Possible fossil Sporae dispersae of Hepaticae and Anthocerotales in the fossil record

Divya Darshan Pant & Rita Singh

Pant, Divya Darshan & Singh. Rita 1991. Possible fossil Sporae dispersaeof Hepaticae and Anthocerotales in the fossil record Palaeobotanist 39 (1) : 20- 36

The paper attemr:s to find the characters which can distinguish the spores of Hepaticae and Anthocerotales from iso- or micro-~poresof pteridophytes and pollen grains of gymnosperms and angiosperms for the indentification of possible fossil Sporae dispersae of reported from different geological horizons. The problem was approached by looking for characters in fossil spores which are (i) exclusively bryophytic, (ii) prevailingly bryophytic, and (iii) closely matching characters of in situ spores of fossil bryophytes in a first hand comparison. Our account also takes into consideration the Sporae dispersae which were described or suspected by previous workers as those of bryophytes but which could equally well belong to pteridophytes.

Key-wor-ds- Palynology, Adherent tetrads, Pseudocolpus, Hepaticae, Anthocerotales.

Divya Darshan Pant & Rita Singh, Department of Botany, University of Allahabad, Allahabad 211 002, India.

'~n:hT

~ ~ 1f'41~(\hfl~~ ~ ~ ~"1f(f *

~

~ ~ ~ ~ ~rrpif nrmu-trJf if ;j.f Fi'1I~d"l"t"1"t~ Gltif

~ <;l1414iI$t)~1~3iT ~ ~ ~?:~ (i)~'~ ~~, ~~, Ilfuct'fu;it (ii) a'!T(iii)

~itu-'l'nT3iTit 'll1~jqo;l$tl41'liT qfUfd' fu;m % 3f'lfiU ~ G!f'fd' fu;m %<9m it t{\il4iI~t)41

AFTER the discovery of hepatic thalli in British stalked sporangia had arisen from a common thallus. Carboniferous rocks (Walton, 1925, 1928), the time Dixon (1927) has referred this fossil to Andreaea. As range of this group was further extended up to the the spores in the spore sacs are not accompanied by Devonian by Hueber (1961) who reported elaters, the attribution of this fossil to Hepaticae Hepaticites devonicus from Onteroa Red Beds of becomes doubtful (see Krassilov & Schuster, 1984). North America which is now renamed as Schuster (1966) has accordingly referred the Pallavicinites devonicus (Hueb.) Schuster (see to an order of its own, the Sporogonitales. Schuster, 1966). There are also a few problematic Another fossil of the same kind called fossils which have been reported from the Lower Lyonophyton was described by Remy and Remy Devonian. One of these was referred by Andrews (1980) from. Rhynie Chert bed as haVing as (1960) ro Sporogonites Halle and it shows a thallus­ "independent and possibly autotrophic gametophyte like carbonaceous mass below a number of parallel which is of approximately equal size and shape with subadjacent, unbranched erect stalks. If unbroken the sporophytes from the same beds" This fossil is they terminate in ovoid spore sacs containing spores described as haVing a cup-shaped gametophore with ca 20-25 J.Lm in diameter. No details of the spore lobed margins. Remy and Remy (1980) have characters are available. Andrews believed that the compared it with Horneophyton on the basis of itS 20 PANT & SINGH-FOSSIL SPORAE DISPERSAE OF HEPATICAE AND ANTHOCEROTAlES 21 anatomy as well as the morphology of fertile parts the environment was transitional between that of and they have further suggested that Lyonophyton water and terrestrial and the shows a rhyniensis seems to be "the gametophyte of a plant combination of characters of both the habitats. He representing an evolutionary stage previous to the further suggests that Protosalvinia was probably separation of land plants into bryophytes and higher "caught in act" of evolving into a plant which was land plants." Till recently all other Devonian forms becoming adapted for living in a desiccating which had a central strand were included among the envi ronment. pteridophytes. In fact, one of these called, Rhynia Parka decipiens is another Lower Devonian major was regarded as one of the best known plant haVing a flat thallus-like body whose surface Devonian pteridophytes but a recent renaming of was covered by flattened disc-shaped structures with this form as Aglaophyton by Edwards (1986) is based numerous cutinized spores ca 28-34 /-.Lmin diameter. on his claim that its central strand lacks secondary The thallus is parenchymatous in nature (Lang, wall thickenings in the longitudinal cell walls. 1937). The affinites of Parka too are uncertain, it has According to him it consists of cells having an alga-like thalloid body and cutinized spores like uniformly thick walls and he raises doubts about its those of land plants (see Walton, 1953). being a vascular plant suggesting that it had a The third uncertain fossil Steganotheca shows a bryophytic level of organisation (Taylor, 1988). plant compression which is comparable with However, the observations of Edwards, on this form, Cooksonia in having terminal sporangia on which is otherwise quite like Cooksonia and Rhynia dichotomously branched axes but their central gwynne vaughanii need confirmation before they strands lack tracheids so that it could be compared can be accepted particularly when they are based on with bryophytes instead of pteridophytes. the negative evidence of thickenings. The above reports of megafossils beginning Beside the above two forms, there are some with Lower Devonian make it at once likely that other uncertain Devonian -like genera, some of the dispersed miospores described by called Protosalvinia Dawson, Parka decipiens Lang different authors from the beds of different and Steganotheca. Out of these, Protosalvinia is geological ages beginning with strata of this period reported from the Upper Devonian Black Shales of could even belong to bryophytes. Indeed, after east-central U.S.A. and its detached sporiferous Walton discovered undoubted structurally preserved organs usually show once forked branches which are bryophytes in the Carboniferous, he suggested to approximately 2-5 mm in width and appear to have Knox (1939) that she should look for spores of been borne on a thalloid plant body. The forks of bryophytes among fossil Sporae dispersae. However, the thallus show sporocarp-Iike structures, each she only mentioned some fossil spores (see Knox, containing a single tetrad of 4 large spores (? 1939; figs 41-51) which were broadly comparable megaspores) ca 200 /-.Lmin the cavity of inner tissues. with those of some liVing bryophytes like Anthoceros The affinities of Protosalvinia are uncertain. It has chilensis, hibernica, Anthoceros punctatus, been assigned to the algae by White and Riccia crystallina, R. beyrichiana, R. jluitans, Stadnichenko (1923) but Kidston and Lang (1924) angulosa and Plagiochasma subplana. believed it to be intermediate between the algae and To the best of our knowledge the next paper lowermost vascular plants. Krausel (1941) created a dealing with bryophytic spores was published by new intermediate class, the Algomycetes, between Lundblad (1954) when she created a new form algae and fungi. Arnold (1954) observed that "it had genus based on the dispersed united spore tetrads evolved to a level comparable to that of the Ricciisporites tuberculatus Lundb lad from the bryophytes" but at the same time he contradicted Jurassic of Scania and attributed them to Ricciaceae himself by saying that there was no apparent reason but later thought that its affinity was dubious for classifying it with the bryophytes. Lacey (1969) (Lundblad, 1959). Krassilov and Schuster (1984) find thought that Protosalvinia could be compared with that R. tuberculatus can be compared with a Riccia-Iike bryophytes which were reported by sphaerocarpalean tetrad more closely. Walton (1949) and Hueber (1961) from the In her account of Upper Mesozoic microfloras Carboniferous but sporogonia of Riccia differ from from south-eastern Australia, Dettmann (1963) those of Protosalvinia in having a number of tetrads compared some fossil miospores with those of some of smaller spores instead of a single one of large extant and fossil bryophytes, e.g., Foraminisporis spores. Stewart (1983) is of the opinion that dailyi (Cookson & Dettmann) Dettmann has been Protosalvinia does not fit well into any group of compared with the spores of Phaeoceros extant plants. The shales in which Protosalvinia has bulbiculosus (Brotero) Prosk., Foraminisporis been found suggest a shoreline (littoral zone) where wonthaggiensis (Cookson & Dettmann) Dettmann 22 THE PALAEOBOTANIST

with spores of NOLOthyLas breutelLi Gottsche, of the spores of some liVing forms of Indian Rouseisporites retieuLatus Pocock, with spores of Hepaticae and Anthocerotales as well as published Riceia beyrichiana Hampe and R eanaLicuLata Hoff. accounts of spores of the eX'1ant members of these and Couperisporites labuLallis Dettmann with spores groups besides looking for possible bryophytic of Naiadita LaneeoLala Buckman as described by spores in published literature of fossil Sporae Harris (1939). Among Other Sporae dispersae, Meh ra dispersae. As a result we can categorically state that (1974) compared some spores described as colpate and porate pollen grains (microspores) of Leisphosphaera rOlunda, Uniporata torosa, living gymnosperms and angiosperms differ radiplly Phosphospbaera obscura by Pichova (1967) from the from the spores of bryophytes and pteridophytes. Early Cambrian of south east Siberia with those of The only exceptions are the pollen grains of extant LophoeoLea and Jungermannia which are pteridosperms, like Potoniea Halle (1933), presently found in wet tropical forests. Mehra (1974) Crossotheea hoeningbausii Brongnt., C. hughesiana merely expressed surprise at the resemblance of the Kidston (Wodehouse, 1935), C. sagittata (Arnold, smooth exined fossil and living spores. 1947) which show prOXimal trilete marks besides Some other authors like Gray (1985), distal col pi and the pollen gr3ins of DoLerotheea, Richardson (1985), Pflug and Reitz (1987), which have a biarmed proximal monolete with a Richardson and Ioannides (1973) have offered vestigeal third arm and also colpi on the distal side compelling evidences to suggest that a few (Schopf, 1949). Disaccate pollen grains of Abies sporomorphs of Early Silurian and Mid-Ordovician nobiLis tOo are reported to show proximal triletes age which occur in the form of monads, diads, and (Wodehouse, 193')). Pant (1954) has even tetrads were produced by bryophytes. However, mentioned some angiospennous pollen grains like most Other authors discount the existence of fossil those of Trapa and some Liliaceae which show bryophytes or their spores during Cambrian, apparent trilete marks. Ordovician or Silurian times due to the absence of However, bryophytic spores are most closely any authentic macrofossils of bryophytes or land comparable with those of pteridophytes whose plants in rocks of these ages. isospores and microspores are usually trilete and Even in the rocks where macrofossiIs of more or less similar in size. In trying to look for bryophytes have been reported, no serious attempt distinctions, between spores of these two groups has been made lately to look for possible bryophytic and for the identification of fossil Sporae dispersae Sporae dispersae. In fact, in this connection Miller of bryophytes from those of pteridophytes we (1984) is of the opinion that in comparison with concluded that the problems could be approached macroscopic fossils, dispersed bryophytic spores in three ways. Firstly by looking for exclUSively have received little organized study, although they bryophytic characters of spores which are nOt found are probably abundantly represented in certain in the spores of Pteridophyta, Gymnospermae and sedimentary environments. To fill up this lacuna the Angiospermae. Secondly, we may try to find out present study has been undertaken. characters of spores which are prevailingly found in spores of living bryophytes and occur only rarely in other plants. Thirdly, we may identify bryophytic CHARACTERS OF HEPATIC AND fossil Sporae dispersae by comparing them first hand HORNWORT SPORES with figures, photographs and descriptions of similar Before we deal with our study of bryophytic looking in situ spores of fossil bryophytes or those spores we have to point Out that in the present of extant bryophytes. article which deals only with the fossil spores of Our account also takes intO consideration the Hepaticae and Anthocerotales, we have left out the Sporae dispersae which were described or suspected fossil spores of foliose hepatics and mosses since by previous workers as those of bryophytes. These limitations of space do not permit us to deal with are mentioned in Table 1 against the names of the spores of these plants in a single article. We comparable living forms. hope to cover this aspect at a later stage. We may also mention that in our search for CATEGORICAL DISTINCTIONS MARKING fossil Sporpe dispersae of hepatics and hornworts we OUT BRYOPHYTIC SPORES have first attempted to find out characters, which can distinguish the spores of these groups from the iso­ Among the categorical distinctions which mark and micro-spores of pteridophytes and pollen grains out bryophytic spores from those of pteridophytes of gymnosperms or angiosperms. To enable us to do (and other land plants) we may mention: so we began our work by making a first hand study 1. The peripheral flanges of some bryophytic PANT & SINGH-FOSSIL SPORAE D1SPERSAE OF HEPATICAE AND ANTI-IOCEROTALES 23

Table 1- Possible fossil bryophytlc sporae dlspersae

FOSSIL SPOROMORPHS AUTHORS AGE & LOCALllY COMPARABLE EXTANT FORMS AlTRlBUTING AFFINITIES

Family-Fossombroniaceae EV5. Fig. 50, Knox, 1939 (PI. XLI, fig. 50) Knox, 1939 Carboniferous, Productive Coal Fossombronia angulosa (Fig. 12) Measures of Fife FamUy-Pallaviclniaceae Mlgula emend. Schuster Fig. 44, Knox, 1939 (PI. XL, fig. 44) Knox, 1939 Carboniferous, Productive Coal Moerckia hibernica (Fig. 10) Measures of Fife Couperisporiles tabulatus Dettmann, 1963 Upper Mesozoic of South Australia Fossil spores of Naiadita lanceolaro (Dettmann, 1963; PI. XXI, (Buckmann) Harris figs 12-19; fig. 6c) FamUy- Sphaerocarpaceae cavers Aequitriradites (Del. & Sprum.) Dettmann, 1963 Upper Mesozoic of ustern Australia Spores of modern Sphaerocarpaceae Cookson & Dettmann, 1961 Coptospora striata Dettmann, 1963 Dettmann, 1963 Upper Mesozoic of South Australia Ceothal/us tuberosus Campbell (PI. XX, figs 1-5) FamUy- Targlonlaceae Endl. Echimonocolpiles rudae Present authors Eocene, Tibu Area, Colombia Cyathodium aureo-nilens Hamm. & Garc. D. M. Dez. 1965 Potonie, 1970 (Taf. 15, fig. 168) FamUy-Aytonlaceae cavers Fig. 51, Knox, 1939 (PI. XLI, fig. 51)Knox, 1939 Carboniferous, Productive Coal Plagiochasma subplana (Fig. 22 a,b) Measures of Fife

Aequilriradites spinulosus Cookson & Present authors Upper Mesozoic of ustern Australia Plagiochasma mtermedium Dettman (Dettmann, 1963; PI XXII, Lindenb. et Gott. figs 1·13) Aequitriradites verrucosus Present authors Upper Mesozoic of ustern Australia Plagiochasma intermedium Cookson & Dettmann Lindenb. et Gott. Perotriletes granulatus Couper, 1953 Present authors Jurassic of New Zealand (very rare) Reboulia hemispherica (PI. 3, figs 28, 29) FamUy-Marchantlaceae (Bisch.) Endl. Hamulalisporis hamulalis Present authors Eocene of Geiseltal, Lutel, Germany Marchanlia palmata Krutzsch, 1959 (Potonie, 1966; Taf. 4, fig. 50) FamUy-Rlcclaceae Dumon. Fig. 47, Knox, 1939 (PI. XU, fig_ 47) Knox, 1939 Carboniferous, Productive Coal Riccia crystal/ina (Fig. 28) Measures of Fife Fig. 48, Knox, 1939 (PI. XU, fig. 48) Knox, 1939 Carboniferous, Productive Coal Riccia beyrichiana (Fig. 26) Measures of Fife Fig_ 49, Knox, 1939 (PI. XU, fig. 49) Knox, 1939 Carboniferous, Productive Coal Riccia !Iuilans (Fig. 27) Measures of Fife Camptotriletes corrugatus (Ibrahim) Present authors Palaeozoic of West Germany Riccia sp. Potonie & Kremp, 1954 (Taf. 7, fig. 25)

Lapposisporiles lapposus Present authors Permian of Netherlands RJccia curlisii Visscher (Potonie, 1970; Taf. 4, fig. 33)

Quadrisporiles horridus Hennelly Present authors Permian-Triassic Transition of Riccia perssonii Khan (Hennelly, 1958, PI. V, figs 6, 7) New South Wales, Australia; (Potonie & Lele, 1959, PI. 1, Carbo-Permian, India fig. 26·36) Contd. 24 THE PALAEOBOTANIST

Table l-Contd.

FOSSIL SPOROMORPHS AUTHORS AGE & LOCALIlY COMPARABLEEXTANT FORMS AlTRIBUTING AFFINITIES

Tigrisporites hallensis Klaus, 1960 Present authors Triassic salsburg, Germany Spores of Ricciaceae (POlonii: 1966, Taf. 4, fig. 51) Perorriletes pseudoreticulatus Present authors Jurassic of New Zealand Riccia cana/fculaUl Couper, 1953 (PI. 3, fig. 30)

Schizosporis reticulatus Cookson & Present authors Upper Mesozoic of Eastern Australia Riccia discolor L. et L. Dettmann, 1959 (Dettmann, 1963; (Dettmann, 1963), Upper Neo· PI. XXVI, figs 20, 21) comian beds of Canada (Pocock, 1962) Cooksonites variabilis Pocock, 1962 Dettmann, 1963 Valanginian·Barremian strata of Modern spores of Ricciaceae (Dettmann, 1963; PI. XXI, figs 8·11) Canada (Pocock, 1962) Upper Mesozoic of S.E. Australia (Dettmann, 1963) Rouseisporites reticulatus Dettmann, 1963 Barremian·Lower strata of Canada Riccia beyrichiana Hamper and Pocock, 1962 (Dettmann 1963, (Pocock, 1962), Aptian·A1bian and Riccia cana/iculaUl Hollm. PI. XXIII, figs 4·9; Potonie, 1966, Turonian beds of Siberia (Samoilo­ fig. 6d) vitch et al., 1961). Upper Mesozoic of Eastern Australia (Dettmann, 1959, 1963). Famlly- Anthocerotaceae Dum. Fig. 45, Knox, 1939 (PI. XU, fig. 45) Knox, 1939 Carboniferous, productive Coal Anthoceros chilensis (Fig. 30) Measures of Fife Fig. 46, Knox, 1939 (PI. XU, fig. 46) Knox, 1939 Carboniferous, Productive Coal Anthoceros punctatus (Fig. 29) Measures of Fife Pseudogravisporites reticulatus Present authors Lower Permian of Netherlands Pbaeoceros laevis Visscher, 1966 (Potonie, 1970, Taf. 7, fig. 70) Foraminisporis dailyi Dettmann, 1963 Upper Mesozoic of South East Phaeoceros bu/biculosus (BrOlero) Dettmann, 1963 (PI. XIV, figs 15·18) Australia Nototbylas breutelli Gottsche Foraminisporis wonthaggiensis Dettmann, 1963 Upper Mesozoic of South East Nototbylas breutelli Gottsche (Cookson & Dettmann) Dettmann, Australia 1963 (PI. XIV, figs 15·18) Anthocerisporis europas Krutzsch, 1963 Krutzsch, 1963 Oligocene-Miocene Anthoceros (Tafel 3, figs 1·8) Germany

Antbocerisporis bohemicus Krtz. & Krutzsch, 1963 Oligo-Miocene Anthoceros Pad. (Krutzsch, 1963; Tafel 4, figs 1·7) Foraminisporis granoverrucatus Krutzsch, 1963 Upper Oligocene Anthoceros tuberculatus Krtz. & Pad. (Krutzsch, 1963; Tafel 2, figs 1·8) Foraminisporis zonaloides KrUlzsch, KrUlzsch, 1963 Miocene? Antboceros spp. 1963 (Tafel I, figs 1·6) Rudolphisporis rudolphi (Krtz.) Krutzsch, 1963 Lower Miocene Anthoceros d. punctatus Krtz. & Pacl. (Krutzsch, 1963; N. Bohemia Tafel 9,. figs 1·7) Rudolphisporis saetosus (Pflanzl.) Krtz. Krutzsch, 1963 Oligocene, Bohemia Anthoceros d. punctatus (Krutzsch, 1963; Tafel 10, fig. 7) Rudolphisporis torgauensis Krtz. Krutzsch, 1963 Middle Oligocene, Bohemia Anthoceros d. punctatus (Krutzsch, 1963; Tafel 10, figs 1·6) Contd. PANT & SINGH-FOSSIL SPORAE DISPERSAE OF HEPATICAE AND ANTHOCEROTALES 25

Saxosporis duebenensis Krtz. Krutzsch, 1963 Lower Miocene, Germany Phaeoceros laevis subsp. (Krutzsch, 1963; Tafel 5, figs 1·8) carolinianus

Saxosporis gracilis Krtz. & Pacl. Krutzsch, 1963 Oligo-Miocene, Germany Phaeoceros laevis subsp. (Krutzsch, 1963; Tafel 6, figs 1·6) carolinianus

Pbaeoceros Form A, 5, C ]arzen, 1979 Upper Cretaceous Phaeoceros laevis, (jarzen, 1979; PI. VI, figs 1·8) Form A resembles to fossil spores of Baculatisporites pseudoprimarius Krtz. & Thiergart and Ceratosporltes couliensis Srivastava

spores like Riccia, Plagiochasma, Reboulia young condition), e.g., spores of Treubia Goeb., which show pores near their trilete laesurae Fossombronia. Raddi, Sewardiella Kash., These pores are not at all comparable with the Calycularia Mitt., Riccardia Raddi, Monoclea germ pores of angiospermous pollen and Schust., Targionia L., Cyalhodium Kunze., possibly they never function like them. Flanges Conocephalum Necker, ExormOlheca Mitt., are present in some pteridophytic spores but Dumortiera Reinw., Riccia billardieri Mont. & none of them shows such pores although pores N., R. discolor L. & L., R. melanospora Kash., etc. of a different kind are found in the spores of Such spores are categorically absent in some species of Asplenium like A. abyssinicum pteridophytes except in modern Equisetum, which reportedly shows a prominent granular whose spores have peculiar hygroscopic elaters. perine with circular or oval non-sculptured areas 4. A characteristic proximal depression is present sometimes replaced by large perforations (see between two folds in spores of Cyathodium as Nayar, 1964) but the size, shape and position of reported by Mehra and Sood (1969) and Mehra these perforations are not figured and therefore and Sokhi t1972). However, such a depression it was not possible for us to compare these cannot be regarded as a colpus since a true spores with the porate spores of Hepaticae colpus has never been observed in the spores of whose position and number is characteristic for thalloid Hepaticae and in Anthocerotales. The a particular species. term col pus sensu Faegri and Iversen (19'50) is 2. Among other exclusive bryophytic characters we typically applied to a thinner area of the exine may mention the occurrence of permanently on the distal side from where the germ tube adherent spore tetrads like those of generally emerges and sometimes ectexine may Cryplolhallus Malmb., LudWig even be absent in the region of a col pus. where the tetrads are tetrahedral and in some '5. Non-adherent monolete spores produced in species of Riccia (Mich) L. like R. curtisii James bilateral tetrads are typically absent among and R. perssonii Khan where the tetrads could hepatics but as mentioned in (2) above, bilateral also be bilateral. No doubt the microspores of spores are found only in Riccia curtisii and R. some species of Selaginella, vi z., S. estrellensis perssonii (Khan, 19'5'5) although in these they Hieron and S. delicatissima are also dispersed in occur in permanently adherent tetrads. tetrads but they are clearly different from the 6. Complete absence of megaspores among the spores of the bryophytes in being enclosed in a species of bryophytes. These are present in some common wall (see Tryon & Tryon, 1982, p. 821 pteridophytes and theoretically also in all seed 822). The pollen grains of some angiosperms, plants. e.g., Ericaceae, Mimosaceae, Droseraceae, Sapindaceae and others (see Erdtman, 19'52) are also jugate but they are typically colpate and IDENTIFICATION OF SPORAE DISPERSAE OF aperturate while the jugate microspores of FOSSIL HEPATICAE bryophytes are neither col pate nor aperturate. Out of the categorical distinctions which are Moreover, the jugate tetrads of bryophytes have mentioned above, we have only come across generally larger elements (ca '5'5 to 60 /-lm), sporomorphs having characters (2) and (4) among whereas the jugate pollen grains of the above hitherto described fossil Sporae dispersae. We find angiosperms are very small. character (1) consisting of pores in the flange near 3. Mature spores of many hepatics become alete the ends of trilete laesurae, is present only in (although they are arranged in tetrads in the Rouseisporites reticulatus Pocock. These spores have 26 THE PALAEOBOTANlST been compared bv Dettmann ( 196::\) with the spores ridges Sullivan and Hibbert (1964) compared of Riccia iJeyricbiana Hampe and Riccia Tetrapterites confidently with the mature spores of canaliclilata Hoff. but we find that such pores are Sphaerocarpos LudWig whose spores remain also present in Riccia crtlciala Kash Accordinglv, we adherent in tetrads even after liberation (see Text­ can assign ROliseisporites to Hepaticae with certainiry fig. 1 M, N). Though these spores are not found in (Text-fig 1 0, P} situ, their possible bryophytic nature cannot be Among earlier reported sporomorphs joined denied firstly since the earliest authentic liverworts tetrads (character 2), the first spore of this kind was are known from the Upper Devonian and secondly, reported by Erdtman ( 19'54) from the Rhaetic Liassic since unusual type of spore tetrads called of north-west Scania who provisionally referred them Tetrapterites Sullivan & Hibbert occur in beds of to Telraslilcites magna. As quoted by Lundblad Menai Straits, Caernarvonshire As pointed Out by (19'19) "Erdtman thought that sulcate grains of a Lacey (1969) these could even belong to a similar type have not been encountered in recent fragmentary thalloid bryophyte which had been plants Slightly similar pollen types occur in certain mistaken by Walton (1928) as minute pinnules of a monocotyledons and monocotyledonoid pteridosperm genus Rhodea Pres!. Lacey thought that dicotdedons However, it would be preposterous to these thalloid fragments could be p~utof a Riccia­ base any conclusions regarding the affinities of like plant which was the parent plant of 'Telraslilcitcs' on such grounds" In the same year Tetrapterites. Thirdly, as already discussed by Lacey Lundblad (19'14) also found similar spores and the are themselves possibly referred them to Ricciisporites tllberculatus. She also represented by Naiadita Buckman from as far back gave a detailed diagnosis and on the basis of her as the Rhaetic of Britain and that genus also has large studies, attributed them for the first time to spores which sometimes persist in tetrads and bryophytes (Ricciaceae) but later (Lundb lad, 19'59) sometimes separate out but always show a thought that their affinities were uncertain though prominent flange on each member of the tetrad. In Krassilov and Schuster (1984) find that R the occasional separation of members of tetrads into luberculatus can be compared to sphaerocarpalean monads and their occasional persistence in tetrads tetrads more closely but on the basis of our Naiadita seems to be comparable with observations we think that it is more closely Haplomitrium gibbsiae (Steph.) Schuster (see comparable to the spore tetrads of Riccia curtisii Sch uster, 1984) and Riccia frostii (our unpublished (see P!. 1, figs 1-4) than those of sphaerocarpalean findings). spores. Potonie (1970) in his synopsis of Sporae A third permanently joined tetrad of the same dispersae has described Lapposisporites lapposus kind called Tetrapterites uisensis Sullivan & Hibbert Visscher from the Permian of Netherlands as having 1964 is described from the Lower Carbon iferous united spore tetrads but these differ reponedly from shales in the Menai Straits, Caernarvonshire. This Ricciisporites tllberclllatlls in the absence of a distal tetrad is about 200 }.lm in diameter and it is enclosed "sulcus". We observed that these sporomorphs by a non-cellular membrane which has a tetrahedral closely resemble the spore tetrads of liVing Riccia shape and possesses a large wing like expansion curtisii James (see Text-fig. 1E, F). Without near each of its four apices. Subsequently on the reference to the previous accounts of united fossil basis of scanning electron microscopic studies tetrads by Erdtman (19'54), and Lundblad (19'54, Hibbert (1967) found that the enclosing membrane 19'59) another similar fossil sporomorph was of spore is highly ornamented with anastomosing described by Hennelly (19'58) as Quadrisporites - PLATE 1

1, 2. Ricciisporites tuberculatus Lundblad: Spore tetrads spore tetrads X 250. showing distal sulcus. 11-13. Tetrahedraletes cr. T medinensis Gray et al. : Permanently 3. Riccia curtisii James: Permanently joined spore tetrads X joined isobilateral tetrads of spores. 1000. 12. Conventional tetrahedral arrangement of spores in a tetrad 4. Sphaerocarpos donnellii Ausl. : Joined spore tetrad. X 1000. 5. Densoisporites: A spore tetrad with one missing spore X 16, 17 Riccia perssonii Khan. : Permanenrly joined isobilateral 500. and cruciate spore tetrads. 6. Lundbladispora : Tetrad showing isobilateral arrangemenr 1,2. after Lundblad, 1954; 4, after Schuster, 1984; 7-10, of spores X 500. after POlOnie & Lele, 1959; 11-13, after Gray et al., 1986; 6-10,14,15. Quadrisporites horridus Henn.: Isobilateral joined 14, 15, after Hennelly, 1958). PANT & SINGH-FOSSIL SPORAE DfSPERSAE OF HEPATICAE AND ANTHOCEROTALES 27

2 3'

4

•• 0,., ~ .~ ... . ,. ;"

." a ~ ,

•

7 8

9

PLATE 1 N 00

/tB~";:;,,::::?,~"'"

,- ..

,;. ' . .('< ~: . ,'.

~2~ ~ ... ~ ., '... () J . ~, ~ A~ ~ ~t:-,..:~: ~ '. '. ~ .- .'" • \,.( ~ ~. ~ _~ :,:~::. :~J N '··""'·"'·\d_.• , ~ ~ PANT & SINGH-FOSSIL SPORAE DlSPERSAE OF HEPATICAE AND ANTHOCEROTALES 29

horridus from the Permian-Triassic transition of New by Kremp (196'5), wherefrom germination takes South Wales, Australia (PI. 1, figs 14, 15). Shortly place although some of the living hepatics have alete afterwards Potonie and Lele (1959) gave an spores, viz., Riccia (mich.) L., Cyathodium Kunze, emended diagnosis of the same species from Talchir Targionia L. which bear a prOXimal conca\'ity but beds and South Rewa Basin (Carbo-Permian age) of that too has not been shown to be meant for Indian Peninsula. These authors neither compared germination, although germinal furrows have been their Quadrisporites horridus with previously reported in most of jungermannialean spores by described adherent fossil tetrads nor with spores of Miyoshi (1966). living bI)'ophytes which have permanently adherent Among spores of bI)'ophytes and pteridophytes tetrads even though their Quadrisporites horridus we can also distinguish two other kinds mentioned (PI. 1, figs 7 -10) is closely similar to tetrads of Riccia below: perssonii Khan (see PI. 1, figs 16, 17). Unlike the above dispersed adherent spore tetrads which are comparable with the united spore Spores having characters more common in tetrads of bI)'ophytes, Chaloner (19'58) reported Bryophytes but rare in Pteridophytes tetrads of Didymosporites having two large and two Among such spores reticulate ornamentation small spores enclosed in ,a common membrane which are attributed to Stauropteris hurntislandica seems to be more common in bI)'ophytes than tuberculate, verrucate or echinate (see Text-fig. lL). Other spore tetrads which are sculpturing particularly among hepatics, whereas the spores joined by tubular connections are also reported in of pteridophytes are more commonly echinate, some extant pteridophytes and angiosperms verrucate and tuberculate and less commonly including some Indian species of Isoetes L. described reticulate except in the families Lycopodiaceae, by Pant and Srivastava (1962) and by Goswami and Sphenophyllaceae and Ophioglossaceae. Echinate Khandelwal (1973) in Ophioglossum costatum and spores of bryophytes like Cyathodium Kunze are o vulgatum L. Wodehouse (193'5) had earlier characterised by haVing a proximal concavity. As far mentioned a report of the occurrence of similar as we can gather from our own study of spores and pollen grains by Stout (1918) in some angiosperms from preViously published accounts fragmenrimurate like Cichorium intyhus. The intines of such spores (broken muri) too are prevailingly more common in are continuous through tubular connections and bryophytes but rare in pteridophytic spores although they have a Single continuous protoplast. Pant and they have been sometimes reported in spores like Srivastava (1962) suggested that the occurrences of those of Lycopodium cernuum L., L. serpentinum occasional joined spores may have been caused by Kunze and in L. uoluhile Frost. the nonseparation or incomplete separation of individual spores of a tetrad due to meiotic irregularities caused by hybridity. Spores having characters rare in Bryophytes The presence of the so-called "sulcus" in but more common in Pteridophytes Lundblad's Ricciisporites tuherculatus remains uncertain since the wall of the distal depressed area Among spores of this kind are those having in the joined fossil grains has never been shown to "Crassimarginate" and "incrassate" trilete laesurae have a thinner wall like that of a sulcus, as defined with thickened lips or unsculptured ridges which

0+- Text-figure I-A, Scbizosporis reticulatus (Cookson & Dettmann) Dettmann: Alete spore showing reticulate ornamentation; B, Riccia discolor L. & L. : Alete reticulate spore X 500; C, Tigrisporites ballensis Klaus; D, Riccia sps. : Spore showing irregular reticulations X 500; E, Lapposisporites lapposus Visscher. : Joined spore tetrad; F, Riccia curtisii James: Permanently joined tetrad of spores X 500; G, Perotriletes pseudoreticulatus Couper; H, Riceia canaliculata Hoffm. : Spore showing rugulate ornamentation X 500; I, Pseudogravisporites reticulatus Visscher; J, Pbaeoceros laevis (L.) Prosk. : Spore showing margin around trilete mark X 500; K, Pbaeoceros bimalayensis Kash. : Spore X 500; L, Didymosporites scottii Chal. : Spore tetrad showing two small and two large spores; M, Tetrapterites visensis Sullivan & Hibbert. N, Cryptotballus mirabilis Malmb. : Spore tetrad; 0, Rouseisporites reticulatus Pocock; P, Riccia cruciata Kash. : both spores (0, P) shOWing pores in the flange X 500; Q, Ecbimonocolpites rudae Hamm. & Garc. 0. M. Dez. : Ecf)inate spore showing a colpus·like depression; R, Cyatbodium aureo-nitens (Griff.) Mitt. : Spore showing a proXimal concaVity and echinate exine sculpturing X 500; S, Hamulatisporis bammulatus Krtzch.; T, Marcbantia polymorpba L: Distal surface of a spore; U, Camptotriletes corrugatus (Ibrahim) Potonie & Kremp : Trilete spore fragmentimurate sometimes anastomosing; V, Riccia fluitans L. : Spore showing fragmentimurate exine X 500; W, Riccia frostii Aust. : Spore X 500; x, Y, Perotriletes granulatus Couper; Z, Z., Plagiocbasma intermedium Undeb_ & Gott. : Distal and proXimal surface of a spore showing granulate flange_ (A, C, M, 0, S, after Potonie, 1966; E, I, Q, after Potonie, 1970; L, after Potonie, 1960; N, after Schuster, 1984; U, after Potonie & Kremp, 1954; G, X, Y, after Couper, 1953.) 30 THE PALAEOBOTANIST occur commonly in pteridophytes (Erdtman, 19'52; PerOlrileles pseudoreliculalus (Couper, 19'53)­ Harris, 19'5'5) but these are seldom reponed in any Riccia canaliclilaia Hoffm. (Text-fig. 1 G, H), bryophytes, the only spores which have a slightly Pseudogravisporiles reticulalus Visscher (Potonie, ridged trilete are those of Phaeoceros (Jimalayensis 1970)-Phaeoceros bimalayensis Kashyap and P. Kashyap and P laevis Prosk. laevis (L.) Prosk (Te),,'!:-fig. 1 I-K) and Tigrisporiles Apan from the characters mentioned above as ballensis Klaus Nov. resemble spores of Riccia sp. in categorical or otherwise for distinguishing their size and ornamentation (Text-fig. I C, D). bryophytic spores from those of other land plants Aequilriradiles ~'errucosus,A. spinulosus one of the most reliable methods of referring a Cookson & Dettmann and Schizosporis reliculailis particular fossi I sporomorph to a blyophyte should Cookson & Dettmann (see Dettmann, (963) which be first hand comparison of spores of living are reponed from the Upper Mesozoic of Eastern bryophytes with a fossil sporomorph as has been Australia can be compared with the spores of the done among others by Knox (1939), Dettmann Plagiochasma inlermedium Lindenb. (PI. 2, figs 1-6) (1963), Nagy (1968), Jarzen (1979) and Krutzsch and with the spores of R. melanospora (PI. 2, figs 7­ (1963 ). 9) (see Gupta & Udar, 1986) respectively and also with spores of R. discolor-Text-fig. lA, B IDENTIFICATION OF SPORAE DISPERSAE OF (determined by Dr K. R. Bapna and collected from FOSSIL HEPATICAE AND ANTHOCEROTALES Mt. Abu, Rajasthan, India) On the basis of the above mentioned categorical and prevalent distinctions between the spores of IDENTIFICATION OF BRYOPHYTIC FOSSIL pteridophytes and bryophytes we found that SPORAE DISPERSAE WITH SIMILAR dispersed fossil spores like Camplolrileles LOOKING IN SITU SPORES OF FOSSIL corrugalus (Ibrahim) Potonie & Kremp 1954 BRYOPHYTES described from the Permo-Carboniferous beds of The only definitely known in situ spores of Germany resemble living spores of Riccia fluilam L fossil bryophytes which have been described so far and Riccia froslii Aust. in having broken muri are those of Naiadila lanceolala (Such.) Harris (fragmentimurate) forming incomplete reticulations 1939; Hepalicites disco ides Douglas 1973; H (Text-fig. 1 U-W), Echimonocolpiles rudae Hamm. nidplirensis Pant & Basu 1978; NOlOlhylacites & Garc., D. M. Dez (see POtonie, 1970) which are filiform is Nemjec & PacltOva (see Krassilov & described from the Eocene of Columbia are quite Schuster, 1983) and NOlOlhyliles nirulai Chitaley & like those of present day Cyalhodium aureo-nilens Yawale 1980 which range in age from the Triassic to (Griff.) Mitt. because they have echinate exine and a the upper·most Cretaceous or Eocene. The concavity on one side (Text-fig. 1 Q, R). The characters of most of the above in situ spores are following spores resemble spores of liVing forms rather generalised and it is difficult to compare them noted against each of them Hamlilalisporis with any bryophytic spores but a fossil sporomorph, hamulalus Krutzsch (see Poronie, 1966)­ Couperisporites labulallis Dettmann is closely Nlarchanlia polymorpha L. (Text-fig. 1 S, T), comparable with in situ spores of Naiadita PerOlrileles granulalus (see Couper, 19'53)­ lanceolala (see Dettmann, 1963). Plagiochasma inlermedium Lindenb. & Gott. and The occurrence of C. labulalus in the Triassic of

Reboulia hemispherica Raddi (Text-fig. 1 X-Z1 ), Australia lends suppon to its relationship with

PLATE 2 --

1, 2. Aequitriradites verrucosus (Cookson & Dettmann): surface X 500. Spores showing dislal surface x 500. 11. Phaeoceros himalayensis Kash. : Proximal surface in spore 3, 4. Aequitriradites spinuloslls (Cookson & Dettmann): X 500. Proximal and dislal surface of spores showing spinules X 12 -15. Foraminisporis wonthaggiensis (Cookson & Dettmann) 500 Dettmann: Spores showing proximal and dislal surfaces 5, 6. Plagiochasma intermedium Undeb. et GOtt.: Proximal of spinate spores X 500. and distal surface of a spore showing spinulate 16, 17, 21-23.NolOthylas breutellii Gottsche.: Spores showing orhamenlation X 500. spinulate and verrucate omamenlation X 500. 7, 8. Schizosporis reticulatus (Cookson & Dettmann): Spore 18-20. Foraminisporis daily; (Cookson & Dettmann) Dettmann: showing polar reticulate view X 250. Proximal and dislal surface of a spore X 500. 9. Riccia melanospora Kash.: Alete spore with reticulate (1-4,7-8, 12-23. after Dettmann, 1963; 9. after Gupla & ornamenlation X 600. Udar, 1986). 10. Phaeoceros laevis (L.) Prosk. : Spore showing proxi mal PANT & SINGH-FOSSIL SPORAE DISPERSAE OF HEPATICAE AND ANTHOCEROTALES 31

I.

• ...

~'M .",' ~ -....?1 f" .... . ~JI .~ .. '. • ~:;.:.;~,:•.0 ';~ . ;J / ~~;';.~...... '.. ...,. ~ ~"';iJ~~ ,r' .... __ ~.~,- . -, :~1?··,:..;:;. I~'_,' '~") : ~, ,.. I'~"'I"7 ~ ~-.~ !...... t__ ...~'.:.".~$ ..\ . fo...e.. ; ;;:." • . ~.::-~~~.:..~_. \...... "*-.- .. ":......

!q:~!Pi~,i

7

20

.8

A~~'~' -t,~,r~~;t. ~m,·~1(;F;r '~"t,~..-\. if::":;"/ .f.." l..• 'H :::., ~~':

r~" ..-::L~/~:;;. I .•..,.' ~,"f.!!?": ~:;'·foI~~ ,.~, .....(..) . :' ~~$/.. 1 .... -,f :_~~~,~=~:~;~~~ .{ .... '(' r ~ ~ • _ .. I ,,-'. ~;.. "" \. l.fJ.'- ~.. :'~ : l f t,··.. '. ,,1,. ./ {:\"',;:~:.~/~~.;~", ~-. 'I .•,(.,...... \~.t.' . .. ~":.,~.;.~ f f ., .... :."'':'''''l "'; ~~;~';>;t'), ','.;;:<::,}:~~~~ ...... -:'-.~~ ...=-:..../'..

PLATE 2 32 THE PALAEOBOTANIST

Naiadita and it may suggest that Naiadita or. allied from Lapposisporites in the presence of a distal plants also existed in Australia, sulcus, Playford (196'5) called jugate tetrads having DISCUSSION AND CONCLUSIONS laevigate exine by the name Lundbladispora and Banerji and Maheshwari (197'5) described similar Our comparative study of the spores of extant spores from Panchet beds (Triassic) under bryophytes and other land plants enables us to say Decisporis variabilis; Tewari and Meena (1988) now that fossil Sporae dispersae of bryophytes can be assigned spores of the same kind to Lundbladispora placed into three broad categories: (i) spores which Playford and Densoisporites Weyland & Krieger can be categorically recognised as belonging to emend Dettmann, whose ornamentation was bryophytes on the basis of the presence of characters reportedly diverse, However, this is not discernible which are indubitably different from those of other from their photographs although the occurrence of land plants, (ii) spores which can be assigned to crusiate and tetrahedral tetrads not reported by these brvophytes with some uncertainiry on the basis of a uthors, can be easily made (see PI. 1, figs 5, 6). more frequent occurrence of their characters in Seemingly unaware of the publications of Gray brvophytes and less so in other land plants, and (iii) (198'5) and others on the abundance of obligate spores which may belong to bryophytes but cannot jugate tetrads in Pre-Triassic strata, Tewari and be distinguished from those of other land plants, Meena (1988) quoted Heslop-Harrison (1973) to because of the presence of characters which occur visualize "a situation (which) appears to have arisen equally frequently in all land plants but some of during the Early Triassic times" where they said that them could belong to bryophytes Although we the enzymic reaction for the separation of spores know it for certain that land plants had come into "fails due to some reasons or the other" and leads to existence during the Upper Silurian and Lower "a condition where a number of tetrads will remain Devonian and the earliest plants attributable to intact and consequently no individual spore gets Bryophyta are also recognised from the Lower released". The fact of the existence of jugate tetrads Devon ian, as far as Sporae dispersae are concerned, in Ordovician, Silurian would push back the categorically distinguishable spores of the group supposed failure of reaction at least by about 2'50 have not been reported from the Silurian-Devonian m.y. in time but we are unable to understand the time, seemingly fashionable necessiry of quoting Heslop­ The present geological agewise search of fossil Harrison (1973) and his enzymic mechanism to bryophytic spores shows that the first categorically explain the occurrence of obligate tetrads in the distinguishable bryophytic miospores of the type Early Triassic. represented by permanently adherent jugate Leaving aside the above digression into the tetrahedral tetrads are traceable up to the Middle enzymic explanation, we may sum up our Ordovician of North America and Saudi Arabia, Late information on the above mentioned permanently Ordo\-ician and Early Silurian of Ghana, united tetrads of variously named spores as Czechoslovakia, North America and Libya from indicating that bryophytes having such generalised where Gray (198'5) reported some obligate tetrads of forms of spores may have come into existence as Tetrahedraletes_ Thereafter such tetrads under the early as the Middle Ordovician or Silurian and name Tetrapterites vissensis are 'reported from Lower thereafter continued to exist during the Devonian, Carboniferous by Sullivan and Hibbert (1964). Carboniferous, Permian, Triassic and Liassic times. Spores of this generalised type are also reponed At this stage it is necessary to pOint out that the from Carbo-Permian beds of India by Potonie and occurrence of similar jugate pollen and spore tetrads Lele (19'59) and from the Permian·Triassic transition in angiosperms and Selagineffa cannot be used as an beds of New South Wales in Australia by Hennelly argument to weaken the evidence for the occurrence (19'58) as Quadrisporites horridus, similar spore of bryophytes in the above mentioned strata since tetrads were later described by Visscher, 1966 (see beside such spores of these plants being radically Potonie, 1970; Tiwari & Meena, 1988) under the different from those of bryophytes, on the basis of names Lapposisporites lapposus and Paralundbladi­ the fossil record of angiosperms it would be wrong spora from the Permian of Netherlands which are to imagine that jugate tetrads of spores occurring reportedly different from the earlier described jugate from Ordovician to Jurassic times could belong to tetrads of Ricciisporites tuberculatus (Lundblad, that group. Even their attribution to heterosporous 19'54, 19'59) from Rhaeto-Liassic beds of north-west pteridophytes like Selagineffa would seem to be Scania. Visscher mentions that Ricciisporites differs equally unlikely unless it is supported by associated PANT & SINGH-FOSSIL SPORAE DISPERSAE OF HEPATICAE AND ANTI-IOCEROTALES 33 megaspores. On the contrary, even though we could tetrads occur in the angiosperms which had come assign jugate tetrads from Pre Cretaceous to Mid­ into existence. Ordovician, to bryophytes on the basis of the Although bryophyte-like jugate spores have nor presence of fossils of this group in such beds or in been recorded in strata after the Liassic, the slightly later ones, it would be difficult to assign cosmopolitan occurrence of bryophytes producing Cretaceous and Post Cretaceous jugate tetrads to such spores at the present time suggests t:lat future bryophytes with cerrainiry since very similar jugate research on Sporae dispersae of strata after the

Table 2-Fossil spores attributable to Hepaticae and Anthocerotales through Geological Times

ERA PERIOD EPOCH FOSSIL SPOROMORPHS Quaternary Recent Pleistocene Pliocene Miocene Baculatisporites pseudoprimarius Knz. & Thier.; Foraminisporis zonaloides Krlz.; Rudolphisporis rudolphi Knz.; Saxosporis duebensis Knz. Cenozoic Terliary Oligocene Bohemiasporis vac/avensis Knz. & Pad.; Anthocerisporis europaeus Knz.; A. bohemicus Knz. & Pad.; CeralOsporites couliensis Srivastava Eocene • Echimonocolpites rudae Hamm. & Gare. D. M. Dez. • Hamulatisporis hamulatis Krutzsch Palaeocene

Cretaceous Upper • Schizosporis reticulatus Cookson & Dettmann Lower Phaeoceros Form A; Phaeoceros Form B; Phaeoceros Form C Jarzen (1979); Aequitriradites sps. (Del. & Sprum.) Cookson & Dettmann; • A verucosus, •A spinulosus (Cookson & Dettmann; Cooksonites variabi/is Pocock; Coptospora striata Dettmann; Couperisporites tabulatus Dettmann; Foraminisporis dailyi Cookson & Dettmann; Foraminisporis wonthaggiensis Cookson & Dettmann; • Rouseisporites reticulatus Pocock Upper Mesozoic Jurassic Middle • Pero/n·letes granulatus Couper; • Perotriletes pseudoreticulatus Lower Couper Upper Ricciisporites tuberculatus Lundblad, • Tigrisporiles ha//ensis Klaus; Triassic Middle Lundbladispora Playford; Densoisporites Weyland & Krieger emend Lower Dettmann; Verrucosisporites Ibrahim emend. Smith et a/.; • Quadrisporites horridus Hennelly

Upper Permian Lower Lapposisporiles lapposus Visscher; • Pseudogravisporites reticulatus Visscher ~ Upper • Quadrisporites horridus Hennelly o Pennsylvanian Middle 0:: tLI & t.r.. Lower • Camptom·lites corrugatus (Ibrahim) Potonie Kremp Z o c:l Mississippian Upper Un·named Sporae dispersae by Knox (1939) Lower Tetrapteriles visensis Sullivan & Hibbert ~

Upper Palaeozoic Devonian Middle Lower Tetrahedraletes medinensis Gray et a/. Upper Silurian Trilete cutinized spores Lower Tetrahedraletes medinensis Gray et al. Upper Ordovician Tetrahedraletes cf. T. medinensis Gray et al. Lower 'Fossil spores which have been anributed here for the first time. .34 THE PALAEOBOTANIST

Liassic may also yield remains of such spores. Pocock (1962). Spores of this kind have not been In this connection Gray (1985) has suggested reported in any pteridophytes. that these earliest jugate tetrads could belong either Coming to the second category of Sporae to anyone or any combination of the following­ dispersae which can be doubtfully assigned to protobryophytes, protohepatics or hepatics bryophytes are alete echinate spores having a unrelated to the later trilete spore producers or to concavity on one side like those of modem common ancestors for bryophytes' and vascular Cyathodium aureo-nitens. This category includes plants. On the contrary, Schuster (1966), Crum dispersed spores which have been reported from the (1972) and Hebant (1977) have suggested that the Palaeocene of Columbia as Echimonocolpites rudae similarities between these fossil spores which are by Hammen and Garcia de Mutis (1965) but we need jugate and those of some extant bryophytes, are two to look for spores of this kind in earlier or later great to be due to parallel evolution. We agree with strata. the latter viewpoint and believe that the occurrence Presently our knowledge of the spores of the of joined spores of Tetrapterites-Quadrisporites type earliest plants attributed to the bryophytes occuring in the rocks of Middle Ordovician age indicates that in Upper Silurian or Lower Devonian strata, is rather out of our two groups of plant amphibians­ scanty but if the spores of these plants are Bryophyta and Pteridophyta, the Bryophyta seem to represented in the Sporae dispersae of these times have come into existence earlier and that even in (see Gray, 1985; Richardson, 1985; Richardson & plants of this group perhaps hepatics like Riccia or Ioannides, 1973; Gensel & Andrews, 1987) they Sphaerocarpos appeared before the foliose forms. If seem to belong to the third category of bryophytic in addition to this evidence the fossil history of early spores which are indistinguishable from those of pteridophytes continues to remain traceable only up other land plants. to the Upper Silurian, as it is at present, it would The sporomorphs which have been attributed to suggest that there would be no question of assuming the bryophytes indicate their lineages, viZ., that the bryophytes have evolved from pteridophytes , Sphaerocarplaes. Marchantiales and as some authors have suggested. Indeed, the Anthocerotales which can be traced back in historical evidence of permanently joined tetrads of geological time up to the Middle Ordovician (see Riccia- and Sphaerocarpos-type being traceable up to Table 2) but such sporomorphs have not so far been the Ordovician could strongly militate against the reported in any earlier strata. Reduction Theory of bryoph ytes or the ideas about Among the sporomorphs which have been their thallose forms having arisen from foliose compared with those of extant forms of bryophytes, forms. the largest number are Riccia-like suggesting that Another type of categorically distinguishable such forms have been in existence not only for a Sporae dispersae of bryophytes which show long time but had also been fairly common. Another reticulate sculpturing and an equatorial flange fact which is emphasized by this study is our haVing pores flanking the distal end of trilete uncertainty about the origin of bryophytes and the laesurae like those of modem Riccia, Plagiochasma evol ution of their various lineages in geological and Reboulia, are shown here to go back only up to time. These problems can be solved only by further Upper Mesozoic (Lower Cretaceous) where they are work on the fossils of the group. exemplified by Rouseisporites reticulatus (see Pocock, 1962; Samoilovitch et a/., 1961; Dettmann, 1959, 1963) and thus it is fairly clear that plants of ACKNOWLEDGEMENTS Marchantiales like those mentioned above had also come into existence in the Lower Cretaceous or Upper Jurassic if not earlier. A search of their The authors are thankful to Professor D. D. existence in subsequent strata is also likely to be Nautiyal for manifold help and research facilities. fruitful. They thank to Dr D. K. Chauhan for his kind help in The third kind of categorically determinable preparation of figures. Furthermore, the junior bryophytic Sporae dispersae which are alete and author (R.S.) wishes to thank the Council of reticulate like those of many species of modem Scientific & Industrial Research, India for awarding Riccia, vi'z., R. billardieri, R. crispatula, R. discolor, R. her the Research Associateship and the senior author gangetica and R. melanospora include Schizosporis (D.O. P.) thanks the INSA for a Senior Scientist reticulatus described from the Upper Mesozoic of Award. Lastly the authors thank Mr S. K. Yadava for Australia (Cookson & Dettmann, 1959; Dettmann, his patience with repeated changes in their 1963) and Upper Neocomian beds of Canada by manuscript and consequent retyping. PANT & SINGH-FOSSIL SPORAE DISPERSAE OF HEPATICAE AND ANTIIOCEROTALES 35

REFERENCES Harris, W. F. 1955, A manual oj the spores oj New Zealand, Pteridophyta. Brilish (Museum) Nalural HislOry, London. Hebant, 1977. The conducting tissues bryophytes. Andrews, H. N. 1960. NOtes on Belgian specimens of Sporogonites. c. oj Palaeobotanist 7 : 85·89. Vaduz: J. Cramer, pp. 157. Hennelly, J. P. F. 1958. Spores and pollen from a Permian·Triassic Arnold, C. A. 1947. An introduction to Palaeobotany. New York. lransilion, New Soulh Wales. Proc. Linn. Soc. New South Arnold, C. A. 1954. Fossil sporocarps of lhe genus Protosalvinia Dawson, Wilh special reference 10 P. jurcata (Dawson) Wales LXXXIII(2) : 363·369. comb. nov. Svensk. bot. Tidskr. 48 : 292·300. Heslop·Harrison, J. 1973, Pollen: development and physiology. Banerji, Jayasri & Maheshwari, H. K. 1975. Palynomorphs from lhe London, BUllerwonh. Panchel Group exposed in Sukri River, Auranga Coalfield, Hibbert, F. A, 1967. The use of scanning eleclron microscopy in Bihar. Palaeobotanist 22(2) : 158·170. lhe sludy of Carboniferous miospores. New Phytologist 66 : Chaloner, W. G. 1958. Isolaled megaspore lelrads of Stauropteris 825·826. burntislandica. Ann. Bot. 22: 197·204. Hueber, F, M. 1961. Hepaticites devonicus: a new fossil liverwort Chilaley, S. & Yawale, N. R. 1980. Notothylites nirulai gen. el sp. from lhe Devonian of New York. Ann. Miss. bot, Cdn 48 : nov., a pelrified sporogonium from lhe Deccan Intertrappean 125·132. beds of Mohgaon Kalan, M.P., India. Botanique 9 : 111·118. Jarzen, D. M. 1979. Spore morphology of some Andiocerotaceae Cookson, I. C. & Dellmann, M. E. 1959. On Schizosporis, a and lhe occurrence of Phaeoceros spores in lhe Cretaceous of new form genus from AUSlralian Crelaceous deposils. Micro· North America, Pollen Spores 21(1,2): 211·231. paleontology S: 213·216. Khan, S, A. 1955. Riccia perssonii, a new and interesling species Couper, R. A. 1953. Upper Mesozoic and Cainozoic spores and from EaSl Pakistan. Svensk. bot. Tidskr. 49 : 433, pollen grains from New Zealand. N. Z. geo/. Sun!. Paleo nt. KidslOn, R. & Lang, W. H. 1924. On lhe presence of lelrads of Bull. 22 : 1·73. resistant spores in lhe lissue of Sporocarpon jurcatum Crum, H. 1972. The geographic origins of lhe mosses of North Dawson from lhe Devonian of America. Trans. R. Soc. Edinb. America's eaSlern deciduous forest I Hattori bot. Lab. S3 : 597·601. 3S : 269·298. Knox, E. M. 1939. The spores of Bryophyta compared Wilh lhose Dellmann, M. E. 1959. Upper Mesozoic microfloras in well cores of Carboniferous age. Trans. Proc. bot. Soc. Edinb. 32(4) : from Woodside and Hedley. ViclOria, Proc. R. Soc. Viet. 477·487. 71 : 99·105. Krassilov, V. A, & Schusler, R. M. 1984. Palaeozoic and Mesozoic Dellmann, M. E. 1963. Upper Mesozoic microfloras from Soulh fossils. In: Schusler, R. M. (Ed.)-New Manual oj Bryology, EaSl Auslralia. Proc. R. Soc. Viet. 77(1) : 1·148. Hallori BOl, LaboralOry, pp. 1172·1190. Dixon, H. N. 1927. Muscineae. In: W. Jongmans (Ed.) Fossi· Krausel, R. 1941. Die sporokarpien Dawsons, eine new Thallo· lium Catalogus. II, Plantae 13 : 1·116. phyten Kalsse des Devons. Palaeontographica 868 : 113·135. Douglas, J. G. 1973. The Mesozoic floras of ViCloria. 3. Ceo/. Sum Kremp, G.O.W, 1965, Morphologic Encyclopedia oj Palynology. Viet. 29 : 185. Tucson : U.A. Press. Edwards, D. S. 1986 Aglaophyton major: a non·vascular land· Krulzsch, W. 1963, Alias der millet·und jungertiaren dispersen planl from lhe Devonian Rhynie Chert. Bot. I Linn. Soc. Lond. sporen und Pollen·sowie der Mikroplanktonformen des 93 : 173·204. nordlichen Milleluropas-Liefg II-Die sporen der Anlhocero· Erdlman, G. 1952. Pollen morphology and plant : Angio· taceae und der Lycopodiaceae-VEB DIsch. Verlag Wiss. sperms. Uppsala. Berlin: 1·141. Erdlman, G. 1954. Some remarks on lerms, diagnoses, classifi· Lacey, W. S. 1969. Fossil bryophytes. Bioi, Rev. 44 : 189·205. calion and melhods in palynology. Svensk bot. Tidskr. 48(2) : Lang, W. H. 1937. On plant remains from lhe Dowmonian of Uppsala. England and Wales. Phi/' R. Soc. Lond. ser. B 227 : 245·291. Lundblad, B. 1954. Contribulions 10 lhe geological hislOry Fageri, K. & Iversen, J. 1950. Text book oj modern pollen analysis. lhe Hepalicae. Fossil Marchamiales from lhe Rhaelic Copenhagen. of Liassic coal mines of Skromberga (Prov, of Scania) Gensel, P. G. & Andrews, H. N. 1987. The evolulion of early land planls. Am. Scient. 7S : 478·489. Sweden. Svensk. bot. Tidskrijt. 48: 381·417. Lundblad, B, 1959. On Ricciisporites tuberculatus and ilS occur· Goswami, H. K. & Khandelwal, S. 1973. Wall layers in normal and abnormal spores of Ophioglossum L. I Indian bot. Soc. S2 : rence in certain Slrata of lhe "Hollviken II" boring in S. W. 206·213. Scania, Crana Palyno/. 2 : 1·10. P. N. Gray, J. 1985. The microfossil record of early land plants: Mehra, 1974. Evolution oj spores through ages. Palynological Advances in understanding of early lerreslrialisalion­ Sociery of India, Nalional Botanic Garden, Lucknow. 1970·84. In: Chaloner, W. G. & Lawson, J. D. (eds)­ Mehra, P. N. & Sood, S. 1969. Sludies on lhe spore morphology of Evolution and environment in the Late Silurian and Early some weSl Himalayan hepalics and AnlhocerOles. Res. Bull. Devonian. Phi/' Trans. R. Soc. Lond. 3098 : 167·195. Punjab Univ. n. ser, 20 (I·II): 71·103. Gray, J., Theron, J. N. & BOUCOI,A. J. 1986. Age of lhe Cedar berg Mehra, P. N. & Sokhi,J. K. 1972. Embryology of Cyathodiumjlabe· Formalion, Soulh Africa and early land plant evolulion. Ceo/. l/atum Mehra, I Hattori bot. Lab. 36: 17·53. Mag. 123(4) : 445·454. Miller, N, G. 1984. Tertiary and Qualernary fossils. In : Schusler, Gupta, A, & Udar, R. 1986. Palyno'laXonomy of selected Indian R. M, (Ed,)-Manual oj Bryology, VoL 2, pp. 1194·1230 liverworts. Bryophytorum Bibliotheca 29: 1·201. Hallori BOlanical Laboralory, Japan. Halle, T. G. 1933. The SlrUClure of certain fossil spore bearing Miyoshi, N. 1966, Spore morphology of Hepalicae in Japan. Bull. organs believed 10 belong pleridosperms. K. Svenska Veten· Okayama College Sci 2 : 1·40. skAkad. Hand. 12(3): 1·103, Nagy, E. 1968. Moss spores in Hungarian Neogene Slrata. Hammen, T. Van der & Garcia de MUlis, C. 1965. The Paleocene Acta bot. Acad. Sci. Hung. 14(1,2) : 113·132. pollen flora of Colombia, Leid. geol, Meded. 3S : 105·116. Nayar, B. K. 1964. Palynology of modern pleridophytes. In: Nair, Harris, T. M. 1939. Naiadita, a fossil bryophyte Wilh reproduclive P. K. K. (Ed.)-Advances in Palynology, pp. 101·141. Nalio· organs. Annis. bryo/. 12: 57·70. nal Botanic Garden, Lucknow. 36 THE PALAEOBOTANIST

Nemjec, F. & Kvacek, Z. 1976. Senonian plant macrofossils from Soc. Lond. 3098: 201-205. the region of Zliv and Huboka (near Ceske Budejovice) in Richardson, J. B. & Ioannides, N. 1973. Silurian palynomorph from South Bohemia, pp. 1·83. University Karlova, Praha. the Tanezzuft and Acacus formations, Tripolitania, North Pant, D. D. 1954. Suggestions for the classification and nomen­ Africa. Micropaleontology 19 : 257-307. clature of fossil spores and pollen grains. Bot. Rev. 20 : 33-60. Samoilovitch, S. R. et al. 1961. Pollen and spores of Pant, D. D. & Srivastava, G. K. 1962. The genus Isoetes in India. Western Siberia; Jurassic to Palaeocene. VNIGRI. Tr. All Proc. natn. Institute Sci., 288(3) : 242-270. Union Petrol. Sci. Res. Geol. Expl. Inst. 177: 657 Pant, D. D. & Basu, N. 1977. Bryophyta through the ages. In : (in Russian) Current trends in life sciences (L. P. Mall Commemoration Schopf, j. M. 1949. Pteridosperm male fructifications: American volume) 9 : 65-74. species of Dolerotheca with notes regarding certain allied Pant, D. D. & Basu, N. 1978. On twO structurally preserved bryo­ forms. Report of investigations, 142, State geol. Sum JIIinois, phytes from the Triassic of Nidpur, India. Palaeobotanist25 : Ulbana. 340-349 Schuster, R. M. 1966. The Hepaticae and Anthocerotae east of the Pichova, N. G. 1967. Microfossils of Lower Cambrian and Precamb· hundredth meridian. Columbia Univ. Press, New York. rian deposits in Eastern Siberia. Rev. Palaeobot. Palynol. Schuster, R. M. 1984. New manual of Bryology, Vol. II. The 5(1-4) : 3138. Hallori bot. l3.b., Japan. pnug, H. D. & Reitz, E. 1987. Palynology in metamorphic rocks: Stewart, W. N. 1983. Palaeobotany and the evolution of plants. Indication of early land plants. Naturwissenschaften 74 : Cambridge Univ. Press. 386-387 Stout, A. B. 1918. Duplication and cohesion in the main axis of Playford, G. 1965. Plant microfossils from Triassic sediments Chichorium intybus. Mem. Brookyn bot. Gdn 1 : 480-485. near Poatina, Tasmania. j. geol. Soc. Aust. 12 : 173-210. Sullivan, H. J. & Hibbert, F. A. 1964. Tetrapterites visensis-a new Pocock, S. A. j. 1962. Microfloral analysis and age determi­ spore bearing structure from the Lower Carboniferous. nation of .strata at the Jurassic Cretaceous boundary in Palaeontology 7 : 64-71. Western Canada plains. Palaeontographica 1118 : 1·95. Taylor, T. N. 1988. The origin of land plants: Some answers, more POlOnie, R. 1960. Synopsis der Gallungen der Sporae dispersae PI. questions. Taxon 37(4) : 805·833. R. S. & Meena, K. L. 1988. Abundance spore tetrads III. Beih. geol. ]b. 39 : 1·189. '~Tiwari, of in POlOnie, R. 1966. Synopsis der Gallungen der Sporae dispersae PI. the Early Triassic sediments of India and their significance. TV. Beih. geol. lb. 72 : 1-244. Palaeobotanist 37(2) : 210·214. POlOnie, R. 1970. Synopsis der Gallungen der Sporae dispersae Pt. Tryon, R. M. & Tryon, A. M. 1982. Fern and allied plants. Springer­ V. Beih. geol. ]b. 87 : 1-222. Verlag, New York Inc. POlOnie, R. & Kremp, G. 1954. Die Gallungen der Palaeozoischen Walton, J. 1925. Carboniferous Bryophyta I. Hepaticae. Ann. Bot Sporae dispersae und ihre stratigraphie. Beih. geol. ]b. 69 : 39 : 563·572. 111·195. Walton, J. 1928. Carboniferous Bryophyta II, Hepaticae and Musci. POlOnie, R. & Lele, K. M. 1959. Studies in the Talchir nora of Ann. Bot. 42: 707·716. India-I. Sporae dispersae from the Talchir beds of South Rewa Walton, J. 1949. A thalloid plant (d. Hepaticites sp.) showing Gondwana Basin. Palaeobotanist 8(1·2) : 22-37. evidence of growth in situ from the Coal Measures at Dollar, Remy, W. & Remy, R. 1980. Lyonophyton rbyniensis nov. gen. et Clackmannanshire. Trans. geol. Soc. Glasgow 21 : 278-280. nov. spec. ein GamelOphyt aus dem Chert von Rhynie (Unter Walton, j. 1953. An introduction to the study of fossil plants. Devon, SChollland). Argumenta Palaeobotanica 6: 37-72. Adams & Charles Black, London. Richardson, j. B. 1985. Lower Palaeozoic sporomorph: Their strati· White, D. & Stadnichenko, T. 1923. Some mother plants of petro­ leum in the Devonian Black shales. Econ. Geol. 18 : 238-252. graphical distribution and possible affinities. In: Chaloner, Wodehouse, R. P. 1935. Pollen grains. McGraw Hill Book Co., w. G. & l3.wson, j. D. (eds)-Evolution and environment in New York. the Late Silurian and Early Devonian. Phil. Trans. R.