iPaleontological Joumal. Vol. 34. Suflpl. I. :WOO.pf'· S/17-S98. :Original Ru.1siarr Text Cop)-righr © 2000 !Jy Krassilov. 'English Translation COJl)Tight © 2000 !Jy MAIK "Nauka/lnterperiodico" (Russia). ======PHYTOGEOGRAPHY======

Permian Phytogeographic Zonality and Its Implications · for Continental Positions and Climates V. A. Krassilov Paleontological Institute, Russian Academy of Sciences, ul. Profsoyuznaya 123, Moscow, 117868 Russia Received December 15, 1999

Abstract-The traditional Permian phytogeographic divisions do not correspond to separate landmasses. Nei­ ther are they floristic divisions for their emblematic leaf genera, such as Cordaites or Glossopteris, are morpho­ types shared by the taxa that are not closely related. A revised scheme of Permian phytogeography is based on the dominant leaf morphologies, taphonomic evidence of seasonality and dead mass (coal) accumulation. It is consistent with climatic zonation including the nemoral, or humid temperate, xerothermic and humothermic zones in the Arcto-Permian realm and their counterparts in the Noto-Permian realm. The Permo-Tethyan realm of mixed Arcto- and Noto-Permian elements corresponds to a global megashear zone of the Permian tropics. This scheme lends no support to either a continental reassembly with submeridional Eurasia, equatorial Euramerian realm (which actually had a Mediterranean type climate) and the Tethys widely open to the east, or a position of the Permian South Pole near Africa (which had a much warmer climate than Antarctica). A west­ ward shear of the eastern Tethyan microcontinents from their pre-Permian position near the Sunda Plate would explain both the constriction of the tropical zone between Tibet and the Himalayas and the origin of mixed flo­ ras. The supposedly glaciated areas of Permian tillites supported a diverse arboreal vegetation suggesting a cli­ mate totally different from that of Pleistocene glaciation.

INTRODUCTION In the 1930s the great Permian geofloras were dis­ I argued (Krassilov, 1972, 1975, 1997, etc.) that tinguished on the basis of dominant leaf morph'ologies, major phytogeographic divisions of the past are vegeta­ such as Callipteris, Cordaites, Gigantopteris and Glos­ tional, rather than floristic, for they are based on a few sopteris, defining the Euramerian, Angaran, Cathaysian dominant life forms defining the aspects of regional and Gondwana realms respectively (Jongmans, 1936, fossil plant assemblages rather than on computed floris­ 1937, Gothan, 1937; Kryshtofovich, 1937; Meyen, tic similarities and the ranks of endemic taxa, as is nec­ 1970). These realms were considered as floristic units essary for a proper floristic classification. The Arcto­ corresponding to separate landmasses. As such they Tertiary "geoflora" is the first widely accepted division played a significant role in the development of the con­ of the kind. It is equivalent to the nemoral, or temperate tinental drift concept and, later, plate tectonics. To this broad-leaved biome that is recognizable in the Meso­ day they are widely used in paleogeographic recon­ zoic (the Phoenicopsis realm: Krassilov, 1972, etc.) and, structions. However the Permian geofloras are not perhaps further back in time. Climatically the nemoral strictly definable in terms of taxonomic distinctions biome corresponds to the humid temperate zone. because of morphological parallelism of their emblem­ South of the nemoral zone there is, and was, a like­ atic leaf types that might belong to different, even not wise persistent realm of evergreen and deciduous arbo­ closely related, taxa. Thus the Cordaites-type leaves real biomes with a prominent xeromorphic element. occur in the typical cordaites, vojnovskyaleans and The climatic conditions are warm temperate to subtrop­ glossopterids (Noeggerathiopsis). Though widely ical of the Mediterranean or Texas types, with a dry spread in Euramerian, Angaran, Cathaysian and Gond­ summer season. The corresponding vegetation realm in wana floras, this morphotype is poor evidence of floris­ the Miocene of North America was called the Madro­ tic affinities. The same is true for the Asiatic and North Tertiary flora (Axelrod, 1958). This vegetation type American gigantopterids, the Gondwana and Angaran extended over Mediterranean, Central Asia and as far representatives of Palaeovittaria-Zamiopteris-Pur­ east as northern China. In the Mesozoic the xerothermal songia morphotypes, etc. Glossopteris associates with zone was marked by the Cycadeoidea assemblages with at least two different types of reproductive organs-cup­ brachyphyllous coniferoids (Krassilov, 1972, etc.). ular, as in Lidgetonia, and strobilar, as in Kendostrobus. On the other hand, the relatively dry northern coni­ This leaf morphotype might, thereby, belong to at least fer forests and tundras, as well as the tropical rain for­ two groups of Paleozoic gymnosperms, one related to ests are of a recent origin, related to the Late Cenozoic pteridosperms and the other closer to cycadophytes glaciation, with no equivalents in the Mesozoic ice-free (Surange and Chandra, 1975; Maheshwari and Srivas­ cli":late, but probably represented during the Late Pale­ tava, 1992). Taxonomic affinities of Gigantopteris and ozmc. its morphological allies are still more intricate. Plants

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Fig. 1. Fertile gigantopterid leaves Gigantotheca with marttioid synangia from the Tontseyan Fonnation, Fujian, Southern China (the material courteously provided by Prof. Yao, Nanjing Institute of Geology and Paleontology, Nanjing): (a) xlO; (b) x25.

with Gigantonoclea-type leaves might have two or defined by Went ( 1971) and the reason for it might ha more different types of reproductive structures. The been the same-horizontal gene transfer over the g{ typical material from Shansi associates with Jiaochen­ graphic realm. gia, a pinnate microsporophyll with reduced lamina In the above interpretation, the Permian "floras" and semisynangiate sporangial clusters (Wang, 1999) "floristic provinces" are essentially the "morphocho while the morphologically similar leaves from South­ ons" defined by a few peculiar morphotypes that spre em China are accompanied by Gigantotheca sporo­ over taxonomic and ecological boundaries. Their cor phylls bearing imbricate synangia on unreduced leaf spondence to floristic or vegetational divisions is not blades (Fig. 1). In the gigantopterid leaves that I col­ be taken for granted, but is a matter of taxonomic a lected from the latest Permian (Changhsingian) of paleoecological analysis. Laibin, South China a structurally preserved axis shows tracheids with biseriate bordered pitting similar to those of the Permian conifers and peltasperms (Fig. 2). GENERAL ON PALEOVEGETATION ANALYSI Why certain leaf morphotypes were so widespread Paleovegetation analysis is based on the plant I in one geographic realm while rare or lacking in the fonns rather than taxa. Life fonns are groups of org< others remains enigmatic. Phylogenetic parallelism is isms adapted to a certain functional role in the bio not a plausible explanation for we are dealing with community. At the lower hierarchical levels th plant groups that are not even distantly related (e.g., the roughly coincide with species (though conspeci marattialean ferns and peltasmermous gymnosperms in organisms can fill more than one niche depending the case of gigantopterids). A biomic interpretation age, sex, etc.). In extinct plants the life form is ess{ (Krassilov and Naugolnykh, 2000) appears consistent tially an interpretation of morphological characte with a convergence of prevailing leaf morphotypes, as Fossil leaves are the most frequent plant macrofoss in the present day sclerophyllous vegetation. However, providing an important life form information, for th a purely ecological parallelism seems improbable for morphology relates to their structural position (in t the morphotypes that, as Glossopteris, might have canopy or understorey), habitat, climate, pollinati spread over a wide range of biotopes. The most similar mode, etc. For the dominant Permian plants we ha recent phenomenon is geographic parallelism as also evidence of their shoot morphology, axial an.

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Fig. 2. Anatomy of an axis with attached gigantoptcrid leaf form the latest Permian of Penlaitan Section. Talung Formation. Laibin. Southern China: (a) specimen no. 47931106-25. leafy shoot with axis partly preserved as compression. xI 0: (b) protoxylem tracheid with helical thickenings. SEM: (c. d) metaxylem tracheids with obligue pit-like series between helical coils and secondary xylem tracheids with biseriate pitting-on the radial walls. SEM.

omy, underground organs, reproductive structures and uous forms, accumulate as leaf mats. Coal is a fossil­ even their pollinators. Seasonal dynamics of plant com­ ized accumulation of plant dead mass and as such is munities is inferred, in addition to leaf morphology, evidence of highly productive wetland (helophyte) from the taphonomy of leaf assemblages that, in decid- communities.

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However this information is useful only in the con­ (Kryshtofovich, 1937; Meyen, 1970) covered Siberia, text of stratigraphic lithofacies sequences. For exam­ Petchora Basin in the northern Fore-Urals, most of ple, in the above mentioned Late Permian plant locality Mongolia and northern China north of 45° N. The of Laibin, South China (see Shen eta/., 1999 for strati­ Angaran flora is typically represented in the Kuznetsk graphic information), the plant bed occurs on the top of Basin, West Siberia, a major coal-producing area in a matine regressive sequence. It is about 50 em thick Asiatic Russia. Cordaites and the morphologically sim­ consisting, from base up, of a thin coal, a root bed with ilar Ru.floria are the dominant leaf genera often depos­ arthtophyte stems and abundant pecopterid fronds, and ited as one-species plant beds. a shale 'with Gigantonoclea and fragmentary conifers. TheAngaran Cordaites (Rujloria)-type leaves might The succession means an expansion of the arthrophyte­ . at least partly belong to the extinct gymnosperm order pecopterid coastal peatbog community immediately Vojnovskyales (Neuburg, 1965). Phylogenetic relation­ following the retreat of the sea. Later it was replaced by ships of Vojnovskyales to the typical Euramerian cor­ the riparian gigantopterid community. The conifers are daites is a matter of discussion (Meyen, 1984). In the rare, but they also occur as a transported material in present author's opinion (Krassilov, 1997) they are only marine strata downsection suggesting a more distant distantly related. The Vojnovskyales are reported from source at higher elevations. few localities outside northern Asia, such as Midtkap in Fossil plant assemblages are defined by the repeated Greenland and Mount Dall in Alaska (Wagner eta/., association of their dominant life forms. In the above 1982; LePage and Pfefferkorn, 1996; Naugolnykh, this example, we have two types of wetland assemblages, of volume). Among the spore-plants an arthrophyte group, which the fern-arthrophyte one is recognized world­ the Tchemovskiales appears characteristic of the, but wide, while the gigantopterid one is characteristic of not necessarily endemic to, the Angaran-type floras. the Cathaysian region. In other regions it was partly or completely substituted by the callipterid, cordaite or The geographic range of "cordaite flora" was glossopterid assemblages. This is a rough picture, but a thought to coincide with the borders of Angaraland. sufficient one to start with. However, the Angaran realm is linked with the Eurame­ rian callipterid realm by a broad zone of mixed floras The precursory Permian-type assemblages sporadi­ extending over the Urals, Volga Basin and to Arctic cally occur in the Late Carboniferous of both Laurasian Canada (Fig. 3). Many European taxa, such as Quadro­ and Gondwana landmasses. In Gondwana realm, a cladus, have spread over this transitional zone to the decline of the arboreal lepidophyte wetlands is tradi­ Petchora Basin, Taymyr (Sadovnikov, 1997) and fur­ tionally explained by climatic changes related to glaci­ ther east. Actually, the Late Permian uplift of Uralian ation. In the Euramerian realm the coeval vegetational Ranges suturing the closed seaway had a minor effect change is, l'i.kewise habitually, ascribed to aridity. How­ ever, the change took place worldwide, with humid on the position of phytogeographic boundaries that in non-glacial Cathaysia no exception. A drop in ground the Mesozoic maintained nearly the same position as in water levels caused by global regression at about the the Permian (Krassilov and Shorokhova. 1975; Krassi­ lov, 1997). Carboniferous-Permian boundary might have been critical for lepidophyte wetlands. On evidence of the bet­ A popular notion of the Angaran "cordaite taiga" is ter-dated tillites, glaciation was restricted by this trans­ incompatible with leaf morphology, for the present day boundary interval. New types of wetlands appeared with typical taiga is formed of evergreen needle-leaved a subsequent sea level rise in the Artinskian. Owing to trees. In the dry permafrost areas east ofYenisey River the role of wetlands as a major source of fossil plant it is replaced by the larch taiga of deciduous needle­ material, this event is commonly taken as the beginning leaved trees shedding needles from persistent spur­ of the fully developed Permian flora. shoots. The Cordaites-Ru.floria type of broad taeniate leaves with parallel venation commonly occurs in monocotyledonous angiosperms in which they are of a ARCTO-PERMIAN REALM phyllodic origin, derived from inflated petioles with the In most Laurasian localities the fern-arthrophyte leaf blade reduced. Phyllody relates to developmental peatland communities are associated with gymnosperm acceleration in tum related to rapid leaf expansion wetlands dominated by vojnovskyaleans, cordaites, required by a short growth season. In the cordaite peltasperms or the related gigantopterids. Despite con­ realm, the monodominant to oligodominant forests siderable geographic differentiation, these types of grew on raised peat bogs in which a considerable part wetlands intergrade in the broad ecotonal zones. There­ of biomass consisted of bryophytes (Durante, 1995). fore we can' speak of the Arcto-Permian realm as anal­ The Ru.floria-type leaves were shed separately, with ogous to the Holarctic or Arcto-Tertiary. clear-cut abscission lines, or clustered on deciduous spur shoots. They formed thick leaf-mats, the common­ est type of fossil plant assemblages in the coal mea­ Temperate Nemoral Zone sures. Since considerable peat accumulation requires a The Permian fossil plant assemblages of northern humid summer-wet climate, a combination of morpho­ Asia. known as the Angaran flora or floristic kingdom logical and taphonomic data indicate the presence of

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o I i!II !!III 0 IV •v • VI e VII o VIII

Fig. 3. Mid-Permian phytogeographic zones-the major biomes on the present day map (prepared with participation ofNaugolnykh: Krassilov and Naugolnykh, 2000). Shading: (J) temperate nemoral, (2) warm temperate meso-xeromorphic, (3) subtropical xero­ thermic, (4) subtropical humothermic, (5) Tethyan tropical and mixed. Symbols of floristic composition: 1-Angaran, 11-mixed Eurangarian, III-mixed Cathangarian, IV-Eauramerian, V-mixed Euramerian/Cathamerian. VI-Cathamerian, VII-mixed Eurogondwana, VIII-Gondwana. Localities: 1-Midtcap, 2-Petchora, 3-Westem Taimyr, 4--Eastern Taimyr, 5-Norilsk, 6- Tunguska, 7-Kuznetsk, 8-Verkhoyansk, 9-Sverdrup, 10-Soyana, 11-Kama, 12-Tchekarda, 13-Karaungir, 14--Fergana, 15-Junggar, 16-Tabun Tologoy, 17-North Korea, 18-Primorye, 19-Appalachian, 20-Cantabria,.21-0robic Alps, 22-Tur­ ingia, 23-Utah, 24--Arizona (Hermit Shale). 25-Kansas, 26-Guadalcanal, 27-High Atlas, 28-Hazro, 29-Saudi Arabia, 30-Iran, 31-Texas, 32-Mexico. 33-Shanxi. 34--Anhui/Henan, 35-Jiangsi, 36-Chubut, 37-Niger, 38-Gabon, 39-Kash­ mir, 40-Lhasa, 41-Yunnan, 42-Phetchabun, 43-Jambi, 44--1rian Jaya, 45-Transvaal, 46-Handapa, 47-Perth, 48~ Bowen, 49-Mount Achemar.

summer-green broad-leaved vegetation ecologically tation extended to Kazakhstan (Karaungir locality that similar to the present-day nemoral zone of the humid yielded seeds similar to Euramerian Salpingostoma: temperate climate. Krassilov, 1999), Central Asia and northern China. On palynological evidence, the warm-temeprate vegetation zone can be traced to Greenland and Arctic Canada Wann-Temperate Ecotone (Utting, 1994). The Permian flora of Fore-Urals and Volga Basin is considered as a broad ecotone of callipterid and cor­ In China, the Permian floras of Junggar-Khingan daite vegetation. It is dominated by Euramerian pel­ (Hinggan) latitudinal area are considered as belonging tasperms, conifers and their derived forms (Zalessky, to Angaran realm (Li and Wu, 1996; Li, 1997; Huang 1927; Naugolnykh, 1998). The latest Permian assem­ and Ding, 1998). Actually they are of ecotonal aspect, blages of this area constitute the so-called Tatarina with the Volgian Tatarbza and Phylladoderma admixed flora named after a highly polymorphic leaf-genus of by the Cathaysian taeniopterids (Wang and Wang, peltasperms the lobate variants of which resemble both 1986). This zone is traced to southern Mongolia and the European Callipteris and American Supaia Primorye in the Rul!sian Far East were the dominant (Gomankov and Meyen, 1986). "Tatarina flora" can be Rufloria-Cordaites morphotypes associate with gigan­ seen just as a local variant of the "Callipteris flora." The topterids, a feature shared with floras of North Korea, Angaran ruflorias and representatives of the fern - tch­ Anhui and adjacent provincies in eastern China (Mei ernovskian wetland communities formed a significant, eta[., 1996) and perhaps typical of the Pacific coastal yet subordinate element. To the east the ecotonal vege- areas. Notably the marine f~tunas of fusulinids and bra-

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chiopods from the same latitudues are also of a mixed range consists of a. number of tectonic blocks at one boreaJ/tropical aspect (Sib and Zhao, 1966). time or another separated by seaways. The widely scat­ tered Middle East lEurope and eastern North America are dominated by niopteroid fertile leaves with marginal ovules (Mamay callipterids (peltasperms with theAutunia-type strobili) et al., 1988; Mei et al., 1992). On the other hand, and conifers, with a highly diverse fern-arthrophyte Supaia, the emblematic North American genus, has components. The boundary with Cathaysia over south­ been recently recorded from China (Wang, 1997). ern Xinjang and along the northern border of Tarim Therefore, the so called Cathaysian flora was actually Basin to the Lesser Khingan (Li, 1997) was easily sur.:. Cathamerian. mountable for these components that were also wide­ spread in Gondwana floras (see below). In China, the Cathaysian (Cathamerian) flora is presently subdivided into the northern and southern The Euramerian peltasperms and conifers had xero­ provinces supposedly confined to disjunct Cathaysian morphic leaves with thick cuticles that are usually asso­ terrains on the eastern periphery of Tethys. The north­ ciated with dry climates. However leaf xeromorphism em Cathaysian flora occurs on the Sino-Korean....:.Tarim is also a typical feature of wetland (helophytic) habi­ block bound from the south by the latitudinal Kunlun - tats. Preservational features of xeromorphic Permian Ointing fault zone traceable to Central Japan (Li, plants suggest their proximity to deposition sites. Their 1997). However the floristic differences between the remains often occur in paralic facies indicating a northern and southern provinces involve a few endemic coastal, perhaps mangrove, community. genera, such as Emplectopteridium and Gigantotheca Another type of wetland vegetation, with abundant (Li et al., 1996; Li, 1997). More significant are quanti­ pecopterids and calamites, is here designated as the tative distinctions, with Otofolium, Gigantopteris and fern-arthrophyte marsh community. These two types Rajahia common in the south in contrast to the predom­ might intergrade in the marginal habitats. Yet in strati­ inantly northern Cathaysiopteris, Emplectopteris, large graphic sequences they often alternate, with the fern­ Taeniopteris and Yuania. Such distinctions sugges~ cli­ arthtophyte plant beds mostly confined to regressive matic differentiation rather than isolation. The Permian parts of sedimentary cycles. The peltasperm-conifer flora of Xu-Huai-Yu region in eastern-central China (at assemblages typically came from estuarine deposits the junction of Jiangsu, Anhui and Henan provinces) is (e.g., the cuticle beds of Nedubrovo: Krassilov et al., considered as a separate floristic subprovince (Mei 1999), whereas the fem-arthrophyte assemblages more et al., 1996), but is, in a sense, transitional between the commonly occur in deltaic deposits. Their stratigraphic northern and southern Cathaysian floras, having both recurrence might have been related to sea level fluctua­ Cathaysiopteris and Gigantopteris as equally common tions. elements. A palaeobotanical standpoint would hold that 'fhe Euramerian plant localities commonly associ­ the northern and southern Chinese blocks had already ate with red or variegated beds that are traditionally collided by the Devon~an (Li et al., 1996; Li, 1997). conceived of as evidence of arid climate. '!Ypical red­ In the Early Permian, the typical Cathaysian flora of beds are molassoids of the Hercynian foldbelts that Shansi (Shanxi) Formation with Lobatannularia and served as a source of ferruginous clastic material. Emplectopteris replaced the essentially lepidophyte A change from ferrous to ferric oxides (green to red flora of the conformably underlying Taiyuan Forma­ beds) relates to frequent fluctuations of ground water tion. This rather abrupt floristic change at about the level reflecting not so much a low mean precipitation, Asselian/Sakmarian boundary (Li, 1997) might reflect as a seasonal dryness of a widely variable duration. a substitution of cosmopolitan lepidophyte wetland Marginal variants of redbed facies associate with either community by the newly appearing arboreal wetlands evaporites or coals. On the sum of evidence, the climate dominated by taeniopterids, emplectopterids and of the Permian xeromorphic zone might have been gigantopterids. In the Lower Shihhotse these. assem­ close to the modem Mediterranean climate. blages occur in the coal measures that in the Upper Shi­ hhotse are replaced by redbeds with Gigantonoclea Humothermic Zone (Wang, 1999). In the latest Permian the Northern Cathaysian province was invaded by European conifers Gigantopterids and emplectopterids are characteris­ and peltasperms (also by the Gondwana glossopterids, tic of the Permian plant localities in China, Korea, see below), while the typical gigantopterid flora sur­ Japan and Laos constituting the Cathaysian flora. To vived in the southern Cathaysia alone. southeast they extend to Thailand, Malaysia, Indonesia and New Guinea. Typical of gigantopterid vegetation In the American southwest, the Proto-Cordillera are also large taeniopterid leaves and diverse pecop­ Province was already differentiated by the Late Penn­ terids, at least partly shared with the xerothermjc zone, sylvanian (Reed, 1947). Hete, as elsewhere, the fem­ lobatannularias and other arthrophytes. Their Asiatic arthrophyte marshes expanded at the expense of lepido-

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phyte communities (Reed and Mamay, 1964). The Per­ wanaland separated by the latitudinal Tethys Ocean mian plant assemblages of Texas and Oklahoma are of from the Laurasian landmass in the north. This scheme a "Cathaysian" aspect owing to the commonly occur­ has encouraged such eminent paleobotanists as Florin ring gigantopterid and taeniopterid morphotypes, the and Meyen to develop separate phylogenies for the fertile variants of which bore marginal ovules (Mamay "northern" and "southern" gymnosperms. Subsequent et al., 1988). Of the gigantopterids, Cathaysiopteris discoveries of Glossopteris or related leaf morphotypes and Gigantonoclea morphotypes are formally shared north of Himalayas were considered as evidence of with northern Cathaysia (though probably convergent), either episodic trans-Tethyan land connections or the while the other three, Gigantopte,ridium, Delnortea and larger Gondwanaland comprising Tibet, Iran, Anatolia, Zeilleropteris are endemic. The North American gigan­ etc. (which, incidentally, would place the Tethys-Hima­ topterid province borders on xeromorphic vegetation of layas inside the Gondwanaland). Utah and Arizona with Supaia and Walchia and of Kan­ sas with Glenopteris (Reed and Mamay, 1964). The classical Gondwana flora came from theNar­ mada-Son-Mahandi graben system that transects the Thus xeromorphic vegetation penetrated both the peninsular India. Boulder beds occur at the base of their Cathaysian and North American parts of gigantopterid basinal sedimentary fill followed by coal measures that realm. Floristic exchanges of the latter with xerother­ contain the bulk of the flora. Himalayan occurrences mic zone indicate similar climatic conditions. However are scattered from Kashmir to Assam, with a few outli­ a widespread coal accumulation suggests a relatively ers in Tibet, Yunnan and the "mixed" floras of Indone­ humid climate of Cathaysia in the Early Permian. The sia and New Guinea. A number of well-defined species Dadoxylon-type fossil woods from this area show are shared by the Indian and Australian floras, but other vaguely marked growth rings (Sze, 1952). Apparently, parts of "Gondwanaland" are floristically less similar the Northern Cathaysian and Euramerian realms had (Archangelsky, 1990, 1996; Maheshwari, 1992). similar temperature conditions but differed in precipita­ tion as the present day arid and humid subtropical A primitive glossopterid morphotype Gangamopt­ areas. In the Late Permian their climatic differences eris lacking distinct midrib appears in the pre-tillite became less pronounced. Xeromorphic vegetation deposits already. The early Gangamopteris plants are eventually replaced the native gigantopterid communi­ commonly envisaged as growing in sparse vegetation ties by the end of the Permian. patches of a glacial landscape. However the fossil The gigantopterids and emplectopterids, prominent plants assemblages found immediately below and in the Cathaysian-type assemblages of eastern Asia and above the Dwykka Tillite in South Africa, Bacchus western North America, are characterized by a compos­ Marsh Tillite in Australia and Talchir boulder beds in ite nature of their fronds in which the entire or simply peninsular India suggest a less dismal picture, with a pinnate leaf blades were formed by marginal fusion of number of Carboniferous survivors (Chandra et al., the ultimate and penultimate segments, with the origi­ 1992; Pant, 1996). The leaf morphology gives no support nal segmentation betrayed by the conservative venation to either the "Gangamopteris tundra" or "Gangamopt­ pattern (Asama, 1960; Krassilov, 1995). The hierachial eris taiga." Gangamopteris is linked to Zamiopteris­ areolate venation of such leaves owes its origin to mer­ Cordaites via Palaeovittaria and Noegerrathiopsis istematic fusion zones acting as plate meristems implicating similar climatic conditions, i.e., temperate, (Krassilov, 1995). This leaf morphology is evidence of of the nemoral zone. prolonged marginal meristematic activity characteristic of megaphyllous life forms of equable climates. An The emblematic morphotype Glossopteris emerged advanced Gigantopteris morphotype, some variants of and diversified in the period of widespread coal accu­ which showing clearly defined drip-tips (Yao, 1986) mulation. Some variants of its highly polymorphic occur in Southern Cathaysia the tropical climate of leaves show distinct abscission scars, the others had which is also indicated by the tree ferns Psaronius and drip-tips, a character of rain-forest foliage. Such mor­ Lepidodendron surviving from the Carboniferous. phological diversity obviously suggests a wide range of habitats (Pant, 1996) which is confirmed by tapho­ Thus, in the Arcto-Permian realm, we can prelimi­ nomic data (Retallack, 1980). Glossopterids are some­ nary recognize the climatically defined zones of the times preserved as leaf mats containing spur-shoots temperate nemoral, xerothermic and homothermic sub­ shed with clusters of intact leaves, as in deciduous tropical vegetation. The thermophilic assemblages of riparian trees. A constant association of Glossopteris Southern China actually belong to the Permo-Thetyan with Vertebraria, an aerenchymatous underground Realm (below). organ, suggests a peat bog habitat. The coriaceous lin­ gulate leaves, as in the modern mangroves, and the uniquely preserved seedlings (Pant and Nat.Itiyal, 1987) NOTO-PERMIAN REALM are also consistent with a helophyte life form of a warm In the 1920s the apparently uniform Late Paleozoic temperate to subtropical climate. The maximal mor­ Glossopteris flora of India and southern continents phological diversity of G{ossopteris (42 species from a prompted the idea of Paleozoic supercontinent Gond- single locality) is reported from the redbed Late Per-

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mian of Handapa, Mahandi Basin (Chandra and Singh, present day arboreal vegetation, only some riparian 1992). plants, such as willows and poplars, have comparable An impression of the floristic integrity of Gondwana latitudinal ranges. is due primarily to ubiquitious glossopterids that are conspicuous and often numerically dominant fossils. PERMO-THETYAN REALM However, the uniformity of glossopterids is only valid at higher taxonomic levels; regional species composi­ The Tetyan phytogeographic realm is a continuous tions might have been highly endemic (McLoughlin, zone of mixed Cathaysian and Gondwana floras 1994). Other components show less uniform distribu­ extending from southern Spain and Morocco over Mid­ tion, with significant differences between the eastern dle East and Anatolia and North Africa (Wagner, 1962; and western Gondwanalands (Maheshwari, 1992) or Broutin, 1977; El-Khayal et a/., 1980; Lejal-Nicol, the Notoafroamerican, Australoindian and Palaeoant­ 1990; Aassoumi, 1994; Broutin et al., 1995) to Tibet, arctic provinces of Li (1997). Yunnan, Thailand (Phetchabun), Sumatra (Jambi) and West New Guinea (Irian Jaya). Ecological equivalents The differentiation primarily pertains to pecopterid­ of the Southern Cathaysian vegetation might occur on arthrophyte assemblages South America have a considerable "Euramerian" com­ The Cathasian gigantopterids occur in the Permian ponent of thermophilic pteridophytes (Archangelsky, floras of Anatolia, Thailand, southern Malaysia and 1990, 1996) indicating climatic conditions n?t too dif­ New Guinea (Kon'no, 1963; Asama et al., 1975; Wag­ ferent from those in the northern xerothermic zone of ner, 1962), whereas the Gondwana elements, prevail in Europe and eastern North America. Fossil woods from the mixed floras of Kashmir, southern Tibet (Lhasa) Chubut often lacks distinct growth rings (Archangel­ and Yunnan where they associate with the Cathaysian sky, 1996). Notably, a thermophilic Southern Cathar­ pecopterids, calamites and sphenophylls (Singh et a/., sian Pectinopteris or a closely allied form occurs m 1982; Pant eta/., 1984; Rigby, 1989, 1996; Li and Wu, South Africa and elsewhere (Wan and Basinger, 1992). 1996; Chandra and Sun, 1997). The Cathaysian and At the same time, the widespread coal accumulation in Gondwana type assemblages sometimes occur in dif­ India and Australia (Retallack, 1982; Balme et a/., ferent localities separated by tectonic fault zones and 1995) suggests a relatively humid variant of warm tem­ interpreted as separate terraines (Li and Wu, 1996). perate to subtropical climate, similar to that of Northern Alternatively, such differentiation can be ascribed to a Cathaysia. mosaic vegetation structure including the glossopterid The Permian plant assemblages of Antarctica differ and gigantopterid communities as distinct ecotypes. form the rest of Gondwana realm in a lower taxonomic Both macrofossil and palynological data show a diversity and a scarcity of pteridophytes (Archangel­ combination of the Euramerian, Cathaysian and Gond­ sky, 1990; Cuneo et al., 1993). The latter feature was wana elements in the Permian western Tethys extend­ .emphasized by Cuneo eta/. (1993) as indicating a lack ing over Northern Africa, Middle East, Iran and Paki­ of undergrowth. As suggested above, the diverse pteri­ stan (Aassoumi, 1994; Broutin, 1977; Lejal-Nicol, dophyte component associates with a special type of 1990; Broutin eta/., 1995; Utting, Piasecki, 1996; Igbal fern-arthrophyte wetlands as different from the gym­ eta/., 1998). In palynological assemblages of Yunnan nosperm (peltasperm, glossopterid) wetlands with the Gondwana elements amount to 80% (Gao, 1998). scarce pteridophytes of a few peculiar species. Inciden­ Glossopterids might penetrate t~e Cathaysian rea~m tally, the Antarctic glossopterid assemblage co~tains a through trans-Himalayan connectiOns over Kashmir­ peculiar fern Skaaripteris, suggestedly aerenchimatous Tibet and along the Pacific coast as far north as Pri­ and probably aquatic or semiaquatic (Galtier and Tay­ morye where both Gangamopteris and Glossopteris lor, 1994). Therefore, the Antarctic Permian climates were found in a single locality (Zimina, 1967). Abrop­ were not too harsh for fern growth in general, but less topteris Mo and the recently found Laibinopteris favorable for the pecopterid-arthrophyte communities Krassilov from the latest Permian of South China are than in the other parts ofNoto-Permian realm. The Ant­ similar to bracteate fructifications of glossopterids that arctic "fossil forest" is a dense growth of small to once more became rather common in the Triassic of medium sized trees with Vertebraria pneumatophores. northern-central China (Meng, 1995). The Gangamopteris and, later, Glossopteris asse.m­ blages are associated with coal seams about 2 ~ thick containing stems 50 em in diameters (McLoughhn and DISCUSSION Drinnan, 1997). The seams contain clastic dikes that A conclusion from the foregoing review is that the might form upon permafrost veins of paisa mires as in Permian "floristic" divisions could be more consis­ the present day cool temperate areas in Canada (Krull, tently described as the ecogeographic vegetational 1999). biomes controlled by climatic zonation. At about the The Glossopteris vegetation thus extended from Carboniferous-Permian boundary the cosmopolitan cool temperate to subtropical climatic zones. In the lepidophyte wetlands were replaced by new types of

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plant commumttes dominated by various gymno­ and climatological grounds. In a glacial climate, the sperms. These vegetational changes are associated with "Gondwanaland" interiors would experience dry cli­ a global climatic change, regression and the lowering of mate, with cyclonic air masses barred from penetration ground water level. Climatic differentiation of new inland by cold atmospheric fronts. Neither coal accu­ plant communities had defined the aspects of the mulation nor permanent ice cover would have been sus­ regional fossil plant assemblages. They are preliminary tainable in such conditions. The apparent floristic assigned to the temperate (Angara and Antarctica homogeneity is due primarily to a broad ecological realms), xeric subtropical (Eurameria and West Gond­ range of glossopterids. However, ecologically the arbo­ wana), humid subtropical (Nqrth Cathaysia and East real broad-leaved glossopterid communities had little Gondwana) and tropical (Tethyan) zones. in common with the Pleistocene to recent periglacial These Permian zones are roughly parallel to the vegetation, suggesting a totally different character of present day latitudinal belts thus scarcely compatible the Permian glacial events. Notably, the Triassic glos­ with paleogeographic reconstructions in which Laur­ sopterid "Gondwana" was a redbed zone extending asia is placed submeridionally, with the Tethys widely from India to northern China. gaping to the east (Scotese, 1984, etc.). Actually their The continental reassemblies implying the Permian position on the present-day geographic maps does not South Pole near Africa and the .Euramerian realm at seem anomalous. Thus the Angaran realm roughly cor­ equator are not compatible with the Permian phyto­ responds to the zone of northen conifer forests and tun­ geography. On both paleovegetational and lithological dras (which does not mean that they are ecologically evidence, central Antarctica experienced a much cooler equivalent; simply there is a phytogeographic realm of Permian climate than South Africa or South America the same extent on the modern map). Its western which implies the Permian South Pole close to its boundary is oblique to the modern latitudes which does present day position. From ecological standpoints, the not necessarily means rotation of the continent for the Euramerian plant assemblages had little in common corresponding boundaries are similarly oblique in the with either humid or arid variants of tropical vegeta­ Mesozoic (Krassilov, 1972, 1997 and elsewhere), Ter­ tion. They were pligodominant, of xeromorphic ever­ tiary and at ~resent even. green (rather than deciduous) aspect and with abundant Likewise, the geographic range of Glossopteris is well preserved conifer remains that obviously came fairly similar to that of the present day "Gondwana" from lowland sources (now conifers are lacking in trop­ family Proteaceae that appeared about 100 Ma after the ical lowlands). There were free floristic exchanges supposed break up of Gondwanaland. Notably, the between the Euramerian and the temperate Angaraland mixed Cathaysian-Gondwana flora of New Guinea realms via a warm-temperate ecotone over Eastern occurs fairly close to the present day WalJace line. Europe and the Urals. A widely held biogeographic opinion links such The Euramerian-Cathamerian-Gondwana "floristic present day geographic features with past distribution mixing" might result from penetration of interzonal of landmasses. But traced back to the Permian they are wetland communities (Fig. 4) in disturbed vegetation of scarcely compatible with a radical reassembly of the the tectonically active Tethys realm. In the high relief continents. areas there might happen also a taphonomic mixing of Neither of the Permian phytogeographic realms was plant remains coming from different altitudinal belts. restricted to a separate landmass. Their expansion over As shown on the modern map, Tethys corresponds the Uralian, Tibetan and Ginling sutures is consistent to a great shear zone that extends over the Sulaiman with intermittent seaways rather than vast oceans. Ranges in Pakistan, Karokaram-Nujiang fault zone in Suturing of the Uralian geosyncline had a minor effect Tibet, Arakan Yoma Range in Burma and Andaman­ on the Angaran biome that preserved its climatic and Nikobar Ridge to Sunda Trensh, Timor and the Owen vegetational identity far beyond its supposed isolation Stanley Range in New Guinea (reviewed in Krassilov, as a landmass. In fact, the Triassic Siberian realm still 1985). The Tethyan ophoilites, that are interpreted as coincided with the "Angaraland" (Krassilov and abducted remains of oceanic crust, occur in narrow Shorokhova, 1975) and even the later vegetational-di­ zones bordering numerous blocks of continental crust matic boundaries still tracked the familiar outline the larger of which are the Mediterranean (divisible in (Krassilov, 1972; 1997). several minor blocks), Taurian, Arabian, Beshte-Lut in In much the same way, the seaways of central and Iran, Helmend in Afganistan, Quangtang in Tibet, southern China had a little effect on floristic similarity Gangdis in India and Sunda in southestern Asia. At the of the northern and southern Cathaysian blocks. Their times of Tethyan transgression most of their area might floristic distinction increased (rather than decreased) in have been covered with epeiric seas. But the Permian the latest Late Permian owing to expansion of the dry floristic mixing is incompatible with the idea of Tethys subtropical belt that brought a wave of European as a broad ocean between Laurasia and "Gondwana­ migrants to northern China. land", with fragments of the latter rafted and coli ided to The integrity of the Gondwana realm as a continu­ mainland Asia at different times during Mesozoic and. ous landmass is questionable on both phytogeographic early Tertiary, as implied by plate tectonics. Floristic

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T ST E ST T Aassoumi, M.H., Les Paleojlores du Permien du Maroc Cen­ tral, These du Dr. de l'Univ. Paris, vol. 6, 1994. Axelrod, D.I., Evolution of the Madro-Tertiary Geoflora, Bot. Rev., 1958, vol. 24, no 7, pp. 433-503. Balme, B.E., Kershaw, A.P., and Webb, J.A., Floras of Aus­ tralian Coal Measures, with Notes on their associated Meso­ . zoic Faunas, in Geology ofAustralian Coal Basins. Ward, C.R., Harrington, H.J., Mallett, C.W., and Beeston, J.W., Eds., Geol. Soc. Australia, 1995, spec. publ., no. I, pp. 41-62. Broutin, J ., Nouvelles donnes sur Ia ftore des bassins Autuno­ Stephaniens des environ de Guadalcanal (province de I Seville-Espagne), Cuademos Geologia Iberica, 1977, vol. 4, I pp. 91-98. Pecopte!ldi-sphen~phyllion 1 Broutin, J., Roger, J., Plate], J.P., Angiolini, L., Baud, A., Bucher, H., Marcoux, J., and AI Hasmi, H., The Permian Pangea Phytogeographic Implications of New Paleontologi­ cal Discoveries in Oman (Arabian Peninsula), C.R. Acad. Fig. 4. Schematic zonal distribution of the typical mid-Per­ Sci. Paris, 1995, vol. 321, ser.lla, pp. 1069-1086. mian plant communities. Vertical columns represent phyto­ climatic zones (biomes): T-temperate, ST-warm temper­ Chandra, S. and Singh, K.J., The Genus Glossopteris from ate, E-equatorial. the Late Permian Beds of Handapa, Orissa, India, Rev. Palaeobot. Palynol., 1992, vol. 75, pp. 183-218. Chandra, S., Srivastawa, A.K., and Singh, K.S., Lawer Per­ links across the ophiolite sutures mean that the Tethyan mian Fossils from India and Early Developmental History of blocks were parts of a continuous fold belt in the Per­ the Glossopteris Flora, Acta Palaeobot. Polon., 1992, mian already. vol. 32, no. 1, pp. 5-19. The tectonic style in the western part of this zone is Chandra, S. and Sun, K., Evolution and Comparison of the Gondwana Flora and the Cathaysia Flora, Palaeobotanist, defined by the large-scale right-lateral shear that might 1997, vol. 46, no 3, pp. 35-46. bt;ought continental blocks of southeastern Asia to the Cuneo, R., Isbell, J., Taylor, E.L., and Taylor, T.N., The Glos­ north of their pre-Permian position close to Sunda sopteris Flora from Antarctica: Taphonomy and Paleoecol­ plate. A displacement of this kind would explain both ogy, C. R. XI/ICC-P(BuenosAires), 1993, vol. 2, pp. 13-40. the Indo-Australian floristic links and the constriction Durante, M.V., Reconstruction of Climatic Changes in the of tropical zone over the junction of Tibet and Himala­ Late Paleozoic of Angarida (on the Basis of Phytogeographic yas. In the Earth's rotation stress field the maximal lat­ Data), Stratigr. Geol. Korrel., 1995, vol. 3, no. 2, pp. 25-37. itudinal shear zone (as reflected in transform faulting of El-Khayal, A.A., Chaloner, W.G., and Hill, C.R., Palaeozoic mid-oceanic ridges) corresponds to the equatorial min­ Plants from Saudi Arabia, Nature, 1980, vol. 285, no. 1, imum of Coriolis force (Tanner, 1964; Krause, 1970). pp. 33-34. This consideration supports a tropical position of the Galtier, J. and Taylor, T.N., The First Record of Ferns from Permo-Tethyan realm. the Permian of Antarctica, Rev. Palaeobot. Palynol., 1994, vol. 83, pp. 227-239. ACKNOWLEDGMENTS Gao, L., On the Discovery of a Gondwana Affinity Microf­ lora from Baoshan, West Yunnan and Its Geologic Signifi­ This work was supported by the Russian Foundation cance, Acta Geoscientia Sinica. Bull. Chinese Acad. Geol. of Basic Research, grant no. 98-04-49010 and by the Sci., 1998, vol. 19, no 48, pp. 105-112. Federal Scientific Technical Program. 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