Lower Cretaceous (Upper Barremian-Lower Aptian?) Palynoflora from ...

Lower Cretaceous (Upper Barremian-Lower Aptian?) Palynoflora from the Kitadani Formation (Tetori Group, Inner Zone of Central Japan) Author(s): Julien Legrand , Denise Pons , Kazuo Terada , Atsushi Yabe and Harufumi Nishida Source: Paleontological Research, 17(3):201-229. 2013. Published By: The Palaeontological Society of Japan DOI: http://dx.doi.org/10.2517/1342-8144-17.3.201 URL: http://www.bioone.org/doi/full/10.2517/1342-8144-17.3.201

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Paleontological Research, vol. 17, no. 3, pp. 201–229, August 1, 2013 © by the Palaeontological Society of Japan doi:10.2517/1342-8144-17.3.201

Lower Cretaceous (upper Barremian-lower Aptian?) palynoflora from the Kitadani Formation (Tetori Group, Inner Zone of central Japan) JULIEN LEGRAND1, 2, DENISE PONS1, KAZUO TERADA3, ATSUSHI YABE4

AND

HARUFUMI NISHIDA2

1UMR 7207 CNRS, Centre de Recherche sur la Paléobiodiversité et les Paléoenvironnements, Université Pierre et Marie Curie (UPMC), Paris 75231, France 2Department of Biological Sciences, Faculty of Science and Engineering, Chuo University, Tokyo 112-8551, Japan (e-mail: [email protected]) 3 Fukui Prefectural Dinosaur Museum, Katsuyama 911-8601, Japan 4National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba 305-0005, Japan

Received May 14, 2012; Revised manuscript accepted November 13, 2012

Abstract. The Tetori Group, which crops out in the Inner Zone of central Japan, has been extensively studied for its rich floral and vertebrate fossil assemblages. The authors provide the first contribution to the palynoflora of the Kitadani Formation, which has been dated as late Barremian to early Aptian on the basis of the freshwater bivalves recorded from it. The studied assemblage yields 45 genera and 41 species of spores and gymnosperm pollen grains, some freshwater algae, one epiphyllous fungus, and various plant fragments. No angiosperm pollen grains were observed. Some forms described here in detail are not yet known from the literature. They probably correspond to new species, but the scarcity of the specimens has caused us to place them temporarily in open nomenclature. This palynological study adds new data to the present knowledge on the Barremian-Aptian Tetori-type Paleoflora of eastern Asia. The authors compare the palynological inventory with recently published data obtained from the Barremian strata of the Choshi Group, the Outer Zone of Southwest Japan. Then, they situate the results among the previous paleoenvironmental reconstructions of the Tamodani Flora, and confirm a warm temperate and moderately humid climate, with locally drier conditions. Even if some elements of the assemblage suggest transportation, all of them are of continental origin and confirm a fluvio-lacustrine environment for the deposition. Key words: Inner Zone of Japan, Kitadani Formation, Lower Cretaceous, Palynology, Tamodani Flora

Introduction During the late Mesozoic period, Japan was separated into two zones by a marine basin: the Inner Zone (northwestern part, along the Sea of Japan) connected to the east of the Korean Peninsula and representing the margin of the Eurasian continent, and the Outer Zone (southeastern part, along the Pacific Ocean) in Southwest Japan represented by oceanic islands located further to the south (Hirooka et al., 1983, 1985; Yaskawa, 1975; Otofuji et al., 1985; Kojima, 1989; Matsukawa and Obata, 1992; Matsuoka et al., 1997; Ishida et al., 2003; Lee and Kim, 2005; Lee, 2008; Hisada et al., 2008; Matsukawa and Fukui, 2009). However, some authors doubt the existence of the Outer Zone terranes, and propose an oceanward growth of all of proto-Japan along the Eurasian

continental margin, until it became an island arc during the Miocene (Isozaki, 1996, 1997; Maruyama et al., 1997; Otoh, 1998; Otoh and Sasaki, 1998; Yamakita and Otoh, 2000; Isozaki et al., 2010). From his extensive study of fossil plants from Japan and eastern Asia, Kimura (1979, 1987) distinguished a “Tetori-(Siberian-)type” Paleoflora in the Inner Zone and a “Ryoseki-(NorthGondwanian-)type” Paleoflora in the Outer Zone, reflecting peculiar conditions of geography and climate and arguing for the terranes hypothesis, with a lowlatitudinal origin of the Outer Zone terranes (Figure 1). He further subdivided the Tetori-type Paleoflora into four stratofloras (Kimura, 1975). However, recent stratigraphic and paleobotanical studies (Yabe et al., 2003; Yabe and Kubota, 2004; Yamada and Uemura, 2008; Yamada, 2009) challenge the uniformity of the Tetori-

202

Julien Legrand et al.

138°E

((1) 1)) Tetori-type Flora

Tetori-type Flora

36°N

ed

pe -ty Flora

40°N Kitadani Dinosaur Quarry

In Ou ner Z ter one Zon e

Kitadani Dinosaur Quarry

x

44°N

(2) N Mi

126°E

34°N

Mixedtype Flora

Mixed-type Flora

Ryoseki-type Flora 500 km

130°E

28°N

Ryoseki-type Flora

140°E

pre-Jurassic Asian continental margin Abukuma and South Kitakami terranes Tamba, Mino and Ashio terranes

20°N

Middle–Late Jurassic terranes Early Cretaceous terranes Late Cretaceous terranes pre-Cambrian (?) - early Paleozoic terranes

South Chichibu terrane

500 km

floras boundaries coastlines faults Localities of Early Cretaceous floras Ryoseki-type Tetori-type Mixed-type

Figure 1. Paleofloristic provinces in Japan and eastern Asia during the Late Jurassic to the Early Cretaceous. 1, paleogeographical map illustrating the relative locations of the Inner and Outer Zones (modified from Golozoubov et al., 1999); 2, a present-day map (from Kimura, 1987).

type Paleoflora throughout the Middle Jurassic to Early Cretaceous time interval, emphasized in the paleofloral provincialism concept of Kimura and coworkers (Kimura, 1958, 1979, 1987; Kimura and Sekido, 1976, 1978; Kimura and Ohana, 1997), and point to the necessity of its revision. The Tetori Group sporadically outcrops in the Hida region of the Tetori Basin (Matsukawa, 1991). It is Middle Jurassic to Early Cretaceous in age and is divided, in ascending order, into the Kuzuryu, Itoshiro and Akaiwa subgroups (Maeda, 1961; Fujita, 2002; Kusuhashi et al., 2002). Microremains from the Tetori Group were studied for the first time by Umetsu (2002), Umetsu and Matsuoka (2003) and Umetsu and Sato (2007), whose samples covered the whole Tetori Group except for its uppermost part (the Kitadani Formation and correlatives). They reported palynomorphs from the Itoshiro and Akaiwa subgroups

(most likely in the Nochino Formation in the Kuzuryu River area, equivalent to the Akaiwa Formation). However, their data were not good enough regarding the preservation state of fossils and stratigraphic range, and they could only identify five genera of fern spores (Appendicisporites, Cicatricosisporites, Cyathidites, Osmundacidites, Schizaeoisporites) and one genus of gymnosperm pollen grains (Classopollis), but no species. The Fukui Prefectural Dinosaur Museum ordered a palynological survey from Palyno Survey Company Co. Ltd. in 1998 at the Dinosaur Quarry of the Kitadani Formation, in which two genera of spores (Cicatricosisporites and Concavisporites) and six genera of pollen grains (Podocarpidites, Araucariacites, Classopollis, Ephedripites, Cycadopites, cf. Exesipollenites) were identified. On the occasion of an excavation project held by the Fukui Prefectural Dinosaur Museum (Shibata and Goto, 2008) at the locality known as the “Dinosaur Quarry”

Palynoflora from the Kitadani Formation Sakhal

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Russia

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Pri m

China

Hokkaido

an

Fukui Prefecture

Honshu

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rea Ko

Japon

36º15'N

Shikoku Kyushu

1000 km 136ºE

137ºE Noto Peninsula

Sea of Japan

36º50'N

TOYAMA PREFECTURE

Kitadani Dinosaur Quarry

ISHIKAWA Mount Haku

Takayama

36ºN FUKUI PREFECTURE GIFU PREFECTURE

SHIGA PREF.

Sugiyama

50 km

Su giy am a

R iv er

Biwa lake

Katsuyama

Takin

ami

Rive

r

marine and non-marine sediments - Upper Pleistocene to Holocene lower terrace - Upper Pleistocene non-alkaline mafic volcanic rocks - Lower Pleistocene non-alkaline mafic volcanic rocks - Lower to Middle Miocene non-alkaline mafic volcanic rocks - Upper Miocene to Pliocene non-alkaline felsic volcanic rocks - Upper Cretaceous non-marine sedimentary rocks (Tetori Group) - Lower Cretaceous Funatsu Granite - Jurassic gneiss - Hida metamorphic rocks 2000 m river Fukui-Ishikawa prefectural boundary

136º50'E Figure 2.

36º10'N

136º55'E

Location map of the quarry and geology of the area (modified from Geological Survey of Japan, AIST, 2010).

along the Sugiyama River (affluent to the Takinami River, northeast of Katsuyama City), we took samples for palynological study in the siltstones and fine-grained sandstones of the Kitadani Formation (Akaiwa Subgroup, Tetori Group), which crops out in the Takinami River area (Figure 2). Our study considerably complements macrofloral data previously reported from the Barremian-Aptian Tamodani Flora (= Myodani Flora: Matsuo and Omura, 1966). It will help to improve the understanding of the paleoenvironment of this area and to enhance knowledge of palynofloras of the Inner Zone of Japan. Another goal of the paper is the compilation of a more complete species list for these units. We also compare the composition of the Kitadani palynoflora

with that reported from the Barremian of the Choshi Group, the Outer Zone of Japan (Legrand et al., 2011).

Geological setting The Akaiwa Subgroup is mainly composed of alluvial sediments deposited in a freshwater deltaic or fluvial environment (Masuda et al., 1991). The Kitadani Formation can be correlated to the Myodani Formation (Kawai, 1961), which represents the uppermost part of the Akaiwa Subgroup in the Tedori River area (Maeda, 1958, 1961). It conformably overlies the Akaiwa Formation and is unconformably overlain by the Omichidani Formation (Upper Cretaceous) (for the relations among units

Materials and methods The samples were taken from four horizons of siltstone and fine-grained sandstone cropping out at the locality known as the “Kitadani Dinosaur Quarry”. Methods for the palynological study are the same as in Legrand et al. (2011). The formation, the horizon, the number of the slide, and the position of fossils under the England Finder™ Graticule are indicated in the legend for each specimen illustrated in this paper. Because it is difficult to compare the fossil spores and

(2)

300 m

Kita-4

Kita-3 Kita-2

3rd Dinosaur Excavation Project - FPDM 2007

Kitadani Fm

(1)

Kita-1 Aka. Fm

and floras mentioned, see Fujita, 2003; Yabe et al., 2003). The freshwater sediments of the formation are mainly composed of alternating beds of siltstone and sandstone, and probably represent deposition in a meandering river environment (Azuma, 2003) (Figure 3). Maeda (1961, 1962), Isaji (1993) and Kozai et al. (2001, 2002) studied the mollusc and bivalve freshwater fauna (the “TPN fauna”) of the formation, among which Nippononaia ryosekiana Suzuki, reported by Isaji (1993), characterizes the late Barremian Sebayashi-type Fauna of the Monobegawa Group (the Outer Zone of Southwest Japan) (Kozai et al., 2002) and suggests assigning a late Barremian to early Aptian age to the formation. Moreover, Kubota (2005) reported a charophyte gyrogonite assemblage dated as Barremian from the Takinami River area. A rich vertebrate fossil assemblage (turtles, crocodiles, dinosaurs, fishes) has been reported from the “Kitadani Dinosaur Quarry” in the past few decades (Azuma and Tomida, 1995, 1997; Kobayashi, 1998; Azuma and Currie, 2000; Goto et al., 2002; Kobayashi and Azuma, 2003; Currie and Azuma, 2006; Shibata and Goto, 2008). Fossil plants from the Tamodani Flora were studied by Matsuo and Omura (1966), Kimura (1975), Kimura and Horiuchi (1979), Yabe et al. (2003) and Yabe and Kubota (2004). However, only a few fossil plants have been reported from the Kitadani Formation, and many plant megafossils still remain unstudied (Goto et al., 2002). The megafossil assemblage from the Kitadani Formation is characterized by a scarcity of Filicopsida (mainly represented by Onychiopsis of Dicksoniaceae and Gleichenites of Gleicheniaceae) and an abundance of Cycadales and Coniferales (represented by cones and twigs). Yabe and Kubota (2004) described a twig of Brachyphyllum obesum Heer (Cheirolepidiaceae), representing the first occurrence of this species in the Inner Zone. Zamiophyllum sp., considered as a “Ryoseki-type” element, was reported from the Myodani Formation, which is stratigraphically correlated to the Kitadani Formation (Matsuo and Omura, 1966).

Omi. Fm


Lower Cretaceous upper Barremian-lower Aptian ?

204

unconformity siltstone alternating beds of siltstone and fine-grained sandstone fine-grained sandstone coarse-grained sandstone

palynoflora plant remain gastropod bivalve dinosaur remain 50 m (1) 10 m (2)

Figure 3. Synthetic stratigraphical column of the Kitadani Formation along the Takinami River (1, modified from Maeda, 1958; 2, modified from Shibata and Goto, 2008). Abbreviations: Aka. Fm, Akaiwa Formation; Omi. Fm, Omichidani Formation.

pollen grains to genera of plants living today, we followed the classification scheme of sporae dispersae, based on palynomorph morphology, defined by Potonié and Kremp (1954, 1955), Dettmann (1963), Potonié (1956, 1958, 1960, 1966, 1970a, 1970b, 1975), and Pflug (1953). However, we tried to determine the presumed botanical affinities based on previous works identifying palynomorphs found in situ in fructifications, or on morphological similarities with present species.

Systematic descriptions We present below, in nomenclatural order, species that

Palynoflora from the Kitadani Formation

205

Table 1. List of all the taxa encountered during our investigation, in nomenclatural order, with their occurrences in the formation and their figurative position in brackets. -, one grain; R, rare; P, present; C, common. Kita-2

Kita-3

Kita-4

R P P

P P

R R P

R R P

– R R P

ANTETURMA PROXIMEGERMINANTES Turma Triletes Azonales Suprasubturma Acavatitriletes Subturma Azonotriletes Infraturma Laevigati, Quasilaevigati Auritulinasporites deltaformis Burger, 1966 Biretisporites potoniaei (Delcourt and Sprumont, 1955) Delcourt, Dettmann and Hughes, 1963 (Figure 4.1) Biretisporites sp. (Figure 4.2) Cibotiumspora paradoxa (Maljavkina, 1949) Chang, 1965 Cyathidites australis Couper, 1953 (Figure 4.6) Cyathidites minor Couper, 1953 Deltoidospora hallii Miner, 1935 Todisporites major Couper, 1958 (Figure 4.3) Todisporites minor Couper, 1958 (Figure 4.4) Infraturma Apiculati Subinfraturma Granulati Scabrati Concavissimisporites punctatus (Delcourt and Sprumont, 1955) Brenner, 1963 (Figure 4.5) Granulatisporites sp. (Figure 4.7) Impardecispora apiverrucata (Couper, 1958) Venkatachala, Kar and Raza, 1969 (Figure 4.9) Osmundacidites wellmanii Couper, 1953 (Figure 4.8) Subinfraturma Verrucati Converrucosisporites sp. in Legrand, Pons, Nishida and Yamada, 2011 Leptolepidites psarosus Norris, 1969 (Figures 4.11a–b, 4.12) Manumia japonica Legrand, Pons, Nishida and Yamada, 2011 (Figure 4.10) Subinfraturma Baculati Baculatisporites comaumensis (Cookson, 1953) Potonié, 1956 (Figure 4.13) Baculatisporites sp. (Figures 4.14a–b) Subinfraturma Nodati Anapiculatisporites cooksonae Playford, 1965 (Figure 4.15) Echinatisporis varispinosus (Pocock, 1962) Srivastava, 1975 Echinatisporis sp. (Figure 4.16) Infraturma Murornati Cicatricosisporites hallei Delcourt and Sprumont, 1955 (Figures 5.7, 5.8, 5.10) Cicatricosisporites hughesi Dettmann, 1963 (Figure 4.17) Cicatricosisporites minor (Bolkhovitina, 1959) Pocock, 1964 (Figures 5.2a–b, 5.4) Cicatricosisporites pseudotripartitus (Bolkhovitina, 1961) Dettmann, 1963 (Figures 5.9a–b) Cicatricosisporites sinuosus Hunt, 1985 Cicatricosisporites sp. 2 in Legrand, Pons, Nishida and Yamada, 2011 Cicatricosisporites cf. C. sp. in Williams and Bujak, 1980 (Figures 5.3a–b, 5.5a–b, 5.6) Cicatricosisporites sp. 1 (Figures 5.1a–b) Cicatricosisporites sp. 2 (Figures 6.1a–b) Cicatricosisporites sp. 3 (Figures 6.2a–b, 6.3a–b) Cicatricosisporites sp. 4 (Figures 6.4a–b) Ischyosporites crateris Balme, 1957 (Figures 6.5–7, 6.10) Reticulatisporites spp. (Figures 7.1, 7.4, 7.5) Retitriletes austroclavatidites (Cookson, 1953) Döring, Krutzsch, Mai and Schulz in Krutzsch, 1963

– R P –

–

R R R R

R R P R

– R R

R R P

P P

R

R P R

R

R

P

R

R R – R R

R R R P R

R R R R R R R

C R R

206

Julien Legrand et al. Table 1.

Continued.

Retitriletes sp. 1 (Figure 6.9) Retitriletes sp. 2 (Figures 6.8, 6.11) Ruffordiaspora australiensis (Cookson, 1953) Dettmann and Clifford, 1992 (Figures 7.2, 7.3, 7.6, 7.10a–b) Turma Triletes Zonales Subturma Auritotriletes Infraturma Appendiciferi Appendicisporites potomacensis Brenner, 1963 (Figures 7.7a–b) Plicatella sp. (Figures 7.8a–b) Subturma Zonotriletes Infraturma Cingulati Cingulatisporites sp. 1 in Legrand, Pons, Nishida and Yamada, 2011 (Figure 8.1) Cingulatisporites sp. (Figure 8.2) Contignisporites sp. (Figures 7.9a–b) Polycingulatisporites reduncus (Bolkhovitina, 1953) Playford and Dettmann, 1965 Subinfraturma Laticingulati Gleicheniidites senonicus Ross, 1949 (Figures 8.3–4) Infraturma Tricrassati Coronatispora valdensis (Couper, 1958) Dettmann, 1963 (Figures 8.5a–b) Subturma Zonolaminatitriletes Infraturma Zonati Aequitriradites spinulosus (Cookson and Dettmann, 1958) Cookson and Dettmann, 1961 (Figure 8.7) Aequitriradites verrucosus (Cookson and Dettmann, 1958) Cookson and Dettmann, 1961 Aequitriradites sp. in Legrand, Pons, Nishida and Yamada, 2011 (Figures 8.8, 8.9) Couperisporites complexus (Couper, 1958) Pocock, 1962 (Figures 8.11a–b) Triporoletes reticulatus (Pocock, 1962) Playford, 1971 (Figures 8.10–b) Turma Monoletes Suprasubturma Acavatomonoletes Subturma Azonomonoletes Infraturma Laevigatomonoleti Laevigatosporites ovatus Wilson and Webster, 1946 (Figure 8.12) Laevigatosporites sp. (Figure 8.13) Incertae sedis Incertae sedis sp. 1 (Figure 8.6) Incertae sedis sp. 2 (Figure 8.16)

Kita-2 P R

Kita-3 P R P

Kita-4 R

R R

R R – R

R R

–

R

R –

R

–

R R R P R

R

R

R

R –

P –

R

– –

ANTETURMA VARIEGERMINANTES (= POLLENITES) Turma Saccites Subturma Monosaccites Callialasporites dampieri (Balme, 1957) Sukh-Dev, 1961 (Figure 10.1) Subturma Disaccites Alisporites thomasii (Couper, 1958) Pocock, 1962 Alisporites sp. in Legrand, Pons, Nishida and Yamada, 2011 Cedripites sp. (Figure 10.2) Vitreisporites pallidus (Reissinger, 1939) Nilsson, 1958

–

– R

R R R

–

Palynoflora from the Kitadani Formation Table 1.

207

Continued.

Turma Aletes and Kryptoaperturates Subturma Azonaletes Infraturma Psilonapiti Inaperturopollenites sp. in Legrand, Pons, Nishida and Yamada, 2011 Infraturma Granulonapiti Araucariacites australis Cookson, 1947 ex Couper, 1953 Balmeiopsis limbatus (Balme, 1957) Archangelsky, 1977 Spheripollenites psilatus Couper, 1958 Taxodiaceaepollenites hiatus (Potonié, 1931) Kremp, 1949 ex Potonié, 1958 Infraturma Circumpollini Classopollis torosus (Reissinger, 1950) Couper, 1958 emend. Burger, 1965 (Figures 10.3, 10.4) Turma Plicates Subturma Costates (= Polyplicates) Infraturma Costati Ephedripites montanaensis Brenner, 1968 Ephedripites sp. 1 (Figure 10.5) Ephedripites sp. 2 (Figure 10.6) Gnetaceaepollenites sp. in Legrand, Pons, Nishida and Yamada, 2011 Gnetaceaepollenites sp. (Figures 10.7, 10.8) Subturma Monocolpates (Monosulcites) and Zonocolpates Infraturma Quasilaevigati and Microsculptati Cycadopites minimus (Cookson, 1947) Pocock, 1970 Cycadopites sp. (Figure 10.11) Subturma Tricolpates, Triptyches Infraturma Heterotricolpati, Praecolpati Eucommiidites minor Groot and Penny, 1960 Eucommiidites troedssonii (Erdtman, 1948) Potonié, 1958 Turma Poroses Subturma Monoporines Exesipollenites tumulus Balme, 1957 (Figure 10.9) Exesipollenites tumulus Balme, 1957 subsp. triangulus Liu in Song, Zheng, Liu, Ye, Wang and Zhou, 1980 (Figures 8.14, 10.15) Incertae sedis Incertae sedis sp. 2 in Legrand, Pons, Nishida and Yamada, 2011 Incertae sedis sp. 3 (Figure 10.12)

Kita-2

Kita-3

Kita-4

R

R

R

P

P R

R – –

R

R

R

R – –

R –

R –

– R

R R

P R

R R

R R

C –

C –

– –

ALGAE CHLOROPHYTA Charophyceae Chomotriletes minor (Kedves, 1961) Pocock, 1970 (Figure 10.10) Ovoidites parvus (Cookson and Dettmann, 1959) Nakoman, 1966 Prasinophyceae Cymatiosphaera sp. (Figures 10.15, 10.16) Prasinophycean spores (Figures 10.13, 10.14) Incertae sedis Schizosporis reticulatus Cookson and Dettmann, 1959

R R R C R

C

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Julien Legrand et al. Table 1.

Continued. Kita-2

Kita-3

Kita-4

FUNGI ASCOMYCOTA Microthyriacites sp. in Legrand, Pons, Nishida and Yamada, 2011

R

Incertae sedis Incertae sedis sp. 4 (Figure 10.17)

R

PLANT FRAGMENTS Cuticle fragments of gymnosperms (Figures 11.1a–b) Cuticle fragments of Cheirolepidiaceae (Figures 11.2–11.5) Cross-fields of Coniferalean woods (Figures 10.18, 10.19) Vascular element of Bennettitales (Figure 10.21) Vessel element with scalariform perforation plates (Figure 10.20)

have been identified during this investigation (Table 1). The preparations are housed in the Collection de Paléobotanique of the Université Pierre et Marie Curie (UPMC), Paris (France). Only taxa in open nomenclature are described. Well known palynomorphs are not described. ANTETURMA PROXIMEGERMINANTES Turma Triletes Azonales

shows similarities to our spore. Botanical affinities.—Filicopsida. Infraturma Apiculati Subinfraturma Granulati Scabrati Genus Granulatisporites Ibrahim, 1933 emend. Potonié and Kremp, 1954 Type species: Granulatisporites granulatus Ibrahim, 1933

Suprasubturma Acavatitriletes Granulatisporites sp. Subturma Azonotriletes Infraturma Laevigati, Quasilaevigati Genus Biretisporites (Delcourt and Sprumont, 1955) Delcourt, Dettmann and Hughes, 1963 Type species: Biretisporites potoniaei (Delcourt and Sprumont, 1955) Delcourt, Dettmann and Hughes, 1963 Biretisporites sp. Figure 4.2 Biretisporites sp. in Legrand, 2009, p. 128, pl. I, figs. 6-7.

Occurrence.—Kitadani Fm (horizons Kita-2, 3, 4). Description.—Psilate trilete microspore. Amb rounded triangular. The laesurae are straight, thin, strongly raised (2 to 6 μm), and extend to the equator of the spore. Exine about 2 μm thick. Equatorial diameter = 20–55 μm. Remarks.—Lygodiumsporites equilabiatus Li, 2000 reported from the Neocomian of northwestern China,

Figure 4.7 Granulatisporites sp. A. in Legrand, 2009, p. 134, pl. II, fig. 9.

Occurrence.—Kitadani Fm (horizons Kita-2, 3). Distribution.—Barremian of the Choshi Group in southwestern Honshu, Japan (Legrand, 2009). Description.—Trilete microspore. Amb rounded triangular. The laesurae are narrow (less than 1 μm), straight, and extend to 3/4 of the spore radius. Both faces are covered by granula uniformly spaced. Exine thin (about 1 μm). Equatorial diameter = 25–45 μm. Botanical affinities.—Filicopsida. Subinfraturma Baculati Genus Baculatisporites Pflug and Thomson in Thomson and Pflug, 1953 Type species: Baculatisporites primarius (Wolff, 1934) Pflug and Thomson in Thomson and Pflug, 1953


209

Figure 4. 1, Biretisporites potoniaei (Delcourt and Sprumont, 1955) Delcourt, Dettmann and Hughes, 1963, proximal face, Kitadani Fm, 2c-M50/3; 2, Biretisporites sp., proximal face, Kitadani Fm, 2a-C51-b; 3, Todisporites major Couper, 1958, proximal face, Kitadani Fm, SEM3f; 4, Todisporites minor Couper, 1958, proximal face, Kitadani Fm, 2-2c-K50/2; 5, Concavissimisporites punctatus (Delcourt and Sprumont, 1955) Brenner, 1963, proximal face, Kitadani Fm, 2c-K67/2; 6, Cyathidites australis Couper, 1953, proximal face, Kitadani Fm, 2b-N34/3; 7, Granulatisporites sp., proximal face, Kitadani Fm, SEM-3e; 8, Osmundacidites wellmanii Couper, 1953, proximal face, Kitadani Fm, 2xK41/2; 9, Impardecispora apiverrucata (Couper, 1958) Venkatachala, Kar and Raza, 1969, proximal face, Kitadani Fm, 2b-S32/3; 10, Manumia japonica Legrand, Pons, Nishida and Yamada, 2011, proximal face, Kitadani Fm, 2a-X48; 11a–b, 12, Leptolepidites psarosus Norris, 1969 (11a–b, proximal and distal faces, Kitadani Fm, 3-3-M44/4; 12, distal face, Kitadani Fm, 3-3-P51/1); 13, Baculatisporites comaumensis (Cookson, 1953) Potonié, 1956, distal face, Kitadani Fm, 2a-M51/2; 14a–b, Baculatisporites sp., proximal and distal faces, Kitadani Fm, 2bN61; 15, Anapiculatisporites cooksonae Playford, 1965, Kitadani Fm, 2a-O61/1; 16, Echinatisporis sp., Kitadani Fm, 2a-V61/2; 17, Cicatricosisporites hughesi Dettmann, 1963, distal face, Kitadani Fm, SEM-3f. Scale bar: 10 μm.

210

Julien Legrand et al. Baculatisporites sp. Figures 4.14a–b

Baculatisporites sp. D. in Legrand, 2009, p. 143, pl. IV, figs. 5-6.

Occurrence.—Kitadani Fm (horizons Kita-2, 4). Description.—Trilete microspore. Amb rounded triangular. The laesurae are straight and extend to 3/4 of the spore radius. The proximal face is psilate. The equatorial area and the distal face are densely covered by small bacula (0.5–1.5 μm high and 0.1–0.5 μm wide). Exine 1– 1.5 μm thick. Equatorial diameter = 28–40 μm. Botanical affinities.—Osmundales, Osmundaceae.

from the same laesura. The distal face is ornamented by three sets of six muri parallel to the equator, among which one goes on with three or four of the muri to the distal pole (Figure 9). The ornamentation of this form corresponds to the type I-B defined by Krutzsch (1963). The muri join near the apices where they form thickenings (Figure 9). Equatorial diameter = 50–60 μm. Remarks.—This form seems to be similar to Cicatricosisporites sp. reported by Williams and Bujak (1980) from the Valanginian of Deep Sea Drilling Project Site 416, located in the Atlantic Ocean northeast of Africa. Botanical affinities.—Schizaeales, Anemiaceae. Cicatricosisporites sp. 1

Subinfraturma Nodati Figures 5.1a–b

Genus Echinatisporis Krutzsch, 1959 Type species: Echinatisporis varispinosus (Pocock, 1962) Srivastava, 1975 Echinatisporis sp. Figure 4.16 Echinatisporis sp. E. in Legrand, 2009, p. 148, pl. V, fig. 13.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Microspore. Amb oval. The laesurae are not seen. Both faces are ornamented by long echinae (4–5 μm high and 1–2 μm in basal diameter) more or less curved. Exine about 1.5 μm thick. Equatorial diameter = 35–40 μm. Botanical affinities.—Lycopsida, Selaginellaceae.

Cicatricosisporites sp. J. in Legrand, 2009, p. 161, pl. X, fig. 4.

Occurrence.—Kitadani Fm (horizon Kita-3). Description.—Small trilete microspore. Amb rounded. The laesurae are straight, bordered by narrow lips, and extend to 2/3 of the spore radius. Exine canaliculate; muri can bifurcate. On the proximal face, three sets of 6– 7 muri (about 1.5 μm wide) depart from a laesura and are parallel to the adjacent one when turning unclockwise. The distal face is ornamented by a set of about eight muri parallel to the opposite side, running from one of the apices until the distal pole; a second set of about 12 muri fans out from the pole to the other two apices (Figure 9). Equatorial diameter = 25 μm. Botanical affinities.—Schizaeales, Anemiaceae. Cicatricosisporites sp. 2

Infraturma Murornati Genus Cicatricosisporites Potonié and Gelletich, 1933 emend. Potonié, 1966 Type species: Cicatricosisporites dorogensis Potonié and Gelletich, 1933 Cicatricosisporites cf. C. sp. in Williams and Bujak, 1980 Figures 5.3a–b, 5a–b, 6 Cicatricosisporites sp. M. in Legrand, 2009, p. 162, pl. XI, figs. 1-2.

Occurrence.—Kitadani Fm (horizon Kita-3). Description.—Large trilete microspore. Amb rounded. The laesurae are straight, bordered by narrow lips, and seem to extend to 1/2 of the spore radius. Exine canaliculate; muri can bifurcate. On the proximal face, three sets of about 10 muri depart from a laesura and are parallel to the adjacent one; among the three sets, two depart

Figures 6.1a–b Cicatricosisporites sp. G. in Legrand, 2009, p. 160–161, pl. X, fig. 5.

Occurrence.—Kitadani Fm (horizon Kita-3). Description.—Trilete microspore. Amb rounded triangular. The laesurae are straight, bordered by narrow lips, and extend to 3/4 of the spore radius. Exine cicatricose. The proximal face is ornamented by three sets of 10 muri (about 0.5 μm wide, separated by furrows of the same width) departing from a laesura and parallel to the adjacent one when turning clockwise. On the distal face, 18 to 20 narrow muri are more or less parallel between them and to one set of the proximal face, fanning out from an apex (Figure 9). The ornamentation of this form corresponds to the type VI-C defined by Krutzsch (1963). Equatorial diameter = 25–50 μm. Botanical affinities.—Schizaeales, Anemiaceae.


211

Figure 5. 1a–b, Cicatricosisporites sp. 1, proximal and distal faces, Kitadani Fm, 3-9-O51; 2a–b, 4, Cicatricosisporites minor (Bolkhovitina, 1959) Pocock, 1964; 2a–b, proximal and distal faces, Kitadani Fm, 2z-D32/3; 4, proximal face, Kitadani Fm, SEM-3f; 3a–b, 5a–b, 6, Cicatricosisporites cf. C. sp. in Williams and Bujak (1980); 3a–b, Kitadani Fm, 3-7-J23; 5a–b, Kitadani Fm, 3-5-G36/1; 6, Kitadani Fm, SEM-3f; 7, 8, 10, Cicatricosisporites hallei Delcourt and Sprumont, 1955; 7, distal face, Kitadani Fm, SEM-3c; 8, proximal face, Kitadani Fm, SEM-3c; 10, proximal face, Kitadani Fm, SEM-3c; 9a–b, Cicatricosisporites pseudotripartitus (Bolkhovitina, 1961) Dettmann, 1963, proximal and distal faces, Kitadani Fm, 3-9-L36/1. Scale bar: 10 μm.

212


Figure 6. 1a–b, Cicatricosisporites sp. 2, proximal and distal faces, Kitadani Fm, 2z-S22/3; 2a–b, 3a–b, Cicatricosisporites sp. 3; 2a– b, proximal and distal faces, Kitadani Fm, 2w-L50/1; 3a–b, proximal and distal faces, Kitadani Fm, 2v-P63; 4a–b, Cicatricosisporites sp. 4, proximal and distal faces, Kitadani Fm, 2v-T57/2; 5a–b, 6, 7, 10, Ischyosporites crateris Balme, 1957; 5a, b, proximal and distal faces, Kitadani Fm, 3-2-N29/1; 6, proximal face, Kitadani Fm, SEM-3f; 7, distal face, Kitadani Fm, SEM-3f; 10, proximal face, Kitadani Fm, SEM3a; 8, 11, Retitriletes sp. 2; 8, lateral view, Kitadani Fm, SEM-3a; 11, proximal face, Kitadani Fm, SEM-3f; 9, Retitriletes sp. 1, distal face, Kitadani Fm, 2c-N56g. Scale bar: 10 μm.

Palynoflora from the Kitadani Formation Cicatricosisporites sp. 3 Figures 6.2a–b, 3a–b Cicatricosisporites sp. H. in Legrand, 2009, p. 161, pl. IX, fig. 6. Cicatricosisporites sp. I. in Legrand, 2009, p. 161, pl. IX, fig. 7.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Trilete microspore. Amb rounded triangular. The laesurae are straight, slightly raised, and extend to 3/4 of the spore radius. Exine cicatricose. On the proximal face, a small psilate contact area is followed by three sets of 3–4 muri parallel to the equator, which link the laesurae. The distal face is ornamented by 8– 9 muri slightly curved and parallel between them and to one set on the proximal face (Figure 9). The ornamentation of this form corresponds to the type VI-B defined by Krutzsch (1963). Equatorial diameter = 25–30 μm. Botanical affinities.—Schizaeales, Anemiaceae. Cicatricosisporites sp. 4 Figures 6.4a–b Cicatricosisporites sp. L. in Legrand, 2009, p. 162, pl. X, fig. 1.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Large trilete microspore. Amb rounded triangular with sides straight to slightly concave and raised tops. The laesurae are straight, slightly raised, and extend to 3/4 of the spore radius. Exine cicatricose. On the proximal face, a small psilate contact area is followed by three sets of three muri parallel to the equator, which join near the apices to form thickenings. The distal face is ornamented by three sets of three muri parallel to the equator, among which 2–3 muri of one set goes on to the distal pole (Figure 9). The ornamentation of this form corresponds to the type I-B defined by Krutzsch (1963). Equatorial diameter = 55–60 μm. Botanical affinities.—Schizaeales, Anemiaceae. Genus Reticulatisporites Ibrahim, 1933 emend. Neves, 1964 Type species: Reticulatisporites reticulatus (Ibrahim in Potonié, Ibrahim and Loose, 1932) Ibrahim, 1933

213

the distal face (muri about 1 μm high; luminae polygonal, 2–8 μm in diameter). Exine about 1.5 μm thick. A thin membrane can be seen at the equator (less than 2 μm wide). Equatorial diameter = 32–45 μm. Botanical affinities.—Schizaeales, Schizaeaceae. Genus Retitriletes van der Hammen, 1956 ex Pierce, 1961 emend. Döring, Krutzsch, Mai and Schulz in Krutzsch, 1963 Type species: Retitriletes globosus Pierce, 1961 Retitriletes sp. 1 Figure 6.9 Retitriletes sp. in Legrand, 2009, p. 154, pl. VII, fig. 7.

Occurrence.—Kitadani Fm (horizons Kita-2, 3, 4). Description.—Trilete microspore. Amb rounded triangular more or less deformed. The laesurae are not well seen. The proximal face is flat and psilate, followed by a reticulum near the equator and on the distal face (mesh polygonal, of various shapes; muri 4–7 μm high); this reticulum is topped by a more or less transparent membrane. Exine about 1.5 μm thick. Equatorial diameter = 40–60 μm. Remarks.—Junggarsporites membranceous Yu (1982), reported by Li (2000) from the Neocomian of the Tarim Basin in Northwest China, is strongly similar to our form. Botanical affinities.—Lycopsida, Lycopodiaceae (Döring et al. in Krutzsch, 1963). Retitriletes sp. 2 Figures 6.8, 6.11

Occurrence.—Kitadani Fm (horizon Kita-3). Description.—Trilete microspore. Amb rounded triangular. The laesurae are straight, slightly raised, and extend to 3/4 of the spore radius. A small psilate contact area is followed by a reticulum (luminae polygonal, 2– 3 μm in diameter; muri about 2 μm high) near the equator and on the distal face. Equatorial diameter = 45–50 μm. Botanical affinities.—Lycopsida, Lycopodiaceae (Döring et al. in Krutzsch, 1963).

Reticulatisporites spp. Figures 7.1, 7.4, 7.5

Turma Triletes Zonales

Reticulatisporites sp. B. in Legrand, 2009, p. 153, pl. VII, fig. 10-11.

Subturma Auritotriletes

Occurrence.—Kitadani Fm (horizons Kita-2, 3). Description.—Trilete microspores. Ambs rounded triangular. The laesurae are straight and extend to the equator of the spore. On the proximal face, a psilate contact area is followed by a reticulum near the equator and on

Infraturma Appendiciferi Genus Plicatella Maljavkina, 1949 emend. Burden and Hills, 1989 Type species: Plicatella trichacantha Maljavkina, 1949

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Figure 7. 1, 4, 5, Reticulatisporites spp.; 1, proximal face, Kitadani Fm, 2-2a-J49/3; 4, distal face, Kitadani Fm, 2-2a-G63m; 5, distal face, Kitadani Fm, SEM-3f; 2, 3, 6, 10a–b, Ruffordiaspora australiensis (Cookson, 1953) Dettmann and Clifford, 1992; 2, lateral view, Kitadani Fm, SEM-3c; 3, lateral view, Kitadani Fm, SEM-3f; 6, proximal face, Kitadani Fm, SEM-3f; 10a–b, proximal and distal faces, Kitadani Fm, 3-2-J66/1; 7a–b, Appendicisporites potomacensis Brenner, 1963, proximal and distal faces, Kitadani Fm, 2b-N60/2; 8a–b, Plicatella sp., proximal and distal faces, Kitadani Fm, 2w-S36; 9a–b, Contignisporites sp., proximal and distal faces, Kitadani Fm, 2y-P35. Scale bar: 10 μm.

Palynoflora from the Kitadani Formation Plicatella sp. Figures 7.8a–b Appendicisporites sp. B. in Legrand, 2009, p. 166, pl. XII, fig. 3.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Trilete microspore. Amb triangular with straight to slightly convex sides; crest (about 10 μm high) at the apices. The laesurae are thin, straight, and extend to 3/4 of the spore radius. Exine canaliculate. On the proximal face, three sets of 2–3 muri (about 1 μm wide) parallel to the equator. The distal face is ornamented by 8–10 muri parallel to one side, extending beyond the apices to form transparent crests. Equatorial diameter = 30–55 μm. Remarks.—The diagnosis of Plicatella baqueroensis (Archangelsky and Gamerro, 1966) Davies, 1985 has similarities with our description, but P. baqueroensis is larger (equatorial diameter 60 to 97 μm) than the Japanese form. Botanical affinities.—Schizaeales, Schizaeaceae (Dettmann and Clifford, 1992).

215

rounded triangular. The laesurae are straight, and extend to 3/4 of the spore radius. Exine canaliculate. The proximal face is psilate. The distal face is ornamented by four muri (about 3 μm wide) departing from the cingulum and parallel to one side. No verruca on the proximal face. All specimens have a ǻ-ratio (Dettmann, 1963) comprised between 1 and 1.5. Equatorial diameter = 25–45 μm; cingulum width = 2 μm. Botanical affinities.—Polypodiales, Pteridaceae (Filatoff and Price, 1988). Turma Monoletes Suprasubturma Acavatomonoletes Subturma Azonomonoletes Infraturma Laevigatomonoleti Genus Laevigatosporites Ibrahim, 1933 emend. Schopf, Wilson and Bentall, 1944 Type species: Laevigatosporites vulgaris (Ibrahim in Potonié, Ibrahim and Loose, 1932) Ibrahim, 1933

Subturma Zonotriletes Infraturma Cingulati Genus Cingulatisporites Thomson in Thomson and Pflug, 1953 Type species: Cingulatisporites levispeciosus Pflug in Thomson and Pflug, 1953 Cingulatisporites sp. Figure 8.2 Cingulatisporites sp. C. in Legrand, 2009, p. 168, pl. XIV, fig. 9.

Occurrence.—Kitadani Fm (horizons Kita-2, 4). Description.—Small cingulate trilete microspore. Amb rounded. The laesurae are straight, slightly raised, and extend to the cingulum. Exine scabrate. Equatorial diameter = 20–25 μm; cingulum width = 2–4 μm. Botanical affinities.—Filicopsida.

Laevigatosporites sp. Figure 8.13 Laevigatosporites sp. B. in Legrand, 2009, p. 178, pl. XVII, fig. 7.

Occurrence.—Kitadani Fm (horizon Kita-3). Description.—Monolete microspore. Amb beanshaped, broadly elliptical in lateral view; proximal face slightly concave. The laesura is straight and runs on about 1/3 of the length of the proximal face. Exine psilate, about 1.5 μm thick. Equatorial diameter = 43 μm; polar diameter = 28 μm. Botanical affinities.—Filicopsida. Incertae sedis Incertae sedis sp. 1 Figure 8.6 Incertae sedis sp. D. in Legrand, 2009, p. 179, pl. XVII, fig. 4.

Genus Contignisporites Dettmann, 1963 Type species: Contignisporites glebulentus Dettmann, 1963 Contignisporites sp. Figures 7.9a–b

Occurrence.—Kitadani Fm (horizon Kita-2, 3, 4). Description.—Cingulate trilete microspore. Amb

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Small trilete psilate microspore. Amb rounded triangular. The laesurae are straight, slightly raised, and extend to the equator of the spore. Exine 1– 1.5 μm thick. Equatorial diameter = 15 μm. Botanical affinities.—Filicopsida.

216


Figure 8. 1, Cingulatisporites sp. 1 in Legrand et al., 2011, proximal face, Kitadani Fm, 3-10-M21/3; 2, Cingulatisporites sp., proximal face, Kitadani Fm, 2c-K69/3; 3, 4, Gleicheniidites senonicus Ross, 1949; 3, proximal face, Kitadani Fm, 3a-O35/2; 4, proximal face, Kitadani Fm, 3cX61b; 5a–b, Coronatispora valdensis (Couper, 1958) Dettmann, 1963, proximal and distal faces, Kitadani Fm, 2c-O32/4; 6, Incertae sedis sp. 1, proximal face, Kitadani Fm, 2a-K50/3; 7, Aequitriradites spinulosus (Cookson and Dettmann, 1958) Cookson and Dettmann, 1961, Kitadani Fm, 2w-D46; 8, 9, Aequitriradites sp. in Legrand et al., 2011; 8, Kitadani Fm, 2w-F44/2; 9, Kitadani Fm, SEM-3d; 10a–b, Triporoletes reticulatus (Pocock, 1962) Playford, 1971, proximal and distal faces, Kitadani Fm, 2w-V49; 11a–b, Couperisporites complexus (Couper, 1958) Pocock, 1962, Kitadani Fm, 2-2c-G53bg; 12, Laevigatosporites ovatus Wilson and Webster, 1946, lateral view, Kitadani Fm, 2w-J66/1; 13, Laevigatosporites sp., lateral view, Kitadani Fm, 3-3c-P72/1; 14, 15, Exesipollenites tumulus Balme, 1957 subspecies triangulus Liu in Song, Zheng, Liu, Ye, Wang and Zhou, 1980; 14, Kitadani Fm, 2a-L32/2; 15, Kitadani Fm, 3a-R61/1; 16, Incertae sedis sp. 2, Kitadani Fm, 2a-K51/3. Scale bar: 10 μm, except 11b: 5 μm.


Cicatricosisporites cf. C. sp. in Williams and Bujak (1980)

Cicatricosisporites sp. 1

Cicatricosisporites sp. 3 Figure 9.

217



Ornamentation of the schizaealean forms corresponding to taxa placed in open nomenclature.

Incertae sedis sp. 2 Figure 8.16 Incertae sedis sp. E. in Legrand, 2009, p. 179, pl. XI, fig. 3.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Large microspore. Amb oval. The laesurae are not seen. Exine canaliculate. The muri of the proximal and distal faces cross each other, and seem to gather to form a thickening. Length × width = 75 × 50 μm. Botanical affinities.—Schizaeales or Gnetales?. ANTETURMA VARIEGERMINANTES = POLLENITES

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Bisaccate pollen grain. Shape is wider than long, with sacci and central body of similar size. Central body is spherical. Sacci are oval, folded, slightly hanging, and wrap the proximal face of the central body. Exine about 4 μm thick. Total width = 55 μm. Central body (length × width) = 40 × 30 μm. Saccates (length × width) = 40 × 20 μm. Botanical affinities.—Coniferales, Pinaceae. Turma Plicates Subturma Costates (= Polyplicates) Infraturma Costati

Turma Saccites Subturma Disaccites Genus Cedripites Wodehouse, 1933 Type species: Cedripites eocenicus Wodehouse, 1933 Cedripites sp. Figure 10.2 Cedripites sp. A. in Legrand, 2009, p. 183, pl. XVIII, figs. 9-10.

Genus Ephedripites Bolkhovitina, 1953 ex Potonié, 1958 Type species: Ephedripites mediolobatus Bolkhovitina, 1953 ex Potonié, 1958 Ephedripites sp. 1 Figure 10.5 Ephedripites sp. A. in Legrand, 2009, p. 190, pl. XX, fig. 17.

Occurrence.—Kitadani Fm (horizons Kita-2, 3). Description.—Polyplicate pollen grain. Amb ellipsoi-

218


Figure 10. 1, Callialasporites dampieri (Balme, 1957) Sukh-Dev, 1961, Kitadani Fm, 2y-X35/4; 2, Cedripites sp., polar view, Kitadani Fm, 2w-M35; 3, 4, Classopollis torosus (Reissinger, 1950) Couper, 1958 emend. Burger, 1965; 3, polar view, Kitadani Fm, 3-3-S24/10; 4, polar view, Kitadani Fm, SEM-3f; 5, Ephedripites sp. 1, lateral view, Kitadani Fm, 2x-R47/4; 6, Ephedripites sp. 2, lateral view, Kitadani Fm, 2-2c-Q62/3; 7, 8, Gnetaceaepollenites sp.; 7, lateral view, Kitadani Fm, 2w-Q56; 8, polar view, Kitadani Fm, 2w-Q58; 9, Exesipollenites tumulus Balme, 1957, polar view, Kitadani Fm, 2c-Q57/3; 10, Chomotriletes minor (Kedves, 1961) Pocock, 1970, Kitadani Fm, 3-6-P55; 11, Cycadopites sp., Kitadani Fm, SEM-3d; 12, Incertae sedis sp. 3, Kitadani Fm, 2-2a-F37/4; 13, 14, prasinophycean spores; 13, Kitadani Fm, SEM-3e; 14, Kitadani Fm, SEM-3e; 15, 16, Cymatiosphaera sp.; 15, Kitadani Fm, 2w-S36; 16, Kitadani Fm, 2a-S28/4; 17, Incertae sedis sp. 4, Kitadani Fm, 2b-K37/1; 18, 19, cross-fields of Coniferales, radial section; 18, Kitadani Fm, 3-9-W30/2; 19, Kitadani Fm, 3-7-D43/1; 20, vessel with scalariform perforation plates and opposite intervessel pits, Kitadani Fm, SEM-3c; 21, bennettitalean vascular element, Kitadani Fm, 2-2a A39/3. Scale bar: 10 μm.

Palynoflora from the Kitadani Formation dal. The exine is ornamented by about 25 longitudinal straight ribs; they are separated by furrows less than 0.5 μm wide, and join at the two ends of the grain. Ribs sections semicircular, about 1–1.5 μm wide. Length = 90 μm; width = 35 μm. Botanical affinities.—Gnetales, Ephedraceae (Ephedra).

219

Occurrence.—Kitadani Fm (horizon Kita-3). Description.—Monosulcate pollen grain. Amb elliptical with apices slightly sharpened. A wide furrow runs along the whole grain. Exine psilate (about 1.5 μm thick). Length = 55 μm; width = 30 μm. Botanical affinities.—Cycadales or Bennettitales (Balme, 1995).

Ephedripites sp. 2 Figure 10.6 Ephedripites sp. B. in Legrand, 2009, p. 190, pl. XX, fig. 21.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Polyplicate pollen grain. Amb oval. The exine is ornamented by about 10 longitudinal straight ribs; they are separated by furrows 1–1.5 μm wide, and join at the two ends of the grain. Ribs sections semicircular, about 3 μm wide. Length = 38 μm; width = 24 μm. Botanical affinities.—Gnetales, Ephedraceae (Ephedra). Genus Gnetaceaepollenites Thiergart, 1938 Type species: Gnetaceaepollenites ellipticus Thiergart, 1938 Gnetaceaepollenites sp. Figures 10.7, 10.8 Gnetaceaepollenites sp. A. in Legrand, 2009, p. 190, pl. XX, fig. 1314, 16, 18.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Polyplicate pollen grain. Amb ellipsoidal. The exine is ornamented by about 12–15 ribs running from one end of the grain to the other. Ribs 1.5– 2.5 μm wide, separated by furrows less than 0.5 μm wide. Ends of the ribs closely gather at each end of the long axe, without joining. Length = 40–45 μm; width = 22–30 μm. Botanical affinities.—Gnetales. Subturma Monocolpates (Monosulcites) and Zonocolpates Infraturma Quasilaevigati and Microsculptati Genus Cycadopites Wodehouse, 1933 ex Wilson and Webster, 1946 Type species: Cycadopites follicularis Wilson and Webster, 1946 Cycadopites sp. Figure 10.11

Incertae sedis Incertae sedis sp. 3 Figure 10.12 Incertae sedis sp. H. in Legrand, 2009, p. 195–196, pl. XXI, fig. 10.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Polyplicate pollen grain. Amb ellipsoidal fusiform, very deformed. The exine is ornamented by many straight narrow ribs (less than 1 μm) that longitudinally run along the grain, separated by furrows of the same width (about 20 ribs can be seen). The ribs merge at each end of the grain. Length = 55–60 μm; width = 25–28 μm. Remarks.—This form shows morphological similarities with Jugella sp. cf. J. sibirica Mtch. and Shakhm reported by Li (2000) and Yu et al. (1982) from the Lower Cretaceous of northern and central China. Botanical affinities.—Gymnosperms, Gnetales. ALGAE Genus Cymatiosphaera Wetzel, 1933 ex Deflandre, 1954 Type species: Cymatiosphaera radiata Wetzel, 1933 Cymatiosphaera sp. Figures 10.15, 10.16 Incertae sedis sp. J. in Legrand, 2009, p. 202, pl. XXIV, figs. 7-8.

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Polygonal-shaped body. The face seen is delimited by a polygonal, smoothly undulating “muri,” extended on its sides by six other polygons. Ornamentation psilate to scabrate. Equatorial diameter = 30 μm. Botanical affinities.—Chlorophyta, Prasinophyceae. Incertae sedis Incertae sedis sp. 4 Figure 10.17 Incertae sedis sp. I. in Legrand, 2009, p. 202, pl. XXIV, fig. 9.

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Table 2. Botanical affinities of all the taxa encountered during our investigation. Phylogenic classification is modified from Hilton and Bateman (2006), Smith et al. (2006), Chase and Reveal (2009) – APG III - and Friis et al. (2009). Abbreviations used for the Tamodani Flora: A, Akaiwa Formation (Kimura, 1975); C, Chinaboradani Formation (Kimura and Horiuchi, 1979); K, Kitadani Formation (Yabe and Kubota, 2004); O, probably upper part of the Akaiwa Subgroup, exposed in the upper stream of the Ogamigo Valley (Yabe et al., 2003). BOTANICAL AFFINITY Class

Subclass [Bryophyta]

FOSSILS

Order

Marchantiidae

Family

Microflora

Unknown

Lycopodiidae

Lycopodiales

Lycopodiaceae

Retitriletes austroclavatidites, Retitriletes sp. 1, Retitriletes sp. 2

Selaginellidae

Selaginellales

Selaginellaceae

Anapiculatisporites cooksonae, Echinatisporis varispinosus, Echinatisporis sp.

Equisetidae

Equisetales

[Lycophyta]

Equisetaceae Polypodiaceae

Equisetites sp. in Kimura, 1975 (A) Baculatisporites comaumensis

Aspleniaceae Polypodiales

Macrofossils from the Tamodani Flora

Aequitriradites spinulosus, Aequitriradites verrucosus, Aequitriradites sp. in Legrand et al., 2011, Couperisporites complexus, Triporoletes reticulatus

Pteridaceae

Asplenium dicksonianum Heer (C) Contignisporites sp., Manumia japonica

Arctopteris sp. in Kimura, 1975 (A)

Dennstaedtiaceae Biretisporites potoniaei Saccolomataceae Cicatricosisporites sinuosus

Cyatheales

Schizaeales [Monilophyta]

Cibotiumspora paradoxa, Concavissimisporites verrucosus, Converrucosisporites sp. in Legrand et al., 2011, Dicksoniaceae or Cyathidites australis, Cyathidites minor, Deltoidospora hallii, Cyatheaceae Impardecispora apiverrucata, Ischyosporites crateris, Leptolepidites psarosus

Anemiaceae

Cicatricosisporites hallei, Cicatricosisporites hughesi, Cicatricosisporites minor, Cicatricosisporites pseudotripartitus, Cicatricosisporites sp. 2 in Legrand et al., 2011, Cicatricosisporites cf. C. sp. in Williams and Bujak, 1980, Cicatricosisporites sp. 1, Cicatricosisporites sp. 2, Cicatricosisporites sp. 3, Cicatricosisporites sp. 4, Ruffordiaspora australiensis

Schizaeaceae

Appendicisporites potomacensis, Ischyosporites crateris, Plicatella sp., Reticulatisporites spp.

Matoniaceae

Auritulinasporites deltaformis

Polypodiidae

Gleicheniales

Gleicheniaceae

Osmundales

Osmundaceae

[Embryophyta]

Unknown

Unknown Ginkgooidae

Ginkgoales

Cycadidae

Cycadales

Cyatheaceae: Birisia onychioides (Vassilevskaja and Kara-Mursa) Samylina (A, C), Coniopteris sp. cf. C. arctica (Prynada) Samylina (C), Cyathocaulis naktongensis Ogura (O) Dicksoniaceae: Onychiopsis elongata (Geyler) Yokoyama (A, C)

Auritulinasporites deltaformis, Gleicheniidites senonicus

Gleichenites nipponensis Oishi (C), Gleichenites porsildi Seward (A)

Baculatisporites comaumensis, Baculatisporites sp., Biretisporites potoniaei, Osmundacidites wellmanii, Todisporites major, Todisporites minor

Osmunda? sp. in Kimura and Horiuchi, 1979 (C), Osmundopsis sp. in Kimura, 1975 (A), cf. Osmundopsis sp. cf. O. efimoviae Samylina (C)

Biretisporites sp., Cingulatisporites sp., Coronatispora valdensis, Granulatisporites sp., Laevigatosporites ovatus, Laevigatosporites sp., Polycingulatisporites reduncus

Adiantopteris sp. in Kimura, 1975 (A), Cladophlebis ex gr. denticulata (Brongniart) Fontaine (A), Cladophlebis sp. cf. C. pseudolobifolia Vakhrameev (A), Jacutopteris sp. in Kimura, 1975 (A), Sphenopteris kochibeana (Yokoyama) Oishi (A)

Incertae sedis sp. 1, Incertae sedis sp. 2 Ginkgoidium? sp. in Kimura, 1975 (A), Ginkgoites sp. in Kimura, 1975 (A), Pseudotorellia sp. in Kimura, 1975 (A), Sphenobaiera? sp. in Kimura, 1975 (A)

Ginkgoaceae Cycadaceae Taxaceae or Cupressaceae

Cycadopites minimus, Cycadopites sp.

Nilssonia sp. in Kimura, 1975 (A)

Exesipollenites tumulus, Exesipollenites tumulus subsp. triangulus, Spheripollenites psilatus, Taxodiaceaepollenites hiatus

Podocarpaceae or Araucariacites australis, Balmeiopsis limbatus, Araucariaceae Callialasporites dampieri Pinaceae Pinidae

Coniferales

Cedripites sp.

Cheirolepidiaceae Classopollis torosus, cuticle fragments and or Voltziaceae cross-fields of Cheirolepidiaceae Inaperturopollenites sp. in Legrand et al., 2011

[Gymnosperms] Unknown

Unknown

Pteridospermales

Unknown

Caytoniales

Caytoniaceae

Bennettitales

Unknown

Alisporites thomasii, Alisporites sp. in Legrand et al., 2011 Vitreisporites pallidus Cycadopites minimus, Cycadopites sp., vascular element of Bennettitales

Erdtmanithecales Erdtmanithecaceae Eucommiidites minor, Eucommiidites troedssonii BEG group

Ephedraceae

Ephedripites montanaensis, Ephedripites sp. 1, Ephedripites sp. 2

Unknown

Gnetaceaepollenites sp. in Legrand et al., 2011, Gnetaceaepollenites sp., Incertae sedis sp. 3

Gnetales

Unknown

Incertae sedis sp. 2 in Legrand et al., 2011

Brachyphyllum obesum Heer (K) Coniferae sp. in Kimura and Horiuchi, 1979 (C), Conites sp. in Kimura, 1975 (A), Pityophyllum lindstroemii Nathorst (A), Podozamites eichwaldi Schimper (A, C), Podozamites reinii Geyler (A)

Palynoflora from the Kitadani Formation Table 2.

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Continued. BOTANICAL AFFINITY

Phylum

FOSSILS

Class

Order

Family

Charophyceae

Zygnematales

Zygnemataceae

Microflora Chomotriletes minor, Ovoidites parvus

Pyramimonadales Cymatiosphaeraceae Cymatiosphaera sp. Algae Prasinophyceae Chlorophyta Unknown Prasinophycean spores Unknown Fungi Dothideomycetes Ascomycota

Microthyriales

Schizosporis reticulatus Unknown

Microthyriacites sp. in Legrand et al., 2011

Occurrence.—Kitadani Fm (horizon Kita-2). Description.—Uniformly reticulated body. Shape polygonal. Flat spines (about 1.5 μm thick, 5–10 μm in basal diameter, 5–10 μm high) surmount the reticulum. Equatorial diameter = 35–40 μm. Botanical affinities.—Unknown.

Results The horizon Kita-1 of the Kitadani Formation, at the lowermost part of the outcrop, is barren of palynomorphs. The horizons Kita-2, 3 and 4 contain a rich and diverse assemblage, among which the best state of preservation is observed in the horizon Kita-2. The Kitadani palynoflora consists of 79 morphospecies, among which we could identify 47 genera and 46 species of spores and pollen grains (Table 1). They are represented by 5 morphospecies of Bryophyta, 6 morphospecies of Lycophyta, 41 morphospecies of Monilophyta (dominated by 15 morphospecies of Schizaeales, 9 morphospecies of Cyatheales, and 6 morphospecies of Osmundales), 24 morphospecies of gymnosperms (dominated by 10 morphospecies of Coniferales, 6 morphospecies of Gnetales, and 2 morphospecies of Bennettitales or Cycadales) (Table 2). Spores of Filicopsida dominate the assemblage. Among them, schizaealean spores are the most abundant and diversified, with many types of Cicatricosisporites (eleven types are observed), followed by Plicatella, Appendicisporites and some rare Reticulatisporites. We note the presence of Cicatricosisporites sinuosus (Saccolomataceae), which was already reported from the Barremian Ashikajima and Kimigahama formations of the Choshi Group (Legrand, 2009; Legrand et al., 2011). Impardecispora apiverrucata (Dicksoniaceae), Ischyosporites crateris, I. estherae (Schizaeaceae or Dicksoniaceae) and Concavissimisporites verrucosus (Pteridaceae, Cyatheaceae or Dicksoniaceae) are common, and Cyathidites (Cyatheaceae or Dicksoniaceae) is rare. Osmundaceous spores are also recorded: Biretisporites potoniaei, Baculatisporites spp., Osmundacidites spp. and Todisporites spp. Gleicheniidites senonicus (Gleicheniaceae) is quite rare. The ornamented spores

Manumia japonica (Pteridaceae?), Leptolepidites psarosus and Converrucosisporites sp. are present. Umetsu and Sato (2007) reported a spore similar to Leptolepidites psarosus from the late Aptian-early Albian Hiraiga Formation (northeastern Honshu, Japan) under the name Multinodisporites sp., but the ornamentation of this genus does not correspond to our observation. Coronatispora valdensis (Filicopsida) is rare. Monolete spores of the genus Laevigatosporites are recorded in all three horizons of the formation, and were observed in groups in the horizon Kita-3. Spores of hepatics are well represented, with common Couperisporites complexus in the three horizons. Aequitriradites spinulosus, A. verrucosus and Triporoletes reticulatus are also common. Spores with lycopsid affinities are common, with Echinatisporis varispinosus and Retitriletes austroclavatidites. Among the gymnosperm pollen grains, Exesipollenites tumulus and Classopollis torosus (Coniferales or Erdtmanithecales: Tekleva and Krassilov, 2009) are most abundant. Eucommiidites (Erdtmanithecales), Ephedripites and Gnetaceaepollenites (Gnetales) are rare. Monosulcate pollen grains of the genus Cycadopites (Bennettitales, Cycadales or Pentoxylales), Araucariacites australis and Balmeiopsis limbatus (Araucariaceae) and Callialasporites dampieri (Podocarpaceae) are common. Bisaccate pollen grains are rare, represented by the genera Alisporites, Cedripites and Pityosporites. Spheripollenites psilatus (Cupressaceae or Taxaceae) is rare. In addition to spores and pollen grains, wood fragments are recorded: cross-fields of Coniferales (Figures 10.18, 10.19), a fragment of bennettitalean vascular element (Figure 10.21), a vessel element with scalariform perforation plates (Figure 10.20). Cuticles with cell walls slightly sinuous sometimes extending into trichomes (Figures 11.1a–b), and cuticles with papillae on the epidermal cells and stomata subsidiary cells (Figures 11.2– 11.5), probably belonging to cheirolepidiaceous plants, are also encountered. Epiphyllous fungi, fungi spores of Microthyriacites, prasinophycean and zygnematacean freshwater algae spores (Figures 10.11–10.14, 10.16) and the species Schizosporis reticulatus are present. We summarize the composition of the palynoflora in a

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Figure 11. 1a–b, Internal view of gymnosperm cuticle fragments, with detail on a trichome (hair), Kitadani Fm, SEM-3a; 2–5, cuticle fragments of Cheirolepidiaceae, with papillae on the epidermous cells and stomata subsidiary cells; 2, Kitadani Fm, SEM-3f; 3, Kitadani Fm, 3-9-T31; 4, Kitadani Fm, 3-2-U51b; 5, stomata with papillae on its subsidiary cells, Kitadani Fm, 3-9-O27/1. Scale bars: 10 μm.

Kita-2

Kita-3

7%

8% 20%

8%

Kita-4 10%

12%

2% 24%

10% 12%

12%

3% 11%

14% 14%

3% 1%

12% 1% 1% 25%

25%

28%

15% 1% 2%

19%

Psilate spores

Bisaccate pollen grains

Exesipollenites tumulus

Cicatricose spores

Monosulcate pollen grains

Araucariaceae

Other ornamented spores

Classopollis torosus

Other gymnosperms

Figure 12.

Percentages of nine palynomorph groups in the Kita-2, Kita-3 and Kita-4 horizons of the Kitadani Formation.

Palynoflora from the Kitadani Formation diagram (Figure 12). The horizon Kita-4 is not as well preserved as horizons Kita-2 and Kita-3, which can partly explain the differences observed in percentages.

Discussion Palynofloral characteristics of the Kitadani Formation The palynoflora from the Kitadani Formation shows a diverse assemblage. Most identified taxa have a long stratigraphic range and/or global distribution during the Cretaceous period. In the absence of angiosperm pollen grains or other biostratigraphically significant microfossils in this terrestrial deposit, we face the same difficulty as previously expressed by several authors concerning the assignation of an accurate age to the Kitadani Formation. The Kitadani palynoflora is dominated by spores of herbaceous plants (50–59%), mostly Filicopsida: Schizaeales, Cyatheales, and Osmundales; gymnosperm pollen is subdominant (21–27%) and consists primarily of Coniferales and Bennettitales, Cycadales or Pentoxylales. The Kitadani palynoflora has few Gleicheniales, bisaccate pollen grains, or pollen of Gnetales and Erdtmanithecales. It shows a general agreement in floristic composition with what has been described for the Tetoritype Paleoflora. It differs in the absence of Ginkgoales and Czekanowskiales pollen, which may be difficult to distinguish from other monosulcate pollen grains. Our palynofloral results showing dominance of Filicopsida spores and absence of matoniaceous spores confirm the Tetori-type macrofloral data of Kimura (1987). Manumia japonica, reported for the first time from the Barremian palynofloras of the Choshi Group (Legrand et al., 2011), is also present in the Kitadani Formation, but in lower proportions. Genera Appendicisporites and Gleicheniidites, as well as species Impardecispora apiverrucata, Ischyosporites crateris and Couperisporites complexus are rare to frequent in the Kitadani Formation, whereas they have not been reported from the assemblages of the Choshi Group (Legrand et al., 2011). The hepatics, common in the palynomorph assemblages, have not been reported from the macroflora. Among pollen grains, the presence of Classopollis is consistent with the report of Brachyphyllum obesum. Classopollis torosus and Exesipollenites tumulus are equally abundant and characterize the assemblage. Among Exesipollenites grains, some show an oval to subtriangular shape and an apparently triangular central body that corresponds to the subspecies E. tumulus subsp. triangulus previously reported from southeastern China (Song et al., 1980; Han and Jiang, 1981; Song et al., 1981, 1995).

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Bisaccate pollen grains are encountered in low percentages in the Kitadani Formation, with the genera Alisporites (Pteridospermales?) and Cedripites (Coniferales, Pinaceae) and the species Vitreisporites pallidus (Caytoniales). Among Ephedripites grains observed, Ephedripites montanaensis characterizes the Lower Cretaceous worldwide. Gnetales, and more particularly the genera Ephedripites and Gnetaceaepollenites, are largely represented in the Lower Cretaceous sediments of Asia, often reported under the genus name Schizaeoisporites. Paleoecology Only continental elements were encountered in the assemblages: spores and pollen grains of terrestrial plants (Marchantiopsida, Lycopsida, Filicopsida and gymnosperms), wood and cuticle fragments of gymnosperms, fungi and freshwater algae. The records of the freshwater green algae Ovoidites parvus and Chomotriletes minor are in accordance with previous reports of freshwater molluscs and bivalves, and indicate a fluvio-lacustrine environment. Fern spores of Cyathidites and Deltoidospora as well as epiphyllous fungi are encountered, implying locally humid conditions (Harris, 1937; Pedersen and Lund, 1980). A number of palynomorphs and wood remains are fragmented, suggesting long maceration or transportation. However, the fine-grained sediments in which microremains are included, with accumulation of woods, refute long-distance transportation (Tschudy, 1961). We can surmise that most of the assemblages probably derived from the local community near the depositional area. Terrestrial elements were carried out by water (runoff) or wind from the adjacent areas, which can explain the rare occurrence of bisaccate pollen grains, for example. The latter may also have not been very abundant in the source vegetation. Among spores, the record of groups of Laevigatosporites ovatus represents immature spores still grouped in the sporangium, and confirms the proximity of the plant producing these spores to the site of deposition. Paleoclimate Inner Japan, as the easternmost part of the Eurasian continent, was attributed to a humid subtropical latitudinal climatic belt in some paleoclimatic reconstructions for the Early Cretaceous (Kimura, 1987; Chumakov et al., 1995; Golozoubov et al., 1999). The genera Cyathidites and Deltoidospora are common. The diversity of schizaealean spores associated to some gleichenialean spores, the low percentages of bisaccate pollen grains, and the abundance of monosulcate pollen grains, are consistent with a warm and humid climate. Cheirolepidiaceous-like wood remains and cuti-

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cle fragments have been recorded, particularly in the horizon Kita-3. Some cuticles show trichomes or papillae, indicating an adaptation to dry (papillae on stomata subsidiary cells, which extend them to protect the substomatal cavity from desiccation) and sunny (papillae on epidermous cells) conditions (Pons, 1979; Watson and Alvin, 1996). Concerning the macroremains previously reported from the Kitadani Formation, only the Coniferales Brachyphyllum obesum has been identified (Yabe and Kubota, 2004), so the cuticle fragments encountered may belong to this species. Cheirolepidiaceous plants are considered to have formed mangroves, and also to have been adapted to arid or semiarid conditions (Srivastava, 1976; Upchurch and Doyle, 1981; Pons and Koeniguer, 1985). In the adjacent regions of South China (semiarid to arid climate), Classopollis is much more abundant (50%) than in Japan (12–15% in the Kitadani Formation of the Inner Zone, and 9–14% in the Choshi Group of the Outer Zone; see Legrand et al., 2011), indicating different climatic conditions. Ruffordiaspora australiensis (Schizaeales, Anemiaceae) is present in the formation; this spore was produced by Ruffordia goepperti (Dunker) Seward, whose macroremains were previously reported from many Lower Cretaceous localities in Europe (England: Seward, 1894; Spain: Diéguez and Meléndez, 2000), China (Deng and Chen, 2001), North America (Skog, 1992) and Japan (Kuwajima Formation: Yabe, 1922; Kimura, 1987). This plant is thought to have lived near lakes and to have tolerated dry conditions (Hughes and Moody-Stuart, 1966). Various paleoenvironments are represented in the site of deposition. Our results indicate a warm temperate and humid climate, with locally drier conditions. They are in accordance with previous hypotheses by Kimura (1979, 1987) and Vakhrameev (1991).

Conclusion We identified 47 genera and 46 species of spores and pollen grains from the late Barremian to early Aptian? Kitadani Formation. This palynofloral description provides new data to the present knowledge of the Inner Zone palynofloras previously studied by Umetsu (2002), Umetsu and Matsuoka (2003) and Umetsu and Sato (2007), who identified seven morpho-genera of spores and pollen grains in a bad state of preservation. Among palynomorphs, only continental elements have been recorded, suggesting a fluvio-lacustrine environment with locally drier conditions. The composition of the assemblages indicates that most of the elements probably come from the local community near the depositional area. Our results are consistent with the hypothesis by Vakhrameev (1991) that the Tamodani Flora belonged to

an ecotone located between the Euro-Sinian (including the Outer Zone) and Sibero-Canadian Provinces.

Acknowledgements This study has been financed by a scholarship for Education and Research offered by the Japanese Government (Monbukagakusho: MEXT), and by the University of Tokyo and Chuo University, Tokyo, Japan. We are grateful to the Fukui Prefectural Dinosaur Museum for the opportunity to collect samples from the “Kitadani Dinosaur Quarry”. Many thanks to Jean Broutin (Université Pierre et Marie Curie) for his kind advice and correction of the manuscript. We are thankful to Dr. Debra Willard and Dr. Hiroshi Kurita for accepting to review this manuscript. Their comments greatly contributed to its improvement.

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