Permian conifers of Western Angaraland

ELS EVIER

Review of Palaeobotany and Palynology96 (1997) 351-447

REVIEW OF PALAEOBOTANY AND PALYNOLOGY

Permian conifers of Western Angaraland Sergei V. Meyen *'*

Abstract Coniferalean genera and species for which fructifications and/or epidermal-cuticular characters are known, are described for material coming from the Kungurian, Ufimian (?), Kazanian and Tatarian stages of the Russian Platform and Fore-Urals. Kungurodendron Meyen, gen. nov. comprises both female and male fructifications and vegetative shoots. The axillary complex is flattened, seed stalks numerous, abaxially situated, with hooked apex, seeds apically attached. Pollen with columellar-like sexine. In Timanostrobus Meyen, gen. nov. lateral polysperms bear numerous seed stalks and sterile scales, pollen asaccate with perforated tectum. Associating leafy shoots of the Brachyphyllum-Pagiophyllum type. Concholepis Meyen, gen. nov. includes compound polysperms having false (?) seed scales that bear numerous abaxial and marginal appendages. Microstrobili associating with polysperms of Sashinia and the leafy shoots Quadrocladus, and producing pollen Scutasporites are referred to Dvinostrobus. The diagnosis of the genus Steirophyllum is emended. This genus has been erroneously merged with Ullmannia, although it appears to be an older synonym of Quadrocladus. New combinations Cyparissidium appressum (Zalessky) Meyen, comb. nov. and Taxodiella bardaeana (Zalessky) Meyen, comb. nov. are proposed. Several species of Pseudovoltzia (?), Cyparissidium, Sashinia, Quadrocladus and Geinitzia are described. With regard to the data on all these taxa and results of the revision of Lebachia, Ernestiodendron and other Euramerican conifers, main aspects of the evolutionary morphology of fructifications, pollen and vegetative shoots are treated. A new phylogeny of the early conifers is proposed. Both the overall system of conifers and names of the extinct suprageneric taxa should be based on genera typified by female fructifications. Instead of the family Walchiaceae (= Lebachiaceae), the new family Lebachiellaceae is proposed. Its type genus Lebachiella Meyen, gen. nov. includes polysperms previously referred to Lebachia or Walchia. The generic name Lebachia is illegitimate. Walchia is regarded as a satellite genus of the family Lebachiellaceae. A new system of the early conifers is suggested. Florogenetic consequences of the study are treated. Keywords: Pinales; Evolutionary morphology; Systematics; Phylogeny; Upper Palaeozoic; Western Angaraland

1. Introduction The current reconstructions of the earlier phases of coniferalean phylogeny and the derivation of the order Pinales from the Cordaitanthales are

* Correspondence should be addressed to A.V. Gomankov or I.A. Ignatiev, Geological Institute, Russian Academy of Sciences, Pyzhevskyper. 7, Moscow 109017, Russia. 0034-6667/97/$17.00 © 1997 Elsevier Science B.V. All rights reserved PII S0034-6667 (96)00046-2

mostly based on the Euramerican late Palaeozoic Cordaitanthus (= Cordaianthus), Lebachia, Ernestiodendron, Pseudovoltzia, Ullmannia and a few Mesozoic conifers. Having examined these taxa, Florin (1938, 1939, 1940, 1944, 1945, 1951) concluded that the coniferalean seed scale is homologous to the female axillary complex of Cordaitanthus. He held that in the lineage from cordaites to advanced conifers, the axillary complex has become dorsiventral, its sterile scales have

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been reduced in number and fused into the seed scale to which the initially free seed stalks and the bract became adnate. The genus Pseudovoltzia with its free seed stalks linked the Lebachiaceae with more advanced conifers having true seed scales. Schweitzer (1963) has shown that in Pseudovoltzia there were no free seed stalks and the seeds were attached directly to lobes of a true seed scale, as Walton (1928) had thought earlier. Schweitzer "has broken the thread of Florin's story" (Harris, 1976, p. 126) and suggested a quite different coniferalean phylogeny. On the other hand, Pant and Nautiyal (1967) and Pant (1977) derived lebachians from the Gondwana Buriadia-like conifers. The origin of taxads stands apart t¥om the aforementioned issues. Florin held that taxads do not belong to the conifers, and constitute an independent taxon of the same rank. However, Harris (1976, p. 131 ) opined that, "the game of comparative morphology ... within the rules" would allow the derivation of taxads from lebachians. The above is only a brief outline of the major controversies in early coniferalean phylogeny. Studies over the last several years (GrauvogelStamm, 1978; Clement-Westerhof, 1984; Mapes and Rothwell, 1984; Meyen, 1976-1978, 1981, 1982a, 1984; Scott and Chaloner, 1983) have widened the circle of better studied early conifers. Simultaneously, serious errors have been revealed in the traditional interpretations of fructifications of the genera Ernestiodendron and Lebachia. Nomenclatural questions entailed by the illegitimate status of the names Lebachia and Lebachiaceae (Clement-Westerhof, 1984) have arisen. The presently described Permian conifers of Western Angaraland (i.e., Russian Platform and Fore-Urals) essentially widen the diversity of female fructifications, pollen and epidermal characters of early conifers. Some of the described conifers (from the Kungurian, Ufimian and Kazanian stages) partly fill the hiatus between West European conifers of the Rotliegend and Zechstein (Fig. 1 ). Many questions of the evolution of conifers of Western Angaraland entail certain phytogeographic and stratigraphic consequences.

The present paper does not provide an exhaustive revision of all earlier described taxa. Below are described mostly taxa tbr which fructifications and/or epidermal-cuticular characters are known. In the author's collection there are also numerous vegetative shoots lacking cuticle and devoid of salient morphological features. A systematization of this material would not be interesting both taxonomically and stratigraphically. Such conifers will be briefly described in Sections 2 and 5.9. The following taxa (in order of treatment) are described and discussed in the systematic part ( Section 5 ).

Kungurodendron Meyen, gen. nov. Kungurodendron sharovii Meyen, sp. nov. Kungurodendron? sp. SVM-1 ()'parissidium Heer, 1874 Cyparissidium appressum (Zalessky) Meyen, comb. nov.

Cyparissidium entsovae Meyen, sp. nov. 7"imanostrobus Meyen, gen. nov. Timanostrobus muravievii Meyen, sp. nov. Concholepis Meyen, gen. nov. Concholepis harrisii Meyen, sp. nov. Sashinia Meyen, 1978 Sashinia aristovensis Meyen Sashinia borealis Meyen Dvinostrobus Gomankov et Meyen, 1986 Dvinostrobus sagittalis Gomankov et Meyen Quadrocladus M~dler, 1957 Quadrocladus (al. Steirophyllum) komiensis Meyen, sp. nov. Quadrocladus dvmensis Meyen Quadrocladus borealis Meyen Quadrocladus schweitzeri Meyen Steirophyllum Eichwald, 1854 Steirophyllum lanceolatum Eichwald Geinitzia Endlicher, 1847 Geinitzia subangarica Meyen Geinitzia sp. SVM-I, SVM-2 Taxodiella Zalessky, 1939 Taxodiella bardaeana (Zalessky) Meyen, comb. n o ~'.

Pseudovoltzia Florin, 1927 Pseudovoltzia? cornuta Meyen, sp. nov.

S. V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447 EQUATORIAL BELT

WESTERN ANGARALAND U//matvKa, Ps~dovoltzia, Quadrocladus, Culmitzschia

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__ Pseudovoltzia ?cornuta Ouadracladu$ (a/. Stelrophyllum) komiensis Tirnanos~robus muravievlJ Concholepis harrisii

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Fig. 1. Stratigraphicposition of Upper Palaeozoic conifers in the Equatorial belt (to left) and Western Angaraland (to right).

The material of the paper has been collected by the author and persons mentioned in Section 3. All the specimens unless otherwise stated are kept in the Geological Institute, Russian Academy of Sciences. Compressions were macerated in Schulze's liquid. Transfer preparations were made both from fructifications and leafy shoots. Drawings were

prepared by the author with a camera lucida, traced from photographs or with the aid of synthetic tracing paper which covered specimens and was moistened with alcohol. When examining specimens, crossed polarizing filters were frequently used. Material was illuminated with polarized light and observed through a "crossed" polaroid filter to avoid flare and enhance contrast.

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S. ~ Meyen / Review1 t~/ Palaeoholany and Pa/vnolog.!' 96 (1997) 351 447

2. Historical sketch

The Permian conifers of Western Angaraland were first described in the middle of the last century together with other plant remains under the generic names Araucarites, Pinus, Steirophyllum, Ullmannia, Voltzia and Walchia (Eichwald, 1854, 1854-1861, 1855, 1859-1860: Goeppert, 1864 1865; Golovkinsky, 1869; Kutorga, 1844: Ludwig, 1863; Schmalhausen, 1887). Most of these conifers came from cuprous sandstones of the Upper Permian and were represented by small lYagments of vegetative shoots. Most of the originals to the listed papers have been lost or their location is unknown. A part of Eichwald's collection is kept in the Museum of the Leningrad University. The specimens are accompanied by labels with his identifications, but the correspondence between the specimens and published figures cannot be established with certainty. Some of these specimens were figured by Zalessky (1927) who noted that the correspondence between Eichwald's specimens and figures is uncertain. Judging from available specimens and published illustrations, the conifers described in the listed papers are divided into two groups. One group includes shoots with short broad leaves and obtuse apices. They were mostly described under the name Ullmannia bronnii although Eichwald called them Steirophyllum lanceolatum Eichwald (see Section 5.8.1). Conifers of the second group have narrower, longer and more crowded leaves. Eichwald described them as Cupressites biarmicus Eichwald. Goeppert (1864-1865) proposed the combination UIImannia biarmica that has been followed in the literature. Walchia./bliosa belongs to the same group. In the 1920-1930s the conifers of Western Angaraland were studied by Zalessky (1927, 1929, 1937, 1939). At the outset he figured several specimens earlier studied by Eichwald and Kutorga as well as some additional ones (Zalessky, 1927). Most of the specimens belong to two aforementioned groups. He also figured a specimen ascribed to Walchia hypnoides Brongn. from the Lower Permian of the middle Fore-Urals (Zalessky, 1927, pl. 33, figs. 5, 5a). This specimen was referred by Florin (1939, p. 224) to a new species Walchia

(Lebachia?) stricta Florin together with one specimen from the Lower Permian of Oklahoma. Zalessky also figured some badly preserved conifers that cannot be identified. Later on, Zalessky (1929) described three new species of conifers from the Kazanian beds of the Kama River embayment. Ullmannia morkvaschica Zalessky includes a shoot with irregularly arranged appendages. The coniferalean affinity of the specimen is not evident. In its gross morphology it resembles shoots of Phylladoderma with persistent leaf bases. Similar shoots were described by Vladimirovich (1984) as Quasistrobus ramiflorus Vladimirovich. On the other hand, U. morkvaschica is similar to dicranophyllalean shoots of the Mostotchkia type (Meyen and Smoller, 1986). Another species, Voltzia mamadyschensis, is even more similar to Mostotchkia. Pityophyllum permiensis Zalessky includes one fragment of a linear leaf with midrib. Its affinity is uncertain and the generic name Pityophyllum is hardly justified. Several species, mostly new ones, were described by Zalessky (1937, 1939) from the Kungurian stage of the middle Fore-Urals. These are Walchia uralica Zalessky, W. appressa Zalessky, W. bardaeamt Zalessky, ~ peremiana Zalessky, Ullmannia bronnii Goeppert, U. bardaeana Zalessky, Bardella splendida Zalessky, Ammatopsis mira Zalessky (Zalessky, 1937), W. appressa, W. densa Zalessky, Brachyphyllum primordiale Zalessky, Voltzia principalis Zalessky, V. prisca Zalessky, Taxodiella recticaulis Zalessky, Uralodendron verticillatum Zalessky, Biarmodendron Joliosum Zalessky (Zalessky, 1939). Some of the species are analyzed in Sections 5.1.1, 5.2.1 and 5.10. Bardella splendida, Ammatopsis mira and Biarmodendron ./oliosum may belong to dicranophylls of the Mostotchkia type. The coniferalean affinity of ~'i)ltzia principalis and V. prisca is even more doubtful. Similar specimens are present in the author's collection. Although they are externally similar to conifers with spatulate or linear leaves, they have been found in organic connection with Peltaspermum-like fructifications. Probably these plants belong to a group ancestral to Cardiolepidaceae. Some of Zalessky's taxa were briefly mentioned by Florin (1940) who has refrained from comments on their systematic position.

S. K Meyen / Review of Palaeobotany and Palynology 96 (1997) 351 447

In the literature of the 1930s and of subsequent decades various conifers are listed among other plant remains coming from different Permian stages of the Russian Platform and Fore-Urals. These are the above mentioned species Ullmannia

biarmica, U. bronnii, Walchia hypnoides, W. foliosa, W. appressa, sometimes W. piniformis, etc. (Moskaleva, 1940; Tikhvinskaya, 1946; Nalivkin, 1949; Tefanova, 1963, 1971; Radczenko, 1966; Radczenko et al., 1973; Zoricheva and Sedova, 1954; Meyen, 1971a; etc.). In the 1970s the author began special, more profound studies of the conifers of Western Angaraland with particular attention to epidermalcuticular characters and fructifications. Presently, only data on conifers of the Tatarian stage have been published, namely on the leafy shoots Quadrocladus and associating female (Sashinia) and male (Dvinostrobus) fructifications, and pollen of the Scutasporites type (Meyen, 1976-1978, 1981, 1982a, 1984; Gomankov and Meyen, 1986). Initially, the author (Meyen, 1976-1978) erroneously referred to the conifers the family Cardiolepidaceae represented by the genera Phylladoderma (leaves), Cardiolepis (female capsules), Permotheca (synangia), Nucicarpus (seeds) and Vesicaspora (pollen). Later on, the affinity of the family with the order Peltaspermales was revealed (Meyen, 1981, 1982a, 1984). The genus Slivkovia previously affiliated with conifers (Meyen, 1976-1978) has been referred to the order Dicranophyllales together with some other plants externally resembling conifers (Meyen and Smoller, 1986). The studies of fructifications and epidermalcuticular characters clearly show that taxa established on gross morphological characters of vegetative shoots alone can be envisaged as merely collective form groups. Therefore, a revision of previously established genera and species is mostly impossible.

3. Stratigraphie setting of the material The conifers described below come from the Kungurian, Ufimian (?), Kazanian and Tatarian stages of the Russian Platform and Fore-Urals.

355

Although the stratotype sections of these stages are located in the same area, the localities of the conifers do not belong to such sections. Nevertheless, stratigraphic knowledge of the area is full enough for reliable assignment of most of the localities to certain stages and substages. The dating of the locality Shapkina in the Timan-Pechora region (northeastern part of Pechora Fore-Urals) is less certain. The correlation between the enumerated standard stages and the West European Permian units remains unresolved (see details in Kozur, 1980a,b). In any case it seems evident that the Kungurian stage is younger than those portions of the West European Autunian and Rotliegend to which localities yielding the conifers Lebachia, Ernestiodendron and Walchiostrobus are confined. There is wide divergence of opinion among stratigraphers as to the correlation of West European Zechstein yielding Pseudovoltzia, Ullmannia and Quadrocladus with the standard Permian stages. Different stratigraphers correlated Zechstein with the Kungurian, Ufimian, Kazanian or Tatarian stages. With conodonts, the correlation with the uppermost Kazanian-Tatarian appears more probable (Kozur, 1980b). Palaeomagnetic observations suggest, however, that the Tatarian stage may correspond to lower portions of the West European section beginning with the uppermost Rotliegend (Menning, 1981 ). One cannot exclude that a part of the Zechstein may correspond to a large regional hiatus between the Tatarian and overlying Triassic beds (on the range of this hiatus see: Gomankov, 1983; Gomankov and Meyen, 1986). The position of the Val Gardena Formation (Alps, Italy) yielding Ortiseia (Clement-Westerhof, 1984) in the succession of the standard stages is also unclear. In the succession of the Tethyan stages, Val Gardena roughly corresponds to the Dzhulfian or Abadehian (Midian) (ClementWesterhof, 1984). By palaeomagnetic and palynological data the Tatarian stage also corresponds to these Tethyan stages (Gomankov, 1983). On the other hand, by palynological data the beds with Ortiseia are correlatable with the West European Zechstein (Clement-Westerhof, 1984). Thus, the Val Gardena Formation undoubtedly belongs to the Upper Permian, probably to its upper part.

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S. V. Meyen / Review (?/Pa&eohotany and Palynoh)gy 96 (1997) 351 447

The oldest known coniferalean fructifications Lebachia h)ckardii Mapes et Rothwell -- were described from the Topeka Limestone Formation of Kansas (Mapes and Rothwell, 1984). The Formation belongs to the Shawnee Group, i.e. the middle part of the Virgilian (Ebanks et al., 1979). Mapes and Rothwell admit a wider age interval for the plant-bearing beds including lowermost Permian, because the conifers are accompanied by plants usually regarded as Permian, e.g.. Dichophyllum moorei Elias sometimes affiliating with Callipteris,[tabellifera Weiss. However, assemblages of miospore and plant megafossils typically Permian in habit have proved to appear in separate layers as low as the middle Stephanian (Doubinger and Langiaux, 1982). Such assemblages characterize in the Stephanian of France, barren members of the section. In non-coal-bearing regions the miospore assemblages of Permian habit range throughout the Upper Carboniferous and enter the upper Middle Carboniferous. This has been observed in sections along the eastern slope of the Urals (Chuvashov et al., 1984) and Dzhungaria. The Shawnee Group of Kansas belongs to deposits of the same type. Therefore, the findings of plants having a Permian habit in this group do not contradict the Late Carboniferous age of the embedding rocks.

3.1. Localities (?]the Kungurian stage The Kungurian conifers come from the central and southern parts of the Sylva Depression. According to the current stratigraphic schemes, the Kungurian of this region is subdivided into the Lekskaya (below) and Koshelevskaya suites belonging, respectively, to the Filippovsky and lrensky horizons ( Unified and correlational stratigraphic schemes of Urals, 1980). These are presumably terrigenous deposits with interbeds o1" limestones, salt and anhydrite. The conifers were collected in localities along the rivers Barda ( localities Krutaya Katushka, Krasnaya Glinka, Matveevo-1 near the village of Matveevo) and Sylva (localities Chekarda-I and Chekarda-2). The localities along the Barda River occur in the center of the Sylva Depression. Chuvashov and Dyupina (1973) provide descriptions of the

pertinent section. The deposits recognized in the unified scheme as the Lekskaya suite, are referred by them to the Krylovskaya suite consisting of three members. The lower member (220 250 m thick) is dominated by yellowish-grey sandstones with partings of sandy limestones. The middle member (50 60 m thick) is composed by dark argillites and marls. The upper member (50 m thick) consists of light-grey and yellowish-grey microlaminated marls. The conifers come from the upper part of the upper member, i.e. from the top of the Lekskaya (Krylovskaya) suite. The overlying Koshelevskaya suite consists of alternating greenish- and yellowish-grey sandstones, siltstones and claystones. The miospore assemblage of the Krylovskaya suite is strikingly dominated by gymnosperm pollen (striated quasisaccates, 27.3%; Vittatina, 19.8%: quasimonosaccates, 10.6%; Azonaletes, 9.1%; Ginkgocycadophytus, 8.1%; data by Dyupina). The section along Barda is correlated with more western type sections of the Filippovsky horizon by tracing members according to lithological features as well as by miospore assemblages (see details in: Porfiriev, 1963; Chuvashov and Dyupina, 1973: Oborin and Khursik, 1973; Zolotova et al., 1973; Dyupina, 1984). Locality Krutaya Katushka. Left bank of the Barda River 2 km above the bridge in the village of Matveevo. Conifers were collected in the lower part of the outcrop (Fig. 2) together with Equisetina magnivaginata Zalessky, Spheno-

phyllum, Paracalamites, Phyllotheca, Callipteris, Mauerites, Biarrnopteris pulchra Zalessky, Sylvia striata Zalessky, Psygmophyllum, Cordaites, Rufloria (with narrow dorsal furrows), Entsovia kungurica Meyen, seeds. Locality Krasknaya Glinka. Right bank of the Barda River 1.3 km above the bride in the village of Matveevo. Conifers were collected in two intervals of the section (Fig. 2) together with

Psygrnophyllum, Mauerites, Callipteris, Asterodiscus, Cordaites, Rufloria (with narrow dorsal furrows), Entsovia, seeds. Layer 1 yielded specimens Nos. 3773/473 494, layer 6 specimens Nos. 3773/498-519. The localities along the Sylva River occur near

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S. I~ Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447 Shapkina BB

m

Sluda

BB

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Fig. 2. Stratigraphic position of beds with conifers in some localities. 1 = sandstones; 2 = argillites; 3 = aleurolites; 4 = pebbles; 5 = marls; 6 = limestones; 7---findingsof conifers. the mouth of the Chekarda River, i.e., in the south of the Sylva Depression. They are confined to the Koshelevskaya suite (Zalessky and Chirkova, 1940; Nalivkin, 1949; Porfiriev, 1963). Deposits presently included in the Koshelevskaya suite, had been referred by Kruglov (1933) to the Tisovskaya (its major upper part) and Suksurskaya suites. The plant-bearing beds were placed into the Tisovskaya suite (Zalessky, 1956, 1957). The Koshelevskaya suite coincides in its stratigraphic range with the Irensky horizon, in the type section of which located on the Ufimian Plateau several members are recognized (in ascending order: Ledyanopeshcherskaya, Nevolinskaya, Shalashninskaya, Elkinskaya, Demidkovskaya, Tyuiskaya, Lunezhskaya). The part of the Koshelevskaya suite

yielding numerous plant remains (Chekardinskaya member of Chuvashov) approximately corresponds to the Elkinskaya member (Dyupina, 1984). Gorsky (1970) correlated the plant-bearing part of the Koshelevskaya suite with the lower part of the Ufimian stage. Palynological studies have shown (Chuvashov and Dyupina, 1973) that Gorsky's correlation is valid only for the section along the Ufa River. As to the sections of the Koshelevskaya suite along the Sylva River, their correlation, even partial, with the Solikamsky horizon of the Ufimian stage has not been admitted (Unified and correlational stratigraphic schemes of Urals, 1980). Locality Chekarda-1. Left bank of the Sylva River immediately below the mouth of the Chekarda

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Review (~/Palaeohotany and Palvno/ogy 96 (1997) 351 447

River. The description of the outcrop yielding plant remains has been published by Permyakov (1938, pp. 70-71). By comparison with this description, the column in Fig. 2 includes the following corrections: ( 1) layers 6 9 (total 0.85 m thick) are united because they are not sufficiently persistent along strike; (2) thicknesses of layers have been defined more accurately; (3) sandstones are shown in the basal part of layer 14; (4) breccialike rock is shown in the basal part of layer 13. Layers 6-9 yielded specimens Nos. 3773/626 648 and the entire collection 3737 (collected by A.G. Sharov, Palaeontological Institute of the USSR Academy of Sciences), and layer 14 specimens Nos. 3773/554-625; specimens Nos. 3773/649 660 come from the talus. Locality Chekarda-2. Left bank of the Sylva River immediately above the mouth of the Chekarda River. The description of the outcrop has been published by Permyakov (1938, p. 70). In 1965 the outcrop proved to be completely covered by talus; its column has been compiled (Fig. 2) according to Permyakov's description. Layers 2 3 yielded specimens Nos. 3773/661-694, and the talus specimen No. 3773/695. Both localities yield rich floras including (apart from conifers) Sphenophyllum, Phyllotheca,

Paracalamites, Annularia, Pecopteris, Callipteris, Peltaspermum, Asterodiscus, Psygmophyllum, Mauerites, Biarmopteris pulchra, Cordaites. Rufloria (with narrow dorsal furrows), Entsovia, Samaropsis triquetra Zalessky and other seeds, Pholidophyllum ornatum Zalessky, etc. 3.2. Locali O, (~['the Ufimian (?)stage Only one locality, Shapkina, has been referred to the Ufimian stage with some doubts. Borehole Shapkina-1 has been drilled on the left bank of the Pechora River 120 km I¥om its mouth ( 70 km to the east-southeast from the city of Narian-Mar) in the axial part of Denisovian Depression of the Pechora Syneclise. Yield of core was very limited ( Fig. 2) and a continuous description of the section is impossible. Conifers were found at a depth of 1508.2 1511.1 m in dark-grey siltstones belonging to the middle part of the Ekushanskaya suite. The

age of the suite remains debatable ( Entsova et al., 1969; Entsova, 1970, 1972; Telnova, 1974; Varyukhina et al., 1981 ). The suite has been initially established on the borehole material in the lower stream of the Pechora River. Its stratotype (70 m thick) is located in the borehole NarianMar-l. In this borehole as well as in other boreholes drilled in lower stream of the Pechora River, the Ekushanskaya suite is composed of grey calciferous sandstones with gravel interbeds in the lower part. Marine fauna occurring in the suite (Komiella rotundata Toula, K. sulaensis V. Barchatova, Svalbardia capitolina Toula, Cancrinella ex gr. cancrini Verneuil, Anidanthus kuliki Fredericks, Muirwoodia mammata Keyserling, Healdia ivanovi Martius, etc) is variously treated as Kungurian, Ufimian or Kazanian. Plant remains have been found in several boreholes. These are Phylladoderma, Rufloria ex gr. recta (Neuburg) Meyen, Mostotchkia gomankovii Meyen et Smoller, Psygmophyllum sp., etc. In the stratotype sections of the Permian stages the genus Phylladoderma appears near the boundary between the Solikamsky and Sheshminsky horizons of the Ufimian stage. Therefore, the correlation of the plant-bearing part of the Ekushanskaya suite with the Kungurian or Solikamsky horizon is hardly possible. Mostotchkia gomankovii was recorded in the lower Kazanian (see below). However, the conifers found in the suite differ from Kazanian ones and, as will be shown below, are more primitive. Therefore the Sheshminsky age of the Ekushanskaya flora appears most probable. This age assignment agrees well with other plant megafossils. The combination of Phylladoderma with R. ex gr. recta is characteristic for that (lower) part of the Pechora series (Pechora basin), the Sheshminsky age of which is widely adopted, The upper subsuite of the overlying Telvisskaya suite yielded a Kazanian ostracode assemblage with an admixture of Ufimian taxa (Kashevarova, 1973). The assemblage is comparable to those from the Kazanian of Timan. The admixture of Ufimian taxa is regarded as evidence for the early Kazanian age of the assemblage. The miospore assemblage of the Ekushanskaya suite is clearly different from those of underlying and overlying beds. According to palynological

S. V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

data the basal part of the suite corresponds to the boundary beds between the Vorkutskaya and Pechorskaya series of the Pechora basin (Telnova, 1974). The assemblage of miospores from the suite is subdivided into two subassemblages, conifers of Shapkina being confined to the lower part of interval yielding the upper subassemblage. Telnova (1974) correlates this part of the section with the Sheshminsky horizon of the Solikamsk region.

3.3. Localities of the Kazanian stage Western Fore-Timan Locality Vym-1. Right bank of the Vym River 4 km below the mouth of the Koin River, lower Kazanian substage. Specimens collected by V.I. Rosanov in 1968, I.S. Muraviev and G.A. Ioffe in 1969, M.G. Minikh in 1971 and A.V. Gomankov in 1979. Locality Vym-2. the same bank 1.7 km below the mouth of the Koin River, lower Kazanian substage. Specimens collected by M.G. Minikh in 1971. The stratigraphic setting of these localities is given in the literature (Plotnikov and Molin, 1969; Molin and Koloda, 1972; Muraviev et al., 1975; Varyukhina et al., 1981). The Kazanian deposits are divided into the Cheviyusskaya and Veslyanskaya suites (beds) that approximately corresponds to the lower and upper Kazanian substages. The lower Kazanian consists of three rhythms, conifers being confined to the lower one. Muraviev and coauthors provide the following characteristics of the lower rhythm (its thickness is 30-35 m). It commences with limestones (up to 0.4 m) in places grading into conglomerate. Overlying yellowish-grey sandstones (20 m) contain one-two clay partings. There are many brachiopods, bivalves and foraminifers (including Nodosaria suchonensis Mikloucho-Maclay and N. cf. pseudoconcinna Mikloucho-Maclay). Upwards occurs a grey clayey-carbonate member (up to 15 m). Its lower part is represented by argillites and aleurolites with thin interbeds of sandstone and marl lenses. Upwards lies an alternation of argillites (with fresh water ostracodes Darwinula aft. persimplex Kotschetkova and Placidea sp.), aleu-

359

rolites and sandstones. The member is crowned by brownish-grey aleurolites with three partings of clayey limestones. The lower parting yields many marine bivalves, less frequent gastropods, and ostracodes. The middle parting corresponds to the locality Vym-1. The locality Vym-2 appears to occupy the same stratigraphic position. The middle rhythm (25-30 m) is a terrigenouscarbonate with marine fauna (Nodosaria suchonensis Mikloucho-Maclay, Geinitzina cf. spandeli Tscherdynzov, Monoceratina cf. parvula Kotschetkova, Cavellina grandis Schneider, Actuaria secunda Kotschetkova, brachiopods, bivalves, bryozoans). The upper rhythm (20-25 m) is mostly terrigenous in its lower part (sandstones, argillites), and carbonate upwards, still higher it is composed by clays and marls. The upper Kazanian is mostly carbonate (up to 100 m). The flora in the localities Vym-1 and Vym-2, as well as in other localities of the same portion of the section, includes Timanostrobus muravievii Meyen, sp. nov., Concholepis harrisii Meyen, sp. nov., Mostotchkia gomankovii, Rhaphidopteris sp., "Odontopteris" rossica Zalessky, Wattia sp., Samaropsis sp., Signacularia sp., etc. The type material of Rhaphidopteris praecursoria Meyen (1979) is roughly of the same age. Locality Sluda. The village of Sluda 60 km northwards from Syktyvkar City, borehole CC-104. Specimens collected by F.I. Entsova in 1960. The plant-bearing beds belong to the lower Kazanian Cheviyusskaya suite briefly characterized above. The plant-bearing interval of the borehole (Fig. 2) has been placed by Molin and Koloda (1972) into the Upper Cheviyusskaya subsuite corresponding to the two upper rhythms of the earlier described section along the Vym River. The plant-bearing interval also yielded marine fauna (preliminary identifications by K.F. Sedykh made in 1960): depth 415.1 m - Aulosteges sp., Camarophoria sp., Rhynchopora geinitziana Verneuil; depth 414.3 m Aulosteges horrescens Verneuil, Productidae; depth 415.5 m - Athyris pectinifera Sowerby, Spirifer sp.; depth 491.1 m - Productus hemisphaericum Kutozga. Plant remains belong to Quadrocladus (al. Steirophyllum) komiensis Meyen, sp. nov., Paracalamites, Phylladoderma, Entsovia rarisulcata

360

S. [~ Meven, Rcvicli ~?! Palaeohotany and Pa(vnolo~o' 96 (1997) 351 447

Meyen, Rhaphidopteris praeeursoria, Cordaites, seeds.

Locality Kityak (Akbatyrevo or Preobrazhenskv Mine). Left bank of the Kityak River (= Kityachka; left affluent of the Burets River which is right affluent of the Vyatka River) across the village of Bolshoi Kityak (Akbatyrevo), Malmyzh District, Kirov Area: specimens collected by the author (collection 3773) and A.G. Sharov (collection 1438). The locality has become known owing to findings of tetrapods, and has been described in the catalogue by Efremov and Viyushkov ( 1955 ). The locality belongs to the Belebeyevskaya suite, i.e. the Upper Permian deposits facially replacing marine deposits of the Kazanian stage in an eastward direction. The position of the boundaries both between the lower and upper Kazanian, as well as the Kazanian and Tatarian in these deposits is not quite clear. Therefore, the position of the locality Kityak in the general section of the Upper Permian is debatable. The locality occurs exactly in the zone of the facial transition from the marine Kazanian deposits to continental ones, where the lower Kazanian is still represented by marine beds belonging to the subzone b 2 of Forsh ( 1955, fig. 4), i.e. alternation of grey clays, sandstones, marls and limestones with a high clay content. The locality Kityak occurs in overlying continental deposits of the subzone v~ of Forsh (1955, fig. 5 ). Tikhvinskaya ( 1946, p. 256) refers the ore-bearing (cuprous) beds with tetrapod remains to the lower part of the Belebeyevskaya suite. Among tetrapods found in cuprous sandstones of the locality the amphibians Pla(vops stuckenbergii Trautschold and Melosaurus, and reptiles (Deinocephalia) Syodon and Moh'bdopygus areanus Tchudinov have been identified. This tetrapod assemblage belongs to the Deinocephalian Zone II of Efremov referred by Chudinov ( 1983 ) to the lower Tatarian. Accordingly, he refers the locality Kityak (Akbatyrevo Mine) to the Lower Tatarian along with all the localities of the same Zone. He refers to the paper of Tikhvinskaya, Chepikov and others where the Tatarian age of localities of the Zone had been established. However, in these papers the Tatarian age of all the localities of Zone II had not been maintained.

Chepikov (1946) noted that the localities Akbatyrevo and Kamskie Polyany belong to the Belebeyevskaya suite. Rachitsky (1969) held that the cuprous sandstones yielding the Tatarian tetrapods are locally entrenched into the Belebeyevskaya suite and can be mistaken for the upper Kazanian: however, he left the stratigraphical position of tetrapods from the Akbatyrevo Mine as an open question. Plant remains occurring in the locality Kityak may help in the resolution of this question. The locality yields numerous Sphenophyllum stouckenbergii Schmalhausen and Pseudovoltzia? eornuta Meyen, ,s)v. nov. along with Pecopteris helenaeana Zalessky, Danaeites, Lobatannularia, Paracalumites, "Odontopteris" rossica, Rhaphidopteris,

Phylladoderma, Perrnotheca, Nucicarpus, Psygmophyllum, seeds. Similar assemblages are known in the Kazanian of the east part of the Russian Platform (Tefanova, 1963, 1971; Meyen, 1971a; Esaulova, 1984; Varyukhina et al., 1981; Vladimirovich, 1984). The association of the large number of Sphenophyllum stouckenbergii and Phylladoderma was found in the locality KzylBairak (right bank of the Volga River south of Kazan City) in the so-called "Transitional Member", i.e. in the uppermost Kazanian. In undoubtedly Tatarian deposits of Western Angaraland, sphenophylls have not yet been recorded. Conifers found in Kzyl-Bairak belong to Steirophyllum lanceolatum, i.e. species characteristic of the lower Tatarian deposits of the Kargala Mines (see below). The conifers in Kityak are different and of older habit. Therefore, considering the similarity of the Kityak flora with other upper Kazanian floras, the locality Kityak can be placed into the upper Kazanian substage below the level of the "Transitional Member". Accordingly, the Kazanian age of the lower part of Zone II can be admitted.

3.4. Localities ~fthe Tatarian stage Conifers come from eight localities of which one (Kargala Mines) belongs to the lower Tatarian, one (Isady) to the Severodvinsky horizon of the upper Tatarian, and the remainder (Aristovo, Viled, Titovo, Dor, Luptyug, Kalinovka) to the middle part of the Vyatsky horizon of the upper

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Tatarian (Fig. 3). The stratigraphic setting and faunal assemblages of the upper Tatarian localities are described in more detail elsewhere (Gomankov and Meyen, 1986).

Locality Kargala Mines (Kuzminovsky Mine). A few available specimens from the Kargala Mines have very poor geographic and stratigraphic control. The accompanying labels say the following: "Kargala cuprous mines. Kuzminovsky pile. Orenburg Area, Belozersk District. Coll. by Bocharov 1946". According to Efremov and Viyushkov (1955, p. 56), the Kuzminovsky Mine is located "on a high watershed 9 km N from the settlement of Gorny on the Verkhnyaya Kargalka River, 4 k m to NE from the Myasnikovsky (Vshivy) Farm and 7 km SE from the Vlasievsky Mine. The bone-bearing lower marl member is penetrated by three big mines occurring in the centre of the mine near the watershed road. The section has not been described. Remains of land Sukhona and Malaya Severnaya Dvina Rivers Suites after Lyutkevich, 1935

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vertebrates occur in a lens of ore-bearing grey compact marl 0.6 m thick and in overlying cuprous slaty grey, faintly sandy marls together with abundant remains of plants, fish, estherias, anthracosia and insects." The authors provide a list of publications dealing with the Kargala Mines. The stratigraphic position of the plant-bearing beds was treated repeatedly in the literature. Tikhvinskaya (1952) carefully analyzed the issue and referred the beds to the lowermost Tatarian. The same conclusion has been drawn by other workers (Chepikov, 1 9 4 6 , 1 9 4 8 ; Rachitsky, 1969; Chudinov, 1983; etc.). The available specimens of Steirophyllum lanceolatum are indistinguishable in gross morphology from those found in Kzyl-Bairak (see above) and belonging to the "Transitional Member" (uppermost Kazanian). Owing to the absence of accurate stratigraphic information for the hand specimens, their age cannot be treated in more detail. Vyatka River Probable ranges

Subdivision after Pa

of localities

Beds

Kornaritskie Ustyuc~skie

I

o

~-~

I I

I I "~

Gremyachevskie

Severodvinskaya

Klimovskie

Mutovinskie Opokskie Nyuksenitskie $ukhonskaya

Dmitrievskie Verkhnetozmenskie Shardinskie

Nizhneust inskaya

Karpogorskie Mar iegorskie

=n Stage

Fig. 3. Subdivision and correlation of sections along Sukhona, Malaya Severnaya Dvina and Vyatka Rivers, and stratigraphic position of localities of Tatarina flora. 1 = conglomerates; 2 = sands and sandstones; 3 = aleurolites; 4 = clays; 5 = marls; 6 = limestones; 7 = gypsiferous rocks; 8 = direct magnetic polarity; 9 = reversed magnetic polarity.

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Locality Isady. Vologda Area, left bank of the Sukhona River about 2 km below mouth of the Mutovka River and 300 m above the village of" Isady. Specimens collected by A.G. Olferiev in 1969, M.G. Minikh in 1971, A.V. Gomankov and the author in 1978. The conifers were collected in the middle part of a large lens composed of grey aleurolites and sandstones and embedded in red argillites and marls. The lower part of the plantbearing interval yielded coniferalean remains (Geinitzia sp. SVM-2) devoid of cuticles and without accompanying fructifications. They are associated with the fern Fefilopteris having Taeniopterislike leaves, and numerous Pursongia. The overlying layer contains mass accumulations of plant remains, mostly of Tatarina and Phylladoderma subgen. Aequistomia. There are Protosphagnum nervatum Neuburg, Fasciostomia delicata Gomankov (leaves of lycopods), stems of articulates, fragments of compressions of F~'l~'lopteris pilosa Gomankov, as well as Peltaspermopsis sp., Tatarina cf. conspicua Meyen, Salpingocarpus bicornutus Meyen, S. variabilis Meyen, Phylladoderma (Aequistomia) annulata Meyen, P. (A.) rastorguevii Meyen, P. (A.) trichophora Meyen, Rhaphidopteris sp., Permotheca striat([bra Meyen et Gomankov, P. vesicasporoides Meyen, Esaulova et Gomankov, Pursongia beloussovae (Radcz.) Gomenko, Dvinostrobus sagittalis Gomankov et Meyen, Quadrocladus schweitzeri Meyen, Arisada densa Meyen, insect wings. Locality Aristovo. Vologda Area, right bank of the Malaya Severnaya Dvina River 100 m above pier near the village of Aristovo. Specimens collected by A.G. Olferiev in 1969, M.G. Minikh in 1971, A.V. Gomankov, I.A. Ignatiev and the author in 1976. The description of the locality is given by Efremov and Viyushkov (1955). The conifers were collected in a small lens of calciferous aleurolites and argillites embedded in a bigger lens of yellowish-green sands that, in their turn, occur within red clays and marls. Dominating are leaves of the Tatarina and Phylladoderma subgen. Aequistomia, there are numerous fructifications of Peltaspermopsis and Permotheca. The total list includes: Pelliothallites tataricus Meyen (liverworths), Protosphagnum nervatum, Rhizinigerites neuburgae

Meyen (and other mosses), Fasciostomia delicata, stems of articulates, Fefilopteris pilosa, Pecopteris sp., Peltaspermopsis buevichia (Gomankov et Meyen) Gomankov, Tatarina conspicua, T. pinnata Meyen et Gomankov, Salpingocarpus bicornutus,

S. variabilis, Phylladoderma (Aequistomia) aequalis Meyen, P. (A.) annulata, Permotheca striat(['era, P. vesicasporoides, P. vittatinifera Meyen et Gomankov, Pursongia beloussovae, Sashinia aristovensis Meyen, Dvinostrobus sagittalis, Quadrocladus dvinensis Meyen, Arisada densa. There are also insects, ostracodes, conchostracs, bivalves, charophytes and vertebrates. Locality Viled. Arkhangelsk Area, borehole 06 in the village of Pavlovskoe near bridge across the Viled River, depth 139 154.5 m. Specimens collected by M.A. Stepanenko in 1965. Plant remains occur in calciferous light-grey aleurolites, dominating are Protosphagnum, Quadrocladus, Tatarina, seeds. The plant assemblage includes: Proto,sphagnum nervatum, Rhizinigerites neuburgae (and other mosses), Fasciostomia delicata, Pelta-

spermopsis buevichia, Salpingocarpus bicornutus, S. variabilis, Phylladoderma (Aequistomia) annulata, Sashinia borealis Meyen, Dvinostrobus sagittalis, Quadrocladus borealis Meyen, Geinitzia sp. SVM- 1, Pseudovoltzia? sp. Abundant ostracodes belong to Darwinula parallela (Spizharskyi), D. inornata Spizharskyi, D. inornata var. macra Lunjak, D. digitalis Mischina, D. daedala Mischina, Darwinuloides tataricus (Posnez), D. svijazhicus (Scharapova), Suchonella cyrta Zekina, Sinusuella ~jatkiensis Posnez, etc. (identifications by A.E. Kalis).

Locality Titovo. Vologda Area, left bank of the Engit River (affluent of Yuza River) 1.5 km north of the village of Titovo, borehole 4 Titovo, depth 138 144.4 m. Specimens collected by A.L. Buslovich in 1967. Plant remains overfill grey sandstones and aleurolites, dominating are Tatarina and Quadrocladus, less frequent are Phylladoderma subgen. Aequistomia, Glossophyllum and Rhaphidopteris. The plant assemblage includes: Tatarina conspicua, Stiphorus biseriatus Meyen, GIossophyllum permiense Meyen, Salpingocarpus bicornutus, S. cf. variabilis, Phylladoderma (Aequistomia) tatarica Meyen, Doliostomia krassi-

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lovii Meyen, Amphorisperum sp., Rhaphidopteris kiuntzeliae Meyen, Permotheca striatifera, P. vesicasporoides, Pursongia beloussovae, Sphenarion? sp., Sashinia aristovensis, Quadrocladus dvinensis. Ostracodes are represented by the "Suchonella typica, S. cornuta" assemblage: S. typica Spizharskyi, S. cornuta Spizharskyi, S. stelmachovi Spizharskyi (dominant), Darwinula inornata, D. parallela, Sinusuella vjatkensis, Tatariella stricta Mischina, Volganella kashevarovi Belousova, Placidea lutkevichi (Spizharskyi), etc. Locality Dor. Kostroma Area, Vokhma District, the village of Dor, borehole 514 Dor at the northwest outskirts of the village, depth 157.5-158 m. Specimens collected by A.G. Olferiev in 1964. Tatarina dominates. The locality yielded T. olferievii Meyen, Pursongia beloussovae, Quadrocladus dvinensis, and ostracodes of the same assemblage as in Titovo, namely Darwinula inornata, D. parallela, Suchonella stelmachovi, S. typica, S. cornuta, S. circularis Mischina, etc. Locality Luptyug. Kirov Area, right bank of the Vetluga River 26 km above mouth of the Vokhma River, borehole 529 Luptyug at the northwest outskirts of the village of Luptyug, depth 108.1-113.7 m. Specimens collected by A.G. Olferiev in 1965. Dominating are Tatarina and Protosphagnum. The flora includes: P. nervatum, Peltasperrnopsis sp., Tatarina olferievii, T. conspicua, Pursongia beloussovae, Geinitzia subangarica Meyen. Ostracodes of the same assemblage as in Titovo and Dor, namely Darwinula inornata, D. parallela, Suchonella stelmachovi, S. typica, S. cornuta, Volganella vetlugensis Belousova, V. magna Spizharskyi, Placidea volgensis Belousova, Unzhiella gemella Mischina, Darwinuloides tataricus, Sinusuella ignota Spizharskyi, etc. Locality Kalinovka. Kostroma Area, Pyshug District, the village of Kalinovka, borehole 361 Kalinovka, depth 153.5-160 m. Specimens collected by V.R. Lozovsky. Dominating are Tatarina and Rhaphidopteris. The flora includes: T. conspicua, R. antiqua Meyen, R. sp., Pursongia beloussovae, Sphenarion? sp., Dvinostrobus sagittalis. The stratigraphic position of the listed upper Tatarian localities is shown in Fig. 3. Thus, the distribution of the Permian conifers

363

of Western Angaraland and of better studied conifers of the Equatorial phytochoria acquires the pattern shown in Fig. 1.

4. Terminology Female fructifications are described below in terms explained in detail elsewhere (Meyen, 1982b, 1984) and presently entering the literature (Anderson and Anderson, 1985; Doludenko and Kostina, 1985). Seed-bearing organs of gymnosperms are termed "polysperms". In conifers and cordaitanthaleans simple axillary polysperms are arranged into a racemose compound polysperm (commonly called "cone" or "megastrobilus"). The seed and its stalk are together termed a "monosperm" (="Samenschuppen" of Florin, 1938, 1939, 1940, 1944, 1945)or "megasporophyll" of Florin, 1951 ). Sterile scales accompanying a polysperm form "circasperm". This terminology permits clear definition of the most important questions relating to coniferalean fructifications. For instance, Florin treated the seed scale as a modified circasperm to which monosperms were secondarily adnate. Schweitzer (1963) and Gurdrs and Dupuy (1974) regarded the seed scale as modified leaves bearing monosperms with reduced stalks. These views will be treated in detail in Section 6. In Kungurodendron and, probably, other Palaeozoic conifers, the female axillary complex includes, in addition to sterile scales inherited from cordaitanthaleans and seed stalks, sterile appendages resembling seed stalks in outward appearance and some epidermal characters, but having either underdeveloped seed scars or no traces of the scars at all. Such appendages can be termed "sterilized seed stalks" or "funiculodia" (by analogy with staminodia of flowering plants). The pollen produced by some Palaeozoic conifers can be called "protosaccate" (following Scheuring, 1974; see also Foster, 1979). This term, however, suggests that such a pollen structure preceded the saccate one, but this is not always so (Meyen, 1984). Therefore, the term "quasisaccate" is proposed instead, as well as "quasisaccus" for "protosaccus". Quasisaccate pollen simulating

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monosaccate and disaccate can be called, respectively, quasimonosaccate and quasidisaccate. A characteristic feature of a number of Palaeozoic conifers, e.g. Walchia (al. Lehachia), Ernestiodendron, Kungurodemtron, Swillingtonia, "Lebachia" lockardii, and Ortiseia, is the presence of small epidermal cells with disproportionately large papillae or unicellular hairs. Such cells are often described as hair bases (trichopodia), although they do not always bear hairs. For these specialized papillose cells a new term "papilliger'" is proposed (from Latin "gero" - bearing). This term can also be applied to such peculiar cells in those cases when we do not know whether they bore papillae or unicellular hairs. As in another work (Gomankov and Meyen, 1986), a special mode of designations is used below for species devoid of specific epithets and accompanied by "sp.". References to descriptions relating to taxa with such "sp." are complicated and inconvenient. To simplify the references the word "'sp.'" may be accompanied by initials of author and number. Numbers belonging to the given author must not be repeated in his or her works. In the above cited paper of Gomankov and Meyen, the species Geinitzia sp. SVM-1 (i.e., belonging to S.V. Meyen) has been described. Below, another type of Geinitzia sp. is described. Accordingly, it obtains the designation "sp. SVM-2". Naturally, such designations must not be regulated by the nomenclatural code and are not guarded by priority. This is but a small addition to the practice of references and citing.

5. Systematic description

leafy twig and bearing helically arranged bracts differing from foliage leaves in large size. Axillary complex dorsiventrally flattened, its basal sterile scales arranged along lax spiral. Upper part of axillary complex occupied by adaxially placed decurrent hooked seed stalks, and apical and abaxial straight funiculodia similar to seed stalks in their epidermal structure and occasionally bearing aborted seed scars. Microstrobili elliptical, with numerous helically arranged crowded microsporophylls. Pollen globose with thin quasisaccus composed by columellar-like sexine and embracing body equatorially and distally. Etymology: From the Kungurian stage and dendron (Greek), tree.Nomenclatural notes: The new genus is introduced for the entire set of organs vegetative shoots of two orders of branching, polysperms, microstrobili and pollen. However, the nomenclatural type of the type species K. sharovii Meyen, .sp. nov. is represented by a polysperm. Detached twigs may also be referred to the genera Walchia sensu Florin (1938) or Cvparissidium sensu Harris (I 969, 1979); see also descriptions of C. appressum and C. ents'ovae below). The correspondence of microstrobili to a certain form-genus (e.g., Masculos'trobus, Walchianthus, etc.) remains unknown. The associated pollen cannot be ascribed to a definite genus established for dispersed miospores. ()mTparison: The new genus differs from Walchia (al. Lebachia) and Ortiseia in having numerous seed stalks bearing seeds apically, much more reduced sterile scales of the circasperm, the presence of funiculodia, and pollen structure. The relations to other genera and the systematic affinity of the new genus will be treated in detail in Sections 6 8.

5.1. K U N G U R O D E N D R O N Meyen, gen. not' 7)'pc: Kungurondendron sharovii Meyen, sT). nov. Diagnosis': Vegetative leafy shoots of penultimate order bear pinnately arranged branches of last order. Leaves lanceolate, mucronate, with two stomatal bands on upper surface and few stomata on lower surface near leaf base. Margin with cuticular microdenticulation or unicellular hairs. Main axis of compound polysperm terminating

5.1. I. Kungurodendron sharovii Meyen sp. nov. (Plates I III; Figs. 4-6, 7 A G , 32B) Holoo,pe: Specimen No. 3737/259 (Plate II, 1, 2; Plate III, 27). 1982a Pseudovoltzia sp. Meyen, pl. 16, fig. 134. T]'pe locality: Chekarda-l. Etymology: After the late Dr. A.G. Sharov who

S. I~ Meyen / Review of Palaeobotany and Palynology 96 (1997) 351 447

collected a part of the type material including the holotype. Diagnosis: Branches of penultimate order up to 8 mm wide, their leaves 3.5 mm wide and 12 mm long. Branches of last order up to 80 mm long, their leaves 1-1.25 mm wide and 4-9 mm long, adpressed to axis or slightly diverging, lanceolate, with apical mucro. Stomatal bands fused apically and grade upwards into papillose band. Nonstomatal cells of stomatal band polygonal, variously orientated, more or less papillose, among them rare papilligers and trichopodia occur. Cells of axial and marginal stomata free bands as well as cells of lower epidermis longitudinally elongated arranged into irregular files. 6-7 (5-8) subsidiary cells with strongly cutinized radial and/or periclinal walls. Stomatal pit round-polygonal overhung by papillae of proximal walls of subsidiary cells. Lower leaf side bears below apex large trichopodium surrounded by files of shorter cells. Compound polysperms up to 70 mm long and up to 30 mm wide. Bracts up to 12 mm long and up to 3.5 mm wide. Axillary polysperms up to 7-8 mm long, their basal flattened part 3-4 mm wide. Sterile scales decurrent in this part of axillary polysperm, their upper surface very short, margin with same microdenticulation as in foliage leaves. Upper part of axillary complex bears adaxially 9 11 seed stalks with hooked backwards apices crowned by seed scars accompanied by cuticular fold. Seed stalk bears rare stomata and solitary short unicellular hairs. Apical and abaxial subapical sides of axillary complex bear 15-20 erect funiculodia occasionally having variously underdeveloped seed scars. Microstrobili about 3 cm long and 8 mm wide. Microsporophylls numerous, elbow-shaped, with distal lamina having same stomata as foliage leaves. Pollen globose or folded and then boatshaped. Columellae-like bodies of sexine arranged into polygonal network. Proximal side finely granulose, proximal slit small, mono-, di- or trilete. Description: The description is based on 32 specimens of sterile shoots from two localities (well preserved cuticle has been obtained from 8 specimens), and 8 fructifications (6 polysperms and 2 microstrobili) from the type locality. Vegetative

365

shoots are mostly represented by unbranched fragments (Fig. 4B, C, D, I). Probably they are either branches of the last order or apices of branches of the penultimate order. Lateral branches of pinnately branching shoots (Fig. 4A, G) vary in length, they are attached at acute angles and bear smaller leaves than branches of the penultimate order. Apices of branches obtuse. Leaves are either closely adpressed to axes (Fig. 4A, B, I) or slightly diverging (Fig. 4C, D, G in the lower part). The lower leaf side is slightly carinate. General epidermal structure described in the diagnosis persists in all leaves. Variations refer to the degree of cell papillosity in stomatal bands (papillae can be well developed or faint and then the periclinal walls bear irregular ridges or fissures; Plate I, 2-4, 7, 11), the degree and type (continuous or marginal of different width) of cutinization of subsidiary cells (PlateI, 3, 4, 7, 11; Fig. 4L, M). In some stomata one or two subsidiary cells lack stronger cutinization (Fig. 4L to left, Fig. 5A). Solitary stomata are totally devoid of the stronger cutinization of the subsidiary cells (Fig. 4M), and then one can better see that the periclinal walls of the subsidiary cells are inclined towards the aperture and form the outer stomatal pit. Leaf mucro is short and strongly cutinized (Fig. 4H). Trichopodia and papilligers in stomatal bands are rare (Plate I, 2). Cells of the non-stomatiferous bands are more elongated than in stomatal bands, and orientated along the leaf axis or obliquely to the leaf margin. The axial non-stomatiferous band gradually disappears towards the apex where stomatal bands merge (Plate I, 8). Still further towards the apex the axial stomatal band grades into a band of papillose cells. Marginal teeth vary from short, even papilla-like (Fig. 4E, F) to very long, hair-shaped with inner cavity (as in Fig. 6I). Sometimes the uniseriate arrangement of the teeth is violated. Lower cuticle shows elliptical, spindle-shaped or slit-like lacunae (Plate I, 12) of unknown nature. They vary both in size and number. Probably they reflect certain organic characters rather than a degree of preservation. In the basal part and sometimes near the margins, the lower side shows a few stomata (Fig. 5B) in which subsidiary cells are less prominent in their cutinization than in

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Fig. 5. Kungurodendron sharovii Meyen, sp. nov. Stomata of bracts of compound polysperms. (A) Unequal cutinization of subsidiary cells, upper epidermis, preparation No. 3737/168, point 3. (B) The same on lower epidermis in basal part of bract, preparation No. 3737/149-B, point 4. Locality Chekarda-l. Scale bar= 300 ~tm.

stomata of the upper side. The large subapical trichopodium (Plate I, 1, 9; Fig. 4H, K) is situated at a distance of 2.5-3.5 mm from the leaf tip. It is surrounded by short cells forming a field of an anomalous cellular structure 250-500 ~tm in length and 200-300 ~tm in width. Axes bearing compound polysperms are 5-6 mm wide. Their leaves are of the same size as those of axes of the penultimate order, and grade upwards into bracts. The latter differ from foliage leaves in more delicate cuticle and more spaced stomata in stomatal bands (Plate II, 15, 19, 21-23). Marginal teeth are usually extended into long hairs (Plate II, 15; Fig. 61). The upper cuticle can be traced down to the base thus suggesting that the bract was free. The gross morphology of the compound polysperm is better seen in the holotype (Plate II, 13, 14)

bearing no less than 60 axillary complexes. The holotype preserved a subtending leafy axis, bracts are well seen laterally and apically, axillary complexes are faced to observer with their abaxial side without seed stalks. Axillary complexes were studied in other specimens by the dissolution of fragments of compound polysperms in hydrochloric acid, there are also detached axillary complexes (Plate II, 16-18, 20; Plate III, 26-31; Fig. 6A-C). The basal part of the axillary complex slightly widens upwards and is covered by distant subtriangular sterile scales (Plate II, 18). Their lower epidermis is composed by longitudinal files of cells gradually passing on to the axis. Owing to a high adnation of the scales to the axis, only a narrow asymmetrical strip is left of the lower surface. Cells are sometimes papillose (Fig. 6D), stomata may

Fig. 4. Kungurodendron sharovii Meyen, sp. nov. Vegetative shoots and their epidermal structure. (A) Branch of penultimate order with long pinnately arranged branches of last order, spec. No. 3737/111. (B) Apex of thick leafy branch, spec. No. 3737/254A. (C) Spec. No. 3737/26. (D) Spec. No. 3737/15A. (E) Leaf margin with teeth and narrow stomata-free band, preparation No. 3737/15A, point 4. (F) Marginal teeth, preparation No. 3737/270, point 3B. (G) Branch of penultimate order with short branches of last order, spec. No. 3737/270. (H) Leaf apex with mucro, subapical trichopodium and surrounding field of anomalous cells (at arrow), preparation No. 3737/240-A. (I) Spec. No. 3737/240. (J) Unfolded cuticle near leaf margin, black band shows cuticle fold along margin, stomata shown by dots, leaf axis at arrow, preparation No. 3737/270, point 3B. (K) Unfolded leaf cuticle, stomata shown by dots, subapical trichopodium with surrounding field of anomalous cells at arrow, preparation No. 3737/110-a. (L) Stomata with strongly cutinized radial walls of subsidiary cells, preparation No. 3737/270, point 3. (M) Stoma with subsidiary cells without additional cutinization, preparation No. 3737/110-a, point 1. Locality Chekarda-1. Scale bar = 10 mm (A), 5 mm (B-D, G, I), 1 mm (H, J, K), 100 ~tm (E, F), 10 lam (L, M).

S. ~ Meyen / Review ~l Palaeobotanv and Palvnology 96 1997) 351 447

368

PLATE I

s. v. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

occur. The margin bears the same teeth or short hairs as foliage leaves, a mucro may be present (Fig. 6G, H). On detached axillary complexes these sterile scales are well seen on the adaxial side. Higher up, the axis shows longitudinal cuticular folds corresponding to decurrent bases of seed stalks and funiculodia. The seed stalks (Plate III, 27-31) are 4 mm long in their free part. Their hooked part is 1 . 5 - 3 m m long, cylindrical (Fig. 6E) or apically widened. The seed occupies nearly the entire apex of the stalk, and is bordered by a circular swelling (Plate III, 28, 29). The apical funiculodia are erect with tongue-shaped or elliptical apices which sometimes show strongly reduced seed scars (Plate III, 27, 30, 31) bordered by thin cuticular flanges that probably belong to aborted ovules. In the better preserved microstrobilus (Plate III, 32) the stalk is torn. The partially longitudinal fracture plane shows a thin axis, and microsporophylls attached at a right angle (Plate III, 33). The distal part of the microsporophyll is elbow-shaped and leaf-like. The preservation of microsporophylls is insufficient to recognize the number and mode of attachment of sporangia. Cuticular fragments show the same stomatal structure as leaves and bracts. Pollen (Plate III, 34-43; Fig. 7 A - G ) globose, 40-55 (50) ~tm in diameter, often folded and then boat-shaped (Plate III, 34) with the distal surface outside. Unfolded grains showing both sides are relatively few. The proximal side, when

369

exposed, shows small (6-10) monolete, dilete or asymmetrically trilete slit (Plate III, 36, 42). The surrounding exine displays faint microgranulation. Near the equator the granulation grades into a reticulate structure. The network is formed by columellae-like elements, the meshes being irregularly polygonal (Plate III, 40, 41, 43). The dimension of the meshes increases towards the distal pole. In places the columellar-like layer forms shallow swellings with a cavity inside (Plate III, 35, 40, 42). The above description is based on observations made with the light microscope. B. Lugardon (Toulouse) studied sections of the pollen with TEM. His photographs show that columellaelike elements are accompanied by residual alveolar (quasisaccate) structure. A detailed description of the T E M observations will be published elsewhere. For the present this pollen can be conventionally called quasisaccate with columellar-like sexine.

5.1.2. Kungurodendron? sp. S V M - 1

(Fig. 7I,J?, 8C) Description: Locality Shapkina yielded conifers probably related both to Kungurodendron and Timanostrobus. There are several vegetative shoots described below as Cyparissidium entsovae, and an

accumulation of seeds, both associated with masses of uniform pollen (Plate IV, 48-53; Fig. 71, J) very similar to that of T. muravievii. The preservation

PLATE I Kungurodendron sharovii Meyen,gen. et sp. nov. Epidermal structure of leaves of vegetativeshoots. Chekarda-1. 1. Largesubapical trichopodium on lower leaf surface. Preparation No. 3737/240-A, point 2. × 300. 2. Uppercuticle, portion of stomatal band with trichopodium in the center. Preparation No. 3737/15A-A, point 2. × 300. 3. Uppercuticle, portions of the same stomatal band with numerous papillae (to left) and without papillae and with fissured cuticle of periclinal walls (to right). Preparation No. 3737/112B, point 2. x 300. 4. Uppercuticle, stomatal band, fissured cuticle of periclinal walls, the same point. × 300. 5, 6. Upper cuticle, stoma at differentfocus. Preparation No. 3737/240-A, point, x 1000. 5. Papillaeon subsidiary cells. 6. Guard cells. 7. Uppercuticle, stomata without intensifiedcutinization of subsidiary cells. Preparation No. 3737/112A, point 18. × 300. 8. Outfoldedcuticle in leaf apex, in the middle merging papillose stomatal bands; black bands are leaf margins. Preparation No. 3737/112A, point 22. x 30. 9. Lowercuticle with subapical trichopodium and surrounding group of cells (to left). Preparation No. 3737/110a. × 100. 10. Leaf margin, to the right lower cuticle with differentiatedcutinization of periclinal walls. Preparation No. 3737/110. x 100. 11. Uppercuticle, stomatal band with strongly papillose cells. Preparation No. 3737/112B,point 13. x 300. 12. Lowercuticle with slit-like lacunae. Preparation No. 3737/112B, point 12. × 100.

S. b: Me.l'en Review ~1 Palaeobotany amt Pa(vnologv 96 (1997) 351 447

370

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Fig. 6. Kungurodendron sharovii Meyen, .V~. nov. Female fructifications• (A) Adaxial view of axillary polysperm, spec. No. 3773/559-2• (B) The same, spec. No. 3773/675-1. (C) Same specimen after removal of most part of compression, on the matrix an imprint of abaxial side of polysperm is seen. (D) Papillose epidermal cells of sterile scales of basal part of polysperm, preparation No. 3737/62A-A, point 5. (E) A part of hooked seed stalk with apical seed scar and short hair (at arrow), preparation No. 3737/149a, point 33. (F) Straight funiculodia with aborted seed scars• preparation No. 3737/149-B. (G) Sterile scale from basal part of axillary complex with distinct mucro, preparation No. 3737/149aC. point 14. ( H ) The same without mucro, preparation No. 3737/149a, point 6. (I) Hairy margin of bract, preparation No. 3737/149aB, point 3C. Locality Chekarda-I. Scale b a r = 1 mm (A-C, E. F ), 300 I~m (G, H ), 100 Iam (D, I).

and amount of material is insufficient for an accurate diagnosis of the genus and a species description. Vegetative shoots are similar to those of K. sharovii in leaf microstructure. One of the vegetative shoots (Fig. 8A) lies side by side with an accumulation of vegetative leaves or scales of indistinct outline, an organ resembling a hooked seed stalk (Fig. 8C, top left), and several seeds (Fig. 8C, to right). The seed compression has broken into pieces during maceration, so that the interrelation of cuticular membranes could not be established. The megaspore membrane is thick. The seeds are more than 5 m m long and not more

than 3.5 m m wide, oval, with elevated middle part and flattened indistinct flange. Pollen of the same type has been found inside the seeds. Comparison: In the structure of the associated pollen, these remains are related to Timanostrobus muravievii, whereas the presence of a hooked seed stalk (?) and the structure of associated leaves (see Section 5.2.2) suggest relationships to Kungurodendron sharovii. This combination of characters agrees well with the intermediate stratigraphic position of these remains. They are younger than K. sharovii and older than T. muravievii. Locality: Shapkina.

S. V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351 447

5.2. Genus Cyparissidium Heer,1974 Diagnosis: Phyllotaxis helical. Free part of leaf gradually narrows from leaf cushion (leaf base). Leaves more or less flattened, adpressed to stem, elongated. Leaf length exceeds width of leaf lamina or leaf cushion. Remarks: General questions of the systematics and nomenclature of older conifers are treated in Section 8. In the same section a system of formgenera for vegetative shoots that cannot be studied in necessary detail, is explained. Following Harris (1969, 1979) and Andersons (Anderson and Anderson, 1985) Cyparissidium is treated as such a genus. Its diagnosis is given above, following Harris with minor changes. According to the diagnosis, Cyparissidium comprises shoots with leaves adpressed to the axis. This character, however, may show variations. It is here adopted that a species fits the generic diagnosis even when the diagnostic characters are observed in a part of its specimens. Obviously, taxonomic decisions in such cases will be conventional. 5.2.1. Cyparissidium appressum (Zalessky) Meyen, comb. nov. (Fig. 9) Basionym: Walchia appressa Zalessky, 1937, Probl. Paleontol., 2-3: 73, figs. 37, 38 (lectotype). Synonyms: 1927? Walchia hypnoides Brongniart, Zalessky, pp. 31, 48, pl. 33, figs. 5, 5a. 1939? Walchia (Lebachia) stricta Florin (pars), Florin, p. 224, pl. 135/136, figs. 11, 12. Description: The following description of the species is given in the protologue (Zalessky, 1937, pp. 73-74): "Empreinte d'une petite branche feuillre d'une longueur d'environ 8 cm et d'une largeur 4.5 mm portant des feuilles lancrolres uninervres 6troitoment strires/t la branche et atteignant une longueur de 7 mm et une largeur de 0.75 mm dans leur partie moyenne (fig. 37 [= Fig. 9A in the present paper]). Je rapporte /l Walchia appressa Zalessky non seulement les fines petites branches plus grosses atteignant une largeur ou une 6paisseur de 10 mm et portant de feuilles serrres/t ces

371

branches comme aux premirres dont une est reproduite sur la fig. 38 [= Fig. 9B in the present paper]". Later on, Zalessky (1939, pp. 362-363) supplemented the description on the basis of additional specimens including an unbranched shoot 20 cm long and 12mm wide having leaves up to 10-20 mm long, as well as a branched shoot 23 cm long with pinnately arranged alternating branches. He mentioned that the leaves show a keel sometimes complicated by a furrow, and numerous thin veins. Obviously imprints of hypodermal strands had been mistaken for veins, because such dense venation is hardly possible in conifers. Among hand specimens coming from the type locality ( Krutaya Katushka) as well as other localities mentioned by Zalessky (Krasnaya Glinka, Chekarda) there are many shoots fitting the above description and accompanying drawings. Their belonging to the present species is beyond doubt. On the other hand, these shoots are indistinguishable in their gross morphology from those associated with Kungurodendron sharovii. However, in some other localities similar shoots are not accompanied by fructifications or such vegetative shoots for which epidermal characters are known. This concerns, e.g. the shoot referred by Zalessky to W. hypnoides and by Florin to W. stricta (see synonymy). The type specimens of C. appressum and K. sharovii may well belong to one natural species. If we merge them, however, the species C. appressum would comprise both sterile shoots and fructifications. In such a case, this taxon comprising well preserved fructifications and pollen will be typified by a fragment of a vegetative shoot which has been lost and is even devoid of epidermal characteristics. That is why it seems unreasonable to apply the epithet "appressum" to shoots having epidermal characteristics, and to fructifications. For the same reasons, Zalessky's specimens and similar ones when they are devoid of epidermal characteristics are given the generic name Cyparissidium. Accordingly, the new combination Cyparissidium appressum (Zalessky) is here proposed (the basionym and lectotype are noted in the synonymy). Unfortunately, in his description of Walchia appressa, Zalessky had not mentioned the margin structure. In all the specimens placed by me into

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X V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

C. appressum, marginal microdenticulation is well seen. This character distinguishes C. appressum from other species of the genus excepting C. entsovae (see below). In the future such shoots having leaves with the marginal microdenticulation may be referred to a separate genus. 5.2.2. Cyparissidium entsovae Meyen sp. nov. (Plate IV, 44-47; Fig. 8A,B,D,E) Holotype: Specimen No. 3775/117-2 (Plate IV, 44-47; Fig. 8A). Type locality: Shapkina. Etymology: after the geologist F.I. Entsova who collected the type material. Description: Vegetative shoots of two types provisionally designated as A and B. The type A (Plate IV, 44-47; Fig. 8A, B) comprises the holotype. This is a fragment of a shoot of the last order (?), covered with tightly adpressed oval leaves with a long mucro (Plate IV, 47). The leaves are 2.2-2.5 mm wide and 5-6 mm long. The cuticle is strongly corroded. One (lower?) leaf side (Plate IV, 44 to left, 45) is covered by wavy files of longitudinally orientated cells. On the opposite side (Plate IV, 44 to right, 46) cells of the same kind form a band along the leaf axis and two marginal bands with a band of papillose cells in between. Stomata can be recognized after groups of papillae belonging to subsidiary cells (Plate IV,

373

46). The leaf margin is microdenticulate, but the cuticle of the teeth is heavily damaged. This shoot lies on the fracture plane side by side with remains described above as Kungurodendron? sp. SVM-1. The same bed yielded 5 specimens of leafy shoots of the type B (Fig. 8D, E). Leafy axis of the penultimate order bears pinnately arranged short (about 3.5 cm long) branches of the last order. Leaves are falcate, 1.5-2 mm long and 0.6-0.7 mm wide, with acuminate apex. The cuticle is heavily corroded so that among diagnostic characters only a long mucro and the marginal microdenticulation (as in the type A) can be recognized. Remains of both types are covered by masses of pollen (Plate IV, 48-53) of the same general type as in Timanostrobus muravievii (see Section 5.3.1). In places the pollen covers the cuticle with a continuous layer. Most of the grains are folded in varying degrees (Plate IV, 48, 49, 53). Unfolded grains (Plate IV, 50-52) show a distinct transversal proximal slit. Presently it is difficult to say whether the types A and B belong to one natural species. Having a rather different gross morphologically, they are united by such common characters as marginal microdenticulation, mucronate leaf apex and type of associated pollen. Probably they represent axes of different order. The same pollen covers seeds of Kungurodendron? sp. SVM-1 and was found inside them. All these parts appear to have belonged to

PLATE I I Kungurodendron sharovii Meyen, gen. et sp. nov. Polysperms. Chekarda-1. 13, 14. Compound polysperm. Holotype. No. 3737/259. 13. xl. 14. x3. 15. Cuticular structure of bract near margin, to left--upper cuticle, to right--lower cuticle with slit-like lacunae, marginal hairs are seen. Preparation No. 3737/149-A, point 25. × 100. 16. Solitary hair on seed stalk (see also Plate III, 29; Fig. 6E at arrow). Preparation No. 149a, point 33. x 300. 17. Cuticular papillae on the edge of the same seed stalk. Same point, x 300. 18. Cuticle of basal part of axillary complex, sterile scale with denticulate upper margin. Same preparation, point 6. x 50. 19. Lower cuticle of bract. Preparation No. 3737/149-B, point 4. × 100. 20. Stoma and cells with cuticular papillae in basal part of seed stalk. Preparation No. 3737/149aA, point 22. × 300. 21. Upper cuticle of bract, stomatal band with papillose cells and intensified cutinization of subsidiary cells, to left trichopodium. Preparation No. 3737/149aB, point 3A. x 300. 22. Upper cuticle of bract, stomata with lesser cutinization of subsidiary cells. Preparation No. 3737/149-B, point 2. x 3. 23. Lower cuticle with slit-like lacunae, to left margin with hairs. Preparation No. 3737/149-A, point 25. × 100. 24. Stomatal files of upper cuticle of sterile leaf belonging to polysperrn-bearing axis. Preparation No. 3737/168, point 10A. × 100. 25. Sinuous sutures along radial walls in basal part of axillary complex. Preparation No. 3737/149aC, point 14. x 1000.

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S. V. Meyen / Review of Palaeobotanyand Palynology96 (1997) 351-447 the same plants closely related to Kungurodendron. These plants differ from K. sharovii and Cyparissidium appressum in the more attenuated apex in leaves o f type A, the more diverging smaller and falcate leaves o f type B, inconspicuous cutinization o f subsidiary cells o f stomata by comparison with non-stomatal epidermal cells. In the presence of papillose stomatiferous bands on one leaf side, the type A is certainly closer to Walchia (al. Lebachia) and K. sharovii, than to T. muravievii, but in the structure o f the associated pollen C. entsovae it is closer to T. muravievii than to K. sharovii, Type B is similar to Walchia (al. Lebachia) in the leaf outline and microdenticulation.

375

thin quasisaccus embracing inner b o d y both equatorially and distally. Monolete proximal aperture m a y be present. Etymology: F r o m the Timan R a n g e and strobus (Latin), cone. Comparison: Timanostrobus differs strikingly from all other conifers in having m u c h more n u m e r o u s m o n o s p e r m s (seed stalks) on lateral polysperms which are remarkably similar to those o f the Vojnovskyales. M o r e detailed comparisons with other genera and the systematic affinity of the genus are considered in Sections 6 8.

5.3.1. Timanostrobus muravievii Meyen ,sp. nov. 5.3. T I M A N O S T R O B U S Meyen, gen. nov.

(Plate IV, 54; Plate V; Plate VI, 66-74; Figs. 7K,L, 8G, 1 0 A - H , 11, 12, 36P)

Type: Timanostrobus muravievii Meyen, sp. nov. Diagnosis:

Genus established for polysperms, although other organs are also known. Main axis o f c o m p o u n d polysperm terminating a leafy twig and bearing helically arranged simple lateral polysperms. Bracts have not been observed. Thick axis o f lateral polysperm densely covered by n u m e r o u s monosperms, replaced near apex by sterile scales. Leafy twigs o f Brachyphyllum or Pagiophyllum type. Male strobili crown leaf twigs. Pollen with

Holotype: Specimen No. 4577/207-2 (Plate V, 55, 61, 62; Fig. 10A). Type locality: Vym-1. Etymology: After Prof. I.S. Muraviev. Diagnosis: C o m p o u n d polysperms up to 11 cm long and 4 c m wide (main axis 10-13 m m wide) terminating leafy twigs o f the Brachyphyllum or Pagiophyllum type. Simple lateral polysperms obconical. Seeds winged, on short decurrent stalks. Megaspore m e m b r a n e thick, 0.8 2.5 m m long,

PLATE llI

Kungurodendron sharovii Meyen, gen. et sp. nov. Fructifications. Chekarda-l. 26. 27. 28. 29. 30. 31. 32. 33. 34 43. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43.

Detached axillary complex with abscissed seeds, stereopair (= Fig. 6A, reconstruction on Fig. 32B). Spec. No. 3773/559. x 5. Apex of funiculodium with scar of aborted ovule. Preparation No. 3737/259-1, holotype, x 30. Apical seed scar on seed stalk. Preparation No. 3737/149a, point 33. x 100. The same seed stalk (= Fig. 6E). x 30. Funiculodium with scar of aborted ocule. Preparation No. 3737/149aB, point 1. x 30. Outfolded cuticle of funiculodium, at top--scar of aborted ovule, Preparation No. 3737/149-B, point 3. x 50. Microstrobilus. Spec. No. 3737/176. x 2. Same specimen, detail of middle part. x 5. Pollen from the same microstrobilus, preparation No. 3737/176. Boat-shaped grain, point 21(1). x 500. Grain with slight exfoliation of sexine at left (=Fig. 7E), point 21(11). x 500. Proximal side with dilete slit (=Fig. 7A), point 17. x 500. Same grain, middle focus. × 500. Same grain, focus at reticulum on distal side. × 500. Fine reticulum near equator, point 31 (1). x 1400. A coarser reticulum of distal side and columellar-like sexine (= Fig. 7B), point 19(I1). x 1400. Reticulum with large meshes, point 28. × 1400. Proximal slit, saccus-like exfoliation of sexine (= Fig. 7D), point 26( 1). x 1400. Reticulum with large meshes and well seen columella-like elements seen as minute black dots, point 31 (III), x 1400.

376

S. V. Meyen / Review o/ Palaeohotany und Palynology 96 (1997) 351 447

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Fig. 7. Schematic drawings of pollen of Kungurodendron sharovii Meyen, ~sT~.nov. ( A G ) , Timanostrobus muravievii Meyen, sp. nov. (K-L) and pollen associated with Kungurodendron? sp. SVM-1 and Cyparissidium entsovae Meyen, sp. nov. (H-J). (A-G) Preparation No. 3737/176, body outline shown by thicker line, points 17 (A), 18 (B), 25 (C), 26 (D), 21-II (E), 4 (F), 23 (G). (H) Preparation No~ 3775/117-5(4, 5, 6), point 3. (I) Preparation No. 3775/115, point 2. (J) Preparation No. 3775/117-5(4,5,6), point 1. (K, L) preparation No. 4100/193-1 points 19 ( K ) and 4 (L). Localities Chekarda- 1 ( ~ G ) , Shapkina ( H J ), Vym-2 ( K, L). Scale bar = 10 ktm.

0.6-1.7 mm wide. Elongated microstrobili, 6-8 mm wide and 3-4 cm long, occupy apical position on repeatedly dichotomizing leafy twig. Pollen 50-70 gm in diameter, round, often folded and then boat-shaped. Quasisaccus with densely spaced minute lumina, gradually disappearing at proximal side near equator and leaving a smooth proximal area with a monolete slit. Foliage leaves from lanceolate to widely rhomboid, adpressed to axis or slightly diverging. Epidermis uniform on both leaf surfaces. Stomata freely scattered or in

indistinct short files, variously oriented. 5-6 subsidiary cells form regular ring around sunken guard cells. Leaf margin microdenticulate. Description: The description is based on several dozens of compound polysperms (Plate IV, 55-60; Fig. 10A-E, Figs. 11, 12). In three of them the compound polysperms crown leafy twigs (Plate V, 55, 56, 60; Fig. 10C). Lateral simple polysperms are usually with shed seeds, but in the holotype (Plate V, 55, 61, 62; Fig. 10A) and several other specimens, middle portions of lateral polysperms

S. V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

yield megaspore membranes obviously belonging to persistent seeds. The compression of this portion of polysperms yields cuticle of axis and attached monosperms (Fig. 10E). The latter are helically arranged. Judging from cuticle preparations, seed stalks were short and decurrent. The epidermal cells of the stalks are longitudinally elongated; stomata have not been found. The same cuticular structure was observed in sterile apical scales (Plate V, 57; Fig. 10F). Sterile scales may be present (?) also in the basal part of lateral polysperm. In the seed compressions only megaspore membranes are well preserved. The integument outer cuticle has not been recognized, its inner cuticle is thin showing longitudinally oriented cells. The nucellar cuticle is also thin along the megaspore margins and thicker above the megaspore. The pollen found near the nucellar apex is the same as in associated microstrobili (Plate V, 61, 62). The same bed yielded many dispersed seeds with a thick megaspore membrane and membranaceous wings of variable outlines. In one of the seeds (Fig. 8G) the same pollen was found. The elongated cylindrical microstrobili (Plate V, 63, 65; Fig. 10H) have not preserved structural details. Both the microsporophyll structure and disposition of sporangia cannot be recognized. Some sporangia and attenuated apices of microsporophylls are seen along the margins of the microstrobili. The pollen found in situ (Plate IV, 54; Plate VI, 66, 71; Fig. 7K, L) varies in outline, depending on the degree of folding. In unfolded grains a short occasionally opened proximal slit and a boundary of the quasisaccate structure are seen (Plate IV, 54; Plate VI, 66, 67, 69-70). Many grains are folded and boat-shaped (Plate VI, 68). They may be easily mistaken for monocolpate pollen identified in the literature as Ginkgocycadophytus, Coniferites, Ginkgaletes, etc. Similarly looking pollen was figured (without description) under the names Acusporidatina reticuloida Koloda and Reticulatina heterobrochata Koloda (Molin et al., 1983, pl. 6, figs. 1-4; pl. 7, figs. 1-6). The leaves of leafy shoots are rhomboid or lanceolate, closely adpressed to the axes or somewhat diverging (Plate V, 56, 64; Fig. 11). The shoots are up to 10 cm long and 2-18 mm wide. Some of them are longitudinally split (Fig. l lB).

377

They branch irregularly at acute angles. Most of the specimens yielded thick, but heavily corroded cuticle with indistinct cell outlines and damaged marginal cuticular teeth. One specimen yielded a better preserved cuticle (Plate VI, 72-74; Fig. 11C, D). Stomatal pits are small and have thickened borders (Plate VI, 72, 73; Fig. liD). Subsidiary cells do not differ from other epidermal cells in either outline or cutinization. Marginal teeth (Plate VI, 74) are fused into a continuous denticulate cutin band. Shoots found together with fructifications (but not in organic connection) can be conventionally called Brachyphyllum (al. Pagiophyllum) sp. SVM-1 until fuller cuticular characteristics are obtained. Localities: Vym-1, Vym-2.

5.4. CONCHOLEPIS Meyen, gen. nov Type: Concholepis harrisii Meyen, sp. nov. Diagnosis: Axis of compound paniculate polysperm continuing from leafy branch. Seed scales and free subtending bracts arranged in a loose helix. Bracts simple and well developed. Seed scales isometric or elongated, with marginal, abaxial and submarginal adaxial projections, bearing more than one seed. Etymology: Greek, concho, shell; lepis, scale. Comparison: Concholepis differs from other coniferalean genera in having numerous projections on the seed scales. Probably, the axillary polysperms of Concholepis have arisen in a different way than true seed scales. This question, as well as the systematic position of the genus and relations to other genera are discussed in detail in Sections 6, 7 and 8.

5.4.1. Concholepis harrisii Meyen sp. nov. (Plate VI, 75-77; Plate VII, 78, 79; Figs. 13, 14, 15A,B, 36Q)

Holotype: Specimens No. 3974/27-1 and counterpart 27A-1 (Plate VII, 79; Fig. 13A). Type locality: Vym-1. Etymology: After the late Professor T.M. Harris. Diagnosis: Compound polysperms up to 12 cm

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S. IL Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

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Fig. 8. Cyparissidium entsovae Meyen, ~p. nov. (A, B, D, E), Kungurodendron? sp. SVM-l (C), 'Walchiostrobus' lodevensis Florin (F) and Timanostrobus muravievii Meyen (G). (A) Sterile shoot, transfer-preparation No. 3775/117-2 (epidermal structure see Plate IV, 44-46). (B) Reconstruction of leaf. (C) Group of seeds (to right) and putative seed stalk (top left), transfer-preparation No. 3775/117-5(1-7). (D, E) Branches with smaller leaves. (F) Polysperms, drawn after photograph of Florin (1940, pl. 155/156, fig. 2). (G) Dispersed seed, spec. No. 3974/33b-3. Localities Shapkina (A-E), Lod6ve, France (F), Vym-1 (G). Scale b a r = 1 mm (A-D, G), 1 cm (E, F).

PLATE IV 44-47. Cyparissidium entsovae Meyen, ap. nov. Holotype. Shapkina. 44. Outfolded cuticle of both leaf sides, to left lower cuticle, to right upper cuticle with two stomatal bands. Preparation No. 3775/117-2, point 2. x 50. 45. Lower cuticle shown from margin (to left) to leaf axis (darker band to right), same point. × 100. 46. Portion of papillose stomatal band (to left), same point, x 300. 47. Mucro at leaf apex. Same preparation, point 5. x 230. 48. Kungurodendron?sp. SVM-1. Shapkina. Pollen at cuticular membrane of ovule. Preparation No. 3775/117(4, 5, 6), point 2. x 230.

49-53. 49. 50. 51. 52. 53. 54.

Pollen associated with Cyparissidium entsovae Meyen, sp. nov. and Kungurodendron? sp. SVM-1. Shapkina. Boat-shaped grain. Preparation No. 3775/115-a, point 5. x 500. Partly infolded grain (= Fig. 7I). Preparation No. 3775/115, point 2. x 500. Same grain focus at proximal slit, differential interference contrast. × 1000. Same grain, focus at distal perforations. Boat-shaped grain (=Fig. 7J), focus at distal side. Preparation No. 3775/115A, point 4. x 1000, Timanostrobus muravievii Meyen, gen. et ~p. nov. Vym-1. Pollen grain turned by smoother proximal side to top (= Fig. 10G). Preparation No. 4577/223-1, point 4. × 500.

S. bq Meyen , Review q/Palaeobotany and Palynology 96 (1997) 351-447

380

A B Fig. 9. Cyparissidium appressum (Zalessky) Meyen. comb. nov. (AI Syntype. (B) Lectotype. Redrawn from Zalessky (1937, figs. 37, 38--1ectotype). Localities Krutaya Katushka (A). Chekarda-1 (BI. Scale bar = 1 cm.

long. Bracts lanceolate, about 2 mm wide and more than 1 cm long. Seed scales sessile, nearly circular or elongated, about 10ram long. Marginal, abaxial and submarginal adaxial projections obtuse, parallel sided, 2 m m long and 1-2 mm wide. Abaxial projections arranged in crossed oblique rows. Seeds at least two per scale: megaspore membrane thick. Vegetative shoots of the Geinitzia type. Description: Two entire (including the holotype) and three fragmentary compound polysperms (Plate VI, 75-77; Plate VII, 79; Figs. 13, 14) associated with a large number of sterile twigs (Plate VI, 78; Fig. 15A, B). In the holotype the more or less median fracture plane exposes the axis, bearing 8-9 axillary complexes on each side. They are obliquely orientated on the fracture plane. Therefore, only distal projections on the seed scales are observed. Another compound polysperm looks similar (Fig. 14). The third specimen (Plate VI, 76, 77; Fig. 13B, C), although fragmentary, shows the general outline and both surfaces of the seed scales. Flattened marginal projections are more than 12 in number, separated by narrow

sinuses. After the powdery compression was removed, the abaxial side exhibited 12 projections arranged in several crossed oblique rows. The adaxial side bears vestigial submarginal projections, alternating with marginal ones. Otherwise this side is smooth, with very feeble oval contours which may correspond to seed scars. A possibility exists, however, that the submarginal projections are vestigial seed stalks, and the oval contours are artefacts of preservation. The bract (Fig. 13D) is hastate, with a semicircular transverse basal fold, and is longitudinally striated. A tentative reconstruction of the axillary complex is shown in Figs. 13E and 36Q. The branching of vegetative shoots is irregular. Leaves are helically arranged, decurrent, falcate or nearly straight, up to 12mm long and up to 2.5 mm wide. Some of the shoots bear strange short lateral branchlets with shorter leaves ( Plate VII, 78, lower part). These may be underdeveloped proliferating polysperms. The cuticle is badly preserved. Localities: Vym-1 (polysperms and vegetative shoots), Vym-2 (vegetative shoots).

PLATE V Timanostrobus muravievii Meyen. gen. et sp. ran', Vym-l, 55-62, 64: Vym-2, 63, 65. 55. Holotype. No. 4577/207-2 (= Fig. 10A). x 1. 56. Leafy part of shoot bearing compound polysperm ( - Fig. 10C). Spec. No. 4577/'207-1. x 2. 57. Cuticle of sterile scales in apical part of lateral polysperm. Preparation No. 4577/207-1. x 10. 58. Lateral polysperms in apical part of compound polysperm (= Fig. 10D). Spec. No. 4577/223-1. x 3. 59. Leafy axis bearing compound polysperm. Spec. No. 4577;201. × 1. 60. Same specimen, detail, x 2. 61. Holotype. Megaspore membrane and fragments of cuticular membranes of nucellus and integument, at top --pollen grain. Prep. No. 4577/207-2, point 2. x 100. 62. The same magnified to show pollen grain. × 5(10. 63. Edge of microstrobilus with protruding microsporophylls and sporangia. Prep. No. 4100/193-1. x 10. 64. Vegetative shoots. Prep. No. 4577/206. x 2. 65. Microstrobili on branching twig. Prep. No. 41(t0193-1. x I.

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Fig. 10. Timanostrobus muravievii Meyen, ~V~-nov. (A H) and l))jm)vskya paradoxa Neub. (1, J ). (A) Holotype No. 4577'207-2, megaspore membranes in black. (B) Scheme of seed stalk structure, ovule shown by broken line, arrow corresponds to projection of the ovule main plane onto plane of drawing. (C) Lower part of compound polysperm, and subtending leafy twig, spec. No. 4577/207-1. (D) Lateral polysperms with seed stalks in their basal parts, and sterile scales apically, spec. No. 4577/223-1. (E) Basal part of lateral polysperm with numerous short seed stalks, drawn after cuticular preparation No. 4577/223A-1. (F) Cuticle of apical sterile scale, preparation No. 4577/201-3, point I. (G) Pollen stuck to lateral polysperm, proximal side to left, preparation No. 4577/223-1, point 4 (see Plate IV, 54). (H) Microstrobilus on leafy twig, spec. No. 4577/104A-2. (I) Flattened edge of lateral polysperm with bent backwards and apically expanded seed stalk, drawn in alcohol and crossed polarizing filters, spec. No. 3039/194e (C). (J) Scheme of seed stalk, ovule shown by broken line, arrow corresponds to projection of the ovule main plane onto plane of drawing. Localities Vym-I ( A H ) and Mine 3 of Khalmeriyu coal-field, Pechora basin, near coat seam J,~ (1, J ). Scale bar = 1 cm ( A C , H ), 1 mm (E, I), 100 pm (F), 10 pm (G}.

5.5. Genus Sashinia Meyen, 1978 Type: Sashinia aristovensis Meyen, 1978. Diagnosis: Compound polysperms consisting of axis bearing helically arranged distant bracts differing from foliage leaves in larger size. Bract subtends lateral simple polysperm. Its axis bears usual foliage leaves (Quadrocladus type) replaced near the apex by a bunch of seed stalks (monosperms) similar to foliage leaves and having the same epidermal structure. Apex of seed stalk thickened into a terminal pad which is abaxially bent and covers the seed attached to the abaxial

side of seed stalk below its inflection. Associated vegetative shoots are of the Quadrocladus type, microstrobili of the Dvinostrobus type, pollen of the Scutasporites and Lueckisporites (?) type. Specific composition: S. aristovensis Meyen and S. horealis Meyen. 5.5.1. Sashinia aristovensis Meyen, 1978 (Plate VII, 82 84; Fig. 16A-E) 1976 1 9 7 8 Sashinia aristovensis Meyen, pp. 304 306, pl. 2, fig. 15; text-fig. 13. 1986 Sashinia aristovensis Gomankov and Meyen, pl. 15, figs. 9 12; pl. 16, figs. 1-5; text-figs. 66, 67a zh.

S. V, Meyen / Review o f Palaeobotany and Palynology 96 (1997) 351-447

383

A Fig. 11. Timanostrobus muravievii Meyen, sp. nov. Leafy shoots and their epidermal structure. (A) Spec. No 3974/27-4. (B) Spec. No. 3974/2-2, 2-3, section of wood shaded. (C) Spec. No. 4577/198-1. (D) Two stomata, preparation No. 4577/198-1, point 1. Locality Vym-1. Scale b a r = l cm (A, B), 1 mm (C), 100 ~tm (D).

Fig. 12. Timanostrobus muravievii Meyen, sp. nov. Reconstruction of compound polysperm and a detached lateral polysperm.

Description (abbreviated, for full description see: Gomankov and Meyen, 1986): Dwarf fertile shoots (lateral polysperms) bear 10-11 monosperms 5 - 6 m m long, helically arranged. Their inflected terminal pads (initially erroneously described as seeds, Meyen, 1976-1978, 1981) are

4-5 mm long and 2-3 mm wide. The ovule is nearly fully covered by the terminal pad (Plate VII, 84; Fig. 16B, C). Seeds after shedding left scars (Plate VII, 85). The megaspore membrane was not observed. The interrelation of cutinized membranes inside the seed is not fully understood. The

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S. V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447 nucellus (Plate VII, 83) is crowned by a rather long salpinx (as in seeds of Pseudovoltzia and Ullmannia, Schweitzer, 1963). The epidermal structure of the distal pad (Plate VII, 86) and of the stalk below the pad is nearly the same as in associated vegetative twigs, although stomata are less numerous. The epidermal cells are arranged in rather regular files, they are more isometric on terminal pads where the cellular files are seen only along margins. The cuticle is thicker on the outer side of the pad. The associated vegetative shoots belong to Quadrocladus dvinensis (see Section 5.7.2), microstrobili to Dvinostrobus sagittalis (see Section 5.6.1 ), and pollen to Scutasporites unicus Klaus. 5.5.2. Sashinia borealis Meyen, 1981 (Fig. 16F-I, K - S ) 1981 Sashinia borealis Meyen, pp. 162-165, pl. 1, figs. 5-7; pl. 2, figs. 19-21; text-figs. 4-6, 8-13. 1986 Sashinia borealis G o m a n k o v and Meyen, pl. 15, fig. 13; text-figs. 67z-k, 68a-d, zh, 69. Description (abbreviated, for full description see: G o m a n k o v and Meyen, 1986): The main axis of the compound polysperm is more than 40 m m long and 2 m m wide. It bears helically arranged distant bracts attached at acute angles 6 - 7 m m long and 1-1.2 m m wide. Each axillary lateral polysperm

385

produces 2 - 6 seed stalks 6-8 m m long. The epidermal structure of bracts, of sterile leaves of the axillary complexes, and of seed stalks is the same as in leaves of the associated vegetative shoots Quadrocladus borealis (see Section 5.7.3). Epidermal cells of the terminal pad are more isometric and form files only along the pad margins. The nucellus was probably crowned by a salpinx. The megaspore membrane is very thin. Comparison: S. borealis differs from S. aristovensis in a lesser number of seed stalks in axillary complexes and in their smaller size. Epidermal differences between these species are the same as between associated twigs Q. borealis and Q. dvinensis (Table 1 ).

5.6. Genus Dvinostrobus G o m a n k o v et Meyen, 1986 Type: Dvinostrobus sagittalis G o m a n k o v et Meyen, 1986. Diagnosis: Unbranched axis bearing helically arranged microsporophylls having distal triangular shield. Sporangia attached in pairs to sporangiophores departing from microsporophyll the slender stalk in its middle part. Pollen of Scutasporites type. Comparison: Relations between Dvinostrobus and

PLATE vI 66 74. Timanostrobus muravievii Meyen, gen. et sp. nov. Vym-2, 66-71; Vym-1, 72 72. Pollen from sporangium, focused partly at perforations on distal side, partly at fine-grainedstructure near equator on proximal 66. side. Preparation No. 4100/193-1, point 30, differential interference contrast, x 1000. Same grain, focused at proximal slit. 67. Variously folded pollen from sporangium. Preparation No. 4100/193-1, point 14. x 300. 68. Partly folded pollen from sporangium (=Fig. 7K). Preparation No. 4100/193-1, point 19. × 500. 69. Same grain, focused at proximal slit. × 1400. 70. Same grain, focused at distal side, differential interference contrast. × 1000. 71. 72-74. Cuticle of vegetative shoots, preparation No. 4577/198-1. Stomata (a stoma near left margin shown in Fig. liD), point 1. x 300. 72. Same point at lower magnification. × 100. 73. Marginal teeth, point 2. x 100. 74. 75-77. Concholepis harrisii Meyen, gen. et sp. nov. Vym-1. Axillary complex with marginal projections. Spec. No. 4100/201A-7. × 5. 75. Axillary complexes and fragments of bracts (=Fig. 13C), coal left intact. Spec. No. 4577/216A-2. × 3. 76. Portion of counterpart of the same specimen (=Fig. liB), imprints of abaxial appendages are seen. Spec. 77. No. 4577/216B-2. × 3.

386

S. K Meyen / Review oJPalaeobotany and Palynology 96 (1997) 351 447

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Fig. 13. Concholepis harrisii Meyen, sp. nov. (A) Holotype No. 3974/27-1, megaspore membrane of one seed in black. (B) Two seed scales with abaxial projections, a part of bract stippled, spec. No. 4577/216B-2. (C) Counterpart of the same specimen with smooth adaxial surface, marginal and submarginal projections of seed scale, parts of bracts stippled, spec. No. 4577/215A-2. (D) Specimen with fully open bract at the apex, spec. No. 4577/204-1. (E) Schematic longitudinal section of bract (stippled), seed scale (in black) and seeds (shown by broken lines). Locality Vym-1. Scale b a r = 1 cm.

PLATE VII Concholepis harrisii Meyen, gen. et sp. nov. Vym-1. Associated vegetative shoots, at bottom--problematical dwarf-shoots. Spec. No. 4100/61. × 1. 78. Holotype. No. 3974/27a-1. x 1. 79. 80, 81. Pollen Scutasporites unicus Klaus from microsporangia of Dvinostrobus sagittalis Gomankov et Meyen. Aristovo. Preparation No. 4552/164-1d, point 1. × 640. 80. Preparation No. 3954/677-1E, point 1. x 640. 81. 82-86. Sashinia aristovensis Meyen. Aristovo, 83, 84; Titovo, 82, 85, 86. 82. Mucro at the apex of the seed stalk (see Fig. 16D, E). Preparation No. 3782/10-1, point 1. x 100. Ovule, cuticular membrane of nucellus with well developed salpinx (below salpinx a group of pollen grains), and a thinner 83. inner cuticle of integument consisting of larger cells. Preparation No. 4552/92-1a, point 1. x 100. 84. Axillary complex. Holotype. No. 4552/296-1 (=Fig. 16A-C). x 3. Seed scar on seed stalk cuticle (see Fig. 16D, E). Preparation No. 3782/10-1, point 2. x 100. 85. Cuticle of the same seed stalk with indistinct sutures of radial walls, corner spines and stoma (top left). Same preparation, 86. point 6. × 300. Quadrocladus dvinensis Meyen. Aristovo. Holotype. No. 4552/277a-1 (= Fig. 20A, left part), x 3. 87. Geinitzia sp. SVM-I. Viled. Marginal teeth (=Fig. 27). Preparation No. 3774/5-1, point 3. × 100. 88.

388

S. Ii Meyen

Review ¢?/Pah~eohotany am/Palvnoloj(y 96 (1997) 351 447

Fig. 14. Concholepis harrisii Meyen, sp. nov. Compound polysperm, seed at arrow, spec. No. 4577/227.1 (above) and counterpart No. 4577/227A-1. Locality Vym-1. Length of the specimen 154 mm.

other genera established for microstrobili are discussed in Sections 6.2 and 8.

5.6.1. Dvinostrobus sagittalis Gomankov et Meyen, 1986 (Plate VII, 80, 81 Plate VIII, 89: Fig. 17)

Holotype: Specimen No. 4552/494-1 (Plate VIII, 89; Fig. 17A-C); Aristovo. 1981 Microstrobili and pollen associated with Sashinia aristovensis and Quadrocladus dvinensis, Meyen, p. 162, text-figs. 1-3. 1986 Dvinostrobus sagittalis Gomankov et Meyen, pl. 8, figs. 5-8; pl. 9, figs. 4, 5; text-figs. 70, 71V, G. Description (abbreviated, for full description see: Gomankov and Meyen, 1986): The holotype is a lax microstrobilus 1 cm wide and 1.5 cm long. Its main axis is 1.5 mm wide, bearing fine longitudinal striations. The microsporophyll stalk is 3.5 mm long, the terminal shield sized 2.5 x 4.5 mm. The sporangia are aggregated in bunches (Plate VIII, 89), in places their paired attachment to sporangiophore branchlets was observed. Sporangia are

2 3 mm long, with a tapering base; the apex is rounded or truncated, showing a thickening. The sporangial wall is composed by longitudinally oriented cells. The sporangia yielded pollen Scutasporites unicus (Plate VII, 80, 81; Fig. 17D). The quasisaccus outline and the distinctness of taenia are very variable. The cuticle of the distal shield is similar on both sides, although somewhat thinner abaxially. The epidermal cells are arranged in regular files, sometimes bearing corner spines. Periclinal walls bear large papillae that may grade into unicellular hairs. The stomata are irregularly arranged and oriented. Proximal thickenings of subsidiary cells are lip-like or absent. The shield margin may show irregular microdenticulation (Fig. 17B). The overall cuticular structure is similar to that of the associated twigs Quadrocladus dvinensis (see Section 5.7.2). The cuticle of the main axis and of the microsporophyll stalks is very thin, showing imprints of regular files of strongly elongated cells; stomata are lacking. There are also remains of more compact (young?) microstrobili.

S. V. Meven / Reviewof Palaeobotanyand Palynology96 (1997) 351 447

389

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A Fig. 15. Leafy sterile shoots associated with Concholepis harrisff Meyen,sp. nov. (A, B) and Pseudovoltzia?cornuta Meyen,sp. nov. (C, D). (A) Spec. No. 3784/185. (B) Spec. No. 4100/58-1. (C) Spec. No. 1438/13. (D) Spec. No. 1438/3A. LocalityVym-1 (A, B) and Kityak (C, D). Scale bar=5 cm (A), 1 cm (B), 1 mm (C, D).

Localities: Aristovo (31 specimens), Isady (29 specimens), Viled ( 1 specimen), Titovo (?; 1 specimen), Kalinovka (1 specimen).

5. 7. Genus Quadrocladus M/idler, 1957 Type: Quadrocladus florinii M~idler, 1957. Diagnosis: Shoots are both dwarf and long, or only long. Leaves helically arranged, round (unifacial), oval or rhomboid (and then bifacial) in cross section, linear, spatulate or subtriangular in outline, with obtuse or acuminate apex, occasionally with a short mucro. Stomata arranged throughout both surfaces, sometimes forming short files 3 to 7 (more often 4-5) subsidiary cells similar to other epidermal cells, sometimes more cutinized, may bear thickenings or papillae faced towards stomatal pit. Guard cells sunken. Leaf margin may bear

microscopic teeth represented by cuticular papillae or unicellular hairs. Specific composition: Quadrocladus solmsii (Gothan et Nagalhard) Schweitzer (Q. florinii M~idler is regarded as a younger synonym of the species) and Q. orobiformis (Schlotheim) Schweitzer from the Zechstein of Western Europe (Schweitzer, 1960; Ullrich, 1964); Q. (al. Steirophyllum) komiensis Meyen, sp. nov., Q. dvinensis Meyen, Q. borealis Meyen and Q. schweitzeri Meyen (the present paper); Q. sibiricus (Neuburg) Meyen from the Permo-Triassic volcanics of the Tunguska Basin and the Maltsevskaya Series of the Kuznetsk Basin (Meyen, 1981); Quadrocladus (or Steirophyllum) may also comprise some conifers from the Zone C of Nan-Shan (Durante, 1980). A synopsis of characters of the above species is given in Table 1.

390

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Rcviel~ ~?/ Palaeol,otan.l' aml Pall'n dogy 96 ( 1997; 351 447

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Fig. 16. Sashinia arismvensis Meyen A El, S. horealis Meyen (l: l, K S) and Quadrocludus horealis Meyen (J). (A) Transferpreparation of holotype No. 4552/296-1 as seen from both sides. I B) The same, apex of one of seed stalks with a seed under apical pad. (C) The same, reconstruction. (DI Detached seed stalk as seen l¥om both sides before maceration, spec. No. 3782/10-l. (E) The same, successive stages of maceration causing increase of the compression size, black dot fungus; alter outfolding of the apex a seed scar contour (at arrow to righl on the outfolded part of cuticle) was exposed. ( F ) General view of holotype No. 3774/I-8 before

S. V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351 447

39l

Table 1 Synopsis of characters of Quadrocladus and Steirophyllum ~._o

2x6

Ouadrocladus solmsii

1:3

1.1-2 x 7

Q. orobiformis

1:3,5

Q. dvinensis

1-2(2.5) x 6 1:3

Q. borea/is

0.6-1.3 x 5 1:4

Q. schweitzeri

1-2x 5-11

(~ sibiricus

0.8--1.5 x 12

0.5-1.5 x 3-6 1:4-6

Q. (al. Steirophyllurn) komiensis

$teirophy/lum lanceo/atum

3 x 10

Legend: (+) - occasional

presence of a character

{ ( + ) ) - very rare presence of a character

transferring. (G) Transfer-preparation of counterpart of apical part of the holotype [crosses on (F) and (G) show corresponding point]. (H) Transfer-preparation of the holotype. (I) Part of counterpart of the holotype [see (F) and (H) at arrow], axillary complex, seed stalk with preserved compression (at arrow). (J) Holotype No. 3774/1-1, vegetative shoot depicted side by side with (F) in the same position as exposed on the fracture plane. (K) same as in (I), magnified apex of the seed stalk after removal of compression, contour of ovule at arrow. (L) The same, scheme of splitting the specimen in longitudinal direction, compression in black, matrix stippled. (M) The same, reconstruction, ovule covered by compressed apical pad. (N-S) Cuticle of holotype. (N) Cuticle of apical pad preparation No. 3774/1-8e( 1), point 1. (O) Cuticle of the seed stalk below apical pad, preparation No. 3774/1-8c. (P, Q) Cuticle of opposite sides of vegetative leaf in basal part of axillary complex, preparation No. 3774/1-8h. (R, S) cuticle of the opposite side of bract, preparation No. 3774/1-8. Localities Aristovo (A, B), Titovo (D, E), Viled (F-S). Scale bar= 1 cm (A, F-H, J), 1 mm (B, D, E, K), 100 ~tm (N-S).

392

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S. ~ Meyen / Review ~4fPalaeohotany and Palynology 96 (1997) 351-447

Comparisons of Quadrocladus with other genera given together with the description of Steirophyllum and in sections Sections 5.4 and 8 (see also: Gomankov and Meyen, 1986).

is

5. 7.1. Quadrocladus(al. Steirophyllum) komiensis Meyen, sp. nov. (Plate VIII, 93-97; Figs. 18, 19A-E)

Holotype: Specimens No. 3742/21-2 and counterpart 21A-2 (Plate VIII, 93, 95-97; Fig. 18A, B, Fig. 19B-E). Type locality: Sluda. Etymology: After Komi ASSR. Diagnosis: Leaves varying in length and angle of attachment, up to 6 mm long and 1.5 mm wide, with obtuse or acuminate apex. Epidermis uniform on both leaf surfaces. Epidermal cells in longitudinal rows. Stomata irregularly arranged, longitudinally or obliquely orientated, less frequent along the central part of leaf. Guard cells sunken. 4 (5 6) subsidiary cells with thickened proximal walls or with proximal papillae, distal walls form a round or (less frequently) irregular contour to stomatal apparatus. Leaf margin with cuticular teeth. Cuticle thickened in leaf apex. Description: The type locality yielded several

393

isolated leaves and three leafy shoots probably representing abscissed small branchlets of the last order. Both length and width of leaves as well as angles of their attachment vary, even within one specimen. Some of the leaves are linear whereas others are short and tongue-shaped. The lower (?) cuticle is thicker and better preserved, showing less frequent stomata. Otherwise, both sides of the leaf are of the same cuticular structure. Stomata are not numerous, less frequent along the leaf axis (Plate VIII, 96). Their subsidiary cells are not distinguished by a stronger cutinization (Plate VIII, 93, 95). Marginal teeth are large and devoid of an internal cavity; they are inclined towards the leaf base or apex. Corner spines and pores in sutures above radial walls (Fig. 19C) may occur. One specimen shows a group of uniform quasidisaccate pollen of the Platysaccus type stuck to the cuticle (Plate VIII, 97). This pollen might have belonged to this conifer. Comparison: The present species may be referred either to Quadrocladus or Steirophyllum (see Section 5.8). The selection of the name Quadrocladus is quite conventional and has been motivated by the following reasons. In Steirophyllum so far we do not know if marginal microdenticulation, strong variations in leaf shape

PLATE VIII Dvinostrobus sagittalis Gomankov et Meyen. Aristovo. Two sporangial groups on transfer preparation of the holotype; stereopair. Spec. No. 4552/494-1. x 10. 90-92. Taxodiella bardaeana (Zalessky) Meyen, comb. nov. Chekarda-l. Mucro at the leaf apex. Preparation No. 3737/64A, point 4. × 50. 90. Cuticle of non-stomatal leaf portion. Same preparation, point 6. × 300. 91. Stomatal band (to right), stuck cuticle of leaf opposite sides. Preparation No. 3737/56, point 7. × 300. 92. 93 96. Quadrocladus (al. Steirophyllum) komiensis Meyen, sp. nov. Sluda. Holotype. Stoma on the side with the thinner cuticle. Preparation No. 3742/21-2A, point 3. × 300. 93. Cuticle of apical portion of the leaf. Preparation No. 3742/21A-6, point 1. x 50. 94. Holotype. Stomata on side with the thicker cuticle (= Fig. 19E). Preparation No. 3742/21-2A, point 3. x 300. 95. Cuticle of the same leaf side, at top left marginal teeth. The same point. × 50. 96. Pollen Platysaccus and Protohaploxypinus (at top left) stuck to the thinner cuticle of the leaf. Same preparation, point 1. × 280. 97. 98-102. Pseudovoltzia? cornuta Meyen, sp. nov. Kityak. Detached axillary complex (= Fig. 29B). Spec. No. 1438/6. × 5. 98. Ovule in attachment (?) to axillary complex (see 101). Spec. No. 1438/23. × 5. 99. Proliferated (?) compound polysperm. Spec. No. 1438/9. × 2. 100. Compound polysperm with attached (?) ovule (in upper left corner; see 99), and subtending leafy twig. Spec. No. 1438/23. × 2. 101. Associated vegetative shoots. Spec. No. 1438/4. × 1. 102.

S. 1~ .llcy~,H Rcvi~'wol Palaeohofanv am/Pa/vnolo~y 96 (1997) 35l 447

394

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Fig. 17. Dvino~stro/~t~.~~a~#ta/i,~ Gomankov ct Meson. I A! Holotspc No. 4552 494-1 belkwe transferring. (B) Holotype, margin or" distal shield with microdenticulation, preparation No. 4552 494-1d, point 3. I(') Hololypc, sporangia and sporangiophores on transfer preparation. (D) Pollen of Scuta.v)oritcs type from sporangium, preparation No. 455Z164-5a, point 1. (E) Dispersed pollen of the same type, preparation No. 4552371-4-51, Locality Aristow~ Scale bar-5 mm (A), 1 mm (C), 1(10l,ma (B). 50 ~tm (D. E).

and size, and stomata with larger n u m b e r of subsidiary cells may occur. The new species is similar to other species of Qua~b'ocladus in having irregularly arranged stomata, the presence of marginal teeth, a n d a u n i f o r m epidermis on both leaf surfaces. It differs fi'om Q. solmsii, Q. orohil~rmi.s a n d Q. sibiricus in having more flattened leaves and well developed marginal teeth, from Q. dvi-

m'nsi~' in having more flattened leaves and better developed papillae on the subsidiary cells as well as the larger n u m b e r of stomata with 6 subsidiaries. from Q. horeali~" in the m u c h better developed marginal teeth. The upper Tatarian and PermoTriassic species of Quadrocladus in Western Angaraland and Siberia associate with Lueckisporites a n d Scutasporites pollen (Meyen,

395

S. I~ Meven / Review ~['Palaeobotany and Palynology 96 (1997) 351 447

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Fig. 18. Quadrocladus (al. Steirophyllum) komiensis Meyen, sp. nov. (A) Holotype No. 3742/21A-2. (B) counterpart of A, spec. No. 3742/21-2. (C) Spec. No. 3742/21-6. (D) Spec. No. 3742/12-1. Locality Sluda. Scale bar= 1 mm.

1981), which is absent in Sluda (and in the Kazanian in general ). Obviously plants with Q. komiensis produced different pollen, perhaps of the Platysaccus type. Locality: Sluda.

5. 7.2. Quadrocladus dvinensis Meyen, 1978 (Plate VII, 87; Figs. 20, 21 )

Holotype: Specimens No. 4552/277-1 and counterpart 277a-1 (Plate VII, 87; Fig. 20A G); Aristovo. 1976 1978 Quadrocladus dvinensis Meyen, pp. 306 308, pl. 2, figs. 13, 14; text-figs. 14-19. 1986 Quadrocladus dvinsensis Gomankov and Meyen, pl. 16, fig. 7; text-figs. 72, 73.

Description (abbreviated, for full description see: Gomankov and Meyen, 1986): Shoots of the last

order are branched at acute angles. Long shoots bear dwarf ones (PlateVII, 87) in the axils of some leaves. Leaves are strongly decurrent, nearly circular in cross section, upper surface flattened, narrower than the lower one which is strongly convex (Fig. 21). Marginal teeth up to 200/am long, variously orientated, irregularly spaced, sometimes solitary. Leaves of long shoots tongueshaped, with an obtuse or acuminate apex; those of dwarf shoots short, with convex margins. Judging from scars on the leaf cushions, leaves abscissed with the aid of an abscission layer. The degree of the order in arrangement of epidermal cells is varying, sometimes they form rather regular files. Corner spines are common. Leaves are amphistomatic, stomata are irregularly orientated, less frequently arranged into short files. Stomata

S. [. 3,h'ven Revie~ o/ Palaeohotan) am/ t~ah'mdo~y 96 (1997) 351 447

396

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Fig. 19. Quadrocladus (al. Steiroph.vlh#~l) komicnsis Meyen, ,V~.m;~. ( A E ) and Gemitzia sp. SVM-2 ( F ). ( A ) Distribution of stomata (large dots) in leaf apex, preparation No. 3742 21A-6, point 2. (B) Unl\)lded cuticle along leaf margin with teeth, marginal fold of cuticle and teeth in black, preparation No. 3742 21-2A. point 2. (C) Holotype, corner spines and pores in sutures along radial walls. preparation No. 3742/21-2A. point 2. (D} Holotype, stoma with proximal papillae on subsidiary cells, preparation No. 3742/21-2A, point 2. (E) Holotype, stoma without proximal papillae, preparation No. 374 21-2A, point 3. (F) Spec. No. 3981/88a-1. Localities Sluda ( ~ E ) and Isady (F). Scale b a r - I cm (F), I mm (A). 100 Ixna (B), I() p.m (C E).

are orientated mostly longitudinally, they are monocyclic or incompletely dicyclic. The proximal cutinization of subsidiary cells is lip-like, less frequently as small papillae. Stomata with 5 (56%) and 4 (32%) subsidiary cells dominate. Comparison: Q. dvinensis differs from Q. sohnsii and Q. orobiJormis in the outline of the leaf cross section, less developed proximal papillae and predominating stomata with 5 subsidiaries (instead of 4). In Q. solmsii the marginal denticulation is

rarely observed in the basal part of leaves, and in Q. oroh([brmis it was not observed. Q. dvinensis differs from Q. sibiricus in the presence of teeth, less developed proximal papillae on subsidiary cells, absence of septated cells and the irregular branching of the shoots. Localities: Several hundred specimens from Aristovo, Titovo and Dot. Detached leaves with long teeth in Viled may represent deviating specimens of Q. horealis.

S. V. Meyen /Review q/Palaeobotany and Palynology 96 (1997) 351-447

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Fig. 20. Quadrocladus dvinensis Meyen. ( A G ) Holotype No. 4552/277a-1. (A) General view. (B) Leaf margin, preparation No. 4552/277a-1 a. (C) Epidermis, the same preparation. (D F ) Stomata, the same preparation. (G) Cuticle of the leaf margin, corner spines directed towards the margin bearing small teeth. (H) Preparation No. 3854/473-1, point 2. (I) Stoma, same point. Locality Aristovo. Scale b a r = 5 mm (A), 100 i,tm (B I).

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Fig. 21. Quadrocla&l,~ dvinensis Meyen. (A, B) Transfer-preparation of leafy shoot, spec. No. 4552,32a. ((7) Stoma, preparation No. 4552/453-a. Locality' Aristovo. Scale b a r = 3 mm (A, B), 20 pm ( ( ' )

5. 7.3. Quadrocladus borealis Meyen, 1981 ( Figs. 16I, 22, 23 )

Holotype: Specimen No. 3774/1-1 (Fig. 16I, Fig. 22 ); Viled. 1981 Quadrocladus borealis Meyen, p. 165, pl. 1, figs. 1-4; text-figs. 7, 14-16. 1986 Quadrocladus horealis Gomankov et Meyen, text-figs. 74 75. Description (abbreviated, for full description

E~

see: Gomankov and Meyen, 1986). Part of branches bear narrow (less than l mm wide) leaves attached at acute angles. Fragments of shorter tongue-shaped leaves probably belonged to dwarf shoots. Leaf apex round or feebly acuminate, leaf base decurrent on a leaf cushion (Fig. 23). An abscission layer was probably present. Leaves are triangular in cross section. The epidermis is uniform on both leaf sides. The margin is poorly distinguished in the epidermal

--

Fig. 22. Quadrocladus horealis Meyen. ( A ) Cuticle of portion of leafy shoot, opening in cuticle shows initial cross section of leaf near its base, preparation No. 3774,/7-la. (B) Cuticle of opposite leaf sides, holotype, preparation No. 3774,,'1-1. Locality Viled. Scale bar = 1 m m ( A ) , t00 I.tm (B).

S. V. Meyen / Review ~f Palaeobotany and Palynology 96 (1997) 351-447

ii

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399

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Fig. 23. Quadrocladus borealis Meyen, bark with leaf cushions (A, B), epidermal structure of leaf cushion (C-F), stomata with preserved guard cells on the leaf cuticle (G). (A) Spec. No. 3774/2a-1. (B) Leaf cushion, spec. No. 3774/2b-12. (C) Preparation No. 3774/2a-1, point 2. (D) Stoma, same preparation, point 1. (E) Stomata, preparation No. 3774/2b-12, point 1. (F) Cuticle of the margin, preparation No. 3774/2b-12, point 2. (G) Preparation No. 3774/1A-26, point 1. Locality Viled. Scale bar= 1 mm (A, B), 100 ~tm (C-G).

structure, or occasionally accompanied by minute teeth. Epidermal cells isometric. Papillae are absent, corner spines are common. Stomata are irregularly arranged, in rare cases forming short files, mostly longitudinally orientated. Stomata with 5 (56%) or 4 (30%) subsidiary cells are dominating. Comparison: Q. borealis differs from other species

in its smaller leaves; from Q. solrnsii, Q. orobiformis, Q. komiensis and Q. sibiricus in less frequently having papillae on the subsidiary cells and triangular cross section of the leaf; from Q. dvinensis in the virtual absence of marginal teeth, smaller corner spines and triangular leaf cross section. Locality: Viled (more than 30 specimens).

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5. 7.4. Quadrocla~h¢sschweitzeri Meyen, 1986 ( Fig. 24)

Holoo'pe: Specimen No. 3981/55-3 ( Fig. 24G-1 ): Isady. 1986 Quadrocludus schweitzeri Gomankov and Meyen, text-fig. 76. Description: There are fragments of branches of the last order. Leaves are dense, attached at acute

angles, less frequently diverging, 5 I I mm long, 1 2 mm wide, linear, with an acuminate apex, decurrent. The epidermis is nearly the same on both leaf surfaces, consisting of a regular system of polygonal cells poorly ordered in indistinct files in the apical part of the leaf: towards the base the cellular files are more regular consisting of rectangular cells up to 25 lain wide and up to 100 gm long. Radial walls are often indistinct, periclinal

Dt.kI

I

1-31H m

tFig. 24. Quach'ucla~h~s .~chweitzeri Meyen. I A) H olotype No. 3981 55-3. (B) Spec. No. 3981/58-1. (C) Spec. No. 3981/49a-14. (D) Spec. No. 3981/66zh-68. ( E ) Spec. No. 3981/66v- I 0. ( F ) Spec. No. 3981 64-1. (G) Cuticle of leaf side 2 of the holotype, preparation No. 3981'55-3, point 2. ( H ) Cuticle of leaf side l, the same preparalion, point I. (I) Stoma on leaf side 2, the same preparation, point 3. IJ ) Stoma on leaf side I, the same preparation, point 1. Localily Isady. Scale b a r = 1 cm ( ~ F ), 100 gm (G J 1.

S. lC Meven / Review qf Palaeobotany and Palynology 96 (1997) 351-447

walls may bear a faint middle thickening. Occasionally corner spines are present. Stomata are rare, guard cells longitudinally orientated. Subsidiary cells may be distinguished by a stronger cutinization, they bear large papillae overhanging the guard cells. 2-5 papillae are observed per stoma. The stomatal pit is occasionally bordered by a cutin ring, Leaf margins are smooth. Teeth occur only in solitary leaves having a shorter lamina of the same width. Comparison: Q. schweitzeri differs from other species in having less distinct radial walls of the epidermal cells; from other species of Western Angaraland - - in the rareness of teeth, absence (or rare occurrence) of lateral dwarf shoots; from Q. borealis - - in having much larger leaves; and from Q. sibiricus - - in less frequent stomata and absence of septae in epidermal cells. Locality: Isady (30 specimens). 5.8. Genus Steirophyllum Eichwald, 1854, emend.

1854 Steirophyllum Eichwald, p. 183. 1855 Steirophyllum Eichwald, p. 237. 1860 Steirophyllum Eichwald, p. 237. 1887 Ullmannia Goeppert, Schmalhausen, p. 21. 1927 Ullmannia Zalessky, pp. 27, 29. Type: Steirophyllum lanceolatum Eichwald, 1854. Remarks: When Eichwald (see synonymy) introduced this genus he had known about Goeppert's (1850) description of the genus Ullmannia. He considered that in Ullmannia the leaves are more swollen and shorter, less distant, and more distinctly striated and larger. Eichwald noted that S. lanceolatum comprises conifers earlier described by Fischer de Waldheim (1847) as Annularia ovata Fischer de Waldheim. Due to this, for formal reasons, the specific epithet "lanceolatum" is invalid, and a new combination Steirophyllum ovatum should be introduced. However, presently it seems better to refrain from this step, because the location of the type material of A. ovata is unknown, hence we cannot ascertain the synonymy of both species. In the first description of the type species S. lanceolatum, Eichwald (1854) mentioned two vari-

401

eties and admitted the possibility that they represent separate species S. latifolium and S. angustifolium. However, in the heading of the description only the epithet "lanceolatum" is mentioned and the two other names are used provisionally. In subsequent description of the same species (Eichwald, 1855, 1859-1860) there is no mention of these two species and their names. Therefore, for the type species the epithet "lanceolatum" can be retained. Goeppert (1864 1865) had placed S. lanceolatum and, hence, the genus Steirophyllum into the synonymy of the genus Ullmannia and the species U. bronnii Goeppert. This viewpoint had been adopted by Schmalhausen (1887) and Zalessky (1927). In subsequent literature the name Steirophyllum was mentioned only as a synonym of Ullmannia. The species U. bronnii was regarded as a very common element of the Upper Permian flora of Western Angaraland, and comprised specimens identical to S. lanceolatum. It is true, the situation regarding the type material of S. lanceolaturn is uncertain. Originals to Eichwald's monograph are kept in the Museum of Leningrad University. Some specimens coming from the same collection and figured by Zalessky (1927, pl. 9, figs. 4, 5) are kept in Tschernyshov Museum (Leningrad). Among all these presently available specimens, those indisputably figured by Eichwald (1854-1861, 1859-1860, pl. 18, figs. 6, 7) are absent. Zalessky held that the two specimens figured by him in his atlas, had been used for Eichwald's drawings, but the reasons for his opinion are unclear. The arrangement of twigs on these specimens and Eichwald's drawings is quite different. Nevertheless, we are undoubtedly dealing with conifers of the same species. Their shoots are very peculiar and can easily be recognized in collections by their short and broad leaves, often with a marginal border; the apex is obtuse (Fig. 25A-F ). Shoots of this very type in the author's collection come from the Kuzminovsky Mine belonging to the group of the Kargala Mines from where many other specimens of this species including one of the originals of Schmalhausen originate (1887, pl. 6, fig. 18). This original is presently kept in the Museum of the Kazan University and has been

402

S. 1". Mel'ell RevielL o/ Palaeohota,y am/Pa/vmdo~y 96 (1997) 351 447

referred to Ullmannia bronnii. It is indistinguishable from Eichwald's specimens. Three of the specimens present in the author's collection yielded leaf cuticle ( Fig. 24D, E, I -M ) that has proved to be quite different from that in U. hronnii (Gothan and Nagalhard, 1921 1923: M'/idler. 1957: Schweitzer, 1960; Ullrich, 1964). In S. lanceolatum, the stomata are not arranged in files, subsidiary cells are less numerous, the leaves are devoid of the characteristic striation (as was rightfully noted by Eichwald). Thus, the independent status of the species lanceolatum is now fully supported by epidermal studies. Moreover, the assignment of the species to the genus Ullmannia is doubtful. In epidermal characters, such as the absence of stomatal files and lesser number of subsidiary cells. S. laneolatum is much closer to Quadrocladus than

Ullmannia. The genus Quadrocladus initially included a single species Q../torinii Mfidler (1957). Schweitzer (1960) referred this species to Q. sohnsii (Gothan et Nagalhard) Schweitzer, and complemented this genus with the species Q. orobf/brmis (Schlotheim) Schweitzer. Both species are characterized by a small number (mostly 4) of subsidiary cells. In the Zechstein flora of Western Europe, this character well distinguishes Quadrocladus from Ulhnannia and Pseudovoltzia where the number of subsidiary cells is larger (6 9, up to 12 in L( hronnii, 5 7 in U. frumentaria, 6 8 in Pseudovoltzia lieheana). I_.. jrumentaria shows characteristically regular stomatal files, in U. bronnii stomatal files are also present, although sometimes, particularly in the leaf apex, they are less pronounced. In Quadrocladus stomatal files are absent (Q. orohf[ormis) or poorly developed (Q. solmsii). If we characterize the genus Quadrocladus according to these European species alone. Steirophyllum lanceolatum will occupy an interme-

diate position between Q. oroh([brmis" and U. bronnit in its epidermal characters, viz. the stomatal liles are absent (as in Q. orob(/brmis), and the number of subsidiary cells is 4 6, that is closer to L. hronnii. If we take into consideration also all the species of Quadrocladus described above, the demarcation between this genus and Steirophyllum becomes impossible. That is why the species komiensis was given a combined generic designation (see Section 5.7.1 ). In other words, two conclusions can be drawn: (1) the genus Steirophyllum cannot be merged with Ulhnannia: {2) the genus Qua~h'ocladus proves to be a younger heterotypic (taxonomic) synonym of

Steirophyllum. It is true that the characters distinguishing 5teirophyllum and Ullmannia may be regarded as insufficient for the independent status of Steirophyllum. In this case, merging Steirophyllum and Ullmannia, we have to merge them also with Qua~h'ocladus. But for the same reasons Ullmannia can be merged with Pseudovoltzia which is very close to U..[i'umentaria in epidermal characters. Such a decision would hardly find support among specialists in the Palaeozoic conifers. Therefore it seems better to admit the aforesaid two conclusions. Thus. according to nomenclatural rules and leaning upon the widely recognized assemblage of taxonomically significant characters, we have to restore the genus Steirophyllum, referring to it all the species hitherto belonging to Quadrocladus. On the other hand, the name Steirophyllum has not been practically in use since the time of its introduction. Curiously, unlike other genera established by Eichwald, Steirophyllum had not been included in Andrews (1970) index of generic names.

Fig. 25, Steirophyllum lanceolatum Eichwald. (A, B)Type specimens, detached leaf [(A) redrawn from Eichwald, 1855, pt. 18, fig. 7 )] and leafy shoots [(B) Eichwald, 1855, pl. 18, fig. 6, lectotype to right]. (C) Specimen No. 1439/4. (D, E) Detached leaves with naturally macerated compression, stomata shown by dots, portions with non-preserved cellular structure shaded, spec. No. 1439/1 (D) and 1439/6 (E). ( F ) Spec. No. 1439/3-2. (G) Shoot apex, spec. No. 1439/3-3. ( l ) Stoma with large proximal papillae on subsidiary cells, preparation No. 1439/2-A, point I [the stoma at two arrows on (J)]. (J) Cuticle of the middle leaf part, the same point. ( K ) Corner spines, preparation No. 1439/2-A, point 3. (L) Pores in sutures, preparation No. 1439/2-A, point 4. ( M ) Stoma with 4 subsidiary cells devoid of proximal papillae, the same point. ( N ) Stoma wilh small proximal papillae [at one arrow on (J)]. Locality Kuzminovsky Mine, Scale bar= l cm ( B, C, H ), I m m ( D G). 1001_tin(J). 1 0 p m ( l . K N).

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S. I'. Mew't* Review ql Pa&eoholanv am/PalynoloKl' 96 (1997) 351 447

Therefore the name Quach'o('ladus can be conserved. Before making final taxonomic and nomenclatural decisions, the following reasons are to be considered. The independent status of Steirophyllum and Ullmannia rests on the following assumptions: ( 1) genus Ullmannia is not collective, i.e., the species E,~ bronnii and U../i'umentaria belong to the same natural genus; (2) species U. hronnii is not collective, i.e., the epidermally studied specimens belong to the same species as all remaining ones: (3) all the species of Quadrocladus belong to a single natural genus, i.e.. associated with largely uniform fructifications and pollen. All the three assumptions, however, are not well founded as indicated by the following circumstantial evidence. Both made and female fructifications, that arc traditionally ascribed to Ulhnanniu. are known only in ~( Jfumentaria, not in the type species /_". hronnii. The species U. Ji'umenlaria is well studied. This species has characteristic and fairly unilk)rm leafy twigs. The pollen obtained from microstrobili of b\ fi'umentaria is operculate, of the Jugasporites type (Foster, 1983), widely spread in deposits yielding leafy twigs. All the twigs yielding cuticle show the same epidermal characters; thus, this is a "'good" species. LL hronnii is quite another matter. This species comprises all Zechstein shoots with short fleshy leaves, but in outward appearance the species do not look homogeneous. The heterogeneity of U. bronnii is striking in the protologue (Goeppert, 1850, pl. 20, figs. 1 26: see also Goeppert, 1864 1865, pl. 45, figs. 1 26). Some of the leafy shoots bear large leaves with an acuminate apex and longitudinal striation, other shoots bear quite different leaves more convex and with round apices. The leaves of the second type are more similar to Steirophyllum lanceolatum than those of the first type. In subsequent papers the name U. bronnii was ascribed to essentially different shoots. The persistence of epidermal characters in specimens referred to U. hronnii is not evident as well. Gothan and Nagalhard (1921 1923, pl. 5, fig. 2) depict regular stomatal files. In photographs presented by Schweitzer (1960, pl. 8, figs. l, 2) and Ullrich (1964, pl. 13, figs. 1-3) the files are much less distinct. The

persistence of other characters cannot be judged fi-om available descriptions and illustrations. A clear understanding both of the bulk and diagnostic characters of the species is prevented by the lack of knowledge of epidermal characters of the type material. From available publications one cannot comprehend how representative the epidermal studies have been. Moreover, magnified photographs of specimens yielding cuticular preparations have not been published. Of the utmost importance is that a lectotype of the species appears has not been selected. The selection of a lectotype is badly needed, because Goeppert referred to U. hronnii, apart from coniferalean leafy twigs, also Peltaspermum-like fructifications (Goeppert, 1864 1865, pl. 45, figs. 24, 25) probably belonging to associated fronds of Lepidopteris martinsii. These fructifications were described in the subsequent literature as Strobilites hronnii (Florin, 1944; Stoneley, 1958). At last, the very nomenclatural validity of the epithet hronnii is not evident. Both Goeppert and subsequent authors merged U. bronnil with the earlier described species Cupressites ulh77annii Bronn, the epithet of which has a priority. The variation of U. hronnii shoots known to Goeppert too, was treated as intraspecific. However, suspicion arises that this is not the case, and that under the binomial U. bronnii essentially different plants have been described. This suspicion is supported, apart from the above mentioned outward habit of shoots, also by the following indirect considerations. As was said above. U. fi'umentaria is accompanied by the pollen Jugasporites. Visscher (1971, p. 34) suggested that ~. hronnii is associated with Lueckisporites pollen, because (a) relative frequency of both taxa coincides particularly when they are dominating, (b) Lueckisporites and Jugasporites are similar in the presence " o f a sexine free area on the proximal face of the corpus". Teratological forms of Jugaal~orites may even approach the taeniate condition of Lueckisporites. Visscher noted a contradiction to Florin's (1944) statement that U. hronnii produced non-taeniate pollen, but considered that the strobilus yielding the pollen had not been found in organic connection to the leafy twig. The latter consideration is incorrect, however, because

S. ~ Meyen / Review of Palaeobotany and Palynology 96 (1997) 351 447

the pertinent photograph provided by Florin (1944, pl. 169/170, fig. 10) shows the attachment of the strobilus. Visscher's conclusion on the association between U. bronnii and Lueckisporites was argued by the author (Meyen, 1981), because in the PermoTriassic of Siberia the association between Lueckisporites and Quadrocladus sibiricus is evident. This pollen is very common in the upper Tatarian of Western Angaraland, where shoots comparable to U. bronnii are absent. Therefore, the author suggested that in Western Europe, as in Siberia, Lueckisporites associates with Quadrocladus. This admission, however, contradicts a drastic discrepancy between occurrences of both kinds of remains in Europe, where Lueckisporites dominates among miospores, whereas Quadrocladus shoots are rare. Visscher's observations and data on Siberia and Western Angaraland would be mutually consistent, if the collective nature of U. bronnii is admitted. Probably among specimens currently referred to U. bronnii, there are (1) conifers closely related to U. frumentaria and having nearly the same cuticle and pollen, and (2) conifers more related to Quadrocladus, i.e., having less pronounced stomatal files, fewer number of subsidiary cells in stomata, and producing Leuckisporites pollen. In any case, the insufficient knowledge of the species U. bronnii and of its relations to U. frumentaria and Quadrocladus (considering all its species, and not only two Zechstein ones) prevents any final decision regarding the synonymy of Quadrocladus and Steirophyllum. Presently S. lanceolatum occupies an intermediate position between U. bronnii and Quadrocladus. Therefore, for the time being the following preliminary solution can be adopted: (1) the independent status of the species lanceolatum is restored; (2) this species is retained in the genus Steirophyllum. It is desirable to attach more natural circumscription to the genera Ullmannia, Steirophyllum and Quadrocladus and not to place them among form-genera selected by Harris for coniferalean vegetative shoots (see Section 8). In this case the circumscription of these genera is to be made with due attention to associated fructification and pollen. None are presently known for the type species of all the three genera.

405

The typification of both U. bronnii and Q. solmsii should be also made. These issues, however, are beyond the scope of the present paper.

5.8.1. Steirophyllum lanceolatum Eichwald, 1854, emend. (Fig. 25)

LectoO,pe: One of the specimens figured by Eichwald (see synonymy) on pl. 18, fig. 6; see Fig. 25B in the present paper. 1854-1861 Steirophyllum lanceolatum Eichwald, p. 183, pl. 18, figs. 6, 7. 1855 Steirophyllum lanceolatum Eichwald. 1859-1860 Steirophyllum lanceolatum Eichwald, p. 237, pl. 18, figs. 6, 7. 1887 Ullmannia bronnii Goeppert, Schmalhausen, p. 21, pl. 6, fig. 18. 1927 Ullmannia bronnii Goeppert, Zalessky, pp. 22, 32, 39, 49, pl. 9, figs. 4, 5; pl. 36, figs. 1 (?), 2.

Diagnosis: Branches of last order up to 7 cm long and up to 2 cm wide. Leaves helically arranged, attached at acute or right angles, up to 10 mm long and up to 3 mm wide, with an obtuse apex. Margins nearly parallel. Leaf base decurrent. In apical part of the shoot, leaves are curved inward. Epidermis of both leaf sides identical, composed of polygonal cells with rounded corners. Cellular files absent. Stomata arranged and orientated irregularly. Subsidiary cells 4-8, more often 4-6. Periclinal walls of subsidiary cells with radial wrinkling. Many stomata show proximal papillae of varying outlines on subsidiary cells. Corner spines may be present. Description: The author's collection contains 9 fragments of leafy shoots and 2 detached leaves. The author also has seen originals of Zalessky, specimens studied by Eichwald, and one original of Schmalhausen (see synonymy), as well as specimens in the collection of N.K. Esaulova coming from the locality Kzyl-Bairak and kept in Kazan University. The latter specimens have not been properly studied and are not considered in the present description. As a whole, the gross morphology of both the shoots and leaves is fairly uniform. Although the leaf sizes and angles of attachment are varying,

41)6

X 1] Mevcn ,' Review ol Pakwobotato' and Palvm)h*gy 96 (1997) 351 447

the overall habit of the shoots is persistent. Characteristic are prominent folds on the leaves that suggest nearly round cross section of the leaf lamina. The leaf margin often shows a wide flattened border ( Fig. 25C, F ) also seen in Eichwald's drawings (Fig. 25A, B). In one of the shoots ( Fig. 25C ) a seed occurs among leaves, but this is probably an accidental co-occurrence. This species probably also includes a shoot apex with smaller falcate leaves (Fig. 24G). The original of Schmalhausen (see synonymy) may give an impression of a fertile shoot, because coarse folds in leaf apices form round-triangular contours simulating seed scars. This specimen is peculiar in leaf attachment at varying angles as properly shown on Schmalhausen's drawings. Compression suitable for maceration is preserved on a single specimen (Fig. 25H) having somewhat smaller and narrower leaves. The same epidermal structure is well seen under the reflected light on the detached leaves having more typical appearance (Fig. 25D, E). Therefore, the specimen which yielded the cuticular preparations can be ascribed to the present species without doubt. The compression is broken into small pieces, and the general topography of large portions of the epidermis cannot be observed. However, all the fragments are of the same structure, hence there are no reasons to consider that the epidermal structure of the opposite leaf sides was different. The stomata are monocyclic, less frequently with one or two encircling cells indistinguishable from surrounding epidermal cells. The stomata are strikingly variable both in the number of subsidiary cells and the degree of the development of their proximal papillae (Fig. 25I, J, N) that can be completely absent ( Fig. 25M ). The subsidiary cells are well distinguished among epidermal cells by their fine wrinkling usually directed towards the cell center. Due to corrosion of the cuticle, the number of subsidiaries was counted only in 20 stomata, of them 5 stomata have 4 subsidiaries (25%), 8 stomata have 5 subsidiaries (40%), 5 stomata have 6 subsidiaries (25%); one stomata was found with 7, and one with 8 subsidiaries (5% each). Corner spines are varying in development. In places they are large and occur at each angle (Fig. 25K). Sutures above radial walls may be

pierced by irregularly arranged pores (Fig. 25L). Marginal teeth have not been recorded. Comparison: The similarity between S. lanceolatum and Quadrocladus has already been mentioned. Relations between S. lanceolatum and different species of Quadrocladus can be seen from Table 1. S. lanceolatum differs from the Subangara species of Quadrocladus in the much lesser polymorphism of the foliage. Shoots of S. lanceolatum are very similar to specimens described by M~idler (1957, pl. 9, figs. 6, 7) as Q. florinii, although in the latter stomata with 4 subsidiary cells lacking wrinkling and having periclinal walls more strongly cutinized than other epidermal predominate. Both the leaf outline and the epidermis topography in the two species are identical. Localio': Kuzminovsky Mine.

5.9. Genus Geinitzia Endlicher 1847

l)'pe: Geinitzia cretacea Endlicher, 1847. Remarks: The genus Geinitzia is here understood following Harris (1969, 1979) as one of specially selected genera for coniferalean vegetative shoots (see Section 8). Harris (1969, p. 249) noted, that the type species G. cretacea comprises the Cretaceous shoots earlier described as Sedites rabenhorstii Geinitz. M.P. Doludenko (pets. commun.), studying shoots of this type from the Cretaceous of the Ukraine, considers that the selection of the genus by Harris is not quite good, because the type material of Geinitzia will probably be connected with certain fructifications that may allow, together with anatomical and epidermal characters of vegetative shoots, the assignment of the genus to the Taxodiaceae. If we treat the genus Geinitzia as a form-genus, the establishment of the natural affinity of the type species does not involve the same affinity of other species of the same genus. Nevertheless, in the system of form-genera used for vegetative shoots, the genus Geinitzia would be better replaced by another genus of less certain natural affinity particularly concerning the type species. Diagnosis (alter Harris, 1979, with changes): Vegetative shoots. Phyllotaxis helical, less frequently with opposed leaves, and then leaves are

S. I4. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

arranged in one plane. Leaves elongated (length to width ratio 5:1 or less), dorsiventrally flattened, diverging from stem. Leaf with a single vein. Leaf base may be petiolate.

5. 9.1. Geinitzia subangarica Meyen, 1986 (Fig. 26) Holotype: Specimen No. 3765/95-3 (Fig. 26B, K-M ); Luptyug. 1986 Geinitzia subangarica Meyen, Gomankov and Meyen, text-fig. 77. Description (abbreviated, for full description see Gomankov and Meyen, 1986): The leaves are linear or spatulate, up to 12mm long and 1-3.5mm wide, lens-shaped in cross section, decurrent. The leaf base is slightly attenuated, with an abscission layer. The apex is semicircular or acuminate. The margin is smooth. Two resin ducts run through nearly the full leaf length. Radial walls of the epidermal cells are indistinctly seen on the leaf cuticle. Files of the epidermal cells are seen only in more elongated leaves. Leaves are hypostomatic. The stomata are not arranged in files and bands, longitudinally orientated, incompletely dicyclic, stomatal pits narrow. Stomata are bracketed by lateral encircling cells. Subsidiary cells bear proximal swellings. Sometimes the guard cells are nearly surficial. Some of the epidermal cells have dark resin-like contents. Comparison: G. subangarica resembles Jurassic conifers described by Florin (1958) as Camptophyllum schimperi Nathorst, but differs in the absence of stomatal bands. It differs from Quadrocladus in its hypostomatic leaves, elongated stomatal pits and the presence of resin ducts. The systematic affinity of the species is unknown. It occurs together with microstrobili resembling Dvinostrobus; other conifers in the same locality being absent. Locality: Luptyug (15 specimens, including 3 leafy shoots). 5.9.2. Geinitzia sp. S V M - 1, S V M - 2 (Plate VII, 88; Figs. 19F, 27) Description (see also Gomankov and Meyen, 1986, pl. 16, fig. 6; text-fig. 78). There are leafy shoots

407

up to 5 cm long. The leaves are linear, up to 25 mm long and 1.5-2 mm wide, with an acuminate apex. The cuticle is very thin and badly preserved so that the outlines both of the epidermal cells and stomata are indistinct. The guard cells feebly sunken, longitudinally orientated, surrounded by 5-6 (?) subsidiary cells. One specimen shows leaf margins with long teeth. The above description concerns conifers designated as Geinitzia sp. SVM-1. The same species may comprise shoots coming from Isady. One of them is shown in Fig. 19F. Their gross morphology is the same, but the epidermal structure is unknown. Therefore, they are designated as Geinitzia sp. SVM-2. Comparison: The conifers described above are externally similar to shoots figured in the literature under the names Ullmannia (al. Cupressites) biarmica Eichwald and Walchia foliosa (Eichwald, 1854, 1859; Schmalhausen, 1887; Zalessky, 1927). These generic affiliations are very doubtful, and the diagnostic characters of both taxa as well as their relations to the described conifers are unclear. Localities: Aristovo (4 specimens), Viled (2 specimens), Isady (numerous specimens without compressions).

5.10. Genus TaxodieUa Zalessky, 1939 Basionym: Taxodiella recticaulis Zalessky, 1939, p. 367. Remarks: Both the generic name Taxodiella and the proposed combination Taxodiella bardaeana (Zalessky) Meyen, comb. nov. are used below without full confidence in the adopted taxonomic and nomenclatural decisions. The Kungurian localities Chekarda-1, Krutaya Katushka and Krasnaya Glinka yielded numerous coniferalean shoots with small leaves. In outward appearance they are rather uniform and, as will be shown below, may well belong to a single species. Similar shoots were described by Zalessky as Walchia bardaeana Zalessky, W. peremiana Zalessky, I,t4. uralica Zalessky (Zalessky, 1937), 14(. densa and Taxodiella recticaulis Zalessky (Zalessky, 1939). As is usual in Zalessky's works, these taxa had

408

S, l,~ Meyen / Review qI'Palaeobotany and Palynology 96 (1997) 351 447

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S.V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

been given very short descriptions neither with explanations of differences between the taxa, nor with differences between them and earlier described genera and species of conifers. Obviously, the generic name Walchia, no matter W. piniformis or W. filiciformis is its nomenclatural type (see Section 8), is unsuitable for the present conifers. These may be referred to one of the form-genera in the system of coniferalean vegetative shoots as proposed by Harris (see Sections 5.9 and 8). Such a decision would mean a rejection of the already established genus Taxodiella. In the meantime, the preservation of this genus is desirable because the described conifers most likely belong to a separate natural genus. As in Kungurodendron, Concholepis, Ernestiodendron and some other Palaeozoic conifers, the leaf sizes in Taxodiella undergo significant changes with transition from one branching order to another. In Taxodiella, branches of three successive orders are known, and accordingly, three dimensional classes of leaves are displayed by a single branching twig. Therefore, a discrimination of separate species on the basis of the leaf sizes alone in these conifers is impossible. The degree of leaf divergence from the axis may also vary within one specimen. Considering variations of these and other characters in the available material (Fig. 28G-Q), one cannot adopt those characters that distinguish the enumerated Zalessky's taxa as reliable. If we merge them, however, a selection of the specific epithet should be made. Obviously it should be selected among bardaeana, uralica or peremiana as published earlier (Zalessky, 1937). The situation with W. uralica is unclear, because the magnification under which the holotype (the only figured specimen) is depicted is unknown. In the caption to his fig. 36, Zalessky had written "1:1", that corresponds to dimensions of the specimen mentioned in the description. In this case, however, leaves on the figure prove to be twice as big as noted in the description. Probably the drawing had been enlarged twice, and the dimensions of the entire specimen had been measured by Zalessky according to the drawing, and those of the leaves according to the specimen. Due to this uncertainty the epithet uralica cannot be used safely. Of the holotypes of W. bardaeana and W.

409

peremiana the former is completer and its identification with the available material is more reliable. Thus, the epithet bardaeana can be adopted. As the specimen Taxodiella recticaulis, being published in 1939, is treated as a younger synonym, the combination Taxodiella bardaeana (Zalessky) Meyen, comb. nov. is proposed (see its basionym in the synonymy of the species). Taxodiella differs from Walchia (al. Lebachia) in having simple, not forked (of Gomphostrobus type) leaves of the shoots of the penultimate order, and from Ernestiodendron--in a less regular branching and a number of epidermal characters (fewer number of subsidiary cells, their partially transverse orientation, absence of papilligers).

5.10.1. TaxodieUa bardaeana ( Zalessky) Meyen, comb. nov. (Plate VIII, 90-92; Fig. 28) Basionym: Walchia bardaeana Zalessky, 1937, Probl. Paleontol., 5: 75, fig. 39 (holotype, the only figured specimen). Synonyms: 1937 Walchia peremiana Zalessky, p. 74, fig. 40. 1937 ?Walchia uralica Zalessky, p. 71, fig. 36. 1939 Walchia densa Zalessky, p. 363, figs. 42, 43. 1939 Taxodiella recticaulis Zalessky, p. 367, figs. 48, 49. Emended diagnosis: Leafy twigs of at least three orders of branching producing leaves of corresponding dimension classes (smallest leaves were produced by branches of the last order). Branching axillary, from distinctly pinnate to poorly ordered, plagiotropic (?). Leaves needle-shaped, flattened, with slightly convex margins, up to 6 mm long and up to 2 mm wide, variously diverging from axes, occasionally adpressed to them. Leaf apex acuminate, mucronate. Margins bear microscopic cuticular teeth. Mesophyll often shows rod-shaped bodies with resin-like contents. Stomata arranged in bands (?), subsidiary cells with proximal papillae, guard cells often orientated at right angle to leaf axis. Description: There are 14 shoots of different dimensions and varying degree of branching (Fig. 27G-Q). The most complete specimen is figured in Fig. 27G-J. The preserved part of its

410

S. li~ Meyen / Review ~dPalaeobotany and Palvnology 96 (1997) 351 447

Fig. 27. Geinitzia sp. SVM-1. (A) Specimen No. 37745-1 before transferring. (B) Transfer-preparation of the same specimen. (C~ Cuticle, preparation No. 3774/5-1+ point 1. Locality Viled. Scale bar l cm (A, B)+ 100 p.m (C).

main axis is nearly 16 cm long. The lower part of the main axis preserved solitary leaves and small branchlets. The absence of larger branches in this part of the specimen is hardly related to preservation. The preserved branches of the penultimate order are progressively shorter towards the twig base. A remarkable feature of this twig is an irregular alternation of branches of different length on the left side of the main axis, branches on the right side are shorter, but also different in length and disordered. The longest branches attached on the left side at the top bear irregularly arranged small branchlets of the last order (Fig. 28I ). Other

specimens (Fig. 28N, P, O) also show disordered branching and small branchlets of the last order. The axillary branching is well seen (Fig. 28M, O). The surface of axes is canaliculate (Fig. 28I). The leaf outline is difficult to judge due to flattening of leaves in various planes. When the leaves are spread in the fracture plane, one can see that they are lanceolate, with convex or nearly parallel margins (Fig. 28J, O). The apex is acuminate bearing a short mucro (Plate VIII, 90). The dependence of leaf sizes from the order of branching is very distinct (Fig. 28G-I, K, M-P). The larger the leaves, the stronger is their divergence from the

Fig. 28. Taxodiella bardaeana (Zalessky) Meyen+ comb. nov. (A F) Specimens described by Zalessky under the name Walchia densa Zalessky, 1939, figs. 42, 43 (A, B), 14( hardaeana Zalessky, 1937, fig. 39, holotype (C), Taxodiella recticaulis Zalessky, 1939, figs. 49 (D) and 48 (E), /,E peremiana Zalessky, 1937, fig. 40 (F). (G) Spec. No. 2009/44 (collected by H.T. Mauer). (H) Same specimen, detail of place at arrow. (I) Same specimen, detail of place near cross. (J) Same specimen, detail of branch of penultimate order. (K) Spec. No. 3737/56. (M) Spec. No. 3737/59, two axillary shoots of last order, subtending leaves stippled. (N) Same specimen, general view. (O) Spec. No. 3737/70. (P) Spec, No. 4560/2 (collected by Yu.M. Zalessky). (Q) Spec. No. 3737/64. Localities Chekarda-1 (A, B, L-O, Q), Krasnaya Glinka (C+ K)+ Krutaya Katushka (D-J, P). Scale b a r = l cm (AG+ K, L, N, P)+ 1 mm (H J, M, O, Q).

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412

S. 1~ Meyen / Review q/Palaeobotany and Palynology 96 (1997) 351 447

axis. In naturally macerated leaf compression, dark rod-shaped or round bodies are observed. They are up to 500 ~tm long and up to 100 ~tm wide. During maceration their resin-like contents are dissolved, but their outlines remain on the cuticle. The compression of most specimens underwent natural maceration in varying degree. The cuticle of the opposite leaf sides is stuck and cannot be separated. Corrosion of the compression is very strong, and the mesophyll cannot be removed from the cuticle. Only very small fragments of cuticle show epidermal structure. In some fragments (Plate VIII, 91) one can only see rectangular cells in more or less regular files; stomata are absent. Other fragments do not allow tracing of the cellular structure due to corrosion, superposition of cuticles belonging to different leaf surfaces, and adhering mesophyll, but groups of papillae allow recognition of stomata (Plate VIII, 92). They seem to be arranged into files, at least in places. Sometimes transverse orientation of stomata is seen. Subsidiary cells are not numerous (5 6). Some fragments show the leaf margin with heavily corroded cuticular teeth. Comparison: Specimens described above as well as those figured by Zalessky (see synonymy) differ significantly, but this difference can be treated as intraspecific because it may be seen within a single specimen. For instance, Zalessky referred to Walchia densa shoots having smaller leaves than in W. bardaeana. However, one of his originals (Fig. 28A) shows leaves of two sizes, larger leaves being of the same dimension as in W. bardaeana. Small branchlets of the last order are figured by Zalessky in W. densa, W. bardaeana, W. uralica and Taxodiella recticaulis. The same branchlets are observed in the author's collection. The peculiar nature of this character allows it to be treated as a species marker. Probably this is a structure preceding the dwarf shoots of younger conifers. The shoot of Taxodiella recticaulis reproduced in Fig. 28E deviates from all the species placed into the synonymy of T. bardaeana. The major part of its main axis does not bear branches and leaves. However, the specimen in Fig. 28G bears in its lower part only rare branches and leaves. Therefore, T. recticaulis may be regarded as an

extreme member in the variation series of this character. Localities: Krasnaya Glinka (specimen identified by Zalessky as W. densa; holotype of W. bardaeana); Krutaya Katushka (7 specimens; holotypes of Taxodiella recticaulis and W. peremiana), Chekarda-1 (6 specimens; holotype of W. uralica, and syntypes of W. densa). All the type material of Zatessky's taxa has been lost. 5.11. Genus Pseudovoltzia Florin, 1927

5.11.1. Pseudovoltzia? cornuta Meyen, sp. nov. (Plate VIII, 98-102; Figs. 15C,D, 29A,B, 30F,G) Holotype: Specimen No. 1438/15 (Plate VIII, 99; Fig. 29A). 1887 ?Ullmannia sp., Schmalhausen, pl. 6, figs. 20, 21; pl. 7, figs. 11-13 (vegetative shoots). Type locality: Kityak. Etymology: cornutus (Latin), horned. Diagnosis: Axis of compound polysperm continuing from leafy shoot, bracts helically arranged, free, linear, more than 6 mm long and 1 mm wide. Seed scale consists of proximal part with parallel margins and distal lamina divided into acuminate lobes. Lateral lobes long and bent outwards, central ones short and straight. Sterile shoots of Geinitzia type. Leaves falcate, up to 6 mm long and 1.5 mm wide. Description: The type locality yielded 4 compound polysperms (Plate VIII, 99 101; Fig. 29A), one isolates seed scale (Plate VIII, 98; Fig. 29B) and numerous sterile shoots (Plate VIII, 102; Fig. 15C, D, Fig. 30F, G). The seed scale structure is better seen in the isolated specimen. In its middle part there is a projection where the seed was probably attached. One seed (in attachment) is seen in the holotype. Dispersed seeds found in the same layer are shown in Fig. 30A-E. They are oval or obovate, 2.5-4.5 mm long and 2-2.5 mm wide, with a marginal border and shallow longitudinal folds. The seed apex is attenuated bicornute or funnel-shaped. The bicornute structure might have arisen as a result of a longitudinal fracture of the funnel-shaped apex. Which of the associated seeds belong to P.? cornuta is unknown. The accompany-

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Fig. 29. (A) Pseudovoltzia? cornuta Meyen, sp. nov., holotype No. 1438/15, compound polysperm, seed in attachment (?) at arrow, locality Kityak. (B) P.? cornuta, detached seed scale, spec. No. 1438/6, locality Kityak. (C) Swedenborgia longifolia Stanisl., seed scale, Upper Triassic, Donets basin (redrawn from Stanislavsky, 1971). (D) Cycadocarpidium pilosum Gr.-Stamm, seed scale and bract, seed scars stippled, Middle Triassic, Vosges (modified from Grauvogel-Stamm, 1978). (E, F) Pseudovoltzia liebeana (Gein.) Florin, seed scale and bract [(E) outer view; (F) longitudinal section)]; seed shown in black, seed scars stippled, vascular bundles shown by broken lines, upper Permian of Western Europe (modified from Schweitzer, 1963). Scale bar= 1 cm (A, C), 1 mm (B, D).

ing sterile shoots are similar to those of Concholepis harrisii, but the leaves are smaller. Comparison: These specimens were referred to Pseudovoltzia with a query, because the bract is free and the seed scale lobes cannot be separated into "sterile" and "fertile" unlike in the type species P. liebeana (Geinitz) Florin. However, its seems premature to establish a new genus for the present species. Homed seed scales are known in the lower Mesozoic genera Aethophyllum, Swedenborgia, Cycadocarpidium and Tricranolepis (Fig. 29C, D; Stanislavsky, 1976; GrauvogelStamm, 1978) all showing a bract partly fused with the seed scale. The three former genera produced different leaves. Sterile shoots similar to those of P.? cornuta were earlier described from the type locality as Ullmannia sp. (see synonymy).

The question of the presence of Ullmannia in Western Angaraland is discussed in Sections 5.8 and 9. Locality: Kityak.

6. Evolutionary morphology of Late Palaeozoic and Early Mesozoic conifers The following comparative analysis concerns only those characters which have been studied in Permian conifers of Western Angaraland, and those Palaeozoic and Mesozoic genera for which relations to conifers of Western Angaraland are more comprehensible.

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!

1

Fig. 30. Seeds ( A E ) and leafy twigs (F, G) associated with Pseudovoltcia". cornuta Meyen, sp. nov. (A) Spec. No. 3773/80A. (B) Spec. No. 3773/96-1. (C) Spec. No. 3773/80 (counterpart of A). (D) Spec. No. 3773/96-1. (E) Spec. No. 3773/93. (F) Spec. No. 3773/81. (G) Spec. No. 3773/97. Locality Kityak. Scale bar 1 crn.

6.1. Femalefructifications Female fructifications of Late Palaeozoic conifers can be divided into two main types. The first type comprises the Buriadiaceae having solitary (not aggregated into polysperms) seeds produced by non-specialized shoots indistinguishable from sterile ones (Buriadia, Walkomiella; Pant and Nautiyal, 1967; Surange and Prem Singh, 1952). The second type comprises compound polysperms: the axis bears helically arranged fertile complexes (simple lateral polysperms), usually subtended by bracts. Among Upper Palaeozoic lateral polysperms at least 10 types can be recognized corresponding to different genera. Lebachia (Fig. 31 ). Florin ( 1938, 194) suggested the following interpretation of the axillary complex of Lebachia pinijbrmis (Schlotheim) Florin. The axil of a forked bract (of the Gomphostrobus type)

bears a fertile dwarf shoot with numerous helically arranged sterile scales and a single orthotropous clavate ovule having an apical notch. The ovule is located in the apical part of the axillary complex on its adaxial side. This interpretation has been unanimously adopted. In the meantime, more careful studies of the published photographs (Florin, 1938), a comparison with other late Palaeozoic conifers, and with associating seeds (Samaropsis delaJbndii Zeiller) demonstrate that Florin's interpretation of the clavate organ as an ovule is erroneous (Clement-Westerhof, 1984). The following arguments can be adduced against Florin's interpretation: ( 1) The clavate outline with a gradually tapering base is very unusual for all late Palaeozoic ovules (seeds) of Pinopsida. The ovules of Pinopsida, when apically attached, always have an expanded base abruptly constricted in the place of attach-

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415

l II

Fig. 31. Lebachiellaflorinii Meyen, sp. nov. ( = Lebachia piniformis sensu Florin), interpretation of lateral polysperm, seed stalk stippled, bract shaded, ovule shown by broken line, seed scar in black. (A) Adaxial view. (B) Viewed from the side of bract. (C) Lateral view.

ment. After the shedding of the seed a distinct scar had been formed at the seed stalk apex. Florin held, that Lebachia is associated with Samaropsis delafondii seeds. The latter have a constricted base and hence, cannot be ontogenetically derived from clavate organs. (2) The identity of the clavate organ with an ovule has not been supported by a proper analysis of cutinized membranes, in particular, the cuticle of the integument in the ovule apex and of the micropyle has not been demonstrated. A membrane identified with the outer cuticle of the integument (Florin, 1938, pl. 21/22, figs. 7-12) is much more similar to cuticle of sterile scales of the axiUary complex. (3) The clavate organ shows, immediately under the apical notch, a distinct contour (Florin, 1938, pl. 19/20, figs. 10, 11) which is strikingly similar to a seed scar. (4) The similarity between the clavate organ of Lebachia and a flattened leaf-like seed stalk of O.rtiseia (Fig. 32C, E, Fig. 33A; ClementWesterhof, 1984) is beyond doubt. Thus, there are grounds to admit the interpretation of the axillary polysperm of Lebachia pini-

form& as given in Fig. 31, and to reject Florin's interpretation. "Lebachia" lockardii (Fig. 32A). This species was established by Rothwell (1982b) and Mapes and Rothwell (1984) on permineralized compound polysperms, microstrobili and leafy twigs coming from the Upper Carboniferous of Kansas. Mapes and Rothwell held that their study of the anatomically preserved material supports Florin's interpretation of Lebachia. In the meantime, in "L." lockardii the seed stalks are thin and hooked, and the apically attached ovule shows an abruptly constricted base. Therefore "L." lockardii is closer in this respect to Kungurodendron (Fig. 32B) and Cordaitanthales (see below) than to Lebachia. Obviously, this species cannot be placed into the genus Lebachia (that is why the generic name is quoted), and there are reasons to separate it into a new genus (see Section 8). An important peculiarity of "L." lockardii is the dorsiventral flattening of the axillary complex and the restriction of seed stalks to its adaxial side. A primitive character of the species is the forked bract (as in Lebachia). Ortiseia (Fig. 32C-F, Fig. 33A). Polysperrns of Ortiseia were studied in detail by Clement-

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Fig. 32. Reconstruction of lateral polysperms of "Lebachia" lockardii Mapes et Rothwell (A), Kungurodendron sharovii Meyen, .Tx nov. (B) and Ortiseia leonardii Florin emend. Clement-Westerhof (C F), ovules and seed scars in black, bracts shaded, seed stalks in (A, C-E) stippled, in (B) seed stalks densely stippled and funiculodia sparsely stippled, in (F) place of seed attachment stippled, in (C) and (E) bract removed; adaxial (C) and abaxial (E) views, seed in (F) shown with chalazsa at top. Upper Carboniferous of Kansas (A), locality Chekarda- 1 (B), Upper Permian of Italy (C F ). Modified from Mapes and Rothwell (1984) (A), and ClementWesterhof (1984) (C-F). Scale bar = 2 ram.

Fig. 33. Differentiated elements of the bract-axillary complex in Ortiseia h,onardii Florin, emend. Clement-Westerhof (A) and differentiated ones in Sashinia borealis Meyen (B); bracts shaded, seed stalks stippled, seed scars in black [in (B) partly shown by broken line], in O. leonardii place of bract attachment shown by broken oval contour. (A) modified from Clement-Westerhof (1984).

S. V. Meyen / Reviewof Palaeobotanyand Palynology96 (1997)351-447 Westerhof (1984). The bract is wide, leaf-like, with an attenuated non-forked apex. The axillary complex is flattened, composed of numerous sterile scales varying in outline depending on their position in the axillary complex. The seed stalk is broad, leaf-like, sometimes with a small apical notch. The seed scar is situated in the middle of the stalk on its abaxial side, but since the seed stalk occupies an adaxial position in the axillary complex, the seed scar appears on the adaxial side of the latter. The ovule (seed) is large, with basal outgrowths and a laterally displaced area of attachment. As a whole, Ortiseia proves to be closely related to Lebachia, but differs in the lower position of the ovule on the seed stalk, the more flattened and more distinct dorsiventrality of the axillary complex, and the simple (non-forked) bract. Kungurondendron (Fig. 32B). Salient features of Kungurondendron essential for the present discussion are: (1) a large number of seed stalks; (2) their adaxial position in the axillary complex; (3) the attachment of the ovule to the hooked apex of the seed stalk; (4) the presence of numerous sterile scales in the basal part of the axillary complex; (5) the presence of funiculodia (sterilized seed stalks) at the apex and on the abaxial side of the axillary complex; (6) the transition from sterile scales to seed stalks; (7) the simple bract indistinguishable from foliage leaves. Timanostrobus (Fig. 12). In the overall habit of the lateral polysperms, Timanostrobus is much closer to Vojnovskyaceae (Cordaitanthales) than to conifers. There are no signs of the flattening of the lateral polysperms. Multiple seed stalks cover the major part of the lateral polysperm, and sterile scales--its apex and base (?). The bract, if present, cannot be recognized among basal appendages of the lateral polysperm. It remains unknown, whether sterile scales of the lateral polysperm are initially sterile (i.e., derived from circasperm members of Cordaitanthales) or are funiculodia (i.e., sterilized seed stalks) as in Kungurodendron. Sashinia (Fig. 16C, Fig. 33B, Fig. 34S, T). This genus is characterized by: (1) a likeness of bracts, elements of the axillary complex (sterile scales and seed stalks) and foliage leaves (Fig. 33B); (2) the

417

aggregation of seed stalks in the apical part of the axillary complex; (3) the abaxial bend of the distal part of the seeds stalks and its pad-like expansion; and (4) the abaxial ovule position under the distal pad. Walchiostrobus, Ernestiodendron (Fig. 34A-R). According to Florin (1939, 1944), the axillary complex of Ernestiodendron filiciforme (Schloth.) Florin was radial and composed of only free seed stalks without accompanying sterile scales. Compound polysperms, where seed stalks were accompanied by sterile scales, he (Florin, 1940) referred them to Walchiostrobus (Ernestiodendron?) fasciculatus Florin. Similar detached axillary complexes were described by him as Walchiostrobus sp. a. The most important specimens had been figured by Florin in diagrammatic drawings and very good photographs, the reciprocal correspondence of which had been explained in the captions. Some of his drawings have become classics, and were repeatedly reproduced in many textbooks. Unfortunately, the fidelity of his drawings has not been checked by subsequent workers. The present author has made a careful comparison between the drawings, photographs and some originals. Regretfully, the drawings do not well illustrate characters observed on the specimens which illustrate Florin's general ideas. As will be shown below, Florin systematically mistook subapical seed scars for the nucelli of orthotropous ovules. There are several features which allow us to distinguish seed scars from ovules. In well-preserved compression-impression material, the ovule microstructure, as seen under a binocular microscope, is quite different from that of the stalk. When we observe a round, oval or rhomboid contour on a stalk, and the microstructure around the contour and downwards along the stalk is continuous and uniform, the pertinent contour can safely be treated as a seed scar. In gymnosperm seeds (ovules) the nuceUar contour is usually connected with the seed apex, with a more or less clear micropyle trace, often underlined by a coaly strand. Besides, in both cordaitanthaleans and conifers the ovules are basaUy constricted, and the seed stalk never grades into the apical ovule of the same width without any discernible bound-

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t

I

d e"

Fig. 34. Monosperms and lateral polysperms of primitive conifers. {A-C) specimens referred by Florin to ErnestiodendronfiliciJorme (Schloth.) Florin. (A) Organs treated by Florin (1944, fig. 36) as monosperms. {B) Same specimen traced after photograph (Florin, 1939, pl. 117/118, fig. 21). (C) Seed stalk with subapical seed scar and subtending bract, drawn after photograph (Florin, 1939, pl. 119/120, figs. 3, 4). (D-G) Specimens referred by Florin to Wak'hia (Ernestiodendron?) germanica Florin. (D) Axillary complex in abaxial view, drawn after photograph (Florin, 1939, pl. 149/150, fig. 2 ). (E) Same axillary complex in interpretation of Florin ( 1944, fig. 35f). (F) Two monosperms with ovules, drawn after photographs (Florin, 1939, pl. 149/150, fig. 3). (G) Same axillary complex in interpretation of Florin (1944, fig. 35e). (H) Specimen of the same type as Florin's Walchiostrobus sp. a, flattened seed scale consists of monosperms bearing seed scars (stippled), circasperm consists of numerous scales, Museum for Naturkunde, Berlin. (I, J ) Seed stalk of specimen referred by Florin to Walchiostrobus (Ernestiodendron?)fasciculatus Florin. (I) Interpretation of Florin (1944, fig. 34a). (J) The same traced after photograph (Florin, 1940, pl. 153/154, figs. 15 17). (K, 1) Detached axillary complex of Florin's Walchiostrobus sp. a. (K) interpretation of Florin ( 1944, fig. 34h). (L) The same traced after photographs (Florin, 1940, pl. 153/154, figs. 19, 20). (M) Walchiostrobus sp. SVM-1, detached seed scale devoid of circasperm, subapical seed scars are seen, Museum ft~r Naturkunde, Berlin. (N) Schematic drawing of specimen shown on (H } with added ovules. (O) Schematic drawing of specimen shown on (M) with added ovule. (P) Walchiostrobus (al. Ernestiodendron), monosperm structure in Florin's interpretation. {Q) Proposed interpretation of the same monosperm, seed scars stippled, ovules in black. (R) Monosperms of some Cordaitanthales and most primitive conifers (lateral view, ovule in black). (S) Axillary complex of Sashinia, ovules (in black) attached abaxially under apical pad bent outward. (T) Monosperm of Sashinia, lateral view, abaxial side turned to right, seed in black. Scale bar= 1 mm.

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ary. On the other hand, in some cordaitanthaleans and conifers aborted seeds are observed (Meyen, 1982a; Rothwell, 1982a); the present author encountered aborted seeds in Sashinia and the Peltaspermales. Such ovules are frequently found in attachment because they have not managed to develop an abscission layer. When the aborted ovule is subapical it overlaps the stalk. With these and other criteria in mind we can now turn to the comparison of Florin's drawings and photographs. Fig. 34A-G, I-L offers a comparison of some of Florin's drawings (A, E, G, I, K) with accurate tracings of the corresponding photographs (B, D, F, J, L). His drawing of Ernestiodendron filiciforme (Fig. 34A) shows two monosperms with orthotropous ovules, having distinct contours of the nucellus and a wide trapeziform micropyle. On the corresponding photograph (see its tracing in Fig. 34B) the apical notch is absent (see also Fig. 34C). The oval contour described as a seed nucellus is obviously a seed scar identical to seed scars of Pseudovoltzia liebeana (see Schweitzer, 1963, pl. 1, fig. 7), but subapical in position. The presence of the seed scars (not seeds!) is quite evident in Walchiostrobus sp. a (Fig. 34L, K). The pertinent drawing of Florin is rather misleading. It shows numerous sterile scales and five monosperms. The latter bear orthotropous ovules with distinct nucelli and wide trapeziform micropyles. The corresponding photograph, very well taken, shows only very feeble traces of sterile scales, only four indisputable monosperms; apical notches (micropyles) are certainly absent, and closed contours, treated by Florin as nucelli, are undoubtedly seed scars. The three-dimensional arrangement of monosperms implied by Florin's drawing cannot be admitted either. In the case of a helical arrangement, the monosperms would hardly all be orientated in the same morphological plane towards the observer. In this specimen all the monosperms bear seed scars facing upwards. Hence the whole organ was initially flattened, again as in Pseudovoltzia liebeana. The present author studied specimens comparable with Walchiostrobus sp. a (Fig. 34H) in the Museum ftir Naturkunde, Berlin. Their monosperms are arranged in a single plane, bear

419

subapical seed scars of the same orientation, and their lower portions are overlapped by sterile scales. In the same collection there are polysperms conventionally here designated as Walchiostrobus sp. SVM-1 (Fig. 34M). They are devoid of sterile scales, and the monosperms are basally fused. A flattened polysperm consisting of monosperms basally fused into a limb is well seen in W. fasciculatus Florin (1939, pl. 153/154, figs. 11-17). Its interpretation by Florin (1944, fig. 34a) is obviously erroneous. For instance, he depicted its monosperms as having orthotropous ovules with a very wide barrel-shaped micropyle (Fig. 34I), but the supporting photographs taken both in xylol and in dry condition exhibit a quite different structure (Fig. 34J). The apical notch is very shallow. An oval contour (nucellus in Florin's treatment) is closed below (not open as in Florin's drawing) and disappears apically. It may be either an aborted ovule or a seed scar as shown in Fig. 34Q. Florin's drawing of Walchia (Ernestiodendron?) germanica Florin (Fig. 34D-G) are not convincing either. They show free monosperms bearing inverted apical ovules. Judging from photographs, the monosperms may well be basally fused, in non of them are the ovule outlines distinct, and the ovules themselves may be subapicaUy attached and underdeveloped. In the Museum for Naturkunde, the present author examined one of the originals of Walchiostrobus gothanii Florin ( 1939, pl. 151/152, figs. 49, 50). This species was affiliated by Florin with Lebachia, instead of Ernestiodendron, on obscure grounds. Judging from this specimen, Florin's drawings of the axillary polysperms of W. gothanii (Florin, 1944, fig. 33F, G) are erroneous. In the middle part of the original the axillary complexes are spread in the plane of the main axis. The monosperms are basally fused into a limb and bear subapical scars treated by Florin as seeds. Thus, in E. filiciforme, W. gothanii, W. fasciculatus, W. sp. a and W. sp. SVM-1, the lateral polysperms (axillary complexes) are actually bifacial seed scales consisting of basally fused monosperms (stalks with subapical abaxial ovules) intermittently accompanied by sterile scales. Some

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of the scales may represent funiculodia, but the basal ones obviously belong to a vestigial circasperm homologous to that of Lebachia, though more reduced. The above interpretation of the axillary polysperms of Walchiostrobus and Ernestiodendron was proposed by the author in 1984 (Meyen, 1984, pp. 83-86, fig. 27). Clement-Westerhof (1984) has arrived at similar conclusions independently. She also considers that oval contours treated by Florin as ovules in E. filiciforme (Fig. 34A-C) are seed scars, but she did not mention that Florin had made the same mistake in the case of W. (E?) fasciculatus, W. gothanii and Walchiostrobus sp. a. Besides, she considered that E. filicijbrme produced solitary seed stalks in the axil of the bract. For the present it remains uncertain whether this species produced solitary axillary seed stalks or if only one seed stalk of a more complex lateral polysperm is exposed by the fracture plane. The bract of Walchiostrobus-Ernestiodendron was free, because ( 1) there are findings of detached axillary complexes of Walchiostrobus sp. a and alike, and (2) in some compound polysperms the bracts are devoid of axillary polysperms (they had abscissed?). Concholepis (Fig. 36Q). As was said above (Section 5.4), the morphological interpretation of the Concholepis polysperms is uncertain. We do not know, whether the seeds were attached directly to a smooth adaxial surface of the axillary complex or to submarginal adaxial projections which would be then seed stalks. Numerous projections along the margin and on the abaxial side of the axillary complex can be homologous to sterile scales and funiculodia of Kungurodendron. Pseudovoltzia (Fig. 29E, F). The drastic changes in Florin's interpretations of the axillary complex of Pseudovoltzia that resulted from Schweitzer's studies were mentioned in Section 1. The seed scale of P. liebeana is comparable to that of Ortiseia in the seed attachment in the middle part of the seed stalk. Fertile lobes of the seed scale of P. liebeana are homologous to seed stalks and back sterile lobes - - to either sterile scales of the above described conifers, or funiculodia. Pseudovoltzia liebeana is close to Ortiseia and

Sashinia in the low position of seeds on the seed scale lobes (seed stalks), but differ from them in having a true seed scale and its basal fusion with the bract. Pseudovoltzia? cornuta described above ( Fig. 29A, B) probably belongs to an independent genus, but is insufficiently known. Its seed scale is free from bract. Ullmannia (Fig. 35A-C). Polysperms are known in U. frumentaria (Schlotheim) Goeppert. The well known better preserved specimen had been erroneously referred by Florin (1944, p. 484) to U. bronnii Goeppert (see Schweitzer, 1963, p. 16). Florin held, that the seed scale of Ullmannia consists of five completely fused scales and a single seed on a long stalk overlapping the seed scale in its lower part (Fig. 35A). The published photographs, however, do not support his conclusion. Schweitzer (1963) studied additional material and suggested another interpretation of the seed scale (Fig. 35C) that agrees much better with all we know about Late Palaeozoic conifers, namely, the ovule is attached adaxially to the middle of the seed scale which show no traces of the fusion of several scales. The organ interpreted by Florin as a seed scale may well be a solitary seed stalk comparable to that of Ortiseia. Then the bract-axillary complex of U. frumentaria will prove to be comparable with that of Ernestiodendronfiliciforme as treated by Clement-Westerhof (see above). The published photographs of the U. frumentaria polysperms do not allow us to judge, whether these lateral seed-bearing organs were subtended by bracts. Photographs presented by Florin ( 1944, pl. 179/180, figs. 17-19; pl. 181/182, figs. 1, 2), drawing of Geinitz (1880) as well as drawings of fertile shoots presented by Weigelt (1928) under the name "Archaeopodocarpus germanicus", do not show bracts. Therefore, a possibility emerges that the repeatedly figured fertile shoot of U. frumentaria corresponds to a detached lateral polysperm of Sashinia (rather than to a compound polysperm), but differing in the helical arrangement of seed stalks throughout the axis instead of their aggregation in an apical bunch. Voltziopsis (Fig. 35D). Detailed descriptions of Voltziopsis polysperms were given by Townrow (1967b) on materials from the Triassic of

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421

C Fig. 35. (A-C) Seed scale of Ullmanniafrumentaria (Schloth.) Goepp. (A, B) Reconstruction by Florin (1944), adaxial (A) and abaxial (B) views, ovules in black, seed stalk stippled, bract shaded. (C) Reconstruction by Schweitzer (1963). (D) Voltziopsis wolganensis Townrow, reconstruction of seed scale, ovules in black, bract shaded.

Gondwana. The axil of a forked bract bear 5-6-lobed seed scale with a basal limb and narrow lobes. In the middle part of the latter, inverted ovules are attached. Voltziopsis differs from Pseudovoltzia in its forked bract and the absence of sterile scale. Voltzia. The female fructification structure of Triassic Voltzia are poorly known. In 1984 L. Grauvogel-Stamm demonstrated to the author both polysperms and detached axillary complexes probably belonging to the type species V. heterophylla Brongniart. Although the preservation of the material is not quite perfect, one can clearly see that the seed scales are of the same overall structure as those of Pseudovoltzia. In his interpretation of the Voltzia polysperms Florin (1944, 1951 ) appears to have made the same mistakes as in the case of Pseudovoltzia. Fructification structure of other Permian and older Mesozoic conifers is given in the literature (Schweitzer, 1963; Stanislavsky, 1971, 1976; Miller, 1977, 1982; Anderson, 1978; Grauvogel-Stamm, 1978; Anderson and Anderson, 1985). Better studied are the genera Swedenborgia (Fig. 28C), Aethophyllum, Cycadocarpidium (Fig. 29D), Tricranolepis and Borysthenia. In all of them the seed scale is lobed with low attachment of the ovules to the lobes; sterile scales are absent, the bract is more or less fused with the seed scale, it appears as a small appendage (Swedenborgia), may be long and narrow (Aethophyllum, Boristhenia) or wider with several parallel veins (a

part of Cycadocarpidium) or a single (?) vein ( Telemachus, Cycadocarpidium pilosum GrauvogelStamm). The Upper Triassic genus Rissikia (Townrow, 1967a) stands apart from the above mentioned early conifers. It has a forked bract (as in Voltziopsis) and the seed scale is subdivided into three deep lobes each bearing one-two ovules. Of utmost importance is that in the ovules Townrow recorded long seed stalks attached to the lobe lower part. The ovules are inverted. Thus, Rissikia demonstrates that type of the lateral polysperm construction that had been erroneously ascribed by Florin to many Permian and Triassic conifers. Both the appearance and homology of the seed stalk in Rissikia is enigmatic. It is true that free seed stalks were figured by Roselt (1957-1958) in Tricranolepis, probably under the influence of Florin's views. Grauvogel-Stamm (1978)p. 162, pl. 21, fig. 2a; text-fig. 47a) figured the Aethophyllum stipulate seed scale having longitudinally split lobes similar to free seed stalks. The free stalks, if they were present in some Triassic conifers, might have arisen secondarily as in living Cryptomeria (Schweitzer, 1963; Bierhorst, 1971). Remaining Permian and Triassic conifers yielding compound polysperms have been insufficiently studied for reliable comparisons, but they probably fit the above described diversity of the bract-axillary complexes. Comparing the above polysperm type with each other and with polysperms of Cordaitanthales (not

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S. V. Meyen/ Reviewof Palaeobotanyand Palynology96 (1997) 351-447 only Cordaitanthaceae, but also Vojnovskyaceae and Rufloriaceae) and Dicranophyllales, and considering the relative stratigraphic ranges of the genera, we may outline primitive and advanced characters of polysperms as well as the main modes of their transformation (Fig. 36): (1) From unflattened axillary complex (nearly all Cordaitanthales and at least a part of Dicranophyllales; Timanostrobus, Sashinia) to flattened ones (remaining conifers). (2) Shift of seed stalks onto the adaxial side of the axillary complex (Lebachia, " L . " lockardii,

Kungurodendron). (3) From a large number of seed stalks or sessile ovules (most of Cordaitanthales; Timanostrobus, Kungurodendron) to a lesser number of them

(Sashinia, Concholepis, Walchiostrobus, Pseudovoltzia, Voltzia, Voltziopsis, Swedenborgia, Aethophyllum, Cycadocarpidium, Boristhenia) and then to a single seed stalk (Lebachia, Ortiseia, Ullmannia?); with variations in this character within species ("Lebachia" lockardii). (4) From the apical position of ovules on seed stalks (most of Cordaitanthales; " L . " lockardii, Kungurodendron, Timanostrobus) to abaxial (Gaussia scutellata Neuburg, Lebachia, Ortiseia, Sashinia,

Walchiostrobus-Ernestiodendron, Pseudovoltzia, Ullmannia, Voltzia, Voltziopsis, Swedenborgia, Aethophyllum, Cycadocarpidium, Borysthenia). (5) From a forked bract (Lebachia, " L . " lockardii, Voltziopsis) to simple one (Cordaithanthales, most of conifers). (6) From free seed stalks (Kungurodendron, Timanostrobus, Sashinia) to their basal fusion into a primitive seed scale (Walchiostrobus-

Ernestiodendron, Concholepis, Pseudovoltzia, Voltzia, Voltziopsis, Swedenborgia, Aethophyllum,

423

Cycadocarpidium, Borysthenia) and further to a fully fused seed scale (most Mesozoic conifers). (7) From the presence of numerous sterile scales in the axillary complex (many Cordaitanthales, Lebachia, " L . " lockardii, Ortiseia, Kungurodendron, Timanostrobus, Sashinia, Walchiostrobus-Ernestiodendron, Concholepis) to a fewer number (Pseudovoltzia) and a full reduction (most conifers, some Cordaitanthales). (8) From a free bract (all Carboniferous and Lower Permian conifers, the Upper Permian

Ortiseia, Sashinia, Concholepis, Pseudovoltzia ? cornuta) to a basal fusion of bract and seed scale (Pseudovoltzia liebeana, all Triassic conifers). An undoubtedly primitive character is the position of compound polysperms at the continuation of leafy twigs corresponding in thickness to branches of the last or penultimate orders. In Cordaitanthaceae, the compound polysperm was located in the same orthostichy with the subtending vegetative leaf, but much higher its axil, sometimes at the same level as the vegetative leaf of the adjacent orthostichy (Grand'Eury, 1877; own observations of the author). The attachment above the axil was also observed in lateral polysperms of Vojnovskya (Meyen, 1984). The location of compound polysperms (and microstrobili) at the continuation of leafy twigs can be treated as an innovation of early conifers. Probably, none of the Late Palaeozoic and Triassic conifers develop a specialized stalk of compound polysperms. Several characters cannot reliably be treated as primitive or advanced, namely: (1) a straight or hooked seed stalk; (2) the degree of the similarity between sterile scales of the axillary complex, seed stalks, bracts, and vegetative leaves; (3) the transition between seed stalks and sterile scales within

Fig. 36. Main types of axillary complexes, lateral polysperms or seed-bearing branchlets in Cordaitanthales (Cr), Dicranophyllales (De) and Pinales (Pn), and probable ways of their transformations; bracts and ovules in black, seed scales (N, R-U), leaf-likeseed stalks (L, O) and funiculodia(M) stippled, sterile scales (circaspermmembers) shown by narrow white triangles, homologousorgans shown identical, helicallyarranged sterile scales and monosperms two ranked. (A) Cordaitanthuspseudofluitans (Kidst.) Crook. (B) C. diversiflorusCrook (sterile scales problematical). (C) C. zeilleri Ren. sp. (D) C duquesnensis Rothw. sp. (E) Vojnovskya. (F) Gaussia cristataNeut. (G) G. scutellataNeub. (H) Suchoviella. (I) Dicranophyllum.(J) 'Lebaehia' lockardiiMapes et Rothwell. (K) Buriadia. (L) Lebaehiellaflorinii Meyen, sp. nov. (M) Kungurodendron. (N) Walehiostrobus,(0) Sashinia. (P) Timanostrobus. (Q) Coneholepis. (R) Pseudovoltzia, Voltzia.(S) Swedenborgia,Aethophyllumand allied Triassic genera. (T, U) more advanced Mesozoic and Cenozoicconifers with partly (T) and completely(U) fused seed scale and bract. (V) taxads.

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the axillary complex (as in Kungurodendron); (4) seed stalk length; (5) mutual position of seed stalks and sterile scales along the axis of the axillary complex. Some of the listed transformations of characters are probably irreversible. These are the shift of seed stalks onto the adaxial side of the axillary complex, the shift of ovules from the apex of seed stalks to their abaxial side, the basal fusion of seed stalks into a primitive seed scale, and the loss of sterile scales in the axillary complex. Other transformations are reversible to varying degrees. These are the flattening of the axillary complex, reduction of number both of seed stalks and sterile scales, the degree of likeness of elements of the bract axillary complex and foliage leaves, forked or simple bracts (a forked bract in the Triassic genera Voltziopsis and Rissikia is suggestive), the degree of fusion of the bract with the axillary complex, straight or hooked seed stalk, length of the seed stalks and lobes of the seed scale. None of the mentioned genera possesses the lull set of either primitive of advanced characters. The maximum number of primitive characters is observed in "Lebachia" lockardii and Timanostrobus, but even they show certain advanced characters (flattening of the axillary complex, a few or a single seed stalk in "L." lockardii, partial or complete reduction of the bract in Timanostrobus). In the most advanced Triassic conifers some primitive characters are preserved, namely a deep lobbing of the seed scale, large dimensions of bracts. fusion of bract and axillary complex for a short distance, the absence of a specialized stalk in the compound polysperms. All said above about the primitive or advanced nature of female fructifications cannot be referred to pertinent genera as a whole, because the estimation of genera in terms of primitivity and advancedness needs consideration of a wider set of characters. The above material allows us to conclude on the origin of the coniferalean seed scale. Florin (1938, 1939, 1940, 1944, 1945, 1951) held that the seed scale had resulted from a fusion of the circasperm members of Lebachia-like conifers. In the meantime, as early as in Walchiostrobus Ernestiodendron there is a fully formed seed scale

consisting of basally fused seed stalks and accompanied by sterile scales. Then the latter are reduced. Thus, the origin of the seed scale by means of seed stalk fusion appears much more plausible. It is essential that the seed-bearing adaxial side of the seed scale corresponds to the fused abaxial sides of seed stalks. Hence the fused seed scale can be treated as a unifacial organ (Meyen, 1984). The nature of the axillary complex in Concholepis is less comprehensible. Numerous abaxial and marginal appendages give an impression that this axillary complex had been formed at the expense of a further flattening of a Kungurodendron-like axillary complex, and further fusion and reduction of its components. In this case sterile elements of the axillary complex might have participated in the formation of the seed scale. A phenomenon of seed stalk sterilization should be also born in mind. Such sterilized seed stalks (funiculodia) forming a kind of a "secondary" circasperm, might have participated in the seed scale formation. The relative share of seed stalks and elements of the circasperm (both secondary and primary) in the seed scale formation might have been different in various coniferalean lineages.

6.2. Male Ji'uct![ications Among Permian conifers of Western Angaraland, male fructifications are known only in Kungurodendron, Timanostrobus and Sashinia (genus Dvinostrobus). From other older conifers, microstrobili were described in Lebachia, "'L.'"

lockardii, Ortiseia, Walchiostrobus-Ernestiodendron, Ullmannia, Voltzia (?), Aethophyllum and Cvcadocarpidium. Microstrobili of Late Palaeozoic conifers are poorly known. Florin (1940) noted, that the microsporophylls in Lebachia are peltate, bearing two pendent sporangia attached to the distal shield on both sides of the stalk insertion. Although this interpretation has been widely adopted in the literature, it had not been supported by convincing photographs. Microsporophylls of Ortiseia, Kungurodendron, Tirnanostrobus and Sashinia also had a distal shield, but the mode of sporangial attachment (with the aid of thin sporangiophores

S. V. Meyen / Reviewof Palaeobotanyand Palynology96 (1997) 351-447 to the middle of the stalk) is known only in Sashinia. Probably, sporangia were attached to the stalk, rather than to the shield, in the three remaining genera, either. According to G. Mapes (pers. commun.), microsporophylls of "L." lockardii were of the same construction. In Ullmannia numerous sporangia were attached to the shield (Potoni6 and Schweitzer, 1960; Schweitzer, 1960). The Upper Pennsylvanian genus Lasiostrobus (Taylor, 1970; Taylor and Millay, 1977) can be tentatively place into conifers. It was described as an isolated microstrobilus. Its sporangia are adnated to the abaxial side of the stalk of a peltate microsporophyll. Microstrobili of Triassic conifers are better known (Grauvogel-Stamm, 1978; GrauvogelStamm and Schaarschmidt, 1979). Among them largely the same types of microstrobili can be recognized. As a whole, the Late Palaeozoic and Triassic microstrobili possessed microsporophyUs of the following types. (1) Sporangia attached to distal shield. (a) Sporangia are paired and free - - Lebachia

(?). (b) Sporangia are paired and adnate to stalk - Amydrostrobus, Cornpsostrobus. (c) Sporangia numerous and free - Willsiostrobus (including microstrobili associated with Ullmannia). (2) Sporangia adnated to stalk with entire length - - Lasiostrobus. (3) Sporangia are attached with the aid of thin sporangiophores to adaxial side of the stalk in its middle part - - Dvinostrobus, Darneya, Sertostrobus, probably also "Lebachia" lockardii, Ortiseia, Kungurodendron and Timanostrobus. Distal shields may be without a heel (Lasiostrobus, Ortiseia, Kungurodendron, Timanostrobus, Dvinostrobus) or with a heel, i.e. peltate (Lebachia?, Darneya, Sertostrobus, Willsiostrobus, Amydrostrobus, Compsostrobus). In the female fructification structure an essential similarity exists between older conifers and Cordaitanthales, whereas in the structure of microstrobili both groups are principally different. Two genera, however, violate this general rule. One of

425

them is Cladostrobus (Rufloriaceae, Upper Permian of Siberia and Mongolia; Maheshwari and Meyen, 1975). Its microsporophylls are similar to those of group 3, particularly Dvinostrobus and Darneya. The Cladostrobus microsporophylls stand apart among other cordaitanthalean male fructifications (Meyen, 1984). Probably, in their structure as well as in comparable coniferalean microsporophylls we observe a recapitulation of characters of archaeopteridalean progymnosperms, with fertile pinna which (of penultimate order) Beck (1981) homologized the microstrobilus of Lebachiaceae. In the Archaeopteridales, the fertile pinna of the last order may be crowned by a sterile pinnule which can be homologized with the distal shield of coniferalean microsporophyll. Sporangia of the Archaeopteridales are inserted along the acroscopical side of the pinna of the last order, that finds a parallel in the adaxial position of sporangia on the microsporophyll stalk in Darneya and Sertostrobus. At last, sporangia of the Archaeopteridales may be born by thin branching sporangiophores, again as in these coniferalean genera. Somewhat similar bunches of sporangia are known in several members of the Cordaitanthales. The second genus is the Upper Triassic Ruehleostachys conventionally referred to conifers or cordaites (Roselt, 1955-1956; GrauvogelStamm and Schaarschmidt, 1979). Here long tubular microsporangia are aggregated in bunches attached directly to the strobilar axis. A direct attachment of solitary sporangia to the strobilar axis is known in the Upper Permian Angara genus Pechorostrobus (Rufloriaceae; Meyen, 1984). Types 1 and 2 of coniferalean microsporophylls having different sporangial arrangement do not intergrade. Probably transitions between them never existed, and the changes between the types happened homoeotically (Meyen, 1984).

6.3. Pollen Among the Permian conifers of Western Angaraland, a globose asaccate pollen is known in Kungurodendron and Timanostrobus, and a quasidisaccate pollen of the Scutasporites type in Sashinia-Dvinostrobus-Quadrocladus. Probably, some conifers having Quadrocladus shoots pro-

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duced Lueckisporites pollen (see Section 5.8). Pollen of the Platysaccus type might have associated with Quadrocladus (al. Steirophyllum) komiensis. Pollen is also known in many older conifers outside Western Angaraland. In Lebachia, " L . " lockardii and Walchiostrobus-Ernestiodendron the pollen Potonieisporites was recorded (Florin, 1938, 1939, 1940, 1944, 1945; Bharadwaj, 1963-1964, 1974), in Ortiseia Nuskoisporites (Clement-Westerhof, 1984), in Ullmanniajrumentaria Jugasporites (Schweitzer, 1960; see also Foster, 1983), in Voltzia Triadispora (Grauvogel-Stamm, 1978 ), in Yuccites- Willsiostrobus, Cycadocarpidium-Willsiostrobus and in Ruehleostachys Alisporites (Grauvogel-Stamm, 1978; Roselt, 1955-1956). Townrow (1967b) suggested that Voltziopsis was associated with striated pollen of the Protohaploxypinus type. Such pollen was found in the Upper Triassic Rissikia (Podocarpaceae; Townrow, 1967a). Both description and comparison of pollen of older conifers involve serious difficulties. Pollen of the Potonieisporites, Nuskoisporites, Lueckisporites, Jugasporites, Triadispora. Alisporites and Scutasporites types is usually described as saccate. But, as was shown by the TEM studies, some of these pollen types (e.g., Lueckisporites and Jugasporites) lack a continuous inner cavity comparable with a true saccus, and their structure is a protosaccus (Scheuring, 1974; Foster, 1979, 1983 ), or quasisaccus (see Section 4). Under the light microscope, pollen of the enumerated genera appear quasisaccate, i.e., instead of a hollow with a thin reticulum on the inner surface of the wall, there is a spongy mass of sporopollenin with numerous lumina of different sizes and orientations. The study of pollen associated with Ortiseia (of the Nuskoisporites type; Clement-Westerhof, 1984) and "Lebachia" lockardii (of the Potonieisporites type; Mapes and Rothwell, 1984) has shown, however, that these pollen types possess a large inner hollow surrounded by a thick layer having the same spongy structure as in the quasisaccus. Such pollen -- it can be called "subsaccate" occupies an intermediate position between saccate (eusaccate; Foster, 1983) and quasisaccate. Having been buried and compressed such pollen will be indistinguishable from quasisaccate because the hollow will be

closed. Thus, the pollen of the listed genera was either quasisaccate or subsaccate. Another organization of pollen was recorded in the Western Angara genera Kungurodendron and Timanostrobus. The proximal side is smooth or with a very fine granulose surface. The body is surrounded both equatorially and distally by a thin flange. In Kungurodendron the flange simulates the columellate structure of angiosperms, and remotely resembles the structure known in ~Tassopollis (Cheirolepidiaceae) and Lasiostrobus. Shallow swellings of the flange are a little similar to saccus-like swellings of Lasiostrobus pollen. The pollen of Timanostrobus is the same as the dispersed pollen figures (without description) by Koloda (Molin et al., 1983) under the names Acusporidatina reticuloida Koloda and Reticulatina heterobrochata Koloda. This pollen lacks a columellar-like layer (or it is unrecognizable under the light microscope), but there is another feature characteristic of angiosperms, namely a perforated outer layer appearing under the light and scanning microscopes as a perforated tectum. This pollen is conventionally termed below "tectate". No such columellar-like or tectate exine has yet been observed in other Palaeozoic and Triassic conifers. However, if the suggested coniferalian affinity of Lasiostrobus is correct, we can say that the columellar-like exine appeared in conifers as early as the Carboniferous. Somehow or other, long before the end of the Permian, conifers produced two major pollen types. The first type is characterized by expansion of the sexine as a quasisaccus, subsaccus or eusaccus having either toroidal (Potonieisporites, Nuskoisporites) or bilobed (Scutasporites, Luecki-

sporites, Jugasporites, Triadispora, Alisporites, lllinites) outlines or transitional between them (some Jugasporites and Triadispora). Usually these two varieties of the first pollen type are described in the literature as mono- and disaccate, respectively. The pollen of the first type may be further complicated by ribbing (taeniae). The second pollen type develops either a columellar-like layer or perforated tectum, whereas a tendency to the exine exfoliation into sacci is nearly absent. There is another character of coniferalean pollen finding a parallel with angiosperm pollen, namely

S. V. Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

an operculum upon the proximal aperture in Jugasporites, i.e. pollen of Ullmannia (Foster, 1983). Foster suggested that the grooves between the taeniae in Lueckisporites and Taeniaesporites might also bear operculate structures. The most primitive coniferalean pollen appears to have been of the Potonieisporites type. It is close to cordaitanthalean pollen, particularly to Felixipollenites having a thick alveolar layer along the inner side of the saccus wall, and a welldeveloped proximal slit (Millay and Taylor, 1974). If, however, Lasiostrobus does belong to the conifers, its pollen with columellar-like exine and small saccus-like swellings may also be considered as primitive according to the stratigraphic position of the genus. From this pollen type the pollen of Timanostrobus and Kungurodendron can be derived. Another possibility is that from the very beginning of the evolution of conifers, a repeated reduction of either a eusaccus or quasisaccus and resulted in independently arising asaccate forms. This possibility is indirectly evidenced by the independent and obviously secondary appearance of asaccate pollen in some Pinaceae and Podocarpaceae. Aberrant asaccate grains are known in species normally producing saccate pollen. As was said above, quasisaccate and eusaccate organizations are linked by transitions. In some Ginkgoopsida (Permotheca, Peltaspermaceae) the sizes of lumina in the quasisaccus may vary significantly even in pollen of one sporangium, so that some grains look typically quasisaccate under the light microscope, where as others look eusaccate (Gomankov and Meyen, 1986). A similar variation is observed in conifers (Aethophyllum stipulare; Grauvogel-Stamm, 1978, pl. 25, figs. 3-7). There are all reasons to consider that in conifers, as in Cordaitanthales and Ginkgoopsida, transformations linking eu- and quasisaccate types of organization were reversible (Meyen, 1984). Therefore none of the types can be regarded as more primitive. It is to be noted that the organization of pollen in Archaeopteridales (Pettitt, 1966) remains quasisaccate. In other progymnosperms a cavate pollen is known. Thus, the exine exfoliation and the formation of a hollow was a fluctuating character as early as in Devonian progymnosperms.

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A subdivision of the eu(quasi)saccus into two lobes with the formation of disaccate or quasidisaccate pollen had happened secondarily. Already in Potonieisporites-like pollen a bilateral symmetry is outlined. Further transformations in this character are also reversible. For example, the quasimonosaccate organization of pollen of some species of Tsuga has undoubtedly arisen secondarily from a disaccate type. The appearance of ribs on the distal side is a relatively advanced character. The ribbed (striated) pollen was produced by the genus Rissikia (Townrow, 1967a). Wide taeniae in the Upper Permian pollen Scutasporites and Lueckisporites are of the same structure as proximal ribs. The genus Taeniaesporites (Permian-Triassic) may well belong to conifers also. In later conifers both ribs and wide taeniae disappeared. But in the Triassic there are conifers producing asaccate pollen with ribs observed on all sides of the grain (pollen of Equisetosporites type from microstrobili Masculostrobus clathratus Ash, Upper Triassic of Arizona; Ash, 1972). A primitive character is the preservation of a proximal aperture characteristic of many Palaeozoic and some Triassic conifers. The gradual disappearance of the proximal aperture proceeded later in conifers than in Cordaitanthales and Ginkgoopsida. The distal aperture of Palaeozoic and Triassic conifers is very poorly studied and its ultrastructure is nearly unknown. The absence of the exine thinning at the distal side in the Upper Carboniferous Potonieisporites associated with "Lebachia" lockardii (Mapes and Rothwell, 1984, pl. 16, fig. 4) and the Upper Permian Nuskoisporites associated with Ortiseia (ClementWesterhof, 1984, pl. 32, fig. 3) suggests a proximal germination of these grains (i.e., the belong to prepollen rather than pollen, see Chaloner, 1970). No traces of a distal aperture are observed also in the pollen (prepollen) of Timanostrobus and

Kungurodendron. 6.4. Vegetativeshoots Material on the morphology of vegetative shoots of Upper Palaeozoic and Mesozoic conifers are

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very abundant. The following analysis concerns only some characters. At the outset, difficulties in distinguishing vegetative leaves of conifers and Dicranophyllales are to be underlined. As has been recently shown (Meyen and Smoller, 1986), dicranophyllalean vegetative leaves are very diversified and moreover, demonstrate an essential parallelism with coniferalean leaves. In gross morphology, shoots of Mostotchkia and Slivkovia resemble coniferalean shoots and differ only in the presence of dorsal stomatiferous furrows. Entsovia leaves with numerous veins are also similar to analogous coniferalean leaves and differ in the same character. Dicranophylls are characterized by the same marginal microdenticulation and papillosity of subsidiary cells of the stomata as many older conifers. There are other common features both of leaves and leafy shoots between the two groups. Therefore, a possibility exists that some genera traditionally placed among conifers (e.g., Carpentieria and Lecrosia) belong to the Dicranophyllales. It is important that some Dicranophyllum produced simple (non-forked) leaves and the stomatal bands may be surficial. Shoots combining both characters would be gross-morphologically indistinguishable from coniferalean. Branching of shoots. The earliest conifers of the Lebachia and Ernestiodendron types differ from cordaitanthaleans in having a distinctly pinnate branching of the shoots of penultimate order, i.e., they are plagiotropic. Shoots with twice pinnate branching in one plane are known. In these older forms certain violations of the pinnate order may be observed. A poorly ordered branching is known in many Permian genera (Sashinia-Quadrocladus,

Pseudovoltzia, Ullmannia, Timanostrobus, Concholepis). Both branching types - - pinnate and poorly ordered - - persist until the present time. This character is at best of a generic value. Interestingly, the shoots with regular pinnate branching are not characteristic for Kazanian and Tatarian conifers of Western Angaraland, and the Upper Palaeozoic conifers of Gondwana. Among older conifers (as in angiosperms) a reduction of branching and a general trend to

somatic reduction is observed. Triassic Aethophyllum produced herbaceous annual aerial shoots (Grauvogel-Stamm, 1978), stems of Yuccites were unbranched (L. Grauvogel-Stamm, pers. commun.), devoid of secondary wood and probably succulent. The maximum diversity of life forms of conifers seems to fall in the Middle Triassic. Also in the Triassic poorly ordered branching, and a differentiation of shoots into dwarf (mostly lateral) and long shoots begins (Sashinia-

Quadrocladus, Concholepis). Leaves. Acquaintance with the Palaeozoic conifers does not answer the question of the origin of their leaves. Admitting the origin of conifers from cordaitanthaleans, and considering the similarity between coniferalean leaves and sterile scales of cordaitanthalean fructifications, one can suggest that the needle-shaped coniferalean leaves had originated as a result of the distribution of the cataphyllary ontogenetic programme on to the vegetative leaves (Rothwell, 1982b; Meyen, 1984). We cannot exclude, however, a common origin of coniferalean and dicranophyllalean leaves. The dicranophyllalean and cordaitanthalean leaves might have arisen through a phyllodization of lagenostomalean petioles (Meyen, 1984; Meyen and Smoller, 1986). Later on, a reduction of the phyllodia proceeded. Nevertheless, it seems suggestive that in the most primitive conifers cataphylls are absent, and the degree of difference between circasperm members and vegetative leaves is insignificant. Therefore, the former viewpoint on the origin of coniferalean leaves appears more plausible. Specialized scales accompanying fructifications, e.g. circasperm members of Willsiostrobus denticulatus (Grauvogel-Stamm) GrauvogelStamm et Schaarschmidt (Grauvogel-Stamm, 1978, p. 120, text-fig. 29a) or scales of dwarf shoots of Pinaceae are obviously of later and secondary origin. The earliest conifers are characterized by dimorphic leaves (Lebachia, Swillingtonia; Scott and Chaloner, 1983): leaves of branches of the last order are simple, whereas those of preceding orders are forked, of the Gomphostrobus type. This dimorphism soon disappears, but many Late Palaeozoic and Triassic conifers display strong intraspecific

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variations both in size and form of leaves. In

Ernestiodendron, Taxodiella, Kungurodendron and, partly, Quadrocladus the leaf dimensions regularly decrease from thicker to thinner branches. A distinct heterophylly may occur within a single branch, it is poorly expressed in Quadrocladus, and conspicuous in Voltzia and Pseudovoltzia. All the above mentioned characters are of a low taxonomic weight, although the regular decrease of the leaf dimensions in thinner branches is a primitive character. In the oldest conifers of the Westphalian (Rothwell, 1982b; Scott and Chaloner, 1983; the author's own observations on the Middle Carboniferous conifers of Middle Asia and Kazakhstan) the leaves are either adpressed to the axis and narrow-triangular, simple and forked, or divergent, awl- or S-shaped. In all of them the base is wide, not tapering or faintly narrowing. The leaves are fiat, or rhomboid and then transversely elongated in cross section. Shorter leaves (of the Brachyphyllum type) first appeared in the Permian (see description of Timanostrobus; White, 1929; Zalessky, 1939). The Upper Permian and Triassic Quadrocladus may display leaves round in cross section, sometimes with a narrow upper surface and much wider and convex lower one. A distinct petiolar narrowing of the leaf base appeared as late as the Triassic (Albertia, Aethophyllum, Dechellyia, some Elatocladus). In the Triassic, the first conifers with bilaterally (not dorsiventrally) flattened leaves appeared (Rissikia; Townrow, 1967a). An undoubtedly advanced character of conifers independently appearing during their phylogeny, is venation consisting of several (sometimes numerous) parallel veins. In the Lower-Middle Triassic Aethophyllum and Albertia the veins enter the lateral margins. In younger Podozamites, Heidiphyllum, Araucariodendron, Agathis, some Podocarpus, etc., all the veins are directed towards the apex where they converge. The same transformation in venation is observed in Dicranophyllales. In the most primitive forms (Dicranophyllum known since the Serpukhovian, i.e. Namurian A) there is only a midrib dichotomizing following the division of the leaf lamina (as in Gomphostrobus-like leaves of Lebachiaceae). In the

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Late Carboniferous, leaves appear (Entsovia rara Gluch.) having numerous veins partly entering the lateral margins (Glukhova, 1978), and in the Early Permian, leaves (E. kungurica Meyen) with all veins directed to the apex. Shoots with leaf cushions and distinct leaf scars suggesting the presence of an abscission layer, are known since the Late Permian (Quadrocladus,

Geinitzia subangarica). A primitive character is the cuticular marginal microdenticulation known in the Westphalian Swillingtonia, and many other Upper Palaeozoic conifers and Dicranophyllales. In Cordaitanthales this character is extremely rare (the author knows a single case of microdenticulation that has been recorded in Cordaites from the Upper Permian of the Kuznetsk basin; this specimen was discovered by M.P. Kushnerev in 1984). In the Permian, there were also conifers with a smooth leaf margin. This character may vary even within one shoot (see Quadrocladus). The microdenticulation may also occur in bracts, circasperm members, and microsporophylls. Epidermal characters. Leaves, bracts, circasperm members and seed stalks show epidermal features with a systematic value. The pertinent set of epidermal characters is known in Lebachia, Ortiseia, Kungurodendron and Sashinia. Epidermal characters of vegetative leaves were described in "L."

lockardii, Ernestiodendron, Timanostrobus, Cyparissidium entsovae, Taxodiella, Pseudovoltzia, Ullmannia, Voltziopsis, Buriadia, Paranocladus, Walkomiella and many younger conifers. We shall deal below with epidermal characters of only older conifers. In the putative ancestors of conifers, whether Cordaitanthales or Dicranophyllales, the leaves are hypostomatic, the stomata are rare or absent on the upper leaf surface. Dicranophyllales and a part of the Cordaitanthales are characterized by compact stomatal bands sometimes sunken into dorsal furrows. In other Cordaitanthales, the stomata form loose bands with a more or less regular stomatal files. A totally disordered arrangement of stomata is unknown among the Cordaitanthales. The stomata of Cordaitanthales and the majority of Dicranophyllales are longitudinally orientated.

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In the oldest conifers Swillingtonia from the Westphalian B (Scott and Chaloner, 1983), the leaves are hypostomatic. The stomata are arranged into two wide bands and are longitudinally orientated. The leaves of "Lebachia'" Iockardii have similar stomatal bands, but the upper leaf side shows additional bands and files of stomata (Mapes and Rothwell, 1984). The number of stomata in a band is lesser than in Swillingtonia. The stomatal bands of Walchia (al. Lebachia) piniformis and related species are located on both leaf surfaces. In Ernestiodendron, the stomata are arranged into files throughout the leaf. A disordered arrangement of stomata without bands and files, as well as a variable orientation of stomata first appear in the Late Permian Quadrocladus-Steirophyllum. Transversely orientated stomata are known in Taxodiella. Older conifers usually show longitudinally orientated stomata. Since the Late Permian to the recent, the conifers preserve the indicated types of the epidermal topography, although in the Mesozoic some additional types appeared. As late as the Jurassic, conifers appeared in which all the stomata are transversely orientated (Podozamites) or such orientation prevails (some Taxodiaceae).

Lebach&, Kungurodendron and other primitive conifers having actinocytic or petalocytic stomata. The published photographs do not convince, however, that the stomata of Swillingtonia are really paracytic. The cells interpreted as subsidiary cells are probably guard cells, and the cutin ridges known in the guard cells of many conifers (see, e.g., guard cells of Sequoia; Dilcher, 1974, p. 122, figs. 19, 20) have been mistaken for the outer contour of the guard cells. In support of their interpretation Scott and Chaloner refer to the alleged paracytic stomata in Drepanophycus (Stubblefield and Banks, 1978) and Alethopteris (Oestry-Stidd and Stidd, 1976). The paracytic structure of stomata in the former genus is also doubtful. As to Alethopteris, the erroneous affiliation of its stomata with the paracytic type has been convincingly shown (Reihmann and Schabillion, 1976; Mickle and Rothwell, 1982), the source of the error having been the same as in the case of Swillingtonia. Florin (1951) noted that the presence of trichomes in Palaeozoic conifers is a primitive feature that is not characteristic of living conifers.

Swillingtonia, Lebachia, Ernestiodendron, Kungurodendron and other primitive conifers are

7. Phylogenetic relations between orders Cordaitanthales, Dicranophyllales and Pinales. Early phylogeny of conifers

characteristic of smaller sizes and a higher papillosity of cells within stomatal bands. These conifers also usually show papilligers, known in the Late Permian only in Ortiseia. Circasperm members of Kungurodendron show a minute sinuosity of radial walls (Plate II, 25). Subsequently this character appeared again in the most advanced conifers, e.g. Pinaceae. The stomata of the earliest conifers may be either monocyelic or dicyclic, with greater or lesser number of subsidiaries, with varying cutinization and papillosity of the latter, the degree of the guard cell immersion may also vary. These characters cannot be organized into a code of primitiveness. Scott and Chaloner (1983) noted paracytic stomata in Swillingtonia. In none of the pinopsid genera have such stomata been recorded so far. Besides, according to the entire assemblage of epidermal characters Swillingtonia is very close to

Phylogenetic relations between cordaitanthaleans, dicranophylls and conifers were treated in earlier papers of the author (Meyen, 1984; Meyen and Smoller, 1986). The derivation of conifers from the Cordaitanthales or, to be more precise, Euramerican Carboniferous Cordaitanthaceae is based on the features of the female axillary complex, pollen, wood and certain epidermal characters (Florin, 1938, 1939, 1940, 1944, 1945, 1951). Inferences on the phylogenetic relations between conifers and dicranophylls are much more difficult due to the inadequate knowledge of the latter group. Nevertheless, the origin of conifers from dicranophylls cannot be excluded (Meyen and Smoller, 1986). The similarity of foliage between the oldest dicranophylls (genus Dicranophyllum) and conifers is striking. At least some dicranophylls seem to have produced the same racemose

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polysperms as more primitive Rufloriaceae. Dorsal furrows on leaves of Mostotchkia, certain Dicranophyllum and older Rufloria are remarkably similar. Older dicranophylls could be very closely related to the Cordaitanthales. In the Late Early Carboniferous both groups could have formed a single group of closely related plants finally diverging as late as the Middle Carboniferous. The rise of conifers also falls in the beginning of the Middle Carboniferous. Thus, another version is possible, namely the origins of the Cordaitanthales, Dicranophyllales and Pinales as independent orders at the beginning of the Middle Carboniferous from a single group of Early Carboniferous plants combining characters of all the three orders. A further phylogeny of the Palaeozoic and Early Mesozoic conifers, if reconstructed on the basis of all the above discussed characters along with the stratigraphic ranges of relevant genera (Fig. 1) appears as follows (Fig. 37). The position of Swillingtonia at the base of the phylogenetic tree is determined by (1) their low stratigraphic position (Westphalian B), (2) hypostomatic leaves as in Cordaitanthales, (3) putative association with pollen of the Potonieisporites type known since the Namurian, (4) a forked lamina of some vegetative leaves. Many characters observed in Swillingtonia also occur in "Lebachia" lockardii. These are the presence of numerous papilligers in stomatal bands, the distinctness of the stomatal bands, the presence of marginal teeth and hypodermis, etc. The forked bracts of "L." lockardii are comparable with forked leaves of Swillingtonia. "L." lockardii yielded the Potonieisporites pollen. "L." lockardii is closely related to Walchia (al. Lebachia) as shown by a wide circle of characters. The most important differences between them are the following characters of "L." lockardii: (1) the distal attachment of seeds; (2) the seed stalks are similar to those of Cordaitanthales and differ from clavate and more leaf-like seed-stalks of Walchia (al. Lebachia); (3) the occasional presence of more than one seed stalk in the axillary complex. The axillary complex structure characteristic of Walchia (al. Lebachia) has been retained in the Late Permian Ortiseia. The subsaccate pollen

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Potonieisporites and Nuskoisporites are also similar. Common features of both genera are the stomatal structure and the presence of papilligers. However, in the general topography of the epidermis and in the absence of forked leaves, Ortiseia is closer to Ernestiodendron than to Walchia (al. Lebachia). Somehow or other, the three genera are obviously related. The Late Permian Sashinia also shows primitive characters. These are numerous free seed stalks at the apex of the axillary complex, the attachment of sporangiophores to the microsporophyll stalk, a leaf-like habit and the absence of a heel in the microsporophyll distal shield. However, Sashinia seems to have undergone a certain despecialization, namely a secondary likeness of elements of the bract-axillary complex and foliage leaves. The advanced characters of Sashinia by comparison with Walchia (al. Lebachia) are the attachment of ovules under the hooked seed stalk apex, quasidisaccate pollen with taeniae, irregular arrangement of stomata throughout both leaf surfaces and their occasional irregular orientation, facultative disappearance of marginal microdenticulation, a round cross section of leaves and the presence of the abscission layer in some species, the appearance of the heterophylly and lateral dwarf shoots. Many primitive characters are also preserved in Kungurodendron, viz. the ovules are apically attached to hooked seed stalks (as in "L." Iockardii), numerous sterile scales in the axillary complex, common features in stomata and stomatal band structure with Walchia (al. Lebachia), absence of the distal aperture in pollen, and absence of the distal shield in the microsporophyll. Relatively advanced characters of Kungurodendron are a pronounced flattening of the axillary complex, a reduced habit of its circasperm members, the displacement of seed stalks on to its adaxial side, a simple (non-forked) bract, epistomatic leaves and bracts, and asaccate pollen with a strongly reduced proximal slit. The columellar-like exine could have been inherited from the Carboniferous Lasiostrobus-like conifers. The most interesting feature of Kungurodendron is the appearance of a secondary transition between seed stalks and circasperm elements. Usually such transitions between organs are treated in terms of

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'ARAUOARIAOEAE

I~XODIAOEAE

~flEIROLEPII}IACEAE~-----

PINAOEAE

6oristhen2u

# d

AE

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their homology and of a semophylogenetic links between pertinent organs. In the case of Kungurodendron, the transition between seed stalks and sterile scales cannot be treated as primary, because in all older members of Pinopsida the differences between these organs are discrete. Obviously we encounter here an introgression of ontogenetic programmes having no relation to the origins of one organ from the other. Kungurodendron can be tentatively linked with the Late Permian Concholepis in which the axillary complex had been transformed into a false (?) seed scale with residual projections on the abaxial side an along the margin. The polysperms of Timanostrobus look more primitive. The lateral polysperms are nearly identical to those ofVojnovskyaceae. A peculiar feature is the absence or a very strong reduction of bracts. In the pollen structure, primitive and advanced characters are combined as in Kungurodendron. But the foliage leaves (of the Brachyphyllum or Pagiophyllum type) are undoubtedly advanced together with their epidermal structure. Another advanced character is the poor order of branching. Obviously, the evolution of Kungurodendron, Concholepis and Timanostrobus independently proceeded form genera having abaxially attached seeds. Whether these three genera are phylogenetically united cannot be judged in the absence of data on the epidermal and pollen characters of Concholepis. In the pollen structure and some other characters, Kungurodendron is close to Lasiostrobus, on the one hand, and Kungurodendron, on the other, as shown on the phylogenetic scheme. The phylogenetic line from Walchia (al. Lebachia) to Walchiostrobus-Ernestiodendron is based on the identity of their pollen, the conservation of sterile scales in the axillary complex, the abaxial attachment of ovules to seed stalks, and common peculiar epidermal characters (stomatal structure, the presence of papilligers). Important

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differences between these genera are the fusion of seed stalks into a primitive seed scale, and an arrangement of stomata in regular files in Walchiostrobus-Ernestiodendron. Further in the line towards Pseudovoltzia, the number of vegetative scales in the axillary complex decreases and they become similar to seed stalks. Here might have proceeded the same process of the introgression of ontogenetic programmes as in Kungurodendron. In Pseudovoltzia the basal fusion of the seed scale and bract occurred. The interpretation of the Ullmannia polysperms remains uncertain (see Section 6.1). However, epidermal characters of Ullmannia and Pseudovoltzia are closely similar. The relationships between Voltzia and Pseudovoltzia in the polysperm structure is evident. On the other hand, both genera are undoubtedly related to the Triassic genera Aethophyllum, Cycadocarpidium and Albertia (i.e., genera with mostly multiveined vegetative leaves), and also Swedenborgia, Boristhenia and Voltziopsis. All these conifers lack sterile scales in the axillary complex, the bracts remains partly are fused with the seed scale, and a forked apex may appear again in the bract (Voltziopsis). This group of Triassic conifers might have given rise to the families Podocarpaceae, Taxodiaceae, Pinaceae, Araucariaceae and Cheirolepidiaceae. Conifers showing features of these families are known from the Upper Triassic. Rissikia can be affiliated with the Podocarpaceae (Townrow, 1967a); Voltziopsis is close to Taxodiaceae (Townrow, 1967b), and the Upper Triassic Compsostrobus to Pinaceae (Delevoryas and Hope, 1973). Conifers very close to the living Araucaria are known since the Jurassic (Stockey, 1982). The appearance of the Araucariaceae in the Triassic is probable. The Classopollis pollen restricted the Cheirolepidiaceae is known since the Upper Triassic. A little is known on the origins of the Buriadiaceae and Taxaceae. The Buriadiaceae are

Fig. 37. Tentative phylogeny of early conifers. K = subfamily Kungurondendroideae; L=subfamily Lebachielloideae; V= family Voltziaceae. Curved line divides groups of genera deserving separation into independent families, more advanced of which gave rise to the main families of Mesozoic and Cenozoic conifers.

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close to primitive conifers in vegetative shoot morphology, but their origin from a still undeciphered group of Palaeozoic Pinopsida cannot be excluded. The suggested origin of Lebachiaceae from Buriadia-like conifers (Pant and Nautiyal, 1967; Pant, 1977) cannot be supported by any concrete character. An opinion on the independent evolution of Taxaceae from other Pinopsida has found a wide recognition. However, many characters suggest a relationship between Taxaceae and Cephalotaxaceae, and the latter are very close to "true" conifers in a large number of characters. Harris (1976) has shown how the fructifications of taxads can be derived from those of Lebachiaceae, although he did not know that Florin had erroneously interpreted the structure of the axiUary complex of Lebachia, but the scheme proposed by Harris can be retained if we insert "Lebachia" lockardii instead of Lebachia.

8. Systematics and nomenclature of early conifers The above phylogenetic analysis implies the subdivision of better known Late Palaeozoic and Triassic conifers into four main groups: (A) Genera with apically attached ovules and numerous sterile scales in the axillary complex. These are "Lebachia" lockardii, Kungurodendron, Timanostrobus and, probably, Concholepis. The same group may comprise Swillingtonia and

Lasiostrobus. (B) Genera with abaxial attachment of ovule to seed stalk; the seed stalks are solitary, if numerous they are free. These are Walchia (al. Lebachia), Ortiseia and Sashinia. (C) Genera with a true seed scale accompanied or not by sterile scales. These are Walchio-

strobus-Ernestiodendron, Pseudovoltzia, Ullmannia (?), and all Triassic genera listed in Fig. 37, i.e., Voltzia, Aethophyllum, Yuccites, etc. (D) Genera with ovules that are not aggregated into polysperms. These are Buriadia, Walkomiella and Paraburiadia. These four groups may be given a status of independent families. However, the groups A and B share many important features. Therefore they can be better treated as subfamilies of a

single family. Group C comprises genera currently referred to the families Lebachiaceae = Walchiaceae (Ernestiodendron, Walchiostrobus), Voltziaceae (Pseudovoltzia, Ullmannia, Voltcia, Voltziopsis) and Cycadocarpidiaceae = Podozamitaceae (Cycadocarpidium, Swedenborgia, Boristhenia). Considering close links between these genera as shown by different characters, they may be united into a single family. Group D stands apart and corresponds to the family Buriadiaceae. Pant (1977) suggested to raise its rank to the order level, but before that we need a fuller knowledge of the relevant genera, particularly those mentioned by Archangelsky (1985) and awaiting a full description and validation. Other previously proposed families and orders of older conifers are omitted from consideration (Ullmanniaceae, Voltziales, etc.), because they had been established on Florin's erroneous treatment of pertinent female fructifications. The selection of names for the suggested families and subfamilies is easy in the case of groups A and D. Within group A the better known and more peculiar is the genus Kungurodendron. Accordingly, the subfamily name would be Kungurodendroideae. For group D the name Buriadiaceae is available. It is to be noted that the name Buriadiales had been proposed for the other (Pant, 1977), but the corresponding family, as often happens in palaeobotany, had not been proposed. The name Buriadiaceae is included in the system of conifers proposed by the author (Meyen, 1984), but whether anybody had introduced this name earlier is unknown to me. The situation with names for the other two groups is much more complicated. According to the nomenclatural rules, the family comprising the groups A and B should be named Walchiaceae (Clement-Westerhof, 1984), instead of Lebachiaceae as adopted in the literature. Then the subfamily name should be Walchioideae. Both the taxa should be typified by the generic name Walchia having a priority, and the name Lebachia should be abandoned. This decision creates serious nomenclatural complications, however. The literature yields more than 60 species of Walchia, of which only 4 can be referred to Walchia as a synonym of Lebachia (Clement-Westerhof, 1984).

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Most of the other species prove to be without a suitable generic name, because their fructifications are unknown. It is true that Clement-Westerhof (1984) proposed to refer a part of the species of Lebachia and one species of Walchia to the genus Culmitzschia on the basis of epidermal characters. Species with unknown epidermal structure prove to be beyond the system of genera. This problem can be resolved in two ways. The least radical decision is a conservation of the names Lebachia and Lebachiaceae widely circulating in the literature. Then the change of the type species of Walchia will be necessary. W. filiciformis has already been proposed for this role. In such a case the generic name Ernestiodendron should be abandoned. For polysperms associated with Ernestiodendron the name Walchiostrobus is available. Therefore the rejection of the name Ernestiodendron will not influence the nomenclature of fructifications, that is agreeable. On the other hand if we want to keep the name Walchia for numerous species currently referred to this genus, we have to exclude epidermal characters known in the type species W. filiciformis = Ernestiodendron filiciforme sensu Florin, from the generic diagnosis. This decision will entail both taxonomical and nomenclatural complications. If the system of the vegetative shoots of Palaeozoic conifers does not involve epidermal characters, it should be put into agreement with the already available analogous system of Mesozoic genera. Epidermal characters have been studied in many Palaeozoic species, and should be used for distinguishing species and genera. Thus, the conservation of the name Lebachia would create some more difficult questions. Besides, the very attempts at conservation are not always successful. Chances that the conservation of Lebachia and Lebachiaceae will be made are not good. One of the arguments against the conservation of Lebachia and Lebachiaceae is that the nomenclatural types (i.e., holotypes of types species) of Walchia, Lebachia and Ernestiodendron are vegetative shoots. Their connection with certain fructifications is known in the case of Walchia-Lebachia and is less evident in the case of Ernestiodendron. The lectotype of Walchia (al. Lebachia)piniformis (Florin, 1938, pl. 1/2, figs. 1, 2) is badly preserved,

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and its cuticle cannot be studied. The lectotype of the type species Ernestiodendron filiciforme has been lost. Unfortunately, other Late Palaeozoic and Triassic genera arranged into the system of conifers according to fructification structure are also typified by sterile shoots devoid of epidermal characteristics, the type material is often missing (or its location is unknown), and the lectotypes of the type species have not been selected. This pertains, among others, to Pseudovoltzia, Ullrnannia and Voltzia, although all three genera are highly important in the systematics of earlier conifers. The separation of nomenclature of vegetative shoots and female fructifications has not been practized for Pseudovoltzia, Voltzia and Ullmannia, whereas the male fructifications of Voltzia and Ullmannia have received separate generic names. This practice has resulted in serious confusion. For instance, the genus Ullmannia was mentioned in many Upper Palaeozoic floras, although both the pollen and polysperm structure of the type species are unknown, being recorded only in U. frumentaria. When we say that Ullmannia is a widespread genus belonging to the family Voltziaceae, we proceed from a dubious admission that all shoots of the Ullmannia type associated with the same polysperms as in U. frumentaria although this may not be so (see Section 5.8). All of this happened with conifers due to the typification of main taxa by vegetative shoots and has parallels in the articulates (Meyen, 1971b; Grauvogel-Stamm, 1978), lycopsids (Meyen, 1976; Thomas and Brack-Hanes, 1984), cordaitanthaleans (Meyen, 1972, 1984; Maheshwari and Meyen, 1975) and other plants (see detail in Meyen, 1978a, 1984). Therefore, a tendency gradually develops towards a rearrangement of the system of extinct supraganaric taxa in order to typify them by genera and species established on fertile parts (Grauvogel-Stamm, 1978; Meyen, 1978a, 1984; Thomas and Brack-Hanes, 1984). To this aim the previously selected type genera should be withdrawn from newly established taxa (to avoid the rejection of their names on the basis of priority) and treated as informally associated (satellite) in relation to a certain order or family (see

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details on satellite taxa in Meyen, 1978c; Thomas and Brack-Hanes, 1984). Apparently, the same procedure should be adopted in the case of conifers. This means that their system (and phylogeny) should be based on genera typified by fertile material. Suprageneric taxa should be typified by such genera and species that have as many characteristics as possible (female and male fructifications, pollen, vegetative shoots and their epidermal characters) and do not include species established on vegetative shoots alone. The question of the classification of vegetative shoots stands apart. When their association with fructifications is quite evident and if both kinds of remains come from the same localities (as in the case of Timanostrobus muravievii and Kungurodendron sharovii), a separate nomenclature for both is unnecessary. However, if in some localities the vegetative shoots are not accompanied by relevant fructifications, we need a separate nomenclature for the vegetative shoots. The latter should be referred to a taxon typified by a vegetative shoot (as in the case of Quadrocladus dvinensis that is not always accompanied by Sashinia aristovensis; for the same reasons some of the earlier described species were referred to Cvparissidium and Geinitzia). Vegetative shoots, the association of which with certain fructifications is unknown or doubtful, should be referred to genera of their own. Principles of classification of such material were treated in detail by Harris (1969, 1979) with presumably Mesozoic conifers in view. Earlier Florin and other palaeobotanists began to establish independent genera of Mesozoic conifers based on epidermal characters. This practice has resulted in problems of demarcation between the genera as well as of the search of suitable genera in the literature. Harris suggested a system of a few genera distinguished by selected gross morphological characters. The epidermal characters were suggested for the demarcation of species. If we adopt the generic diagnoses proposed by Harris, the species of the genus Walchia available in the literature can be distributed in the same system of genera. For instance, W. appressa Zalessky can be referred to Cyparissidium (see Section 5.2.1 ), W.

laxi)blia (Florin) N~mejc, to Geinitzia, etc. Such passing of all species of Walchia through Harris' system of genera would be reasonable. A separate approach can be adopted to species having epidermal characteristics. Harris rejected epidermal features as a basis for establishing genera. However, epidermal characters can be treated in the same way as Harris approached the gross morphological characters; that is we can use a limited number of better defined (mostly topographical) characters. Both their selection and introduction of the relevant system of genera are beyond the scope of the present paper. Still, considering the very possibility of such an approach, the genera Quadrocladus and Taxodiella are retained independently. Thus, the system of better studied Palaeozoic and, partly, Triassic conifers can be built in the following way. First of all, the family Buriadiaceae (genera Buriadia and Walkomiella) can be established. On the basis of epidermal characters and phytogeographic affinity, Paranocladus can be regarded as a satellite genus of this family. Paraburiadia (Zimina, 1983) is another satellite genus. The genus Walchia with the type species W. pinijbrmis (Schlotheim) Sternberg is retained for vegetative shoots alone, because such is the type material. The most important diagnostic characters of this genus in this understanding are pinnate branching of shoots and forked leaves on the shoots of penultimate and higher orders. All species of Walchia having only simple leaves should be distributed among genera according to Harris' system. The polysperms described and figured by Florin and referred by him to this species should be referred to a separate genus, for which the name Lebachiella Meyen, gen. nov. with the type species Lebachiella florinii Meyen, sp. nov. is here proposed. The associated microstrobili can be placed into the genus Walehianthus Florin (1940). The genus Ernestiodendron comprises only vegetative shoots, including those having epidermal characteristics. The associated female fructifications are referred to the genus Walchiostrobus Florin (1940) with the type species W. gothanii Florin. Polysperms associated with Pseudovoltzia,

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Ullmannia and Voltzia should be also ascribed to independent genera. The genera Lebachiella, Ortiseia and Sashinia compose the subfamily Lebachielloideae that, together with the subfamily Kungurondendroideae, constitute the family Lebachiellaceae. It is true that the genus Ortiseia is better studied and can be selected for typification, but its nomenclatural type (holotype of O. leonardii Florin) is a vegetative shoot. The family Voltziaceae is retained for the present in the above explained bulk. Its name needs to be changed, because the type genus Voltzia has unsatisfactory type material and can be at best a satellite genus of a family. If the emended family includes the genus Cycadocarpidium, the name Cycadocarpidiaceae can be adopted, and the genus Voltzia, retained for vegetative shoots alone, will be a satellite genus of this family. But before doing so, we need more reliable data on the type species Cycadocarpidium erdmannii Nathorst (1886), the interpretation of which by Florin (1944) adopted in the literature, is doubtful. Probably, this species did not have free seed stalks as figured by Florin (1944, text-fig. 60a-e; compare with his pl. 183/184, figs. 4-11). For polysperms associated with Voltzia a new genus is needed. Perhaps it will be the genus comprising polysperms currently referred to Pseudovoltzia. Thus the system of early conifers acquires the following view. Family LEBACHIELLACEAEMeyen, fam. nov. Type genus: LEBACHIELLA Meyen, gen. nov. Diagnosis: Leaves simple or forked. Axis of compound polysperm continuing leafy twig, bears helically arranged bracts (occasionally probably reduced) similar to foliage leaves. In bract axils lateral polysperms (axillary complexes) are located, they bear numerous sterile scales and one to many monosperms. Microstrobili continuing leafy twigs, consist of microsporophylls with distal shield. Subfamily LEBACHIELLOIDEAEMeyen, subfam, nov. Diagnosis: Monosperms numerous or one per axillary complex. Ovules attached abaxially to the

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upper part of seed stalk. Generic composition: Lebachiella Meyen, gen. nov.; Ortiseia Florin, 1964, emend. Clement-Westerhof, 1984; Sashinia Meyen; Dvinostrobus Gomankov et Meyen. Genus LEBACHIELLA Meyen, gen. nov. 1938 Lebachia Florin (pars)

Type: Lebachiella florinii Meyen, sp. nov. Diagnosis: Genus established for compound polysperms. Axis of compound polysperm continuing leafy shoot of the last order. Bract forked, helically arranged. The axillary complex consists of the short axis bearing numerous sterile scales, and one clavate seed stalk on adaxial side. Inverted ovule attached subapically on abaxial side of seed stalk. Specific composition: The genus Lebachiella seems to comprise fertile shoots described by Florin (1939) as Lebachia hypnoides (Brongniart) Florin, L. garnettensis Florin, L. goeppertiana Florin. These shoots may constitute separate species of the genus Lebachiella, but their separation requires additional studies of the type material. Comparison: Lebachiella differs from Ortiseia in a lesser degree of flattening of the axillary complex, a forked bract, a notched seed stalk apex, a higher position of the ovule on the seed stalk, and the presence of stomatal bands on the bract, and from Sashinia in having a single monosperm without hooked apical pad, a forked bract, and a lesser similitude between bracts, circasperm members and vegetative leaves. Lebachiella flovinii Meyen, sp. nov. Holotype: Specimen figured by Florin (1938, pl. 19/20, figs. 1-20; see diagrammatic drawing of Florin (1944, text-fig. 32b-e) and Fig. 31 in the present paper. Remarks: Some of Florin's figures, particularly the drawings, were repeatedly reproduced in the literature. References on this literature (including later publications of Florin) where the cited figures are simple reproduced, have not been included in the synonymy.

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1938 Lebachiella piniformis (Schlotheim) Florin (pars), Florin, p. 60, pl. 7/8, figs. 6 (?), 7 (?), 9 11 (?), pl. 9/10, figs. 6-10, 13 (?), 16 17, pl. 11//12, figs. 4 (?), 5-7, pl. 15/16, figs. 1-30, pl. 17/18, figs. 11-24, pl. 19/20, figs. 1-30, pl. 21/22, figs. 1 19, 20-24 (?), 25 (?), pl. 27/28, figs. 7 8 (?), pl. 29/30, figs. 6-7 (?). 1944 Lebachiella piniJormis (Schlotheim) Florin (pars), Florin, text-fig. 32b-e. Etymology: After the late Prof. R. Florin. Diagnosis: Sterile leaves of fertile twigs as in Walchia piniformis sensu Florin (1938). Compound polysperms cylindrical, 4 10 cm long and 7-14 cm wide, continuing shoots of the last order. These shoots are arranged in two ranks on axes of penultimate order. Bracts of the Gomphostrobus type (same as leaves on branches of penultimate order), 7-11 mm long, 1.4 3 (up to 6 ) m m wide near the base, forked, bifacial, helically attached, diverging, widened towards the base, with one forked vein, each vein branch directed to the lobe of the forked lamina. Bracts amphistomatic, bear adaxially two papillose stomatal bands merging basally and directed to lamina lobes, one band per lobe. Stomata mostly longitudinally orientated. Upper epidermis with numerous papilligers (or trichopodia). Stomata the same as in foliage leaves. Lower epidermis without stomata in middle and apical parts, and with two groups of distant and deformed stomatal files associated with numerous papilligers (or trichopodia) near the base. Margin with cuticular teeth or unicellular hairs. Sterile scales of axillary complex numerous, become smaller towards its base and apex, the largest in its middle part, up to 5 mm long and 1.5 3.2 mm, narrow triangular to oval, with a blunt or acuminate apex, amphistomatic. Stomatal files more or less regular. Epidermal cells papillose, there are trichopodia. Seed stalk strongly flattened, clavate, with notched apex under which the seed scar is located. Epidermis with stomata arranged in short files, otherwise similar to epidermis of sterile scales. Remarks: A more detailed diagnosis concerning the vegetative leaves, bracts and scales of the axillary complex is given by Florin (1938, pp. 60-62) who described the seed stalk as an ovule (Samenanlage: Florin, 1938, pp. 61-62).

Male fructifications and pollen are not included in the diagnosis of the species. Subfamily

KUNGURODENDROIDEAE

Meyen,

subfam, nov.

Type genus: KungurodendronMeyen, gen. nov. Diagnosis: Seed stalks with apically attached ovules. Generic composition: Kungurodendron Meyen, gen. nov., Timanostrobus Meyen, gen. nov., Concholepis Meyen, gen. nov. (?). Herein also belongs "Lebachia" lockardii Mapes et Rothwell, 1984 and, probably, Lasiostrobus. The former should be referred to a new genus. Satellite genera of the Jamily Lebachiellaceae: Walchia Sternberg, 1825, non sensu Florin; Quadrocladus M~idler, 1957; Carpentieria N6mejc et Augusta, 1934; Culmitzschia Ullrich (sensu Clement-Westerhof, 1984); Lecrosia Florin, 1940; Swillingtonia Scott et Chaloner, 1983; Lasiostrobus Taylor, 1970. Family VOLTZIACEAEArnold, 1947

Diagnosis: Compound polysperms on leafy twigs. Bracts helically arranged, free or basally fused with base of seed scale, needle-shaped or wider and then with numerous veins, less frequently forked. Axillary seed scales lobed in varying degree or entire (?); in more primitive forms accompanied by sterile scales belonging to axillary complex. Ovules attached adaxially to lobes of seed scale or to middle part of entire seed scale (?). Microsporophylls with thin stalk and distal shield, often peltate (when shield has a heel). Microsporangia attached to the distal shield under stalk attachment or near the stalk, or to branched sporangiophores inserted on the stalk in its middle part. Leaves needle-shaped or wider, with one or numerous veins. When numerous, veins directed to leaf apex or entering lateral margins. Generic composition: Voltzia Brongniart, 1828; Walchiostrobus Florin, 1940; Pseudovoltzia Florin, 1927; Voltziopsis H. Potoni6, 1899; Swedenborgia Nathorst, 1876; Aethophyllum Brongniart, 1828, emend. Grauvogel-Stamm, 1978; Cycadocarpidium Nathorst, 1886; Boristhenia Stanislavsky, 1976.

s. v. Meyen / Reviewof Palaeobotanyand Palynology96 (1997)351-447 Satellite genera: Ullmannia Goeppert, 1850; Yuccites Schimper et Mougeot, 1844; Willsiostrobus Grauvogel-Stamm et Maubeuge, 1969, emend. Grauvogel-Stamm, 1978; Sertostrobus Grauvogel-Stamm, 1969; Albertia Schimper, 1837; Podozamites (Brongniart) C.F.W. Braun, 1843; Lindleycladus Harris, 1979. Remarks: The present family can be subdivided into two subfamilies (or even independent families). One of them may comprise more primitive forms with free (or slightly fused with axillary complex) bracts, vestigial elements of the circasperm, and more closely related to the Lebachiellaceae in epidermal features (particularly in the presence of papilligers), marginal microdenticulation and the pollen structure. This concerns, first of all, Walchiostrobus and Pseudovoltzia. These subfamilies cannot be established right now, because many genera, particularly the genus Voltzia, have been poorly studied. As was said earlier, the family should be renamed and typified by a genus established on female fructifications. In this case the genus Voltzia will be a satellite of the new family.

9. Florogenetic conclusions. The Subangara area in the system of Late Palaeozoic phytochoria

Most Permian conifers of Western Angaraland were traditionally referred to genera common in Western Europe (Ullmannia, Walchia), including Mesozoic genera (Voltzia, Pityophyllum, Brachyphyllum). Endemic genera established by Zalessky (1937, 1939) are rare. They were based on gross morphological features of vegetative shoots, and their relations to West European conifers is uncertain. Results of the present study allow us to conclude that the systematic composition of conifers of Western Angaraland (Subangara area) differs drastically from that of Western Europe (Euramerican area of the Earliest Permian, and Atlantic area or kingdom of the remainder part of the Permian - Meyen, 1978b). We should bear in mind, however, that this comparison pertains to partly asynchronous assemblages of conifers (Fig. 1 ). In Western

439

Angaraland there are no conifers coeval to the Autunian, and in Western Europe - conifers coeval to the Kungurian, Ufimian and, probably, Kazanian. In coeval deposits of both areas there are the common Upper Permian genera Quadrocladus and Pseudovoltzia (?). An essentially closer relationship between Euramerican + Atlantic and Western Angara (Subangara) conifers is indirectly evidenced by a largely similar composition of the dispersed gymnosperm pollen (Visscher, 1971; Chuvashov and Dyupina, 1973; Hart, 1974; Chuvashov et al., 1984). As well seen in Fig. 1, the presently described Western Angara conifers fill the hiatus in the general succession of Western European and North American conifers. One can infer from the previous sections that the conifers of Western Angaraland are unexpectedly primitive. The most advanced among them is Pseudovoltzia ? cornuta that is essentially more primitive than P. liebeana from the Western European Zechstein. The primitiveness of Sashinia dominating among conifers of Western Angaraland at the end of the Permian is really surprising. The coniferalean phylogeny shows (Fig. 37) that the Subangara genera had not come into the mainstream of the evolution of Pinales and belong to lateral lines. The main coniferalean evolution during the Permian appears to have proceeded within the Equatorial belt. The ecotonal regions between the Angara area and the Equatorial belt in the Carboniferous, i.e. Urals, Kazakhstan and partly Middle Asia, seem to have played a more important role in coniferalean evolution. This is evidenced by both palynological data and megafossil findings. A large number of quasi(sub)saccate pollen probably belonging to the Pinales (including pollen of the Potonieisporites type) appear on the western slope of Urals since the middle of the Moscovian (Chuvashov et al., 1984), i.e. since the Westphalian C - D (Wagner and Bowman, 1983). Conifers similar to Ernestiodendron and Walchia (al. Lebachia) appear in Middle Asia since the upper part of the Moscovian stage. T.A. Bykovskaya (Tashkent) demonstrated to the author numerous conifers from the Takhtek locality (South Ferghana) belonging to the Upper Moscovian substage (Sixtel et al., 1975). Savitskaya and Iskandarkhodzhaev

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(1984) refer the appearance of conifers in Middle Asia to the Moscovian stage. M.I. Radtchenko (Alma-Ata) demonstrated to the author both vegetative shoots and compound polysperms of conifers (they seem to belong to a new genus) from the Zhamanbulakskaya suite of Dzhungaria. This suite yields a diversified marine fauna of the upper part of the Moscovian stage, i.e., also corresponds to the Westphalian C-D. The Westphalian D of Oklahoma also yields Walchia-like conifers (Rothwell, 1982b). From this level and onwards, conifers (both vegetative shoots and putative coniferalean pollen) are particularly abundant in non-coal-bearing regions (such as the Urals, Kazakhstan, Dzhungaria, southeast of Kansas, east of Oklahoma) and barren members of coal-bearing basins (Donets basin, some basins of Central French Massif, Czechoslovakia). Apparently the flourishing of conifers was promoted by more arid environments and better drained biotopes. It seems suggestive in this respect that the oldest coniferalean megafossils from the Westphalian B of England are allochthonous and had been transported from more elevated, hence drier and better drained habitats (Scott and Chaloner, 1983). Interestingly, in the Upper Palaeozoic of Gondwanaland, the conifers do not belong to plants avoiding coal measures. Buriadia and Walkomiella occur in India in coal-bearing formations. Northern conifers become more common in plant assemblages of coal-bearing deposits only from the middle part of the Triassic (from the Lower Keuper), and become one of the dominating group in such assemblages only since the Jurassic, i.e., when the rise all the main families had been completed (Florin, 1963 ). For the present it is difficult to judge as to how widespread were the conifers described in this paper within the Subangara area or elsewhere. Such judgements should rest on fructification findings rather than on the external similarity of vegetative shoots. Such data on the Subangara conifers outside the Russian Platform and ForeUrals are practically absent (Meyen, 1982a). For the same reason we cannot draw other geographical parallels, e.g., with numerous coniferalean remains known in the Lower Permian of Cathaysia (Bohlin, 1971, 1976; Gu and Zhi, 1974), the Far

East, Kazakhstan, Middle Asia and the Pechora Fore-Urals. In the coal-bearing Upper Palaeozoic of Siberia indisputable conifers are unknown. Affinity with conifers of Ullmannia bugarichtaensis Chachlov coming from the Upper Permian of Tunguska basin (Rasskazova, 1963) has not been proved. Some Upper Palaeozoic Siberian plants previously placed among conifers belong to the articulates ( Meyen, 1982a). A large number of conifers appear in Siberia as late as the Permotriassic in a flora which can be called Korvunchanskaya. It comes from volcanics and other coalfree deposits. Here again we observe, however, mostly primitive conifers with Quadrocladus foliage and Lueckisporites pollen (Meyen, 1981). It is true, that from the same beds, Voltzia and the fructifications Darneya, Willsiostrobus and Sertostrobus were noted (Dobruskina, 1984), but the pertinent material is not sufficiently well preserved or has not been studied in detail. The above identifications of the European genera need further support. The identification of the genus Voltzia cannot be supported by epidermal studies, because the epidermal structure of its type material is unknown. Even more significant is the fact that the available palynological data (Obonitskaya, 1974) contradict the alleged parallels between the assemblage of Siberian Permotriassic conifers and the Triassic conifers of Europe, and rather suggest links with the Permian Western European (Zechstein) and Western Angara conifers. Thus, the cradle of conifers was located in more drained habitats of the Equatorial belt. The maximum diversity of living conifers also falls in areas with a relatively dry and warm climate (Florin, 1963). The living conifers are most abundant in number of individuals in Northern extraequatorial humid areas, but their taxonomic diversity in such areas is not high. Strikingly dominating are a few genera of Pinaceae and Taxodiaceae, both families having been of equatorial origins. From phylogenetic and florogenetic viewpoints, the coniferalean forests of the recent Holarctics are rather primitive. A large number of conifers present in the Permian of the Subangara area can be offered as an argument against the affiliation of this area with the Angara kingdom. As was said earlier,

S. K Meyen / Review of Palaeobotany and Palynology 96 (1997) 351-447

conifers are absent from the typical Angara flora of Siberia. On the other hand, the Subangara conifers differ essentially from Euramerican and Atlantic ones. They are mostly represented by different genera and are less advanced. In the Late Early and Late Permian, the Subangara area contained many typically Angara taxa particularly those in common with the Pechora province (Meyen, 1982a). The phytogeographic affinity of earlier floras which inhabited the eastern part of the Russian Platform and Fore-Urals might have been different. Unfortunately, these floras can be judged by palynological data alone and suggest a more pronounced affinity with the Euramerican flora. The Carboniferous miospore assemblages of the Urals share many features with Euramerican ones, mostly at the expense of quasisaccate (or subsaccate, "quasisaccate" is used below to cover both) gymnosperm pollen which dominates in the assemblages of the Urals and is of subordinate role in Europe. The Upper Middle and Upper Carboniferous assemblages of the Urals are dominated by striated and non-striated quasisaccate pollen and are more similar to the Permian assemblages of Europe and Fore-Urals, than to coeval European assemblages (Chuvashov and Dyupina, 1973; Chuvashov et al., 1984). This used to involve stratigraphic errors. Miospore assemblages of the same habit are traced further to Kazakhstan. The already mentioned Zhamanbulakskaya suite yielding Middle Carboniferous marine fauna, had erroneously been assigned by palynologists to the Artinskian-Kungurian. Obviously the pertinent territory can be regarded as an independent phytochorion (province or area) that may be related to the Euramerican area in the same way as the Subangara area of the later period--to the Angara area. Unfortunately the miospore data do not allow us to judge regarding the presence of Angara taxa in this Carboniferous phytochoria. Both the miospore assemblages of the same composition and plant megafossils are known in the Zhamanbulakskaya suite of Dzhungaria, where undisputable Angara plants are lacking. For the present, this phytochorion cannot be named "Subangara". The name "Ural-Kazakhstanian"

441

earlier used by Radczenko (1966) for the Early Permian is available and can temporarily be used until fuller data on the extent of the phytochorion are obtained. The role of conifers in the Ural-Kazakhstanian flora was probably significant, but the quasisaccate pollen dominating in the pertinent miospore assemblages might have also been produced by Cordaitanthales, Dicranophyllales and older Peltaspermales (Particularly Trichopityaceae). Some of the plants characteristic of the Permian Subangara area, including conifers, might have been derivates of the Ural-Kazakhstanian flora of the Carboniferous. For the present it is merely a guess. The available data on the floras occurring between typically Angara and Euramerican floras are very scanty, and filling this gap in our knowledge is an exciting topic of future palaeofloristic studies.

10. Conclusions

The results of the present study coupled with the latest data on the European and North American material widely extend the diversity of early conifers. This pertains particularly to female fructifications and pollen. The interpretation of the female axillary complex suggested by Florin and widely circulating in the literature, is to be modified. The most important correction is that the seed scale has originated from fused seed stalks rather than at the expense of sterile parts of the ancestral axillary complex. The process of seed scale formation commenced very early among primitive conifers of the WalchiostrobusErnestiodendron type. A separate group of conifers having numerous free seed stalks persisted throughout the Permian and was particularly characteristic of Western Angaraland. Some of its members produced a peculiar asaccate pollen hitherto unknown in early conifers. The newly obtained data are not in conflict with the currently adopted derivation of the Pinales from the Cordaitanthales. On the other hand, the Pinales prove to be closely related to the DicranophyUales. Conifers might have originated from an ancestral Early Carboniferous group com-

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bining characters of all three orders. Further evolution of conifers proceeded in several lineages diverging during the Permian. One lineage led to the Voltziaceae and further to the majority of the living families. Another lineage gave rise to taxads. The remainder were blind alleys expired before the end of the Palaeozoic or during the Early Triassic. The general system of fossil conifers requires an essential revision. The suprageneric taxa should be based on genera established on female fructifications when the associated sterile shoots, male fructifications and pollen are also known. As a first step in this direction the family Lebachiellaceae with two subfamilies is proposed. The family Voltziaceae should be replaced by a newly established family typified by female fructifications. Some genera occupying a key position in coniferalean phylogeny, particularly Ullmannia, Pseudovoltzia and Voltzia, also need both emendation and appropriate typification. Conifers first appeared in well drained habitats of the humid climatic zone. From the Middle Carboniferous until the Mid-Cretaceous they dominated in arid and semiarid areas where coal accumulation processes were suppressed. Conifers became common elements of wet habitats and humid climatic zones as late as the Triassic. The Permian conifers of Western Angaraland (Subangara area) differ from coeval equatorial conifers in their systematic composition being less advanced. Some of them might have been derivates of the Ural-Kazakhstanian flora of the Carboniferous.

Acknowledgements I am grateful to Dr. A.L. Buslovich, Dr. A.V. Gomankov, Dr. F.I. Entsova, Mrs. G.A. Ioffe, Dr. V.R. Lozovsky, Dr. M.G. Minikh, Professor I.S. Muraviev, Dr. A.G. Olferiev, Dr. V.I. Rozanov, Professor A.G. Sharov, and Mrs. M.A. Stepanenko for providing collections, and to Dr. A.V. G o m a n k o v and Mr. I.A. Ignatiev for the aid in collecting. Thanks are due to Professor Dr. M. Barthel for this help during my work in the Museum for Naturkunde and providing photo-

graphs of some specimens. I thank Miss T.A. Bykovskaya, Dr. N.K. Esaulova, Mr. T.A. Iskandarkhodzhaev, Dr. M.I. Radtchenko, Dr. K.Z. Salmenova and Mrs. L.A. Savitskaya for showing their collections. I am thankful to Dr. L. Grauvogel-Stamm and Professor H.-J. Schweitzer for fruitful discussions and unpublished information on European conifers, and to Dr. G. Mapes for providing data on some North American conifers. Special thanks are extended to Dr. B. Lugardon for the TEM study of the pollen of Kungurodendron. I owe my thanks to for going through the manuscript, correcting my English and valuable suggestions. The technical aid of Mrs. M.A. Meyen is gratefully acknowledged.

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