permian plant macrofossils from fossilryggen, vestfjella...

Introduction

The distribution of fossil Glossopteris leaves on disparatesouthern lands represented key evidence in early argumentsto support plate tectonics and the notion of a formersupercontinent: Gondwana. The discovery of Glossopterisleaves on Mount Buckley at 85°S by Robert Scott's1910–13 Antarctic expedition (Seward 1914) confirmedthat Antarctica hosted this widespread SouthernHemisphere taxon and that during the Permian, robustvegetation was able to survive close to the South Pole. Since1912, glossopterids and associated Permian plants havebeen recorded from numerous localities in theTransantarctic Mountains, Antarctica (Edwards 1928,Cridland 1963, Rigby & Schopf 1969, Schopf 1970,Lambrecht et al. 1972, Lacey & Lucas 1981, Taylor et al.1992, Cúneo et al. 1993), the Ellsworth Mountains (Gee1989, Taylor et al. 1989), and the Prince Charles Mountains(White 1969, Neish et al. 1993, McLoughlin et al. 1997).

Currently, the only plant macrofossils described andillustrated from Dronning Maud Land are those collectedfrom three beds exposed in Milorgfjella, the north-easternextension of Heimefrontfjella, by the British AntarcticSurvey in the late 1960s (Juckes 1972, Plumstead 1975).The Milorgfjella assemblages are dominated byglossopterids in which the central veins are poorlyaggregated (representing the ‘Gangamopteris condition’).Other Milorgfjella fossils include a conifer twig,sphenophyte axes, possible lycophyte axes and fragmentaryfern fronds, and various gymnosperm axes including somewith attached spur shoots (Table I). These assemblages aretypical of low-diversity Early Permian floras from southern

Gondwana. Recent palynological studies have supported anEarly Permian age for the Milorgfjella succession (Larssonet al. 1990, Lindström 1995b). Hjelle & Winsnes (1972)first reported the presence of similar fossil plants atFossilryggen in Vestfjella and, based on photographs of thematerial, Dr E.P. Plumstead suggested that the fossilsrepresented an Early Permian assemblage, althoughyounger than that preserved at Milorgfjella. However, morerecent palynological studies (Lindström 1995a, 1996) havesuggested a Middle Permian age for at least some of thesedimentary rocks at Fossilryggen. The new assemblagefrom the northern exposures of Fossilryggen represents thefirst Middle Permian macrofossil flora described fromDronning Maud Land.

Geological setting

The scattered ridges and isolated nunataks of the Vestfjellamountain range are situated along the inner part of the iceshelf embracing the western extent of Dronning MaudLand. Nunataks in this region predominantly exposeJurassic basalts and dolerite intrusions but olivine-gabbrodominates the southern part of the mountain range.Fossilryggen is situated in the north-eastern part of themountain range and is the only locality where Palaeozoicsedimentary rocks crop out. The geology of the area hasbeen described by Hjelle & Winsnes (1972), Olaussen(1985), Larsson (1990, 1991) and Lindström (1995a, 1996),and it will only be summarized briefly herein.

The approximately 2 km long, low and narrow ridge ofFossilryggen is the main nunatak in the area (Fig. 1). It is

Antarctic Science 17 (1): 73–86 (2005) © Antarctic Science Ltd Printed in the UK DOI: 10.1017/S0954102005002464

73

Permian plant macrofossils from Fossilryggen, Vestfjella,Dronning Maud Land

STEPHEN MCLOUGHLIN1*, KENT LARSSON2 and SOFIE LINDSTRÖM2

1School of Natural Resource Sciences, Queensland University of Technology, PO Box 2434, Brisbane, QLD 4001, Australia2GeoBiosphere Science Centre, Lund University, Sölvegatan 12, SE-223 62 Lund, Sweden

*[email protected]

Abstract: A low diversity plant macrofossil assemblage described from the northern section ofFossilryggen, Vestfjella, Dronning Maud Land, Antarctica, is dominated by matted leaf impressions ofGlossopteris sp. cf. G. communis Feistmantel and Glossopteris sp. cf. G. spatulata Pant & Singh, and in situfinely branched Vertebraria indica Royle rootlets. Equisetalean stems, rhizomes and leaf whorls(Phyllotheca australis Brongniart emend. Townrow), isolated seeds, scale-leaves, and fragmentarygymnosperm axes represent minor components of the assemblage. The fossils are preserved in fine-grained,floodbasin sediments and dark palaeosols. Although lacking definitive biostratigraphical indices, the flora isconsidered to represent a Middle Permian assemblage based principally on lithological and palynologicalcorrelation with the southern section at Fossilryggen and broad similarities to mid-Permian plantassemblages elsewhere in Gondwana.

Received 2 January 2004, accepted 23 August 2004

Key words: East Antarctica, Glossopteris, Middle Permian, palynostratigraphy, Phyllotheca, Vertebraria

flanked to the west by a smaller nunatak, the NW Nunatak(informal name), and to the north-west by three smalloutliers (Fig. 1). Palaeozoic sedimentary rocks crop out nearthe southern and northern limits of Fossilryggen, in the"NW Nunatak" and in the middle outlier (Fig. 1). Thesouthern section at Fossilryggen hosts the most completesedimentary succession where approximately 60 m ofsandstones and shales are exposed. The southern section istransected by six dolerite dykes and one normal fault, whichdownfaults the northern portion of the outcrop. The centraland northern parts of Fossilryggen consist mainly ofbasaltic rocks, although black shales are exposed intopographical lows. The north-western margin of the ridgeconsists of a 30 m thick sequence of plant-rich black shales,minor sandstones and mudstones, here called the ‘northernsection’.

Due to the intervening basaltic rocks, the relationshipbetween the southern and northern sections at Fossilryggenis not clear. The northern section is situated topographicallylower than the southern section (Lindström 1995a, textfig. 2), but since the sedimentary strata at Fossilryggen dipslightly towards the north-west it is possible that thenorthern section is somewhat younger than the southernone.

Material and methods

The material described herein was collected from thenorthern section of Fossilryggen and the adjacent NorthwestNunatak by Professor Kent Larsson during the SwedishAntarctic Research Programme (SWEDARP) expeditions

to the area in 1988/89 and 1989/90. Macrofossils arepreserved as impressions in dark siltstone. Specimens werephotographed under strong unilateral light from the upperleft. All illustrated specimens (prefixed LO) are lodged inthe collections of the Geology Department, GeoBiosphereScience Centre, Lund University. In the systematicdescriptions OD = original designation, M = by monotype,SD = subsequent designation.

Palynology

Lindström (1994, 1995a, 1996) recognized sixty-six taxaincluding eight algae or acritarch species when analysingthe palynostratigraphy of the southern sedimentary sectionat Fossilryggen and nearby "NW Nunatak". Theassemblages also included thirty-five gymnosperm pollentaxa, seventeen fern spores, four lycophyte species, and tworegarded as sphenophytes (Lindström 1996). Theassemblages are dominated by bisaccate pollen, includingboth taeniate forms (viz. Protohaploxypinus amplus,P. limpidus, Striatopodocarpidites cancellatus and S. fusus)and non-taeniate species (viz. Scheuringipollenitesmaximus, S. ovatus and Alisporites spp.). Fern spores arecommon, especially Leiotriletes directus, Horriditriletestereteangulatus, H. filiformis and Osmundaciditeswellmanii. Didecitriletes ericianus, D. uncinatus,Microbaculispora villosa, Indospora laevigata andDictyotriletes labyrinthicus appear in the upper part of thesouthern section, together with Guttulapolleniteshannonicus and Protohaploxypinus rugatus (Lindström1996).

74 STEPHEN MCLOUGHLIN et al.

Table I. Revised identifications of Lower Permian plant fossils collected by the British Antarctic Survey from Milorgfjella, Dronning Maud Land andoriginally described by Plumstead (1975).

Taxon identified by Plumstead (1975) from Milorgfjella Illustrations of Plumstead (1975) Revised identification

Lycopodiopsis pedroanus (Carr.) Edwards pl. Id Possibly a lycophyte or conifer axisParacalamites sp. cf. Phyllotheca australis Brong. pl. Ia (1), Ib Paracalamites australis RigbyIndeterminate small branched stem pl. Ic Possibly a fern frond fragmentStem with small round scars pl. Ia (2) Unidentifiable axisHollow mound with grooved spiral markings pl. Ie Unidentifiable axiscf. Cordaitales pl. IIa–c, IIIa, IIId Unidentifiable gymnosperm axescf. Glossopteridae pl. IIa, IIIb, IIIc, VIIa, VIIIa, Unidentifiable gymnosperm axescf. Ginkgophyta long and short shoots pl. Vd–g, VIa–d Glossopterid long shoots bearing lateral spur shootscf. Coniferophyta pl. VIIb, VIIc Unidentifiable gymnosperm axesWood fragments pl. IVa–g Gymnosperm xylem tracheids with bordered pitsGangamopteris obovata Carr. pl. IXa–d Gangamopteris sp. cf. G. obovata (Carruthers) WhiteGangamopteris cyclopteroides Feist. pl. Xa–d Gangamopteris sp. cf. G. spatulata McCoyGangamopteris sp. pl. XIVa (top) Gangamopteris obovata Carr.Walikalia juckesii Plumstead pl. XIa–f Gangamopteris sp. cf. G. spatulata McCoyEuryphyllum antarcticum Plumstead pl. XIIa–d, XIIIa–b Gangamopteris sp. cf. G. angustifolia (McCoy) McCoyEuryphyllum sp. pl. XIVc–d Glossopteris sp. cf. G. douglasii Rigby & ChandraNoeggerathiopsis hislopi (Bunb.) Feist. pl. XIVe Gangamopteris sp. cf. G. obovata (Carruthers) WhiteNoeggerathiopsis spathulata (?) Dana pl. XIVf–g Gangamopteris sp. cf. G. obovata (Carruthers) Whitecf. Palaeovittaria sp. pl. XIVh Gangamopteris sp.Coniferous twig pl. XIVa–b cf. Walkomiella transvaalensis le RouxMixed foliage pl. XV Gangamopteris sp. cf. G. obovata (Carruthers) WhiteFungi pl. IXc Indeterminate imprints and weathering patterns

Lindström (1996) suggested that the successive firstappearances of taxa within the southern section atFossilryggen and the "NW Nunatak" were more related topreservational factors than age differences. Thepalynofloras were, therefore, correlated with AustralianUpper Stage 5 (late Wordian–Lopingian) microfloras(Lindström 1996, Foster & Archbold 2001). However, ifone considers the possibility that the successive firstappearances of taxa actually reflect age differences withinthe southern section, it is possible that the upper part of thesection is equivalent to Australian Lower Stage 5b/c (lateRoadian–early Wordian) microfloras (Lindström 1995a,Foster & Archbold 2001), the lower part being somewhatolder. The co-appearance of Microbaculispora villosa andDidecitriletes ericianus is a pattern that differs from thetypical Australian palynostratigraphic succession. In theKaroo Basin, South Africa, which during the Permian was

juxtaposed to Dronning Maud Land, these taxa also appearat the same level (Anderson 1977). In the Prince CharlesMountains M. villosa first appears at a slightly lower levelthan D. ericianus (Lindström, personal observation), andthis is also the pattern in India (Tiwari & Tripathi 1992).

Lindström (1996) found no identifiable palynomorphs insamples from the northern section at Fossilryggen. For thisstudy, the samples were reprocessed according to standardpalynological methods. One sample, F90-83, contains a fewidentifiable palynomorphs but they are generallyfragmentary, and of high thermal maturity, black tobrownish-black even after treatment with Schultze’ssolution. The following taxa have been identified:Horriditriletes tereteangulatus, Lunatisporites sp.,Osmundacidites wellmanii, Protohaploxypinus amplus andScheuringipollenites maximus. These few taxa areunfortunately not sufficient to date the northern section

VESTFJELLA PERMIAN PLANT FOSSILS 75

Fig. 1. Locality details of the fossilassemblage: a. Map of Antarcticashowing Dronning Maud Land, b. Mapof Dronning Maud Land showing theposition of Fossilryggen, c. Map ofFossilryggen, Vestfjella, showing thesource of the described fossilassemblage (after Larsson 1991).

more precisely than to the Permian.

Systematic palaeobotany

Phylum SphenophytaClass SphenopsidaOrder Equisetales

Family Gondwanostachyaceae Meyen 1969Phyllotheca Brongniart emend. Townrow 1955

Type species: Phyllotheca australis Brongniart emend.Townrow 1955 (OD/M); ?Newcastle Coal Measures; Upper

Permian; Hawkesbury River area, northern Sydney Basin,Australia.

Phyllotheca australis Brongniart emend. Townrow 1955Fig. 2a–g

Lectotype: No. 3388, Museum National d'HistoireNaturelle, Paris (Townrow 1955, text fig. 1a).

Specimens: LO 9379t, LO 9380t, LO 9382t, LO 9383t, LO9390t, and one unfigured specimen.

Diagnosis: See Townrow (1955, pp. 42–43).


Fig. 2. Phyllotheca australis Brongniart emend. Townrow 1955. a. b. [LO 9379t], c. [LO 9380t]: Longitudinally ribbed, segmented axes. d. [LO 9381t]: Longitudinally ribbed axis with fine roots emerging from a node, e. g. [LO 9382t], f. [LO 9383t]: Whorls or partially fused,linear leaves attached to slender segmented axes (e) or detached (f. g.). Scale bars = 10 mm for a–f; 5 mm for g.

Description: Material assigned to this species includesfragmentary rhizomes bearing fine roots (Fig. 2a, b, d),slender aerial stems bearing leaves at nodes (Fig. 2c, e, f),and detached leaf whorls (Fig. 2g). Axes are preservedpredominantly parallel to bedding. Rhizomes reach > 260 mm long and up to 16 mm wide with 15 mm long and< 0.5 mm wide roots emerging from nodes spaced up to

50 mm apart. External longitudinal ridges and grooves areindistinct on root-bearing axes; ridges are opposite acrossnodes. Leaf-bearing axes are generally narrower (< 4 mmwide) and shorter (longest available fragment 30 mm long)than root-bearing axes. They bear longitudinally striateridges 0.5–1 mm apart. Nodes on these axes are 4–8 mmapart and bear whorls of around 18 linear leaves that are


Fig. 3. Glossopteris sp. cf. G. communis Feistmantel 1879. a. Falcate-linear leaf [LO 9384t]. b. Linear leaf with well-developed petiole [LO 9385t]. c. Venation details of very small, linear leaf [LO 9386t]. d. Very narrow obovate-linear leaf [LO 9387t]. e. Venation details ofan intermediate-sized leaf [LO 9388t]. f. Enlargement of the petiole of a small leaf [LO 9385t]. g. Intermediate-size, narrowly obovate leaf[LO 9389t]. h. Intermediate-size narrowly obovate leaf [LO 9390t]. i. Large leaf [LO 9391t]. j. Central portion of large leaf [LO 9392t]. k. Enlargement of intermediate-size leaf with a slightly emarginate apex [LO 9389t]. l. Enlargement of the base of a large leaf [LO 9393t].Scale bars = 10 mm.

fused to form a proximal sheath for about one-third to two-thirds of their length (Fig. 2e & g). The 3–7 mm long sheathis closely appressed to the stem (Fig. 2f). Distally, the leavesdiverge with a free length of 5–8 mm. Leaves are0.5–0.75 mm wide at the limit of fusion and apices arepointed acute.

Comments: Isolated, leafless, segmented axes from thePermian of Gondwana with longitudinal ribs arranged inopposition at the nodes may belong to a range ofsphenophyte families and most have been assigned to theform species Paracalamites australis Rigby (1966).However, some Vestfjella axes bear leaf whorls (Fig. 2e)traditionally assigned to Phyllotheca australis Brongniartemend. Townrow 1955 in the Gondwanostachyaceae(Meyen 1969). Based on comparisons to extant Equisetumspecies, these plants are assumed to have been rhizomatous,herbaceous, and restricted to relatively moist habitats.

Geographic and stratigraphical range: Commonthroughout the Permian of Gondwana (Pant & Kidwai1968); most abundant in Middle to Upper Permian strata(Anderson & Anderson 1985).

Phylum ?ConiferophytaOrder Glossopteridales sensu Pant 1982

Glossopteris Brongniart 1830

Type species: Glossopteris browniana (Brongniart)Brongniart 1830 (SD of Brongniart 1830); ?Newcastle CoalMeasures; Upper Permian; Gulgong-Maitland districts,Sydney Basin, Australia.

Glossopteris sp. cf. G. communis Feistmantel 1879Figs. 3a–l

Specimens: LO 9384t, LO 9385t, LO 9386t, LO 9387t, LO 9388t, LO 9389t, LO 9390t, LO 9391t, LO 9392t, LO 9417t, and ten unfigured specimens.

Description: Leaves oblanceolate to linear (length:width =3.5:1 to 10:1), typically 60–115 mm long (possibly up to150 mm), 6–42 mm wide (typically 17–25 mm wide). Thewidest point of the leaf is three-quarters to four-fifths alongits length. The leaves have a gently tapering cuneate basewith a 3–4 mm wide truncate abscission point (Fig. 3a, b, d,f, g, l). In some cases the leaf base may be slightly flared atthe abscission point. The apex is generally rounded but in afew specimens it is slightly emarginate (Fig. 3k); the marginis otherwise entire. The midrib varies from indistinct toprominent, around 3 mm wide at the base, taperingthroughout, persisting to the apex where it is < 0.5 mmthick, or in smaller forms evanescing in the distal 5 mm ofthe leaf (Fig. 3c). The veins emerge from the midrib at10–20° and arch gently and continuously throughout thelamina (Fig. 3e, h, l). Venation angles midway between themidrib and margin are 30–50° near the base and centre ofthe leaf but reduce to 20–25° towards the apex. The

marginal venation angle reaches 60–70° near the base andcentre of the leaf but reduces to 40–50° towards the apex.Vein density is around 21–25 per 10 mm at the margin in thecentral part of the leaf.

Comments: This species shows substantial variation in formfrom small, narrow, linear leaves (Fig. 3a–g) to moderatelylarge, oblanceolate leaves (Fig. 3h–l). Despite this broadrange in shape and size the leaves are considered toconstitute a single species because they are represented by amorphological continuum. Small linear forms mayrepresent young, incompletely expanded or exposed(canopy) leaves. Broad forms may represent mature shadeleaves. All are characterized by continuously arched,relatively dense, anastomosing secondary veins with mid-leaf marginal venation angles of 60–70° and narrowlycrescentic areolae.

Three Indian Glossopteris species are closely comparableto these Antarctic specimens. Glossopteris communisFeistmantel (1879) from the Lower Permian KarharbariFormation is the most similar in shape and venation style.The larger Antarctic specimens are particularly similar tothe type specimen of G. communis; smaller specimens tendto have a much greater length:width ratio and fall outsidethe range traditionally recognized for the Indian species(Chandra & Surange 1979). Glossopteris intermittensFeistmantel 1881 from the Lower to Middle PermianBarakar Formation is comparable in size and venationcurvature to Glossopteris sp. cf. G. communis but the formertypically has a length:width ratio of only 3:1 and sparsesecondary vein anastomoses (Chandra & Surange 1979).Glossopteris tenuifolia Pant & Gupta 1968 from the UpperPermian Raniganj Formation has a shape and size moreclosely comparable to the narrower examples of theAntarctic leaves but it tends to have straighter secondaryveins intersecting the margin at < 45°.

Lower Permian leaves from the Karoo Basin attributed byAnderson & Anderson (1985) to Hirsutum intermittensPlumstead, Arberia madagascariensis (Appert) Anderson &Anderson, and A. leeukuilensis Anderson & Anderson havesimilar venation styles to Glossopteris sp. cf. G. communisFeistmantel but these South African forms are typicallymuch larger (160–200 mm long) with acute, pointed apices.Amongst other South African forms, the Fossilryggenspecimens are closely comparable to Upper Permian sterileleaves from the Karoo Basin attributed by Anderson &Anderson (1985) to Lidgettonia africana Thomas. The latterare oblanceolate, 80–150 mm long, with narrowlycrescentic areolae, vein marginal angles of around 50°, andin some cases an emarginate apex. In the absence ofcuticular information or attached reproductive structures theFossilryggen specimens and L. africana leaves appear todiffer only in their degree of vein curvature. Amongsteastern Australian Glossopteris leaves, the Upper PermianG. curvinervosa McLoughlin (1994a) has the most similar


venation style but it tends to have a more acute apex that theFossilryggen leaves.

Geographic and stratigraphic range: Glossopteriscommunis sensu stricto is apparently restricted to the

Karharbari Formation (Lower Permian) of north-easternpeninsular India (Chandra & Surange 1979) but numeroussuperficially similar leaves have been reported throughoutGondwana from the Lower, Middle and Upper Permian(Lacey et al. 1975, Rigby et al. 1980, Rayner & Coventry


Fig. 4. Glossopteris sp. cf. G. spatulata Pant & Singh 1971. a. Leaf with abbreviated base [LO 9394t]. b. Large leaf with a short, tapered,base [LO 9395t]. c. Central portion of a leaf [LO 9396t]. d. Leaf with ill-defined, predominantly straight, secondary veins [LO 9397t]. e. Details of venation near the leaf apex [LO 9398t]. f. High-angle secondary venation from the central region of a large leaf [LO 9399t].g. Enlargement of the central portion of a leaf showing predominantly straight secondary veins in the outer part of the lamina [LO 9394t].h. Enlargement of the central portion of a leaf showing the venation pattern [LO 9400t]. Scale bars = 10 mm.

1985). Determination of the relationships between thesewidely separated fossils must await cuticular details orattached reproductive structures.

Glossopteris sp. cf. G. spatulata Pant & Singh 1971

Figs. 4a–h

Specimens: LO 9394t, LO 9395t, LO 9396t, LO 9397t, LO9398t, LO 9399t, LO 9400t, and one unfigured specimen.

Description: Leaves narrowly elliptic to narrowlyoblanceolate (length:width = 3:1 to 4.5:1), reaching around170–200 mm long and up to 70 mm wide (Fig. 4a–d). Thewidest point is two-thirds to three-quarters of the distancefrom base to apex. The margin is entire, the apex broadlyrounded and the base cuneate (Fig. 4a, b, e). The midrib isrobust at the base, up to 4 mm wide, tapering throughout butpersisting to the apex where it is < 0.5 mm wide. The veinsemerge from the midrib at 10–20°, arch gently in the innerhalf of the lamina but only slightly in the outer part

(Fig. 4f–h). The mid-lamina venation angle is 50–65° nearthe base and centre of the leaf and reduces to 25–30° nearthe apex. The marginal venation angle reaches 65–70° nearthe base and centre of the leaf but reduces to 30°–40°towards the apex. Areolae are very narrowly crescentic tolinear. The marginal venation density is 20–28 per 10 mm inthe central part of the leaf.

Comments: Glossopteris sp. cf. G. spatulata Pant & Singh(1971) represents leaves that are typically twice as wide andsubstantially longer than Glossopteris sp. cf. G. communisfrom the same locality. Additionally, the secondary veins ofGlossopteris sp. cf. G. spatulata tend to arch near the midriband have a relatively straight course in the outer part of thelamina (Fig. 4e–h). In this respect, Glossopteris sp. cf. G. spatulata is similar to G. acutifolia McLoughlin (1994a)and G. kingii McLoughlin (1994b) from the Upper Permianof eastern Australia. However, the Australian species havedifferent suites of accessory morphological characters


Fig. 5. Vertebraria indica Royle 1834; Root impressions. a. Large root with two rows of chamber infillings [LO 9401t], b. Large root with asingle, central row of chamber infillings flanked by featureless secondary wood impressions [LO 9402t], c. Intermediate-size root with alarge lateral branch [LO 9403t], d. Intermediate-size root with three longitudinal rows of chamber infillings [LO 9404t], e. Small rootsshowing minimal segmentation, giving off ultimate rootlets at nodes [LO 9405t], f. Intermediate and fine rootlets with multiple orders ofbranching [LO 9406t], g. Intermediate-size, bedding-parallel, root giving off several orders of lateral roots [LO 9407t]. Scale bars = 10 mm.

including smaller size, more acute apices or a lessermarginal venation angle. Large leaves from the AustralianUpper Permian [e.g. G. ampla Dana (1849) and G. grandisMcLoughlin (1994a)] tend to have broader areolae and lessdense venation.

The Antarctic leaves are closest in shape, size andvenation pattern to Glossopteris spatulata from the UpperPermian Raniganj Formation of India (Chandra & Surange1979). However, Pant & Singh (1971) erected G. spatulataon both macromorphological and cuticular characters. Asthe Antarctic specimens lack cuticle it is unclear whetherthey are conspecific with the Indian forms. Glossopterisfeistmantelii Rigby, reported from the Upper Permian ofIndia also has a similar venation style but tends to be aproportionately broader leaf (length:width = 2:1) with acordate base (Rigby 1964, Chandra & Surange 1979).

Some South African Lower Permian Glossopteris leavesassociated with Ottokaria buriadica Plumsteadsuperficially resemble Glossopteris sp. cf. G. spatulata butthey have a truncate base and lack a well-defined midrib(Plumstead 1956, Anderson & Anderson 1985). MiddlePermian leaves co-preserved with Arberia allweyensisAnderson & Anderson fruits are distinguished by theirtapered acute apices. Upper Permian leaves from the easternKaroo Basin associated with Lidgettonia mooiriverensisAnderson & Anderson share many features withGlossopteris sp. cf. G. spatulata. They differ only in havinga more attenuate (petiolate) base. As no fruits were foundattached to, or associated with, Glossopteris sp. cf. G. spatulata at Fossilryggen, an affinity with Lidgettoniacan not be confirmed. The Fossilryggen leaves are alsostrikingly similar to specimens assigned to G. communisfrom the Lower Permian of Madagascar, although the lattermay reach 300 mm long and 100 mm wide (Appert 1977).

Geographic and stratigraphic range: Glossopterisspatulata is notionally restricted to the Raniganj Formation(Upper Permian) of north-eastern peninsular India (Chandra& Surange, 1979). The closely comparable forms fromFossilryggen have not been reported elsewhere. Theirrelationship to G. spatulata sensu stricto can not bedetermined until cuticular details become available.

Vertebraria Royle emend. Schopf 1982

Type species: Vertebraria indica Royle 1834 (M); RaniganjFormation; Upper Permian; Raniganj Coalfield, DamodarValley, India.

Vertebraria indica Royle 1834Fig. 5a–g

Holotype: V4189, Natural History Museum, London.

Specimens : LO 9401t, LO 9402t, LO 9403t, LO 9404t, LO9405t, LO 9406t, LO 9407t, and four unfigured specimens.

Diagnosis: See Schopf (1982, p. 41).

Description: Root impressions reaching in excess of 270 mm long and up to 38 mm wide, with one to four rowsof rectangular, alternating segments (Fig. 5a–d). The largestspecimens show segments restricted to the central part ofthe root; the outer parts, up to 10 mm wide, are relativelysmooth or possess fine longitudinal striae or transverse cleatmarkings (Fig. 5b). Roots show up to three orders ofbranching. Lateral rootlets emerge (typically at 20–45°)from the transverse partitions in larger roots (Fig. 5f & g).The ultimate rootlets are typically 2–4 mm long, 1–1.5 mmwide, with a single vascular strand (Fig. 5e & f). Roots areorientated predominantly parallel to, or at low angle to,bedding.

Comments: Gould (1975) demonstrated a completeanatomical transition between permineralized young rootsof Vertebraria indica and mature root or stem woodreferable to Araucarioxylon. The smooth or longitudinallystriate outer region in the larger root impressions in theFossilryggen assemblage (Fig. 5b) probably represents azone of continuous secondary xylem forming completegrowth rings around the inner (segmented) zone withschizogenous cavities. Pant (1958) assigned very small,univeined rootlets from a Permian, glossopterid-rich,assemblage of Tanzania to Lithorhiza Pant but such rootletsare clearly connected to larger, segmented/chambered rootsreferable to Vertebraria indica (Fig. 5e & f) and theirtaxonomic differentiation seems superfluous. Additionally,Neish et al. (1993) showed a wide variation in the anatomyof Vertebraria roots, especially in the ontogeneticdevelopment and number of schizogenous cavities, whichwould likely give rise to variably segmented or non-segmented forms when preserved as impressions/compressions.

Vertebraria indica roots are preserved in situ within flat-laminated, dark siltstones, and clearly display several ordersof delicate branching. In a few cases they transect beddingplanes at a low angle but they are predominantly orientatedparallel to bedding suggesting that the parent plantspossessed relatively shallow but broadly spreading rootsystems. Root-rich beds typically contain few leaves andprobably represent organic-rich, immature palaeosols infloodbasin environments with a high water table.

Geographic and stratigraphic range: Common throughoutthe Permian of Gondwana.

Glossopterid scale leavesFigs. 6a–f

Specimens: LO 9408t, LO 9409t, LO 9410t, LO 9411t, LO 9412t, LO 9413t, and one unfigured specimen.

Description: Detached, elliptical-rhombic, concavo-convex,entire-margined, scale-leaves that are 12–18 mm wide and15–19 mm long, including a 2–3 mm long acuminate apex(Fig. 6a, d, f). A single small (6 mm wide, > 8 mm long)


specimen (Fig. 6e) falls outside this range. Althoughtypically poorly preserved, the bases of all scales areapparently truncate (broadly attached). Several veinsemerge from the scale base and gently diverge and archacross the lamina. The veins dichotomize and anastomosethroughout the lamina to maintain a density of around 24per 10 mm (measured across the lamina); no distinct midribis developed.

Comments: Isolated scale leaves of this type may representdetached sterile bracts originally protecting the dormant tipsof branches or they may constitute the foliar component ofmicrosporangiate glossopterid fruits such as Eretmonia duToit emend. Lacey, van Dijk & Gordon-Gray (White 1978,Anderson & Anderson 1985, Holmes 1995). Both sterileand fertile scale leaves typically show a broad variation insize and shape. The Vestfjella bracts do not bear sporangia.


Fig. 6. a–f. Rhombic glossopterid scale leaves with gently divergent reticulate venation [a. LO 9408t, b. LO 9409t, c. LO 9410t, d. LO 9411t, e. LO 9412t; f. LO 9413t]. g–j. Samaropsis sp. [g. LO 9414t, h. LO 9415t, i. LO 9416t, j. LO 9417t]. k–n. Rotundocarpus sp.cf. R. ovatus Maithy 1965 [k. LO 9418t, l. LO 9419t, m. LO 9420t, n. LO 9421t]. Scale bar = 5 mm for a–f; 1 mm for g–n.

Such bracts with very few distinctive morphologicalcharacters have traditionally been assigned to informalcategories such as 'scale leaves' or 'squamae' or, in somecases, to established form genera (Chandra & Surange1977a, 1977b, Banerjee 1984). It is likely that scale leavesproduced by any glossopterid plant ranged from entirelysterile, functionally protective bud scales to stronglyexpanded microsporophylls (White 1978). Additionally,some Gondwanan scale leaves show evidence of eitherbasal or apical expansion transitional with normalvegetative leaves (Anderson & Anderson 1985, Holmes1995). Consequently, it is difficult to confidently assignthese fossils to any established taxon in the absence ofattached reproductive structures.

Geographic and stratigraphic range: Sterile glossopteridscale leaves are common throughout the Permian ofGondwana but are especially abundant in Upper Permianstrata (Anderson & Anderson 1985).

Dispersed seedsSamaropsis Goeppert 1864

Type species: Samaropsis ulmiformis Goeppert 1864 (OD);rock unit uncertain; Permian; Braunau, Bohemia, CzechRepublic.

Samaropsis sp.

Figs. 6g–j

Specimens: LO 9414t, LO 9415t, LO 9416t, and oneunfigured specimen.

Description: Circular to broadly elliptical seeds 7–8 mmlong, 6–7 mm wide. The central body is generally indistinct,flattened, elliptical to ovate, 7–8 mm long and 3–4.5 mmwide (Fig. 6g, i, j), but it is slightly raised on one impression(Fig. 6h). The central body is flanked by crescentic marginalwings 1–2 mm wide that are, in some cases, extended intodeep lobes adjacent to the chalazal end of the seed (Fig. 6h).The chalazal end of the central body is rounded to truncate;the micropylar end is rounded. The central body bearsindistinct, longitudinal striations; the wings are generallyfeatureless. All seeds are detached from other organs.

Comments: Seeds of this type are probably attributable toglossopterids based on the lack of other gymnosperm fossilsin the assemblage. Seeds associated with glossopterids inother parts of Gondwana generally fall into three sizecategories. Very small seeds (< 5 mm long) have been foundattached to capitate fruits such as Dictyopteridium,Senotheca, and Plumsteadia (Banerjee 1969, Surange &Chandra 1973, Chandra & Surange 1976, Benecke 1976,Rigby 1978). Medium-sized seeds (5–15 mm long) havebeen found attached to, or associated with, a broad array ofcupulate and loosely branched fruits such as Arberia,Rigbya, Cometia and Lidgettonia (Rigby 1972, Lacey et al.1975, Anderson & Anderson 1985, McLoughlin 1990).

Large seeds (15–80 mm long) have not been found attachedto glossopterid fruits but they are likely affiliates based oncommon association. Samaropsis sp. from Fossilryggenfalls into the medium-sized category of seeds. Most fruitsassociated with seeds of this size category have Middle toLate Permian ranges. The exception is Arberia, whichmostly ranges from Early to earliest Middle Permian.However, attached Arberia seeds lack strongly developedwings (Rigby 1972, Appert 1977) and the seed body isusually much longer than wide (ovate to narrowlyelliptical).

Seeds attached to Lidgettonia lidgettonioides (Lacey, vanDijk & Gordon-Gray) Anderson & Anderson 1985 from theUpper Permian of the Karoo Basin, South Africa, resemblethe Fossilryggen Samaropsis species in shape, dimensionsand surface features but the proximal lobes of the wings areless developed in the former. Samaropsis rigbyi Millan1977 from the Tubarão Group (Early Permian), Brazil, hassimilar broad wings with prominent proximal lobes but it istypically twice the size of the Fossilryggen seeds. The fewmorphological characters available from platyspermic seedimpressions prohibit further comparisons.

Geographic range: Not known beyond Fossilryggen.

Rotundocarpus Maithy 1965

Type species: Rotundocarpus striatus Maithy 1965 (OD);Karharbari Formation; Lower Permian; Jogtiabad Pit,Karharbari Colliery, Giridih Coalfield, India.

Rotundocarpus sp. cf. R. ovatus Maithy 1965Fig. 6k–n

Specimens: LO 9418t, LO 9419t, LO 9420t, LO 9421t, andtwo unfigured specimens.

Description: Very small, ovate to elliptical seed, 3 mm longand 2 mm wide (Fig. 6k–n). The seeds are preserved asconvex impressions or, in some cases, the centre of the seed(nucellus), or the seed as whole, may have decayed andbeen filled with fine sediment to produce an ovate toelliptical cast. These seeds either lack a lateral wing or thewing is extremely reduced. A narrow (0.1 mm) indentationin the sediment is evident around some specimens. Theouter surface bears fine longitudinal striations but the seedsare otherwise featureless.

Comments: These seeds are relatively common in beds atFossilryggen in association with Glossopteris leaffragments, Samaropsis sp., and wood fragments, but theymay be easily overlooked owing to their diminutive size.The dearth of diagnostic features or attached fruits prohibitsconclusive determination of their affinities but they aresimilar in size, shape, and in the lack of an obvious wing, toRotundocarpus ovatus Maithy 1965 from the KarharbariFormation (Lower Permian), Giridih Coalfield, DamodarValley, India. A few other Gondwanan Palaeozoic fossils,


e.g. Carpolithus circularis Walkom 1935 from the UpperPermian of eastern Australia, appear to be similar-sizedradiospermic seeds but they generally differ in havingextended micropyles or ridged ornamentation.

Geographic and stratigraphic range: Not known beyondFossilryggen.

Gymnosperm axesFig. 7b & c

Specimens: LO 9423t, LO 9422t.

Comments: Several axis impressions are present in theFossilryggen assemblage. They reach > 140 mm long andshow either discontinuous longitudinal striations (Fig. 7c)or irregular cleat patterns (Fig. 7b). Their size suggests thatthey are derived from gymnosperms but they lack additionaldiagnostic features.

Discussion

The macroflora from Fossilryggen represents a typical low-diversity, high-latitude, Gondwanan, Permian plantassemblage. Only two species of glossopterid foliage arerepresented and neither can be confidently attributed to anestablished species. Glossopterids have conservative root

morphology and it is likely that both Glossopteris specieswere borne on plants possessing roots of Vertebraria indicatype. Similarly, protective scale leaves and micro-sporophylls have relatively conservative morphologiesamongst glossopterids, differing mainly in size and apexshape. Previous schemes to generically segregateglossopterid scale leaves have not been widely acceptedoutside the immediate study area (Chandra & Surange1977b, Banerjee 1984, Anderson & Anderson 1985). Allorgans in the Vestfjella assemblage are dissociated, hence itis not possible to link the scale leaves or dispersed seedswith either of the Glossopteris species.

None of the Fossilryggen taxa provides a precise agecontrol on the host rocks. The fossils have morphologicalsimilarities to both Early and Late Permian species fromother parts of Gondwana and this may be consistent with anintermediate (Middle Permian) age deduced frompalynoassemblages in the adjacent (southern section)sedimentary rocks at Fossilryggen (= early Late Permian ofthe traditional two-part Permian timescale used byLindström 1995a). The Fossilryggen assemblage lacksGangamopteris leaves that are typical of Early Permianfloras. Such leaves are well-represented in the nearbyMilorgfjella flora (Plumstead 1975, Table I).

Phyllotheca australis, the only non-gymnospermous plant


Fig. 7a. Matted Glossopteris sp. cf. G. communis Feistmantel 1879 leaves on a bedding plane [LO 9387t]. b. Irregular axis fragment withremnant conjugate cleat sets [LO 9422t]. c. Large gymnosperm axis impression with discontinuous longitudinal striae [LO 9423t]. Scalebars = 10 mm.

in the assemblage, is a common element in GondwananPermian assemblages. This plant probably occupiedunderstorey niches in consistently wet habitats based on thesmall stature of the Permian forms and their requirement formoist conditions to enable gamete exchange. Glossopterissp. cf. G. communis is locally represented by denseaccumulations of foliage (Fig. 7a) that possibly representmats of deciduous leaves. Glossopterids are generallyconsidered to have been large trees occupying floodbasinsettings and they provided the principal organic constituentsof Gondwanan Permian coals (Gould & Delevoryas 1977,Retallack & Dilcher 1988). The segmented roots ofglossopterids contain schizogenous cavities (Neish et al.1993) that probably represent adaptations for gas exchangein dysaerobic, waterlogged environments. The abundanceof Vertebraria preserved in dark palaeosol horizons atFossilryggen suggests that the plants existed in relativelyuniform stands growing on consistently wet substrates.

If the exposures in the northern and southern sections ofFossilryggen are correlative then palynofloras of the MiddlePermian succession indicate a ‘hidden’ diversity of fernsand lycophytes not represented in the macroflora. Weinterpret the relatively complete organs represented in theassemblage to reflect essentially autochthonouspreservation of plant remains from a moist floodbasincommunity. It is likely that the coeval palynoassemblagesfrom these deposits contain pollen and spores derived froma broader source area than the macrofloras and providegreater insights into the diversity of the regional MiddlePermian biota.

Acknowledgements

This project was partly funded by an Australian ResearchCouncil Large Grant to S.M. The Swedish Polar ResearchSecretariate is acknowledged for logistic support during theSWEDARP 1988/89 and 19989/90 expeditions. Financialsupport K.L. and S.L. was obtained by several grants fromthe Swedish Science Council.

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