Systematics, biostratigraphy and paleoecology of the genus Toddalia Jussieu (rutaceae) in the European tertiary

Review o f Palaeobotany and Palynology, 28 (1979): 311--363 311 © Elsevier Scientific Publishing Company, Amsterdam -- Printed in The Netherlands

SYSTEMATICS, BIOSTRATIGRAPHY AND PALEOECOLOGY OF THE GENUS TODDALIA JUSSIEU (RUTACEAE) IN THE EUROPEAN TERTIARY

HANS-JOACHIM GREGOR

Institut fur Paliiontologie und historische Geologic, Mi~nchen (Germany GFR) (Received December 8, 1977; revised version accepted November 13, 1978)

ABSTRACT Gregor, H.-J., 1979. Systematics, biostratigraphy and paleoecology of the genus Toddalia Jussieu (Rutaceae) in the European Tertiary. Rev. Palaeobot. Palynol., 28: 311--363. The genus Toddalia Jussieu, which has a Recent distribution from West Africa to Southeast Asia (between 30 ° northern and 30 ° southern latitude) is present in the European Tertiary (between 35 ° and 55 ° northern latitude). In nearly all browncoal deposits, clay pits and in many fissure-fillings one can find hard and boat-shaped seeds with rutaceous characteristics. The new species T. maerkeri, T. rhenana and T. thielei, as well as the new combination T.exeavata, are added to the taxa which were known up to now. Some new varieties of T. latisiliquata, T. maii and T. rhenana complete the picture of this variable group. The stratigraphical range is from Eocene to Pliocene, with the most abundant occurrence in the Miocene. The Recent species grow as lianas in areas with subtropical to tropical conditions n o t below 14°C mean annual temperature and about 2000 m m mean annual precipitation, mostly in evergreen forests. The paleoecology of the Tertiary sites was probably similar to the Recent ones and the accompanying mastixioidean floras support these conclusions. A key for the fossil taxa and graphs for comparison help to identify the fossil groups -the first one with small seeds from Eocene to the Pliocene and the second with large seeds mainly in the (Middle) Miocene. INTRODUCTION F o r m o r e t h a n a h u n d r e d years, r e n i f o r m , hard, shining, black seeds have b e e n f o u n d in m a n y T e r t i a r y d e p o s i t s in E u r o p e a n d there have b e e n m a n y a t t e m p t s b y s p e c i a l i s t s t o c l a s s i f y t h e m , m o s t l y as l e g u m e s . C h a n d l e r ( 1 9 6 3 ) m a d e the first reference to an organ genus, Toddaliospermum, b u t did n o t c o v e r t h e R e c e n t m a t e r i a l . I n 1 9 7 3 Mai c o m p a r e d t h e fossil s e e d s f o r t h e f i r s t t i m e w i t h t h e s e e d s o f R e c e n t T o d d a l i a species. S i n c e t h e n t h i s m o n o g r a p h o n fossil T o d d a l i a h a s b e e n i n p r e p a r a t i o n b y t h e a u t h o r .

312 MATERIAL T h i s r e s e a r c h is b a s e d o n t h e e x a m i n a t i o n o f n e a r l y 2 0 0 R e c e n t f r u i t s a n d a b o u t 1 5 0 0 R e c e n t s e e d s f r o m all o v e r t h e w o r l d , t o g e t h e r w i t h a p p r o x i m a t e l y 5 0 0 fossil s e e d s f r o m t h e w h o l e o f E u r o p e , p a r t l y c o l l e c t e d b y t h e M u n i c h researchers, partly loaned from many persons and institutions. The following institutions have sent Recent material for comparison with t h e fossils. T h e list is i n a l p h a b e t i c a l o r d e r o f t h e cities, i n w h i c h t h e i n s t i t u t i o n s are s i t u a t e d : Antwerp (Belgium) -- Koninklijke Maatschappij voor Dierkunde van Antwerpen Bangkok (Thailand) -- Department of Agriculture Berlin--Dahlem (Germany GFR) -- Botanischer Garten und Botanisches Museum Budapest (Hungary) -- Termes Zettudomanyi Muzeum Canton (Peoples Republic of China) -- Institutum Botanicum Austro-Sinense Coimbatore (India) -- Botanical Survey of India Copenhagen (Denmark) -- The Botanical Museum, University of Copenhagen Florence (Italy) -- Herbarium Universitatis Florentinae, Instituto Botanico Halle-Saale (Germany GDR) -- Botanischer Garten, Sektion Biowissenschaften H o n g k o n g - Agriculture and Fisheries Department, Kowloon Jamaica Plain (Mass., U.S.A.) -- The Arnold Arboretum of Harvard University Kew (Richmond, Great Britain) -- Royal Botanic Gardens London (Great Britain) -- British Museum of Natural History Munich (Germany GFR) -- Bayerische Botanische Staatssammlung Nago (Okinawa, Japan) -- Okinawa Forest Experiment Station, by courtesy of Government Forest Experiment Station, Meguro West Beltsville (Md., U.S.A.) -- Seed Collection and Identification Plant Taxonomy Laboratory Ziirich (Switzerland) -- Botanischer Garten und Institut fiir systematische Botanik der Universit~t T h e f o l l o w i n g is a list o f c o l l e c t i o n s o f fossil s e e d s i n a l p h a b e t i c o r d e r o f t h e cities, i n w h i c h t h e y are l o c a t e d : Berlin (Germany GDR) -- Museum fiir Naturkunde, Pal~ontologisches Museum Cologne (Germany G F R ) - Geologisches Institut der Universit~t (Coll. Weyland) Essen (Germany GFR) -- Ruhrland-Museum Haarlem (The N e t h e r l a n d s ) - Rijks Geologische Dienst Krakov ( P o l a n d ) - Instytut Botaniki (Coll. J. Zablocki) Lisbon (Portugal) -- Universidade nova de Lisboa, Depto. de Ciencias de Terra London (Great Britain) -- British Museum of Natural History Munich (Germany GFR) -- Bayerische Staatssammlung fiir Paliiontologie und historische Geologie Prague(Czechoslovakia) -- Ustredni Ustav Geologicky, and Narodni Muzeum Sessenheim (France) -- Coll. F. Geissert Utrecht (The Netherlands) -- Rijksuniversiteit Utrecht, Laboratorium voor Palaeobotanie en Palynologie Warsaw (Poland) -- Muzeum Ziemi Wiirzburg (Germany GFR) -- Geologisch-Pal~/ontologisches Institut der Universit~t (Coll. F. Kirchheimer). L i s t o f sites, y i e l d i n g fossil s e e d s o f Toddalia:

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Czechoslovakia Cheb and Sokolov Basin -- Middle Miocene, Karpathian Chomutov--Most--Teplice Basin, Cernovice, Boring Cv 62 d -- L ow er to Middle Miocene Hradek Basin, o p e n mine K r i s t i n a - Middle Miocene, Karpathian Moravian Basin, Mikulov -- Upper Miocene, Badenian

England Cliff End, Mudeford -- Highcliff Sands, Upper Eocene, Bartonian

France Bischweiler--Kaltenhouse, Boring -- L ow e r Pliocene, Brunssumian sands Haguenau, Alsace -- L o w e r Pliocene, Brunssumian sands and clays Pont-de-Gall, Cantal -- L o w e r Pliocene, Susterian sands Schwighouse, Alsace, boring -- L o w e r Pliocene, Brunssumian sands Soufflenheim, Alsace -- L ow er Pliocene, Reuverian sands

Germany (GFR) Burgmagerbein, Bissingen, fissure filling -- Low er to Middle Oligocene Eschweiler, o p en mine Zukunft~West -- Middle Miocene Sands (equivalents of the " N e u r a t h e r Sands"), floral zone VI Haag, Treuchtlingen, fissure-filling -- L ow e r to Middle Oligocene Habichtswald, Kassel -- Upper Miocene coal ?Haidhof-Ponholz, Oberpfalz -- Upper Miocene clays and coals (material could n o t be f o u n d ! ) Harburg, Ries -- u p p e r m o s t Middle Oligocene, Websteri browncoal Hessenbriicker H a m m e r near Laubach, Wetterau -- Middle Miocene coal, floral zone VI t o VIII? Langenau, Ulm -- Upper Miocene clay, Upper Freshwater Molasse MShren, Treuchtlingen, f i s s u r e - f i l l i n g - L ow er Oligocene Niederpleis, KSln -- Upper Oligocene or L o w e r Miocene clays ?Salzhausen, Vogelsberg -- Upper Miocene (material surely belongs to the Hessenbriicker Hammer, see above) Schwandorf, o p en pits, Oder, Briickelholz, H o f e n s t e t t e n , Nordfeld -- Middle Miocene sands, clays and quarzites, floral zone VI Viehhausen, Regensburg -- Upper Miocene coal ZiSlpich, open mine Victor-Rolff -- Upper Miocene sands ( L o w e r Pliocene c a n n o t be excluded)

Germany (GDR) Hartau, Lausitz -- Middle Miocene clay, floral zone VI Holzweissig, Bitterfeld -- L ow er Miocene, " B i t t e r f e l d e r D e c k t o n " , floral zone III Klettwitz -- Upper Miocene clay, floral zone X Nerchau -- Middle Oligocene (?), argillaceous sands

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The Netherlands Bouwberg, Brunssum -- Lower Pliocene clays, Brunssumian Poland Turow, Bogatynia, open mine--- Middle Miocene coal, floral zone VI Wieliczka, saltmine -- Middle M i o c e n e ' , floral zone XII Portugal Povoa de Santarem, Lisbon -- Middle to Upper Miocene, Vindobonian GENERAL OBSERVATIONS ON RECENT TODDALIA

Systematic position The family Rutaceae is divided into the subfamilies Rutoideae, Flindersioideae, Spathelioideae, Citroideae, Rhabdodendroideae and the Toddalioideae. The latter can be separated into further groups and one of them is named Toddalieae. The Toddalieae are further divided into Phellodendrinae, Pteleinae, etc., and the Toddaliinae. The Toddaliinae contain the following genera: Acronychia Forster, Skimmia Thunberg, Teclea Verdoorn, Toddalia Jussieu, Toddaliopsis Engler and Vepris Jussieu (compare to all mentioned systematic determinations in Engler, 1931, Bd. 19 a). According to Verdoorn (1926, p p . 3 9 3 , 4 0 0 ) the genus Toddalia is m o n o t y p i c and has only one Recent s p e c i e s - Toddalia asiatica (L.) Lamarck (synonym: T.aculeata Persoon). In contrast to this opinion, a b o u t 25 species from Africa and Asia (descriptions mostly w i t h o u t mentioning fruits or seeds) and, astonishingly enough, two species from the Rio Grande Region (Brazil), have been described (Don, 1831, pp.805, 806). The latter " S o u t h American" species do n o t belong to Toddalia, b u t perhaps to the genus Scopolia Smith (ibid. p.805). This theory is kindly confirmed by M. Emmerich (Rio de Janeiro) and should be of interest to recent botanists. It is m y opinion, that the seeds of Toddalia show that there are more Recent species b u t it is n o t the intention here to produce a monograph on the Recent genus Toddalia. Illustrations of T. asiatica (leaves, flowers and fruits) are given in Figs.9--11 (by courtesy of Zen'Iti Kisi, Government Forest Experiment Station, Meguro, T o k y o and Kei-Ichi Shimabuku, University of the R y u k y u s , Shuri, Naha City, Okinawa, both Japan).

Morphology and histology of the fructifications The fructifications of Toddalia asiatica (L.) Lamarck are berries with 3--8 seeds (Figs.l, 2, 12--15). i Formerly thought to be Upper Miocene but shown by geochemical research to be Middle Miocene (compare J~hnichen et al., 1977, p.352; and L. Stuchlik and H. Walther, pers. comm.).

315 (1) The fruit. The fruit is a berry or a drupe, more or less rounded to elongate, with a sarcocarp rich in oil. The fleshy exocarp consists of 3--8 locules, each with one s u b c a m p y l o t r o p o u s seed (Figs.l, 2). The size of the fruit in fresh state is a b o u t 10 mm (Fig.15), in a dried state a b o u t 5 mm (Figs.12--14). (2) The seeds. The seeds are reniform, small (2.5 X 2.0 mm) to large (6.5 X 4.5 mm) with a hard and b o n y testa, finely pitted (Fig.16) and mostly shining yellowish or brownish. The hilum lies on the ventral side with the micropyle at the apical end and the raphe and the chalaza (inner part) at the basal end (Figs.l, 3--7). According to Corner (1976, p.236), the vascular bundle of the raphe is often curved over the chalaza (Figs.6, 7), with a small vascular plexus around the chalaza and two very short branches to the antiraphe from the chalazal flexure. This description is confirmed in Recent as well as fossil seeds of our collection. Corner (1976, p.236, figs.494, 495) provides new and interesting data on the embryology and histology of Toddalia asiatica. The subcampylotropous seeds have three layers in the testa (Fig.8), an outer palisade cell wall (Fig.8a); an outer mesophyll of thick-walled, lignified and angular cells (Fig.8b); and an inner mesophyll with cells in rows, not lignified, b u t parenchymatous (Fig.8c; and ibid. p.236). The tegmen is thin-walled with fine criss-crossed spiral lignification (Figs.8d, 55--58). The similarities between the Recent and the fossil Toddalia seeds are illustrated in Figs.54--64 (SEM), showing the inner and outer surfaces of seeds and seed-casts. The s o m e w h a t similar seeds of Z a n t h o x y l u m L. can be distinguished from the seeds of Toddalia by having a long and even hilum all over the straight ventral side, a mostly rugose to reticulate surface sculpture and a blackshining outer seed-testa (also compare Boesewinkel, 1977). As a guide to the use of the key and the descriptions, the measurements were taken in the following manner (compare Figs.3--5). In the literature the length is often also called breadth; it is dependent on the way of measuring seeds. The indices give the ratio between the measurements, i.e. the ratios height--length and breadth--length, the first showing the shape in a mathematical way, the second giving information on the number of seeds per fruit (for the indices and the morphology compare Table V, Appendix). Distribution pa ttern and ecology The Recent genus Toddalia shows a SW African to SE Asian distribution. In general the following regions can be mentioned (see Fig.70): S Africa, W Africa, Madagascar, Mauritius, Mascarene Islands, Seychelles, India, Himalayan Range, Sumatra, Java, Borneo, the Philippines, the Moluccas, Taiwan, S Japan and nearly the whole of China (partly information from C.G.G.J. van Steenis, written communication, 1977). The following paragraphs list more detailed work on the geographical distribution, ecological

316

and sociological characteristics of the environment of Toddalia, with special references to the Asiatic region. For Africa, some authors mention T.asiatica from Transvaal, Natal, Rhodesia, Nyasaland, Cameroun, Tanzania, Kenya, Uganda, Kongo-Brazzaville, the Central African Republic, Gabon, Sudan and Bourbon (after Hooker, 1875 p.306; Verdoorn, 1926, pp.400--401; Harvey and Sonder, 1959--60, p.447; Aubreville, 1963a, p.107; 1963b, p.75). It must be pointed out that the "Toddalia" fruits from Herbarium Sieber (Antwerp and Halle) belong to the genus Vepris Jussieu. They were collected on Mauritius. In India and Burma, T.asiatica occurs in Bengal, Bombay, etc. (Kirtikar and Basu, 1918, p.465), and especially in the subtropical Himalayan region from Kumaun eastwards to Bhutan, ascending to 1700 m elevation, the Khasi Mountains ascending to 2000 m, and throughout the western peninsula

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(after Hooker, 1875, p.497; Hara, 1966, p.171). According to Champion and Seth (1968, p.168), the creeper-swamp forest (Brahmaputra Valley), where Toddalia grows, contains Magnolia, Litsea, Quercus and Calamus. A richer type of forest (Nilgiri hills, Madras) yields Toddalia, Ternstroemia, Gordonia, Ilex, Meliosma, Cinnamomum, Symplocos, Turpinia, Rubus and many more (ibid. p.281). Trimen (1893, pp.215, 216) mentions T.aculeata Persoon from Ceylon, Kurz (1877, p.183) the same species from tropical and moister hill forests in Burma. In Thailand, according to Cralb (1931, p.220) and Guillaumin in Humbert (1946, pp.613, 614). T.asiatica grows in shrub jungle and evergreen forests (on river banks) in many provinces, T. tonkinensis Guillaumin in Tonkin. On Java, the species T.aculeata Persoon can be found all over the island at 1000--2300 m altitude and together with Cissus species (Vitaceae) it forms 25 m long lianas (Koorders, 1912, p.422). Backer (1965, p.101) suggested that Toddalia effusa Turczaninow is in fact T.asiatica. This species is to be found in humid forests (1000--2300 m elevation), monsoon forests, mixed lowland and hill rain forests in mountain swamps and along lakesides on Java. On the Philippine Islands the liana grows on Luzon, Rizal, Laguna, Nueva Vizcaya and Palawan in thickets at low and medium altitudes up to 1700 m (Merrill, 1923, p.333 and Vidal and Soler, 1886, p. 75). On Taiwan, T.asiatica is found in Nantou, Mt. Pahsienshan, Mt. Alishan, Taichung, Taitung, Kaohsiung and Hengchun in secondary forests and waste lands near the seashore (Li, 1963, p.385). In Japan, the climber is distributed in the evergreen broad-leaved forests of the Ryukyu Floral Region and has its northern limit of occurrence on the island of Amani-oshima (Hara, 1959, p.85). The sociological character of the evergreen forests in this region is shown by taxa like Mallotus, Wikstroemia, Symplocos, and many other subtropical plant species. Sargent (1916, pp.137, 138) defines T.asiatica as a semiscandent scrub (2--6 m tall) in W Hupeh at altitudes around 30--300 m, and in Szechwan and Yunnan (1600 m). In the same provinces and some others -- Shensi, Hupei, Fig. 1. Berry of To ddalia asia tica in longitudinal section with two seeds and vascular bundles. Fig.2. Cross-section of a berry with eight seeds lying concentrically around the central vascular strand. Fig.3--5. S c h e m a t i c drawings of Toddalia seeds with the m e a s u r e m e n t s length (l), height (h) and breadth (b) and hilum (hi), raphe (r) and m i c r o p y l e (m). Fig.3. F r o m the side. Fig.4. F r o m above, d marks the dehiscing line. Fig.5. F r o m the front. Fig.6a--e. Various raphe excrescences on Toddalia seeds; the raphe normal (a), with a small curved bundle (b), with a long curved bundle over the chalaza (c), with a long bundle reaching to the antiraphe (d), and a bundle projecting basalwards (e). Fig.7. Longitudinal section through a Toddalia seed with raphe (r), chalaza (ch), a second curved bundle over the chalaza (c) and the vascular strand going to the stalk (v). Fig.8. Cross-section of the testa of a Toddalia seed (fossil, changed after Corner, 1976, fig.495), a. Outer epidermal layer (o.e.) -- palisade of thin-walled, n o t lignified cells. b. O u t e r m e s o p h y l l -- lignified, thick-walled cells, c. Inner m e s o p h y l l - - n o t lignified, thickwalled cells. Inner epidermal layer (i.e.) -- unspecialized cells, d. T e g m e n -- thin-walled cells with fine criss-crossed spiral lignification (striate thickening after Boesewinkel, 1977, p.199).

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Hunan, Chekiang, Fukien, Kwangtung (Canton), Kwangsi and Kweichow -Handel-Mazzetti, (1929--36, p.628) and the Iconographia Cormophytorum Sinicorum (1972, vol.II, p.552) mention the same species, especially from mixed-mesophytic and evergreen forests. These few notes provide only a general impression of the geographical distribution and the ecological environment of the Toddalia lianas. In summary, one can say that the climber Toddalia asiatica and the other species of the genus (?) grow abundantly in most regions with subtropical to tropical climates, except the New World, Australia and New Guinea. All references also indicate that the Recent genus is restricted to the tropical--subtropical area between 30°N and 30°S. This means that the ecological distribution is restricted to the tropical and paratropical rain forest and the subtropical forest (sensu Van Beusekom, 1971, p p . 3 8 8 , 3 8 9 ) . In Table IV (p. 353) climatological data concerning Toddalia asiatica (and related species) are given, and it is clear that it lives in regions with mean annual temperatures of 14--28°C and 500--3500 mm (up to 12000 mm!) annual precipitation. If all data are taken together, we have a " m o d e l climate" of subtropical character with 20°C mean annual temperature and approximately 2000 mm mean annual rainfall. These data vary according to elevation, floral region {monsoon, etc.) and many other factors. The above-mentioned conditions which we suggest, occurred in Tertiary times, when Toddalia was abundant. The absence of the seeds of this genus, for example in the Pliocene, may have been due to a decrease of temperature to below 14°C. FOSSIL EVIDENCE

Introduction and general considerations As we have seen above, fossil reniform seeds of the Toddalia type have often been found and described, mostly under "Leguminosae". With the knowledge of the generic determination, specimens of these fossils could be found in nearly every European collection of fossil fructifications -- and in this way it was easy to compare the existing fossils with one another. There have only been reports of fossil seeds, never of fossil leaves, except the "subfossil to recent" leaves of Toddalia aculeata mentioned by Menzel Fig.9. Appearance of a Toddalia asiatica shrub (photo: Zen'Iti-Kisi, Government Forest Experiment Station, Meguro, Tokyo, Japan). Fig.10. Flowers and leaves of Toddalia asiatica (L.) Lamk. (photo: Kei-Ichi Shimabuku, University of the Ryukyus, Shuri, Naha City, Okinawa, Japan). F i g . l l . A branch of Toddalia asiatica (L.) Lamk. with fruits (photo: Kei-Ichi Shimabuku, Okinawa, Japan). Fig.12. Dried fruit of Toddalia with eight seeds and partly rotten sarcocarp from apical. (T.asiatica from Tsui, Canton, China); x 4. Fig.13. Dried fruit (five seeds) of Toddalia asiatica with oil cells (Hiniduma, Distr. Galle, Ceylon); x 4. Fig.14. Broken up fruit of Toddalia asiatica with the left seed "in situ" and a basal raphe excrescence (Tsui, Canton); X 4.

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321 ( 1 9 1 3 ) f r o m C a m e r o u n . N e i t h e r have fossil pollen grains b e e n f o u n d u p to now. I am glad t o say h e r e t h a t m y colleague H. Thiele-Pfeiffer will s h o r t l y publish fossil Toddalia p o l l e n grains f r o m W a c k e r s d o r f (1979). It will be o f interest t o revise some fossil findings o f " T o d d a l i a " and t o p o i n t o u t t h a t t h e y b e l o n g to a closely related f o r m -- Clausenopsis Engler. T o d d a l i o s p e r m u m o r n a t u m (Chandler, 1 9 6 3 ) and m y Toddalia f r o m A r j u z a n x (S F r a n c e ) (in Gregor, 1 9 7 5 b , p p . 1 2 7 , 128, fig.41) b e l o n g t o this genus. NEW TAXA Toddalia excavata (Chandler) nov. c o m b . Basionym and holotype: 1963 Toddaliospermum excavatum Chandler, The Lower Tertiary Floras of Southern England III. Flora of the Bournemouth Beds; the Boscombe, and the Highcliff Sands, p.93, pl.14, figs.31--34. Inv. No.V 43397 (Br. Mus. Nat. Hist.). E m e n d e d diagnosis: Small r e n i f o r m seed, b i s y m m e t r i c , with d e e p l y e x c a v a t e d triangular hilar scar. F i n e l y p i t t e d surface with equiaxial cells (0.017 m m ) . Obscure c o n c e n t r i c ridges parallel to the dorsal margin. Size: L e n g t h : 1.55 m m ; height: 1.15 m m ; b r e a d t h : 0.8 m m ; b r e a d t h - - l e n g t h index: 0.51; h e i g h t - - l e n g t h index: 0.71. T y p e : Seed. L o c u s typicus: Cliff End, M u d e f o r d , England. S t r a t u m t y p i c u m : Highcliff Sands or Cliff E n d Beds, u p p e r m o s t Auversian, Eocene. R e m a r k s : T h e v e r y small seed u n d o u b t e d l y belongs t o the genus Toddalia, as I have seen m y s e l f in the British Museum. T h e m e a s u r e m e n t in C h a n d l e r (1963, p . 9 3 ) is d i f f e r e n t f r o m t h a t o n the pictures (ibid. pl.14, figs.31--34). T h e e r r o r is a b o u t 16% and is c o r r e c t e d in this diagnosis. T h e seed belongs t o a m a i i - t y p e (see p.338).

Fig.15. A fleshy fruit of Toddalia asiatica with oil cells in the sarcocarp and the vascular strands going to the seeds (fruit from Zen'Iti-Kisi, Meguro, Japan); x 4. Fig.16. Recent seed of Toddalia asiatica (L.)Lamk. with the raphe to the left (Madras, India, Br. Mus. Nat. Hist., London); x 10. Fig.17. X-ray photo of a fossil Toddalia seed, showing the seed cast, the compact hilar scar and the raphe channel with a small "sack" at the chalazal point; × 10. Fig.18. X-ray photo of a fossil Toddalia seed, partly abraded and denuding the raphe channel at the most basal end, imitating a legume seed (compare Martyia naviculaeformis E.M. Reid, 1923); x 10. Fig.19--27. Populations of recent and fossil Toddalia seeds in natural size for comparisons with newly found material. Fig.19. T.asiatica (L.)Lamk. -- Hiniduma, Distr. Galle, Ceylon. Fig.20. T.asiatica ( L . ) L a m k . - Hongkong, China. Fig.21. T.aculeata P e r s o o n - Madras, India. Fig.22. T.maii Gregor var. minima Gregor. Fig.23. T. maii Gregor. Fig.24. T. thielei Gregor. Fig.25. T.rhenana Gregor. Fig.26. T.latisiliquata (Ludwig) Gregor. Fig.27. T. naviculaeformis (E.M. Reid) Gregor. (All seeds are property of the Bayerische Staatssammlung fiir Pal~ontologie und historische Geologie, Miinchen.)

322

Toddalia latisiliquata (Ludwig) Gregor var. gracilis nov. var. (Fig.29) Holotype: Inv. No.3481, Museum fiir Naturkunde, Pal~iontologisches Museum, Berlin. Diagnosis: Seeds typical of T. latisiliquata, b u t smaller and more graceful {tiny boats). Size: Length: 4.0--5.5 mm; height: 3.4--3.8 mm; breadth: 2.0--2.5 mm; height--length index: 0.72; breadth--length index: 0.44. Locus typicus: Hessenbrficker Hammer near Laubach, Wetterau (Germany GFR). Stratum typicum: Middle Miocene, floral zone VI (to VIII?); oldest Wetterau browncoal. Remarks: Especially in material from Hessenbriicken, a population of this variety can be seen (compare Gregor 1978a, pl.IV, figs.la, b, 2a, b, pl.V, figs.la, b). Toddalia latisiliquata (Ludwig) Gregor var. viehhausenensis nov. var. (Figs. 30--32) Holotype: Inv. No. 101--501, Geologisch-Pal/iontologisches Institut der Universit/it Wiirzburg. Diagnosis: Seeds typically T. latisiliquata type, b u t with a more narrow hilum and some shallow furrows vertical to the ventral ridge. Size: Length: 5.3--6.0 mm; height: 3.3--4.0 mm; breadth: 2.3--2.6 mm; height--length index: 0.63; breadth--length index: 0.43. Locus typicus: Mine Ludwigszeche, Viehhausen near Regensburg (Germany GFR). Stratum typicum: Upper Miocene browncoal, Lower Tortonian. Remarks: These typical seeds were found only in the material from Viehhausen (Inv. No.101--501 to 101--506, Geol.-Pal~iontol. Inst. Wiirzburg) and were given for study by courtesy of W. Trapp (Wfirzburg). The age of the browncoal from Viehhausen can be confirmed as (Lower) Tortonian, as the faunal character shows the following species: Mastodon angustidens Cuvier, Lagomeryx meyeri Hofmann, Dorcatherium crassum Edwards, Hyotherium soemmeringi v.Meyer, Steneofiber jaegeri Kaup and many more (Wappenschmitt, 1936). Toddalia maerkeri nov. spec. (Figs.33, 34) Misidentification Miiller 1972: Epipremnum crassum C. and E.M. Reid. In: Die Oligoz~nAblagerungen im Gebiet des NSrdlinger Rieses, p.77.

Diagnosis: Seeds very small (3.5 X 1.9 mm), elongate with a very strongly curved vascular bundle over the chalaza, making an extensive raphe excrescence (to the middle of the dorsal face). Deep, triangular hilum with extruding micropyle. Closely related to Toddalia maii Gregor.

323 Size: Length: 3.5 mm; height: 1.9--2.1 mm; breadth: 1.9--2.0 mm; breadth-length index: 0.57; height--length index: 0.55. Type: Seed. Holotype: 1972 XXI 17, Bayerische Staatssammlung fiir Pal~iontologie und historische Geologie, Mfinchen. Isotype: 1972 XXI 84 (ibidem). Derivatio nominis: maerkeri -- from the M~irker Cement Company in Harburg, Ries, S Germany GFR. Locus typicus: Quarry of M~rker Cement, Harburg, Ries, S Germany GFR. Stratum typicum: Websteri Browncoal (gray and black, clayish coal) in "RiesTrfimmermassen" and "Bunter Breccie" (coloured breccia) -- uppermost Middle Oligocene (Chattian, Antoingt). Remarks: The seed was first described as Epipremnum crassum and was found (in fissure-filling-like lenses) together with Stratiotes, Brasenia, Spirematospermum and Cladium (Mfiller, 1972, p.77). In the quarry of the M~rker Cement one can see different clay- and coal-lenses in the so-called "Riestrfimmer-Massen" (impact crater ejecta, breccia), struck by the Ries Meteorite in the Tortonian. The associated pollen flora (ibid. p.140, Harburg -- M~ker-4) indicates Chattian age (Antoingt). Toddalia maii Gregor vat. minor nov. var. (Fig.22) Holo type: Inv. No. 1970 X 823; Bayerische Staatssammlung ffir Paliiontologie und historische Geologie, Miinchen. Diagnosis: Seeds typical of T.maii in shape, b u t smaller (in population). Size: Length: 2.0--3.2 mm; height: 1.6--2.2 mm; breadth: 1.2--1.8 mm; height--length index: 0.73; breadth--length index: 0.57. Locus typicus: Open mine Brfickelholz, Bayer. Braunkohlen-Industrie AG Schwandorf (Oberpfalz), Germany GFR. Stratum typicum: Middle Miocene, sandy browncoal, floral zone VI. Remarks: The small seeds are surely a modification of the c o m m o n T.maii. The more a u t o c h t h o n o u s strata bearing these tiny seeds, are located in Schwandorf (open mine Briickelholz, BBI. Upper Coal seam, sandy coal; Middle Miocene, floral zone VI), Langenau near Ulm (grayish-black clay of the Upper Freshwater-Molasse, Helvetian to Tortonian age), and in the clay-pit Niederpleis near KSln (Lower Coal-seam, Upper Oligocene or Lower Miocene). Toddalia maii Gregor var. abnormis nov. var. (Figs.36, 37) Holotype: Inv. No.G-1976-4; Coll. F. Geissert, Sessenheim, Alsace, France. Diagnosis: Seeds typical of T. maii, b u t with some strange distortions and flattened areas, caused by accumulation of (mostly two) more seeds in the place of one seed (per locule) or disturbance of t h e surrounding seeds by this "twin-occurrence". Size: Length: 3.5--4.5 mm; the other measurements are too atypical and have no significance.

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Locus typicus: Schwighouse near Haguenau, Alsace (France), boring. Stratum typicum: Lower Pliocene, Brunssumian clay and sand. Remarks: At the first glance, the seeds bring some confusion into the study of fossil Toddalia seeds, b u t the reason for the atypical specimens can be seen very easily in Recent fruit material. The occurrence of this type is fairly restricted to Late Miocene and especially Pliocene times (Klettwitz -- floral zone X; environment of Haguenau, Alsace, France, Brunssumian) (compare Gregor, 1978a, pl.VI, figs.5 and 6). For the loan of this material I have to thank F. Geissert, Sessenheim and D.H. Mai, Berlin. Toddalia rhenana nov. spec. (Figs.38, 39) Diagnosis: Seeds reniform, elongated, shining black, with a triangular, symmetrical hilum. Resembling T. maii Gregor, b u t bigger (4.0 X 2.8 mm), longer, and with a concave ditch below the hilar scar parallel to the dorsal margin. Prominent raphe canal and apical micropyle. Size: Length: 3.3--4.5 mm; height: 2.2--2.7 mm; breadth: 1.4--2.0 ram; breadth--length index: 0.43; height--length index: 0.64. Type: Seeds. Holotype: 1976 XVII 1, Bayerische Staatssammlung fiir Pal~ontologie und historische Geologie, Miinchen. Isotypes: 1976 XVII 2-3, 1976 VII 2, Bayerische Staatssammlung fiir Pal~iontologie und historische Geologie, Miinchen. Derivatio nominis: From rhenanus, Rhenish. Locus typicus: Open mine Zukunft-West near Eschweiler, Rheinische Braunkohlenwerke AG KSln, Germany GFR. Stratum typicum: Equivalents of the so-called "Neurather Sande" between the coal seams Frimmersdorf and Garzweiler; Middle Miocene, floral zone VI. Remarks: The tiny seeds are u n d o u b t e d l y related to T.maii from many Miocene localities, but have another shape and size. This new species is found at different sites of Middle Miocene to Pliocene age. The Middle Miocene Mastixioideae flora from Eschweiler, which accompanies this species, resembles that from Schwandorf (Gregor, 1975b). The Pliocene flora types associated Fig.28 and 29. Toddalia latisiliquata (Ludwig) Gregor -- Laubach, Wetterau, Middle Miocene; × 10. Fig.28. No.3477, Museum fiir Naturkunde--Pal/iontologisches Museum, Berlin. Fig.29. Var. gracilis Gregor (No.3481, ibidem). Fig.30---32. Toddalia latisiliquata (Ludwig) Gregor var. viehhausenensis Gregor -- m i n e Ludwigszeche, Viehhausen near Regensburg; X 10. Fig.30. a. From the side. b. From above (No.101--501, Coll. Kirchheimer, Geologisch-Palh'ontologisches Institut Univ. Wiirzburg). Fig.31. Other seed (No.101--502, ibidem). Fig.32. Piece of browncoal w i t h o n e seed in situ (Fig.31, No. 101--502). Fig.33 and 34. Toddalia maerkeri Gregor -- M~irker Cement Quarry, Harburg, Ries, Websteri browncoal, Middle Oligocene; × 10. Fig.33. Holotype (Inv. No.1972 XXI 17, Bayerische Staatssammlung fiJr Palh'ontologie und historische Geologie, Miinchen). a. From the side, raphe excrescence to the right, b. From above. Fig.34. Well-preserved seed with raphe excrescence going basal to the antiraphe (Inv. No.1972 XXI 84, ibidem), a. From the side, raphe left. b. From above.

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with T. rhenana are more humid-temperate and less rich in species. The material from Eschweiler could be studied by courtesy of J.v.d. Burgh (Utrecht). Further sites: (1) Clay pit Bouwberg (I), The Netherlands, Brunssum A, Lower Pliocene (Material in the Rijks Geologische Dienst, Haarlem -- for permission to study it I have to thank W.H. Zagwijn). (2) Open mine VictorRolff, Eschweiler (Rhenish browncoal), sand over coal seam Garzweiler, Pliocene or uppermost Miocene? (material in the Geol. Inst. KSln, by permission of K. Kempf). (3) Open mine Zukunft-West, Eschweiler (Rhenish browncoal), coal seam Frimmersdorf, Middle Miocene {material could be studied by courtesy of K. Kilpper, Essen). Toddalia rhenana Gregor vat. major nov. vat. Basionym and holotype : 1959 Cytisus latisiliquata Ludwig vel Martyia naviculaeformis E.M. Reid -- Zagwijn, Fortschr. Geol. Rhld. Westf., 4, pl.1, fig.6 (without No., Rijks Geol. Dienst, Haarlem, The Netherlands).

Diagnosis: Seeds of T.rhenana type, elongated with a concave ditch, but bigger than the normal seeds. Size: Length: 4.5--5.4 mm; height: 2.5--3.2 mm; breadth: 1.8--2.0 mm; height--length index: 0.64; breadth--length index: 0.41. Locus typicus: Bouwberg, clay pit (The Netherlands) Fig.35. Toddalia mail Gregor -- open mine Oder II (Bayer. Braunkohlen-Industrie AG Schwandorf), Middle Miocene, floral zone VI; × 10. (Inv. No.1970 X 132, Bayerische Staatssammlung ftir Pal~iontologie und historische Geologie, Miinchen). Fig.36. Toddalia mail Gregor var. abnormis Gregor -- boring Schwighouse near Haguenau, Alsace, France, Lower Pliocene, Brunssumian; x 10. (No.G-1976-4, Coll. F. Geissert, Sessenheim near Haguenau, France) Fig.37. Toddalia mail Gregor var. abnormis Gregor -- open mine Klettwitz (GDR), "Spezialton", Upper Miocene, floral zone X; x 12. a. From above, b. From the side. Fig.38--39. Toddalia rhenana Gregor -- open mine Zukunft-West near Eschweiler, equivalents of the "Neurather Sande", Middle Miocene, floral zone VI; X 10. Fig.38. Holotype (1976 XVII 1 ). a. From the s i d e - raphe right, b. From above. Fig.39. Isotype (1976 XVII 2) from the side. Fig.40--41. Toddalia thielei Gregor -- open mine Oder (Bayer. Braunkohlen-Industrie AG Schwandorf), Middle Miocene, floral zone VI; x 10. Fig.40. Holotype (1970 X 133). a. From the side -- raphe right, b. From above. Fig. 41. Isotype (1970 x 546) from the side. Fig.42. Toddalia sp. 2 -- open mine Oder II (Bayer. Braunkohlen-Industrie AG Schwandorf), Middle Miocene, floral zone VI; x 10. (1970 X 128), a. Abraded seed from above, raphe channel is seen as a hole to the extreme right, b. From lateral. Fig.43--49. Toddalia sp. 3 (div. spec.? ) from southern Germany. All seed-cast fillings from fissure-fillings in Upper Jurassic limestones; all Lower to Middle Oligocene; × 10. Fig.43. Type 1 -- Burgmagerbein 3 (1972 XIV 44). Fig.44. Type 1 -- Burgmagerbein 3 (1972 XIV 45). Fig.45. Type 2 -- Burgmagerbein 3 (1972 XIV 47 ). Fig.46. Type 4 -- Burgmagerbein 3 (1972 XIV 43). Fig.47. Type 5 -- Haag 2 (1975 XXIII 41). Fig.48. Type 3 -Burgmagerbein 3 (1972 XIV 41). Fig.49. Type 3 -- Burgmagerbein 3 (1972 XIV 42). Figs.38--49. Material is property of the Bayerische Staatssammlung fiir Paliiontologie und historische Geologie, Miinchen.

328 Stratum typicum: Lower Pliocene, Brunssumian clay. Remarks: This type occurs rarely in open mine Zukunft-West (Eschweiler, Middle Miocene), but mainly at Bouwberg I (Brunssum, Lower Pliocene, Brunssumian). Toddalia thielei nov. spec. (Figs.40, 41) 1975b 1978a

Toddalia maii Gregor, Die mittelmioz~ne Mastixioideen-Flora aus dem Braunkohlen-Tagebau Oder IIbei Wackersdorf (Oberpfalz), p.131, pl.7, figs.2a, b. Toddalia maii Gregor, Acta Palaeobotanica, XIX, p. 26, pl. V, figs. 3a, b.

Diagnosis: The seeds are bigger than those of T. maii Gregor, measuring 4.5 X 2.8 mm, robust, thick, with a coarsely pitted surface of equiaxial cells. The ventral margin is straight with a triangular, short and asymmetrically lying hilum. In transverse section the outline is dorsally rounded and ventrally flattened, not round as in T. mail Size: Length: 3.9--4.4 mm; height: 2.4--3.0 mm; breadth: 1.8--2.2 mm; height--length index: 0.65; breadth--length index: 0.49. Type: Seeds. Holotype: Inv. No.1970 X 133 Bayerische Staatssammlung fiir Pal~iontologie und historische Geologie, Mfinchen. Isotypes: 1970 X 545, 546 (ibidem). Derivatio nominis: named after H. Thiele--Pfeiffer, who found the first fossil Toddalia pollen grain in the survey of the Tertiary browncoal of open mine Oder II (Schwandorf) {doctoral thesis in preparation at the University of Munich for 1979). Locus typicus: Open mine Oder II, BBI Schwandorf (Oberpfalz), Germany GFR. Stratum typicum: Main interbed, sand; Middle Miocene, floral zone VI. Remarks: The seeds were first identified as T.maii, but now can be distinguished from this species using more material. T. thielei occurs at the following sites: (1) open mine Oder, Schwandorf area, Oberpfaiz -- main interbed, sand, Middle Miocene, floral zone VI; (2) open mine Bdickelholz, Schwandorf, Oberpfalz -- upper coal seam, sand and sandy coal, Middle Miocene, floral zone VI, (3) open mine Turow near Bogatynia (Poland) -- coal seam, Middle Miocene, floral zone VI; (4) salt mine Wieliczka, Poland -- Middle Miocene, floral zone XII; (5) clay pit Niederpleis near KSln -- Upper Oligocene or Lower Miocene (one seed Inv. No.F 213 in the Geol. Inst. Univ. KSln, by courtesy of K. Kempf). At least some seeds can be mentioned which show an intermediate status and cannot be referred to any definite species. Perhaps they are new species or atypical seeds of already known types. More material must be collected to decide this question.

329

Toddalia spec.1 (Figs.52, 53) Seeds from Holzweissig and Nerchau (both Germany GDR) are p u t together into this species. They are partly worn by transport and measure a b o u t 3.8--4.1 mm in length, 2.3--2.6 mm in height and 1.6--1.8 mm in breadth. The height--length ratio is a b o u t 0.63 and the breadth--length ratio approximately 0.42. They have a prominent raphe (like T. rhenana) and a long and flat hilar scar. Nerchau has Middle Oligocene argillaceous sand and Holzweissig, the so-called "Bitterfelder D e c k t o n " , a clay of floral zone III (Lower Miocene). I have to thank D.H. Mai, Berlin, for his kindness in letting me study this material.

Toddalia spec.2 (Figs.18, 42) One seed from open mine Oder II (BBI Schwandorf) is larger than those of

T. thielei, smaller than those of T.latisiliquata and has a small hilum, a small raphe excrescence and a smooth surface (Inv. No.1970 X 128, Bayerische Staatssammlung fi]r Pal~iontologie und historische Geologie, Mfinchen). It resembles a seed from open mine Zukunft-West near Eschwefler with nearly the same measurements. Both localities yielded sands of Middle Miocene age, i.e. floral zone VI.

Toddalia spec.3 (div. spec.) (Figs.43--49) Seeds of this (these) Oligocene species cannot be referred to any fossil species, because they are not actual seeds, b u t seed casts, composed mainly of phosphate (other elements are A1, Si, Ca, Fe and K, Ti, S -- this research was kindly done by K. Weber-Dieffenbach from the Institut ffir Allgemeine und angewandte Geologie, University of Munich with the energy dispersive X-ray fluorescence analysis system (Ortec Tefa 6111). They clearly show the size and roughly the shape of the actual seeds, b u t no further identification marks, such as raphe, micropyle or hilar scar are present. The infillings are reniform, curved, elongate and the seeds must have had a size of 2.2--4.8 mm in length and 1.2--2.5 mm in height. The surface corresponds to the inner wall of the actual seeds (polygonal structure Figs.61, 62). They are known only from some fissure-fillings in Upper Jurassic Limestone of the "Schw~ibische Alb" near the Ries Crater. Animal fossils like Pseudosciurus suevicus Hensel, Anoplotherium pompeckji Dietrich and Plagiolophus fraasi (H.v. Meyer), all from MShren 19, and Palaeotherium duvali Pomel, Protapirus priscus Filhol and Sciurodon cadurcense Schlosser, the latter from MShren 20 (compare to b o t h localities Heissig and Schmidt-Kittler, 1975 and 1976) are associated with the plant remains. The above mentioned Lower to Middle Oligocene faunas are not y e t published. Five different types (species?) can be separated:

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Type 1: This first type is very small, 2 × 2 mm, strongly curved and reniform like a U, mostly symmetrical (Figs.43, 44). Type 2: The second is closely related to the first, but bigger, 2.5 X 2.0 mm, more concave and not so strongly curved (Fig.45). Type 3: This is the biggest type with a length of 3.5--4.5 m m and a height of 1.5--2.0 mm, less curved to elongated, round in transverse section and sometimes apical-laterally flattened (Figs.48, 49). Type 4: This form measures 3.0 X 2.0 mm, is larger and thicker than type 2, but must be closely related to type 3. There are only three specimens from Burgrnagerbein 3 (Fig.46). Type 5: Three typical, fish-hook-like specimens are derived from the fissure-filling Haag 2. The shape is analogous to type 3 but with a sharp apical end like a needle. The size is 3.2 X 1.4 mm. Perhaps this type belongs to type 3 (Fig.47). The material of the following fissure-fillings is in the Bayerische Staatssammlung fiJr Pal~iontologie und historische Geologie MiJnchen. (I have to thank R. Dehm and K. Heissig of this Institution for the possibility to work with their material): Burgmagerbein 3 (Bissingen), Inv. No.1972 XIV 41--47 Burgmagerbein 5 (Bissingen), Inv. No.1975 XXV 1 Haag 2 (Trenchtlingen), Inv. No.1975 XXIII 41 MShren 19 (Treuchtlingen), Inv. No.1974 XXV 31 MShren 20 (Treuchtlingen), Inv. No.1975 XII 41

Lower Lower Lower Lower Lower

to Middle Oligocene to Middle Oligocene to Middle Oligocene Oligocene Oligocene

Annotated enumeration o f fossils, included in the genus Toddalia Jussieu The following list contains effective and non-effective publications of different Toddalia types. Because of a lack of clear definitions about "effective" publications, the works of Gregor (1975a and b), Hol:~ (1974) and Knobloch and Konzalova {1977) are thought here as non-effective publications, in order to make the descriptions of the different species clearer. (1) Carpolithus latisiliquatus (Ludwig) Mai (1964), Pal~iontol. Abh., B, 2, 1, p.l18, pl.XVI, fig.12. Fig.50. Toddalia naviculaeformis (E.M. Reid) Gregor -- open mine Briickelholz (Bayer. Braunkohlen-Industrie AG Schwandorf), Middle Miocene, floral zone VI; X 10. Seed from the side, raphe to the left (Inv. No.1970 X 129, Bayerische Staatssammlung fiir Pal~iontologie und historische Geologie, Miinchen). Fig.51. Toddalia turovensis (Czeczott and Skirgiello) Gregor -- open mine Briickelholz (Bayer.Braunkohlen-Industrie AG Schwandorf), Middle Miocene, floral zone VI; × 10. Seed from lateral, raphe to the left (Inv. No.1970 X 459, Bayerische Staatssammlung fiir Pal~iontologie und historische Geologie, Miinchen). Fig.52. Toddalia sp.1 -- Nerchau (GDR), sands, Middle Oligocene; x 12. a. From the side, raphe to the right, b. From above. Fig.53. Toddalia sp.1 -- Holzweissig (GDR), "Bitterfelder Deckton", floral zone III, Lower Miocene; x 12. a. F r o m the side, raphe left. b. From above.

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333 Middle Miocene, floral zone VI; clay pit of brickyard Hartau, Oberlausitz, Germany GDR. Remarks: The single seed measures 6 X 4 mm and is the only fossil of this type found up to n o w in the Lausitz (see No.19). (2) Cytisus latisiliquata Ludwig (1860), Palaeontographica, B, 8, 2--4, p.145, pl.58, figs.14, 17. Middle Miocene, floral zone VI (to VIII); oldest part of the Wetterau browncoal; Hessenbrficker Hammer near Laubach (Vogelsberg), Germany GFR. Remarks: Ludwig described husks and seeds under the same name, b u t mentioned that the seeds looked t o o small for the fruits (see No.19). (3) Cytisus latisiliquata Ludwig vel Martyia naviculaeformis E.M. Reid-Zagwijn (1959), Fortschr. Geol. Rhld. Westf., 4, pl.1, fig.6. Pliocene, Brunssum clay; Brunssum, The Netherlands. Remarks: The 5 mm long seed belongs to T.rhenana Gregor (see p.325). (4) cf. Cytisus latisiliquata Ludwig: Bu~ek and Hol:~ (1964), Sb. Geol. Ust. Csl., Paleontol., 4, p.120, pl.6, fig.5, Text-fig.2--9a--c. Lower to Middle Miocene, Burdigalian--Helvetian; Boring Cernovice Cv62 d, Chomutov--Most--Teplice Basin, C.S.S.R. Remarks: The seeds, measuring 3.0 X 2.2 mm, u n d o u b t e d l y belong to T. mail Gregor from the Schwandorf browncoal (see Nos.22, 23, 25). (5) Epipremnum crassum C. and E.M. Reid: Mfiller (1972), Die Oligoz~inAblagerungen im Gebiet des NSrdlinger Rieses, p.77. Middle Oligocene, Websteri-browncoal horizon; M ~ k e r Cement Company, Harburg i.Ries, Germany GFR. Remarks: The single specimen is described on p.322 as Toddalia maerkeri nov. spec. The original Epipremnum material from the Dutch--Prussian border (Reid and Reid, 1915, p. 71, 72) has no similarities with Toddalia seeds, (6) Leguminosae g.(?): E.M. Reid (1920), Bull. Soc. G~ol. Fr., Ser. 4, 20, p. 69, 70, pl.III, figs. 29a, b. Lower Pliocene; Pont-de-Gall, Cantal, France. Fig.54--58. SEM (Institut ffir Raster-Elektronenmikroskopie, H. Klingele, Munich). Fig.54. A half seed of a fossil T.maii in full sight showing the shell-wall and the inner surface, poorly preserved without tegmen (compare Fig.63). The square shows the hypothetical tegmen (taken from a recent Toddalia asiatica compare Figs.55, 56); X 40. Fig.55. Tegmen of a recent T.asiatica (Tsui, Canton, China) with criss-crossed spiral lignification; X 90. Fig.56. See Fig.55, but X 580. Fig. 57. Tegmen of a fossil T. mail (Schwandorf browncoal) with criss-crossed spiral lignification; × 580. Fig.58. Inner testa wall with remnants of the criss-crossed spiral lignification (worse preservation, see Fig.63); x 580.

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C,O 4~

335

Remarks: In 1923 the seeds were named Martyia naviculaeformis Reid (1923) (see N o . l l ) . (7) Leguminosae: Kirchheimer (1936), Zentralbl. Min. etc., 1936, pp. 345-346. Upper Miocene; browncoal mine H a i d h o f between Regensburg and S c h w a n d o r f (Oberpfalz, Ger m any GFR). Remarks: Some seeds of Toddalia t y p e have been found at this site, but the materiN is lost (?). T he 6 m m long seeds perhaps belonged to T.latisiliquata (see No.19). In m y opinion t h e y were f ound in Viehhausen (No.9). (8) Leguminosae: Kirchheimer ( 1957), Die L aubge w~ichse der Braun kohlenzeit, pp.211, 412. Remarks: Th e aut hor links Cytisus latisiliquata with the Dalbergieae and expresses suspicion a b o u t the c o n n e c t i o n of fossil Martyia naviculaeformis with the legumes. (9) Leguminosae: Kirchheimer (1937), Grundziige einer Pflanzenkunde der deutschen Braunkohlen, p.75, Abb.86. Remarks: Here the a u t h o r compares different types of leguminosae-like seeds and states the character of the Cantal, Viehhausen and Habichtswald fossils. The seeds f r o m Viehhausen are 5.8--6.4 m m long. (10) Leguminosae: Zeidler (1938), Palaeontographica, B.83, 4--6, p.197. Upper Miocene, T or t oni an; browncoal mine Viehhausen near Regensburg, G e r m a n y GFR. Remarks: The 4 m m long seeds (after Zeidler) are in reality 6 m m long and 4 m m high and u n d o u b t e d l y related to T. latisiliquata. On the n e x t pages, the remaining material from Wiirzburg is described as T.latisiliquata var. viehhausenensis (see p.337). (11) Martyia naviculaeformis E.M. Reid (1923), Bull. Soc. G~ol. Fr., S~r. 4., 23, p p .3 2 7 - - 3 3 1 , fig.5a. L o w e r Pliocene; Pont-de-Gaff, Cantal, France. Remarks: The original material was b o r r o w e d from the British Museum Nat. Hist. ( L o n d o n ) and newly c o m b i n e d with T.naviculaeformis (E.M. Reid) Gregor (see No.30). Fig.59. Surface of a fossil Toddalia mail (Schwandorf browncoal); × 500. Fig.60. Surface of a recent T.asiatica (L.)Lamk. (Mt. Elgon, Kenya); x 500. Fig.61. Fossil T. sp.3--type 1 (1972 XIV 45) with a typical "honeycomb"-structure, corresponding with the structure of the inner surface of the seed shell (Fig.63); x 280. Fig.62. The same surface as in Fig.61, but magnified x 750. Fig.63. Inner surface (bad preservation by fossilization) of a seed shell of fossil T. maii below the tegmen with "honeycomb"-structure; x 750. Fig.64. Cross-section of the fossil shell wall of T. maii; with lower inner mesophyll and upper outer mesophyll (compare Fig.8); × 140. Figs.59--64. SEM (Institut fiir Raster-Elektronenmikroskopie, H. Klingele, Munich).

336 (12) Robinia hesperidum Unger (1864), Denkschr. Akad. Wiss. Wien, Math.Nat. CI., 22, p.21, pl.IV, fig.14. Oliogocene; Parschlug (Steiermark), Austria. Remarks: Unger combined fruits and seeds under the same name, which is wrong in the nomenclatural sense. The seeds of Fig.14 could be of Toddalia type, but without original material there is no possibility of saying anything about it. According to Kirchheimer (1957, p.291) they have no botanical value. (13) Sapoticarpum turovense Czeczott and Skirgiello (1975), Prace Muzeum Ziemi, 24, pp.44--45, 55--56, pl.XI, figs.l--14. Middle Miocene, floral zone VI; open mine Turow near Bogatynia, Poland. Remarks: In the Polish text the holotype is signified on pl.XI, figs.l--14, in the English text on pl.XI, figs.l--9 (?). The enlargement of figs.4--14 is given as natural size, but actually it is times two. The species was newly combined to T.turovensis (Czeczott and Skirgiello) Gregor (Acta Palaeobot., 1978). The raceboat-shaped seeds are very typical (see No.33). (14) Toddalia: Knobloch and Konzalova (1977), Progress in Cenophytic Palaeobotany of Czechoslovakia, In: Advances in Angiosperm Palaeobotany, pp.12, 17, 19. Eggenburgian -- north Bohemian browncoal. Karpathian -- Cypris clay in the Cheb and Sokolov Basins. Badenian -- Mikulov, south Moravian Basin. Remarks: New investigations in the C.S.S.R. identified many sites with fossil Toddalia-seeds, which will be published separately (uneffective publication). (15) Toddalia: Knobloch and Konzalova, Cour. Forsch.-Inst. Senckenberg, 34, pp.45, 49, 50, 51. Eggenburgian -- north Bohemian browncoal. Karpathian -- Cypris clay in the Cheb and Sokolov Basins. B a d e n i a n - Mikulov, south Moravian Basin. Remarks: In contrast to the same article (No.14) this is an effective publication. No species are mentioned. (16) cf. Toddalia aculeata Persoon: Menzel (1913), Beitr. Geol. Erforsch. Dtsch. Schutzgeb., 18, II, p.27. Pleistocene, Holocene, young basaltic tuffites, Etome, Cameroun, Africa. Remarks: The fossil leaf imprints resemble the recent leaves of T.aculeata Persoon. They are in the collection of the Naturhistoriska Riksmuseet, Stockholm (Sweden). (17) Toddalia latisiliquata (Ludwig) Holy (1974), The Neogene Mastixioideae flora from the upper layers of open pit Kristina (Hradek n.N.), pp.64, 65, pl.12, figs.8, 9.

337 Middle Miocene, floral zone VI, Karpathian (?); open cast mine Kristina, Hradek-Basin, C.S.S.R. Remarks: The new combination of this species was done independently from m y work in 1974, b u t in a non-effective publication. (18) Toddalia latisiliquata (Ludwig) Gregor (1975a), Cour. Forsch.-Inst. Senckenberg, 13, p.126. Middle Miocene, floral zone VI (to VIII ?), oldest Wetterau browncoal; Hessenbrficker Hammer near Laubach (Wetterau), Germany GFR. Remarks: Gregor's fig.6 on p.126 belongs to this new combination and not to "T. naviculaeformis". (19) Toddalia latisiliquata (Ludwig) Gregor (1978a), Acta Palaeobot., XIX, pp. 25, 26, pl.II, figs.3--5, pl.III, figs.l--6, pl.IV, figs.I--6, pl.V. fig.1. Middle Miocene, floral zone VI (to VIII ?), oldest Wetterau browncoal; Hessenbriicker Hammer near Laubach (Wetterau), Germany GFR. Remarks: Original material from the collections of A1. Braun and R. Ludwig could be surveyed by courtesy of H. J~ihnichen (Museum ffir Naturkunde -Pal~iontologisches Museum, Berlin). T.latisiliquata is found at following sites: (1) Haidhof (?), Regensburg (Oberpfalz); Germany G F R -- Upper Miocene (material?). (2) claypit Hartau, ZittauBasin, Germany GDR -- Middle Miocene, floral zone VI. (3) Salt mine Wieliczka, Poland -- Middle Miocene, floral zone XII (material was given for the research by the courtesy of J. Zablocki, Torun, and M. LancuckaSrodoniowa, Krakow, Instytut Botaniki). (4) Mine Ludwigszeche near Viehhausen, Regensburg -- Upper Miocene, Lower Tortonian. (5) Schwandorf browncoal (Oberpfalz), Germany G F R -- Middle Miocene, floral zone VI; open pits Oder and Brfickelholz (main interbed--sand and upper coal seam-sand). (6) Open mine Kristina, Hradek Basin C.S.S.R.; Middle Miocene, floral zone VI (Karpathian?). (7) Povoa de Santarem, Lisbon, Portugal -- Middle Miocene. (8) Open cast mine Zukunft-West near Eschweiler, Germany G F R (equivalents of the Neurather Sands -- Middle Miocene, floral zone VI. (20) Toddalia latisiliquata (Ludwig) Gregor (1978b), Palaeontographica, B, 167, 1--3, p.43, Taf. 9, fig. 10. Middle Miocene, floral zone VI; open mine Oder II near Schwandorf (Oberpfalz). Remarks: This seed was wrongly determined as T.naviculaeformis (see No.18). (21) Toddalia latisiliquata (Ludwig) Gregor (1979), Palaeontographica, B, in press, fig.1. Upper Miocene, Lower Tortonian; Mine Ludwigszeche near Viehhausen, Regensburg, Germany GFR. Remarks: The seeds lie in a piece of coal and belong to the Coll. Kirchheimer (Wfirzburg). They are described above (p.322) as var. viehhausenensis nov.var.

338 (22) Toddalia maii Gregor (1975a), Cour. Forsch. Inst. Senckenberg, 13, p.125, fig.5. Middle Miocene, floral zone VI; open mine Oder II near Schwandorf (Oberpfalz), Germany GFR. Remarks: This species is published in another non-effective work (1975b; an effective publication is in press for 1978 (see No.26). (23) Toddalia maii Gregor (1975b), Die mittelmioz~ine Mastixioideen-flora aus dem Braunkohlen-Tagebau Oder II bei Wackersdorf (Oberpfalz), pp.130-132, pl.7, figs.l, 2. Middle Miocene, floral zone VI, open mine Oder II near Schwandorf (Oberpfalz), Germany GFR. Remarks: This work is thought to be non-effective like No.22. (24) Toddalia mail Gregor--Gregor and Jung (1977), Bayerischer Braunkohlen Bergbau, 102, S.9, Taf.I, fig.11. Middle Miocene, floral zone VI; open mine Oder near Schwandorf. Remarks: The picture shows the holotype. (25) Toddalia maii Gregor (1978a), Acta Palaeobot., XIX, pp. 26, 27, figs. 2--4, pl.VI, figs.5--7. Middle Miocene, floral zone VI; open mine Oder II near Schwandorf (Oberpfalz), Germany GFR (Fig.35). Remarks: The description is the same as in Nos.22 and 23. T.maii occurs at the following sites: (1) Grandes Carri~res, Soufflenheim, Alsace, France -- Lower Pliocene, older Reuverian. (2) Haguenau, Alsace, France (borings Bischwiller--Kaltenhouse, Airport Haguenau and Schwighouse, + 32 m) -- Lower Pliocene, Brunssumian. {3) open mine Turow, Poland, coal-- Middle Miocene, floral zone VI. (4) Open clay-pit Niederpleis near KSln, Germany GFR, lower browncoal -- Upper Oligocene or Lower Miocene. (5) Autobahn-pit Langenau near Ulm, Germany GFR -- Upper Freshwater Molasse (Helvetian--Tortonian), Upper Miocene. (6) Salt mine Wieliczka, Poland -- Middle Miocene, floral zone XII. (7) Schwandorf browncoal (Oberpfalz), Germany GFR -- Middle Miocene, floral zone VI; open pits Bri]ckelholz (upper coal seam--sands), Hofenstetten (main interbed--clay) and Nordfeld (upper coal seam--quartzite). (8) Boring Cv 62d, Cernovice, Chomutov-Most--Teplice Basin, C.S.S.R. -- Lower to Middle Miocene (Burdigalian-Helvetian). (9) Povoa de Santarem, Lisbon, Portugal -- Middle Miocene. (10) Klettwitz, Germany GDR, Spezialton -- Upper Miocene, floral zone X. (11) Mine Ludwigszeche near Viehhausen, R e g e n s b u r g - Upper Miocene, Lower Tortonian. (12) Undorf near Regensburg -- Middle Miocene (?), Karpathian (?), Tortonian (?). (26) Toddalia mail Gregor (1978b), Palaeontographica, B, 167, 1--3, p.43, Taf. 9, fig. 9. Middle Miocene, floral zone VI; open mine Oder II near Schwandorf (Oberpfalz), Germany GFR.

339

Remarks: Here the holotype is shown (see No.25). (27) Toddalia mail Gregor (1979), Palaeontographica, B, in press. Upper Miocene (Lower Tortonian) -- Viehhausen near Regensburg. Middle Miocene (Karpathian?Tortonian?) -- Undorf near Regensburg. Remarks: Two fragmentary seeds were found at Undorf and Viehhausen in browncoal, some at Hofenstetten in clays. (28) Toddalia naviculaeformis (E.M. Reid) Gregor, Cour. Forsch.-Inst. Senckenberg, 13 (1975a), p.125, 126, Abb.7. Middle Miocene, floral zone VI; open cast mine Oder and Briickelholz, Schwandorf (Oberpfalz), Germany GFR. Remarks: The species occurs abundantly in many European sites, but was not newly combined on the holotype. Fig.6 is not T.naviculaeformis, but T.latisiliquata (see No.18 above). (29) Toddalia naviculaeformis (E.M. Reid) Gregor (1975b), Die mittelmioz~ine Mastixioideen-Flora aus dem Braunkohlen-Tagebau Oder II bei Wackersdorf (Oberpfalz), pp.132--135, pl.7, figs.4,5, text-figs.44, 46, 47 (not pl.7, fig.3). Middle Miocene, floral zone VI; open mine Oder II near Schwandorf (Oberpfalz), Germany GFR. Remarks: This work, like in No. 28, is thought to be non-effective and the new combination depends on material from other sites than the one to which the holotype belongs. The seed (Inv. No.1970 X 127) is not T.naviculaeformis but belongs to T.latisiliquata (compare No.18). (30) Toddalia naviculaeformis (E.M. Reid) Gregor (1978a), Acta Palaeobot., XIX, pp. 27, 28, pl.V, figs.5--7, pl.VI, fig.1. Lower Pliocene, Pont (Susterian); Pont-de-Gail, Cantal, France. Remarks: The original material, which is housed in the British Museum (Nat. Hist.), London, could be studied by the courtesy of C.H. Shute and H.W. Ball (Inv. No.: V25667 and V25585) T.naviculaeformis is present at: (1) Wieliczka in Poland, salt mine, Middle Miocene, floral zone XII. (2) Open mine Zukunft-West near Eschweiler, equivalents of Neurather Sands; Middle Miocene, floral zone VI. (3) Habichtswald near Kassel, lower strata under coal seam; Upper Miocene (material ?); (4) Schwandorf, Oberpfalz; Middle Miocene, floral zone VI; open mine Oder II (main interbed--sand); open mine Brfickelholz (upper coal seam--sand). (31) Toddalia naviculaeformis (E.M. Reid) Gregor (1978b), Palaeontographica, B, 167, 1--3 p. 43, Taf. 9, fig. 12. Middle Miocene, floral zone VI; open mine Briickelholz near Schwandorf (Oberpfalz), Germany GFR. Remarks: This type of seed is very common in Brfickelholz.

340 (32) Toddalia turovensis (Czeczott and Skirgiello) Gregor (nom. nud.) -Gregor and Jung (1977), Bayerischer Braunkohlen Bergbau, 102, Taf.I, fig.12. Middle Miocene, floral zone VI; open mine Briickelholz near Schwandorf. Remarks: Only one seed is shown. (33) Toddalia turovensis (Czeczott and Skirgiello) Gregor (1978a), Acta Palaeobot., XIX, pp. 29, 30, pl.VI, figs.2--4. Middle Miocene, floral zone VI; open cast mine Turow near Bogatynia, Poland. Remarks: The raceboat-shaped species is closely related to T.naviculaeformis and occurs in Turow, Schwandorf (Fig.51) and Eschweiler. For other remarks see no.13. (34) Toddalia turovensis (Czeczott and Skirgiello) Gregor (1978b) Palaeontographica, B167, 1--3, p.43, Taf.9, fig.ll. Middle Miocene, floral zone VI; open mine Briickelholz near Schwandorf (Oberpfalz), Germany GFR. Remarks: Badly preserved specimens of this species can mistakenly be taken for T. naviculaeformis. (35) Toddalia sp.: Geissert, Menillet and Farjanel (1977), Bad. Landesver. Naturk. Natursch. Freiburg i. Br., in press. Pliocene, Reuverian; boring Airport Haguenau (Alsace), France. Remarks: These T. maii type seeds occur in many deposits in Alsace (France). (36) Toddalia sp.: Gregor (1979), Palaeontographica, B, in press. Middle Miocene, floral zone VI; open mine Hofenstetten near Schwandorf (Oberpfalz), Germany GFR. Remarks: The badly preserved seeds perhaps belong to T. latisiliquata or T. naviculaeformis.

(37) Toddaliospermum excavatum Chandler (1963), The Lower Tertiary Floras of Southern England III, Flora of the Bournemouth Beds; the Boscombe, and the Highcliff Sands p.93, p1.14, figs.31--34. Highcliff Sands, Auversian, Eocene; Cliff End, Mudeford, England. Remarks: This species is newly combined with Toddalia excavata in this work (see p.321). List of fossils, excluded from the genus Toddalia Jussieu

(1) Toddalia maii Gregor (1975b), Die mittelmioziine Mastixioideen-Flora aus dem Tagebau Oder II bei Wackersdorf (Oberpfalz), pp.125, 127, Taf.10, abb.41 (Arjuzanx). Middle Miocene, Helvetian--Tortonian; Arjuzanx near Morcenx, S France. Remarks: This lonely seed belongs to the genus Clausenopsis Engler and will be published separately.

341 (2) Toddaliospermum: Krumbiegel (1968), Dtsch. Ges. Geol. Wiss., Das Geiseltal, p.62. Eocene browncoal; Geiseltal near Halle-Saale, Germany GDR. Remarks: The fossils are only related to Rutaceae (?), but have no similarities with any Toddalia species (compare Toddaliospermum leonhardii and Toddaliospermum neomarcense, no.3 and 4). (3) Toddaliospermum leonhardii Mai (1976), Abh. Zentr. Geol. Inst., 26 p.116, pl.V, figs.17--23. Middle Eocene, upper middle coal; open mine Leonhardt, Geiseltal near Halle-Saale, Germany GDR. Remarks: This species does not belong to the genus Toddalia b u t perhaps to a related form. (4) Toddaliospermum neomarcense Mai (1976), Abh. Zentr. Geol. Inst., 26, p . l 1 7 , pl.V, figs.10--16. Middle Eocene, middle coal; open cast mine Neumarkt-Siid, Geiseltal near Halle-Saale, Germany GDR. Remarks: This species, like the foregoing (No.3), is perhaps a rutaceous form, but not Toddalia. (5) Toddaliospermum ornatum Chandler (1963), The Lower Tertiary Floras of Southern England III, Flora of the Bournemouth Beds; the Boscombe, and the Highcliff Sands, pp.92, 93, pl.14, figs.29, 30. Upper Eocene, Barton (Led), Highcliff Sands and Boscombe Sands, Cliff End, Mudeford and Southbourne, England. Remarks: The seeds do not resemble those of any Toddalia species, but surely are closely related. Especially the genuses Vepris Jussieu or some Fagara Linn~ types (Zanthoxyleae) have to be examined; b u t Clausenopsis Engl. is the genus with which it can be best compared. SPECIAL PART

Presentation of data In the following part the results are given of the comparisons between the Recent and the fossil species. They clarify the biostratigraphic and systematic relationships of the fossils and demonstrate the whole paleoecological setting, including lithological aspects, sociology, climatology and geography. All the reconstructions are made under the assumption, that the distributions of Recent and fossil Toddalia plants in a biocenosis are not significantly different. In order to compare the most important Recent and fossil species with one another and to coordinate newly discovered seeds with these species, a series is shown in natural size (Figs.19--27).

342

C o m p a r i s o n o f f o s s i l m a t e r i a l w i t h s p e c i a l data I n T a b l e I, s p e c i e s a r e e n u m e r a t e d w i t h t h e i r m e a s u r e m e n t s ( l e n g t h , h e i g h t , breadth), notes about the vascular bundle over the chalaza, the reconstructed number of seeds per fruit, and some special remarks. Fig.65 shows the relation height--length. TABLE I Fossil Toddalia species and their morphology Species

Length

Height

Breadth

Vascular bundle

Seeds per fruit

Special remarks

T.excavata

1.55

1.11

0.8

--

4.5

very small squat

T.latisiliquata

5.5--7.0 6.2

3.8--4.5 4.1

2.5--4.0 3.2

--

4.5

boatshaped

4.0--5.5 5.0

3.4--3.8 3.6

2.0--2.5 2.2

_

5.6

var. gracilis

boat-shaped squat, pitted surface

T.latisiliquata vat. viehhausenensis

5.3--6.0 5.5

3.3--4.0 3.5

2.3--2.6 2.4

5.6

boat-shaped

T. maerkeri

3.5

1.9

1.9

+

4.5

long-rapheexcrescence

T.maii

2.5--3.8 3.2

2.0--2.5 2.3

1.5--2.2 1.9

--

2--4

squat

T.maii var. minor

2.0--3.2 2.6

1.6--2.2 1.9

1.2--1.8 1.5

3.4

squat

T. naviculaeformis

6.0--10.0 3.0--5.5 7.5 4.2

2.5--3.5 3.1

6.7 (reduction!)

boat-shaped

T.rhenana

3.3--4.5 3.9

2.2--2.7 2.5

1.4--2.0 1.7

--

5.6

very elongated

4.5--5.4 4.8

2.7--3.2 2.9

1.8--2.0 1.9

_

5.6

T. thielei

3.5--4.4 3.9

2.4--3.0 2.5

1.8--2.2 1.9

--

4.5

T.turovensis

7.5--9.5 8.7

4.5--5.0 4.8

2.5--2.8 2.6

(+)--

8 (reduction ! )

very slender raceboatshaped

T. s p a

3.8--4.1

2.3--2.6

1.6--1.8

--

6.7

boat-shaped

T. sp.2

4.9--5.5

2.9--3.1

2.3

+

5.6

short rapheexcrescence

T.latisiliquata

T.rhenana var. major

-

-

-

-

-

-

big, elongated symmetric squat, pitted surface

343

Height 5.0

/,.0

2.0

cP i 2.0

,

,

l,.O

l

o

62

o

~

8,O

,

,

Length

I0,0

Fig.65. Graph, showing the relation between length and height of various fossil T o d d a l i a species. The numbers indicate the following species: 1 = T . e x c a v a t a ; 2 = T.latisiliquata; 3 = T.latisiliquata var. gracilis; 4 = T.latisiliquata vat. v i e h h a u s e n e n s i s ; 5 = T . m a e r k e r i ; 6 = T . m a i i ; 7 = T . m a i i var. m i n o r ; 8 = T . n a v i c u l a e f o r m i s ; 9 = T . r h e n a n a ; 10 = T . r h e n a n a var. m a j o r ; 11 = T. thielei; 12 = T. turovensis.

C o m p a r i s o n s o f the fossil T o d d a l i a species f r o m various localities with the d i f f e r e n t r e c e n t species o f T o d d a l i a a r e m a d e in Table II. TABLE II List of fossil T o d d a l i a species with comparable Recent species (with locality) Fossil species

Recent species (locality)

T excavata

no such small seeds yet known

T. la tisiliq ua ta

T.asiatica (Tananarive, Madagascar) T.asiatica (Phetchabun, Thailand) T . a c u l e a t a (Dehra Dun, India) T. asia tica ( K w a i Noi River, Thailand )

(Madras, India)

T. m a e r k e r i

T.asiatica var. gracilis

T. maii

T.asiatica T.asiatica T.asiatica T.asiatica T. asia tica

(Hiniduma, Ceylon) (Hongkong, China) (Kwangtung, China) (Nepal, India) (Kyimbila, Nyassa)

T. n a v i c u l a e f o r m i s

T.asiatica

(Coonoor, Madras, India)

T. r h e n a n a

T.asiatica

(Kivu, Congo)

T. thielei

T.asiatica var. o b t u s i f o l i a

T. t u r o v e n s i s

no direct comparison possible but close are T.asiatica (see from Madras, India)

(Kukul Betta, Madras, India)

344 Key to the Toddalia types Size, shape, hilum, raphe, and indices (HLI = height--length index; BLI = breadth--length index, Table III) are taken as morphological characteristics for differentiating the fossil Toddalia species in a key. A. i.

.

S. I. .

.

C. I. 2.

Seeds small, 1.5--3.8 m m long, hilum triangular Without curved vascular bundle over chalaza, thick, reniform, HLI over 0.70; BLI over 0.50 a. Seeds very small, 1.55 m m long . . . . . . . . . . T.excavata b. Seeds larger, 3.2 × 2.4 m m . . . . . . . . . . . . . T.maii With curved vascular bundle over chalaza, r o u n d in transverse section, HLI and BLI a b o u t 0.55 a. Seeds curved, with raphe-excrescence 3.5 X 1.9 m m . . T.maerkeri Seeds medi um size, 3.9--7.0 m m long, hilum triangular to elongate Seeds distinctly elongated, 4.0 × 2.5 mm, boat-shaped, hilum symmetrical with p r o m i n e n t raphe, BLI 0.43; HLI 0.64 .... T.rhenana Seeds stout and thick with flattened lateral sides, straight ventral margin and asymmetrical hilum, 3.9 X 2.5 mm; HLI 0.65; BLI 0.51 . . . . . . . . . . . . . . . . . . . . . . . . . T. thielei Seeds large and very variable, with elongated hilum, oft en thick and round, 6.2 × 4.1 mm; HLI 0.63--0.71; BLI 0.36--0.44 . T.latisiliquata Seeds of large size, 7.0--11.0 mm, hilum extending nearly the whole length of the seeds. Seeds robust, with very p r o m i n e n t raphe (often abraded), 7.5 X 4.2 mm; BLI 0.41 . . . . . . . . . . . . . . T.naviculaeformis Seeds raceboat-shaped, with straight ventral margin, 8.7 X 4.8 mm; BLI 0.29 . . . . . . . . . . . . . . . . . . T.turovensis

Graphs and indices On the graphs one can see the populations of fossil Toddalia seeds in relation to one another. Fig.66 shows the mean height--length and breadth-length indices o f the different species. The term " i n d e x " means the percentage relation o f two measurements, in this case the smaller is used as the dividend, the larger as the divisor. The indices are shown in Table III and on Fig.66. Biostratigraphy In Fig.67 the di f f er e nt fossil species of Toddalia are shown in stratigraphic position and duration. At the first glance it is clear t hat the genus is very old (Eocene) and extends in E u r o p e to Early Pliocene, having a m a x i m u m of occurrence in Middle Miocene times. Some species like T. maii have a long duration f r o m Late Oligocene to Early Pliocene times, others like T.excavata flourished for only a short time. Some time-gaps with missing fossils surely

345

depend on rarity of localities and seeds will be found in the future (especially in the Lower Miocene).

Breadth-Length-Index

0.60

o

@

0.50

0./,0

@

0,30

, 0.50

, 0.60

,

,

,

0.70

, Heighf-Lengfh-lndex 0.80

Fig.66. T h e d i f f e r e n t Toddalia-species a n d t h e i r indices. T h e n u m b e r s are t h e same as in Fig.60. T A B L E III T h e fossil Toddalia species a n d t h e i r indices

T.excavata T. latisiliqua ta T. latisiliquata var. gracilis var. viehhausenensis T. m a e r k e r i T.maii var. m i n o r T. n a v i c u l a e f o r m i s T. rhenana vat. m a j o r T. thielei T. turovensis T. sp.1 T. sp.2

Height--length index

Breadth--length index

0.71 0.66 0.63 0.72 0.63 0.57 0.71 0.73 0.56 0.64 0.64 0.65 0.55 0.64 0.58

0.51 0.51 0.36 0.44 0.43 0.55 0.59 0.57 0.41 0.43 0.41 0.49 0.29 0.42 0.44

346

Epoch

Species

Eocene

Oligocene

L

M

Miocene

U

T. excovoto I tofisiliqafo T. moerkeri

m Illll

T. moil

II II II

T. navicutoeformis T. rhenono T. thietei

7

mill

T. turovensJs T. spec. I T. spec. 2 T. spec. 3

Fig.67. Stratigraphical scheme, showing the different fossil Toddalia species in their occurrence in the Tertiary. Some reflections may be allowed here a b o u t Toddalia fossils missing where t h e y could be expected. For example in Salzhausen or in some i m p o r t a n t i nt er beds of the Rhenish browncoal (Diiren or Herzogenrath) the fossils are missing. The same occurs in Wiesa (GDR), where Toddalia, in m y opinion, must have grown in Middle Miocene times. Whether these are stratigraphic questions or problems of the facies is n o t possible to say at the m o m e n t . Following Mai (1965, 1967) and Ahrens and Lotsch (1967), the same graph is p u t forward for the floral zones (Fig.68) and the occurrence of Toddalia in these zones. It is clear t hat zone VI which yields the richest floras, must also have the mo s t Toddalia fossils. T.maii is abundant in Neogene sediments of all floral zones and does n o t indicate a special one, whereas T. turovensis, for example, is restricted to zone VI and T. thielei to zones VI, XII and probabl y zones I--III. The species com pos i t i on is also very i m p o r t a n t for different ages. Thus for e x amp le, the Middle Miocene zone VI is characterized by m any different species whereas in the Upper Miocene T.latisiliquata is c o m m o n e s t , only accompanied by o t h e r rare occurrences. In the Lo wer Pliocene, only three species exist, two with small seeds (T.maii and T.rhenana), f o u n d only in Alsace, and at Brunssum and one with large seeds (T.naviculaeformis), f o u n d at Pont-de-Gail. Here I have neither the space nor the intention to write a b o u t the problems of stratigraphy, especially the Upper Miocene--Pliocene b o u n d a r y (compare Steininger et al., 1976). It

347

Epoch

Otigocene

Miocene

F[orazones

Species

7

I IlI

I

Ill

1

I

IV I

vii ~vIII IX

x

Xl

XIII

T. [atisiliquQtn

-i

T. mQii

T novicu[Qeformis

I

--

T. rhenano T, fhietei

? J ?

T. furovensis

?

--F--__L__

I spec. 1 _

I spec. 2

Fig.68. The "Biostratigraphical Zones" (after Mai, 1965, 1967) and their content of fossil Toddalia species (black = sure fossil findings). can be that Pont-de-Gail is uppermost Miocene and n o t Pliocene and so the before-mentioned idea would be wrong and only a small-seeded group would remain in Pliocene times. In Middle Oligocene times the genus must have been very c o m m o n (T. sp.3 from fissure-fillings), while T. maerkeri is a rare species in the uppermost Middle Oligocene. Morphogenetic relationships The a t t e m p t to interpret the relationships of Toddalia species is, like every morphogenetic idea, very hypothetical (Fig.69). One can see different groups of similar species (similar in size, shape, etc.), but the precise relationships are obscure and must be reconstructed. Thus, the small T. maii, T. thielei and T. rhenana are grouped together, likewise the large-seeded T.naviculaeformis, T. turovensis and T. latisiliquata. The Oligocene species of Toddalia may be t h o u g h t to be two groups, ancestral to the Miocene species. The only fossil of the Eocene, T.excavata, could be the missing link with older types. Consequently, it may be expected that the whole group of Toddalia lianas existed in Palaeogene or Cretaceous times, but so far no fossils have been found. The hypothetical ancestors of Toddalia are t h o u g h t to have been unspecialised ecotypes, lianas or semiscandent scrubs, hardly dependent on a special soil, other plants or the climate (tropical to subtropical).

348

ToddQlin div. spec.

Hotocene

I I

I I I

I I

q

PLiocene

Miocene

/

\

I

%

/

/

% 7

,'7

I i ,i

Otigocene

Eocene

Paleocene

I I

ancmnf member

F i g . 6 9 . T h e m o r p h o g e n e t i c r e l a t i o n s o f t h e d i f f e r e n t fossil Toddalia s p e c i e s to o n e a n o t h e r in t h e c o u r s e o f t i m e ( b r o k e n lines s h o w lack o f d a t a ) .

349

Paleoecology In this wide field we can try to find the environmental conditions under which our fossil Toddalia species grew.

Lithological conditions Strata of different lithologies contain different species of Toddalia. Three main lithofacies will be discussed in respect of Toddalia fossils. Sands: In these allochthonous sediments the big seeds of T.naviculaeformis, T. turovensis and the small ones of T.maii are the c o m m o n e s t types. All seeds are more or less abraded, especially in T.naviculaeformis (compare Gregor, 1978a, pl.VI, fig.1 and Gregor, 1975b, p.134, fig.47), which often lack the raphe and thus resemble legume seeds. They indicate a long transportation and the plants surely belonged to dense forests surrounding the swampy vegetation. These mesophytic to evergreen forests are demonstrated by thousands of fruits and seeds of the so-called mastixioidean floras. Clays: Clays of semi- and h y p a u t o c h t h o n o u s character yield only few fossils of mainly one species -- T. maii. The accompanying vegetation indicates the existence of sparse thickets, often with Glyptostrobus and Nyssa, Spirematospermum and Magnolia. These poor floras are similar to the modern North American Pocosin woods. Coal: Especially in the coal seams of Hessenbriicken and Viehhausen many seeds of T.latisiliquata were found. This species must have grown in swamp and wetland vegetation. Perhaps Rubus-like thickets of Toddalia line the lakes, ponds, and rivers of giant swamps, sprinkled with bushy islands. Sociological reconstruction All the fossil seeds of Toddalia were found together with other seeds and fruits of various characters. In the following section the most important associated seed floras are mentioned to show the vegetational conditions, under which these Toddalia lianas grew. In Eocene times Toddalia excavata was a member of a forest type vegetation, including Calamus, conifers, rutaceous forms, Gordonia, Eurya, with Mastixia and Epacridicarpum -- the composition being of tropical to subtropical character (Chandler, 1963). In the Lower and Middle Oligocene the genus was abundant together with Vitis on the Jurassic plain of the "Schw~bische Alb" near the Ries Crater. The fossils are f o u n d especially in fissure fillings. In the same area, but in the uppermost Middle Oligocene times, there were small swamps with poor vegetation: Stratiotes websteri, Brasenia, Spirematos p e r m u m and Cladium (Mfiller, 1972). In the Miocene there are some differences in the floras between the Lower, the Middle and the Upper Miocene. From the lower part, only a few data are

350 available. The most important flora from the Chomutov--Most--Teplice Basin ( Bu~ek and Hol:~, 1964) yields Fungi, Taxodium, Sequoiadendron,

Glyptostrobus, Eurya, Myrica, Nyssa, Stratiotes, Spirematospermum wetzleri and Calamus and "Epipremnum cristatum." Here T. maii is the only species of Toddalia. In the Middle Miocene all types of Toddalia seeds are found (Tomaii, T. naviculaeformis, T. turovensis, T. rhenana, T. latisiliquata) together with rich, subtropical mastixioidean floras, composed of Mastixiaceae, Symplocaceae, Styracaceae, Theaceae, Rosaceae, and many more. The floras from Schwandorf, Eschweiler, Hartau and Turow) (compare Gregor, 1975b; Czeczott and SkirgieUo, 1975; Mai, 1964; Van der Burgh, in preparation) axe typical of this vegetation. On the other hand, the autochthonous floras with T. maii yielded species like Magnolia burseracea, Meliosma and Distylium uralensis together with Zanthoxylum, Symplocos, Carya, Nyssa and Manglietia (open mine Hofstetten, BBI Schwandorf, Gregor, 1979, in press). The upper part of the Middle Miocene, e.g. in a salt mine in Wieliczka in Poland has many new taxa accompanying T.maii, T.latisiliquata and

T.naviculaeformis: Pinus, Pterocarya, Carya, Fagus, Liquidambar, Vitis, Engelhardtia, Fortunearia, Magnolia, Pterostyrax and Olea, all of humid character (Zabtocki, 1928 and 1930). Especially the flora from the Hessenbriicker Hammer near Laubach had many Toddalia seeds together with Carya, Cinnamomum, Glyptostrobus, Magnolia, Meliosma, Nyssa, Prunus langsdorfi, Sapindoidea margaritifera and Vitis. T. latisiliquata may have been a species climbing on trees in gallery forests and swampy environments (Mai, 1964). The same facies can be postulated for Viehhausen near Regensburg, where the seeds of this species occur together with Spondiaecarpum, Chionanthus, Sapindoidea, Myrica, Stratiotes, Zanthoxylum wemdingense and Spirematospermum wetzleri. The age of this locality is Upper Miocene (Tortonian), higher than that of Hessenbriicken. In the lower part of the Upper Miocene, T. maii is found together with Juniper, Chionanthus kornii, Zanthoxylum and Spirematospermum, mainly in a u t o c h t h o n o u s horizons like in Langenau near Ulm (not y e t published). The last three mentioned floras indicate a vegetation type, similar to the Pocosin woods in the USA (see above and Wells, 1928, p.234). In the Rhenish area T.rhenana is found in horizons from the uppermost Miocene to lowermost Pliocene with a flora of Stuartia--Cyclocarya-Arctostaphyloides--Symplocos character (open mine Victor-Rolff near Ziilpich, Schultz, 1962). Typical Lower Pliocene vegetation is found near Haguenau in Alsace (France) with Picea, Carpinus, Fagus, Corylopsis, Styrax, Vitis and Symplocos, Only T. maii (Geissert, 1972) is found here. In the Cantal area (S France), big seeds of T.naviculaeformis seem to have found a relict niche as in Alsace. The temperate flora consists of taxa like

Sparganium, Menispermum, Magnolia, Crataegus, Phellodendron, Meliosma,

351

Symplocos, Sambucus, and Trichosanthes. It is clearly an allochthonous flora (E.M. Reid, 1920, 1923). As regards the stratigraphic problems, compare p.346. Paleogeographical distribution The map (Fig.71) shows the occurrence of the fossil Toddalia species in Europe from Eocene to Early Pliocene times. It can be clearly seen that there is a geographical fluctuation in the course of time. In the Pliocene some relictniches exist in Europe (France and Netherlands), where Toddalia persisted perhaps under relatively bad conditions until Late Pliocene. From that time on the genus has retreated eastward and southward to Asia and Africa. As we will see, this rapid disappearance of Toddalia must have been due to climatic conditions. In Early Pliocene times there were subtropical to humid conditions in Europe, becoming colder in the Late Pliocene. For a theoretical model, the limitation lines of Toddalia today and in the Tertiary (hypothetically) are shown on the map (Fig.70). The "limitation line" is the line of the extreme occurrence of Toddalia lianas to the north and the south. Nowadays it is bounded by latitudes 30°N and 30°S, which means a region of 60 ° of latitude in tropical and subtropical areas. If the Tertiary sites of Toddalia are t h o u g h t to lie on the limit lines of the genus, and the Tertiary climate-zones are assumed to be identical with recent ones, then the equator may be reconstructed on a different line from that of today and moved about 20 ° to the north. More likely the Tertiary climate as

•

~j~

....

~-

.

Y

Fig.70. Geographical distribution of recent (lined) and fossil (black) Toddalia plants. Line 1 m e a n s the northern limit of Toddalia asiatica (= 35°latitude), line 2 the c o n s t r u c t e d "fossil" limit of Tertiary Toddalia species (55°latitude).

352

C

r

/ Fig.71. European distribution of fossil Toddalia species in the Tertiary. 0 = Eocene; • = Oligocene; • -- Miocene; • = Pliocene. a w h o l e was w a r m e r a n d c o l d e r areas w e r e missing or r e s t r i c t e d o n l y to far n o r t h a n d south. P e r h a p s t h e r e was h a r d l y a n y i c e s h e e t on the p o l e s ( c o m p a r e A x e l r o d a n d Bailey, 1 9 6 9 a n d V a n der Burgh, 1 9 7 3 , p . 2 0 6 ) . A t t h e end o f t h e P l i o c e n e the c l i m a t e grew colder.

Paleoclimatological data T a b l e I V ( A p p e n d i x ) gives s o m e c l i m a t o l o g i c a l d a t a on t h e o c c u r r e n c e o f R e c e n t Toddalia asiatica. I t gives an i m p r e s s i o n o f t h e c o n d i t i o n s u n d e r w h i c h Toddalia grows, a n d t h e s e d a t a m a y also be e x t r a p o l a t e d t o t h e fossils. By c o m p a r i s o n w i t h t h e a c c o m p a n y i n g fruits a n d seeds o f t h e fossil floras, it s e e m s c o n v i n c i n g t h a t t h e c l i m a t o l o g i c a l c o n d i t i o n s o f t h e fossil Toddalia o c c u r r e n c e w e r e v e r y similar to t h o s e o f t h e R e c e n t ones. A t r u e r e c o n s t r u c t i o n o f a fossil c l i m a t e is n o t possible here, b u t as is seen in T a b l e IV, s o m e d a t a are available a b o u t m e a n a n n u a l t e m p e r a t u r e s a n d m e a n a n n u a l p r e c i p i t a t i o n , c o n c e r n i n g regions w i t h R e c e n t Toddalia plants. R o u g h l y it can be said, t h a t severe frosts c a n n o t h a v e o c c u r r e d in t h e fossil climates; a p o s s i b i l i t y o f m i s t a n d high h u m i d i t y is given, seasonal changes w e r e n o t as m a r k e d as t o d a y in t h e C f b c l i m a t e o f Middle E u r o p e ( a c c o r d i n g t o the s y s t e m o f K S p p e n in Bltithgen, 1 9 6 6 , p p . 5 2 0 - - 5 2 6 ) .

J

353

The recent Toddalia lives between 14 and 27°C (mean of 20°C) mean annual temperatures and 500 to 10 000 mm annual precipitation (mean 2000 mm). No such exact data are available for the different Tertiary times, but there were subtropical to warm-humid conditions, and the mean annual temperature sometimes perhaps went to the limit of 14 ° and in Late Pliocene times below 14°C. It is only in connection with other fossils found with Toddalia seeds that a better reconstruction of climate becomes possible. APPENDIX TABLE IV Geographical distribution of recent T o d d a l i a with climate data. Geographical distribution o f Toddalia

Mean annual temperature (°C) ~

Brahmaputra Valley, India Nilgiri Hills, India Nilgiri Hills, India Khasi Hills, India New Delhi, India Hengchun, Taiwan Manila, The Philippines Borneo, Indonesia Bangkok, Thailand Singapore, Malaysia Naha, Okinawa/Japan

26.3 17.0 14.1 17.2 25.0 25.0 27.2 27.2 27.7 26.6 22.2

1600 1512 1500 10797 640 2153 2082 3571 1397 2413 2103

Tananarive, Madagascar Cape Town, South Africa Canton, China Hongkong, China T'eng-ch'ung, China Meng-tzu, China Shanghai, China Sian, China Chungking, China Lashio, China

18.3 16.6 22.2 22.2 15.5 20.0 16.1 15.5 18.8 21.6

1358 508 1643 2161 1242 965 1135 490 1092 1572

~ 20.0

~ 2000

Mean data

J Data after Meteorological Office (1966).

Precipitation per year (mm) ~

Special remarks

6 m above sea level 1749 m 2300 m

Monsoon region

Malayan trop. floral zone

(Bangkok)

(Hongkong)

(Canton)

U.S. Nat. Herb. (West Beltsville)

Steward, Chiao and Cheo 78

Larsen 10270 (Copenhagen)

Shimizu et al. 11344 (Copenhagen)

Shimizu 11393 (Copenhagen)

Thailand

Hongkong, China

Canton, China

Tsui, Canton, China

Kweichow, China

Ratchasima, Khao Yai, 800 m Thailand

Phetchabun, Phu-Miang 12--1300 m, Thailand

Hualien, 1500 m, Taiwan

Toddalia sp.

Toddalia asiatica (L.) Lam.

T. asiatica (L.) Lain. (= T.aculeata Persoon)

Toddalia asiatica (L.) Lam.

Toddalia asiatica (L.) Lam.

Toddalia asiatica

Toddalia asiatica

Toddalia asiatica

Herbarium No.

Locality

Species

5 dried fruits

4 dried fruits

3 dried fruits

1 dried fruit

13 dried fruits

10 dried fruits

many seeds

10 dried fruits

Material

Comparison of recent Toddalia material with special data

TABLE V

5.5 5.0

4.5--5.0

3.5

5.0--6.0 6.0

5.0--6.0

4.5--5.0

4.0-4.5

Length

4.2 4.0

3.0

2.5

3.5--4.0 3.0

3.0--3.5

3.0--3.5

2.0--3.0

Height

2.5 2.5

1.5

1.5

5--7 6

7

5

8

6(5)

(8.7)5(6)

?

6

No. o f s e e d s per ~ u i t

unripe, very small

1.5--2.0 2.0

2.0

2.0

2.0

Breadth

0.45 0.50

0.31

0.42

0.30

0.36

0.42

0.47

Breadth-length index

+

+--

+

Vascular bundle curved over chalaza

¢J1

Steward and Chev. 808A (Jamaica Plain)

(Berlin--Dahlem) 10 dried fruits

H.Y. Liang 63027 (Munich)

Bernardi 15404 (Munich)

Johnson et al. 8314 A (Jamaica Plain)

(Budapest)

Kwangsi, China

Kwangtung, China

Hainan, Ngai Distr., China

Hiniduma near Nelue, Distr. Galle, Ceylon

Nuwara, Ceylon

Palawan, The Philippines

Palawan, The Philippines

Toddalia asiatica

Toddalia asiatica

Toddalia asiatica

Toddalia asiatica

Toddalia asiatica

Toddalia asiatica

Toddalia asiatica

(L.) Kurz

Elmer 12161 A (Jamaica Plain)

Siam A 310 (Jamaica Plain)

Kwai Noi River, Siam

Toddalia asiatica

(L.) Lam.

5 dried fruits

Sic Law 5905 A (Jamaica Plain)

Yai Hsien dish, Hainan, China

Toddalia asiatica

5 dried fruits

3 dried fruits

3 dried fruits

8 dried fruits

3 dried fruits

3 dried fruits

4 dried fruits

5 dried fruits

Ngong T'inho TAAM Shah, 381 A Kwangtung, (Jamaica Plain) China

Toddalia asiatica

4.5

4.0--5.0

5.0

3.5

4.0

4.5

5.5

4.5 4.5

4.0--4.5

5.0--5.5

3.0

3.0

3.0--3.5

3.0

3.0

3.0

3.0

3.8 3.2

3.0--3.2

3:0--3.5

1.6

1.5--2.0

2.2

2.0

1.5

1.8

1.5

1.5--1.7 1.4

1.7

1.7--1.9

5(4)

(4) 5

5,3,2,1

7

6,7

6

4 (4--6) (6)

6(7)

0.35

0.40

0.44

0.5?

0.37

0.38

0.27

0.35 0.31

0.40

0.34

+

--+

+

+(-)

(+)

Wall. Cat. No.1206/1 (Munich)

Hook and Thomson (Munich)

Williams 519 (London)

Hohenack 577 (Munich)

Herb. Wight 359 (Munich)

Eamk L. 18483 (London)

Dupl. No.47159 (Coimbatore)

Napalia, The Philippines

Khasi Hills India 900-1800 m

Nepal, India

Canara, East India

Peninsulae Ind. orientalis

Madras, India

Madras, India Guindy Park

Toddalia floribunda Wall.

Toddalia floribunda Wall.

Toddalia asiatica

Toddalia aculeata Persoon

Toddalia asiatica

Toddalia aculeata Persoon

Toddalia asiatica (L.) Lamk. var gracilis Gamble

Loher (1906) from Kew (Munich)

Elmer 12767 (Zurich)

Palawan, The Philippines

Toddalia asiatica (L.) Kurz

Toddalia aculeata Montalban, Persoon Rhizal, The Philippines

Herbarium No.

Locality

Species

TABLE V ( c o n t i n u e d )

5 fruits

1 seed

1 dried fruit

1 dried fruit

4 seeds

1 dried fruit

1 dried fruit

2 dried fruits

2 dried fruits

Material

3.5

6.5

3.5

6.0

5.0

5.0--6.0

3.5--4.0

4.5--5.0

4.5

Length

2.2--2.6

4.0

2.5

3.5

2.5--3.0,

3.0

2.2--2.5

2.0--2.2

2.5

Height

2.0

2.5

2.0

2.5

1.8

2.0

1.5

1.7

1.7

Breadth

3(5)

?

3

3

4

5

5

4--5

4

No. of seeds per fruit

0.57

0.38

0.57

0.41

0.36

0.36

0.40

0.44

0.37

index

Breadth-length

+ long

+

+

+

+

Vascular bundle curved over chalaza

co

Sieber 72 (Halle-Saale)

J. & M.S. Clemens 30210 A (Jamaica Plain)

Dehra Dun, NW India

Khasia Hills, 900--1800 m India

Mauritius

Mauritius

Mr. Kinabalu, Borneo

Lourenzo Marques

Tananarive, Mt. Angavokely, Madagascar

Kaapsche Hoop, S Africa

Toddalia aculeata Persoon

Toddalia as&tica

Toddalia sp.

Toddalia paniculata Lam.

Toddalia asiatica

Toddal& lanceola~ G. Don

Toddal& asmtica

Toddalm na~lens~ Sond.

(Budapest)

(Berlin--Dahlem)

Rogers 22473 (Zurich)

Herb. Sieber (Antwerp)

Hooker and Thomson (Copenhagen)

(Budapest)

Dupl. No.37396 (Coimbatore)

Madras, India Kukal Betta, 1925 m

Toddalia asiatica (L.) Lamk. var. obtusifolia Gamble

Dupl. No.39112 (Coimbatore)

Madras, India Coonoor, 1700 m

Toddalia asiatica (L.) Lamk. vat. floribunda Gamble

3 dried fruits

12 dried fruits

2 dried fruits

3 dried fruits

1 dried fruit

8 dried fruits

1 dried fruit

1 seed

10 fruits

2 fruits

4.0--5.0 4.5

5.0-5.5 5.5

=

=

5.5

5.5

4.8 4.0

7.0-8.0

3.0-3.2 4.5

3.5 3.4

3.5

3.5

2.8 3.0

4.0-4.5

6

--

4(3) 2

4

Toddalia ?

5 (2--3) 2

Vepris lanceolata

4 6

Vepris sp.

Vepris sp.

2.0--2.7 2.5

2.0 1.9

2.0

2.2

2.5 2.0

3.0-3.3

0.53 0.55

0.38 0.34

0.36

0.40

0.52 0.50

0.41

(+)

(+)

(+)

+

+

--2

¢,O

2 dried fruits

Stolz 1984 (Copenhagen)

Kyimsala, 800 m, Nyassa

Kyimbila, (Munich) 1500--1600 m, Nyassa

Kivu Lake, Congo

Manica e Sofala, G orongosa, Mozambique

Pangani, Bushiri, Tanzania

Nyassa Highlands, Africa

Toddalia asiatica

Toddalia asiatica

Toddalia aculeata Persoon

Toddalia asiatica

Toddalia lanceolata

Toddalia aculeata Persoon

Stolz 1984 (Ziirich)

(Munich)

Torre and Paiva 12267 (Munich)

(Berlin--Dahlem)

4 dried fruits

Leonhard 2625 (Copenhagen)

Kivu, Walihali, Kashebere, Congo

Toddalia asiatica

2 dried fruits

4 dried fruits

7 dried fruits

3 dried fruits (1 Vepris)

4 dried fruits

1 fruit

Lucas 133 (Florence)

Kakamega Forest, Kenya

Toddalia asiatica (L.) Lain.

Material

Herbarium No.

Locality

Species

TABLE V (continued)

4.0

4.0-4.5

3.0

3.5--4.0

3.5

3.5 3.2

2.5--3.0

Length

2.8

3.0

2.1

2.5--3.0

2.5

2.6 2.5

2.0

Height

5

4(3)

(4--5) 6

No. of seeds per fruit

4(5)

unripe

2.0--2.2

2.1

1.3

1.9

2.0

1.9 1.8

1.0---1.5

Breadth

0.52

0.50

0.43

0.51

0.57

0.54 0.56

0.48

Breadth-length index

Vascular bundle curved over chalaza

O1 00

c~

Herb. Pretoria 1153 (London)

Transvaal, Soutpansberg, South Africa

Nago, Okinawa, Japan

Toddalia asiatica

Toddalia asiatica (L.)

Lam.

Lam.

(Nago)

S.Taiti 594 (Kew)

Mt. Elgon, Kenya, Nat. Park

Toddalia asiatica (L.)

Lam.

Stauffer 361 (Ziirich)

Kivu--Nord Virunga Chain, Congo

Toddalia asiatica (L.)

many seeds and fruits in alcohol

4 dried fruits

4 dried fruits

2 dried fruits

3.5--4.0

4.5--5.0

5.0-5.5

4.0

2.5--3.0

2.8--3.0

3.0-3.5

3.0

1.0--1.5

2.0

2.0--2.2

1.9

6--7

4(5)

5

5

0.45

0.42

0.40

0.47

+--

+--

¢.n ¢o

360 ACKNOWLEDGEMENTS This study was financed by the Deutsche Forschungs Gemeinschaft, Bad Godesberg. I heartily t ha nk this institution for the o p p o r t u n i t y of carrying o u t this research -- as well as Prof. W. Jung from the Institut ffir Pal~iontologie und historische Geologie, Miinchen, who p e r m i t t e d me to work in his departm e n t o f Palaeobotany. I thank m y colleagues H. Schauderna, H. Thiele and D. Mfiller f r o m the same institute for their co-operation and interest. I wish to emphasize the unique t e a m w o r k with m y friends D.H. Mai, F. Hol~, J.v.d. Burgh, H. J~ihnichen and m y colleagues K. Kilpper, F. Geissert and M. Lancucka-Srodoniowa. The photographs were kindly made by M. Dressier, the drawings by K.J.W. Dossow (both f r om the Institut ffir Pal/iontologie und historische Geologie, Miinchen), the stereoscan-pictures (Mk IIa, Cambridge Instr., G.B.) by the Institut ffir R a s t e r - E l e k t r o n e n m i k r o s k o p i e H. Klingele (Mfinchen) and the X-ray p h o t o s (Siemens-Mammomat) by U. Schulze (Neu-Perlach Hospital). The Bayerische Braunkohlen-Industrie AG S c h w a n d o r f kindly managed the possibilities to collect m os t of the fossil Toddalia populations. Moreover, this study would n o t have been possible w i t h o u t the generous c o o p e r a t i o n o f m a n y colleagues and institutions, especially in G e r m a n y GDR, Czechoslovakia and Poland, but also the People's Republic of China, the U.S.A., Great Britain and T he Netherlands. Here I have to t hank among o t h er persons, S.P. Lau, C.H. Shute, Zen'Iti Kisi and Kei-Ichi Shimabuku, W.H. Zagwijn, I.C. Verdoorn, W. Trapp and Chow Han Chuan. Finally, m y grateful thanks to m y friend and colleague M. Collinson (British Museum of Natural History), who correct ed the English manuscript and p u t forward some good criticism. I sincerely t h a n k m y wife Uta for her help in every way during the five m o n t h s of this project. REFERENCES Ahrens, H. and Lotsch, D., 1967. Die geologischen Grundlagen der Aufstellung der Florenzonen im jiingeren Terti~r der Lausitz. Abh. Zentr. Geol. Inst., 10: 39--54. Aubreville, A., 1963a: Flore du Gabon, 6. Rutac~es. Mus. Nat. Hist. Nat., Paris, 122 pp. Aubreville, A., 1963b. Flore du Cameroun, 1. Rutac~es. Mus. Nat. Hist. Nat., Paris, 174 pp. Axelrod, D.I. and Bailey, H.P., 1969. Paleotemperature analysis of Tertiary floras. Palaeogeogr., Palaeoclimatol., Palaeoecol., 6: 163--195. Backer, C.A., 1965. Flora of Java, vol.II, Faro.133. Rutaceae, Groningen, 641 pp. Bliithgen, J., 1966. Allgemeine Klimageographie. Walter de Gruyter, 720 pp, Berlin. Boesewinkel, F.D., 1977. Development of Ovule and Testa in Rutaceae I: Ruta, Zanthoxylum and Skimmia. Acta Bot. Neerl., 26(3): 193--211. Bu~ek, C. and Hol~z,F., 1964. Small-sized plant remains from the coal formation of the Chomutov--Most--Teplice Basin. Sb. Geol. Usted. Csl., Paleontol., 4: 105--138. Champion, H.G. and Seth, S.K., 1968. A Revised Survey of the Forest Types of India. Delhi, 402 pp. Chandler, M.E.J., 1963. The Lower Tertiary Floras of Southern England. 3. Flora of the Bournemouth Beds, the Boscombe, and the Highcliff Sands. Br. Mus. Nat. Hist,, London, 169 pp.

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