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Home , Liriodendron, Magnoliaceae

: a review of literature

By RICHARD S. PEIGLER

The family Magnoliaceae is fairly Acceptable classifications of extant well represented in the fossil record of and animals can be reached by the Tertiary and Quaternary Periods of studies of morphology, chromosome the Cenozoic Era (see Fig. 1). Some numbers, biochemical analyses, use of questionable records have been cladistic methods, etc. in groups where ascribed to the Period of little or no fossil record is available. the Mesozoic Era. The botanists and However, we would know compara- horticulturists who have developed the tively little about a group such as the classification of Magnoliaceae have redwoods and sequoias (Taxodiaceae), done so almost entirely by studying containing only three extant relict living (extant) species, largely ignoring species, if we did not study the the fossil record. Currently, paleo- abundant fossil material which shows botanists are re-evaluating the group to have been widespread in assigned to Magnoliaceae; in fact, the the Northern Hemisphere in past ages. literature all through the past century For the Magnoliaceae, new finds of has examples of where fossil names fossils coupled with more reliable and have been transferred into or out of the sophisticated evaluation (Spicer 1986, Magnoliaceae. For the extant species, Thomas 1986) of these and earlier finds there likewise has been a certain ensure that this data base on phylogeny amount of transfer of species from one of the family will be increasingly to another and a refinement of useful to those working on of the family concept: e.g. genera such as living Magnoliaceae. I/lie/am /fadsura, and Schisandra, The current distribution of formerly considered to be - MagnoTiaceae is restricted largely to ceous, have been placed under separate eastern Asia, eastern , familiew I am not certain if the generic and Mesoamerica. The past distribution boundaries defined by Dandy (1971) or included Europe, western Asia, western intrageneric classification of two North America, and Greenland. These subgenera and eleven sections within past and present ranges are shared by Magnolia as outlined by Seitner (1968) other familiar plants: Ls//urdamher, are yct stable and widely accepted. The Awe/ras and Linderr, to name a few. taxonomic debates in ktagnolia Genera currently restricted to eastern literature at the levels of species, Asia such as Ailanrhus (tree of subspecies, varieties, and putative heaven), Cinnamomum (camphor and hybrids will continue long into the cinnamon), and 2e/hove were formerly future, and I doubt that fossil data will present in western North America, have any value at these lower Europe, and western Asia (Zhilin 1984 taxonomic levels. and others). In general, extinction

MAGNOUA ysll, 19s9 among the fauna and flora of these taxonomically as seen in the study by latter regions was caused by cooling and Page (1984). She described a new fossil drying of the climates through the genus of plant judged to likely be Tertiary followed by episodes of Magnoliaceae from the Upper glaciation (Ice Ages) during the Cretaceous of California. Comparisons Pleistocene (Tidwell 1975, Tiifney 1977, of cambium, secretory cells, primary Wolfe 1978, 1980, etc. ). and secondary xylem, etc. were made to This is a review article based mainly those of living Alagnolia Linbdeadmn on historical and current literature, and and Aiiehehh as well as to hiagnrohh- it contains no original research. eerz~lon/srnoehensir (Page)Wheeler, Scott, ht Barghoorn. The Miocene Plant fragments ss fossils AdagnoNxy/on aeandens Schflnf. was Leaawx Many fossils were also described from a species of assigned to the wrong genus or even petrified wood (Nemejc 1975: 96). plant the wrong family by early workers. I fossil genera based on fragments of cannot say whether the figured wood or stem have traditionally ended specimen of Alagnohh //o/z'raan/Jeo/a with the suffix -arv/on. Comparisons (Fig. 2) is correctly placed. Such between wood fossils must be made re-evaluation would be befler left to a with caution because certain features of trained paleobotanist. Tidwefl (1975) wood anatomy are thought to represent gave the following characteristics to evolutionary grades, rather than clades. identify fossil Adagnoila : Pollen andiyoue/x Pollen of unlobed, entire (unserrated) margins; Magnoliaceae may have useful midvein tapering from thick near base taxonomic characters. Huang (year?) to thinner toward apex, 4 - 15 provided detailed descriptions of the secondary veins of alternate to pollen grains of AAgnolia asico subopposite ones curving apically near (Lour. )DG, Ad/ehelia a/ha DG, margins often connecting to ones 8/ehelia lo/znowna (Kanehira) above; tertiary veins at right angles to Masam. , and Aireheliogazf'haehdaeh/lai secondaries. Kvacek (1978) said that (Kanehira tk Yam. )H. Keng. leaves of Magnoliaceae have "internal Unfortunately, as Tiffney (1977) secretory bodies, paracytic stomata with pointed out, pollen of Magnoliaceae is guard cells slightly elevated above quite susceptible to chemical and subsidiary cells, uniseriate trichomes. " biological degradation, so fossil pollen He went on to describe the leaf hairs of this family is virtually non-existent. and to discuss how to distinguish the Baghai (1988) mentioned reports of various genera within Magnoliaceae Lrlzb dead/on fossil pollen. (see Figs. 3-4). I found no figures or descriptions of $/ems Although Tidwefl (1975) Adagnolia in the literature. provided photographs of many fossil Although magnoliaceous flowers readily species of xylem, none are for shatter with fafling to the Magnoliaceae. Taylor (1981:87) ground, fossilization of entire flowers illustrated a "scalariform pitted vessel seems possible since branchlets bearing of Aiagnolia" with a 275X magnifl- intact flowers are often blown to the cation. The detailed stem structure of ground during storms. In their diagrams Magnoliaceae is undoubtedly useful of the proposed reconstruction of their

MAGNOVA Fall, 1999 Fig. I GEOLOGIC TIME CHART

Millions of Years ERA PERIOD EPOCH Start Duration

Holocene = Recent Quaternary Pleistocene 1.8 1.8

Pliocene 10

Cenozoic Miocene 25

Tertiary Oligocene 40 15

Eocene 60 20

Paleocene 70 10

(Last period of "Age of Reptiles" Mesozoic Cretaceous 135 65 following and )

Arorer a4rer arfen nre nafnnuhanre. She llokaene b oronl ro the bar laeeesenra

extinct magnoliaceous genus Arc- 4/agnoliaesrrobus nomtvi (Jabl. ) haeanihus Dilcher & Crane (1984) Rdsky, hfzgnoliaesrrobus hungaricus showed leaves, stems, , and Rdsky (both cited from Hungary by mature aggregate . This remarkable Hably 1985), Magnoliaesov bus plant may represent a species of gilmourii Seward & Conway from Magnoliaceae which existed prior to the Cretaceous of Greenland, and Magnolia divergence of the two tribes (seusu nrlliynim Stanilovsky from the Eocene Dandy 1971) of the family. of the Ukraine (P. Dorofeev & N, Cones of kfaguoila and imchanitzkaja iu Takhtajan 1974). allied genera are found in fossil Figures of these latter two were given deposits and are generally more readily by Nemejc (1975). Also from identified as belonging to this family Greenland are fossil cones of kfagnolia than are the fossils of other parts of the iuglefieldii Heer (L Rnffle, personal plant. Because cones of Linhfendrva communication). The 'Maguo/ia sp. shaner into separate when ripe, fructus" described and figured by generally these samaras are known as Shvaryeva (1983) appears to be a few fossils for that genus. Some species separate carpels on fragments of stone. described from remains of cones are Cones of cycads and conifers (Piuus

MAGNOLIA Fall, 19S9 ASeraaepuora, etc. ) are well known itself. Z. Kvacek (pers. comm. ) from the fossil record, many of which considers the so-called Upper are beautifully preserved (Tidwell 1975, Cretaceous itdagnoira species of Taylor 1981, Mai 1987a, etc. ). Europe such as itd. speeiaca Heer and getsds The magnoliaceous of id. smplifolu Heer to be doubtful. the tribe Magnoliaceae has a brightly LaMotte (1952) cited several records colored fleshy outer layer (sarcotesta) for the genus from the Paleocene and on a stony interior portion (sclero- Eocene, but these will require testa), the latter of which is abundantly re-confirmation in future studies. preserved in fossil deposits in many Reports from Oligocene, Miocene, and locales of the Northern Hemisphere. later epochs (refer to Fig. 1) are Tiffney (1977) has reviewed the plentiful and most of these are likely literature and drawn many new valid. Tiffney (1971:311) wrote: conclusions pertaining to the "Although itdagnoirh seeds are relationships between earlier fossil common throughout the European finds and extant taxa within Magnolia- Tertiary, forms comparable to modern ceae. Another significant discussion species do not become prevalent until which relied mainly on seed fossils was the Late Miocene, when they supplant by Tralau (1963: 37-40). Seminal earlier, now extinct, forms. " remains are apparently reliably A fossil plant from North America identifiable to the generic level. Several named itdagnoiraoborwra Newberry is paleobotanists confine their studies considered to be a junior synonym of primarily to fossil seeds (e.g. Mai 1971, the fossil iVyxw ueruara Newberry 1975, 1987a, b). Many diagrams and (Hollickin Newberry 1898). photographs of magnoliaceous seeds Interestingly, this name is also a junior are given in the references cited above primary homonym of Magnoila obotwra in this paragraph as well as by Nemejc Thunberg, which in turn is considered a (1975) and Takhtajan (1974). Figure 5 subjective junior synonym of the extant shows a fossil seed of Megnoira Japanese itd. /iypoleuea Siebold dt girtrrbensis Szafer (1961) from the Zuccarini. Hably (1985) compared Miocene of Poland. Seventeen species certain fossil finds to h/ odors, but of itdagnoira based on seeds from the he was referring to the modern London clay (Eocene) deposits were Japanese species, not the North listed by Chandler (1964), although American fossil name. Tiffney (1977) suggested that several of Another nomenclatural error is these names may be synonymized after found in Andrednszky (1966). He new information he developed on seeds compared some Hungarian fossils to of itiagnoira is applied to these the living itd. grnndil/onr L, AX specimens. acuminara L, and iV. punduana Waflich with no problem (although the Fossils of Magnolia latter is now considered to be a species Records of itdagnoils from the of itdreireliaf. However, in the same Cretaceous of North America are paper he compared another leaf doubtful (J. Wolfe, G. Upchurch, pers. imprint to the "Japanese AX deaiitara comm. ). The family may have arisen Zuoc. ", and yet Ad dealdara is a living that early, but probably not the genus Mexican species. Based on the name, stAONOUA Fall, 19s9 pra. 2 Fioiobpe of Magnolia noriaaaalioola Cooterei/ From piorataanx Trit r Co, Coioraoh Vpperhlaaenc. Specimenin Vnir. Cob. ltloreom Boulder. wc might guess that Andrednszky Pliocene of Poland; Pliocene of France, meant Atf denudaia Dear. , a synonym Germany, and ; and Miocene and of Ai heplapeia (Buc'hoz) Dandy. Oligocene of the USSR. Tralau's map 9 However, thc fossil leaf is about 20 proposed a past distribution of Al centimeters long and 10 wide, kobus across Europe and Asia. comparable to the Mexican Al. Therefore, it seems likely that dealbanz Hably (1985: 19) again cited representatives of the section Buergeria the fossil material as "Ad cf. deaibara of the genus were widespread and Sieb. dt Zucc. " in his catalog. common in Eurasia for the past 35 A detailed analysis of fossils from million years, becoming extinct only Eurasia was considered to represent the during the last 1 1/2 million years extant Japanese Al. kobus DC. by except in the Far East. Tralau (1963). He listed numerous The excellent work by Tiffney literature refcrenccs of AE eor Ludwig, (1977) is the most comprehensive and Af. ultima Kirchheimer, and W kobus insightful paper I have found on fossil all under the synonymy of Af. kobus Magnoliaceae, the detailed discussions All these fossils were considered to be of which are not indicated by the title referable to A'. kobus itself or taxa that of the article. He described two ncw were virtually indistinguishable from it. species of Magnolia from the Brandon They included records from the Lower Lignite (Middle Oligocene?) of Pleistocene in The Netherlands; Upper Vermont, based on seeds. His Adagnolia Miocene, Miocene, Upper Pliocene, selrrenrrionalis is perhaps nearest to

StAGNOUA Fall, 19S9 M. grrndiliorn, but he went on to four species in his catalog of North explain why none of the European American fossil species: L novpetz'9 comparisons to M. grnndilorn are Berry from the Miocene of Washington valid. Mngnolia uwitociri Tiffney shares State; L itstloittlot Wodehouse from characteristics of the seeds of section the Eocene of Garfield County, yytooritodon (particularly the Mexican Colorado; L tulipiiorn L from the M. scliiMiinn Schlechtendal) and Holocene and Pleistocene of Maryland, section 7)titihsstrtttn such as M. Alabama, North Carolina, and ttcumdtatn and M. Puinpnoitotn Louisiana; and the European L (Buc'hoz)Dandy. Tiffney proposed itrotncothii Unger from Paleocene various hypotheses of relationships of localities of Greenland and Iceland. his two new species to other fossil and Nemejc (1975) listed L Sap. extant species. The affinities between also from Iceland and Greenland.~ the various Eurasian fossils called M. Baghai (1988) compared L iteritorth to nitdna M. cor, and M. gotiur were the two extant species in considerable discussed in greater detail than was detail; her work appears to be the most given by Tralau (1963), and new current and reliable for fossils of this comparisons were made to well known genus. living species. A final paragraph deals Arnold (1947) stated that with separation of the North American Liniedondron in Europe was and European floras which were terminated by Pleistocene glaciation, apparently connected by a land bridge but "up to that time a species during Early Eocene (see also Wolfe indistinguishable from the American L 1978, 1980). tulipiFea existed in the lowlands of Two further signiTicant works on western Europe. " Some of these fossil fossil Mngnoitlt must be mentioned. finds of France, Poland, and Germany These are the study by Mai (1975) and have been called L tnijtttlbrn Rails the 14 species of Mngnoith treated by Reid (Nemejc 1975). Mai (1987a) Dorofeev & Imchanitzkaja (in described L itttgiir based on material Takhtajan 1974). Here one can find from various German Miocene locales. complete synonymies, original A well known European tuliptree is L descriptions, redescriptions, and good itrotncnuii (mentioned above from figures. Pneva (1986) has since added Greenland). It is cited by Hably (1985) the Upper Oligocene Mngnoitn from deposits in Hungary and by tnitittnihnii to the list of Asian fossil Givulescu (pers. comm. ) from . This species had large leaves Romania. A junior synonym is L and was compared by Pneva to modern itoiteticum Heer from Switzerland. Asian and American representatives of Kirchheimer named L gotndtntn from the section Ryudotitortnotn. seeds from Pliocene in Poland (E Zastawniak, pars. comm. ). The Fossils of Lirlodendron Oligocene L bnueri Ettingshausen was Remains of this genus are known described in 1869 from Czechoslovakia from leaf and seed compressions. As (Z. Kvacek, pers. comm. ). Two further mentioned earlier, individual samaras Oligocene species were named by are generally fossilized because ripe Dorofeev (dt Takhtajan 1974): L cones shatter. LaMotte (1952) cited ttnitthum and L uralense

StAGttOIIA Fall, 19S9 isa 3 Fomilirafrmwsfcrao(ahaamlia ksialiaae shifvasek Fiom /Vzrrkerasforf'ffavaoh' f'mrmaoy. Loaerhfsoseoe. Sfsmmeo wishfsersoiefs~ the lxabrffse. Pboree fsom Dr. 2. Xvasek

Liriodendron meefu' Heer from the the fossil maple Aeerminuufolium Cretaceous of Nebraska has been Chancy (Wolfe & Tanai 1987). demonstrated to be a leaf imprint Obviously unrelated plant groups may belonging to the legume genus share the peculiar leaf shape seen in Dalhergia according to Arnold (1974); true Lii~endron. Thus, none of the Cretaceous specimens from Greenland, species mentioned in this paragraph are Kansas, and Nebraska were cited as true Magnoliaceae. Dafhergifessimp!er (Newberry)Sew. & Con. (Fabaceae) which had also been Fossils of Talaulna, Manglietia. ascribed to by the and original author (Nemejc 1975). These The present-day distributions of the fossils are now believed to be plants genera ltdrehelia and ltfanglieria are which are not legumes either (G. restricted to southeastern Asia, whereas Upchurch, pers. comm. ). hfenispermo- extant species of lafauma grow in phyllum eelakovdii (Velenovsky) tropical America and Asia (Dandy Velenovsky was originally called a 1971, Nemejc 1975). Like Magnolia Liriodendion (Z. Kvacek, pers. and Liriodendron, these three genera comm. ). Liriodendron irilohafa Chancy ranged in other parts of the world is now considered to be a specimen of during the Tertiary. Fossil and extant

MAGNOLIA Fall, 1999 seeds of these genera can be identified present in Japan during the Holocene by reference to the distinguishing and Bulgaria during the Eocene characters outlined by Tiffney (1977). (Tiffney 1977). I have been unable to The extinct magnoliaceous genus obtain the original references for the Alagnoliaespermum Kirchheimer, very few finds of this genus in the fossil based on Upper Oligocene seeds from record, but they are cited in Tiffney's Poland and Germany, has certain bibliography. characters like the seeds of and others more like hlauglienh Acknowledgments (Tiffney 1977). Leaf fossils of Talauma I have received cordial help in the and bgan~liena can be identified by the form of literature, photographs, or characteristic hairs and epidermis discussions from the following persons. described by Kvacek (1978). I am particularly gratified by the Chandler (1964) described Talauma generous help I was given by our rurlkinsonii from the London Clay flora friends in eastern Europe. I thank Dr. (Eocene) of southern England, based R. Givulescu (Romania), Dr. Lilla on seeds. Tahruma egemnsis And- Hably (Hungarian Natural History rednszky (1955), described from fossil Museum, Budapest), Mr. Harold leaves from Hungaty, was likened to Hopkins (Maryland), Dr. Wieslaw the extant T. Pubeseens Merrill from Krzeminski and Dr. Ewa Zastawniak the Philippines. Another leaf fossil (Polish Academy of Sciences, Cracow), from Hungary assigned to this genus Dr. Zlatko Kvacek (Czechoslovak was cited by Hably (1985: 22). Academy of Sciences, Prague), Mai (1971, 1987b) described three Librarians (Denver Botanic Gardens), new fossil species of blangilenh and Dr. Peter Robinson (Univ. of Colorado transferred Carpolirhusarhkersenii to Museum, Boulder), Dr. L Riiflle and the genus. These descriptions are based Dr. Dieter Hans Mai (Museum fur on seeds from Germany as follows: Naturkunde der Humboldt-UniversiQt, bganglielia bereynrha Mai (1971), Berlin), Dr. A. K. Shakryl (Academy of Middle Paleocene, likened to the living Sciences of Georgian SSR, Sukhumi), bg kuungrungensa (Merr. )Dandy of Dr. G. R. Upchurch, Jr. (National ; bl. zinke~nii (Geinitz)Mai Center for Atmospheric Research, (1971), Lower Oligocene; bV. germanic Boulder), Dr. Jack A. Wolfe (U. S. Mai (1971), Miocene, considered to Geological Survey, Denver), Dr. Sergy reSemble the eXtant atl glauos Blume G. Zhilin (Academy of Sciences of from Java; and hf. mulrrmsrara Mai USSR, Leningrad). My colleague (1987b), Upper Paleocene. Other fossil Michael J. Weissmann (Univ. of finds from USSR have been classified Colorado Museum) kindly photograp- under Manglierrh (Shakryl 1987: 118). hed the holotype of bgagnoila Some of the many species of italagnoirh liorrssanricnia shown in Figure 2. ~ described from fossil seeds from the London Clay (Chandler 1964) may be found to actually belong under Or. Peigleris Research Assoeiare Alanglieua after being re-examined ~krgv Oeparrmenr, OenserMuseum (Tiffney 1977). orbiarum/History, City Park centaur, The genus Wicheirk was possibly CO kKBfy

MAGNOLIA FalL 1989 Fvp 4 f oner cuticle oftenffosvilof Magnolia liblavenaLv ftvfrnnsctatt Nreytsnd) ffancel: Fium fihtvr Mine ne, vr Cotopnc Rhiachvnd, Cseimsny. Mtoca ne. Photo fium Dr. L RhttBa:

Pig. S Tnu vvansofholotypeseedof Magnolia gliwicensis Szvfar. FiomStvn GImtce tipper Silesis. Pohmd. Afiddle Fsvcene. Photo fium Dr. E. Zvstvwnsvh

MAGNOLIA Fall, 1969 Uterattsrts Cited

Andrednszky, G. 1955. Neue und interessante tertizre Pflanzenarten aus Ungarn. Ann. Hnr. /tlarur. Sf' /lung 6: 37-50. Andrednszky, G. 1966. On the Upper Oligocene flora of Hungary: analysis of the site at the Wind Brickyard, Eger. Srud. Diol FIT@ 5: 1-151. Andrews, H. N. , Jr. 1961. S/adit iir pa/ktoho/any. J. Wiley & Sons, New York. xii + 487 pp. Arnold, G A. 1947. Aninrroducrrhn rope/eohoraay. McGraw-Hill, New York. Ix + 433 pp. Baghai, N. L 1988. Lii/odenditzn (Magnoliaceae) from the Miocene Clarkia flora of Idaho. Amer. y. Box 75(4): 451-464. Chandler, M. E. J. 1964. Tyre Lorver 7'err/sky//oras olsourhernMg&ndr vol. 4, A summary and survey of findings in the light of recent botanical observations. Brit. Mus. Nat. Hist. , London. 151 pp. , 4 pls. Dandy, J. E. 1971. The classification of Magnoliaceae. Aervs/ Amer. /a/age. Soc 8(1): 3-6. Dilcher, D. L & P. R. Crane. 1984. Archaeanrhusr an early angiosperm from the Cenomanian of the western interior of North America. Aun. /tlzfrourr'//or. Gard. 71(2): 351-383. Givulescu, R. 1975. Fossile Pflanzen aus dern Pannon von Delureni, Rumgnien. Pa/amnrogrsphiar 153B: 150-182. Hably, L 1985. Catalogue of the Hungarian Cenozoic leaf-flora. Srud. //oz F/kg 28: 5-58. Huang, T. -C. (year?) Pollen//ora o/'Tarlvan. NatL Taiwan Univ. Bot. Dept. Press. Kvacek, Z. 1978. Some members of Magnoliaceae from the European Tertiary. Palmnr. Konf. '77-Uaiv. /farl Praha pp 169-182. LaMotte, R. S. 1952. Catalogue of the Cenozoic plants of North America through 1950. /totem. Geol Soc Amer. 51: 381 pp. Mai, D. H. 1971. Fossile Funde von /rlangilerra Blume (Magnoliaceae). Peddes Reperlorium spectrum novarum regni vegetzhilis 82(6): 441-448. Mai, D. H. 1975. Beitrgge zur Bestimmung und Nommenklatur fossiler Magnolien. Ped. Rep 86(9-10): 559-578. Mai, D. H. 1987a. Neue Arten nach Frilchten und Samen aus dern Tertigr von Nordwestsachsen und der Lausitz. Red. Rep 98(1-2): 105-126. Mai, D. H. 1987b. Neue Friichte und Samen aus palaoztnen Ablagerungen Mittcleuropas. Fed. Rep. 98(3-4): 197-229. Nemejc, F. 1975. Pa/coho/sudra, vol. 4: Systematicld csst, rostliny krytosemennd. Academia, Praha. 566 pp. , 40 pls. Newberry, J. S. 1898. The later extinct floras of North America. Monogr. U. S Geol Surv. , vol. 35. Washington, D.C.

MAGNOLIA 10 Fall, l 909 Page, V. M. 1984. A possible magnoiioid floral axis, Loishoglia betteneourui from the Upper Cretaceous of central California. L Arnold Arb. 65(1): 95-104. Pneva, G. P. 1986. New view of Magnoiia in the Late Oligocene flora of Ahshutasa (eastern Kazakhstan), pp. 112-119in L Yu. Budantsyev, S. G. Zhilin, et aL, editors, Pioblemsin pa/coho/any. Nauka, Leningrad. 176 pp. , 58 pls. (in Russian) Seitner, P. G. 1968. A taxonomic diagram of the genus Magnolia. Aelvsl Amer. Maga. Sou 5(1): 3-5. Shakryl, A. IC 1987. New view of the genus Magnolia L. from the Pliocene of Abkhazia. Trudy Sukltum. //ot. Sade 31: 112-119.(in Russian) Shvaryeva, N. Ya. 1983. Miocene/lors ofdieLorver Carpathians Naukova Dumka, Kiev. 160 pp. , 80 pls. (in Russian) Spicer, R. A 1986. Comparative leaf architecture analysis of Cretaceous radiating angiosperms, pp. 221-232in R. A. Spicer & B. A. Thomas, eds. , Systemaircand taxonomic approaehesin pataeobotany. Clarcndon Press, Oxford. x + 321 pp. Szafer, W. 1961. Miocenska flora ze Starych Gliiwic na Slasku. Prate Inst. Geol, Warsratva 33: 1-205. Takhtajan, A. , cd. 1974. Fossil//orverthg plants or thte USSR, voL 1: Magnolia- ccae —Eucommiaceae. Nauka, Leningrad. 188 pp. , 124 pls. (in Russian) Taylor, T. N. 1981. Paleobotanyt an thtroduction to /ossilplant bio/os. McGraw- Hill, Ncw York. xiii + 589 pp. Thomas, B. A. 1986. The biochemical analysis of fossil plants and its use in taxonomy and systematics, pp. 39-51 lu Spicer & Thomas, op. eik Tidwcll, W. D. 1975. Common louilplants ot'western berth Amer/ha Brigham Young Univ. Press, Provo, Utah. 197 pp. , 48 pls. (16 col. ), 1 chart. Tiffney, B. H. 1977. Fruits and seeds of the Brandon Lignite: Magnoliaceae. Bot. / Linn. Soe 75: 299-323. Tralau, H. 1963. Asiatic dicotyledonous affinities in the Cainozoic flora of Europe. Kung/ Svenska Vet. Akad' Hand/, 4th scr. , vol. 9: 1-87. Wolfe, J. A. 1978. A paleobotanical intcrprctation of Tertiary climates in the Northern hemisphere. Amer. Scient/st 66(6): 694-703. Wolfe, J. A. 1980. Tertiary climates and floristic relationships at high latitudes in thc Northern Hemisphere. Palaeogeogr. , Pa/aeocfimato/, Pa/aeoeeo/ 30: 313-323. Wolfe, J. A & T. Tanai. 1987. Systematics, pyhlogeny, and distribution of Arer (maples) in the Cenozoic of western North America. J. Fae. Sei, //oklaido Univ. 22(1): 1-246. Zhilin, S. G. 1984. Grades of principal flora formations of the temperate forest of Kazakhstan during the Oligocene to Early Miocene. Komarov Lecture Series ' Aead. Set. USSR voL 33: 1-112. (in Russian)

MAGNGUA Pall, 19S9