIary v~natlOn mation in Budapest-Obuda are very. well thin and comprising series of approximately identifiable with those of the recen t Codtaeum transverse veins between the mldnb, secon­ variegatum (L.) BI. Codiaemn varief5,atum is daries and subsecondaries, forked and anas­ a shrub or small tree of coastal areas III Java, tomosing, their enclosed areas interwov.en at 5-1500 m. The very variable leaf forms by very fine reticulation. Texture cona­ are characterized principally by the vena­ ceous. tion. The small-leaved recent herbanum Remarks and comparison - The fossil specimens, serving for comparison, origin­ leaves can be best compared to those of the ate from Luzon (PL. 3, FIG. 8), and the recent montanum (Willd.) largeleaved one is from New Guinea (PI.. 3, Mull. Arg. This recent shrub is very fre­ FIG. 12). quent in the forests of the tropical Hlm~laya, Locality - Budapest-Obuda, marl depo­ in the forests of Java and India, and III the sits Upper Eocene. monsoon areas. Generotype - PI. 3, Fig. 9.- Coil. No. The marginal dentation in our fossil leaves 65.32.1.- and the counterpart PI. 3, Fig. 10. appear to be more uniformly or bluntly - Coil. No. 65.33.1. dentate and the base more distinctly trun­ Paratype - PI. 3, Fig. 11.- ~oll. No. 65. cate or scutate, than in any of the Euphcr­ 34.1.- Palaeobotanical CollectIOn, Depart­ biace~l1ls species found in the Upper Eocene ment of Botany, Hungarian Natural History deposits in Hungary. Museum, Budapest. These types of fossil leaves are no~ ra:e in the Tertiary and Cretaceous deposits m Omphaleaephyllum weylandi gen. et sp. nov. Alaska. Acer inaequale Heer and Vt~urnum PI. 2, Fig. 6 aequale Hollick (1936) from the Tertl.ary of Alaska are very much like our fossil l~af Diagnosis gen. et sp.- Leaves obovate, remains, but in details differences eXist broadest near the apex. Gradually and between them. smoothly decurrent at the base.. Petiol~ RASKY - SOME PLANT REMAINS FROM THE TERTIARY OF HUNGARY 267

not preserved. Margin entire. Midrib fr~it is not determinable with absolute •. thick. Secondaries stoute widely spaced certainty. This species is represented by spreading at right angles from the midrib only the figured specimen and its counter­ and directed towards leaf margin, connected part. I am reasonably satisfied that it inside the margin by prominent arches, represen ts one Alchortua species. The camptodrome. Tertiaries •thin, mostly fruit of the recent Alchornea castaneifolia poorly visible, thus rendering the details (Willd.) Juss. from South America (Orinoco, oh'\r.11re. Texture coriaceous. Amazonas), can be compared with the fossil Remarks and comparison - These leaf imprints. Fruib of similar habit charac" impressions are similar to those of the living terize other genera, especially in the family Omphalea biglandulosa (Pers.) Baill. species Flacourtiaceae. Other similar modern fnlit from Madagascar. These are shrubs, rarely is Distylium stellare O.K. from Malaysia of trees, but not rarely climbers. The genus the family Hamamelidaceae. Omphalea is distributed now in Central Locality -- Budapest-Obuda, marl depo­ and South America (Brasil, Peru), in south­ sits, Upper E0cene. eastern Asia (Philippines, Borneo), in Aus­ Holotype - Pl. 2, Fig. 7.- Coil. No. 63. I~ _~ tralia and Madagascar. 1026.1. and the counterpart. Palaeobota­ Othtrwi~e the leaf remains of Ompha­ nical Collection, Department of Botany, leaephyllum weylandi resemhle with those Hungarian Natural History Museum, of the recent Tetraplandra ridelii Mull. Arg. Budapest. from Brasil and West Indies, but the leaf base of Tetraplandra ridelii deviates from DISCUSSION our fossil leaves. Fossil leaf remains of Omphalea patagonica The Euphorbiaceae is one of the largest Berry (1938, p. 85, PL. 25, FIG. 1) from existing family of trees, shrubs and herbs. Argentina and from Chile (ENGELHARDT, The trees are widely distributed in various 1891, p. 672, PL. 9, FIG. 1), have already parts of the Tropics. been described, which may be compared A considerably but relatively imigni­ with the existing form of the tropical Ame­ ficant number are recorded of the Euphor­ rican Omphalea diandra L. But these South biaceae as fossil remains from the Upper American fossil leaf remains are readily Creataceous and the Tertiary deposi ts. distinguishable from those of Omphaleae­ These fossil·remains consist of leaves, fruits, phyllum weylandi. They are present in the flowers and woods. Fossil representatives Tertiary tuff deposits of Ipolytarnoc. Om­ of the Euphorbiaceae were recorded from phaleaephyllum weylandi is not abundant Europe, North and South Africa, India, in the tuff deposits of lpolytarnoc, hut it Japan and North and South America. is a VfTy characteristic impression in the Though difference of opinion regarding fossil flora. the determination of some fossil records is Locality - I polytarnoc, North Hungary, justifiable, I regard - for exampIe - the tuff deposits, Tertiary. leaf remain or flex gigas Engelhardt (1885, Generotype - PI. 2, Fig. 6.- Coil. No. 65. p. 357, PL. 23, FIG. 4) from the Jesuiten­ 6.1.- Pala~obotanical Collection, Depart­ graben near Kundratitz (Bohemia) from the ment of Botany, Hungarian Natural History LOWer Miocene, as Euphorbiaceae, which Museum, Budapest. is very sim dar, and almost identical, with the leave" of th~ existing East Australian Carpolithus alcharneaeformis sp. nov. shrub Caelebogyne ilicifolia J. Smith. (The PI. 2, Fig. 7 leaves of the recent Caelebogyne ilicifolia J. Smith are also som~what similar to those Diagnosis - Fruit elliptic in outline, 11 of somo fossil Qu~rcus cruciata A. Br.) On mm. in length by 8·0 mm. in maximum width the other hand, the leaf remain of Sapindus midway betwee!1 the apzx and the base. lin~aYlI)lius Bony (1930, p. 101, PL. 14, Apex cuspidate. Measurable and curved FIG. 19) from the Wilcox flora (Hollv peduncle 8·0 mm. in length. Fruit-lobes Spring,). is a bilam[nate leaf, like that of longitudinally grooved. Texture apparently certain rec':n t species of Euphorbiaceae. coriaceous. A' closdy related modern species Codiaeum Remarks and comparison - The botanical variegatum (L.) Blume f. appendiculatum affinity of this characteristic and interesting Celak also furnishes excellent comparison THE PALAEOBOTANIST with these fossil leaf remains. The genus near the sea, and also indicate perhaps a Codiaeum is distributed now in Malaysia tropical-subtropical rainforest, with various and in the Island of the South Sea. altitudinal zones in the Upper Eocene time 1. From the Upper Eocene of the envi­ of Budap~t-Obuda, not unlike to those, in rons of Budapest-Obuda presented the fossil which their modern alliances in the recent leaf remains of Baloghiaephyllum miocenicum. vegetation live· today. It is n9t possible This fossil sr~cies indicates a close relation­ to reconstruct exactly the fossil plant com­ ship with the Jiving Baloghia lucida from munities from this fossil flora and to com­ New Cal<>donia. Agrostistachyophyllum tom­ pare exactly with living plant communities. harrisi, Codiaeophyllum palaeovariegatum and 2. The leaf remains of Omphaleaephyllum Baliospermophyllum kraeuseli have been weylandi from the younger Tertiary tuff comparerl wilh the existing southeastern deposits at Ipolytarn6c, North Hungary, Asiatic species: Agrostistachys massoana, and the leaf imprints of Centroplacophyl­ Codiaeum variegatum and Baliospermum lum palaeoglaucinum from the same locality montanum. Alchornea floribunda the most (RASKY, in Press), are tropical in their similar living species of A lchorneaephyllum modern affinities ancl suggest the possibility chandleri, is a member of the West African that their ancesters were also tropical in tropical forests. On the other hand, Alchor­ character. neaephyllum grambasti shows close relation­ ship with the Jiving species Alchornea iricu­ ACKNOWLEDGEMENTS rana in Brasil. M acarangaephyllum palaeomonandrum and The author wishes to express his sincere JVI allotophyllu.m palaeomiquelianum are appreciation to Mrs. Dr. Savitri Sahni, described from the Upper Eocene of Buda­ the President at the Birbal Sahni Institute pest-Obuda too (Risky, in Press). of Palaeobotany and the Palaeobotanical The occurrence cf these genera and Society Lucknow, for the invitation to con­ species of the Euphorbiaceae in the fossil tribute thig paper, and to Professor Dr. K. flora, com pared with their living alliances R. Surange, Director at the Birbal Sahni and their distribution, indicate a moist Institute of Palaeobotany, Lucknow, for and warm (tropical) climate of costal areas the rea~ing of the manuscript.

REFERENCES

ANDREWS, H. N. (1961). Studies in Paleobotany. HURUSAvA, ISAO (1962). Can't the so-called Eu­ New York & London. phorbiaceae (sensu lato) keep his rank as a true AXELROD, D. I. (1963). Fossil floras suggest compact taxon) Acta phytotax. Kyoto 2: 82-83. stable. not drifting continents. ]. geophys. HOLLICK, A. (1936). The Tertiary floras of Alaska. Res. 68: 3257-3263. U.S. geol. 5lISY~J +1 "'lOA 'LSINvl.OrrOavT"d 3:IIL R.\SKY -- l'rXrE 2

U'l -,

t[ '"l0A 'LSf;-;VLOnO:;IV"lVd 3Hl. .\ 'TSVH RASKY - SOME PLANT REMAINS FROM THE TERTIARY OF HUNGARY 269

MULLER, JOANNES. ARGOVIENSIS (1873-1874). South Arcot District, Madras. J. Indian bol. Euphorbiaceae in C.F.Ph. Martius: Flora Brasi­ Soc. 35(3): 284-307. liensis, 11 (2) Monachii. Idem (1960). Silicified woods from the Tertiary NAVALE, G. K. B. (1960). Phyllantinium banga­ rocks of South India. Palaeontographica, 106B' lamodense: a new species of fossiL Euphor­ 99-140. biaceous wood from the "Cuddalore series" RASKY, K. (1960). Pfl.anzenreste aus dem Ober­ of India. Palaeobotanist, 9(1-2): 11-16. eozan Ungarns. Senck. lelh. 41: 423-449. PAX, F. (1912-19r4). Euphorbiaceae in A. Engler: Idem (1962). Tertiary plant remains from Hun­ Das Pflanzenreich, IV. 147. I-VII, Leipzig. gary. Upper Eocene and Middle Oligo­ PAX, F. & HOFFMANN, K. (1919-1924). Euphor­ cene. Atm. Rist.-nat. Mus. Nat. Rung., 54: biaceae in A. Engler: Das Pflanzenreich, IV. 31-55. 147. IX-XVII, Leipzig. Idem (1964). Studies of the Tertiary plant remains Idem (1931). Euphorbiaceae in A. Engler und from Hungary. Ibid. 56: 63-96. K. Prantl: Die natiirlichen Pflanzenfamilien, Idem (1965). Contribution to the study of the 2. Aufl., 19/c: 11-233, Leipzig. Tertiary plant remains from Hungary. Ibid. PRAKASH, U. (1959). Studies in the Deccan Inter­ 57, in press. trappean flora 3. On a new species of fossil WEYLAND, H. (1964). Lehrbuch del' Palaobotanik. woods of Euphorbiaceae from the Intertrappean 2. Auf!. Berlin. beds of Madhya Pradesh. Palaeobotanist, 6(2): ZAKLINSKAJA, E. D. (1963). Importance of Angios­ 77-81. perm pollen for the stratigraphy of Cretaceous PURl, G. S. (1960). Indian forest ecology I & 2. and Paleogene deposits and botanical-geog­ New Delhi - Calcutta. raphical provinces at the bundary between the RAMANUJAM, C. G. K. ,(1956). Fossil woods of Cretaceous and Tertiary systems. DoM. Euphorbiaceae from the Tertiary rocks of S.tovj. Palynol. 9: 105-112.

EXPLANATION OF PLATES

(All photographs are umetouched)

PLATE 7. Carpolithus alchorneaeformis sp. nov. X 2. 1. Agrostistachyophyllum tomharrisi gen. et sp. nov. XI. PLATE 3 2. Alchorneaephyllum grambasti (Risky) comb. nov. xl. 8. Codiaeum variegatum (L.) Blume, recent leaf 3. Alchorneaephytlum chandleri sp. nov. x I. for comparison from Luzon. x 1. 9. Codiaeophyllum palaeovariegatum gen. et sp. PLATE 2 nov. X I. 10. Codiaeophyllum palaeovariegafum gen. et 4. Baliospermophy/lum kraeuseli gen. et sp. nov. sp. nov. fragment of the counterpart from Fig. 9, X 2/3. slightly enlarged. 5. Baliospermophyllum kraeuseli gen. et sp. nov. II. Codiaeophyll..m palaeovariegalum gen. et sp. fragment of the counterpart. X I. nov. X I. 6. Omphaleaephytlum weylandi gen. et sp. nov. 12. Codiaeum variegatum (L.) Blume, recent xl. leaf for comparison from New Guinea. X 1.

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