Fossil Flora from the Kazanian (Middle Permian) Iva ... - Springer Link

ISSN 00310301, Paleontological Journal, 2015, Vol. 49, No. 11, pp. 1193–1205. © Pleiades Publishing, Ltd., 2015.

Fossil Flora from the Kazanian (Middle Permian) IvaGora Locality, Soyana River, Arkhangelsk Region, Russia S. V. Naugolnykh Geological Institute, Russian Academy of Sciences, Pyzhevskii per. 7, Moscow, 119017 Russia Kazan Federal University, Kremlevskaya ul. 18, Kazan, 420008 Russia email: [email protected] Received September 29, 2014

Abstract—A floral assemblage from the IvaGora locality (middle reaches of the Soyana River, Arkhangelsk Region) is described. The flora includes lycopsids, sphenophytes, ferns, peltasperms (order Peltaspermales, families Peltaspermaceae and Angaropeltaceae), conifers, and vojnovskians (order Vojnovskyales). A new genus of isolated seeds, Megasylvella gen. nov., is established. Praephylladoderma sojanaeana (Zalessky, 1937) Naugolnykh, comb. nov., emend. nov. is newly combined. The new species Megasylvella ivagorica Naugol nykh, sp. nov. and Radicites trimorphus Naugolnykh, sp. nov. are described. Keywords: pteridophytes, gymnosperms, new taxa, seeds, roots, paleobotany, morphology, Permian, Kazanian DOI: 10.1134/S0031030115110076

INTRODUCTION A great number of Permian sections have been rec ognized and studied in the Russian Platform by many generations of paleontologists and stratigraphers, although these sections still allow essential expansion of our knowledge of the organic world in the Permian Period. Special attention of experts was paid to the IvaGora locality, which is situated at the middle reaches on the right bank of the Soyana River in the Arkhangelsk Region, 50 km upstream from the village of Soyana (Fig. 1). The IvaGora locality was discovered by M.B. Edemsky in 1926 (for more detail, see Shcherbakov, 2007). The Permian sections on the Soyana and Kuloi rivers were studied in detail by J.D. Zekkel in 1935 (Zekkel, 1939; Shcherbakov, 2007). The outstanding Russian paleobotanist Zalessky (1937) published the first descriptions of fos sil plants from the IvaGora locality: Pecopteris attenta Zalessky, P. spiculosa Zalessky, P. conserrata Zalessky, Odontopteris sojanaeana Zalessky, and Meristophyllum sojanaeanum Zalessky. Ignatiev and Naugolnykh (2001) reported on the IvaGora Flora in a review of Kazanian floras of Eurasia. The present study is devoted to the general charac teristics of the flora from the IvaGora locality and description of some most interesting fossil plants of the IvaGora Floral Assemblage. MATERIAL AND METHODS This paper is based on plant remains collected by the geologist V.N. Kuleshov (Geological Institute,

Russian Academy of Sciences) in the IvaGora local ity in 2004 and transferred to me for examination (for more detail, see Naugolnykh and Kuleshov, 2005). The collection comprises more than 70 imprints and phytoleims. The holotype of Megasylvella ivagorica Naugolnykh gen. et sp. nov. was transferred to me by A.V. Pakhnevich (for more detail, see Pakhnevich and Savinov, 1998). The specimens are stored in the Geo logical Institute of Russian Academy of Sciences (GIN), collection no. 4851. Additional material includes the previously examined ferns (Naugolnykh, 2013) from the IvaGora locality from collected by A.G. Sharov, collection no. 4547. The epidermal microstructure of ferns of the Iva Gora Flora was studied under a Stereoscan 600 scan ning electron microscope in the Geological Institute of the Russian Academy of Sciences. STRATIGRAPHY OF ENCLOSING BEDS Plant remains of the IvaGora locality are confined to the middle part of the section, an interbed of light beige platy marls (socalled IvaGora Beds), which also enclose remains of the hingeless brachiopod Lin gula sp., bivalves, merostomates (Paleolimulus sp.: see Shcherbakov, 2007, p. 2), and insects. Platy marls overlie with conformity red sandstones of the Shesh minskian Horizon of the Ufimian Stage. Along the strike, the IvaGora Beds are replaced easterly by limestones and dolomites with remains marine inver tebrates, the taxonomic composition of which is indic ative of the Early Kazanian age of the IvaGora Beds (Likharev and MiklukhoMaklai, 1977).

1193

1194

NAUGOLNYKH

16 15 14 13 12 400 0 400 1200 km

30°

11 10 KZ UF

40°

50°

60°

9 8 7 6

70°

Vorkuta

Arkhangelsk

60° 10 m

1

St. Petersburg

2

Perm Kazan

3 1m 5

Moscow

4

Samara

5 4 3 2

70°

6 Volgograd

7 8

RostovonDon

1 Fig. 1. Geographical and stratigraphic position of the IvaGora locality (marked by asterisk). Designations: (1) limestone; (2) sandy limestone; (3) platy marl; (4) sandstone; (5) sand (on the lower left) and loose sandstone (on the lower right); (6) clay; (7) plant remains (disperse cuticles on the left; plant macrofossils, leaf flora, in the center; and root remains on the right); (8) marine invertebrates (hinged brachiopods on the left and hingeless brachiopods on the right).

GENERAL CHARACTERISTICS OF THE FOSSIL FLORA OF THE IVAGORA LOCALITY Taking into account distinctive features of the flora from the IvaGora locality, it seems plausible to estab lish on this basis the new IvaGora Floral Assemblage, which is dated Early Kazanian. Regarding cryptogams, the IvaGora Floral Assemblage is particularly rich in leaf verticils and foli ated shoots of Annularia aff. carinata Gutbier (Pl. 3, figs. 2–8). Plant remains of the fern Pecopteris ex gr. leptophylla Bunbury (Pl. 4, figs. 1–10, Fig. 2) are rather common (for more detail, see below).

The material includes phylloids and sporophylls of heterosporous lycopsids, the preservation of which prevents reliable identification. They are probably close to the lycopsid Signacularia Zalessky, which is characteristic of Kazanian deposits (Zalessky, 1937). Ferns of the morphological group of pecopterids, which are common in the IvaGora locality (Fig. 2), deserve particular attention. They are usually repre sented here by small fragments of the last order pinnae or, rarely, fragments of the penultimate order pinnae. Based on the material from the IvaGora locality, Zalessky initially established three new pecopterid species: Pecopteris attenta Zalessky, P. spiculosa

PALEONTOLOGICAL JOURNAL

Vol. 49

No. 11

2015

FOSSIL FLORA FROM THE KAZANIAN (MIDDLE PERMIAN) IVAGORA LOCALITY

1195

(c)

(a)

(b)

(f)

(d)

(e)

(g)

(h)

(l)

(i) (m) (j)

(k)

Fig. 2. Fern leaves of the pecopterid morphological group: Pecopteris ex gr. leptophylla Bunbury (a–h, j–m); Pecopteris sp. (i): (a) last order pinna described by Zalessky (1937, textfig. 62) as Pecopteris attenta Zalessky; (b) last order pinna described by Zalessky (1937, textfig. 63) as Pecopteris spiculosa Zalessky; (c) last order pinna described by Zalessky (1937, textfig. 64) as Pecopteris conserrata Zalessky; (d–m) specimens: (d) GIN, no. 4547/28; (e) GIN, no. 4851/188; (f) GIN, no. 4851/187; (g) GIN, no. 4547/29; (h) GIN, no. 4547/50 (epidermal cuticular structure was examined, see here Pl. 4, figs. 6, 7, 9, 10); (i) GIN, no. 4851/NN01; (j) GIN, no. 4851/300; (k) GIN, no. 4851/190; (l) GIN, no. 4851/186; (m) GIN, no. 4547/NN02. IvaGora locality; Middle Permian, Kazanian Stage, IvaGora Beds. Scale bars: (ac, e, f, l, m) 1 cm; (l, k) 5 mm; (d, g, h) 2 mm; (j) 1 mm. PALEONTOLOGICAL JOURNAL

Vol. 49

No. 11

2015

1196

NAUGOLNYKH

Zalessky, and P. conserrata Zalessky (Zalessky, 1937; see here Figs. 2a–2c). When studying a more repre sentative sample of pecopterid leaves from the Iva Gora locality, it was shown that the three species established by Zalessky fit into morphological varia tion range of leaves of one natural or botanical species and they are very close and sometimes indistinguish able from leaves of the gleichenian fern Oligocarpia leptophylla (Bunbury) GrauvogelStamm et Doubin ger, which is widespread in the Lower Permian of Euramerica (Wagner and Lemos de Sousa, 1985; Bar thel and Rössler, 1995). However, since fertile pecop terid leaves have not yet been found in the IvaGora locality and the taxonomic identity of pecopterid leaves from IvaGora and the Euramerian species Oli gocarpia leptophylla remains questionable, they are determined in open nomenclature as Pecopteris ex gr. leptophylla Bunbury. In addition to Pecopteris ex gr. leptophylla, the Iva Gora locality has yielded rare pecopterid leaves with unicoherently fused pinnules and rounded apices, although the preservation of these remains (Fig. 2i) prevents exact identification. Externally, these leaves resemble the Euramerian species P. unita Brongniart belonging to the order Marattiales. It should be noted that the IvaGora Assemblage includes one fragment of the middle part of a bipin nate leaf, tentatively determined as Lobatopteris sp. (Ignatiev and Naugolnykh, 2001, pp. 64, 67, pl. II, fig. 4, textfig. 2d; Naugolnykh, 2013, textfig. 30h). The rachis of a penultimate order pinna of Lobatopteris sp. from the IvaGora Flora is thick, mas sive, welldeveloped, 5 mm wide, with a thin longitu dinal central groove on the adaxial side of the rachis. Rachises of the last order pinnae diverge from the rachis of the penultimate order pinna at an angle of 50°. The last order pinnae are long, with subparallel margins. Judging from available fragment and the rate of pinnule shortening towards the apex of the last order pinna, the complete length of the last order pinnae of Lobatopteris sp. was probably 12–13 cm. The maxi mum observed width of the last order pinnae is 17 mm. The pinnules attached to the rachis of the last order pinna are fused by their edges for threefourths of the extent. The pinnules are relatively short, isometric in outline, with even edges and rounded apex. The mid veins of pinnules diverge from the rachis of the last order pinna at a sharp angle unusual for pecopterids, ranging from 25° to 35°; however, almost at once after

deviation from the rachis of the last order pinna, the midvein curves abruptly proximally and passes along the longitudinal axis of the pinnule at an angle of 45°–50° relative to the rachis of the last order pinna, reaching the edge of pinnule apex. A basiscopic lateral vein is the first to diverge from the midvein base of the pin nule, then, it bifurcates once or twice or, less fre quently, remains simple. Then, the midvein gives rise to an acroscopic lateral vein, which is usually simple or bifurcating one time. Farther, two or three more pairs of lateral veins diverge in alternate order from the mid vein and usually dichotomize once or, less frequently, remain simple. The edges of pinnules are complicated with fine rugosity, which coincides in direction with the veins. Similar remains of a fern leaf from the Inta Forma tion, which were also initially determined as Lobatopt eris sp., was figured in a study devoted to the paleobo tanic characteristics of Permian floras of the Pechora Basin (Pukhonto and Fefilova, 1983, pl. IX, figs. 1, 1a). Subsequently, this specimen was cited as a new spe cies, Lobatopteris vorcutensis Pukhonto, in manuscr. (Pukhonto and Naugolnykh, 2009, pl. V, figs. 1, 4). Since detailed venation of the Pechora form remain uncertain, it is presently difficult to compare Lobatop teris sp. from the Soyana Flora with the externally sim ilar fern from the Pechora Basin. The IvaGora Floral Assemblage includes rather frequent foliated shoots of sphenophytes determined as Annularia aff. carinata Gutbier (Pl. 3, figs. 2–8). Similar plant remains also occur in other fossil floras of the Kazanian Age of European Russia (Esaulova, 1986, pl. VII, figs. 1–3, pl. VIII, figs. 1, 2; Naugolnykh et al., 2014). The IvaGora locality has also yielded a verticil fragment probably belonging to a sphenophyte reproductive organ (Pl. 3, fig. 1). Among gymnosperms from IvaGora, it is impor tant to mention pinnate peltasperm leaves (of the cal lipterid morphological group, family Peltasper maceae), which were described by Zalessky (1937, p. 98, textfigs. 74, 75) as Odontopteris sojanaeana Zalessky and, according to the modern nomenclature, should be assigned to the species Permocallipteris wan genheimii (Fischer) Naugolnykh. In addition to callipterids, the IvaGora locality has yielded leaves of angaropelts of the genus Prae phylladoderma Naugolnykh (family Angaropeltaceae, order Peltaspermales), which are described in detail in the present paper.

Explanation of Plate 3 Fig. 1. Sphenophyta incertae sedis, specimen GIN, no. 4851/302. Figs. 2–8. Annularia aff. carinata Gutbier, specimens: (2) GIN, no. 4851/223; (3) GIN, no. 4851/224; (4, 6) GIN, no. 4851/220; (5, 7) GIN, no. 4851/221; (8) GIN, no. 4851/222. IvaGora locality; Middle Permian, Kazanian Stage, IvaGora Beds. Scale bars: (1) 0.5 cm and (2–8) 1 cm. PALEONTOLOGICAL JOURNAL

Vol. 49

No. 11

2015


1197 Plate 3

1

2

3

4

5

6

7


8

Vol. 49

No. 11

2015

1198

NAUGOLNYKH Plate 4

1

2

3

6

5 4

7

8

9

10

Explanation of Plate 4 Figs. 1–10. Pecopteris ex gr. leptophylla Bunbury, specimens: (1) GIN, no. 4851/186; (2) GIN, no. 4851/187; (3, 6, 7, 9, 10) GIN, no. 4547/50; (4) GIN, no. 4547/29a; (5) GIN, no. 4547/28; (8) GIN, no. 4851/189. IvaGora locality; Middle Permian, Kaza nian Stage, IvaGora Beds. Scale bars: (1, 2, 8) 1 cm; (4, 5) 2 mm; (7) 200 µm; (6, 10) 100 µm; (3) 50 µm. PALEONTOLOGICAL JOURNAL

Vol. 49

No. 11

2015


Foliated shoots of conifers (see, for example, Ignatiev and Naugolnykh, 2001, pl. 1, fig. 1 on the lower left; see here Pl. 4, fig. 6, lower left) occur in the IvaGora locality; exact taxonomic position of these forms remains uncertain. The IvaGora Floral Assemblage includes leaves of Rufloria (Alatorufloria) S. Meyen, Entsovia S. Meyen (Pl. 5, figs. 4, 7, 9), and also bracts of Nephropsis Zalessky and scaly leaves–cataphylls of Lepeophyllum Zalessky, which I regard as gymnosperms of the class Vojnovskyopsida (Naugolnykh, 2010). Some seeds with a welldeveloped wing described below also belong to this group. SYSTEMATIC PALEOBOTANY DIVISION PINOPHYTA REVEAL, 1996 CLASS PE LTASPERMOPSIDA CRONQUIST, 1981 Order Peltaspermales Taylor, 1981 Family Angaropeltaceae (Doweld, 2001, orf. incorr.) Naugolnykh, 2012 Genus Praephylladoderma Naugolnykh, 2007 Praephylladoderma sojanaeana (Zalessky, 1937) Naugolnykh, comb. nov., emend. nov. Plate 5, figs. 1–3, 5, 6, 8

Meristophyllum sojanaeanum Zalessky: Zalessky, 1937, p. 99, textfig. 76. Phylladoderma sp.: Ignatiev and Naugolnykh, 2001, pl. II, fig. 1. Basionym: Meristophyllum sojanaeanum Zalessky: Zalessky, 1937, p. 99, textfig. 76.

H o l o t y p e. Figured by Zalessky (1937, text fig. 76); IvaGora locality; Middle Permian, Kazanian Stage. D i a g n o s i s. Narrow lanceolate leaves with attenuate base and round apex. Two veins coming into leaf base. Veins relatively thick, dichotomizing one to two times, extending towards leaf apex, and never coming into leaf lateral margins. Apex of welldevel oped leaves occasionally having small middle sinus of dissection. D e s c r i p t i o n. The leaves are narrow lanceolate in outline, usually at most 4–5 cm long at the maxi mum width of 6 mm. The leaf base is usually attenuate wedgeshaped and the apex is rounded or blunted. The maximum leaf width is in its upper onethird. Some leaves are shortened, about 20 mm long; however, their greatest width is 6 mm, as in long welldeveloped leaves. Two veins enter the leaf base, which almost at once repeatedly dichotomize. Each vein usually bifur cates at most twice; however, some veins dichotomize up to three times. Veins are relatively thick, up to 150– 200 µm wide, very slowly becoming thinner towards the leaf apex. Veins are frequently constant in thick ness throughout the leaf extent. Some shoots have sev eral leaves preserved in attachment to the brachyblast (Ignatiev and Naugolnykh, 2001, pl. 2, fig. 2). PALEONTOLOGICAL JOURNAL

Vol. 49

No. 11

1199

C o m p a r i s o n. P. sojanaeana is most similar to the species Praephylladoderma leptoderma Naugol nykh occurring in the Kungurian Stage of the Fore Urals, which is probably ancestral to P. sojanaeana, and differs from it in the much smaller size, the thicker veins, and the presence of an apical notch, which is nonetheless not always present in P. sojanaeana. R e m a r k s. The leaves referred in the present study to the genus Praephylladoderma Naugolnykh were previously described by Zalessky (1937) and assigned to the genus Meristophyllum Zalessky. As Zalessky established the genus Meristophyllum, he assigned to it two species, Meristophyllum indivisum from the Krutaya Katushka1 locality (middle Fore Urals, left bank of the Barda River, upstream from the village of Matveevo) and M. sojanaeanum from the IvaGora locality; however, he did not designate the type species of this genus. Subsequently, it became evident that the two species not only do not belong to one genus, but also represent different gymnosperm groups. The mistake resulted from the superficial sim ilarity between double dorsal grooves of Meristophyl lum indivisum and the veins of M. sojanaeanum posi tioned close to each other after bifurcation. The author designated M. indivisum as the type species of the genus Meristophyllum. This species is rather similar to species of the genus Entsovia S. Meyen, which is closely related to the genus Meris tophyllum. Peculiar narrowleafed phylladoderms are rela tively frequent in the Ufimian and Kazanian stages (Esaulova, 1986, pl. XII, figs. 2–5; Fefilova, 1981, pl. IV, fig. 5). Some of these leaves were determined in open nomenclature as Meristophyllum cf. sojanaeanum (Esaulova, 1986, pl. XIX, fig. 2) or even as Meristo phyllum sojanaeanum (Fefilova, 1981, pl. IV, fig. 5). These leaves possibly also belong to the genus Prae phylladoderma. O c c u r r e n c e. Kazanian Stage of the Russian Platform and ForeUrals. M a t e r i a l. Seven satisfactory and wellpreserved leaves. C L A S S VO J N O V S K Y O P S I D A NAUGOLNYKH, 2010 Order Vojnovskyales Neuburg ex Emberger, 1968 Family Vojnovskyaceae Neuburg, 1963 Genus Megasylvella Naugolnykh, gen. nov.

E t y m o l o g y. From the Latin mega (large) and the generic name Sylvella. Ty p e s p e c i e s. Megasylvella ivagorica Naugol nykh, sp. nov. D i a g n o s i s. Platyspermic seeds ranging from ovoid to subtriangular, with subtriangular apical wing with acute apex. Wing slightly asymmetric. Widest part of wing in middle portion of seed. Micropilar tube conical, basically wide, gradually narrowing to its dis

2015

1200

NAUGOLNYKH Plate 5

1

2

3

4

5

6

7

8


9

Vol. 49

No. 11

2015


tal part. Nucellus obovate, with acute base; largest part of nucellus in its upper onethird portion of nucellus. C o m p a r i s o n. The new genus differs from the closest genus Sylvella Zalessky in the much larger seeds (up to 6 cm long in Megasylvella and even more in contrast to 2.5 cm long in Sylvella). The second important difference is the general shape and extent of development of the wing in Megasylvella which sur rounds the nucellus and, thus, is developed to an even greater extent than the apically excessively developed wing of seeds of the genus Sylvella, although this con cerns relative rather than absolute length. The third essential difference between these genera is the shape of the micropilar tube, which is very well developed in Megasylvella in the shape of a long conical apical out growth. R e m a r k s. Large anisopterous seeds found in the Novyi Kuvak locality (Naugolnykh et al., 2013, pl. IV, figs. 1–4, textfigs. 2, 3) probably belong to the genus Megasylvella. They differ somewhat from the type spe cies and possibly represent a separate species of the genus Megasylvella. The collection from the Novyi Kuvak locality includes remains of a reproductive organ, which is a clustering association of seeds of Megasylvella pre served in natural attachment to the common axis (Naugolnykh et al., 2013, pl. IV, fig. 2; Figs. 2a, 2b). This organ is very similar to seedbearing organs of Suchoviella Ignatiev et S. Meyen (Ignatiev and Meyen, 1989) belonging to the order Vojnovskyales sensu Naugolnykh (2010). Thus, it is highly probable that seeds of the genus Megasylvella belong to the gymnosperm order Vojnovskyales. The close genus Sylvella has previously been referred to the family Vojnovskyaceae (Meyen, 1982, 1984, 1987, 1988). O c c u r r e n c e. Middle Permian, Kazanian Stage, European Russia. S p e c i e s c o m p o s i t i o n. Type species. Megasylvella ivagorica Naugolnykh, sp. nov. Plate 6, fig. 1

E t y m o l o g y. From the IvaGora locality. H o l o t y p e. GIN, no. 4851/229 (PB 121/1); Iva Gora locality, Middle Permian, Kazanian Stage, Iva Gora Beds (Pl. 6, fig. 1; Fig. 3). D i a g n o s i s. The same as in the genus. D e s c r i p t i o n (Fig. 3). The seeds are flattened, platyspermic, from rounded obovate to subtriangular,

1201

with a welldeveloped apical wing. The seed (including wing) is 6 cm long or slightly greater and 3 cm wide. Juvenile poorly developed seeds can be smaller. The wing is usually slightly asymmetrical due to a greater development of one side and weak curvature of the ter minal wing part, which is not always manifested. The wing almost completely surrounding the nucellus and micropilar tube is thin filmy, covered with very thin shading (microfolds) directed perpendicular or slightly obliquely relative to the seed edge. The wing has rare large, but low folds, which coincide in direc tion with fine shading. The wing margin is mostly even, but the middle and lower seed parts sometimes have wavy margins, occasionally complicated with weak lobes. The wing apex is somewhat pointed. The widest wing part is located in the middle part of the seed. The micropyle is conical, with expanded base, nar rowing gradually distally. The micropilar tube is 35 mm long and 7 mm wide. The nucellus is obovate, with a pointed base and the larger part located in the upper onethird of the nucellus. The nucellus is 29 mm long, with the maximum width of 13 mm. O c c u r r e n c e. Middle Permian, Kazanian Stage, European Russia. M a t e r i a l. Six wellpreserved seeds. CLASS INCETAE SEDIS Genus Radicites Potonie, 1893 Radicites trimorphus Naugolnykh, sp. nov. Plate 6, figs. 2 and 5

E t y m o l o g y. From the Latin trimorphus (trimor phous), based on peculiar morphology of root branches of the first, second, and third orders. H o l o t y p e. GIN, no. 4851/230; IvaGora local ity; Middle Permian, Kazanian Stage, IvaGora Beds (Pl. 6, fig. 2). D i a g n o s i s. Isolated roots with three orders of branching. Axes of first order thick, straight, having longitudinal ribs. Axes of second order thinner, pos sessing small round or ovoid bodies, probably with supplementary tissues. Axes of third order very small and thin, having absorbing function. D e s c r i p t i o n. The roots are up to 20–30 cm long, as the first order axes are up to 3 mm wide, branching up to three times. The first order axes are straight or slightly curved. Their surface usually has

Explanation of Plate 5 Gymnosperm leaves of the IvaGora Floral Assemblage. IvaGora locality; Middle Permian, Kazanian Stage, IvaGora Beds. Figs. 1–3, 5, 6, 8. Praephylladoderma sojanaeana (Zalessky, 1937) Naugolnykh, comb. nov., emend. nov.; (1) specimen GIN, no. 4851/232; (2, 6) specimen GIN, no. 4851/233; (3) holotype, after Zalessky (1937, textfig. 76); (5) specimen GIN, no. 4851/234; (8) specimen GIN, no. 4851/235. Figs. 4, 7, 9. Entsovia sp., specimen GIN, no. 4851/226. Scale bars: (1–5, 8, 9) 1 cm and (6, 7) 1 mm. PALEONTOLOGICAL JOURNAL

Vol. 49

No. 11

2015

1202

NAUGOLNYKH

Fig. 3. Megasylvella ivagorica Naugolnykh, gen. et sp. nov., holotype GIN, no. 4851/229 (PB 121/1). IvaGora locality; Middle Permian, Kazanian Stage, IvaGora Beds. Scale bar, 1 cm. PALEONTOLOGICAL JOURNAL

Vol. 49

No. 11

2015


1203 Plate 6

2

1

3

4

5

Explanation of Plate 6 Fossil flora from the IvaGora locality; Middle Permian, Kazanian Stage, IvaGora Beds. Scale bars: (1–3, 5) 1 cm and (4) 1 mm. Fig. 1. Megasylvella ivagorica Naugolnykh, gen. et sp. nov., holotype GIN, no. 4851/229 (PB 121/1). Figs. 2 and 5. Radicites trimorphus Naugolnykh, sp. nov.: (2) holotype GIN, no. 4851/230, (5) specimen GIN, no. 4851/nn03. Fig. 3. Samaropsis sp. 1, specimen GIN, no. 4851/231. Fig. 4. Samaropsis sp. 2, specimen GIN, no. 4851/228. PALEONTOLOGICAL JOURNAL

Vol. 49

No. 11

2015

1204

NAUGOLNYKH

distinct longitudinal ribs. The axes of the second order diverge from the first order axes at a right angle; they are usually at most 15–20 mm long at the maximum width of 1.8 mm. The axes of the second order may have small spherical or ellipsoidal structures up to 2– 3 mm of maximum dimension. These structures are presumably root nodules with storage tissue. The sec ond order axes usually apically dichotomize at a sharp angle (about 50°–60°), with the formation of the third order axes. The third order axes are very thin, curved. They are usually at most 0.8 mm wide. The maximum visible length of the third order axes is 19 mm. They most likely performed the absorbing function. C o m p a r i s o n. The new species differs from the closest species Radicites sentjakensis Esaulova (1986) in the presence of terminal expansions formed of stor age tissue on the second order roots and also in the presence of longitudinal ribbing of the first order axes. Externally similar fossil roots have been described in open nomenclature as Radicites sp. 3 (Arefiev and Naugolnykh, 1998) from the Salarevo Formation (Vyatkian Stage, Upper Permian) in the Northern Dvina River Basin. R e m a r k s. The presence of numerous roots in the basal part of the platy marl member (Bed 8; see Fig. 1) is evidence that, as the initial deposits were formed, the substrate was exposed to intense influence of the biota in aerial or subaerial conditions, in partic ular, it was colonized by higher plants producing the roots Radicites trimorphus. Thus, Bed 8 of the Iva Gora section is in essence a FPS profile (about the ter minology, see Naugolnykh, 2014; etc.) or poorly developed paleosol. O c c u r r e n c e. Middle Permian, Kazanian Stage; European Russia. M a t e r i a l. Five satisfactory and wellpreserved specimens. ACKNOWLEDGMENTS I am sincerely grateful to V.N. Kuleshov (Geologi cal Institute, Russian Academy of Sciences, Moscow) and A.V. Pakhnevich (Borissiak Paleontological Insti tute, Russian Academy of Sciences, Moscow) for an opportunity of studying the plant remains described in the present paper. This study was supported by the Russian Govern ment Program for Competitive Growth of Kazan Fed eral University among World’s Leading Scientific and Education Centers. REFERENCES Arefiev, M.P. and Naugolnykh, S.V., Isolated roots from the Tatarian Stage of the basin of the Sukhona and Malaya North ern Dvina rivers, Paleontol. Zh., 1998, no. 1, pp. 86–99.

Barthel, M. and Rössler, R., RotliegendFarne in weißen VulkanAschen“Tonsteine” der DohlenFormation als paläobotanische Fundschichten, Veroff. Mus. Naturk. Chemnitz., 1995, vol. 18, pp. 5–24. Esaulova, N.K., Flora kazanskogo yarusa Prikam’ya (Flora from the Kazanian Stage of the Kama Region), Kazan: Kazan. Gos. Univ., 1986. Fefilova, L.A., Macroflora, in Korrelyatsiya raznofat sial’nykh razrezov verkhnei permi severa Evropeiskoi chasti SSSR (Correlation of Heterofacies Sections of the Upper Permian of the North European Part of the USSR), Lenin grad: Nauka, 1981, pp. 16–18. Ignatiev, I.A. and Meyen, S.V., Suchoviella—gen. nov. from the Permian of Angaraland and a review of the systematics of Cordaitanthales, Rev. Palaeobot. Palynol., 1989, vol. 57, pp. 313–339. Ignatiev, I.A. and Naugolnykh, S.V., Early Kazanian flora of the Soyana River and its position among floras and plant zones of Angaraland of the same age, Stratigr. Geol. Korre lyatsiya, 2001, vol. 9, no. 3, pp. 1–94. Likharev, B.K. and MiklukhoMaklai, K.V., IvaGora Beds (“facies”), in Stratigraficheskii slovar’ SSSR. Karbon, perm’ (Stratigraphic Dictionary of the USSR: Carboniferous, Permian), Leningrad: Nedra, 1977, p. 152. Meyen, S.V., The Carboniferous and Permian floras of Angaraland: A synthesis, Biol. Mem., 1982, vol. 7, pp. 1– 109. Meyen, S.V., Basic features of gymnosperm systematics and phylogeny as shown by the fossil record, Bot. Rev., 1984, vol. 50, no. 1, pp. 1–111. Meyen, S.V., Fundamentals of Palaeobotany, London: Chapman and Hall, 1987. Meyen, S.V., Gymnosperms of the Angara flora, in Origin and Evolution of Gymnosperms, Beck, Ch.B., Ed., New York: Columbia Univ. Press, 1988, pp. 338–381. Naugolnykh, S.V., Gymnosperms of the class Vojnovsky opsida: New insight into old problems, in Paleontologiya i stratigrafiya permskoi sistemy v muzeinykh ekspozitsiyakh i chastnykh kollektsiyakh. Sb. nauch. rabot (Paleontology and Stratigraphy of the Permian System in Museum Expositions and Private Collections: Collected Works), Kungur–Perm: Poligraf Siti, 2010, pp. 10–18. Naugolnykh, S.V., Permian ferns of western Angaraland, Paleontol. J., 2013, vol. 47, no. 12, pp. 1379–1462. Naugolnykh, S.V., Devonian paleosoils of the Andoma Mountain, Sci. Russ., 2014, no. 4, pp. 12–18. Naugolnykh, S.V. and Kuleshov, V.N., Iskopaemaya flora reki Soyany: okno v permskii period (Fossil Flora from the Soyana River: Breakthrough into the Permian Period), Arkhangelsk: Izd. Tsentr SGMU, 2005. Naugolnykh, S.V., Morov, V.P., Varenov, D.V., and Varen ova, T.V., Flora of the Kazanian Stage of the Isakly locality (Samara Region) as a reflection of hydrophilous vegetative communities of the middle of the Permian Period, in Pale ontologiya v muzeinoi praktike (Paleontology in Museum Practice), Moscow: MediaGrand, 2014, pp. 66–78. Naugolnykh, S.V., Sidorov, A.A., Varenov, D.V., and Varen ova, T.V., Permian fossil plants from the Novyi Kuvak and Buzbash localities (Samara Region): Taxonomic diversity, in Ob’’ekty paleontologicheskogo i geologicheskogo naslediya i rol’ muzeev v ikh izuchenii i okhrane (Objects of Paleonto logical and Geological Heritage and the Role of Museums


Vol. 49

No. 11

2015

FOSSIL FLORA FROM THE KAZANIAN (MIDDLE PERMIAN) IVAGORA LOCALITY in Their Study and Protection), Kungur: Kungur. Istori. Arkhitekt. Khudozh. Muz.Zap., 2013, pp. 46–61. Pakhnevich, A.V. and Savinov, I.A., Catalogue of fossil plants collection at the Biological–Chemical Faculty of Lenin Moscow Pedagogical State University, in Sbornik stu dencheskikh rabot po biologii (Collected Student’s Works in Biology), Moscow: Mosk. Pedag. Gos. Univ., 1998, pp. 5–16. Pukhonto, S.K. and Fefilova, L.A., Makroflora, in Paleon tologicheskii atlas permskikh otlozhenii Pechorskogo ugol’nogo basseina (Paleontological Atlas of Permian Deposits of the Pechora Coal Basin), Leningrad: Nauka, 1983. Pukhonto, S.K. and Naugolnykh, S.V., Evolution of higher plants of the ForeUrals in the Permian Period, in Nauka i prosveshchenie. K 250letiyu Geologicheskogo muzeya RAN (of higher plants of the ForeUrals in the Permian Period), Moscow: Nauka, 2009, pp. 319–352.


Vol. 49

No. 11

1205

Shcherbakov, D.E., Za paleozoiskimi nasekomymi na Soy anu (Search for Paleozoic Insects on the Soyana River), Arkhangelsk: Izd. Tsentr SGMU, 2007. Wagner, R.H. and Lemos de Sousa, M.J., Oligocarpia lepto phylla (Bunbury), nomenclatorial history and description of the lectotype, Contrib. Carbon. Geol. Palaeontol. Iber. Penin. An. Cienc. Porto. Suppl., 1985, vol. 64, pp. 481–490. Zalessky, M.D., Sur la distinction de l’Etage Bardien dans le Permien de l’Oural et sur sa flore fossile, Probl. Paleontol., 1937, vols. 2–3, pp. 37–101. Zekkel’, Ya.D., Geological studies in the area of Zimnii Bereg of the White Sea, Kuloy Basin, and lower reaches of the Mezen River, Tr. Severn. Geol. Upravl., 1939, no. 3, pp. 1–94.

2015

Translated by G. Rautian