Permian ferns of western Angaraland

ISSN 0031�0301, Paleontological Journal, 2013, Vol. 47, No. 12, pp. 1379–1462. © Pleiades Publishing, Ltd., 2013.

1379

CONTENTSINTRODUCTION 1380MATERIAL AND METHODS 1380HISTORY OF THE STUDY 1383DESCRIPTIVE MORPHOLOGY AND TERMINOLOGICAL SYSTEM 1386PALEOBOTANICAL DESCRIPTION 1388RELATIONSHIPS BETWEEN THE EVOLUTION OF FERNS AND CLIMATICAND GEODYNAMIC EVENTS 1453 SIGNIFICANCE OF FERNS FOR STRATIGRAPHY OF THE PERMIAN STRATA 1457CONCLUSIONS 1458ACKNOWLEDGMENTS 1458REFERENCES 1458

Permian Ferns of Western AngaralandS. V. Naugolnykh

Geological Institute, Russian Academy of Sciences, Pyzhevskii per. 7, Moscow, 119017 Russiae�mail: [email protected]

Received February 6, 2012

Abstract—Permian ferns of Angaraland, particularly their taxonomy and systematics are considered. Thefollowing new genera and species of Permian ferns are described: Acrogenotheca ramificata Naugolnykh, gen.et sp. nov., Convexocarpus Naugolnykh, gen. nov., Todites emarginatus Naugolnykh, sp. nov., T. coronatusNaugolnykh, sp. nov., Tumidopteris clavata Naugolnykh, gen. et sp. nov., and Geperapteris rotunda Naugol�nykh, sp. nov. The history of the study of Permian ferns of Angaraland is discussed in detail. Descriptive ter�minology for compound pinnate leaves of ferns is proposed. Exact data on the geographical and stratigraph�ical position of the sections and localities of plant remains are provided. Detailed descriptions and (or)emended diagnoses of earlier described taxa, such as Convexocarpus distichus (Naugolnykh) Naugolnykh,comb. nov., Corsinopteris dicranophora (Naugolnykh) Doweld, C. semilibera (Naugolnykh) Doweld,C. petschorica (Fefilova) Doweld, Todites lobulatus Naugolnykh, Dvinopteridium edemskii Zalessky, emend.nov., Prynadaeopteris karpovii (Radczenko) Radczenko, emend. nov., Myriotheca cf. scaberrima Lesquereux,Pecopteris anthriscifolia (Goeppert) Zalessky, emend. nov., Pecopteris ex gr. leptophylla Bunbury in Ribeiro,and Lobatopteris sp. are provided. The stratigraphical range and significance of Permian ferns for generalpaleophytogeographical reconstruction are considered.

Keywords: Pteridophyta, Polypodiopsida, ferns, systematics, new taxa, Permian, Angaraland

DOI: 10.1134/S0031030113120010

1380

PALEONTOLOGICAL JOURNAL Vol. 47 No. 12 2013

NAUGOLNYKH

INTRODUCTION

In the modern flora, ferns are rather diverse inregard to life (forms and ecological preferences andthey are comparable in taxonomic diversity only toangiosperms.

In the geological past, ferns were also rather impor�tant and, in certain periods and certain communities,for example, in the Late Paleozoic and Middle Meso�zoic, they played a leading role.

Fern remains in the Permian beds are usually rare,but diverse. By the early 1980s, from the Permian ofthe Fore�Urals (which belongs paleophytogeographi�cally to western Angaraland, along with adjacentregions of the Russian Platform), a total of about50 species that are more or less reliably assigned toferns had been described. A large part of these speciesare represented by sterile leaves; however, some spe�cies were established based on specimens with pre�served fertile organs, which provided the determina�tion of their taxonomic position. These species prima�rily include Asterotheca (?) kojimensis (Zalessky)Fefilova, Corsinopteris petschorica (Fefilova) Doweld,Oligocarpia permiana Fefilova, and Todites sp.(Fefilova, 1973). Subsequently, Naugolnykh (1995,1998a, 1998b, 1998c, 2002) described other fertileferns from the Permian of the Fore�Urals, includingthe marattialean ferns Ptychocarpus distichus Naugol�nykh, Corsinopteris dicranophora (Naugolnykh) Dow�eld, C. semilibera (Naugolnykh) Doweld, and theosmundacean fern Todites lobulatus Naugolnykh.

Further studies of Permian floras from the Fore�Urals and adjacent regions, paleophytogeographicallybelonging to western Angaraland sometimes regardedas the independent Subangara Ecotonal Belt (Meyen,1987), have shown that they include new fern speciesof both eusporangiate and leptosporangiate membersof this group. The present study is devoted to the

description of these new ferns and new interpretationof previously established taxa.

In addition to fern remains from the Permian of theFore�Urals, the data on ferns from the Kazanian of theIva�Gora locality (Arkhangelsk Region), Severodvin�ian of the Nikulino locality (Vologda Region), andPermian of the Pechora and Kuznetsk coal basins areconsidered; because on the one hand, they are taxo�nomically close to Permian forms from the Fore�Uralsand, on the other hand, they complete the generalsequence of fern assemblages.

The study provides diagnoses and some additionaldata (comparison, remarks, etc.) only on new generaand genera with essentially changed diagnosis.

MATERIAL AND METHODS

The present study is based on plant remains whichI collected during the last years in a number of Per�mian localities (Figs. 1–3) situated within the middleand southern Fore�Urals (Perm and Sverdlovskregions, Bashkiria), Fore�Urals near the PechoraRiver (Kozhim River, Komi Republic), and theArkhangelsk and Vologda regions. Additional materialincludes certain specimens collected by A.G. Sharov(Moscow, Paleontological Institute, Academy of Sci�ences of the USSR; presently the Borissiak Paleonto�logical Institute of the Russian Academy of Sciences:PIN) and several specimens transferred to me byV.N. Kuleshov (Moscow, Geological Institute, Rus�sian Academy of Sciences: GIN), A.A. Shkolin (Mos�cow, PIN), V.A. Tsimbal (Moscow, independentresearcher), P.A. Sofronitskii, V.G. Novokshonov(Perm, Permian State University), and M.P. Arefiev(Moscow, GIN). In 2006, S.K. Pukhonto (Moscow,Vernadsky State Geological Museum of the RussianAcademy of Sciences) transferred to me for studying arather representative collection of plant remains fromthe Permian of the Pechora Coal Basin, includingferns described in the present study. I am sincerelygrateful to all colleagues who provided me with speci�mens for studying.

I in general adhere to the point of view accepted bymany stratigraphers concerning the necessity of pres�ervation of the traditional division of the Permian Sys�tem into two (Upper and Lower) series, with theUfimian Stage in the Upper series (Lozovsky et al.,2009). The Ufimian Stage is treated in its former vol�ume (with the Solikamskian and Sheshminskian hori�zons), in agreement with available biostratigraphicand geohistorical data. In this improved stratigraphicalscheme, the terms Cisuralian, Biarmian, and Tatarianare retained to designate subseries of the Permian Sys�tem. The lower series consists of the only subseries(Cisuralian), and the upper series consists of two sub�series (lower and upper, Biarmian and Tatarian). TheUrzhumian and Severodvinian regional stages are pro�posed to be retained in the status presently existing de

12

400 0 400 1200 km

3

Fig. 1. Geographical position of locality groups cited in thepaper: (1) Fore�Urals near Perm, (2) Fore�Urals near thePechora River and the Pechora Coal Basin, and (3) Kuz�netsk Basin.


PERMIAN FERNS OF WESTERN ANGARALAND 1381

facto, that is, corresponding to the correlation poten�tial and volume of the Urzhumian and Severodvinianhorizons. The status and the nomenclature of theVyatkian Stage require specification (for more detail,see Lozovsky et al., 2009). The necessity of the recog�nition of the uppermost stage of the Permian System(Vladimirian) was previously substantiated by me(Naugolnykh, 2005b). It should be noted that theVladimirian Stage is used in works of both Russian andforeign paleontologists and stratigraphers (Ozhgibesovet al., 2009; Pukhonto and Naugolnykh, 2009; Yanet al., 2010, Yang et al., 2011; Singh et al., 2012).

At the same time, despite certain eclecticism of thepresently existing stratigraphic scheme of the Per�mian System, as it was adopted by the Permian Sub�committee of the Interdepartmental StratigraphicalCommittee of Russia, I use here a somewhat modi�fied tripartite division of the Permian System into the

lower (Cisuralian) series (comprising the Asselian,Sakmarian, Artinskian, and Kungurian stages); mid�dle (Biarmian) series (Ufimian, Kazanian, andUrzhumian stages), and upper (Tatarian) series(Severodvinian, Vyatkian, and Vladimirian stages).

The stratigraphic relations and nomenclature ofthe groups and formations of the Permian of the Kuz�netsk Basin follow Betekhtina et al. (1988).

The microscopic study of the sporangia, conduct�ing elements, topography of the epidermis and sporespreserved in the sporangia in situ was performed usinga JEOL JSM�6060A scanning electron microscope(Faculty of Soil Sciences, Moscow State University),Vega Tescan MV 2300 scanning electron microscope,and Carl Zeiss Axiostar digital optical microscope inthe Geological Institute of the Russian Academy ofSciences, Moscow (GIN).

0 70 140

km

1

2

3

4

5

KirovPerm

Kazan

Vorkuta

Ufa

Fig. 2. Geographical position of locality groups understudy: (1) Chekarda 1; (2) Rakhmangulovo, Kriulino;(3) Fore�Urals near the Pechora River and Pechora CoalBasin (Kozhim 5, IK�886, IK�678, IK�657, IK�385,IK�687, IK�631, VK�22, VK�33, N�228, K�2400,KhK�1056, KhK�1134, 11374); (4) Soyana (Iva�Gora);and (5) Nikulino.

30 0 30 60 km

1

2

3

4

56

89 10

1112

Pechora

Kozhim

Inta

Vorkuta

Khalmer�Yu

7

Fig. 3. Geographical position of localities under study:Fore�Urals near the Pechora River and Pechora CoalBasin: (1) IK�886; (2) Kozhim 5; (3) IK�678; (4) IK�657;(5) IK�385; (6) IK�687; (7) IK�631; (8) VK�22; (9) VK�33;(10) N�228; K�2400; (11) KhK�1056; (12) KhK�1134,1137.

1382


NAUGOLNYKH

The sporangia were macerated in concentratednitric acid without potassium chloride. After the treat�ment with ammonia and washing in distilled water, thecuticle and spores extracted from sporangia were pre�pared and then mounted on the object stubs for study�ing by a scanning electron microscope.

The data on localities mentioned in the presentstudy, their geographic and stratigraphic position, age,and the list of fern taxa included in the taxonomic partare given below.

Lower Permian

Rakhmangulovo 1. Left bank of the Ufa River; steepof the basal terrace near the village of Rakhmangulovo(Krasnoufimskii District, Sverdlovsk Region); LowerPermian, Kungurian Stage. The locality has yieldedPecopteris uralica Zalessky, Pecopteris spp., and trunkfragments of treelike ferns.

Rakhmangulovo 2. Left bank of the Ufa River,quarry in the upper part of the basal terrace, 1 km eastof the village of Rakhmangulovo (Krasnoufimskii Dis�trict, Sverdlovsk Region); Lower Permian, KungurianStage. The locality has yielded Pecopteris anthriscifolia(Goeppert) Zalessky, P. uralica, Pecopteris spp., andtrunk fragments of treelike ferns.

Kriulino. Left bank of the Ufa River; trenchesunder a gas pipeline in the eastern outskirts of the townof Krasnoufimsk (Sverdlovsk Region); Lower Per�mian, Kungurian Stage. The locality has yielded Con�vexocarpus distichus (Naugolnykh) Naugolnykhcomb. nov., Pecopteris uralica, Pecopteris spp., andtrunk fragments of treelike ferns.

Matveevo Group of localities. The localities situ�ated near the village of Matveevo (Matveevo, KrasnayaGlinka, Krutaya Katushka (sections 1 and 2) locali�ties; for detailed correlation, see Naugolnykh, 1998c).Steeps on both banks of the Barda River near the vil�lage of Matveevo, Lysvenskii District, Perm Region;Lower Permian, Kungurian Stage. These localitieshave yielded Pecopteris anthriscifolia, P. uralica,Pecopteris spp., Myriotheca cf. scaberrima Lesquereux,and Sphenopteris sp.

Chekarda 1, Bed 10. Left bank of the Sylva River,50 m downstream from the mouth of the ChekardaRiver, Suksunskii District, Perm Region; Lower Per�mian, Kungurian Stage. This locality has yielded Con�vexocarpus distichus, comb. nov., Corsinopteris dicran�ophora, Acrogenotheca ramificata Naugolnykh, gen. etsp. nov., Pecopteris uralica, and P. anthriscifolia as wellas remains of other ferns: P. synica Zalessky, P. helena�eana Zalessky, Pecopteris spp., Sphenopteris sp., Ovop�teris sp., and Myriotheca cf. scaberrima, which werecharacterized and figured in the previous studies(Naugolnykh, 1995, 1998a, 1998b, 1998c, 2007).

Vorkuta 1 (outcrop 37). Right bank of the VorkutaRiver near the village of Rudnik; Lower Permian,

Kungurian Stage, Lekvorkutskaya Formation. Pecopt�eris anthriscifolia, sterile leaves.

Middle and Upper Permian

Kozhim 5. Pechora Coal Basin. Left bank of theKozhim River, 5 km downstream from the railroadbridge near the village of Kozhim–Rudnik (Bed 96;for details of the stratigraphic column, see Naugol�nykh, 1998b, text�fig. 4, pp. 162–163), Komi Repub�lic; Middle Permian, Ufimian Stage; Inta Formation,Vorkuta Group. This locality has yielded the holotypesof Todites lobulatus, Corsinopteris semilibera, andPrynadaeopteris (?) sp.

Borehole VK�22 (exact positions of boreholesVK�22, VK�33, KhK�1056, KhK�1134, KhK�1137,IK�385, IK�426, IK�426 IK�631, IK�657, IK�678,IK�686, IK�687, IK�886, K�228, and K�2400 areshown in Fig. 2 and in work of Pukhonto, 1998), depthof 746.5 m. Fore�Urals near the Pechora River, Vor�gashorskoe Coal Field, Komi Republic; Middle Per�mian, Ufimian Stage; Inta Formation, VorkutaGroup. Pecopteris sp., sterile leaves.

Borehole VK�33, 753.2 m of depth, VorgashorskoeCoal Field, Komi Republic; Middle Permian, Ufim�ian Stage; Inta Formation, Vorkuta Group. Tumidopt�eris clavata, gen. et sp. nov., fertile and sterile leaves.This locality has also yielded sterile leaves ofPrynadaeopteris venusta Radczenko (Pukhonto andFefilova, 1983, pl. VI, fig. 2).

Borehole KhK�1056, 96 m of depth. Fore�Uralsnear the Pechora River, Khalmer’yusskoe Coal Field,Komi Republic; Middle Permian, Urzhumian Stage;Talbei Formation, Pechora Group. Pecopterisanthriscifolia, sterile leaves.

Borehole KhK�1058, 4336 m of depth. Fore�Uralsnear the Pechora River, Khalmer’yusskoe Coal Field,Komi Republic; Middle Permian, Kazanian Stage.Corsinopteris petschorica.

Borehole KhK�1137, 244.2 m of depth. Fore�Uralsnear the Pechora River, Khalmer’yusskoe Coal Field,Komi Republic; Middle Permian, Kazanian Stage;uppermost part of the Inta Formation, Vorkuta Group(see also borehole IK�385–IK�886). Tumidopteris clav�ata, gen. et sp. nov., the holotype of which is a fertilefrond fragment.

Borehole IK�385, 104 m of depth. Fore�Urals nearthe Pechora River, Intinskoe Coal Field. T. clavata,gen. et sp. nov., sterile leaves.

Borehole IK�426, 729.4 m of depth. Fore�Uralsnear the Pechora River, Intinskoe Coal Field. T. clavata,gen. et sp. nov., sterile leaves.

Borehole IK�426, 533.25 m of depth. Fore�Uralsnear the Pechora River, Intinskoe Coal Field. Corsi�nopteris petschorica.

Borehole IK�631, 344.5 m of depth. Fore�Uralsnear the Pechora River, Intinskoe Coal Field.C. petschorica.



Borehole IK�657, 864.8 m of depth. Fore�Uralsnear the Pechora River, Intinskoe Coal Field. Pecopt�eris anthriscifolia, sterile leaves; at 765.7 m of depth,fertile leaves of Corsinopteris petschorica are recorded.

Borehole IK�678, 960 m of depth. Fore�Urals nearthe Pechora River, Intinskoe Coal Field. Pecopterisanthriscifolia.

Borehole IK�686, 140 m of depth. Fore�Urals nearthe Pechora River, Intinskoe Coal Field. Tumidopterisclavata, gen. et sp. nov., sterile leaves.



Exact stratigraphic position of localities of Bore�holes IK�385, IK�426, IK�426, IK�631, IK�657, IK�678, IK�686, IK�687, IK�886 is uncertain. They aretentatively assigned here to the Kazanian (Middle Per�mian).

Borehole K�228, 254.2 m of depth. Fore�Uralsnear the Pechora River, Vorkutskoe Coal Field, KomiRepublic; Middle Permian, Ufimian Stage, Inta For�mation, Vorkuta Group. A trunk fragment of a tree�like fern.

Borehole K�2400, 300 m of depth. Vorkutskoe CoalField, Komi Republic; Middle Permian, UfimianStage; Inta Formation, Vorkuta Group. T. clavata,gen. et sp. nov., sterile leaves.

Iva�Gora. Arkhangelsk Region, right bank of theSoyana River, 50 km upstream from the village of Soy�ana; Middle Permian, Kazanian Stage. Pecopteris ex gr.leptophylla Bunbury, Lobatopteris sp. (the determina�tion of Lobatopteris was initially questionable: Ignatievand Naugolnykh, 2001).

Nikulino. Right bank of the Sukhona River, 2 kmdownstream from the village of Poldarsa, VologdaRegion; Upper Permian, Severodvinian Stage. Thislocality has yielded Dvinopteridium edemskii Zalessky,emend. nov.

Geographical and stratigraphic correlation of fossilPermian ferns which are represented by isolated spec�imens or involved for comparison are given at the firstciting.

HISTORY OF THE STUDY

The historical sketch provides data on the mostimportant studies devoted to the morphology and tax�onomy of Late Paleozoic (Upper Carboniferous andPermian) ferns. Particular attention is paid to worksdealing with Permian ferns of Angaraland.

One of the best works devoted to the morphologyand taxonomy of Paleozoic ferns published in the 19thcentury is the review of Stur (1883). This work placesprimary emphasis on fern fertile fronds and male

reproductive organs of pteridosperms (which were atthat time regarded as spore�producing organs belong�ing to ferns) coming from the Carboniferous beds ofWestern and Central Europe; however, in addition toCarboniferous forms, it also described fossil ferns fromthe European Autunian, i.e., from the Lower Permian.The study of Stur developed the basis for the modernunderstanding of several paleobotanically importantgenera, such as Discopteris Stur, Asterotheca Presl, Oli�gocarpia Goeppert, Senftenbergia Corda, and Dan�aeites Goeppert. Stur (1883, p. 785) published a rathersignificant table providing comparisons of a number ofextant fern genera (mostly eusporangiates) with theirCarboniferous predecessors and showing direct phylo�genetic relationships of these groups. The genusCalymmotheca Stur was considered to include speci�mens presumably belonging to both male reproductiveorgans of pteridosperms and fern sori or sporophores:Calymmotheca minor Stur, C. schatzlarensis Stur,C. avoldensis Stur, and C. frenzlii Stur. Stur consider�ably contributed to the understanding of morphologi�cal nature of the small circular structures, which weretreated both before and after Stur’s studies as sporan�gia or even seed scars; however, these structures are infact fruit bodies of fungi, which he described as Phylla�chora diplothmematis Stur.

The ideas of Stur concerning the morphology andgeneric taxonomy of Paleozoic ferns (Pl. 1, figs. 1–14)were to a great extent accepted by the followers as abasis for the development of taxonomy of this plantgroup.

Ferns from Permian localities of the Fore�Uralsinclude both Euramerian forms related to thosedescribed by Stur and other paleobotanists from Euro�pean localities, and typical Angaraland forms. Someimportant records of fern leaves and mineralizedtrunks from Permian deposits of the Fore�Urals weredescribed and figured in works of Kutorga (1842,1844) and Eichwald (1854), pioneers of Russian pale�ontology.

Fern leaves from the Permian of the Pechora Basinand Fore�Urals were figured and characterized byI.F. Schmalhausen. In his widely known work,Schmalhausen (1879) described and figured severalfern species, a large part of which (contrary to theopinion of Schmalhausen), were Permian rather thanJurassic. Many species of Permian ferns described bySchmalhausen were erroneously referred to the extantgenera Asplenium L. and Cyathea J.E. Smith. From theKuznetsk Basin, Schmalhausen described Aspleniumwhitbyense Brongniart, A. petruschinense Heer,A. argutulum Heer, Pecopteris recta Schmalhausen,Rhisomopteris sp., and Cyathea tchihatchevii Schmal�hausen; leaves of the last fern are very similar in mor�phology to the later described synonymous mor�phospecies Pecopteris hyperborea Zalessky (Zalessky,1934a) and P. leninskiensis (Chachlov) Radczenko(Radczenko, 1955).

1384


NAUGOLNYKH

Fossil remains of the vojnovskyalean gymnospermRhiptozamites goeppertii leaves, which were found inco�occurrence with the fern Cyathea tchihatchewii,support the Permian age of the last species.

In addition, Schmalhausen described Aspleniumwhitbyense, A. petruschinense, and Cyathea tchi�hatchevii from the Pechora Basin and Asplenium(Euasplenium) tunguscanum Schmalhausen, A. whit�byense, A. petruschinense, A. czekanowskii Schmal�hausen, Acrostichum sibiricum Schmalhausen, andPecopteris recta from the Tunguska Basin.

Plate VII, fig. 21 of the same study (Schmalhausen,1879) shows a leaf under the name Filicites sp., whichis actually a leaf of the gymnosperm (praeginkgo�phyte) Psygmophyllum sp., which comes from the Ora�nets River (Pechora River Basin). In spite of its name,this form is unrelated to ferns.

Among the fern species described by Schmal�hausen, Asplenium tunguscanum is of particular inter�est. The original description of this species is based onsix specimens of bipinnate leaf fragments (Schmal�hausen, 1879, pl. XIII, figs. 1, 2, 5) and pinnae of thelast order (Schmalhausen, 1879, pl. XIII, figs. 3, 4).The specimen with the entire margined pinnules hav�ing rounded apices, figured by Schmalhausen (1879,pl. XIII, fig. 7) and also assigned to “Asplenium” tun�guscanum, essentially differs from other specimens,which are characterized by pointed apices and pres�ence of well�developed lobes. Another specimen fig�ured by Schmalhausen (1879, pl. XIII, fig. 5) also dif�fers from other specimens of “A”. tunguscanum. Thelateral veins of pinnules of this fern dichotomize onceor, sometimes, twice, while the lateral veins of otherspecimens are mostly simple. However, the differencein venation could be easily explained by the differentextent of vascularization of young or apical parts andbetter developed or basal position of some partsbelonged to compound pinnate leaf.

Mineralized trunks of osmundacean ferns from thePermian beds of the Fore�Urals were described by theEnglish paleobotanists Kidston and Gwynne�Vaughan(1907).

The large�scale studies of Permian ferns from theFore�Urals were started by M.D. Zalessky. In 1925, hepublished a remarkable paper devoted to the sproutanatomy of the osmundacean ferns Thamnopteris kid�stonii Zalessky, T. gwynne�vaughanii Zalessky, andZalesskya uralica Zalessky from the Permian of theFore�Urals (Zalessky, 1925).

In addition to many endemic species, Zalessky(1918) determined in the Permo–Carboniferous ofSiberia a fern species from the Stephanian of Westernand Central Europe, Pecopteris (Asterotheca) oreopte�ridia (Schlotheim) Renault et Zeiller. Subsequently,Neuburg (1948) corroborated the presence of leaves ofthis type in the Upper Carboniferous of Siberia. Adetailed drawing of this leaf venation type was laterpublished by Meyen (1982b, 1990).

Zalessky (1933a, 1934b) and Zalessky and Tschirk�ova (1937) described several species of sphenopteroidfern leaves from the Carboniferous and Permian ofSiberia: Sphenopteris sibirica Zalessky, S. comptulaZalessky, S. ischanovensis Zalessky, S. batschatensisZalessky, S. praestans Zalessky, S. izylensis Zalessky,S. bellatula Zalessky, S. odontopteroides Zalessky,S. jagunovana Zalessky, S. asiatica Zalessky, S. eureaZalessky, and S. praesalairica Zalessky. Subsequently,Neuburg (1948) refigured and partially redescribedthese species, fairly indicating that, in many cases,comparison of these species with each other and pre�viously established species is complicated because thetype material is rather fragmentary. In addition to theredescription of species established by Zalessky, Neu�burg described the new species Sphenopteris intermediaNeuburg, S. imitans Neuburg, S. incrassata Neuburg,S. kumpanii Neuburg, S. erosilobata Neuburg, andremains of pinnate fernlike leaves determined in opennomenclature as Sphenopteris sp. “a” (for more detail,see discussion below).

In addition to sphenopteroid leaves, the UpperPaleozoic beds of Siberia have yielded the followingpinnate leaves, most of which apparently belong toferns: Pecopteris angaridensis Zalessky, P. abensisZalessky, P. alsophiloides Zalessky, P. martia Neuburg,P. comptula Zalessky, and P. leninskiensis (=P. hyper�borea) (Zalessky, 1933a, 1934b; Zalessky and Tschirk�ova, 1937). It should be noted that some of the abovepecopterid species (for example, Pecopteris abensis)were described based on apical frond fragments, inwhich the pinnule bases are usually wider than in themiddle part or at the base of fronds; therefore, someAngaraland species of Pecopteris Brongniart from theabove list may belong to the formal genus SphenopterisBrongniart.

In the paper describing plant remains from the Per�mian beds outcropping in the Pechora River Basin,Zalessky described several more fern species basedmostly on sterile leaves (Zalessky, 1934a): Pecopterishelenaeana Zalessky, P. synica (when characterizingthis species, Zalessky figured and briefly describedboth sterile and fertile leaves), P. tchernovii Zalessky,P. kojimensis Zalessky, P. comiana Zalessky, P. borealisZalessky, P. (Sphenopteris) ripensis Zalessky, and Isiol�opteris serrata Zalessky. The exact taxonomic positionof these species was not discussed at that time.Zalessky assigned them, along with the description ofpinnate pteridosperm leaves, to the section Filices etPteridospermae. Later, Zalessky (1939) described fernleaves from the Kungurian Stage of the middle Fore�Urals and referred them to Pecopteris uralica andP. suksunensis Zalessky.

Sphenopteroid leaves described by Zalessky(1937b) as Sphenopteris pennaticisa Zalessky from theKungurian beds of the Chekarda locality apparentlybelong to a gymnosperm plant of psygmophylloidmorphology rather than fern.



In a fundamental work of Neuburg (1948) devotedto the Late Paleozoic flora of the Kuznetsk Basin,ferns received much attention, although almost alldescriptions concern sterile leaves assigned to the for�mal genera Pecopteris and Sphenopteris. Neuburgdescribed the following species previously establishedby Schmalhausen, Zalessky, Radczenko, and otherpaleobotanists: Sphenopteris sibirica, S. comptula,S. ischanovensis, S. batschatensis, S. praestans,S. izylensis, S. eurina Zalessky, S. bellatula, S. tungus�cana (Schmalhausen) Zalessky, S. odontopteroides,S. jagunovana, S. asiatica, S. eurea, S. praesalairica,S. (Oligocarpia?) karpovii Radczenko, P. angaridensisZalessky, Pecopteris oreopteridia (Schlotheim) Renaultet Zeiller, P. abensis, P. alsophiloides, P. comptula,P. anthriscifolia, P. leninskiensis (Chachlov) Radc�zenko, along with several new taxa, i.e., S. intermediaNeuburg, S. imitans, S. incrassata, S. kumpanii,S. erosilobata, S. laxifolia Neuburg, and Pecopterismartia Neuburg and forms described in open nomen�clature (Sphenopteris sp. “a”). It is highly probablethat a large part of species established based on sphe�nopteroid leaves are synonyms; however, to state thiswith confidence a detailed study of variation and fer�tile organs is required. The species Sphenopteris odon�topteroides and S. kumpanii apparently belong to thepteridosperm group of Paragondwanidium rather thanferns, as Neuburg (1948) assumed in her remarks, andMeyen (1982b) corroborated this opinion, proposingnew combinations, Paragondwanidium odontopteroi�des (Zalessky) S. Meyen and Paragondwanidium kum�panii (Neuburg) S. Meyen.

Simultaneously with the improvement of fern sys�tematics based on the structure of fertile leaves, theformal generic taxonomy of sterile leaves was alsodeveloped. Special attention was paid to the venationand ontogenetic development of the leaf lamina. Inparticular, within a wide pecopterid morphologicalgroup, it was possible to recognize the genus Lobatop�teris Wagner (Wagner, 1958), which was initially con�sidered to be characteristic only of the Upper Carbon�iferous and Lower Permian beds of Western Europeand North America, but was later recorded in theUpper Permian beds of the Pechora Coal Basin(Pukhonto and Fefilova, 1983) and ArkhangelskRegion (Ignatiev and Naugolnykh, 2001; see hereFig. 22h).

In an extensive monograph devoted to the descrip�tion of ferns and fernlike plants from the Permian ofthe Fore�Urals near the Pechora River (Fefilova,1973) and a detailed report on the Permian System ofthe Fore�Urals near the Pechora River (Pukhonto andFefilova, 1983), the following fern species aredescribed and figured in photoplates: Asterotheca (?)kojimensis, A. (?) pluriseriata Fefilova, Asterotheca sp.,“Orthotheca” petschorica Fefilova (probable synonymsof this species include Pecopteris helenaeana), Todites sp.,Prynadaeopteris ambigua Fefilova, P. karpovii Radc�

zenko, P. silovaensis Fefilova, P. tschernovii (Zalessky)Fefilova, P. venusta, P. vorcutana (Zalessky) Fefilova,P. (?) alifera Fefilova, Pecopteris anthriscifolia, P. bore�alis, P. compta Radczenko, P. helenaeana, P. hyper�borea, P. micropinnata Fefilova, P. nelynensis Fefilova,P. niamdensis Zalessky, P. obtusa Radczenko, P. synicaZalessky, P. varsanofievae Fefilova, P. verecundaeFefilova, P. aff. hyperborea, P. (?) dissimilaris Fefilova,Cladophlebis aff. nystroemii Halle, Lobatopteris sp.,Sphenopteris cuneata Neuburg et Fefilova, P. pumilaFefilova, and S. stenophylla Fefilova. Later, Pukhonto(1998) figured some of these species (Prynadaeopterisvorcutana, P. venusta, “Orthotheca” petschorica, etc.).

Fefilova (1973) established for the first time thepresence of the marattialean genus Asterotheca(Corda, 1845) in Angaraland (Subangaraland) floras.It should be noted that the genus Asterotheca in thepresently accepted sense comprising star�shapedsynangial aggregations of sporangia belonging toMarattiaceae is used in the description of both Paleo�zoic and Mesozoic floras. Moreover, the genus Aster�otheca is often assigned to the family Asterothecaceaeinstead of the family Marattiaceae (Holmes, 2001). Insome Triassic floras, members of Asterotheca are ratherdiverse. For example, the Middle Triassic megafossilflora from the Basin Creek Formation of the Nym�boida coal measures, New South Wales, Australia con�tains four species of Asterotheca (Asterotheca trullensisHolmes, A. nymboidensis Holmes, A. chevronerviaHolmes, and A. diameson Holmes) and four more speciesare established in open nomenclature: Asterotheca sp. A,Asterotheca sp. B, Asterotheca sp. C, and Asterotheca sp. D(Holmes, 2001).

Holmes (2001, p. 43) provided the following char�acteristics of ferns included in the genus Asterotheca:“Asterotheca is a genus erected for fertile ferns withpecopteroid pinnules which bear a line of adjacentsynangia on the abaxial surface between the midveinand the margin. The synangia are composed of groupsof sporangia conjoined at the base and dehiscing alongthe apical suture line. An annulus is absent. The genusincludes species from both the northern and southernhemispheres which range in age from Carboniferousto Upper Jurassic. It is most probably not a naturalgenus.”

In such a wide sense, the name Asterotheca isapplied as a formalized classification unit to ferns fromCarboniferous and Permian deposits almost through�out the world, including the Permian of Gondwana(Cuneo et al., 2000).

In particular, remains of a fertile fern frond fromthe Chekarda locality (apparently Chekarda 1) wereinitially assigned to the genus Asterotheca by Meyen(1982b, 1990, pl. XXXVII, fig. 5). At present, I refermorphologically similar forms from the same localityto the species Convexocarpus distichus, comb. nov. (seedescriptions below).

1386


NAUGOLNYKH

In a detailed review of Carboniferous and Permianfloras of Angaraland, Meyen (1982b) provided thecharacteristics of higher plant groups and the data onthe morphology and taxonomic position of Late Pale�ozoic ferns of Angaraland (this work was subsequentlypublished in Russian translation: Meyen, 1990).Meyen tentatively divided all ferns from the Carbonif�erous and Permian of Angaraland into six groups:(1) Early Carboniferous Chacassopteris Radczenko;(2) endemic species of the cosmopolitan morphogen�era Sphenopteris and Pecopteris with unknown fertileorgans; (3) endemic Angaraland genera SiberiellaRadczenko (non Siberiella K.B. Korde, 1957, a greenalga genus, junior homonym of the genus SiberiellaRadczenko, 1955), Prynadaeopteris Radczenko(emendation of this genus is given below), GeperapterisS. Meyen (this genus was validly established in a differ�ent work of Meyen which was also published in 1982,see below); (4) Asterotheca, Danaeites, and Oligocar�pia, which Meyen considered to be typical genera ofthe Permo–Carboniferous of the Euramerian Realm;(5) petrified sprouts and trunks of Osmundaceae (heremarked that, at the moment of publication of thiswork, imprints of their fronds had not been recordedin this region); and (6) “mesophytic” genera from theterminal Permian and Permo–Triassic, including theosmundaceans Todites Seward and CladophlebisBrongniart. Meyen believed that the species describedby Fefilova as A. (?) pluriseriata should also be assignedto the genus Todites. I adhere to this point of view.

In the paper devoted to the establishment of thenew genus Geperapteris and characteristics of its sporespreserved in situ, Meyen excluded some species(Prynadaeopteris silovaensis, P. tchernovii) from thegenus Prynadaeopteris and transferred them, alongwith the species Sphenopteris imitans, to the genusGeperapteris (Meyen, 1982c). Geperapteris imitans(Neuburg) S. Meyen was designated as the type spe�cies of Geperapteris.

Gomankov (in Gomankov and Meyen, 1986,pp. 36–38) established the new genus FefilopterisGomankov and referred it to ferns based on “theabsence of cuticle resistant to maceration and generalmorphology” (Gomankov and Meyen, 1986, p. 37).The material described by Gomankov comes from twolocalities, Aristovo (holotype) and Isady. Both out�crops are in the Sukhona River Basin (VologdaRegion). In the part Remarks, the type species Fefilop�teris pilosa Gomankov is compared mostly with thegymnosperm genera Glossopteris Brongniart andRhabdotaenia Pant, which frequently have a thin cuti�cle nonresistant to maceration (K.J. Singh, personalcommunication). In 2012, Karasev (PIN) showed metaeniopteroid leaves from the Isady locality, which areexternally very similar to the leaves described byGomankov as Fefilopteris pilosa. In my opinion, thereis no reason to assign these leaves to ferns. They aremuch more similar to leaves of the cycadophyte Tae�niopteris eckardti Germar known from the Upper Per�

mian beds (Zechstein) of Western Europe (Weigelt,1930, pp. 647–649, text�figs. 13–16; Stoneley, 1958,pl. 37, figs. 3a, 3b). “Wheels” (according to the termi�nology of Gomankov in Gomankov and Meyen, 1986,p. 37), which were interpreted in the original descrip�tion as “places of attachment of sporangia,” are in myopinion fruit bodies of the fungi Phyllachora (seeabove). Unfortunately, description of the genusFefilopteris is not accompanied with the section Com�parison, preventing comparative analysis of this genusin the sense of its author with other genera.

Esaulova (1996) described fertile remains of fernsfrom the Kazanian Stage of the Volga and Lower Kamaregions: Acitheca gigantea Esaulova, Todites gracilisEsaulova, and Oligocarpia kamiensis Esaulova. Sheindicated co�occurrence of leaves of Todites graciliswith trunks of Thamnopteris schlechtendalii (Eich�wald) Schimper. Unfortunately, Esaulova did notdescribe in detail the fertile organs of those ferns and,therefore, the generic assignment of these forms isdoubtful.

A great achievement in the study of Paleozoic fernswas comprehension of the fact that the fern leafchanges in the course of ontogenetic development,with the transformation of both morphology and orderof pinnation pattern in the segments of the compoundleaf. It is necessary to use the character of thesechanges in descriptions of species and diagnoses ofgenera of fossil ferns (Danze, 1955).

I described in the previous studies the marattialeanfern species Ptychocarpus distichus (Naugolnykh,1995), Orthotheca semilibera Naugolnykh (Naugol�nykh, 1998a), Orthotheca dicranophora Naugolnykh(Naugolnykh, 1998c), and the osmundacean speciesTodites lobulatus (Naugolnykh, 2002). Subsequently,in the monograph (Naugolnykh, 2007), in text�fig. 50showing the morphology and anatomy of Convexocar�pus (earlier Ptychocarpus) distichus, comb. nov., thescale bar is only provided to text�fig. 50A; that of text�fig. 50B is 1 mm. In text�fig. 52 of the same work, thescale bar is 2 mm (see also Naugolnykh, 2009, p. 18,text�fig. 1d). Fortunately, this inaccuracy does notresult in considerable difficulties, because the samespecimens are shown in plates and accompanied withexact data on magnification.

At present, it becomes particularly important forthe study of the morphology and taxonomy of Permianferns of the Fore�Urals to apply new taxonomicapproaches, primarily the study of variability, particu�larly variability of pinnule outlines and venation withinindividual fronds.

DESCRIPTIVE MORPHOLOGY AND TERMINOLOGICAL SYSTEM

Despite the general unification of descriptive mor�phology of compound pinnate leaves typical for fossilferns, but also observed in various seed plant groups (in



particular, pteridosperms, which often co�occur inCarboniferous and Permian localities with remains oftrue ferns), some terms are differently understood bypaleobotanists. In addition, the same morphologicalstructures are sometimes designated by differentterms. The glossary of the main morphological termsused in the present study is given below.

The frond is a flattened foliar organ with a pinnatedesign. It may be a true leaf, or has a cladodial nature.The order of pinnation can change (usually increases)in the frond of the same plant, as the frond grows in thecourse of ontogeny and also due to seasonal changes inenvironments.

The pinna (penultimate order segment) is a frondsegment of the penultimate and all preceding orders.The frond was monopinnate, bipinnate, or, less fre�quently, tripinnate or even quadripinnate. The higherpinnation orders are extremely rare. In the frondstructure, I recognize the terminal pinna correspond�ing to the frond apex and having pinnules, instead ofpinnae, in contrast to the middle and basal frond parts(Fig. 4, TP) and lateral pinnae (pinnae of the lastorder, or, to put it differently, segments of the penulti�mate order) positioned on the main rachis of the frondin its middle and basal parts. The lateral pinnae areusually narrower than the terminal pinna of the samefrond.

The pinnule is the last order segment of the frond.It is either entire or lobate. In the course of frond mor�phogenesis, some pinnules (primarily apical) trans�form from entire into lobate and, then, into evendeeper dissected and, finally, becoming young pinnae(segments of the penultimate order). Laminae of theformer pinnule are transformed into new segments ofthe last order (young pinnules). The pinnal apex fre�quently has a terminal pinnule (Fig. 4, TPL), whichdiffer in morphology from ordinary pinnules of thesame frond.

The rachis is the axial part of the frond, where thesegments are attached to. It is rounded or subtriangu�lar in cross section, grooved, naked or winged. In somespecies, it has additional morphological elements inthe shape of hairs or scales.

The limb (wing) is foliaceous projections posi�tioned on the margins of the rachis and usually formedby the decurrent part of the leaf lamina of pinnules. Inaddition, the term limb frequently designates therachis wings.

The coalescence is fusion of pinnules at the base oreven by the margins. This is frequently observed in api�cal pinnules and, in some species, within the entirefrond.

The sporangium is a depository of spores, which isusually rounded or elliptical. The sporangia are singleor, growing together, form aggregations named sori (ifsporangia have pedicels (sporangiophores), which areattached to the common base) or synangia (if sporan�

gia are fused by their margins to form linear or rosette�like aggregations).

The midvein is the axial vein of pinnules (=segmentsof the last order), which diverges directly from the pin�nal rachis (=segment of the penultimate order). Themidvein is frequently termed the main or primary vein.

Lateral veins are the veins deviating at variousangles and in various order from the midvein of thepinnule. They are either simple or dichotomize severaltimes. They are frequently designated side (lateral) orsecondary veins.

Topological homology is the genetic correspon�dence between morphologically different structures,primarily, between segments of the last and penulti�mate orders in forms with a reduced leaf lamina, seg�ments of the last order of which correspond to the lat�

TPL

TP

AL

KL

LP

TPL

FR

~

Fig. 4. Morphology of the compound pinnate leaf of fernsand terminological system used in the present work:(FR) frond rachis; (LP) lateral last order pinnae;(TP) frond apex, sometimes representing a modified ter�minal last order pinna; (TPL) terminal pinnule;(AL) anadromic lobe; and (KL) catadromic lobe.

1388


NAUGOLNYKH

eral veins of pinnules of the initial (ancestral) form,while segments of the penultimate order correspond topinnules, in contrast to forms with a nonreduced leaflamina. Each case of the nonstandard topologicalhomology is indicated in descriptions.

PALEOBOTANICAL DESCRIPTION

D I V I S I O N P T E R I D O P H Y T A S C H I M P E R , 1 8 7 9

C L A S S M A R A T T I O P S I D A TA K H T A D J A N , 1 9 8 6

SUBCLASS EUSPORANGIATAE GOEBEL, 1882

Order Marattiales Prantl, 1874

Family Marattiaceae Bertchtold et J. Presl, 1820

Genus Acrogenotheca Naugolnykh, gen. nov.

E t y m o l o g y. From the Latin acrogenus (formedat the top).

Ty p e s p e c i e s. Acrogenotheca ramificataNaugolnykh, sp. nov.

D i a g n o s i s. Fertile pinnules without leaf lam�ina, possessing radially symmetrical synangia com�posed of four sporangia almost completely fused bytheir margins. Each synangium having basal blister.Spores monolete, with smooth sporoderm.

C o m p a r i s o n. The genus Acrogenotheca gen.nov. is most similar in morphology to the genus Syd�neia Psenicka and Bek, Zodrow, Cleal et Hemsley(Psenicka et al., 2003) and differs from it in the radi�

ally symmetrical rather than bilaterally symmetricalsynangia, the significantly fewer sporangia in thesynangium (Acrogenotheca gen. nov. has four, whileSydneia has up to 25), and in the presence of a basalblister of the synangium. In addition, the rachis andsynangiophores of Sydneia are covered with fine tri�chomes, while Acrogenotheca gen. nov. lacks them.

S p e c i e s c o m p o s i t i o n. Type species.

Acrogenotheca ramificata Naugolnykh, sp. nov.

Plate 2, figs. 1–6; Plate 3, figs. 1–6; Plate 4, figs. 1–6; Plate 5, figs. 1–5; Plate 6, figs. 1–5

Calymmotheca (?) sp.: Naugolnykh, 1995, text�fig. 6G; Gruntet al., 1998, pl. XXXIII, fig. 5.

E t y m o l o g y. From the Latin ramificatus(branching), after branching lateral veins of fertile seg�ments with a reduced leaf lamina.

H o l o t y p e. GIN, no. 3773(11)/318(92), middlepart of a bipinnate leaf, Chekarda 1 locality, Bed 10,designated here (Figs. 5–7; Pl. 2, figs. 1–6; Pl. 3,figs. 1–6; Pl. 4, figs. 1–6; pl. 5, figs. 1–5; Pl. 6, figs. 1–5).

D i a g n o s i s. Homosporous eusporangiate fernwith compound pinnate leaves with reduced leaf lam�ina or even without leaf lamina. Lateral veins withhydathods. Some lateral veins of last order segments(on average, two or three of nine–ten veins) havingterminally positioned synangia, consisting of fourmarginally fused sporangia. Spores rounded or ovoidin outlines, monolete, with smooth sporoderm, about25 µm in diameter.

D e s c r i p t i o n (Figs. 5–7). The fronds are usu�ally compound pinnate with at least two orders of pin�nation. Fertile leaves have many synangia at the endsof synangiophores. In my opinion, synangiophores aretopologically homologous to lateral veins of fertilepinnules of the phylogenetically initial ancestral form(see above in the section Terminology). The specimenlacks a trace of the leaf lamina. The frond fragment is10 mm long and 8 mm wide.

The frond rachis is slightly curved, almost straight;it is 0.8 mm wide. The axial part of the rachis has a0.3–0.4�mm�wide groove complicated by hardly dis�cernible shallow folds.

Segments of the penultimate order diverge from therachis at an angle of 55°; they are apparently topolog�ically homologous to the midveins of pinnules with areduced leaf lamina. Five segments of the penultimateorder are completely preserved. In addition, the basesof two more segments of the penultimate order areobserved. Segments of the penultimate order are 4.5–5 mm long at the maximum width of rachises of0.3 mm.

Each segment of the penultimate order has seg�ments of the last order, which are topologically homol�ogous to the lateral veins of fertile pinnules. Most ofthe segments of the last order are sterile, although fer�tile segments with terminal synangia are also present;

Fig. 5. Morphology of a frond fragment of the marattialeanfern Acrogenotheca ramificata Naugolnykh, gen. et sp.nov., holotype GIN, no. 3773(11)/318(92). Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10; LowerPermian, Kungurian Stage. Scale bar, 0.5 cm.



segments with synangia (synangiophores) are usuallyanatropously curved. Each segment of the penultimateorder has two or three synangia. Some segments of thelast order expand apically to form fusiform thickeningof hydathods. Two apical segments of the last orderform a regular fork. In other words, the apex of eachsegment of the penultimate order regularly dichoto�mizes. The length of segments of the last order rangesfrom 2 mm (segments of the last order positioned inthe basal part of segments of the penultimate order) to0.5 mm (segments of the last order located in the api�cal part of segments of the penultimate order). Seg�ments of the last order are attached to segments of thepenultimate order at an angle of 40°–45°.

The conducting system consists of thin elements ofthe protoxylem surrounded by thicker metaxylemformed of elements with ring and spiral thickenings ofwalls. The external zones of a conducting bundle arecomposed of mechanical tissues.

Fertile segments of the last order (synangiophores)are strongly curved in the apical part and possess synan�gia. The synangia are 1 mm long and 0.5–0.8 mm wide.Each synangium consists of four sporangia almostcompletely fused by the margins. Only the very distalmargins of sporangia are free. The wall of sporangiawas multilayered and consisted of at least two or threecell layers.

In almost all observable synangia, the base has aspherical or ellipsoidal bulge (blister), which eitherdirectly adjoins the fused sporangia or is positioned ata distance of 0.5–0.6 mm from them. The bulge size ison average 0.3 × 0.4 mm.

Many spores are preserved inside the sporangia.One of the best preserved sporangia contained at least95 spores, although, in the living plant, the number ofspores was probably even greater, because this spo�rangium was already open through a longitudinalcrack and some spores could have dropped out beforeit was buried.

The spores are rounded in outline, spherical, orellipsoidal, 22–25 µm in diameter. Occasionally, thespores are larger, up to 30 µm in diameter. Many sporesare distorted, shaped as a boat or depressed, as thoughcontorted at the center. The spores are monolete; how�ever, some specimens have an additional short ray mak�ing the scar asymmetrical triradiate in outline. Thesporoderm surface is smooth, but sometimes compli�cated with small weakly developed granules.

R e m a r k s. In my opinion, the extreme rarity ofAcrogenotheca ramificata gen. et sp. nov. in orycto�coenoses probably results from the fact that these fernsgrew at a large distance from the water basin (lagoon),where the thanatocenosis was formed, comparably tothe plants of near�shore communities, which prevailin some interbeds of the Chekarda 1 locality. Judging

Fig. 6. Morphology of a frond fragment of the marattialean fern Acrogenotheca ramificata Naugolnykh, gen. et sp. nov., holotypeGIN, no. 3773(11)/318(92), Detailed drawing of the synangial structure. Middle Fore�Urals, Perm Region, Chekarda 1 locality,Bed 10; Lower Permian, Kungurian Stage. Scale bar, 1 mm.

1390


NAUGOLNYKH

from the morphology of the object (holotype), it seemsplausible that, by the moment of maturation, synangiahung down because of the absence of any other sup�port, except for a thin synangiophores, because theleaf lamina was completely reduced. The basal blisterof synangia resembles in extern shape brightly coloredaril of some extant angiosperm fruits (for example, thespindle�tree Euonymus L.), intended for attraction ofanimals serving as disseminators.

The synangia of A. ramificata gen. et sp. nov. havemuch in common with the synangia of Burnithecapusilla Meyer�Berthaud et Galtier from the LowerCarboniferous of Scotland (Meyer�Berthaud andGaltier, 1986). The main difference is the muchgreater number (eight versus four) of sporangia in each

synangium of Burnitheca. In addition, the synangia ofB. pusilla are distinctly asymmetrical, so that on one ofside of the synangium, they are much longer than onthe other side, although they gradually change inlength from one side of the synangium to the other. InAcrogenotheca ramificata, all sporangia of onesynangium are approximately equal in length.

The spores found in situ in the sporangia of Burni�theca pusilla are also similar to spores of Acrogenothecaramificata gen. et sp. nov., but differ distinctly in thetrilete pattern and well�developed sculpting in theshape of small and widely spaced spines and tubercles.The spores similar to spores of A. ramificata gen. et sp.nov. are known in many species of extinct and extantMarattiaceae. Very similar monolete spores have been

(a)

(b)(c)

(d) (e) (f)

Fig. 7. Morphology of a frond fragment of the marattialean fern Acrogenotheca ramificata Naugolnykh, gen. et sp. nov., holotypeGIN, no. 3773(11)/318(92): (a, c, f) structural details of synangia with a basal blister, (d) spores preserved in situ, (e) synangiumfrom which the in situ spores were extracted, and (b) reconstruction of fertile pinna fragment with two synangia. Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10; Lower Permian, Kungurian Stage. Scale bar, 1 mm (except for the left lowerfigure, in which scale bar is 0.5 mm.



Plate 1

1 2 3 4

5 67

8

10

11 12 13 14

E x p l a n a t i o n o f P l a t e 1

Figs. 1–14. The main morphological types of fertile organs of ferns from the Carboniferous and Lower Permian of Europe (afterStur, 1883): (1, 2) Asterotheca sternbergii Goeppert; (3) Oligocarpia gutbierii Goeppert; (4) Danaeites sarepontanus Stur;(5) Calymmotheca minor Stur; (6) Calymmotheca schatzlarensis Stur; (7, 8) Oligocarpia lindsaeoides Ettingshausen; (9, 13) Discop�teris schumannii Stur; (10) Scolecopteris arborescens Schlotheim; (11) Scolecopteris polymorpha Brongniart; (12) Discopteris kar�winensis Stur; and (14) Scolecopteris cyathea Schlotheim. Not to scale.

1392


NAUGOLNYKH

Plate 2

1

2

3

4 5 6

E x p l a n a t i o n o f P l a t e 2Figs. 1–6. Frond morphology of the marattialean fern Acrogenotheca ramificata Naugolnykh, sp. nov., holotype GIN,no. 3773(11)/318(92): (1) general morphology; (2, 3, 5, 6) structure of individual synangia: (2, 5) synangium that provided in situspores (see fig. 4 on this plate and Pls. 5 and 6); (3, 6) synangia with distinct basal blister (positioned above the synangium);(4) spores preserved in situ. Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10. Lower Permian, Kungurian Stage.Scale bars: (1) 5 mm, (2, 3, 5, 6) 1 mm, and (4) 100 µm.



Plate 3

1

2

3

4

65


Figs. 1–6. Structure of synangia and venation of the marattialean fern Acrogenotheca ramificata Naugolnykh, sp. nov., holotypeGIN, no. 3773(11)/318(92): (1, 2, 4–6) synangia: (2, 4–6) preserved in natural attachment to synangiophores; (3) character ofvenation, with distinct middle and lateral veins. Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10; Lower Permian,Kungurian Stage. Scale bar, 500 µm.

1394


NAUGOLNYKH

Plate 4

1

2

4

3

5 6

VE

P

VE


Figs. 1–6. Structure of the rachis of the last order segment of the marattialean fern Acrogenotheca ramificata Naugolnykh, sp. nov.,holotype GIN, no. 3773(11)/318(92); (P) micropores in the wall of a conducting element and (VE) conducting vascular element.Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10. Lower Permian, Kungurian Stage. Scale bars: (1, 6) 200 µm,(2, 4) 100 µm, (3) 20 µm, and (5) 500 µm.



Plate 5

1

2

3

4 5


Figs. 1–5. The marattialean fern Acrogenotheca ramificata Naugolnykh, sp. nov., holotype GIN, no. 3773(11)/318(92), sporespreserved in situ. Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10. Lower Permian, Kungurian Stage. Scale bars,100 µm.

1396


NAUGOLNYKH

Plate 6

1

2

3

4 5


Figs. 1–5. The marattialean fern Acrogenotheca ramificata Naugolnykh, sp. nov., holotype GIN, no. 3773(11)/318(92), sporespreserved in situ. Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10. Lower Permian, Kungurian Stage. Scale bars,20 µm.



extracted from the sporangia of the marattialean fernQasimia lanceolata Wang et Yang, 1996 (Wang andYang, 1996; pl. 1, figs. 7–9).

The exact taxonomic position of Burnitheca pusillaremains uncertain. The authors of this taxon areinclined to assign it to either the family Marattiaceaeor male fructifications of pteridosperms, proposing,however, that this plant is more likely related to ferns(Meyer�Berthaud and Galtier, 1986).

M a t e r i a l. Holotype.

Genus Convexocarpus Naugolnykh, gen. nov.

E t y m o l o g y. From the Latin convexus (convex),based on the convex ornamentation of the synangia.

Ty p e s p e c i e s. Convexocarpus distichus(Naugolnykh) Naugolnykh, comb. nov.

D i a g n o s i s. Fertile pinnules pecopteroid, spo�rangia arranged in relatively short radially symmetricalsynangia with elements of bilateral symmetry, formedby fusion of proximal side of synangium with leaf lam�ina of fertile pinnule. Synangia arranged in two rowsalong midvein of fertile pinnule, on its abaxial surface.Spores round, trilete. Rays of trilete scar occupyingtwo�thirds of spore radius. Sporoderm covered by finescabrate ornamentation.

C o m p a r i s o n. The new genus is most similar toEoangiopteris Mamay (Mamay, 1950), GrandeuryellaStur (Stur, 1883), and Millaya Mapes et Schabilion(Mapes and Schabilion, 1979b) and differs from themin the orientation of sporangia in the synangium; inConvexocarpus, they are inclined or even parallelrather than perpendicular to the abaxial surface of theleaf lamina. The new genus differs from Danaeites Stur(Stur, 1883) in the radially symmetrical synangia. Itdiffers from the genus Ptychocapus Weiss (in the initial,narrow sense) (Weiss, 1869) in the biserial arrange�ment of synangia on the abaxial surface of fertile pin�nules; from the genera Acitheca Schimper (Schimper,1879) and Scolecopteris Zenker (Zenker, 1837) in thefusion between the distal synangial side and abaxialsurface of the leaf lamina of the fertile pinnule.

R e m a r k s. The type species of the genus Con�vexocarpus was originally referred to the genus Pty�chocarpus Weiss (Naugolnykh, 1995). The formsdetermined as Ptychocarpus sp. were recorded in thesame beds (Vladimirovich, 1985, 1986), although theyhave not been described. At present, the volume andmorphological diagnostics of the genus Ptychocarpusare essentially differently understood by researchers.

In particular, the well�known handbook Funda�mentals of Paleontology provides the following charac�teristics of the genus Ptychocarpus: “The fossil recordprovides fertile and sterile leaves. The leaves are com�pound pinnate; the rachis is finely tuberculate. Thepinnae are alternating, linear or linear–lanceolate,adjoining or slightly overlapping each other. The pin�nules are alternating, with parallel margins and

rounded apex, with an entire or dentate margin, fusedwith each other to a greater or lesser extent. Venationis pinnate; the midvein reaches the pinnule apex,where it is divided into two; the lateral veins are alwayssimple. The synangia are in the shape of a truncatecone, consist of five or six sporangia around the pla�centa (receptacle), which encloses a vascular bundle”(Vakhrameev et al., 1963, p. 567).

This description is undoubtedly based on the mor�phology of the species Scolecopteris unitus, sinceVakhrameev et al. (1963) designated Ptychocarpus uni�tus (Brongniart) Zeiller as the type species of the genusPtychocarpus; however, this is incorrect, because, asthe genus was established, it included the only speciesP. hexastichus Weiss, which should be taken for thetype species. Certain minor discrepancies between thecharacteristics and figures in the same work (in partic�ular, in text�fig. 66, each synangium contains seven oreight sporangia, in contrast to five or six sporangia, asindicated in the description) look insignificant againstthis background.

Some forms resembling P. hexastichus wererecorded in other regions, for example, Ptychocarpussp. from the Upper Carboniferous of Canada (Zodrowand McCadlish, 1980). However, in contrast to classi�cal P. hexastichus, in Fore�Ural Ptychocarpus�likeforms, the abaxial surface of fertile pinnules had onlytwo longitudinal rows of synangia instead of six, as isreflected in the species epithet given by the author.Based on a number of morphological features (seebelow description of the species Convexocarpus disti�chus), the species Ptychocarpus distichus is transferredin the present study to a new genus, Convexocarpus,gen. nov.

The structure of complete fertile fronds of the ferngenus Convexocarpus is not known; however, judgingfrom available fragments, they were at least tripinnate(probably quadripinnate) and subtriangular in outline.

S p e c i e s c o m p o s i t i o n. Type species.O c c u r r e n c e. Lower Permian, Kungurian Stage

of the middle Fore�Urals.

Convexocarpus distichus (Naugolnykh, 1995) Naugolnykh, comb. nov.

Plate 7, figs. 1–7; Plate 8, figs. 1 and 3

Ptychocarpus distichus Naugolnykh: Naugolnykh, 1995,pp. 57–60, pls. I, II, figs. 5A–5C; Naugolnykh, 1998c, pp. 58–60,text�figs. 20a–20c, pl. VIII, figs. 1–4; Grunt et al., 1998,pl. XXXIII, fig. 3; pl. XXXIV, figs. 1–3 and 5; Naugolnykh, 2007,text�figs. 50A, 51F; pl. VIII, fig. 7; pl. IX, fig. 4; pl. XIII, fig. 4;pl. XIV, figs. 3 and 7; pl. XVI, fig. 4.

Ptychocarpus (?) sp.: Naugolnykh, 1998c, text�figs. 20F–20H;pl. VIII, fig. 1.

H o l o t y p e. GIN, no. 3773(11)/27(89), middlepart of a fertile frond; Chekarda 1 locality, Bed 10; itwas figured by Naugolnykh (1995, text�fig. 5A–5C,pl. I, figs. A–D, pl. II, figs. A–C.

D e s c r i p t i o n (Fig. 8). The collection understudy contains eight variously preserved fertile leaves

1398


NAUGOLNYKH

2

3 4

5

6 7 8

1

Plate 7



and ten sterile leaves, which probably belong to thesame species. The most representative remains of fer�tile leaves belong to either the middle part of bipinnatefronds, or apical parts of similar fronds. The frond api�ces are typically wide triangular in outline and treatedas the terminal pinnae (see Fig. 4 and TerminologicalSystem), the pinnules of which become lateral pinnaeof the last order in the course of the development offronds. In contrast to the wide triangular outline of theapical frond pinna, the lateral pinnae of the last orderhave subparallel margins. The lateral pinnae of theapical frond part are relatively widely spaced, althoughthe spaces between them are less than the pinna width.In the middle frond part (pinna of the penultimateorder), the lateral pinnae (pinnae of the last order) arepositioned more closely, almost adjoin each other bythe basal margins, but never overlap a neighboringpinna. Judging from the subparallel margins of pinnaeof the penultimate order of well�developed fronds,they apparently formed a pinnate structure of a higher(third) order, and this frond could have been rather

large, reaching at least 1 m in length, as is judged fromthe rate of shortening the pinnae towards the frond tip.

The frond apical pinna (pinna of the ultimate orderor the last order pinna) is sometimes transformed intoa weakly developed terminal pinnule formed by coher�ent fusion of small undeveloped apical pinnules.

The frond rachis is straight, sometimes slightlycurved. A longitudinal groove extends along the axialpart of the rachis. The rachis is 3–4 mm wide and themaximum groove width is 2 mm. The rachis of the api�cal part of pinnae of the penultimate order usually hasa thin limb (wing) formed of decurrent bases of apicalpinnules (pinnules of the apical pinna of the frond)and lateral pinnae. The rachis of the middle and basalparts of the frond is naked; in this case, the limb isabsent.

Pinnae of the last order (lateral pinnae) divergefrom the rachis of the frond (pinna of the penultimateorder) at an angle of 70°–75°. Pinnae of the last ordergradually increase in length towards the frond base and

1 2 3 4

Fig. 8. Morphology of a frond fragment of the marattialean fern Convexocarpus distichus (Naugolnykh) Naugolnykh, comb. nov.,reconstruction based on the drawing of specimen GIN, no. 4856/141 (Pl. 7, figs. 2, 5, 6). Right upper figure is schematic longi�tudinal section through synangium (magnified). Designations: (1) conducting tissues in pinnule; (2) conducting bundle extend�ing to receptacle of synangium; (3) cavities of sporangia; (4) tissues of mesophyll in fertile pinnule and multilayered wall of spo�rangia. Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10; Lower Permian, Kungurian Stage. Scale bar, 1 cm.


Figs. 1–8. Marattialean ferns of the Fore�Urals: (1) in situ spore of Verrucosisporites–Punctatisporites intermediate morphologi�cal type, extracted from (3) sporangium of Convexocarpus distichus (Naugolnykh) Naugolnykh, comb. nov., specimen GIN,no. 3773(11)/286(91); (2–7) morphology of fertile leaves of the marattialean fern Convexocarpus distichus comb. nov.: (2) speci�men GIN, no. 4856/141, structure of fertile last order pinna, direct light; (3) specimen GIN, no. 3773(11)/286(91), well�devel�oped fertile last order pinna; (4) , specimen GIN, no. 3773(11)/255(90), middle part of fertile last order pinna, with the apex lostbefore fossilization; (5) specimen GIN, no. 4856/141, structure of four synangia, with the free ends of sporangia turned down�wards; (6) specimen GIN, no. 4856/141, structure of the middle part of fertile last order pinna in slanting light; (7) specimenGIN, no. 4856/245, middle part of fertile last order pinna without apex; (8) specimen GIN, no. 4856/142, sterile last order pinnaof Pecopteris uralica Zalessky, probably belonging to Convexocarpus distichus comb. nov. (see also Fig. 21f). Middle Fore�Urals,Perm Region, Chekarda 1 locality; Lower Permian, Kungurian Stage: (1, 3, 4) Bed 7 and (2, 5–8) Bed 10. Scale bars: (1) 10 µm,(2–4, 6–8) 1 cm, and (5) 1 mm.

1400


NAUGOLNYKH

reach 70 mm. Well�developed pinnae of the last orderare almost constant in width, 10–12 mm, except forthe pinna apex. The rachis of the last order pinnae is atmost 1 mm wide. The apex of the last order pinnae isrounded. Apical pinnules of the last order pinna areusually coherently fused to form a single subtriangularterminal pinnule with rounded tips of blades, each ofwhich corresponds to the initial fused pinnule.

Synangial structure (Pl. 7, figs. 3, 5, 6; Fig. 8). Thesynangia are radially symmetrical, positioned close toeach other in two parallel rows extending along themidvein of fertile pinnule. The synangia vary from 1.3to 1.8 mm in diameter. Each synangium consists ofeight to ten sporangia fused at the base. The apicalparts of sporangia are attenuated, pointed. Theyremain free. The synangia are usually markedly asym�metrical, because one side is fused with the leaf laminaof fertile pinnule. The most developed fertile pinnuleshave six or seven pairs of synangia. The synangia areattached to the basal part of lateral veins, usually at adistance of 0.3–0.4 mm from the midvein of fertilepinnule. Each synangium consists of eight or, lessoften, nine or ten sporangia, which are fused by mar�gins almost throughout the length, except for the api�cal part. The central part of the synangium containedthe receptacle, providing attachment for sporangia.The sporangia are elongated, 0.7 mm long and 0.2 mmof maximum width, with rounded bases and pointedor extended apex. The sporangial wall is multilayered,consisting of four or five cell layers. Since the synangiaare usually preserved lying on one side, only four spo�rangia located on the side of the synangium turned tothe observer are usually seen. The synangium is0.7 mm long and of 0.6 mm of maximum width.

New material has provided improved morphologi�cal characteristics of in situ spores of this fern. Thespores are round in outline, with a distinct trilete tet�rad scar, 40–50 µm in diameter. The scar rays do notreach the spore margin (equator), terminating at a dis�tance of one�third of the spore radius. The spores areslightly bordered, slightly flattened in the equatorialpart. The sporoderm surface is shagreen, with smallgrooves and tubercles about 0.01 µm in diameter,sometimes with rare verrucae.

R e m a r k s. The formal species Pecoperis uralica,which was apparently established from a sterile pinnaof the last order belonging to the marattialean fernConvexocarpus distichus, comb. nov., has much incommon with the species Pecopteris unita Brongniart

from the Upper Carboniferous and Lower Permian ofEuramerica (Jennings and Millay, 1979), as Zalessky(1939) marked already in the original description ofthis species. This species is also similar to other pecop�terids from the Upper Paleozoic Euramerica. Forexample, P. uralica is very similar to juvenile pinnae ofP. hispanica Wagner from the Stephanian of Spain(Wagner, 1985); however, it is distinguished by simplelateral veins, while in P. hispanica, they bifurcate oncein the middle and basal parts of the last order pinnae(Wagner, 1985, pl. I, fig. 3; venation is distinct in theright lower pinna of the last order).

A sporangium of Convexocarpus distichus, comb.nov. [specimen GIN, no. 3773(11)/286(91); see herePl. 7, figs. 1, 3; also figured by Naugolnykh (1995,text�figs. 5D, 5E; 1998c, text�figs. 20D, 20E)] hasyielded spores preserved in situ and very similar tospores from the sporangium of other specimen ofC. distichus, comb. nov. (Pl. 7, fig. 4) and previouslyfigured by the author (Naugolnykh, 1998c, pl. VII,fig. 1, pl. X, fig. 6). Very similar spores are characteris�tic of the marattialean fern genus Acitheca (Zodrowet al., 2006). Spores of this type in disperse conditionare assigned in to the genus Verrucosisporites Ibrahim(Ibrahim, 1933), if they have well�developed verrucaewhich cover most of the surface, or to the genus Punc�tatisporites Ibrahim (Ibrahim, 1933), if their extine isdotted. Spores of both types occur in situ in the spo�rangia of marattialean ferns (see Balme, 1995).

M a t e r i a l. Eight satisfactory and well�preservedfragments of fertile leaves and ten sterile leaves fromthe Krasnaya Glinka, Krutaya Katushka 1, KrutayaKatushka 2, Rakhmangulovo 1, Rakhmangulovo 2,and Chekarda 1 localities.

O c c u r r e n c e. Lower Permian, Kungurian Stageof the middle Fore�Urals.

Genus Corsinopteris Doweld, 2001, emend. nov.

Ty p e s p e c i e s. Corsinopteris saraepontana(Corsin) Doweld, Upper Carboniferous and LowerPermian of Western Europe.

D i a g n o s i s. Fertile pinnules having well�devel�oped synangiophores divided into two branches andarranged in two rows extending along midvein. Biseri�ate synangia consisting of sporangia fused by theirmargins. Synangia always positioned above point ofmain dichotomy of lateral veins.


Figs. 1–4. Morphology of fertile leaves of marattialean ferns from the Permian of the Fore�Urals: (1, 3) Convexocarpus distichus(Naugolnykh) Naugolnykh, comb. nov.: (1) specimen GIN, no. 4856/140, apical part of bipinnate frond; (3) specimen GIN,no. 3737/130, last order pinna; (2) Corsinopteris semilibera (Naugolnykh) Doweld, specimen GIN, no. 4846/132, fertile last orderpinnae with well�developed hydathods; (4) Corsinopteris petschorica (Fefilova) Doweld, specimen GIN, no. 4851/198, middlepart of the last order pinna. Localities: (1, 3) Middle Fore�Urals, Perm Region, Chekarda 1, Bed 10; Lower Permian, KungurianStage; (2) Kozhim River, Kozhim 5 locality; Middle Permian, Ufimian Stage; (4) Fore�Urals near the Pechora River, boreholeIK�657, 765.7 m of depth; Middle Permian, Kazanian Stage. Scale bars, 1 cm.



Plate 8

1 2

3 4

1402


NAUGOLNYKH

C o m p a r i s o n. Corsinopteris is similar to thegenus Sydneia, differing from it in the leaf laminawhich is only partially reduced in some species, forexample, Corsinopteris semilibera. It differs from thegenus Eoangiopteris and formal genus AcaulangiumMillay (Millay, 1977) with bilaterally symmetricalsynangia in the considerably greater number of spo�rangia in the synangium (18–20 and more versus sevenor eight in Eoangiopteris and Acaulangium). It differsfrom the genus Gemellitheca Wagner, Hill et El�Khayal(Wagner et al., 1985) in the successive fusion of all spo�rangia by their margins for at least three�fourths oftheir extent; it differs from the genus Qasimia Hill,Wagner et El�Khayal (Hill et al., 1985) in the fewersporangia per synangium, the synangia detached fromeach other to a greater extent, and in the well�devel�oped synangiophores, which are characteristic ofCorsinopteris.

R e a s o n f o r e m e n d a t i o n. Although thediagnosis of the genus Corsinopteris in the originaldescription (Doweld, 2001) is rather verbose, it con�tains a number of essential inaccuracies, the mostimportant of which is the statement that “Synangiabiseriate, attached laterally to the dichotomousnerves” (Doweld, 2001, p. 1098). From this formula�tion it is impossible to understand the arrangement ofsporangia in synangia. The text provided by Doweldimplies that the biserial synangia were attached on oneside to dichotomizing veins, apparently, throughouttheir extent (i.e., above and below the point of dichot�omy, because other arrangement is not provided in theinitial diagnosis). In fact, the synangia of the threeAngaraland species (Corsinopteris dicranophora,C. semilibera, and C. petschorica (Fefilova) Doweld)referred by Doweld to the genus Corsinopteris sensuDoweld show a different structure. Their synangia arealways positioned above the point of the first (main)dichotomy of the lateral veins of fertile pinnule. Theexact structure of synangia of the type species C. sarae�pontana and their arrangement on the lateral veins arenot known; however, available data (Corsin, 1951)suggest that it was similar in structure to the synangiaof Angaraland species. At the same time, it is notinconceivable that Angaraland (Subangaraland)members of the genus Corsinopteris should be trans�ferred to a separate genus in the future.

Certain less important inaccuracies concern theincorrect or too generalized characteristics of otherfeatures of the genus included by Doweld in the diag�nosis of Corsinopteris. In this situation, I decided toreduce the diagnosis, retaining only improved charac�teristics of synangiophores, which distinguish thegenus Corsinopteris from other related genera ofmarattialean ferns.

R e m a r k s. The Upper Carboniferous beds ofNova Scotia (Canada) have yielded a peculiar plantfossil (Psenicka et al., 2003); in the original descrip�tion, it was treated as a fertile frond fragment of a

marattialean fern and assigned to a new species andgenus, Sydneia manleyi Psenicka and Bek, Zodrow,Cleal et Hemsley. This plant is characterized by almostcomplete reduction of the leaf lamina of fertile pin�nules and aggregation of sporangia in bilateral synan�gia, which produced monolete spores. A similar struc�ture is also observed in some other Carboniferous andPermian marattialean ferns, in particular, Corsinopt�eris semilibera, with partial reduction of the leaf lam�ina on fertile pinnules, which was described from theUfimian beds of the Fore�Urals near the PechoraRiver (Kozhim section: Grunt et al., 1998; a detaileddescription of this species see below); however, manycharacters of the same set are characteristic of malefructifications of lyginopterid pteridosperms (Cros�sotheca Zeiller; see review in Millay and Taylor, 1979;Stubblefield, 1984).

S p e c i e s c o m p o s i t i o n. Four species: Corsi�nopteris saraepontana, C. dicranophora, C. semilibera,and C. petschorica.

O c c u r r e n c e. Upper Carboniferous and Per�mian of the Northern Hemisphere.

Corsinopteris dicranophora (Naugolnykh, 1998) Doweld, 2001

Plate 9, figs. 1, 2, and 4

Orthotheca sp. (sp. nov.): Naugolnykh, 1995, text�fig. 6E.Orthotheca dicranophora Naugolnykh: Naugolnykh, 1998c,

pp. 57–58, text�fig. 19, pl. VII, fig. 8.Corsinopteris dicranophora (Naugolnykh) Doweld: Doweld,

2001, p. 1098.

H o l o t y p e. GIN, no. 3773(11)/291(91); Chekar�da 1 locality, Bed 10 (Pl. 9, figs. 1, 2, 4).

D i a g n o s i s. Frond at least tripinnate, medium�sized. Rachis with fine longitudinal ribs. Elongateddecurrent pinnules alternating, with pecopteroidbases. Pinnules with entire margins and round apex.Rachis of last order pinnae with narrow limb. Fertilepinnules having 10–20 synangia positioned on oncedichotomizing lateral veins. Midvein decurrent, pass�ing into axial part of pinnule base or occasionally posi�tioned closer to anadromic margin of pinnule. Lateralveins alternating, dichotomizing once at base.

D e s c r i p t i o n. I published a detailed descrip�tion of this species in a previous study (Naugolnykh,1998c, pp. 57–58). Below a modified and reducedvariant is provided.

The total length of fertile fronds is several tens cen�timeters. The maximum observable width of the lastorder fertile pinna is 25 mm in an incomplete 56�mm�long pinna (with the torn off apical part). The pinnalrachis is slightly curved, has on the adaxial surface alongitudinal groove, which corresponds to a low rib onthe abaxial surface. The maximum width of the pinnalrachis to the last order is 1.4 mm.

The pinnules are pecopteroid, entire, elongated,decurrent, attached to the pinnal rachis at an angle of65°. The apex is rounded. The maximum pinnulelength is 14 mm at the width of 6 mm. The pinnal



Plate 9

2

1

3

4 5 6


Figs. 1–6. Morphology of (1, 2, 4–6) fertile and (3) sterile leaves of marattialean ferns from the Permian of the Fore�Urals:(1, 2, 4) Corsinopteris dicranophora (Naugolnykh) Doweld, emend. nov., holotype GIN, no. 3773(11)/291(91); (3) Pecopteris hel�enaeana Zalessky, specimen GIN, no. 4856/191a, middle part of the last order pinna (counterpart of specimen GIN,no. 4856/191); (5, 6) Corsinopteris petschorica (Fefilova) Doweld, imprint and counterpart of a fertile pinnule: (5) specimen GIN,no. 4851/201, lateral light from the right; (6) specimen GIN, no. 4851/202, lateral light from the left. Middle Fore�Urals: (1–4),Perm Region, Chekarda 1 locality, Bed 10; Lower Permian, Kungurian Stage; (5, 6) Pechora Coal Basin, borehole KhK�1058,433.6 m of depth; Middle Permian, Kazanian Stage. Scale bars, 1 cm.

1404


NAUGOLNYKH

rachis of the last order is winged due to decurrent basi�scopic parts of the pinnule. The limb is 1.5 mm wide.The pinnule midvein is well�developed, reaches themargin of the pinnule apex. Lateral veins diverge fromthe midvein at an angle of 50° and dichotomize oncein the lower part (Pl. 9, fig. 4). The branches formed ofthe lateral vein have biserial synangia above the pointof dichotomy. The synangia are formed of sporangiafused by their margins for on average two�thirds of thelength. The maximum length of synangia is 2 mm atthe width of 0.9 mm. The sporangia are 0.4 mm longand 0.15–0.20 mm wide.


Corsinopteris semilibera (Naugolnykh, 1998) Doweld, 2001, emend. nov.

Plate 8, fig. 2.

Orthotheca semilibera Naugolnykh: Grunt et al., 1998, pp. 253,255, text�fig. 48, fig. 9; text�fig. 70 and 71, pl. XLIII, fig. 2.

Corsinopteris semilibera (Naugolnykh) Doweld: Doweld, 2001,p. 1098. Naugolnykh, 2009, p. 18, text�fig. 1c, pl. I, fig. 2.

H o l o t y p e. GIN, no. 4856/127; Kozhim 5 local�ity; figured by Grunt et al. (1998, text�figs. 70 (1) and71 (3, 6).

D i a g n o s i s. Fronds at least tripinnate. Sterilepinnules subtriangular, fused at base, and having twoor three pairs of lateral veins. Midvein well developed,with two or three pairs of lateral veins. Fertile pinnulesrather diverse, ranging from those resembling sterilepinnules, but with synangia, to pinnules with stronglyreduced leaf lamina. Synangia biserial, positioned dis�tally along lateral veins, distal to main dichotomy oflateral veins. Lateral veins dichotomizing once ortwice and possessing hydathods. Sporangia rangingfrom fusiform to ovoid.

D e s c r i p t i o n (Fig. 9). The collection includesboth fertile leaves and leaves combining fertile andsterile segments.

The fronds are at least tripinnate. The last orderpinnae are in alternating positions, attached to thefrond rachis at an angle of 50°–80°. The frond rachisis well developed, about 2 mm wide, usually smooth,but sometimes with thin longitudinal folds and ribs.Sterile pinnae of the last order are medium�sized(about 30 mm long) at the maximum width of 10 mm.One pinna has on average about seven pairs of pinnules(segments of the last order). Pinnules are subtriangu�lar, attached to the rachis of the last order pinna at anangle of 40°–60°, rarely, at 80°. Margins of sterile pin�nules slightly crenate. The pinnule apices are usuallyrounded. Pinnules are coalescently fused from thebase to half length or sometimes even more to form aunicoherent leaf lamina. The pinnule midvein has afusiform hydathod in the distal part. Each pinnulecontains one or two pairs of lateral veins. Some lateralveins, particularly those of pinnules positioned in theproximal part to the last order pinna, also havehydathods. In the apical part of the last order pinna,the sequence of the formation of pinnules may be

irregular. This results in the development of a relativelylarge terminal pinnule, with unicoherent venation.The leaf lamina is very thin and nonresistent to macer�ation. Sterile and fertile pinnae are usually present inthe same fronds. Some pinnae are partially fertile, verysimilar in morphology to completely sterile pinnae.Modified, completely fertile pinnae with a partiallyreduced leaf lamina are less frequent. Sporangia arearranged in the synangium, from six to eight in eachsynangium. In the typical case, the midvein divergesfrom the pinnal rachis, passing in the fertile pinnuleand, in turn, gives rise to the lateral veins, whichdichotomize once. Those branches of lateral veinswhich were formed after dichotomy have synangia.The synangia are two rows of elongated sporangiapositioned almost perpendicular to the axis bearingthem. The sporangia are fused by the margins foralmost the entire extent. Lateral veins, which bearsynangia, sometimes have hydathod in the distal part.The basal part of fertile pinnules is more complex instructure, containing lateral veins which sometimesdichotomize twice rather than once; the seconddichotomy is only formed in the anterior anadromicbranch of the preceding dichotomy. Rarely, an addi�tional branch is formed in the catadromic instead ofanadromic branch of the second dichotomy and givesrise to an additional middle vein, which also hassynangia on each of these branches.

The sporangia are elongated oval, elliptic. In somecases, they slightly expand apically, becominginversely obovate in outline. The sporangia are onaverage 0.8–0.9 mm long and 0.4 mm wide.

C o m p a r i s o n. Corsinopteris semilibera is inter�mediate between the Kungurian species C. dicrano�phora from the Middle Fore�Urals (Naugolnykh,1998c) and C. petschorica, which is characteristicmostly of the Kazanian of the Fore�Urals near thePechora River (Fefilova, 1973). It differs fromC. dicranophora in the wider synangia (on averageabout 1.5 mm wide versus 0.9 mm) and in the complexpattern of the basal parts of fertile pinnules with thelateral veins dichotomizing twice or thrice. In addi�tion, fertile fronds with a reduced leaf lamina have notbeen recorded in C. dicranophora. C. semilibera differsfrom C. petschorica in the absence of pairwise rap�prochement of pinnules and lateral veins and in thereduced leaf lamina in the fertile leaf part.

M a t e r i a l. Eleven fragmentary preserved fertileand sterile pinnae.

Corsinopteris petschorica (Fefilova, 1973) Doweld, 2001, emend. nov.

Plate 8, fig. 4; Plate 9, figs. 5 and 6; Plate 10, figs. 1–3

Orthotheca petschorica Fefilova: Fefilova, 1973, pp. 36–40,text�fig. 6, pl. VI, figs. 1–3; pl. VII, figs. 1–3; pl. VIII, figs. 1–5;Pukhonto and Fefilova, 1983, pl. V, figs. 1 and 6; Pukhonto, 1998,pl. 3, fig. 3, pl. 39, fig. 4.

Corsinopteris petschorica (Fefilova) Doweld: Doweld, 2001,p. 1098.



H o l o t y p e. GIN, no. 3048/275; southwesternPai Khoi, Nadota River; figured by Fefilova (1973,pl. VI, fig. 1).

E m e n d e d d i a g n o s i s. Fronds tripinnate orquadripinnate. Sterile pinnules pecopteroid, some�times lanceolate. Midvein robust, with four to ninecouples of once dichotomizing lateral veins. Apicalparts of fertile pinnules sometimes partly sterile. Pin�nule apex round. Lateral veins once dichotomizingclose to their attachment to midvein. Synangia biseri�ate, positioned distally along lateral veins just abovepoint of main dichotomy of lateral veins. Sporangiaovoid.

D e s c r i p t i o n. I examined two representativespecimens of fertile leaves referred to this species. Abetter preserved specimen (Pl. 10, fig. 1) is the apical

part of a large fertile frond. All pinnules observed inthis specimen are fertile. Seven pinnule pairs are pre�served, so that pinnules on the right side of the frondare preserved completely. Pinnules vary in shape fromrounded isometric in apical part to elongated lan�ceolate in more proximally positioned pinnules. Pin�nules gradually decrease in length distally. The maxi�mum observable length of pinnules is 40 mm at 9 mmof width. The angle of attachment of pinnules to thepinnal rachis of the last order varies from 45° (in apicalpinnules) to 55°–60° (in proximal pinnules). Pinnulesare decurrent, coalescently fused at the base. The mid�vein of pinnules is slightly curved at the base, distinctlydecurrent. Lateral veins dichotomize at the base, justafter deviation from the pinnule midvein. Biserialsynangia are positioned above the point of dichotomyon the lateral veins. In other words, lateral veins are

(a)

(b)

(c)

(d) (e)

SA

SP

SPH

SN

Fig. 9. Morphology of the marattialean fern Corsinopteris semilibera (Naugolnykh) Doweld, emend. nov.: (a, c, d) structure ofsynangia: (a) holotype GIN, no. 4846/127; (c) syntype GIN, no. 4846/130; (d) specimen GIN, no. 4846/110; (b) generalizedscheme of synangium structure; and (e) reconstruction of synangial aggregations. Designations: (SA) synangial apex,(SN) synangium, (SP) sporangia, (SPH) synangiophore. Fore�Urals near the Pechora River, Kozhim 5 locality. Middle Permian,Ufimian Stage. Scale bar, 1 mm.

1406


NAUGOLNYKH

Plate 10

1

4 5

3

6

2



E x p l a n a t i o n o f P l a t e 1 0

Figs. 1–6. Morphology of (1–3) fertile and (4–6) sterile leaves of marattialean ferns from the Permian of the Fore�Urals: (1–3)Corsinopteris petschorica (Fefilova) Doweld: (1) specimen GIN, no. 4851/177, apical part of a fertile frond; (2, 3) last order pinnawith sterile pinnules in the upper part and fertile pinnules in the lower part: (2) specimen GIN, no. 4851/178 and (3) specimenGIN, no. 4851/179; (4) Pecopteris cf. filatovae Sixtel, specimen GIN, no. 3773(11)/292(91), sterile last order pinna with well�developed hydathods at the ends of lateral pinnule veins; (5) Pecopteris helenaeana Zalessky, specimen GIN, no. 3773(11)/13(89),middle part of a last order pinna; and (6) Pecopteris uralica Zalessky, specimen GIN, no. 3773/375, sterile last order pinna. Local�ities: (4–6) Middle Fore�Urals, Perm Region, Chekarda 1 locality; Lower Permian, Kungurian Stage: (5) Bed 7 and (4, 6) Bed10; (1–3) Pechora Coal Basin; Middle Permian, Kazanian Stage: (1) borehole IK�426, 533.25 of depth; (2, 3) borehole IK�631,344.5 m of depth. Scale bars, 1 cm.

(possibly transformed into) dichotomizing synangio�phores, which have biserial synangia on the branches.Lateral veins are attached to the midvein at 50°. Pin�nules have from four (in apical pinnules) to nine(in basal pinnules) pairs of lateral veins. The synangiaconsist of seven or eight pairs of elongated clavate spo�rangia, which are fused by the margins along almostthe entire length. The synangia are biserial. Both rowsof sporangia are positioned symmetrically relative toeach other along the branches of lateral veins above thepoint of dichotomy of the lateral vein (synangio�phore).

M a t e r i a l. Apical part of a fertile frond and par�tially fertile pinna of the last order.

O c c u r r e n c e. Middle Permian, Kazanian Stageof the Fore�Urals near the Pechora River.

C L A S S O S M U N D O P S I D A TA K H T A D J A N , 1 9 8 6

Order Osmundales Bromhead, 1840

Family Osmundaceae Bertchtold et Presl, 1820

Genus Todites Seward, 1900

Ty p e s p e c i e s. Todites williamsonii (Brongn�iart) Seward, Jurassic of the Northern Hemisphere.

D i a g n o s i s. Tripinnate fertile fronds having pin�nules abaxially covered by free stalked sporangia. Spo�rangia with cluster of thick�wall cells above slitlike sto�mium. Spores of Osmundacidites Couper and Todis�porites Couper types.

C o m p a r i s o n. Todites is most similar to thegenus Osmundopsis Harris and differs from it in themorphological similarity between fertile and sterileleaves and in the co�occurrence of fertile and sterilepinnae in the same frond.

R e m a r k s. The original diagnosis of the genusTodites is supplemented by me with information onspores of Osmundacidites, which were recorded in situin the sporangia of Todites lobulatus (Naugolnykh,2002). Spores of this type were also marked in thegenus Todites by other authors (see review in Balme,1995), although this was not included in the genericdiagnosis (see, e.g., Harris, 1961; Vakhrameev et al.,1963).

Protoleptosporangiate ferns in localities of theeastern part of the Russian Platform and Fore�Urals

are represented by fertile leaves of the genus Todites,sterile leaves of Cladophlebis, and petrified trunks ofthe fern family Osmundaceae.

The presence of protoleptosporangiate ferns in thePermian beds of the Fore�Urals and eastern RussianPlatform has initially shown by the records of mineral�ized trunks assigned to several species: one specimenfrom cupriferous sandstones in the vicinity of Permwas incorrectly determined as “Lepidodendron”(Kutorga, 1842, pl. II, fig. 2), Tubicaulis rhomboidalisKutorga (Kutorga, 1844), Sphallopteris schlechtendaliiEichwald, Chelepteris gracilis Eichwald, Bathypterisrhomboidalis Eichwald, and Anomorhoea fischeriiEichwald (Eichwald, 1854). Zalessky (1927) providedadditional well�illustrated material of mineralizedstems of osmundaceans from the same beds. Subse�quently, Fefilova (1973) described in open nomencla�ture fragments of fertile pinnae from the Ufimian andKazanian of the Fore�Urals near the Pechora Riverand referred them to the genus Todites.

Based on the material from the Inta Formation(Lower Ufimian) on the Kozhim River, I described anew species, Todites lobulatus (Naugolnykh, 2002),which is distinguished from the majority of congenersby the sphenopteroid rather than pecopteroid fertilepinnules. Further studies have shown that the Middleand Upper Permian beds of the Fore�Urals near thePechora River enclose at least three other types ofosmundacean ferns: (1) with small entire pecopteroidfertile pinnules; (2) fertile and sterile pecopteroid pin�nules with lobate margins and compressed bases com�bined in one pinna; and (3) with large subtriangularsterile pinnules coalescently fused by their bases. Thefirst type is tentatively determined as Todites sp.(Pl. 14, fig. 4; Pl. 15, fig. 4); the second type isdescribed below as the new species Todites emarginatusNaugolnykh, sp. nov.; and the third type is the newspecies T. coronatus Naugolnykh, sp. nov.

Anderson and Anderson (2008) proposed a newclassification system for fertile ferns of the genusTodites, in which some species were attributed to thegenera Rooitodites Anderson et Anderson, BirtoditesAnderson et Anderson, and Elantodites Anderson etAnderson. However, it seems that the characters usedfor the establishment of Rooitodites, Birtodites, andElantodites are sometimes observed within one mor�

1408


NAUGOLNYKH

phologically diverse species (Anderson and Anderson,2008, pl. 20, figs. 1–5, pl. 21, figs. 1–5).

A natural question arising in the study of fern leavesof pecopteroid morphology, which are widespread inthe Permian beds of Western Angaraland (Pl. 11) anddisplay a wide variation range, is connected with thepossibility of differentiation between sterile leaves ofosmundacean and marattialean ferns. In my opinion,sterile leaves of Permian osmundacean show a morecomplex venation (see, e.g., Fig. 10j) and, usually,wavy or lobate margin; however, certain conclusionsconcerning the taxonomic position of particular leavesof pecopteroid morphology require the study of asso�ciated fertile organs.

S p e c i e s c o m p o s i t i o n. Fifty species.O c c u r r e n c e. Permian–Cretaceous, every�

where.

Todites lobulatus Naugolnykh, 2002

Plate 12, figs. 1–7; Plate 13, figs. 1–10

Todites lobulatus Naugolnykh: Naugolnykh, 2002, pp. 470–473, text�figs. 3–7; Naugolnykh, 2009, p. 19, text�figs. 2a–2d and2g; Pukhonto and Naugolnykh, 2009, text�fig. 2.

H o l o t y p e. GIN, no. 4846/102, Fore�Urals nearthe Pechora River, Kozhim 5 locality (Pl. 12; figs. 1–5;Figs. 10a–10d, 10k).

D i a g n o s i s. Ferns with tripinnate fronds. Pin�nules deeply dissected, with well�developed lobes.Pinnule apex round. Sporangia free, located on abax�ial leaf surface. Sporangia small (0.5 mm in diameter),round or ovoid, with group of thick�walled cells onsporangial tip or side. Spores round, with trilete tetradscar; scar radii simple, straight; sporoderm with gran�ulate or scabrate ornamentation. Spores ranging insize from 20 µm to 45 µm.

D e s c r i p t i o n (Figs. 10a–10c, 10k). The frondis at least tripinnate. The last order pinnal rachises are1.5 mm wide. The pinnules are distinctly sphe�nopteroid, with a deeply dissected margin and well�developed laminae. The pinnule leaf lamina is dis�sected for on average one�third of the pinnule width;in the basal part of the pinnule, it is sometimes evengreater. Well�developed pinnules vary in size from 13.3to 15.8 mm, with the maximum pinnule width of8 mm. The pinnule apex is round. The sporangia arefree, numerous, attached to the abaxial surface of fer�tile pinnules.

The number of separate sporangia per well�devel�oped fertile pinnule is sometimes greater than 200.The sporangia are spherical or ellipsoidal, have a shortsporangiophore, lateral slitlike stomium (Naugol�nykh, 2002, text�fig. 5) open in a group of thick�walledcells on the sporangium apex. The sporangia are onaverage 0.5 mm in diameter. The sporangia enclosedspores of the Osmundacidites type, ranging in diameterfrom 20 to 45 µm, with small spines and granules onthe distal surface.

C o m p a r i s o n. Todites lobulatus is similar to thespecies Todites emarginatus sp. nov., differing from it inthe fertile pinnules of sphenopteroid appearance anddeeply dissected into lobes, the simpler venation, andin the absence of simple entire�margined pinnules.


Todites emarginatus Naugolnykh, sp. nov.

Plate 14, figs. 1–3; Plate 15, fig. 5

E t y m o l o g y. From the Latin emarginatus (emar�ginate, sinuate), based on the dissected leaf lamina ofpinnules into round lobes.

H o l o t y p e. GIN, no. 4851/173, partially fertilefrond fragment, with three variously preserved pinnaeof the last order; Pechora Coal Basin, Pechora River,outcrop no. 22 (after Zalessky, 1934a); Upper Per�mian, Severodvinian Stage; designated here (Pl. 14,figs. 1–3; Pl. 15, fig. 5; Figs. 10j 10l, 11, 12).

D i a g n o s i s. Fronds at least bipinnate. Sterilepinnules ranging from pecopteroid to sphenopteroid,with lobate margins and round apices. Pinnule mid�veins slightly undulating. Lateral veins dichotomizingup to two times. Fertile pinnules possessing isolatedfree sporangia, sometimes arranged in compact clus�ters. Thick�walled cells specialized for opening sto�mium located above stomium and forming short row.Each fertile pinnule with 80–100 sporangia.

D e s c r i p t i o n (Figs. 10j, 10l, 11, 12). The frondis at least bipinnate. The rachis of the penultimateorder pinna is thick, massive, 7–8 mm wide, coveredwith thin, but distinct longitudinal ribs and folds.Rachises of the last order pinnae are attached to therachis to the penultimate order pinna at an angle of60°–67°. Rachises of the last order pinnae are straightor slightly curved towards the apex of the penultimateorder pinna. The axial part of rachises of the last orderpinnae contains a shallow, but distinct longitudinalgroove semiround in cross section. The rachis surfaceis even, trichome structures have not been recorded.Alternating pinnules are attached to the rachis of thelast order pinnae at an angle of 65°. Sterile pinnulesrange from pecopteroid (pecopteroid pinnules arelocated in the middle and apical parts of the last orderpinna) to sphenopteroid (sphenopteroid pinnules arelocated in the basis of the last order pinnae). Pecop�teroid pinnules are linguiform in outline, with a roundor blunted apex and distinctly lobate or, less often,crenate margin. Venation is pinnate. The midveinpasses strictly along the axial part of pinnules, slightlycurves. The ends of the midvein and lateral veins donot reach the pinnule margin. Four or five pairs of lat�eral veins diverge from the midvein at an angle of 45°.Each lateral vein dichotomizes close to the base. Theanterior (acroscopic) branch dichotomizes one moretime. In well�developed pinnules, the posterior (basis�copic) branch of lateral veins also sometimes dichoto�mizes. In extremely rare cases, the secondary anteriorbranch of the acroscopic vein dichotomizes one more



time. The pinnule margin changes from wavy (in api�cal pinnules) to crenate (in pinnules of the middle partof the last order pinna) and, then, to lobate (in basalpinnules). In the middle and apical parts of the lastorder pinna, neighboring pinnules are connected by anarrow limb, which forms a thin wing of the rachis. Inthe basal part of the rachis of the last order pinna, thelimb is absent.

Fertile pinnules are similar in size and outline tosterile pinnules, but have many spherical or ellipsoidalsporangia on the abaxial surface. Each completelydeveloped fertile pinnule can bear from 80 to 100 spo�rangia. Sporangia are positioned separately, but some�times form dense aggregations of 20–25 sporangiaeach. These aggregations are sometimes displaced orconfined to lobes of fertile pinnule. In the axial part of

(a)

(b)

(c)

(d)

(e) (f) (g) (h)

(i)

(k)

(l)

(j)

Fig. 10. Morphology of osmundacean ferns from the Permian of the Fore�Urals: (a, b) spore structure preserved in situ, (c) spo�rangia, (d) fertile leaf with sphenopteroid pinnules, (e–h) scheme showing increasing specialization of sporangial cells, and(i, j, k, l) morphological diversity of fertile leaves; (a–d, k) Todites lobulatus Naugolnykh, holotype GIN, no. 4846/102: (a) sporesof Osmundacidites sp. preserved in sporangia in situ, (b) sporoderm ornamentation, (c) structure of individual sporangium,(d) middle part of a fertile leaf of the last order, (e–h) morphogenetic trend in the change of sporangial structure in (e–g) euspo�rangiate and (h) leptosporangiate ferns (after Pant and Khare, 1974, modified): (e) Stauropteris oldhamia Binney, Middle Car�boniferous of Western Europe; (f) Botryopteris globosa Darrah, Middle Carboniferous of the United States; (g) Senftenbergia pen�naeformis (Brongn.) Stur, Middle Carboniferous of Western Europe; (h) Osmunda sp.; (i–j, k) Todites emarginatus Naugolnykh,sp. nov.: (i) scale bar for fig. j; (j) lateral last order pinna combining sterile and fertile pinnules; (k) structure of individual lobe ofa fertile pinnule: functionally upper (adaxial) side on the left) and functionally lower (abaxial) side on the right; (l) sterile andfertile pinnules of the same last order pinna; Localities: (a–d, k) Kozhim 5, Middle Permian, Ufimian Stage; (j, l) outcrop ofT.A. Dobrolyubova no. 22; Kazanian Stage, Pechora River (see Zalessky, 1934, p. 288). Scale bars: (a) 50 µm; (b) 5 µm;(c, e⎯h) 0.5 mm; (d, k, l) 5 mm; and (i, j) 10 mm.

1410


NAUGOLNYKH

Plate 11

1 2 3

4

5

6 7



pinnules, under the pinnule midvein, sporangia areusually fewer or absent. Perfectly preserved sporangiadisplay a group of specialized apical cells with thick�ened walls, which were responsible for disclosing thesporangium after maturation. The cell size is on aver�age 50 × 70 µm; some cells are larger. This group ofcells forms a curving row above the stomium, which

projects significantly above the sporangium and makesit somewhat asymmetrical.

An interesting feature of the species Todites emar�ginatus sp. nov. is the combination of sterile and fertilepinnules within one pinna of the last order, with thefertile pinnules usually located in the basal part of thelast order pinna (Pl. 14, figs. 1, 3; Fig. 10j, 10i),


Figs. 1–7. Morphology of various ferns from the Permian of Angaraland assigned to the conditional informal “parataxonomical”groups of (1–4, 6, 7) pecopterids and (5) sphenopterids: (1) Pecopteris anthriscifolia (Goeppert) Zalessky, specimen GIN,no. 4851/206, well�developed last order pinna; (2) Pecopteris aff. anthriscifolia (Goeppert) Zalessky, specimen GIN,no. 4851/180, penultimate order pinna with four partially preserved last order pinnae; (3) Prynadaeopteris aff. alifera Fefilova,specimen GIN, no. 4851/181, middle part of a penultimate order pinna, which combines fertile (top) and sterile (bottom) lastorder pinnae; (4, 6) Pecopteris sp., specimen GIN, no. 4851/182, middle part of a tripinnate frond photographed in (4) slantingand (6) direct light at different angles; (5) Tumidopteris clavata Naugolnykh, gen. et sp. nov., specimen GIN, no. 4851/183, sterilepinna; and (7) Pecopteris anthriscifolia (Goeppert) Zalessky, specimen GIN, no. 4851/207, last order pinna with lobate pinnules.Localities: Pechora Coal Basin; Middle Permian, Kazanian Stage: (1) borehole IK�612, 777 m of depth; (2) borehole IK�627,164.5 m of depth; (3) borehole IK�636, 380.1 m of depth; (4, 6) borehole VK�22, 746.5 m of depth; (5) borehole IK�686, 349.10 mof depth; and (7); borehole IK�627, 163.5 m of depth. Scale bars, 1 cm.

(а) (b)

(c) (d)

Fig. 11. Morphology of fertile pinnules of Todites emarginatus Naugolnykh, sp. nov., holotype GIN, no. 4851/173: (a, c) detaileddrawing of sporangial positions and (b, d) photographs with linear retouch used for producing drawing. Locality: Pechora CoalBasin, Pechora River, outcrop of T.A. Dobrolyubova no. 22 (see Zalessky, 1934, p. 288); Middle Permian, Kazanian Stage. Scalebar, 1 mm.

1412


NAUGOLNYKH

although they sometimes occur in its middle part(Pl. 15, fig. 5).

C o m p a r i s o n. Two more species of the genusTodites, T. lobulatus (see above) and T. coronatus sp.nov. (see below), are described from the Permian of theFore�Urals near the Pechora River; in addition,remains of osmundacean ferns determined in opennomenclature as Todites sp. have been described andfigured from the same beds (Fefilova, 1973; Pukhontoand Fefilova, 1983, pl. V, fig. 2; pl. VII, fig. 6). The newspecies differs from T. lobulatus in the considerablyweaker dissection of the leaf lamina into lobes, fromT. lobulatus and T. coronatus sp. nov. in the combina�tion of sphenopteroid and pecopteroid pinnules in thesame last order pinna, the more complex venation,and in the round or blunted pinnule apices. The spec�imens of Todites sp. figured by Fefilova (1973) andPukhonto and Fefilova (1983) differ from bothT. emarginatus sp. nov. and T. lobulatus in the entirepinnules with considerably fewer sporangia.

R e m a r k s. I found the specimen designated asthe holotype of T. emarginatus sp. nov., among plant

remains of Zalessky’s collection stored in the GIN.Later, a part of this collection was transferred to theVernadsky State Geological Museum (Moscow). Thespecimen was not labeled, nor accompanied withinformative material; however, the specimen itself hasan inscription “Pechora 22(7).” Published works ofZalessky do not contain a figure or description of thisspecimen; however, in the paper published in 1934,locality no. 22 on the Pechora River is mentioned;thus, the specimen in question probably comes fromthis locality, with individual number �7�. Since thespecimen lacks a unified collection number of GIN, itreceived a new number (GIN, no. 4851/173) andplaced in the collection with other type specimensconsidered in the present study.


Todites coronatus Naugolnykh, sp. nov.


E t y m o l o g y. From the Latin coronatus(crowned), based on the presence of well�developed

(a)

(b)

(c) (d)

Fig. 12. Pinnule venation in Todites emarginatus Naugolnykh, sp. nov., holotype GIN, no. 4851/173 (a–d); (d) relative positionsof sterile and fertile pinnules. Conducting tissues are filled with black; dotted lines are presumable commissures between lobes.Locality: Pechora Coal Basin, Pechora River, outcrop of T.A. Dobrolyubova no. 22 (see Zalessky, 1934, p. 288; Middle Permian,Kazanian Stage). Scale bar, 1 mm.



1

Plate 12

2 3

4

5

6 7


Figs. 1–7. Spore structure and macromorphology of the osmundacean fern Todites lobulatus Naugolnykh, holotype GIN,no. 4846/102: (1–4) spores of Osmundacidites sp., preserved in sporangia in situ; (5) middle part of a fertile last order pinna, smallround sporangia are particularly distinct on the pinnule located on the upper left part of the pinna; (6, 7) spores of Osmundaciditessp. extracted from matrix just near the holotype of Todites lobulatus. Middle Permian, Kozhim 5 locality; Ufimian Stage. Scalebars: (1–4, 7) 50 µm, (6) 40 µm, and (5) 5 mm.

1414


NAUGOLNYKH

1

Plate 13

2 3

4 5 6

7 8

9 10


Figs. 1–10. Spores of Osmundacidites sp.: (1–5, 7–10) extracted from matrix and (6) a spore preserved in situ in a sporangium ofthe osmundacean fern Todites lobulatus Naugolnykh, holotype GIN, no. 4846/102: (1, 2) spores with weak granulate sporodermornamentation, proximal view; (3–5) spores with distinct granulate ornamentation and small spines on sporoderm: (3) distortedspore and (4, 5) distal view; (6) poorly developed spore preserved in situ in sporangium; (7–10) sporoderm ornamentation pat�tern; distinct granules and spines particularly well developed on the distal spore side. Pechora Coal Basin, Kozhim 5 locality;Middle Permian, Ufimian Stage. Scale bars: (1–3, 5) 30 µm, (4) 20 µm, (6) 50 µm, and (7–10) 10 µm.



3

Plate 14

4

2

1

E x p l a n a t i o n o f P l a t e 1 4Figs. 1–4. Macromorphological leaf structure of osmundacean ferns: (1–3) Todites emarginatus Naugolnykh, sp. nov., holotype GIN,no. 4851/173 and (4) Todites sp., specimen GIN, no. 4851/174: (1) sterile (upper) and fertile (lower) pinnules positioned on a last order pinna;(2) sterile (on the left) and fertile (on the right) pinnules located on two neighboring last order pinnae; (3) last order pinna with three fertilepinnules located in the basal part of catadromic (basiscopic) pinnule row (at the bottom right on the figure); (4) Todites sp., middle part of afertile penultimate order pinna with small pinnules subtriangular in outline. Localities: Pechora Coal Basin, Pechora River; Middle Permian,Kazanian Stage: (1–3) outcrop no. 22, after Zalessky (1934); (4) borehole VK�19, 650 m of depth. Scale bars: (1, 2) 5 mm and (3, 4) 10 mm.

1416


NAUGOLNYKH

Fig. 13. Frond morphology in Todites coronatus Naugolnykh, sp. nov., holotype GIN, no. 4851/203a. Locality: Pechora CoalBasin, borehole IK 657, 532.0 m of depth; Middle Permian, Kazanian Stage. Scale bar, 1 cm.

specialized cells with thickened walls on the spo�rangium apex.

H o l o t y p e. GIN, no. 4851/203a, middle part ofa bipinnate frond (middle part of a penultimate orderpinna), with three variously preserved fertile and par�tially fertile pinnae of the last order; Pechora CoalBasin, borehole IK 657, 532.0 m of depth; Middle

Permian, Kazanian Stage; designated here (Pl. 16;Pl. 17, figs. 1–4; Figs. 13, 14).

D i a g n o s i s. Fern with fertile or partly fertile lastorder pinnae with pecopteroid pinnules. Frond rachisthick, robust, having central adaxial furrow. Sterilepinnules pecopteroid, connected by their bases andforming limb (wing), surrounding rachis of last order


Figs. 1–7. Macromorphological and anatomical structure of trunks and leaves of osmundacean ferns from the Permian of theFore�Urals: (1–3) imprint of a large trunk presumably belonging to an osmundacean fern possibly associated with fertile leavesof Todites sp. and sterile leaves Cladophlebis sp., specimen GIN, no. 3003/60; (4) Todites sp., specimen GIN, no. 4851/174, mid�dle part of a fertile penultimate order pinna with small pinnules subtriangular in outline, with frontal (flashing) light; (5) Toditesemarginatus Naugolnykh, sp. nov., holotype GIN, no. 4851/173, fertile pinna; (6) Bathypteris rhoboidea Eichwald (= ? Sphallop�teris schlechtendalii Eichwald), specimen TsNIGR Museum, no. 45/1543 (exposition of the Monographic Department), crosssection of a large trunk of an osmundacean fern; (7) trunk imprint of Thamnopteris sp., specimen GIN, no. 4851/204, many pet�iole bases of frond rachises located on trunk in dense spiral order. Localities: Pechora Coal Basin; Middle Permian, KazanianStage: (1–3) borehole K�228, 254.2 m of depth; (4) borehole VK�19, 650 m of depth; (5) Pechora River, outcrop no. 22;(6) former Orenburg Region (in 1854), Kamskovotkinskii copper mine; and (7) Pechora Coal Basin, Adz’va River, outcrop 4�40,Bed 3. Scale bars, 1 cm.



pinna. Pinnule midveins distinct, extending for three�fourths of pinnule extent and then dichotomizing onceor twice (in basal part of pinna), only once (in middlepart of pinna), or simple (in apical part of pinna). Pin�

nule apex round or blunted. Fertile pinnules pecop�teroid, but sometimes constricted at base. Each fertilepinnule with up to 50 sporangia positioned on pinnuleabaxial surface. Sporangia clavate or fusiform, consist�

1

Plate 15

3

2

4

5

6 7

1418


NAUGOLNYKH

ing of two cell types. Basal and middle parts of spo�rangium consisting of relatively small elongated cellswith thin walls. Apical part of sporangium consistingof larger cells with thick walls. Thick�walled apicalcells forming crownlike structure surrounding sto�mium of sporangium and responsible for opening sto�mium. Spores in situ of Osmundacidites type, rangingfrom round to subtriangular, up to 50 µm in diameter,with exine having small spines.

D e s c r i p t i o n (Figs. 13–15). Fronds of this fernspecies are very large, tripinnate. The main rachis ofthe tripinnate frond (observed in specimen GIN,no. 4851/203b: Pl. 17, fig. 6; Fig. 15) is 2 cm thick;moreover, this thickness does not correspond to themost proximal part of the main rachis, which wasprobably even thicker. Available rachis fragment onlyslightly decreases in thickness towards the frond apex,becoming less than 1 mm thinner over 50 mm of the

rachis length. Based on this, the total frond length wasprobably about 1 m.

In specimen GIN, no. 4851/203b (see Remarks),the base of the penultimate order pinna is preserved inattachment to the frond rachis (Fig. 15, on the right).The rachis of the penultimate order pinna is 5 mmwide and attached to the main rachis of the frond at anangle of 68°. The surface of the main rachis of thefrond and rachis of the penultimate order pinna is cov�ered with thin longitudinal rugosity. The rachis of thelast order pinna is attached to the rachis of the penul�timate order pinna at an angle of 63°. The greatestthickness of the rachis of the last order pinna of thisspecimen is 1 mm. Pinnules are attached to the rachisof the last order pinna at an angle from 60° to 65°. Theinclination of pinnules to the pinna rachis increasestowards the apex of the last order pinna. Pinnules onthe left side (according to the position in Fig. 15) of the

(a) (b)

(c) (d) (e)

(f) (g) (h)

Fig. 14. Sporangial structure in Todites coronatus Naugolnykh, sp. nov., holotype GIN, no. 4851/203a (a–h); (e) reconstructionof an individual sporangium. Locality: Pechora Coal Basin, borehole IK 657, 532.0 m of depth; Middle Permian, Kazanian Stage.Scale bar, 100 µm.



1

2

3

4

5

6

8 9 10

7

Plate 16

E x p l a n a t i o n o f P l a t e 1 6Figs. 1–10. Frond morphology in Todites coronatus Naugolnykh, holotype GIN, no. 4851/203a: (1) general structure of the mid�dle part of a partially fertile frond; (2–10) spore structure of Osmundacidites sp., preserved in situ in sporangia. Locality: PechoraCoal Basin, borehole IK 657, 532.0 m of depth; Middle Permian, Kazanian Stage. Scale bars: (1) 1 cm and (2–10) 20 µm.

1420


NAUGOLNYKH

last order pinna are positioned at a considerablysmaller angle (on average about 40°), although thisdistinction is probably accounted for by diageneticdeformation of the leaf lamina in the course of com�pression of deposits enclosing plant remains.

The pinnules are pecopteroid, coalescently fused atthe base, with the formation of a limb (wing border),which is identical to the limb of sterile pinnules (holo�

type, see below). Both anadromic (acroscopic) andcatadromic (basiscopic) margins of pinnules have rel�atively small, but distinct lobes. The pinnule midveinsare well�developed, about 0.4 mm thick, pass for two�thirds of the pinnule length and, then, dichotomize.The acroscopic branch produced by dichotomy of themidvein sometimes dichotomizes one more time. Themidvein has two or three pairs of lateral veins, which

Fig. 15. Frond morphology in Todites coronatus Naugolnykh, sp. nov., syntype GIN, no. 4851/203b. Locality: Pechora CoalBasin, borehole IK 657, 532.0 m of depth; Middle Permian, Kazanian Stage. Scale bar, 1 cm.


Figs. 1–6. Frond morphology in Todites coronatus Naugolnykh: (1–4) holotype GIN, no. 4851/203a: (1) arrangement of spo�rangia on apical pinnules of a fertile pinna, pinnules are coalescently fused at the base, (2) apical part of sporangium, with well�pronounced slitlike stomium, (3) individual sporangium, lateral view, (4) apical part of a fertile last order pinna, fine structuralfeatures of which are shown in (1); (5, 6) syntype GIN, no. 4851/203b: (5) sterile last order pinna with pinnules coalescently fusedat the base; (6) part of a large tripinnate frond with a thick main rachis. Locality: Pechora Coal Basin, borehole IK 657, 532.0 mof depth; Middle Upper Permian, Kazanian Stage. Scale bars: (1, 4) 2 mm, (2, 3) 100 µm, and (5, 6) 1 cm.



1

Plate 17

3 4

5 6

2

1422


NAUGOLNYKH

are also well developed and, at the base, as thick as themidvein. Lateral veins dichotomize once. In one case,the second dichotomy of acroscopic branches of thelateral vein is observed.

There is a fragment of the middle part of the penul�timate order pinna, which is treated as either the mid�dle part of a bipinnate frond or the middle part of thelateral second order pinna of a tripinnate frond. Sinceit is established with certainty that this species hastripinnate fronds (see above), the second variant seemsmore plausible.

The rachis of the penultimate order pinna is well�developed, 2.5–2.6 mm wide, and has a distinct longi�tudinal groove. Three pinnae of the last order are pre�served in attachment to the rachis of the penultimateorder pinna, the lower of these pinnae is completelyfertile, and two upper are partially fertile. The point ofattachment of the lower completely fertile pinna of thelast order to the rachis of the penultimate order pinnais not preserved; however, the relative position of thelast order pinnae is definite evidence that theybelonged to the same frond. In addition to three lastorder pinnae located to the left (according to the posi�tion in Fig. 13) of the rachis of the penultimate orderpinna, this frond fragment retains the rachis base ofone more pinna of the last order, which is located tothe right of the rachis of the penultimate order pinna.A pinnule base with partially preserved midvein andtwo lateral veins is observed in attachment to thisrachis. The relative positions of rachises of the lastorder pinnae on the rachis of the penultimate orderpinna show that the last order pinnae are in alternatingarrangement.

The rachises of the last order pinnae are relativelythin, up to 1 mm thick, smooth, slightly curvedtowards the frond apex. They are attached to the rachisof the penultimate order pinna at an angle of 65°. Theobserved last order pinnae range from 36 to 40 mm oflength.

The last order pinnae have up to eight pairs of pin�nules. The apex of the last order pinna has an apicalpinnule with two or three basal lobes located on thepinnule margins and corresponding morphogeneti�cally to rudimentary pinnules fused by their margins.

Sterile pinnules are subtriangular in outline,pecopteroid, with coalescently fused bases, whichform a well�developed border (limb) of the rachis.Pinnules are attached to the rachis of the last orderpinna at an angle of 60°. Sterile pinnules are 4.5 mmwide, 7.5 mm of the maximum observable length, anddisplays rather distinct venation. Each sterile pinnulehas a well�developed midvein, which extends along thepinnule for three�fourths of its length and then bifur�cates. Resulting branches are simple or one or both ofthem bifurcate one more time.

Each well�developed sterile pinnule contains threeor four pairs of lateral veins, the base of which is asthick as the midvein, rapidly becoming narrower

towards the tip. The basal lateral veins bifurcate one ortwo times. Repeated bifurcation is usually onlyobserved in the acroscopic branch of the lateral vein;both branches of the lateral vein bifurcate less often.The lateral veins in the middle part of the pinnulebifurcate once. Rarely, the acroscopic branch gives riseto additional branches. One or two simple lateral veinsare located in the pinnule apex below (proximal to) theveins formed by the branching apex of the midvein.

The margins of sterile pinnules range from wavy toslightly crenate, although the basal parts of pinnulessometimes contain small marginal lobes, identical tomarginal lobes of sterile pinnules of the syntypedescribed above. The midvein of pinnules is archedtowards the base of the last order pinna. The angle ofattachment of pinnules to the rachis of the last orderpinna gradually increases towards the apex of the lastorder pinna.

Fertile pinnules are mostly pecopteroid; however,pinnules located in the middle and proximal parts offertile pinna of the last order rarely have slightly con�stricted margins. Partially fertile pinnae of the lastorder have two or three fertile pinnules at the base ofthe basiscopic pinnule row. Fertile pinnules are similarin habitus to sterile pinnules, but somewhat longer andnarrower. The maximum size of fertile pinnules is8 × 3 mm. Towards the apex of fertile pinna of the lastorder, pinnules gradually decrease in length; however,the width remains almost constant, slowly decreasingto 2.7 mm. The apices of fertile pinnules are round; thelateral margins of pinnules are subparallel.

Venation of fertile pinnules is very weak. The mid�vein is in the middle part of the pinnule and passes forthree�fourths of its length.

The abaxial surface of fertile pinnules has manysporangia. The sporangia are isolated, not assembledin sori or other complex structure. One fertile pinnuleusually has at least fifty sporangia.

The sporangia range in shape from rounded ellipticto clavate and in size from 200 × 100 to 400 × 200 µm.The sporangial base is extended cuneate and the apicalpart is widened. The sporangial wall is single�layered,consists of elongated polygonal (usually tetragonal orpentagonal) cells. The cells extend along the spo�rangium from the base to tip. At the sporangium base,they are more extended, becoming more isometrictowards the tip, except for specialized cells surround�ing the stomium (see below). The wall cells of the spo�rangium are on average 50 × 20 µm. The apical part ofthe sporangium contains a slitlike stomium up to 80µm long. The stomium is surrounded by one completerow or two incomplete rows of specialized cells, whichare larger than wall cells of the sporangium and havethickened cell walls. The specialized cells provided themechanism for opening the stomium. At the spo�rangium apex, they range from 50 × 20 to 100 × 30 µm.

Spores preserved in situ have been extracted fromthe sporangia (Pl. 16, figs. 2–10). They are from round



to subtriangular in outline, with fine�tuberculate(“shagreen”) surface ornamentation and distincttriradiate tetrad scar. The scar rays (= trilete mark)closely approach the spore equator. In well�developedspores, the scar margins sometimes have a laesuralthickening (Pl. 16, figs. 4, 6, 10). Spores range from 15to 25 µm in diameter.

C o m p a r i s o n. The new species is most similarto Todites emarginatus sp. nov., differing from it in theabsence of well�developed constriction of basal mar�gins of sterile pinnules, the subtriangular shape of pin�nules, the simpler venation of sterile pinnules, and inthe much stronger developed limb of the rachis of thelast order pinnae. It differs from T. lobulatus in thepecopteroid rather than sphenopteroid fertile pinnulesand the significantly fewer sporangia per fertile pin�nule (50 versus 100–120).

R e m a r k s. The collection under study includestwo fern specimens referred to this species. Bothspecimens come from the same core piece, but differ�ent planes separated by 9 mm. A more representativefragment of the middle part is an at least bipinnatefrond, which is designated as the holotype (GIN,no. 4851/203a); the second specimen (GIN,no. 4851/203b) is the basal part of a very large tripin�nate frond.

M a t e r i a l. Holotype and syntype.

Genus Dvinopteridium Zalessky, 1937, emend. nov.

Dvinopteridium Zalessky: Zalessky, 1937a, p. 17.

Ty p e s p e c i e s. Dvinopteridium edemskiiZalessky, 1937; northern Russian Platform; UpperPermian, Severodvinian Stage.

D i a g n o s i s. Fronds tripinnate. Thick robustrachis of penultimate pinna bearing last order pinnaeattached to rachis of penultimate order pinna at 45°–55°in regular alternating order. Neuro�cladophleboidpinnules attached to rachis of last order pinna at50°–55°. Pinnule margins lobate. Lobes curvingtowards pinnule apex. Venation varying from pinnate(with midvein) to neuropteroid (without midvein).Pinnule apex from round to acute. Pinnules slightlycurving towards last order pinna apex. Sporangiaround, free, densely spaced on abaxial surface of fertileor partly fertile pinnules.

D e s c r i p t i o n. Regularly alternating rachises ofthe last order pinnae are attached to a thick robustrachis of the penultimate order pinna at an angle of45°–55°. Neuro�cladophleboid pinnules with a lobatemargin are attached to rachises of the last order pinnaeat an angle of 50°–70°. The angle of attachment ofpinnules gradually decreases towards the apex of thelast order pinna. Lobes are always significantly curvedtowards the pinnule apex. The angle of attachment ofbasal pinnules of both anadromic (acroscopic) andcatadromic (basiscopic) rows sometimes reaches 70°.Venation ranges from pinnate to flabellate neu�

ropteroid (in apical pinnules). The pinnule apicesrange from rounded to pointed. Pinnules slightly curvetowards the frond apex. The sporangia are round, iso�lated, not forming sori or synangia, positioned on theabaxial surface of fertile or partially fertile pinnules.

C o m p a r i s o n. The main distinctive feature ofthe genus Dvinopteridium from the formal genusPecopteris Brongniart (Brongniart, 1822) is the combi�nation of neuropteroid and pecopteroid venationwithin the same frond. Additional morphological dis�tinction of the genus is the presence of the specificmarginal lobes, which are sometimes excessivelydeveloped and curved towards the pinnule apex. Thesemorphological features have not been recorded inother fern genera of the pecopterid morphologicalgroup.

Judging from the sporangial structure, the genusDvinopteridium should be referred to osmundaceanferns (family Osmundaceae). This genus differs fromother genera of the family in the structure of sterileleaves, which are sufficiently distinct to take Dvinopte�ridium for a valid taxon.

R e m a r k s. In the initial study, the generic diag�nosis has not been provided. Zalessky restricted him�self to the statement that this fern is similar in mor�phology to the genus Gondwanidium, indicating that“M.D. Edemsky et E.M. Lutkievicz ont fourniencore, comme provenant probablement des dépôtsdu méme étage Tatarien sur la rivière Soukhona, desempreintes d’une nouvelle fougère pluripinnée dontj’ai donné la description sous la dénomination de Dvi�nopteridium edemskii Zalessky ressemblant par lecaractère de ses pinnules et par leur nervation auxreprésentants du genre Gondwanidium” [M.D. Edem�sky and E.M. Lutkevicz also provided the imprints ofa new fern possibly originated from the deposits of thesame Tatarian Stage of the Sukhona River; this fern isdescribed by me under the name Dvinopteridiumedemskii Zalessky, which is similar in the character ofits pinnules and venation to representatives of thegenus Gondwanidium] (Zalessky, 1937a, p. 17, transla�tion by S.V.N.].

S p e c i e s c o m p o s i t i o n. Type species.O c c u r r e n c e. Upper Permian, Severodvinian

Stage; northern Russian Platform.

Dvinopteridium edemskii Zalessky, 1937, emend. nov.


Dvinopteridium edemskii Zalessky: Zalessky, 1937a, p. 17, text�figs. 3 and 4; Naugolnykh, 2004, pl. XV, fig. 1.

Pecopteris sp. AVG�1: Gomankov and Meyen, 1986, pp. 38–39,text�fig. 11.

Pecopteris sp. (“Dvinopteridium edemskii” Zalessky): Goman�kov and Meyen, 1986, pl. IV, fig. 4.

Pecopteris sp. AVG�1 (“Dvinopteridium edemskii” Zalessky):Arefiev and Naugolnykh, 1998, pl. VI, fig. 11.

H o l o t y p e. Has not been designated in the orig�inal description.

1424


NAUGOLNYKH

1

Plate 18

2

3 5

4

6 7



L e c t o t y p e. Designated by the present author(figured by Zalessky, 1937a, p. 18, text�fig. 3);Sukhona River, village of Porog. The syntype figuredby Zalessky comes from the section on the SukhonaRiver the mouth of the Strel’na River (Zalessky, 1937a,text�fig. 4).

D i a g n o s i s. The species belongs to a monotypicgenus.

D e s c r i p t i o n (Figs. 16, 17). The collectionunder study contains representative fragments ofbipinnate fronds (Pl. 18, fig. 6; Pl. 19, figs. 1, 2, 4;

(a)

(b)(c)

(d) (e)

(f)

(g)

(h)

Fig. 16. Structure of sterile and fertile leaves of Dvinopteridium edemskii Zalessky: (a) specimen GIN, no. 4851/193, three roundsporangia with slitlike apical stomium surrounded by large cells; (b) specimen GIN, no. 4851/195, middle part of the last orderpinna with pinnules of the neuro�cladophleboid type; (c) specimen GIN, no. 4851/193, fragment of a fertile last order pinna,with round apices of two pinnules and sporangia attached to the abaxial surface of pinnules shown on the right; (d) specimenGIN, no. 4851/193, partially disrupted fertile pinnule, with many round sporangia; (e) specimen GIN, no. 4851/193, middle partof the last order pinna with neuro�cladophleboid pinnules; (f) hypothetical structure of sporangium, apical part of sporangiumdisplays slitlike stomium surrounded by specialized cells with thickened walls; (g) specimen GIN, no. 4851/197, middle part ofbipinnate frond; and (h) specimen GIN, no. 4851/194, last order pinna with neuro�cladophleboid pinnules. Locality: Nikulino,Vologda Region; Upper Permian, Severodvinian Stage. Scale bars: (a, f) 0.5 mm, (b, d, e, g, h) 1 cm, and (c) 0.5 cm.


Figs. 1–7. Leaf morphology Dvinopteridium edemskii Zalessky: (1) specimen GIN, no. 4851/195, middle part of a last order pinnawith neuro�cladophleboid pinnules; (2) specimen GIN, no. 4851/193, three round sporangia with apical slitlike stomium sur�rounded by large cells; (3) specimen GIN, no. 4851/193, partially disrupted fertile pinnule with many round sporangia; (4) spec�imen GIN, no. 4851/193, fragment of a fertile last order pinna, with apices of two pinnules and round sporangia located on theabaxial surface of the leaf lamina seen on the right; (5) specimen GIN, no. 4851/193, middle part of a last order pinna with neuro�cladophleboid pinnules; (6) specimen GIN, no. 4851/197, middle part of a bipinnate frond; and (7) specimen GIN,no. 4851/194, last order pinna with neuro�cladophleboid pinnules. Locality: Vologda Region, Nikulino; Upper Permian,Severodvinian Stage. Scale bars: (1, 3, 5–7) 1 cm, (4) 0.5 cm, and (2) 0.5 mm.

1426


NAUGOLNYKH

Figs. 16g, 17) and isolated last order pinnae. In addi�tion, there is one specimen, the basal part of a largetripinnate frond, the rachis of which is 15 mm wide.Judging from the measurements of the frond frag�ments, a complete tripinnate frond of this fern specieswas more than 70–80 cm long. The total length of thepenultimate order pinnae remains uncertain, althoughavailable fragments suggest that these pinnae wereabout 30–40 cm long at 12–15 cm of width. The max�imum width of the rachis of the last order pinnae isabout 2 mm. The rachis is covered with fine longitudi�nal rugosity. A distinct longitudinal groove is absent.

Rachises of the last order pinnae are attached torachises of the penultimate order pinnae at an angleranging from 45° to 55° (it sometimes reaches 90°) inthe alternating order. Rachises of the last order pinnaehave pinnules attached to rachises at an angle of 50°–70° in regular alternating order. Pinnules are neuro�cladophleboid, pecopteroid (Pl. 18, figs. 6, 7; Pl. 19,figs. 2, 4; Figs. 16g, 16h, 17), or, less often, classic cla�dophleboid (Pl. 18, figs. 1, 5; Pl. 19, figs. 1, 3;Figs. 16b, 16e), with crenate or lobate margin. Well�developed pinnules are usually 5 × 12 mm. The acro�scopic margin of pinnules usually has deeper incisionsthan the basiscopic margin (Fig. 17). Lobes of well�developed pinnules are usually distinctly inclinedtowards the pinnule apices.

Venation of pinnules ranges from pinnate, with awell�pronounced midvein, to neuropteroid, almostwithout a midvein. The midvein is well�developed inpinnules of the basal or middle parts of the last orderpinna. It extends along the middle of the pinnule,reaching the last one�third of its extent and bifurcates.Then, the acroscopic branch produced by this bifurca�tion dichotomizes regularly (isotomically) two ormore times and the basiscopic branch dichotomizesonce. In some cases, the order of branching changesseveral times, so that the basiscopic branch forms onemore bifurcation. The midvein is thin, but distinct,usually slightly undulating. A well�developed pinnule

usually has four or five pairs of lateral veins, whichdiverge from the midvein of the pinnule in the alter�nating order. Each pinnule lobe generally contains twoends of lateral veins.

The sporangia are attached to the abaxial surface offertile pinnules. They are round, from 0.5 to 0.9 mm indiameter, with slitlike stomium, surrounded by large spe�cialized cells (Pl. 18, figs. 2–4; Figs. 16a, 16c, 16d, 16f).One fertile pinnule usually has more than 20 sporan�gia. A pinna usually combines both fertile and sterilepinnules (Pl. 18, fig. 4; Fig. 16c).

M a t e r i a l. Five representative fragments ofbipinnate and tripinnate fronds.

C L A S S G L E I C H E N I O P S I D A D O W E L D , 2 0 0 1

Order Gleicheniales Frank, 1877

Family Gleicheniaceae Presl, 1825

Genus Tumidopteris Naugolnykh, gen. nov.

E t y m o l o g y. From the Latin tumidus (swollen),based on the voluminous, as though swollen sporangia.

Ty p e s p e c i e s. Tumidopteris clavata Naugol�nykh, sp. nov.

D i a g n o s i s. Fronds at least bipinnate. Pinnulessphenopteroid, pinna rachis with well�developed limb.Pinnule apex acute or pointed. Elliptical sporangiaarranged in rosette�like sori, with 8–11 sporangia persorus. Sporangial bases very slightly fused and attachedto round receptacle. Apical part of sporangium withsmall but distinct sinus.

C o m p a r i s o n. The new genus is most similar toPrynadaeopteris, emend. nov. and differs from it in theweakly fused bases of sporangia (Prynadaeopteris haswell�fused sporangial bases), the fewer sporangia persorus (8–11 versus 14–20), the presence of an apicalsinus in sporangia, and in the distinctly sphenopteroidfertile and sterile pinnules. The new genus is also sim�ilar to the genus Hapalopteris Stur (Stur, 1883), differ�ing from it in the well�developed rachial limb in fertilepinnules, pointed rather than rounded apices of pin�nules and lobes, and in many sporangia per sorus, atleast eight (versus four or five sporangia in sori ofHapalopteris). Structural details of the sporangia ofHapalopteris or exact taxonomic position of this genusremain uncertain. Ferns with similar morphology areknown from the Upper Carboniferous of the DonetsBasin (A.K. Shchegolev (Institute of Geological Sci�ences, Kiev, Ukraine), personal communication,1989). Sterile pinnules of Tumidopteris differ from themorphologically similar sterile pinnules of the genusGeperapteris (and Prynadaeopteris) in the distinctlysphenopteroid shape.

S p e c i e s c o m p o s i t i o n. Type species.O c c u r r e n c e. Middle and Upper Permian of

the Pechora Coal Basin, Fore�Urals near the PechoraRiver, and, probably, Siberia and Mongolia.

Fig. 17. Pinnule morphology in Dvinopteridium edemskiiZalessky, specimen GIN, no. 4851/191; Nikulino locality,Vologda Region; Upper Permian, Severodvinian Stage.Scale bar, 1 cm.



1

Plate 19

3 4

2


Figs. 1–4. Leaf morphology of Dvinopteridium edemskii Zalessky: (1) specimen GIN, no. 4851/191, middle part of a bipinnatefrond; (2) specimen GIN, no. 4851/191, basal frond part; (3) specimen GIN, no. 4851/192, last order pinna with cladophleboidpinnules; and (4) specimen GIN, no. 4851/191, general morphology of the middle frond part. Locality: Vologda Region,Nikulino; Upper Permian, Severodvinian Stage. Scale bar, 1 cm.

1428


NAUGOLNYKH

Tumidopteris clavata Naugolnykh, sp. nov.

Plate 20, figs. 1–6; Plate 21, figs. 1–6; Plate 22, figs. 1–4

E t y m o l o g y. From the Latin clavata, based onthe clavate shape of sporangia.

H o l o t y p e. GIN, no. 4851/167, fertile frondfragment; Pechora Coal Basin, Khalmer’yusskoe coalmine, borehole KhK�1137, 244.2 m of depth; desig�nated here (Pl. 20, figs. 1–6; Pl. 21, figs. 1–6;Figs. 18–21).

D i a g n o s i s. The species belongs to a monotypicgenus.

D e s c r i p t i o n (Figs. 18–23). Fertile leaves aresphenopteroid, with a well�developed leaf lamina. Therachis of fertile pinnae of the last order curves slightlytowards the frond apex. The rachis has a shallow, butdistinct longitudinal groove on the adaxial surface. Awell�developed wing of the rachis (limb) is alwayspresent. Pinnules are attached to the rachis of the lastorder pinna at an angle of 45°–55°. They are sphe�nopteroid, with an extended base and pointed apex.Each pinnule is dissected into four or five lobes,which, in turn, have lobes of the second order. Thedepth of dissection of the pinnule leaf lamina intolobes of the first order is usually at most half distancefrom the lobe apex to midvein of the pinnule. The pin�nule midvein is distinct, relatively thin, and straight. In

Fig. 19. Morphology of the leptosporangiate fern Tumidopteris clavata Naugolnykh, gen. et sp. nov., holotype GIN, no. 4851/167,basal part of fertile pinnule with four preserved sori. Locality: KhK�1137, Khalmer’yusskoe Coal Field; Middle Permian, Kaza�nian Stage. Scale bar, 1 mm.

Fig. 18. Morphology of the leptosporangiate fernTumidopteris clavata Naugolnykh, gen. et sp. nov., holo�type GIN, no. 4851/167, middle part of fertile penultimateorder pinna. Locality: KhK�1137, Khalmer’yusskoe CoalField; Middle Permian, Kazanian Stage. Scale bar, 1 mm.



the apical pinnule part, the midvein is sometimesweakly undulating. Lateral veins extend from the pin�nule midvein into the first order lobes at an angle of45°–50° and successively dichotomize up to threetimes. The ends of lateral veins of fertile pinnules haverosette�like sori composed of 8–11 almost free ellipticsporangia slightly fused at their bases and attached towell�developed round receptacle. The apical part ofsporangia has a relatively small sinus, probably corre�sponding to the point of opening the sporangium aftermaturation of spores. Sori vary in diameter from 0.7 to1 mm. The sporangium varies from 400 to 450 µm oflength at the maximum width of 300 µm. The sporan�gial wall is thin, consisting of one cell layer. Cells of thesporangial wall are elongated, on average 25 × 100 µm.Some cells are excessively well developed, up to210 µm long. Cells extend along the sporangium axis;only at the apex, near the apical sinus, they sometimesslightly curve towards the sinus. The cell walls of spo�rangia are thin, straight or slightly curved.

The epidermal–cuticular structure of the abaxialside of the leaf lamina of the fertile pinnule was exam�ined (Pl. 20, figs. 3–6; Fig. 21b). Costal (supracostal)

Fig. 20. Morphology of the leptosporangiate fernTumidopteris clavata Naugolnykh, gen. et sp. nov., holo�type GIN, no. 4851/167, sorus. Locality: KhK�1137,Khalmer’yusskoe Coal Field; Middle Permian, KazanianStage. Scale bar, 100 µm.

(a)(b)

Fig. 21. Morphology of the leptosporangiate fern Tumidopteris clavata Naugolnykh, gen. et sp. nov., holotype GIN, no. 4851/167:(a) sorus and (b) epidermis of fertile pinnule. Locality: KhK�1137, Khalmer’yusskoe Coal Field; Middle Permian, KazanianStage. Scale bar, 100 µm.

1430


NAUGOLNYKH

cells are very long (up to 300 µm long) and narrow (onaverage about 10–15 µm wide). Intercostal cells areisometric, round or slightly extended along the veins,hexagonal or heptagonal. Cell walls are thin, straight

or slightly curved. The stomata are relatively regularlydistributed over the intercostal fields. The stomatalapertures directed uniformly, along neighboring veinsof the pinnule which border the intercostal field.

(a)

(b)

(c)

(d)(e)

Fig. 22. Morphology of sterile leaves and a partially fertile leaf presumably belonging to the leptosporangiate fern TumidopterisNaugolnykh, gen. nov.: (a) specimen GIN, no. 4851/168; (b) specimen GIN, no. 4851/169; (c) specimen GIN, no. 4851/170;(d) specimen GIN, no. 4851/171; and (e) specimen GIN, no. 4851/172. Localities: Pechora Coal Basin; Middle Permian, Kaza�nian Stage: (a) borehole IK�687, 102 m of depth; (b) borehole IK�426, 729.4 m of depth; (c) borehole IK�686, 104 m of depth;(d) borehole VK�33, 753.2 m of depth; and (e) borehole IK�686, 94 m of depth. Scale bar, 1 cm.



Guard cells are 40–50 µm long and 15–20 µm wide.Subsidery cells are nonspecialized. Trichome struc�tures are absent.

In addition to fertile pinnae, there are sterile pen�ultimate order pinnae (Pl. 22, figs. 1–4; Figs. 22, 23).The distinctive sphenopteroid shape of pinnules, with

(a)

(b)

(c)

(d)(e)

Fig. 23. Morphology of sterile leaves and a partially fertile leaf presumably belonging to the leptosporangiate fern TumidopterisNaugolnykh, gen. nov., shadow diagrams: (a) specimen GIN, no. 4851/168; (b) specimen GIN, no. 4851/169; (c) specimenGIN, no. 4851/170; (d) specimen GIN, no. 4851/171; and (e) specimen GIN, no. 4851/172. Localities: Pechora Coal Basin; Mid�dle Permian, Kazanian Stage: (a) borehole IK�687, 102 m of depth; (b) borehole IK�426, 729.4 m of depth; (c) borehole IK�686,104 m of depth; (d) borehole VK�33, 753.2 m of depth; and (e), borehole IK�686, 94 m of depth. Scale bar, 1 cm.

1432


NAUGOLNYKH

1

Plate 20

2

3

4

5 6

E x p l a n a t i o n o f P l a t e 2 0Figs. 1–6. Morphology of the leptosporangiate fern Tumidopteris clavata Naugolnykh, gen. et sp. nov., holotype GIN, no. 4851/167:(1) group of five sori; (2) structure of an individual sorus; (3, 4) structure of the midvein of a fertile pinnule, with distinct longitudinalribbing corresponding to the arrangement of conducting elements; (5) deviation of the lateral vein (top) from the midvein of a fertilepinnule; (6) epidermal structure of a fertile pinnule (SEM), showing common epidermal cells (see also Fig. 21b). Locality: KhK�1137,Khalmer’yusskoe Coal Field; Middle Permian, Kazanian Stage. Scale bars: (1, 4) 200 µm, (2) 500 µm, and (3, 5, 6) 100 µm.



1

2

3

4

5 6

Plate 21


Figs. 1–6. Morphology of the leptosporangiate fern Tumidopteris clavata Naugolnykh, gen. et sp. nov., holotype GIN,no. 4851/167: (1) group of three sori; (2, 3, 5) individual sori; (4) two sori of different size; in smaller sorus, sporangia are spacedwider; and (6) sporangium with notched apex. Locality: Khalmer’yusskoe Coal Field, KhK�1137; Middle Permian, KazanianStage. Scale bars: (1) 500 µm and (2–6) 200 µm.

1434


NAUGOLNYKH

1

Plate 22

2

3 4


Figs. 1–4. Sterile leaves presumably belonging to the leptosporangiate fern Tumidopteris clavata Naugolnykh, gen. et sp. nov.:(1) specimen GIN, no. 4851/168; (2) specimen GIN, no. 4851/170; (3) specimen GIN, no. 4851/172; and (4) specimen GIN,no. 4851/169. Localities: Pechora Coal Basin; Middle Permian, Kazanian Stage: (1) borehole IK�687, 102 m of depth; (2) bore�hole IK�686, 104 m of depth; (3) borehole IK�686, 94 m of depth; and (4) borehole IK�426, 729.4 m of depth. Scale bar, 1 cm.



a well�developed limb of rachises of the last order pin�nae allow us to suggest that these pinnae belonged tothe same parent plant.

R e m a r k s. The new species is similar to Oligocar�pia brongniartii Stur (Stur, 1883, p. 688, text�fig. 16) inthe general design of sphenopteroid fertile pinnules(particularly in the arrangement of sori at the end oflateral veins) and differs from it in the absence ofannulus, the elongated shape of sporangia, and in theposition of sporangia within the same plane.

Some insignificant structural variations of the leaflamina (in the pinnule width and depth of dissection ofthe pinnule margin) of Tumidopteris clavata sp. nov.are probably accounted for by the effect of intraspe�cific variation.

M a t e r i a l. Middle part of a fertile bipinnatefrond, three fertile pinnae, and four sterile penulti�mate order pinnae.

Genus Geperapteris S. Meyen, 1982Geperapteris rotunda Naugolnykh, sp. nov.

Plate 23, figs. 1–5

E t y m o l o g y. From the Latin rotunda (round),based on the round shape of sori.

H o l o t y p e. GIN, no. 4851/175; middle part of afertile frond; Pechora Coal Basin, borehole VK�12,807.5 m of depth; Middle Permian, Kazanian Stage;designated here (Pl. 23, figs. 1–5; Fig. 24).

D i a g n o s i s. Fronds bipinnate or tripinnate.Frond rachis undulating. Pinna rachis straight. Pin�nules pecopteroid, with lobate margins. Sori capitate,round, hemispherical, consisting of 14–18 sporangiaattached to receptacle. Spores round to subtriangular,30–40 µm in diameter, with trilete scar. Rays of triletescar reaching one�third of spore radius. Proximal sur�face of spore smooth, distal surface ornamented byfine scabrae.

D e s c r i p t i o n (Fig. 24). The leaves are bipin�nate to tripinnate. Rachises of the last order pinnae arestraight. Pinnules are wide, pecopteroid, with a lobedmargin. Sori are rounded hemispherical, consist onaverage of 14–18 elongated sporangia, radiallyarranged around the common base (receptacle).

I examined two fragments of the middle part ofbipinnate fertile leaves, which come from differentplanes of the same sample (the planes are separated bya 14�mm�thick carbonaceous claystone interbed).The specimen designated as the holotype is the middlepart of a penultimate order pinna, with five fertile pin�nae preserved in attachment to the rachis of the penul�timate order pinna. The rachis has a weakly developedshallow longitudinal groove. The rachis of the penulti�mate order pinna is distinctly undulating, curvestowards the base of the last order pinnae.

The regularly alternating last order pinnae areattached to the rachis of the penultimate order pinnaat an angle of 75°–80°. Rachises of the last order pin�

nae are straight or slightly curved towards the base ofthe last order pinna or, on the contrary, towards theapex of the last order pinna. The relatively well�pre�served last order pinnae of the syntype are more than30 mm long. The maximum observable width ofrachises of the last order pinnae is 0.9 mm.

Pinnules are attached to rachises of the last orderpinnae at an angle of 50°–60°. They are alternating,pecopteroid, with a wide base, on average 14–15 mmlong and 5 mm wide. The base of sterile pinnules dis�tinctly descend along the rachis. Pinnules located inthe apical part of the last order pinnae are attached tothe rachis at a sharper angle than pinnules of the mid�dle and basal parts to the last order pinnae. The angleof pinnule attachment gradually decreases towards theapex of the last order pinna. Venation is pinnate. Themaximum observable width of the midvein is 0.3 mmand lateral veins are 0.1 mm wide. Lateral veins divergefrom the midvein at an angle of 45°. They dichotomizeonce. Each pinnule has four pairs of lateral veins.

The end of each branch of lateral veins and thedichotomizing end of the midvein have sori on theabaxial surface of fertile pinnules. Each pinnule has upto 18–20 sori.

The sori are round, about 0.9 mm in diameter, con�sist of 14–18 elongated clavate sporangia, connectedby their narrow end to the common receptacle. Theyare 0.4–0.5 mm long and 0.07–0.08 mm wide. Sporeshave been extracted from sporangia. They range fromround to subtriangular in outline, are trilete, 30–40 µm in diameter. Rays of the trilete scar terminateshort of reaching the spore equator, at a distance oftwo�thirds of the spore radius. The proximal spore sur�face is smooth and the distal surface is covered by finescabrate ornamentation.

C o m p a r i s o n a n d r e m a r k s. The new spe�cies differs from the type species Geperapteris imitansin the considerably wider angle of attachment of thelast order pinnae to the rachis of the penultimate orderpinna (in G. imitans, it is 30°–55°; in G. rotunda sp.nov., 75°–80°) and the considerably fewer sporangiaper sorus (G. imitans has 60 or more and G. rotunda sp.nov. has 14–18). In G. rotunda sp. nov., bipinnateleaves are known; however, it is probable that completefronds of this species were tripinnate or even quad�ripinnate.

M a t e r i a l. Holotype and syntype.

Genus Prynadaeopteris Radczenko, 1955, emend. nov.

Ty p e s p e c i e s. Prynadaeopteris karpovii Radc�zenko, emend. nov.; Upper (Middle) Permian of theKuznetsk Basin and Pechora Coal Basin.

D i a g n o s i s. Fronds at least tripinnate, rachisrelatively thin, straight or slightly curved, could bewinged. Sterile pinnules pecopteroid, sometimes withconstricted bases, acute apices, and lobate lateral mar�gins. Midvein well developed, slightly curved, having

1436


NAUGOLNYKH

(a)

(b)

(c) (d) (e)

Fig. 24. Geperapteris rotunda Naugolnykh, sp. nov., holotype GIN, no. 4851/175: (a) venation in individual pinnule, (b) macro�morphology, and (c, d) spores preserved in sporangia of the holotype in situ. Locality: Pechora Coal Basin, borehole VK�12, 807.5 mof depth; Middle Permian, Kazanian Stage. Scale bars: (a) 0.5 cm, (b) 1 cm, and (c, d) 20 µm.

E x p l a n a t i o n o f P l a t e 2 3Figs. 1–5. Geperapteris rotunda Naugolnykh, sp. nov., holotype GIN, GIN, no. 4851/175: (1) macromorphology, middle part ofa fertile frond; (2) venation in an individual pinnule; (3–5) spores preserved in sporangia in situ. Locality: Pechora Coal Basin,borehole VK�12, 807.5 m of depth; Middle Permian, Kazanian Stage. Scale bars: (1) 1 cm, (2) 0.5 cm, and (3–5) 20 µm.



1

Plate 23

2

3 4 5

1438


NAUGOLNYKH

from three to six lateral veins bifurcating once or twice.Fertile pinnules similar to sterile pinnules, but pos�sessing sori. Sori located on abaxial leaf surface nearpinnule margin. Each sorus containing 14–20 radiallypositioned obovate or clavate free sporangia attachedby their bases to well�developed round or subcylindri�cal central receptacle. Annulus absent.

S p e c i e s c o m p o s i t i o n. Prynadaeopteriskarpovii Radczenko, P. sibirica (Zalessky) Radczenko,P. eurina (Zalessky) Radczenko, P. dymovii Radc�zenko, and P. vorcutana (Zalessky) Fefilova.

C o m p a r i s o n. The genus Prynadaeopteris ismost similar to Tumidopteris described in the presentstudy and differs from it in the greater number of spo�rangia per sorus (14–20 in Prynadaeopteris and 8–11in Tumidopteris), the absence of an apical sinus (apicalnotch) at the sporangial apex, and in the pecopteroidrather than sphenopteroid fertile and sterile pinnules.

R e m a r k s. The taxonomic status of the genusPrynadaeopteris, which is frequently included in thelists of Permian floras of the Fore�Urals, and its spe�cies composition are debatable.

Meyen (1982c) has shown that the genusPrynadaeopteris in the initial sense is essentially hetero�geneous. In addition to forms with rosette�like sori con�sisting of relatively small number (usually at most 20) ofrelatively large sporangia, the genus included formswith capitate sori consisting of many (up to 60 or evenmore) relatively small sporangia. Meyen transferredthese forms to a separate genus, Geperapteris, mem�bers of which are characteristic primarily of the UpperPaleozoic beds of central Angaraland; however, as hasbeen shown later (see above the description of Geper�apteris rotunda Naugolnykh, sp. nov.), they also occurin the Permian of the Pechora Coal Basin and arecharacterized by fewer sporangia per sorus (14–18),but have clearly capitate sori.

I examined a well�preserved specimen of the typespecies Prynadaeopteris karpovii, which is the middlepart of a well�developed fertile frond (Pl. 24, figs. 1–6;Figs. 25c, 25d), coming from the Middle Permian(=Biarmian) coal�bearing beds of the Pechora Basin,which are assigned to the Kazanian Stage. Based onthe study of this material, I provided above the newformalized diagnosis of the genus Prynadaeopteris,which is accompanying by the table showing changesin the species composition of this genus in the courseof investigation (see below). The morphology of fertileand sterile leaves that were assigned by differentauthors in different years to the genus Prynadaeopterisis shown in Figs. 25a, 25b, 26a–26m.

It is proposed that the genus comprises the follow�ing species: Prynadaeopteris karpovii, P. sibirica,P. eurina, P. dymovii, and P. vorcutana. Some of thesespecies may be synonymous.

Table 1 shows a new combination: Geperapterisalifera (Fefilova) Naugolnykh, comb. nov. Basionym:Prynadaeopteris (?) alifera Fefilova (1973, pp. 63–65;

Pl. 15, figs. 1–4; Fig. 14). Holotype: GIN, no. 302/1(179�24/1), it was figured by Fefilova (1973, pl. XV,figs. 1, 2).

O c c u r r e n c e. Upper Carboniferous and Per�mian of northeastern Eurasia and Greenland (Wagneret al., 1982, 1999, 2002).

Prynadaeopteris karpovii (Radczenko, 1936) Radczenko, 1955, emend. nov.


Sphenopteris (Oligocarpia?) karpovii Radczenko: Radczenko,1936, p. 4, pl. I, figs. 3–5, text�figs. 2–5; Neuburg, 1948, pp. 105–107, pl. XX, figs. 3 and 4.

Prynadaeopteris karpovii (Radczenko) Radczenko: Radc�zenko, 1955, pp. 49–51, text�figs. 51–53; Radczenko, 1956,pp. 145–146, pl. XXV, figs. 6 and 7, pl. XXVI, fig. 1; Gorelova andRadczenko, 1962, pp. 96–97, pl. VIII, figs. 1–5; Fefilova, 1973,pp. 49–51, text�fig. 9, pl. XI, figs. 1–3; Pukhonto and Fefilova,1983, pp. 42–43, pl. VI, fig. 8; Pukhonto, 1998, pl. 37, fig. 4.

Pecopteris (Prynadaeopteris) aff. karpovii Radczenko: Pukh�onto and Fefilova, 1983, pl. VI, fig. 8.

H o l o t y p e. Apical part of a sterile frond figuredby Radczenko (1955, text�fig. 51) and reproducedhere in Fig. 20c; it comes from the Upper Permian ofthe Kuznetsk Basin, left bank of the Tom River nearthe village of Nikol’skoe, downstream from the mouthof the Verkhnyaya Ters’ River. Unfortunately, thedepository of the holotype of Prynadaeopteris karpovii isnot known. I tried to find the type material of the genusPrynadaeopteris in the Chernyshev Central ResearchGeological Museum, St. Petersburg (TsNIGRMuseum) and Karpinsky All�Russia Research Insti�tute of Geology, St. Petersburg (VSEGEI); however,they have not been found in these collections, nor inother collections examined by G.P. Radczenko.

N e o t y p e. Specimen GIN, no. 4851/166, largefragment of a fertile frond from the Pechora CoalBasin, Leur�Yaga locality; Middle Permian, KazanianStage is proposed to be taken for the neotype ofPrynadaeopteris karpovii (Pl. 24; figs. 1, 3–6;Figs. 25c, 25d).

D i a g n o s i s. Fronds tripinnate. Pinnules pecop�teroid. Sori consisting of 18–20 radially positionedclavate sporangia fused by their bases and attached towell�developed round central receptacle. Annulusabsent. Sori located near pinnule margin, sometimesforming groups of four or five sori.

D e s c r i p t i o n (Figs. 25b–25d). The collectionexamined contains one large fertile frond (neotype)and two sterile last order pinna.

In the fertile frond, the apex and probably the baseare not preserved; however, available fragment providesufficient data about the leaf structure as a whole.

The frond is at least tripinnate. The rachis of thepenultimate order pinna is thick, on average 2.5 mmthick, naked, virtually lacks a limb. Sometimes, a verythin border which is probably formed due to postdi�agenetic deformation of cover rachidial tissue ratherthan by the leaf lamina, extends along the rachis.



Table 1. Changes in the taxonomic composition of the genus Prynadaeopteris

Radczenko, 1955, 1956 Fefilova, 1973 Meyen, 1982 Naugolnykh, this work:

Prynadaeopteris karpovii Radczenko, 1955; type species

Prynadaeopteris karpovii Radczenko, 1955; type species

– Prynadaeopteris karpovii Radczenko, 1955; type species

Prynadaeopteris sibirica (Zalessky) Radczenko, 1955

– – ?

Prynadaeopteris eurina (Zalessky) Radczenko, 1955

– – ?

Prynadaeopteris dymovii Radczenko, 1955

– – ?

Prynadaeopteris maneichensis (Zalessky) Radczenko, 1955

– ?=Geperapteris imitans (Neuburg) S. Meyen, 1982

=Geperapteris imitans (Neuburg) S. Meyen, 1982

Prynadaeopteris tunguscana (Schmalhausen) Radczenko, 1955

– ?=Geperapteris imitans (Neuburg) S. Meyen, 1982

=Geperapteris imitans (Neuburg) S. Meyen, 1982

Prynadaeopteris venusta Radczenko, 1956

Prynadaeopteris venusta Radczenko, 1956

– Prynadaeopteris venusta Rad�czenko, 1956

Prynadaeopteris vorcutana (Zalessky) Fefilova, 1973

– ?

Prynadaeopteris (?) alifera Fefilova, 1973

– Geperapteris alifera (Fefilova) Naugolnykh comb. nov.

Prynadaeopteris irregularis Radczenko, 1956

– – ?

Prynadaeopteris silovaensis Fefilova, 1973

Geperapteris silovaensis (Fefilova) S. Meyen

Geperapteris silovaensis (Fefilova) S. Meyen

Prynadaeopteris tchernovii (Zalessky) Fefilova, 1973

Geperapteris tchernovii (Zalessky) S. Meyen

Geperapteris tchernovii (Zalessky) S. Meyen

Prynadaeopteris anthriscifolia (Goeppert) Radczenko, 1955

Pecopteris anthriscifolia (Goeppert) Zalessky 1918

– Pecopteris anthriscifolia (Goeppert) Zalessky, 1918

Judging from the rate of decreasing thickness of therachis towards the frond apex, complete frond wasprobably about 0.5 m long and 25–30 cm wide.

Rachises of the last order pinnae are attached in thealternating order to the rachis of the penultimate orderpinna at an angle of 65°–70°. They are 2.5–3 timesthinner than the rachis of the penultimate order pinna,at most 1 mm wide. Judging from the preserved struc�tures, all last order pinnae seen in the frond fragment(except for one; for more detail, see the text below) arecompletely fertile, although in the majority of the lastorder pinnae, the apices are partially torn off.

Fertile pinnules are attached to the rachis of the lastorder pinnae and fused by the margins for at least halfof their extent. Pinnules are subtriangular in outline,on average 7–8 mm long and 5 mm wide. The midveinof pinnules departs from the rachis of the last orderpinna at an angle of 45°. In the upper part of the pin�nule, one or two pairs of lateral veins diverge from themidvein. It is impossible to establish with certainty thestructure of lateral veins of fertile pinnules, since theyare partially or completely covered by the sori attached

to them and positioned above them. The sori areround, have a small depression in the center, whichcorresponds to the position of the receptacle, by whichthe sorus was attached to the vein of the fertile pinnule.In imprints, the depression looks like a distinct smallprominence (see, e.g., Pl. 24, figs. 1, 4, 5). The sori areon average 2.5 mm in diameter, consist of 15–18 radi�ally positioned clavate sporangia, which are widenedtowards the apex and narrowed towards the base. Thesporangia are on average 1 mm long at the maximumwidth of 0.3–0.4 mm.

Sterile pinnae of the penultimate order referred tothis species show in general the same structure as fertileleaves, but lack sori. Pinnules of sterile pinnae are sub�triangular in outline. Pinnules of well�developed sterilepinnae of the last order are on average 7 × 14 mm. Thepinnules are usually pecopteroid, although their basesare sometimes slightly constricted. The pinnule mar�gins are distinctly lobate. The lobe apices are pointed.One lateral vein enters each lobe. Each side of well�developed pinnules sometimes has up to five lobes.

1440


NAUGOLNYKH

(a)

(b)

(c) (d)

Fig. 25. Fertile leaves of various members of the genus Prynadaeopteris Radczenko: (a) morphology of fertile last order pinna ofPrynadaeopteris natalianae Gorelova (after Gorelova and Radczenko, 1962); (b) morphology of Prynadaeopteris karpovii Radc�zenko (type species of the genus Prynadaeopteris according to the figure of the type material in the protolog after Radczenko,1955); and (c, d) Prynadaeopteris karpovii Radczenko, examined specimen GIN, no. 4851/166, surface of leaf lamina marked bypoints. Localities: (a) Kuznetsk Coal Basin, village of Kamyshikha, borehole 1, 92 m of depth; Middle Permian, Kazanian Stage;Il’inka Formation, Leninskaya Subformation; (c, d) Leur�Yaga locality; Middle Permian, Kazanian Stage. Scale bars: (a, bupper) 1 cm and (c, d) 0.5 cm; structure of sorus and fertile pinnule of Prynadaeopteris karpovii Radczenko are given after Rad�czenko (1955) not to scale.



1

Plate 24

2

3 4

5 6


Figs. 1–6. Fertile (1, 3–6) and sterile (2) leaves of Prynadaeopteris karpovii Radczenko, neotype GIN, no. 4851/166: (1) partial fertilelast order pinna, with distinctly visible rosette�like sori; (2) sterile last order pinna; (3) middle part of a fertile frond with three last orderpinnae; (4) relative positions of sori on a fertile pinnule; (5) middle part of a fertile last order pinna; and (6) fertile last order pinna withthe apex partially torn off. Locality: Pechora Basin, Leur�Yaga locality; Middle Permian, Kazanian Stage. Scale bars, 1 cm.

1442


NAUGOLNYKH

C o m p a r i s o n a n d r e m a r k s. The speciesPrynadaeopteris karpovii is similar to P. venusta anddiffer from it in the greater number of sporangia persorus and pointed rather than rounded apices of pin�nules and individual lobes. The generic assignment ofthe species P. venusta remains questionable, becausestructural details of its fertile organs are uncertain.Fefilova (1973) marked that P. venusta is characterizedby the sori consisting of three to five sporangia(Fefilova, 1973); however, in the figures provided byher to show the morphology of fertile pinnules of thisspecies, the sori consist of six or seven sporangia(Fefilova, 1973, text�fig. 12c). The only partly fusedsporangia of P. venusta, like pecopteroid pinnules ofthis fern, suggest that it should be assigned to the genusPrynadaeopteris, whereas a relatively small number ofsporangia per sorus make this species similar to thespecies Tumidopteris clavata sp. nov. Since the speciesPrynadaeopteris venusta combines characters of thegenera Prynadaeopteris and Tumidopteris and isrecorded beginning the Ufimian Stage, it is probablyan ancestral form that corresponds to the base of thedivergent event which resulted in the formation of thegenera Prynadaeopteris and Tumidopteris in the Mid�dle Permian (Kazanian).

In one last order pinna of P. karpovii, the basal partis fertile (Pl. 24, fig. 3) and two�thirds of the pinnalocated distal to the fertile part correspond in mor�phology to sterile leaves of the holotype and syntypefigured by Radczenko (1955) long with the originaldescriptions of the species Prynadaeopteris karpovii(Radczenko, 1955, text�fig. 51, holotype; text�fig. 53,upper part, syntype). Two isolated sterile pinnae showthe same structure as these leaves and, hence, are alsoreferred in the present study to this species. One ofthem is figured in Pl. 24, fig. 2.

The aggregation of sporangia in groups of six orseven in one sorus, which is sometimes observed inPrynadaeopteris karpovii, probably resulted in a toogeneralized and perhaps inaccurate picture of thesorus of P. karpovii in the protolog (Radczenko, 1955,text�fig. 53, lower part; reproduced in the presentstudy, Fig. 19b, lower part).

Sterile leaves of P. karpovii are very similar (may beconspecific) to the species Pecopteris anthriscifolia.

M a t e r i a l. Large fragment of a fertile tripinnatefrond and two isolated sterile pinnae of the last order.

O c c u r r e n c e. Middle Permian, Kazanian Stageof the Pechora Coal Basin, Fore�Urals near thePechora River, Kuznetsk Basin, and Tunguska Basin.

Order incertae sedis

Family Incertae sedis

Genus Myriotheca Zeiller, 1883Myriotheca cf. scaberrima Lesquereux, 1870

Plate 30, fig. 2

Myriotheca cf. scaberrima Lesquereux: Naugolnykh, 2007,text�fig. 53; Naugolnykh, 2009, text�fig. 1a.

D e s c r i p t i o n. The most completely preservedspecimen is the middle part of a bipinnate frond withseven preserved last order pinnae, five of which aremorphologically complete. The frond fragment is35 mm long and 25 mm wide. The rachis is thick, 1.5mm wide, with a fine longitudinal ribbing and distinctgroove on the adaxial surface of the rachis. The lastorder pinnae diverge from the rachis at an angle rang�ing from 70° to 90°. Pinnules range from almost sphe�nopteroid to pecopteroid. Sphenopteroid pinnules arelocated in the basal part of the last order pinnae. Thefirst (basal) pinnule of the acroscopic row is usuallyexcessively developed, 1.5–2 times larger in linearmeasurements than succeeding pinnules.

The sporangia are arranged in round sori, whichare located on the abaxial surface of fertile pinnules.The sori are 1.5 mm in diameter, gradually decreasingin size from the basal part of the pinnule to its apex. Awell�developed fertile pinnule usually has nine sori.Young apical pinnules usually have two or three smallsori (about 1 mm in diameter). Each sorus consists of30–35 round sporangia apparently attached to thecentral receptacle. The largest well�developed sporan�gia located in marginal parts of sori are on average150 µm long.

R e m a r k s. Fertile fern leaves of this species wereinitially determined as Discopteris sp. (Vladimirovich,1985); however, the study of additional material (seeNaugolnykh, 2007, p. 127, text�fig. 53; and a photo�graph of another specimen attributed to the same spe�cies see here, Pl. 30, fig. 2) has shown that it should bereferred to the genus Myriotheca.

M a t e r i a l. Two specimens from the Chekarda 1locality.

Genus Pecopteris Brongniart, 1825Pecopteris anthriscifolia (Goeppert, 1845) Zalessky, 1918, emend. nov.

Plate 25, figs. 1–5; Plate 26, figs. 1–6; Plate 27, figs. 2 and 4; Plate 28, figs. 1–5; Plate 30, fig. 1

For synonymy, see Fefilova, 1973, p. 66.

H o l o t y p e. Figured by Goeppert (1845, pl. 28,fig. 9), reproduced by Neuburg (1948, pl. XIX, fig. 4)


Figs. 1–6. Sterile fern leaves from the Lower Permian beds of the Fore�Urals: (1–5) Pecopteris anthriscifolia (Goeppert) Zalessky:(1) specimen GIN, no. 4856/294, terminal part of a last order pinna with well�preserved venation; (2) specimen GIN, no. 4856/295,imprint of a last order pinna; (3) specimen GIN, no. 4856/293, last order pinna with partially preserved compressed tissues; (4) spec�imen GIN, no. 3773(11)/415(96), middle frond part, with preserved bases of a last order pinnae; (5) specimen GIN,no. 3773(11)/190(91), middle part of a bipinnate leaf; (6) Ovopteris sp., specimen GIN, no. 3773(11)/288(91), last order pinna.Locality: Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10; Lower Permian, Kungurian Stage. Scale bars, 1 cm.



2

Plate 25

3 4

1

5 6

1444


NAUGOLNYKH

(a) (b) (c)

(d)

(e)

(f) (g)

(h)

(i)

(k) (l)

(j)

(m)

and Radczenko (1955, pl. IX, fig. 1); Permian of theKuznetsk Basin.

D i a g n o s i s. Fronds tripinnate. Last order pin�nae alternating, attached to penultimate pinna rachisat 65°–70°. Pinnules attached to last order pinnarachis at 65°–75°. Pinnules pecopteroid, lobate, fusedby their bases, with acute apices. Venation pinnate,

midvein reaching pinnule apex. Lateral veins dichoto�mizing; their acroscopic branches dichotomizing onemore time.

D e s c r i p t i o n (Figs. 27, 28, 29a–29e). Rachisesof the penultimate order pinnae are well�developed,with a shallow, hardly discernible longitudinal adaxialgroove. Rachises of the penultimate order pinnae



Fig. 26. Leaf morphology in species of the genus Prynadaeopteris Radczenko according to the initial understanding of this genus(after Radczenko, 1955): (a) Prynadaeopteris sibirica (Zalessky) Radczenko; (b, h) P. dymovii Radczenko; (c, f) P. karpovii Rad�czenko, (c) holotype; (d) P. tunguscana (Schmalhausen) Radczenko; (e, g, j) P. maneichensis (Zalessky) Radczenko; (i) P. irreg�ularis Radczenko; (k, l) P. eurina (Zalessky) Radczenko; and (m) P. anthriscifolia (Goeppert) Radczenko. Localities: KuznetskCoal Basin; Permian: (a) right bank of the Mungat River near the village of Krapivino, Alykaevskaya Formation, BalakhonskayaGroup); (b) Tom’–Usinskii District of the Kuznetsk Basin, left bank of the Tom River, 3 km upstream from the village of Korai,lower part of the Intermediate Formation, Balakhonskaya Group; (c) left bank of the Tom River near the village of Nikol’skoe,downstream from the mouth of the Verkhnyaya Ters’ River, Tailuganskaya Formation, Erunakovskaya Subformation, Kol’chu�ginskaya Group; (d) Tom’–Usinskii District of the Kuznetsk Basin, right bank of the Usu River, downstream from the NizhnyayaRiver Lower, middle part of the Intermediate Formation, Balakhonskaya Group; (e, j) Tom’–Usinskii District of the KuznetskBasin, right bank of the Usu River, Chelbakh�Kaya locality, downstream from the Nizhnyaya River, middle part of the Interme�diate Formation, Balakhonskaya Group; (f) Uskatskii District of the Kuznetsk Basin, Sokolovskoe Coal Field, ErunakovskayaGroup; (g) right bank of the Tom River upstream from the Poryvaiskaya mine, lower part of the Ishanovskaya Formation, Bala�khonskaya Group; (h) Tom’–Usinskii District of the Kuznetsk Basin, left bank of the Tom River, 2 km upstream from the villageof Korai, lower part of the Intermediate Formation, Balakhonskaya Group; (i) Shushtulepskoe Coal Field, southern slope ofKarachiyakskaya Mountain, Kemerovo Formation, Balakhonskaya Group; (k, l) Zav’yalovskoe Coal Field, Alykaevskaya For�mation, Balakhonskaya Group; and (m) left bank of the Inya River upstream from the village of Erunakovo, Erunakovo Group.Scale bar, 1 cm.

sometimes have a limb, a narrow bordering band(Pl. 26, fig. 4); however, they are more frequentlynaked (Pl. 26, fig. 5; Pl. 27, fig. 4).

The most completely preserved penultimate orderpinna (in my collection) is 19 cm long and 6 cm wide;however, judging from available fragments, the penul�timate order pinnae of this fern species could havebeen much larger. The fronds of tripinnate design wereprobably more than 50–60 cm long. The proportionsof the penultimate order pinnae apparently varydepending on the position of particular pinnae on thefrond rachis. In many extant ferns (for example, Pte�ridium aquilinum (L.) Kuhn et Decken), the basalpenultimate order pinnae are relatively wider than api�cal ones.

Rachises of the last order pinnae are attached inalternating order to the rachis of the penultimate orderpinna at an angle of 65°–70°. Rachises of the last orderpinnae usually distinctly curve towards the apex of thepenultimate order pinna. The rachis of the last orderpinnae is usually half as wide as that of the penultimateorder pinna. The maximum observable width of therachis of the last order pinna is 1.5 mm. Rachises of thelast order pinnae always have a well�developed limb,which is particularly distinct in specimens with rela�tively widely placed pinnules (Pl. 26, fig. 3; Figs. 29c,29e). The limb is usually as wide as the rachis of the lastorder pinna; thus, as the axial part of the rachis is1 mm wide, the rachis with the bordering limb is 3 mmwide (limb extends for 1 mm on each side of therachis). Well�developed last order pinnae are usuallyabout 8–10 cm long and 2–2.5 cm wide.

Pinnules are attached in alternating order to rachisesof the last order pinnae at an angle of 65°–75°. They arepecopteroid, with a lobed margin. The base of basalpinnules is sometimes slightly constricted, so thatthese pinnules become more similar in morphology tosphenopteroid pinnules. The most developed largepinnules are on average 8 × 16 mm. Pinnules graduallydecrease in size towards the apex of the last order

pinna. The angle of attachment of pinnules to therachis decreases in the same direction. One well�developed last order pinna has on average 12–14 pairsof pinnules.

Venation is pinnate. Lateral veins diverge in alter�nating order from a well�pronounced midvein, whichis usually slightly undulating and extends to the pin�nule apex. Each pinnule usually contains four or fivepairs of lateral veins. All lateral veins dichotomize; theacroscopic branch produced by this dichotomy bifur�cates one more time. In the basal lateral veins, thebasiscopic branch also sometimes bifurcates. Threeends of lateral veins usually enter each pinnule lobe,the middle end passes directly into the lobe apex.

In one specimen, the leaf lamina is partially miner�alized, allowing the study of the epidermal–cuticularstructure of the abaxial side of a sterile leaf (Pl. 28, figs.1–5; Figs. 28b, 28c, 28e). Common cells of the epi�dermis extend along the pinnule midvein. They areelongated trapezoidal or elongated polygonal (usuallyhexagonal and heptagonal) in outline and on average40 × 90 µm in dimensions. The stomata are simple,lack modified subsidery cells, unsunken. The stomatalapertures are narrow, slitlike, turned in the same direc�tion as common cells of the epidermis. Guard cellshave liplike thickenings (Pl. 28, fig. 5; Fig. 28e).

Unfortunately, available collections do not containfertile leaves to compare with sterile leaves of Pecopt�eris anthriscifolia. Therefore, following Zalessky, thisspecies is assigned to the formal genus Pecopteris.

C o m p a r i s o n. P. anthriscifolia is most similar tothe other pecopterid species P. leptophylla Bunbury,which is characteristic primarily of Lower Permianfloras of Western and Central Europe and occurs in theLower Permian of western Angaraland (Subangara�land Ecotonal Realm), in the Artinskian and Kun�gurian beds (Vladimirovich, 1986; Naugolnykh,1998c), and in open nomenclature (P. ex gr. Lepto�phylla) in the Kazanian beds (Ignatiev and Naugol�nykh, 2001; Naugolnykh and Kuleshov, 2005). It wasalso proposed that the species P. anthriscifolia and

1446


NAUGOLNYKH

P. leptophylla are synonyms (for discussion, see Neu�burg, 1948, p. 118). Judging from the initial figure ofthe type specimens of P. leptophylla (Ribeiro, 1853,

pl. VII, figs. 11a, 11b; reproduced by Wagner andLemos de Sousa, 1985), the main differences of P. lep�tophylla from P. anthriscifolia include the narrower

(a)

(b)

(c)

Fig. 27. Pecopteris anthriscifolia (Goeppert) Zalessky, leaf structure after Zalessky and Tschirkova (1938): (a, b) Pecopterisanthriscifolia (Goeppert) Zalessky f. petschorensis Zalessky (Zalessky and Tschirkova, 1938, text�figs. 38, 39, respectively);(c) Pecopteris anthriscifolia (Goeppert) Zalessky f. vorcutensis Zalessky (Zalessky and Tschirkova, 1938, text�fig. 41). Localities:Pechora Coal Basin, Vorkuta River: (a) Rudnitsky settlement; (b) Vorkuta mines, coal�bearing strata; and (c) Vorkuta mines, nearCoal Bed 8 (outcrop 36, T. Ponomarev). Scale bar, 1 cm.



1 2 3

5

4

6

Plate 26


Figs. 1–6. Sterile leaves of Pecopteris anthriscifolia (Goeppert) Zalessky: (1) specimen GIN, no. 4856/295; (2) specimen GIN,no. 4851/184; (3) specimen GIN, no. 4856/144; (4) specimen GIN, no. 4856/192; and (5, 6) specimen GIN,no. 3773(11)/4(89). Localities: (1, 3–6) Middle Fore�Urals, Perm Region, Chekarda 1, Bed 10; Lower Permian, KungurianStage; (2) Pechora Coal Basin, borehole IK�657, 864.8 m of depth; Middle Permian, Kazanian Stage. Scale bars, 1 cm.

1448


NAUGOLNYKH

1

2 3

4

Plate 27



proportions of pinnules, the stronger fusion betweentheir margins (usually for at least half of the pinnulelength), the less developed lobes, and the simplervenation, which are characteristic of P. leptophylla. Inthis connection, the identification of fern in the study

of Vladimirovich (1986, pl. 138, fig. 1) is doubtful;contrary to the opinion of Vladimirovich, it should bereferred to P. anthriscifolia rather than P. leptophylla.

R e m a r k s. This fern species is apparently mostfrequently mentioned in the paleobotanic literature

(a) (b)

(c)(d)

(e)

LV

LV MV

GC

LV

LV

Fig. 28. Leaf morphology of Pecopteris anthriscifolia (Goeppert) Zalessky: (a, d) macromorphology and (b, c, e) epidermal–cuticular leaf structure: (a) middle part of the last order pinna with two lobate pinnules with simple once or dichotomizing lateralveins (dotted line shows orientation of the longitudinal axes of extended common epidermal cells; (b) epidermal structure withcommon epidermal cells extending along lateral veins; (c) epidermal topography, in distinct costal fields, cover cells are moreelongated and, in the intercostal fields, cells are more isometric: (MV) pinnule midvein and (LV) pinnule lateral veins; (d) middlepart of the penultimate order pinna with two almost completely preserved last order pinnae; and (e) structure of two closely posi�tioned stomata: (GC) guard cells); (a–c, e) specimen GIN, no. 4851/218 and (d) specimen GIN, no. 3737/116. Localities:Pechora Coal Basin, Vorkuta 1 locality, right bank of the Vorkuta River, outcrop 37; Lower Permian, Kungurian Stage (a–c, e);Chekarda 1, Bed 10; Middle Fore�Urals (d). Scale bars: (a) 1 mm, (b) 50 µm, (c) 200 µm, (d) 2 mm, and (e) 10 µm.


Figs. 1–4. Sterile leaves of (2, 4) Pecopteris anthriscifolia (Goeppert) Zalessky compared to that of the extant leptosporangiateferns (1) Athyrium filix�femina L. and (3) Dryopteris filix�mas L.: (1) middle (on the left) and apical parts of the penultimate orderpinna from my private herbarium; (2) specimen GIN, no. 4856/221, juvenile pinna curved in the shape of a “bischof crozier”;(3) GIN, no. P5 (S.V. Naugolnykh’s herbarium), two juvenile bipinnate fronds with the apices twisted in the shape of a helix toprotect the meristematic zones; (4) specimen GIN, no. 4851/185, middle part of a penultimate order pinna. Localities: (2) Mid�dle Fore�Urals, Perm Region, Chekarda 1, Bed 10; Lower Permian, Kungurian Stage; (4) Pechora Coal Basin, Er�Yaga River,outcrop T�Ya 109; Middle Permian, Kazanian Stage; (1) Moscow Region, forest near Lake Hydra, Ramenskoe; Recent; (3) Mos�cow, Bitsevskii Park; Recent. Scale bars, 1 cm.

1450


NAUGOLNYKH

devoted to Permian floras of Angaraland and adjacentregions of the Subangaraland Ecotonal Belt. The tax�onomy and nomenclature of this fern raise moreheated debates among paleobotanists than any otherLate Paleozoic ferns of Angaraland.

Comparative material that I used for the descrip�tion of this species includes sterile fern leaves referredto P. anthriscifolia, which were collected in the south�ern, middle, and Pechora Fore�Urals and in thePechora Coal Basin (see the section Material). Inaddition to isolated last order pinnae (Pl. 25, figs. 2, 3;Pl. 26, figs. 2, 3) and variously preserved bipinnatefronds (penultimate order pinnae: Pl. 25, figs. 4, 5;Pl. 26, figs. 4–6), the collection includes tripinnatefrond fragments (Pl. 27, fig. 4), which strongly suggestthat leaves of this species were at least tripinnate. Thisstatement is supported by leaves of P. anthriscifoliafrom the Permian of the Pechora Coal Basin, whichwere previously described by Zalessky and Tschirkova

(1938; see here Fig. 27). The tripinnate design ofleaves of P. anthriscifolia from the Permian of the Kuz�netsk Basin was noted by Neuburg (1948). Amongleaves of this species from the Permian of the PechoraBasin which were later figured by Fefilova, tripinnatefronds are absent; however, the tripinnate design ofleaves of P. anthriscifolia was marked in the descriptionand included in the diagnosis as a diagnostic character(Fefilova, 1973, p. 66).

In Fore�Ural Permian floras, fern leaves assignedto the species Pecopteris anthriscifolia (if only sterileleaves are found) are particularly frequent. In the casethat similar leaves with fertile organs are found, theyare usually determined as Prynadaeopteris anthriscifo�lia (Goeppert) Radczenko. Structural details of fertileorgans of Pecopteris (Prynadaeopteris) anthriscifolia(Goeppert) Zalessky required for the establishment ofthe taxonomic position of this fern remain uncertain.This important question is a task of future studies of

(a)

(b)

(c)

(d) (e)

(f)

Fig. 29. Sterile leaf morphology: (a–e) Pecopteris anthriscifolia (Goeppert) Zalessky in comparison with the morphology of(f) isolated sterile pinna of the marattialean fern Pecopteris uralica Zalessky: (a) specimen GIN, no. 4856/221; (b) specimen GIN,no. 4856/220; (c) specimen GIN, no. 4856/229; (d) specimen GIN, no. 4856/147; (e) specimen GIN, no. 4856/144; and(f) specimen GIN, no. 4856/142. Middle Fore�Urals, Perm Region, Chekarda 1 locality, Bed 10; Lower Permian, KungurianStage. Scale bar, 1 cm.



(a)

(b)

(c) (d)

(e) (f) (g)

(h)

Fig. 30. Epidermal–cuticular and macromorphological leaf structure in ferns from the Iva�Gora locality (Soyana River, Arkhan�gelsk Region): (a–g) Pecopteris ex gr. leptophylla Bunbury: (a) stoma; (b) zone of contact between the leaf lamina epidermis (cos�tal field; on the lower right) and epidermis located above the pinnule midvein (on the upper left); (c) costal field epidermis;(d) zone of contact between the intercostal (on the left) and costal (on the right) fields; (a, b, d) preparations of specimen GIN,no. 4547/50; (e) specimen GIN, no. 4547/50, terminal last order pinna with coalescently fused pinnules (lower part of this spec�imen was used for preparations shown in a–d; (f) specimen GIN, no. 4547/28, well�developed last order pinna; (g) specimenGIN, no. 4547/29, middle part of pinna; and (h) Lobatopteris sp., specimen GIN, no. 4547/30, middle frond part with bases ofthe last order pinnae. Iva�Gora locality, Soyana River, Arkhangelsk Region; Middle Permian, Kazanian Stage. Scale bars:(a) 40 µm, (b) 200 µm, (c, d) 100 µm, and (e–h) 2 mm.

Permian ferns of the Fore�Urals. To date, it is onlyknown that fertile organs of this fern are sori (Radc�zenko, 1955), which resemble somewhat in structurethe sori of Oligocarpia, a fern genus that frequentlyoccurs in the Upper Carboniferous (Stephanian) andLower Permian (Autunian) of Western and CentralEurope (Stur, 1883; Grauvogel�Stamm and Doubin�ger, 1975; Wagner and Lemos de Sousa, 1983; Bartheland Rossler, 1995; Psenicka and Bek, 2001; Barthel,2005), North America (Zodrow and McCandlish,1982), China (Wang et al., 1999; Wang and Wu, 1999),and South America (Cesari et al., 1998). Remains ofOligocarpia are particularly typical for the areas wherelow�latitude termophilic vegetation characterized byfloras of the Euramerian type occurred at the end ofthe Paleozoic. The genus Oligocarpia is a typical repre�sentative of leptosporangiate gleicheniacean ferns

(family Gleicheniaceae). If the similarity betweenPrynadaeopteris anthriscifolia and some species of Oli�gocarpia does not result from convergent develop�ment, it is possible to assume that P. anthriscifolia(along with other members of the genus Prynadaeopt�eris) is close to the family Gleicheniaceae and, thus,belongs to leptosporangiate ferns.

M a t e r i a l. Fifteen specimens represented by iso�lated pinnules and fragments of the last and penulti�mate order pinnae, one complete penultimate orderpinna, and a tripinnate frond fragment from the Lowerand Middle Permian of the Fore�Urals.

O c c u r r e n c e. Permian of Angaraland and theSubangaraland Ecotonal Belt; Artinskian, Kungurian,Ufimian, and Kazanian stages of the Fore�Urals,Pechora Coal Basin, Siberia, Mongolia, and, proba�bly, northern China.

1452


NAUGOLNYKH

1

2

3

4 5

Plate 28


Figs. 1–5. Leaf morphology of Pecopteris anthriscifolia (Goeppert) Zalessky, specimen GIN, no. 4851/218: (1) macromorpho�logical structure of two neighboring pinnules; (2–5) epidermal–cuticular leaf structure: (1) middle part of the last order pinnawith two lobate pinnules with simple or once dichotomizing lateral veins: (2, 3) epidermis with common cells extending alonglateral veins; (4) pinnule fragment with distinctly seen veins; and (5) two neighboring stomata. Right bank of the Vorkuta River,Vorkuta 1 locality, outcrop 37; Lower Permian, Kungurian Stage. Scale bars: (1) 1 mm, (2, 3) 50 µm, (4) 200 µm, and (5) 10 µm.



Pecopteris ex gr. leptophylla Bunbury in Ribeiro, 1853


Pecopteris attenta Zalessky: Zalessky, 1937a, p. 92, text�fig. 62.Pecopteris spiculosa Zalessky: Zalessky, 1937a, pp. 92–93, text�

fig. 63.Pecopteris conserrata Zalessky: Zalessky, 1937a, p. 93, text�

fig. 64.Pecopteris ex gr. leptophylla Bunbury: Ignatiev and Naugol�

nykh, 2001, p. 64, pl. I, figs. 2a and 2b, text�figs. 3a–3d.

D e s c r i p t i o n (Figs. 30a–30g). Judging fromthe maximum observable width of the penultimateorder pinnae (4 cm), the total leaf length could havebeen several tens of centimeters. The rachis of the pen�ultimate order pinnae is thick, well�developed. It is upto 1.5–2 mm wide. Rachises of the last order pinnaeare attached to the rachis of the penultimate order pin�nae at an angle of 30°–35°. Pinnae are usually subtri�angular in outline (Pl. 29, figs. 1, 4–7; Fig. 30f); how�ever, the collection contains a frond fragment with thelast order pinnae with subparallel margins (Pl. 29,fig. 2). The largest and most developed last order pinnais 25 mm long and 8 mm wide. Pinnules are attachedto rachises of the last order pinnae, coalescently fusedby the margins usually for at least half of their length.In the apical part of the penultimate order pinnae,poorly developed last order pinnae presumablybecome more similar in morphology to usual pinnules;however, in contrast to standard pinnules, theseunderdeveloped last order pinnae show a more com�plex venation (compare a juvenile last order pinna(Fig. 30e) with a pinnule of a well�developed last orderpinna (Fig. 30f)).

Pinnules are pecopteroid, distinctly subtriangularin outline, with a pointed apex and lobate margins.The midvein is well pronounced, undulating, closelyapproaches the pinnule apex. Lateral veins diverge inthe alternating order from the midvein at an angle of30°. Well�developed pinnules positioned at the base ofthe last order pinna usually contain three pairs of lat�eral veins. Apical pinnules have up to one pair of lat�eral veins. Lateral veins are simple, usually slightlycurved towards the pinnule apex. Each marginal lobeof pinnules contains one lateral vein, which is some�times displaced towards the acroscopic (anterior) lobemargin.

C o m p a r i s o n. See description of the speciesPecopteris anthriscifolia.

R e m a r k s. The structure of sterile fern leaves ofthis morphotype fits well in the morphological charac�teristics of the species Pecopteris leptophylla; however,since available specimens are rather fragmentary, thisform is described in open nomenclature as <ex gr.>.

M a t e r i a l. Six fragments of pinnae of the last andpenultimate orders.

Genus Lobatopteris Wagner, 1958Lobatopteris sp.

Lobatopteris sp.: Ignatiev and Naugolnykh, 2001, pp. 64, 67,pl. II, 4; fig. 2.

D e s c r i p t i o n (Fig. 30h). The rachis of the pen�ultimate order pinna is well�developed, thick, massive,5 mm wide, with a narrow longitudinal central grooveon the adaxial side of the rachis. Rachises of the lastorder pinnae diverge from the rachis of the penulti�mate order pinna at an angle of 50°. The last order pin�nae are long, with subparallel margins.

The total length of the last order pinnae is notknown. Judging from the available fragment and rateof decrease in pinnule length towards the apex of thelast order pinna, it was probably 12–13 cm long. Themaximum observable width of the last order pinnae is17 mm. Pinnules are attached to the rachis of the lastorder pinna, coalescently fused by the margins forthree�fourths of their length. Pinnules are relativelyshort, isometric in outline, with even margins androunded apex. The pinnule midveins unusually divergefrom the rachis of the last order pinna at an angle thatis uncommonly acute for pecopterids, ranging from25° to 35°; however, close to the point of divergencefrom the rachis of the last order pinna, the midveincurves considerably posteriorly and, then, passes alongthe longitudinal pinnule axis at an angle of 45°–50°relative to the rachis of the last order pinna, reachingthe pinnule apex margin. From the base of the pinnulemidvein, the basiscopic lateral vein is the first todiverge; it dichotomizes once or twice, or, less fre�quently, remains simple. Then, the midvein gives riseto the acroscopic lateral vein, which is usually simple,or dichotomizes once, and, then, two or three pairs ofthe lateral veins diverge in the alternating order andusually dichotomize once; less frequently, they aresimple. The pinnule margins are complicated by finerugosity, which shows the same direction as the veins.

R e m a r k s. Similar remains of a fern leaf from theInta Formation, which were also initially determinedas Lobatopteris sp., were reported in the study devotedto the paleobotanic characteristics of Permian floras ofthe Pechora Basin (Pukhonto and Fefilova, 1983,pl. IX, figs. 1, 1a). Subsequently, this specimen wasregarded as a new species, Lobatopteris vorcutensisPukhonto (in Naugolnykh and Pukhonto, 2009, pl. V,figs. 1, 4). Unfortunately, since fine venation featuresof the form from the Pechora Basin remain uncertain,so that it is presently difficult to compare it with theexternally similar fern Lobatopteris sp. from the Soy�ana Flora.

M a t e r i a l. One fragment of the middle part of abipinnate frond.

RELATIONSHIPS BETWEEN THE EVOLUTION OF FERNS

AND CLIMATIC AND GEODYNAMIC EVENTS

A decrease in the proportion of Marattiaceae andsimultaneous appearance of some groups traditionallyconsidered to be typical Mesozoic (Osmundaceae) inthe Permian of the Fore�Urals was accounted for by

1454


NAUGOLNYKH

2 3 4

5 6 7

1

Plate 29



both the evolutionarily morphogenetic processes andregional climatic changes caused by the geodynamicprocesses (primarily by the raise of the Paleourals andaccomplishment of the last phase of the Hercynianorogeny) and the general global trend towards anincrease in aridity, which occurred in the latter half ofthe Permian and beginning of the Triassic.

In the modern flora of the Northern Hemisphere,the Marattiaceae do not grow north of 30° N and, inthe Southern Hemisphere, they are also confined tothe regions of very warm and humid climate and frostfree winters (New Zealand, Australia). Based on thesedata, it is proposed that, in the geological past, theMarattiaceae showed the same ecological climaticpreferences, therefore, this fern group comprises indextaxa suitable for paleoclimatic reconstruction. In thePermian Period, the Paleourals occupied a sublatitu�dinal position. Ferns of the family Marattiaceae havenot been recorded in Permian floras of Siberia, sug�gesting that the Ural Mountains were just the northernboundary between the tropical (along with subtropi�cal) and boreal climatic zones in the Permian Period(for more detail, see Naugolnykh, 2004).

Abundant Marattiaceae are known from the UpperCarboniferous and Permian of Cathaysia (Asamaet al., 1968; Kon’no et al., 1971; Wan and Basinger,1992; Wang and Yang, 1996; Yang et al., 1996) andEuramerica (Jennings and Millay, 1979; Mapes andSchabilion, 1979a, 1979b; Stubblefield, 1984; Lesni�kowska and Millay, 1985; Lesnikowska and Galtier,1992; Lesnikowska and Willard, 1997; Millay, 1997;Barthel, 2005). Localities of Cathaysian and Eurame�rian Marattiaceae, which are frequently representedby the same or very similar genera, form a wide paleo�sublatitudinally extended band, which corresponds tothe position of equatorial and two tropical belts in theLate Paleozoic.

In the context of discussion concerning the signifi�cance of Late Paleozoic ferns, the taxonomic compo�sition of the Permian Godare�Gachal Flora (CentralIran), containing Euramerian and Cathaysian ferns, isindicative.

In its taxonomic composition, the Early PermianGodare�Gachal Flora (Leven et al., 2011) is a typicalmixed flora combining Euramerian (Annularia cari�nata Gutbier, Pecopteris monyi Zeiller, and P. cyathea(Schlotheim) Brongniart) and Cathaysian (Fascipteriscf. robusta (Kawasaki) Broutin and Taeniopteris cf.crassicaulis Jongmans et Gothan) components. Thegenus Discinites Feistmantel, which is present in the

Godare�Gachal Flora, occurs in both Euramerica andCathaysia.

Many researchers noted that, in the Late Paleo�zoic, the peripheral regions of Euramerica andCathaysia were apparently connected by migrationgateways (Laveine et al., 1989; Wang, 1996; etc.). Inthe Carboniferous Period, the two paleofloristicrealms showed a great similarity, so that the unitedAmerosinian Equatorial–Tropical Belt is often recog�nized at that period (Wagner, 1962). In the PermianPeriod, particularly in the Late Permian, the propor�tion of endemics in Euramerian and Cathaysian florasincreased considerably (Meyen, 1987); however, at thebeginning of the Permian, plants could migrate east�erly from Paleo�Europe along the northern coast ofthe Tethys. These migration pathways apparentlypassed through the area of the development of theGodare�Gachal Flora, which is supported by the glo�bal palinspastic reconstructions for the Early Permian.

According to the scheme of paleofloristic zonation(paleophytogeographic zonation based on the floralprinciple), the Godare�Gachal Flora is situated withinthe boundary band between Euramerian–“Atlantic”floras on the southwest and Subangaraland floras onthe northeast (Meyen, 1987). In the map showing dis�tribution of plants belonging to different paleophyto�geographic units at the end of the Early Permian andbeginning of the Late Permian, (Meyen, 1987, p. 323,fig. 76), the Khazro and Ga’ara localities as well as theterritory of Iran which has yielded the Godare�GachalFlora are referred to the Cathaysian Realm, which isbordered according to this scheme by the “Atlantic”(Euramerian) Empire without a transition. This isunderstandable through the other paleophytogeo�graphic scheme provided by Meyen (1987, p. 315,text�fig. 75b), which distinctly shows that the Eurame�rian Realm and Cathaysian Realm are separated by anextensive oceanic basin.

However, available data contradict these schemes.Even the taxonomic composition of the Khazro andGa’ara floras shows a mixture of Euramerian andCathaysian elements and, in the Godare�GachalFlora, Euramerian elements even prevail. Thisstrongly suggests that, during the first half of the Per�mian, the Euramerican and Cathaysian floras werelinked by certain migration gateways.

Some formalities of the paleophytogeographicschemes based on the floral principle, that is, compar�ative analysis of the taxonomic composition of con�


Figs. 1–7. Leaf morphology of Pecopteris ex gr. leptophylla Bunbury: (1, 4, 6) specimen GIN, no. 4851/187, apical part of a younglast order pinna, photograph taken in (1) neutral, (4) lateral slanting, and (6) frontal light; (2) specimen GIN, no. 54851/186,middle part of a bipinnate frond with two preserved last order pinnae; (3) specimen GIN, no. 4851/188, middle part of a younglast order pinna; (5) specimen GIN, no. 4851/189, terminal part of a last order pinna; (7) specimen GIN, no. 4851/190, coun�terpart of specimen GIN, no. 4851/189. Arkhangelsk Region, Soyana River, Iva�Gora locality; Middle Permian, Kazanian Stage.Scale bars, 1 cm.

1456


NAUGOLNYKH

1

2

3

Plate 30


Figs. 1–3. Leaf morphology in pecopterids from the Permian of (1, 3) the Kuznetsk Coal Basin and (2) Fore�Urals near Perm:(1, 3) Pecopteris anthriscifolia (Goeppert) Zalessky, originals: (1) TsNIGR Museum, no. 59/573 (the same specimen was figuredby Zalessky, pl. XXXIII, fig. 2); (3) TsNIGR Museum, no. 168/573 (the same specimen was figured by Zalessky, 1918, pl. XXXV,figs. 1, 1a); (2) Myriotheca cf. scaberrima Lesquereux, specimen GIN, no. 4856/218. Localities: (1, 3) Kuznetsk Basin; LowerPermian, Kungurian Stage: (1) right bank of the Inya River, upstream from the village of Khmelevaya; (3) Inya River, village ofMeretskaya; (2) Middle Fore�Urals, Perm Region, Chekarda 1, Bed 7; Lower Permian, Kungurian Stage. Scale bars, 1 cm.



temporaneous floras was repeatedly marked in the lit�erature (for more detail, see Naugolnykh, 2005a).

To produce an alternative scheme of paleophyto�geographic zonation of the Earth in the PermianPeriod, I proposed to use primarily physiognomicalcharacters of vegetation rather than the taxonomiccomposition of floras (Naugolnykh, 2005a, 2007).The following main biomes have been recognized andbased on vegetation types and mapped: (1) impover�ished boreal vegetation of the tundra or forest–tundratypes; (2) typical Angaraland deciduous vegetation(Permian analogue of modern boreal forests, the so�called Angaraland taiga); (3) tropical and equatorialdroughty semiarid zones, with vegetation ecologicallysimilar to the present day the Mediterranean vegeta�tion type; (4) equatorial and tropical evergreen vegeta�tion (Permian analogue of modern rain forests);(5) xerophilous vegetation of drainless inland depres�sions; (6) biome of the zone of mixed vegetation of theMediterranean ecological physiognomical type,which is characterized by the Euramerian floral type,and notal vegetation of the southern moderate andcold moderate biome; and (7) notal deciduous vegeta�tion dominated by glossopterids (dictyopteridians).

The Godare�Gachal Flora is considered to belongto Early Permian vegetation of the Mediterraneantype, which grew on both northern and southerncoasts of the Paleotethys. However, the absence in thisflora of xerophilous forms, such as callipterids andwalchian conifers combined with the predominance ofrelatively hydrophilous pecopterids are evidence ofmore humid conditions in this region, which, in turn,could have been caused by relatively high humidity ofthe biotopes directly adjacent to the marine coasts.More favorable conditions in this region compared todroughty semiarid winter�wet climate of westerly areasof Euramerica probably promoted expansion of someEuramerian plants along the northern coast of theTethys to northern Cathaysia and provided penetra�tion of migrator plants from Cathaysia into this region.

Mixed floras containing Euramerian and Cathay�sian elements occur almost throughout the northernperiphery of the Peritethys, including southern Spain(Guadalcanal Flora: Broutin, 1974, 1977, 1985,1986), Turkey (Khazro Flora: Wagner, 1962; Archan�gelsky and Wagner, 1983), Saudi Arabia (UnayzakhFlora: El�Khayal et al., 1980; Lemoigne, 1981; El�Khayal and Wagner, 1983, Hill et al., 1985), Iraq(Ga’ara Flora: Ctyroky, 1973), and Afghanistan(Meyen, 2002). In almost all of these floras, Cathay�sian elements are represented by spore�bearing plants,the fern Fascipteris Dong and the sphenophyte (possi�bly calamite) Lobatannularia Kawasaki. Due to dis�tinctive reproductive features of spore�bearing plants,which require moisture in the reproductive season totransfer reproductive products, develop the gameto�phyte, and germinate spores, it is evident that, within

the entire relatively narrow seaside band, the climatewas at least seasonal humid or semihumid.

The fact that plants migrated from Euramerica toCathaysia was explained by the effect of monsoons(Wang, 1996); however, this may be treated essentiallydifferently. Since in the Permian, the entire easternTethys was extremely shallow and contained a chain ofislands and microcontinents, which extended from thesouth to the north and facilitated migration of Gond�wanan plants within Cathaysia, the oceanic northerntrade wind current should turn to the north up to lati�tude 50 and, then, deviate westerly. Therefore, theeffect of monsoons was observed much more northerlythan at the present time, mostly in northern Cathaysiaand southern Angaraland (where coal�bearing strataintensely accumulated in the Permian Period). How�ever, the effect of monsoons abruptly weakened furtherwesterly and completely disappeared, because theystroke against the Paleo�Urals, which were rather highat the end of the Paleozoic. In the region of the mon�soon shadow of the Paleo�Urals, which covered theentire Fore�Urals, Russian Platform, and a large partof Western Europe, the climate gradually became drierbecause of a global increase in aridity intensifiedlocally by the monsoon shadow, which provoked to agreat extent northeastern migration of hygrophilousplants along the northern coast of the Tethys into theregions with a more humid climate.

SIGNIFICANCE OF FERNS FOR STRATIGRAPHY

OF THE PERMIAN STRATA

Remains of Permian ferns of the Fore�Urals areextremely poorly used in stratigraphy. Fern species areat best recorded along with other fossil plants to pro�vide general characteristics of floral assemblages.However, in my opinion, the high evolutionary rate,wide and rather rapid dispersal of this plant groupwithin the region under study allow the use of fern spe�cies (or at least some of them) for stratigraphy.

I proposed a scheme of vertical distribution of fernremains in the Kozhim section from the Sakmarian toUfimian inclusive (Naugolnykh in Grunt et al., 1998,p. 208, text�fig. 48). The Sakmarian beds have yieldedPecopteris aff. denticulata�cristata Brongniart, P. aff.alata Brongniart, and P. aff. ripensis Zalessky; theArtinskian beds have Pecopteris sp. and Sphenopterissp.; the Kungurian beds have Pecopteris (? Asterotheca)kojimensis Zalessky and P. cf. borealis Zalessky; andthe Ufimian beds have Corsinopteris semilibera andPrynadaeopteris sp. Notwithstanding the fact thatmany of the above listed forms are determined in opennomenclature, it is evident that each fern assemblagecorresponding to a particular stage of the PermianSystem within the Kozhim section displays a distinc�tive stratigraphic pattern. Preliminary data on the dis�tribution of ferns in the Permian beds of other regionsof the Fore�Urals suggest that, in the future, it will be

1458


NAUGOLNYKH

possible to recognize for them sequences of fernassemblages, a fern zonal scale of a sort.

CONCLUSIONS

To resolve the questions related to taxonomy ofPermian ferns of the Fore�Urals, particularly at thespecies and generic levels, and establish the taxonomicposition of species with known (at present or in thefuture) fertile organs, it is necessary to use newmethodical approaches developed in modern paleo�botany. The importance of studying variation in bothsterile and fertile leaves has already been markedabove. Combined use of light and electron scanningmicroscopy for the study not only of the synangia andsporangia, but also spores preserved in situ and ana�tomical characters of leaves and mineralized stemsseems rather promising. The records of morphologi�cally complete fertile organs of ferns in the Lower Per�mian beds of the middle Fore�Urals, particularly car�bonate nodules of the Urminskaya Formation (Sargin�skii Horizon of the Artinskian Stage), look ratherencouraging. Thorough examination of these remainsis an interesting topic of future studies.

To date, it has been shown with certainty that thePermian beds of the Fore�Urals enclose remains ofeusporangiate, protoleptosporangiate, and leptospo�rangiate ferns, which belong to both previously estab�lished genera and species and new taxa (species andgenera), some of which are described in the presentpaper. The specificity of fern assemblages characteriz�ing different stages of the Permian System in the Fore�Urals and eastern Russian Platform suggests that thestudy of these plants will provide a means for widepaleogeographical reconstructions and stratigraphicpurposes.

ACKNOWLEDGMENTS

I am grateful to all colleagues who provided mewith specimens for studying, to G.N. Aleksandrova(Geological Institute, Russian Academy of Sciences)for help in maceration of fertile fern leaves and discus�sions concerning the taxonomy of disperse fern spores,to S.A. Inozemtsev (Faculty of Soil Science, MoscowState University) for an opportunity to use the equip�ment of the Laboratory of Electron Microscopy of theFaculty of Soil Science of Moscow State University.

This study was supported by the Russian Founda�tion for Basic Research, project no. 11�05�92692�a.

REFERENCES

Anderson, H.M. and Anderson, J.M., Molteno Ferns: LateTriassic Biodiversity in Southern Africa, Pretoria: Strelitzia,2008, vol. 21.

Archangelsky, S. and Wagner, R.H., Glossopteris anatolicasp. nov. from uppermost Permian strata in south�east Tur�

key, Bull. Brit. Mus. Nat. Hist. (Geol.), 1983, vol. 37, no. 3,pp. 81–91.Arefiev, M.P. and Naugolnykh, S.V., Isolated roots from theTatarian Stage of the basin of the Sukhona and MalayaNorthern Dvina rivers, Paleontol. Zh., 1998, no. 1, pp. 86–99.Asama, K., Iwai, J., Veeraburas, M., and Honghusonthi, A.,Permian plants from Loei, Thailand: Contributions to theGeology and Paleontology of Southeast Asia: XLV, in Geologyand Palaeontology of Southeast Asia, Tokyo: Tokyo Univ.Press, 1968, vol. 4, pp. 82–98.Balme, B.E., Fossil in situ spores and pollen grains: Anannotated catalogue, Rev. Palaeobot. Palynol., 1995, vol. 87,pp. 81–323.Barthel, M., Die Rotliegendflora des Thuringer Waldes:Teil 3. Farne, Veroff. Naturhist. Mus. Schleusingen., 2005,vol. 20, pp. 27–56.Barthel, M. and Rossler, R. Rotliegend�Farne in weisenVulkan�Aschen�"Tonsteine" der Dohlen�Formation alspalaeobotanische Fundschichten, Veroff. Mus. Naturk.Chemnitz., 1995, vol. 18, pp. 5–24.Bertchtold, F.G. and Presl, J.S., O prirozenosti rostlin anebrostlinar, obsahugjcj popsany a wyobrazenj rostlin podle raduprirozenych zporadane, s zewrybnym wyznamenanjm wlast�nostj, uzitecnosti a skodliwostj, obzwlaste wywodin a zplodin,spusobu wydobywanj poslednjch dobroty a porusenosti ney�gistegsjho poznanj a skausenj, tez spusobu uzitecnych sazenj,chowanj a rozmnozovanj, Praha: J. Krause, 1820.Betekhtina, O.A., Gorelova, S.G., Dryagina, L.L., et al.,Verkhnii paleozoi Angaridy. Fauna i Flora (The Upper Pale�ozoic of Angaraland: Fauna and Flora), Novosibirsk:Nauka, Sib. Otd., 1988.Bromhead, E.F., Remarks on the botanical system of Pro�fessor Perleb, Mag. Natur. Hist. J. Zool., Bot., Mineral.,Geol., Meteorol., 1840, Ser. 2, no. 4, pp. 329–338.Brongniart, A., Sur la classification et la distribution desvégétaux fossiles, Mém. Mus. Hist. Natur., 1822, vol. 8,pp. 203–240.Broutin, J., Sur quelques plantes fossiles du bassin Autun�Stephanien de Guadalcanal (province de Seville, Espagne),Lagascalia, 1974, vol. 4, no. 2, pp. 221–237.Broutin, J., Nouvelles données sur la flore des bassinsAutun–Stephaniens des environs de Guadalcanal (prov�ince de Seville–Espagne), Guad. Geol. Iber., 1977, vol. 4,pp. 91–98.Broutin, J., Ginkgophyllum boureaui, nouvelle espèce deginkgophyta du Permien inférieur du Sud�Ouest del’Espagne, Bull. de la section des sciences, vol. 8: Paléobota�nique recherches nouvelles sur l’évolution végétale, Paris:Com. Trav. Hist. Sci., 1985, pp. 125–132.Broutin, J., Etude paéobotanique et palynologique du passageCarbonifère–Permien dans le Sud–Oest de la PeninsulaIbérique, Paris: Com. Trav. Hist. Sci., 1986.Cesari, S.N., Cuneo, N.R., and Archangelsky, S., Unanueva probable Gleicheniaceae del Permico Inferior dePatagonia, Argentina, Rev. Espan. Paleontol., 1998, vol. 13,no. 1, pp. 81–92.Corda, A.J., Flora Protogaea: Beiträge zur Flora der Vorwelt,Berlin: S. Calvary and Co., 1845.Corsin, P., Bassin houiller de la Lorraine. Memoire publie surl’initiative et avec le concours de Houilleres du Bassin de Lor�raine et de la Regie des Mines de la Sarre, vol. 1: Flore fossile,



fasc. 4: Pecopteridees: Etudes gites mineraux de la France,Lille: Loos�Nord, 1951.Ctyroky, P., Permian flora from the Ga’ara region (westernIraq), Neues Jahrb. Geol. Palaontol. Mon. Krefeld Schweiz.,1973, no. 1, pp. 383–388.Cuneo, N.R., Archangelsky, S., and Cesari, S.N., Asteroth�eca frenguellii (Archangelsky y de la Sota) nov. comb.,helecho permico de Paragonia, Argentina, Ameghiniana(Rev. Asoc. Paleontol. Argent.), 2000, vol. 37, no. 3, pp. 363–367.Danze, J., Role du development ontogenique dans la diag�nose paleobotanique, Ann. Soc. Geol. Nord., 1955, vol. 75,pp. 83–93.Doweld, A.B., Corsinopteris, a new substitute name forOrthotheca (Marattiales–Marattiopsida), Taxon, 2001,vol. 50, pp. 1097–1099.Eichwald, E.I., Paleontologiya Rossii: Drevnii period:I. Flora grauvakkovoi, gornoizvestkovistoi i medistoslantsevoiformatsii Rossii (Paleontology of Russia: Ancient Period:I. The Flora of Graywacke, Calcareous, and CupriferousShaly Formations of Russia), St. Petersburg: J. Ionson,1854.El�Khayal, A.A., Chaloner, W.G., and Hill, C.R., Palaeo�zoic plants from Saudi Arabia, Nature, 1980, vol. 285, no. 1,pp. 33–34.El�Khayal, A.A. and Wagner, R.H., Upper Permian stratig�raphy and megafloras of Saudi Arabia: Palaeogeographicand climatic implications, Compte Rendu, 10e Congres Car�bonifere, Madrid, 1983, vol. 3, pp. 17–26.Esaulova, N.K., New fern species from the Kazaniandeposits of the Kama Region, Paleontol. Zh., 1996, no. 4,pp. 121–126.Fefilova, L.A., Paporotnikovidnye permi severa Priural’skogoprogiba (Pteridophyta of the Permian of the Northern Fore�Ural Foredeep), Leningrad: Nauka, 1973.Goeppert, H.R., in: Tchihatcheff, P., Voyage scientifiquedans l’Altai oriental, Paris: Chez Gide, 1845, pp. 378–392.Gomankov A.V. and Meyen, S.V., Tatarian flora (composi�tion and distribution in the Late Permian of Eurasia,Tr. Geol. Inst. Ross. Akad. Nauk, 1986, no. 401, pp. 1–174.Gorelova, S.G. and Radczenko, G.P., Main Late Permianplants of the Altai–Sayany mountain region, Tr. Vsesoyuz.Nauchno–Issled. Geol. Inst. Nov. Ser., 1962, vol. 79 (Mate�rials to phytostratigraphy of the Upper Permian of theAltai–Sayany mountain region), pp. 39–242.Grauvogel�Stamm, L. and Doubinger, J., Deux fougeresfertiles du Stephanien du Massif Central (France), Geobios,1975, vol. 8, pp. 409–421.Grunt, T.A., Esaulova, N.K., and Kanev, G.P., Eds., Biotavostoka evropeiskoi Rossii na rubezhe rannei i pozdnei permi:kollektivnaya monografiya (Biota of Eastern European Rus�sia at the Early–Late Permian Boundary: Collective Mono�graph), Moscow: Geos, 1998.Harris, T.M., The Yorkshire Jurassic flora: I. Thallophytaand Pteridophyta, Ann. Brit. Mus. Natur. Hist., 1961, pp. 1–212.Hill, C.R., Wagner, R.H., and El�Khayal, A.A., Qasimiagen. nov., an early Marattia�like fern from the Permian ofSaudi Arabia, Scr. Geol., 1985, vol. 79, pp. 1–50.Holmes, W.B.K., The Middle Triassic megafossil flora ofthe Basin Creek Formation, Nymboida coal measures, New

South Wales, Australia: Part 2. Filicophyta, Proc. Linn. Soc.New South Wales, 2001, vol. 123, pp. 39–87.

Ibrahim, A.C., Sporenformen des Agirhorizonts des Ruhr�Riviers, Berlin: Triltsch, 1933.

Ignatiev, I.A. and Naugolnykh, S.V., Early Kazanian floraof the Soyana River and its position among contemporane�ous floras and vegetative zones of Angaraland, Stratigr. Geol.Korrelyatsiya, 2001, vol. 9, no. 3, pp. 58–75.

Jennings, J.R. and Millay, M.A., Morphology of fertilePecopteris unita from the Middle Pennsylvanian of Illinois,Palaeontology, 1979, vol. 22, part 4, pp. 913–920.

Kidston, R.W. and Gwynne�Vaughan, D.T., On the fossilOsmundaceae: Part 1, Transact. Roy. Soc. Einburg, 1907,vol. 45, pp. 759–780.

Kon’no, E., Asama, K., and Rajah, S.S., The Late PermianLinggieu flora from the Gunong Blumut area, Johore,Malaysia: Contributions to the Geology and Palaeontologyof southeast Asia, XC, in Geology and Palaeontology ofSoutheast Asia, Tokyo: Univ. Tokyo Press, 1971, vol. 9,pp. 1–85.

Kutorga, S.S., Beitrag zur Palaeontologie Russlands, inVerhandlungen der Russisch–Kaiserlichen MineralogischenGesellschaft zu St. Petersburg, St. Petersburg: Gretsch, 1842,pp. 1–33.

Kutorga, S.S., Zweiter Beitrag zur Palaeontologie Russ�lands, in Verhandlungen der Russisch–Kaiserlichen Miner�alogischen Gesellschaft zu St. Petersburg, St. Petersburg:Gretsch, 1844, pp. 62–104.

Laveine, J.�P., Zhang, S., and Lemoigne, Y., Global paleo�botany, as exemplified by some Upper Carboniferous pteri�dosperms, Bull. Soc. Belge Geol., 1989, vol. 98, no. 2,pp. 115–125.

Lemoigne, Y., Flore mixte au Permien superieur en ArabieSaoudite, Geobios, 1981, no. 14, part 5, pp. 611–635.

Lesnikowska, A.D. and Galtier, J., Permineralized Maratti�ales from the Stephanian and Autunian of central France:A reinvestigation of Grandeuryella renaultii (Stur) Weissemend., Rev. Palaeobot. Palynol., 1992, vol. 72, pp. 299–315.

Lesnikowska, A.D. and Millay, M.A., Studies of PaleozoicMarattialeans: New species of Scolecopteris (Marattiales)from the Pennsylvanian of North America, Am. J. Bot.,1985, vol. 72, no. 5, pp. 649–658.

Lesnikowska, A.D. and Willard, D.A., Two new species ofScolecopteris (Marattiales), sources of Torispora securisBalme and Thymospora thiessenii (Kosanke) Wilson et Ven�katachala, Rev. Palaeobot. Palynol., 1997, vol. 95, pp. 211–225.

Leven, E.Ja., Naugolnykh, S.V., and Gorgij, M.N., Newfindings of Permian marine and terrestrial fossils in CentralIran (the Kalmard block) and their significance for correla�tion of the Tethyan, Uralian and West European scales,Riv. Ital. Paleontol. Stratigr., 2011, vol. 117, no. 3, pp. 355–374.

Lozovsky, V.R., Minikh, M.G., Grunt, T.A., et al., TheUfimian Stage of the East Europe scale: Status, validity,correlation potential, Stratigr. Geol. Korrelyatsiya, 2009,vol. 17, no. 6, pp. 46–58.

Mamay, S.H., Some American Carboniferous fern fructifi�cations, Ann. Mo. Bot. Gard., 1950, vol. 37, pp. 409–477.

1460


NAUGOLNYKH

Mapes, G. and Schabilion, J.T., A new species of Acitheca(Marattiales) from the Middle Pennsylvanian of Oklahoma,J. Paleontol., 1979a, vol. 53, no. 3, pp. 685–694.Mapes, G. and Schabilion, J.T., Millaya gen. n., an UpperPaleozoic genus of Marattialean synangia, Am. J. Bot.,1979b, vol. 66, no. 10, pp. 1164–1172.Meyen, S.V., New data on spore�bearing organs of UpperPaleozoic ferns of Angaraland, Paleontol. J., 1982a, no. 4,pp. 108–112.Meyen, S.V., The Carboniferous and Permian floras ofAngaraland (a synthesis), Biol. Mem., 1982b, vol. 7, no. 1,pp. 1–109.Meyen, S.V., New data on fertile organs of Upper Paleozoicferns of Angaraland, Paleontol. Zh., 1982c, no. 4, pp. 108–112.Meyen, S.V., Osnovy paleobotaniki (Fundamentals of Pale�obotany), Moscow: Nedra, 1987.Meyen, S.V., Carboniferous and Permian floras of Angara�land: Review, in Teoreticheskie problemy paleobotaniki(Theoretical Problems of Paleobotany), Moscow: Nauka,1990, pp. 131–223.Meyen, S.V., On the Subangara palaeofloristic area of thePermian, Sbornik pamyati chlena�korrespondenta AN SSSR,professora Vsevoloda Andreevicha Vakhrameeva (k 90�letiyuso dnya rozhdeniya (Collective Woks in Memory of the Cor�responding Member of the Academy of Sciences of theUSSR, Professors Vsevolod Andreevich Vakhrameev: To the90th Anniversary of the Birthday), Moscow: Geos, 2002,pp. 232–246.Meyer�Berthaud, B. and Galtier, J., Une nouvelle fructifi�cation du Carbonifere inferieur d’Escosse: Burnitheca, Fili�cinee ou pteridospermale?, CR Acad. Sci. Paris, 1986,vol. 303, ser. 2, no. 13, pp. 1263–1268.Millay, M.A., Acaulangium gen. n., a fertile marattialeanfrom the Upper Pennsylvanian of Illinois, Am. J. Bot., 1977,vol. 64, no. 2, pp. 223–229.Millay, M.A., A review of permineralized EuramericanCarboniferous tree ferns, Rev. Palaeobot. Palynol., 1997,vol. 95, pp. 191–209.Millay, M.A. and Taylor, T.N., Paleozoic seed fern pollenorgans, Bot. Rev., 1979, vol. 45, no. 3, pp. 301–375.Naugolnykh, S.V., New species of Ptychocarpus Weiss fromthe Lower Permian of the Middle Cisurals with remarks onsystematics and morphology of some associated features,Paleontol. J., 1995, vol. 29, no. 1A, pp. 44–62.Naugolnykh, S.V., Description of plant remains of Orthoth�eca semilibera Naugolnykh, sp. nov., in Biota vostokaevropeiskoi Rossii na rubezhe rannei i pozdnei permi (kollek�tivnaya monografiya) (Biota of Eastern European Russia atthe Early–Late Permian Boundary: Collective Works),Esaulova, N.K., Grunt, T.A., and Kanev, G.P., Eds., Mos�cow: Geos, 1998a, pp. 253–255.Naugolnykh, S.V., Comparative analysis of Permian floralassemblages of the Kozhim section (Fore�Urals near thePechora River) and stratotypic region (Middle Fore�Urals),Tr. Geol. Inst. Ross. Akad. Nauk, 1998b, no. 500 (UralMountains: Fundamental problems of geodynamics andstratigraphy), pp. 154–182.Naugolnykh, S.V., Flora of the Kungurian Stage of theMiddle Fore�Urals, Tr. Geol. Inst. Ross. Akad. Nauk, 1998c,vol. 509, pp. 1–201.

Naugolnykh, S.V., A new species of Todites (Pteridophyta)with in situ spores from the Upper Permian of Pechora Cis�Urals (Russia), Acta Palaeontol. Polon., 2002, vol. 47, no. 3,pp. 469–478.

Naugolnykh, S.V., Paleophytogeography of the PermianPeriod, Tr. Geol. Inst. Ross. Akad. Nauk, 2004, vol. 550 (Cli�mate during the epochs of great biosphere changes),pp. 194–220.

Naugolnykh, S.V., Permian Calamites gigas Brongniart,1828: The morphological concept, paleoecology, andimplications for paleophytogeography and paleoclimatol�ogy, Paleontol. Zh., 2005a, no. 3, pp. 94–103.

Naugolnykh, S.V., Upper Permian flora of Vjazniki (Euro�pean part of Russia), its Zechstein appearance, and thenature of the Permian/Triassic extinction, New MexicoMus. Natur. Hist. Sci., 2005b, bull. 30 (The Nonmarine Per�mian: Albuquerque), pp. 226–242.

Naugolnykh, S.V., Permian floras of the Urals, Tr. Geol.Inst. Ross. Akad. Nauk, 2007, vol. 524, pp. 1–322.

Naugolnykh, S.V., Permian ferns of the Fore�Urals: Oldproblems and new approaches, in Stratigrafiya i regional’nayageologiya Russkoi platformy i Zapadnogo Urala. Sborniknauchnykh trudov k 110 letiyu so dnya rozhdeniya professoraN.P. Gerasimova (Stratigraphy and Regional Geology of theRussian Platform and the Western Ural Mountains: Pro�ceedings to the 110th Anniversary of the Birthday of Profes�sor N.P. Gerasimov), Perm: Perm. Gos. Univ., 2009,pp. 17–32.

Naugolnykh, S.V. and Kuleshov, V.N., Iskopaemaya florareki Soyany: okno v permskii period (Fossil Flora of the Soy�ana River: A Window to the Permian Period), Arkhangelsk:Severn. Gos. Med. Univ., 2005.

Neuburg, M.F., Paleontologiya SSSR (Paleontology of theUSSR), vol. 12, part 3, no. 2: Verkhnepaleozoiskaya floraKuznetskogo basseina (An Upper Paleozoic Flora from theKuznetsk Basin), Moscow–Leningrad: Akad. Nauk SSSR,1948.

Osnovy paleontologii. Vodorosli, mkhi, psilofity, plaunovye,chlenistostebe’nye, paporotniki (Fundamentals of Paleontol�ogy: Algae, Mosses, Psilophytes, Club Mosses, Spheno�phytes, and Ferns), Moscow: Gos. Nauchno–Tekhn. Izd.Lit. Geol. Okhran. Nedr, 1963.

Ozhgibesov, V.P., Tereshchenko I.I., and Naugolnykh, S.V.,Permskii period: organicheskii mir na zakate paleozoya (ThePermian Period: Organic World at the End of the Paleo�zoic), Perm: Art�Dizain, 2009.

Prantl, K., Lehrbuch der Botanik fär mittlere und höhereLehransteller, Leipzig: W. Engelmann, 1874.

Psenicka, J. and Bek, J., Oligocarpia lindsaeoides (Etting�shausen) Stur and its spores from the Westphalian of CentralBohemia (Czech Republic), Acta Mus. Nat. Pragae. Ser. BHist. Natur., 2001, vol. 57, nos. 3–4, pp. 57–68.

Psenicka, J., Bek, J., Zodrow, E.L., et al., A new Late West�falian fossil marattialean fern from Nova Scotia, Bot.J. Linn. Soc., 2003, vol. 142, pp. 199–212.

Pukhonto, S.K., Stratigrafiya i floristicheskaya kharakteris�tika permskikh otlozhenii Pechorskogo basseina (Stratigraphyand floral Characteristics of Permian Deposits of thePechora Basin), Moscow: Nauchn. Mir, 1998.

Pukhonto, S.K. and Fefilova, L.A., Macroflora, in Paleon�tologicheskii atlas permskikh otlozhenii Pechorskogo ugol’nogo



basseina (Paleontological Atlas of Permian Deposits of thePechora Coal Basin), Leningrad: Nauka, 1983, pp. 28–92.Pukhonto, S.K. and Naugolnykh, S.V., Evolution of higherplants of the Fore�Urals in the Permian Period, in Nauka iprosveshchenie. K 250�letiyu Geologicheskogo muzeya RAN(Science and Education: To the 250 Anniversary of theGeological Museum of the Russian Academy of Sciences),Moscow: Nauka, 2009, pp. 319–352.Radczenko, G.P., Some plant fossils from the area of theOstashkiny Mountains in the Kuznetsk Basin, in Mat. Geol.Zap.�Sib. Kraya, 1936, no. 35, pp. 1–24.Radczenko, G.P., Index forms of Upper Paleozoic floras ofthe Sayany–Altai region, in Atlas rukovodyashchikh formiskopaemykh fauny i flory Zapadnoi Sibiri (Atlas of IndexForms of the Fossil Fauna and Flora of Western Siberia),vol. 2. Moscow: Gos. Nauchno–Tekhn. Izd. Lit. Geol.Okhran. Nedr, 1955, pp. 42–154.Radczenko, G.P., Index forms of fossil plants from coal�bearing deposits of the Kuznetsk Basin, in Atlas rukovod�yashchikh form iskopaemykh fauny i flory permskikh otlozhe�nii Kuznetskogo basseina (Atlas of Index Forms of the FossilFauna and Flora of Permian Deposits of the KuznetskBasin), Moscow: Gos. Nauchno–Tekhn. Izd. Lit. Geol.Okhran. Nedr, 1956, pp. 110–202.Ribeiro, C., On the Carboniferous and Silurian formationsof the neighbourhood of Bussaco in Portugal, Quart. J. Geol.Soc. London, 1853, vol. 9, pp. 135–161.Seward, C., The Jurassic Flora: Part 1. The Yorkshire Coast,London: Brit. Mus. Natur. Hist., 1900.Schimper, W.P., Handbuch der Palaeontologie, Teil II.Palaeophytologie, Lief. 1, Leipzig: Oldenbourg, 1879.Schmalhausen, J., Beiträge zur Jura�Flora Russlands,Mem. l’Acad. Imper. Sci. St. Petersbourg, 1879, ser. 7, vol. 27,no. 4, pp. 1–96.Singh, K.J., Naugolnykh, S.V., and Saxena, A., Permianand Triassic plant assemblages from the Tatapani–RamkolaCoalfield (India), in Palaeontology and Evolution of theBiodiversity in the Earth History (in Museum Context), Mos�cow: Geos, 2012, pp. 99–108.Stoneley, H.M.M., The Upper Permian flora of England,Bull. Brit. Mus. (Nat. Hist.) Geol., 1958, vol. 3, no. 9,pp. 295–337.Stubblefield, S.P., Taxonomic delimitation among Pennsyl�vanian marattialean fructifications, J. Paleontol., 1984,vol. 58, no. 3, pp. 793–803.Stur, M.D., Zur Morphologie und Systematik der Culm�und Carbonfarne, Stitzb. Naturwis. Kl., 1883, vol. 88, part 1,pp. 633–846.Takhtadjan, A.L., Higher taxa of vascular plants, excludingflowering plants, in Problemy paleobotaniki (Problems ofPaleobotany), Leningrad: Nauka, 1986a, pp. 135–142.Takhtadjan, A.L., Upper taxa of higher plants, excludingflowering plants, in Problems of Palaeobotany, Leningrad:Nauka, 1986b, pp. 135–142.Vakhrameev V.A., Vasilevskaya I.A., Vladimirovich, V.P.,et al., Phylum Pteropsida (pteridospermophytes), inOsnovy paleontologii. Vodorosli, mkhi, psilofity, plaunovye,chlenistostebe’nye, paporotniki (Fundamentals of Paleontol�ogy: Algae, Mosses, Psilophytes, Club Mosses, Arthro�phytes, and Ferns), Moscow: Gos. Nauchno–Tekhn. Izd.Lit. Geol. Okhran. Nedr, 1963, pp. 526–616.

Vladimirovich, V.P., Typical Kungurian Flora of the MiddleUral Mountains, Available from VINITI, no. 377�V86(Moscow, 1985).Vladimirovich, V.P., Higher plants: Telomophyta, Tr. Vse�soyuz. Nauchno–Issled. Geol. Inst. Nov. Ser., 1986, vol. 331(Atlas of characteristic assemblages of the Permian faunaand flora of the Ural Mountains and Russian Platform),pp. 32–38.Wagner, R.H., Lobatopteris alloiopteroides, una nuevaespece de Pecopteroidea del Estefaniense A espanol, Estud.Geol., 1958, vol. 14, no. 38, pp. 81–107.Wagner, R.H., On a mixed Cathaysia and Gondwana florafrom S.E. Anatolia (Turkey), in Compte Rendu, 4�e Congr.Carbonifere, Heerlen: Maestricht, 1962, vol. 3, pp. 745–752.Wagner, R.H., Pecopteris hispanica sp. nov., an UpperStephanian fern from the Cinera–Matallana Coalfield, NWSpain, An. Fac. Cienc. Porto. Suppl., 1985, vol. 64, pp. 1–4.Wagner, R.H., Hill, C.R., and El�Khayal, A.A., Gemellith�eca gen. nov., a fertile pecopterid fern from the Upper Permianof the Middle East, Scr. Geol., 1985, vol. 79, pp. 51–74.Wagner, R.H., Lausberg, S., and Naugolnykh, S.V., ThePermian Angara Flora from North Greenland: A progressreport, in XIV International Congress on the Carboniferousand Permian, Calgary: Univ. Calgary Press, 1999, p. 150.Wagner, R.H., Lausberg, S., and Naugolnykh, S., The Per�mian Angara flora from North Greenland: A progressreport, Carboniferous of the World, Hills, L.V.,Henderson, C.M., and Bamber, E.W., Eds., Can. Soc.Petrol. Geol. Mem., 2002, vol. 19, pp. 382–392.Wagner, R.H. and Lemos de Sousa, M.J., Oligocarpia lepto�phylla (Bunbury), nomenclatorial history and description ofthe lectotype, in Contrib. Carbonif. Geol. Palaeontol. Iber.Penins. An. Cienc. Porto. Suppl., 1985, vol. 64, pp. 481–490.Wagner, R.H., Soper, N.J., and Higgins, A.K., A Late Per�mian flora of Pechora affinity in North Greenland, Groenl.Geol. Unders., 1982, vol. 108, pp. 5–13.Wan, Zhihui and Basinger, J.F., On the fern PectinangiumLi et al., emend. (Marattiales), with spores in situ from thePermian of southern China, Rev. Palaeobot. Palynol., 1992,vol. 75, pp. 219–238.Wang, H. and Yang, G., Microscopical study of Qasimiafrom the Permian of western Henan Province, CentralChina, Palaeobotanist, 1996, vol. 45, pp. 255–258.Wang, Y., Guignard, G., and Barale, G., Morphologicaland ultrastructural studies on in situ spores of Oligocarpia(Gleicheniaceae) from the Lower Permian of Xinjiang,China, Int. J. Plant Sci., 1999, vol. 160, no. 5, pp. 1035–1045.Wang, Y. and Wu, X., Oligocarpia kepingensis sp. nov. fromthe Lower Permian of the northern Tarim basin, Xinjiangand its in situ spores, Chin. Sci. Bull., 1999, vol. 44, no. 2,pp. 108–111.Wang, Z.�Q., Past global floristic changes: The Permiangreat Eurasian floral interchange, Palaeontology, 1996,vol. 39, part 1, pp. 189–217.Weigelt, J., Neue Pflanzenfunde aus dem MansfelderKupferschiefer, Leopoldina, 1930, vol. 6, pp. 643–668.Weiss, C.E., Fossile Flora der jungstein Steinkohlenformationund des Rothliegenden in Saar–Rhein–Gebiete, Bonn: Ver�lag A. Henry, 1869, vol. 1.

1462


NAUGOLNYKH

Yan, T., Naugolnykh, S.V., and Sun, G., Upper Permianflora of Jiefangcun (northern China): Taxonomical compo�sition and importance for paleophytogeography, in Paleon�tologiya i stratigrafiya permskoi sistemy v muzeinykhekspozitsiyakh i chastnykh kollektsiyakh (Paleontology andstratigraphy of the Permian System in museum expositions andprivate collections), Perm: Poligr. Siti, 2010, pp. 36–42.Yang, G., Wang, H., and Sheng, A., Morphological andmicroscopical study on Scolecopteris from China, Palaeo�botanist, 1996, vol. 45, pp. 238–246.Yang Tao, Naugolnykh, S.V., and Sun, G., A new represen�tative of Neocalamites Halle from the Upper Permian ofnortheastern China (Jiefangcun Formation), Paleontol. J.,2011, vol. 45, no. 3, pp. 335–346.Zalessky, M.D., Flore paléozoïque de la série d’Angara,Mem. Com. Géol. Nouv. Sér., 1918, vol. 147, pp. 1–76.Zalessky, M.D., On new species of Permian Osmundaceae,Izv. Glavn. Bot. Sada, 1925, vol. 1, pp. 1–15.Zalessky, M.D., Permian flora of the Ural marginal areas ofAngaraland: Atlas, Tr. Geol. Kom. Nov. Ser., 1927, no. 176,pp. 1–52 (Zalessky, M.D., Flore Permienne des limitesOuraliennes de l’Angaride. Atlas, Mem. Comité Géol. Nouv.Sér., 1927, vol. 176).Zalessky, M.D., On subdivision and age of the anthra�colithic system of the Kuznetsk Basin based on extinctflora, Izv. Akad. Nauk, Otd. Mat. Estestv. Nauk, 1933a,no. 4, pp. 607–630.Zalessky, M.D., On new fossil plants of the anthracolithicsystem of the Kuznetsk Basin: I, Izv. Akad. Nauk, Otd. Mat.Estestv. Nauk, 1933b, no. 8, pp. 1213–1258.Zalessky, M.D., Observations sur les végétaux permiens dubassin de la Petchora: I, Bull. l’Acad. Sci. l’URSS. Classe Sci.Math. Natur., 1934a, nos. 2–3, pp. 241–290.Zalessky, M.D., Observations sur les végétaux nouveaux duterrain permien du Bassin de Kouznetzk: II, Izv. Akad.Nauk Otd. Matem. Estest. Nauk, 1934b, no. 5, pp. 743–775.

Zalessky, M.D., Description de quelques végétaux fossilesde l’étage kamien du Permien de l’Oural et de la rivière Soi�ana, Probl. Paleontol., 1937a, vols. 2–3, pp. 88–101.Zalessky, M.D., Flores permiennes de la plaine Russe,de l’Oural et du bassin de Kosnetzk et les correlations desdép ts qui les contiennent, Probl. Paleontol., 1937b,vols. 2–3, pp. 9–35.Zalessky, M.D., Végétaux du Permien du Bardien del’Oural, Probl. Paleontol., 1939, vol. 5, pp. 329–374.Zalessky, M.D. and Tchirkova, H.T., PhytostratigraphischeUntersuchungen im Bereich der Kohlenfuhrenden Schich�ten der Permischen Becken von Kuznetzk und vonMinussinsk in Sibirien, Palaeontographica, 1937, vol. 82,part B, Lief. 5–6, pp. 172–203.Zalessky, M.D. and Tschirkova, E.F., Permskaya floraPechorskogo Urala i khrebta Pai�Khoi (Permian Flora of thePechora Ural Mountains and Pai Khoi Mountain Range),Moscow–Leningrad: Akad. Nauk SSSR, 1938.Zenker, J.C., Scolecopteris legans, ein neues fossilsFarngewächs mit Fructificationen, Linnaea, 1837, vol. 11,pp. 509–512.Zodrow, E.L. and McCandlish, K., Upper Carboniferousfossil flora of Nova Scotia in the collections of the Nova ScotiaMuseum, with special reference to the Sydney Coalfield, Hal�ifax: Nova Scot. Mus., 1980.Zodrow, E.L. and McCandlish, K., Oligocarpia belliisp. nov. from the middle Upper Carboniferous of Cape Bre�ton Island, Nova Scotia, Canada, Palaeontographica, 1982,vol. 181, Lfg. 4–6, pp. 109–117.Zodrow, E.L., Simunek, Z., Cleal, C.J., Bek, J., andPsenicka, J., Taxonomic revision of the Palaeozoic maratti�alean fern Acitheca Schimper, Rev. Palaeobot. Palynol.,2006, vol. 138, pp. 239–280.

Translated by G. Rautian

o

Date post:	26-Nov-2023
Category:	Documents
Upload:	independent
View:	0 times
Download:	0 times

Permian ferns of western Angaraland

Documents