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The Pailleoboiailisl 55(2006). 29-40 0031-0174/2006 $2.00

Non-geniculate coralline from the Uttatur Group (Early ), south India

P.K. MISRAl, A.K.]AUHRF, S.K. SINGHl, S. KISHOREl AND A. RA]ANIKANTH3

IDepartment of Botany, University of Lucknow, Lucknow 226007, India. l Department of Geology, University of Lucknow, Lucknow 226007, India. J Birbal Sahni Institute of Palaeobotany. 53 University Road, Lucknow 226 007, India

(Received 3 August. 2005; revised version accepted 8 May, 2006)

ABSTRACT

Misra PK, Jauhri AK, Singh SK, Kishore S & Rajanikanth A 2006. Non-geniculate from the Uttatur Group (Early Cretaceous), south India. The Palaeobotanist 55 (1-3): 29-40.

The present paper records 8 species of non-geniculate coralline algae from the Early Cretaceous of the Uttatur Group, Tiruchirapalli District, Tamil Nadu, south India. Of these, six species are distributed among three genera of the family: three species belong to Amphiroa. one species is assigned to Lithothamnion and two species are placed with Lithophyllum. Two species are referable to Sporolithon ofSporolithaceae. Taxonomic differentiation is based on growth form, cell fusions and nature of conceptacle pore.

The species of Amphiroa are recorded for the first time from the study area while Amphiroa kaskaella is recorded for the first time from India. Amphiroafo/iacea and A. guatemalense are documented for the first time from the Uttatur Group (Cretaceous) of the Cauvery Basin. The algal association suggests depositional environments ranging from shallower to deeper parts of sea. The abundance of coralline algae indicates reefal environment.

Key-words-Calcareous algae, Rhodophyceae, Kallakudi Limestone, Uttatur Group, Early Cretaceous.

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INTRODUCTION Significant work on stratigraphy, micropalaeontology, depositional environment and THE present paper deals with the morphotaxonomic tectonic evolution ofthe Cauvery Basin has been carried .. description ofnon-geniculate coralline algae from out by Rama Rao (1956), BaneJji (1972), Chiplonkar the Lower Cretaceous Kallakudi Limestone (Uttatur and Tapaswi (1975), Ramanathan (1979), Sundaram Group), Cauvery Basin, south India exposed in the and Rao (1986), Jafar and Rai (1989), Govindan et areas around Olaipadi Mine near Govindarajapatnam al. (1998), Madhavaraju and Ramasamy (1999,2002), ofTiruchirapalli District, Tamil Nadu (Fig. 1). The Slilldaram et al. (2001), Madhavaraju et al. (2002, Cretaceous sediments are well exposed in isolated areas 2004) and Nagendra et al. (2002). The Uttatur Group (Pondicherry, Vridhachalam, Ariyalur, Tanjore and is important from the palaeontological point ofview as Sivaganga) ofthe Cauvery Basin (BaneJji, 1972). The it is characterised by rich assemblages ofanimal and Cauvery Basin is the southemmost basin along the plant fossils. Calcareous algae reported earlier from the eastern margin ofPeninsular India and is nearly 400 Coralline Limestone of the Uttatur Group were km long and 130 km wide. It is located between represented by Cayeuxia sp., C. fruticulosa, Latitude 12° and 9° 30' North and Longitude 78° and Acicularia antiqua, Neomeris occidentalis, 80° 30' East. The Cretaceous rocks ofthe Timchirapalli Neomeris sp., Halimeda sp., Solenoporajurassica, area were first recognised and described by Blanford S coromandelensis, S sahnii and Lithothamnion sp. (1862). He divided Cretaceous succession of the (Narayan Rao, 1944, 1946; Rama Rao & Prasanna Cauvery Basin into three groups: Uttatur Group, Kumar, 1932; Rama Rao & Gowda, 1954; Trichinopoly Group and Ariyalur Group. The present Rajanikanth, 1988, 1992; Gowda, 1978). Mi~raet al. calcareous algal assemblage is reported from the (2004) reported eight species ofcalcareous algae fi'om Kallakudi Limestone ofthe Uttatur Group. This group the Kallakudi mines ofthe Uttatur Group. is named after the village Uttatur. It extends over a 70 METHODOLOGY km long, 4-6 krn wide area and beds dip 10° due east. It is divisible into three formations, viz., Grey Shale, The material was collected from the Kallakudi Kallakudi Limestone and Karai ShalelMaruvattur Clay Limestone of the Olaipadi Mine (near (Govindan et al., 1998; Fig. 2). ~vindarajapatnam),TiruchirapaUiDistl;ct, TamilNadu, The outcrops ofthe Kallakudi Limestone (Aptian­ and the calcareous algae were studied in random thin Albian) occur along the western margin of the basin sections; about 130 slides were prepared by Logitech from the Olaipadi Mine exposed near Machine. Thin sections used in studying the fossil Govindarajapatnam (Fig. 3), Asur-Kallal in the north, coraBines are oftwo types: one parallel to the direction Kudikkadu-Varagupadi in the middle to Tirupattur in of filament growth and perpendicular to the thallus the southwest and Kallakudi in the southeast. It is a surface and the other perpendicular to the direction of conspicuously pink, hard, compact limestone with filament growth for measuring cell diameters. The stromatactoid structures, large irregular cavemous voids, taxonomic differentiation ofthe coralline algae in the vugs fille9 with coarse, crystalline quartz and calcite at present work, however, has been made on the basis of places. The fossils include , corals, bryozoa, certain diagnostic anatomical features oflivingcoraUines, echinoids, ostracods, bivalves and foraminifera which are preserved even in the fossil material. A brief (Govindan et al., 1998)~ summary ofthese features and the related telminology MISRA £T AL.-NON-GENICULATE CORALLINE ALGAE FROM THE UTTATUR GROUP. SOUTH INDIA 31

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Fig. I-Geological map ofTiruchirapalli area (after Govindan et at., 1998). 32 THE PALAEOBOTANIST

SYS SER STAGE FORMATION GROUP Growth form warty to fruticose, peripheral region TEM IES is well developed with distinct zonation. Filaments radially organized inside the protuberances, cell fusions KALLAMEDU Sandstone Maastrichtian conspicuous. Tetralbisporangial conceptacles KALLANKURICHCHI multiporate.

ARIYALUR Lithothamnion sp. GROUP

Campanian SILLAKUDI Qj Q. (PI. 2.5, 6) m Q. ~ ~ 0 Description-Growth form encrusting, thallus W Santonian GARUDAMANGALAM TRICHINOPOLY U organisation monomerous. Core filaments non-coaxial, Coniacian GROUP « PARAVAY core portion usually 250 Ilm thick. Cell fusions present. I- Turonian MMUVAnUR Cells 15-25 Ilm in length and 10-14 Ilm in width. w Clay 0:: Cenomanian Peripheral cells 8-12 Ilm in length and 10-12 Ilm in U KAMI width. Asexual conceptacles present, cavity 110-140 Shale UTIATUR GROUP Ilm in height and 500-600 Ilm in width. Sample no.-VL/2. Albian s\· Qj ~ ~ Slide no.-56. 0 ...J Locality--Olaipadi Mine, Govindarajapatnam.

UPPER Occurrence--Uttatur Group. Pre Albian TERANI GONDWANA GROUP Discussion-The present specimen resembles ARCHAEAN EASTERN GHAT COMPLEX Lithothamnion cf. L. lacroixi Johnson and Kaska (1965) reported from the lower Miocene of the Fig. 2-Cretaceous lithostratigraphy (outcrops) Cauvery Guatemala in growth form, cell dimensions ofcore and Basin, India (after Govindan el aI., 1998). peripheral filaments ofthe thallus. In addition, the tetra! based on Rasser and Piller (1999), Bassi (1997, 1998), bisporangial conceptacles also show similarity in shape, Braga et al. (1993), etc. were presented in Misra et size and the nature ofdevelopment ofthe conceptacles. al. (200 I). The studied material is preserved at the Algology Subfamily-LITHOPHYLLOIDEAE Setchell, 1943 Laboratory, Department of Botany, University of Lucknow, Lucknow. Genus-LTTHOPHYLLUM Philippi, 1837

SYSTEMATICS Growth form crustose to fruticose, composed entirely of protuberances. Crustose portion of plants Division-RHODOPHYTA Wettstein, 1901 and lamellae dorsiventral and dimerous or monomerous, or both in the same plant. Cell fusions absent. Tetral Class-RHODOPHYCEAE Rabenhorst, 1863 bisporangial conceptacles uniporate and clearly delimited. Order-CORALLINALES Silva & Johansen, 1986 Lithophyllum sp. 1 Family~ORALLINACEAELamouroux, 1812 (PI. 2.7, 8) Subfamily- Bizzozero, 1885

Genus-LITHOTHAMNION Heydrich, 1897 Description-Growth form encrusting and MISRA ET AL.-NON-GENICULATE CORALLINE ALGAE FROM THE UTTATUR GROUP. SOUTH INDIA 33

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Fig. 3- The litho-biostratigraphic representation of the Olaipadi mines (Govindarajapatnam) showing position of the fossil­ yielding samples (indicated by vertical line), strongly branching, thallus organisation monomerous, Description-Growth form encrusting, thallus Core filaments coaxial, cells ofcore filaments two types­ organisation monomerous. Core filaments coaxial, cells long cells 30-40 ~min length and 8-12 ~min width; of core filaments 15-18 ~min length and 10-12 ~min short cells 18-24 ~min length and 8-12 ~min width. width. The marginal peripheral filaments present, ranging The marginal peripheral filaments present, ranging from from 8-1 0 ~m in width, cells not measurable. Tetra/ 250-300 ~min width. Cells of peripheral filament 12­ bisporangial conceptacle uniporate, showing cells 180­ 16 ~min length and 8-12 ~min width. Tetralbisporangial 200 ~min length and 300-350 ~min width. conceptacle uniporate, with cells 80-1 00 ~min length Sample no.-VL/2. and 200-400 ~min width. Slide no.-6. Sample no.-VL/3-4. Locality---Olaipadi Mine, Govindarajapatnam. Slide no.-33. Occurrence-Uttatur Group. Locality-Olaipadi Mine, Govindarajapatnam. Discussion-Thallus organization and uniporate Occurrence-Uttatur Group. conceptacle morphology suggestthat these specimens Discussion-Thallus organization, cell size ofcore are broadly similar to Lithophyllum. The present filaments (protuberance), peripheral filaments and specimens are not comparable with any known species uniporate nature ofthe conceptacles ofthe specimens of the genus Lithophyllum. indicate their resemblance to Lithophyllum altemicellum Johnson (1964). Johnson reported this Genus-AMPHIROA Lamouroux, 1812 species from the Miocene of Guam. However, conceptacles are larger in the present specimen and Previously, Amphiroa was assigned to differs from L. altemicellum. geniculate coralline algae. Recently, Bailey (1999) transferred this genus into non-geniculate group on the Lithophyllum sp. 2 basis 18S rRNA gene sequencing and indicated its relationship with Titanoderma of subfamily (PI. 2.9) Lithophylloideae. Hence, in the present study we have 34 THE PALAEOBOTANIST followed the Bailey's (1999) classification placing 1954lshijima, p. 53, pI. 37, fig. 8. Amphiroa with subfamily Lithophylloideae ofnon­ 1957 Johnson, p. 238, pI. 37, fig. 2. geniculate coralline algae. This genus has segments that 2003 Kundal and Dharashivkar, p. 249, pI. 1, fig. 4. are cylindrical to flattened in shape. The medullary tissue Description-lntergenicula cylindrical, flattened is characterized by one or more rows of long cells with atapering end. The medullary rows ofcells arched. alternating with a single row of short cells and is Medulla consists ofthree rows oflong cells alternating surrounded by a distinctly layered cortical tissue. with one row ofshort cells; medullary filaments 1.1mm Conceptacles are marginal and lateral in position. long and up to 0.25 mm broad; long cells ofmedullary filaments 40-50 ~min length and 8-12 ~min width, Amphiroa guatemalense Johnson and Kaska, short cells 20-35 ~mlong and 8-12 ~mbroad. 1965 Sample no.-VL/ 1. Slide no.-33. (PI. 1.1, 2) Locality---Olaipadi Mine, Govindarajapatnam. Occurrence-Uttatur Group. 1965 Johnson and Kaska, p. 52, pI. 24, figs 1,2 Discussion-This specimen seems to be and pI. 25, fig. 1. comparable with Amphiroafoliacea in its medullary Description-Segments broadly cylindrical, 1.5 filaments showing alternation ofthree rows oflong cells mm long and up to 1.1 mm broad; medullary filaments with one row ofshort cells in the thallus. showing one row oflong cells alternating with one row ofshort cells; long cells 25-35 ~m in length and 8-12 Amphiroa kaskaella Johnson and Kaska, 1965 ~min width; short cells 15-25 ~m long and 8-12 ~m broad; peripheral filaments about 50 ~m in diameter (PI. 1.5-8) and cells 7-10 ~min length and 7-10 ~min width. Sample no.-VL/I. 1965 Johnson and Kaska, p. 53, pI. 25, figs 2, 3. Slide no.-27. Description-Segments long and wide, 1.0 mm Locality---Olaipadi Mine, Govindarajapatnam. long and up to 0.5 mm broad, medullary filaments show Occurrence-Uttatur Group. two rows oflong cells alternating with one or two rows Discussion-The present specimen is comparable ofshort cells; long cells 60-80 ~mlong and I0-12 ~m with Amphiroa guatemalense Johnson and Kaska in broad; short cells 20-25 ~m long and 10-12 ~mbroad. shape and size ofshort and long cells ofmedullary region Marginal peripheral filaments thin, about 60 ~mwide; of the thallus. Johnson and Kaska (1965) reported cells 10-12 ~m in length and 8-10 ~min width. Amphiroaguatemalense from the Palaeocene of Sample no.-VL/3, 4. Guatemala. Slide no.-48, 50, 71. Locality---Olaipadi Mine, Govindarajapatnam. Amphiroafoliacea Lamouroux, 1812 Occurrence-UttaturGroup. Discussion-The present specimen is referable to (PI. 1.3,4) Amphiroa kaskaella Johnson and Kaska reported

PLATE 1 "/ I. Amphiroa guatemalense. x 50. 5. Amphiroa kaskaella. x 50. 2. Amphiroa guatemalense. x 130. 6. Amphiroa kaskaella. x 130. 3. Amphiroafoliacea. x 50. 7. Amphiroa kaskaella. x 130. 4. Amphiroafoliacea. x 130. 8. Amphiroa kaskaella. x 50. MISRA ET AL.-NON-GENICULATE CORALLINE ALGAE FROM THE UTTATUR GROUP, SOUTH INDIA 35

PLATE 1 36 THE PALAEOBOTANIST from the Palaeocene ofGuatemala (Jolmson & Kaska Description-Growth fonn encrusting to crustose 1965) in having regular alternation oflong and short and lobate, 4.5 mm long and 3.0 mm in width. Thallus cells in the medullary region. Besides this character, the organisation monomerOliS.Core filaments non-coaxial. cells ofmedullary region also show similarity in shape, Cells ofcore filaments 18-20 /Jm in length and 8- J 2 size and their arrangement in the thallus. /Jmin width. The peripheral region ofencrusting portion restricted to dorsal part ofthe thallus with cells 15-20 Family-SPOROLITHACEAE Verheij, 1993 /Jmin length and 8-10 /Jm inwidth. Cell fusions present. Tetrasporangial conceptacles arranged in sori. Individual Genus-SPOROLITHON Heydrich, 1897 sporangial compartments rectangular, elliptical or ovoid in longitudinal section and circular in transverse section; The genus is characterized by epithallial cells with 60-80 Ilm in height and 35-50 Ilm in width. Sori usually flattened and flared cells and tetra/bisporangial arise from a layer of elongated cells. Filaments conceptacles separated by interspersed calci fied (paraphyses) are interspersed between the sporangial filaments (paraphyses) (Woelkerling, 1988). According compartments. to Moussavian and Kuss (1990), Sporolithon is the Sample no.-VLlI. correct generic name for the corallines earlier included Slide no.-34. in Archaeolithothamnium Rothpletz, 1891, since the Locality-Olaipadi Mine, Govindarajapatnam. latter name was not validly pUblished. Verheij (1993) Occurrence-Uttatur Group. proposed a new family, Sporolithaceae to separate Discussion-The present specimen is comparable Sporolithon from the rest ofthe corallines on account with Sporolithon lugeoni (Pfender) Moussavian and ofsimultaneous cruciate cleaving ofits tetrasporangia. Kuss recorded by Misra and Kumar (1988) and Ghosh Tetrasporangial chambers are surrounded by calcified and Maithy (1996) in shape and size ofsporangia and paraphyses (filaments). Townsend et al. (1995), in peripheral cells. Misra and Kumar (1988) reported this order to avoid the confusing terminology ofreproductive form from the Cretaceous ofVaragur, Tiruchirapalli structures, regarded the family Sporolithaceae as having District, Tamil Nadu. Ghosh and Maithy (1999) also tetrasporangia which show cruciately arranged spores reported possibly a very similar form from the within calcified sporangial compartments (Aguirre & Maastrichtian of the Kallankurichchi Formation, Braga, 1998). AriyalurGroup, Tamil Nadu.

Sporolithon lugeoni (Pfender) MOllssavian and Sporolithon sp. Kuss, 1990 (PI. 2.4) (PI. 2.1- 3)

1926 Pfender, p. 324, pI. 9, 13. Description-Growth form encrusting with 1988 Misra and Kumar, p. 46, pI. 4, figs 7, 9, 11. protuberances, thickness ofencrusting thalli up to 1.7 1990 Moussavian and Kuss, p. 929-942. mm. Thallus organization monomerous. Core filaments 1996 Ghosh and Maithy, p. 68, pI. 1, figs 1-4. non-coaxial. Cells regular, multilayered. Medullary core 1999 Ghosh and Maithy, p. 37, pI. 1, fig. a. filaments quite regular. Cells 18-221lm long and 8-12

PLATE 2 -,\..

1. Sporolithon lugeoni. x SO. 6. Lithothamnion sp. x 130. 2. Spo~olithonlugeoni. x SO. 7. Lithophyllum sp. I. x SO. 3. Sporolithon lugeoni. x SO. 8. Lithophyllum sp. I. x 130. 4. Sporolithon sp. x 130. 9. Lithophyllum sp. 2. x SO. 5. Lithothamnion sp. x 130. MISRA ET AL-NON-GENICULATE CORALLINE ALGAE FROM THE UTTATUR GROUP, SOUTH INDIA 37

PLATE 2 38 THE PALAEOBOTANIST

~mwide. Peripheral filament marginal with cells 12-18 (Lithophylloideae) prefers wann, shallow waters at ~min length and 8-1 0 ~min width. Sporangia ovoid, depths < 20 m (Braga & Martin, 1988). Ecological 40-60 ~m long and 30-40 ~m in width. Filaments data indicate that Amphiroa generally lives in water (paraphyses) interspersed between the sporangial less than 30 m (Cloud, 1952) but is common at depths compartments. between 20 and 25 m (Johnson, 1957). Though poorly Sample no.-VL/4. known from the Albian sediments ofthe world, the Slide no.-50. species of Amphiroa in the present assemblage are Locality-----Olaipadi Mine, Govindarajapatnam. dominant. Occurrence-Uttatur Group. The present investigation suggests that the Kallakudi Discussion-The present specimen differs from Limestone deposited on a carbonate platform in which Sporolithon lugeoni in shape and arrangement oftetral a number of algal biofacies could possibly be bisporangial compartments and cell size. However, distinguished and represent depositional environments orientation ofthallus ofpresent specimen is similar. ranging from shallower « 20 m) to deeper (> 30 m) parts of sea. DISCUSSION Acknowledgements-Thanks are due to the Head, Botany The present paper records eight species of the and Geology departments, Lucknow University, Lucknowand the Director, Birbal Sahni Institute of Palaeobotany, families Corallinaceae and Sporolithaceae of class Lucknow for facilities and encouragement. Dr M. Rasser Rhodophyceae from the LowerCretaceous succession (Vienna, Austria) and Prof loan Bucur (Babes-Bolyai of the Uttatur Group. These taxa assigned to four University, Romania) are gratefully thanked for reviewing genera are Amphiroa kaskaella, A. foliacea, A. the original manuscript and offering helpfid commentsfor its guatemalense, Sporolithon lugeoni, Sporolithon sp., improvement. The financial assistance from the Department of Science and Technology, New Delhi through Project No. Lithothamnion sp., Lithophyllum sp. 1 and SR/S4/ES-12/2002 for carrying out the present work is Lithophyllum sp. 2. The taxonomic features such as thankfully acknowledged. growth form, cell fusions and nature ofconceptacle pore have been used in discriminating between different REFERENCES genera. The species of Amphiroa documented here are recorded for the first time from the study area. Of Adey WH, Townsend RA & Boykins WT 1982. The crustose these, the presence ofAmphiroa kaskaella is recorded coralline algae (Rhodophyta: Corallinaceae) of the Hawaiian for the first time in the Lower Cretaceous successions Islands. Smithon. Contribution Marine Science 15 : 1-74. Aguirre J & Braga JC \998. Redescription of Lemoine's (1939) ofIndia. Types of coralline algal species from Algeria. Palaeontology Studies on the variable morphology ofthe present­ 41 :489-507. day coralline algae and their preferences to different Bailey JC 1999. Phylogenetic positions of Lithophyllwn climates and water depths have provided useful data incrustans and Titanoderma pustulatum (Corallinaceae, for palaeoenvironmental inferences. The abundance of Rhodophyta) based on 18S rRNA gene sequence analyses with a revised classification ofLithophylloideae. Phycologia coralline algae in the Kallakudi Limestone indicates 38/3: 208-216. reefal environment. Reefs and adjacent facies are Banerji RK 1972. Conflicting stratigraphical ranges observed characterised by corals and different coralline algal in the Miocene planktonic foraminifera from the Cauvery fonns which carry out cementing and framework­ Basin, South India. Proceedings of the II Indian Colloquium binding function in this environment. hnport..ant coralline on Micropalaeontology and Stratigraphy: 40-49. Bassi D 1997. Vegetative anatomy and palaeoecology of genera in the present assemblage are Lithothamnion, Polystrata alba Pfender, 1968 (Cryptonemiales, Sporollthon and Lithophyllum. Presence of Peyssonneliaceae) from the Upper Eocene of northern Italy. Sporolithon and Lithothamnion is indicative of depths Review Paleobiol Geneve 16: 309-320. more than 20 m (Adey et at., 1982; Minnery, 1990; Bassi D 1998. Coralline red algae (Corallinales, Rhodophyta) Perrin et al., 1995), while Lithophyllum from the upper Eocene Calcare di Nago (Lake Garda, MISRA ET AL.-NON-GENICULATE CORALLINE ALGAE FROM THE UTTATUR GROUP, SOUTH INDIA 39

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