Antarctic Science 17 (1): 73–86 (2005) © Antarctic Science Ltd Printed in the UK DOI: 10.1017/S0954102005002464 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, Australia 2GeoBiosphere 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 of Fossilryggen, Vestfjella, Dronning Maud Land, Antarctica, is dominated by matted leaf impressions of Glossopteris sp. cf. G. communis Feistmantel and Glossopteris sp. cf. G. spatulata Pant & Singh, and in situ finely branched Vertebraria indica Royle rootlets. Equisetalean stems, rhizomes and leaf whorls (Phyllotheca australis Brongniart emend. Townrow), isolated seeds, scale-leaves, and fragmentary gymnosperm 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 is considered to represent a Middle Permian assemblage based principally on lithological and palynological correlation with the southern section at Fossilryggen and broad similarities to mid-Permian plant assemblages elsewhere in Gondwana.
Received 2 January 2004, accepted 23 August 2004
Key words: East Antarctica, Glossopteris, Middle Permian, palynostratigraphy, Phyllotheca, Vertebraria
Introduction Gondwana. Recent palynological studies have supported an The distribution of fossil Glossopteris leaves on disparate Early Permian age for the Milorgfjella succession (Larsson southern lands represented key evidence in early arguments et al. 1990, Lindström 1995b). Hjelle & Winsnes (1972) to support plate tectonics and the notion of a former first reported the presence of similar fossil plants at supercontinent: Gondwana. The discovery of Glossopteris Fossilryggen in Vestfjella and, based on photographs of the leaves on Mount Buckley at 85°S by Robert Scott's material, Dr E.P. Plumstead suggested that the fossils 1910–13 Antarctic expedition (Seward 1914) confirmed represented an Early Permian assemblage, although that Antarctica hosted this widespread Southern younger than that preserved at Milorgfjella. However, more Hemisphere taxon and that during the Permian, robust recent palynological studies (Lindström 1995a, 1996) have vegetation was able to survive close to the South Pole. Since suggested a Middle Permian age for at least some of the 1912, glossopterids and associated Permian plants have sedimentary rocks at Fossilryggen. The new assemblage been recorded from numerous localities in the from the northern exposures of Fossilryggen represents the Transantarctic Mountains, Antarctica (Edwards 1928, first Middle Permian macrofossil flora described from Cridland 1963, Rigby & Schopf 1969, Schopf 1970, Dronning Maud Land. Lambrecht et al. 1972, Lacey & Lucas 1981, Taylor et al. 1992, Cúneo et al. 1993), the Ellsworth Mountains (Gee Geological setting 1989, Taylor et al. 1989), and the Prince Charles Mountains (White 1969, Neish et al. 1993, McLoughlin et al. 1997). The scattered ridges and isolated nunataks of the Vestfjella Currently, the only plant macrofossils described and mountain range are situated along the inner part of the ice illustrated from Dronning Maud Land are those collected shelf embracing the western extent of Dronning Maud from three beds exposed in Milorgfjella, the north-eastern Land. Nunataks in this region predominantly expose extension of Heimefrontfjella, by the British Antarctic Jurassic basalts and dolerite intrusions but olivine-gabbro Survey in the late 1960s (Juckes 1972, Plumstead 1975). dominates the southern part of the mountain range. The Milorgfjella assemblages are dominated by Fossilryggen is situated in the north-eastern part of the glossopterids in which the central veins are poorly mountain range and is the only locality where Palaeozoic aggregated (representing the ‘Gangamopteris condition’). sedimentary rocks crop out. The geology of the area has Other Milorgfjella fossils include a conifer twig, been described by Hjelle & Winsnes (1972), Olaussen sphenophyte axes, possible lycophyte axes and fragmentary (1985), Larsson (1990, 1991) and Lindström (1995a, 1996), fern fronds, and various gymnosperm axes including some and it will only be summarized briefly herein. with attached spur shoots (Table I). These assemblages are The approximately 2 km long, low and narrow ridge of typical of low-diversity Early Permian floras from southern Fossilryggen is the main nunatak in the area (Fig. 1). It is
73 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 and originally 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 axis Paracalamites sp. cf. Phyllotheca australis Brong. pl. Ia (1), Ib Paracalamites australis Rigby Indeterminate small branched stem pl. Ic Possibly a fern frond fragment Stem with small round scars pl. Ia (2) Unidentifiable axis Hollow mound with grooved spiral markings pl. Ie Unidentifiable axis cf. Cordaitales pl. IIa–c, IIIa, IIId Unidentifiable gymnosperm axes cf. Glossopteridae pl. IIa, IIIb, IIIc, VIIa, VIIIa, Unidentifiable gymnosperm axes cf. Ginkgophyta long and short shoots pl. Vd–g, VIa–d Glossopterid long shoots bearing lateral spur shoots cf. Coniferophyta pl. VIIb, VIIc Unidentifiable gymnosperm axes Wood fragments pl. IVa–g Gymnosperm xylem tracheids with bordered pits Gangamopteris obovata Carr. pl. IXa–d Gangamopteris sp. cf. G. obovata (Carruthers) White Gangamopteris cyclopteroides Feist. pl. Xa–d Gangamopteris sp. cf. G. spatulata McCoy Gangamopteris sp. pl. XIVa (top) Gangamopteris obovata Carr. Walikalia juckesii Plumstead pl. XIa–f Gangamopteris sp. cf. G. spatulata McCoy Euryphyllum antarcticum Plumstead pl. XIIa–d, XIIIa–b Gangamopteris sp. cf. G. angustifolia (McCoy) McCoy Euryphyllum sp. pl. XIVc–d Glossopteris sp. cf. G. douglasii Rigby & Chandra Noeggerathiopsis hislopi (Bunb.) Feist. pl. XIVe Gangamopteris sp. cf. G. obovata (Carruthers) White Noeggerathiopsis spathulata (?) Dana pl. XIVf–g Gangamopteris sp. cf. G. obovata (Carruthers) White cf. Palaeovittaria sp. pl. XIVh Gangamopteris sp. Coniferous twig pl. XIVa–b cf. Walkomiella transvaalensis le Roux Mixed foliage pl. XV Gangamopteris sp. cf. G. obovata (Carruthers) White Fungi pl. IXc Indeterminate imprints and weathering patterns flanked to the west by a smaller nunatak, the NW Nunatak to the area in 1988/89 and 1989/90. Macrofossils are (informal name), and to the north-west by three small preserved as impressions in dark siltstone. Specimens were outliers (Fig. 1). Palaeozoic sedimentary rocks crop out near photographed under strong unilateral light from the upper the southern and northern limits of Fossilryggen, in the left. All illustrated specimens (prefixed LO) are lodged in "NW Nunatak" and in the middle outlier (Fig. 1). The the collections of the Geology Department, GeoBiosphere southern section at Fossilryggen hosts the most complete Science Centre, Lund University. In the systematic sedimentary succession where approximately 60 m of descriptions OD = original designation, M = by monotype, sandstones and shales are exposed. The southern section is SD = subsequent designation. transected by six dolerite dykes and one normal fault, which downfaults the northern portion of the outcrop. The central Palynology and northern parts of Fossilryggen consist mainly of basaltic rocks, although black shales are exposed in Lindström (1994, 1995a, 1996) recognized sixty-six taxa topographical lows. The north-western margin of the ridge including eight algae or acritarch species when analysing consists of a 30 m thick sequence of plant-rich black shales, the palynostratigraphy of the southern sedimentary section minor sandstones and mudstones, here called the ‘northern at Fossilryggen and nearby "NW Nunatak". The section’. assemblages also included thirty-five gymnosperm pollen Due to the intervening basaltic rocks, the relationship taxa, seventeen fern spores, four lycophyte species, and two between the southern and northern sections at Fossilryggen regarded as sphenophytes (Lindström 1996). The is not clear. The northern section is situated topographically assemblages are dominated by bisaccate pollen, including lower than the southern section (Lindström 1995a, text both taeniate forms (viz. Protohaploxypinus amplus, fig. 2), but since the sedimentary strata at Fossilryggen dip P. limpidus, Striatopodocarpidites cancellatus and S. fusus) slightly towards the north-west it is possible that the and non-taeniate species (viz. Scheuringipollenites northern section is somewhat younger than the southern maximus, S. ovatus and Alisporites spp.). Fern spores are one. common, especially Leiotriletes directus, Horriditriletes tereteangulatus, H. filiformis and Osmundacidites wellmanii. Didecitriletes ericianus, D. uncinatus, Material and methods Microbaculispora villosa, Indospora laevigata and The material described herein was collected from the Dictyotriletes labyrinthicus appear in the upper part of the northern section of Fossilryggen and the adjacent Northwest southern section, together with Guttulapollenites Nunatak by Professor Kent Larsson during the Swedish hannonicus and Protohaploxypinus rugatus (Lindström Antarctic Research Programme (SWEDARP) expeditions 1996). VESTFJELLA PERMIAN PLANT FOSSILS 75
Fig. 1. Locality details of the fossil assemblage: a. Map of Antarctica showing Dronning Maud Land, b. Map of Dronning Maud Land showing the position of Fossilryggen, c. Map of Fossilryggen, Vestfjella, showing the source of the described fossil assemblage (after Larsson 1991).
Lindström (1996) suggested that the successive first juxtaposed to Dronning Maud Land, these taxa also appear appearances of taxa within the southern section at at the same level (Anderson 1977). In the Prince Charles Fossilryggen and the "NW Nunatak" were more related to Mountains M. villosa first appears at a slightly lower level preservational factors than age differences. The than D. ericianus (Lindström, personal observation), and palynofloras were, therefore, correlated with Australian this is also the pattern in India (Tiwari & Tripathi 1992). Upper Stage 5 (late Wordian–Lopingian) microfloras Lindström (1996) found no identifiable palynomorphs in (Lindström 1996, Foster & Archbold 2001). However, if samples from the northern section at Fossilryggen. For this one considers the possibility that the successive first study, the samples were reprocessed according to standard appearances of taxa actually reflect age differences within palynological methods. One sample, F90-83, contains a few the southern section, it is possible that the upper part of the identifiable palynomorphs but they are generally section is equivalent to Australian Lower Stage 5b/c (late fragmentary, and of high thermal maturity, black to Roadian–early Wordian) microfloras (Lindström 1995a, brownish-black even after treatment with Schultze’s Foster & Archbold 2001), the lower part being somewhat solution. The following taxa have been identified: older. The co-appearance of Microbaculispora villosa and Horriditriletes tereteangulatus, Lunatisporites sp., Didecitriletes ericianus is a pattern that differs from the Osmundacidites wellmanii, Protohaploxypinus amplus and typical Australian palynostratigraphic succession. In the Scheuringipollenites maximus. These few taxa are Karoo Basin, South Africa, which during the Permian was unfortunately not sufficient to date the northern section 76 STEPHEN MCLOUGHLIN et al.
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. more precisely than to the Permian. Permian; Hawkesbury River area, northern Sydney Basin, Australia. Systematic palaeobotany Phyllotheca australis Brongniart emend. Townrow 1955 Phylum Sphenophyta Fig. 2a–g Class Sphenopsida Lectotype: No. 3388, Museum National d'Histoire Order Equisetales Naturelle, Paris (Townrow 1955, text fig. 1a). Family Gondwanostachyaceae Meyen 1969 Phyllotheca Brongniart emend. Townrow 1955 Specimens: LO 9379t, LO 9380t, LO 9382t, LO 9383t, LO 9390t, and one unfigured specimen. Type species: Phyllotheca australis Brongniart emend. Townrow 1955 (OD/M); ?Newcastle Coal Measures; Upper Diagnosis: See Townrow (1955, pp. 42–43). VESTFJELLA PERMIAN PLANT FOSSILS 77
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 of an 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.
Description: Material assigned to this species includes 50 mm apart. External longitudinal ridges and grooves are fragmentary rhizomes bearing fine roots (Fig. 2a, b, d), indistinct on root-bearing axes; ridges are opposite across slender aerial stems bearing leaves at nodes (Fig. 2c, e, f), nodes. Leaf-bearing axes are generally narrower (< 4 mm and detached leaf whorls (Fig. 2g). Axes are preserved wide) and shorter (longest available fragment 30 mm long) predominantly parallel to bedding. Rhizomes reach than root-bearing axes. They bear longitudinally striate > 260 mm long and up to 16 mm wide with 15 mm long and ridges 0.5–1 mm apart. Nodes on these axes are 4–8 mm < 0.5 mm wide roots emerging from nodes spaced up to apart and bear whorls of around 18 linear leaves that are 78 STEPHEN MCLOUGHLIN et al. fused to form a proximal sheath for about one-third to two- marginal venation angle reaches 60–70° near the base and thirds of their length (Fig. 2e & g). The 3–7 mm long sheath centre of the leaf but reduces to 40–50° towards the apex. is closely appressed to the stem (Fig. 2f). Distally, the leaves Vein density is around 21–25 per 10 mm at the margin in the diverge with a free length of 5–8 mm. Leaves are central part of the leaf. 0.5–0.75 mm wide at the limit of fusion and apices are pointed acute. Comments: This species shows substantial variation in form from small, narrow, linear leaves (Fig. 3a–g) to moderately Comments: Isolated, leafless, segmented axes from the large, oblanceolate leaves (Fig. 3h–l). Despite this broad Permian of Gondwana with longitudinal ribs arranged in range in shape and size the leaves are considered to opposition at the nodes may belong to a range of constitute a single species because they are represented by a sphenophyte families and most have been assigned to the morphological continuum. Small linear forms may form species Paracalamites australis Rigby (1966). represent young, incompletely expanded or exposed However, some Vestfjella axes bear leaf whorls (Fig. 2e) (canopy) leaves. Broad forms may represent mature shade traditionally assigned to Phyllotheca australis Brongniart leaves. All are characterized by continuously arched, emend. Townrow 1955 in the Gondwanostachyaceae relatively dense, anastomosing secondary veins with mid- (Meyen 1969). Based on comparisons to extant Equisetum leaf marginal venation angles of 60–70° and narrowly species, these plants are assumed to have been rhizomatous, crescentic areolae. herbaceous, and restricted to relatively moist habitats. Three Indian Glossopteris species are closely comparable Geographic and stratigraphical range: Common to these Antarctic specimens. Glossopteris communis throughout the Permian of Gondwana (Pant & Kidwai Feistmantel (1879) from the Lower Permian Karharbari 1968); most abundant in Middle to Upper Permian strata Formation is the most similar in shape and venation style. (Anderson & Anderson 1985). The larger Antarctic specimens are particularly similar to the type specimen of G. communis; smaller specimens tend Phylum ?Coniferophyta to have a much greater length:width ratio and fall outside Order Glossopteridales sensu Pant 1982 the range traditionally recognized for the Indian species Glossopteris Brongniart 1830 (Chandra & Surange 1979). Glossopteris intermittens Type species: Glossopteris browniana (Brongniart) Feistmantel 1881 from the Lower to Middle Permian Brongniart 1830 (SD of Brongniart 1830); ?Newcastle Coal Barakar Formation is comparable in size and venation Measures; Upper Permian; Gulgong-Maitland districts, curvature to Glossopteris sp. cf. G. communis but the former Sydney Basin, Australia. typically has a length:width ratio of only 3:1 and sparse secondary vein anastomoses (Chandra & Surange 1979). Glossopteris sp. cf. G. communis Feistmantel 1879 Glossopteris tenuifolia Pant & Gupta 1968 from the Upper Figs. 3a–l Permian Raniganj Formation has a shape and size more closely comparable to the narrower examples of the Specimens: LO 9384t, LO 9385t, LO 9386t, LO 9387t, Antarctic leaves but it tends to have straighter secondary LO 9388t, LO 9389t, LO 9390t, LO 9391t, LO 9392t, veins intersecting the margin at < 45°. LO 9417t, and ten unfigured specimens. Lower Permian leaves from the Karoo Basin attributed by Description: Leaves oblanceolate to linear (length:width = Anderson & Anderson (1985) to Hirsutum intermittens 3.5:1 to 10:1), typically 60–115 mm long (possibly up to Plumstead, Arberia madagascariensis (Appert) Anderson & 150 mm), 6–42 mm wide (typically 17–25 mm wide). The Anderson, and A. leeukuilensis Anderson & Anderson have widest point of the leaf is three-quarters to four-fifths along similar venation styles to Glossopteris sp. cf. G. communis its length. The leaves have a gently tapering cuneate base Feistmantel but these South African forms are typically with a 3–4 mm wide truncate abscission point (Fig. 3a, b, d, much larger (160–200 mm long) with acute, pointed apices. f, g, l). In some cases the leaf base may be slightly flared at Amongst other South African forms, the Fossilryggen the abscission point. The apex is generally rounded but in a specimens are closely comparable to Upper Permian sterile few specimens it is slightly emarginate (Fig. 3k); the margin leaves from the Karoo Basin attributed by Anderson & is otherwise entire. The midrib varies from indistinct to Anderson (1985) to Lidgettonia africana Thomas. The latter prominent, around 3 mm wide at the base, tapering are oblanceolate, 80–150 mm long, with narrowly throughout, persisting to the apex where it is < 0.5 mm crescentic areolae, vein marginal angles of around 50°, and thick, or in smaller forms evanescing in the distal 5 mm of in some cases an emarginate apex. In the absence of the leaf (Fig. 3c). The veins emerge from the midrib at cuticular information or attached reproductive structures the 10–20° and arch gently and continuously throughout the Fossilryggen specimens and L. africana leaves appear to lamina (Fig. 3e, h, l). Venation angles midway between the differ only in their degree of vein curvature. Amongst midrib and margin are 30–50° near the base and centre of eastern Australian Glossopteris leaves, the Upper Permian the leaf but reduce to 20–25° towards the apex. The G. curvinervosa McLoughlin (1994a) has the most similar VESTFJELLA PERMIAN PLANT FOSSILS 79
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. venation style but it tends to have a more acute apex that the Karharbari Formation (Lower Permian) of north-eastern Fossilryggen leaves. peninsular India (Chandra & Surange 1979) but numerous superficially similar leaves have been reported throughout Geographic and stratigraphic range: Glossopteris Gondwana from the Lower, Middle and Upper Permian communis sensu stricto is apparently restricted to the (Lacey et al. 1975, Rigby et al. 1980, Rayner & Coventry 80 STEPHEN MCLOUGHLIN et al.
Fig. 5. Vertebraria indica Royle 1834; Root impressions. a. Large root with two rows of chamber infillings [LO 9401t], b. Large root with a single, central row of chamber infillings flanked by featureless secondary wood impressions [LO 9402t], c. Intermediate-size root with a large lateral branch [LO 9403t], d. Intermediate-size root with three longitudinal rows of chamber infillings [LO 9404t], e. Small roots showing minimal segmentation, giving off ultimate rootlets at nodes [LO 9405t], f. Intermediate and fine rootlets with multiple orders of branching [LO 9406t], g. Intermediate-size, bedding-parallel, root giving off several orders of lateral roots [LO 9407t]. Scale bars = 10 mm.
1985). Determination of the relationships between these (Fig. 4f–h). The mid-lamina venation angle is 50–65° near widely separated fossils must await cuticular details or the base and centre of the leaf and reduces to 25–30° near attached reproductive structures. the apex. The marginal venation angle reaches 65–70° near the base and centre of the leaf but reduces to 30°–40° Glossopteris sp. cf. G. spatulata Pant & Singh 1971 towards the apex. Areolae are very narrowly crescentic to Figs. 4a–h linear. The marginal venation density is 20–28 per 10 mm in the central part of the leaf. Specimens: LO 9394t, LO 9395t, LO 9396t, LO 9397t, LO 9398t, LO 9399t, LO 9400t, and one unfigured specimen. Comments: Glossopteris sp. cf. G. spatulata Pant & Singh Description: Leaves narrowly elliptic to narrowly (1971) represents leaves that are typically twice as wide and oblanceolate (length:width = 3:1 to 4.5:1), reaching around substantially longer than Glossopteris sp. cf. G. communis 170–200 mm long and up to 70 mm wide (Fig. 4a–d). The from the same locality. Additionally, the secondary veins of widest point is two-thirds to three-quarters of the distance Glossopteris sp. cf. G. spatulata tend to arch near the midrib from base to apex. The margin is entire, the apex broadly and have a relatively straight course in the outer part of the rounded and the base cuneate (Fig. 4a, b, e). The midrib is lamina (Fig. 4e–h). In this respect, Glossopteris sp. cf. robust at the base, up to 4 mm wide, tapering throughout but G. spatulata is similar to G. acutifolia McLoughlin (1994a) persisting to the apex where it is < 0.5 mm wide. The veins and G. kingii McLoughlin (1994b) from the Upper Permian emerge from the midrib at 10–20°, arch gently in the inner of eastern Australia. However, the Australian species have half of the lamina but only slightly in the outer part different suites of accessory morphological characters VESTFJELLA PERMIAN PLANT FOSSILS 81 including smaller size, more acute apices or a lesser Description: Root impressions reaching in excess of marginal venation angle. Large leaves from the Australian 270 mm long and up to 38 mm wide, with one to four rows Upper Permian [e.g. G. ampla Dana (1849) and G. grandis of rectangular, alternating segments (Fig. 5a–d). The largest McLoughlin (1994a)] tend to have broader areolae and less specimens show segments restricted to the central part of dense venation. the root; the outer parts, up to 10 mm wide, are relatively The Antarctic leaves are closest in shape, size and smooth or possess fine longitudinal striae or transverse cleat venation pattern to Glossopteris spatulata from the Upper markings (Fig. 5b). Roots show up to three orders of Permian Raniganj Formation of India (Chandra & Surange branching. Lateral rootlets emerge (typically at 20–45°) 1979). However, Pant & Singh (1971) erected G. spatulata from the transverse partitions in larger roots (Fig. 5f & g). on both macromorphological and cuticular characters. As The ultimate rootlets are typically 2–4 mm long, 1–1.5 mm the Antarctic specimens lack cuticle it is unclear whether wide, with a single vascular strand (Fig. 5e & f). Roots are they are conspecific with the Indian forms. Glossopteris orientated predominantly parallel to, or at low angle to, feistmantelii Rigby, reported from the Upper Permian of bedding. India also has a similar venation style but tends to be a proportionately broader leaf (length:width = 2:1) with a Comments: Gould (1975) demonstrated a complete cordate base (Rigby 1964, Chandra & Surange 1979). anatomical transition between permineralized young roots Some South African Lower Permian Glossopteris leaves of Vertebraria indica and mature root or stem wood associated with Ottokaria buriadica Plumstead referable to Araucarioxylon. The smooth or longitudinally superficially resemble Glossopteris sp. cf. G. spatulata but striate outer region in the larger root impressions in the they have a truncate base and lack a well-defined midrib Fossilryggen assemblage (Fig. 5b) probably represents a (Plumstead 1956, Anderson & Anderson 1985). Middle zone of continuous secondary xylem forming complete Permian leaves co-preserved with Arberia allweyensis growth rings around the inner (segmented) zone with Anderson & Anderson fruits are distinguished by their schizogenous cavities. Pant (1958) assigned very small, tapered acute apices. Upper Permian leaves from the eastern univeined rootlets from a Permian, glossopterid-rich, Karoo Basin associated with Lidgettonia mooiriverensis assemblage of Tanzania to Lithorhiza Pant but such rootlets Anderson & Anderson share many features with are clearly connected to larger, segmented/chambered roots Glossopteris sp. cf. G. spatulata. They differ only in having referable to Vertebraria indica (Fig. 5e & f) and their a more attenuate (petiolate) base. As no fruits were found taxonomic differentiation seems superfluous. Additionally, attached to, or associated with, Glossopteris sp. cf. Neish et al. (1993) showed a wide variation in the anatomy G. spatulata at Fossilryggen, an affinity with Lidgettonia of Vertebraria roots, especially in the ontogenetic can not be confirmed. The Fossilryggen leaves are also development and number of schizogenous cavities, which strikingly similar to specimens assigned to G. communis would likely give rise to variably segmented or non- from the Lower Permian of Madagascar, although the latter segmented forms when preserved as impressions/ may reach 300 mm long and 100 mm wide (Appert 1977). compressions. Vertebraria indica roots are preserved in situ within flat- Geographic and stratigraphic range: Glossopteris laminated, dark siltstones, and clearly display several orders spatulata is notionally restricted to the Raniganj Formation of delicate branching. In a few cases they transect bedding (Upper Permian) of north-eastern peninsular India (Chandra planes at a low angle but they are predominantly orientated & Surange, 1979). The closely comparable forms from parallel to bedding suggesting that the parent plants Fossilryggen have not been reported elsewhere. Their possessed relatively shallow but broadly spreading root relationship to G. spatulata sensu stricto can not be systems. Root-rich beds typically contain few leaves and determined until cuticular details become available. probably represent organic-rich, immature palaeosols in floodbasin environments with a high water table. Vertebraria Royle emend. Schopf 1982 Geographic and stratigraphic range: Common throughout Type species: Vertebraria indica Royle 1834 (M); Raniganj the Permian of Gondwana. Formation; Upper Permian; Raniganj Coalfield, Damodar Valley, India. Glossopterid scale leaves Vertebraria indica Royle 1834 Figs. 6a–f Fig. 5a–g Specimens: LO 9408t, LO 9409t, LO 9410t, LO 9411t, Holotype: V4189, Natural History Museum, London. LO 9412t, LO 9413t, and one unfigured specimen. Specimens : LO 9401t, LO 9402t, LO 9403t, LO 9404t, LO Description: Detached, elliptical-rhombic, concavo-convex, 9405t, LO 9406t, LO 9407t, and four unfigured specimens. entire-margined, scale-leaves that are 12–18 mm wide and 15–19 mm long, including a 2–3 mm long acuminate apex Diagnosis: See Schopf (1982, p. 41). (Fig. 6a, d, f). A single small (6 mm wide, > 8 mm long) 82 STEPHEN MCLOUGHLIN et al.
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. specimen (Fig. 6e) falls outside this range. Although Comments: Isolated scale leaves of this type may represent typically poorly preserved, the bases of all scales are detached sterile bracts originally protecting the dormant tips apparently truncate (broadly attached). Several veins of branches or they may constitute the foliar component of emerge from the scale base and gently diverge and arch microsporangiate glossopterid fruits such as Eretmonia du across the lamina. The veins dichotomize and anastomose Toit emend. Lacey, van Dijk & Gordon-Gray (White 1978, throughout the lamina to maintain a density of around 24 Anderson & Anderson 1985, Holmes 1995). Both sterile per 10 mm (measured across the lamina); no distinct midrib and fertile scale leaves typically show a broad variation in is developed. size and shape. The Vestfjella bracts do not bear sporangia. VESTFJELLA PERMIAN PLANT FOSSILS 83
Such bracts with very few distinctive morphological Large seeds (15–80 mm long) have not been found attached characters have traditionally been assigned to informal to glossopterid fruits but they are likely affiliates based on categories such as 'scale leaves' or 'squamae' or, in some common association. Samaropsis sp. from Fossilryggen cases, to established form genera (Chandra & Surange falls into the medium-sized category of seeds. Most fruits 1977a, 1977b, Banerjee 1984). It is likely that scale leaves associated with seeds of this size category have Middle to produced by any glossopterid plant ranged from entirely Late Permian ranges. The exception is Arberia, which sterile, functionally protective bud scales to strongly mostly ranges from Early to earliest Middle Permian. expanded microsporophylls (White 1978). Additionally, However, attached Arberia seeds lack strongly developed some Gondwanan scale leaves show evidence of either wings (Rigby 1972, Appert 1977) and the seed body is basal or apical expansion transitional with normal usually much longer than wide (ovate to narrowly vegetative leaves (Anderson & Anderson 1985, Holmes elliptical). 1995). Consequently, it is difficult to confidently assign Seeds attached to Lidgettonia lidgettonioides (Lacey, van these fossils to any established taxon in the absence of Dijk & Gordon-Gray) Anderson & Anderson 1985 from the attached reproductive structures. Upper Permian of the Karoo Basin, South Africa, resemble the Fossilryggen Samaropsis species in shape, dimensions Geographic and stratigraphic range: Sterile glossopterid and surface features but the proximal lobes of the wings are scale leaves are common throughout the Permian of less developed in the former. Samaropsis rigbyi Millan Gondwana but are especially abundant in Upper Permian 1977 from the Tubarão Group (Early Permian), Brazil, has strata (Anderson & Anderson 1985). similar broad wings with prominent proximal lobes but it is Dispersed seeds typically twice the size of the Fossilryggen seeds. The few Samaropsis Goeppert 1864 morphological characters available from platyspermic seed impressions prohibit further comparisons. Type species: Samaropsis ulmiformis Goeppert 1864 (OD); rock unit uncertain; Permian; Braunau, Bohemia, Czech Geographic range: Not known beyond Fossilryggen. Republic. Rotundocarpus Maithy 1965 Samaropsis sp. Type species: Rotundocarpus striatus Maithy 1965 (OD); Figs. 6g–j Karharbari Formation; Lower Permian; Jogtiabad Pit, Specimens: LO 9414t, LO 9415t, LO 9416t, and one Karharbari Colliery, Giridih Coalfield, India. unfigured specimen. Rotundocarpus sp. cf. R. ovatus Maithy 1965 Description: Circular to broadly elliptical seeds 7–8 mm Fig. 6k–n long, 6–7 mm wide. The central body is generally indistinct, Specimens: LO 9418t, LO 9419t, LO 9420t, LO 9421t, and flattened, elliptical to ovate, 7–8 mm long and 3–4.5 mm two unfigured specimens. wide (Fig. 6g, i, j), but it is slightly raised on one impression (Fig. 6h). The central body is flanked by crescentic marginal Description: Very small, ovate to elliptical seed, 3 mm long wings 1–2 mm wide that are, in some cases, extended into and 2 mm wide (Fig. 6k–n). The seeds are preserved as deep lobes adjacent to the chalazal end of the seed (Fig. 6h). convex impressions or, in some cases, the centre of the seed The chalazal end of the central body is rounded to truncate; (nucellus), or the seed as whole, may have decayed and the micropylar end is rounded. The central body bears been filled with fine sediment to produce an ovate to indistinct, longitudinal striations; the wings are generally elliptical cast. These seeds either lack a lateral wing or the featureless. All seeds are detached from other organs. wing is extremely reduced. A narrow (0.1 mm) indentation in the sediment is evident around some specimens. The Comments: Seeds of this type are probably attributable to outer surface bears fine longitudinal striations but the seeds glossopterids based on the lack of other gymnosperm fossils are otherwise featureless. in the assemblage. Seeds associated with glossopterids in other parts of Gondwana generally fall into three size Comments: These seeds are relatively common in beds at categories. Very small seeds (< 5 mm long) have been found Fossilryggen in association with Glossopteris leaf attached to capitate fruits such as Dictyopteridium, fragments, Samaropsis sp., and wood fragments, but they Senotheca, and Plumsteadia (Banerjee 1969, Surange & may be easily overlooked owing to their diminutive size. Chandra 1973, Chandra & Surange 1976, Benecke 1976, The dearth of diagnostic features or attached fruits prohibits Rigby 1978). Medium-sized seeds (5–15 mm long) have conclusive determination of their affinities but they are been found attached to, or associated with, a broad array of similar in size, shape, and in the lack of an obvious wing, to cupulate and loosely branched fruits such as Arberia, Rotundocarpus ovatus Maithy 1965 from the Karharbari Rigbya, Cometia and Lidgettonia (Rigby 1972, Lacey et al. Formation (Lower Permian), Giridih Coalfield, Damodar 1975, Anderson & Anderson 1985, McLoughlin 1990). Valley, India. A few other Gondwanan Palaeozoic fossils, 84 STEPHEN MCLOUGHLIN et al.
Fig. 7a. Matted Glossopteris sp. cf. G. communis Feistmantel 1879 leaves on a bedding plane [LO 9387t]. b. Irregular axis fragment with remnant conjugate cleat sets [LO 9422t]. c. Large gymnosperm axis impression with discontinuous longitudinal striae [LO 9423t]. Scale bars = 10 mm. e.g. Carpolithus circularis Walkom 1935 from the Upper morphology and it is likely that both Glossopteris species Permian of eastern Australia, appear to be similar-sized were borne on plants possessing roots of Vertebraria indica radiospermic seeds but they generally differ in having type. Similarly, protective scale leaves and micro- extended micropyles or ridged ornamentation. sporophylls have relatively conservative morphologies amongst glossopterids, differing mainly in size and apex Geographic and stratigraphic range: Not known beyond shape. Previous schemes to generically segregate Fossilryggen. glossopterid scale leaves have not been widely accepted Gymnosperm axes outside the immediate study area (Chandra & Surange Fig. 7b & c 1977b, Banerjee 1984, Anderson & Anderson 1985). All organs in the Vestfjella assemblage are dissociated, hence it Specimens: LO 9423t, LO 9422t. is not possible to link the scale leaves or dispersed seeds Comments: Several axis impressions are present in the with either of the Glossopteris species. Fossilryggen assemblage. They reach > 140 mm long and None of the Fossilryggen taxa provides a precise age show either discontinuous longitudinal striations (Fig. 7c) control on the host rocks. The fossils have morphological or irregular cleat patterns (Fig. 7b). Their size suggests that similarities to both Early and Late Permian species from they are derived from gymnosperms but they lack additional other parts of Gondwana and this may be consistent with an diagnostic features. intermediate (Middle Permian) age deduced from palynoassemblages in the adjacent (southern section) sedimentary rocks at Fossilryggen (= early Late Permian of Discussion the traditional two-part Permian timescale used by The macroflora from Fossilryggen represents a typical low- Lindström 1995a). The Fossilryggen assemblage lacks diversity, high-latitude, Gondwanan, Permian plant Gangamopteris leaves that are typical of Early Permian assemblage. Only two species of glossopterid foliage are floras. Such leaves are well-represented in the nearby represented and neither can be confidently attributed to an Milorgfjella flora (Plumstead 1975, Table I). established species. Glossopterids have conservative root Phyllotheca australis, the only non-gymnospermous plant VESTFJELLA PERMIAN PLANT FOSSILS 85 in the assemblage, is a common element in Gondwanan Proceedings of a symposium on evolutionary botany and Permian assemblages. This plant probably occupied biostratigraphy. A.K. Ghosh Commemoration Volume. New Delhi: understorey niches in consistently wet habitats based on the Today and Tomorrows Printers and Publishers, 29–59. BENECKE, A.K. 1976. Several new forms of Glossopteris fructifications small stature of the Permian forms and their requirement for from the Beaufort Daptocephalus-zone (Upper Permian) of Natal, South moist conditions to enable gamete exchange. Glossopteris Africa. Palaeontologia Africana, 19, 97–125. sp. cf. G. communis is locally represented by dense BRONGNIART, A. 1828–38. Histoire des végétaux fossiles on reserches accumulations of foliage (Fig. 7a) that possibly represent botaniques et géologiques sur les végétaux renfermés dans les diverses mats of deciduous leaves. Glossopterids are generally couches du globe. Paris: G. Dufour & Ed. D'Ocagne, 1, 1–136. CHANDRA, S. & SURANGE, K.R. 1976. Studies of the reproductive organs of considered to have been large trees occupying floodbasin Glossopteris Part 1. Dictyopteridium feistmanteli sp. nov. attached on settings and they provided the principal organic constituents Glossopteris tenuinervis. Palaeontographica 156B, 87–102. of Gondwanan Permian coals (Gould & Delevoryas 1977, CHANDRA, S. & SURANGE, K.R. 1977a. Fertile bracts and scales of Retallack & Dilcher 1988). The segmented roots of Glossopteris fructifications from the Lower Gondwana of India. glossopterids contain schizogenous cavities (Neish et al. Palaeobotanist, 24, 195–201. CHANDRA, S. & SURANGE, K.R. 1977b. Some scale leaves and sporangia 1993) that probably represent adaptations for gas exchange from the Raniganj Coalfield, India. Palaeobotanist, 24, 245–253. in dysaerobic, waterlogged environments. The abundance CHANDRA, S. & SURANGE, K.R. 1979. Revision of the Indian species of of Vertebraria preserved in dark palaeosol horizons at Glossopteris. Lucknow: Birbal Sahni Institute of Palaeobotany, Fossilryggen suggests that the plants existed in relatively Monograph 2, 301 pp. uniform stands growing on consistently wet substrates. CRIDLAND, A.A. 1963. Glossopteris flora from the Ohio Range, Antarctica. American Journal of Botany, 50, 186–195. If the exposures in the northern and southern sections of CÚNEO, R., ISBELL, J., TAYLOR, E.L. & TAYLOR, T.N. 1993. The Fossilryggen are correlative then palynofloras of the Middle Glossopteris flora from Antarctica: taphonomy and paleoecology. 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