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© Publications scientifi ques du Muséum national d’Histoire naturelle, Paris, 2018 ISSN (imprimé / print) : 1280-9659/ ISSN (électronique / electronic) : 1638-9395 Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin, East coast of India

Chopparapu H. CHINNAPPA Andhra Loyola College, Vijayawada, Andhra Pradesh (India)

Annamraju RAJANIKANTH Birbal Sahni Institute of Palaeobotany, 53 University Road, Lucknow, Uttar Pradesh (India)

Submitted on 13 October 2017 | accepted on 8 February 2018 | published on 21 June 2018

urn:lsid:zoobank.org:pub:2AE40BC8-437B-4B1B-A0E2-282EE3D876FD

Chinnappa C. H. & Rajanikanth A. 2018. — Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin, East coast of India. Geodiversitas 40 (12): 259-281. https://doi.org/10.5252/ geodiversitas2018v40a12. http://geodiversitas.com/40/12

ABSTRACT Th e Early Cretaceous plant diversity and palaeoecology of the Krishna-Godavari Basin fl ora is studied. Th e study is based on the plant fossils collected by the authors during the recent years and published reports of past work. Nature and mode of preservation of the leafy fossils were considered to under- stand the vegetation relationship. Similarly, Nearest Living Equivalent method and palaeoecological information of diverse plant groups from the published sources, along with sedimentological inputs are adopted to draw the palaeoenvironnement. Th e results show that the fl ora was dominated by KEY WORDS Flora, bennettitaleans. Th e vegetation includes plant fossils from the parauto- to allo-chthonous sources. taphonomy, Th e association of the plant fossils with marine fauna indicates the vegetation was growing near to palaeoecology, the sea. Th e phytogeographical correlation of the fl ora shows its similarity with that of Antarctica phytogeography, Early Cretaceous, and Australia in the generic composition but greatly diff er in specifi c composition. Th e composite Krishna-Godavari Basin. fl ora indicates the prevalence of warm and humid conditions.

RÉSUMÉ Analyse fl oristique et implications paléoécologiques des séquences du Crétacé inférieur du bassin de Krishna- Godavari, Côte Est de l’Inde. La diversité et la paléoécologie de la fl ore du Crétacé inférieur du bassin de Krishna-Godavari sont étudiées. Ce travail exploite des récoltes récentes et des rapports préliminaires faits par les auteurs. La nature et le mode de préservation des feuilles fossiles sont étudiés afi n de comprendre la relation entre les végétaux. Pour reconstituer les paléoenvironnements, des données sédimentologiques et des informations paléoécologiques sur des groupes variés ont été utilisées. Cette étude a aussi été complétée MOTS CLÉS par une approche fondée sur la méthode actualiste. Les résultats montrent que la fl ore est dominée Flore, taphonomie, par les bennittitales. La végétation comprend des plantes d’origines parautochtone à allochtone. paléoécologie, L’association de plantes à des éléments de faune marine indique un développement proche de la mer. phytogéographie, L’analyse de corrélation phytogéographique montre des similitudes avec l’Antarctique et l’Australie Crétacé inférieur, Bassin de Krishna- dans sa composition générique, mais cette fl ore diff ère dans sa composition spécifi que. Cette fl ore Godavari. composite indique une prévalence de conditions climatiques chaudes et humides.

GEODIVERSITAS • 2018 • 40 (12) © Publications scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 259 Chinnappa C. H. & Rajanikanth A.

an area of 28 000 km2 on shore and 24 000-49 000 km2 off shore (Rangaraju et al. 1993). Th e basin has been classifi ed as a N major intra-cratonic rift within Gondwanaland until the Early Jurassic period and it later transformed in to peri-cratonic rift basin (Biswas 1992). Sediments co-relatable to those of the Early Cretaceous (‘Upper Gondwana’) are exposed near the western and northwestern fringe of the Krishna (Budavada and Vemavaram formations) and Godavari (Golapalli and Rajasthan South Rewa Raghavapuram formations) depressions. Th e sedimentation Basin Basin Rajmahal in these Cretaceous successions was linked with the faulting of Kutch Basin Basin basement blocks as a result of reactivation of NE-SW trending Satpura Basin Precambrian faults (Biswas 1992). Pranhita- Palaeobotanical studies from the Krishna-Godavari basin Godavari have been conducted for more than a century (Feistmantel Basin Krishna-Godavari Basin 1879; Seward & Sahni 1920; Sahni 1928; Baksi 1964, 1967, 1968; Bose & Jain 1967; Jain 1968; Mahabale & Satyanarayana INDIA 1979; Vagyani 1984, 1985; Vagyani & Zutting 1986; Vag- Palar Basin yani & Jamane 1988; Pandya & Sukh-Dev 1990; Pandya et al. 1990; Prasad & Pundir 1999; Chinnappa et al. 2014, 2015; Chinnappa 2016). Th ese studies were mostly restricted to the Cauvery Basin 500 km taxonomy; however, no attempt has been made to discuss the taphoceonosis, diversity and palaeoecological implications FIG. 1 . — Locality map showing the fossil collection sites and the localization of the fl ora. Moreover, these studies were restricted either to of the of the Krishna-Godavari Basin (East Coast of India). macro- or microfl ora. Macro- or microfl oral investigations of ancient sediments can provide a partial picture; both must be considered together INTRODUCTION to get a complete picture of the vegetation (Chinnappa & Rajanikanth 2017). Th erefore, the present study aims to analyze Fluvial-paralic sediments from the East Coast of India host both the micro- and macrofl oras from the Early Cretaceous some of the important Early Cretaceous (Neocomian-Aptian) sediments of Krishna-Godavari Basin, to provide a detailed fossil fl oras recording a range of vegetation types preserved in account of taphoceonosis and to determine palaeoecological variety of depositional settings during the rifting event that diversity patterns. fragmented eastern Gondwana. Plant fossil studies from these basins (e.g. Mehrotra et al. 2012; Rajanikanth & Chinnappa 2016 and references therein) also show that sedimentary basins GEOLOGICAL SETTING AND STRATIGRAPHY: host information that contributes to our understanding of the southern Gondwana palaeogeography during the Early Th e Krishna-Godavari (KG) Basin contains about 5 km of Cretaceous period (Rao & Venkatachala 1972; Baksi 1977). sediments recording several cycles of deposition, ranging in age Geographically and tectonically, these sedimentary units are from Late Carboniferous to Pleistocene. Th e basin came into grouped under ‘Coastal Gondwanas’, which are distributed in existence following rifting along eastern continental margin of Cauvery (CV), Palar (PL), Krishna-Godavari (KG), Pranhita- Indian craton in the Early Mesozoic (Sastri et al. 1973). Faults Godavari (PG) and Mahanadi (MH) basins (Fig. 1). Th e penetrating to the basement defi ne the series of horst and grabens segmentation of these East Coast Indian sedimentary basins cascad down towards the ocean and are aligned NE-SW along is owing to the intrinsic subcrustal distinction along the east Precambrian eastern Ghat trend (Sastri et al. 1973, 1981; Rao coast. Each one of the subcrustal blocks behaved diff erently 2001). Th e fi rst attempt to describe the geology of the basin and mimicked the inherent subcrustal mosaic during the was made by Blandford et al. (1856), followed by King (1880) Gondwana period and carved out for itself an exclusive basin who carried out detailed geological studies of the outcrops. of Mesozoic-Cenozoic age. Th ese basins, however, are separated Subsequently, these successions were studied in detail by several by major tectonic elements. A major fault separates Bengal researchers (Sastri et al. 1973, 1981; Rao 1993, 2001; Laksh- from Mahanadi; Visakhapatnam high separates Mahanadi minarayana et al. 1992; Prabhakar & Zutshin 1993; Raju & from Krishna-Godavari; Nayudupeta high separates Krishna- Misra 1996; Prasad & Pundir 1999; Lakshminarayana 2002). Godavari from Palar; Chingleput high separates Palar from Th e Early Cretaceous Golapalli, Raghavapuram, Budavada and Cauvery Basins (Lal et al. 2009). Vemavaram sediments are exposed towards the western and Th e Krishna-Godavari Basin (KG) has received much atten- northwestern fringes of the basin (Fig. 2). Th ese sediments tion in recent times due to its high petroliferous/hydrocarbon were distributed in two depressions namely Godavari (Golapa- source rock potential (Mehrotra et al. 2012). It is one of the lli and Raghavapuram formations) and Krishna (Budavada and most important petroliferous basins of India and occupies Vemavaram formations). Lakshminarayana et al. (1992) revised

260 GEODIVERSITAS • 2018 • 40 (12) Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin (India)

Hyderabad Ramanujapuram Khammam

Rajahmundry 17°S Sobhanadripuram Kakinada

Vijayawada

Guntur

Machilipatnam

16° Chirala INDIA Ommevaram

Nandyal Bay of Bengal Ongole

79°E 80° 81° 82°

FIG. 2 . — Map of the Krishna-Godavari Basin (East Coast of India; after Sastri et al. 1973) (map made with http://www.simplemappr.net). the stratigraphy of these ‘Upper Gondwana’ sediments of the Sinha 1986; Venkatachala & Rajanikath 1987). Th ese forma- KG Basin and considered the Golapalli Formation under Kota tions are correlated with each other based on fl oral and faunal Formation (Jurassic) at Musnuru, Nehrunagaram and Soma- data (Sastri et al. 1973) Among these formations Golapalli, varam areas and similar rocks around Golapalli-Nuzividu areas Raghavapuram, Budavada and Vemavaram are dated as the were included under the Tirupati Formation (Late Cretaceous). Early Cretaceous and Tirupati and Pavalur are Late Cretaceous However, Prasad & Pundir (1999) and Rao (2001) treated based on mega and micro-fl oral and faunal evidences such these rocks under Golapalli Formation as originally identifi ed as ammonites, brachiopods, lamellibranchs, fi sh scales and by the King (1880). Th e tectonic framework and depositional mammalian ribs (Spath 1933; Venkatachala & Sinha 1986; environment of the basin was studied by Sastri et al. (1973, Venkatachala & Rajanikath 1987; Prasad & Pundir 1999). Th e 1981), Baksi (1977), Vasudeva Rao & Krishna Rao (1977), generalized lithostratigraphy of the basin is given in the Table 1. Venkatachala & Sinha (1986) and Rao (2001). Th e Early Cretaceous sediments exposed in the Godavari Depression are divided into the Golapalli, Raghavapuram MATERIAL AND METHODS and Tirupati formations (Fig. 3A). Th e Golapalli Formation comprises white to pale white and light brown sandstone with Plant fossil material was collected from the mudstone succes- siltstone partings. Th e Raghavapuram Formation sandwiches sion of Raghavapuram Formation, exposed as a small hillock between the Golapalli and Tirupati formations, characterized by at about 1 km towards the South East of the village Ramanu- white pale to reddish shale and red ferruginous claystone, light japuram (17°13’27’’N, 81°19’11’’E), near Jangareddigudam, buff to grayish white, medium grained glauconitic sandstone. West Godavari District, Andhra Pradesh, India (Fig. 2). Th e Th e Tirupati Formation overlies the Raghavapuram Forma- fossiliferous section is characterized by white pale to reddish tion and composed of purple red to light brown, medium mudstone/shale and red ferruginous claystone, and light buff grained sandstone. to grayish white, medium grained glauconitic sandstone. Th e Early Cretaceous sediments exposed in the Krishna Plant fossils were preserved in pale-reddish mudstones/earthy Depression are also divided into the Budavada, Vemavaram shales (Fig. 3A). Th e plant fossils were also collected from the and Pavaluru formations (Fig. 3B). Th e sedimentary rocks of Vemavaram Formation, exposed at road cutting near about Budavada Formation is composed of sandstone. Th e Vemavaram 0.5 km towards the North West and North East of the vil- Formation lies between the Budavada and Pavalur formations lage Ommevaram (15°41’17’’N, 80°09’01’’E), near Ongole, and is composed primarily of shale containing carbonaceous Prakasam District, Andhra Pradesh, India (Fig. 2). Th ese plant matter. Th e Pavalur Formation comprises of medium to coarse fossils were preserved in shale (Fig. 3B). Th e fossil leaves were grained clay and lateritic sandstone and it is overlying by the preserved as impressions, and cuticules have not been recov- Deccan intertrappeans (Rao 2001). ered. Examination was done using an Olympus SZH 10 stereo Th e Golapalli, Ragahavapuram and Tirupati formations are dissecting microscope. All specimens were photographed with considered, equivalent to the Budavada, Vemavaram and Pav- Canon SX 150 IS digital camera using either polarized light alur formations respectively (Sastri et al. 1973; Venkatachala & or low angle lighting to reveal surface details.

GEODIVERSITAS • 2018 • 40 (12) 261 Chinnappa C. H. & Rajanikanth A.

TABLE 1 . — Lithostratigraphy and ages of the various lithounits in the KG Basin (data from Sastri et al. 1973; Lakshminarayana et al. 1992; Prasad & Pundir 1999) .

Group Formation Lithology Age Recent alluvium Tertiary Rajahmundry Red, feldspathic, ferruginous, laterized, Miocene-Pliocene crossbedded and conglomeratic sandstone Infra-Inter-Trappeans Basaltic lava Late Cretaceous-Early Paleocene Unconformity - Upper Gondwana Tirupati/Pavalur Purple red-light brown sandstone/clay and Late Cretaceous caleritic sandstone Raghavapuram/ White pale-reddish earthy shale, red Early Cretaceous Vemavaram ferruginous claystone, light buff -grayish white glauconitic sandstone/shale containing carbonaceous matter Unconformity - Golapalli/Budavada White-pale white and light brown sandstone Early Cretaceous with siltstone partings/sandstone Unconformity - Lower Gondwana Chintalapudi/Kamthi Coarse grained feldspaathic sandstone, Permain alternating calcareous claystone Unconformity/Fault - Proterozoic Khondalite Igneous and metamorphic rocks Precambrian

To isolate spores and pollen, sediments from all the localities fl ora is diverse and composed of bryophytes, pteridophytes, that have yielded macrofossils were also treated with hydro- gymnosperms and angiosperms (Appendices 1, 2). Th e diversity chloric, hydrofl uoric and nitric acid (HCl-HF-HNO3) and and abundance patterns of these plant groups, however, are not sieved (mesh width 25 mm). However, only samples from the consistent between the macro- and microfl oras. Raghavapuram were productive. Th e slides were studied under an Olympus BH 2 microscope, fi tted with a digital camera. All BRYOPHYTES samples and slides (BSIP 40156-40192 and 40243-40285) are Macrofossils belonging to this group have not (yet) been recorded deposited in the repository of Birbal Sahni Institute of Palaeo- to date. In contrast, spores are well represented and consti- botany for future reference. tute members of Anthocerataceae (Foraminisporis Krutzsch), Th e taxonomic affi nities at family level of the spore/pol- Sphaerocarpaceae (Aequitriradites Delcourt & Sprumont, len obtained are after Ramanujam & Rajeshwar Rao (1979). emend. Cookson & Dettmann and Coptospora Dettmann) Similarly, the taxonomic affi nity of the pteridophytic fronds and Reillaceae (Cooksonites Pocock, and Staplinisporites Pocock) at family level follows Harris (1969) and Barbacka & Bodor (Ramanujam 1957; Kar & Sah 1970; Venkatachala & Sinha (2008). However, the taxonomic affi nities at family level are not 1986; Prasad & Pundir 1999; Mehrotra et al. 2012). Th e certain for many gymnosperm taxa; many of these taxa could qualitative and quantitative composition of the group is less be related to more than one family. Macro-and microfl oral spe- when compared to the other spore producing plants such as cies diversity was analysed separately by considering the total pteridophytes (Appendix 2). number of taxa known in the fl ora. Th e reports of the present study and previous studies are taken into consideration for this PTERIDOPHYTES purpose. Th e taxonomic diversity of the fl ora is presented in Pteridophytes are represented in both macro-and microfl oras, pie diagrams as simple percentage representation of each group especially the microfl ora. Fossil fronds belong to the Marattiaceae, (at order level), generated by using MS Excel. Similarly, the Osmundaceae, Gleicheniaceae and Dicksoniaceae and a few Sphe- abundance of the various plant taxa was calculated by count- nopteris Sternberg type fossils of unknown affi nity were recorded ing the number of samples for the given taxa. Th e diff erence (Fig. 4). Many families such as the Lycopodiaceae, Selaginel- between the micro-and macrofl ora is explained with regard to laceae, Matoniaceae, Schizaeaceae, Marsileaceae, Cyatheaceae taphonomy and natural variations. and Polypodiaceae are only represented in the microfl ora. Among these families, the Schizaeaceae and Cyatheaceae display highest species diversity (Fig. 5). Th e Schizaeaceae include six genera COMPOSITION OF THE FLORA encompassing a total of 17 species and Cyatheaceae constitute 23 species in eight genera (Appendix 2). A few more pterido- Th e synthesis of macro-and microfl ora from the Early Creta- phytic spore taxa, cannot be assigned to any family with certainty ceous sequences of the Krishna-Godavari Basin shows that the and they are placed in pteridophyte incertae sedis (Appendix 2).

262 GEODIVERSITAS • 2018 • 40 (12) Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin (India)

A Tirupati Formation

B Recent alluvial

Pavalur Formation Raghavapuram Formation

Conglomeratic Sandstone

Shale Vemavaram Formation Vemavaram

Fossiliferous Bed

Sandy shale

Claystone

3 m 0.5 m Silty sandstone

Budavada Sandstone Golapalli Formation Formation

FIG. 3 . — Litho-column of fossiliferous sections exposed at Ramanujapuram (A) and Ommevaram (B), Krishna-Godavari Basin (East Coast of India).

GEODIVERSITAS • 2018 • 40 (12) 263 Chinnappa C. H. & Rajanikanth A.

Marattiaceae fossil leaves are rare in the Krisha-Godavari Th innfeldia Ettingshausen and Pachypteris (Brongniart) Harris fl ora, there are only a couple of reports of these fern fronds with single species in each genus (Appendix 1). Of the two from the Golapalli Formation (Feistmantel 1876, 1877). genera the former was reported from the Vemavaram and Th e family is represented by a single genus with one species: Raghavapuram formations (Feistmantel 1879; Baksi 1968; Marattiopsis macrocarpa (Oldham & Morris) Seward and Chinnappa 2016) and the latter is known from the Vema- Sahni. Spores related to this family have not been reported. varam and Golapalli formations (Feistmantel 1876; Pandya & Macrofossils of Osmundaceae are represented by fi ve species Sukh-Dev 1990) belonging to two genera: Cladophlebis Brongniart and Todites Traditionally, Th innfeldia type leaves reported from the early Seward emend. Harris (Appendix 2). Within the two gen- Cretaceous of India were variously assigned to Dicroidium era, the former is known from the both Raghavapuram and Gothan and Th innfeldia Ettingshausen (Feistmantel 1879, Golapalli formations but latter is known only from Golapalli 1882; Lele 1962; Rao & Lele 1963; Baksi 1968; Jain 1968). Formation (Pandya & Sukh-Dev 1990; Prasad & Pundir Many of these specimens were originally described under 1999). From the Vemavaram Formation the macrofossils of Th innfeldia by Feistmantel (1879, 1882). Later, however, Lele the Osmundaceae are altogether absent. All the species were (1962) and Rao & Lele (1963) re-evaluated these fossil leaf preserved as pinnae fragments, whole fronds are unknown. types and transferred them from Th innfeldia to Dicroidium. Pinnules were well preserved with a distinct venation pattern. While doing so, Lele (1962) and Rao & Lele (1963) also Th e Krishna-Godavari fl ora includes spores related to Osmun- considered the similar fossil leaves were also in Dicroidium daceae in four genera – Baculatisporites Pfl ug & Th omson, despite their origin from the Early Cretaceous succession and Biretisporites (Delcourt & Sprumont) Delcourt et al., Osmun- morphological variations (Townrow 1957). Subsequently, dacidites Couper, and Todisporites Couper, altogether with fi ve Baksi (1968) and Jain (1968) also placed the leaf fossils species (Appendix 2). Osmundacidites type of spore is known resembling Th innfeldia in Dicroidium instead. Dicroidium as produced by Todites (Naugolnykh 2002). Although, the is characterized by dichotomized rachis with frequent sphe- relationship of rest of the spores with Cladophlebis and Todites nopteroid to taeniopteroid venation in the pinnules whereas is not known, their spores are considered to be produced by Th innfeldia lacks the dichotomized rachis and shows frequent Osmundaceae members (Kustatscher et al. 2010). odontopteroid and alethopteroid venation. Further, Th inn- Gleicheniaceae includes a single genus, Gleichenia Smith, feldia is considered as geologically younger (Jurassic) than the with two species (Appendix 1), which are reported from the Dicroidium (Triassic). Chinnappa (2016) has considered the Golapalli Formation (Feistmantel 1876; Prasad & Pundir 1999). leaves described under Dicroidium from the Early Cretaceous Microfossils are represented by three genera: Gleicheniidites sediments of India under Th innfeldia based on the morphology Ross, Ornamentifera Bolkhovitina and Plicifera Bolkhovitina, and stratigraphic distribution. Recently, Cleal & Rees (2003) altogether with seven species. considered Th innfeldia as taxonomic synonym of Pachypteris; From Dicksoniaceae a single genus Onychiopsis Yokoyama, nevertheless, we believe that the Indian specimens referred to represented by a single species (Appendix 1) was found from Th innfeldia needs a critical re-examination before considering the Golapalli Formation (Prasad & Pundir 1999). Th e genus them under Pachypteris. corresponds well with the living fern Onychium Kaulf in its morphology and spore morphology. Only a fragmentary CYCADALEANS specimen is known in the studied fl ora, not enough mor- Foliage resembling that of modern-day cycads has long pre- phological details were observed hence the specimen is placed sented palaeobotanists with major problems, especially when under the comparative form O. psilotoides Stokes & Webb on dealing with foliage lacking cuticles. Taeniopteris Brongniart the basis of gross morphological similarity (Prasad & Pundir leaves are one such type that has a poorly understand systematic 1999). A number of spore genera are known to originate from position (Pott & Launis 2015; Van Konijnenburg-Van Cittert Cyatheaceae/Dicksoniaceae and they are the most abundant et al. 2017; Chinnappa & Rajanikanth 2017). Th e affi nity of in the present microfl ora (Appendix 2). leaves of this type can be clarifi ed only when they associated with reproductive structures or preserved with cuticle (Cleal & GYMNOSPERMS Rees 2003; Pott & Launis 2015; Van Konijnenburg-Van Cit- Gymnosperms are important components of the Mesozoic tert et al. 2017). However, in the studied locality the leaves fl oral ecosystems, dominating until the Late Cretaceous are known only as imprints, with no cuticles or associated (Vakhrameev 1991). However, they started to decrease in their reproductive structures having been found to date. Leaves of diversity and abundance, with the appearance of angiosperms this type are here represented by a single species Taeniopteris in the Early Cretaceous and their subsequent expansion in spatulata McClelland known from all the three litho-units: the Late Cretaceous (McLoughlin 2001; Friis et al. 2011). Vemavaram (Feistmantel 1879), Raghavapuram (Baksi 1968; In the Krishna-Godavari fl ora, the gymnosperms constitute Chinnappa 2016) and Golapalli (Feistmantel 1876; Pandya & a major share. Sukh-Dev 1990; Prasad & Pundir 1999). Because the pre- sent leaves are preserved as impressions and there is a large PTERIDOSPERMS uncertainty regarding the systematic affi nity of the genus, we Pteridosperms are rare components in the Early Cretaceous placed them under incertae sedis within the gymnosperms as Krishna-Godavari fl ora and are represented by two genera: suggested by Cleal & Rees (2003) and Pott & Launis (2015).

264 GEODIVERSITAS • 2018 • 40 (12) Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin (India)

A BC D EF

G HI

JKL M

FIG. 4. — Plant material preserved from the KG basin: A, Ptilophyllum cutchense (Morris) Bose & Kasat; B, Ptilophyllum acutifolium (Morris) Bose & Kasat; C, Anomo- zamites sp.; D, Taeniopteris sp.; E, Otozamites vemavaramensis Bose & Jain; F, Ptilophyllum rarinervis (Feistmantel) Bose & Kasat; G, Dictyozamites ommevaramensis Chinnappa, Rajanikanth & Rao; H, Dictyozamites feistmantelii Bose & Bano; I, Ginkgo sp.; J, Pityospermum godavarianum Chinnappa, Rajanikanth & Rao; K, Araucarites raghavapurensis Chinnappa, Rajanikanth & Rao; L, Elatocladus loyolii Chinnappa, Rajanikanth & Rao; M, Brachyphyllum sehoraensis Bose & Maheshwari. Scale bars: 1 cm.

GEODIVERSITAS • 2018 • 40 (12) 265 Chinnappa C. H. & Rajanikanth A.

A B

CD

E F

G

FIG. 5 . — Plant material preserved from the KG basin: A, Thinnfeldia vemavaramensis (Feistmantel) Chinnappa, Rajanikanth & Rao; B, Brachyphyllum expansum (Sternberg) Seward; C, Pterophyllum footeanum Feistmantel; D, Elatocladus loyolii Chinnappa, Rajanikanth & Rao; E, Pagiophyllum ommevaramensis Chinnappa, Rajanikanth & Rao; F, Pagiophyllum cf. rewaensis Bose & Sukh-Dev; G, Elatocladus loyolii. Scale bars: 1 cm.

266 GEODIVERSITAS • 2018 • 40 (12) Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin (India)

Upland podocarpean and taxavean vegetation

Low/wet-land bennettitalean-ginkgolean and araucarian vegetation

Ocean

Legend Herbaceous angiospermic vegetation Riverbank bryophytes, fern Ferns and cycadalean vegetation Coastal pteridospermalean, bennettitalean and araucarian vegetation

Pteridospermalean Cycadalean Bennettitaleans Bennettitaleans Bennettitaleans (Pachypteris/Thinnfeldia) (Taeniopteris) (Otozamites/Anomozamites) (Ptilophyllum) (Pterophyllum)

Araucarians Bennettitaleans Podocarpians Taxacean Ginkgo (Bracyphyllum/ Angiosperm (Dictyozamites) (Elatocladus) (Taxites/Torreyites) Pagiophyllum)

FIG. 6 . — The macrofl oral diversity of the various plant groups in the Krishna-Godaveri Basin (East Coast of India).

BENNETTITALEANS being mostly found in Vemavaram Formation (Feistmantel Bennettitalean leaves are most common in the Early Cre- 1879; Chinnappa et al. 2015; Chinnappa 2016). Th ese gen- taceous Krishna-Godavari fl oras. Six genera: Anomozamites era are represented by six species each (Appendix 1). Speci- Schimper, Dictyozamites Oldham, Otozamites Braun, emend. mens belonging to Dictyozamites are rarely observed in the Watson & Sincock Pterophyllum Brongniart, Ptilophyllum Raghavapuram and Golapalli formations and are represented Morris, and Dictyozamites Oldham were identifi ed fol- in the fl ora by only a few fragmentary leaves (Baksi 1964; lowing the criteria given by Harris (1969) and Watson & Pandya & Sukh-Dev 1990). Otozamites is altogether absent Sincock (1992). from Golapalli Formation. Th e other bennettitalean foliages Among these, Ptilophyllum fronds are the most common such as Anomozamites and Pterophyllum are comparatively and widely distributed throughout all the three formations. less common. From each of these genera three species are Twelve species have been reported to date (Appendix 1), documented (Appendix 1). Th e former is reported only from of which six are assigned to the comparative forms such as the Vemavaram, whereas the latter known from all the three P. distans (Feistmantel) Bose & Kasat, P. cf. institacallum areas (Feistmantel 1879; Chinnappa et al. 2015). Bose, P. amarjolense (Bose) Bose & Kasat, P. cf. gladiatum Little fertile bennettitalean material has been described Bose & Sukh-Dev, P. cf. horridum (Roy) Bose & Kasat, P. cf. from Gondwana fl oras to date (Cantrill 2000). However, a jabalpurense (Jacob & Jacob) Bose & Kasat (Appendix 1). few specimens of Cycadolepis Saporta and Williamsonia Car- Ptilophyllum acutifolium (Morris) Bose & Kasat, and P. cutch- ruthers are present in the fl ora studied here (Seward & Sahni ense (Morris) Bose & Kasat are most common in the studied 1920; Baksi 1968; Pandya & Sukh-Dev 1990). Cycadolepis plant assemblage. Th e identifi cation of these species based is generally presumed to represent the bract from the base of only on the gross morphology. To date, cuticle has not been a female cone (Cantrill 1997a), while Williamsonia is con- prepared from the Krishna-Godavari fl oras. Dictyozamites and sidered to be a bennettitalean strobilus (Harris 1969) and Otozamites leaves are nearly as common as Ptilophyllum, but they are represented here by two species each (Appendix 2).

GEODIVERSITAS • 2018 • 40 (12) 267 Chinnappa C. H. & Rajanikanth A.

GINKGOALEANS Th e other signifi cant conifers found in the fl ora are taxa Members of the Ginkgoales are most commonly encoun- belonging to Elatocladus where the genus is represented by tered from the Krishna-Godavari Basin; however, they are six species (Appendix 1). Among them E. plana (Feistmantel) restricted to the Raghavapuram Formation (Feistmantel Seward is common and widely distributed. Th e specimens of the 1877; Baksi 1967, 1968; Vagyani 1985; Chinnappa et al. genus from Vemavaram show comparatively large sized leaves 2015). Th is indicates that the plants were growing in compared to material from the Raghavapuram and Gangapur. abundance in the basin. Mesozoic leaves resembling mod- In the Krishna-Godavari fl ora, the genus is represented only ern ginkgo leaves are usually placed in the genus Ginkgo as impressions of vegetative leafy twig, the cuticles and repro- Linnaeus or Ginkgoites Seward. Seward (1919) and Tralau ductive structures have so far not been recovered. Th erefore (1967) maintained the distinction between the two gen- the precise taxonomic affi nity of the genus is unclear. Since era based on the nature of lobes in leaf. Leaves of Ginkgo the genus Elatocladus from India has already been related to are divided into two or more lobes by shallow notches Podocarpaceae (Chinnappa & Rajanikanth 2017), it is treated which never reach the basal part of the lamina, whereas here under the same family i.e., Podocarpaceae. Pollen allied in Ginkgoites leaves are deeply and symmetrically divided to the Podocarpaceae is known under four genera: Micro- into narrow segments (Tralau 1967). However, Harris & cachrydites Cookson, emend. Couper, Platysaccus Naumova, Millington (1974) considered these distinctions are not emend. Potonié, Podocarpidites Cookson, emend. Couper, and applicable in reality as trees of Ginkgo biloba can produce Podosporites Rao and constitute a great deal of the microfl ora, many deeply divided leaves. Chinnappa (2016) merged all both qualitatively as well as quantitatively. Among them, the Indian specimens previously described under Ginkgoites Podocarpidites shows the highest species diversity with eight with Ginkgo, thus considering the recommendations of species (Appendix 2). Th e quantitative representation of the Harris & Millington (1974). Th e genus here represented genus is also very high and alone represents more than 20% in by four species which gives 5% of species diversity in the the pollen spectra (Venkatachala & Sinha 1986). Less abun- Krishna-Godavari basin (Appendix 1). Th e pollen grain dant are the genera Podosporites represented by three species Ginkgocycadophytus Samoilovitch (with two species) is the respectively. Microcachrydites and Platysaccus are represented only possible genus represented in the Krishna-Godavari by a single species each (Appendix 2). fl ora probably produced by ginkgoaleans (Appendix 2). Th e Taxales are comparatively rare with only a single genus and species of macro-remains Torreyites constricta CONIFERALEANS Sahni being known only from the Vemavaram Forma- Although the abundance of the conifer remains is low in the tion (Sahni 1931). A very few plant fossil records can be fl ora they show considerable taxonomic diversity (Appendices 1, assigned to this genus with confi dence (Sahni 1931). Th e 2). Eight genera are represented by macroremains: Araucarites vegetative shoots bearing spirally arranged and distichously Presl, Brachyphyllum Brongniart, Pagiophyllum Heer, Elatocladus placed linear leaves characterized by two well marked sto- Halle, Torreyites Seward, Conites Sternberg, Harrisiophyllum matal grooves on the lower surface and absence of distinct Pant, Srivastava & Pant, and Pityospermum Nathorst. Th e midrib resembling in habit those of Torreya were usually number of species identifi ed within each genus is given in assigned under Torreyites (Seward 1919). However, because Appendix 1. the bands represent grooves to which the stomata were Th e genus Araucarites, belonging to the Araucariaceae, confi ned referral to Torreyites is not always possible when includes four species (Appendix 1) and is distributed to cuticles are not preserved (Seward 1919; Sahni 1931). Th e all the three formations: Vemavaram (Feistmantel 1879) only reliable character is absence of a distinct midrib and Raghavapuram (Chinnappa et al. 2015) and Golapalli (Feist- the characteristic leaf shape. Although the fossil leaves were mantel 1876). Th e genus Araucarites is based on an ovuliferous identifi ed based on the characters of the extant genus, with- cone scale that resembles those of recent Araucariaceae. Ovu- out reproductive organs it is impossible to determine the liferous scales with a single ovule/seed and a free distal ligule precise systematic position of shoots of this common form are indicative of its affi nity with the Araucariaceae (Cleal & and these generic names do not imply any direct relation- Rees 2003). Our specimens of Araucarites are preserved with ship with the extant members (Seward 1919). Th e records a woody bract and a centrally fused seed mark. of taxalean woods (Taxaceoxylon) in India during Mesozoic Leafy axes belonging to Brachyphyllum and Pagiophyllum are times (Rajanikanth & Sukh-Dev 1989) however, suggest common among the conifer remains in the Krishna-Godavari their possible existence as suggested by Sahni (1928). fl ora (Sahni 1931; Baksi 1968; Pandya & Sukh-Dev 1990; Genera like Harrisiophyllum Pant, Srivastava & Pant and Chinnappa et al. 2015). Th ese genera both encompass fi ve Pityospermum Nathorst are known from a single isolated species each (Appendix 1). Th e taxonomic relationship of specimen each (Chinnappa et al. 2015). Although cuticles Pagiophyllum and Brachyphyllum is a unclear at the family level of the Harrisiophyllum were described from Bansa, South (see Chinnappa & Rajanikanth 2017). Th ese taxa are herein Rewa Formation by Pant et al. (1983), they did not provide provisionally placed under the Araucariceae after Bose & enough information to narrow down the taxonomic affi nity Maheshwari (1975). Pollen belonging to the Araucariaceae is of the genus to the family level. Records of Pityo spermum, known as Araucariacites Cookson and Callialasporites Sukh- a winged seed, are not common from the Early Cretaceous Dev (Appendix 2). sediments of India. Pityospermum sp., from the Sriperum-

268 GEODIVERSITAS • 2018 • 40 (12) Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin (India)

budur Formation, Palar Basin located in Tamil Nadu, India (Jeyasingh & Kumarasamy 1994) and from P. godavarianum 2% the Raghavapuram Formation (Chinnappa et al. 2015) were 14% 2% the only reports known to date. Th e precise taxonomic affi n- ity of the seeds of this type is not clear. Th e striking resem- blance between such seeds and seeds of recent Pinus and other 35% Abietineaceae (Seward 1919) suggest their possible affi nities Pteridophytes with Pinaceae or/and Abietineaceae. However, until more Pteridosperms evidence is found it is not possible to confi rm their family 42% Cycadaleans affi nity. Th erefore, here we consider them under the broad Bennettitaleans group Coniferales. Recovery of Abietineaepollenites pollen 5% belonging to Abietineaceae from the Krishna-Godavari Basin Ginkgoaleans (Ramanujam 1957) is noteworthy. Coniferaleans Classopollis Pfl ud emend. Pocock & Jansonius are abundant in the Early Cretaceous successions of India (e.g. Ramanu- jam & Rajeshwara Rao 1979; Venkatachala & Sinha 1986) FIG. 7 . — The macrofl oral diversity of the various plant groups in the Krishna- Godaveri Basin (East Coast of India). and other Gondwanan and non Gondwanan land masses (Vakhrameev 1991). Th ey are represented here by three spe- cies (Appendix 1). In spite of rich records of Classopollis from 4% the Early Cretaceous sediments of India, nothing is known 8% about their parent plants. Th e type of pollen is generally pre- 11% sumed to be produced by Cheirolepidiaceae (Venkatachala 1966). However, such pollen is shown to be produced by wide Bryophytes variety of fossil-taxa including Pagiophyllum and Brachyphyl- 9% Pteridophytes lum type fossils (Kendall 1949; Couper 1955; Venkatachala 52% 1966; Srivastava 1976; Tosolini et al. 2013). Such leaf types Ginkgoaleans Coniferaleans belong to the Cheirolepidiaceae (Tosolini et al. 2013), the 15% Podocarpaceae and Araucariaceae (Harris 1979). In India, Incertae sedis gymnosperms Classopollis pollen has mostly been recovered from the same Incertae sedis sediments yielding the species of Pagiophyllum and Brachy- Angiosperms phyllum (Ramanujam & Rajeshwara Rao 1979). However, 1% it has never been recovered from in situ cones attached with Pagiophyllum and/or Brachyphyllum. Th e taxonomic affi ni- FIG. 8 . — The microfl oral diversity of the various plant groups in the Krishna- ties of this foliage in India (see Chinnappa & Rajanikanth Godaveri Basin (East Coast of India). 2017), and the parent taxa of Classopollis necessitates awaiting further evidence. Conifer remains in the Early Cretaceous Krishna-Godavari FLORAL DIVERSITY fl ora include two species of detached strobili placed under Conites (Appendix 2). Although there can be little doubt that MEGAFLORA it is a conifer reproductive organ not much is known about Taxonomic analysis of the megafl ora, which includes leaves and their structure and their precise affi nity. A number of pollen cone scales (Figs 4; 5) demonstrate the presence of pteridophytes, forms of unknown conifer affi nity also occur in the Krishna- and gymnosperms. A total of 81 species under 20 genera have Godavari basin. Th ey are listed in the Appendix 2. been reported to date. Th e pteridophytes constitute 11 species under six genera representing 14% in the fl ora. Th e gymnosperms ANGIOSPERMS are a major element in the fl ora, they include pteridosperms, Angiosperm macrofossil remains from the Early Cretaceous bennettitaleans, cycadaleans, ginkgoleans and conifers. With two sequences of India are rare (Chinnappa & Rajanikanth 2017). species and two genera the pteridosperms comprise 2%. Th e ben- Microfossil reports from subsurface and surface data from nettitaleans are dominant elements of the fl ora and they include the Early Cretaceous sequences of India clearly indicate their 34 species under six genera with 42% of the species share. Th e occurrence by this time (Mehrotra et al. 2012). Th e Early conifers constitute 28 species under 8 genera and they represent Cretaceous microfossil assemblage from Krishna-Godavari 35% of the total diversity. Th e ginkgoleans and cycadaleans Basin includes about thirteen species of angiosperm pollen are minor components in the fl ora in terms of diversity. Th e (Appendix 2). A small assemblage of megafossils resembling former include four species under a single genus and make up fl owering plants were recovered from the Early Cretaceous 5%, and the latter consists of two species and one genus with sequences Krishna-Godavari Basin (Chinnappa 2016). Th ese 2%. Although, the species diversity of the ginkgoleans is less, fossil taxa mostly resemble an aquatic plants with ribbon they are extremely abundant in the Raghavapuram Formation. shaped and dissected leaves. Th e percentage share of each group has shown in the Figure 7.

GEODIVERSITAS • 2018 • 40 (12) 269 Chinnappa C. H. & Rajanikanth A.

MICROFLORA of the fern leaves can be explained by their delicate structure Th e Early Cretaceous microfl oras (spore/pollen) from the Krishna- and poor preservation potential (Gastaldo 1988, 1992; Spicer Godavari Basin are rich and diverse (Ramanujam 1957; Kar & 1991). However, spores of this group are diverse and abun- Sah 1970; Venkatachala & Sinha 1986; Prasad & Pundir 1999; dant suggesting these plants were probably not from the far Mehrotra et al. 2010, 2012; and present study). Th ese studies distance and their source might be close to the depositional reported a range of taxa affi liated to the bryophytes, pterido- site (Venkatachala & Sinha 1986; Spicer 1991).Th e leaves of phytes, gymnosperms and angiosperms (Appendix 2). Although, Taeniopteris are rare and are recovered as isolated fragments the conifer pollen dominates the assemblage, pteridophytes (Pandya & Sukh-Dev 1990; Prasad & Pundir 1999; Chin- spores are also very abundant. Th e conifers include 23 species nappa et al. 2015) suggesting that perhaps they were produced under seven genera and it comprises 15% in the fl ora. Other by plants that did not grow in the immediate vicinity of the gymnospermous taxa include ginkgoaleans and taxa of uncertain depositional environment. Th e bennettitaleans are common affi nity (incertae sedis). Th e ginkgoaleans consists of two species and well preserved in many cases the leaves preserved most under a single genus and make up a small percentage (1%) in of its length (Fig. 4B). Th ese compound leafy rachises with the fl ora. Th e taxa placed in incertae sedis include 14 species in intact pinnae are suggesting their rapid burial and minimal nine genera and represents 9%. Th e pteridophytes, composed transportation (Ferguson 1985). Th erefore, the parent plant of 82 species, are grouped under 39 genera and occupy 52% of that produced these leaves is here considered as a local and was species diversity. Th e presence of diverse and abundant spores probably growing within or very near the depositional envi- suggests the luxurious growth of the group (Appendix 2). Th e ronment. Ginkgo leaves are common in the Krishna-Godavari percentage share of each group is shown in the Figure 8. fl ora and are locally more abundant in the Raghavapuram Th e signifi cant contributions of the microfl ora are the reports Formation. Th ese leaf types are well preserved with petioles, of bryophytic spores and angiosperm pollen (Mehrotra et al. however, in a few cases both the base and apex of the leaf are 2010, 2012). Presence of bryophytic spores suggests exist- missing (Fig. 4I). Th is suggests leaves might be exposed to ence of the group in the Krishna-Godavari fl ora, although short distance transportation. Th e pollen of this group is also the megafl oral elements have not yet been reported. Th e well preserved. It is rational to consider them as local con- bryophyte spores are comparatively rare and they include stituents of the fl ora based on the preservational attributes. only seven species in six genera and comprise 4% of species Th e conifers in the fl ora include araucarians, podocarps and diversity. Th e records of the angiospermous pollen are of special members of the Taxaceae. Th e araucarians constitute moderately interest here. Th ey are here represented by 13 species under preserved leafy axes with branching preserved (Fig. 5B, E), and 10 genera, which occupy 8% of species diversity in the total cone scales with clear seed marks (Fig. 4K). Th e preservation fl ora. Th ere are a few more taxa of uncertain affi nity, which mode of these taxa suggests considerable transportation before are here referred to incertae sedis, they include 18 species in their fossilization, consequently they are considered here as 13 genera and constitute 11% of species diversity. regional elements. Th e podocarps occur sporadically in the fl ora; they are mostly represented by isolated fragmentary leaf axes and a winged seed (Figs 4L; 5D, G). Th ey possibly TAPHONOMIC IMPLICATIONS travelled into the depositional site from regional uplands. High frequency of saccate pollen produced by Podocarpaceae Th e concentration and degree of fragmentation of the plant is also indicative of its upland habitat (Venkatachala & Sinha fossils has been often used as indicative of original vegita- 1986; Abbink et al. 2004). Th e saccate pollen must have been tional relationship (auto-allochthonous). Plant taphonomic fl ew from the uplands and entered into the depositional sites. studies by Ferguson (1985), Spicer & Greer (1986), Gastaldo Th e megafl oral records of bryophytes are not known. Th e (1988), Spicer (1991) and others suggested that the degree presence of the spores such as Foraminisporis, Coptospora, of fragmentation can be related to the distance travelled by a Cooksonites and Stereisporites indicates that the bryophytes given plant material before its fossilization. Th is fragmenta- also constitute as part of the fl ora (Mehrotra et al. 2012). tion determines that leaves in general cannot be transported Th e absence of macrofossil remains is probably because of long distances before their burial and fossilization (Spicer their poor preservation capacity and taphonomic constraints 1991). Additionally, the fragmentation of the plant parts in (Reader & Stewart 1972; Spicer 1991). Th e bryophytes in the fossil fl oras is also due to other physical and biological general include small herbs with delicate foliages that make factors (Ferguson 1985). Generally, the fossils also get badly them diffi cult to preserve in fossil record. mutilated during the process of recovery from sediments like Th e early angiosperms are mostly thought to be herbs and/ highly cracked and jointed mudstone (Baksi 1968). or small shrubs (Taylor & Hickey 1996; Sun et al. 2002; Th e fragmentary nature of the recovered specimens here Field et al. 2004, 2009). Th e herbaceous nature might be a indicates that the plant fossils were subjected to transportation possible reason for the dearth of their macrofossils (Spicer (Behrensmeyer & Hook 1992). Th e degree of fragmentation 1991), yet their pollen can be preserved. Angiosperms are among the various plant assemblages is not uniform, thus sug- rare in the Indian Early Cretaceous fl ora and represented by gesting distances travelled by these plant fossils diff er. Ferns a few poorly preserved leaves which are ribbon shaped and were mostly recovered as isolated fragmentary pinnules and dissected (Chinnappa 2016). Th e characteristic shape of these their concentration is poor. Th e paucity and fragmentation leaves suggest their aquatic nature.

270 GEODIVERSITAS • 2018 • 40 (12) Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin (India)

Late Jurassic Early Cretaceous

Neotethys West Gondwana Neotethys SA Af India Ma India

East Gondwana An Au

FIG. 9 . — Global Palaeogeography map during the Jurassic and Cretaceous periods (after Chatterjee et al. 2013).

Th e above account suggests that many of the plant fossils Ferns have mostly been considered to grow ‘‘under moist, were most likely exposed to some pre-depositional sorting and rather warm conditions either in marshes, along riverbanks abrasion suggesting that perhaps they were produced by plants or as understorey in forests’’ (Van Konijnenburg-Van Cittert that did not grow in the immediate vicinity of the deposition 2002). Only a few taxa (e.g. Gleichenia [Gleicheniaceae], environment. Th is could be the probable reason for the low Phlebopteris and Weichselia [Matoniaceae]) can tolerate full diversity and low number of specimens in the samples and par- sunlight and are able to adapt to the stress related conditions ticularly explain the lack of delicate remnants such as sporangia (Van Konijnenburg-Van Cittert 2002; Abbink et al. 2004; and frond fragments of bryophytes in the megafossil fl ora. Schrank 2010). However, the majority of the Jurassic-Cretaceous Nonetheless, the study also indicates the fl ora is taxonomically ferns are considered to be elements of moist lush vegetation less diversifi ed as it is (Chinnappa 2016). Experimental observa- (Harris 1961) often occurring near river banks (Pelzer et al. tion of leaf transportation in fl uvial systems by Spicer & Greer 1992). Th e group is generally believed to attain high abun- (1986) indicated that the maximum distance travelled in ideal dances under relatively humid conditions optimal for plant conditions is less than 1.5 km. Th us, the fl oral components growth. Th is is highlighted by relations between lithologies were probably drifted from local to regional sites. Th erefore, indicative of moist environments with high spore abundances the present assemblage is considered to represent parauto- to (Maheshwari & Jana 2004). Th e ferns in the Krishna-Godavari allo-chthonous elements and the fl ora mostly includes the local lithologically associated with claystone/mudstone and silty to regional vegetation. It has also been demonstrated that a sandy stone. Th is facies association indicates overbank and low energy condition within a fl uvial system was inferred on backswampy depositional settings (Boggs 2006) the basis of grain size parameters (Rao 2001). Fluvial settings Th e pteridosperms such as Pachypteris indica (Oldham & generally refl ect the local fl ora though the herbaceous com- Morris) Bose & Roy and Th innfeldia vemavaramensis (Feist- ponent is rare (Burnham 1989). mantel) Chinnappa, Rajanikanth & Rao are associated with the sandstone and shale/mudstone facies respectively. Th e former is also known from the other basins like Kutch, Sat- PALAEOECOLOGICAL IMPLICATIONS pura, Rajmahal and Mahanadi (Rajanikanth & Chinnappa 2016), while the later is exclusive to the Krishna-Godavari Extant bryophytes mostly grow in moist and shady areas, as Basin. Pachypteris indica is considered to produce large sized they require water to complete their life cycle. Only a few bushes that formed mangrove thickets along the river mouth species are known to withstand water stress but even these inundated by tides (Banerji 2004). Th e frequent association taxa require water during the reproductive stage (Abbink et al. of Th innfeldia vemavaramensis with marine fossil fauna and 2004; Schrank 2010). Th erefore, this group is here considered leathery leaves suggest its coastal habitat. to grow at river banks and back-swampy settings as indicated Mesozoic cycadaleans are considered to occupy the low-lands by their association with silt and mud (Boggs 2006). Th e and share the forest fl oor with ferns and favour fairly boggy con- swampy environments is also supported by an unequal dis- ditions (Abbink et al. 2004). Taeniopteris spatulata McClelland tribution of phytoplanktons (Venkatachala & Sinha 1986) is the only possible cycadalean taxa reported from the Krishna-

GEODIVERSITAS • 2018 • 40 (12) 271 Chinnappa C. H. & Rajanikanth A.

Godavari Basin. Th is taxon is considered to be grown on the ecologically saturated environments such as wetlands (Royer river fl oodplain, distal to the river channel, but occupied fairly et al. 2003; Zhou 2009). Th e low diversity of the genus in the boggy conditions in clearings or near the forest margins (Chin- modern fl ora also suggests this ecologically restricted distribu- nappa & Rajanikanth 2017). Where light was reduced from tion (Alvin & Watson 1996). Th e Ginkgo leaves reported from a closed canopy of conifers, cycadaleans colonized the forest the Krishna-Godavari Basin are associated with mudstone fl oor in patches and together with fern thickets and presumably facies, which is indicative of fl oodplain deposits (Boggs 2006). preferred areas of the canopy with less dense stands of trees. Th e coniferous taxa such as Brachyphyllum regularis Borkar & Th e members of bennettitaleans were considered to grow Chiplonkar, B. sehoraensis Bose & Maheshwari, Pagiophyllum in diverse niches, ranging from delta areas to coastal margins gollapallensis Pandya & Sukh-Dev, and Pagiophyllum cf. grantii or in upland areas (Vakhrameev 1991; Abbink et al. 2004). Bose & Banerji are found in an assemblage consists of ferns However, it is generally assumed that most species favored and cycadophytes, suggesting they share similar habitats such humid environment (Pott et al. 2008) and preferred to grow as lowlands. Th e presence of thin cuticle and amphistomatic at coastal margins (Vakhrameev 1991) and lowlands (Krassilov condition in Brachyphyllum sehoraensis and Pagiophyllum cf. 1975). Th e spatial heterogeneity in the distribution pattern of grantii (Bose & Maheshwari 1973; Bose & Banerji 1984; various taxa in the fossil assemblages of the studied fl ora and also indicate their growth at high water availability. Yet some their association with diff erent sedimentary facies support the species of Brachyphyllum such B. feistmantelii (Halle) Sahni, above views. Th e taxa like Otozamites acutifolius Feistmantel, B. rhombicum Feistmantel, Pagiophyllum cf. marwarensis Bose & O. exhislopi Bose, O. imbricatus Feistmantel, O. gondwanensis Sukh-Dev and P. ommevaramensis Chinnappa, Rajanikanth & Bose, D. hallei Sahni & Rao, D. ommevaramensis Chinnappa, Rao preferred to grow at coastal regions as indicated by their Rajanikanth & Rao, Pterophyllum footeanum Feistmantel, lithofacies association with shale and frequent mixing with P. incisum Sahni & Rao and Ptilophyllum heterophylla Chin- marine fauna. Th e fertile parts reported here probably produced nappa, Rajanikanth & Rao are rich and exclusive to the near by Brachyphyllum and/or Pagiophyllum. Extant members of sea deposits and frequent association of the leaves with marine araucarians frequently grow near shores as they can withstand fauna suggests coastal habitat. Presence of recurved leaf margin, the infl uence of salt wind (Vakhrameev 1991). Th eir leathery strongly cutinized guard cells and sunken stomata as observed leaves and thick cuticles are adaptations to such type harsh in the species of Otozamites (Bose 1974; Bose & Banerji 1984) conditions. Evidence from the megafossil record indicates that and characteristic leathery nature of the leaves is considered araucarian trees usually grow in lowlands, probably coastal to be a strategy to minimize the water loss in the coastal zone margins and preferred cooler environments (Venkatachala (Vakhrameev 1991; Villar de Seoane 2001; Wright et al. 2005). 1966; Ramanujam 1980; Vakhrameev 1991). Th e association of these leaves with shale is also in congru- Th e members of podocarpaceae are numerically less repre- ence with above interpretations (Boggs 2006). Similarly, the sented in the megafl ora when compared to any other members, association of the other bennettitaleans such as Dictyozamites but they are abundant in the microfl ora. In spite of their robust indicus Feistmantel, Pterophyllum kingianum Feistmantel, Pti- nature, the week representation of podocarps in the fl ora, in lophyllum deodikarii Mahabale & Satyanarayana, P. tenerrimum terms of their abundance suggest they probable growing away (Feistmantel) Bose & Kasat, Ptilophyllum cf. institacallum Bose, from the depositional site and fi nd diffi cult to incorporate in the Ptilophyllum cf. amarjolense Bose, Ptilophyllum cf. gladiatum fossil assemblages (Spicer 1991). Being able to fl y with wind the Bose & Sukh-Dev, Ptilophyllum cf. horridum Roy and Ptilophyl- pollen might had reached the depositional site and preserved lum cf. jabalpurense Jacob & Jacob with cycadaleans and ferns in good number. Today podocarps are particularly common indicates that, they are dwelling in the lowland areas. Th eir in mountain areas of the tropics and subtropics. During the sedimentary association with claystone and clay sandstone also Mesozoic, the family inhabited relatively dry areas of upland suggests fl ood deposits (Boggs 2006). Th e taxa like Dictyozamites forest (Vakhrameev 1991) or cool, wet upland forest (Doyle feistmantelii Bose & Bano, Ptilophyllum acutifolium (Morris) et al. 1982). High frequency of saccate pollen produced by Bose & Kasat, P. cutchense (Morris) Bose & Kasat have wide Podocarpaceae is also indicative of its upland habitat (Abbink distribution and they are common in all type of lithologies et al. 2004). Th e association of taxaceans with podocarps is and found in every fossil assemblage. Th is suggests that these indicative of their similar distribution. taxa might favor the wide ecological niches. Th e palaeoeocology of the angiosperms in the present fl ora Species of Ginkgo occur frequently in the fl ora; they repre- is not clear due to the limited megafossils evidance and lack sent mostly deciduous moist loving plants, growing alongside of knowledge regarding their systematic affi nities. Th e char- conifers. Although, the extant species is restricted to China, acteristic shapes of these leaves (ribbon shaped and dissected) the ginkgoleans had wide distribution during the Mesozoic suggest their aquatic nature (Chinnappa 2016). It has been (Royer et al. 2003). Mesozoic ginkgoaleans seem to have been believed that the early angiosperms were herbs/shrubs and able to tolerate a wide range of climates from warm and wet mostly preferred to grow near water bodies (Feild et al. 2009). temperate (or even cool) in coastal plain and lowland to inland Th e angiosperms pollen reported here, possibly originated from riparian/swamp environments, respectively. In spite of their the plants of the similar habitat. broad adaptability, however, it appears that the ginkgoaleans From the above discussion that the spore-producing bryo- on the whole were more abundant and diverse in mesic, warm phytes/pteridophytes are generally believed to prefer river banks temperate to temperate climates and inhabited stable and and still some may have been underground cover under ben-

272 GEODIVERSITAS • 2018 • 40 (12) Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin (India)

INDIA ANTARTICA AUSTRALIA

Krishna- Godavari

Pranhita-Godavari Basin Basin Cauvery Basin Palar Basin Mahanadi Basin Kutch Basin Rajasthan Basin Satpura Basin South Rewa Basin Rajmahal Basin Alexander Island Livingston Island Larsen Basin Perth Basin Carnarvon Basin Canning Basin Basin Maryborough Surat Basin Basin Eromanga Otway/Gippsland Basin

TP Mc Fm Fm All Mst Albian WB Fm GCK Tlb Fm Bur. Sst N.G. Fm Fre.

KP Fm Osborne Fm Ss.

Ss.

Windalia Radiolarite k Parda Wal. Fm Wal. Fm Wal. Aptian An Fm

Mud Eumeralla Fm Fm Womnhaggi LP Fm CN Fm

Pluto Glacier Fm Sh Dan Ss. Bir. Ss. Barremian Pedersen Fm Strzelecki Group Bungil Fm Bullsbrook Fm Bullsbrook Leederville Fm Cronin Ss. Cronin Warnbro Group Warnbro Cadna-owie Fm EARLY CRETACEOUS EARLY Hauterivian Spartan Glader Fm Fossil Bluff Group Fossil Bluff Gangapur/Chikiala Fm Budavada Fm Fm Vemavaram Golapalli Fm Raghavapuram Fm Sivaganga Fm Sripernumbudur Fm Athgarh Fm Bhuj Fm Fm Gardeshwar Pariwar Fm Jabalpur Fm Bansa Fm Rajmahal Fm CP Beds Rintoul Ck Fm Kumbarilla Beds Broome Ss. Broome South Perth Shale Crayfish Subgroup Valanginian Neocomian Mooga Ss. CC Fm Ora Fm Hooray Sst. Yarraloola Fm Yarraloola Nanutarra Fm Callawa Fm Sst. Berriasian Gob. Tyers Cong. Tyers Parmelia Fm FmGrahams Creek Fm Maryborough Piliga Sst. JUR. Tithonian Himalla Ridge Fm PB Fm W Fm Nordenskjold Fm Nordenskjold

FIG. 10 . — Correlation chart of the diff erent Early Cretaceous basins from India, Antarctica and Australia, deducted from the information available in this article. Abbreviations: All Mst, Allaru Mudstone; Ank Ss, Anketell Sandstone; Bir Ss, Birdrong Sandstone; Bur, Burrum Coal Measures; CC Fm, Chester Cone Formation; CN Fm, Cerro Negro Formation; Cong, Conglomerate ; CP Beds, Crabeater Point Beds; Dan Ss, Dandaragan Sandstone; Fre Ss, Frezier Sandstone; GCK, Griman Ck Formation; Gob. Sst, Gobb Sandstone; KP Fm, Kotick Point Formation; LP Fm, Lagrelius Point Formation; Mc Fm, Mackunda Formation; Mud Sh, Muderong Shale; N.G. Fm, Neptune Glacier Formation; Ora Fm, Orallo Formation; PB Fm, President Beaches Formation; Sst, Surat Siltstone; Tlb Fm, Toolebuc Formation; TP Fm, Triton Point Formation; Wal. Fm, Wallumbilla Formation; WB Fm, Whisky Bay Formation; W Fm, West Formation.

nettitaleans and conifers. Th e cycadaleans probably grew on environmental conditions. During the Early Cretaceous, Indian the river fl oodplain, distal to the river channel, but preferred subcontinent along with Australia and Antarctica constitute fairly boggy conditions in clearings or on the outskirts of the an east Gondwana (Fig. 9). Th e palaeoclimatic maps of the forest. Th e bennettitaleans are considered to have inhabited Scotese (2000) have shown that these land masses were under lowlands to coastal regions. Th e ginkgoaleans are wetland lov- the infl uence of warm temperate conditions. Th erefore, it is ers. Th e members of araucarian conifers are interpreted to dwell expected to be fl oral similarities among the Indian subconti- in lowlands yet some species may extend into coastal margins. nent, Australia and Antarctica. Th e palaeogeographic maps Th e podocarps and taxaceans possibly occupied the upland of the Jurassic and Cretaceous periods have shown that the regions. Palaeoecological reconstruction of the Early Cretaceous east coast of India is closely associated with Western Australia Krishna-Godavari Basin fl ora is illustrated in Figure 6. and east Antarctica. Hence the comparison is here restricted to the fl oras of these regions. Th e poor dating of the Indian Mesozoic sediments is one PHYTOGEOGRAPHICAL CORRELATION of the signifi cant constraints to compare the Indian Early Cretaceous fl ora with the other Gondwana fl oras. Unlike the To a large extent type and distribution pattern of the land plants Early Cretaceous successions of Australia and Antarctica, there is constrained by its immediate physical environment (Spicer are no precise age determinations for the Early Cretaceous et al. 1994). Th is constraint has operated thorough out land successions of India (Fig. 10). Th e Indian Early Cretaceous plant evolution and has repeated morphologies under similar successions were assigned age ranging from Neocomian to

GEODIVERSITAS • 2018 • 40 (12) 273 Chinnappa C. H. & Rajanikanth A.

Aptian almost covering the entire Early Cretaceous period. cutchense Feistmantel. Th e fossil fl ora from adjoining intra- Palaeobotanical studies around the globe, however, indicated cratonic rift basins i.e., Pranhta-Godavari (PG) diff ers in the that except the appearance of angiosperms there are not many dominance of conifers. Except Ptilophyllum other bennetti- changes in compositions of fl ora during the intervals of the talean genera like Dictyozamites, Otozamites, Anomozamites, Early Cretaceous. Th is permits a broad scale comparison of and Pterophyllum are rare and Ginkgo is totally absent from the Indian Early Cretaceous fl ora with that of Australia and the PG fl ora. Among the other Early Cretaceous fl oras of Antarctica. India, the KG fl ora is closely comparable with the fl ora of Th e President Head fl ora, Antarctica, is distinctive, and Rajmahal (Banerji 2000). Both these fl oras are predomi- appears to be characteristic of a high latitude fl ora (Cantrill nated by bennettitalean foliages and share many taxa. Th e 2000). Bryophytes and hepatophytes are abundant and diverse, taxa common to both the fl oras are: Dictyozamites falcatus, a feature unique to the Cretaceous Antarctica (Drinnan & D. indicus Feistmantel, Otozamites imbricatus, Ptilophyllum Chambers 1986; Cantrill 1997a, b). Th e President Head fl ora distans (Feistmantel) Bose & Kasat, P. incisum, Ptilophyllum shares two species, Dictyozamites falcatus Medlicott & Banford acutifolium, P. cutchense, P. rarinervis (Feistmantel) Bose & and Pachypteris indica (Oldham & Morris) Bose & Roy and Kasat, P. tenerrimum, Williamsonia blandfordii Feistmantel, also eleven genera: Lophosoria Presl (= Gleichenia/Gleichenites W. indica Seward, W. kakadbhitensis Pandya & Sukh-Dev Goeppert), Sphenopteris, Pachypteris, Taeniopteris, Ptilophyl- and Cycadolepis indica Gupta, Brachyphyllum expansum lum, Dictyozamites, Cycadolepis, Araucarites, Pagiophyllum, (Sternberg) Seward, Pagiophyllum cf. marwarensis Bose & Conites and Elatocladus. However, at species level they diff er Sukh-Dev, Elatocladus confertus Seward & Sahni, E. plana, from the Indian taxa. Th e most similar generic and possibly E. jabalpurensis (Feistmantel) Sahni and E. tenerrimus (Feist- even specifi c comparison appears to be with the fl ora of the mantel) Sahni. Th e major diff erence between these fl oras Western Australia. Similar to the Early Cretaceous KG fl ora, is in presence and/or absence of pentoxylaleans, which are the Neocomian Western Australian fl ora is also dominated by common from the Rajmahal. bennettitaleans (McLoughlin 1996). Both these fl oras share several genera (Gleichenites [= Microphyllapteris], Cladophlebis, Sphenopteris, Th innfeldia, Taeniopteris, Ptilophyllum, Araucarites Acknowledgements [araucarian cone scale], Pityospermum [winged seed], Elato- Th e author Rajanikanth is thankful to the Prof. Sunil Bajpai, cladus) and some species like Ptilophyllum acutifolium and Director, Birbal Sahni Institute of Palaeosciences, Lucknow P. cutchense in common. Th e principle diff erence between and Chinnappa is thankful to management of Andhra Loyola the two fl oras is in the presence or absence of Dictyozamites. College, Vijayawada for their encouragement and support While it is absent from the Western Australian fl ora, it is more and Dr Van Konijnenburg-Van Cittert, J. H. A (emeritus common in the Krishna-Godavari fl ora. professor at the University of Leiden) and Dr Spicer, R. A., Th e phytogeographical distribution and diversity of the for their suggestions and for attention to the language of an Indian East Coast fl ora during the Early Cretaceous was initial draft of the paper. We thank Dr Maria Barbacka and infl uenced by drifting of the Indian subcontinent. During an anonymous reviewer for their valuable, constructive com- this period initiation and separation of East Coast from the ments and suggestions on the manuscript. 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Submitted on 20 July 2017; accepted on 29 January 2018; published on 21 June 2018.

278 GEODIVERSITAS • 2018 • 40 (12) Floristical and Palaeoecological implications of the Early Cretaceous sequences of Krishna-Godavari Basin (India)

APPENDIX 1 . — List of the macrofl ora reported from the Krishana-Godavri Basin.

PTERIDOPHYTES BENNETTITALEANS (continuation) Cladophlebis medlicottiana (Oldham) Pascoe, 1959 Ptilophyllum heterophylla Chinnappa, Rajanikanth & Rao, 2014 C. polypodioides Brongniart, 1849 P. rarinervis (Feistmantel) Bose & Kasat, 1972 Cladophlebis cf. longipennis Seward, 1925 P. tenerrimum (Feistmantel) Bose & Kasat, 1972 Cladophlebis sp. Ptilophyllum cf. distans (Feistmantel) Bose & Kasat, 1972 Gleichenia bosahii (Bose) Pant & Srivastava, 1977 Ptilophyllum cf. institacallum Bose, 1959 G. nordenskioeldii Heer, 1874 Ptilophyllum cf. amarjolense Bose, 1953 Marattiopsis macrocarpa (Oldham & Morris) Seward & Ptilophyllum cf. gladiatum Bose & Sukh-Dev, 1958 Sahni, 1920 Ptilophyllum cf. horridum Roy, 1963 Onychiopsis cf. psilotoides (Stopes & Web) Ward, 1905 Ptilophyllum cf. jabalpurense Jacob & Jacob, 1954 Todites indicus (Oldham & Morris) Bose & Sah, 1968 Williamsonia blandfordii Feistmantel, 1876 Sphenopteris specifi ca (Feistmantel) Roy, 1968 W. indica Seward, 1917 Sphenopteris sp. GINKGOALEANS GYMNOSPERMS Ginkgo crassipes (Feistmantel) Chinnappa, 2016 G. feistmantelii (Bose & Sukh-Dev) Chinnappa, 2016 PTERIDOSPERMS Ginkgo sp. A Th innfeldia vemavaramensis (Feistmantel) Chinnappa, Rajani- Ginkgo sp. kanth & Rao, 2015 Pachypteris indica (Oldham & Morris) Bose & Roy, 1968 CONIFERALEANS Araucarites cutchensis Feistmantel, 1877 CYCADALEANS A. fi brosa Sukh-Dev & Bose, 1972 Taeniopteris spatulata (McClelland) Bose & Banerji, 1981 A. macropteris Feistmantel, 1877 Taeniopteris sp. A. minutus Bose & Maheshwari, 1973 Brachyphyllum expansum (Sternberg) Seward, 1904 BENNETTITALEANS B. feistmantelii (Halle) Sahni, 1928 Anomozamites sp. B. regularis Borkar & Chiplonkar, 1973 Cycadolepis indica Gupta, 1954 B. rhombicum Feistmantel, 1879 Cycadolepis sp. B. sehoraensis Bose & Maheshwari, 1973 Dictyozamites falcatus Medlicott & Banford, 1879 Brachyphyllum sp. D. feistmantelii Bose & Bano, 1978 Conites sessilis Sahni, 1928 D. hallei Sahni & Rao, 1933 Conites sp. D. indicus Feistmantel, 1876 Elatocladus confertus Seward & Sahni, 1920 D. ommevaramensis Chinnappa, Rajanikanth & Rao, 2014 E. jabalpurensis (Feistmantel) Sahni, 1928 D. sahnii Gupta & Sharma, 1968 E. loyolii Chinnappa, Rajanikanth & Rao, 2014 Otozamites acutifolius Feistmantel, 1879 E. plana (Feistmantel) Seward, 1919 O. bengalensis Schimper, 1870 E. tenerrimus (Feistmantel) Sahni, 1928 O. exhislopi Bose, 1974 E. vemavaramensis Pandya Pandya, Srivastava & Sukh-Dev, 1990 O. gondwanensis Bose, 1974 Elatocladus sp. O. imbricatus Feistmantel, 1879 Harrisiophyllum hacketioides Pant, Srivastava & Pant, 1983 O. vemavaramensis Bose & Jain, 1967 Pagiophyllum feistmantelii Halle, 1913 Otozamites sp. P. gollapallensis Pandya & Sukh-Dev, 1990 Pterophyllum footeanum Feistmantel, 1879 Pagiophyllum cf. grantii Bose & Banerji, 1984 P. incisum Sahni & Rao, 1933 Pagiophyllum cf. marwarensis Bose & Sukh-Dev, 1972 P. kingianum Feistmantel, 1877 P. ommevaramensis Chinnappa, Rajanikanth & Rao, 2014 Pterophyllum sp. Pagiophyllum sp. Ptilophyllum acutifolium (Morris) Bose & Kasat, 1972 Pityospermum godavarianum Chinnappa, Rajanikanth & P. cutchense (Morris) Bose & Kasat, 1972 Rao, 2015 P. deodikarii Mahabale & Satyanarayana, 1979 Torreyites constricta (Feistmantel) Seward & Sahni, 1920

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APPENDIX 2 . — List of the microfl ora reported from the Krishana-Godavri Basin.

BRYOPHYTES Foraminisporis dailyi Dettmann, 1963 PTERIDOPHYTES (CONTINUATION) F. wonthaggiensis Dettmann, 1963 Impardecispora sp. Coronatispora sp. Ischyosporites punctatus Cookson & Dettmann, 1958 Aequitriradites spinulosus Cookson & Dettmann, 1961 Ischyosporites sp. Coptospora cauveriana Venkatachala, 1973 Klukisporites foveolatus Pocock, 1965 Cooksonites variabilis Pocock, 1962 K. scaberis Dettmann, 1963 Staplinisporites caminus Pocock, 1962 Laevigataletes sp. Laevigatazonaletes sp. PTERIDOPHYTES Laevigatimonoletes sp. Alsophyllidites grandis Sah & Jain, 1965 Laevigatisporites sp. Appendicisporites sellingii Pocock, 1964 Leptolepidites major Couper, 1953 Baculatisporites comaumensis Potonié, 1953 L. verrucatus Couper, 1953 B. baculatus Krutzsch, 1959 Leptolepidites sp. Biretisporites potoniae Delcourt & Sprumont, 1955 Liratosporites sp. Ceratosporites equalis Cookson & Dettmann, 1958 Lycopodiumsporites crassimacerius Hedlund, 1966 Cicatricosisporites australiensis Potonié, 1956 L. eminulus Dettmann, 1963 C. hughesii Dettmann, 1963 L. regulatus Semenova, 1970 C. lodbrokiae Dettmann, 1963 L. reticulum Venkatachala & Kar, 1968 Cicatricosisporites sp. L. reticulumsporites Dettmann, 1963 Conbaculatisporites densibaculatus Sharma, Jain & Venka- Lycopodiumsporites sp. tachala, 1977 Matonisporites sp. Concavissimisporites punctatus Pocock, 1964 Murospora fl orida Pocock, 1961 C. variverrucatus Brenner, 1963 Neoraistrickia truncatus Potonié, 1956 Concavissimisporites sp. Ornamentifera granulosa Sharma, Jain & Venkatachala, 1977 Contignisporites cooksoniae Dettmann, 1963 Ornamentifera sp. C. glebulentus Dettmann, 1963 Osmundacidites wellmanii Couper, 1953 C. multimuratus Dettmann, 1963 Plicifera senonicus (Ross) Bolkhovitina, 1966 Contignisporites sp. Polycingulatisporites reduncus Playford & Dettmann, 1965 Crybelosporites punctatus Dettmann, 1963 Reticulatazonalesporites sp. C. striatus Dettmann, 1963 Reticulatisporites sp. C. stylosus Dettmann, 1963 Retitriletes austroclavatidites (Cookson) Krutzsch, 1963 Crybelosporites sp. R. circolumenus Backhouse, 1978 Cyatheacidites tectifera Archangelsky & Gamerro, 1965 R. eminulus Srivastava, 1975 Cyathidites asper Dettmann, 1963 Sestrosporites pseudoalveolatus Dettmann, 1963 C. australis Couper, 1953 Striatella balmei Filatoff & Price, 1988 C. cutchensis Singh, Srivastava & Roy, 1964 Taurocusporites segmentatus Stover, 1962 C. jurassicus Kar & Sah, 1970 Th ymospora sp. C. minor Couper, 1953 Todisporites minor Couper, 1953 C. pseudopunctatus Singh, Srivastava & Roy, 1964 Triletes verrucosus Faddeeva, 1965 C. punctatus Delcourt, Dettmann & Hughes, 1963 Triletes sp. C. rajmahalensis Sah & Jain, 1964 C. trilobatus Sah & Jain, 1964 Cyathidites sp. Deltoidospora diaphana Wilson & Webster, 1946 D. juncta Singh, 1964 Densoisporites microregulatus Brenner, 1963 D. velatus Dettmann, 1963 Dictyophyllidites harrisii Couper, 1958 Gleichinidites circinidites (Cookson) Dettmann, 1963 G. senonicus Dettmann, 1963 Impardecispora purverulenta (Verbitzkaya) Venkatachala, Kar & Raza, 1969 I. trireticulosa Venkatachala, Kar & Raza, 1969 I. tribotrys (Dettmann) Venkatachala, Kar & Raza, 1969

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APPENDIX 2 (continuation). — List of the microfl ora reported from the Krishana-Godavri Basin.

GYMNOSPERMS ANGIOSPERMS Abietineaepollenites ellipticus Kar & Sah, 1970 Asteropollis asteroides Hedlund & Norris, 1968 A. robustus Kar & Sah, 1970 Clavatipollenites hughesii Couper, 1958 Alisporites grandis Dettmann, 1963 Clavatricolporites leticiae Leidelmeyer, 1966 Alisporites sp. Liliacidites reticulatus Doyle, 1975 Araucariacites australis (Cookson, 1947) Cooper, 1953 Polybrevicolpites sp. A. indicus Sukh-Dev, 1961 Polycolpites sp. Callialasporites dampieri Sukh Dev, 1961 Racemonocolpites facilis González-Guzmán. 1967 C. monoalasporus Sukh Dev, 1961 R. ramonus González-Guzmán. 1967 C. segmentatus Srivastava, 1963 Rousea georgensis Dettmann, 1973 C. triletus Singh, Srivastava et Roy, 1964 Spinizonocolpites echinatus Muller, 1968 C. trilobatus Sukh Dev, 1961 Tricolpites georgensis (Brenner) Dettmann, 1973 Callialasporites sp. Tubulifl oridites lilleie (Couper) Farabee & Canright, 1986 Cedripites cretaceus Pocock, 1962 Turonipollis helmegii Van Ameron, 1975 C. nudis Kar & Sah, 1970 Classopollis classoides Fensome, 1983 INCERTAE SEDIS C. torosus Burger, 1965 Apiculatasporites sp. Entylissa sp. Apiculatimonoletes sp. Florinites sp. Apicultaletes sp. Ginkgocycadophytus srivastavae Kar & Sah, 1970 Bhujiasporites sp. G. nitidus Venkatachala, 1969 Complexiopollis complicatus Góczán, 1964 Granuloperculatipollis fl avatus Kar, 1970 Coniatisporites telata Singh, 1972 G. subcircularis Kar & Sah, 1970 Dictyotosporites complex Cookson & Dettmann, 1958 G. triletus Kar & Sah, 1970 Dictyotosporites speciosus Cookson & Dettmann 1958 Indusiisporites microsaccatus Kar & Sah, 1970 Erdtmannipollis sp. Laricoidites indicus Singh, Srivastava & Roy, 1964 Odontochitina operculata Defl andre & Cookson, 1955 Microcachrydites antarcticus Couper, 1953 Periplecosporites sp. Pityosporites sp. Ramanujamiaspora reticulata Ramanujam, 1957 Platysaccus sp. Regulatisporites sp. Podocarpidites alareticulosus Sah & Jain, 1965 Setosisporites sp. P. cristiexinus Sah & Jain, 1965 Singhipollis rudis Kar & Sah, 1970 P. ellipticus Cookson, 1947 Singhipollis triletus Singh, Srivastava & Roy, 1964 P. grandis Sah, 1965 Striatotuberculatisporites sp. P. multisimus (Bolkhovitina) Pocock. Venkatachala, Kar & Raza, 1969 P. rarus Singh, Srivastava & Roy, 1964 P. typicus Sah, 1965 Podocarpidites sp. Podosporites raoi Singh, Srivastava & Roy, 1964 P. tripakshii Rao, 1943 Podosporites sp.

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