A R E V I S I O N O F T H E O V U L I F E R O U S F R U C T I F I C A T I O N S OF G L O S S O P T E R I D S F R O M T H E P E R M I A N O F S O U T H A F R I C A V O L U M E I I Rosemary Adendorff A thesis submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Doctor of Philosophy Johannesburg, 2005 ii LIST OF TABLES APPENDIX I Table A.I.1. Catalogue numbers and localities for Rigbya. Table A.I.2. Catalogue numbers and localities for incertae sedis Table A.I.3. Catalogue numbers and localities for Arberia. Table A.I.4. Catalogue numbers and localities for Vereenia. Table A.I.5. Catalogue numbers and localities for Bifariala. Table A.I.6. Catalogue numbers and localities for Estcourtia. Table A.I.7. Catalogue numbers and localities for Elatra. Table A.I.8. Catalogue numbers and localities for Ottokaria. Table A.I.9. Catalogue numbers and localities for Scutum. Table A.I.10. Catalogue numbers and localities for Gladiopomum. Table A.I.11. Catalogue numbers and localities for Plumsteadia. Table A.I.12. Catalogue numbers and localities for Gonophylloides. Table A.I.13. Catalogue numbers and localities for Dictyopteridium. Table A.I.14. Catalogue numbers and localities for Lidgettonia. APPENDIX II Table A.II.1. Quantitative data collected for Rigbya arberioides & incertae sedis. Table A.II.2. Quantitative data collected for Arberia spp. & Vereenia leeukuilensis. Table A.II.3. Quantitative data collected for Bifariala, Estcourtia & Elatra. Table A.II.4. Quantitative data collected for Ottokaria spp. Table A.II.5. Quantitative data collected for Scutum leslii. Table A.II.6. Quantitative data collected for Gladiopomum spp. Table A.II.7. Quantitative data collected for Plumsteadia spp. Table A.II.8. Quantitative data collected for Gonophylloides spp. Table A.II.9. Quantitative data collected for Dictyopteridium spp. Table A.II.10. Quantitative data collected or referenced for Lidgettonia spp. PLATES Table P.1. Alphabetical list of glossopterid ovulate fructifications with chapter and plate references. 364 APPENDIX I CATALOGUE NUMBERS AND LOCALITIES OF ORIGIN OF ALL SPECIMENS REFERRED TO IN THIS INVESTIGATION RIGBYACEAE Table A.I.1. Catalogue numbers and localities for Rigbya. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS BP/2/11847a BP/2/13047 BP/2/1644a BP/2/8173 BP/2/8174 BP/2/8178a&b BP/2/8179 BP/2/8180 BP/2/8181a&b BP/2/8182 BP/2/8183 BP/2/8184 BP/2/8185a&b BP/2/8187a&b BP/2/8188 BP/2/8191 BP/2/8193 BP/2/8196 BP/2/8205 BP/2/8207 NM/1641a NM/1642a&b NM/1647a NM/1648 NM/1649 NM/1650 NM/1651 NM/1653 NM/1656 NM/1658 NM/1660 NM/1666 NM/1669a spn1 NM/1670a Mooi River 20 14 BPI NM R. arberioides BP/2/11386c BP/2/11727 BP/2/11844a&b BP/2/11845a&b BP/2/11846 BP/2/11847b BP/2/11848a Bulwer 7 BPI 365 Table A.I.2. Catalogue numbers and localities for incertae sedis SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS incertae sedis (previously known as Arberia allweyensis; Anderson & Anderson, 1985) PRE/F/8380a&b Lawley 1 PRE ARBERIACEAE Table A.I.3. Catalogue numbers and localities for Arberia. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS A. hlobanensis BP/2/15893 BP/2/15914(cp) Hlobane 1 BPI A. madagascarensis GSP/H/102 Hammanskraal 1 GSP A. allweyensis Lawley 1 PRE Table A.I.4. Catalogue numbers and localities for Vereenia. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS V. leeukuilensis BP/2/14282 BP/2/14283 BP/2/14284 BP/2/14285 BP/2/14286a BP/2/14287a Vereeniging 6 BPI 366 DICTYOPTERIDIACEAE Table A.I.5. Catalogue numbers and localities for Bifariala. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS B. intermittens BP/2/13692b BP/2/13939 BP/2/13963 BP/2/13964 BP/2/13966b BP/2/13967a&b BP/2/13979 BP/2/13980 BP/2/13982 BP/2/13987 BP/2/13989 BP/2/13993 BP/2/13994 BP/2/13998 BP/2/14001 BP/2/14002 BP/2/14003 BP/2/14004 BP/2/14008 BP/2/14010 BP/2/14011 BP/2/14012a BP/2/14014 BP/2/14015 BP/2/14018 BP/2/14019 BP/2/14026a&b BP/2/14031 BP/2/14034 BP/2/14049 VM/03/3205/59 VM/03/3205/60 VM/03/3205/61 VM/03/3205/62 VM/03/3205/63 VM/03/3205/64 VM/03/3205/65 VM/03/3205/67 VM/03/3205/68 VM/03/3205/69 VM/03/3205/70 Vereeniging 30 11 BPI VM Table A.I.6. Catalogue numbers and localities for Estcourtia. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS E. conspicua BP/2/8137a BP/2/8172a&b NM/1276a&b Mooi River 3 BPI NM 367 Table A.I.7. Catalogue numbers and localities for Elatra. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS BP/2/15701 Vereeniging 1 BPI E. leslii BP/2/- BP/2/7080 BP7092 BP7092 spn2 BP7093b BP7102 BP7124 BP7127 BP7129 spn1 BP7129 spn2 BP7132a BP7140a spn1 BP7140a spn2 BP7158 BP7163 BP7176a BP7214 Rspn2 BP7214 BP7220 spn1 BP7220 spn2 BP7221a BP7222a BP7318 BP7348 spn1 BP7395 BP7396 BP7400 spn1 BP7400 spn2 BP7401 BP7402 BP7426 BP7433 BP15701 BP16188 GSP/HI/3 GSP/HI/37a GSP/HI/37b spn1 GSP/HI/37b spn2 GSP/HI/60 GSP/HI/124 GSP/HI/154 GSP/HI/155 GSP/HI/157 GSP/HI/213 GSP/HI/258 spn1 GSP/HI/258 spn2 GSP/HII/65 GSP/HII/82 GSP/HII/133b GSP/HII/175 PRE/F/15a spn1 PRE/F/28 PRE/F/36a R Hammanskraal 34 16 3 BPI GSP PRE 368 Table A.I.8. Catalogue numbers and localities for Ottokaria. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS O. buriadica BP/2/13599a BP/2/13599b BP/2/13630 BP/2/13632 BP/2/13633 BP/2/13635a&b BP/2/13636 BP/2/13638 BP/2/13639a BP/2/13639b VM/03/3205/122 VM/03/3205/124 VM/03/3205/127 VM/03/3205/135 Vereeniging 10 4 BPI VM O. hammanskraalensis BP/2/7221b BP/2/7291 BP/2/7415 GSP/HI/152 spn1 GSP/HI/152 spn2 Hammanskraal 3 2 BPI GSP O. transvaalensis BP/2/13069 BP/2/13600a BP/2/13600b BP/2/13603 BP/2/13605 BP/2/13606 BP/2/13607a BP/2/13607b BP/2/13611 BP/2/13613? BP/2/13615 BP/2/13621 BP/2/13622 BP/2/13624 BP/2/13634 BP/2/13643 BP/2/13645 BP/2/13646 BP/2/13674 VM/03/3205/117 VM/03/3205/126 VM/03/3205/128 VM/03/3205/129 VM/03/3205/130 VM/03/3205/132 Vereeniging 19 6 BPI VM 369 Table A.I.9. Catalogue numbers and localities for Scutum. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS BP/2/- BP/2/16136a BP/2/16139 BP/2/16152a Ermelo 4 BPI S. leslii BP/2/1424 BP/2/13539 BP/2/13543 BP/2/13729a BP/2/13730a BP/2/13731 BP/2/13732 BP/2/13733 BP/2/13735 BP/2/13736 BP/2/13737 BP/2/13739 BP/2/13740 BP/2/13741 BP/2/13744 BP/2/13745 BP/2/13746 BP/2/13747 BP/2/13749 BP/2/13750 BP/2/13751 BP/2/13753 BP/2/13758 BP/2/13760 BP/2/13762 BP/2/13763 BP/2/13765 BP/2/13769 BP/2/13774 BP/2/13783 BP/2/13799 BP/2/13800 BP/2/13803 BP/2/13808a BP/2/13810 BP/2/13814 BP/2/13820 BP/2/13822 BP/2/13826 BP/2/13829 BP/2/13834 BP/2/13837a BP/2/13865 BP/2/13866 BP/2/13867 BP/2/13868 BP/2/13872 BP/2/13874 BP/2/13880 BP/2/13883a&b BP/2/13884 BP/2/13885 Vereeniging 56 18 BPI VM 370 SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS BP/2/13888a BP/2/13889a BP/2/13890 BP/2/13891a VM/03/3205/85 VM/03/3205/86 VM/03/3205/87 VM/03/3205/90 VM/03/3205/96 VM/03/3205/98 VM/03/3205/99 VM/03/3205/97 VM/03/3205/101 VM/03/3205/103 VM/03/3205/105 VM/03/3205/108 VM/03/3205/110 VM/03/3205/111 VM/03/3205/112 VM/03/3205/115 VM/03/3205/164 VM/03/3205/165 Table A.I.10. Catalogue numbers and localities for Gladiopomum. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS BP/2/13754 BP/2/13936 BP/2/13937 BP/2/13941 BP/2/13945 BP/2/13971 VM/03/3205/64 Vereeniging 6 1 BPI VM G. dutoitides BP/2/16037 Hlobane 1 BPI G. acadarense BP/2/16329a&b BP/2/16330a BP/2/16331a&b BP/2/16332 spn1 BP/2/16332 spn2 BP/2/16333a&b BP/2/16334 BP/2/16335a BP/2/16337 BP/2/16341 BP/2/16342a BP/2/16343a&b BP/2/16344a&b BP/2/16345 BP/2/16346 BP/2/16347a&b BP/2/16348 BP/2/16349 BP/2/16350 BP/2/16351a&b Cedara 23 1 BPI GSP 371 BP/2/16352a BP/2/16353 BP/2/16354a&b BP/2/16355a&b GSP/- G. elongatum BP/2/28880a&b BP/2/28876a Rietspruit 2 BPI Table A.I.11. Catalogue numbers and localities for Plumsteadia. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS P. lerouxii BP/2/13550 BP/2/14149a&b BP/2/14151 BP/2/14152 BP/2/14153 BP/2/14156 BP/2/14157 BP/2/14158 BP/2/14159 BP/2/14160 BP/2/14162 BP/2/14165 BP/2/14166 BP/2/14169 BP/2/14170a&b BP/2/14171 BP/2/14173a BP/2/14174 BP/2/14175a&b BP/2/14176b BP/2/14178 BP/2/14179 BP/2/14181a BP/2/14182 BP/2/14183 BP/2/14186 BP/2/14189 BP/2/14190 BP/2/14191a&b BP/2/14205 BP/2/14210 BP/2/14211 BP/2/14220 BP/2/14221 VM/03/3205/167 VM/03/3205/50 VM/03/3205/52 Vereeniging 34 3 BPI VM NM/1266 spn1 NM/1266 spn2 NM/1266 R Mooi River 3 NM BP/2/11188a&b Inhluzane 11 BPI P. gibbosa BP/2/12985b R BP/2/12521a&b BP/2/12536 BP/2/12537 BP/2/12984a Loskop 14 BPI 372 SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS Rspn1 BP/2/12984a Rspn3 BP/2/12984a BP/2/12987 BP/2/13000a&b BP/2/12995b BP/2/12994a&b BP/2/13004a&b BP/2/13011a BP/2/13008a spn2 Table A.I.12. Catalogue numbers and localities for Gonophylloides. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS BP-? Hammanskraal 1 BPI G.strictum BP/2/14222a&b BP/2/14223 BP/2/14224 BP/2/14225 BP/2/14228a&b BP/2/14230a BP/2/14231a&b BP/2/14233 BP/2/14235b BP/2/14236 BP/2/14245 VM/03/3205/78 VM/03/3205/79 VM/03/3205/82 Vereeniging 11 3 BPI VM G. waltonii BP/2/14238 BP/2/14239b VM/03/3205/80 VM/03/3205/81 Vereeniging 2 2 BPI VM Table A.I.13. Catalogue numbers and localities for Dictyopteridium. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS D. flabellatum BP/2/12525a spn1 BP/2/12525a spn2 BP/2/12525a spn3 BP/2/12525a spn4 BP/2/12525b BP/2/12529a BP/2/12529a spn2 BP/2/12533 BP/2/12534 BP/2/12539 BP/2/12968 Loskop 18 BPI 373 SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS BP/2/12969 BP/2/12970 BP/2/12971 BP/2/12972 BP/2/12973 BP/2/12974a BP/2/12985a&b D. natalensis BP/2/8147a BP/2/8148b BP/2/8151b BP/2/8152b BP/2/8154a spn2 BP/2/8154b BP/2/8159a BP/2/8160a&b BP/2/8162 BP/2/8167a BP/2/8167b spn2 BP/2/8168a&b BP/2/8169 BP/2/8171 BP/2/12544 BP/2/13049 BP/2/13100a&b NM/1241a spn2 NM/1254 NM/1244a&b NM/1243b NM/1242a&b NM/1246a NM/1247b NM/1274a NM/1260 NM/1257 NM/1256(9) NM/1251b NM/1280a NM/1280b NM/1273b NM/1261 NM/1252a&b NM/1250a NM/1249a&b NM/1248a&b NM/1672 NM/1258 NM/1274b NM/1275b NM/1683a&b Mooi River 25 17 BPI NM cf. D. sporiferum BP/2/13056 Mooi River 1 BPI 374 LIDGETTONIACEAE Table A.I.14. Catalogue numbers and localities for Lidgettonia. SPECIES CATALOGUE NUMBERS LOCALITY NUMBER OF SPECIMENS INSTITUTIONS L. lidgettonioides As per Lacey et. al. (1975), Anderson & Anderson (1985) Mooi River Bulwer Bergville 50 2 5 BPI, NM BPI BPI As per Thomas (1958), Lacey et. al. (1975), Anderson & Anderson (1985) Mooi River Lidgetton Loskop Estcourt 50 10 20 4 BPI, NM BP11103a BP11106b BP11107b BP11108b BP11112b BP11116b BP11146a Inhluzane 7 BPI L. africana BP/2/13950a&b spn2 BP/2/19351a&b BP/2/28274 BP/2/28356a BP/2/28934a&b BP/2/28935a&b spn1 BP/2/28935a&b spn2 BP/2/28935a&b spn3 PRE/F/8393a&b spn1 Lawley 8 1 BPI PRE L. elegans As per Lacey et. al. (1975) Mooi River 16 BPI, NM 375 APPENDIX II QUANTITATIVE DATA SUMMARY 376 377 378 379 380 381 382 383 384 385 386 387 388 389 390 391 392 393 APPENDIX III GLOSSARY OF BASIC TERMS Below is a list of some of the terms used in the description of the glossopterid fructifications and their subtending leaves. FRUCTIFICATIONS Acropetal: maturing/produced sequentially from the base to the apex. Basipetal: maturing/produced sequentially from apex to base. Campylodromous venation: several primary veins or their branches, originating at or near a single point and running in strongly recurved arches that converge apically (e.g. Leaf Architecture Working Group, 1999). Capitulum (pl. capitula): a simple derivation of the Latin ?caput ? meaning ?head?, and applied here to members of the Dictyopterideaceae and Lidgettoniaceae, in reference to the receptacle and its wing. Cicatrix (pl. cicatrices): from the Latin word for scar; used here for the central, often raised (in impressions) tubercle in the centre of a seed scar, which represents the point of hilar detachment from the fructification (used by Schopf, 1976); Cupule: the Oxford English Dictionary defines a cupule as a cup-shaped organ, receptacle, etc. The word is derived from the Latin cupula, or cupola, which is a rounded dome.The term is broadly used in botanical, mycological and zoological fields, and has been used by some authors (e.g. Lacey et al., 1975) to refer to the microfructi attached to scale leaves in the Lidgettoniaceae. Some authors have objected to the term on the grounds that it infers homologies with northern hemisphere pteridosperms. Here the fertile structures of the Lidgettoniaceae are called capitula to avoid confusion. Dentate wing margin: a scalloped margin, where the section of wing corresponding to a marginal seed scar is broadly indented, the ?teeth? aligned with the junctions between seed scars; compare with a lobed wing. Fertiliger: a term favoured by some authors for the fertile unit comprising a glossopterid fructification and its subtending leaf (introduced by Schopf, 1976, but not extensively used). Fluting: shallow, rounded grooves in the wing, perpendicular to the receptacle margin and bounded by sharp creases (in impressions of the fertile surface), or ridges (in impressions of the sterile surface) that correspond to veins on the sterile surface of the receptacle. 394 Lobed wing: characteristic of Ottokaria, where the wing is divided into lobes that exhibit varying degrees of fusion; the mid-line of each lobe corresponds to the mid-line of a single marginal seed scar; compare with a dentate wing margin. Pedicel: in this study refers to the stalk of the ovulate fructifications. Polysperm: multiovulate fructification, bearing multiple seeds; term introduced by Schopf (1976) and used by Anderson & Anderson (1985) but not by many other authors. Primary wing: Bifariala intermittens and Elatra leslii, previously assigned to the genus Hirsutum, have been found to possess at least two superposed wings. The more robust, prominent wing, characterised in both taxa by a tapered base and elongated apex, has been referred to here as the ?primary wing?. In both B. intermittens and E. leslii, the primary wing was the only wing to have been included in the original diagnoses of the two species. The more obscure, apparently more fragile, secondary wing, is recognised as such for the first time in this study. The terms ?primary? and ?secondary? have been assigned on the basis of the prominence of the wing forms. Receptacle: the central, seed-bearing part of both the capitate and cupulate fructifications (sensu McLoughlin, 1990a); the receptacle is bifacial, with seed scars on one surface and venation on the opposing surface. Secondary wing: see ?Primary wing?. Seed scar: an elliptical to circular, raised cushion-like structure, usually with a central depression containing a raised tubercle, representing a point of seed attachment; the cushion represents a hollow in which the seed base would have rested, and the tubercle represents the cicatrix, or vascular connection to the fructification. Seed scar density: the number of seed scars counted within a 25 mm2 area of the receptacle. Wing: the flange of tissue along the periphery of the receptacle; may be variously fluted, lobed, scalloped, broad, narrow etc. depending on the taxon; always finely striated; fluting corresponds to veins on sterile surface of fructification and seed scars on fertile surface in capitate polysperms; thinner than the fleshy receptacle, apparently membranous and delicate in some cases. LEAVES Although glossopterid leaf taxonomy has only been touched upon, descriptions of the subtending Glossopteris leaves of attached fructifications have been included and certain descriptors used in the elucidation of the leaves need to be clarified (see text-fig. 3.1.5, p. 49). 395 Leaf sectors: three length-wise sectors or regions are referred to in leaf descriptions, viz. apical, medial and basal; the lamina has also been divided into three lateral regions, from midrib to margin, viz. proximal (adjacent to midrib), mid-laminal and marginal. Marginal vein angle: the angle formed between the medio-longitudinal leaf axis and a line drawn through a vein near the margin of the leaf, in the medial section of the leaf. Marginal vein density: the number of veins along a 5 mm section of the leaf margin. Mesh: an area of leaf lamina bounded by anastomosing and bifurcating veins. Mid-laminal vein angle: the angle formed by the medio-longitudinal leaf axis and a line drawn through a vein in the mid-laminal portion of the medial section of a leaf. Midrib: the longitudinally orientated bundle of veins extending along part, or the entire medial length of some Glossopteris leaves, and which may resemble a strong, central mid-vein; there is no single, large vein but rather an aggregate of closely spaced veins; well developed hypodermal tissue is often associated with the midrib, emphasising its prominence. 396 APPENDIX IV TAXONOMIC ISSUES SURROUNDING THE GENUS HIRSUTUM Under the impression that Hirsutum was an illegitimate and confusing generic title which had been discredited by several authors in the past (e.g. Mukherjee et al., 1966; Rigby, 1978; Banerjee, 1984), Adendorff et al. (2002) unwittingly created a stir recently by reassigning the type species of this genus to Gladiopomum dutoitides. This publication was spurred on by the discovery of a number of exciting new morphological features amongst the fructifications historically grouped within Plumstead?s (1958a) genus Hirsutum, and represents the first of three intended papers re-defining these taxa. Some workers still defend the legitimacy of the name Hirsutum (e.g. Anderson & Anderson, 1985; Edith Taylor, IOPC 2004, Argentina, pers. comm.), and this Appendix has been included to explain why the decision was made to disband such a historic genus. Strictly speaking, the name ?Hirsutum? is illegitimate in terms of Article 20.2 of the ICBN (Tokyo Code, Greuter et al. 1994), which states that a descriptive term should not be used as a generic title. ?Hirsutum? is an adjective used in formal taxonomic diagnoses (p. 338; Stearn, 1973), and therefore the name is nomenclaturally invalid. It is for this very reason that the name ?Lanceolatus?, another of Plumstead?s (1952) glossopterid fructification taxa, has been declared illegitimate in terms of the code (?Lanceolatus? is cited as an example of an illegal name in the Toyko Code; ICBN Art. 20.2, ex. 4). Of course, nomenclatural correctness does not supersede the need for a taxonomic system that is useful, practical and consistent. There are several other reasons which weigh heavily in favour of discontinuing the use of this generic title. The following is a summary and discussion of the complex events which have created the taxonomic challenge with which we are faced today 397 (see section 3.2, p. 77 for more information about the morphological features mentioned here). In 1952 Plumstead created the genus Scutum, erecting five species to accommodate the variation she noticed within this group. Scutum dutoitides was described as being longer and narrower than members of the other species, and as having a wing which was smoother and narrower, with poorly defined fluting and an entire margin. She noted that the wing in this species tapered away at the base, and was ?finely striated in outward and upward curves looking like hairs?. Later she interpreted these striations as hair-like pollenate organs, and on this basis, Plumstead (1958a) placed all specimens residing in Scutum dutoitides in a new genus Hirsutum. In the same paper Plumstead created a new species, Hirsutum intermittens to accommodate specimens of Scutum dutoitides which: - were larger and broader; - had a receptacle with a broad, square base and pointed apex; - had a wing which tapered abruptly at the base; - were closely associated with Glossopteris intermittens. In creating H. intermittens, Plumstead (1958a) transferred all specimens with wings ?finely striated in outward and upward curves looking like hairs?, out of the type species H. dutoitides. Yet, Plumstead?s primary reason for the creation of the genus Hirsutum, was to acknowledge the presence of this particular feature, which she interpreted to represent pollenate organs. At this point, the type species of the genus Hirsutum did not possess the key diagnostic characters specified by the generic diagnosis. Plumstead (1958a) had only inferred the presence of pollenate organs in the type species H. dutoitides based on similarities in the wing structure with the secondary or scutoid wing of H. intermittens, which did possess these striations on its primary wing. Plumstead (1958a) apparently considered these taxa to be distinguished only on differences in receptacle shape and dimensions, and she explained the lack of pollenate structures in some specimens of both taxa in terms of these structures 398 being ephemeral or transient. In fact the ?pollenate structures? were missing in every single one of the H. dutoitides specimens. In effect, Plumstead (1958a) gave no valid reasons for the separation of the type species, H. dutoitides from Scutum. The only distinguishing feature she cited for the new genus was that the fertile organs bore hair-like pollenate organs. Since we know that none of the fructifications actually bore pollenate structures, the diagnosis was never valid. At the time, most authors viewed the diagnosis with suspicion, and challenged the wisdom in creating a new genus on the basis of this feature alone. Rigby (1978), Mukherjee et al. (1966) and Banerjee (1984) all concurred that Hirsutum was an unacceptable taxon, and they did not accept the evidence provided by Plumstead (1956a, 1958a) for the existence of pollen bearing organs. They effectively sank the genus, recognising the component species as members of Scutum. But the fact of the matter is that Plumstead?s (1958a) H. dutoitides does belong in a separate genus, and is a distinct taxonomic entity, even though for reasons other than those proposed in the original diagnosis. It could be argued that whatever Plumstead?s reasons were for singling out these fructifications as a separate taxon, her recognition of this entity at a generic level should be honoured, and the name she chose retained, but with hefty emendation of the genus. Perhaps with this in mind, Anderson & Anderson (1985) perpetuated use of the name Hirstum, and included H. dutoitides, H. intermittens, and two new species, H. leslii and H. acadarense within the genus. Anderson & Anderson?s (1985) revised diagnosis of Hirsutum characterised it as having a wing with an extended apex, tapered base, and a ?hirsute appearance?. The type species, H. dutoitides had none of these features, and there were no diagnostic characters unifying the group of fructifications the genus was purported to encompass. This study has revealed that members of H. intermittens and H. leslii have complex wing structures that set them apart from any other known glossopterid fructification, and which should free them from any obligation to the name Hirsutum. 399 However, there is one additional complication involving the relationship between H. dutoitides and H. intermittens. We have already discussed how H. dutoitides has certain features in common with H. intermittens. The secondary wing in H. intermittens (which Plumstead (1958a) referred to as the ?real wing?) is virtually identical to the wing of H. dutoitides, which is presumably why Plumstead initially grouped these fructifications within the same genus. The complication involves the specimen that Plumstead (1958a) assigned as the type for H. dutoitides. Adendorff et al. (2002) reluctantly retained this specimen as the type for Gladiopomum dutoitides, despite the fact it was poorly preserved and the counterpart (showing the sterile surface of the fructification) had been misplaced. After carefully studying the photographs of the counterpart in Anderson & Anderson (1985; p. 250, pl. 75, figs 1&2), and subsequent to the discovery of the elaborate wing structure of Hirsutum intermittens, I am now convinced that the type for H. dutoitides is a specimen of H. intermittens in which the secondary wing is fully exposed, and the primary wing is only visible in the apical part of the fructification, creating the effect of a particularly broad apical spine, the primary diagnostic feature of the genus Gladiopomum. Without damaging the type specimen, there is no way of confirming or disproving this point, but if it is in fact a specimen of H. intermittens, it has far-reaching implications regarding the description of the attached leaves in both species, which in turn will affect perceptions regarding the validity of leaf taxonomy in glossopterid studies. This presents us with a sticky situation. If the type specimen of H. dutoitides is in fact a specimen of H. intermittens, then strictly speaking we should assign a new type specimen and a new specific epithet for H. dutoitides, and all those specimens currently residing within H. intermittens should be transferred to H. dutoitides under the existing type specimen. This taxonomic fiasco would contribute disastrously to an already very confusing situation. I propose, for the sake of clarity and utility and in the spirit of meaningful taxonomy, that we should entirely disband the genus Hirsutum and assign its 400 various species to separate genera. The name Bifariala is proposed for the genus encompassing Hirsutum intermittens and Hirsutum leslii is a junior synonym of Appert?s (1977) genus Elatra, although it should remain within a distinct species. 401 APPENDIX V PUBLICATIONS PRODUCED (Please find copies in back cover). Adendorff, R., Bamford, M.K. and McLoughlin, S. 2003. Liknopetalon: a review of a rare gondwanan, Permian pteridophyte. Review of Palaeobotany and Palynology 126:83-101. Adendorff, R., McLoughlin, S. and Bamford, M.K. 2002. A new genus of ovuliferous glossopterid fructifications from South Africa. Palaeontologia africana 38:1-17. 402 REFERENCES Adendorff, R., McLoughlin, S. & Bamford, M.K. 2002. A new genus of ovuliferous glossopterid fructifications from South Africa. Palaeontologia africana 38:1-17. Adendorff, R., Bamford, M.K. & McLoughlin, S. 2003. Liknopetalon: a review of a rare gondwanan, Permian pteridophyte. Review of Palaeobotany and Palynology 126:83-101. Aitken, G.A. 1994. Permian palynomorphs from the number 5 seam, Ecca Group, Witbank/Highveld coalfields, South Africa. Palaeontologia africana 31:97-109. Alvin, K. & Chaloner, W.G. 1970. Parallel evolution in leaf venation: an alternative view of angiosperm origin. Nature 226:662-663. Anderson, H.M. & Anderson, J.M. 1997. Towards new paradigms in Permo- Triassic Karoo Palaeobotany (and associated faunas) through the past 50 years. Palaeontologia africana 33:11-21. Anderson, J.M. 1977. The biostratigraphy of the Permian and Triassic. Part 3. A review of Gondwana Permian palynology with particular reference to the northern Karoo Basin, South Africa. Memoirs of the Botanical Survey of South Africa 41:1-67. Anderson, J.M. 1999. Towards Gondwana Alive. Gondwana Alive Society, Pretoria. 139 pp. Anderson, J.M. & Anderson, H.M. 1985. Palaeoflora of southern Africa. Prodomus of southern African megafloras Devonian to Lower Cretaceous. A.A. Balkema, Rotterdam. 423 pp. Anderson, J.M. & Anderson, H.M. 2003. Heyday of the gymnosperms: systematics and biodiversity of the Late Triassic Molteno fructifications. Strelizia 15. National Botanical Institute, Pretoria. Anderson, J.M., Anderson, H.M., Archangelsky, S., Bamford, M., Chandra, S., Dettmann, M.E., Hill, R.S., McLoughlin, S. & R?sler, O. 1999. Patterns of plant colonization and diversification on Gondwana and its dispersed continental fragments. African Journal of Earth Sciences 28:145-167. Andrews, H.N. 1970. Index of generic names of fossil plants 1820-1965. Bulletin of the U.S. Geological Survey 1300:1-354. Appert, O. 1977. Die Glossopterisflora der Sakoa in s?dwest-Madagaskar. Palaeontographica 162B:1-50. Arber, E.A.N. 1905. On the sporangium-like organs of Glossopteris browniana. Quarterly Journal of the Geological Society 61:324-338. 403 Archangelsky, S. & Bonetti, M. 1963. Fructificacione de glossopterideas del Permico del Bajo de la Leona, Provincia de Buenos Aires. Ameghiniana 3:29-34. Ash, S. 1981. Glossopterid leaves from the Early Mesozoic of northern Mexico and Honduras. Palaeobotanist 28-29:201-206. Bajpai, U. 1992. Morphological trends in the Gondwana plants. In: Venkatachala, B.S. & Singh, H.P. (eds.), Symposium on four decades of Indian Palaeobotany, Palaeobotanist 40. Bajpai, U. & Singh, V.K. 1986. Araucarioxylon kumarpurensis, a new gymnospermous wood from the Upper Permian of West Bengal. Palaeobotanist 35:53-56. Bajpai, U. & Tewari, R. 1990. Plant fossils from the upper beds of Raniganj Formation in Jharia Coalfield. Palaeobotanist 38:43-48. Bamford, M.K. 1999. Permo-Triassic fossil woods from the South African Karoo Basin. Palaeontologia africana 35:25-40. Bamford, M.K. 2004. Diversity of the woody vegetation of Gondwanan southern Africa. Gondwana Research 7(1):153-164. Bamford, M.K. & Philippe, J.L. 2001. Jurassic-Early Cretaceous Gondwanan homoxylous woods: a nomenclatural revision of the genera with taxonomic notes. Review of Palaeobotany and Palynology 113(4):287-297. Banerjee, M. 1968. On Scutum stowanum Plumstead, the fructification borne by Glossopteris decipiens Feistm. from the Raniganj Stage of India and cuticular study of the fertile and vegetative leaves of G. decipiens Feistm. Bulletin of the Botanical Society of Bengal 22:165-168. Banerjee, M. 1973. Glossopteridean fructifications: 1. Dictyopteridium sporiferum Feistmantel. Bulletin of the Botanical Society of Bengal 27:77-84. Banerjee, M. 1978. Genus Glossopteris Brongniart and its stratigraphic significance in the Palaeozoics of India. Part 1 - a revisional study of some species of the genus Glossopteris. Bulletin of the Botanical Society of Bengal 32:81-125. Banerjee, M. 1984. Fertile organs of the Glossopteris flora and their possible relationship in the line of evolution. Evolutionary Botany and Biostratigraphy, A.K. Ghosh Commem. Vol., pp. 29-59. Benecke, A.K. 1976. Several new forms of Glossopteris fructifications from the Beaufort Daptocephalus-zone (Upper Permian) of Natal, South Africa. Palaeontologia africana 19:97-125. 404 Bennets, K.P. 1965. The flint clay deposits of the area between Pretoria and Belfast, Transvaal. Bulletin of the Geological Survey of South Africa 45:2. Benton, M.J. 1995. Diversification and extinction in the history of life. Science 268 (5207):52-58. Blume, C.L. 1825. Bijdragen tot de Flora van Nederlandsche Indi?. Batavia. p. 293. Botha, B.J.V. & Linstr?m, W. 1978. A note on the stratigraphy of the Beaufort Group in north-western Natal. Transactions of the Geological Society of South Africa 81:35-40. Bredell, J.H. 1974. pH as a guide to the genesis and quality of Transvaal refractory clays. Annals of the Geological Survey 10:23-32. Bredell, J.H. 1979. The nature and distribution of the Karoo sequence on the East and West Rand with special reference to refractory clay deposits. Geokongres 77: Geological Society of South Africa Special Publications 6:151- 160. Brongniart, A. 1828. Prodrome d?une histoire des v?g?taux fossiles. Paris. 223 pp. Bunbury, C.J.F. 1861. Notes on a collection of fossil plants from Nagpur, central India. Quarterly Journal of the Geological Society of London 17:325-346. Cairncross, B. & Cadle, A.B. 1988. Depositional palaeoenvironments of the coal-bearing Permian Vryheid Formation in the east Witbank Coalfield, South Africa. South African Journal of Geology 91:1-17. Chaloner, W.G. 1999. Plant and spore compression in sediments. In: Jones, T.P. & Rowe, N.P. (eds.), Fossil Plants and Spores: modern techniques. The Geological Society of London, Bath. pp 36-40. Chandra, S. 1974. Glossopteris and allied genera ? cuticular studies. In: Surange, K.R., Lakhanpal, R.N., & Bharadwaj, D.C. (eds). Aspects and appraisal of Indian palaeobotany. Birbal Sahni Institute of Palaeobotany, Lucknow. pp. 144-153. Chandra, A. & Srivastava, A.K. 1981. A new species of Arberia from the Lower Gondwana of South Rewa Gondwana Basin, India. Palaeobotanist 28-29:40-45. Chandra, S. & Surange, K.R. 1976. Studies of the reproductive organs of Glossopteris Part 1. Dictyopteridium feistmanteli sp. nov. attached on Glossopteris tenuinervis. Palaeontographica 156B:87-102. 405 Chandra, S. & Surange, K.R. 1977a. Cuticular studies of the reproductive organs of Glossopteris Part 3. Two new female fructifications -Jambadostrobus and Venustostrobus - borne on Glossopteris leaves. Palaeontographica 164B:127-152. Chandra, S. & Surange, K.R. 1977b. Cuticular studies of the reproductive organs of Glossopteris Part 2. Cistella-type fructification Plumsteadiostrobus ellipticus gen. et sp. nov. attached on Glossopteris taenioides Feistmantel. Palaeobotanist 23:61-174. Chandra, S. & Surange, K.R. 1977c. Cuticular studies of the reproductive organs of Glossopteris Part 4. Venustostrobus indicus sp. nov. Palaeobotanist 24:149-160. Chandra, S. & Surange, K.R. 1977d. Fertile bracts and scales of Glossopteris fructifications from the Lower Gondwana of India. Palaeobotanist 24:195-201. Chandra, S. & Surange, K.R. 1979. Revision of the Indian species of Glossopteris. Monograph 2. Birbal Sahni Institute of Palaeobotany, Lucknow. 301 pp. Chatterjee, S. & Sen, J. 1963. A glossopteridean fructification from India. Nature 200:1124. Cridland, A.A. 1963. Glossopteris flora from the Ohio Range, Antarctica. American Journal of Botany 50:186-195. Dana, J.D. 1849. Geology: Appendix L. In: Wilkes' United States Exploring Expedition 1838-1842. C. Sherman, Philadelphia. pp. 714-720. Delevoryas, T. & Person, C.P. 1975. Mexiglossa varia gen. et sp. nov., a new genus of glossopteroid leaves from the Jurassic of Oaxaca, Mexico. Palaeontographica 154B:114-120. Dolianiti, E. 1954. A flora do Gondwana Inferior em Santa Catarina. Notas Preliminaries Estudos Div. Geol. Mineralog. Rio de Janeiro 81:1-5. Dolianiti, E. 1971. A flora do Gondwana Inferior em Santa Catarina VII ? O g?nero Ottokaria. Ottokaria santa-catarinae n. sp. An. Acad. Bras. Cienc. 43(Suplemento):337-342. Doweld, A.B. 2003. New Syllabus of Plant Families (A Plant World System). Geos, Moscow. 700 pp. Doyle, J.A. & Donoghue, M.J. 1986. Seed plant phylogeny and the origin of angiosperms; an experimental cladistic approach. Botanical Review 52:321-431. du Toit, A.L. 1927. The fossil flora of the Upper Karoo beds. Annals of the South African Museum 22:289-420. 406 du Toit, A.L. 1932. Some fossil plants from the Karoo System of South Africa. Annals of the South African Museum 28:370-393. du Toit, A.L. 1937. Our Wandering Continents. Oliver & Boyd, London, 366 pp. Erwin, D.H. 1993. The great Palaeozoic crisis: life and death in the Permian. Columbia University Press, New York, 327 pp. Erwin, D.H. 1994. The Permo-Triassic extinction. Nature (London) 367 (6460):231-236. Erwin, D.H. 1999. Biospheric perturbations during Gondwanan times: from the Neoproterozoic-Cambrian radiation to the end-Permian crisis. Journal of African Earth Sciences 28:115-128. Etheridge, R. Jr. 1894. On the mode of attachment of the leaves or fronds to the caudex in Glossopteris, with remarks on the relation of the genus to its allies; with a note on its stratigraphical distribution in Australia, by T.W. Edgeworth David. Proceedings of the Linnean Society of New South Wales, 1894, 2nd ser. 9:228-258. Fairon-Demaret, M., Hilton, J. & Berry, C.M. 1999. Surface preparation of macrofossils (d?gagement). In: Jones, T.P. & Rowe, N.P. (eds.), Fossil Plants and Spores: modern techniques. The Geological Society of London, Bath. pp 33-35. Falcon, R.M.S. 1986. A brief review of the origin, formation, and distribution of coal in Southern Africa. In: Anheusser, C.R. & Maske, S. (eds). Mineral Deposits of Southern Africa. Vols I & II. Geological Society of South Africa, Johannesburg. Feistmantel, O. 1879. The fossil flora of the Gondwana System - The flora of the Talchir-Karharbari beds. Memoirs of the Geological Survey of India, Palaeontologia Indica (12) 3(1):1-48. Feistmantel, O. 1881. The fossil flora of the Gondwana System. The flora of the Talchir - Karharbari beds. Memoirs of the Geological Survey of India. Palaeontologia Indica (12), 3 (1) (Supplement):49-64. Florin, R. 1944. Die Koniferen des Oberkarbons und des unteren Perms. Pt. 7. Palaeontographica 85B:457-654. Francis, J.E., Woolfe, K.J., Arnot, M.J. & Barrett, P.J. 1993. Permian forests of Alan Hills, Antarctica: the palaeoclimate of Gondwanan high latitudes. Special Papers in Palaeontology 49:75-83. Geertsema, H., van Dijk, D.E. & van den Heever, J.A. 2002. Palaeozoic insects of Southern Africa: a review. Palaeontologia africana 38:19-25. 407 Gould, R.E. 1975. A preliminary report on petrified axes of Vertebraria from the Permian of eastern Australia. In: Campbell, K.S.W. (ed.). Gondwana Geology. A.N.U. Press, Canberra. pp. 109-115. Gould, R.E. & Delevoryas, T. 1977. The biology of Glossopteris: evidence from petrified seed-bearing and pollen-bearing organs. Alcheringa 2:387-399. Greuter, W. et al. 1994. International Code of Botanical Nomenclature (Tokyo Code). Regnum Vegetabile 131. Koeltz Scientific Books, K?ningstein. Groenewald, G.H. 1984. Stratigrafie en sedimentologie van die Groep Beaufort in die noordoos Vrystaat. M.Sc. dissertation, Rand Afrikaans University, Johannesburg (unpubl.). Groenewald, G.H. 1989. Stratigrafie en sedimentologie van die Groep Beaufort in die noordoos Vrystaat. Geological Survey of South Africa Bulletin 96. Guerra-Sommer, M. & Cazzulo-Klepzig, M. 2000. As Floras Gondu?nicas do Paleoz?ico Superior do Rio Grande do Sul. In: Holz, M. & De Ros, L.F. (eds). Paleontologia do Rio Grande do Sul. Edi??o CIGO/UFRGS. 398 pp. Holmes, W.B.K. 1974. On some fructifications of the Glossopteridales from the Upper Permian of New South Wales. Proceedings of the Linnean Society of New South Wales 98:131-141. Holmes, W.B.K. 1990. Austroglossa walkomii Holmes, a glossopterid ovulate fructification from the Late Permian of New South Wales. Proceedings, 3IOP Conference, Melbourne 1988:67-73. Holmes, W.B.K. 1992. Glossopteris-like leaves from the Triassic of eastern Australia. Geophytology 22:119-125. Holmes, W.B.K. 1995. The Late Permian megafossil flora from Cooyal, New South Wales, Australia. In: Pant, D.D. (ed.). Global Environment and Diversification of Plants through Geological Time. Birbal Sahni Centenary Volume, Allahabad. pp. 123-152. Jeyasingh, D.E.P. 1987. A plea for precise descriptions of glossopterid leaf compressions. Journal of the Swamy Botanical Club 4 (4):173-179. Johnson, M.R. (ed.). 1994. Lexicon of South African Stratigraphy. Part 1: Phanerozoic units. South African Committee for Stratigraphy, Council for Geoscience, Pretoria. 56 pp. Johnson, M.R., Van Vuuren, C.J., Hegenberger, W.F., Key, R. & Shoko, U. 1996. Stratigraphy of the Karoo Supergroup in southern Africa: an overview. Journal of African Earth Sciences 23:3-15. 408 Johnson, M.R., Van Vuuren, C.J., Visser, J.N.J., Cole, D.I., Wickens, H. de V., Christie, A.D.M. & Roberts, D.L. 1997. The Foreland Karoo Basin, South Africa. In: R.C. Selley (ed.), African Basins. Sedimentary Basins of the World, 3, Elsevier Science B.V., Amsterdam. pp. 269-317. Kerr, R.A. 1993. The greatest extinction gets greater. Science 262 (5138):1370-1371. Kerr, R.A. 2000. Biggest extinction hit land and sea. Science 289 (5485):1666- 1667. Keyser, N. (compiler). 1997. Geological Map of the Republic of South Africa. Council for Geoscience, Pretoria. Kitching, J. 1977. The distribution of the Karoo vertebrate fauna. Memoir of the Bernard Price Institute for Palaeontological Research, University of the Witwatersrand 1:1-131. Kov?cs-Endr?dy, ?. 1976. Notes on some Glossopteris species from Hammanskraal (Transvaal). Palaeontologia africana 19:67-95. Kov?cs-Endr?dy, ?. 1977. The systematic position of the genus Gangamopteris McCoy. Geological Survey of South Africa Bulletin 60:19-71. Kov?cs-Endr?dy, ?. 1979. A re-evaluation of the venation structure of Glossopteris. Annals of the Geological Survey of South Africa 12:107-141. Kov?cs-Endr?dy, ?. 1981. "Broad sense interpretation" of Glossopteris leaves: a critique. Palaeontologia africana 24:35-38. Kov?cs-Endr?dy, ?. 1984. Notes on Glossopteris conspicua Feistmantel and on its assumed stratigraphic significance. Annals of the Geological Survey of South Africa 17:69-85. Kov?cs-Endr?dy, ?. 1991. On the Late Permian age of Ecca Glossopteris floras in the Transvaal Province with a key to and description of twenty five Glossopteris species. Memoir 77, 111p. Geological Survey of South Africa. Kr?usel, R., Maithy, P.K. & Maheshwari, H.K. 1961-1962. Gymnospermous woods with primary structures from Gondwana rocks ? a review. Palaeobotanist 10:97-107. Kyle, R.A. 1974. Plumsteadia ovata n. sp., a glossopterid fructification from South Victoria Land, Antarctica. New Zealand Journal of Geology and Geophysics 17:719-721. Lacey, W.S. 1959. Occurrence of presumed Glossopteridean fructifications in Rhodesia and Nyasaland. Nature 184:1592-1593. 409 Lacey, W.S. 1974. New Permian Glossopteris flora from Natal. South African Journal of Science 70:154-156. Lacey, W.S. 1978. A review of the Upper Permian Glossopteris flora in western Natal. Palaeobotanist 25:185-189. Lacey, W.S. & Huard-Moine, D. 1966. Karroo floras of Rhodesia and Malawi- Part 2. The Glossopteris flora in the Wankie district of Southern Rhodesia. Symposium on Floristics and Stratigraphy of Gondwanaland. Birbal Sahni Institute of Palaeobotany, Lucknow. pp. 13-25. Lacey, W., van Dijk, D.E. & Gordon-Gray, K.D. 1975. Fossil plants from the Upper Permian in the Mooi River district of Natal, South Africa. Annals of the Natal Museum 22:349-420. Lambrecht, L.L., Lacey, W.S., & Smith, C.S. 1972. Observations on the Permian flora of the Law Glacier area central Transantarctic Mountains. Bulletin of the Society for Belgian Geolology, Paleontology and Hydrology 81(3-4):161-167. Leaf Architecture Working Group. 1999. Manual of Leaf Architecture - morphological description and categorization of dicotyledonous and net-veined monocotyledonous angiosperms. Smithsonian Institution. 65 pp. Le Roux, S.F. 1976. On some 'northern' elements in the Lower Gondwana flora of Vereeniging, Transvaal. Palaeontologia africana 19:59-65. Le Roux, S.F. & Anderson, H.M. 1977. A review of the localities and flora of the Lower Permian Karoo strata at Vereeniging, South Africa. Palaeontologia africana 20:27-42. Leslie, T.N. 1904. The fossil flora of Vereeniging. Transactions of the Geological Society Vol. VI:82-88. Leslie, T.N. 1921. Observations on some fossil plants from the Permo-Carboniferous of Vereeniging. Proceedings of the Geological Society of South Africa 25:19-30. Lindstr?m, S., McLoughlin, S. & Drinnan, A,N. 1997. Intraspecific variation of taeniate bisaccate pollen within Permian glossopterid sporangia, from the Prince Charles Mountains, Antarctica. International Journal of Plant Science, 158:673-684. Lindstr?m, W. 1973. Die geologie van die sisteem Karoo wes van Mooirivier, Natal. M.Sc. thesis, University of the Orange Free State, Bloemfontein (unpubl.) Lindstr?m, W. 1987. Die geologie van die gebied Durban: Toeligting van blad 2930. Departement van Mineraal- en Energiesake, Pretoria. pp. 19-21. 410 Lundqvist, G. 1919. Fossil Pflanzen der Glossopteris flora aus Brasilien. Kungliga Svenska Vetenskapsakademiens Handlingar 60 (3):1-36. Maheshwari, H.K. 1965a. Studies in the Glossopteris Flora of India ? 31. Some remarks on the genus Glossopteris Sternb. Palaeobotanist 14:36-45. Maheshwari, H.K. 1965b. Studies in the Glossopteris flora of India - 23. On two fructifications from Raniganj stage of the Raniganj Coalfield, Bengal. Palaeobotanist 13:144-147. Maheshwari, H.K. 1968. Gonophylloides nom. nov. Taxon 17:238-239. Maheshwari, H.K. 1972. Permian wood from Antarctica and revision of some Lower Gondwana wood taxa. Palaeontographica 138B:1-43. Maheshwari. H.K. 1976. Floristics of the Permian and Triassic Gondwanas of India. Palaeobotanist 23(2):145-160. Maheshwari, H.K. 1990. The glossopterid fructifications: an overview. In: Douglas, J.G. & Christophel, D.C. (eds). Proceedings of the 3rd International Organization of Palaeobotany Conference (Melbourne, 1988). pp. 11-15. Maheshwari, H.K. & Tewari, R. 1992. Epidermal morphology of some Indian species of the genus Glossopteris Brongniart. Palaeobotanist 39:338-380. Maithy, P.K. 1965. Studies in the Glossopteris flora of India - 18. Gymnospermic seeds and seed-bearing organs from the Karharbari beds of the Giridih Coalfield, Bihar. Palaeobotanist, 13:45-56. Maithy, P.K. 1970. Studies in the Glossopteris flora of India - 39. On some new plant fossils from the Karharbari Beds, Giridih Coalfield, India. Palaeobotanist 18:167-172. McLoughlin, S. 1990a. Some Permian glossopterid fructifications and leaves from the Bowen Basin, Queensland, Australia. Review of Palaeobotany and Palynology 62:11-40. McLoughlin, S. 1990b. Late Permian glossopterid fructifications from the Bowen and Sydney Basins, eastern Australia. Geobios 23(3):283-297. McLoughlin, S. 1992. Late Permian plant megafossils from the Bowen Basin, Queensland, Australia: Part 1. Palaeontographica 228B:105-149. McLoughlin, S. 1993a. Glossopterid megafossils in Permian Gondwanic non-marine biostratigraphy. In: Findlay, R.H., Unrug, R., Banks, M.R. & Veevers, J.J. eds. Gondwana Eight: Assembly, Evolution and Dispersal. Rotterdam: A.A. Balkema, pp. 253-264. McLoughlin, S. 1993b. Plant fossil distributions in some Australian Permian non-marine sediments. Sedimentary Geology 85: 601-619. 411 McLoughlin, S. 1993c. Late Permian plant megafossils from the Bowen Basin, Queensland, Australia: Part 2. Palaeontographica 231B:1-29. McLoughlin, S. 1995. Bergiopteris and glossopterid fructifications from the Permian of Western Australia and Queensland. Alcheringa 19:175-192. McLoughlin, S. & Drinnan, A.N. 1996. Anatomically preserved Noeggerathiopsis leaves from east Antarctica. Review of Palaeobotany and Palynology 92:207-227. McLoughlin, S., Drinnan, A.N. & Lindstr?m, S. 1997. Gondwanan floristic and sedimentological trends during the Permian-Triassic transition: new evidence from the Amery Group, northern Prince Charles Mountains, East Antarctica. Antarctic Science 9:281-298. McManus, H.A., Taylor, E.L., Taylor, T.N. & Collinson, J.W. 2002. A petrified Glossopteris flora from Collinson Ridge, central Transantarctic Mountains: Late Permian or Early Triassic? Review of Palaeobotany and Palynology 120:233- 246. Melville, R. 1960. A new theory of the angiosperm flower. Nature 188:14-18. Melville, R. 1962. A new theory of the angiosperm flower. Kew Bulletin 16:1-63. Melville, R. 1969. Leaf venation patterns and the origin of the angiosperms. Nature 224:121-125. Melville, R. 1970. Links between the glossopterideans and the angiosperms. Proceedings and Papers, 2nd Gondwana Symposium, Johannesburg. Geological Society of South Africa, Johannesburg. pp. 585-588. Melville, R. 1983a. Two new genera of Glossopteridae. Botanical Journal of the Linnaean Society 86:275-277. Melville, R. 1983b. Glossopteridae, Angiospermidae and the evidence for angiosperm origins. Botanical Journal of the Linnaean Society 86:279-323. Men?ndez , C.A. 1962a. Prencia de Ottokaria en la flora de Glossopteris de la Provincia de Buenos Aires. Review of the Geological Association of Argentina 17(1-2):5-9. Men?ndez , C.A. 1962b. Hallazgo de una fructificacion en la flora de Glossopteris de la Provincia de Buenos Aires (Lanceolatus bonariensis sp. nov.), con consideraciones sobre la nomenclatura de fructificaciones de Glossopteris. Ameghiniana 2:175-182. Meyen, S.V. 1984. Basic features of gymnosperm systematics and phylogeny as evidenced by the fossil record. Botanical Review 50(1):1-111. 412 Meyen, S.V. 1987. Fundamentals of Palaeobotany. Chapman & Hall, London, 432 pp. Millan, J.H. 1967. Novas fructifica??es na flora Glossopteris do Gondwana inferior do Brasil. Brasil Departamento nacional da producioen mineral, Divis?o de geologia e mineralogia, Notas preliminares e estudos 140:1-19. Millsteed, B.D. 1994. Palynological evidence for the age of the Permian Karoo coal deposits near Vereeniging, northern Orange Free State, South Africa. South African Journal of Geology 97:15-20. Mukherjee, S., Banerjee, M., & Sen, J. 1966. Further glossopterid fructifications from India. Palaeontographica 117B:99-113. Nannfeldt, J.A. 1932. Studien ?ber die Morphologie und Systematik der nichtlichenisierten Operculaten Discomyceten. Nova Acta Regiae Soc. Scient. Upsal. Ser. IV, 8: 265 et seq. Neish P.G., Drinnan A.N. & Cantrill D.J. 1993. Structure and ontogeny of Vertebraria from silicified Permian sediments in East Antarctica. Review of Palaeobotany and Palynology 79(3-4):221-244. Nishida, H., Kudo, K., Pigg, K.B. & Rigby, J.F. 2002. New characters of reproductive organs of Glossopteris in relation to its phylogeny. Abstract from the Sixth Conference of the International Organisation of Palaeobotany, Qinhuandao, Hebei Province, China. Nishida, H., K., Pigg, Kudo, K.B. & Rigby, J.F. 2004. Zooidogamy in the Late Permian genus Glossopteris. Journal of Plant Research 117:323-328. Nishida, H., Pigg, K.B. & Rigby, J.F. 2003. Swimming sperm in an extinct Gondwanan plant. Nature 422:396-397. Pant, D.D. 1958. The structure of some leaves and fructifications of the Glossopteris flora of Tanganyika. Bulletin of the British Museum (Nat. Hist.) Geology 3:125-176. Pant, D.D. 1962. Some recent contributions towards our knowledge of the Glossopteris flora. In: Maheshwari, P., Johri, B.M. & Vasil, I.K. (eds), Proceedings of the Summer School of Botany ? Darjeeling, 1960. pp. 302-319. Pant, D.D. 1977. The plant of Glossopteris. Journal of the Indian Botanical Society 56:1-23. Pant, D.D. 1982. The Lower Gondwana gymnosperms and their relationship. Review of Palaeobotany and Palynology 37:55-70. Pant, D.D. 1996. The biogeography of the late Palaeozoic floras of India. Review of Palaeobotany and Palynology 90 (1-2):79-98. 413 Pant, D.D. 1999. Dominant gymnosperms of the Glossopteris flora. Palaeobotanist 48:111-123. Pant, D.D. & Gupta, K.L. 1968. Cuticular structure of some Indian Lower Gondwana species of Glossopteris Brongniart. Part 1. Palaeontographica 124B:45-81. Pant, D.D. & Gupta, K.L. 1971. Cuticular structure of some Indian Lower Gondwana species of Glossopteris Brongniart. Part 2. Palaeontographica 132B:130-152. Pant, D.D. & Nautiyal, D.D. 1960. Some seeds and sporangia of the Glossopteris flora from Raniganj Coalfield, India. Palaeontographica 107B:41-64. Pant, D.D. & Nautiyal, D.D. 1965. Seed-bearing Ottokaria-like fructifications from India. Nature 207:623-624. Pant, D.D. & Nautiyal, D.D. 1966. On two peculiar fossils of Karharbari Stage, India. Symposium on the floristics and stratigraphy of Gondwanaland. Birbal Sahni Institute of Palaeobotany, Lucknow. pp. 98-101. Pant, D.D. & Nautiyal, D.D. 1984. On the morphology and structure of Ottokaria zeilleri sp. nov. - a female fructification of Glossopteris. Palaeontographica 193B:127-152. Pant, D.D. & Pant, R. 1987. Some Glossopteris leaves from Indian Triassic beds. Palaeontographica 205B:165-178. Pant, D.D. & Singh, R.S. 1971. Cuticular structure of some Indian Lower Gondwana species of Glossopteris Brongniart. Part 3. Palaeontographica 135B:1-40. Pant, D.D. & Singh, R.S. 1974. On the stem and attachment of Glossopteris and Gangamopteris leaves Part 2 - structural features. Palaeontographica 147B:42-73. Pant, D.D. & Singh, V.K. 1987. Xylotomy of some woods from Raniganj Formation (Permian), Raniganj Coalfield, India. Palaeontographica 203B:1-82. Philippe, M. 1993. Nomenclature g?n?rique des trach?idoxyles fossiles m?sozo?ques ? champs araucario?des. Taxon 42:72-80. Pigg, K. B. 1990. Anatomically preserved Glossopteris foliage from the central Transantarctic Mountains. Review of Palaeobotany and Palynology 66:105-127. Pigg, K.B. & McLoughlin, S. 1992. Comparison of anatomically preserved glossopterid leaves from the Bowen Basin, Queensland, and Sydney Basin, New South Wales. 4th International Organization of Palaeobotany Conference, Paris 1992. (Abstract). 414 Pigg, K.B. & McLoughlin, S. 1997. Anatomically preserved Glossopteris leaves from the Bowen and Sydney basins, Australia. Review of Palaeobotany and Palynology 97:339-359. Pigg, K.B. & Taylor, T.N. 1990. Permineralized Glossopteris and Dicroidium from Australia. In: Taylor, T.N. & Taylor, E.L. (eds), Antarctic Palaeobiology: Its Role in Reconstructing Gondwana. Springer-Verlag, New York. pp. 164-172. Pigg, K.B. & Taylor, T.N. 1993. Anatomically preserved Glossopteris stems with attached leaves from the central Transantarctic Mountains, Antarctica. American Journal of Botany 80:500-516. Pigg, K. B. & Trivett, M. L. 1994. Evolution of the glossopterid gymnosperms from Permian Gondwana. Journal of Plant Research 107:461-477. Plumstead, E.P. 1952. Description of two new genera and six new species of fructifications borne on Glossopteris leaves. Transactions of the Geological Society of South Africa 55:281-328. Plumstead, E.P. 1956a. Bisexual fructifications borne on Glossopteris leaves from South Africa. Palaeontographica 100B:1-25. Plumstead, E.P. 1956b. On Ottokaria, the fructification of Gangamopteris. Transactions of the Geological Society of South Africa 59:211-236. Plumstead, E.P. 1957. Coal in Southern Africa. Witwatersrand University Press, Johannesburg. 25 pp. Plumstead, E.P. 1958a. Further fructifications of the Glossopteridae and a provisional classification based on them. Transactions of the Geological Society of South Africa 61:1-58. Plumstead, E.P. 1958b. The habit of growth of Glossopteridae. Transactions of the Geological Society of South Africa 61:81-96. Plumstead, E.P. 1962a. Fossil floras of Antarctica. Trans-Antarctica Expedition 1955-1958, Scientific Reports 9:1-154. Plumstead, E. P. 1962b. Possible angiosperms from Lower Permian coal of the Transvaal. Nature 194:594-595. Plumstead, E.P. 1962c. Vannus gondwanensis, a new Gangamopteris fructification from the Transvaal, South Africa. Palaeobotanist 11(1,2):106-113. Plumstead, E.P. 1969. Three thousand million years of plant life in Africa. Alex L Du Toit Memorial Lectures 11. Geological Society of South Africa. 72 pp. Plumstead, E.P. 1975. A new assemblage of fossil plants from Milorgfjella, Dronning Maud Land. British Antarctic Survey Scientific Reports 83:1-30. 415 Potter, P.E., Maynard, J.B. & Pryor, W.A. 1980. Sedimentology of shale. Springer-Verlag, New York. 301 pp. Raup, D.M. & Sepkoski, J.J. Jr. 1982. Mass extinctions in the marine fossil record. Science 215 (4539):1501-1503. Rayner, R.J. & Coventry, M.K. 1985. A Glossopteris flora from the Permian of South Africa. South African Journal of Science 81(1):21-32. Retallack, G.J. 1980. Late Carboniferous to Middle Triassic megafossil floras from the Sydney Basin. In: Herbert, C. & Helby, R.J. (eds.), A Guide to the Sydney Basin. Geological Survey of N.S.W. Bulletin 26:384-430. Retallack, G.J. 1995. Permian-Triassic life crisis on land. Science 267 (5194):77-80. Retallack, G.J. & Dilcher, D.L. 1981. Arguments for a glossopterid ancestry of angiosperms. Palaeobiology 7:54-67. Retallack, G.J. & Dilcher, D.L. 1988. Reconstructions of selected seed ferns. Annals of the Missouri Botanical Gardens 75:1010-1057. Retallack, G.J., Veevers J.J. & Morante, R. 1996. Global coal gap between Permian-Triassic extinction and Middle Triassic recovery of peat-forming plants. Geological Society of America Bulletin 108 (2):195-207. Rex, G.M. 1986. Experimental modelling as an aid to interpreting the original three-dimensional structures of compressions. In: Spicer, R.A. & Thomas, B.A. (eds). Systematic and taxonomic approaches in palaeobotany. The Systematic Association Special Volume no. 31. Clarendon Press, Oxford. pp. 17-36. Riek, E.F. 1973. Fossil insects from the Upper Permian of Natal, South Africa. Annals of the Natal Museum 21(3):513-532. Riek, E.F. 1976. New Upper Permian insects from Natal, South Africa. Annals of the Natal Museum 22(3):755-789. Rigby, J.F. 1963. On a collection of plants of Permian age from Baralaba, Queensland. Proceedings of the Linnaean Society of New South Wales 87:341-351. Rigby, J.F. 1967. On Gangamopteris walkomii sp. nov. Records of the Australian Museum 27:175-182. Rigby, J.F. 1969. The conservation of Plumsteadia Rigby 1963 over Cistella Plumstead 1958. Bolm. Soc. Bras. Geol. 17: 93. Rigby, J.F. 1971. A revision of some plants from the Permian of the Bowen Basin, Queensland. Geological Survey of Queensland Publication 349, Palaeontological Paper 23:1-8. 416 Rigby, J.F. 1972a. On Arberia White, and some related Lower Gondwana female fructifications. Palaeontology 15:108-120. Rigby, J.F. 1972b. The flora of the Kaloola Member of the Baralaba Coal Measures, central Queensland. Geological Survey of Queensland Publication 352, Palaeontology Paper 26:1-12. Rigby, J.F. 1978. Permian glossopterid and other cycadopsid fructifications from Queensland. Geological Survey of Queensland Publication 367, Palaeontology Paper 41:1-21. Rigby, J.F. 1984. The origin of the Glossopteris flora - some thoughts based on macrophyte remains. In: Sharma, A.K. (ed.), Proceedings of a symposium on evolutionary botany and biostratigraphy. A.K. Ghosh Commemoration Volume, Today and Tomorrows Printers and Publishers, New Delhi. pp. 19-28. Rigby, J.F. & Schopf, J.M. 1969. Stratigraphic implications of Antarctic palaeobotanical studies. In: Amos, A.J. (ed.), Gondwana Stratigraphy. UNESCO, Paris, pp. 91-106. Rubidge, B.S. 1995. Biostratigraphy of the Beaufort Group (Karoo Supergroup), South African Committee for Stratigraphy, Biostratigraphic Series, No. 1. 46 pp. SACS See ?South African Committee for Stratigraphy? below. Schopf, J.M. 1967. Antarctic fossil plants collected during the 1966-67 season. United States Antarctic Journal 2:114-116. Schopf, J.M. 1968. Reinterpretation of the glossopterid sporophyll. American Journal of Botany 55 (6):726. Schopf, J.M. 1970. Petrified peat from a Permian coal bed in Antarctica. Science 169:274-275. Schopf, J.M. 1975. Modes of fossil preservation. Review of Palaeobotany and Palynology 20:27-53. Schopf, J.M. 1976. Morphologic interpretation of fertile structures in glossopterid gymnosperms. Review of Palaeobotany and Palynology 21:25-64. Seward, A.C. 1897. On the association of Sigillaria and Glossopteris in South Africa. Quarterly Journal of the Geological Society of London 53:315-340. Seward, A.C. 1898. Note on the plant remains from Vereeniging. Quarterly Journal of the Geological Society of South Africa 54:92-93. Seward, A.C. 1903. Fossil floras of Cape Colony. Annals of the South African Museum 4:1-222 417 Seward, A.C. 1904. Report on collections of Natal fossil plants. From 1: The ecca coal series of Umhlali on the North-East coast of Natal. 2: The Drakensberg range in West Natal. Report of the Geological Survey of Natal and Zululand 2:97-104. Seward, A.C. 1907. On a collection of Permo-Carboniferous plants from the St. Lucia (Somkele) Coalfield, Zululand, and from the Newcastle District, Natal. Transactions of the Geological Society of South Africa 10:81-89. Seward, A.C. 1917. Fossil Plants 3. Pteridospermae, Cycadofilicales, Cordaitales, Cycadophyta. Cambridge University Press, Cambridge. 656 pp. Seward, A.C. & Sahni, B. 1920. Indian Gondwana plants: a revision. Memoirs of the Geological Survey of India. Palaeontologia Indica, N.S. 7(1):1-55. Singh, S.M. 2000. Taxonomy and diversity of the genus Glossopteris. Palaeobotanist 49:333-352. Singh, G. & Shah, S.C. 1966. Fossil plants from the Sakoa Series, Madagascar. C.R. Sem. Geol., Madagascar 1966:17-19. Smithies, S.J., 1978. Studies in a Middle Ecca (Lower Permian) flora from Hammanskraal, Transvaal, South Africa with emphasis on the glossopterid fructification Ottokaria Zeiller. MSc. Dissertation (unpubl.), University of the Witwatersrand, Johannesburg. Smithies, S.J., 1985. Glossopteridales: Hirsutum leslii. In: Anderson, H.M. & Anderson, J.M. (eds.). The Palaeoflora of Southern Africa: Prodromus of Southern African Megafloras, Devonian to Lower Cretaceous. A.A. Balkema, Rotterdam. pp. 7, 121. South African Committee for Stratigraphy (SACS). 1980. Stratigraphy of South Africa. Part 1: Lithostratigraphy of the Republic of South Africa, South West Africa/ Namibia, and the Republics of Bophuthatswana, Transkei and Venda: Handbook of the Geological Survey of South Africa 8, 690pp. Srivastava, A.K. 1956. Glossopteris, Gangamopteris and Palaeovittaria from the Raniganj Coalfields. Palaeobotanist 5:1-45. Srivastava, A.K. 1978. Studies in the Glossopteris flora of India - 43. Some new plant fossils from the lower Gondwana sediments of Auranga Coalfield, Bihar. Palaeobotanist 25:486-495. Stearn, W.T. 1973. Botanical Latin. David & Charles, Newton Abbot. Stebbins, G.L. 1974. Flowering plants: evolution above the species level. Arnold, London. Stewart, W.N. & Rothwell, G.W. 1993. Palaeobotany and the evolution of plants. 2nd ed., Cambridge University Press. 418 Surange, K.R. & Chandra, S. 1973a. Dictyopteridium sporiferum Feistmantel - female cone from the Lower Gondwana of India. Palaeobotanist 20:127-136. Surange, K.R. & Chandra, S. 1973b. Denkania indica gen. et sp. nov., a glossopteridean fructification from the Lower Gondwana of India. Palaeobotanist 20:264-268. Surange, K.R. & Chandra, S. 1973c. Partha, a new type of female fructification from the Lower Gondwana of India. Palaeobotanist 20:356-360. Surange, K.R. & Chandra, S. 1974a. Fructifications of Glossopteridae from India. Palaeobotanist 21:1-17. Surange, K.R. & Chandra, S. 1974b. Lidgettonia mucronata sp. nov: a female fructification from the Lower Gondwana of India. Palaeobotanist 21:121-126. Surange, K.R. & Chandra, S. 1974c. Further observations on Glossotheca Surange & Maheshwari: a male fructification of Glossopteridales. Palaeobotanist 21:248-254. Surange, K.R. & Chandra, S. 1974d. Some male fructifications of Glossopteridales. Palaeobotanist 21:255-266. Surange, K.R. & Chandra, S. 1975. Morphology of the gymnospermous fructifications of the Glossopteris flora and their relationships. Palaeontographica 149B:153-180. Surange, K.R. & Chandra, S. 1978. Morphology and affinities of Glossopteris. Palaeobotanist 25:509-524. Surange, K.R. & Lele, K.M. 1956. Studies in the Glossopteris flora of India - 6. Plant fossils from Talchir Beds of South Rewa Gondwana Basin. Palaeobotanist 5: 82-90. Surange, K.R. & Maheshwari, H.K. 1970. Some male and female fructifications of Glossopteridales from India. Palaeontographica, 129B:178-192. Surange, K.R. & P.N. Srivastava. 1956. Studies in the Glossopteris flora of India - 5. Generic status of Glossopteris, Gangamopteris and Palaeovittaria. Palaeobotanist 5:46-49. Taverner-Smith, R., Cooper, J.A.J. & Rayner, R.J. 1988. Depositional environments in the Volksrust Formation (Permian) in the Mdlatuze River, Zululand. South African Journal of Geology 91:198-206. Taylor, E.L. 1996. Enigmatic gymnosperms? Structurally preserved Permian and Triassic seed ferns from Antarctica. Review of Palaeobotany and Palynology 90:303-318. 419 Taylor, E.L. & Taylor, T.N. 1992. Reproductive biology of the Permian Glossopteridales and their suggested relationship to flowering plants. Proceedings of the National Academy of Science, USA 89:11495-11497. Taylor, E.L., Taylor, T.N. & C?neo, N.R. 1992. The present is not the key to the past: A polar forest from the Permian of Antarctica. Science 257:1675-1677. Taylor, T.N. 1981. Paleobotany: an introduction to fossil plant biology. McGraw- Hill Book Company, New York. 561 pp. Taylor, T.N. & Taylor, E.L. 1993. The Biology and Evolution of Fossil Plants. Prentice Hall, Englewood Cliffs, New Jersey. 982 pp. Thomas, H. H. 1921. An Ottokaria-like plant from South Africa. Quarterly Journal of the Geological Society vol. lxxvii, pt. 4. pp 285-288. Thomas, H.H. 1952. A Glossopteris with whorled leaves. Palaeobotanist 1:435-438. Thomas, H.H. 1958. Lidgettonia, a new type of fertile Glossopteris. Bulletin of the British Museum (Natural History) 3:179-89. van Dijk, D.E. 1981. A study of the type locality of Lidgettonia africana Thomas 1958. Palaeontologia africana 24:43-61. van Dijk, D.E. 1998. Insect Faunas of South Africa From the Upper Permian and the Permian/Triassic Boundary. Palaeontologia africana 34:34-48. van Dijk, D.E. 2000. Contributions to knowledge of some southern African fossil sites and their fossils. M.Sc. dissertation (unpubl.), University of Stellenbosch, Stellenbosch. van Dijk, D.E. & Geertsema, H. 1999. Permian insects from the Beaufort Group of Natal, South Africa. Annals of the Natal Museum 40:137-171. Veevers, J.J., Cole, D.I. & Cowan, E.J. 1994. Southern Africa: Karoo Basin and Cape Fold Belt. In: Veevers, J.J. & Powell, C.McA. (eds.), Permian-Triassic Pangean basins and foldbelts along the Panthalassan margin of Gondwanaland. Boulder, Colorado, Geological Society of America Memoir 184. Walkom, A.B. 1922. Palaeozoic floras of Queensland. part 1. The flora of the Lower and Upper Bowen Series. Geological Survey of Queensland Publication 270: 1-65. Weaver, L., McLoughlin, S. & Drinnan, A.N. 1997. Fossil woods from the Upper Permian Bainmedart Coal Measures, northern Prince Charles Mountains, East Antarctica. AGSO Journal of Australian Geology & Geophysics 16 (5):655- 676. Wesley, A. 1963. The status of some fossil plants. Advances in Botany 1:1-72. 420 White, D. 1908. Report on the fossil flora of the coal measures of Brazil. Relatorio final Comisseo de Estudos das Minas de Carveo de Pedra do Brazil. Imprensa Nacional Rio de Janeiro 3:336-617. White, M.E. 1961. Plant fossils in core samples from AAO 5 Roma Bore, north Queensland. Bureau of Mineral Resources of Australia Record 1961/18:1-4. (unpubl.). White, M.E. 1963. Reproductive structures in Australian Upper Permian Glossopteridae. Proceedings of the Linnean Society of N.S.W. 88:392-396. White, M.E. 1978. Reproductive structures of the Glossopteridales in the plant fossil collection of the Australian Museum. Records of the Australian Museum 31 (12):473-504. White, M.E. 1986. The Greening of Gondwana. Reed Books, Frenchs Forest, N.S.W., Australia. 256 pp. Zeiller, M.R. 1896. Etude sur Quelques plantes fossiles en particulier Vertebraria et Glossopteris des environs de Johannesburg (Transvaal). Bulletin of the Geological Society of France 24:349-378. Zeiller, R. 1902. Observations sur quelques plantes fossiles des Lower Gondwanas. Memoirs of the Geological Survey of India, Palaeontologia Indica. N.S. 2:1-40. Zhao, L., Taylor, T.N. & Taylor, E.L. 1995. Cupulate glossopterid seeds from the Permian Buckley Formation, central Transantarctic Mountains. Antarctic Journal of the United States 30:54-55. 421 422 PLATES Table P.1. Alphabetical list of glossopterid ovulate fructifications with chapter and plate references. TAXON CHAPTER PAGE PLATES Arberia hlobanensis 6.1 171 10 a, b; 11 h, i Arberia madagascarensis 6.1 167 10 e, f; 11 j, k Bifariala intermittens 7.1 186 12 ? 18 Dictyopteridium flabellatum 7.9 305 87 - 88; 89 e-g Dictyopteridium natalensis 7.9 301 83 d-r; 84 - 86; 89 c, d cf. Dictyopteridium sporiferum 7.9 299 83 a-c; 89 a, b Elatra leslii 7.3 204 23 ? 42 Estcourtia conspicua 7.2 195 19 ? 22 Gladiopomum acadarense 7.6 254 61 - 66; 69 a, b, e(ii) Gladiopomum dutoitides 7.6 249 59 - 60; 68; 69 d, e(i) Gladiopomum elongatum 7.6 257 67; 69 c, e(iii) Gonophylloides strictum 7.8 284 79 - 80; 81 a-d; 82 b-d Gonophylloides waltonii 7.8 287 81 e-I; 82 a incertae sedis (prev. Arberia allweyensis) 5.2 157 10 c, d; 11 d, e, f Lidgettonia africana 8.1 316 90 - 95; 100 a-c Lidgettonia elegans 8.1 321 98 - 99; 100 f, g Lidgettonia lidgettonioides 8.1 319 96 - 97; 100 d, e Ottokaria buriadica 7.4 221 45 - 46; 47 a-c; 48 c Ottokaria hammanskraalensis 7.4 219 47 d, e; 48 b Ottokaria transvaalensis 7.4 216 43 - 44; 48 a Plumsteadia gibbosa 7.7 271 74 - 77; 78 a, b Plumsteadia lerouxii 7.7 268 70 - 73; 78 c-f Rigbya arberioides 5.1 151 2 ? 9 Scutum leslii 7.5 234 49 ? 58 Vereenia leeukuilensis 6.2 179 10 f-I; 11 a-c Summary of plate contents Plate 1: Intra-familial seed scar morphology. Plates 2-9: Rigbya arberiodes Plates 10-11: Aberia spp. & incertae sedis Plates 12-18: Bifariala intermittens Plates 19-22: Estcourtia conspicua Plates 23-42: Elatra leslii Plates 43-48: Ottokaria spp. Plates 49-58: Scutum leslii Plates 59-69: Gladiopomum spp. Plates 70-78: Plumsteadia spp. Plates 79-82: Gonophylloides spp. Plates 83-89: Dictyopteridium spp. Plates 90-100: Lidgettonia spp. A NOTE REGARDING SCALE: All fructifications have been represented at approximately twice life-size, with some enlargements of important features. The scale bar towards the bottom of the plate applies to all photographs which do not have their own scale bar inserted somewhere on the photograph. However, in some cases, where there is an enlargement of the part and counterpart of a specimen, a scale bar may be included on only one of the photographs. PLATE 1. Seed scar details of members from each of the four families of ovulate glossopterid fructifications recognised here. Note how each specimen has a seed scar with a finely striated distal wing or scale-like extension. In the Dictyopteridiaceae and Lidgettoniaceae, the wing is divided into units delimited by grooves passing on either side of each scar. Each unit is comparable to the arrangement of seed scar to distal scale/wing seen in the Arberiaceae and Rigbyaceae. (a) Rigbyaceae: Rigbya arberioides (BP/2/13047); (b) Arberiaceae: Arberia hlobanensis (BP/2/15893); (c) Dictyopteridiaceae: Scutum leslii (BP/2/13735); (d) Lidgettoniaceae: Lidgettonia lidgettonioides (NM/1576b). For scale comparisons, see relevant plates. PLATES 2 - 9: Rigbya arberioides PLATE 2. Rigbya arberioides fructifications from Mooi River, displaying a variety of branching patterns; note the specimen in fig. (j), which undergoes a series of dichotomies, producing fourth-order branches; figs (s) & (t): enlargements of the specimen in fig (c), illustrating the presence of seed scars along the distal margin of the fan-shaped lamina; each scar is proximal to a longitudinally striated scale/ wing-like structure with a scalloped distal margin. PLATE 3. Further examples of Rigbya arberioides; figs (d) & (e) are enlargements of figs (a) & (b), and illustrate the bifacial nature of the fructifications; the sterile surface in (e) bears longitudinal striations that continue into the wing-like scales; the fertile surface bears a seed scar at the base of each scale. PLATE 4. Specimens of Rigbya arberioides; figs (a)-(c): part and counterpart of an incomplete fructification, with an enlargement of the primary dichotomy in (c); note the prominent striations on the pedicel, particularly the medial striation running from the dichotomy; each primary branch undergoes two further dichotomies, resulting in eight ultimate branches; figs (d) & (e): a particularly fascinating specimen, with the left side having suffered an injury early on in its development, or possibly a viral infection, resulting in stunting of all branches to the left of the primary dichotomy; figs (f) & (g): a specimen with an elliptical lamina and a single rank of branches along the periphery; the enlargement in fig. (g) illustrates the seed scar morphology very clearly; note the radially striated scars at the branch termini, each with a distal wing. PLATE 5. Specimens of Rigbya arberioides. Figs (a) & (b): winged seeds closely associated with the holotype of R. arberioides; fig. (c): enlargement of the holotype in fig. (f); several of the branches have a seed attached at the base of the wing; fig. (d): enlargement of the attached seed in fig. (g); fig (e): specimen with a broad, fan-shaped lamina without differentiation of individual branches; fig. (h): fructification with a small fan-shaped lamina, and a single rank of long, thin terminal branches. PLATE 6. Enlargements of Rigbya arberioides specimens. Fig. (a): a close-up of the specimen in pl. 2, fig. (q); the morphology of the seed scars is particularly well illustrated in this specimen; fig. (b): an enlargement of the specimen in pl. 2, fig. (p); this specimen has strange triangular seed scars, and although the terminal branches are clearly demarcated, they appear to be connected laterally by a thin lamina; fig. (c): enlargement of specimen in pl. 2, fig. (r) illustrating the smooth, striated sterile surface of a R. arberioides fructification. PLATE 7. Rigbya arberioides from the Bulwer locality; note how all specimens from this locality have a very long pedicel, and a fan-shaped lamina bearing seed scars directly on the margin, without the development of branches. Figs (a) & (b): enlargements of the specimens in figs (d) & (e); note the smooth, striated sterile surface in (b), and the fertile surface bearing seed scars at the base of the scale/ wing-like structures in (a); figs (c) & (f): longitudinally striated sterile surface; figs (g)-(i): further examples of fructifications with very long pedicels. PLATE 8. Drawings and a reconstruction of Rigbya arberioides. PLATE 9. Figs (a)-(d): primary morphological forms of Rigbya arberioides from Mooi River; fig. (a): small basal fan-shaped lamina gives rise to single order branches each terminating in a seed scar with a distal scale/wing; fig. (b): large basal fan-shaped lamina bearing seed scars with scale/wing directly along the distal margin, without development of individual branches; fig. (c): a primary dichotomy gives rise to two primary branches, each of which gives rise to three secondary branches terminating in a seed scar with distal scale/wing; fig. (d): and elliptical lamina gives rise to a series of single-order branches along the periphery, each terminating in a seed scar with distal scale/wing; fig. (e): silhouette drawings of fructifications from Mooi River, giving an indication of the diversity of branching patterns seen in these fructifications. PLATES 10 - 11: Arberia spp. & incertae sedis PLATE 10. Figs (a) & (b): part and counterpart of the holotype and only known specimen, of Arberia hlobanensis; the specimen may well have originally been attached to the associated axis, but was clearly not in organic attachment at the time of preservation; figs (c) & (d): part and counterpart of the holotype and only known specimen, of Arberia allweyensis; fig. (e): two partially overlapping, incomplete specimens of Arberia madagascariensis, the only examples to have been collected in South Africa; fig. (f): holotype of Vereenia leeukuilensis; figs (g)-(i): further examples of V. leeukuilensis. PLATE 11. Drawings and reconstructions of members of the Arberiaceae and of the taxon previously known as ?Arberia allweyensis?, now regarded as incertae sedis. Figs. (a)-(c): Vereenia leeukuilensis; figs (d)-(f): drawings of the part and counterpart of the single known specimen of ?Arberia allweyensis?, and a reconstruction; figs (g)-(i): drawings of the part and counterpart of the Arberia hlobanensis specimen, and a reconstruction; figs (j) & (k): reconstruction and drawings of Arberia madagascariensis. PLATES 12 - 18: Bifariala intermittens PLATE 12. Fig. (a): holotype of B. intermittens fertile surface, with primary wing in apex and along left side of receptacle, and secondary wing exposed to the left of the receptacle; fig (b): fertile surface, secondary wing only visible in basal region (note basal lobes); fig. (c): fertile surface, primary wing visible on left, secondary wing exposed to the right of the receptacle; fig. (d): fertile surface, only the primary wing is visible; fig. (e): fertile surface, only primary wing visible; fig. (f): fertile surface, secondary wing is completely obscured. PLATE 13. Figs. (a), (c), (e), (g) to (j): further examples of fertile surface impressions, with primary wing visible; fig (b): fertile surface primary wing visible on the left and apex, secondary wing visible to the right and in basal area, note basal lobes; fig. (d), (f): impressions of the fertile surface, with primary wing in apex, and secondary wing lobes visible in base; fig. (h): impression of fertile surface with primary wing present laterally and towards the apex, but with a secondary wing lobe visible to the right of the pedicel; fig. (i): fertile surface with secondary wing exposed (note the pronounced peripheral groove and fragment of primary wing in the apex). PLATE 14. Morphology of the sterile surface of Bifariala intermittens. Fig. (a), (b), (h), (l), (n): impressions of the sterile surface with complete exposure of the primary wing, which is continuous with the surface of the receptacle; fig (c): primary wing is exposed in apical portion of fructification, but the basal lobes of the secondary wing are visible in the basal part, at a higher level in the sediment; fig. (d): the primary wing is visible at the apex, but the radially fluted secondary wing is visible along the left side of the receptacle; note the groove along the edge of the receptacle alongside the segment of secondary wing; fig. (e): a key specimen contributing to Plumstead?s (1958) insistence on the presence of pollenate organs in this taxon; this impression of a sterile surface has a heavy coating of reddish iron oxide which has exaggerated the venation on the receptacle and the fluting of the primary wing in the apex of the fructification, creating the illusion of individual strands; fig. (f): fertile counterpart of (h), with prominent peripheral groove between receptacle and primary wing; fig. (g): Glossopteris leaf with B. intermittens attached right near base; enlagement of fructification in fig. (g); figs (i) & (j): part and counterpart illustrating primary and secondary wing morphologies; in fig. (i) the impression of the fertile surface has the primary wing to the left of receptacle and radially striated, fluted secondary wing on the right and in fig. (j) the sterile surface has the primary wing to the right, secondary on the left, with a pronounced groove between receptacle and secondary wing; fig. (k): an impression of the sterile surface of a fructification with remnants of the secondary wing to the left and right of the receptacle; fig. (m): sterile surface with only basal lobes of secondary wing visible; fig. (o): sterile surface with basal lobe of secondary wing visible on left side. PLATE 15. Further examples of Bifariala intermittens displaying dual wing morphologies. Fig. (a) impression of sterile surface has an overlying secondary wing to the left of the concave receptacle, note clear basal lobes of the secondary wing (enlargement in fig. (c)); fig. (b) counterpart of (a); fertile surface with primary wing in apical region overlying smooth secondary wing which is exposed in lateral and basal areas; fig. (d): enlargement of exposed secondary wing (with fluting and striations) in fig. (b); fig. (e): sterile surface with secondary wing lying at a higher level in the sediment than the receptacle; figs (f), (g), (o): fertile surfaces with primary wing developed laterally and in apex, with secondary wing lobes in base; fig. (h): sterile surface with secondary wing at higher level in sediment than small section of primary wing at apex; figs (i), (j): apex of sterile surface of fructification with small section of fluted, striated secondary wing on left side of receptacle; note how secondary wing is discontinuous with receptacle edge, and overlies the primary wing; figs (k), (l): fertile surface of small fructification with primary wing in apex and on right side of fructification, and radially fluted, striated secondary wing to left side of receptacle; note the ridge of primary wing remaining on the secondary wing, adjacent to the receptacle on the left side; figs (m), (n): part and counterpart of a fructification with the primary wing exposed at the apex, secondary wing in the base; fig. (p): sterile surface with primary wing. PLATE 16. Serial photographs of dissections: two impressions of the fertile surface of Bifariala intermittens, revealing the presence of a radially fluted and striated secondary wing beneath the impression of the primary wing. Figs (a)- (c): the primary wing along the left side of the receptacle is removed to reveal the secondary wing beneath; note how the faint fluting corresponds to the junctions between the marginal seed scars of the receptacle; figs (d)-(f): some of the primary wing is removed in the apical region on the left side of the fructification in (e), and in the middle of the fructification on the right side, in fig. (f). PLATE 17. Details of the fructification dissected in pl. 16, figs (a)-(c). Fig. (a): and enlargement of the fructification; figs (b), (c): close-ups of the left side of the fructification illustrating the striations and faint fluting orientated perpendicular to the receptacle edge; note how the fluting corresponds to the positions of the marginal seed scars in fig. (c); the secondary wing has been damaged immediately adjacent to the receptacle on the left side. PLATE 18. Drawings and reconstructions of Bifariala intermittens. Fig. (a): drawing of the specimen in pl. 13, fig. (b); fertile surface with a primary wing remnant in upper left part of fructification, and a well-preserved secondary wing in the base and along the right side of the fructification; this specimen has a particularly long pedicel; fig. (b) HOLOTYPE of B. intermittens, with primary wing in apex and on right side, and secondary wing to the left; fig. (c): apex of the sterile surface of a fructification, with a section of secondary wing preserved on the left; figs (d), (e): drawings of the part and counterpart of a fructification with elements of both the primary and secondary wings; figs (f)-(h): fossil reconstructions of B. intermittens based on the drawings of VM/03/3205/62 and VM/03/3205/63 in figs (d) and (e); figs (g) and (h) represent fructifications with exposure of the entire primary and secondary wing respectively; fig. (i) is a reconstruction of the apex of the original fructification illustrating the relative positions of the wings. PLATES 19 - 22: Estcourtia PLATE 19. Estcourtia conspicua: part and counterpart of the holotype, NM/1276b. PLATE 20. Estcourtia conspicua: part and counterpart of specimen BP/2/8172. PLATE 21. An incomplete specimen of Estcourtia conspicua, with the apical portion of the subtending Glossopteris leaf preserved. PLATE 22. Line drawings of BP/2/8172a, fig. (b) and the holotype (NM/1276a), fig. (c), with a reconstruction of Estcourtia conspicua in fig. (a). PLATES 23 - 42: Elatra leslii PLATE 23. Fig. (a): Holotype of E. leslii from Vereeniging; impression of the sterile surface of the fructification; counterpart of type specimen first described by Thomas (1921), which is housed in the Natural History Museum, London (NHMV20742); fig. (b): fertile surface of specimen with long, acuminate primary wing, and well-developed secondary wing lobes in base (see pl.33 for dissection of this specimen); figs (c), (e): impressions of sterile surface; note continuous passage of veins from receptacle into primary wing; fig. (d): impression of fertile surface, with wedge of sediment bearing impressions of well-defined seed-scars lying above the impression of the primary wing. PLATE 24. Fig. (a): impression of fertile surface of fructification, where cleavage plane has passed close to the covering hood, with sediment bearing seed scar impressions only present in the base and on the left side of the fructification; the secondary wing lobe is visible in the base, on the left; fig. (b): fertile surface with narrow rim of hood visible (depressed region proximal to primary wing); figs (c), (d): part and counterpart of fructification with very elongated primary wing; the hood is visible in (d) as the depressed region proximal to but continuous with the primary wing, around and beneath the wedge of sediment bearing the impression of the fertile surface of the receptacle. PLATE 25. Figs (a)-(c): impressions of the sterile surface; note the depression around the receptacle, at the base of the primary wing, which may represent the line of fusion with the hood; fig. (d): impression of a fertile surface, with very poor seed scar definition. PLATE 26. Further examples of impressions of the sterile surface of Elatra leslii fructifications. PLATE 27. Figs. (a)-(d): impressions of the sterile surface of Elatra leslii; fig. (e): impression of the fertile surface; the wedge of sediment with impressions of seed scars also bears secondary imprints of seeds which lay between the receptacle surface and the hood at the time of preservation; the secondary wing is visible in the base, on the right side of the fructification. PLATE 28. Figs (a)-(c): impressions of the fertile surface of Elatra leslii fructifications; note the prominent basal lobes of the secondary wing in the slightly distorted specimen in fig. (b); fig. (d): this specimen, although damaged and , appears to represent a view of the primary wing and hood; the edge of the aperture of the hood is visible on the left side, but the wing appears to have been torn prior to preservation, with a glossopterid leaf visible protruding through the tear on the right; there is an impression of a seed on the right side of the fructification. PLATE 29. Figs (a)-(c): impressions of the fertile surface of Elatra leslii fructifications with impressions of seeds visible in the wedge of sediment obscuring the hood. PLATE 30. Figs (a), (b): impression of the fertile surface of a specimen of E. leslii with a particularly elongated primary wing; compressions of seeds are visible in the wedge of sediment bearing the impression of the fertile surface of the receptacle (enlargement of seeds in (b)); secondary wing is visible in the base of the fructification; figs (c), (d): impression of fertile surface illustrating the discontinuity of the receptacle and primary wing impressions; note the carbonaceous layer beneath the wedge of sediment bearing the impression of the receptacle (enlargement in fig. (d)); this is the compression fossil of the hood; figs (e), (f): a further example of an impression of the fertile surface of an E. leslii fructification, with particularly well-preserved seed scars (enlargement in fig. (f)). PLATE 31. Figs (a)-(c): fertile surface of Elatra leslii with partial exposure of the hood along the periphery of the impression of the receptacle; note the characteristic, radial creasing of the hood; (d): sterile surface of an E. leslii fructification preserved in attachment to its subtending leaf; note the robust aggregation of veins in the base of the fructification, and the campylodromous pattern of vein divergence. PLATE 32. Specimen BP/2/7401: the fructification is still attached to its subtending leaf, but only remnants of the leaf material remain in the fossil. In fig. (a) the darkly shaded area represents the impression of the subtending leaf, with typical Glossopteris venation and a broad, well-developed midrib; the medium grey shading highlights an impression of the hood; the light grey is the primary wing and sterile surface of the fructification. Note the carbonaceous phytoleim or compressions between the layers of sediment bearing impressions of the various structures (black shaded areas in fig. (a)). Fig. (b) is a close-up of a portion of the specimen illustrating the different layers. Note the glossopterid venation in the impression of the leaf to the lower left of the image. Between the leaf impression and the next layer of sediment, is a layer of black carbon representing the compression fossil of the hood. The layer of matrix below this carbonaceous layer bears an impression of the hood. Below this layer of matrix is another carbonaceous deposit, this time representing the receptacle and primary wing of the fructification. The lowermost layer of sediment bears an impression of the primary wing and sterile surface of the receptacle. Fig. (c) is a view of the whole fructification, and fig. (d) is another close-up of the fructification illustrating the layering of matrix and carbonaceous material. PLATE 33. A closer look at BP/2/7220, and the results of a dissection to reveal the hood beneath the wedge of sediment bearing an impression of the fertile surface of the receptacle. The striations and fluting on the hood are continuous with those of the primary wing. The line of divergence of the hood from the primary wing is gently scalloped, as seen in fig. (c). The hood dips down into the sediment, and is creased due to compression during preservation; it ends in a triangular, tent-shaped opening which spans the lower two thirds of the fructification. In figs (b) and (c), dissection of the basal lobe of the secondary wing on the left reveals an extension of the hood. PLATE 34. Details of the dissection of BP/2/7318 to expose the hood; fig. (a): the impression fossil before dissection; fig. (d): the impression fossil following removal of the wedge of sediment bearing beautifully preserved impressions of seed scars; figs (b)-(g): close views of the edge of the impression of the receptacle, illustrating the presence of a carbonaceous compression between the sediment bearing an impression of the receptacle, and the underlying hood; in figs (b), (e) and (g), the reverse side of the dislodged sediment is shown to bear the impressions of fine striations from the hood beneath; the carbonaceous remnants of the hood lie beneath the impression of the receptacle; the exposed distal edge of the hood (along the aperture) is visible in fig. (f). PLATE 35. Details of the dissection of BP/2/7220, illustrating how removal of the impression of the fertile surface of the receptacle reveals the hood beneath; the reverse surface of some of the fragments of sediment that were removed, have been enlarged to illustrated the presence of fine striations which represent an impression of the inner surface of the hood. PLATE 36. Specimen BP/2/7146a: an unusual specimen of Elatra leslii with three-dimensional preservation; one can follow the phytoleim as it arches away from the receptacle on the left side of the fructification, illustrating the continuity between the covering and primary wings; the two layers of phytoleim merge in the apex of the fructification, where only the primary wing is present; although the orientation of the fructification is unclear, the specimen suggests that the outer margin of the secondary wing may be continuous with the hood. PLATE 37. Fig. (a): a possible view of a fully exposed hood; the wedge-shaped feature may represent the aperture of the hood, or may be a crease in the sterile surface of the fructification; fig. (b) impression of a leaf base with the edge of the sterile surface of an attached fructification visible to the right; fig. (c): a view of a primary wing and fully exposed hood; note the prominent tent- shaped aperture in the hood, and the characteristic creasing of the hood-like wing due to compression during preservation; the slight indentation in the wing running approximately parallel to the aperture represents the line of divergence of the hood from the primary wing; fig. (d): a Glossopteris leaf of the same morphological type as those found with attached specimens of Elatra leslii; the leaf base has a characteristically expanded, bulbous base. PLATE 38. Examples of Elatra leslii fructifications preserved in attachment to a subtending Glossopteris leaf. PLATE 39. Examples of Elatra leslii fructifications preserved in attachment to a subtending Glossopteris leaf. PLATE 40. Examples of Elatra leslii fructifications preserved in attachment to a subtending Glossopteris leaf. PLATE 41. Drawings of Elatra leslii. Fig. (a): drawing of specimen in pl. 24, fig. (a); with an almost fully exposed hood, only some of the matrix bearing an impression of the fertile surface of the receptacle remaining in the lower left of the fructification; figs (b), (c): part and counterpart of the fructification in pl. 24, figs (c) and (d); note continuous surface of the impression of the sterile side of the fructification (c) and the separate wedge of sediment overlying the primary wing on the impression of the fertile side; fig. (d): drawing of the specimen in pl. 25, fig. (a), an impression of the sterile surface with venation passing uninterrupted from the receptacle into the primary wing; note the unusual lobes in the base of the fructification; fig. (e) counterpart of specimen figured in (d), with well-preserved seed-scar detail on wedge of sediment overlying the hood; fig. (f): drawing of specimen with impressions of subtending Glossopteris leaf, hood and sterile surface (figured and described in detail in pl. 32). PLATE 42. Reconstructions of Elatra leslii. Fig. (a): a full fossil reconstruction of the fructification, with pedicel; fig. (b): a fossil reconstruction without pedicel, illustrating the position and structure of the secondary wing, with radial fluting and striations associated with the marginal seed scars at the base of the receptacle; the dotted line at the base of the primary wing represents the line of divergence of the hood, beneath the wedge of sediment bearing the impression of the seed scars; note how the elongated primary wing bears fluting and striations that are apically inclined; the fluting in the primary wing also corresponds to the positions of the marginal seed scars; fig. (c): a reconstruction of a fully exposed hood with its tent-like basal aperture; this is the view that results from complete removal of the impression of the fertile surface and secondary wing. PLATES 43 - 48: Ottokaria spp. PLATE 43. Figs (a)-(f): specimens of Ottokaria transvaalensis from Vereeniging. Figs (a), (b): part and counterpart of the holotype, illustrating the long, narrow pedicel of even width, and the deeply incised wing lobes distinctive of the species. PLATE 44. Additional examples of O. transvaalensis from Vereeniging, illustrating the variation in receptacle shape, wing width and degree of lobe incision. The part and counterpart in figs (h), (i) very clearly show the division of the wing into blunt, rounded lobes. PLATE 45. Ottokaria buriadica from Vereeniging. Figs (a)-(c): the holotype of O. buriadica in attachment to the base of a gangamopterid Glossopteris leaf; figs (d)-(f): additional examples from Vereeniging, showing the round receptacle, narrow wing and pedicel which is broadly expanded at insertion into the receptacle. PLATE 46. Further examples of O. buriadica from Vereeniging. PLATE 47. Figs (a)-(c): O. buriadica from Vereeniging; figs (d) & (e): part and counterpart of the best-preserved specimen of O. hammanskraalensis from Hammanskraal. PLATE 48. Figs (a)-(c): reconstructions of O. transvaalensis, O. hammanskraalensis and O. buriadica respectively. PLATES 49 - 58: Scutum leslii PLATE 49. Scutum leslii specimens from Vereeniging. One of Plumstead?s (1952) syntypes for ?S. rubidgeum? is figured in (a) & (b), and the two syntypes she assigned to S. leslii are figured in (c), (e) & (f). PLATE 50. Additional examples of S. leslii, illustrating the huge variety in size and shape of specimens from Vereeniging. The specimen in fig. (a) was assigned by Anderson & Anderson (1985) to ?S. draperium?, perhaps on the basis of its large size. PLATE 51. S. leslii from Vereeniging; note the very high wing width to receptacle ratio of the specimens in figs (j), (l) & (m), and the slightly cordate bases of those in figs (a), (c), (d) & (h). PLATES 52-53. S. leslii fertiligers from Vereeniging, illustrating the variation in form of the subtending leaves. Figs (d)-(f) illustrate Plumstead?s (1952) syntype for the seed-bearing surface of ?S. rubidgeum?. The specimen figured in pl. 53, figs (g), (h) is an example of ?S. draperium? as per Plumstead (1952, 1956a) and Anderson & Anderson (1985). PLATE 54-55. The controversial, bract-bearing S. leslii specimens from Vereeniging. The ?bracts?, interpreted by Plumstead (1956a, 1958a) as pollenate structures, are here considered to be the elongate wings of attached seeds. PLATE 56. Line drawings of S. leslii fertiligers from Vereeniging. Note the scar on the midrib in fig. (c), at the site of pedicel divergence, and how the midrib is more robust below this point. Specimens such as these lend support to the theory that the pedicel is an axillary structure which is adnate to the leaf. PLATE 57. Figs (a), (b): line drawings of two S. leslii specimens with attached seeds; fig. (c): drawing of a particularly beautifully preserved impression of the fertile surface of an S. leslii fructification; figs (d)-(f): drawings of venation patterns observed on the sterile surfaces of several S. leslii fructifications. PLATE 58. Silhouette drawings of Scutum specimens from Vereeniging, illustrating the wide range of intergrading wing and receptacle morphologies. Members of both S. rubidgeum and S. draperium, as categorised by Anderson & Anderson (1985), have been included (all drawings at approximately life-size). PLATES 59 - 69: Gladiopomum PLATE 59. Gladiopomum dutoitides (Plumstead) Adendorff et al. from Vereeniging. Figure (a): a small specimen with broad, prominent apical spine; fig. (b): a laterally compressed fructification, originally described by Plumstead (1958) as Pluma thomsonii; figs (c), (d): part and counterpart of a fructification, clearly showing the characteristic apical spine, broad wing and elongated elliptical receptacle; fig. (e): a particularly large specimen (fertile surface), with well-defined apical spine and wing fluting; fig. (f): the only specimen of G. dutoitides found at the Hlobane locality (sterile surface), showing a broad wing and narrow receptacle. PLATE 60. Fig. (a): holotype of Gladiopomum dutoitides (Plumstead) Adendorff et al. from Vereeniging; fig. (b): detail of the fructification; fig. (c): distal portion of the attached leaf, showing venation with parallel meshes. PLATE 61. Specimens of Gladiopomum acadarense (Anderson & Anderson) Adendorff et al. from Cedara with gladiate receptacles, broad wings and long petioles. Figures (c), (d): Holotype of G. acadarense: although the base of this specimen?s pedicel overlaps the base of a Glossopteris petiole, a physical connection is not convincing; fig. (a): detail of the associated leaf; fig. (b): fertile surface counterpart of (d). PLATE 62. Figure (a) - fertile surface of a large, broad specimen of G. acadarense with an incomplete wing (tapered wing base?); fig. (b): lanceolate specimen with wing that tapers abruptly at the apex; fig. (c): sterile surface of a fructification with a well-defined apical extension; fig. (d): fertile surface of a particularly robust, broadly lanceolate fructification. PLATE 63. Figures (a), (b): the fertile and sterile surfaces of a lanceolate fructification of G. acadarense with a particularly high wing width: receptacle width ratio and a pointed apex; figs (c), (d): part and counterpart of a large, oblong fructification with rounded apex. PLATE 64. Figures (a), (b): further examples of G. acadarense; fig. (c): fructifications were regularly found in close association. PLATE 65. Figures (a), (f): additional specimens of G. acadarense, illustrating the wide range in size and morphology of the fructifications; figs (a), (b): one of the smallest specimens from Cedara. PLATE 66. Figures (a) - (i): specimens of G. acadarense. Note the tendency of the receptacle to be falcate in figs (d), (h) & (i). PLATE 67. The holotype of Gladiopomum elongatum Adendorff et al. from the Rietspruit locality. Figures (a), (b): part and counterpart, note the very long, narrow receptacle and broad wing. PLATE 68. Drawings of key specimens of Gladiopomum dutoitides from Vereeniging and Hlobane highlighting seed scars, wing features, apical spines and pedicels. Figure (e): drawing of the holotype [pl. 59, fig. (d)]. PLATE 69. Figure (a): drawing of the holotype of Gladiopomum acadarense [pl. 61]; fig. (b): drawing of another specimen of G. acadarense, clearly illustrating the apical extension; fig. (c): line drawing of the holotype of Gladiopomum elongatum [pl. 67]; fig. (d): reconstruction of Gladiopomum dutoitides, based on BP/2/13936 [pl. 59, fig. (d)]; fig. (e): comparative reconstructions of G. dutoitides (i), G. acadarense (ii), and G. elongatum (iii). PLATES 70 - 78: Plumsteadia spp. PLATE 70. Figs (a)-(i), Plumsteadia lerouxii: holotype part and counterpart in figs (a), (b), (e) and (f) (specimens BP/2/14211 and BP/2/14179). Note the slightly hastate base in BP/2/14191 (figs (c) and (d)), which may be due to the presence basal wing lobes. PLATES 71-73. Further examples of Plumsteadia lerouxii and subtending Glossopteris leaves. PLATE 74. Plumsteadia gibbosa from Loskop: part and counterpart of holotype in figs (a)-(c) and (f), illustrating the bulbous nature of the seed scars, as well as partially obscured, attached seeds; figs (d) and (e) are part and counterpart of two adjacent, partially overlapping specimens. PLATE 75. Figs (a)-(i). Specimens of Plumsteadia gibbosa from Loskop. Note wing detail in enlargement of BP/2/12536, fig. (b), on lower right side of the fructification. PLATE 76. Further examples of Plumsteadia gibbosa from Loskop. Figs (e) and (f) are part and counterpart of a fructification attached to the petiole of a Glossopteris leaf attached to a branch bearing other leaves. This specimen has a broad wing with prominent fluting. Wings are evident in most of the specimens figured here. Plumsteadia fructifications are, in many cases, found in close association, as illustrated in fig. (g). The nature of the seed scars and fluted, striated wing is clearly apparent in the enlargements of BP/13000a&b (figs. (k) and (l)). PLATE 77. Figs (a)-(e): Plumsteadia gibbosa from Mooi River. These specimens have a low L:W, and a slightly cordate base, with the prominent, bulbous seed scars typical of P. gibbosa. Figs (f)-(g): Plumsteadia gibbosa (?) from Inhluzane. A mass of closely associated fructifications preserved on a single slab from Inhluzane. The poor preservation of the specimens made conclusive identification impossible, but the shape, dimensions and bulbous seed scars most closely affiliates these fructifications with P. gibbosa. PLATE 78. Reconstructions and line drawings of Plumsteadia gibbosa (figs (a) and (b)); and P. lerouxii (figs (c)-(f)). PLATES 79 - 82: Gonophylloides PLATE 79. Specimens of Gonophylloides strictum, from Vereeniging; the holotype is illustrated in figs (a) and (b). Fig. (f) is the only isolated specimen of G. strictum that has been found. PLATE 80. Additional specimens of Gonophylloides strictum. Specimen BP/2/14230a&b, in figs (a) and (b) has a particularly high length to width ratio for the taxon. PLATE 81. Figs (a)-(d): Gonophylloides strictum Fig. (d) is a reproduction of a photograph from Smithies (1978; p. 90-91, fig. 16, specimen number 41 a/h), illustrating the only known specimen of G. strictum from the Hammanskraal locality. The specimen could not be located in the BPI collections. Figs (e)-(i): Gonophylloides waltonii, with the part and counterpart of the holotype illustrated in figs (e) and (f). PLATE 82. Drawings and reconstructions of Gonophylloides waltonii (fig. (a)) and G. strictum (figs (b), (c) & (d)). Fig. (d) illustrates the shape of the subtending leaf of G. strictum and the relatives sizes of the fructification and leaf (life-size). PLATES 83 - 89: Dictyopteridium spp. PLATE 83. Figs (a)-(c): two examples of fructifications from Mooi River which are probably referable to Dictyopteridium sporiferum; note the lanceolate shape with long, tapering apex and particularly smooth receptacle with tubercles. Figs (d)-(r): Dictyopteridium natalensis from Mooi River; figs (m), (n): impression of a D. natalensis specimen in apparent attachment to a Glossopteris leaf; note the prominence of the secondary imprint or raised boss on the glossopteris leaf, marking the prior position of the fructification ? only a remant of the impression of the fructification remains in the lower left portion of the boss, as best seen in the enlargement (fig. (n)). PLATE 84. Enlargements of examples of D. natalensis illustrated in pl. 83; fig. (a): apical region of specimen in pl. 83, fig. (h), showing details of the fertile surface of the receptacle, and the nature of the narrow, fluted wing; fig. (b): apical portion of specimen in pl. 83, fig. (p), with surface detail of fertile surface of receptacle, and illustration of the short wing with fluting corresponding to the positions of the marginal seed scars; figs (c), (d): enlargement of the part and counterpart illustrated in pl. 83, figs (e), (f), showing the veined nature of the sterile surface of the receptacle, with a strong medio-longitudinal concentration of veins, and the fertile surface of the receptacle with its poorly defined seed scar cushions and small tubercles. PLATE 85. Further examples of D. natalensis from Mooi River. Fig. (b): sterile surface of a D. natalensis fructification; figs (d), (e): an unusual specimen with a clearly dentate wing margin; this is not the result of irregular breaking of the matrix or any other artefact, as the marginal scallops correspond to the wing fluting in the same manner as seen in other fructifications such as Scutum. PLATE 86. Figs (a)-(g): D. natalensis fructifications from Mooi River, in apparent attachment to Glossopteris leaves; figs (h), (i): an aggregation of structures which appear to be poorly preserved examples of D. natalensis. PLATE 87. Members of Dictyopteridium flabellatum, from the Loskop locality. Figs (a)-(c): a particularly small specimen, with a relatively broad wing with well- defined fluting; figs (d)-(g): paratype specimen clearly illustrating the pronounced venation pattern on both the sterile (fig. (d)) surface and fertile surface (fig. (g)) of the fructification; note the inclination of the wing into the sediment in figs (f), (g); fig. (h): holotype specimen, representing the fertile surface of a fructification; note how the wing has was folded over the fertile surface, and is only exposed towards the uppermost left side of the impression, where the sediment bearing the impression of the receptacle surface has been removed; the raised areolae along the margin of the receptacle are particularly well illustrated. PLATE 88. Additional examples of D. flabellatum from Loskop. The wing is particularly well preserved in the paratype specimen in figs (e), (f). Fig. (i) illustrates an example of two fructifications preserved in opposition. PLATE 89. Drawings and reconstructions of the three forms of Dictyopteridium from South Africa. Figs (a), (b): a drawing and reconstruction respectively of a fructification from Mooi River which belong within D. sporiferum; fig. (c): a drawing of a fragment of D. natalensis in apparent attachment to a glossopteris leaf; fig. (d): a reconstruction of D. natalensis; figs (e), (f): drawings of a paratype and holotype respectively of D. flabellatum; fig. (g): a reconstruction of D. flabellatum based on a drawing of the holotype. PLATES 90 - 100: Lidgettonia spp. PLATE 90. Specimens of Lidgettonia africana from Mooi River. Fig. (h) is the specimen assigned by Anderson & Anderson (1985) as the holotype of their species L. mooiriverensis, here synonymised with L. africana. (The 10 mm scale bar applies to figs (a)- (j); figs (f)-(k) are same-scale enlargements of the capitula and associated seeds in figs (e), (h) and (j) respectively). PLATES 91-92. Additional specimens of L. africana from Mooi River, illustrating the variation in scale leaf and capitulum morphology. Figs (m) & (n) on pl. 92, are enlargements of capitula in figs (l) & (o). Note how a seed scar is visible at the base of each lobe/segment of the capitulum in (n), whereas the striations are continuous from the wing into the receptacle in (m) which illustrates the sterile surface of the capitulum. PLATE 93. Specimens of L. africana from the Lawley locality. Note the variation in capitulum wing morphology, ranging from entire in figs (a)-(d), to scalloped with sharply pointed lobes, as in figs (j)-(l). PLATE 94. Figs (a)-(j): additional specimens of L. africana from Lawley. Note in fig. (a) how several specimens have been preserved in close proximity on the same slab. These aggregated specimens (#1 - #3) are illustrated in pl. 93 (figs (c), (d), (g)-(l)). The uppermost, un-numbered arrow indicates an additional specimen of L. africana, the arrow near the centre of the photograph is pointing to a beautifully preserved insect wing, and the lowermost arrow is indicating the position of a detached capitulum. Anderson & Anderson (1985) placed the specimens from Lawley in a separate species, L. lawleyensis, with PRE/F/8393b (figs (b)-(d)) as the holotype. These specimens are here considered to fall within the circumscription of L. africana. Figs (k)-(p): examples of L. africana from Inhluzane. Anderson & Anderson (1985) placed these specimens in a separate species, L. inhluzanensis, with BP/2/11146 as the holotype (figs (n) & (o)), but they are regarded here as being synonymous with L. africana. PLATE 95. Figs (a)-(i): L. africana from the type locality, Lidgetton. Note the attached seeds in figs (a), (b), with enlargements in figs (c), (d). Figs (j)-(q): specimens of L. africana from the Loskop locality. PLATE 96. Lidgettonia lidgettonioides from Mooi River. Note the beautiful example of an attached seed in figs (e), (f), with an enlargement in fig. (g). Figs (k) and (l) illustrate the deep scalloping of the wing, and the lateral insertion of the pedicel into the capitulum. PLATE 97. Figs (a)-(j): additional specimens of L. lidgettonioides from Mooi River. Figs (k)-(m): L. lidgettonioides from Bulwer; figs (n), (o): L. lidgettonioides from Bergville. PLATE 98. Lidgettonia elegans from Mooi River. The holotype, figured in (a)-(d) is a magnificent specimen, clearly illustrating the capitulum morphology and with an attached seed. Figs (i)-(l) illustrate another specimen with attached seeds. Figs (g), (h) provide the clearest example of the spatulate capitulum. PLATE 99. Further examples of L. elegans from Mooi River. Specimens illustrated in (a)-(i) each have more than one pair of capitula per scale leaf. Figs (m), (n) illustrate a clear example of a seed which appears to be attached. PLATE 100. Drawings and reconstructions of the three species of Lidgettonia recognised here from the Upper Permian of South Africa. Figs (a)-(c): reconstructions of Lidgettonia africana; fig. (a) fossil reconstruction of the fertile surface of a single, dorsiventrally compressed capitulum, and a seed of the type found attached to L. africana capitula; fig. (b) plant reconstruction of a single capitulum with two attached seeds; fig. (c) plant reconstruction of a L. africana scale leaf with three pairs of attached, umbrella- like capitula. Figs (d), (e): a drawing and a reconstruction of Lidgettonia lidgettonioides and a seed of the type found in attachment. Figs (f), (g): Lidgettonia elegans; fig. (f) a drawing of a single, dorsiventrally compressed capitulum of L. elegans; fig. (g) a reconstruction of L. elegans and its unusually shaped seed.