Borbola, Andrea

Kedves, Miklós

Priskin, Katalin

Kedves Miklós

Borbola Andrea

Priskin Katalin

University of Szeged

LM and TEM investigations on the Upper Cretaceous Ajkaite of Hungary II.

2. LM AND TEM INVESTIGATIONS ON THE UPPER CRETACEOUS AJKAITE OF HUNGARY III. M . KEDVES, A . BORBOLA a n d K . PRISKIN Cell Biological and Evolutionary Micropaleontological Laboratory of the University of Szeged, H-6701, P.O. Box 993, Szeged, Hungary . Abstract Light and transmission electron-microscope investigations were implemented on some trilete spores char-acteristic of the amber layers of the Ajka Brown Coal Basin. Thé presumed functions of the appendices of the Appendicisporites tricuspidatus spore type, with it's restricted regional and stratigraphie occurrences, are discussed in comparison with the elater bearing fossil and extant spores of the genus Equisetum. The results of the TEM Ajkaite trilete spore studies are compared with those LM data of the "adriennis" type spore mas-sula. Key words: Upper Cretaceous, Ajkaite, spores, LM, TEM. Introduction The Upper Cret'aceous Carpathian Basin spore assemblages have several local or re-gional restricted elements (KEDVES and PINIZ,1983). Some of these data have been pre-viously published (KEDVES, SZÓNOKY, MADARÁSZ and KOVÁCS, 2000), in particular the Eunormapolles type Hungaropollis GÓCZÁN 1964a; and the Schizaeaceous fern spore with long appendices, described by WEYLAND and GREIFELD (1953). The pre-sumed function of the appendices of this type of spore and its stratigraphical and geo-graphical distributions are the subject of this LM study. In addition, the ultrathin sec-tions from the amber have produced the first results concerning the fossil sporoderm. The aim of this paper is the following: 1. Investigate the spores of "tricuspidatus type" as one of a more or less local ele-ment of the European Normapolles Province, together with other local elements, that are found in the Carpathian Basin. 2. Present new ultrastructure TEM data from fossil Schizaeaceae spore type recov-ered from the Ajkaite. Materials and Methods Material from previously published LM studies (KEDVES, BORBOLA and PRISKIN, 2001; KEDVES and ALVAREZ RAMIS, 2002) were used in this investigation. The Ajkaite (sample KG-99) was embedded in Aral-dite (Durcupan, Fluka). The TEM photographs were taken with the Hungarian Academy of Sciences De-partment of Biophysics, Biological Research Center's Tesla BS-540 at a resolution of 6-7Á. 11 Results LM Results: Nomenclatural question of Appendicisporites tricuspidatus WEYLAND and GREIFELD 1 9 5 3 (Plate 2 . 1 . , figs. 1 - 4 ) After the publication of THIERGART (I949>and WEYLAND and GREIFELD (1953) a large number of Lower and Upper Cretaceous spores were assigned to the form-genus Appendicisporites. These spores also included a large number that had thickened appendices. Many of these types of spores were placed in the form-genus Plicatella MALYAVKINA adding to the nomenclatural confusion. DEÁK (1963, 1965), tried to re-solve this taxonomic problem. However, the nomenclature of the fossil Schizaeaceous spores has still not been satisfactorily resolved. For example, DAVIES (1985) in his monograph of the Anemiacean, Schizaeacean and related spores introduced the follow-ing taxa: Plicatella tricuspidata (WEYLAND and GREIFELD 1953, p. 12, Table 3, fig. 18) comb. nov. Early Senonian: formerly Appendicisporites, Anemia. GRANZOW and HELMERICH (1992, p. 439) wrote the following: "1949 wurd von THIERGART "Sporites appendicifer" und 1953 von WEYLAND and GREIFELD "Appendicisporites tricuspidatus" veröffentlich. Ein gegenüberstellung der beiden Sporen aus dem Senon von Deutschland bzw. Polen ergibt, dass sie eindeutung zur gleichen Formart gehören". These spores were classified as Plicatella appendicifera. This present study was restricted to only those spores in which the lengh of the appendices is longer than the spore radius. In re-gards to the current nomenclatural problems, it is agreed in this paper to accept the con-cept of WEYLAND and GREIFELD (1953, p. 43) citing the following: "THIERGART bezeichnet die Spore als eine stark spezialisierte kurzlebige Form". The most important occurrences of the spores of "tricuspidatus type" are as follows: Alb-Cenomanian. - A. tricuspidatus WEYL. and GR., Vojvodina and near Belgrade, Yugoslavia, PANTIC and DULIC ( 1 9 9 4 ) . Cenomanian. - A. tricuspidatus WEYL. et GREIF., Pont Bati, Vendée, France, D U R A N D , TERS and VERGER ( 1 9 6 3 ) , Appendicisporites tricuspidatus W E Y L . et GREIF. 1 9 5 3 , Charente-Maritime, France, DEÁK and COMBAZ ( 1 9 6 7 ) , Anemia tricuspidata ( W E Y L A N D et KRIEGER) BOLKHOVITINA 1 9 6 1 , Laudun Gard, France, MÉDUS et TRIAT ( 1 9 6 9 ) , A. tricuspidatus W E Y L et GREIF., Peruc Formation, Czech Rep., PACLTOVÁ and SVOBODOVÁ ( 1 9 9 2 ) , A. tricuspidatus WEYL. et GREIF., Blansko Graben, Czech Rep., SVOBODOVÁ ( 1 9 9 7 ) , Appendicisporites tricuspidatus, Poganovo Paralic Series South-east Serbia, PANTIC and DULIC ( 1 9 9 9 ) . Cenomanian/Turonian. - A. tricuspidatus WEYLAND et GREIFELD, León, Northern Spain, V A N AMEROM ( 1 9 6 5 ) , A. tricuspidatus W E Y L . et GREIF. Vocontian Basin, Czech R e p . , SVOBODOVÁ, M É O N a n d PACLTOVÁ ( 1 9 9 8 ) . Turonian. - A. tricuspidatus W E Y L . et GREIF. Sabran, Gard, France, DUCREUX, GAIL-LARD and SAMUEL (1982). Plate 2.1. 1-4. Appendicisporites tricuspidatus WEYLAND et GREIFELD 1953., slide: KG-99-9, Cross-table number: 22.7/141.2, figs. 1,2. 500x, figs. 3,4, 1 .OOOx. 5. Schizaeaceae spore of "adriennis type"., slide: KG-99-19, Cross-table number: 20.3/140.8. 6. Ultrastructure of a laevigate trilete spore in the amber. Block No.: 99-KG-5, Negative No.: 8103, S.OOOx. 7. Detail of the ultrastructure of a massula of laevigate trilete spores. Block No.: 99-KG-5, Negative No.: 8106, 15-OOOx. 12 Senonian. - A. tricuspidatus WEYL. et GREIF. Quedlinburg Germany, WEYLAND and GREIFELD ( 1 9 5 3 ) . Santonian. - A. tricuspidatus WEYL. et GREIF. 1953, Ajka, Hungary, G O C Z A N (1964a). Santonian/Campanian. - A. tricuspidatus W E Y L A N D et G R E I F E L D (15), Bakony, Hun-gary, G6CZAN (1964a,b). As regards the spores from the brown coal of Ajka we cite from the paper of G6CZAN (1964a); p. 233: "3. In zone "C" the spores with verrucate structure (13), Appendici-sporites tricuspidatus WEYLAND et GREIFELD (15), 40-45 p. sized Oculopollis (12), as well as tetracolporate (11) and inaperturate (14) 60 to 100 p in diameter, are dominant (Figs. 11 to 15)". • Campanian. - Gruppe 3, PI. I , fig. 21 only, Aachen, Germany, K R U T Z S C H (1957). Ful-velian. - Appendicisporites tricuspidatus, Saon, France Drome, I P E R T (1976). LM data the of the spores isolated from the brown coal amber bearing layers of Ajka Amb triangular with convex sides (Plate 2.1., figs. 1,2). The laesurae of the tetrad mark reach the base of the appendices (Plate 2.1., figs. 1,2,4). Sculpture canaliculate, the size of four isolated striae and muri is about 20 pm (Plate 2.1., fig. 4). Diameter of the spore without appendices 48-60 pm, the appendices are 45-55 pm long so the size of the appen-dices is larger than the radius of the spore. Sometimes the appendices are bended in the direction of the proximal pole (Plate 2.1., figs. 1,2). It may be presumed that in the sporan-gia, the appendices were close to the spore body as is found in extant spore genus Equise-tum. The appendices are characteristic twisted well shown in figs. 2,4 in Plate 2.1. The new TEM results of the spores in the Ajkaite A massula of laevigate spores of Schizaeaceae of "adriennis type" (Plate 2.1., fig. 5) occurred in our slides. TEM data from similar mass of spores was observed (Plate 2.1., fig. 7). The surface of the exospore is uneven and its inner part is, in general, electron dense. Similar mass of spores from the Paleozoic was published by WELLMAN, ED-WARDS and A X E (1998), and WELLMAN (1999). Another spore, without protoplasm, occurred in our ultrathin sections of the amber (Plate, 2.1., fig. 6). The ultrastructure of the laesurae is also well shown with the inner part of the exospore being electron dense. Discussion and Conclusions The long appendices of A. tricuspidatus may be comparable to the elater bearing fossil and extant Equisetum spores. The fossil forms from the Pennsylvanian with three circinately coiled elaters were decribed by WILSON ( 1 9 4 3 ) and assigned to Elaterites triferens. There are several papers concerning this subject with the most important ones, in our view, are as follows: POTONIE ( 1 9 5 6 ) emphasized the following, p. 5 6 : "Die Gat-tung zeigt Beziehung zu den mesozoischen Gattung Appendicisporites WEYLAND & KRIEGER 1 9 5 3 " . G O O D and TAYLOR ( 1 9 7 4 ) published LM and SEM photographs of E. triferens spores isolated from Calamocarpon insignis microsporangia. Fig. 6, in P. 149, illustrates a twisted elater with a striate surface. GOOD ( 1 9 7 5 ) described several Calam-itean cones with associated spores and a wind dissemination was emphasized for the function of the elater in Equisetum. "In the case of calamitalen spores it is also possible that elaters functioned as a means of propelling the spores from the sporangium and in doing so became detached" (GOOD, 1 9 7 5 , p. 8 4 ) . The Mesozoic fossil elater bearing spore - Equisetosporites DAUGHERTY 1941 probably has four elaters following the 14 monograph of POTONIÉ (1956). KEDVES (1979) reviewed the most important basic re-sults of the spores of the extant genus Equisetum through LM and SEM investigations of fourteen species. The striate surface of the elaters of these fourteen species was observed and the fine ornamentation was discussed. It is worth of mentioning that there are four elaters in the spores of Equisetum in contrast to those of the previously mentioned Pa-leozoic calamitean spores. After being subjected to high temperatures, the twisting of the elaters is more charactertistic. It can be presumed that the function of the Schizaeaceae appendices is similar to those of the Equisetaceae elaters. It also is noteworthy that the number of these appendices is three and different, in contrast, to the more or less globu-lar Sphenopsida spores that are within the fossil forms of Schizaeaceae. This latter men-tioned group, an isolated one of limited stratigraphical and geographical distribution, may have similar morphological features for their dispersion. This type of spore was discussed herein as an addition to the local előments of the Upper Cretaceous flora of the Carpathian Basin. For example, the genus Hungaröpollis GÓCZÁN 1964a has a simi-lar restricted distribution. All these data are important for the understanding the fossil vegetation contained in the "amber tree" The new data presented herein are important for the understanding the ultrastructure of fossil spores. It is hoped that, in spite of the fact that protoplasm was not preserved in the spores of this study, the information will be useful to future investigators. 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LM and TEM investigations on the Upper Cretaceous Ajkaite of Hungary III.

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LM and TEM investigations on the Upper Cretaceous Ajkaite of Hungary II.

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