Turkish Journal of Earth Sciences (Turkish J. Earth Sci.), Vol. 15, 2006, pp. 155-180. Copyright ©TÜB‹TAK
Stratigraphic and Palaeoenvironmental Significance of Bartonian–Priabonian (Middle–Late Eocene) Microfossils from the Baflçeflme Formation, Denizli Province, Western Anatolia
MEHMET SERKAN AKK‹RAZ1, FUNDA AKGÜN1, SEFER ÖRÇEN2, ANGELA ANNELIESE BRUCH3 & VOLKER MOSBRUGGER3
1Dokuz Eylül Üniversitesi, Mühendislik Fakültesi, Jeoloji Mühendisli¤i Bölümü, Bornova, TR–35100 ‹zmir, Turkey (E-mail: [email protected]) 2Yüzüncü Y›l Üniversitesi, Mühendislik-Mimarl›k Fakültesi, Jeoloji Mühendisli¤i Bölümü, TR–65080 Van, Turkey 3Institute and Museum of Geology and Palaeontology, Sigwartstr. 10, D–72076 Tübingen, Germany
Abstract: This study explains the stratigraphical and palaeoenvironmental significance of Bartonian–Priabonian (Middle–Late Eocene) fossils, the fauna and flora obtained from the Baflçeflme formation of the Çardak–Tokça basin (western Anatolia). The studied sequence is an outcrop from the Baflçeflme formation, deposited in a shallow- marine to coastal environment without stratigraphic breaks. Forty genera and 58 species have been recognized in the palynological assemblage of the Baflçeflme formation. A well-preserved diverse palynomorph and foraminiferal assemblages yield the Middle–Late Eocene age for the Baflçeflme formation. In western Anatolia, mangrove elements Nypa and Pelliciera have been first recorded in this study. The pollen of Mauritia and Acrostichum occur in the back-mangrove environment. Lowland–Riparian and montane elements are characterized by dominance of Myricaceae, Betulaceae, Engelhardia, Fagaceae, Myrtaceae, Anacardiaceae and Taxodiaceae, Pinus, Abies, Picea, Cathaya, Quercus and Castanea, respectively. Fresh-water elements are represented by Sparganiaceae, Pediastrum sp. and Aglaoreidia cyclops. The palynological data for samples taken from the lower part of the section of the Baflçeflme formation indicate a back-mangrove environment. The presence of poorly preserved dinoflagellate cysts suggest that sedimentation occurred in a mangrove environment in the upper part of the section. The well-preserved foraminiferal data, along with corals, bivalves and gastropods, indicate that sedimentation ceased in the shallow-marine environment. In this paper, terrestrial climatic conditions of the Baflçeflme formation are also discussed on the basis of the coexistence approach.
Key Words: Middle–Late Eocene, mangrove, western Anatolia, benthic foraminifer, palynomorph, palaeoenvironment, palaeoclimate
Denizli Yöresindeki Bartoniyen–Priyaboniyen (Orta–Geç Eosen) Mikrofosilleri içeren Baflçeflme Formasyonu’nun Stratigrafik ve Ortamsal Önemi, Bat› Anadolu
Özet: Bu çal›flma, Çardak–Tokça havzas›n›n (Bat› Anadolu) Baflçeflme formasyonunundan elde edilen Bartoniyen–Priyaboniyen (Orta–Geç Eosen) yafll›, fauna ve floran›n stratigrafik ve paleoortamsal önemini aç›klar. Çal›fl›lan istif, stratigrafik kesiklik olmaks›z›n, s›¤ deniz, k›y› ortam›nda çökelmifl Baflçeflme formasyonundan bir yüzlektir. Baflçeflme formasyonunun palinolojik toplulu¤unda 40 cins ve 58 tür tan›mlanm›flt›r. ‹yi korunmufl çeflitli palinomorf ve foraminifer topluluklar›, Baflçeflme formasyonun yafl›n› Orta–Geç Eosen olarak vermektedir. Bat› Anadolu’da, mangrove elementleri olan Nypa ve Pelliciera ilk kez bu çal›flmada kaydedilmifltir. Mangrov gerisindeki ortamda Mauritia ve Acrostichum polenleri mevcuttur. Alçak alan-Irmak kenar› elemanlar›, Myricaceae, Betulaceae, Engelhardia, Fagaceae, Myrtaceae, Anacardiaceae ve Taxodiaceae, da¤ elemanlar› ise Pinus, Abies, Picea, Cathaya, Quercus ve Castanea ile karakterize edilir. Tatl›su elamanlar› Sparganiaceae, Pediastrum sp. ve Aglaoreidia cyclops ile temsil edilir. Elde edilen palinolojik veriler, Baflçeflme formasyonun alt kesimlerinden al›nan örneklerin mangrov gerisi ortam› belirtti¤ini göstermektedir. ‹stifin üst kesimlerinde kötü korunmufl dinoflagellatler›n varl›¤› tortullaflman›n
155 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
mangrove ortam›nda gerçekleflti¤ini göstermektedir. Mercan, bivalvia ve gastropodlu iyi korunmufl foraminifer verileri, tortulaflman›n s›¤ denizel ortamda son buldu¤unu göstermektedir. Bu makalede, ‘ coexistence approach’ yöntemine dayal›, Baflçeflme formasyonunun karasal iklimsel koflullar› da tart›fl›lm›flt›r.
Anahtar Sözcükler: Orta–Geç Eosen, mangrov, Bat› Anadolu, bentik foraminifer, palinomorf, paleoortam, paleoiklim
Introduction types of conglomerates, these belonging to the Eocene, The Palaeocene–Eocene sedimentary successions of Lower Oligocene and Oligocene, respectively. fienel western Anatolia are claimed to have developed on the (1997) collected the findings of Göktafl et al. (1989) and different tectonostratigraphic units, such as the Lycian reinterpreted the age of the Baflçeflme formation as Late Nappes (Poisson 1976; Yalç›nkaya et al. 1986; Göktafl et Lutetian–Priabonian. Sözbilir et al. (2001) studied the al. 1989; Özkaya 1991; fienel 1991, 1997; Collins & stratigraphic and tectonic properties of the Eocene in the Robertson 1997, 1998, 1999; Bozkurt & Park 1999; Baklan succession, located 10 km from the study area, Sözbilir et al. 2001; Sözbilir 2002), the Menderes Massif (Figure 1b) and reported a well-preserved marine fauna (e.g., Poisson 1976; Özkaya 1990, 1991; Bozkurt & which yields a Bartonian (Middle Eocene) age. Park 1994; Özer et al. 2001; Koralay et al. 2004) and This study focuses on the Baflçeflme formation, the Bey Da¤lar› carbonate platform (Özkaya 1991; Collins composed of both marine and coal-bearing lacustrine & Robertson 1998; Sar› & Özer 2002) (Figure 1a, b). sediments. The objective of this paper is to provide for The non–metamorphosed Palaeocene–Eocene the first time palynological and foraminiferal evidence sedimentary successions of western Anatolia generally from the Maden and the Asar members of the Baflçeflme consist of conglomerate, sandstone, turbiditic formation, to obtain precise ages, to ascertain sandstone–mudstone alternations, bioclastic limestone depositional environments, to begin to understand lenses, blocks of limestones and volcanic rocks, and these qualitative palaeoclimatic conditions for these units, and have been interpreted to be of the supra-allochthonous also to analyze the similarities to and differences from basin type, and to have developed over the Lycian Nappe correlative Eocene basins in Turkey. package (Sözbilir 2002). Limited coal-bearing Eocene outcrops have been observed in western Anatolia. The coal-bearing Eocene sediments of the Çardak-Tokça Stratigraphy basin, which stratigraphically overlie the Lycian Nappes, In the area, pre-Eocene basement consists of the are exposed 35 km east of Denizli (Figure 1a, b). Triassic–Lower Eocene Lycian Nappes and generally Micropalaeontological and stratigraphical studies on comprises metaconglomerate, metasandstone, the Eocene formations of the Çardak-Tokça basin have recrystallized limestone, metavolcanites, dolomite, been either neglected or carried out by Mineral Research dolomitic limestones, and ophiolitic-rock matrix and of Exploration Institute (M.T.A.). The unpublished report blocks (Göktafl et al. 1989). The Baflçeflme formation of Göktafl et al. (1989) was the first comprehensive unconformably overlies the Lycian Nappes and is made up stratigraphic and palaeontological study of the Tertiary of four different members which are terrestrial and sediments of the Çardak-Tokça basin. The Baflçeflme shallow marine in character; these are (from bottom to formation was formerly subdivided into fours members top) the Dazlak, Beflparmak reef, Maden and Asar (from bottom to top), the Dazlak, Beflparmak reef, members (Göktafl et al. 1989; fienel 1997) (Figures 1b & Maden and Asar members. That study reported that the 2). In the study area, the Dazlak, Maden and Asar age of the Baflçeflme formation is Late Eocene members occur the sequence (Figure 2). Here, their (Priabonian) on the basis of unillustrated benthic lithological properties are described briefly, in ascending foraminifers, mollusks and corals. fiahbaz & Görmüfl order. (1992) examined the stratigraphic and sedimentological The Dazlak member, which is barren of microfossils, properties of the conglomerates that crop out as the generally comprises a reddish conglomerate and Çardak-Tokça basin fill and recognized three different sandstone alternation of transgressive character. The
156 M.S. AKK‹RAZ ET AL. 1000 m 0 ÇARDAK Upper Miocene - Pliocene sediments Avdan Geological map of the Hayrettin (b) Oligocene sediments me ı ş çe BOZKURT ş formation Armutalan undiff. Ba BAKLAN aziçi ğ Bo Asar mem. 1987; Seyito¤lu & Scott 1996). N et al. b
Maden mem. ACIGÖL 20 km
1985; Konak TURKEY Figure 1b me formation 0 ş et al. Figure 1a Area in Middle-Upper Eocene çe parmak reef mem. ş ş Be Ba İ ZL 29° İ DEN Dazlak mem. REGIONAL LOCATION R İ EH Neogene sediments Ş 1989). ALA LA
et al. Ğ basins MU section line Oligo-Miocene 28° AYDIN fault zmir-Ankara Zone İ R İ ZM İ Simplified geological map of western Anatolia (fiengör & Y›lmaz 1981; fiengör thrusts
Reactivated Çardak–Tokça basin (modified from Göktafl Lycian Nappes N SEA AEGEAN 27° 37° 38° 39° a Figure 1. (a) Quaternary sediments Menderes Massif
157 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA A17 A22 A20 A19 A18 A21
530 534
538 542 fine conglomerate mudstone covered coral medium conglomerate coarse sandstone fine sandstone sandy limestone coal limestone benthic foraminifer bioclast bivalve channel fill planar bedding cross lamination bioturbation coarse conglomerate gastropod barren sample gradational erosive productive sample EOCENE LATE
Age ASAR
02B/38 02B/31 Member
ME Ş ÇE Ş BA EXPLANATIONS Formation 02/49 02/48 02/47 02B/04 A6 A5 A4 A3 A2 A1 A13 A12 A7 A15 A11 A10 A16 A14 A9 A8 02B/05 02B/07 02B/06 02/58 02B/08 02/53 02/54 02/55 02/57 02/56 x
404 432 416 444 424 448 452 420 468 460 EOCENE ?LATE - MIDDLE 408 514 440 502 456 412 526 510 522 436 506 464 428 518 AEEOCENE LATE
Age
MADEN ASAR
Member ME Ş ÇE Ş BA
Formation 02/43 M10 M11 M9 M8 02/44 02/42 02/39 02/40 02/41 02/38 02/46 02/45 02/37
300 364 368 328 396 360 312 308 372 320 356 304 392 376 384 348 316 352 324 332 340 388 380 336 344 IDE-?AEEOCENE ?LATE - MIDDLE
Age MADEN
Member ME Ş ÇE Ş BA
Formation 02B/03 01B/24 02B/28 02B/26 02B/27 02B/23 01B/20 02B/33 02B/32 01B/11 02B/34 01B/33 02B/35 M7 01B/44 02B/31 02B/29 02B/02 02B/01 02B/30 02B/25 02/36 02B/24 M4 M6 M5
260 296 208 220 284 216 268 288 280 248 272 276 264 228 212 204 256 224 200 240 252 236 292 232 244 IDE-?AEEOCENE ?LATE - MIDDLE
Age MADEN
Member ME Ş ÇE Ş BA
Formation 02/21 02/20 02/22
108 104 128 136 140 132 164 160 188 184 124 148 120 192 116 152 176 196 100 112 168 172 156 144 180 IDE-?AEEOCENE ?LATE - MIDDLE
Age DAZLAK MADEN
Member ME Ş ÇE Ş BA
Formation
4 8 Metres
Detailed measured stratigraphic section from the Baflçeflme formation. See Figure 1 for the location of the section. 20 12 16 84 80 32 88 92 40 44 68 60 48 36 64 56 52 96 28 24 72 76 IDE-?AEEOCENE ?LATE - MIDDLE
Age DAZLAK
Member ME Ş ÇE Ş BA
Formation Figure 2.
158 M.S. AKK‹RAZ ET AL.
conglomerates are coarse-grained and poorly sorted. Anatolia, statistical analyses were done using the PAST Channel fills occur at some levels in the sandstones programme developed by Ryan et al. (1995). Non-metric (Figure 2). The Dazlak member was interpreted as multidimensional scaling (MDS), using the Hamming alluvial-fan deposits by fiahbaz & Görmüfl (1992). distance and Raup-Crick index of unweighted pair-group The Maden member, transitional with the underlying average (UPGMA) cluster analysis have been applied Dazlak member, generally consists of yellowish sandstone (Figures 3 & 4). Thin sections were prepared in order to and mudstone alternations and includes conglomerate analyze the foraminifers. and reefal limestone lenses (Figure 2). The Maden In addition, the palynoflora has also been analyzed member, deposited in intertidal environment, also quantitatively using the coexistence approach proposed contains coal seams and lenses. Moreover, shallow- by Mosbrugger & Utescher (1997) for climatic analysis. marine macrofossils – such as gastropods and bivalves – The aim of the coexistence approach is to find the are abundant in the sandstones (Figure 2). intervals of various climate parameters for a given fossil The last member of the Eocene transgressive flora in which a maximal number of nearest living sequence is the Asar member, which generally comprises relatives (NLR) of this flora may coexist. These cream-coloured reefal limestones (Figures 1b & 2). In coexistence intervals are considered the best description some places, the member includes conglomerate, of the palaeoclimatic situation under which the fossil flora sandstone, sandy limestone and mudstone (Figure 2). It lived. The application of the coexistence approach is has abundant macrofossils and microfossils, such as coral, facilitated by use of a computer programme, ClimStat, benthic foraminifers, gastropods and bivalves. It was and the Palaeoflora database. Here, mean annual deposited in an intertidal environment, including an temperature (MAT) was calculated using this method. ecologic reef complex (Göktafl et al. 1989). Lower–“Middle” Oligocene formations unconformably Palynological Data overlie the Asar member in the study area (Göktafl et al. 1989; fienel 1997; fiahbaz & Görmüfl 1992; Sözbilir In this section, the age of the palynological assemblage of 2002; Akkiraz & Akgün 2005). All of these units are the Maden member is discussed. There is no fossil record overlain by Miocene to Quaternary continental deposits in the Dazlak member because of its coarse-grained, (Koçyi¤it 2005; Westaway et al. 2005; Sözbilir 2005). clastic nature. All palynological outcrop samples were obtained from the Maden member (Figure 2). Thirteen of 49 samples investigated for their pollen content contain Materials and Methods spores and gymnosperm and angiosperm pollen grains, A detailed stratigraphic section was measured in the which are fairly well preserved. The flora is characterized Baflçeflme formation; it contained the Dazlak, Maden and by 58 taxa and is here recorded (Table 1). The Asar members (Figures 1b & 2). A total of 49 clay, angiosperm pollen count is always higher in than spores carbonaceous clay and lignite samples were collected and gymnosperms. In addition, Pediastrum spp. and from the Maden member, the only suitable lithology with poorly preserved dinoflagellate cysts were also observed. respect to palynological study (Figure 2). Thirty samples Statistical analyses yield a long list of stratigraphically were also taken from the section for foraminiferal unimportant, long-ranging taxa, but with a few index investigation, from both the Maden and Asar members taxa, for which stratigraphic ranges are given in Table 2. (Figure 2). Ten grams of each sample were treated with In addition, selected palynomorphs recorded in this study
HCL, HF and HNO 3 for palynological preparation. The are also illustrated in Plates I–III. organic residue was screened through an 8 µm mesh Some of the characteristic Early Eocene taxa of screen and 2 and 6 slides per sample of the >8 µm Normapolles, such as Basapollis, Interpollis and fraction were prepared for transmitted light microscopy. Urkutipollenites, are poorly represented in the Lower Pollen counts were carried out at 400X using an Olympus Eocene and do not occur in the Middle Eocene at microscope. Palynological counts range between 94 and Hungarian localities (Kedves 1986). These pollen have 253 grains/specimen (Table 1). To reconstruct the never been recorded in the samples of the Maden palaeoenvironment of the Baflçeflme formation in western member. The stratigraphic distributions of
159 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA 2 2 3 1 1 2 2 12 1 1 1 22 1 MADEN MEMBER 1 113 2 1 1 72 2 51 1 40 2 1 1 23 3 1 1 62 1 1 1 1 5 ) )1119 haploxylon) 47 2 27 9 1 Lygodium )23
Lygodium
)10 Pinus )1 )4 s sp.) 1 )1 ) ) )1 22 ) )1
Osmunda Osmunda Abies alba Abies Pinus silvestris Picea Cathaya Cycas Cycas Cycas Sequoia (Lycopodiaceae: (Lygodiaceae: Schizaeacea( ( Pteridaceae Pteridaceae 1Polypodiaceae 2( (Pinaceae: 2 ( 5( Cupressaceae TaxodiaceaeTaxodiaceae 1 1 ( ( ( ( SparganiaceaeLepidocaryoideae: Palmae 2 1 1 1 1 1 1 1 sp. 1 2 3 2 2 1 1 sp.
sp. Schizaeaceaabsolutus 1 1 Counting results (quantitative) of palynomorphs encountered in the Baflçeflme formation of the Çardak-Tokça basin. cf. sp. ( sp. ( sp. Selaginellaceae 1 sp. sp. 2 sp. sp. sp. Cycadaceae 6 1 4 1 1 1 2 5 1 sp. sp. 2 3 9 29 2 1 36 7 34 1 sp. Arecoideae Table 1. SAMPLESTAXA SPORES 01B/20 01B/44 02B/03 02B/01 02B/02 02B/31 01B/24 01B/11 01B/33 02B/04 02B/05 02B/07 02B/08 GYMNOSPERMOUS Retitriletes Leiotriletes adriennisLeiotriletes minor Leiotriletes wolffii Leiotriletes microadriennis Leiotriletes Triplanosporites microsinuosus Baculatisporis ovalis Baculatisporites nanus (Acrostichum) Baculatisporis Cicatricosisporites Polypodiaceoisporites microconcavus Polypodiaceoisporites kedvesii Polypodiaceoisporites muricinguliformis Polypodiaceoisporites Echinatisporis Laevigatosporites haardti Abiespollenites Abiespollenites Pityosporites microalatus Pityosporites labdacus Pityosporites Piceapollenites Cathayapollis pulaënsis Cathayapollis Inaperturopollenites dubius Inaperturopollenites concedipites Inaperturopollenites magnus Inaperturopollenites ANGIOSPERMOUS MONOCOTYLEDONEAE Cyacadopites gracilis Cyacadopites instasructus Cycadopites ?minimus Cycadopites Arecipites Sequoiapollenites polyformosus Sparganiaceoisporites polygonalis Longapertites discordis Longapertites
160 M.S. AKK‹RAZ ET AL. 2 1116 3 21 42 2 1 1 1 1 1 1 11 1 2 21 1 1 MADEN MEMBER 32 1 2 2 3 5 345 77 132 5 1 2 5 3 3 4 12 1 16 14 2 5 2 4 1 1 206 210 182 200 194 139 253 203 201 183 204 205 94 111 142 5 185 87 102 204 152 170 48 171 161 76 )9651 3)2 1 1111 )2)11 111 111 )4)1 41251 1 sp.) 3 1
Engelhardia Engelhardia )2 111
Myrica Myrica
) Quercus Quercus )16) 111 )111 )11 )36 22 )2 211 )121 ) )1
Nypa ?Carya Tilia Mauritia Ulmus Carpinus Pterocarya Alnus Salix/Platanus Castanea Trigonabalanus Castanea Monocotyledonopsidae (Myricaceae: (Myricaceae: JuglandaceaeJuglandaceae(Juglandaceae: (Juglandaceae: Betulaceae 3( ( 3( Olacaceae 5( ( 6( ( Chenopodiaceae( (Fagaceae: (Fagaceae: Fagaceae 1 Myrtaceae 6 ( 1 1 ( ( Cyrillaceae 1CyrillaceaeOleaceae 2 1 1 1 1 1 1 1 1 3 2
nanus fallax
minor typicus ssp. ssp. ssp. ssp.
stellatus
sp.costatus Pelliciera 9 cf. sp. 2 1 3 1 1 sp. ( sp. cf. sp. spp. sp. sp. Table 1. (continued) SAMPLES 01B/20 01B/44 02B/03 02B/01 02B/02 02B/31 01B/24 01B/11 01B/33 02B/04 02B/05 02B/07 02B/08 DICOTYLEDONEAE Spinizonocolpites Kopekipollenites transdanubicus Triatriopollenites rurensis Triatriopollenites bituitus Triatriopollenites excelsus Triatriopollenites excelsus Plicatopollis lunatus Plicatopollis plicatus Plicapollis Momipites punctatus Momipites quietus Momipites Triporopollenites robustus Labrapollis labraferus Triporopollenites Trivestibulopollenites betuloides Subtriporopllenites anulatus Subtriporopllenites constans Intratriporopollenites indubitalibis Mauritiidites franciscoi Olaxipollis matthessi Polyporopollenites undulosus Polyporopollenites carpinoides Polyporopollenites Polyvestibulopollenites verus Pentapollenites pentangulus Chenopodipollis multiplex Tricolpopollenites retiformis Tricolpopollenites microhenrici Tricolpopollenites asper Tricolpopollenites liblarensis Tricolpopollenites Polycolpites Myrtaceidites mesonesus Tricolporopollenites cingulum oviformis Tricolporopollenites cingulum fusus Tricolporopollenites cingulum pusillus Tricolporopollenites megaexactus Tricolporopollenites pseudocingulum Tricolporopollenites microreticulatus Tricolporopollenites Psilatricolporites crassusPsilatricolporites Nowemprojectus tumanganicus Pediastrum INCERTAE CEDIS Dinoflagellate cysts Pelliciera Total
161 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
02B/04 + 02B/05 A + 02B/07 ++ 02B/08 + + 01B/33 + + 02B/01 + + 01B/24 + + B 02B/31 + + 02B/02 + + 01B/11 + + 01B/44 + + C 01B/20 + 02B/03 + Lowland-Riparian iolglaecysts Dinoflagellate rcihwater Brackish Freshwater Mangrove Montane
1 2
KEY: > 0,5 %: + 0,5-5%: 5-10%: 10-25%: 25-40%: < 40%:
Figure 3. Dendograms for UPGMA cluster analysis of palaeocommunities (bottom) and samples (side), using the Raup-Crick Index. See Figure 2 for the locations of the samples.
Triatriopollenites excelsus, Subtriporopollenites anulatus recorded in sediments older than the Middle Eocene ssp. nanus, Subtriporopollenites constans, Plicatopollis (Nickel 1996; Chateauneuf 1980; Hochuli 1978; Vinken lunatus and Nowemprojectus tumanganicus are restricted 1988). to the Palaeocene and Eocene the world over (Table 2). In addition to the taxa mentioned above, mangrove Nowemprojectus tumanganicus is here recorded for the and back-mangrove elements such as Psilatricolporites first time from the Eocene sediments of Turkey. crassus (Pelliciera), Spinizonocolpites group ( Nypa) and Additionally, Aglaoreidia cyclops appears in the Middle Leiotriletes adriennis (Acrostichum) and Mauritiidites Eocene and seems have its last occurrence within or at franciscoi (Mauritia) were first recorded in western the top of the ‘Middle’ Oligocene. Thus, this species is not Anatolia. According to Rull (1998a, 1999), the Early
162 M.S. AKK‹RAZ ET AL.
0,3
Marine samples 0,2 02B/08 A1- 22 Cluster A 0,1 02B/04
02B/07 02B/05 0 01B/33 02B/31 02B/02 02B/01 Cluster C
Axis 2 01B/24 01B/11 Cluster B -0,1 01B/20
01B/44 -0,2 02B/03
-0,2 -0,1 0 0,1 0,2 0,3 Axis 1
Figure 4. Scattergram of taxa ordination, from a two-dimensional MDS using the Hamming distance. Stress = 0.3028.
Eocene occurrence of Psilatricolporites crassus (Pelliciera) Comparison of Palynological Data with Other Coeval and Spinizonocolpites echinatus (Nypa) is always sparse Basins in Turkey as compared to the Middle Eocene of the Venezuelan To date, only limited palynological studies on the Eocene Maracibo basin; that observation is consistent with the units have been made in the Yozgat and Çorum areas of data of this study. Most European Nypa pollen central Anatolia (Nakoman 1966; Akyol 1980; Akgün et appearances are in Ypresian–Cuisian deposits of France al. 2002; Akgün 2002). The Turkish Eocene microflora (Gruas-Cavagnetto 1977), except for Spanish was first examined by Nakoman (1966). That author occurrences that range from Cuisian to Early Lutetian accepted that the age of the sediments in the Yozgat area (Haseldonckx 1972) in age. Cavagnetto & Anadón (1996) (Sorgun lignites) was Early Eocene based on the presence describe complex mangrove-swamp elements, such as of important biostratigraphic forms such as Nypa, Avicennia and Pelliciera from the Middle Bartonian Triatriopollenites excelsus, Tetracolporopollenites of the eastern Ebro basin (northern Spain). According to biconus, Inaperturopollenites echinatus and Riegel et al. (1999), the mangrove association is Monocolpopollenites granulatus. widespread and diverse in the Middle Eocene seam group relative to the Lower Eocene seam group at Helmstedt, Akyol (1980) studied the palynological aspects of the northern Germany. The oldest biodiversification age of Bayat lignites (nearby Çorum area of central Anatolia). the mangroves on the European sea-shores is Middle That author determined a rich flora, including some Eocene (Plaziat et al. 2001). Based on the palynological forms that have biostratigraphic significance (see Akgün data, the Maden member should have been deposited et al. 2002 for discussion). A Middle–Late Eocene age is during the Middle–?Late Eocene period (Figure 2). suggested on the basis of palynological and foraminiferal data. Although Nakoman (1966) and Akyol (1980) recognized some biostratigraphically significant species
163 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
Table 2. Stratigraphic distribution of selected taxa encountered in the Baflçeflme formation.
Fossil Age References
Leiotriletes adriennis Middle Eocene–Middle Miocene Stuchlik et al. 2001
Leiotriletes wolffii Late Eocene–Pliocene Nickel 1996; Stuchlik et al. 2001
Aglaoreidia cyclops Middle Eocene–Middle Oligocene Nickel 1996 Early–Middle Oligocene Roche 1988; Hochuli 1978; Ollivier-Pierre et al. 1993
Spinizonocolpites Group Eocene Vinken 1988 Middle Eocene–Early Oligocene Elsik 1974; Frederiksen 1973
Triatriopollenites excelsus Paleocene–Late Eocene Thomson & Pflug 1953; Kedves 1969, 1982; Akyol 1978;Gruas-Cavagnetto 1978; Nickel 1996
Plicatopollis lunatus Eocene–Middle Oligocene Nickel 1996 Paleocene–Early Eocene Frederiksen 1980a Middle Eocene Thiele-Pfeiffer 1988 Late Paleocene–Eocene Akgün 2002
Subtriporopollenites anulatus ssp. nanus Late Palaeocene–Late Eocene Thomson & Pflug 1953; Krutzsch 1957, 1970; Krutzsch & Vanhoorne 1977 Subtriporopollenites constans Palaeocene–Early Eocene Kedves 1970; Krutzsch & Vanhoorne 1977 Middle Eocene Thiele-Pfeiffer 1988 Middle Eocene–Middle Oligocene Krutzsch 1970 Palaeocene–Early Oligocene Gruas-Cavagnetto 1978
Nowemprojectus tumanganicus Paleocene–Middle Eocene Bolotnikova (1979) Eocene Frederiksen et al. (2002)
for the Turkish Eocene, they did not document any of the determined as individual grains from the Maden member mangrove species which have been determined in the (western Anatolia) (sample 02B/07). Diversification of present study. pteridophytic spores, tricolpate and tricolporate pollen After those palynological studies at different localities grains is high in samples from the Yoncal› formation in central Anatolia, Akgün et al. (2002) and Akgün (central Anatolia) compared to the Maden member (2002) obtained a rich species diversification. They also (western Anatolia). The species Nowemprojectus obtained new Eocene palynological data and suggested tumanganicus has been determined in samples of the the presence of different kinds of mangrove and back- Maden member (Table 1) but does not occur in the mangrove elements, such as Avicennia, Psilatricolporites Yoncal› formation. The mangrove element (Pelliciera), Spinizonocolpites (Nypa), Mauritiidites Psilatricolporites crassus (Pelliciera) is abundant in almost franciscoi (Mauritia) and Leiotriletes adriennis all samples from the Maden member (54–90%) (Table (Acrostichum) from the Middle–?Upper Eocene Yoncal› 1), but is less abundant in samples from the Yoncal› formation in central Anatolia (Yozgat and Amasya areas). formation. Although the percentages of mangrove The species Spinizonocolpites echinatus, S. prominatus, S. elements are different in central and western Anatolian baculatus, S. gemmatus, S. microgemmatus, S. samples, the Middle–?Upper Eocene coal occurrences of bulbospinosus S. cf. wodehousei, S. indicus, S. cf. central Anatolia can be correlated with the data of this adamenteus and Spinizonocolpites spp. were abundantly study on the basis of the presence of mangrove elements. recorded in samples of the Yoncal› formation (central The difference in relative frequencies of the species is Anatolia). In contrast, Spinizonocolpites spp. has been likely related to the palaeoecological factors which persisted during the deposition of these coals.
164 M.S. AKK‹RAZ ET AL.
Foraminiferal Data and Age Determination summary, the Beflparmak reef, Maden and Asar members Thirty samples were collected from both the Maden and were deposited during Bartonian–Priabonian time in the Asar members of the Baflçeflme formation along the study area. However, it is difficult to interpret the age of section (Figure 2). However, only 22 samples were found the Dazlak member, which is located at the base of the to be micropalaeontologically productive; samples from Baflçeflme formation, due to the lack of fossils. According the limestones of the Maden member are barren (the to fienel (1997), Eocene sedimentation in the Çardak- samples of M4–M11) (Figure 2). Tokça basin began in the Late Lutetian, an age which may be accepted doubtfully for the Dazlak member (Figure 2). However, Sözbilir et al. (2001) studied the microfauna of the Eocene sediments cropping out in the southeastern part of the Baklan area, located 10 km Reconstruction of the Palaeoenvironment northeastern of the study area (Figure 1b). Those The palynological assemblage of the Baflçeflme formation authors obtained well-preserved foraminiferal data from in the Çardak-Tokça basin is limited due to conditions of the Beflparmak reef member, which is conformably sedimentation, and exhibits some differences in the overlain by the Maden member. The well-preserved fauna frequency of the same taxa in different samples (Table 1). comprises Asterigerina rotula, Nummulites perforatus, To reconstruct the palaeoenvironment during the Nummulites beaumonti, Fabiania cassis, Europertia deposition of the Baflçeflme formation, definite ecological magna, Chapmanina gassinensis, Halkardia minima, marker taxa have been selected from the published Silvestrialla tedraedra, Rotalia sp., Globigerina sp., literature. The ecological characteristics of species have Operculina sp., Heterostsegina sp., Assilina sp., Gypsina been grouped under generic headings, such as sp., Planorbulina sp., Textularia sp., Discocyclina sp., “mangroves” ( Nypa and Pelliciera) and “fresh-water Hauerinidae, Rotaliidae and algae. A Bartonian age was elements” ( Aglaoreidia cyclops, Sparganiaceae and suggested by those authors, and is in agreement with our Pediastrum spp.). The rhizomatic palm Nypa is an palynological data for the Maden member. ecological marker of mangrove vegetation (Germeraad et In this study, samples (A1–A22) collected from the al. 1968; Jiménez 1984; Frederiksen 1985; Asar member are rich in benthic foraminifers (Figure 2). Thanikaimoni 1987; Westgate & Gee 1990; Srivastava & A Nummulites assemblage, including Nummulites fabianii Binda 1991; Graham 1995; Rull 1998a; Lenz & Riegel and Nummulites striatus , has been determined. In 2001). Pelliciera is another mangrove element (Jiménez addition, Fabiania cassis, Eorupertia magna, Halkyardia 1984; Rull 1998a). According to Frederiksen (1985), minima, Baculogypsinoides tetraedra, Asterigerina rotula, Aglaoreidia cyclops was found to favour fresh-water Quinqueloculina sp., Asterigerina sp., Discocyclina sp., habitats; e.g., Azolla, Pediastrum, Botryococcus, Heterostegina sp., Eponides sp., Alveolina sp. Potamegaton, Sparganium and Typa. Thus, the species Praebulalveolina sp., Neoalveolina sp., Anomalina sp., was thought to have been produced by fresh-water Mississippina sp., Pararotalia sp., Pyrgo sp. and habitats (Fowler 1971; Frederiksen 1985; Riegel et al. Orbitolites sp. have also been recorded in the samples. 1999). Rotaliidae, Textulariidae, Miliolidae, algae, bivalves, From the palaeovegetational point of view, the gastropods and echinoderms have been documented as palaeocommunities and samples can be divided into two well. The foraminiferal fauna suggests a Late Eocene or three assemblages (Figure 3). In the (Priabonian) age, indicating a carbonate-shelf depositional palaeocommunities dendogram, assemblage 1 consists of environment for the Asar member. mangroves, brackish-water elements and dinoflagellate On the basis of foraminiferal data, Sözbilir et al. cysts. Assemblage 2 includes fresh-water, lowland- (2001) suggested that the age of the Beflparmak reef riparian and montane elements (Figure 3). member is Bartonian. In addition, a In the sample dendogram, assemblage A is Bartonian–?Priabonian age has been suggested for the characterized by a dominance of mangroves, brackish- Maden member on the basis of palynostratigraphic data. water elements and low frequencies of dinoflagellate Also, a Priabonian age is assigned on the basis of well- cysts and montane elements, corresponding to the constrained foraminiferal data from the Asar member. In palaeocommunity cluster 1 (Figure 3). The samples of
165 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
cluster A coincide with samples taken from the upper part (Table 1). The combination of Psilamonocolpites sp., of the section (Figure 2). The presence of poorly Mauritiidites franciscoi (Mauritia) and Leiotriletes preserved dinoflagellate cysts and the abundance of adriennis (Acrostichum) , which are back-mangrove mangrove elements in these samples indicate close swamps, have been observed as individual grains in some proximity to a marine environment (Figures 3 & 5). The samples. Their presence in the Tertiary of Venezuela has samples taken from the lower part of the section lack been interpreted as representative of the wetlands behind dinoflagellate cysts (Figures 2, 3 & 5). mangroves near the limit of tidal influence (Rull 1992, Sample cluster B is represented by the dominance of 1997a, b, 1998a, b). A Mauritia palm forest associated mangrove and the absence of fresh-water elements and with ferns indicates a zone of fresh-water (to locally dinoflagellate cysts. The lowland-riparian and montane brackish) swamps and marshes in the Eocene of Colombia elements in sample cluster B are higher than in cluster A (González-Guzmán 1967). The pollen of Mauritia grew (Figures 3 & 5). back of the coastline (Van Der Hammen & Wijmstra 1964); its presence in the sediments is a reliable indicator The samples of cluster C were taken from the lowest of a warm tropical lowland environment flooded by fresh part of the section (Figure 2) and are represented by the (sometimes oligohaline) waters (Rull 1998b). In addition, presence of fresh-water elements, high frequencies of Mauritiidites may also be observed in more inland mangrove, lowland-riparian and montane elements, and associations. The scarce presence of Nowemprojectus the absence of dinoflagellate cysts (Figures 3 & 5), tumanganicus in samples 02B/03, 2B/04 and 02B/05 corresponding to cluster 2 in the palaeocommunity should be indicative of peaty substrates (Frederiksen et dendogram (Figure 3). There are low numbers of al. 2002) (Table 1). The pollen of Chenopodiaceae may mangrove elements in sample cluster C as compared to endure droughts in salt marshes (Figure 5). clusters A and B (Figure 5). Pteridophytic spores of Polypodiaceae, Schizaceae, All micropalaeontological samples have also been Selaginellaceae, Lygodiaceae and Osmunda are dispersed subjected to non-metric multidimensional scaling (MDS) in the samples indicating the prevalence of perennial using the Hamming distance method (Figure 4). The MDS water in the brackish-water palaeocommunity (Figure 5). shows results similar to those of the cluster analysis. The The presence of Sparganiaceaepollenites (Sparganiaceae), MDS of the samples (Figure 4) shows that groups Pediastrum spp. and Aglaoreidia cyclops in samples identified in the cluster analysis have also been recognized 02B/03, 01B/20 and 01B/44 correspond to sample in this plot. The samples, which are situated at the cluster C and may be considered as the fresh-water positive part of the second axis, show a close proximity to palaeocommunity (Figures 3 & 5). Almost all samples the marine environment based on the presence of marine include lowland-riparian elements such as Juglandaceae, samples (A1–A22); also, cluster A includes dinoflagellate Betulaceae, Engelhardia, Fagaceae and Myrtaceae. cysts, mangroves and brackish-water elements (Figures 3 & 4). The samples at the negative part of the second axis Montane elements are represented by Pinus, Abies, Picea, indicate relatively terrestrial conditions. Cathaya, Quercus and Castanea (Figure 5). The frequencies of lowland-riparian and montane elements Mangrove element Pelliciera (Psilatricolporites and also the presence of fresh-water elements in crassus) is found in abundance in all samples (Table 1). palaeocommunity cluster 2 (which corresponds to the According to Jiménez (1984), Pelliciera is more sensitive sample cluster C) may indicate low sea levels (Figures 3, to high soil salinities than other more widespread 4 & 5). The deposition of these samples in cluster C may neotropical mangroves. It develops best on wet soils, have occurred in a non-marine, back-mangrove shallowly inundated at high tides (Collins et al. 1977). It environment. The abundance of mangrove brackish- also thrives on firm, sandy slightly elevated soils which water elements and the presence of poorly preserved are located in inter-channel areas (Fuch 1970; Jiménez dinoflagellate cysts in sample cluster A indicate high sea 1984). Nypa, pollen of which is observed as individual levels (Figures 3 & 5). Samples of clusters A and B must grains in sample 02B/07, occurs in permanently brackish have been deposited in a mangrove environment (Figure humid soil on the proximal fringes of mangrove fronts 5). (Blasco 1977; Chapman 1976; Srivastava & Binda 1991)
166 M.S. AKK‹RAZ ET AL.
me fm. Marine ş S M BW LR Mo D F PALAEOECOLOGY çe ş Transgression Ba
SHALLOW MARINE Priabonian Asar member
SHALLOW MARINE MANGROVE BRACKISH LOWLAND-RIPARIAN MONTANE Nypa Ferns Quercus Pelliciera Palms Castanea Pinus Fagaceae Myricaceae Taxodiaceae 02B/08
02B/07 Cluster A 02B/05
02B/04
01B/33 SHALLOW MARINE MANGROVE BRACKISH LOWLAND-RIPARIAN MONTANE Pinus Ferns Quercus Castanea 01B/11 Pelliciera Palms Mauritiidites franciscoi Fagaceae Fagaceae Myricaceae Taxodiaceae 01B/24
Maden member 02B/31 Cluster B
02B/02 Bartonian - ?Priabonian
02B/01
SHALLOW MARINE MANGROVE BRACKISH LOWLAND-RIPARIAN MONTANE FRESH WATER Pinus Sparganiaceae Quercus Castanea Picea Ferns Pelliciera Pediastrum Fagaceae Abies Palms 02B/03 Aglaoreidia cyclops Chenopodiaceae Myricaceae Fagaceae Taxodiaceae 01B/44
01B/20 Cluster C
Dinoflagellate cysts Rotaliidae Textulariidae Miliolidae Nummulites Bivalves Gastropods Echinoderms Corals Algae coal S: sample, M: Mangrove, BW: brackish water, LR: lowland-Riparian, Mo:Montane, D: Dinoflagellate cysts, F: Fresh water
% 20 80 20 40 20 20 60 5 3
Figure 5. Relative abundance of ecological groups obtained from the Maden member and palaeoenvironmental reconstruction of the Çardak- Tokça basin (western Anatolia) during the deposition of the coal-bearing Middle–Upper Eocene Baflçeflme formation (Maden and Asar members).
In summary, it is clear that Middle–Late Eocene were attained during deposition of the Asar member transgression is recorded from cluster C to cluster A (Figure 5). The marine transgression is also well (Figure 5). The presence of marine limestones in the documented by the presence of reefal limestones study area indicates that maximum sea-level conditions containing rich marine fossils, such as coral reefs, benthic
167 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
foraminifers and echinoderms (Figure 5). In western 6). The calculated coexistence intervals result in a mean Anatolia, there has been no study focused on the annual temperature (MAT) range mainly between 24.8 Bartonian–Priabonian transgression. However, on a and 25 °C, but intervals between 17.2 and 21.1 °C also regional scale, the Palaeocene–Eocene successions of occur (Figure 6). Thus, two different mean annual western Anatolia were probably associated with temperatures (24.8–25 °C and 17.2–21.1 °C) have been depressions (Sözbilir 2002). obtained by applying the coexistence approach proposed by Mosbrugger & Utescher (1997). For this reason, Nix’s (1982) terms megatherm and mesotherm are quite Palaeoclimate relevant. Nix (1982) recognized plant groups for Quantitative terrestrial palaeoclimatic analysis based on characteristic temperature response models for Australia the palynological assemblage of the Maden member and New Guinea. The megatherm element is dominant (Baflçeflme formation) was carried out using the where the mean annual air temperature exceeds 24 °C coexistence approach proposed by Mosbrugger & and corresponds to the environment of the mangrove Utescher (1997); this technique is based on the ‘nearest association. The mesotherm element is dominant where living relative’ philosophy, the assumption that climatic mean annual air temperature ranges between 14 °C and requirements of Tertiary plant taxa are similar to those of 21 °C and coincides with the slope and montane-forest their living relatives (Mosbrugger & Utescher 1997). The association. An intermediate zone is indicated as an area palaeoclimatic reconstruction of the palynoflora of the where the mean annual temperatures are less than 24 °C Baflçeflme formation is based on a total of 19 taxa (Figure but greater than 21 °C. The obtained climatic values
MAT C Fossil Taxa Nearest Living Relatives Polyvestibulopollenites verus Alnus sp. Arecipites sp. Arecoideae sp. Inaperturopollenites concedipites Taxodiaceae Inaperturopollenites dubius Cupressaceae Momipites punctatus Engelhardia sp. Triatriopollenites rurensis Myrica sp. Tricolpopollenites henrici Quercus sp. Plicatopollis plicatus Juglandaceae Tricolpopollenites microhenrici Fagaceae Cyacadopitessp . Cycadaceae Longapertites discordis Lepidocaryoideae Tricolporopollenites microreticulatus Oleaceae Abiespollenites sp. Abies sp. Chenopodipollis multiplex Chenopodiaceae Piceapollis sp. Picea sp. Tricolporopollenites cingulumssp. fusus Trigonabalanus sp. Trivestibulopollenites betuloides Betulaceae Cathayapollis pulaënsis Cathaya sp. Polyporopollenites carpinoides Carpinus betulus, C. coraliniana -20 -10 0 10 20 30 Bordering taxa Coexistence interval(s) 17.2 - 21.1 C Trigonabalanussp. - Carpinus betulus, Carpinus caroliniana 24.8 - 25 C Lepidocaryoidae - Taxodiaceae
Figure 6. Application of the coexistence approach to the palynoflora of the Maden member in the Çardak-Tokça basin (western Anatolia). The shaded boxes mark the climatic requirements of the taxa, the vertical lines delimit the widths of coexistence intervals (MAT: mean annual temperature).
168 M.S. AKK‹RAZ ET AL.
correspond to the terminology of Nix (1982). Climatic Early Eocene is marked by some decrease in the mean conditions should be 24.8–25 °C near the coast, including annual temperatures in what is present-day Belgium, the megathermal elements (mangrove association). The Germany and the East European platform, later second MAT value (17.2–21.1 °C) corresponds to Nix’s succeeded in Europe by a warm tropical and subtropical mesotherm zone. The intermediate zone is characterized climate (Buchardt 1978; Yasamanov 1982). by a temperature of 21.1–24.8 °C. Nix (1982) used Palaeobotanical determinations of the Middle Eocene these terms to do away with the confusion that is usually palaeoclimate (Gray 1960; Dilcher 1973; Wolfe 1978; associated with the use of tropical, subtropical and Upchurch & Wolfe 1987; Greenwood & Wing 1995) and temperate, since these have both geographic and thermal of the Late Eocene palaeoclimate (Frederiksen 1980b; connotations. On the other hand, according to Wolfe Wolfe 1992) for the northern Gulf Coast, U.S.A. are all (1979), the moist tropical forests have an MAT higher winter-dry tropical to humid paratropical climates on than 25 °C; the 20–25 °C isotherms lie in the region of lowlands, and most infer no freezing. Frederiksen development for the paratropical forest; broad-leaf (1980a, b) suggested a regime of winter-dry tropical evergreen forests grow in a subtropical climate (13–20 climate on lowlands adjacent to the coastline and a °C). The occurrence of Nypa pollen suggests that tropical marginal humid paratropical climate on the upper coastal to subtropical climatic conditions existed during plain. In conclusion, we have identified the presence of a deposition of the Baflçeflme formation. If the habitat mixture of subtropical/paratropical climatic zones, requirements of Nypa during the Eocene were similar to corresponding to an interval having mean annual those of extant Nypa, water temperatures must have temperatures 17.2° and 25 °C, and from coast to been warmer than 24 °C. Nypa cannot survive in montane environments. This climatic discrepancy is temperatures less than 20 °C (Fechner 1988). The related to the effects of the tropical Tethys Sea on species of Nypa, Avicennia and Pelliciera are among the terrestrial environments. megathermal taxa (Cavagnetto & Anadón 1995, 1996). Psilatricolporites crassus (Pelliciera) in particular has been recorded with great frequency in our samples as a Palaeogeographic Data megathermal species. The mangroves need the tropical On the basis of previous studies and the data of the and humid climate to develop (Frederiksen 1985; present study, the present-day and Eocene geographic Westgate & Gee 1990). In addition, the presence of the distributions of mangrove elements Nypa and Pelliciera zooxanthellate coral fauna with larger foraminifera in the have been plotted on maps (Figure 7). The pollen of Nypa carbonate rocks indicates the tropical temperatures of the (Spinizonocolpites) and Pelliciera (Psilatricolporites Tethys Sea. The zooxanthellate coral fauna and reef crassus), which were first recorded from Middle–?Upper growth with larger foraminifera control the minimum Eocene deposits in central Anatolia (Yozgat and Amasya sea-surface temperature (Schmiedl et al. 2002). Ediger et areas) by Akgün (2002) and Akgün et al. (2002), have al. (1990) studied the palynological properties of the been cited in the section on palynological data. Nypa was Tertiary of the northern Thrace Basin (northwestern present on all continents during the Eocene but, at Turkey), and discussed the palaeoclimate of the Eocene present, only occurs in the Indo-Malaysian region (Figure period. According to those authors, the increased 7). Conversely, Pelliciera (Psilatricolporites crassus) percentages of thermophilous elements, such as occurred in the Caribbean area and on the Atlantic coasts Cyacadopites, Monocolpopollenites and Dicolpopollenites of Guyana and Brazil during Eocene–Oligocene time (Rull kalewensis, indicate that the temperature was higher in 1998a); it was also recorded from the Middle–Late the Oligocene than in the Eocene, probably resulting in a Eocene sediments of the Ebro basin (northeast Spain) by temperate climate in the Eocene and a subtropical climate Cavagnetto & Anadón (1995, 1996). Although its in the Oligo–Miocene. However, they also noted that the presence in Africa is uncertain during Eocene time, it has Eocene part of the climate curve should be drawn with been reported from Middle–Upper Eocene sediments in caution on the basis of previous palaeoclimatic studies both central (Yoncal› formation) and western Anatolia (Wolfe 1978; Hochuli 1984). Throughout the Eocene (Baflçeflme formation) (Figure 7). It was also reported epoch, the climate of the Mediterranean region from the Tertiary of the Guiana basin (Van Der Hammen underwent fluctations (Aleksandrova et al. 1987). The & Wijmstra 1964), and from Early Miocene sediments of
169 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
Spinizonocolpites (Present)
? ? Pelliciera (Present)
Figure 7. Present (dashed areas) and Eocene (black squares) geographic distributions of Spinizonocolpites spp. ( Nypa) and Psilatricolporites crassus (Pelliciera) . After Germeraad et al. (1968), Müller (1980, 1981), Thanikaimoni et al. (1984), Frederiksen (1980a, 1985, 1988, 1994), Thanikaimoni (1987), Westgate & Gee (1990), Srivastava & Binda (1991), Cavagnetto & Anadón (1995), Graham (1995), Nickel (1996), Pole & Macphail (1996), Rull (1998a, 1999), El Beialy (1998), Riegel et al. (1999), Akgün (2002), Akgün et al. (2002).
Panama (Graham 1977). According to Graham (1995), waters. At the global scale, since the tropical Tethys free its presence persisted into the Quaternary Gulf/Caribbean seaway connected the Palaeo-Atlantic and Indian oceans, region (Mexico, the Antilles, Central America, and the mangroves should have invaded southeastern Asian, northern South America). However, at present, Pelliciera African, North and South American and European shores. (Psilatricolporites crassus) is observed in a restricted area of central and northern South America (Figure 7). In this study, the present Atlantic mangrove ( Pelliciera) and General Conclusions present Indo-Pacific mangrove ( Nypa) elements have not The results of this study are as follows: (1) Palynological been reported as fossils from western Anatolia. Their and benthic foraminiferal assemblages indicate a presence in Middle–Upper Eocene sediments of western Bartonian–Priabonian age for the Maden and Asar and central Anatolian implies warm mid-latitude Tethys members of the Baflçeflme formation. (2) An overall rise
170 M.S. AKK‹RAZ ET AL.
in sea level from the Bartonian to the Priabonian is 17.2–21.1 °C in the mesotherm zone. These climatic indicated by the distribution of both microfaunal and intervals indicate subtropical/paratropical climatic palynological associations from the back-mangrove to conditions during deposition of the Baflçeflme formation. shallow-marine environment. The lower part of the (4) The circulation of warm Tethys waters was probably Maden member was deposited in a non-marine, back- responsible for the growth of mangroves in both central mangrove environment. The palaeoenvironment changed and western Anatolia during the Middle–Late Eocene. from back-mangrove to mangrove or front-mangrove environment in the upper part of the Maden member and also to a shallow-marine environment in the Asar Acknowledgements member, which contains marine fossils. Based on the This study was supported by research project grants from palynological and microforaminiferal data, sea-level rise the Scientific and Technical Research Council of Turkey likely was related to basin subsidence as a result of (TÜB‹TAK), No. 101Y133, and from Dokuz Eylül tectonic development. (3) The mangrove taxa and coral University, Graduate School of Natural and Applied fauna with larger foraminifera in the carbonate strata of Sciences, No. AFS–02. KB. FEN.046 awarded to Mehmet the Baflçeflme formation indicate tropical water Serkan Akkiraz and Funda Akgün, and by a DAAD temperatures during the Bartonian–Priabonian interval. short–term fellowship awarded to Mehmet Serkan Palaeoclimatic conditions are characterized by 24.8–25 Akkiraz; all are gratefully acknowledged. Valenti Rull, °C MAT values in the mangrove environment, whereas Zühtü Bat› and the editor, Erdin Bozkurt, are thanked for there was a mean annual temperature of 21.1–24.8 °C in their constructive criticisms of the manuscript. the megatherm/mesotherm intermediate zone and of
References AKGÜN, F. 2002. Stratigraphic and paleoenvironmental significance of BOZKURT, E. & PARK, R.G. 1999. The structure of the Palaeozoic schists Eocene palynomorphs of the Çorum-Amasya area in the central in the southern Menderes Massif, western Turkey: a new Anatolia, Turkey. Acta Palaeontologica Sinica 41, 576–591. approach to the origin of the main Menderes metamorphism and its relation to the Lycian Nappes. Geodinamica Acta 12, 25–42. AKGÜN, F., A KAY, E. & E RDO⁄AN, B. 2002. Terrestrial to shallow marine deposition in central Anatolia: a palynological approach. Turkish BUCHARDT, B. 1978. Oxygen isotope paleotemperatures from the Journal of Earth Sciences 11, 1–27. Tertiary period in the North Sea area. Nature 275, 121–123.
AKK‹RAZ, M.S. & A KGÜN, F. 2005. Palynology and age of the Early CAVAGNETTO, C. & A NADÓN, P. 1995. Uno mangrove complexe dans le Oligocene units in Çardak-Tokça basin, southwest Anatolia: Bartonien du basin de le l’Ebre (Ne de l’Espagne). paleoecological implications. Geobios 38, 283–299. Palaeontographica Abteilung B Ionnides 236, 147–165.
AKYOL, E. 1978. Stratigrafik palinoloji, kömür iflletmecili¤i ve bir örnek CAVAGNETTO, C. & A NADÓN, P. 1996. Preliminary palynological data on [Stratigraphic palynology, coal mining and a case study]. Jeoloji floristic and climatic changes during the Middle Eocene- Early Mühendisli¤i Dergisi 6, 31–38 [in Turkish with English abstract]. Oligocene of the eastern Ebro Basin, northeast Spain. Review of Palaeobotany and Palynology 92, 281–305. AKYOL, E. 1980. Etude palynologique de l'Eocene de Bayat (Çorum- Turquie) et essai de correlation entre Karakaya et Emirflah. CHAPMAN, V.J. 1976. Mangrove Vegetation. J. Cramer, FL-9490 Vaduz, Mineral Research and Exploration Institute of Turkey (MTA) 447 p. Bulletin 91, 39–53. CHATEAUNEUF, J.J. 1980. Palynostratigraphie et Paléoclimatologie de l’ ALEKSANDROVA, A.N., P ROZOROV, Y.I. & Y ASAMANOV, N.A. 1987. Climatic Eocène Supérieur et de l’ Oligocène du Bassin de Paris. Mémorie and floristic zonation of the Mediterranean region during the du Bureau de Recherches Géologiques et Minières 116, 1–360. Early Cenozoic time. International Geology Review 29, 503–514. COLLINS, J., BERKELHAMER, R.C. & MESLER, M. 1977. Notes on the natural BLASCO, F. 1977. Outlines of ecology, botany and forestry of the history of the mangrove Pelliciera rhizophorae Tr. & Pl., mangals of the Indian subcontinent. In: C HAPMAN, V.J. (ed), (Theaceae). Bresnia 10/11, 17–29. Ecosystems of the World. Wet Coastal Ecosystems 1, 241–260. COLLINS, A.S. & ROBERTSON, A.H.F. 1997. The Lycian Mélange, southwest BOLOTNIKOVA, M.D. 1979. Spore-Pollen Complexes of Tertiary Deposits Turkey: an emplaced accretionary complex. Geology 25, of the Western Coast of the Sea of Japan. “Science” Publishing 255–258. House, Moscow. COLLINS, A.S. & R OBERTSON, A.H.F. 1998. Processes of Late Cretaceous BOZKURT, E. & PARK, R.G. 1994. Southern Menderes Massif: an incipient to Late Miocene episodic thrust sheet translation in the Lycian metamorphic core complex in western Anatolia, Turkey. Journal Taurides, SW Turkey. Journal of the Geological Society, London of the Geological Society, London 151, 213–216. 155, 759–772.
171 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
COLLINS, A.S. & R OBERTSON, A.H.F. 1999. Evolution of the Lycian GÖKTAfl, F., ÇAKMAKO⁄LU, A., TAR›, E., SÜTÇÜ, Y. F. & S AR›KAYA, H. 1989. allochthon, western Turkey, as a north-facing Late Palaeozoic to Çivril–Çardak Aras›n›n Jeolojisi [Geology of Çivril-Çardak Region]. Mesozoic rift and passive continental margin. Geological Journal Mineral Research and Exploration Institute of Turkey (MTA) 34, 107–138. Report 8701, 107 p [in Turkish, unpublished].
DILCHER, D.L. 1973. A palaeoclimatic interpretation of the Eocene floras GRAHAM, A. 1977. New records of Pelliceria (Theaceae/Pelliceriaceae) in southeastern North America. In: GRAHAM, A. (ed), Vegetation and the Tertiary of the Caribbean. Biotropica 9, 48–52. Vegetational History of Northern Latin America. Elsevier GRAHAM, A. 1995. Diversification of Gula/Caribbean mangrove Publishing Company, 39–59. communities through Cenozoic time. Biotropica 27, 20–27. ED‹GER, V.fi., B AT›, Z. & A L‹flAN, C. 1990. Paleopalynology and GRAY, J. 1960. Temperate pollen genera in the Eocene (Claiborne) flora, paleoecology of Calamus like disulcate pollen grains. Review of Alabama. Science 132, 808–810. Palaeobotany and Palynology 62, 97–105. GREENWOOD, D.R. & W› NG, S.L. 1995. Eocene continental climates and EL BE›ALY, S.Y. 1998. Stratigraphic and palaenvironmental significance of Eocene palynomorphs from the Rusayl Shale Formation, Al latitudinal temperature gradients. Geology 23, 1044–1048. Khawd, northern Oman. Review of Palaeobotany and Palynology GRUAS–CAVAGNETTO, C. 1977. Étude palynologique de l’Eocéne du Bassin 102, 249–258. Anglo-Parisien. Thèse de Doctorate d’État és Sciences Naturelles, Universite de Pierre Marie Curie VI, 287 p. ELS›K, W.C. 1974. Characteristic Eocene palynomorphs of the Gulf Coast, U.S.A. Palaeontographica Abteilung B Ionnides 149, GRUAS-CAVAGNETTO, C. 1978. Étude palynologique de l’Eocéne du Bassin 90–111. Anglo-Parisien. Mémoire de la Société Géologique de France
FECHNER, G.G. 1988. Selected palynomorphs from the Lower to Middle 131,1–64. Eocene of the South Atlas Border Zone (Morocco) and their HASELDONCKX, P. 1972. The presence of Nypa palm in Europe: a solved environmental significance. Palaeogeography, Palaeoclimatology, problem. Geologie en Mijnbouw 51, 645–650. Palaeoecology 65, 73–79. HOCHULI, P.A. 1978. Palynologische Untersuchungen im Oligozän und FOWLER, K. 1971. A new species of Aglaoreidia from the Eocene of Untermiozän der Zentralen und Weslichen Paratethys. Beiträge southern England. Pollen et Spores 13, 135–147. Paläontologie Österreich 4, 1–132.
FREDER›KSEN, N.O. 1973. New mid–Tertiary spores and pollen grains HOCHULI, P.A. 1984. Correlation of the Middle and Late Tertiary from Mississipi and Alabama. Tulane Studies in Geology and sporomorph assemblage. Paléobiologie continentale 14, Paleontology 10, 65–86. 301–314. FREDER›KSEN, N.O. 1980a. Sporomorphs from the Jackson Group (Upper JIMÉNEZ, J.A. 1984. A hypothesis to explain the reduced distribution of Eocene) and adjacent strata of Mississipi and western Alabama. the mangrove Pelliciera rhizophorae Tr. & Pl. Biotropica 16, Geological Survey Professional Paper 1084, 1–75. 304–308. FREDER›KSEN, N.O. 1980b. Mid–Tertiary climate of southeastern United KEDVES, M. 1969. Etudes palynologiques des couches du Tertiaire States: the sporomorph evidence. Journal of Paleontology 54, inferieur de la region Parisienne. IV. Pollens des Normapolles. 728–739. Pollen et Spores 11, 385–396. FREDER›KSEN, N.O. 1985. Review of early Tertiary sporomorph KEDVES, M. 1970. Etudes palynologiques des couches du Tertraire palaeoecology. American Association of Stratigraphical inferieur de la region Parisienne. Pollen et Spores 12, 83–97. Palynologists Contribution Series 19, 1–92. KEDVES, M. 1982. Palynology of the Thanetian layers of Ménat. FREDER›KSEN, N.O. 1988. Sporomorph biostratigraphy, floral changes, Palaeontographica Abteilung B Ionnides 182, 87–150. and paleoclimatology, Eocene and earliest Oligocene of the eastern Gulf Coast. United States Geological Survey Professional Paper KEDVES, M. 1986. Paleogene fossil sporomorphs of the Bakony 1448, 1–68. Mountains IV. Studia Biologica Hungarica 21, 1–120.
FREDER›KSEN, N.O. 1994. Middle and Late Paleocene angiosperm pollen KOÇY‹⁄‹T, A. 2005. The Denizli graben–horst system and the eastern from Pakistan. Palynology 18, 91–137. limit of western Anatolian continental extension: basin fill, structure, deformational mode, throw amount and episodic FREDER›KSEN, N.O., E DWARDS, L.E., A GER, T.A. & S HEEEHAN, T.P. 2002. evolutionary history, SW Turkey. Geodinamica Acta 18, Palynology of Eocene strata in the Sagavanirktok and Canning formations on the north slope of Alaska. Palynology 26, 59–93. 167–208. KONAK, N., A KDEN‹Z, N. & Ö ZTÜRK, E.M. 1987. Geology of the South of FUCH, J.R. 1970. Ecological and palynological notes on Pelliciera rhizophorae. Acta Botanica Neerlandica 19, 884–894. Menderes Massif. Guide Book for the Field Excursion Along Western Anatolia, IGCP Project No. 5, 42–53. GERMERAAD, J.H., H OPPING, C.A. & M ÜLLER, J. 1968. Palynology of Tertiary sediments from tropical areas. Review of Palaeobotany KORALAY, O.E., D ORA, O.Ö., C HEN, F., S AT›R, M. & C ANDAN, O. 2004. and Palynology 6, 189–348. Geochemistry and geochronology of orthogneisses in the Derbent (Alaflehir) area, eastern part of the Ödemifl–Kiraz submassif, GONZÁLEZ-GUZMÁN, A.E. 1967. A Palynological Study on the Upper Los Menderes Massif: Pan-African magmatic activity. Turkish Journal Cuervos and Mirador Formations (Lower and Middle Eocene, Tibu of Earth Sciences 13, 37–61. Area, Colombia). E.J. Brill, Leiden.
172 M.S. AKK‹RAZ ET AL.
KRUTZSCH, W. 1957. Sporen–und Pollengruppen aus der Oberkreide und POLE, M.S. & M ACPHA›L, M.K. 1996. Eocene Nypa from Regatta Point, dem Tertiär Mitteleuropas und ihre stratigraphische Verteilung. Tasmania. Review of Palaeobotany and Palynology 92, 55–67. Zeitschrift für Angewandte Geologie 3, 509–548. R›EGEL, W., B ODE, T., H AMMER, J., H AMMER–SCHIEMANN, G., L ENZ, O. & KRUTZSCH, W. 1970. Atlas der mittel–und jungtertiären dispersen WILDE, V. 1999. The palaeoecology of the Lower and Middle Sporen–und Pollen- sowie der Mikroplanktonformen des Eocene at Helmstedt, northern Germany – a study in contrasts. nördlichen Mitteleuropas. Lieferung VII, VEB Gustav Fischer Acta Palaeobotanica Supplement 2, 349–358. Verlag, Jena, 175 p. ROCHE, E. 1988. Pollen and spores (compilation). In: V INKEN, R. (ed), KRUTZSCH, W. & V ANHOORNE, R. 1977. Die pollen flora von Epinois und The Northwest European Tertiary Basin. Geologisches Jahrbuch Loksbergenin Belgien. Palaeontographica Abteilung B Ionnides (Hannover), Reihe A 100, 288–320. 163, 1–110. RULL, V. 1992. Paleoecología y análisis secuencial de una sección deltaica LENZ, O.K. & R IEGEL, W. 2001. Isopollen maps as a tool for the Terciaria en la Cuenca de Maracaibo. Boletín Sociedad Venezolana reconstruction of a coastal swamp from the Middle Eocene at Geólogos 46, 16–26. Helmstedt (northern Germany). Facies 45, 177–194. RULL, V. 1997a. Sequence analysis of western Venezuelan Cretaceous to MOSBRUGGER, V. & U TESCHER, T. 1997. The coexistence approach – a method for quantitative reconstructions of Tertiary terrestrial Eocene sediments using palynology; chrono-paleoenvironmental palaeoclimate data using the plant fossils. Palaeogeography, and paleovegetational approaches. Palynology 21, 79–90. Palaeoclimatology, Palaeoecology 134, 61–86. RULL, V. 1997b. Oligo–Miocene palynology of the Rio Chama sequence
MÜLLER, J. 1980. Fossil pollen records of extant angiosperms. Botanical (western Venezuela), with comments on fossil algae as Review 47, 1–142. paleoenvironmental indicators. Palynology 21, 213–229.
MÜLLER, J. 1981. Palynological evidence for Paleogene climatic changes. RULL, V. 1998a. Middle Eocene mangroves and vegetation changes in Mémoires du Museum National d’Historie Naturelle, Nouvelle the Maracibo Basin. Palaios 13, 287–296. Séries, B Botanique XXVII, 211–218. RULL, V. 1998b. Biogeographical and evolutionary considerations of NAKOMAN, E. 1966. Analyse sporopollinique des lignites eocenes de Mauritia (Arecaceae), based on palynological evidence. Sorgun (Yozgat–Turquie). Mineral Research and Exploration Palaeogeography, Palaeoclimatology, Palaeoecology 100, Institute of Turkey (MTA) Bulletin 67, 68–88. 109–122.
N›CKEL, B. 1996. Palynofazies und Palynostratigraphie der Pechelbronn RULL, V. 1999. Paleofloristic and paleovegetational changes across the Schihten im nördlichen oberrhein talgraben. Palaeontographica Paleocene–Eocene boundary in northern South America. Abteilung B Ionnides 240, 1–151. Palaeogeography, Palaeoclimatology, Palaeoecology 107, 83–95.
N›X, H.A. 1982. Environmental determinations of biogeography and RYAN, P.D., H ARPER, D.A.T. & W HALLEY, J.S. 1995. PALSTAT, Statistics evolution in Terra Australis. In: BARKER, W.S & GREENSLADE, P.J.M. for palaeontologists . Chapman & Hall, London (now Kluwer (eds), Evolution of the Flora and Fauna of Arid Australia. Peacock Academic Publishers). Publication, Adelaide, 47–66. fiAHBAZ, A. & G ÖRMÜfl, S. 1992. Çardak (Denizli) kuzeyindeki Eosen ve OLLIVIER-PIERRE, M.F., MAUPIN, C., ESTÉOULE-CHOUX, J. & SITTLER, C. 1993. Oligosen yafll› konglomeralar›n stratigrafik ve sedimantolojik Transgression et paleoenvironnement à l’ Oligocène en Bretagne. incelenmesi. [Stratigraphic and sedimentalogical investigation of Sédimentologie, micropaléontologie palynologie et palynofaciès du Eocene and Oligocene conglomerates in the north of Çardak Rupélien du Bassin de Rennes. Palaeogeography, Palaeoclimatology, Palaeoecology 103, 223–250. (Denizli)] 9th Petroleum Congress of Turkey2, 62–74 [in Turkish with English abstract]. ÖZER, S., SÖZB‹L‹R, H., ÖZKAR, ‹., TOKER, V. & SAR›, B. 2001. Stratigraphy of Upper Cretaceous-Palaeogene sequences in the southern and SAR›, B. & Ö ZER, S. 2002. Upper Cretaceous stratigraphy of the Bey eastern Menderes Massif (western Turkey). International Journal Da¤lar› carbonate platform, Korkuteli area (Western Turides, of Earth Sciences 89, 852–866. Turkey). Turkish Journal of Earth Sciences 11, 39–59.
ÖZKAYA, ‹. 1990. Origin of Lycian Nappes, SW Turkey. Tectonophysics SCHM›EDL, G., S CHERBACHER, M., B RUCH, A.A., J ELEN, B., N EBELS›CK, J.H., 177, 367–379. HEMLEBEN, C., M OSBRUGGER, V. & R› FELJ, H. 2002. Palaeoenvironmental evolution of the Paratethys in the Slovenian ÖZKAYA, ‹. 1991. Evolution of a Tertiary volcanogenic trough in SW Basin during the Late Paleogene. International Journal of Earth Turkey – the Alakaya basin of the Lycian belt. Geologische Rundschau 80, 657–668. Sciences 91, 123–132. fiENEL, M. 1991. Palaeocene-Eocene sediments interbedded with PLAZ›AT, J.C., C AVAGNETTO, C., K OEN›GUER, J.C. & B ALTZER, F. 2001. History and biogeography of the mangrove ecosystem, based on volcanics within the Lycian Nappes: Faralya formation. Mineral a critical reassessment of the paleontological record. Wetlands Research and Exploration Institute of Turkey Bulletin 113, 1–14.
Ecology and Management 9, 161–179. fiENEL, M. 1997. Denizli–J9 Quadrangle, 1:100,000 Scale Geological PO›SSON, A. 1976. Essai d’interprétation d’une transversale Map and Explanatory Text. Mineral Research and Exploration Korkuteli–Denizli (Taurus ouest-anatolien-Turkuie). Bulletin of Institute of Turkey (MTA) Publications, Ankara. the Geological Society of France 2, 499–509.
173 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
fiENGÖR, A.M.C. & Y›LMAZ, Y. 1981. Tethyan evolution of Turkey: a plate THOMSON, P.W. & P FLUG, H. 1953. Pollen und Sporen des tectonic approach. Tectonophysics 75, 181–241. mitteleurpäischen Tertiärs. Palaeontographica Abteilung B Ionnides 94, 1–138. fiENGÖR, A.M.C., GÖRÜR, N. & fiARO⁄LU, F. 1985. Strike–slip deformation, basin formation and sedimentation: Strike–slip faulting and UPCHURCH, G.R. & WOLFE, J. A. 1987. Mid–Cretaceous to early Tertiary related basin formation in zones of tectonic escape: Turkey as a vegetation and climate: evidence from fossil leaves and woods. In: case study. In: B›DDLE, K.T. & C HR›ST›E–BUCK, N. (eds), Strike–slip FRIIS, E.M., C HALONER, W.G. & C RANE, P.H. (eds), The Origins of Faulting and Basin Formation . Society of Economic Angiosperms and Their Biological Consequences. Cambridge Paleontologists and Mineralogist, Special Publications 37, University Press, 75–105. 227–264. VAN DER HAMMEN, T. & W IJMSTRA, T.A. 1964. A palynological study on SEY‹TO⁄LU, G. & S COTT, B. 1996. The cause of N-S extensional tectonics the Tertiary and Upper Cretaceous of British Guiana. Leidse in western Turkey: tectonic escape vs back-arc spreading vs Geologische Mededelingen 30, 183–241. orogenic collapse. Journal of Geodynamics 22, 145–153. VINKEN, R. 1988. The Northwest European Tertiary Basin. (IGCP 124). SÖZB‹L‹R, H. 2002. Revised stratigraphy and facies analysis of Geologisches Jahrbuch (Hannover) Reihe A 100, 1–508. Palaeocene–Eocene supra–allochthonous sediments (Denizli, SW WESTAWAY, R., G U›LLOU, H., Y URTMEN, S., D EMIR, T., S CA›LLET, S. & Turkey) and their tectonic significance. Turkish Journal of Earth ROWBOTHAM, G. 2005. Constraints on the timing and regional Sciences 11, 87–112. conditions at the start of the presence phase of crustal extension SÖZB‹L‹R, H. 2005. Oligo–Miocene extension in the Lycian orogen: in western Turkey, from observations in and around the Denizli evidence from the Lycian molasse basin, SW Turkey. Geodinamica region. Geodinamica Acta 18, 209–238. Acta 18, 255–282. WESTGATE, J.W. & G EE, C.T. 1990. Palaeoecology of a Middle Eocene SÖZB‹L‹R, H., Ö ZER, S., S AR›, B. & A VflAR, N. 2001. Supra–allochton mangrove biota (vertebrates, plants and invertebrates) from sedimentary succession in western Anatolia: new stratigraphic southwest Texas. Palaeogeography, Palaeoclimatology, data and tectonic results. Fouth International Turkish Geology Palaeoecology 78, 163–177. Symposium, Abstracts, 36. WOLFE, J.A. 1978. A paleobotanical interpretation of Tertiary climates in SR›VASTAVA, S.K. & B› NDA, P.L. 1991. Depositional History of the Early northern hemisphere. American Scientist 66, 694–703. Eocene Shumaysi Formation, Saudi Arabia. Palynology 15, WOLFE, J.A. 1979. Temperature parameters of humid to mesic forests 47–61. of eastern Asia and relation to forests of other regions of the STUCHL›K, L., Z› EMB›´NSKA-TWORZYDLO, M., K OHLMAN-ADAMSKA, A., Northern Hemisphere and Australasia. United States Geological . GRABOWSKA, I., W AZYNSKA´ , H., S LODKOWSKA, B. & S ADOWSKA,A. Survey Professional Paper 1106, 37 p. 2001. Atlas of pollen and spores of Polish Neogene. W. Szafer WOLFE, J.A. 1992. Climatic, floristic and vegetational changes near the Institute of Botany, Polish Academy Sciences 1, 1–69. Eocene/Oligocene boundary in North America. In: PROTHERO, D.R. THAN›KA›MON›, G. 1987. Mangrove palynology. Institut français de & B ERGGREN, W.A. (eds), Eocene–Oligocene Climatic and Biotic Pondichéry, Travaux de la Section Scientifique et Technique XXIV, Evolution. Princeton University Press, 421–436. 1–100. YALÇ›NKAYA, S., ENG‹N, A., TANER, K., AFflAR, Ö.P., DALK›L›Ç, H. & ÖZGÖNÜL, THAN›KA›MON›, G., CARAT›N›, C., VENKATACHALA, B.S., RAMANUJHAM, C.G.K. & E. 1986. Bat› Toroslar›n Jeolojisi [Geology of Western Taurides]. KAR, R.K. 1984. Selected Tertiary angiosperms pollen from India Mineral Research and Exploration Institute of Turkey (MTA) and their relationship with African Tertiary pollens. Institut Report 7898, 132 p [in Turkish, unpublished]. français de Pondichéry, Travaux de la Section Scientifique et YASAMANOV, N.A. 1982. The problem of global changes in temperature Technique XIX, 1–192. regime of the earth’s surface during Cenozoic time. Izvestiya AN TH›ELE-PFE›FFER, H. 1988. Die Mikroflora aus dem mittelozänen SSSR, seriya geologicheskaya 10, 106–110. ölschiefers von Messel bei Darmstadt. Palaeontographica Abteilung B Ionnides 211, 1–86.
Received 20 April 2005; revised typescript accepted 05 January 2006
174 M.S. AKK‹RAZ ET AL.
Plate I
(all photomicrographs x 500)
Figure 1, 2 Leiotriletes adriennis (R. Potonié and Gelletich) Krutzsch 3 Leiotriletes minoris (Krutzsch)Ziembi´nska-Tworzydlo 4 Leiotriletes wolffii Krutzsch 5 Leiotriletes microadriennis Krutzsch 6 Leiotriletes sp. 7 Cicatricosisporites sp. 8 Triplanosporites microsinuosus Pflanzl 9 Polypodiaceoisporites microconcavus Krutzsch 10 Polypodiaceoisporites kedvesii Stuchlik 11 Polypodiaceoisporites muricinguliformis Nagy 12-15 Polypodiaceoisporites sp. 16 Retitriletes sp. 17, 18 Laevigatosporites haardti (R. Potonié & Venkatachala) Thomson & Pflug 19, 20 Pityosporites labdacus (R. Potonié) Thomson & Pflug 21-24 Pityosporites microalatus (R. Potonié) Thomson & Pflug 25 Abiespollenites cf. absolutus Thiergart
175 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
1 2 3 4
5 6 7 8 9
10 11 12 13 14
15 16 19 17 18
21 20 22
25 23 24
176 M.S. AKK‹RAZ ET AL.
Plate II
(all photomicrographs x 500)
Figure 1 Abiespollenites cf. absolutus Thiergart 2 Cathayapollis pulaënsis (Nagy) Ziembi´nska-Tworzydlo 3 Cathayapollis sp. 4,5 Cyacadopites gracilis (Wodehouse) Krutzsch 6 Cyacadopites intrasructus Krutzsch 7,8 Cyacadopites cf. minimus Krutzsch 9 Psilamonocolpites sp. 10 Sequoiapollenites polyformosus Thiergart 11a, b Spinizonocolpites sp. 12 Longapertites sp. 13 Kopekipollenites transdanubicus Kedves 14 Sparganiaceaepollenites polygonalis Theiergart 15 Triatriopollenites bituitus (R. Potonié) Thomson & Pflug 16-19 Triatriopollenites excelsus (R. Potonié) Thomson & Pflug ssp. typicus Pflug 20, 21 Triatriopollenites excelsus (R. Potonié) Thomson & Pflug ssp. minor Pflug 22-24 Plicatopollis plicatus (R. Potonié) Krutzsch 25-29 Plicatopollis lunatus Kedves 30 Triporopollenites robustus (Pflug) Thomson & Pflug 31 Triporopollenites sp. 32-35 Momipites quietus (R. Potonié) Nichols 36, 37 Trivestibulopollenites betuloides Pflug in Thomson & Pflug 38 Subtriporopollenites anulatus (Pflug) ssp. nanus Pflug & Thomson in Thomson & Pflug 39 Subtriporopollenites constans (Pflug) Thomson & Pflug 40 Olaxipollis matthesii Krutzsch 41a,b Mauritiidites franciscoi (Van Der Hammen) Van Der Hammen & Garcia de Mutis 42 Intratriporopollenites indubitalibis (R. Potonié) Pflug & Thomson in Thomson & Pflug 43, 44 Polyporopollenites undulosus (Wolff) Thomson & Pflug 45 Polyvestibulopollenites verus (R. Potonié) Thomson & Pflug
177 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
1 3
2
4 6 7 8 5
9 10 11a 11b 12
13 14 15 16 17 18
19 20 22 23 24 25 21
26 27 28 29a 29b 30 31
32 33 34 35 36 37 38 39
41a 41b 42 43 40 44 45
178 M.S. AKK‹RAZ ET AL.
Plate III
(all photomicrographs x 500)
Figure 1 Myrtaceidites mesonesus Cookson & Pike 2 Pentapollenites pentangulus (Pflug in Thomson & Pflug) Krutzsch 3, 4 Tricolpopollenites microhenrici (R. Potonié) Thomson & Pflug 5 Tricolpopollenites liblarensis (Thomson) Thomson & Pflug ssp. fallax (R. Potonié) Thomson & Pflug 6 Tricolpopollenites liblarensis (Thomson) Thomson & Pflug spp. liblarensis (Thomson) Thomson & Pflug 7 Tricolpopollenites parmularius (R. Potonié) Thomson & Pflug 8 Tricolpopollenites retiformis (Pflug & Thomson) Thomson & Pflug 9, 10 Tricolporopollenites cingulum (R. Potonié) Thomson & Pflug ssp. fusus (R. Potonié) Thomson & Pflug 11 Tricolporopollenites cingulum (R. Potonié) Thomson & Pflug ssp. pusillus (R. Potonié) Thomson & Pflug 12, 13 Tricolporopollenites megaexactus (R. Potonié) Thomson & Pflug ssp. brühlensis (Thomson in Potonié) Thomson & Thiergart 14 Tricolporopollenites cf. pseudocingulum (R. Potonié) Thomson& Pflug 15-19 Psilatricolpites crassus Van Der Hammen & Wymstra 20 Psilatricolporites cf. costatus Dueñas 21 Chenopodipollis multiplex (Weyland & Pflug) Krutzsch 22-24 Nowemprojectus tumanganicus (Bolotnikova) Frederiksen 25, 26 undifferentiated dinoflagellate cysts 27, 28 Pediastrum spp.
179 MIDDLE–LATE EOCENE MICROFOSSILS FROM DEN‹ZL‹ AREA
1 2 3 4 5 6 7 8
15 9 10 11 12 13 14
16 17 18 19
20 21 22 23 24a 24b
25 26 27 28
180