Review of Palaeobotany and Palynology, 62 (1990): 97-105 EIsevier Science Publishers B.V., Amsterdam - Printed in The Netherlands 97 Paleopalynology and Paleoecology of Ca/amus-like Disulcate Pollen Grains VOLKAN S. EDIGER, ZÜHTÜ BATI and CENGIZ ALISAN Turkish Petroleum Corp., Research Center, 06420 Ankara (Turkey) (Received January 3, 1989; revised and accepted July 10, 1989) Abstract Ediger, V.S., Bati, Z. and Alisan, C., 1990. Paleopalynology Rev. Palaeobot. Palynol., 62: 97-105. and paleoecology of Calamus-like disulcate pollen grains. Disulcate pollen grains belonging to the genus Dicolpopollis have a distinct morphology. The pollen of this genus has characteristics in common with pollen of the modern plant Calamus which is a climbing rotan palm growing on the riparian margins of peat swamps. Dicolpopollis-type pollen are shown to have been abundant especially in the coal seams. Dicolpopollis is mostly found in the Tertiary rocks forming acme zones especially in the Oligocene except for a few occurrences and indicates probably a subtropical dimate. Paleopalynological taxonomy Genus Dicolpopollis Pflanzl1956 (emend.) Type Species: D. kockelii Pflanzl 1956 1956 Dicolpopollis Pflanzl, in: Notizblatt d. Hessischen Ladesamtes Bodenf. Wiesbaden, Bd. 84, p.241 Type Species: D. kockelii Pflanzl 1956 1960 Disulcites Erdtman 1947 ex Potonie, in: Senckenbergiana Lethaea, Bd. 41, p.464 Type Species: D. kalewensis Potonie 1960 1966 Disulcipollenites Nakoman, in: Ann. Geol. du Nord, v. 86, p.87 Type Species: D. kalewensis (Potonie 1960) Nakoman, 1966 1970 Disulcipollis Krutzsch, in: Atlas, v. 7, p.42 Type species: D. cuddalorensis (Ramanujam) Krutzsch 1970 deseription: Microspores with two sulci which are usually parallel to each other (Fig.1). Sulci sometimes converge or even touch each other toward the short side. Outline is irregularly elliptical to trapezoidal. Emended 0034-6667/90/$03.50 Exine is two-Iayered, relatively thick and the sculpture is vermiculate, rugulate or reticuIate. The grains are usually flattened along the meridional plane. Sulci are seen on opposite sides as notches on the equator in the equatorially flattened grains. Discussion: Dicolpopollis Pflanzl 1956 is a valid name for a genus with two species. Potonie (1966)designated the species Dicolpopo Ilis kockelii as lecto-type for the genus. As the genus description is rather inadequate, a new emendation is required. it is the first valid genus name for fossil pollen grains resembling Calamus polIen. The genus Dicolpites Vimal 1952 is validly published (Index Nominum Genericorum, 1979) but apparently based on dicotyledonous material from lignites of the West Punjab (Pakistan). The pictures show features very unlike those of the pollen grains described as Dicolpo-' pollis kockelii which are comparable with those of recent Calamus polIen. Dicolpites Van der Hammen, 1954 is also validly published, but © 1990 Elsevier Science Publishers B.V. 98 PLATE V.S. EDIGER, Z. BATI AND C. ALiSAN i CALAMU8-LIKE Equator D1SULCATE POLLEN GRAINS 99 Botanical ~ reported occur in severalother families such as Araceae and Dioscoreaceae, the sub-family Lepidocaryoideae of the Arecaceae is the most probable taxon for comparison, because megafossils are found together with pollen (Frederiksen, 1985). Chandler (1957) and Nagy and Palfalvy (1963) found spines, fruiting axes, flowers and seeds together with Dicolpopollislike pollen. Calamus is the most possible genus among other genera belonging to Lepidocaryoideae (Nagy, 1963;Gruas-Cavagnetto, 1968; Kedves, 1968, 1979; Muller, 1968; Krutzsch, 1970; Anderson and Muller, 1975; Hochuli, 1978). However, at least eleyen other palm genera in the Lypiddaryoideae, namely in the Calameae Bejaudia, Ceratolobus, Cornera, Dae- ,>/;f/1. Pole Pole EQUATORIAL VIEW POLAR Fig.1. Terms used in describing Dicolpopollis-type grains. VIEW pollen has to be rejected because it is alater homonym of Dicolpites Vimal 1952, which is not allowed under the rules of the ICBN (Art. 64.1).Both genera are not synonyms as theyare based on different types. Disulcites Potonie 1960is a validation of the name of a pollen type of Erdtman (1947). The type of this genus is D. kalewensis (I.N.G., 1979) and Potonie suggested at the same time that this pollen type represents Calamus pollen. His pictures seem to emphasize this idea too. The name is therefore superfluous, being alater synonym of Dicolpopollis Pflanzl, as this genus also represents fossil Calamus pollen. Disulcipollenites Nakoman 1966 is alater, obligate synonym of Disulcites Potonie 1960 and has to be rejected for reasons of priority. Disulcipollis Krutzsch 1970 is a validly published genus with the species D. cuddalorense as the nomenclatural type. The difference between Disulcipollis and Dicolpopollis is, however, so small, that it is suggested here to combine both genera. Krutzsch himself noted the similarity between the two genera already. PLATE affinity: Although Krutzsch (1970) that Dicolpopollis-like pollen also PoIe monorhops, Myrialepsis, Plectocomia, Plectocomiopsis, in the Korthalsieae Korthalsia and in the Metroxyleae Metroxylon and Salacca Schizospatha also produce disulcate pol- len grains (Thanikaimoni, 1970; M. Kedves, 1988, pers. commun.; W. Punt, 1989, pers. commun.). Some other authors suggested different botanical affi.nities for Dicolpopollislike pollen (Demonorhops-Hochuli, 1978; Metroxylon-Krutzsch, 1970; Potamegoton-Piel, 1971). Dicolpopollis kalewensis (Potonie 1960) comb. nov. (emend.) Basionomy: Disulcites kalewensis Potonie 1960 (Plate I, 1-4; Plate II, 1-29) Emended deseription: Disulcate pollen grains with a trapezoidal outline. The grains are usually flattened meridionally. Sulci are slightly converging to the short side and may touch each other. Exine is vermiculate-rugu- i SEM photomicrographs of the D. kalewensis from Thrace Basin, Turkey. 1, 2. Meridionally flattened grains. Sulci are shown by arrows. Note the formation of false-reticulum by the vermiculate-rugulate scultural elements in contact. x 5000. 3. Sulcus ends on the lateral side of the grains. Note the details of vermiculate sculpture. x 3500and x 12,500. 4. Details of sulci. x 10,000. 100 V.S. EDIGER, Z. BATI AND C. ALISAN PLATE II 7 13 8 9 14 15 20 19 25 LO 16 22 21 26 27 17 23 18 24 29 DlSULCATE POLLEN GRAINS CALAMUS·LIKE 10i 21 Calamus species studied by Kedves (1979) are reticulate, others are rugulate, clavate and verrucate. The pollen grains produced by Calamus palustris Griffith are considerably similar to pollen grains of Dieolpopollis kalewensis in having a rugulate sculpture, similar size, and nearly trapezoidal shape (Kedves, 1979,plate IX, figs.17, 18). n=50 20 19 ~18 16.4 x 20.4 " ;: 17l ". .~" ~"5 ~16L~' 14 16 14 15 17 18 L, LENGTH Fig.2. Length-width Basin, Turkey. 19 20 21 22 23 24 25 26 pm. diagram of D. kalewensis from Thrace Iate and a false reticulum may form where these sculptural elements are in contact. Size range is (14.4-19.8)x (16.2-25.2) ~m (50 specimens). The mean size is 16.4± 1.6 x 20.4± 2.0 ~m (Fig.2). Diseussion: The species which are published und er the genera Disuleites and Disuleipollenites should be transferred to Dieolpopollis. These species are Disuleites kalewensis Potonie 1960, Disuleites euddalorense Ramanujam 1966, Disuleites lutetieus Gruas-Cavagnetto 1967. The following species belong to the same genus: Dieolpopollis ealamoides Nagy 1963, Dieolpopollis elegans Muller 1968, Dieolpopollis malesianus Muller 1968, Dieolpopollis fragilis Salujha, Kindra and Rehman 1972, Dieolpopollis proprius Salujha, Kindra and Rehman 1972 and Dieolpopollis retieulatus Salujha, Kindra and Rehman 1980. Some of these species are very similar to each other but no further study is carried out here to revise them because of impossibility of examining their holotypes. For example, Dieolpopollis ealamoides published by Nagy (1963)seems to be monosulcate and should be included probably in Areeipites. Botanical affinity: Seven of the eleven recent Paleogeography Except for the Malaysian occurrence of Dieolpopollis by Muller (1968) and Bande and Prakash (1986),all other species of Dieolpopollis occur in Tertiary strata (Fig.3). Dieolpopollis-type pollen grains are recorded mostly in Eocene and Oligocene strata. The Eocene occurrences are from North America (Tschudy, 1973; Frederiksen, 1973), Burma (Potonie, 1960), France (Gruas-Cavagnetto, 1967, 1968), Germany (Krutzsch, 1970), Hungary (Nagy, 1963;Kedves, 1967)and North India (Salujha et aL.,1972,1974).The Oligocene occurrences are from Turkey (Nakoman, 1966, Akyol, 1971; Ediger, 1982), England (Boulter and Craig, 1979) and North India (Salujha et aL., 1972, 1974). The Neogene occurrences are rather limited and from Germany (Pflanzl, 1956),India (Ramanujam, 1966; Salujha et aL., 1980) and questionably from Turkey (Ediger, 1982).These occurrences show that Dieolpopollis spp. are frequently found in Eocene-Oligocene rocks all over the world. Paleoelima tology forms an acme zone in the Upper Oligocene rocks in the Northern Thrace Basin, Turkey (Fig.4). The only record about the climate of the Turkish Oligocene is by Maedler and Steffens (1979)who believe that a "mild warm climate" occurred in the Late D. kalewensis PLATE II Transmitted light microscope photomicrographs of D. kalewensis from Thrace Basin, Turkey. x 850. 102 V.S. EDIGER, Z. BATI AND C. ALiSAN Fig.3. Recorded occurrences of Dicolpopollis-type pollen on the Late Eocene paleocontinental map. Map is tak en from Smith and Briden (1979). (1) Texas, Mississippi, Alabama, Louisiana, Arkansas, U.S.A., Eocene (Tschudy, 1973; Frederiksen, 1973). (2) Columbia, Maastrichtian (Van der Hammen, 1954). (3) Bristol Channel, England, Middle Oligocene (Boulter and Craig, 1979). (4) Paris Basin, France, Sparnacien, Upper Cuisian (Kedves, 1968). Paris Basin, France, Sparnacien (GruasCavagnetto, 1967, 1968). (5) Belgium, Oligocene (Roche and Schuler, 1976) (6) Hirschberg, Germany, Middle Miocene (Pflanzl, 1956). Germany, Pal eo gene (Krutzsch, 1970). (7) Dorog Basin, Hungary, Lower Eocene (Kedves, 1967). Dorog Basin, Hungary, Lower Miocene (Nagy, 1963). Eger, Hungary, Late Oligocene (Nagy, 1979). (8) Istanbul, Turkey, Lower Oligocene (Akyol, 1971). Thrace Basin, Turkey, Oligocene-Miocene (Ediger, 1982). (9) Yozgat, Turkey, Eocene-Lower Oligocene (Nakoman, 1966). (10) India, Miocene (Ramanujam, 1966). (11) Burma, Eocene (Potonie, 1960). (12) Sarawak, Malaysia, Senonian-Recent (Muller, 1968). Indo-Malayan Region, Pal eo gene-Neo gene (Bande and Prakash, 1986). (13) N. India, Eocene-Oligocene (Salujha et aL., 1972, 1974), Neogene (Salujha et aL., 1980). Oligocene-Early Miocene, but their samples were poor in the number of taxa. The relative percentages of the thermophilous spore-pollen are used as indicators of the dimatic changes in this basin. The dimate curve for the Northern Thrace Basin differs somewhat from the climatic changes in other parts of the world for the Eocene-Oligocene time period recorded in the literature. The dimate usually underwent fluctuations in the Eocene and there is a marked decrease in temperature resulting in a change in dimate from tropical to subtropical during the EoceneOligocene epoch. The increased percentages of thermophilous elements in FigA indicate that the temperature was relatively higher in the Oligocene than in the Eocene, resulting probably in a temperate dimate in the Eocene instead of a subtropical dimate in the Oligo- cene-Miocene. However, according to Wolfe (1978),Hochuli (1984)and Aleksandrova et aL. (1987), the mean annmil temperature of the Late Eocene changed between 25°C at the beginning to 15°C at the end of this period. This means that the dimate was tropical at the beginning and subtropical at the end of the Late Eocene. This discrepancy may be caused by the low representation of spore-pollen in marine sediments deposited in the ?MiddleLate Eocene in the Northern Thrace Basin. For that reason, the Eocene part of the curve should be considered with caution. This situation mayaIso be the cause for the transitional character of the Eocene-Oligocene boundary from temperate to subtropical dimates. However, the well-documented cooling episode at the Eocene-Oligocene boundary caused the extinction of many tropi- MIOOLE LU - , 10 40 30 50 ii60 Thermaphile 70kalewensis ilDeO % i ? i LATE ••• T L ATE U '" , "OR --EAR -,. %SUB LY z.•.•.. u o io3 DlSULCATE POLLEN GRAINS CALAMUS·LIKE T i i ? ? E Elements R C ~ A T A PLlOCENE FigA. Stratigraphic distribution of D. kalewensis and thermophilous elements in the Northern Thrace Basin, Turkey. See Fig.3 for location of the basin. Relative frequencies of D. kalewensis are based on the means of each st age including only spore·pollen. Thermophilous elements include 22 bryophytic and pteridophytic spore taxa belonging to genera Camarazonotriletes, Gleieheniidites, Leiotriletes, Undulatisporites, Triplanosporites, Baeulatisporites, Triletes, Polypodiaeeoisporites, Retitriletes, Cieatrieosisporites, Corrusporis, Laevigatosporites and Verrueatosporites, 7 monocotyledonous angiosperm pollen taxa belonging to genera Cyeadopites, Monoeolpopollenites and Dieolpopollis and 15 dicotyledonous angiosperm pollen taxa belonging to genera Triatriopollenites, Trieolporopollenites and Tetraeolporopollenites. The solid line is drawn based on the percentages of thermophilous taxa, except some facies elements introduced by Hochuli (1978), including D. kalewensis, Laevigatosporites haardti and Trieolporopollenites margaritatus. cal elements and a southward shifting of climatic zones in the Mediterranean region in the Early Oligocene (Aleksandrova et aL., 1987). Except for the Eocene part of the diagram, a marked increase in the relative percentages of thermophilous elements is observed in the Late Oligocene and Oligo-Miocene (probably equivalent to Egerian). Such an increase has also been recorded in Hungary where the climate of the Oligocene-Miocene boundary (Egerian) was definitely subtropical according to the latest palynological information (E. Nagy, 1988,pers. commun.). The mean annual temperature of the Oligocene was about 15°Cat the beginning and about 10°C at the boundary with Miocene (Hochuli, 1984; Aleksandrova et aL., 1987). There is a marked cooling trend from Early Oligocene to Late Oligocene resulting in a climate change from subtropical to temperate-subtropical. However, this decrease is not linear and the temperature suddenly increased up to about 20°C somewhere in the Late Oligocene (Wolfe, 1978;Aleksandrova et aL., 1987and references therein). As the Late Oligocene is believed mainly to be humid (Hochuli, 1984) such a warm period seems quite possible for the Late Oligocene. Other sedimentological and paleontological data from the Thrace Basin also support this. During the Late Oligocene a huge continental area stretching from Yugoslavia to Turkey was present in the Paratethys between the Afro-Arabian and Eurasian continents (Steninger and Rogl, 1984). This Indo-PacificAtlantic seawayacross the Mediterranean may have caused a pronounced warming trend in the Late Oligocene (Haq et aL., 1977). Paleoecology The modern plant Calamus is a climbing rotan palm (liana or yine) which is most abund ant on the riparian margins of peat swamps beyond the marine influence in tropical-subtropical regions (Anderson and Muller, 1975; Bande and Prakash, 1986). The fossil Calamus-type of pollen has been reported in many strata, but in most of the reported occurrences, the maximum abundance seems to be always in coal (Frederiksen, 1985 and references therein). For this it is regarded as facies element (Hochuli, 1978) and it cannot be used as a paleotemperature indicator (C. Gruas-Cavagnetto, 1989,pers. commun.). it is usually found in values of 10-20% (Muller, 1968; Ediger, 1982), but sometimes reaches up to 26-50% or even more than 50% (Akyol, 1971). The apparent preference of Dieolpopollis-producing plants for peat swamps is well shown in Oligocene strata from Turkey by Corsin and Nakoman (1967)and Ediger (1982). 104 V.S. EDIGER, Z. BATI AND C. ALiSAN E ..>il D. kalewensis &: •.. c.. o Rela "o Ö 5 t i 18 io. % 15 ::....-'- •. 10.0' -,,_o."(1. o" '(1.:~"~;: material and the permission to publish the results. Their appreciations are also due to Assoc. Prof. Dr. D. Altiner, The Middle East Technical University, Ankara, Turkey, Prof. Dr. E. Nagy, Hungarian Geological Institute, Budapest, Hungary, Dr. C. Gruas-Cavagnetto, Universite des Sciences et Techniques du Languedoc, Montpellier, France, Prof. Dr. M. Kedves, University of Jozsef Atila, Szeged, Hungary for critically reading and editing the manuscript. M. Bülbül helped in the scanning electron microscopy. References Fig.5. Relative frequencies of D. kalewensis in the Oligocene strata from the Thrace Basin, Turkey (Data from Ediger, 1982). See Fig.3 for location of the basin. They have reported that the maximum relative frequencies are clearly in the coals rather than in the marly intercalations (Fig.5). Chandler (1957,p.75) also reports "plenty of pollen-grains of Calamus indicating a dense Calamus jungle" in the Oligocene lignites of S.W. England. Elsik (1978) reports that the Upper Paleocene coal from East Texas, U.s.A., contains mainly a CalamuspollenitesArecipites association. Roche and Schuler (1976)found pollen of Calamus always associated with spores of Acrostichum in Oligocene peats formed in brackish-water swamp forest in Belgium. A close relationship in the relative frequency distribution between D. kalewensis and Baculatisporites spp., Tricolporopollenites margaritatus, Polyvestibulopollenites verus (Alnus), and some fungal spores is also observed in the Thrace Basin, Turkey (Ediger, 1981, 1982;Ediger and Alisan, 1989). Acknowledgements The authors thank the Turkish Petroleum Corporation Research Center for providing Akyol, E., 1971. Microflore de l'Oligocene Inferieur recoltee dans un sondage pres d' Avcikoru, Sil e-IstanbuL. Pollen Spores, 13: 117-133. Aleksandrova, A.N., Prozorov, Yu.I. and Yasamanov, N.A., 1987. Climatic and floristic zonation of the Mediterranean region during early Cenozoic time. Int. Geol. Rev., 29: 503~514. Anderson, J.A.R. and Muller, J., 1975. Palynological study of Holocene peat and Miocene co al deposit from NW Borneo. 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