Review of Palaeobotany and Palynology Elsevier Publishing Company, Amsterdam - Printed in The Netherlands
T H E FOSSIL SPORES OF T H E F A M I L Y G L E I C H E N I A C E A E (MORPHOLOGY AND TAXONOMY) N. A. BOLCHOVITINA
Geological Institute, Academy of Sciences of the U.S.S.R., Moscow (U.S.S.R.)
(Received August 30, 1966)
SUMMARY
Studies of the spores of 25 Recent species of the subfamily Sticheroideae and 16 species of the subfamily Gleichenioideae were carried out in order to compare them with the fossil spores. The fossil spores of this family are divided into four morphological groups. The spores of the new organ genus Plicifera BOLCHOVITINAgen. nov. (type species Gleichenia delicata BOLCHOVmNA) are attributed to a first group. The spores of this species have the segment folds on the distal surface of the grains. To a second group is attributed the organ genus Gleicheniidites Ross, with spores which have distal folds and equatorial bulges in the exine. To a third group are attributed the spores of the new genus Clavifera BOLCHOVITINAgen. nov. (type species Gleichenia triplex BOLCHOVITINA). This genus has maceform projections on the corners of the spores. To a fourth group belong the spores of the new organ genus Ornamentifera BOLCHOVITINAgen. nov. (type species Gleichenia echinata BOLCHOVITINA), which apart from distal folds and equatorial bulges have a torulose exine.
INTRODUCTION
The Recent tetrahedral spores of five genera of the subfamily Sticheroideae as well as the spores of sixteen species of the subfamily Gleichenioideae (family Gleicheniaceae) have been studied, in order to make a comparison between these Recent and related fossil spores. As a result of these studies it was established that the most characteristic features of this family, distinguishing them from other families, are the distal folds, between the rays of the scar which are the projections of the cavity and give a convex outline to the central part of the distal side. These spores are also characterised by equatorial thickenings of the exine,
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discontinuous at the corner,,. These features, being the most stable, make it possible to attribute tbssil ~pores to this family. Fossil spores of the family Gleicheniaceae from .lurassic and Cretaceous deposits have been described by Ross (1949), BOL('HOVlrINA(1953, 1966), COOKSON (1953), ROUSti (1957), Cot Pt!R (1958). MAHAVKINA (1958), KRUTZSCH (1959), CHhONOVA (1960, 1961), ()m
MORPHOLOGICALGROUPS The present author believes that the Gleicheniaceae spores which have been described thusfar may be divided into four morphological groups based on their characteristic features (Plate I). The first group includes spores with a smooth exine and folds on the distal side. Their Recent analogues are the spores of the subfamily Gleichenioideae, tribes Gleichenieae-Gleichenia polypodioides SMITH and Diplopterygieae-Diplopterygium giganteum WALL (Plate IA I-4). To the second group spores with equatorial thickenings are attributed, discontinuous at the ends of the scar and with folds outlining the convex central part of the distal side. The Recent analogues of this type of fossil spores are representatives of the tribe Gleicheniae-Calymella alpina (R. BROWN) PRESL, D. dicarpa (R. BROWN) PRESL and Gleicheniastrum circinnatum (SWARTZ) NAKM. To this group also belongs the fossil genus Gleicheniidites Ross (Plate IA, B1-4). The species Gleicheniidites senonicus Ross was described by N. E. Ross, based on very scanty data from Sweden; therefore, that author did not give a generic diagnosis. Later, on the basis of the original description and illustration by Ross, DELCOURT and SPRUMONr (1955)formally proposed the genus G&icheniMites with G. senonicus as its type species. in 1964, SKARBY revised the species and type of spores of Gleicheniidites senonicus Ross, on the basis of the original specimens of N. E. Ross, containing sporangia with spores, but also taking into account dispersed spores from the same deposits in the Swedish Lower Campanian from where Ross had derived his material. Skarby believes that the thickening of the exine along and near the equator, which is discontinuous at the corners, is typical for this genus as opposed to the other genera. The exine is smooth and slightly granular. The genus is known from Jurassic to Oligocene and comprises the species Gh, icheniidites senonicus Ross, G. laetus (BOLCHOVITINA) BOLCHOVITINA, G.
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Rev. Palaeobotan. Pal.vnol., 3 (1967) 59-64
PLATE I
]]Z
~/
line 1~
I;ine 2~2
llne 1-1
line 2 - 2
line %1
line 2-2
Z£
<> ,'
/
line 1-1
ine 2 - 2
Structural schemes of the fossil spores of the family Gleicheniaceae. A. Plicifera delicata (BOLCHOVITINA)BOLCHOVITINA. B. Gleicheniidites senonicus Ross. C. Clavifera triplex (BOLCHOVITINA)BOLCHOVITINA. D. Ornamentifera echinata (BOLCHOVITINA)BOLCHOVITINA. 1. Proximal view; 2. distal view; 3. equatorial view; median section along one of the rays along the line 1-1 ; 4. equatorial view; cross section along the line 2-2; 5. proximal view, photograph; magnification × 500; 6. distal view, photograph; magnification × 500.
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[al(/¢)]ius D()RING, G. cclritullus (Botx'ttovlJl,,,\) BOLCIt()VIrIN.~X. G. radiatu,~ (BoLCHOVIIINA) BOLCHOVI'IINA. (;. rasilis (B()I.CIIOVITINA) BOLCIIOVIIINA. (i. umbonatus (BoLcHOVITINA) BOLCHOVIIINA. (i. compositus (BOI.CttOVITINA) BOLCHOVITINA, G. elegans NAt;v, G. minor DOR1NG. G. de/courti DORING, (; toriconcavus KRUTZSCtt.
To the third group are assigned the fossil spores which, besides the folds on the distal side and equatorial thickenings along the sides between the rays of the scar, have clavate processes or projections of the exine at the corners of the spore. The Recent spores have no such projections, but on certain species, for example, Gleicheniastrum microphyllum (R. BROWN) PRESL, Dicranopteris rigida (KuNzE) NAKAI, small processes are observed at the corners of the spore. (Plate IA, C1-4). To the fourth group are attributed the fossil spores which have the features of the spores of the subfamily Gleichenioideae, i.e., distal folds and equatorial thickenings. The sculptured surface, however, is granular, prickly or torulose which distinguishes these from other groups of fossil spores having similar structures. The Recent spores Dicranopteris linearis (R. BROWN) PRESL, having a finegranular surface of the exine, are similar to these fossil spores.
GENERIC REVISIONS The revision of the previously mentioned fossil spores of the genera Gleichenia Sw. and Gleicheniidites Ross allows the distinction of new organ genera, corresponding to three of the four morphological groups described above.
Plicifera BOLCHOVITINAorgan gen. nov., 1966 Gleieheniidites GRIGORIEVAet MALIAVKINA(pars), t961, p.50. Type species: Gleichenia delicata BOLCHOVITINA, 1953, p.22, pl.II, fig.l-4, Moscow region, Aptian. Diagnosis: Triangular spores with rounded corners and slightly convex straight or concave sides. The scar is simple, with rays being either equal to the radius or shorter. On the distal side there are three arcuate folds, reaching or not reaching the ends of the scar, and being curved towards the centre. The exine is smooth. BasisJbr the distinction of a new genus: The dispersed spores, having arcuate folds, differ from the other types of spores and, therefore, should be distinguished as a separate genus. It cannot be named Gleichenia, because to this genus also smooth spores without folds belong. Arcuate folds are also found in Recent spores of both the genus Gleiehenia and of other genera--Diplopterygium and Dicranopteris. 62
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This feature is characteristic of the spores of the subfamily Gleichenioideae; hence the spores of similar type may be attributed to the new organ genus Plicifera and assigned to the family Gleicheniaceae. Comparison and remarks: The genus Plicifera differs from the other fossil genera of the family in having arcuate folds on the distal side, as observed in the spores of Recent representatives of the subfamily Gleichenioideae (Plate 1,5,6). The genus contains two species: Plicifera delicata (BOLCHOVITINA) BOLCHOVITINA, and G. decora (CHLONOVA)BOLCHOVlTINA,with a geochronological distribution from Middle Jurassic to Oligocene.
Clav(fera 1 BOLCHOVITINAorgan gen. nov., 1966 Gleichenia BOLCHOVITINA,1953, p.54. Gleicheniidites GRIGORIEVA et MALIAVKINA (pars), 1961, p.63-64, non Gleicheniidites Ross, 1949. Type species: Gleichenia triplex BOLCHOVITINA,1953, p.54, pl.VIII, fig.10-13, Aptian, Moscow region. Diagnosis: Triangular spores with folds on the distal side possessing a broad equatorial thickening, often wavy along the margin and with processes or clavate projections at the corners. The exine is smooth. Basis Jbr the distinction of a new genus: See the description of the third morphological group. Comparison and remarks: Spores of Clavifera differ from those of Plicifera and Gleicheniidites Ross in having a broad equatorial thickening with uneven waves, processes and projections at the corners. The genus contains only one species Clavifera triplex (BOLCHOVITINA) BOLCHOVITINA (Plate IC5) distributed in Cretaceous and Paleocene deposits of the U.S.S.R. Ornamentifera BOLCHOVITINAorgan gen. nov., 1966 Gleicheniidites GRmORmVA et MALIAVKINA(pars), 1961. Type species: Gleichenia echinata BOLCHOVITINA,1953, p.55, pl.VIII, fig.17; Upper Albian, eastern slope of the southern Urals. Diagnosis: Triangular spores, more seldom rounded-triangular with equatorial thickenings and a torulose or prickly sculpture. Basis for the distinction of a new genus: See the description of the fourth morphological group. Comparison and remarks: The spores of the organ genus Ornamentifera combine equatorial thickenings of the subfamily Gleichenioideae with folds on 1 Clavifera = a spore with a club. Rev. Palaeobotan.PalynoL, 3 (1967) 59-64
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the distal side and tubercles or small prickles on the surface o f the exine. T h c y differ f r o m Plicff~,ra BOI,CHOVtIINA gen. nov. and Gleictzeniidites Ross in tile sctdptured surface o f the exine. T h e genus Ornament(/~,ra is i n t r o d u c e d as a new genus to replace Gleicheniidite.~
GRmORI~VA. T h e speciiic n a m e o f the latter c a n n o t be
retained for spores with a s c u l p t u r e d exine, as it has been used before. Besides. this t e r m was not i n c l u d e d by the I n t e r n a t i o n a l C o m m i t t e e on N o m e n c l a t u r e into the list " n o m i n a g e n e r i c a c o n s e r v a n d a " . T h e genus c o n t a i n s f o u r species: O. echinata (Bot.CHOVlXlNA) BOL(;tq()VII'INA, O. gt'anulata (GRIGORIEVA) Bt)I~CHOVITINA, O. tuberculata (GRIGORIEVA) BOLCHOVIT1NA, O. per(qrina (BOLCHOWrINA) BOLCttOVITINA, f r o m Lower Cretaceous deposits in the U . S . S . R .
REFERENCES
BOLCHOVITINA, N. A., 1953. Spores and pollen characteristic of Cretaceous deposits in the central regions of the U.S.S.R. Tr. Inst. Geol. Nauk, Akad. Nauk S.S.S.R., Geol. Ser., 145(61 ): 1-150 (in Russian). BOLCHOVmNA,N. A., 1966. The fossil spores of the ferns of the family Gleicheniaceae (taxonomy and distribution). [n: M. I. NEJSHTADT(NEUSTADT)(Editor), The Importance of Palynological Analysis for Stratigraphic and Palaeofloritic Investigations. Nauka, Moscow, pp. 65-75 (in Russian). CHLONOVA, A. V., 1960. Species of pollen and spores in the Upper Cretaceous sediments of the Chulym-Yenisey depression. Tr. Inst. Geol. Geofiz., Sibirsk. Otd. Akad. Nauk S.S.S.R., 3:1 105 (in Russian). CHLONOVA, A. V., 1961. Spores and pollen of the upper part of the Upper Cretaceous sediments in the eastern part of the West-Siberian depression. Tr. Inst. Geol. Geofiz., Sibirsk. Otd. Akad. Nauk S.S.S.R., 7:1 99 (in Russian). COOKSON, I. C., 1953. Difference in microspore composition of some samples from a bore at Comaum, South Australia. Australian J. Botany, 1(3): 462-472. COUPER, R. A., 1958. British Mesozoic microspores and pollen grains. Systematic and stratigraphic study. Palaeontographica, B, 103: 75-179. DELCOURT, A. et SPRUMONT, G., 1955. Les spores et grains de pollen du Wealdien du Hainaut. Bull. Soc. Beige G~ol., Pal~ontol., Hydrol., 5: 5-73. GRJGOmEVA, K. N. and MALIAVKINA,V. S., 1961. In: Spores and pollen of western Siberia, Jurassic-Paleocene. Tr. Vses. Nauehn. lssled. Geologorazved. Inst., 177:44-59 (in Russian). KRUTZSCH, W., 1959. Mikropal~iontologische (sporenpal~iontologische) Untersuchungen in der Braunkohle des Geiseltales. Geologie (Berlin), Beih., 21-22: 1-428. NAGY, E., 1963. Spores et pollens nouveaux d'une coupe de la Briqueterie d'Eger (Hongrie). Pollen Spores, 5(2): 397-412. PococK, S. A. J., 1962. Microfloral analysis and age determination of strata at the JurassicCretaceous boundary in the western Canada plains. Palaeontographica, B, 3(1-3): 1-95. Ross, N. E., 1949. On a Cretaceous pollen-and-spore bearing clay deposit of Scania. Bull. Geol. Inst. Univ. Uppsala, 34: 2543. RousE, G. E., 1957. The application of the new nomenclatural approach to Upper Cretaceous plant microfossils from western Canada. Can. J. Botany, 35(3): 359-375. SKARBY,A., 1964. Revision of Gleicheniidites senonicus Ross. Acta Univ. Stockholm, 11(3): 59-77.
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