ISSN 00310301, Paleontological Journal, 2015, Vol. 49, No. 3, pp. 326–336. © Pleiades Publishing, Ltd., 2015. Original Russian Text © S.V. Naugolnykh, A.P. Pronin, 2015, published in Paleontologicheskii Zhurnal, 2015, No. 3, pp. 103–112.
A New Matoniaceous Fern from the Upper Triassic of the Caspian Depression in the Context of Florogenetic Processes of Transition from the Paleozoic to Mesozoic S. V. Naugolnykha and A. P. Proninb a
Geological Institute, Russian Academy of Sciences, Pyzhevskii per. 7, Moscow, 119017 email:
[email protected] b Company “Zhakhan”, Atyrau, Kazakhstan Kazan Federal University, ul. Lenina 18, Kazan, 420008 Tatarstan, Russia Received February 13, 2014
Abstract—A new matoniaceous fern species of the genus Phlebopteris Brongniart from the Upper Triassic of the Caspian Depression (Zhylyoiskii District, Atyrau Region, Kazakhstan) is described. It is represented by both sterile and fertile leaves preserved in situ, including the structure of sporangia and spores. The main trends in the evolution of ferns at transition from the Paleozoic to Mesozoic are discussed. Keywords: Matoniaceae, ferns, Phlebopteris, new taxa, taxonomy, evolution, Triassic, Kazakhstan DOI: 10.1134/S0031030115030090
INTRODUCTION In the study of fossil ferns, fertile leaves are of great importance. Wellpreserved specimens that allow detailed examination of reproductive structures and in situ spores are particularly significant. If fertile leaves with sori or synangia investigated in detail are reliably associated with sterile leaves of a certain type and the plant is assigned to a certain order and family, it is possible to use such data for wider taxonomic or florogenetic generalizations. The present study is devoted to detailed description of a Late Triassic matoniaceous fern, which is referred to a new species of the genus Phlebopteris Brongniart. The second purpose is analysis of the major evolution ary trends in the development of various fern groups at the Paleozoic–Mesozoic transition and, particularly, in the Triassic Period. MATERIAL AND METHODS Samples described in the present study come from the core of borehole “S.Nurzhanov509” (Fig. 1), sit uated in the eastern part of the Caspian Depression (Zhylyoiskii District, Atyrau Region, Kazakhstan), at 3223.0 m of depth. Lipatova (1985) referred this part of the section to the Kusankuduk Horizon, which cor responds to the Ashchitaipak Group of the Upper Tri assic (Lipatova, 1985). The material includes frag ments of fertile and sterile pinnae. The specimens are phytoleims (compressions) and imprints. The sporan
gia and spores obtained from the macerated sori were examined using a Vega Tescan MV 2300 scanning elec tron microscope (SEM) and Meiji Techno MT 9930 light microscope at the Geological Institute of the Russian Academy of Sciences, Moscow (GIN). Spores were extracted from sporangia by two comple mentary methods. Initially, sporangia were separated from fertile leaves using preparation needles under a binocular microscope and macerated for three days in glacial acetic acid. Products of maceration were washed in distilled water and placed on a specimen stage of SEM. For deeper oxidation of coaly matter, second method was used; maceration was processed in concentrated nitric acid for two days; then, phytoleims were washed in distilled water. Products of oxidation were removed by ammonia; cuticles of sporangial walls and spores were washed in distilled water and mounted in permanent preparations, with glycerin–gelatin medium for the study under a light microscope. The specimens described are stored in the Geolog ical Institute of the Russian Academy of Sciences, Moscow (GIN), collection no. 4851. Stratigraphy of Enclosing Deposits Based on palynological and structural–geological data, the enclosing deposits are referred with certainty to the Upper Triassic. In the Caspian Depression, Upper Triassic beds occur locally. They are of lacus trine genesis, filling depressions in underlying Per mian deposits. In this region, the Triassic beds are on
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average 100 m thick. In the area where Permo–Trias sic evaporites and salt domes pinch out (salt diapirs), Upper Triassic deposits are thicker, reaching 500 m of thickness. This type of Triassic deposits is frequently named “Prorva Beds” after the section name. In these beds, among redbed and speckled clayey deposits, there are sandstone beds, which are oilbearing in some places. The Prorva section has yielded many palynomorphs (lower part of the Upper Triassic: Ara trisporites sp., Cyathidites triangularis Rom., presum able spores of the fern Dictyophyllum nilssonii (Brongniart) Goeppert emend. Kruch., Duplex isporites gyrates Playford et Dettmann., Kyrtomisporis speciosus Mädl., K. laevigatus Mädl., Stereisporites sp., PALEONTOLOGICAL JOURNAL
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Ovalipollis sp., Gnetaceaepollenites sp., Ginkgocycado phytus sp., Sulcatisporites sp., Alisporites sp., and Chor dasporites singulichorda Kl.; upper part of the Upper Triassic: Zebrasporites interscriptus (Th.) Kl., Cinguli zonates spp., Kyrtomisporis speciosus Mädl., Trianco raesporites ancorae (Reinch.) E. Sch., Lycopodiacid ites rhaeticus E. Sch., Polycingulatisporites circulus Sim. et Kedv., Cornutisporites laevigatus E. Sch., C. seebergensis E. Sch., Conbaculatisporites longdonen sis Clarke, etc.) and vegetative remains of Late Triassic age: Neocalamites carcinoides Harris, Clathropteris sp., Podozamites lanceolatus (Lindley et Hutton) Braun, P. angustifolius (Eichwald) Heer; Glossophyllum (?) sp., and Sphenobaiera sp. (Lipatova, 1985).
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Fig. 2. Phlebopteris hazarensis Naugolnykh et Pronin, sp. nov., specimen GIN, no. 4851/275 C, venation of ster ile pinnule: (a) original figure of sterile leaf; (b) interpreta tion graphic decoding of morphology. Scale bar, 1 cm.
SYSTEMATIC PALEOBOTANY DIVISION PTERIDOPHYTA S C H I M P E R, 1 8 7 9 C L A S S PO L Y P O D I O P S I D A C R O N Q U I S T , TA K H T A D J I A N ET ZIMMERMANN, 1966 Order Matoniales Reveal, 1993 Family Matoniaceae Presl, 1847 Genus Phlebopteris Brongniart, 1828 Phlebopteris hazarensis Naugolnykh et Pronin, sp. nov. Plate 12, figs. 1–8; Plate 13, figs. 1–10
E t y m o l o g y. From the Khazarian Sea (ancient Arabian name of the Caspian Sea: BahralHazar). H o l o t y p e. GIN, no. 4851/275 A, fertile pinna; GIN, no. 4851/275 B, fragment of the same fertile
pinna; GIN, no. 4851/276 A, counterpart of fertile pinna; Caspian Depression, Kazakhstan, Atyrau Region, Zhylyoiskii District; Upper Triassic, Rhaetian Stage (Figs. 5a, 5c–5e). D i a g n o s i s. Fertile pinnae long, with subparallel margins and thick straight rachis. Pinnules attached to pinnal rachis at right (90°) angle. Fertile pinnules long, narrow, lanceolate, straight or slightly curved toward pinnal apex. Each welldeveloped fertile pin nule having more than thirty sori arranged in two lon gitudinal rows on abaxial surface of pinnule along midvein, on each of its side. Sori round, consisting of ovoid basal disc (receptaculum) and 9–12 sporangia attached to basal disc at acute angle. Each sporangium having distinct annulus formed by large specialized cells. Spores in situ subtriangular, with trilete (tetrad) mark. Proximal surface of spore smooth, distal surface finely vermiculate. Rays of trilete mark not reaching spore equator. Sterile pinnules slightly wider than fer tile pinnules, with pinnate to reticulate venation and lanceolate or prolonged subtriangular outline. D e s c r i p t i o n (Figs. 2–5). The pinnae are long and relatively wide, with long lanceolate entiremar gin pinnules. The maximum known length of fertile pinnae is 99 mm; however, judging from the general leaf proportions, a welldeveloped fertile pinna was more than 200 mm long at more than 100 mm of width. The maximum rachis width is 2 mm. The rachis is smooth, with very weak longitudinal folds. A distinct groove is absent. The mean length of welldeveloped pinnules ranges from 40 to 55 mm, with the maximum width of 4.5 mm. Pinnules are lanceolate in outline, with very gradually tapering distal part (Fig. 2; Pl. 12, fig. 4). The tip is slightly blunted. Venation ranges from pinnate to pinnate–reticu late, with a welldeveloped midvein reaching the pin nule tip (Figs. 2, 3). The lateral veins of sterile pinnules form vague clusters, indirectly suggesting the initially coherent nature of pinnules, which were formed in the taxon ancestral to Phlebopteris hazarensis sp. nov. by the fusion of individual segments of the last order, each of which corresponds to one cluster of lateral veins of Ph. hazarensis sp. nov. The most strongly developed pinnules display basal arches, although they are poorly pronounced. The of Phlebopteris hazarensis sp. nov. are rosette like sori, consisting of on average 11 (9–12) circular or oval sporangia attached to a common basal disk (recep taculum) at an scute angle (Fig. 4, Pl. 12, figs. 6, 7). The receptaculum is oval or circular in shape, with a flat tened distal surface; it was probably attached to the lateral vein base of fertile pinnules. The sori only slightly varies in diameter from 0.8 to 1 mm. The indu sium is absent. Each sporangium has an annulus, consisting of large specialized cells (Pl. 13, figs. 2, 3, 7, 10). The annulus borders threefourths of the sporangium perimeter, so PALEONTOLOGICAL JOURNAL
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Fig. 3. Phlebopteris hazarensis Naugolnykh et Pronin, sp. nov., specimen GIN, no. 4851/275 C, structure of middle part of sterile pinna. Scale bar, 1 cm.
that only the basal part of the sporangium remains free. The annulus is 80–120 µm wide. The sporangia have yielded spores preserved in situ. The spores are mostly triangular (Pl. 12, fig. 3; Pl. 13, figs. 5, 6, 9), although rounded triangular forms also occur (Fig. 5; Pl. 13, fig. 8). The spores vary in diame ter from 20 to 30 µm. The sporoderm surface is smooth on the proximal spore side (Pl. 13, figs. 5, 6). The sporoderm of the distal side has small, but distinct vermicular ornamentation (Pl. 13, figs. 8, 9), which is gradually flattened towards the spore equator. C o m p a r i s o n. The new species differs from the closest species Phlebopteris polypodioides Brongniart (Brongniart, 1828) in the longer and narrower pin nules, the larger sori, and larger sporangia. For exam ple, in the specimens from the Lower Jurassic of West Hubei (China) referred to Ph. polypodioides, the spo rangia are at most 420 µm in diameter (Wang and Mei, PALEONTOLOGICAL JOURNAL
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1999). Judging from the figure of venation in the basal part of the pinnule of Ph. polypodioides accompanying the protolog (Brongniart, 1828, pl. 83, fig. 1A; Fig. 6), the lateral veins of this species formed relatively regu lar trapezoid loops, whereas in Ph. hazarensis sp. nov., such loops are very rare and rounded in outline. Addi tional material, which was the basis for extended treat ment of the species Ph. polypodioides, provided by Harris (1961), shows that, in this species, the basal loops (basal arches) always remain trapezoid or elon gated polygonal in outline (Harris, 1961, text figs. 33C, 33F–33I). The lateral veins of Ph. hazaren sis sp. nov. usually deviate directly from the midveins in the pinnate order (Fig. 3, Pl. 12, fig. 2). In the present study, this character is regarded as archaic, indicative of a greater evolutionary primitiveness of Ph. hazaren sis sp. nov. in comparison with Ph. polypodioides and other Jurassic members of this genus.
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Although spores of the new species show the struc ture typical of matoniaceous ferns, they differ from spores of Ph. muensterii (Schenk) Hirmer et Hoer hammer in the absence of a distinct border (Brick et al., 1955, textfig. 9). According to the formal sys tematics, spores of Ph. hazarensis sp. nov. can be referred to the genera Dictyophyllidites Couper (forms with a thinner sporoderm and without a bordering equatorial thickening) or Matonisporites Couper (forms with a bordering thickening of the sporoderm). Pinnules of Ph. muensterii, are usually crescentic, curving towards the pinnal tip (Brick et al., 1955, pl. VII, figs. 1, 2); however, this character is not observed in the new species. Fertile pinnae of Ph. hazarensis sp. nov. differ from that of the other close species Ph. smithii (Daugherty) Arnold emend. Ash et al. in the much larger size and much more numerous cells in the annulus; Ph. haza rensis sp. nov., always has more than 20 and up to 25, whereas Ph. smithii on average has 12 and the maxi mum number is 19 (Litwin, 1985). R e m a r k s. Note that, as the genus Phlebopteris was established, the author understood it rather widely (Brongniart, 1828), assigning to it the species Ph. poly podioides, Ph. propingua (Lindley et Hutton) Brongn iart, Ph. phillipsii Brongniart, and Ph. (?) undans Brongniart from the Jurassic of Yorkshire, Ph. schouwii Brongniart from the Jurassic of Bornholm, and Ph. nilsonii Brongniart from the Lower Jurassic of Hoer (Scanie). Phlebopteris hazarensis sp. nov. fits completely in the initial understanding of the genus by Brongniart. Phlebopteris hazarensis sp. nov. is rather similar in the structure of fertile pinnae and sori to Nathorstia pectinata (Goeppert) Krassilov. The last species is also assigned to the family Matoniaceae (Krassilov, 1967). However, they differ considerably in the venation and sporangial structure. The lateral veins of Nathorstia pectinata form distinctive terminal forks with repeated dichotomy (Krassilov, 1967, textfig. 14b), which are absent in Phlebopteris hazarensis sp. nov. The sporan gia of Nathorstia pectinata are trapezoid and have a distinctive marginal induration or thickening at the annulus base (Krassilov, 1967, textfig. 14c), in con trast to the sporangia of Phlebopteris hazarensis sp. nov. Ph. hazarensis sp. nov. is somewhat similar in spo rangial structure to extinct cyathean ferns, for exam ple, Cyathea tyrmensis (Seward) Krassilov (Krassilov, 1978), although pinnules of both fertile and sterile leaves of this species are distinctly lobate and the distal
Fig. 4. Phlebopteris hazarensis Naugolnykh et Pronin, sp. nov., holotype GIN, no. 4851/275 A, structure of sorus. Scale bar, 1 mm.
side of spores is smooth in contrast to that of Ph. haz arensis sp. nov. Phlebopteris hazarensis sp. nov. is very similar in sporangial structure to both extant and extinct polypo dioid ferns (Fedotov, 1970, pl. XV, figs. 13, 14), but dif fers from it in the considerably weaker cuticularization of annulus cells. A similar structure of sporangia and in situ spores is observed in some ancient Late Paleozoic leptosporangiate ferns, for example, Doneggia Roth well from the Upper Pennsylvanian of Ohio (Roth well, 1978). O c c u r r e n c e. Upper Triassic, Rhaetian Stage; Caspian Region. M a t e r i a l. Imprint and counterpart of a fertile pinna fragment (holotype), two fragmentary preserved fertile pinnae, and four sterile pinna fragments. CHARACTER OF TRANSITION FROM THE PALEOZOIC TO MESOZOIC BASED ON THE ANALYSIS OF FOSSIL FERNS The author (Naugolnykh, 2013) analyzed the diversity of Permian ferns of the Russian Platform and ForeUrals (including the Pechora Coal Field) in a special paper, which summarizes all available data on the distribution of ferns in this region and describes new genera and species.
Explanation of Plate 12 Figs. 1–8. Phlebopteris hazarensis Naugolnykh et Pronin, sp. nov.: (1) holotype GIN, no. 4851/276 And, fertile pinnule; (2) spec imen GIN, 4851/275 C, sterile pinna, coalescent fusion of pinnule bases is distinct; (3) holotype GIN, no. 4851/275 A, spore extracted from sporangium; (4) holotype GIN, no. 4851/275 A, fertile pinnule; (5) holotype GIN, no. 4851/275 B, structure of middle part of fertile pinnule; (6, 7) holotype GIN, no. 4851/275 B, structure of sori; (8) holotype GIN, no. 4851/275 B, three neighboring fertile pinnules. Scale bars: (3) 10 µm, (5, 6, 7) 1 mm, and (1, 2, 4, 8) 1 cm. PALEONTOLOGICAL JOURNAL
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Fig. 5. Phlebopteris hazarensis Naugolnykh et Pronin, sp. nov., morphological features of fertile leaves: (a) holotype GIN, no. 4851/275 C, three neighboring fertile pinnules; (b) holotype GIN, no. 4851/275 A, spores extracted from sporangium, (c) holo type GIN, no. 4851/276 A, fertile pinna, (d) holotype GIN, no. 4851/275 A, fertile pinnule, (e) holotype GIN, no. 4851/275 A, general structure of fertile pinna. Scale bars: (b) 10 µm, (d) 1 mm, (a, c, e) 1 cm. PALEONTOLOGICAL JOURNAL
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Undoubted dominants among Late Paleozoic ferns are various eusporangiate taxa, of which Marattiaceae were particularly abundant and diverse (Millay, 1997). These were the Marattiaceae which gave rise to large treelike forms in the Carboniferous and Permian. The second most abundant group of Paleozoic ferns were botryopterids, which are mostly characteristic of the Carboniferous (Good, 1979; Millay and Taylor, 1980; Thomas and Taylor, 1993). Late Paleozoic leptosporangiate ferns were not abundant; however, they included rather unusual highly specialized taxa, for example, lianalike Sphy ropteris obliqua (Marrat) Kidston (Van Amerom, 1990). Organs of attachment of Sphyropteris were affixing thorns located on the rachis. Of other Carbon iferous leptosporangiate ferns, we should pay attention to the genera Discopteris Stur (Brousmiche, 1977, 1979; Pfefferkorn, 1978), Grambastia Brousmiche (Brousmiche, 1978), and Tenchovia Psenicka et Bek (Psenicka and Bek, 2004). Within Gondwana, several endemic fern lineages were formed in the Late Paleozoic and became extinct to the end of the Permian. Among Gondwanan endemics, we should pay attention to the genera Damudopteris Pant et Khare (Pant and Khare, 1974), Cuticulatopteris Pant et Misra (Pant and Misra, 1983), and Skaaripteris Galtier et Taylor (Galtier and Taylor, 1994). Some Gondwanan endemics formed treelike PALEONTOLOGICAL JOURNAL
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growth forms; an example is provided by the species Dernbachia brasiliensis Rössler et Galtier from the Permian of Brazil (Rössler and Galtier, 2002). In the Northern Hemisphere, within Angaraland, a number of endemics also appeared, i.e., the Permo– Carboniferous genus Prynadaeopteris Radczenko emend. Naug. and also the Permian genera Acro genotheca Naug., Convexocarpus Naug., Geperapteris S. Meyen, and Tumidopteris Naug. (Meyen, 1982; Naugolnykh, 2013). The genus Prynadaeopteris was recorded in the Permian beds of Cathaysia (Huang, 1983). In addition to abundant and diverse Maratti aceae, Cathaysian floras, which were assigned to the tropical zone of the Permian Period, included several genera of gleicheniacean ferns (Yang et al., 1997). At the Paleozoic–Mesozoic boundary, the general taxonomic diversity of floras changed very rapidly and dramatically on the global scale. Certainly, these changes involved ferns as well. A peculiarity of Triassic ferns, which inherited only slightly from Late Paleozoic predecessors, manifested itself even in the Early Triassic. For example, classical Early Triassic (Buntsandstein) floras of Western and Central Europe contain Anomopteris Brongniart, an unusual genus of uncertain phylogenetic relation ships, although it is provided with a perfect and exhaustive comprehensive description, which follows the best botanical traditions (GrauvogelStamm and
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Explanation of Plate 13 Figs. 1–10. Phlebopteris hazarensis Naugolnykh et Pronin, sp. nov., holotype GIN, no. 4851/275 A, SEM: (1) epidermal–cutic ular structure of abaxial side of fertile pinnule; (2) individual sporangium with completely preserved annulus, lateral view; (3) individual sporangium, view of annulus side; (4) in situ spore, view of equatorial side; (5, 6) spores preserved in situ, view of proximal side; (7) two sporangia belonging to one sorus attached at angle to basal disk of sorus; (8, 9) spores preserved in situ, distal view; (10) annulus of welldeveloped sporangium. Scale bars: (4–6, 8–10) 10 µm and (1–3, 7) 100 µm.
Grauvogel, 1980). It is highly probable that Anomopt eris belongs to a particular phyletic lineage of osmun dacean ferns, which appeared for the first time in the middle of the Permian Period (Naugolnykh, 2002) and reached the phase of idioadaptive radiation only at the beginning of the Mesozoic (Shorokhova, 1975). At the same time, even in the Triassic, the osmundacean forms belonging to the extant genus Osmunda L. already appeared (Phipps et al., 1998). The Upper Triassic beds have yielded rather reli able records of hymenophyllacean ferns referred to a new genus, Hopetedia Axsmith, Krings et Taylor (Axsmith et al., 2001). The most ancient hymenophyl laceans are known from the Carboniferous beds, where these plants are extremely scarce. At the end of the Triassic, the climate of the Euro– Sinean Phytogeographical Realm, which included the Caspian Region (Vakhrameev, 1990), was warm and humid, ranging from humid subtropical to humid tropical. Almost throughout this zone, vegetation was dominated by cycadophytes and thermophilic ferns, which probably include members of the genus Phle bopteris. In lowlatitude Late Triassic vegetation, wide distribution was characteristic not only of ferns of the matoniaceous genus Phlebopteris, but also the osmun dacean Todites Seward, presumably the gleicheni acean Wingatea Ash, and the dipteridian Clathropteris Brongniart (Litwin, 1985). In some regions, for exam ple, the Caucasus, the Marattiaceae already belonging to extant habitual types became even more widespread (Delle et al., 1986). The beginning of the Jurassic period was marked by intense divergence of schisean ferns, which had reached the acme in the Early Cretaceous (Krassilov, 1977). In the middle of the Mesozoic, treelike taxa appeared among cyathean ferns (Tidwell et al., 1989). The fern genus Phlebopteris is a rather common, although not abundant, component of Late Triassic floras of the Northern Hemisphere, particularly in the low latitudes, for example, in Keuper European floras (Kelber and Hansh, 1995); however, the contempora neous Gondwanan beds almost lack of plants of this group (Anderson and Anderson, 2008). In the Euro– Sinean Realm, matoniaceous ferns participated in shore vegetative communities and even dominated in some ecotopes (Wang, 2002). Summing up, it is possible to conclude that, at the Paleozoic–Mesozoic boundary, the evolutionary trend of ferns shifted abruptly. The role of eusporan giate ferns, which dominated in the Paleozoic and were mostly represented by arborescent taxa, was PALEONTOLOGICAL JOURNAL
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sharply reduced at the end of the Permian, although they remained in lowlatitude communities as rather rare relicts. Protoleptosporangiate ferns (osmun daceans), which appeared in the middle of the Per mian, sharply increased in abundance and diversity to the latter half of the Triassic became flourishing in the Jurassic and Early Cretaceous. In the modern flora, leptosporangiate ferns, which provide the basis of tax onomic diversity of Pteridophyta, are only inferior in species number to angiosperms. Leptosporangiate ferns appeared in the Late Paleozoic as a rare group and became extremely widespread throughout the world in the Triassic with the appearance and further evolution of certain taxa, such as Phlebopteris hazaren sis sp. nov. ACKNOWLEDGMENTS We are sincerely grateful to O.P. Yaroshenko (Geo logical Institute, Russian Academy of Sciences) for valuable discussions concerning the systematics of Tri assic palynomorphs and V.A. Krassilov (Haifa Univer sity, Israel) for reviewing the paper and valuable remarks. The study was supported by the State Program for Supporting Competitive Growth of Kazan Federal University among World’s Leading Scientific–Educa tional Centers and the Russian Foundation for Basic Research. REFERENCES Anderson, H.M. and Anderson, J.M., Molteno ferns: Late Triassic biodiversity in southern Africa, Pretoria: S. Afr. Nat. Biodiv. Inst., 2008. Axsmith, B.J., Krings, M., and Taylor, T.N., A filmy fern from the Upper Triassic of North Carolina (USA), Am. J. Bot., 2001, vol. 88, no. 9, pp. 1558–1567. Brik, M.I., Kopytova, E.A., and TurutanovaKetova, A.I., Some Mesozoic ferns of the southwestern ForeUrals and their spores, in Materialy po geologii i poleznym iskopaemym (Materials of Geology and Minerals), Moscow: Gos geoltekhizdat, 1955, pp. 131–176. Brongniart, A., Histoire des végétaux fossiles ou recherches botaniques et géologiques sur les végétaux renfermés dans les diverses couches du globe, Paris: G. Dufour et E. d’Ocagne, 1828. Brousmiche, C., Considérations sur Discopteris occidentalis Gothan 1954 (Pteridophyta du Westphalien d’Europe occi dentale), Géobios, 1977, vol. 10, no. 2, pp. 251–73. Brousmiche, C., Grambastia (Sphenopteris) goldenbergii (Andrae) nov. comb., espècetype d’un nouveau genre de
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Translated by G. Rautian PALEONTOLOGICAL JOURNAL
Vol. 49
No. 3
2015
