ALBERT KR. MANN LIBRARY AT CORNELL UNIVERSITY Cornell University The original of this book is in the Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http://www.archive.org/details/cu31924085826117 CAMBRIDGE BIOLOGICAL SERIES FOSSIL PLANTS CAMBRIDGE UNIVERSITY PRESS C. F. CLAY, Manacer Hondon: FETTER LANE, E.C. 4 London: H. K. LEWIS AND CO, Lrp., 136, GOWER STREET, W.C. 1 Dondvon: WILLIAM WESLEY AND SON, 28, ESSEX STREET, STRAND, W.C. 2 few Work: G. P. PUTNAM’S SONS Bombay, Calcutta and fladras: MACMILLAN AND CO,, Lrtp. Toronto: J. M. DENT AND SONS, Lip. Tokyo: THE MARUZEN-KABUSHIKI-KAISHA All rights reserved (‘oO 'V) ‘puvpsuoon’ ‘nq Ivan ‘ypimprg Diwwonnsy FOSSIL PLANTS A TEXT-BOOK FOR STUDENTS OF BOTANY AND GEOLOGY BY A. C. SEWARD M.A., F.R.S., Hon. Sc.D. Dusiin PROFESSOR OF BOTANY IN THE UNIVERSITY ; MASTER OF DOWNING COLLEGE AND HONORARY FELLOW OF EMMANUEL COLLEGE, CAMBRIDGE WITH 190 ILLUSTRATIONS VOLUME IV GINKGOALES, CONIFERALES, GNETALES CAMBRIDGE: AT THE UNIVERSITY PRESS 1919 5 GOS” nevi /898 y. 4 PREFACE i ie the Preface to Volume. ITI I expressed my thanks for help given to me by many friends in the course of the preparation of the subject-matter of Volumes III and IV, but Dr Scott has again earned my gratitude by very willingly and to very good purpose continuing the tedious task of reading the proofs. It is also a pleasure to acknowledge the ao received from the Staff of the University Press. Since the publication of Volume III Palaeobotany has been deprived of the services of three senior investigators, Professor C.,E. Bertrand of Lille, Monsieur Grand’Eury, and Mr Clement Reid, men whose researches along different lines of inquiry have played a prominent part in the progress of the science during the last few decades. By the death of Miss Ruth Holden, a graduate of Harvard University and a Fellow of Newnham College, Cambridge, Palaeobotany has lost an unusually gifted and promising worker: though a citizen of a country which was then neutral her strong sense of duty led her to lay aside, temporarily as we hoped, botanical research for work with a British Medical Unit in Russia where she died in April of last year. Miss Holden’s last contribution to Palaeobotany (‘On the Anatomy of two Palaeozoic stems from India’; Annals of Botany, vol. xxxt. p. 315, 1917) was published too late to be considered in Volume ITI. If it is possible to carry out my intention of supplementing the descriptive treatment of plants, which forms the basis of Volumes I-IV, by a general review of the Floras of the Past the results will be published as an independent work more intelligible, I hope, to the general reader than the text-book which, with a certain sense of relief, is now brought to a conclusion. A. C. SEWARD. .- Botany SoHooL, CAMBRIDGE, May, 1918. POSTSCRIPT THROUGH the death of Dr Newell Arber in June of last year at the comparatively early age of forty-seven Botany and Geology have lost an able and indefatigable investigator. Since 1901, the date of publication of his first paper, he laboured incessantly and with success to advance palaeobotanical science. To his activity the Sedgwick Museum of Geology at Cambridge is deeply indebted, and through his personal influence several younger men acquired something of their teacher’s enthusiasm. The proofs of this Volume were passed for press in June, 1918, but owing to the exigencies of war conditions publication has been unexpectedly and inevitably delayed. A. O.. 8: June 18, 1919. TABLE OF CONTENTS CHAPTER XL GINKGOALES. Pp. 1—60. PAGE 1-8 A. RECENT 8-60 B, Fossin ; ‘ : ‘ : : : : 1. Petrified wood 8-10; ii. Leaves 10-50, 58-60; GINK- GOITES 10-33; Ginkgocladus 33; BAIERA 33-50; ili. Flowers and Seeds 50-58 ; STENORACHIS 54-58 ; ERETMOPHYLLUM. 58-60. CHAPTER XLI GENERA BELIEVED TO BELONG TO THE GINKGOALES BUT WHICH ON THE AVAILABLE EVIDENCE CANNOT BE REFERRED WITHOUT HESITATION TO THAT GROUP. Pp. 61—%5. : GINKGODIUM 61, 62; CZAEKANOWSKIA 62-68; PEILD- ENIA 68, 69; PHOENICOPSIS and DESMIO- PHYLLUM 69-75. CHAPTER XLII GENERA OF UNCERTAIN POSITION. Pp. 76—105. GLOTTOPHYLLUM 76; GINKGOPSIS 77; NEPHROPSIS 77, 73; PSYGMOPHYLLUM 79-90; RHIPIDOPSIS 90-92; SAPORTAEA 92, 93; DICRANOPHYLLUM 93-101; TRICHOPITYS 101-103 ; SEWARDIA 103-105. CHAPTER XLIII CONIFERALES (RECENT). Pp. 106—164. CHAPTER XLIV CONIFERALES (FOSSIL). Pp. 165—44. Fosstz GymyosrERMous woop (Coniferales) . . CLASSIFICATION OF FosstL CoNIFEROUS WOOD. I. DADOXYLON (and Araucarioxylon) 176-186 ; II. CU- PRESSINOXYLON 186-202; II]. TAXOXYLON 202, 203; IV. MESEMBRIOXYLON 203-212; V. PARA- CEDROXYLON 212, 213; VI. CEDROXYLON 213-219; 167-172 172-176 x CONTENTS VIL PITYOXYLON 219-231; VIII. PROTOPICE- OXYLON 231-234; IX. WOODWORTHIA 234, 235; X. ARAUCARIOPITYS 235, 236; XI. PROTO- CEDROXYLON 236-238; XII. XENOXYLON 238- 242; XITI. ANOMALOXYLON 242, 243; XIV. THYL- LOXYLON 213; XV. PLANOXYLON 244. CHAPTER XLV CONIFERALES (conTINvED). Pp. 245—301. ARAUCARINEAE 4 i Oe DAMMARITES, PROTODAMMARA 245- 252 ; Conites 252-256; ARAUCARITES 256-270 ; ELATIDES 270- 274; PAGIOPHYLLUM 274-277 PALAEOZOIC CONIFERS EXHIBITING CERTAIN FEATURES SUG- GESTIVE OF ARAUCARIAN AFFINITY BUT WHICH CANNOT BE DEFINITELY ASSIGNED TO THAT OR TO ANY OTHER FAMILY oF CONIFERALES ON THE EVIDENCE AT PRESENT AVAILABLE WALCHIA 277-282; SCHIZODENDRON ( Tylodendron) 282-286 ; HAPALOXYLON 286, 287; GOMPHOSTRO- BUS 287, 288; VOLTZIA 289-295 ; SWEDENBORGIA 295, 296; Strobilites 296; ULLMANNIA 296-299 ; ALBERTIA 300, 301. CHAPTER XLVI CONIFERALES (contTinuep). Pp. 302—335. CUPRESSINEAE CUPRESSINOCLADUS 303, 304, 307 -310; THUITES 305; CUPRESSITES 305- 307 ; ATHROTAXITES 311-315; BRACHYPHYLLUM 315-322, 324-328; BRACHYOXYLON 322-324; TAXODITES 328-331; CUNNINGHAMIOSTROBUS 331, 332; MORICONIA 332, 333; CRYPTOMERITES 334, 235; CRYPTO- MERIOPSIS 335. CHAPTER XLVII CONIFERALES (contrnuEp). Pp. 336—367. CALLITRINEAE z : : , 2 : CALLITRITES : 337- 342 ; FRENELOPSIS 342-346. SEQUOIINEAE SEQUOIITES 349- 351, 352 357; Conites 351, 352: GEL NITZIA 357-361; ; BUGEINITZIA 361, 362; PSEUDO- GEINITZIA 362; SPHENOLEPIDIUM 36 3-365. ScrADOPITINEAE < 3 SCIADOPITYTES 366, 367, PAGE 245-277 > 277-301 302-335 336-316 346-365 CONTENTS CHAPTER XLVIII CONIFERALES Un eaeane Pp. 368—404. ABIETINEAE PITYITES 370- 376; PITYOCLADUS 377- 380; PITYO- PHYLLUM 380, 381; PITYOSTROBUS 381-395, PITYANTHUS 395, 396 ; PITYOSPERMUM 396-398 ; PITYOSPORITES 398, 399; ENTOMOLEPIS 399; CROSSOTOLEPIS 400; PREPINUS 400-404. CHAPTER XLIX CONIFERALES (continuEp). Pp. 405—423. PODOCARPINEAE ‘ PODOCARPITES 406- 409; STACHYOTAXUS 410- 419; Strobilites 412, 413; Saxegothopsis 413. xi PAGE 368-404 405-413 PHYLLOCLADINEAE , : 413-417 PHYLLOCLADITES 413, 414; Pr otophyllogiadus 415, 416; Palaeocladus 417 ; Phyllocladopsis 417. TAXINEAE “ 417-423 TAXITES 417- 419; TORREYITES 419- 421; ; VESQUIA 422 ; CEPHALOTAXOPSIS 422, 423. CHAPTER L CONIFERALES INCERTAE SEDIS. Pp. 424—446. TRIOOLEPIS 424; MASCULOSTROBUS 424-426; PA- LISSYA 426-428; ELATOCLADUS 429-432, 435, 436; RETINOSPORITES 432, 133; ANDROVETTIA 436- 438; DACTYOLEPIS 438; RARITANIA 438, 439; SCHIZOLEPIS 439-442; DREPANOLEPIS 442; SCHI- ZOLEPIDELLA 442, 443; CYPARISSIDIUM 443-445 ; BENSTEDTIA 445, 446. CHAPTER LI PODOZAMITES AND NAGEIOPSIS: GENERA INCERTAE SEDIS. Pp. 447—458. PODOZAMITES and CYCADOCARPIDIUM 447-456; NAGEIOPSIS 456-458. CHAPTER LII GNETALES. Pp. 459—472. I. Recenr . 2... 49-468 II. Fossin . ; : , : : ; : . P 469-472 LIST OF ILLUSTRATIONS Frontisriece. Avraucaria Bidwillit. FIG. 630, 631. Ginkgo biloba . 632. 633. 634. 635. 636. 637. 638. 639. 640. 641. 642. 643. 644. 645. 646. 647. 648. Ginkgoites obovata; G. antarctica ‘3 é Ginkgoites digitata var. Huttoni. A. G. Nathorst Ginkgoites digitata. A. G. Nathorst Ginkgoites adiantoides; G pluripartita; G digtiatae G. stbintea G. multinervis ; G. moltenensis ‘ ‘ Ginkgoites digitata ; Ginkgo biloba Ginkgoites digitata . Ginkgoites digitata. Manchester Phil. Lit, Society] Ginkgoites digitata. [Trustees of the British Museum. ] Ginkgoites digitata var. polaris Ginkgoites sibirica; Buaiera Brawndana. [Royal Society eS Scotland.] 3 Ginkgoites Obrutschewt. [Geclog. survey, Russia . Ginkgoites lobata; G. crassipes. [Geolog. Survey, India.] Ginkgoites adiantoides Baiera Raymondi ; Ginkgoites Geinite’ Baiera paucipartita Baiera spectabilis ; B. engin - Baiera Simmondsi. [Australian Museum, Biden 1 649, 650. Bazera gracilis 651. 652. 653. 654. 655. 656. 657. 658. 659. 660. 66]. 662. 663. 664. 665, 666. Psygmophyllum flabellatum Baiera gracilis forma M en M. s. Baiera Phillipsi Baiera Lindleyana . Antholithus sp. . : 4 Antholithus Zeilleri. A. G. Nathorst Stenorachis scanicus Stenorachis lepida. [Geolog. Sanveys; Tesi and Hasan: 1 Eretmophyllum saighanense,; EL. pubescens Ginkgodium Nathorsti Czekanowskia Murrayana. Pruatves of ie British fawn ] Czekanowskia microphylla. [Trustees of the British Museum.] Desmiophyllum Solmst Phoenicopsis speciosa Ginkgopsis Czekanowskit ; N oer cdunceits M. Zalesaley PAGE . 82, 52 LIST OF ILLUSTRATIONS xill ria. PAGE 667. Psygmophyllum Kidstoni. T. N. Leslie . 85 668. Psygmophyllum Haydenr. [Geolog. ai India. i 86 669. Psygmophyllum Grasserti 87 670. Rhipidopsis ginkgoides. M. Palosdlcy ‘ 91 671, 672. Dicranophyllum gallicum - 95, 97 673. Trichopitys heteromorpha . 102 674. Sewardia latifolia 104 674*, Sequoia sempervirens. A. Henry 106 675. Phyllocladus trichomanoides,; P. pot; P. - pina 107 676. Podocarpus latifolia ; 109 677. Cryptomeria japonica 109 678. Araucaria excelsa; A. Rulei. [Royal Boziaty: ] 110 679. Araucaria Cookii var. luxurians; A. Cookii; A. M sitios A. Montana. [Royal Society.] 111 680. Araucaria brasiliensis. [Royal Society.] 112 681. Araucaria Cunninghami. [Royal Society.] . 112 682. Agathis Moorei ; A. loranthifolia. [Royal Society.] » 118 683. Araucaria Cookii; A. imbricata; A. Bidwillii; A. excelsa. [Royal Society.] 114 684. Microspores, microsporophylls, cones na cone- agile: dad eed of various Conifers 116 685. Saxegothaea conspicua. [New Bhytalagist, J , 119 686. Pinus monophylla; Taxodium mucronatum ; T. distichum 119 687. Saxegothaea conspicua.- [New Phytologist. ] 120 688. Torreya nucifera. [New Phytologist. ] 121 689. Araucaria imbricata. G. BR. Wieland F 125 690. Sequoia sempervirens; Abies sp. E. C. Jeffrey : . 1380 691. Araucaria Bidwillii; A. Cooki ; Agathis bornensis ; Larix ameri- cana. R. B Thomson 132 692. Saxegothaea conspicua. [New Phytologist J 133 693. Medullary rays and tracheids of various Conifers 137 — 694. Sections of various Coniferous leaves 141 695. Agathis Mooret; A. australis; A. vitiensis ; Podosarpas Me otleyt. [Royal Society.] F . . 144 696. Agathis Moorei; A. australis. TRayal Susieky] 144 697. Araucaria Biduillii. [Royal Society.] 145 698. Araucaria excelsa 146 699. Cupressus Macnabiana. [Camb. Unig, Bree } 147 700. Libocedrus decurrens. [Camb. Univ. Press.] 148 701. Athrotaxis cupressoides F : . j 150 702. Sequoia gigantea . . . 152, 153 703. Actinostrobus pyrami iahas Tiackiais amin: W. T. Saxton. ‘ : 154 Xiv LIST OF ILLUSTRATIONS FIG. PAGE 704. Pinus eacelsa . : et . 154 705. Pseudolariz Fnnnnpeets Hiies bracteatins A. concolor; A. Fraseri; Larix Griffiths : 157 706. Branches of Abietineae. [Camb. indy, — 157 707. Podocarpus formosensis. R. Diimmer 159 708. Dacrydium elatum; D. araucarioides 160 709. Cephalotaxus Henryr 161 710. Drimys Winterr 162 711. Veronica Hectori; Callitris calseivaving Voor tou satiate Athrotaxis cupressoides; Crassula lycopodioides 163 712. Section of Amethyst Mountain. [Camb. Univ. Press.] 171 713. Dadoxylon (Araucarioxylon) breveradiatum ‘ 184 714. Dadoxylon (Araucarioxylon) kerguelense . 185 715. Cupressinoxylon sp. : 188 716. Cupressinoxylon vectense. C. A. Hashes 191 717. Cupressinoxylon Hortii. [Trustees of the British ‘irwee: ]} . 193 718. Cupressinoxylon polyommatum ; C. Holdenae; C. Koettlitz 196 719. Cupressinoxylon tener . : . 3 ‘ 198 720. Cupressinoxylon (Taxadioxy sien Taxodii; Cupressinoxylon (Taxodioxylon) Sequovanum i 200 721. Mesembrioxylon bedfordense. [Trustees of the British Nfuseum, } 208 722. Mesembrioxylon Schwendae; Mesembrioxylon sp. . ¥ . 209 723. Cedroxylon transiens; Cedroxylon blevillense . : 2 > 215 724. Brachyphyllum eathiense; Cedroxylon Hornet . . : . 216 725. Pityoxylon eiggense , : : é ‘ . ‘ . 222 726. Pityoxylon Sewardi. [Trustees of the British Museum.] . » “224 727. Pityoxylon Benstedi. [Trustees of the British Museum.]. « 226 728. Protopiceoxylon arcticum . : : ; : : , . 233 729. Protopiceoxylon Hdwardsi. [Trustees of the British Museum.] 233 730. Xenoxylon phyllocladoides; X.latipcrosum . . é . 240 731. Anomaloxylon magnoradiatum : ‘ : ‘ : . 242 732. Dammarites Tolli . r e 7 , 7 . 248 733. Dammarites borealis. A. G. Nathonst : : : ‘ . 249 734-736. Conites Juddi. [Royal Society of Scotland.] . . 252-254 737. Araucarites sphaerocarpus. [Camb. Univ. Press.] . i . 257 738. Araucarites ooliticus. [W. G. Carruthers.] . » 257 739. Araucariées ooliticus. [Trustees of the British Hfvseue: i » 258 740. Araucarites Milleri; A. Jeffreyt; A. Rogersi; A. Brodiet; A. Haberleinit; A. cutchensis; A. Phillipsi; Araucarites sp. . 263 741. Araucarites Sternbergit . - r . . é 2 . 268 742. Hlatides Williamsonis . 2... Ds, . 272 743. Elatides curvifolia . ‘ ; é : : ; ‘ . 273 744. Pagiophyllum peregrinum ‘ : , ‘ : . 2 276 FI@. 745. 746. 747, 748. 749. 750. 751. 752. 753. 754. 755, 757. 758. 759. 760. 761. 762. 763. 764. 765. 766. 767. 768. 769, 771. 772, 774. 775. 776. 777. 778. 779. 780. 781. 782. 783, 785. 786. 787. 788. 789. 790. LIST OF ILLUSTRATIONS Walchia piniformis Schizodendron speciosum Gomphostrobus bifidus Voltzia heterophylla; V. Liebeana Swedenborgia cryptomeroides Ullmannia frumentaria; U. Bronni Cupressites taxiformis : Cupressinocladus salicornoides Aihrotaxites Ungeri; A. lycopodioides Brachyphyllum expansum . 756. Brachyphyllum expansum. [Geolog. Snnvey Indio) Brachyphyllum obesum 317, Brachyoxylon notabile ; Areiouconsichtys americana; Drain. mara speciosa; Brachyphyllum crassum. KE. C. Jeffrey Taxodites europaeus Moriconia cyclotoron Callicrites curta : Callitrites Brongniart« ; C. heteclicass °C. ewropaeus Freneiopsis Hoheneggeri ; F. ramosissima Sequoia magnifica. [Camb. Univ. Press. ] Sequoiites concinna. A. G. Nathorst Conites ovalis. [W. G. Carruthets.] Sequotites Langsdorfit . . F Sequoiites Couttsiae. |Camb. Univ. Press.] 770. Geinitzia Reichenbachir Sphenoleprdium Kurrianum 773. Piiyites Solmsi Pityites (Pinites) eirensis Pityocladus longifolius. A. G. Mathonst Pityophyllum longifolium Pityocladus kobukensis [Geolog. Barveys Riteda 1: Pityostrobus Dunkert Pityostrobus Leckenbyi. [W. G. Cuandt hers! i. Pityostrobus Benstedi. [Trustees of the British Museum. ] Pityostrobus oblongus. [Trustees of the British Museum.] Pityostrobus (Pinites) Andraet 784. Pityostrobus (Pinites) ‘icsionenialies Pinites Plutonis Pinus Timleri; K ‘aldo Fouie Picea excelsa. [Camb. Univ. Press. ] Pityospermum Lundgreni. A. G. Nathorst Pityospermum Nanseni; Pityospermum sp. Pityosporites sp. 374, LIST OF ILLUSTRATIONS . Prepinus statensis. E. C. Jeffrey . Podocarpites eocaenica Podocarpites elegans Sequoites Tournali ; Padscarpltes danerta . Podocarpites Campbelli . Stachyolarus elegans. A. G. nihore Strobilites Milleri. [Royal Society of Scotland: ] . Protophyllocladus subintegrifolius; Drepanolepis rotundifolia: Phyllocladopsis heterophylla Masculostrobus Zeilleri; Masculostrobus sp. (Eoapal Sucinty of Scotland. ] 3 ee sphenolepis Elatocladus heterophylla . Elatocladus plana. [Geolog. suvey Tndia; i| eee indica Hlatocladus elegans? E. one . LElatocladus elegans : : . Androvettia statensis ; Gentle Raicienbachit ‘E. C. Jeffrey Raritania gracilis . Schizolepis Moelleri; 8. Brauhide Schtestepiise pemafiens i . Schizolepidella gracilis Cyparissidium gracile; C. minimum . Podozamites lanceolatus. [Trustees of the British Reeve ] 2. Cycadocarpidium Erdmanni; C. Swabii; Podozamites distans . . Podozamites lanceolatus? [Geolog. Survey, India.] Podozamites Reinti ‘ Nageiopsis anglica. ['Trustces a ii British Museura, lee . Nageiopsis longifolia Gnetum africanum . : 2 . Welwitschia mirabilis. A. A. Church PACE 401 406 408 408 409 411 413 416 425 428 430 431 433 434 437 437 439 441 443 444 448 450 453 . 456 457 458 463 466 CHAPTER XL. GINKGOALES. A. RECENT. In the account of the Coniferae contributed to Die Natiir- lichen Pflanzenfamalien+ the genus Ginkgo, in accordance with the prevailing custom, was included in the Taxeae with Taxus, Cephalo- taxus, and Torreya. Hichler had previously referred Ginkgo, or Salisburia, to a separate family, the Salisburyeae?. Hirase’s discovery of motile antherozoids in the pollen-tube of Ginkgo biloba in 1896, ‘the most remarkable event in plant morphology during the last decade of the 19th century,’ confirmed the sus- picion that the association of this ‘unicum de la création actuelle’ with Taxus and other Conifers was inconsistent with a natural scheme of classification. At a later date Engler adopted the family-name Ginkgoaceae, and in his survey of the Embryophyta | ‘Siphonogama the isolation of Ginkgo is emphasised by the refer- ence of the Ginkgoaceae to a special class, the Ginkgoales. Ginkgo biloba L. (Salisburia adiantifolia Smith) has a preeminent claim to be described in Darwin’s words as a, ‘living fossil.’ It is sometimes said to occur in China as a wild plant, but there appears to be no sufficient reason to believe that it would have escaped extinction had it not been carefully tended as a sacred tree in the gardens of temples*. Since its introduction into Europe in 1730, Ginkgo has become familiar in cultivation in the northern hemisphere and thus through man’s agency this monotypic genus thas been restored to regions where it survived as late as the Tertiary epoch. In habit Ginkgo® resembles many Conifers and its long and short shoots recall those of Cedrus and Larix: the short shoots may also be compared with the main trunk of a 1 Hichler (89) p. 108. 2 Seward and Gowan (00) B. p. 113. 3 Engler (97) pp. 19, 341. 4 Elwes and Henry (06) Vol. 1. p. 58. 5 For a full account of the genus, see Sprecher (07). s. IV 1 XL] : GINKGO| 3 -Cycad. The large thin leaves with long and slender petioles are scattered on long shoots or crowded on slow-growing branches covered with leaf-scars. These short shoots are occasionally . branched! and, as Tupper? states, they may branch within the wood of the axis out of which they grow, a feature exhibited by the Triassic Conifer Woodworthia. The deciduous leaves are usually more or less deeply bilobed (fig. 630, A, D, F) but those on short shoots are often smaller, and the margin may be entire or uneven (fig. 630, C). On young and vigorous shoots or on seed- lings the lamina is often deeply divided into several cuneate segments (fig. 630, E). In exceptional cases the lamina may reach a breadth of 20 cm. (fig. 630, A) though as a rule it seldom exceeds half that size. The leaf-scars show two small cicatrices. The considerable range in size and form of the foliage-leaves is an important consideration in connexion with the determination of fossil specimens. Two vascular bundles pass up the petiole: at the summit each divides and the two outer branches follow the outer edge of the lamina, giving off a succession of forked veins. Objection is taken by Prof. Johnson? to the statement that there are two marginal veins on the lower edge of the lamina; he regards the ‘marginal’ vein as the product of the successive fusions of the forked veins of the lamina as they pass towards the leaf-base. Whatever interpretation is adopted, the presence of two broadly diverging marginal veins is a noteworthy feature, and the correct explanation is probably that each is derived from one of the two strands in the petiole and gives off a succession of dichotomously branched veins as it passes along the margin of the leaf-blade. The presence of short secretory tracts at intervals between the veins is a characteristic feature sometimes recognis- able in fossil examples. Throughout the greater part of their course in the lamina the veins are accompanied by a small number of reticulate transfusion-tracheids (fig. 631, G, ¢): these increase in amount near the distal end of each vein and the water-con- ducting elements may be eventually replaced by a group of short, pitted, tracheids*. A group of large cells with brown contents occurs above and below each collateral endarch bundle. The 1 Seward and Gowan (00) B. Pl. rx. fig. 42. ? Tupper (11) p. 376. $ Johnson (14) p. 171. 4 Sprecher (07) pp. 68—71; Bertrand, C. E. (74). 1—2 4 GINKGOALES [cH. stomata, irregularly scattered over the lower epidermis, consist of two guard-cells surrounded by 4—6 accessory cells which pro- _ject towards the centre of the stoma as blunt cuticularised papillaet (fig. 636, C). The epidermal cell-walls are slightly undulate®. The distinctive form of Ginkgo leaves renders almost negligible the danger of confusion with those of other Gymnosperms; but impressions of certain Fern fronds, e.g. Lindsaya reniformas Dry., Pterozonium (Gymnogramme) reniforme Mart., Trichomanes reni- forme Forst., and Scolopendrium nigripes Hook. might be mistaken for imperfectly preserved specimens of Ginkgo. Ginkgo is dioecious. The male flowers occur in loose catkins (fig. 631, B) borne on short shoots in the axil of a scale-leaf: each microsporophyll consists of a short, slender, filament with a small terminal scale or knob bearing as a rule 2 but not infrequently 3 or 4 elliptical microsporangia (fig. 631, A, A’). The microspores recall those of Cycads. Jeffrey? has recently called attention to the occurrence of ‘wings’ on the microspores of Ginkgo: these are _ very slightly developed and hardly warrant the use of the term wing; they present the appearance of very small shoulders giving the spores a form similar to that of a brachiopod shell. The same author expresses the view that Ginkgo presents striking resem- blances to the Abietineae. It has recently been pointed out that the microsporangia have a hypodermal layer of cells with thick- ening bands comparable with the fibrous layer in the anthers of Angiosperms‘. Jeffrey and Torrey® claim that certain anatomical features in the microsporangia of Ginkgo indicate a closer affinity to the Abietineae than to any other section of the Gymnosperms. The vascular bundles of the microsporophylls end in transfusion- tissue which passes almost imperceptibly into the mechanical elements of the sporangial wall : a similar distribution of mechanical tissue occurs in Abietineous microsporangia and there is the same intimate relationship as in Ginkgo between the tracheary and mechanical tissues. The female flower consists of a compara- tively long peduncle borne in the axil of a foliage-leaf, with two 1 Strasburger (66) figs. 139—142; Sprecher (07) figs. 79—81. ? Bertrand, C. E. (74) Pl. rv. figs. 9, 10. 3 Jeffrey (14) Pl. xx. figs. 7, 8. + Starr (10). 5 Jeffrey and Torrey (16). XL] GINKGO ; 5 ovules at the summit, one on each side of the actual apex. Fre- quently one of these is larger than the other. The occasional occurrence of abnormal female flowers is interesting from the point of view of palaeobotanical comparison. In extreme cases So) OVOO® vh Vas i) 2 {) GO > OKC Fig. 631. Gingko biloba. A, A’, B. Microsporophylls and sporangia. C. Seed with the outer flesh removed showing an exceptional, tricarinate (radiospermic), form. D. Abnormal megasporophyll; a, vegetative bud; c, collar. E, F. Tracheids from a stem. G. Transverse section of a leaf-vein; px, proto- xylem, ¢, transfusion-tracheids. (A, B, after Seward and Gowan; D, after Fujii.) the partially modified lamina of a foliage-leaf may bear marginal ovules, the lamina being continuous with the collar (fig. 631, D, ¢) at the base of the ovule. In other cases the peduncle may give off several stalked ovules, as in the specimen described by Fujiit 1 Fujii (96); Seward and Gowan (00) B. Pl. rx. figs. 1—5; Sprecher (07) p. 138. 6 GINKGOALES [CH. and reproduced in fig. 631, D: the apex of the shoot is seen at a. Fertilisation is said to occur after the fall of the ovule, but Hirase states that some seeds contain an embryo while still attached to the tree. The seeds are comparable in size with large cherries; the broad integument consists externally of a thick sarcotesta rich in secretory tissue but without a vascular supply, and an ‘inner sclerotesta which is usually two-angled (platyspermic) but occasionally three-angled and radiospermic (fig. 631, C). An account has recently been published! of some remarkable examples of Ginkgo seeds gathered from one tree: the stony coat showed 2, 3 and 4 ribs and many transitional forms. The sclerotesta is lined by a few layers of loose cells which form a papery membrane in ripe seeds. The absence of vascular tissue in the sarcotesta is a character in which the seeds differ from those of Cycads and. Taxads. At the base of the ovule is a single concentric strand which splits into two branches and these pass through the shell and divide into several bundles on the inner face of the integument forming a continuous mantle? of short reticulate tracheids as in the Palaeozoic seed Stephanospermum®. The base of the ovule is enclosed in a shallow cup or collar, a structure that is probably homologous with the lamina of a foliage-leaf but which has re- ceived various interpretations. A suggestion has been made that the collar may be homologous with the cupule of Lagenostoma?. The nucellus is joined to the integument except at the apex where it forms a prominent cone in which a pollen-chamber is developed: this chamber becomes roofed over by nucellar tissue and at a later stage a blunt outgrowth is produced from the summit of the prothallus, serving as a ‘tent-pole’ to support the roof of the pollen-chamber. There are two or more archegonia at the apex of the prothallus differing from those of Conifers and Cycads in the shorter and more spherical form of the egg-cell and similar to those in some Palaeozoic seeds. Fujii® draws attention to-the _ remarkable capacity for pollination exhibited by Ginkgo and speaks of the conveyance of microspores over a distance of 500 to 1000 metres. Another fact worthy of remark in view of the wide distribution of the Ginkgoales in the Mesozoic era, is the 1 Affourtit and La Riviére (15). Sprecher (07) figs. 120, 147, 148. 3 See Vol. m1. p. 326. 4 Shaw, F. J. F. (08). 5 Fujii (10) p. 216. XL] . GINKGO 7 germination of Ginkgo seeds after 45 days’ immersion in sea- water’, The embryo has normally two hypogean cotyledons though three are not uncommon. Velenovsky? mentions a pecu- liarity, another indication of the isolated position of the genus, in which seedlings of Ginkgo differ from those of other: Phanero- gams; the cotyledons are succeeded by two elongated scales with a forked apex; the next higher leaves, in.which a small bilobed lamina is a characteristic feature, show at the base two divergent prongs representing the fork of the lower scales. The lamina of the foliage-leaf thus arises in the angle of the V-shaped distal end of the earlier scale-leaf. A microspore on germination developes 2—3 prothallus-cells and the generative cell forms two large (110m x 80,) spirally ciliated antherozoids. After fertilisation the egg-nucleus divides, as in some Cycads, until 256 free nuclei are formed?®, but in Ginkgo the subsequent production of walls results in a tissue, called by Lyon‘ the protocorm, which completely fills the egg; whereas in Cycas this tissue is massed at the base and in Zamia wall-formation is also restricted. In Conifers the number of nuclei is much less and the proembryo still further reduced. It is probably legiti- mate to deduce from these facts that Ginkgo is in respect of its embryogeny the most primitive of the Gymnosperms: in this and other characters it is allied more closely to the Cycads than to the Conifers. Saxton® who has described the later stages in the embryogeny of Encephalartos draws attention to certain features shared by that genus and Ginkge. The leaf-traces arise from the stele as a pair of collateral bundles, as in the Palaeozoic genus Mesoxylon, which pass up the petiole. Annual rings are fairly distinct though, as Nicol® recognised, less obvious than in Conifers. The walls of the late-summer tracheids are hardly thicker than those of the spring-elements and the diffe- rence between the early and late wood is often slight’. Circular - bordered pits occur either in a single or double row on the radial walls of the tracheids and are fairly common on the tangential walls. The pits may be separate or in contact, occasionally 1 Ewart (08) p. 78. * 2 Velenovsky (07) p. 457, fig. 291a. 5 Coulter and Chamberlain (03). 4 Lyon (04). 5 Saxton (104). § Nicol (34) A. p. 147. 7 Nakamura (83) p. 25; Fujioka (13) Pl. xvi. 8 GINKGOALES : [cH. slightly flattened and alternate, but usually opposite (fig. 631, E, F). The pores of the pits are often crossed. Rims of Sanio form a well-marked feature on the tracheal walls, and Jeffrey? points out that they are clearly shown in the mature wood but not in close proximity to the primary xylem or in the wood of the reproductive shoots and leaves. True bars of Sanio frequently occur on the tracheids?. The secondary phloem consists of dis- continuous rows of fibres in addition tosieve-tubes and parenchyma. Characteristic features are presented by the uniseriate medullary rays: these are often 1—2 or 1—5 cells deep and do not appear to exceed 11 cells in depth; they are comparatively large and in tangential sections of the wood present an inflated appearance. There are 2—7 elliptical pits in the field of the ray-cells. Xylem- parenchyma though not abundant occurs here and there among the tracheids; the cells have thin walls and are larger than the medullary-ray cells and characterised by the occurrence of stellate calcium oxalate crystals?. B. Fossitt. 1. PETRIFIED WOOD REFERRED TO THE GINKGOALES. The characters of the wood of Ginkgo biloba summarised above are in general agreement with those of many Conifers, and such anatomical features as have been described by authors as more or less distinctive of the genus do not afford very trust- worthy guides to the identification of fossil wood. The compara- tively large size and rounded contour of the medullary-ray cells, as seen in tangential section, though worthy of note as character- istic features, are hardly satisfactory criteria when applied to wood that may have undergone partial decay and been exposed to influences affecting the original form of the more delicate tissues before petrification. The untrustworthy evidence afforded by the size of the medullary rays has been emphasised by Essner* who states that the ray-cells of Ginkgo are larger than those in any genus of Conifers. It has been claimed by Felix® that Ginkgo is 1 Jeffrey (12) p. 548. 2 Miller (90) Pl. xzv. ’ For anatomical details, see also Kleeberg (85); Essner (86); Strasburger (91); Seward and Gowan (00) B.; Penhallow (07); Sprecher (07); Burgerstein (08); Tupper (11). 4 Essner (86). 5 Felix (94). XL] PHYSEMATOPITYS 9 an exception to the general truth of Essner’s conclusions and that the large dimensions and rounded form of the ray-cells are features of diagnostic value, though in the Tertiary specimens compared by him with the wood of Ginkgo the ray-cells do not appear to differ appreciably in size or form from those of true Conifers. Given well-preserved material, it is not improbable that in favour- able cases the characters of fossil wood might furnish adequate grounds for referring it to Ginkgo: the numerous obliquely ellip- tical pits in each ‘field,’ the swollen medullary-ray cells, and the frequent crossing of the pores of the tracheal pits are the features mentioned by Gothan! who considers that the wood of Ginkgo— though difficult to define precisely in an ane tytioal key—may be distinguished from that of Conifers. Among the specimens of wood assigned to Ginkgo there are none, so far as I am aware, that can safely be accepted as entirely above suspicion. In 1850 Goeppert? proposed the generic name Physematopitys® for some Tertiary wood that he believed to possess the anatomical characters of Ginkgo biloba. Kraus* subsequently recognised resin-cells in the wood of Goeppert’s type-species, Physematopitys salisburioides, and identified the specimens as the root-wood of a Cupressinoxylon: he did not, therefore, include Physematopitys in the list of woods contributed by him to Schimper’s Tratté de Paléontologie, but mentioned it as a synonym of Cupressinozylon. Beust® and Barber® among other authors adopt the same course. It has more recently been stated by Krausel’ that Goeppert’s genus Physematopitys has the characters of Protopiceoxylon. Goeppert® afterwards described a second species, Physematopitys succinea, founded on a tangential section — of a piece of Oligocene wood from the Baltic amber, but the data are clearly insufficient to justify its identification as Ginkgo: Conwentz® includes the specimen in Pinus succinifera. Schroeter! described some wood from beds on the Mackenzie river in North Canada, referred to the Miocene period, as Ginkgo sp. 1 Gothan (05) p. 103. ’ 2 Goeppert (50) p. 242, Pl. xurx. figs. 1—5. 3 gtonua, that which is blown out. 4 Kraus in Schimper (72) A. p. 370; Kraus (83); Schenk in Zittel (90) A. p. 871. 5 Beust (85). 6 Barber (98). 7 Krausel (13). 8 Goeppert and Menge (83) A. p. 32, Pl. x. fig. 74. ® Conwentz (90) A. p. 26. 1 Schroeter (80) p. 32, Pl. m1. fig. 27—29. 10 * @INKGOALES [CH. on the ground of the large size of the medullary cells: no pits are described either on the tracheids or on the medullary-ray cells and the unusual size of the ray-cells may well be a pathological or post-mortem phenomenon. The species Physematopitys ex- cellens described by Felix! from beds, probably Eocene in age, in the Caucasus agrees with Cupressinozylon in the presence of rows of resin-parenchyma in the wood, and the depth of the rays greatly exceeds that in Ginkgo biloba. Penhallow? described some calcified wood from Upper Cretaceous beds in the Queen Charlotte Islands as Ginkgo pusilla, but the reasons for assigning it to that genus are not convincing. A fuller description of another specimen regarded as the wood of a Ginkgo has been published by Dr Platen® under the name Physematopitys Goepperti from material collected in Miocene beds in Milam County, Texas. The relatively large size of the medullary-ray cells is mentioned as the chief character on which the determination was based. : It may be said that such fossil specimens as have been referred to Physematopitys or Ginkgo have very little value as records of the occurrence of the genus Ginkgo: in view of the abundance of leaves in Mesozoic and Tertiary strata that are hardly distinguish- able from those of the surviving type it is remarkable—if the anatomical characters of the genus afford in themselves a trust- worthy basis of identity--that more satisfactory specimens have not been found. ii. LEAVEs. GINKGOITES. Gen. nov. It has been customary to use the generic name Ginkgo both for the recent species and for fossil leaves from Mesozoic and Tertiary strata, and in a few cases for Palaeozoic leaves. In certain instances, for example such leaves as those from the Island of Mull and other Tertiary localities referred to Ginkgo adiantoides (fig. 644) there can be no doubt as to generic identity with the recent species and indeed, so far as concerns form and venation, the Eocene leaves might well belong to Ginkgo biloba. On the other hand even in the case of Ginkgo adiantoides we lack 1 Felix (94) p. 107, Pl. rx. fig. 4. ; * Penhallow (02) p. 43, Pls. xu., xm. % Platen (08) p. 143. XL] GINKGOITES 11 the confirmatory evidence of flowers and seeds. From Wealden and Jurassic. rocks numerous leaves have been described that in some cases are practically identical with those of the living species, but for the most part they are characterised by certain features denoting at least a specific difference. For these and for other Ginkgo-like leaves it would seem desirable to follow the usual custom and adopt a designation that does not necessarily imply even generic identity. A few examples of seeds and male flowers are known from Jurassic strata bearing a close resemblance to those of Ginkgo biloba, but such specimens are not common and some of the few that have been found, though probably belonging to the Ginkgoales, may not be correctly included in Ginkgo. I therefore propose to employ the name Ginkgoites for leaves that it is believed belong either to plants generically identical with Ginkgo or to very closely allied types. It is impossible in some cases to draw a sharp line between the genera Ginkgo and Baiera: typical examples of the latter genus are easily recognised by their narrow, relatively longer, and more numerous segments, but it is obvious that characters based on the degree of division of a lamina and on the breadth ‘of the segments are at best unsatisfactory, and the inclusion of certain specimens in one or other genus is purely arbitrary. A difficulty is presented by several types of Palaeozoic leaves assigned by many authors to the Ginkgoales and referred to Ginkgophyllum, Psygmophyllum, and other genera which, while bearing a general resemblance to the leaves of Ginkgo, cannot be regarded as evidence of the occurrence of the class that is now. represented by Ginkgo biloba. It has been suggested that Psygmo- phyllum, Ginkgophyllum, Rhipidopsis, and certain other genera should be included in a distinct group, the Palaeophyllalest, a - group of which the affinities are unknown. Though the adoption of a distinctive group-name has the advantage of indicating the absence of any trustworthy evidence of relationship to the Ginkgo- ales, it is open to question whether anything substantial is gained by the use of a term suggestive of relationship between different leaves that in themselves afford no clue as to the position of the parent-plants. 1 Arber, E. A. N. (12) p. 405. 12 GINKGOALES [cH. The name Ginkgoites as used in this chapter is restricted to leaves that are regarded as records of the Ginkgoales, while the genera referred by Dr Arber to the Palaeophyllales are briefly described as fossils that may or may not be closely related to one another but which cannot as yet be assigned to any place in a natural system of classification. The leaves discovered by Grand’Eury in Permian Uralian beds and described by Saporta as Salisburia primigenia1 should pro- bably be referred to the genus Psygmophyllum: like many other supposed Palaeozoic species assigned to the Ginkgoales or to Ginkgo they afiord no satisfactory evidence of affinity to the surviving genus. Other examples of leaves from Palaeozoic rocks described as species of Ginkgo or Salisburva on inadequate grounds are described in the latter part of this chapter. The Rhaetic leaves described by Brauns as C'yclopteris crenata and afterwards referred by Nathorst, with some doubt, to Ginkgo are described in the account of Psygmophyllum?. Ginkgoites obovata Nathorst. Fig. 632 A shows the form of the specimen from the Rhaetic beds of Scania on which Nathorst® founded the species Ginkgo Fic. 632. A. Ginkgoites obovata. B. Ginkgoites antarctica. (Nat. size; A, after Nathorst; B, drawn from a specimen in the University Museum, Brisbane. ) ? Saporta and Marion (85) p. 145, fig. 74 2 See »p. 145, fig. 74. 88. * Nathorst (86) p. 93, Pl. xx. fig. 5. ora XL} GINKGOITES 13 obovata: the obovate lamina is 5:6 cm. long with a maximum breadth of 2-5cm.; the upper edge is partially torn and the forked veins are about 3 mm. apart. Some dark patches between the veins are probably, as Nathorst suggests, secretory sacs similar to those in the leaves of Ginkgo. In shape but not in venation this type resembles Ginkgodium Nathorsti Yok. (fig. 659, p. 62); except in the absence of a deep median sinus it is, how- ever, nearer to a species from the Jurassic of Dzungaria described originally as Ginkgo Obrutschewi? (tig. 642, p. 26). Ginkgoites Geinitzi Nathorst. The leaf from the Rhaetic beds of Scania on which this species was founded was originally referred by Nathorst3 to Ginkgo but subsequently transferred by him to Baiera; it consists of a slender stalk and a sub-triangular lamina deeply divided into 4—6 linear truncate segments with 2—4 veins dichotomously branched near the base (fig. 645, B, p. 38). An examination of the original specimen leads me to prefer the designation Ginkgoiles to Baiera. This species like many others from Rhaetie rocks is hardly dis- tinguishable from some Jurassic types. Ginkgoites antarctica Saporta. Under the name Salisburta antarctica Saporta* described a single leaf from Australia believed to be of Lower Lias age, but no precise information is given with regard to the locality. Shirley® has also figured a specimen as Ginkgo antarctica from rocks that are probably of Rhaetic age at Denmark Hill, Ipswich (Queens- land). The lamina of Saporta’s specimen is broadly obcuneate and 3-5 cm. broad, characterised by the presence of two marginal veins like those in Ginkgo biloba from which forked branches are given off. This leaf is practically identical with some of the smaller, entire, examples on the short shoots of the recent type. The rather larger specimen figured by Shirley does not present so striking a similarity to those of the existing species. The lack of definite information as to the provenance of the type-specimen 1 See page 61. 2 Seward (11) p. 46, Pls. mr.—vz. 3 Nathorst (78) B. p. 26, Pl. xu. fig. 17. 4 Saporta and Marion (85) p. 142, fig. 71, A.; Ratte (88) Pl. mr. fig. 1; Renault (85) A. Pl. 1. fig. 19. 5 Shirley (98) Pl. 1. fig. 1. 14 GINKGOALES [CH. is unfortunate, but whether or not Shirley’s fossil is identical with Saporta’s specimen there would seem to be no reasonable doubt that it should be included in the genus Ginkgoites. Fig. 632 B is drawn from a photograph of a specimen in the Brisbane Museum which I recently had an opportunity of examin- ing: it is from the Ipswich beds and is undoubtedly specifically identical with Saporta’s type. Ginkgoites digitata (Brongniart). This widely spread Jurassic species founded on leaves from . the Yorkshire coast, was first figured by Phillips in 18291 as Sphenopteris latifolia, but under the same name Brongniart? had a year previously recorded a Carboniferous Fern. In 1830 Bron- gniart? figured and described another Yorkshire specimen as Cyclo- pleris digitata: the generic name Cyclopteris was adopted by Dunker and other authors until Heer* drew attention to the very close agreement between the Jurassic leaves and those of the Maidenhair tree, a similarity that led him to adopt the generic designation. Ginkgo. Leaves hardly distinguishable from the Jurassic impres- sions had previously been recorded from Tertiary rocks as species of Salisburza or Ginkgo. It is impossible to define precisely the several species of Gink- - goites founded on leaves: in the account of the recent species attention is called to the range in leaf-form and its bearing on the determination of fossils. All that can be done is to adopt certain specific names as a matter of convenience, recognising that the differences on which the classification is based are not either sufficiently sharply defined or morphologically important to be regarded as criteria of true specific distinctions. Many authors have employed the specific name Huttoni, first used by Sternberg®, for leaves identical in size and outline with G. digitata but characterised by a deeply-lobed lamina; this difference is, however, not greater than or even as great as differences met with within the species Ginkgo biloba. To facilitate description the designation Hutton: is retained as a form-designation for the more deeply lobed examples included in the species @. digitata (e.g. 1 Phillips (29) A. Pl. vir. fig. 18. See Fontaine in Ward (05) B. p. 121. * Brongniart (28) A. p. 51. * Brongniart (28?) A. p. 219, Pl. uxr bis, figs. 2, 3. 1 Heer ($1?); (77) i. p. 40. ° Seward (00) B. p. 256. XL] GINKGOITES 15 fig. 633). The number of Ginkgoites leaves from Jurassic strata is considerable and the student who attempts to classify specimens in a large collection under specific heads soon finds himself confronted in an acute form with the constantly recurring difficulty of fixing boundaries. As Knowlton! says, ‘In dealing with such an abun- dance of specimens and multiplicity of forms one must needs make either many “species” to accommodate this diversity, or only one or two, and in view of the known variation exhibited by the single living species, the latter plan seems preferable.’ In advocating this use of specific names in a liberal sense I admit the probability or in- deed the certainty that forms specifically distinct will be grouped Fic. 633. Ginkgoites digitata var. Huttoni. A leaf from the Upper Jurassic of Helmsdale, Scotland. (Stockholm Museum; nat. size.) under one designation. It is, however, clearly impossible in the case of impressions of leaves of Ginkgoites to impose limitations based on the form of the lamina, the degree of dissection, and similar variable features that cannot be accepted as trustworthy criteria of true specific distinctions. As material accumulates data may be furnished that will enable us to recognise characters of morpho- logical significance : in carbonised impressions from which cuticular preparations can be made the form of the epidermal cells and the structure of the stomata may supply a valuable aid to more accurate diagnosis. The spacing of the veins is a feature worthy of attention in the description of well-preserved specimens. 1 {Knowlton (14) p. 55. . 16 GINKGOALES (cH. There is also a further difficulty in regard to terminology : the employment of the two generic names Ginkgo or Ginkgowtes and Buiera reveals a striking lack cf uniformity among authors, and the artificial nature of the characters determining the use of one or other generic name necessarily lead to diversity in practice. As with the definition of species within the genus Ginkgowtes, so also the adoption of Ginkgoites or Buera is to a large extent the Fic. 634. Ginkgoites digitata. Leaf from Kap Boheman, Spitzbergen. (Stockholm Museum; nat. size.) result of individual preference and merely expresses an attempt to classify in an arbitrary fashion the numerous types of leaves that in themselves afford no sure guide as to precise affinity. The South African, Rhaetic, specimen shown in fig. 635, L was originally described as Barera moltenensis! but it might equally well be referred to Ginkgoites. 1 Seward (08) B. Pl. u. fig. 4 XL] GINKGOITES 17 The leaves of Ginkgo digitata have a long slender petiole (fig. 634) ; the lamina is semiorbicular or obcuneate, entire, or more or less deeply divided into equal lobes, or irregularly divided into several SP RUN BORE DO > Fic. 635. Ginkgoites. (4 nat. size.) Ginkgoites adiantoides, Tertiary, Island of Mull. AAAAARAARAAAS . pluripartita, Wealden, North Germany (after Schenk). . digitata, Jurassic, Japan (after Yokoyama). digitata var. Huttoni, Jurassic, Australia (after Stirling). sibirica, Jurassic, Siberia (after Heer). digitata, Jurassic, Turkestan. multinervis, Lower Cretaceous, Greenland (after Heer). digitata, Jurassic, Oregon (after Fontaine). digitata, Jurassic, Yorkshire. digitata, Jurassic, Scotland (after Stopes). digitata, Jurassic (or Wealden), Franz Josef Land (after Nathorst). moltenensis, Rhaetic, South Africa. digitata, Jurassic (or Wealden), Spitzbergen (after Heer). segments; the number and size of the segments and the form of their distal ends, truncate or obtuse, vary within wide limits (figs. 635, 637, 639, etc.). Numerous. dichotomously branched Ss. IV 2 18 GINKGOALES [CH. veins spread from the base of the lamina, the veins in the middle of the leaf being generally about 0-8—1 mm. apart. The stomata are practically confined to the lower surface of the lamina. The epidermal cells are polygonal and the walls slightly sinuous as in the recent species, and over the veins the cells are longer and narrower (fig. 636, A). Many of the larger epidermal cells have a cuticular ridge in the middle of the outer - wall, represented in the figure by a black line. The stomata agree closely with those of Ginkgo biloba; the two guard-cells are sur- rounded by a group of subsidiary cells characterised by their Fic. 636. Ginkgoites digitata; epidermal cells (A) and stoma (B). C. Stoma of Ginkgo biloba in two planes. (A, B, drawn by Miss N. Bancroft.) papillose heavily cuticularised walls overarching the stoma (fig. 636, B). The features shown in fig. 636 are not brought out in drawings from cuticles of the same specimen reproduced by Dr Stopes’ in her account of fossil plants from Brora: this may be due in part to a difference in the level at which the stomata were drawn. The stoma of Ginkgo biloba represented in fig. 636, C illustrates the considerable difference produced by viewing a stoma in slightly different planes?. No seeds have been found attached to stems bearing leaves of G. digitata, but seeds closely resembling those of the recent species 1 Stopes (07) p. 380. * Seward (11) p. 47, Pl. v. fig. 62. XL] GINKGOITES 19 occasionally occur in association with the foliage of this and other Jurassic species. Male flowers! similar in habit to those of Ginkgo biloba are also found in beds containing impressions of Ginkgoites. The abundance and wide geographical range of Ginkgoites digitata precludes anything more than a brief reference to some representative types selected in illustration of the range in form and the widespread occurrence of the species in Jurassic floras. The leaf represented in fig. 637 is an unusually complete example from the Middle Jurassic beds of Scarborough; the lamina is 3-8 cm. deep and 6 cm. broad, the venation agrees with that of Ginkgo biloba. A very similar type of leaf is figured by Heer from Upper Jurassic (or Wealden) strata of Spitzbergen as G. integriuscula?, but with the proviso that it may be merely a variety of G. digitata, a view that Nathorst® has wisely adopted. The latter author in speaking of the occurrence of G.. digitata in Spitzbergen states that ‘sometimes the surface of the schists [shales] is as completely covered with the leaves of Ginkgo as the soil beneath a living Ginkgo tree may be in autumn‘.’ In some specimens from the Yorkshire coast the lamina is practically entire as in a leaf from Scarborough in the York Museum figured in 19005. An exceptionally large form is shown in fig. 638; the lamina, 8 cm. broad, is divided into several short and comparatively broad obtuse or truncate lobes*. Fig. 639 shows a leaf from the Stonesfield Slate, now in the Cirencester Museum; the lamina is deeply divided into two broad cuneate lobes as in some forms of the recent species. The Stonesfield Slate specimens were origin- ally named by Buckman Noeggerathia (?) and later Stricklandia Fic. 637. Ginkgoites digitata. (3% nat. size.) M.S. 1 See page 51. 2 Heer (77) i. p. 44, Pl. x. figs. 7—9. 3 Nathorst (97) p. 15; for a discussion of the age of the Spitzbergen beds, see Nathorst (137). 4 Nathorst (11°) p. 221. > Seward and Gowan (00) B. Pl. x. fig. 54. 6 Seward (00) Pl. 1. fig. 5. 20 GINKGOALES Fic. 638. Ginkgoites digitata. (Manchester Museum; nat. size.) [CH. Fia. 639, Ginkgoites digitata. (Cirencester Museum 3 nat. size.) XL] GINKGOITES 21 acuminata’, A deeply divided obcuneate leaf, only 2—4 cm. broad, is figured by Dr Stopes? from Jurassic strata (Lower Oolite) on the Sutherland coast at Brora (fig. 635, J). The specimen shown in fig. 633, also from the Sutherland coast, a few miles north of Brora and of Kimeridgian age, illustrates the type of leaf that may be conveniently referred to Ginkgoites digitata. var. Huttoni®. Heer’s Jurassic species, Ginkgo Jaccardi*, from Switzerland is almost certainly Ginkgoites digitata. Arctic regions and northern Europe and Asia. The leaf reproduced in fig. 634 from a drawing made for me in the Stockholm Museum through the kindness of Prof. Nathorst was originally figured by Heer® from Cape Boheman, Spitzbergen, from beds assigned by Nathorst to the Upper Jurassic series. Nathorst includes in this series strata agreeing in their fossil plants with the Wealden of England and North Germany; the ‘Ginkgo beds’ may be Portlandian or even younger. The veins in this specimen are approximately 1 mm. apart. The leaf represented in fig. 640 is one of several specimens collected by Dr Koettlitz in Franz Josef Land trom beds that are probably Ju- rassic. The lamina is 2:5 cm. in depth and the veins are about 0-8 mm. apart; the lamina between the veins shows a fine transverse striation, not shown in the drawing, a feature occasionally BOE Fara Sarah anand in impressions of Ginkgottes and due in tb aioe: Beane set all probability to the shrinkage of the Land. (Museum of the Geo- mesophyll tissue into transverse bands®. _logical Survey, London; This form of leaf has been described as 0 Size), M.S. Gingko polaris Nath.7 ; itis smaller than most examples of Ginkgowtes 1 Seward (04) B. p. 98. 2 Stopes (07) Pl. xxvu. fig. 4. For a map of the Sutherland Coast, see Seward (11*). 3 Seward and Bancroft (13) p. 886. 4 Heer (76) PI. uviir. fig. 20. 5 Heer (77) i. Pl. x. fig. 2. 6 For similar transverse wrinkling, see Schenk (71) B. Pl. xxrv. figs. 7, 8; Seward (11) Pl. xr. fig. 74; Heer (77) ii. Pl. x1. fig. 1 5. 7 Newton and Teall (97) Pl. xxxvitt. figs. 1, 2; (98) Pl. xx1x. fig. 3; Nathorst (99) Pl. 1. fig. 8 and Thomas (11) Pl. 1v. fig. 8. ) GINKGOALES [CH. digitata and the veins are more crowded. For this type I suggest the designation G. digitata var. polaris. Solms-Laubach? figures an incomplete leaf from Franz Josef Land with a broader lamina divided into several broadly rounded segments as in many British specimens but with rather closer venation. A similar leaf from the same locality is figured by Newton and Teall’. Fig. 635, K shows a lobed leaf similar to that represented in fig. 640, described by Nathorst? from Franz Josef Land and compared by him with G. sibirica Heer and G. flabellata Heer from the Jurassic of East Siberia. A very small specimen similar in form to the larger example shown in fig. 635, K was figured by Nathorst*, also from Franz Josef Land, as Ginkgo polaris var. pygmaea. From the west coast of Greenland Hartz described a leaf very similar to some of the Yorkshire examples as Ginkgo (Baiera) Hermelini®: Hartz regarded the beds as Liassic or Rhaetic, and the occurrence of shells pointing to a Kellaways horizon immediately above the plant-beds suggests that the latter may belong to the Middle Jurassic series*. Ginkgo digitata is represented also in Jurassic strata in the New Siberian Islands by a leaf figured by Nathorst — as Ginkgo sp. which agrees with the type @. digitata var. Huttoni’, and Krystofovié® has described G. digitata from Jurassic beds in Ussuriland at the northern end of the Muravjev-Amursky] penin- sula. Some good specimens are recorded from Bornholm’, of Middle or Lower Jurassic age, as G. Huttoni which are identical in form and size with British specimens. North America. Several examples of leaves of the G. digitata type and some of the form Huttont have been figured from Middle Jurassic rocks in Oregon? (fig. 635, H). Some particularly large examples are named by Fontaine G. Huttont var. magnifolia but these are not 1 Solms-Laubach (04) Pl. 1. fig. 10. 2 Newton and Teall (97) Pl. x11. fig. 10. 3 Nathorst (99) p. 11, Pl. 1. figs. 8—19. 1 Ibid. Pl. 1. figs. 20, 21. 5 Hartz (96) Pl. xix. fig. 1. § Johnstrup (83). 7 Nathorst (07) Pl. 1. fig. 20. 8 Krystofovié (10) Pl. m1. fig. 1. ® Bartholin (94) p. 96, Pl. xit. figs. 1—3; (10) Pl. mt. figs. 9, 10. 10 Fontaine in Ward (05) B. Pls. XXX.—XNXIL., XLVI. XL} GINKGOITES 23 specifically distinct from G. digitata. G@. digitata is represented in Upper Jurassic or Wealden beds in Alaska (Cape Lisburne)!. Other Localities. From Southern Russia Thomas? has recently described good specimens of Ginkgo digitata of Middle Jurassic age some of which agree closely with the large leaves figured by Fontaine from Oregon as G. Huttoni var. magnifolia; the specimens previously figured by Kichwald® as Cyclopteris incisa from the same district are examples of G. digitata. The species is recorded also from Jurassic rocks in Turkestan? (fig. 635, F), Chinese Dzungaria® on the west border of Mongolia, from the region to the east of Lake Baikal®, and from Afghanistan’. The incomplete specimen figured by Feistmantel® from the Jabalpur group of India as Ginkgo lobata, part of which is shown in fig. 643, A, agrees in the form of the lamina and in venation with G. digitata: the veins in the middle of the lamina are from -8 to 1 mm. apart. Feistmantel compares his species with G. digitata, and an examination of one of his figured specimens leads me to assign it to that type; it is indistinguishable from the Afghan specimen already quoted. The piece of a leaf figured by Feistmantel as Ginkgo sp. belongs to a'similar leaf, but the venation is finer and it may be identical with Ginkgo crassipes Feist.® Leaves of the G. digitata type are recorded from Jurassic beds in Victoria? (fig. 635, D) and from Japan" (fig. 635, C). Tuzson has figured a bilobed petiolate leaf from Jurassic rocks in Hungary as Ginkgo parvifolia! ; it is similar in form to most of the specimens referred to G. digitata, but has relatively broader segments: it is interesting as being the first recorded example of Ginkgoites from Hungary. 1 Knowlton (14) Pl. xxiv. 2 Thomas, H. H. (11) p. 73, Pl. iv. fig. 7; Pl. vii. fig. 2. 3 Eichwald (65) Pl. 1Vv. fig. 6. 4 Seward (07?) Pl. vit. 5 Seward (11) Pl. 11. fig. 40. ® Krasser (05) Pl. 1. fig. 3. 7 Seward (12) PI. rv. fig. 51. ‘ 8 Feistmantcel (774) Pl. 1. fig. 1. 9 See page 28. 19 Seward (04) B. fig. 35. 11 Yokoyama (88) B. Pl. x11, fig. 2. 12 Tuzson (14) Pl. xrv. fig. 1. 24 GINKGOALES [CH. Ginkgoites sibirica Heer. The specific name sibirica was given by Heer’ to one of the most abundant forms in the rich plant-beds at Ust Balei near Irkutsk in Siberia; the specimens figured from Siberia as Ginkgo sibirica (fig. 635, E), G. Schmidtiana, and G. lepida cannot be regarded as well-defined species; they agree in the deep division of the lamina into several linear segments with obtuse or in some cases more pointed apices. Heer draws attention to the re- semblance between @. sibirica and G. pluripartita from Wealden rocks but, as he says, the venation is rather coarser in the Siberian leaves and the segments are generally narrower in G. sibirica. Fontaine in his description of Jurassic leaves from Oregon appa- rently identical with Heer’s @. sibirica states that G. Schmidtiana is a smaller form of the same species?, but Ward’ points out that as G. Schmidtiana is described on one page and @. sibirica is defined on the following page the former designation must be preserved. It may be urged that as the name sibirica is the more widely used and familiar term, considerations of convenience should override this meticulously strict interpretation of the rule of priority. A revision of Heer’s Siberian material would, I have no doubt, result in the reduction of his specific terms; on comparing several specimens in the Museums of Copenhagen and Stockholm with the illustrations in the. Flora Fossilis Arctica I found that several of the published figures are far from accurate. For the present the most convenient course would seem to be the retention of Gink- gottes sibirica for leaves similar to some of the more deeply divided forms of G@. digitata and to G. pluripartita, but normally character- ised by a lamina divided almost or quite to the base into oblong, obtuse or more or less acute segments. Leaves of the G. sibirica type, using the term in the wider sense and including Heer’s other species Ginkgo Schmudtiana and G. lepida, are fairly common in Jurassic rocks and occur also in Cretaceous floras; they are recorded from Kimeridgian beds in Scotland? (fig. 641, Knowlton (00) p. 31, Pl. tv. figs. 7—10; PI. v. fig. 5. * Dawson (93) p. 56, Pl. vi. figs. 1I—14. Heer’s species Ginkgo pusilla is founded on Jurassic leaves; Heer (77) ii. p. 61. 7 Penhallow (02) p. 48, Pls. xu., xm. 8 Depape (13). ® Engelhardt and Kinkelin (08) p. 196, Pl. xxmm. fig. 18, 10 Ettingshausen (79) p. 392. xu] BAIERA 33 from the London clay of Sheppey as Salisburia eocenica and afterwards figured by Gardner as Ginkgo? eocenicat, The specimens (11 x 9mm.) are smaller than the seeds of the recent species but in shape and in the keeled shell there is a fairly close resemblance. The hard sclerotesta forms the surface of the fossils. Similar seeds have been assigned to Ginkgoites adiantoides from the Upper Pliocene of the Frankfurt district®, but neither the German nor English specimens possess any interest as records of Ginkgoites seeds. Ginkgocladus. Ettingshausen. An imperfect leaf-like impression described by Ettingshausen® from Eocene beds in New Zealand was made by him the type of a new genus Ginkgocladus and interpreted as a stalked phylloclade similar to those of the recent Conifer Phyllocladus, but because of the presence of a slender stalk and the resemblance of the lateral veins to the venation of Ginkgo Ettingshausen suggested an affinity to that genus. The existence of a midrib is, however, an important difference. Neither the New Zealand species nor similar fragments from Eocene strata in New South Wales (Ginkgocladus austra- hensis*) are of value as botanical records. BAIERA. Braun. This generic name was first used by Braun® for some Triassic and Jurassic leaves agreeing in shape with those of Ginkgoites but distinguished by the greater number and less breadth of the linear segments. Braun’s definition states that the primary veins are dichotomously branched while between them secondary veins form irregular hexagonal meshes. Schenk® examined Braun’s Rhaetic specimens and failed to discover any indication of the presence of secondary veins. In 1877 Heer? emended the original definition of the genus: he refers to the presence of finer veins between the main vascular strands and this feature is shown in 1 Gardner (86) p. 46, Pl. rx. figs. 31—34. 2 Engelhardt and Kinkelin (08) p. 196, Pl. xxim. figs. 16—-18. 3 Ettingshausen (87) p. 39, Pl. vu. fig. 19. 4 Ibid. (88) p. 103, Pl. vim. fig. 32. 5 Braun, C. F. W. (43) p. 20. 6 Schenk (67) A. p. 42. Schenk includes some of Braun’s species of Baiera in Unger’s genus Jeanpaulia which has since been discarded. ? Heer (77) ii. p. 51. 8. IV 34 GINKGOALES [cH. some of his figures: these interstitial ‘veins’ probably mark the position of hypodermal strands of stronger cells, a feature that is not represented in recent or fossil Ginkgo leaves and is by no means generally characteristic of Baiera. Both Braun and Heer describe male and female reproductive organs: Braun interpreted some small specimens as sporocarps but these were recognised by Schenk as young foliage leaves. The male organs are described by Heer as ‘amenta staminifera pedunculata, nuda, filamenta filiformia, antherae loculis 5—12, verticillatis. Semen drupae- forme, basi cupula carnosa cinctum’ and compared with Schenk’s Stachyopitys Preslii which that author afterwards regarded as microstrobili of Batera Muensteriana. Reference is made to these and similar fossils in the account of examples of male flowers!. Seeds have been referred to Bazera on evidence furnished by their occasional association with leaves and by their resemblance to those of Ginkgo. It has been suggested that specimens described from the Potomac group of Virginia and Maryland as Carpolithus ternatus® and other species may be seed- bearing organs of Bazera, but there is no satisfactory evidence in support of this view. In all probability some of the associated seeds belong to Baiera, also some of the microstrobili, e.g. Leuthardt’s Swiss specimens described on a later page, but in the present state of our knowledge it is preferable to regard these specimens as reproductive organs that cannot be assigned with certainty to any particular species of Baiera or Ginkgottes. Leaves assigned to Batera vary within wide limits as regards size, the number of linear segments and their angle of divergence. In many cases the leaves are petiolate though in several instances the petiole is represented by a narrow basal region of the lamina as in Psygmophyllum. It is stated by some authors that the veins are undivided, but though dichotomy is less frequent in Bavera and may be absent in narrow parallel-sided segments it is by no means rare. The difference in venation between such leaves as Ginkgoites digitata and typical species of Baiera, e.g. Baiera gracilis, is mainly the result of the different form and degree of dissection of the lamina. The choice between Baiera or Ginkgoites as the 1 Page 51, ® Fontaine (89) B. pp. 265 etc., Pls. 134 ete. ; Berry (11) p. 372. XL] BAIERA 35 more suitable name for certain forms of leaf is not governed by any definite criterion: specimens described as Ginkgo sibirica, G. lepida, G. concinna!, etc. are indistinguishable from leaves referred by authors to Baiera. Similarly such a species as B. Iandleyana differs very slightly from some forms usually in- cluded in the genus Czekanowskia. Although leaves of Ginkgoites and Baiera are abundantly represented in plant-bearing beds we know very little of the habit of the foliage-shoots; in a few cases there is evidence of the occurrence of several leaves on a single short shoot (fig. 646); in Baiera paucipartita, for example?, the habit is the same as that of Czekanowskia and Phoenicopsis, but in view of the frequent preservation of Czekanowskia leaves still attached to an axis it is surprising that the leaves of Bazera almost always occur as detached specimens. The explanation may be that in Ginkgoites and Baiera the foliage-leaves were borne on long and dwarf-shoots as in the recent species Ginkgo biloba, whereas in Czekanowskia the leaves were confined to shoots of limited growth asin Pinus. Some specimens described by Salfeld? . from the Solenhofen beds of Bavaria as Bazera? longifolia Heer are interesting in this connexion; they consist of fairly stout branches bearing alternate leaf-like organs having the habit of Baiera longifolia but subtended by a short and thick recurved spinous process. There is no means of deciding from the avail- able material whether the resemblance of the leaves to those of Baiera is an expression of relationship or merely a case of parallel development, nor have we any means of determining the morpho- logy of the leaves and the subtending spines. The Solenhofen plant agrees in habit with Sewardia latifolia* from the Wealden of England and is included under that genus. While relationship between Sewardia and Baiera is by no means excluded, it is clear that the species of the former genus differ considerably from typical representatives of Baiera and Ginkgoites. Certain species of Baiera exceed in size the leaves of any example of Ginkgoites, notably B. Simmondsi, B. spectabilis, and others. The specimens described by Fontaine and Berry from the 1 Heer (77) ii. Pl. xm. figs. 6—8. 2 Nathorst (787) B. Pl. xx1. 3 Salfeld (07) B. p. 195, Pl. xx. fig. 3; Pl. xx. fig. 1. 4 Page 105. 36 GINKGOALES [cH. Potomac group in Virginia as Baiera foliosat, consisting of an axis bearing crowded leaves with a deeply and rather irregularly divided lamina and a comparatively broad and flat basal region, are not typical examples of the genus but agree more closely in habit with the older genus Dicranophyllum: their precise position cannot be definitely determined. The structure of the cuticles is known in a few species from accounts published by Schenk, Nathorst? and Thomas‘; the epidermal cells are sometimes characterised by fairly prominent papillae on the outer walls (fig. 647) and the stomata, more abundant on the lower surface but present also on the upper surface, closely resemble those of Ginkgo and Ginkgoites; the guard-cells are slightly depressed and are surrounded by 5—6 sub- sidiary cells with strongly cuticularised and projecting walls (fig. 647). Nathorst®> has drawn attention to the presence in B. spectabilis of traces of some secreted substance in the mesophyll recalling the secretory tracts in the leaves of Ginkgo. Braun and other authors have included Baiera in the Filicales, and attention has been called to the danger of confusing true Fern-fronds with leaves of Baiera. Berry’s discovery of sporangia on the linear segments of Bavera-like leaves from the Potomac beds, originally referred by Fontaine to his genus Baveropsis and regarded by him as Ginkgoaceous, illustrates the possibilities of error in determinations founded on leaves alone. The fertile examples of Baveropsis have been made the type-of a new genus Schizaeopsis®, other species of Fontaine’s genus being transferred to the genus Acrostichopteris; they differ from Bavera in their attachment to slender axes and are no doubt portions of compound Fern-fronds. It is impossible to define with confidence the precise geological range of the genus; leaves from Permian and Upper Carboniferous strata agreeing with Bavera in the deep dissection of the lamina have been assigned to the genus Ginkgophyllum (Psygmophyllum) and compared with Saporta’s Ginkgophyllum Grasserti (fig. 669, p. 87). In imperfect specimens it is not always possible to draw a sharp line between Baiera and species of Psygmo- 1 Fontaine (89) B. p. 213, PL. xcly. fig. 13; Berry (11) p. 372, Pl. ux. * Schenk (67) A. PL vr. figs. 1, 2; Pl. rx. figs. 11—13. 2 Nathorst (06). + Thomas (13) p. 244, fig. 5. ° Nathorst (06) p. 9, fig. 9. 8 Berry (11) p. 214; (112). XL] ” BAIERA 37 phyllum. There are, however, certain Permian leaves that are legitimately included in Baiera; the genus appears to have been widespread in Triassic floras, though more especially in those of the Rhaetic and Jurassic age. Bavera shares with Ginkgoites an important position in the Jurassic vegetation of both hemispheres, but in the Cretaceous period Baiera appears to have been a com- paratively rare genus and in the Tertiary floras it was entirely replaced by members of the Ginkgoales with leaves of the type that still survives. Bavera is clearly an older form than Ginkgovtes ; it is not recorded from India and it has not been found in the Permo-Carboniferous rocks of Gondwana Land. Baiera though unknown in a petrified condition may con- fidently be included in the Ginkgoales; the habit of the leaves, the structure of the epidermal cells and such evidence as there is with regard to the fertile shoots favour this conclusion. It must, however, be added that the position of the Palaeozoic examples is less firmly established. Baiera virginiana Fontaine and White. This species, from Permian beds in Virginia}, is based on im- perfect portions of laminae deeply divided into bifurcate segments with truncate apices and several parallel veins; it agrees in the form of the lamina and in the linear divisions with the type- specimen of Brongniart’s Fucoides digitatus? from Permian beds of Mansfeld, a species which Geinitz® also recorded, but under the generic name Zonarites. Potonié* and other authors, following Heer, transferred the species to Bazera. Heer’s combination, B. digitata, had, however, already been used by Schimper? for the Jurassic species usually called Ginkgottes digitata. Leaves of the form represented by B. virginiana may be closely allied to Saporta’s Permian species Ginkgophyllum (Psygmophyllum) Grasserti®. In the absence of the basal part of the lamina a com- plete diagnosis or accurate identification is impossible. Some .authors have referred fragments of similar leaves to the genus 1 Fontaine and White (80) B. p. 103, Pl. xxxvir. figs. 11, 12. 2 Brongniart (28?) A. p. 69, Pl. rx. fig. 1. 3 Geinitz (62) Pl. xxxu. figs. 1, 2. 4 Potonié (93) A. p. 237, Pl. xxxm. nS 2; Pl. xxxur. fig. 6. 5 Schimper (69) A. p. 423. & Page 87. 38 GINKGOALES _ [cH. Schizopteris, e.g. S. Guembeli from Upper Carboniferous and Permian strata}, but other Permian specimens assigned to Schizopteris, e.g. S. dichotoma and 8. trichomanoides® are generically distinct and probably belong to Pteridosperms. B. virginiana and the leaves referred to Baiera digitata—possibly specifically with Fon- taine’s type—represent Permian forms that agree closely with the larger species B. Simmondsi, Heer’s Keuper species B. furcata, and with the larger examples of B. Muensteriana (Presl) from Rhaetic beds. Baiera Raymondi Renault. This French, Permian (Autunien), species? (fig..645, A) differs but little from B. virginiana; the narrow cuneate leaf is divided Fic. 645. A, Baiera Raymondi. B, Ginkgoites Geinitzt. (Nat. size; A, after Renault; B, after Nathorst.) into very regularly bifurcate narrow linear segments diverging at a small angle. The largest specimen is 10:8 cm. long, the basal 1 Weiss (69) B. Pl. xu. fig. 7; Goeppert (64) A. Pl. rx. figs. 6, 7. * Zeiller (06) B. Pl. 1. figs. 7, 8. * Renault (88) p. 324, fig. 48. XL] BAIERA 39 portion of the lamina is 5 mm. broad and the ultimate segments. « with obtuse apices have a breadth of 25mm. The venation is imperfectly shown in the specimens figured by Renault and Zeiller?. Barera furcata Heer. A type similar to B. multifida Font., but characterised by the more uniformly narrow segments (2—2-5 mm.), is described by Heer?and Leuthardt? from Keuper beds of Switzerland. Leuthardt’s figures show a single vein in the segments, a feature which may be a peculiarity of the species. It was in association with this species that Leuthardt found the male flowers referred to on another page (p. 53). Baiera paucipartita Nathorst. The leaves of this Swedish, Rhaetic, species? may reach a length of 10 cm.; the lamina is narrow and cuneate, deeply divided into bifurcate linear segments with obtuse apices. It differs from Fig. 646. Baiera paucipartita. (After Nathorst; } nat. size ) 4 Zeiller (06) B. p. 202, Pl. xivu. figs. 1, 2. 2 Heer (76) A. Pls. Xx1x., XXX., XXXVI. 3 Leuthardt (03) p. 7, Pls. 1.—1v. 4 Nathorst (86) p. 94, Pls. xx.—xxII. 40 GINKGOALES [CH. B. longifolia Heer and other similar species in the smaller number of segments. There is no petiole but the leaves are attached by a narrow basal portion to a short axis (fig. 646). The occurrence of several leaves on a very short scale-covered dwarf-shoot is an interesting feature which affords evidence of relationship with Czekanowskia and Phoenicopsis. Baiera pauctpartita is described by Yokoyama! from Rhaetic beds in Japan and an imperfect specimen from Rhaetic strata in New Zealand has been assigned by Arber? to this species. In the case of imperfect leaves it is impossible to distinguish specifically between many ‘species’ of Baiera characterised by the division of the lamina into hilobed linear segments. Baiera spectabilis Nathorst. The leaves of this Rhaetic species from the south of Sweden? reach a length of 25cm.; the coriaceous lamina is obcuneate and fan-like but narrower than the leaves of such a species as B. Sam- mondst. There is no well-defined petiole; the lamina is deeply divided into two symmetrical halves each of which is further sub- divided into bifurcate linear segments, and the strongly contracted ultimate segments are a characteristic feature (fig. 647). The basal region forms a stalk-like portion varying in length and, in the specimens so far obtained, not exceeding 3cm. The veins are parallel to the sides of the segments and on the average 1 mm. apart; they are occasionally dichotomously branched. Stomata occur on both sides of the lamina but are more numerous on one surface, presumably the lower; the guard-cells are slightly de- pressed and surrounded by 5—6 subsidiary cells with thickly cuticularised papillose walls (fig. 647, B) as in Ginkgo. The epi- dermal cells on the lower surface are also papillose like those described by Thomas in B. longifolia*. Several dark spherical and spindle-shaped patches found by Nathorst between the cuticular- ised layers are believed to be the remains of some resinous or other substance comparable with that formed in the secretory tracts in the leaves of Ginkgo and some species of Ginkgoites. Baiera spectabilis is one of the larger forms of the species; it 1 Yokoyama (05) Pl. 1. fig. 5. ® Arber, E, A. N. (13) Pl. vam. figs. 2, 3. 3 Nathorst (06). 4 Thomas (13) Pl. xxv. figs. 3, 4. XL} BAIERA 4] ws . 1 Fic. 647. A, Baiera spectabilis; B, stoma CC, D, epidermal cells and stoma of Baiera longifolia. (A, B, after Nathorst A, 3nat. size C, D, after Thomas.) 42 GINKGOALES [cH. resembles B. pulchella, a Jurassic species described by Heer! from East Siberia and by Bartholin? from Bornholm, but in typical examples of B. pulchella the lamina is divided into two segments only. Comparison may be made also with B. longifolia a Jurassic species distinguished by its narrower segments. A specimen from Bornholm referred by Moller? to B. pulchella is probably, as Nathorst suggests, a piece of a B. spectabilis leaf. Baiera Simmondsi (Shirley). The leaves described by Shirley4 as Ginkgo Simmondsi from Denmark Hill near Ipswich in Queensland, from rocks that are probably of Rhaetic age, are of the same type as the leaf on which Ratte® founded his species Jeanpaulia (?) palmata, which he after- wards transferred to Salisburia, from the Wianammata beds (Trias) _ near Sydney. The precise age of the rich flora from Ipswich is difficult to determine: a recent examination of several specimens in the Brisbane collections led me to regard the plants as Rhaetic, but further light on this question will be afforded by Mr Walkom who is engaged in an investigation of the material. The Australian léaves agree closely with Fontaine’s Triassic species, Baiera multi- fida®, from Virginia: the plant-beds of the Richmond coalfield are correlated with the Lunz plant-beds in Austria’, the flora of which has never been adequately illustrated. The specimens from Virginia on which Fontaine founded his species do not afford any evidence of a true petiole and the basal portion of the cuneate lamina is narrower than in the Australian leaves: it is, therefore, not improbable that B. multifida is a distinct though very similar species. Ratte’s name B. palmata cannot be retained as Heer had previously employed the same name for a Jurassic Siberian ~ form’: I have adopted Shirley’s designation in the belief that there are no differences of specific value between the Sydney and Ipswich specimens. The leaf reproduced in fig. 648 is Ratte’s type-specimen in the Australian Museum, Sydney: the whole leaf is nearly 30 cm. > Heer (77) ii. p. 114, Pl. xx. fig. 3¢; Pl. xxu. fig. la; Pl. xxvm. fig. 3. * Bartholin (94) Pl. x1. fig. 5. 5 Mller (03) Pl. rv. fig. 19. 4 Shirley (98) p. 12, Pl. x. > Ratte (87) Pl. xvi; (88). 5 Fontaine (83) B. p. 87, Pls. xtv.—xLvu. ? Berry (12). 8 Heer (77) ii. p. 115, Pl. xxvumn. fig. 2. XL] BAIERA 43 long and 23cm. in breadth; there are nearly 60 ultimate linear segments with obtuse apices and, in the smaller subdivisions, Fic. 648. - Baiera Simmondsi. (Australian Museum, Sydney; 2 nat. size.) 3—5 veins. There is a well-defined petiole and in outline the whole leaf is identical with typical examples of Ginkgo biloba. 44 GINKGOALES [cH. Fontaine speaks of the lamina of his species as reaching a length of 25cm.; both in the method of division and in the form of the segments, B. multifida agrees closely with the specimen shown in fig. 648. A similar form of leaf is figured by Solms-Laubach? from Rhaetic beds in Chile as Batera? Steinmann, but the lamina only is preserved. Schenk’s B. taeniata? from the Rhaetic flora of Franconia is another similar type. Baiera stormbergensis Seward. The specimens described from the Stormberg series (Rhaetic) of South Africa’ are portions of leaves that must have reached a length of 12 cm. or more and a breadth of 10cm. The lamina is deeply divided into broad linear segments which are further subdivided into narrower distal segments. In the lower part of the lamina the venation is comparatively coarse, but as the result of repeated dichotomy the veins are much more numerous in the upper portion. This species may be merely a larger form of Feistmantel’s B. Schenki4 from the same beds, in which the lobes are narrower as in B. longifolia Heer. B. stormbergensis resembles Nathorst’s B. spectabilis from the Rhaetic of Scania, but the segments of the South African leaves have a coarser venation. Barera Muensteriana (Pres)). This Rhaetic species, originally figured by Presl as Sphaero- coccites Muensterianus and subsequently described by Braun as Baiera dichotoma, was named by Schenk Jeanpaulia Muen- steriana. Schenk® examined Braun’s specimens from Franconia and identified the supposed sporocarps as partially expanded segments of foliage-leaves. The leaves are petiolate and the fan- like lamina is deeply dissected into bifurcate linear segments; the veins are numerous and dichotomously branched. The epi- dermal cells are elongate over the veins and elsewhere polygonal ; their walls are straight or slightly sinuous. The stomata are of the usual type met with in Ginkgoaceous plants. Baiera Muensteriana cannot be distinguished by any definite character from leaves that are referred to B. gracilis: in the 1 Solms-Laubach (99) Pl. xiv. fig. 1. ® Schenk (67) A. p. 26, Pl. v. figs. 1—4; Pl. vi. figs. 1, 2. 3 Seward (03) B. p. 64, Pl. vurr. fig. 3. 4 Feistmantel (89) p. 72, Pl. m1. 5 Schenk (67) A. p. 39, Pl. 1x. XL] BAIERA 45 type-specimen of the latter species the segments are fewer than in B. Muensteriana, but in some Jurassic forms (e.g. fig. 651) this difference no longer holds good. This is only one among several instances where Rhaetic and Jurassic ‘species’ cannot be separated by any constant differentiating feature. B. Muenste- riana is recorded also from Persia! and several European localities, but it is impossible to determine its per Pines sane apart from that of Baiera gracilis. Baiera gracilis Bunbury ex Bean ms. The type-specimen of this species from the Middle Jurassic rocks of Yorkshire, as shown in fig. 649, is an impression of an Fia. 649. Baiera gracilis. Type- Fic. 650. Batera gracilis. (British Museum, 39208.) specimen of Bunbury (Bun- bury Collection, Botany School, Cambridge). imperfect leaf with a fan-shaped lamina deeply divided into forked linear segments?: a better exampleis reproduced in fig. 650. Bean 1 Schenk (87) B. Pl. vumr. fig. 44; Zeiller (05) p. 194. 2 Bunbury (51) A. p. 182, Pl. x1. fig. 3. 46 GINKGOALES [cH. referred this type to Schizopteris but Bunbury, while adopting Bean’s Ms. specific name, substituted the generic designation Baiera. The leaves are petiolate and the lamina is divided almost or quite to the base into a varying number of linear segments with obtuse apices. The veins, frequently indistinct, run parallel to the edges of the lamina and there are several in each segment. Leaves identical with or very similar to Baiera gracilis are very widely distributed among Jurassic floras in both hemispheres. Some of the specimens described by authors as Ginkgo lepida are hardly distinguishable from Bunbury’s species; G. concinna! Heer from the Siberian Jurassic flora is another very similar form; also Baiera incurvata Heer? from the Lower Cretaceous of Green- land, B. angustiloba Heer, as figured from Siberia and China, B. bidens (Ten.-Woods)4 from Queensland, B. australis McCoy and B. delicatula Sew. from Jurassic rocks in Victoria®, also leaves recently referred by Halle® to B. australis from the Lower Cretaceous plant-beds of Patagonia. Baiera gracilis is recorded from Upper Jurassic (or Lower Cretaceous) beds in Alaska’, but leaves of this Fic. 651. Baiera gracilis forma Mucnsteriana Fe (British Museum, # nat. size.) M.S. type are rare in the Jurassic strata of North America. The Rhaetic species B. Muensteriana® (Presl) described by Schenk from Franconia and by other authors is a closely allied type which 1} Heer (77) ii. Pl. xun. figs. 6—8. ° Ib-d. (82) B. Pl. xu. fig. 6. % Jb d. (78) ii. Pl. vir. fig. 2; Krasser (05) Pl. 1. fig. 10; Schenk (83) A. Pl. yi. fig. 1. + Tenison-Woods (83) A. Pl. tv. fig. 3, 5 Seward (04°) B. figs. 36—38. ® Halle (13) Pls. rv., v. ” Fontaine in Ward (05) B. Pl. xxrv. fig. 2. 8 Schenk (67) A. PL. rx. XL] BAIERA 47 cannot always be distinguished from B. gracilis. The example _ shown in fig. 651 from the Yorkshire coast has been named B. gra- cilis forma Muensteriana to denote its close resemblance to the Rhaetic species. B. Guilhawmali? described by Zeiller from Rhaetic rocks in Tonkin is another similar form but the leaves are narrower and the apices of the segments more obtuse. On the one hand Baiera gracilis approaches close to B. Lindleyana, a species charac- terised by still narrower segments, and on the other it shades into leaves agreeing with Ginkgottes sibirica. Baiera longifolia (Pomel). Pomel’ described this Jurassic species as Dicropteris longifolia and Heer substituted the generic name Baiera*. The leaves re- semble those of B. Simmondsi.in the division of the lamina into narrow linear segments 2—9 mm. in breadth, but the leaf is narrower and cuneate; the segments have obtuse apices. Heer describes the veins as parallel and simple, 3-7 in each segment. With this species Heer associates some male flowers similar to those shown in fig. 654, also some detached seeds, but in neither case is there any convincing evidence of connexion. The Siberian species B. Czekanowskiana®, recorded also by Méller from Bornholm, is probably not a distinct type. Thomas® records B. longifolia from the Middle Jurassic series of Yorkshire and gives new facts with regard to the structure of the epidermal cells: one of his specimens of an incomplete lamina is 12 cm. long, the whole leaf being at least 18cm. in length. The epidermal cells have a very thick cuticle; those on the lower surface are arranged in longi- tudinal rows and most of them have a prominent papilla; on the lower face the cells are more rounded or hexagonal and the stomata are much more numerous; each pair of guard-cells is surrounded by a group of 5—6 subsidiary cells (fig. 647, C, D) as in Ginkgo. Krasser’ records this species from Jurassic rocks 1 Seward (00) B. p. 264. 2 Zeiller (03) B. Pl. u. figs. 16—19. 3 Pomel (49) p. 9. 4 Heer (77) ii. p. 52, Pl. va figs. 2, 3; Pl. vir; Pl. rx. figs. 1—11; Pl. x. figs. 6, 7; Pl. xv. fig. 110. — : 5 Ibid. p. 56, Pl. x. figs. 1—5; Pl. vu. fig. 1; (82) B. Pl. 1. figs. 4—8; Méller (03) Pl. v. fig. 3. 6 Thomas (13) p. 243, Pl. xxv. figs. 3, 4. 7 Krasser (05) p. 18, Pl. 1. fig. 16. 48 GINKGOALES [CH. in China but the photographic reproduction is unfortunately too obscure to afford any indication as to the nature of the specimen. Baiera Phillipst Nathorst. This Jurassic species! (fig. 652) illustrates the absence of any definite dividing line between Baeira and Ginkgoites; it agrees very closely with G. sebirica and with leaves assigned to G. lepida and other ‘species.’ Fig. 652 is drawn from Phillips’ type-speci- men? which he named Sphenopteris longifolia and afterwards trans- ferred to Cyclopteris; his specific name is not retained because Pomel adopted it for a type subsequently called by Heer Barera longifolia’. Krasser records B. Phillipsi from Jurassic strata in Sardinia’. This species shades into B. gracilis and the very similar B.australisMcCoy, one = ; iia 652. Baiera Phillipsi. (York especially resembling some leaves ~ ypysoum; 4 nat. size.) M.S. included by Halle® in the latter species. Baiera Lindleyana (Schimper). Leaves of this type were first figured by Lindley and Hutton® as Solenites? furcata and transferred by Braun to Bavera. Schimper’ subsequently substituted Jeanpaulia and proposed the specific name Lindleyana on the ground that Heer had employed the designation furcata for a Rhaetic species of Baiera. Saporta included this species in Trichopitys. Baiera Lindleyana is characterised by the deep dissection of the lamina into very narrow, filiform, segments and by the presence of a long and slender petiole (fig. 653). Some forms of this.type with rather broader segments are hardly distinguishable from Bavera gracilis. 1 Nathorst (80) A. p. 76. ° Phillips (75) A. Pl. vit. fig. 17. 3 Seward (00) B. p. 270. 1 Krasser (13) p. 5. 5 Halle (13) Pls tv., v. ® Lindley and Hutton (37) A. Pl. 209. 7 Schimper (69) A. p. 683. XL] BAIERA . 49 In a former account of this species! I included the specimen repro- duced in fig. 661 (p. 66), also a similar specimen figured by Phillips? as a distinct type, Baiera micro- phylla. The examination of addi- tional material collected from the Yorkshire coast by Mr Hamshaw -Thomas leads me to substitute Czekanowskia for Baiera as the more appropriate name for the bunch of leaves represented in fig. 661 which is in all probability identical with B. microphylla as figured by Phillips. In the case of \ incomplete leaves it is by no means easy to distinguish B. Lindleyana from Czekanowskia microphylla ; but in the latter the branches of the Mz lamina are separated by a smaller A B angle and if cuticular preparations are available the stomata afford a / means of differentiation: in Bacera the guard-cells are surrounded by Fic. 653. Baiera Lindleyana. a circular group of cells, while in eee ae fs GURUES Bs Czekanowskia the subsidiary cells = are longer and narrower, forming a more oblong group. Baiera Lindleyana is recorded also from Middle Jurassic rocks in Chinese? Dzungaria and from Upper Jurassic rocks in Scotland?. Some specimens described by Fontaine® from the Black Hills (Lower Cretaceous) as Czekanowskia nervosa Heer are, as Berry® points out, probably leaves of a Baiera, and I am disposed to refer ‘them to B. Lindleyana. Baiera Brauniana (Dunker). This species’, represented by leaves from Wealden and Upper Jurassic rocks, agrees in the form and dissection of the lamina 1 Seward (00) B. p. 266, fig. 46 (p. 268). 2 Phillips (75) A. p. 200, fig. 9. 3 Seward (11) Pl. rv. fig. 44. 4 Seward (11?) Pl. v. fig. 105. 5 Fontaine in Ward (99) B. p. 685, Pl. 169, figs. J, 2. - ® Berry (11) p. 374. 7 Dunker (46) A. p. 11, Pl. v. fig. 4. s. IV 4 50 GINKGOALES [CH. with B. gracilis but is distinguished by the smaller dimensions. The imperfect example shown in fig. 641, B from the Kimeridge beds of Sutherland (Scotland)! illustrates the unsatisfactory char- acters on which specific distinctions are drawn in the case of Baiera leaves agreeing in habit with B. gracilis. Better examples are figured by Schenk? from the Wealden of North Germany. Baiera spetsbergensis Nathorst. This species, one of the smallest representatives of the genus, is described by Nathorst? from Upper Jurassic rocks of Spitz- bergen; it is characterised by the very narrow but apparently cylindrical segments and, except in its smaller size, resembles B. Lindleyana. iii. FLowers anp SEEDs. Our knowledge of seeds assigned to Mesozoic and Tertiary representatives of Ginkgo or to Baiera is limited to casts and impressions of detached examples: no reproductive organs have been discovered either in a petrified state or in connexion with a foliar shoot. Reference has already been made to some small Ginkgo-like seeds from the Eocene beds of Sheppey described by Gardner as Ginkgo ? eocenicu. Many similar seeds are figured by Heer from Jurassic strata in Siberia and elsewhere, in most cases as detached seeds but in a few instances borne singly or in pairs on an axis resembling the peduncle of Ginkgo biloha*. Heer’s seeds are correlated with G. digitata, G. sibirica and other species but only on evidence afforded by association with leaves; they are preserved as oval nuts, sometimes enclosed in a carbonaceous envelope possibly representing an outer flesh, and resemble Ginkgo seeds in shape though they differ from them in their smaller size (S—9 mm. long and 6—8 mm. in diameter): in some of the Jurassic specimens the nuts have an apical beak. All that can be said is that seeds similar except in their smaller size to those of the recent species are not infrequently found in association with different species of Ginkgottes. 1 Seward (112) p. 680. ° Schenk (71) B. p. 224, Pl. ut. figs. 9—14. 3 Nathorst (97) p. 53, Pl. ut. figs. 6—12. 4 Heer (82) ii. A. p. 16, Pls. rv, v; (77) ii. p. 57, PL x3 XL] MALE FLOWERS 51 ‘ It has been suggested that the seed-bearing shoots, which Carruthers named Beania, from Jurassic beds on the Yorkshire coast may have belonged to a member of the Ginkgoales, but it is at least equally probable that Beania is Cycadean and possibly the seed-bearing axis of Nilssonia. The genus is described in Ch. xxxvit. It is possible that specimens from Cretaceous and Jurassic rocks regarded by Heer as male flowers of Ginkgoites sibirica and other species, also specimens described by him as Antholithus Schmidtianus?, may be fertile shoots, which bore seeds and not microsporangia, belonging to Ginkgoites or some other member of the Ginkgoales: the nature of these fossils is, however, uncertain and they are described under the generic name Stenorachis. a. Male Flowers. As with seeds so also with regard to the microsporophylls our information is scanty and indecisive. Nathorst? first suggested that some small carbonised bodies from Yorkshire Jurassic beds figured by Phillips*+ as ‘unknown leaves’ are probably fragments of male flowers of some species of Ginkgoites. The specimen of which Phillips figured a small portion is shown in fig. 654, B; it consists of a slender axis with several short and partially broken lateral branches bearing terminal groups of oblong bodies 4 mm. long and 1 mm. wide, 2—4 in each group: these suggest comparison with pollen-sacs with longitudinal dehiscence, and the habit of the whole fertile shoot agrees with that of a male flower of Ginkgo biloba. In the recent species the microsporangia are only about 2mm. long, but in the occurrence of two to four microsporangia on a single microsporophyll the resemblance between the fossil and recent form is fairly close’. Unfortunately it has not been possible to make any preparations of the cuticularised remains showing microspores, and while the probability is that the oblong bodies are microsporangia it is not impossible that they are small seeds. A collection of identical bodies showing what appears to be a median line of dehiscence is illustrated in Part I of The Jurassic Flora of Yorkshire’. A larger specimen is shown in fig. 654, A; 1 Vol. m1. p. 502. 2 Heer (82) A. p. 21, Pl. rx. 3 Nathorst (80) A. p. 75. * Phillips (29) A.; (75) A. Pl. vin. fig. 23. 5 See page 5. 5 Seward (00) B. p. 260, fig. 45. 4—2 52 GINKGOALES [cH. the axis is 2-3 cm. long and some microsporangia are seen in their original position, while others are detached}. It is by no means unlikely that these specimens are portions of male flowers of Ginkgoites digitata or of some other species, but this cannot be definitely settled until better material is available. Some Rhaetic fossils described by Nathorst? as Antholithus Zeillert present a certain resemblance to these supposed male flowers. One of Fic. 654. Antholithus sp. (Sedgwick Museum, Fic. 655. Antholithus Zeilleri. Cambridge; A,ca. x 14; B,ca. x 2.) A, drawn (After Nathorst; x 2%.) by M. Seward; B, drawn by L. -D. Sayers. Nathorst’s specimens from Scania is reproduced in fig. 655 twice natural size; the photograph, for which I am indebted to Prof. Nathorst who published it in 1908, shows a cuticular preparation of the axis and microsporangia. The axis of Antholithus is dichotomously branched and bears terminal clusters of micro- sporangia about 3-5 mm. long, usually eight in a cluster; several of them have dehisced longitudinally and the apices show a slight separation of the two halves. In some of the sporangia Nathorst found microspores with an average length of 40—43y agreeing closely with the spores of Ginkgo and recent Cycads. Nathorst considered that Antholithus Zeilleri may be a male flower of some Ginkgoaceous plant though a correlation with a Cycadean type is by no means excluded. There is, however, a general resemblance between the English Jurassic specimens shown in fig. 654 and the Rhaetic species; the latter is distinguished by a greater tendency + Seward and Gowan (00) B. Pl. rx. fig. 28. ? Nathorst (08) p. 20, Pl. rv. XL] MALE FLOWERS 53 towards a dichotomous habit of the axis, and in the Jurassic specimen we have no proof as to the nature of the ‘ microsporangia.’ Arguments have recently been brought forward! in favour of regarding Antholithus Zeilleri as the male organ of the plant which bore the fronds known as Lepidopteris Ottonis (Gopp.), originally described by Goeppert as Alethopterts Ottonis and made by Schimper the type of a new genus Lepidopteris. Various statements have been made by authors with regard to the occur- rence of sori on this interesting Rhaetic species, but Antevs believes that the evidence hitherto adduced in favour of a fern-like type of fructification is untrustworthy. It is now sug- gested that these Rhaetic compound fronds with thick linear pinnules belong to some seed-bearing plant and that Antholithus Zeillert represents the microspore-bearing organ: there is no proof of connexion, but there is a very close resemblance in the epidermal characters of Lepidopteris and Antholithus and the latter is only found in beds containing the fronds. Nathorst called attention to resemblances between the cuticle of Antholithus and that of Bazera leaves, but according to the later investigations of Antevs, with which it would appear that Nathorst isin sympathy, there is a closer correspondence as regards cuticular structure with Legidopteris. . Fossils regarded by Nathorst as closely allied to his species are described by Leuthardt? from the Keuper of Basel as male flowers of Baiera furcata Heer; these appear to be almost identical with the English Jurassic specimens; the specimens reproduced by Leuthardt consist of long axes—in one of those shown on his plate the axis is 4cm. long—with short lateral branches bearing terminal groups of three or four microsporangia 4—5 mm. long and 1-5—2 mm. broad very similar to those shown in figs. 654, 655. A comparison may also be made with Schenk’s Stachyopitys Preslit? from the Rhaetic of Franconia which he subsequently regarded as the male flower of Batera Muensteriana Heer, a com- parison previously made by Heer*. This type consists of an axis bearing short lateral appendages terminating in oval bodies opening 1 Antevs (14). 2 Leuthardt (03) p. 9, Pl. m1. 3 Schenk (67) A. p. 185, Pl. xxiv. figs. 9—12. 4 Heer (77) ii. p. 52; Schenk (90) A. p. 261. 54 GINKGOALES [CH. at maturity into 10—12 spreading lobes each of which resembles in appearance a microsporangium of Antholithus Zeillert. No, spores have been isolated and, as Nathorst points out, the agree- ment with the Scanian specimens is to a large extent superficial. The Australian specimens, probably of Rhaetic age, described by Shirley! from Ipswich, Queensland, as Stachyopitys annularwides and S. Simmondsi require further investigation; they may be allied to Stachyopitys Preslii Sch., though neither their morpho- logical nature nor systematic position can be settled without fresh data. Halle? describes some examples of a similar kind from the Jurassic beds of Graham Land as Stachyopitys, cf. annularioides Shir. and thinks it probable that they are portions of some Gymno- spermous male strobilus, but, as he points out, the absence of any member of the Ginkgoales in these southern beds is noteworthy. Specimens similar to those described by Schenk, Shirley, and Halle are also figured from Rhaetic beds in South America as Sphenolepis rhaetica? and from the Stormberg (Rhaetic) series of South Africa as Stachyopitys sp.* The generic name Ginkgoanthus has been adopted by Nathorst®5 for a fragmentary specimen from the Upper Jurassic of Franz Josef Land which he considers may be a male flower of a Gink- goites; but the preservation is too imperfect to admit of satis- factory determination. As regards terminology, in the present state of our knowledge it is preferable to use the non-committal designation Antholithus® for the English, Scanian, Swiss, and Franz Josef Land fossils, leaving Schenk’s Stachyopitys Preslii as a type apart. As regards the English and Swiss specimens, the probability would seem to be that they are the microstrobili of some members of the Ginkgoales. STENORACHIS. This generic name’ was first used by Saporta® for Nathorst’s Zamiostrobus scanicus from Rhaetic and Liassic rocks in Scania® 1 Shirley (98) p, 13, Pl. xv. ® Halle (13) p. 88, Pl. vr. fig. 13. 3 Geinitz (76) B. p. 12, Pl. a. * Seward (03) B. p. 66, Pl. rx. fig. 2. 5 Nathorst (99) p. 13, Pl. 1. figs: 33, 49. ° Used by Nathorst in Linnaeus’s and not Brongniart’s sense; Nathorst (08 p. 23. * crevés, narrow; pdxis, the backbone. 8 Saporta (75) A. Pl. oxvm.; (79) A. p. 193. ® Nathorst (75); (97) p. 20; (02) Pl. x. pp. 16, 17. XL] STENORACHIS 55 and for a Liassic species from Belgium, S. Ponceleti. Nathorst : subsequently adopted Saporta’s genus. I have elsewhere suggested! the application of Stenorachis to various species described by Heer from Jurassic and Cretaceous beds as male flowers of Ginkgoaceae. Although there is no proof as to the morphological nature of the specimens included in this genus some of them, e.g. S. scanicus, present the appearance of seed-bearing shoots though, as Nathorst is careful to point out, the seed-like bodies may not be true seeds. I am inclined to regard Heer’s supposed male flowers? (fig. 657) as possibly fertile shoots of some members of the Ginkgoales which originally bore seeds, but this view is merely tentative. Stenorachis is employed as a designation for specimens consisting of a central axis, generally fairly stout, bearing lateral appendages, whether axial or foliar cannot be definitely determined, either simple or forked and in some cases with terminal seed-like bodies but usually with a small distal swelling or a few spreading lobes as in S. Schmidtianus (Heer)®. Some at least of the specimens included in this genus probably belong to Ginkgoaceous plants, though in regard to others, e.g. S. scanicus, it should be remembered that Nathorst inclines towards a Cycadean affinity. The genus Beania was founded on specimens similar in general habit to species of Stenorachis but in Beania the appendages have a comparatively large terminal shield bearing on its adaxial side two seeds. Stenorachis scanicus (Nathorst).- The type-specimen, first described in 1875 as Zamiostrobus scanicus* and afterwards transferred to Stenorachis, is represented by a comparatively slender axis 10cm. long bearing, at a wide angle, several lateral appendages, spoken of by Nathorst as sporophylls; these are split into two divergent arms each of which bears on the side away from the fork an oval, longitudinally striated, body described as thick and woody (fig. 656). The nature of these bodies is uncertain and Nathorst is inclined to think they are not seeds; he suggests as an alternative interpre- tation that they are laminar structures in which microsporangia are embedded. The morphology of this Rhaetic and Liassic 1 Seward (12) p. 23. 2 Heer (77) ii. Pl. x1.; (78) ii. Pl. vi. fig. 8; (82) ii. A. pp. 18, 21, Pls. vr., 1x. 3 Heer (82) ii. A. p. 21. 4 Nathorst (75). 56 GINKGOALES [CH. species referred by Nathorst to the Cycadophyta? is, therefore, uncertain. It is by no means certain that it has not an equal | claim to inclusion in the Ginkgoales; there are no substantial Fic. 656. Stenorachis scanicus. (After Nathorst; A, nat. size; B, x2.) grounds for such relationship, but the resemblance of this and other species of Stenorachis to abnormal seed-bearing shoots of Ginkgo biloba may be significant (cf. fig. 631, D, p. 5). A similar but rather smaller type was described by Heer? from Upper Jurassic rocks in Spitzbergen as Carpolithes striolatus. Nathorst® examined Heer’s figured specimens and recognised one of them as an example of Stenorachis, agreeing in the possession of forked appendages with S. scanicus and bearing seed-like bodies. The fossils described by Shirley* from Rhaetic(?) beds in Queensland as Beania geminata are similar in habit to Stenorachis scanicus and differ from Beania gracilis Carr. in the absence of distally expanded sporophylls. Another Rhaetic species is described by Nathorst® as Stenorachis Solmsi in which the ‘sporophylls’ have a different form and are characterised by a distal, erect, laminar expansion deeply divided into two segments: no seeds or microsporangia have been found. Stenorachis lepida (Heer). The species for which this name has been suggested was originally regarded by Heer as the male flower of the Jurassic species Ginkgo 1 Nathorst (02) p. 16. ® Heer (77) i. p. 47, Pl. rx. fig. 17. 3 Nathorst (97) p. 20, Pl. 1. fig. 15. 4 Shirley (98) p. 16, Pl. xx. 5 Nathorst (02) p. 17, Pl. 1. figs. 18—21. XL] STENORACHIS 57 lepida (= Ginkgoites sibirica) and described by him under that name?. Similar specimens are correlated by Heer with other species of Ginkgo leaves, and Ginkgo grandiflora? Heer is repre- sented by supposed male flowers only. Similar though rather larger examples are described by Heer from Jurassic beds in Siberia as Antholithus Schmidtianus® and regarded as male flowers of some member of the Ginkgoales, possibly Phoenicopsis; what- ever the parent-plant may have been it is clearly a type closely allied to those he refers to different uae of snide Fig. 657, B Fic. 657. Stenorachis lepida. A, from Amurland; B, C, from Afghanistan. (A, B, nat. size.) shows a specimen of Stenorachis lepida from Jurassic beds in Afghanistan? which is undoubtedly of the same type as Heer’s European examples. One of Heer’s specimens from the Jurassic beds of Amurland® is shown in fig. 657, A: a curved and fairly stout axis bears numerous, spirally disposed, appendages with slightly expanded ends which in a few cases are more or less definitely bilobed. No remains of seeds or microsporangia are preserved, but the swollen ends of the appendages suggest the former presence of some reproductive organs: some of the ap- pendages are bilobed as in the Afghan example. Heer states that some of his specimens bear 2—3 pollen- sacs at the tips of the appendages, but the published figures afford no 1 Heer (77) ii. Pl. x1. 2 Heer (82) A. p. 18, Pl. vr. figs. 1—6. 3 Ibid. p. 21, Pl. rx. 4 Seward (12) p. 23, Pl. rv. fig. 52. 5 Seward (12) p. 28, Pl. 1. fig. 8. 58 GINKGOALES [CH. confirmation of this and an examination of some of Heer’s material lent to me through the kind offices of Dr Zalessky failed to reveal any indication of spores or sporangia. In Heer’s Antholithus Schmidtianus the lateral appendages are said to bear 3—4 pollen- sacs in a terminal whorl, but Heer also suggested the possibility that these bodies are the segments of a calyx-like envelope, a more probable interpretation. It may be that the terminal bodies are homologous with the slightly expanded distal ends of the append- ages in S. lepida and possibly with the collar at the base of the ovules of Ginkgo biloba, which in the case of S. Schmidtianus has the form of a lobed cupular organ which enclosed a seed. It is noteworthy that Heer’s figures show a central scar surrounded by the spreading lobes. The incomplete Jurassic specimens from Victoria (Australia)! described as possibly parts of a female shoot of a Ginkgoaceous plant resemble Stenorachis lepida and should be referred to the same genus. ; A specimen like those represented in fig. 657 has been figured by Krystofovié? from Jurassic rocks in Ussuriland as Ginkgo sp. An imperfect fossil described from Jurassic beds in Australia as - possibly a seed-bearing shoot of a Ginkgoaceous plant® should be included in Stenorachis, as also Raciborski’s Ixostrobus Siemirad- zkui* from Rhaetic beds of Poland. In no case have we any decisive evidence with regard to the parentage or morphological nature of the specimens referred to Stenoyachis, but any material that may represent fertile shoots belonging to Ginkgoales or Cycadophyta should be described in the hope that additional facts may be obtained. ERETMOPHYLLUM. Thomas. A genus founded® on some well-preserved leaves from the Middle Estuarine (Middle Jurassic) beds of Gristhorpe Bay on the Yorkshire coast, and named Eretmophyllum from the paddle-like form of the lamina’. Leaves oblanceolate to linear reaching a length of 12cm. and a breadth of 2cm.; in the type-species, 1 Seward (04?) B. p. 179, figs. 39, 40. ® Krystofovic (10) Pl. 1. fig. 5. 3 Seward (042) B. Pl. xrx. 4 Raciborski (92) Pl. 1. figs. 5—8. 5 Thomas (13). ® éperudv, a paddle. XL} ERETMOPHYLLUM 59 E. pubescens Thom., the leaf is from 7 to 10 cm. long and 1-3 cm. broad; the apex is rounded or retuse (fig. 658, B), the base tapering gradually towards the petiole. Veins 1---1-5mm. apart, dicho- tomously branched in the proximal part of the lamina and usually parallel and simple except where they converge at the apex. The epidermal cells (preserved in the Yorkshire species) are polygonal, with or without papillae (fig. 658, D); the stomata are characterised by an enclosing group of subsidiary cells as in Ginkgo. Eretmophyllum pubescens Thomas. Secretory tracts occur between the veins of the smooth lamina (fig. 658, C) like those in the leaves of Ginkgo. The polygonal cells with straight or slightly undulate walls _ are characterised by papillae (fig. 658, D), one on each cell: these are particularly conspicuous on the lower epidermis to which the sto- mata are confined; the slightly depressed stomata are in regular rows and the guard-cells are sur- rounded by 4—7 subsidiary cells. In another species, E. whitbiense from Whitby, the surface of the lamina, which may be 7cm. longand 1-2. cm. broad and slightly falcate, is rough, and between the veins are strands of elongated cells, pos- sibly denoting the presence of hypo- dermal stereome. Stomata occur on both surfaces and the papillae Fic. 658. A, Hretmephyllum saigha- are confined to the subsidiary cells. nense. B—D, BE. pubescens. D, A leaf figured by Ettingshausen? epidermal cells. (A, after Seward ; from Wealden rocks as Cyclopteris B—D, after Thomas.) squamata, which Schenk? suggests may be a segment of a Gunk- gottes, should probably be included in Eretmophyllum. 1 Ettingshausen (52) Pl. 1v. fig. 1, 2 Schenk (71) B. p. 213. 60 GINKGOALES [cH. XL Eretmophyllum saighanense (Seward). This species (fig. 658, A), originally referred with some misgiving to Podozamites! and compared with Yokoyama’s Ginkgodium, is from Jurassic beds in Afghanistan. There can be little or no doubt as to its generic identity with the Yorkshire leaves The broadly linear lamina tapers gradually to a slender petiole and the veins, 1 mm. apart, are simple except at the proximal end. Mr Thomas is certainly justified in his opinion that Hretmo- phyllum is a member of the Ginkgoales. In shape the leaves resemble Ginkgodium and differ but little from some Jurassic specimens referred to Ginkgoites. They agree in venation, in the presence of short secretory tracts, in the structure of the epidermal cells and stomata with Ginkgo and species of Ginkgoites. Eretmo- phyllum is distinguished from Feildenia by its larger leaves, a coarser venation, and a more definite petiole. Some leaves figured by Fontaine? from Jurassic-Cretaceous rocks of Alaska as Nagewopsis longifolia? Font. have little claim to be included in that genus* and may perhaps be allied to Hretmophyllum. 1 Seward (12) p. 35, Pl. rv. fig. 53. 2 Fontaine in Ward (05) B. Pl. xiv. figs. 1—5. 3 Berry (10) p. 190. CHAPTER XLI. GENERA BELIEVED TO BELONG TO THE GINKGOALES BUT WHICH ON THE AVAILABLE EVIDENCE CANNOT BE RE- FERRED WITHOUT HESITATION TO THAT GROUP. GINKGODIUM. Yokoyama. Yokoyama? defined the genus as follows: ‘Leaf coriaceous, entire or lobed, gradually narrowed towards the base which is thickened at its margin and gradually passes into a short petiole ; veins numerous, simple, parallel; interstitial veins very fine.’ He draws attention to the thickening of the lower margin of the lamina, a feature reminiscent of Ginkgo and to the course of the veins which run parallel to the median axis of the lamina instead of spreading from the base as in Ginkgoites and Barera. Gunk- godium resembles the Palaeozoic genus Whittleseya in the position of the veins but the genera are unlikely to be confused; a com- parison may also be made with the Jurassic genus Hretmophyllum (fig. 658) which has longer and narrower leaves with a coarser venation. We have no information with regard to the cuticular structure, the nature of the supporting axes or reproductive organs. The supposed affinity to Ginkgo rests therefore on leaf-form alone. Ginkgodium Nathorstt Yokoyama. The type-species was founded on specimens from strata in Japan assigned by Yokoyama? to the Middle Jurassic series, but the flora suggests a somewhat higher horizon in the Jurassic system. Some of the leaves are entire, obovate, and have a truncate distal end; others are cuneate and broader at the apex which may be lobed, while in some forms the leaf is divided by a 1 Yokoyama (89) B. p. 56. 2 Ibid. p. 57, Pls. W1., W., VII., IX., XI. 62 GINKGODIUM AND CZEKANOWSKIA [cH. deep median sinus into divergent obtuse segments (fig. 659). One leaf is described as 6-6 cm. long, 2-1 cm. broad with thirty veins and an interstitial ‘vein’ between each pair: the interstitial ‘vein’ is due to the presence of an inter- costal stereome strand. Thomas} records this species from the Bathonian series of Kamenka in the south of Russia (fig. 659, B). The specimens from Alaska named by Fontaine? Ginkgodium? alaskense agree more closely with Ginkgoues. CZEKANOWSKIA. Heer. Heer? gave this name (after Fic. 659. Ginkgodium Nathorsti. (Nat. Czekanowski who discovered the Ha a Mokeyunieg 2s etter specimens) to fascicules of long , and narrow, filiform, leaves with a simple or occasionally forked lamina borne on a short supporting axis covered with broader and shorter scale-leaves. The deciduous fascicules or dwarf-shoots are similar to those of Phoenicopsis. Bunches of Czekanowskia leaves with their short scale-covered supporting axes resemble the dwarf-shoots of Pines’. Heer assigned to this genus some seeds associated with the leaves, also what he believed to be a male flower®, an example of a reproductive shoot of the type described on page 57 as Stenorachis. There is, however, no conclusive evidence as to the nature of the reproductive organs. The venation is seldom shown on the carbonised laminae; some leaves are finely striated while on others there may be one or two narrow ridges that represent veins, but as a rule the impressions afford no indication of the venation. Czekanowskia was placed by the author of the genus in the Ginkgoales, the short shoots being compared with those of Ginkgo though, except in the larger number of the leaves, they closely resemble the foliar spurs of 1 Thomas (11) p. 75, Pl. rv. figs. 9—11; Pl. vim. fig. 3. 2 Fontaine in Ward (05) B. p. 168, Pl. xxiv. figs. 3, 4. 3 Heer (77) ii. p. 65. 4 Cf. Pinus flexilis; Bot. Mag. Tab. 8467. 5 Heer (82) ii. Pl. vi. fig. 7. XLI] CZEKANOWSKIA 63 Pines. The dichotomous branching of the lamina in some forms is another feature in which Czekanowskia resembles’ Baiera and Ginkgo, a resemblance which derives a certain significance from the occurrence of stomata of the Ginkgoaceous type. Nathorst} has described the cuticular membranes of the superficial layers: the epidermal cells have straight walls and the stomata, more numerous on the lower surface, are accompanied by four or five subsidiary cells: these do not form a circular group as in Baiera and Ginkgoites, but, as the result of elongation in the direction of the long axis of the leaf, the group is relatively long and narrow. Cuticular preparations can often be made from the well-preserved leaves that occur in great abundance in the shales of Gristhorpe Bay and elsewhere on the Yorkshire coast?, and some particularly good examples were collected by Prof. Obrutschew from Jurassic rocks in the Djair Mountains in Chinese Dzungaria*®; these occur with carbonised remains of Ginkgoites leaves in papery masses similar in the manner of preservation to the Palaeozoic paper coal.from Russia*. The epidermal cells of the Dzungaria Czek- anowskia, possibly identical with C. rigida but too incomplete to be determined with certainty, have straight walls and are relatively long ; the stomata are scattered and appear as dark patches, their darker colour being due to the thick cuticles of the two or three flanking cells on the sides of the stoma; the epidermal features are similar to those described by Nathorst®, but in the Rhaetic specimens - from Scania the heavily cuticularised accessory cells are generally more numerous. It has been suggested by Jeffrey® that Czekanowskia may be Araucarian in its affinities, but this opinion rests on the slender evidence of association of Czekanowskia-like leaves in Middle Cretaceous rocks with a stem described as Araucariopitys so named because of the association of Araucarian and Abietineous features. Such evidence of affinity as we have would seem to be in favour of relationship with Baiera and Ginkgo though decisive data are not as yet available. The genus is very widely spread in 1 Nathorst (06); see also Seward (00) B. p. 278. 2 Seward (00) B. p. 278, fig. 48. 3 Seward (11) p. 49, Pls. Iv., V., VI., VIT. 4 Vol. 1. p. 68. 5 Nathorst (06). 6 Jeffrey (07); Hollick and Jeffrey (09) B. p. 63, Pl. vr. figs. 1 —-3. 64 CZEKANOWSKIA [CH. Jurassic floras and a few examples are recorded from Cretaceous strata. zekanowskia Murrayana (Lindley and Hutton). On the specimen shown in fig. 660, A Lindley and Hutton! founded the species Solenites Murrayana which they compared A Fic. 660. A, B, Czekanowskia Murrayana. A, The type-specimen of Solenites Murrayana, Lind. and Hutt., from the Middle Jurassic plant-beds of York- shire. (British Museum, no. 3685, no. V. 3684.) with /soetes and Pilularia. The type-specimen is from the Middle Jurassic plant-bed at Gristhorpe Bay on the Yorkshire coast. They describe the narrow leaves as converging to a common ? Lindley and Hutton (34) A. PL. oxxz. XLI] CZEKANOWSKIA 65 point, but the actual axis is not preserved; the lamina is longi- tudinally striated but no veins are shown either on the carbonised lamina or in the magnified cuticle figured in the original description. In the specimen reproduced in fig. 660, B the leaves are seen to be attached to a short and relatively broad axis covered with scale-leaves, one of which is shown bent over on one side of the dwarf-shoot. The comparatively large size and the pendulous position of the scales are characteristic features of the genus which are well seen in figures of Czekanowskia published by Nathorst'. This species was included by Saporta in Jeanpaulia and later transferred to Trichopitys, while Zigno? and some other authors regarded Solenites Murrayana as a species of Isoetes. The com- parison with Isoetes suggested by the form of the leaves is not borne out by the structure of the epidermal cells. Phillips® figured a specimen in 1829 as Flabellaria ? viminea: this specific name though employed before the publication of Murrayana has not been adopted by authors. Some of the specimens included by Heer in his account of the species C. rigida should be referred to C. Murrayana, but in a previous description* of the species I went too far in uniting C. Murrayana and C. rigida. In Czeka- nowskia Murrayana the leaves, usually about !mm. broad but sometimes narrower, reach a length of more than 17 cm.; they are unbranched and in this respect and in their slightly greater breadth differ from C. rigida. The species is characteristic of Middle Jurassic floras. Czekanowskia microphylla (Phillips). The specimen figured by Phillips® from the Yorkshire coast as Baiera microphylla is undoubtedly identical specifically with that reproduced in fig. 661, and both were formerly included in Baiera Lindleyana®. The chief reasons for transferring them to Czekanowskia are the more acute angle of divergence of the filiform segments, the difference in the shape of the leaves, the absence of a petiole, and the occurrence of the Jeaves in a fascicle, a habit not shown by any typical examples of B. Lindleyana though not 1 Nathorst (06) Pl. it. 2 Zigno (56) A. p. 216. For other references, see Seward (00) B. p. 280. 3 Phillips (29) A. Pl. x. fig. 12. 4 Seward (00) B. p. 279. 5 Phillips (75) A. p. 200, fig. 9. ® Seward (00) B. p. 266. Ss. IV 5 66 CZEKANOWSKIA [cH. unknown in the genus (e.g. B. paucipartita). Some specimens which may be identical with this type were obtained several years ago by Dr Nathorst from Yorkshire but never fully described: an examination of his unpublished drawings and of specimens col- lected by Mr Hamshaw Thomas convinced me that some forms of Czekanowskia are more freely branched and exhibit more varia- tion in the breadth of the lamina than I had formerly supposed. In specimens of the type first noticed by Nathorst some of the segmentsare comparatively broad and fern-like, a feature that is not seen in the leaves shown in fig. 661. This species affords a striking contrast to Czekanowskia Murrayana in which the long ernie olan mente e/a, leaves are unbranched, and as 283; nat. size.) Nathorst+ suggests the name Solenites might be revived for the unbranched type; but in the absence of any difference in the epidermal characters, it would seem undesirable to raise to generic rank a feature depending on the simple or branched habit of the leaves of otherwise similar leaf-fascicles. Czekanowskia rigida Heer. This species, founded on specimens: from Srberian Jurassic rocks, is characterised by its branched filiform leaves borne on short shoots enclosed by scale-leaves, triangular or lanceolate in form and in some specimens pendulous on slender stalks. Nathorst® states that the epidermal structure of the scale-leaves is similar to that of the scale-leaves on short shoots of Ginkgo. The characters of the stomata are mentioned in the account of the 1 Nathorst (06) p. 11. . ? Heer (77) ii. p. 70; (78) ii p. 7; (82) A. p. 19. 3 Nathorst (06). XLT] ‘ CZEKANOWSKIA 67 genus. The leaves are generally slightly narrower than the unbranched needles of C. Murrayana, but the habit of the dwarf- shoots is the same. The leaves often show fine striations; in most specimens there is no indication of clearly marked veins though two or three vascular strands are sometimes visible. Heer on very slender evidence refers to this species some seeds and a ‘male flower.’ It is not always easy to distinguish between imperfect examples of C. rigida and Batera Lindleyana: the leaves of the latter type are petiolate and the segments diverge at a wider angle. Two leaves with spreading bifurcate segments figured by Fontaine? from Lower Cretaceous rocks in the Black Hills as Czekanowskia nervosa Heer afford no indication that they were borne in clusters on dwarf-shoots but resemble the petiolate leaves of Baiera Iandleyana. Berry? points out a similarity between Fontaine’s fossils and Barera foliosa Font. Heer’s type-specimens of C. nervosa from the Wealden of Portugal® are more like Czekanowskia leaves. The leaves described from Siberia as C. setacea Heer, though narrower than some forms of C. rigida, are probably not specifically distinct. Czekanowskia rigida is characteristic of Jurassic strata, and occurs in Europe, including Greenland, also in Siberia, China, and Japan. Czekanowskia dichotoma Heer and C. capillaris Newberry. The branched leaves described under these names> from Cretaceous rocks in Greenland and North America are in most cases not sufficiently complete to be assigned with certainty to the genus Czekanowskia; the examples figured by Hollick and Jeffrey® from Middle Cretaceous beds as C. capillarts occur in closely packed groups, but no specimens have been discovered showing any scale-covered supporting axis. While admitting the probability that these species and C. nervosa’ from Wealden strata 1 Fontaine in Ward (99) B. p. 685, Pl. cLx1x. figs. 1, 2. 2 Berry (11) p. 374. 3 Heer (81) Pl. xvi. 4 Heer (77) ii. p. 68; (78) ii. p. 26; (82) A. p. 18. 5 Heer (82) A. p. 8. Newberry and Hollick (95) p. 61. 6 Hollick and Jeffrey (09), B. p. 63, Pl. v1. figs. 1—3. 7 Heer (81) Pl. xvi. : 68 FEILDENIA : [cH. may be allied to Czekanowskia rigida, such evidence as is available points to a maximum development of Czekanowskia in the Jurassic period. FEILDENIA. Heer. In 1870 Heer? described some small linear leaves from Tertiary strata in Spitzbergen for which he proposed the generic name Torellia, defining it as follows: ‘Folia rigida coriacea, basin versus angustata, articulata, tenuiter costata, costis interstitiisque sub- tilissime striatis.’ On the discovery by Capt. Feilden of additional specimens in Miocene beds in Grinnell Land (81° 46’ N.) Heer published a further account of the genus and substituted Feddenia for Torellia because of the previous use of the latter name by Zoologists. Heer compared Fetldenia with Podocarpus, Araucaria, and other Conifers but, mainly because of the occurrence of a leaf with a lobed lamina, he provisionally included the genus in the Taxineae?. The leaves are usually found as detached speci- mens but in one case several are spirally disposed on a stout axis and one imperfect example shows at the base what appears to be a scale-leaf, suggesting that leaves were also borne on short shoots like those of Phoenicopsis and Czekanowskia. Heer lays stress on the ribbing and striation of the surface of the lamina as dis- tinguishing features between Feildenia and Phoenicopsis, but Nathorst®, in his revision of the genus, expresses the opinion that it is only in the tendency to a sickle-like form and a feeble expansion of the slightly curved base that Fedldenia, at best an ill-defined genus, can be distinguished from Phoenicopsis. : Feildenia rigada Heer. This species, from Miocene beds of Spitzbergen* and Grinnell Land§, is represented by linear leaves 6—8 cm. long and 5—8 mm. broad at the widest part, usually rather nearer the apex than the base; the lamina is often slightly falcate and tapers gradually to a narrow base. There are 8--11 veins for the most part parallel but occasionally feebly convergent at the bluntly rounded apex. 1 Heer (71) iii. p. 44. 2 Heer (78) i. p. 20. 3 Nathorst (97), p. 55. 4 Heer (71) ii. p. 44, Pl. vi. figs. 3—12; Pl. xvi. fig. 1b. 5 Heer (78) i. p. 20, Pls. 1, 0, Vim. XLT] PHOENICOPSIS 69 Feildenia Nordenskéldi Nathorst. A species from Upper Jurassic rocks in Spitzbergen founded by Nathorst! on leaves similar to those of F’. rigida but smaller; the lamina is generally 3—4 mm. broad and may reach a length of 45cm. There are usually six veins and as in other species finer longitudinal lines occur between the true veins. A few small leaves very similar to F. Nordenskéldi are described by Nathorst as Feildenia sp.” from Franz Josef Land, probably of Wealden age. Until further evidence is available it is impossible to fix pre- cisely the position of the genus. Though often distinguished by the sickle-shaped lamina and the broad apical region from leaves of Podozamites it is not always possible to separate the leaves of the two genera. PHOENICOPSIS. Heer. DESMIOPHYLLUM. Lesquereux. Phoenicopsis was founded by Heer? on linear leaves from Middle Jurassic strata in Siberia ; the leaves, in extreme cases 20cm. long and varying in breadth from 2 mm. to 2 cm., occur in clusters of six or more and even as many as twenty on very short and relatively broad axes covered with small scale-leaves. These dwari-shoots were deciduous: the lamina is fairly uniform in breadth but tapers gradually towards the slender base and is usually obtusely rounded at the apex; the veins are parallel and very rarely dichotomously branched. The features on which species are founded are often of little systematic value: they are the breadth of the lamina, the arrangement of the veins, the presence or absence of interstitial ‘veins.’ It is very doubtful whether much confidence can be placed on the occurrence of the so-called interstitial veins: in some species of Phoenicopsis the parallel veins show no trace of a smaller ‘vein’ between them, but occasionally in a leaf of the same species there are indications of interstitial ‘veins.’ The breadth of the lamina is also an uncertain guide: well preserved specimens show that the leaves may reach a considerable length and that the lamina gradually decreases in breadth towards the narrow base. Species have been needlessly multiplied particularly 1 Nathorst (97) p. 56, Pl. mr. figs. 16—27. 2 Nathorst (99) p. 15, Pl. 1. figs. 25—30, 32. 3 Heer (77) ii. p. 49. 70 PHOENICOPSIS [cH. in the case of detached leaves which it is often impossible to determine even generically. The characters usually employed for the separation of different forms are conveniently shown in a table published by Krasser!. There is no information available as to the epidermal structure of the various types of Phoenscopsis leaves, nor have we any data with regard to the reproductive organs. The genus is generally included in the Ginkgoales: the dwarf-shoots agree closely with those of Czekanowskia which, in the structure of the epidermis and in the bifurcation of the leaves, resembles Ginkgo and Baiera. The precise position of Phoenicopsis cannot be regarded as settled. The only evidence with regard to ana- tomical structure is that furnished by Solms-Laubach? who described petrified leaves from Jurassic rocks in Franz Josef Land which are probably examples of Phoenicopsis; but, assuming that they belong to this genus, the anatomical data are insufficient to determine the position of the genus within the Gymnosperms. Incomplete and detached leaves agreeing in their venation and in the form of the lamina with those of Phoenicopsis cannot as a rule be distinguished from leaves of Podozamites, Feildenia, or even from narrow forms of Cordactes. The Jurassic specimens from North Germany on which Salfeld? founded the genus Phyl- lotentia should probably be assigned to Phoenicopsis. Solms- Laubach refrained from assigning the imperfect Franz Josef Land leaves to Phoenicopsis because no dwarf-shoots were found; he employed the non-committal generic name Desmiophyllum, a designation that might with advantage be more frequently used for specimens which cannot be proved to belong to Phoenicopsis, Podozamites or other genera with similar leaves. Desmiophyllum. Lesquereux established the genus Desmio- phyllum4 for some narrow sublinear leaves from the Coal Measures of Pennsylvania similar to those of Poacordaites and attached to an imperfectly preserved axis either singly or in small groups. The type-species D. gracile is probably a species of Cordattes: the name Desmiophyllum never came into general use until its revival by Solms-Laubach in 1904 as a convenient term to apply to linear 1 Krasser (05) p. 612. 2 Solms-Laubach (04) Pls. 1., 1. 8 Salfeld (09) B. p. 26, Pl. rv. fig. 3. 4 Lesquereux (78) p. 322; (80) A. p. 556, Pl. 82, fig. 1. XLI] DESMIOPHYLLUM- 71 leaves that in the absence of evidence as to the habit of the shoots cannot be assigned to more precisely defined genera such as Phoenicopsis or Podozamites. Nathorst’ employs Desmiophyllum for some Phoenicopsis-like leaves from Jurassic rocks in the New Siberian Islands which may be specifically identical with those described by Graf Solms from Franz Josef Land, which I propose to call Desmiophyllum Solmsi. In view of the probability that these leaves belong to a species of Phoenicopsis, a brief description of their structure may be conveniently inserted here. Desmiophyllum Solmsi sp. nov. The collection of plants obtained by the Jackson-Harmsworth Expedition (1894—96) to Franz Josef Land includes several specimens of matted linear leaves some of which are figured by Newton and Teall?. Similar leaves collected by Nansen are described by Nathorst®. Subsequently Solms investigated the structure of the leaves figured by Newton and Teall and the photographs reproduced in fig. 662 were taken from sections of the silicified material in the Museum of the Geological Survey. The largest specimens reach a length of 10 cm. and are 5—10 mm. broad; the veins are unbranched and there are six in a breadth of 3mm. of lamina. In transverse section the lamina is of fairly uniform breadth or, owing to the partial collapse of the intercostal mesophyll, it is characterised by prominent ribs (fig. 662, . The lamina is 1 Dawson (63) Pl. xv. fig. 6. 2 Seward and Woodward (05) B. 3 Seward (075) B. 4 Arber (12) p. 400 (footnote). 5 Seward (075) B. XLIt] PSYGMOPHYLLUM 87 divided by a deep median sinus into two bilobed segments and agrees closely with some species of Baiera. By some authors this species would be included in Ginkgophyllum but, as already stated in the account of the genus, the degree of dissection of the leaves is too variable and unimportant a character to be made the basis -of a generic differentiation. Psygmophyllum Girasserti (Saporta). This Permian species from Lodéve}, France, agrees closely with P. flabellatum in the size and outline of the leaves as also in their Fic. 669. Psygmophyllum Grasserti. (After Saporta; § nat. size.) method of attachment to the axis, but it is distinguished by the division of the lamina into linear segments (fig. 669). A species with similar leaves, from the Permian beds of the Ural mountains, is figured by Saporta as Ginkgophyllum kamenskianum?. J ohnston® describes a species, Ginkgophyllum australe, from ‘Mesozoic’ rocks 1 Saporta (75) p. 1018; (84) Pl. 152, fig. 2. 2 Saporta and Marion (85) p. 144, fig.73, B; Saporta (82). 3 Johnston (86) p. 178. 88 GENERA OF UNCERTAIN POSITION [CH. in Tasmania which he compares with G. Grasserti Sap., but as he gives no illustration of the fossil no opinion can be formed of its true nature. Psygmophyllum kiltorkense Johnson (= Kiltorkensia devonica John.). A species! recently described from the Upper Devonian grits of Kiltorcan, Ireland, characterised by fan-shaped leaves 7 cm. long and 5 cm. broad, deeply divided into two symmetrical halves each of which is again divided into two ribbon-like segments with a Ginkgo-like venation. The leaves agree closely with those of P. Grassertt in their general form and in the lobing of the lamina. Johnson believes this type to be an ancestral form of Ginkgo though there are no adequate grounds for such a view. Psygmophyllum ? crenatum (Brauns). The close resemblance presented by the Rhaetic leaves from near Braunschweig, originally described by Brauns? as Cyclopterzs crenata and subsequently figured by Nathorst® as Ginkgo ? crenata, to some Permo-Carboniferoys leaves included in Psygmophyllum suggests affinity with that genus. The obovate lamina, approxi- mately 12cm. long, is slightly lobed on the upper margin and contracted below into a stalk-like base; the forked veins are nearly 3mm. apart. Nathorst compares the species with Psygmo- phyllum flabellatum Lind. and Hutt., but the Rhaetic specimens differ from the English type in their much coarser venation: in the lobing of the lamina and in the coarse venation there is a much greater similarity to the broader leaves described by Arber* as | Psygmophyllum majus. In view of the incomplete nature of the material it is inadvisable to adopt the name Psygmophyllum without reservation. An examination of Nathorst’s specimen in the Stockholm Museum led me to regard it as more probably an example of Psygmophylium than of Ginkgo. 1 Johnson (14). Since this account was written Prof. Johnson has described additional material including stems and foliage [Johnson (17)] demonstrating the occurrence of repeatedly forked filamentous leaves [or leaflets] attached to slender axes bearing also the broader form of lamina. The plant, which he now refers to a new genus Kiltorkensia, may well be a Pteridosperm with compound fronds and dimorphic pinnules. 2 Brauns (66) p. 52, Pl. xm, fig. 8. 3 Nathorst (78). + Arber (12) p. 392, Pls. xnm.—xtiv. XLIt] PSYGMOPHYLLUM 89 Other records of Psygmophyllum. Psygmophyllum primigenium (Saporta). Some leaves dis- covered by Grand’Eury in Permian rocks of the Urals were described by Saporta as Salisburia primigenia and regarded as ‘the prototype of the surviving species. The original specimens are unfortunately not available, but from the published figures it would seem that the species is of the same general type as P. flabellatum. Reference has already been made to an American Devonian species referred by Dawson to Cyclopteris and recently transferred to Psygmophyllum. A leaf or leaflet described by Dawson from Gilboa, New York, as Noeggerathia gilboensis® affords a good example of a specimen that may be a Psygmophyllum leaf ora leaflet of a frond of the Noeggerathia type. Lesquereux® considers Dawson’s specimen to be a pinnule of Palaeopteris. The same remark applies to Lesquereux’s species Noeggerathia obtusa* from the Coa] Measures of Pennsylvania included by Arber in Psygmo- phyllum; it is probably a pinnate frond. The Russian species P. expansum and P. cunedfolium are dis- cussed in the account of the genus. The species Psygmophyllum Delvali> Camb. and Ren. from the Westphalian of Belgium is now admitted to be a leaf of Cordaites. A species described by Sand- berger® from the Permian of the Black Forest as Ginkgophyllum minus has been assigned by Sterzel’ to Dicranophyllum. A leaf figured by Schmalhausen® from the Permian of Kast Russia as Baiera gigas is no doubt a Psygmophyllum allied to P. Kidston. A fragment figured by Schenk from China as Ginkgophyllum sp. is too imperfect to determine, and the specimens from the same locality described as Psygmophyllum angustilobum® are, as Zeiller points out, pirinules of a frond of the Hremopteris type. A sufficient number of examples have been described to illustrate the range of the genus and the unsatisfactory nature of 1 Saporta (82); Saporta and Marion (85) p. 145, fig. 74. 2 Dawson (63) p. 463, Pl. xvu. fig. 6; (71) A. p. 46, Pl. xvr. fig. 172. 8 Lesquereux (80) A. p. 305. 4 Ibid. Pl. xix. figs. 6, 7. 5 Cambier and Renier (10) Pl. vi. fig. 1. 8 Sandberger (90) p. 101. 7 Sterzel (07) p. 820. 8 Schmalhausen (87) Pl. v. fig. 10. ® Schenk (83) A. p. 221, figs. 7, 8; Pl. xu. figs. 22—24. 90 . GENERA OF UNCERTAIN POSITION [cH the material from a botanical point of view. Failing reproductive organs or petrified specimens some useful evidence might be afforded by an examination of the cuticular structure of well preserved leaves. RHIPIDOPSIS. Schmalhausen Schmalhausen! instituted this genus for large petiolate oval leaves from the Permian rocks of the Petschora district, charac- terised by the division of the lamina into several obcuneate or obovate segments closely resembling in their form and venation some forms of Psygmophyllum especially P. Haydeni®. We have no definite information as to the systematic position of the parent- plant; the genus has usually been regarded as a representative of the Ginkgoales on the ground of similarity in the leaves, but while admitting that a relationship between Rhipidopsis and Ginkgo is not improbable it is the safer course to regard Rhiprdopsis as a genus of Gymnosperms of uncertain affinity. Schmalhausen attributes to Rhipidopsis some Samaropsis seeds* found in associa- tion with the leaves, and Kurtz‘ states that he has found leaves and ‘fruits’ in the Argentine. No proof of any connexion between leaves and seeds has so far been discovered. The genus is recorded from Russia, South America, and India from strata that are Permian or approximately Permian in age. Rhipidopsis (fig. 670) is distinguished from Psygmophyllum by the presence of a petiole and from most forms of that genus by the deeper dissection of the lamina, as also by a more pronounced difference in form and size between the several segments of the lamina. Zeiller> has drawn attention to a close resemblance between Rhipidopsis and a specimen figured by Schmalhausen from the Artinsk Permian beds as Psygmophyllum expansum®. Rhipidopsis ginkgoides Schmalhausen. The type-species (fig. 670) is characterised by the large size of the leaves which, according to Schmalhausen’, may reach a 1 Schmalhausen (79) A. p. 50, Pls. vi., viz. 2 See p. 86, fig. 668. 3 Schmalhausen (79) A. Pl. vim. figs. 9—11. 4 Zeiller (96) A. p. 467. 5 Ibid. p. 471. ® Schmalhausen (87) Pl. m1. fig. 10. ? Schmalhausen (79) A. p. 50, Pls. v1., vim. XLIt] RHIPIDOPSIS 91 length including the petiole of 30cm. and a breadth of 11 cm. The segments, 6—10 in number, are often free to the summit of the petiole; they vary considerably in shape and size, the median segments are obcuneate with a broad rounded truncate margin, Fic. 670. Rhipidopsis ginkgoides. (From a photograph of the original of one of Schmalhausen’s figures supplied by Prof. Zalessky.) while the lateral lobes are obovate asymmetrical. The repeatedly forked veins are 1—1-5 mm. apart in the lower part of the lamina but much more crowded in the apical region. The slender petiole 92 GENERA OF UNCERTAIN POSITION {cn.. reaches a length of 10 cm. This species has been recorded from the Argentine but no figures have been published. It is not improbable that the seeds of the Samaropsis type associated with the leaves in the Russian and Argentine localities may belong to this genus, but proof is lacking. Rhipidopsis densinervis Feistmantel. This Indian species from the Raniganj group of the Damuda series! (Lower Gondwana) is founded on some leaf-impressions very similar in size and form to Rhipidopsis ginkgoides. The presence of a petiole is shown on one of the figured specimens: the lamina is deeply divided into about six obcuneate segments that appear to be irregularly lobed on the truncate margin. Rhipi- dopsis densinervis is distinguished by its dense venation and by a difference in size between the lateral and median segments less than in the leaves of R. ginkgoides. Kurtz? states that some specimens found by him in Permo-Carboniferous beds in the Argentine may belong to this species. Rhipidopsis gondwanensis sp. nov. The specimens for which this name is proposed were described by Feistmantel as Rhipidopsis ginkgoides® from the Barakar group of the Damuda series. My examination of the type-specimens confirms Feistmantel’s statement that they agree closely with Schmalhausen’s Russian leaves except in their much smaller size: the Indian leaves reach a length of 3 cm. while in Schmalhausen’s species the lamina may be 14cm. in length. In view of this difference and the wide geographical separation of the two forms it would seem preferable to adopt a distinctive name. The lamina is divided, almost to the base, into 6—10 segments; the larger are cuneate and the smaller obovate and obtuse. SAPORTAEA. Fontaine and White. Fontaine and White? instituted this generic term for some in- complete impressions of large leaves from Permian rocks in Virginia having a broadly cuneate or suborbicular lamina characterised by 1 Feistmantel (80) B. p. 121, Pl. xnvi. A. 2 Zeiller (96) A. p. 467. 5 Feistmantel (81) p. 257, Pl. 1. fig. 1; (86) Pl. m1. A. figs. 1, 2. + Fontaine and White (80) B. p. 99, Pl. xxxvim. XL] DICRANOPHYLLUM 93 a thickened lower margin extending horizontally a short distance on either side of the petiole and presenting the appearance of beimg formed by the bifurcation of the summit of the leaf-stalk at right-angles to its long axis. The lamina is irregularly dissected, but from the published figures it is difficult to distinguish between original lobing and divisions due to tearing. The dichotomously branched veins spread through the lamina from the centre of the base and are given off at a wide angle from the thick lower edge of the lamina. In Saportaea grandifolia the petiole has a length of 10 cm. and the rest of the incomplete leaf is 8 cm. long and 9-5 cm. broad: the second species S. salisburioides is represented by por- tions of similar but smaller leaves with a slender petiole. While comparing these fossils with Ginkgo the authors of the genus call attention to the peculiar features of the lower edge of the lamina and of the venation. The general resemblance in leaf-form between Saportaea and Ginkgo is hardly sufficient to warrant any definite statement as to relationship and this Permian genus must for the present be relegated to the class of Plantae incertae sedis. DICRANOPHYLLUM. Grand’Hury. This genus was first described by Grand’Eury! who, before the publication of the full description of the type-species, suggested the substitution of Hotaxites for Dicranophyllum? the name finally adopted’. The genusis fairly abundant in the Upper Carboniferous rocks of France and occurs also in Portugal, Belgium, and Germany ; it has recently been recorded from England and is represented in the Coal Measures of the United States and Canada. It occurs in Permian strata in Germany but with a few exceptions the genus _is characteristic of Stephanian beds. The systematic position of Dicranophyllum is far from settled ; by many authors it is considered to be a member of the Ginkgoales and is compared also with the Taxeae. In all probability the genus is allied to the Cordaitales, though, as stated in the case of Trichopitys, it cannot be assigned to a definite position in the Gymnosperms until we possess fuller information with regard to the reproductive organs or the anatomical structure. 1 Compt. Rend. Vol. yxxx. p. 1021, 1875. 2 dikpavos, two-pointed. 3 Grand’Eury (77) A. p. 272, Pls. x1v., Xxx. 94 GENERA OF UNCERTAIN POSITION [cH. In habit Dicranophyllum resembles Lepidodendron; it is an arborescent plant sparsely and irregularly though sometimes dichotomously branched; the leaves are crowded and spirally disposed, in some species persistent—in the sense in which the leaves of Araucaria are persistent—while in others they probably fell at an earlier stage. The leaves (fig. 671) exhibit a wide range in size, in the amount of lobing and the angle of divergence of the segments; there is no differentiation into a lamina and petiole nor are there any short foliage-shoots as in Ginkgo; the whole leaf is represented by a narrow lamina, in some species almost spinous, which consists in the basal portion of a simple linear ‘stalk’ reaching in extreme cases a breadth of about 7mm., attached by a decurrent base which persists as an elongated cushion closely resembling the leaf-base on some Lycopodiums or the projecting cushions of Picea (cf. fig. 140, Vol. 1. p. 94). The cushions are contiguous and cover the surface of a branch as in Lepidodendron, but they are distinguished by the occurrence of the leaf-scar at the apex of the cushion in contrast to its sub- apical position in Lepidedendron. The typical form of the leaf- base is shown in fig. 671, A, but in Dicranophyllum Beneckianum Sterz. the transversely elongated leaf-scars are almost contiguous as in some species of Szgillaria. At a distance from the base varying in different species the lamina is divided into two, generally equal, branches that diverge at an acute or wide angle, and in most species each arm undergoes one or more bifurcations in a single plane. The whole leaf may reach a length of over 20 cm. In the basal portion of the lamina there are two or more parallel veins, but in branches in which the leaf-scars are well preserved there is only a single vascular-bundle scar indicating a single leaf-trace up to the base of the lamina. Each segment of the leaf has usually two veins and the acutely pointed ultimate segments have a median vein. The so-called secondary or inter- stitial veins are no doubt due to the presence of hypodermal stereome strands. The narrower Dicranophyllum leaves are very similar to the deeply divided pinnae of Macrozamia heteromera (fig. 671; cf. fig. 396, F, Vol. 11. p. 26). The branching of the lamina is generally regular but in several instances the subdivision is_ irregular (fig. 671, D). On young shoots the leaves may be XL] DICRANOPHYLLUM 95 nearly vertical but in most species they become widely extended and on older branches may be reflexed as in some Lycopods (fig. 121, B, Vol. 1. p. 35). There is some evidence that the pith was discoid as in Cordaites!. The microsporophylls are said to be borne in small ovoid strobili in the axils of foliage leaves, but Fic. 671. Dicranophyllum gallicum. A, piece of a stem showing leaf- cushions. B—D, foliage-shoot and leaves. (After Grand’ Eury.) the only evidence as to their structure so far adduced is furnished by an imperfectly preserved specimen described by Zeiller? associated with a shoot of Dicranophyllum robustum but not actually attached; this consists of a small axis expanded into 1 Renault and Zeiller (88) A. Pl. uxxt. figs. 3, 4. 2 Zeiller (78) Pl. x. figs. 1 @ and 3. 96 GENERA OF UNCERTAIN POSITION [cH. a radially segmented distal portion bearing some imperfectly preserved écaid bodies on its lower face which are probably microsporangia. No spores are recorded. Zeiller compares this sporangiophore with the microsporophyll of a Taxus. On some stems small scale-covered buds occur immediately above the attachment of a leaf; these are probably fertile shoots but we have no definite information with regard to their structure. Some specimens from Commentry! demonstrate the occurrence of small oval ovules or seeds, 4 x 3mm., along the length of ordinary leaves (fig. 672), and seeds are sometimes found associated with the basal portions of foliage leaves though not in organic connexion with them, except in an example described by Renault from Autun as Dicranophyllum gallicum var. Parchemineyi?. Some leaves of D. striatum’ are described ‘as enlarged at the base and slightly concave as if to hold a seed, but if this supposition is correct it involves the admission of two types of seed-bearing organs within the genus. The more probable conclusion is that the seeds were borne along the length of the lamina of the sporo- phylls and on the expanded bases. Some specimens from Lower Cretaceous beds in Virginia described as Baiera foliosa* resemble Dicranophyllum, but in view of the vast chronological gap between these beds and those in which Dicranophyllum occurs it is unlikely that the similarity has any significance. Dicranophyllum gallicum Grand’ Hury. This species, one of the two described by Grand’ Kury in 18775, is the commonest representative of the genus; it is characterised by persistent leaves with a base that is unbranched for a distance of 15—20 mm. and then bifurcates into two equal or approximately equal segments at an angle of about 30°; these reach a length of 10—15 mm. and divide into two acute segments 8---10 mm. long. There are three veins in the basal portion of the lamina, one of which branches below the dichotomy, and each of the divergent arms has two veins. The leaf-cushions are 2—3 times as long as 1 Renault and Zeiller (88) A. Pl. uxxz. fig. 5. * Renault (96) A. p. 375. 3 Renault and Zeiller (88) A. p. 632. 4 Fontaine (89) B. p. 213, Pl. xcrv. fig. 3; Berry (11) Pl. nix. 5 Grand’ Eury (77) A. p. 272, Pls. xtv., xxx. Zeiller (80) A. Pl. txxvu. figs. 1, 2. XLU] ; DICRANOPHYLLUM 97 broad and the median vein of the lamina is continued as a keel in the middle of the persistent base. The sporophylls have the form of foliage leaves and bear numerous ovules (fig. 672). This species is recorded from the coal-fields of the Loire, Com- mentry, Gard, Brive! and elsewhere; it occurs also in the Coal Measures of Portugal?. The specimens described by Grand’- Eury from Gard as D. tripartitum®, which T had an opportunity of examining in the Ecole des Mines, Paris, are not specific- ally distinguishable from D. gallicum.— cae A large decorticated stem of D. gallicum *™* bsg 2, Dereon haus gate : é icum. (After Zeiller from in the Paris Collection recalls a decor- Renault; } nat. size.) ticated stem of Lepidodendron. Some : imperfect specimens described by White* from the Coal Measures of Missouri as Dicranophyllum sp. are compared by him with D. gallicum; one of them consists of a forked foliage-shoot with short and repeatedly bifurcate leaves illustrating the superficial resemblance between Dicranophyllum and Lepidodendron. An- other specimen shows an irregularly branched leaf which might equally well be referred to Trichopitys. Dicranophyllum lusitanicum (Heer). This species was first figured by Gomes® as Cyperites ? sp. on the ground of the similarity of the lamina to some fragments, probably of Lepidodendroid leaves, described by Lindley and Hutton® as Cyperites bicarinata and subsequently included by Heer’ in his genus Distrigophyllum. In a note to his account of Mesozoic plants from Portugal Heer® renames the plant Dzstrigo- phyllum lusitanicum and compares it with Dicranophyllum gallicum Grand’Eury. -De Lima® recognised the true nature of the speci- mens from the Stephanian of Portugal and published a full 1 Zeiller (92?) A. p. 96. 2 Lima, de (88) Pls. 1, mm. 3 Grand’ Eury (90) A. p. 335, Pl. vi. figs. 12, 13. 4 White (99) B. p. 272, Pl. xt. fig. 10; Pl. yxxmr. fig. 1. 5 Gomes (65) p. 32, Pls, 1., v. 6 Lindley and Hutton (32) A. Pl. xxii. 7 Heer (76) A. p. 39. 8 Ibid. (81) p. 11, footnote 1. » Lima, de (88). 8. IV 7 98 GENERA OF UNCERTAIN POSITION [cH. description of the species. The leaves are 14—16-5 cm. long and 2—4 mm. broad at the base; the lamina is once forked and the forks diverge at a very small angle as in D. longifolium Ren. Exceptionally good specimens figured by Gomes and de Lima show numerous long leaves spreading radially in the matrix from a comparatively slender axis. In D. longifolium the leaves are given off at a much more acute angle. Dicranophyllum robustum Zeiller. This type? is similar to D. gallicum but the leaves are only preserved in their basal portions; the lamina is 5—6 mm. broad and bifurcates at a distance of about 15 mm. from the base at an angle of 20°—30°. Ovoid buds occur in the axils of some of the leaves. It was in association with this specimen from the Gard coal-field that Zeiller found the microsporophylls already described. The surface shows particularly well-preserved large and slightly depressed cushions 3—4 cm. long and 4—5 mm. broad. Dicranophyllum Beneckianum Sterzel?. In the form of the leaves this Permian species from Baden closely resembles D. gallicum; it is chiefly of interest because of the almost complete absence of leaf-cushion; the leaf-scars, characterised by their acute lateral angles, are almost contiguous as in some species of Szgillaria. Dicranophyllum Richiri Renier’. In this Belgian Westphalian species the leaves are dichotomously branched into two linear segments at an angle of about 60°; it differs from D. gallicum in the single bifurcation of the lamina, the wider angle of divergence, and in the feebler relief of the leaf-cushions. Dicranophyllum anglicum Kidston. This, the only British species, has recently been described from the Westphalian beds of Staffordshire’. The crowded leaves, 3-50 em. long, are dichotomously branched three or four times into slightly spreading linear rigid segments with a maximum breadth 1 Zeiller (78). 2 Sterzel (07) p. 381, Pls. x1v., Xv. 3 Renier (07) p. 186, Pl. xvur. figs. 3—7; (10?) Pl. cxvu. 1 Kidston (14) p. 170, Pl. xtv. figs. 3, 3a. XL] DICRANOPHYLLUM 99 of 125mm, The undivided portion of the lamina is about 7 mm. long. Kidston speaks of the rhomboidal outline of the leaf and the repeated dichotomy of the lamina as distinguishing features. Some fragments of forked leaves are figured by Schenk! from the Coal Measures of China as Dicranophyllum latum, but the material is too meagre for accurate determination. It is note- worthy that the broader type of Dicranophyllum leaf may easily be confused with an impression of a branched Stigmarian rootlet. The narrower specimens described by Schenk from China as D. angustifolium? are also too fragmentary to be accepted as trustworthy evidence of the occurrence of the genus in the southern flora. Dicranophyllum striatum Grand’Eury. This species like several others is founded on detached leaves?, a circumstance that has led some authors to draw a distinction between species with persistent leaves and those with caducous leaves. There is, however, no good reason for assuming that all species were not of the evergreen type. The leaves of this species are characterised by their great length which may be 24 cm.; the lamina is once or twice forked and is 5—6 mm. broad at the -base, which contains 4—7 veins. Dicranophyllum longifolium Renault. In this Commentry species* the leaves, which reach 14 cm. in length, are characterised by the very small angle of the divergence of the segments, 3° as contrasted with a divergence of 30° in D. gallicum. The leaves are almost erect and twice bifurcate. In addition to Dicranophyllum Beneckianum Sterzel has de- scribed a second species, D. latifolium®, from the Lower Permian of Baden characterised by leaves similar to those of D. striatum but generally longer. The species is founded on leaves and is not a well-defined type. : Two species are recorded by Lesquereux® from the Coal Measures 1 Schenk (83) A. p. 222, Pl. xm. figs. 11, 12. 2 Ibid. Pl. xtra. figs. 17, 18. 3 Grand’Eury (77) A. p. 275. Renault and Zeiller (88) A. Pl. Lxx1. fig. 2. 1 Renault and Zeiller (88) A. p. 631, Pl. Lxxt. fig. 1. 5 Sterzel (07) p. 391, Pl. xv. figs. 9—11. 6 Lesquereux (80) A. pp. 553, 554, Pls. LXxXXIM., LXXXVLL. 4a L 100 GENERA OF UNCERTAIN POSITION [CcH. of Pennsylvania, but neither is represented by very satisfactory specimens: Dicranophyllum dichotomum Lesq. is founded on a dichotomously branched shoot bearing long and narrow leaves in the apical region only and very similar in appearance to Lepido- dendron except in the branched lamina. The second species D. dimorphum Lesq. is represented by leaves and branches, which however are not well preserved. The peculiar subdivision of the apical portion of the laminae suggests a simple leaf wjth a frayed termination. Dicranophyllum glabrum (Dawson). Under this name Dr Stopes! has recently described a well- preserved leaf from the Westphalian series of New Brunswick. The specific name was first applied by Dawson? to specimens which he referred doubtfully to Pstlophyton. The type-specimen is 9 cm. long and 3mm. broad at the base and the lamina is repeatedly branched. This specimen bears a close resemblance to the leaf from Autun described by Renault as Trichopitys Milleryensis®. The imperfect specimens described by Dawson* from Devonian rocks in Queensland as Dicranophyllum australicum and sub- sequently figured by Jack and Etheridge® consist of a slender axis, 3mm. wide, with elongate leaf-bases bearing leaves 3 mm. long with two widely divergent apical segments like those charac- teristic of the sporophylls of Gomphostrobus. The fragments have no claim to be included in Dicranophyllum. There has been confusion between Dicranophyllum and Gom- phostrobus’: as shown by drawings reproduced by Potonié’ of specimens of Gomphostrobus from the Permian of Thuringia, there is a close resemblance in habit between the two genera, but in Gomphostrobus the foliage-leaves are faleate and entire, while the bifurcate sporophylls differ from the leaves of Dicranophyllum in their widely divergent and small apical fork. , 1 Stopes (14) p. 79, Pl. xv. fig. 47. 2 Dawson (62) p. 315: for other references, ‘see Stopes loc. cit. 3 Renault (96) A. p. 378; (93) A. Pl. xxx. fig. 2. 4 Dawson (81) A. p. 306, Pl. xi. figs. 15, 16. 5 Jack and Etheridge (92) B. p. 49. 6 Schenk (90) A. erroneously includes Sigillariostrobus bifidus Geinitz, (73) Pl. m1. figs. 5—7, in Dicranophyllum. See also Sterzel (93) A. p. 111. 7 Potonié (93) A. Pl. xxvim. figs. 1, 2. XLit] TRICHOPITYS 101 A foliage-shoot described by Renault! from Autun as Pinites permiensis, though too imperfect to be identified, is worthy of notice as possibly an example of Dicranophyllum or Trichopitys ; it consists of an axis 3 mm. in diameter bearing numerous spirally disposed leaves 3 cm. long, barely 1 mm. broad and triangular in section, at an angle of 45°. The leaf-cushions are elongate and slightly prominent. It is, however, impossible to decide whether this fossil should be referred to the Lycopodiales or to the Gymno- sperms. There is no evidence that the leaves are attached to short shoots and the use of the generic name Pinites cannot be justified by any trustworthy test. TRICHOPITYS. Saporta. Saporta? proposed the generic name Trichopitys in 1875 for some shoots from the Permian beds of Lodéve bearing long, narrow, and deeply divided leaves; he defined the genus as follows: “Folia verosimiliter rigida cartilagineaque, dichotome partita etiamque pedato-partita, petiolo plus minusve elongato, sursum in lacinias 4—6, anguste lineares, uninerviasque dissecta®.’ Many palaeobotanists have followed Saporta in regarding Trichopitys as a member of the Ginkgoales, but the evidence in support of this view is by no means conclusive. The only species so far described that affords any information with regard to the habit or fertile shoots of the plant is the type-species 7. heteromorpha (fig. 673). A fairly stout branched axis bears leaves varying considerably in size and form; they may be long and filiform, apparently rigid, simple or deeply divided, or short and entire, and in some cases resembling the leaves of certain smaller species of Baiera except in the less regular forking of the lamina. In the axil of some foliage leaves are short, simple or branched, axes bearing seed-like bodies originally described as buds and afterwards regarded as seeds. A specimen figured by Zeiller* from Lodéve (fig. 673) shows a branched axillary shoot bearing several small ovules comparable with an abnormal ovuliferous shoot of Ginkgo (cf. fig. 631, D). 1 Renault (96) A. p. 377; (93) A. Pl. pxxxm. fig. 1. 2 Saporta (75) p. 1020. 3 Ibid. (84) p. 263, Pl. cium. fig. 1. 4 Zeiller (002) B. p. 254, fig. 182. 102 GENERA OF UNCERTAIN POSITION [CH. Renault has figured a portion of a large leaf from Autun as Trichopitys Milleryensis which may belong to the closely allied genus Dicranophyllum; it is 12 cm. long and 3 mm. broad at the base; the narrow basal part: of the lamina has three parallel veins and forks into two arms, each of which again branches into divergent linear segments. The leaf is larger and broader than the leaves of T. heteromorpha and agrees very closely with those of some species referred to Dicranophyllum: the fact that the branch- ing of the lamina is not absolutely regular cannot be accepted as a constant difference between Dicranophyllum and Trichopitys : the leaves shown in fig. 671, which were found attached to un- doubted Dicranophyllum branches, are no more regular in the I vi Hy Fic. 673. Trichopitys heteromorpha. (After Zeiller; 2 nat. size.) forking of the segments than T. Milleryensis, and a leaf recently figured by Dr Stopes? from New Brunswick as Dicranophyllum glabrum (Daws.) bears a close resemblance to Renault’s figure. In some cases the more regular dichotomy of the leaves is a charac- teristic of Dicranophyllum, but it is not a constant feature. Some imperfectly preserved specimens figured by White® from the Coal Measures of Missouri as Dicranophyllum sp. bear irregularly branched leaves which are hardly distinguishable from some of those on Saporta’s type-specimen of Trichopitys heteromorpha. 1 Renault (93) A. Pl. yxxxur. fig. 2; (96) A. p. 378. 2 Stopes (14) p. 79, Pl. xvut. fig. 47. % White (99) B. p. 272, Pl. xx. fig. 10; Pl. uxxut. fig. 1. XLIt] SEWARDIA 103 In the present state of our knowledge it is impossible to give a satisfactory definition of the genus or to state precisely on what grounds it is separated from Dicranophyllum. In Trichopitys, as represented by J’. heteromorpha, the leaves are more variable in form than in Dicranophyllum and less regular in the subdivision of the lamina; there are no persistent leaf-bases like those of Dicranophyllum, but this is a character that could not be seen in imperfectly preserved or partially decorticated specimens. A more important difference would seem to be that in Trichopitys the seeds are borne on special axillary shoots, while in Dicranophyllum they occur on ordinary leaves. Such evidence as we have suggests that Trichopitys is a Gymnosperm possibly allied to the Cordaitales and Ginkgoales, but the facts hardly justify its inclusion in either group. Its affinity to Dicranophyllum cannot be definitely deter- mined though in all probability the two genera are closely related if not indeed generically identical. Saporta included in Trichopitys two Jurassic species, T’. lacini- ata, originally referred to the genus J. eanpaula, and T. Lindleyana? ; in the latter species he included the specimens doubtfully assigned by Lindley and Hutton to Solenites as Solenites ? furcata®. These and other Jurassic leaves that are referred by some authors to Trichopitys are usually regarded as examples of Baiera®; there are no adequate grounds for believing them to be closely related to the Permian Trichopitys. Zeiller* records a fossil from Triassic beds in Madagascar that he thinks may be an example of Trichopitys. SEWARDIA. Zeiller. This generic name was proposed by Zeiller in place of Withamia which, in ignorance of its previous use, I employed for some specimens from the Wealden rocks of Sussex. The inclusion of a second species, Sewardia longifolia, necessitates an extension of the definition of the genus to include spinous branches bearing spirally disposed leaves or leaf-like organs, either orbicular and entire or fan-shaped and deeply divided, in the axil of recurved spinous processes. . 1 Saporta (84) Pl. crv. figs. 1—9. 2 Lindley and Hutton (37) A. Pl. 209. 3 Seward (00) B. p. 266. 4 Zeiller (11?) p. 234. 104 GENERA OF UNCERTAIN POSITION (cH. Sewardia latifolia (Saporta). 1849. Otozamites latifolia Brongniart, Tableau, p. 106. 1872. Sphenozamiies latifolius Schimper’, Traité, Vol. 1. p. 163. 1875. Cycadorachis armata Saporta, Plant. Jurass. p. 196, Pl. 117, fig. 1. 1895. Withamia armata Seward, Wealden Flora, Vol. m1. p. 174, Pl. 1. figs. 1,2; Pl. v. fig. 1. 1900. Sewardia latifolia Zeiller, Eléments Paléobot. p. 233. This species is represented by woody axes, about lcm. in breadth reaching a length of 50 cm., from the Wealden beds of Sussex, bearing more or less orbicular, entire, leaves or leaf-like A. Fia. 674. Sewardia latifolia. Axis (A) and single leaf (B). (British Museum; } nat. size.) organs, 6cm. or more long, in the axil of stout recurved spinous processes (fig. 674, A, B). The leaves are sessile and the venation is of the Cyclopterts type. Spinous axes of the same form had previously been described by Saporta from Kimeridgian rocks in France as Cycadorachis armata, but these show no indication of leaves and were regarded as Cycadean. In a letter written to me in 1895, shortly before his death, the Marquis of Saporta? suggested 1 See also Saporta (75) A. p. 188, Pls. 112. 113. 2 Seward (95) A. p. 175. XLT] SEWARDIA 105 the generic name Acanthozamites as a substitute for Cycadorachis in view of the new data afforded by the English material, but it seemed preferable to adopt some provisional name which did not imply affinity to the Cycads. The leaflets described by Saporta as Sphenozamites latifolius are apparently identical with those found in the English beds, but none of the French specimens were attached to a supporting axis. The relation of spines and ‘leaves’ suggests that the latter may be‘phylloclades borne in the axil of modified spinous leaves, but their morphological nature cannot be determined. In this connexion attention may be called to Dioncophyllum Tholloni Baill. a West African shrub which bears on the long shoots leaves 2—3} inches long each of which has a pair of strong revolute hooks at the apex: in the axils of these leaves are short shoots with larger leaves without hooks. It is suggested! that the apparent lamina of the hooked leaves is a winged petiole, the hooks representing lateral leaflets. Sewardia longifolia (Salfeld). This species was described by Salfeld? from the Solenhofen beds (Upper Jurassic) of Bavaria as Baiera ? longifolia Heer: it is founded on branches nearly 30cm. long bearing large fan- shaped deeply divided leaves, or leaf-like organs, in the axils of recurved spines similar to those in S. latifolia. The ‘leaves’ are identical in habit with those of some species of Baiera, but we have no information with regard to the structure of the epidermis. In view of the uncertainty as to the morphological nature of the leaves or their relationship to leaves of Baiera, it is inadvisable to adopt a generic title that implies affinity to the Ginkgoales. 1 Sprague (16). 2 Salfeld (07) B. p. 195, Pls. xx., xx1. Fic. 674*. Sequoia sempervirens. Near Crescent City, California. (From a photograph by Professor A. Henry.) CHAPTER XLITI. CONIFERALES (RECENT). Tue Coniferales, by far the largest section of the Gymnosperms, present considerable difficulty to the student of fossil plants. There is great divergence of opinion with regard to the relative antiquity of the several families, and tlieir position in an evolu- tionary series. The Abietineae are by some botanists regarded as the most primitive; on the other hand, and this is the view that in my opinion receives most support from the available evidence, it is held that the Araucarineae are both the most primitive and the oldest representatives of the Coniferales. Until recent years the study of fossil Conifers has suffered neglect and little help has been afforded by palaeobotanists to the solution of the morphology of the ovulate shoots of the different genera, a CH. XLIIT] MORPHOLOGY 107 problem that has long exercised the ingenuity of investigators. The view expressed by Jeffrey! that recent work on fossil Conifers corroborates the interpretation of the seed-bearing scales as metamorphosed shoots is based on facts furnished by a study of vegetative organs, which in themselves do not afford any decisive Fic. 675. A, B, Phyllocladus trichomanoides. C, P. hypophyllu. D, EK, Megastro- bilus and seed of Phyllocladus alpina; a, arillus. [A—C from specimens in the British Museum; D, E, after Miss Robertson (Mrs Arber.)] evidence as to the morphology of reproductive shoots. In view of these considerations it is important that an attempt should be made, even at the risk of disproportionate treatment, to give a general account of recent genera which, though necessarily far 1 Jeffrey (10) p. 331. 108 CONIFERALES (RECENT) [cH. from complete, may afford assistance to students prepared to undertake a critical study of the fragmentary records of the rocks. Conifers are trees or shrubs exhibiting a fairly wide range in habit; the ‘great ones of the forest’ such as the Sequoias (fig. 674*), the sugar Pines (Pinus Lambertiana) and Douglas Firs (Pseudotsuga Douglasii) of the Rocky Mountains, Taxodium mucronatum! of Mexico, remarkable for its enormous bulk, the tall and slender Cypresses, the less formal Podocarps of the southern hemisphere, the shrubby Junipers, the dwarf Dacrydium laxifolium? of New Zealand afford examples of recent types. In most species the leaves are small and crowded, not infrequently dimorphic, and in Phyllocladus (fig. 675) reduced to inconspicuous and caducous scales subtending phylloclades. Agathis is exceptional in having narrow ovate leaves reaching a length of nearly 20 cm. (fig. 695) and a similar but smaller Jeaf is characteristic of some species of Podocarpus (fig. 676). The presence of long and short shoots is a striking feature of Pinus, Larix, Pseudolariz, Cedrus, and Sciadopitys: the short shoot, as Goebel says ‘takes no part in the construction of the permanent skeleton of the tree®.’ The whorled arrangement of leaves characteristic of several Cupres- sineae and the Callitrineae is not a constant feature and, as in Lycopodium, both whocled and spiral foliage may occur on the same shoot. Conifers are monoecious or dioecious; the microsporophylls and megasporophylls are borne spirally or in whorls on separate shoots, . and in some genera on separate trees, except in the case of abnormal bisporangiate strobilit. Proliferous cones are not uncommon in some genera: the prolonged axis of the cone of Cryptomeria japonica shown in fig. 677 bears microstrobili in the axils of the small leaves. The microstrobili are for the most part constructed on a uniform plan; they are usually short-lived, small shoots, and each microsporophyll often consists of a slender axis bearing two microsporangia on its lower surface and prolonged as a small upturned distal expansion. In Pinus the sporangia dehisce longitudinally, while Abies (fig. 684, E) affords an example of 1 Gard. Chron. Nov. 26, 1892, p. 648. 2 Hooker, J. D. (52) Pl. 815. *3 Goebel (05) p. 444. * For examples see Sterzel (76); Eichler (82); Worsdell (04); Bartlett (13); Shaw, W. R. (96); Robertson (06); Renner (04); Bayer (08); etc. XLII} FERTILE SHOOTS : 109 transverse dehiscence. The microstrobili of Cedrus are similar but longer. In Torreya (fig. 684, D), Taxodium, Widdringtonia, Fic. 676. Podocarpus latifolia. (% nat. size.) M. 8. Fic. 677. Cryptomeria japonica. Proliferous cone. (Nat. size.) and some other genera there are 4—6 microsporangia on each sporophyll. In Araucaria and Agathis there may be as many as 110 CONIFERALES (RECENT) [cH. 10—20 sporangia, longer and relatively narrower than in other genera and attached by one end, in contrast to the more complete union of sporophyll and sporangium in Pinus. In Araucaria Muellert and A. Rulet (fig. 678) the microstrobili reach a length of 25 cm.: in A. excelsa and A. Cookw they are much smaller (fig. 679, A, B). In Taxus 4—7 spor- angia are radially disposed on the inner face of a flat distal expansion. The microstrobili of Cunninghamial, Pseudolariz?, and Keteleeria are borne in umbels, while in Cryptomeria and Taxodium? they occur in spikes. The microspores may be winged or wing- less: in the Abietineae there are as a rule two conspicuous wings or bladders (fig. 684, B), but the spores of Pseudo- tsuga are wingless and in Tsuga both types occur. In Mrerocachrys* the wings vary from 2 to 6 (fig. 684, C) and in Dacrydium® and Podocarpus* we. 678, Microstrobilus of (fig. 684, A) there are 2 or 3 small Araucaria excelsa (A) and Araucaria Rulei (B). (After bladders. In Taxus, Cephalotaxus, Ea Pee ey anes Torreya, Sciadopitys, the Cupressineae, size; B, } nat. size.) and some other Conifers there are no prothallus cells: the microspores of the Abietineae are character- ised by the occurrence of 2, or occasionally 3 or 4’, evanescent prothallus cells (fig. 684, B) ; in Dacrydium there are 4—-6 prothallus cells; in Microcachrys 3 or 4; in Podocarpus (fig. 684, A) as many as 8, while in Araucaria 15 cells have been recorded and as many as 30 nuclei. The two male gametes are non-motile. The term Conifer though appropriate as regards the majority of the plants so styled is misleading in the case of several genera 1 Siebold (70) Pl. cr. 2 Bot. Mag. Jan. 1908. 3 Gard. Chron. Nov. 25, 1898, p. 659. 4 Thomson (09). 5 Young (07). 6 Jeffrey and Chrysler (07); see also Thibout (96); Burlingame (08); (13); (15); Sinnott (13). 7 Hutchinson (14). XLII} : FERTILE SHOOTS lll which possess ovulate shoots differing widely from cones as the term is generally understood. The cones of Araucaria (figs. 680, 681) and Agathis reach a considerable size; those of Araucaria Bidwillu}, similar to some cones of Encephalartos, may be 28 cm. in diameter and in some species of Agathis® they exceed 11 cm. microstrobilus. D, E, Araucaria Mueller’, part of a microstrobilus and a single sporophyll; ps, microsporangia. F, Araucaria Montana, branch. (After Seward and Ford.) : and are 14cm. long. The cone-scales of Agathis are flat, woody structures bearing»a single ovule (fig. 682): in Araucaria the single seed is embedded in the scale, and a more or less prominent appendage, the ligule, forms a characteristic feature (fig. 683, 1). 1 Gard: Chron. April 14, 1894, p. 465. 2 Seward and Ford (06) B. 112 CONIFERALES (RECENT) (cn. Fic. 680. Araucaria brasiliensis, cone. (é nat. size; from a specimen in the Royal Gardens, Kew.) Fic. 681. Araucaria Cunninghamii, cone. (After Seward and Ford; $¢ nat. size.) XLUI] MORPHOLOGY OF CONES 113 The cone-scales of some species, e.g. A. Cookii, A. excelsa (fig. 683, A, D) are flat and laterally winged, while in A. brasiliensis the thick distal ends closely resemble those of the seed-scales of some Pines: the cone-scale of A. imbricata is larger and deeper, and that of A. Bidwillii broad and woody (fig. 683, B, ©). In Pinus, with cones reaching a length of 2 feet, the mature scales. are apparently simple like those of the Araucarineae: the distal end is broad and rounded (P. silvestris) with a central umbo or, Fic. 682. A, Agathis Mooret, cone-scales; s, scar of seed. B, Agathis loranthifolia, section of ovule showing integument, nucellus, and megaspore with three archegonia, also part of the cone-scale with a projection close to the base of the ovule. (After Seward and Ford.) as in P. Coultert, the umbo is prolonged as a strong recurved spine, while in P. excelsa (fig. 704) and P. Cembra the scales are flatter like those of Picea. In the young Pine cone each scale is clearly a double structure consisting of a lower portion, the bract or carpellary scale, and an upper portion, the ovuliferous scale, bearing two ovules. In the course of development the seminiferous scale alone increases in size, and the bract-scale is hardly visible in the ripe cone or is represented by a small remnant. In Ales, Lariz, Pseudotsuga the dual nature of the scales is obvious at 8. IV 8 114 CONIFERALES (RECENT) [cH. maturity, the bract-scale usually extending beyond the edge of the seminiferous scale (fig. 705), in Abies bracteata reaching a length of 5cm. In Sequoia (fig. 702 B) the cone-scales show no outward sign of a double structure, but each scale contains two Fic. 683. Cone-scales of Araucaria Cookii (A), A. imbricata (B), A. Bidwillit (C), and A. excelsa (D, E); 1, ligule. (After Seward and Ford.) sets of bundles, the lower being normally orientated as in a leaf and the upper vascular strands inversely orientated (cf. fig. 684, R). The occurrence of these two sets of bundles is often quoted in support of the view that the double cone-scale of the Abietineae XLOI] MORPHOLOGY OF CONES 115 represents the lowest term of a series, the upper end of which is represented by the scales of Agathis which have lost all external signs of their supposed dual nature and retain only the inversely orientated bundles as evidence of their descent from an ancestral type in which the ovuliferous and bract-scales were separate organs!. The scales of such genera as Sciadopitys, Athrotawis, Cryptomeria (fig. 684, 8S, N, M), on this hypothesis, occupy an intermediate position. The seminiferous scale of the Abietineae is considered by many botanists to be a leaf or leaf-like organ borne on an axillary shoot subtended by a bract, and it is believed that the simple scale of Agathis has been produced by the gradual fusion of two originally distinct organs. The ligule of Araucaria is held to be the outward and visible sign of the semi- niferous scale that has almost lost its individuality, and with this ligular relic are homologised the upper half of the scale of Sequoia, the deeply toothed upper part of the scale of Cryptomeria (fig. 684, M), the rounded ridge on the abaxial side of the seeds in Athrotazis (fig. 684, N), the membranous outgrowth on the scales of Cunninghamia (fig. 684, K, m), and the seminiferous scale of the Abietineae. It has been pointed out in support of this hypothesis that two vascular bundles are given off from the axis of a Pine cone, one of which forms the bract-scale bundles and the other the vascular supply of the seminiferous scale?. In a recently published paper on the vascular anatomy of the megasporophylls of Conifers by Miss Aase* additional facts are given with regard to the origin and behaviour of the vascular bundles of the cone-scales. In the upper part of a cone of Pinus maritima the bract-supply arises as a single bundle at the base of a gap in the stele, and the bundles of the seminiferous scale are given off from the sides of the gap above the point of origin of the bract-bundle: in the lowest sporophylls, on the other hand, the bract and scale-bundles have a common origin. Stiles (08); Norén (08); Tison (09); Bot. Mag. Tab. 8664 (1916). 3 Robertson (04); (07). 4 Oliver (02); (03). XLU1] SEEDS 121 the nucellus.of Cycadean seeds. The portion of the Torreya seed (fig. 684, V) below the free part of the nucellus has, according to Oliver, been produced by the intercalation of a new basal region that has pushed up the chalaza. Cephalotaxust has plum-like seeds similar to those of Torreya. In Phyllocladus? (fig. 675, E) an ovule enclosed in a papery epimatium occurs in the axil of a succulent bract, and in Taxus a terminal ovule is borne on a short shoot. oe OO 1, ) 0 — Fic. 688. Torreya nucifera, transverse section of seed; a. arillus; v, vascular tissue; sc, outer zone of integument in which the hard shell will be formed ; i, inner part of integument; mz, wall of megaspore; n, nucellus; ps, pro- thallus. (After Oliver.) without any megasporophylls. In Podocarpus* the megasporo- phylls are few and a part or whole of the reproductive apparatus is fleshy: the strobilus of P. Totara (fig. 684, G) consists of 3—4 bracts two of which are usually fertile. Similarly the strobilus of P. imbricata is formed of 2—4 bracts with swollen bases forming the receptacle; in the section shown in fig. 684, I, one bract is 1 Gard. Chron. Oct. 14, 1895, p. 717. 2 Robertson (06). 8 Pilger (03); Brooks and Stiles (10); Stiles (12); Gibbs (12); Sinnott (13). 122 CONIFERALES (RECENT) [cH. fertile and its apex overtops the ovule, while a second bract is sterile. Fig. 684, H shows the strobilus of P. neritfolia in which the lowest bracts are leaf-like and the swollen bases of the upper bracts are fused with the axis and each other to form a receptacle analogous to the flower-axis of Anacardium. P. spicata has a long loose strobilus bearing several ovules (fig. 684, F); and in P. Nagi a single seed occurs on an axillary branch bearing small decussate scales: the flesh of the seed is formed from the epi- matium, and the sclerotesta from the integument. ’ The seeds of Conifers vary greatly in size and shape: those of some species of Pinus and several other Abietineae have a large wing while others are wingless. The absence of a wing in a fossil seed does not necessarily denote an original feature. The seeds of Cedrus, Abies, Picea, Pseudotsuga, Keteleeria and Tsuga are winged. Glick! calls attention to differences in the relation of seed to wing in certain Abietineae: in Picea the base of the wing covers the seed like a spoon; in Abies, Larix, and Cedrus the seed hes in a pocket formed by the enclosing wing, while in Pinus the wing embraces the seed like a pair of pincers. In Agathis (fig. 682, A) there are two very unequal wings. The much smaller seeds of many Conifers have 2—3 wings: in Cupressus they are more or less equal (fig. 684, Q), in Libocedrus and Fokienia markedly unequal; in Fitzroya and Cunninghamia there are 2—3 wings. Our knowledge of the relative vitality of Conifer seeds is meagre’, and from the point of view of the possibilities of dispersal further research is desirable. The viviparous habit of Rhizophora and other Mangrove Dicotyledons, believed by Guppy? to be a primitive feature, is recorded in Podocarpus Makoyt'. The relation between nucellus and integument is less uniform in Conifers than in Cycads. In some genera, e.g. Agathis (fig. 682, B), Dacrydium, Phyllocladus, Fitzroya, Callitris and a few others the nucellus is free from the integument to the base; in Podocarpus the relation is variable; in Pinus and other Abietineae, in Torreya (fig. 684, V) and some other genera the nucellar apex alone is free. The free summit often has the form of a steep cone: in Araucaria® and to a greater degree in Saxegothaea (fig. 685, st) 1 Gliick (02) p. 402. 2 Coker (09). 3 Guppy (06). + Lloyd (02). 5 Seward and Ford (06) B. XLUr] CLASSIFICATION 123 this protrudes through the micropyle: in Fitzroya patagonica the prolongation of the integument as a micropylar tubé with a stigma- like terminal expansion is particularly striking. There is no _ regular pollen-chamber as in Cycads and Ginkgo, but in Pseudo- tsuga* a two-storied chamber, analogous to the pollen-chamber, is formed by a knee-like bend in the integument. A peculiar type of pollination characterises Araucaria: the microspores germinate on the ligule or on the megasporophyll and their tubes grow over or into the scale-tissues on their way to the ovule?. The archegonia of Conifers usually occur at the apex of the prothallus and are few in number, they are separated by a few layers of cells (Abietineae) or form a compact group (Cupressineae). In some Podocarps there may be as many as 14, in Tazodium 34, in Agathis 60, - irregularly distributed on the sides of the prothallus. In Widdring- tonra® 100 archegonia are recorded occupying a lateral position ; in Actinostrobus* Saxton has discovered groups of laterally placed archegonia. In Sequoia® the archegonia are also numerous and not confined to the apex. It is an open question whether or not the greater number and irregular disposition of the archegonia are primitive features. The occasional occurrence of lateral archegonia in Pinus may be a revival of an older habit. Classification. The result of recent research into the morphology and life- histories of genera demand certain changes in the generally adopted grouping. The following classification is an attempt to give clearer expression to the inter-relationships of existing genera®. Arnoldi’ proposed to withdraw Sciadopitys from Sequoia, Taxodium, and other members of the Taxodineae as the type of a separate family; he also suggested the isolation of Sequoia. The more recent work of Coker’ and Lawson® favours the removal of Taxodium and Cryptomeria.to the Cupressineae. Miyake’s re- searches!” point to a similar affinity in the case of Cunninghamia. The genera Athrotaris, Fokienia, and Taiwania are placed tenta- tively in the Cupressineae. The family-name Callitrineae, first 1 Lawson (09). 2 Thomson (07); Eames (13); Burlingame (13); (15). 3 Saxton (10). 4 Tbid. (13). 5 Shaw, W. R. (96); Arnoldi (01). 6 See also Saxton (137). 7 Arnoldi (01); Lawson (10); Radais (94). 8 Coker (03). ® Lawson (04). 10 Miyake (10). 124 CONIFERALES (RECENT) [CH.. used by Masters, has been revived by Saxton! to give expression to the distinctive characters of Callitris, Widdringtonia, and Actinostrobus. Saxton’s work on Tetraclinis leads him to assign it to the Cupressineae. Pilger? makes Phyllocladus the sole genus of Phyllocladoideae, while Miss Robertson? includes it in the Podocarpeae though recognising leanings towards the Taxineae. It should be stated that the changes in classification suggested are based mainly on characters of the gametophyte though anatomical and vegetative features have not been entirely neglected +. I. ARAUCARINEAH. Agathis, Araucaria. II. CUPRESSINEAE. Cupressus, Chamaecyparis, Labo- -cedrus, Thuya, Juniperus, Fitzroya, Diselma, Thujopsis, Taxodium, Glyptostrobus, Cryptomeria, Cunninghamia, Taiwania, Fokienia, Athrotaxis, Tetraclinis. III. CALLITRINEAE. Callitris, Actinostrobus, Widdring- tonia. IV. SEQUOIINEAE. Sequoia. © V. SCIADOPITINEAE. Sciadopitys. VI. ABIETINEAE. Pinus, Cedrus, Larix, Pseudolariz, Picea, Tsuga, Abies, Pseudotsuga, Keteleerva. VII. PODOCARPINEAE. Podocarpus, Dacrydium, Micro- cachrys, Acmopyle, Pherosphaera, Saxegothaea. VII. PHYLLOCLADINEAE. Phyllocladus. IX. TAXINEAE. Tazus, Torreya, Cephalotaxus. The order of the families is not intended to indicate their natural sequence in an evolutionary series, though the Arau- carineae are considered to be the most primitive. As certain authors have suggested, Saxegothaea is probably closely allied to Araucaria, but this is not indicated in the order adopted. Geographical Distribution. The distribution of the Conifers, though too wide a subject for more than a brief notice, is of great interest from a palaeonto- logical point of view. The ABIETINEAE, comparable in their present dominant réle with the Polypodiaceae among the Ferns, 1 Saxton (107); (13°). 2 Pilger (03). 3 Robertson (06). 4 For other views on classification. see Vierhapper (10). > Drude (90); Engler (89); Graner (94); Hildebrand (61); Vierhapper (10). XL] DISTRIBUTION 125 are the most widely spread; for the most part restricted to the northern hemisphere, they are not unrepresented south of the equator. Pinus reaches the tree-limit in the north and extends as far south as Formosa}, Siam, the Malay region, the Philippines’, S. Africa, and the West Indies. Picea has a similar distribution in the north and reaches to the temperate regions of the southern hemisphere. Abies ranges from Europe and Algeria to Siberia, the Himalayas, Japan, and Formosa. Larix flourishes in northern Europe and Siberia, Canada and the northern United States, the Himalayas, and Japan. Tsuga is more especially a North American Fic. 689. Araucaria imbricata on the Andes, Argentina. (From a photo- graph by Dr Wieland.) genus, but it occurs in the Himalayas and in Japan. Pseudotsuga is characteristic of N.W. America and is recorded from Formosa. Pseudolarix is a native of N.E. China and Formosa. Cedrus? occurs in Algeria, Morocco, Syria, Cyprus, and the western Hima- layas. The distribution of the ARAUCARINEAE affords a striking example of the contrast between the present and past range of a family. Araucaria occurs in Brazil, Chile and Argen- tina, in Australia, New Caledonia, New Guinea, the Pacific islands. Agathis is confined to the Australian and Malay region, New Zea- 1 Hayata (10). 2 Foxworthy (11). 3 Hooker, J. D. (62). 126 CONIFERALES (RECENT) [cH. land, New Caledonia, and the Queen Charlotte Islands. The two trees of Araucaria Bidwillii shown in the Frontispiece are survivors of a forest on the hills of Queensland. The photograph reproduced in fig. 689, for which I am indebted to Dr Wieland}, illustrates the habit of Araucaria imbricata on the eastern slopes of the Andes in South-West Argentina where the trunks reach a diameter of two metres. There are few existing trees comparable with these venerable types in the impression they produce of the lapse of ages and the vicissitudes of a dwindled race. CUPRESSINEAE. Cupressus occurs in North America in- cluding the Californian coast and Mexico, in S.E. Europe, temperate Asia, China, and Japan. Chamaecyparis extends to the Sitka Sound and flourishes in China, Japan, and Formosa. Lvzbocedrus, one of the few genera met with in both hemispheres, has a dis- continuous distribution; it occurs in California, Chile, Japan, Australia, New Zealand, New Guinea, and New Caledonia. Thuya flourishes over a wide area in North America and occurs in the Far East. Juniperus is characteristic of temperate regions in both the old and new world and is represented in the Canaries, the Azores, Somaliland, and Mexico. Fitzroya is confined to Patagonia and Chile; Dvrselma to Tasmania. Thujopsis is ex- clusively Japanese. Taxodium is a native of Texas and Mexico, while Glyptostrobus is a closely allied genus in China. The mono- typic Cryptomeria lives in China and Japan; Cunninghamia in China and Formosa. Tatwania and Fokienia have recently been described from Formosa and East China respectively. Athrotaxis is confined to Tasmania and Tetraclinis to North Africa. CALLITRINEAE. Callitris occurs in Australia and New Caledonia; Widdringtonia grows in equatorial and South Africa and in Madagascar; Actinostrobus is restricted to West Australia. SCIADOPITINEAE. Sciadopitys is confined to South Japan. SEQUOIJINEAE. Sequoia is confined to the Pacific coast of North California; S. sempervirens the species with ‘the stronger hold upon existence’ extends into Oregon, while S. gigantea forms groves in the valleys of the Sierra Nevada. PODOCARPINEAEH. Podocarpus, one of the more successful genera, is essentially a southern type: in Africa it extends from 1 Wieland (16), p. 224. XLII] DISTRIBUTION 127 Cape Colony through East Africa to Abyssinia; it occurs in §. America from Patagonia to Brazil and replaces Pinus on the mountains of Costa Ricat; in the West Indies, Malaya, in the Himalayas, China, Japan, Formosa?, Tasmania, New Zealand, New Caledonia, the Fiji Islands and New Guinea. Dacrydiwm has also a fairly wide range in the southern hemisphere, but like the other members of the family, except Podocarpus, it does not cross the equator; it is abundant in the Malay Archipelago® and occurs in New Zealand, Tasmania, New Caledonia, New Guinea, and one species grows in the Chilean swamps. Sazegothaea is a monotypic genus in Chile and Acmopyle an imperfectly known New Caledonian genus. Pherosphaera* occurs in New South Wales, Victoria, and Tasmania; Microcachrys, like Athrotaxis, is Tasmanian. Phyllo- cladus has a wide range in Tasmania, New Zealand, Borneo, New Guinea, and the Philippines. Some questions of exceptional interest from the point of view of the geographical distribution of Conifers in the Pacific region are ably discussed by Mr Guppy in the second volume of his admirable book Observations of a Naturalist in the Pacific between 1896 and 1899. He deals especially with Agathis, Podocarpus, and Dacrydium, and his remarks illustrate the. importance of taking into account palaeobotanical data in any general discussion of the problems suggested by the present and often discontinuous range of existing genera. ‘If,’ he says, ‘there is a real difficulty in applying our canons of plant-dispersal to the distribution of Dammara (Agathis], it is merely the same difficulty that has so often perplexed the botanist. with other Coniferous genera in continental regions, such as, for instance, the occurrence of Pinus excelsa on the far-removed mountains of the Himalayas, and the existence of the Cedar in its isolated homes on the Atlas, the Lebanon mountains, and the Himalayas. Such difficulties largely disappear if we regard the present distribution of the Coniferae as the remnant of what it was in an ancient geological period®.’ TAXINEAE. Tazus is chiefly a northern hemisphere genus; it occurs also in North Africa, Persia, India, the Philippines and the Far East, and extends from Newfoundland to Pacific North 1 Harshberger (11) p. 304. 2 Diimmer (12). 3 Stapf (96). 4 Groom (16). 5 Guppy (06) p. 300, 128 CONIFERALES (RECENT) (cH. America, Mexico, and Florida. Torreya has a more restricted and less continuous range in China, Japan, Florida, and California. Torreya taxifolia is almost extinct; it is separated by over 3000 miles from the other American species Z’. californica and the Pacific separates the latter from the two species in China and Japan’. Cephalotaxus lives in central China, Japan, and India. Anatomical features. The anatomy of Conifers, more especially fron the point of view of the identification of families and genera, has long occupied the attention of botanists, and although much has been done in the direction of more intensive study, the limits within which anatomical features may be safely used are still but vaguely defined. Jeffrey? goes so far as to claim for the anatomical characters of plants a taxonomic value equal to that assigned by zoologists to the anatomical features of animals. Though often extremely useful, in many cases anatomical characters do not reveal more than an affinity between a fossil specimen and a group of recent genera. Statements are often based on insufficient data and many authors have not appreciated the range of variation . in the vegetative shoots of a single tree. Attention has been drawn to the fact that anatomical features are especially variable in branches, and several authors have shown that characters to which importance has been attached are much less constant than has usually been supposed: many features, frequently accepted as trustworthy criteria from the point of view of identification, occur sporadically in other genera than those with which they are usually associated. In the following summary attention is directed to the comparative value of different characters, and prominence is given to possible sources of error in inferences based on anatomical features. The wood of a Conifer consists only of tracheids, with or without resin-canals, and xylem-parenchyma and is characterised by narrow medulla:y rays usually one-cell broad. For convenience in description it is proposed to speak of the wood of the Conifer type as pycnoxylic® in distinction to the Cycadean type of wood which is styled manoxylic. The presence or absence of 1 Berry (08?) p. 648. ® Jeffrey (05) p. 1. 3 ruxvds, compact; pavds, porous, loose in texture, XL] ANATOMY 129 well defined rings of growth should be noted and attention paid to the breadth of the late summer (‘autumn’) wood: Goeppert? considered the breadth of annual rings a character of importance, but Kraus? and others have shown that this is of little significance. In the Cupressineae (in the more restricted sense) it is probably true that the rings are generally though not invariably narrower than in Abietineae: in roots the later wood is smaller in amount and there is a more sudden transition to the spring-wood than in stems®. Though as a rule there is a considerable difference in the thickness between the walls of the spring and summer tracheids, in Podocarpus Nagi* the difference is slight. In some species of Araucaria the rings are absent or feebly marked, a fact noticed long ago by Nicol®, and in other Conifers, e.g., Widdringtonia juniperoides and Tetraclinis® there may be no definite rings; in Libocedrus macrolepis® there is but little difference in the thickness of the spring and summer tracheids. It is, however, impossible to say to what extent this is an inherent tendency and how far it reflects the influence of external conditions: it may be that the frequent absence of rings in Araucarian wood is explicable on the hypothesis that this family is the oldest and most closely related to Palaeozoic types, which are almost invariably characterised by an absence of rings: the habit of forming well defined spring- and late summer-wood may have been acquired at a later stage®. The interest of annual rings is rather biological than taxonomic and it is chiefly in connexion with fossil plants as tests of climate that attention has been directed to this feature. The genus Taxus is peculiar in having no resin-ducts in the cortex or stele of stem and root or in the leaves. In some genera resin-canals are a constant feature in the secondary wood, e.g., Pinus, Picea,’ Larix, Pseudotsuga; while in other Abietineae canals do not usually occur in the xylem. This distinction is, however, by no means constant and, as Jeffrey’® has shown, the 1 Goeppert (50). 2 Kraus (64) p. 146. 3 Gothan (10) p. 11; Penhallow (07) p. 31; von Mohl (62). 4 Fujioka (13). 5 Nicol (34) A. p. 139. § Conwentz (90) A. p. 33. 7 Fujioka (13) p. 213. 8 Thomson (13) p. 33; Gothan (07) p. 25. 9 Seward (92) B; Gothan (087); Antevs (16); (17). 10 Jeffrey (03); (05); etc.; Penhallow (07) pp. 123 et seg.; Jones (13°). s.IVO | 9 130 CONIFERALES (RECENT) _ [CH. great majority of Conifers which are normally without resin-canals in the wood have the power of producing them in response to traumatic stimuli. In Cedrus, Pseudolariz, and Tsuga resin- canals are usually confined to the primary xylem of the root but wounding induces the development of canals in other parts of the wood. In Cedrus, however, both horizontal and vertical trau- matic canals may occur whereas in other Abietineae the traumatic canals are only vertical!, Resin-canals may occur in the first-year wood of some species of Abies (fig. 690, B) and in Sequoia gigantea they are present in the first-year wood of vigorous branches and in the peduncles of cones, but do not normally occur in the later wood. In S. sempervirens canals are as a rule absent and are developed only after wounding (fig. 690, A). In the Araucarineae A B Fic. 690. A, Sequoia sempervirens. B, Abies sp. showing traumatic canals in the wood. (After Jeffrey.) : resin-canals are absent nor are they produced in injured stems: this failure to produce canals in response to disturbances set up by wounds is considered by Jeffrey to be an indication of the relatively late evolution of the family. Pinus, with abundant canals, is regarded as one of the more primitive types; Abies, with very few canals in healthy specimens but readily producing them on wounding (fig. 690, B), is regarded as a slightly later product of evolution, while Sequoia sempervirens (fig. 690, A) in which traumatic canals alone occur is still further removed from the original stock, and the Araucarineae, which are considered to 1 Jeffrey (05) p. 25. XLiIn] ANATOMY’ _ 131 have lost the power of reversion retained by Sequoia and Abies are placed higher in the evolutionary series. The vestigial signi- ficance of resin-canals is by no means generally admitted. Pen- Beno’ hallow? holds, and I believe rightly, that they are not primitive Mt features; their occurrence in the young shoots of certain species and in the peduncles of cones but not in the older wood may, as Gothan? suggests, be correlated with a greater need of protection. Kirsch? considers that the development of canals in young wood and in peduncles may be connected with the relatively greater abundance of food in those regions which, in his opinion, would induce a greater production of parenchyma and secretory passages. Moreover, if the occurrence of canals in the axis of a female cone of Sequoia gigantea is attributed to the retention of an ancestral | character, why do not canals also occur in the axis of the micro-. strobili? The facts demonstrated by Jeffrey and his pupils are of great interest, but considered by themselves they may equally well be interpreted as favouring the greater specialisation and more recent development of those genera in which the production of resin-canals is a normal character. The structure of the epithelial cells is employed as a taxonomic character though, as Conwentz* suggests, it is not a very satis- factory criterion and in petrified tissues it is often difficult to distinguish between true thick walls and walls thickened by secondary deposits. In Pinus the walls of the cells lining the canals are frequently thin®, but in some species thick; Larix and Picea have thick-walled epithelial cells. The occurrence of tyloses,— the parenchymatous cells that invade the cavities of water-con- ducting elements,—has generally been regarded as the monopoly of Angiosperms: though unknown in recent Ferns they occur in some extinct types. Chrysler® has shown that tyloses are produced in the tracheids of Pinus, apparently as a consequence of wounding. Tyloses have also been found in some fossil coniferous woods. The arrangement of the bordered pits on the radial walls of the tracheids is the character to which most attention has .been given. In the Abietineae they form eithe1 single or double, 1 Penhallow (07) p. 150. 2 Gothan (07) p. 40. 3 Kirsch (11); Thomson (13) p. 38. See also Burlingame (15%). 4 Conwentz (90) A, p. 45. 5 Groom and Rushton (13). ® Chrysler (08) B. p. 204. 9—2 132 CONIFERALES (RECENT) Fic. 691. A, Araucaria Bidwillii, tracheids from the cone-axis. B—D, F, Agathis bornensis ; B, wood of branch, multiseriate pits and, at the ray, scalariform pits ; C, tangential section; D, radial section of branch showing transition from alternate and opposite to scalariform pits; F, pits and rudimentary Sanio’s rims. E, Araucaria Cookii, radial section of root. G, H, Larix americana, radial section of root. (After Thomson.) XLUI] ANATOMY 133. and occasionally three or even four, rows (fig. 691, H); they are circular and not contiguous and the pits of a double row are as a rule on the same level; they are opposite and not alternate. This type of pitting occurs also in all the other families except the Araucarineae though in Agathis opposite pits are not unknown! (fig. 691, D) and Conwentz? states that he has seen separate and circular pits in the tracheids of recent species. The occasional occurrence of pits in clusters and not in opposite pairs has been described in Pinus Merkensis*: this is a feature characteristic of the tracheids of some fossil types, e.g., Cedroxylon transiens Goth. In Agathis and Araucaria there may be 1—3 rows and as many as 5 rows on the tracheids of cone peduncles (fig. 691, A). The pits are contiguous and flattened, and those of adjacent rows are alternate and hexagonal*. Thomson has called attention to the occasional occurrence, especially in the region of the rays, of transversely elongated or scalariform pits in the tracheids of Araucaria. A single series of flattened pits and even the occasional occurrence of alternate hexa- gonal pits are not infallible criteria of an Arau- carian affinity: in Dacrydium the pits of a double row may be alternate though rarely contiguous, and this is the case in some other genera, while in Saxegothaea® (fig. 692) the pits are as a rule uni- seriate and often flattened. Worsdell® describes circular and separate pits in the cone-scales of Araucaria and Thomson records alternate bi- seriate pits in the cone-axis and early wood of the Abietineae. Flattened pits are described in Podocarpus polystachya” and I have seen similar pits in the wood of Torreya californica and several Fre. 692. Tracheids 5 = of Saxegothaea con- other conifers other than the Araucarineae. The aia (After size of the bordered pits though worthy of notice Stiles.) is not in itself a feature of much value. As Nicol first pointed out, in Araucaria they are larger than in Tazus; in Pinus they are larger than in Araucaria: Kraus® speaks of the OLN {55 O_o AS 0; 1 Jeffrey (12) Pl. vr. fig. 5. 2 Conwentz (92) p. 35. 3 Groom and Rushton (13). 4 For good figures, see especially Thomson (13). 5 Stiles (08). . 6 Worsdell (99). 7 Gerry (10). 8 Kraus (83). 134 CONIFERALES (RECENT) [cH. Araucarian pits as small (9—12-8 ) in contrast to the broader pits (up to 21p) of the Abietineae, those in Cupressineae being inter- mediate in size. There may, however, be considerable difference’ in the size of the pits in a single typet. The occurrence of spiral thickening bands in addition to bordered pits is characteristic of the Taxineae, but spiral bands occur sporadically in the secondary tracheids of other Conifers, e.g., Phyllocladus, Larix leptolepis, species of Abies and other Abietineae*, also in some species of Cupressus®, In Pseudotsuga spiral bands may occur in all the tra- cheids of an annualring. Some authors assert that the arrangement and grouping of the bands in a tracheid constitute a character of generic value, but there is not complete agreement on this point?. The walls of tracheids frequently exhibit well marked spiral patterns®, due to an entirely different cause, which, especially in some petrified woods, closely simulate spiral bands. In the process of decay enzyme-action may etch into prominence the striation or spiral method of wall-construction; but the spirals are steeper than those of the true thickening bands. The presence of xylem-parenchyma, though of diagnostic value, is too uncertain and, variable a character to be used with great confidence. In young shoots of Sequota xylem-parenchyma may be absent though it is present in older branches*. Such parenchyma occasionally occurs in Abietineous wood’, but it is generally considered a charac- teristic feature of the Cupressineae though in the wood of some members of that family it is not always obvious. The presence of drops of resin in the cells which form vertical series in different parts of the wood, or only in the late summer wood, may render the xylem-parenchyma conspicuous both in transverse and longi- tudinal sections. Rows of parenchyma occur in the wood of Abies pectinata®, also in Podocarpus and Dacrydium. In Taxodium® the thick horizontal walls of the cells are a characteristic feature. Wood-parenchyma is rare in the Araucarineae and, as Penhallow” 1 Schenk in Schimper and Schenk (90) A. p. 848. 2 Bailey (09). 3 Jones (12); (13). * Gothan (05) p. 54; Penhallow (07) p. 41; Burgerstein (08) p. 104; Kraus (83) p. 103; Nakamura (83). 5 Kraus (88); Gothan (05). ® Conwentz (92) p. 35. * Burgerstein (06); Bailey (09). 8 Kny (10). ® Schroeter (80) p. 30. 10 Penhallow (04). XLII] ANATOMY 135 pointed out, tracheids with horizontal patches of resin may be mistaken for resiniferous parenchyma; but true parenchyma occasionally occurs}. Attention has been called to the diagnostic value of the hori- zontal thickening bands which on staining, and often in fossil wood, stand out as conspicuous features on the tracheids of the great majority of Conifers (fig. 693, C). Many authors speak of these bands as bars of Sanio?, apparently overlooking the fact that this term (Sanio’s ‘Balken’) was used by Miiller® for the horizontal bars previously described by Winkler‘ on the tracheids of Araucaria brasiliensis (fig. 693, I). Groom and Rushton® have also called. attention to the inaccurate use of the term Sanio’s bars and they suggest the more appropriate expression Sanio’s rims for the persistent margins of the primordial pit-areas which appear as horizontal lines between the bordered pits. An American author goes so far as to claim that ‘by far the most reliable criterion for diagnosing coniferous wood is the occurrence of the bars [rims] of Sanio®.’ But if, as Jeffrey and his pupils assert, Sanio’s rims are present on the tracheids of all Conifers except the Araucarineae the diagnostic value of this feature is exceedingly small. Jeffrey’ has shown that in the first-year wood of Araucaria and in the cone of A. Bidwillit the pits are not always contiguous and rims of Sanio may then be present. Moreover, as Thomson® states, the darkly stained lines between contiguous pits on some Araucarian tracheids (fig. 691, F) may be regarded as feebly marked rims of Sanio. It is not surprising that in the case of tracheids with 2—3 series of contiguous hexagonal pits, which leave no free surface’, ' Sanio’s rims are not represented”. _ The most recent contribution to our knowledge of the rims of Sanio is by Mr Sifton1! who describes them in petioles of Cycas 1 Jeffrey (12) p. 536. 2 Gerry (10). 3 Miiller (90). 4 Winkler (72). 5 Groom and Rushton (13). See also Rushton (16). 6 Holden (13). p. 252; (137). ? Jeffrey (12) Pl. v1. fig. b. 8 Thomson (13) p. 22. ® Gothan (10) p. 32. 10 In a recent paper entitled ‘Gliding growth and bars of Sanio’ (Grossenbacher, Amer. Journ. Bot. vol. 1. no. 10, 1914) the expression ‘ bars of Sanio’ is employed in an unusual sense and not in accordance with the ordinary usage of the term bars, or rims, of Sanio. 11 Sifton (15). 136 CONIFERALES (RECENT) [CH. revoluta. Jeffrey regards the occurrence of Sanio’s rims in the cone-axis of Araucaria as a vestigial phenomenon. He failed to find any rims of Sanio in the cone-axes of Cycads and this negative evidence was regarded as favourable to his view that the rims in the Araucarineae are derived from the more fully developed rims in the Abietineae. Sifton shows that the rims on the tracheids of Cycas revoluta agree closely with those in the xylem of the Araucarian cone-axes and with those in the cone-axis and root of certain Pines. On the assumption that roots and cone-axes are likely to retain ancestral characters, the resemblance of their rims to those found in the Araucarineae supports the view that the Abietineae are descended from ancestois which had rims of Sanio of the Araucarian or Cycadean type. The conclusion is that the shorter rims in the Araucarineae and on the tracheids of the cone- axis and root of the Abietineae represent the primitive form, the broader rims met with in the Abietineae and most other Conifers being later developments. The pitting on the walls of medullary-ray cells has in recent years received special attention: in some Conifers the horizontal: and tangential walls are strongly pitted (fig. 693, A, G), and this feature is clearly seen in both radial and tangential sections as also, in the case of the horizontal walls, in transverse sections (fig. 693, D, E, F). In most of the Abietineae the pits on the horizontal and tangéntial walls are a prominent feature while on the other hand in some Abietineae the pitting of these walls is feebly developed: to this type of pitting Gothan' has given the name Abietineous pitting. Tg the great majority of recent genera other than members of the Abietineae the horizontal and © tangential walls are smooth (fig. 693, L, O); but there are exceptions. The ray cells in the cone-scales of Agathis are pitted and species of Juniperus®, Libocedrus decurrens and Fitzroya also exhibit a form of Abietineous pitting. Gothan points out that in some Junipers and a few other Cupressineae the pits in the tangential walls differ in detail from the typical Abietineous form and that the pits in the horizontal walls are much less distinct than in the Abietineae: there is, however, no very clear distinction between 1 Gothan (05) p. 43. 2 Ibid. pp. 48, 45, fig. 7; Stopes (15) p. 63. ANATOMY @ @ @ OO MO10| O10 Sete] Ss (© o 2} 10} ©le}| % 44 1QD1@ 014, fodfee} oo] S| vse o0f88] Firs M y [Ef Olo! pele A Fie. 693. A, Abies Veitchii, medullary ray. B, Pinus silvestris, medullary-ray tracheid. C, Abies balsamea, pits and Sanio’s rims. D, E, Pits in medullary-ray cells in Abies homolepis. F, Pits in tangential wall of ray cell of Juniperus virginiana. G, H, K, L, M, N, O, Pits in medullary-ray cells (radial view) in Cedrus atlantica (G); Taxodium distichum (H); Podocarpus andina (K); P. salici- folia (L); Glyptostrobus (M); Sciadopitys (N); sp, spring wood; s, summer wood ; Thuya gigantea (O). I, Bars, b, in tracheids of Araucaria brasiliensis. P. Spiral bands in tracheids of Torreya nucifera. (A, N, after Nakamura; B. D—H, K, L, P, after Gothan; C, after Gerry; I, after Winkler; M, after Kraus; O, after Penhallow.) 138 CONIFERALES (RECENT) _ [CH his Juniperoid and Abietineous types. Each medullary-ray cell is longer in a radial direction than the breadth of a single tracheid and statements as to the number of pits on the radial wall of a ray cell have reference to the area bounded laterally by the vertical walls of a tracheid: this area may be designated the field (‘Kreuzungsfeld’; ‘aire mitoyenne’). It is, however, incorrect in many cases to speak of pits on the radial walls of medullary-ray cells, and if pits occur they are never bordered: in the Abietineae the walls are pitted, but in most other Conifers the pits seen in the field belong to the tracheids in contact with the rays. But in view of the general use of the expression medullary-ray pitting it would be inconvenient to discontinue the current terminology. There is a difference of opinion as to the value of medullary-ray pitting as a criterion of affinity, and it is probable that Gothan over- estimates the taxonomic significance of this character. Within certain limits the pitting on the walls of medullary-ray cells is undoubtedly important, but a comparison of sections of the wood of a collection of genera shakes one’s confidence in the conclusions based by some authors on the form and number of the pits in the field. In the Abietineae a single large simple. pit (‘Hipore’) occupies the field in some species of Pinus but the same type occurs also in Sciadopttys!, species of Podocarpus? (fig. 693, K, N), Microcachrys, Dacrydium, and Phyllocladus*. The Araucarineae are usually described as having several (2—6) oblique pits in each field and as a rule no pits in the horizontal and tangential walls of the ray cells, but Thomson‘ states that the bordered pits in the field of Araucaria and Agathis are confined to the tracheids and an examination of macerated tissue confirms the absence of pits on the walls of the ray cells. A similar absence of pits characterises some other Conifers. Gothan has suggested the term Cupressoid pitting for Conifers in which the field in the spring-wood contains ' bordered pits with a fairly broad pore in a more or less horizontal position, a type of pitting found in some Cupressineae as also in Sequoia and certain other genera. He applies the name Podo- carpoid pitting to woods in which the field shows bordered pits 1 Nakamura (83); Saporta (84) Pl. cxxxvit.; Fujioka (13). 2 Kleeberg (85). 3 Schenk in Schimper and Schenk (90) A. p. 855. 4 Thomson (13) p. 30. XLOI] * ANATOMY 139 with a narrower pore occupying an obliquely vertical position, a form of pit well shown in some species of Podocarpus. In the Podocarpineae and in most of the Cupressineae the tangential and horizontal walls of the ray cells are unpitted. An examination of sections of species of Thuya and some species of Cupressus reveals the presence of pits in the field with an almost vertical pore, and the variation in the breadth of the border and in the posi- tion of the pore is too great to admit of more than a restricted and cautious use of this anatomical feature as a means of distinguishing gene1a or even families. It, is by no means easy even in sections of recent woods to observe with accuracy the structure of the ray pits: in many cases they are more or less bordered, but the greater distinctness of the pore often leads to the neglect of the fainter border. Moreover the small medullary-ray pits may be con- verted into large pits by the action of fungal hyphae. The large pits of some Pines, Sciadopitys, etc., represent one extreme; intermediate types are represented by Cedrus, Taawodium, and Glyptostrobus, while in Juniperus and several other genera the pits are smaller and more numerous!. The depth of the rays as seen in tangential section is a feature to which much attention has been paid, but this is a very variable and comparatively unimportant character?. In a single species of Abies the depth varies from 1 to 63 cells?. Many authors in describing fossil wood state the number of rays per square milli- metre of a tangential section. Characters such as these may undoubtedly be useful in certain cases if used in conjunction with others, due allowance being made for the range of variation within the limits of a single stem. A more important feature is the occurrence of broad rays containing horizontal resin-canals such as those of Pinus, Picea, Lariz, and Pseudotsuga. Another useful criterion is afforded by the association of horizontal tracheids (fig. 693, B) with the parenchyma of a ray usually at the upper and lower margin but sometimes, e.g., Pinus canariensis*, mm the middle. The occurrence of such tracheids was formerly regarded as a trustworthy distinguishing feature of the Abietineae with the exception of Abies and Pseudolarix, but they are now known to 1 Penhallow (07). 2 Essner (86); Barber (98). 3 Fujioka (13). 4 Strasburger (91) p. 21. 140 CONIFERALES (RECENT) (cH. occur in Abies! and several members of other families. Chrysler? states that Cedrus differs from Abies in having ray tracheids mixed with marginal parenchyma, and at the limit of an annual ring the marginal tracheids may be replaced by shadowy cells or ghosts of cells. The tracheids may have smooth walls as in Cedrus, Tsuga, Larix, Pinus Strobus, or, as in other species of Pinus (‘Hard Pines’), their walls are characterised by irregular ingrowths or pegs. Wettstein® states that in Picea omorica horizontal tracheids though common in the main stem do not occur in the rays of branches. De Bary* recognised tracheids in the rays of Sciadopitys; they have been recorded also in Juniperus, Cupressus, Thuja, Sequoia® and, as the result of wounding, in Cunninghamia®, The occurrence of idioblasts in the form of irregular thick- walled elements is characteristic of the pith and cortex of Arau- caria, but similar cells are found in the pith of Torreya nucifera, Podocarpus neriifolia, Dacrydium cupressinum? and Cryptomeria. In some cases, e.g., Abies magnifica’, Picea omorica, horizontal rows of thick-walled cells form diaphragms in the pith. The structure of the secondary phloem has received relatively little attention and owing to its comparatively rare preservation in fossils it is less important to the palaeobotanist. In the Cupres- sineae the regular alternation of tangential rows of hard and soft bast is a characteristic feature, while in the Abietineae the phloem consists of sieve-tubes and parenchyma with a few scattered stone-cells. The absence of albuminous cells in the medullary rays of the phloem region in the Araucarineae is noteworthy and Thomson® states that he found none in Podocarpus. . The structure of the stomata in the leaves of Conifers is fairly uniform: their distribution should be noted though this in itself is not of much value as a distinguishing feature. They may be confined to regular bands (Abies, etc.) or grooves (Torreya, etc., fig. 694, B), or irregularly distributed. The position and number of resin-ducts is often a useful guide: to quote one example only, in the leaves of Agathis and Araucaria (fig. 694, C) the ducts Thompson (12). 2 Chrysler (15). Wettstein (90) p. 511. . De Bary (84) A. p. 490. See also Tassi (05) quoted by Vierhapper (10). Gordon, M. (12); Jones (132). ® Jeffrey (08). Kubart (11%) 8 Jeffrey (05). 8 Thomson (13) p. 31. 1 3 4 5 7 XLII] ANATOMY OF LEAVES 141 occur between the veins, but in the leaves of some species of Podocarpus (fig. 695, D), externally indistinguishable from those of Agathis, the ducts are below the veins’. Caution must be ex- ercised in using the number of resin-ducts as a diagnostic character. Schroeter? draws attention to the occasional absence of ducts in Picea excelsa leaves while in others 1 or 2 are present. In most leaves there is some mechanical tissue immediately below the epidermis either as scattered fibres or a continuous layer, but in Fic. 694. Leaves in transverse section. A, Cephalotarus Fortunei. B, Torreya myristica; t, transfusion-tissue. C, Araucaria imbricata; t, transfusion-tissue D, Pinus monophylia; e, endodermis. E, Cryptomeria japonica. F, Cedrus Libani; e, endodermis; m, medullary ray. Taxus and Torreya there is none. The occurrence of branched idioblasts is a striking feature in the mesophyll of Agathis, Arau- caria, Sciadopitys, and Podocarpus. The infoldings of the walls of the chlorenchyma are especially characteristic of Pine leaves: (fig. 694, D) and they occur also in Cedrus (fig. 694, F), Pseudo- lariz and some other genera. The structure, extent, and position of short isodiametric tracheids in association with the conducting ~ tissue is an important feature. This tissue, the elements of which 1 Seward and Ford (06) B. 2 Schroeter (97). 142 CONIFERALES (RECENT) [cH. are usually termed transfusion-tracheids’, is regarded by some authors as homologous with the centripetal wood of Cycadean leaves (Vol. 111. p. 31) and Bernard definitely adopts the term centri- petal xylem. Jeffrey and, more recently, Takeda’ do not attach a similar morphological significance to the short tracheids, and they are probably justified in their sceptical attitude. The transfusion tracheids are often reticulately pitted: in many leaves they form conspicuous flanges on the sides of the vascular bundles (fig. 694, B, C, t) or they may more or less encircle the vein. In some leaves, e.g., Araucaria, it is noticeable that the amount of transfusion tissue (fig. 694, C, ¢) increases as the vein is traced towards the leaf-apex until the long and narrow elements may be entirely replaced by a group of short transfusion-tracheids. Another type of accessory tracheid is occasionally met with, namely elongated tracheids traversing the mesophyll between the veins and the edge of the lamina. This is seen in the long and narrow leaves of some Podocarps?. The anatomy of cone-scales is too wide a subject for adequate treatment in this sketch. The general rule is that in each scale there are two sets of vascular strands, a lower set of normally orientated bundles and an upper inversely orientated series. In some cone-scales, e.g., Araucarva, concentric vascular strands are a prominent feature. The taxonomic significance of the ana- tomical characters of cone-scales has been discussed by Radais and more recently by Miss Aase®, to whose accounts the student is referred. It has been shown that characters which it has been customary to associate with a definite type of wood may occur sporadically in several other Conifers; but this does not invalidate conclusions based on the prevalent occurrence of such features in a given specimen. It is untrue to say that contiguous and alternate pits are the monopoly of the Araucaiineae and it is incorrect to assert that in Araucarian wood the pits are never separate. Similarly 1 So named by von Mohl; see Wordsell (97); Bernard (04) B.; Carter (11). 2 Jeffrey (08°). 3 Takeda (13); see also Thomson (13). + For further details with regard to leaf-anatomy, see Thomas, F. (66); Ber trand, C. E. (74); Mahlert (85); Striibing (88); Daguillon (90). 5 Radais (94); Sinnot (13); Aase (15). = XL] FAMILY CHARACTERS 143 the pits on the medullary-ray cells, whatever relative value we may assign to this character, are in many cases of considerable assistance even though we are not prepared to follow Gothan to the full extent of his trust in the taxonomic importance of medul- lary-ray pitting. Dr Groom? has described the northern evergreen Conifers as architectural xerophytes having xeromorphic leaves with a xero- phytic structure. He discusses in his remarks on the Ecology of Conifers the correlation of the characteristic wood-structure and the xeromorphic leaves, the cause of the survival of the Coniferae in competition with Dicotyledons, and other questions of interest to the student of the evolution and past history of the group. Short summary of the. characteristics of recent Conifers?. ARAUCARINEAE. There is a close agreement in the structure of the wood in the two members of thisfamily. Attention has already been called to the normal type of pitting of the tracheids and to certain exceptional forms. The occasional tendency towards a scalariform type of pitting (fig. 691, D) is an interesting point. Pits are fairly abundant on the tangential walls of ‘ew! xylem-elements. In the wood of Agathis robusta vertical rows of parenchyma are said to be fairly abundant. Annual rings not infrequently absent or feebly developed. Medullary rays one-cell broad, rarely double, usually 7—15 cells deep; but in Araucaria the depth may reach 26 cells: the ray cells occasionally present a distended appearance in tangential sections of the wood (cf. Ginkgo biloba). The persistence of the leaf-traces in the old wood of Araucaria is a striking feature considered by Lignier to possess diagnostic importance: Jefirey® states that traces are much less persistent in some young stems. In Araucaria each trace arises as a single strand, but in Agathis it leaves the perimedullary region as a double bundle®. Agathis (fig. 695). Leaves sessile or slightly petiolate, opposite, sub- opposite or, on the main axis, spiral; ovate, broadly lanceolate (A. loranthi- folia, 13'x 5em.; A. macrophylla 17 x 5cm.). Resin-canals between the veins ; transfusion-tracheids fairly abundant, but less prominent than in Araucaria. The almost spherical megastrobili (fig. 696) are very characteristic; they 1 Groom (10). 2 In addition to the text-books on Conifers by Beissner, Veitch, and the account in Die Natiirlichen Pflanzenfamilien the student should consult the Report of the Conifer Conference, Journal of the Royal Horticultural Society, Vol. xv. 1892. 3 Noelle (10). 4 Thiselton-Dyer (01%); Seward and Ford (06) B. 5 Jeffrey (12) p. 565. 6 Thomson (13) p. 15. 144 CONIFERALES (RECENT) [cH. Fic. 695. A, Agathis Moorei. B, E, Agathis australis. C, Agathis vitiensis. D, Podocarpus Motleyi. (After Seward and Ford; } nat. size.) B Fic. 696. Piths f, pericycle fibres; ; c, cambium. cheids and the cells are more elongated 1 Groppler (94); Solereder (99) p. 34; (08) p. 5. See also Jeffrey and Cole (16). of Mil XLII]: SOURCES OF ERROR 163 vertically than in the rays of a Conifer. Attention has already been called to the difficulty of distinguishing between the foliage-shoots of some Conifers, Dicotyledons and Lycopodiaceous plants. The Fig. 711. A, Veronica Hectori; B, Callitris calcarata; C, Veronica cupressoides ; D, Athrotaxis cupressoides; E, Crassula lycopodioides. M.S. twigs reproduced in fig. 711 are examples of misleading resemblances, and similar instances are cited by Bommer. The Conifer Podocarpus dacrydioides was described by Banks and Solander as Lycopodium 11—2 164 CONIFERALES (RECENT) [CH. XLIIL arboreum; Dacrydium Bidwillii1, D. Franklini and D. araucarioides (fig. 708, B) recall some species of Lycopodium, and D. cupressinum?, a dimorphic species, may simulate Lycopodium tetragonum. The long, linear, distichous leaves of some species of Podocarpus and Cephalotaxus might, as fossils, be confused with the pinnate leaves of Cycas; further, as Bommer points out, the leaves of Podocarpus Blumei resemble those of Agathis and the seeds are similar to those of Dehaasia media (Lauraceae). The Cupressus type of shoot occurs in Baccharis scolopendra as in other Dicotyledonous plants (fig. 711). The presence of a midrib in a linear Tarus-like leaf though usually easy to recognise is not always obvious, e.g., in Torreya nucifera the midrib cannot always be distinguished on the upper face of the leaves. The recurrence of a similar habit in many Conifers renders difficult the identification of vegetative shoots, particularly as in fossil specimens the precise method of attachment of the leaves, their texture, and other features are frequently unrecognisable. The tendency to dimorphism in many genera is another difficulty: examples of dimorphic shoots _ are afforded by Dacrydium laxtfolium, D. Kirkwi, D.elatium (fig. 708, A), Juniperus chinensis, Araucaria excelsa, Callitris glauca, Podo- carpus cupressina®, P. imbricata, Thuya occidentalis, etc. Allusion has been made to the considerable variation in the length of leaves on a single branch of different Conifers: in such a form as Crypto- meria japonica var. spiralis Sieb. and in similar varieties of other genera the spirally twisted leaves, reminding one on a small scale of the ‘wind-blown’ Acanthus leaves on a Byzantine capital, constitute a feature which might be regarded as of taxonomic importance. The investigation of the cuticular membranes of Conifer leaves, as yet but little attempted, may supply useful criteria as in the case of Cycadean fronds. 1 Pilger (03) fig. 4 A. 2 Kirk (89) Pls. xvu., XIx. 3 Bennett and Brown (52) Pl. x CHAPTER XLIV. CONIFERALES (FOSSIL). Tue task of deciphering the fragmentary remains of Conifers is particularly difficult and no branch of palaeobotanical research makes greater demands upon the patience and self-control of the student. As Saporta says, ‘Aucune étude n’ouvre des perspec- tives plus étendues, mais aucune aussi n’exige plus de réserve et de tatonnements!.’ The determination of impressions of ill-preserved vegetative shoots is often impossible and it is regrettable that many authors have been too ready to employ generic names denoting identity or close relationship with recent types on wholly inade- quate grounds. A recent writer thus sums up the situation - created by an excessive faith in superficial resemblances and a lack of familiarity with existing representatives of the group :— _ ‘Where a knowledge of reproductive parts is lacking, chaos reigns supreme.’ It may be added that impressions or casts of cones in many cases do not afford any real assistance. A comparison of the various forms of foliage-shoots. and strobili met with among recent Conifers demonstrates the danger: of placing confidence in external resemblance as a guide to affinity. It is seldom that reproductive organs are well enough preserved to enable us to recognise features of primary systematic value. Though little has so far been done to test the value of epidermal characters as aids to identification, such results as have been obtained? favour the conclusion that this line of investigation promises to be less fruitful for Conifers than for Cycadean plants. Petrified wood of the Conifer type is abundant in plant-bearing strata from the later Palaeozoic rocks upwards, and considerable pains have been taken to utilise to the full this source of information. Within wide limits anatomical characters are undoubtedly valuable, but the recent tendency to subdivide comprehensive genera, which are 1 Saporta (62) p. 309. 2 Holden, R. (157). 166 CONIFERALES ; [cH. recognised as embracing several recent genera, into genera implying a limitation of affinity within narrower bounds has, I venture to think, been carried too far. The investigation of fossil coniferous wood, in spite of the disappointing quality of the data from the systematist’s point of view is well worth attention. An examina- tion of fossil wood from different geological horizons brings to light many striking instances of a mixture in single plants of features now characteristic of distinct genera. It is the generalised forms that throw light on the nature of the changes produced in anatomical structure in the course of evolution. The older and more generalised types are of special importance to the student of phylogeny. The very difficult question as to the stock from which the Conifers are derived is too wide to be adequately discussed in a general treatise. It is probable that the Coniferales are mono- phyletic, the Araucarineae being the oldest representatives of the group while the Podocarpineae are a closely related series. The widely held view that the Araucarineae are descended from Cordai- talean ancestors is by no means definitely established; it rests mainly on anatomical evidence and the arguments based on a comparison of the reproductive shoots are far from convincing. On the other hand those who favour a Lycopodiaceous ancestry for the Coniferales are confronted with difficulties which, though I venture to think they are not insurmountable, have not been adequately met!. The suggested linking up of the Cordaitales, through types in which the cylinder of secondary xylem is sup- plemented by separate primary strands of vascular tissue, with Lyginopteris and other Pteridosperms leads to the inclusion of the Coniferales among the descendants of an ancient Filicinean stock, but here too the chain of evidence is incomplete particularly as regards the lack of data as to the nature of the reproductive organs of several Palaeozoic genera founded on anatomical characters. The problem is still unsolved: the discovery of additional types and a more thorough comparative study of such data as we possess may enable us to see more clearly the paths along which evolutionary tendencies have operated, but the absence of records of the vegetation of pre-Devonian times deprives us of the means of following to their common source the different phyla of vascular 1 For a useful summary of arguments see Burlingame (15). XLIv] GYMNOSPERMOUS WOOD 167 plants which in the Permo-Carboniferous era had already advanced far beyond the simple ancestral forms which the botanist seeks but rarely finds. The various examples of fossil genera founded on the anatomical features of vegetative organs are dealt with in a separate section and not included with impressions in the descriptions of the several families, partly on the ground that it is rarely possible to demon- strate a connexion between the two sets of records and in part with a view to give a more connected account of the results so far derived from a study of petrified wood. Cross-references to anatomical structure are given in the descriptions of vegetative and repro- ductive organs when there appear to be sound reasons for assuming a generic or family connexion. The classification of woods is at best provisional and the generic characters are far from constant. The main point is that the student cannot afford to neglect this line of enquiry if he desires to obtain a comprehensive view of the changing combinations of structural features preceding their distribution among existing genera. A comparison of recent Conifers and Cycads with their Mesozoic representatives brings out very clearly the fact that while on the one hand the modern Cycads differ widely from the Cycadean type which played a prominent part in Mesozoic floras, recent Conifers on the other hand agree closely in their main features with their Mesozoic ancestors. The Cycads as we know them now are a more recent product of evolution than the Conifers though it by no means follows that the Conifers in the wide sense are ' the more ancient group. Fossit GyMNOSPERMOUS woop (Coniferales). The earliest attempts to identify petrified wood are summarised by Goeppert!, Knowlton*, and other authors. Luidius (Lhwyd)? at the end of the seventeenth century employed the general desig- nation Lithozylon, and the termination -zxylon is still used in generic names applied to fossil wood in conjunction with some prefix im- plying agreement in the more important anatomical features with some recent genus or family. For woods exhibiting a combination of characters unknown in existing genera a distinctive 1 Goeppert (50). 2 Knowlton (892); Gothan (05). 3 Luidius (1699) A. PP 7 168 CONIFERALES [cH. prefix is employed, e.g., Xenoxylon, and some authors make use of a name, e.g., Woodworthia, which does not indicate that the diagnosis is based on anatomical features. Conwentz! adopted the method of adding the prefix Rhizo- to generic terms for wood believed to belong to roots, and Felix? and Lignier? have employed the prefixes Cormo- and Clado- for stem- and branch-wood respec- tively. It is, however, seldom that such differentiation is possible- and it is questionable whether it is wise to attempt refinements of this kind. Barber, in his critical paper on a species of Cupressino- cylon, calls attention to Strasburger’s description of an old mori- bund stem of Larix with root-like characters and Gothan® speaks of a branch of Pinus silvestris with root-attributes. The differences between branches and the main stem are not sufficiently known even in the more familiar types to justify the use of the prefixes Cormo- and Clado- in descriptions of fossil specimens. The scientific study of fossil wood began with Nicol® and Witham whose work was rendered possible by methods of section- cutting first employed, according to Nicol, by a Mr Sanderson, a lapidary. Opinions expressed by Nicol on methods of investi- gating petrified wood are still pertinent afte1 a lapse of 80 years :— ‘To pronounce with certainty whether a fossil Conifer be essen- tially different from any known individual of the recent kind, it would be requisite to have a thorough knowledge of the structure at least of all the different tribes of recent Coniferae; and yet several distinct fossil genera have been indicated by a person who has examined, and that too very superficially, only three slices of three recent Pines, differing not essentially from one another.’ In recent years the tendency has been towards a more detailed study of anatomical characters such as the distribution and form of the pits on medullary-ray cells. The facts recorded in the Chapter on Recent Conifers illustrate the difficulty of arriving at a thoroughly satisfactory classification of anatomical features that may serve as criteria in the identification of recent genera: even in the case of well-preserved fossil wood we have as a rule to rest content with a generic name denoting a combination of characters met with in more than one existing genus. Moreover, as already 1 Conwentz (80) A. 2 Felix (82). 3 Lignier (072). + Barber (98). 5 Gothan (05) p. 19. ® Nicol (34) A. p. 141. XLIV] GYMNOSPERMOUS WOOD 169 shown, recent work has tended to reduce the taxonomic value of certain characters such as the occurrence of ray-tracheids which are more widely distributed than has generally been supposed. In this connexion a word may be added with regard to some common sources of error in anatomical investigation. There is the obvious danger of confusion between features due to petrifying agents or to decay before petrification and those present in the living tree: the thickening of cell-walls, e.g., those of medullary rays, has been shown in some cases to be a pathological pheno- menont. ‘The partial obliteration of bordered pits by decay may cause them to appear separate though originally in contact (fig. 475, B, Vol. 11. p. 257). The recognition of pits on the tangential and horizontal walls of medullary-ray cells is often very difficult, and negative evidence may be misleading. It is by no means always a simple matter to distinguish between true canals and canal-like spaces formed by the destruction of groups of tracheids (e.g., Pityoxylon eiggense; fig. 725). In one case it has been shown that leaf-traces traversing broad medullary rays were mistaken for horizontal resin-canals*. The spiral lines frequently seen on the walls of petrified tracheids caused by the directive influence on the structure of the membrane of the course of enzyme-action may simulate the spiral bands characteristic of Taxus, Torreya, and Cephalotaxus. These are a few of the pitfalls in the path of the palaeobotanist, but despite the difficulties and the frequency with which imperfect preservation prohibits complete diagnosis, the investigation of fossil wood is well worth the attention of students equipped with an intimate knowledge of recent Conifers. The unpromising nature of the material may be a deterrent, though lignitic and other specimens not thoroughly petrified are amenable to special treatment, In the account of recent Conifers attention is called to the sig- nificance of rings of growth: the subject has recently been exhaus- tively treated by Antevs4 and students should consult his memoir in the Progressus rei botanicae for references to the literature. The 1 Gothan (07), p. 25. 2 Penhallow (00) p. 76; Thomson and Allin (12). 8 For methods, see Jeffrey and Chrysler (06); Hollick and Jeffrey (09) B. Gothan (09); Sinnott (09). 4 Antevs (17). 170 CONIFERALES — [CH. subject is interesting and beset with difficulties but well worthy of more thorough treatment than it has so far received. Though petrified Coniferous stems are usually represented by the secondary wood only, the phloem and cortical tissues are sometimes preserved ° and afford useful information. Examples of petrified phloem and other extra-xylem tissues are described by Lignier and other authors. In his description of silicified plants from Franz Josef Land, of Lower Cretaceous or Upper Jurassic age, Solms-Laubach? includes some pieces of Coniferous bark showing patches of peri- derm alternating with secondary phloem consisting of sieve-tubes, phloem-parenchyma, and fibres, also some stone-cells. Some of the sieve-tubes are shown in fig. 718, B, with well-preserved sieve- plates, a feature very rarely preserved. There is not enough wood associated with the phloem and periderm to serve as a means of identification, but Solms-Laubach speaks of the bordered pits on the tracheids and the pits of the medullary rays as indicating Pityoxylon or Cedroxylon. A new generic name Vectia has been instituted by Dr Marie Stopes? for a mass of petrified phloem which she compares more especially with the phloem of recent Conifers: while recognising that the specimen cannot be assigned with confidence to a particular group of Gymnosperms I venture to think it is almost certainly a portion of a Cycadean stem. Reference has already been made in the section devoted to the anatomy of recent Conifers to the relative importance and con- stancy of different characters from a taxonomic point of view and this question need not be further considered. The method of classifying coniferous wood in general use is based on a scheme proposed by Kraus?. A modified form of this scheme was pub- lished by Schenk* and more recently Penhallow', Jeffrey, Lignier, Gothan, Dr Stopes, and other authors have considerably extended our knowledge. Dr Gothan*, whose memoir on the anatomy of Conifers contains much valuable information, employs several generic names denoting identity with recent types, and while admitting the great advance made by him and other workers in 1 Solms-Laubach (04). ° Stopes (15) p. 247. See p. 419, Vol. 1. 3 Kraus in Schimper (72) A. p. 363. 1 Schimper and Schenk (90) A. p. 860. > Penhallow (07). ® Gothan (05). XLIv] PETRIFIED TREES 171 this field, it is difficult to avoid a suspicion of overstraining the significance of certain anatomical minutiae beyond the limits of safety. The great abundance of petrified wood in strata ranging from the late Palaeozoic through the Mesozoic and Tertiary formations, Fic. 712. Section of the north face of Amethyst Mountain, Yellowstone Park. (After Holmes. ) often in places which have yielded few other plant fossils, is in itself a strong incentive to research in a department of palaeobotany that has suffered from superficial work and hasty conclusions. Petrified tree-trunks are among the most impressive monuments 172 CONIFERALES [CH. of former ages: the petrified forests in the desert east of Cairo}; the vast accumulation of Triassic stems, some reaching a length of 200 ft. and 7—10 ft. in diameter, over an area of 10 square miles in Arizona County?; the succession of Tertiary forests in 2000 ft. of voleanic sediment exposed on the sides of Amethyst Mountain in the Yellowstone Park? (fig. 712); and on a smaller scale the Jurassic trees in the Portland quarries are a few of many striking examples of the wealth of material. CLASSIFICATION OF FOSSIL CONIFEROUS WOOD. A. Araucarian pitting on the tracheids. I. DADOXYLON. (Including Araucariorylon of Authors.) Bordered pits on the radial walls of the tracheids, if uniseriate flattened above and below, when in two or more rows alternate and polygonal; separate and circular pits, though rare, may occur. Rims of Sanio usually absent though their occasional occurrence on the secondary tracheids in the cone-axis of recent species of the Araucarineae shows that they are not entirely foreign to wood of the Araucarian type. Xylem-parenchyma absent or rare and may be represented by resiniferous tracheids. Medullary rays uniseriate, rarely double; horizontal and tangential walls smooth; there may be 1—15 small pits in the field, though whether they actually belong to the walls of the ray cells or to the adjacent tracheids has not been definitely determined in fossil species. The pits are simple or bordered, circular or elliptical. Resin-canals are absent both from normal and wounded wood. Palaeozoic to Recent. B. Pitting on the radial walls of the tracheids of the common Coniferous type ; the pits are separate and circular and, if in two or more rows, opposite. Contiguous and more or less flattened pits occur spasmodically on the tracheids of the wood of the genera included in this section. Well-developed rims of Sanio usually occur on the tracheids. 1 Unger (59). ® A good example of an Arizona tree-trank is exhibited in the Plant-Gallery of the British Museum. See Ward (008). * Knowlton (99); Holmes (78); Seward (113) p. 60, fig. 6. XLIV] CLASSIFICATION OF CONIFEROUS WOOD 173 II. CUPRESSINOXYLON. (Including Glyptostroborylon and Taxodioxylon.) . Xylem-parenchyma scattered through the wood and not con- fined to any particular region, often containing resin; the trans- verse walls may be thick and pitted. Medullary rays uniseriate, horizontal and tangential walls smooth (unpitted); there are generally several small pits in the field though in some species referred to this genus there may be a single pit. In the region of the spring-wood the pore of the apparently bordered ray-pits is more or less horizontal; but the form and position of the pore are variable. Resin-canals absent except in wounded parts of the wood. Jurassic to Recent. | Ill. TAXOXYLON. The same anatomical features as in Cupressinoxylon except that the tracheids of the secondary xylem have spiral thickening bands. Tertiary to Recent. IV. MESEMBRIOXYLON. Gen. nov. (Including Podocar- poxylon, Phyllocladoxylon and Paraphyllocladoxylon.) Xylem-parenchyma usually present and scattered, but it is not so characteristic a feature as in Cupressinozylon. Medullary rays usually uniseriate; the pitting is confined to the radial walls as in Cupressinoxylon but in the region of the spring- wood the pore is oblique or more or less vertical; in the summer- wood the pits in the field are indistinguishable from those of Cupressinorylon. There are often several pits in the field but in some species there may be one or two large simple pits in the field. Resin-canals present only in wounded parts of the wood. Jurassic to Recent. V. PARACEDROXYLON. _A genus of doubtful affinity. Bordered pits on the tracheids usually separate; no rims of Sanio. Xylem-parenchyma confined to wounded regions. Medullary- ray cells pitted only on the radial walls except in the injured parts of the wood where the other walls may be pitted. There are 4—6 circular, apparently bordered, pits with an oblique pore in the 174 CONIFERALES [CH. field. In the characters of the normal wood this genus agrees most closely with Cupressinoxylon and Mesembrioxylon. Cretaceous. VI. CEDROXYLON. In some species included in this genus contiguous and flattened bordered pits are fairly common on the radial walls of the tracheids. Xylem-parenchyma, if present, confined to the late summer- wood. Medullary rays uniseriate; all the walls are pitted; tracheids may be present in the rays though they are usually absent. There are 1—6 or rarely more pits in the field, either simple or apparently bordered. Resin-canals confined to wounded regions. Jurassic (Triassic species doubtful) to Recent. VII. PITYOXYLON. (Including Piceorylon and Pinuzylon.) Though in the great majority of cases the tracheids of the secondary xylem have no spiral bands, the presence of such bands in the recent genus Pseudotsuga shows that this feature may occur in wood of the Pityoxylon type. Resin-canals present in the normal wood. Medullary rays of two kinds, uniseriate rays consisting of parenchyma and, in many cases, ray-tracheids, also fusiform rays with horizontal resin-canals. The walls of the ray-tracheids are either smooth or irregularly dentate. All the walls of the medullary-ray cells are pitted; there may be one large simple pit in the field or several small, apparently bordered, pits. ; Jurassic to Recent. VIII. PROTOPICEOXYLON. Similar to Pityorylon except in the absence of horizontal resin-canals in the normal wood; such horizontal canals as occur are regarded as traumatic. Cretaceous to Tertiary. C. Genera in which Araucarian features, especially as regards the tracheal pitting, occur in association with characters met with in recent Abietineae. The genera included in this section afford examples of generalised types and do not resemble recent forms so closely as do the other genera. XLIv] CLASSIFICATION OF CONIFEROUS WOOD 175 IX. WOODWORTHIA. Araucarian tracheal pitting; annual rings feebly marked. Short shoots and a subtending leaf are present in the secondary wood. Resin-canals absent. Medullary rays uniseriate; pits confined to the radial walls. Triassic. X. ARAUCARIOPITYS. Similar to Woodworthia in the possessfon of short shoots and in the pitting of the tracheids though separate and circular pits also occur. Vertical resin-canals abundant in wounded regions. Medullary rays uniseriate; all the walls pitted. Cretaceous. XI. PROTOCEDROXYLON. (Including Metacedroxylon.) Bordered pits on the radial walls of the tracheids in 1—3 rows, usually of the Araucarian type but separate pits also occur. No rims of Sanio and no resin-canals. Xylem-parenchyma usually absent. Medullary rays generally uniseriate; all the walls pitted; 1—3 circular, simple, pits in the field. Jurassic. XII. XENOXYLON. Tracheal pits large, generally flattened above and below though not always, often transversely elongated. Resin-canals absent. Medullary rays uniseriate; pitting confined to the radial walls; usually one large simple pit in the field. Jurassic (Triassic ?). XIII. ANOMALOXYLON. When uniseriate the pits on the tracheids are usually contiguous and flattened; if biseriate the pits are opposite; separate pits also occur. True resin-canals absent, but canal-like spaces lined with small cells occur in some large medullary rays and constitute a charac- teristic feature. Medullary rays uniseriate; pits confined to the radial walls; 2—3 circular simple pits in the field. Jurassic. 176 ‘ CONIFERALES (CH. XIV. THYLLOXYLON. Tracheal pitting partially Araucarian. Xylem-parenchyma at the end of a year’s growth. No true resin-canals, but the central parenchyma of some of the broader rays is replaced by a canal-like space often filled with tyloses. Medullary rays uniseriate; all the walls pitted. Jurassic. o XV. PLANOXYLON. Wood with well-marked annual rings, resin-canals usually absent; tracheids with 1—3 rows of alternate hexagonal bordered pits on the radial walls and in the late wood there may be a single row of separate pits. Xylem-parenchyma occurs only between the spring elements and the last-formed tracheids of the previous year. Medullary rays almost entirely uniseriate; all the walls pitted. Lias to Cretaceous. I. DADOXYLON [and Araucariozylon]. The anatomical characters implied by the expression ‘wood of the Araucarian type’ are enumerated in the Chapter on Recent Conifers, and in Chapter xxx11l. reference is made to the impossi- bility of drawing a clear line of division between the wood of Araucarian plants and that of certain members of the Cordaitales?!. The name Araucarites was used by Goeppert? for fossil wood of the Araucarian type, but the previous use of this name by Presl for impressions of foliage-shoots and cones renders unsuitable its application to wood apart from the fact that Araucarites at once commits an author to a determination implying an affinity which in many cases cannot be demonstrated. Endlicher’s non-committal genus Dadoxylon® has been widely used, especially for Palaeozoic wood having the characters of Araucaria or-Cordaites. This designation leaves open the question of precise systematic position. In 1882 Kraus? instituted the genus Araucarioxylon, a name which has been widely adopted for fossil wood both from Palaeozoic and later formations. The practice of limiting Dadoxylon to Palaeozoic species and reserving Araucarioxylon for Mesozoic and Tertiary 1 See p. 248, Vol. m. 2 Goeppert (45). 3 Endlicher (47). 4 Kraus in Schimper (72) A. p. 370. XLIv] DADOXYLON 177 wood has already been criticised!: it is pointed out that the appli- cation of an age-test is scientifically unsound and cannot fail to be misleading. Although it is probably true that species of Dadoxylon from strata later than the Rhaetic series are in the majority of cases Araucarian, we have no adequate grounds for definitely naming such types Araucariorylon in the sense of membership of the Araucarineae. Similarly some Palaeozoic species of Dadoxylon may well be more closely allied to the Arauca- rineae than to Cordaites: Dadozrylon valdajolense (Moug.)? and D. Rhodeanum (Goepp.)® have both been referred to Walchia: the latter species is included by Tuzson in the genus Ullmannites. We do not know the lower geological limit of the Araucarineae, nor do we know when the Cordaitales became extinct. Tuzson* subdivides wood of the Dadozylon type into several genera in- eluding Pycnophyllum, to which is referred Dadoxylon Brandlingit (Lind. and Hutt.), Ullmannites applied to the type recently named Eristophyton Beinertianum by Zalessky and previously in- cluded by Scott in Calamopitys, also to Dadoxylon Rhodeanum and D. saxonicum, Pagiophyllites, including P. keuperianus (Goepp.), and for Tertiary species the term Araucarites is used. This nomenclature, based partly on age and in part on a supposed connexion between the wood and foliage-shoots, is opposed to sound principles and in some cases is at variance with the true character of the species. When evidence is available in support of a reference either to the Araucarineae or to the Cordaitales the qualifying terms Cordaioxylon or Araucariorylon may be added after Dadoxylon. Such a species as Dadoxylon permicum Merck.® is one of many examples of a Dadozxylon that cannot be more pre- cisely identified. Dadoxylon australe Crié® from Triassic rocks in New Caledonia must be included in the same category. This species is founded on wood showing well-marked rings of growth; there are two or more rows of alternate polygonal pits on the radial walls of the tracheids and the medullary rays are uniseriate and . 3—15 cells in depth. Crié’s species should not be confused with 1 See p. 249, Vol. mm. See also Gothan (05) p. 14; Potonié (02) p. 229. 2 Fliche (03); Mougeot (52) A. 3 Potonié (99) B. p. 294; Gothan (07) p. 17; Tuzson (09) p. 34. 4 Tuzson (09) p. 17. 5 Mercklin (55) p. 53. 6 Crié (89) p. 5, Pls. 1., m1, Vv. Ss. IV 12 178 CONIFERALES [cH. Dadoxylon australe Arber! based on Palaeozoic wood from Australia: the substitution of Dadoxylon for Araucarioxylon, the name used by Crié for his New Caledonian species, necessitates a new specific designation for Arber’s type, which it is proposed to rename Dadozylon Arberi. From Liassic beds in Yorkshire Miss Holden? has described a species as Araucariorylon sp. which she suggests may be one of the oldest representatives of the Arau- carineae or perhaps a Jurassic example of Cordaites: the latter identification is supported by a reference to the recorded occurrence by Lignier of an Artesia? in French Jurassic beds, a test of affinity that cannot be accepted as satisfactory. There remains for consideration the debated question as to the value to be attached to the occurrence of contiguous and flattened pits as an index of Araucarian affinity when this feature is associated with a type of medullary-ray pitting foreign to Dadoxylon. In Gothan’s genus Xenorylon? the tracheids have usually large flattened pits, but the pits on the radial walls of the medullary- ray cells are very different from those characteristic of Arauca- rineous wood. Itis, moreover, not uncommon to find instances of contiguous and alternate pits on the tracheids of a stem in which the more usual type is the Abietineous arrangement. Gothan® lays greater stress on the nature of the pitting on the walls of medullary cells, but Jefirey® has discovered typical Abietineous ray cells in the cone-axis of Araucaria and Agathis. Miss Holden’ goes so far as to maintain that the only feature which holds absolutely is provided by the rims of Sanio: these are invariably absent in Conifers with Araucarian affinities except on the first few tracheids of the cone-axis of Araucaria and Agathis. This author records as Araucariorylon sp.6 a wood (described on a later page) from New Jersey possessing opposite pits on some of the tracheids, also rims of Sanio. We cannot lay down any definite rules with regard to the sporadic variation in tracheal or medullary pitting or as to the relative value to be assigned to one or other character. The statement by Thomson® that the ray cells of such 1 Arber (05) B. p. 191, text-figs. 40—43. 2 Holden, R. (13°) p. 540, Pl. xx. fig. 28. 3 See p. 248, Vol. m1. 4 See page 248, Vol. mm. 5 Gothan (05). 8 Jeffrey (12). * Holden (14) (13?) p. 544. See also Sifton (15). 8 Holden (14) p. 171. ® Thomson (13). XLIV] DADOXYLON | 179 Araucarias as he examined have no pits even on the radial walls leads one to suspect the accuracy of some of the many recorded instances of fossil Araucarian wood with medullary-ray pits; but this does not affect the value to be attached to the presence or absence of well-defined pits on the vertical or horizontal walls of medullary-ray cells. It is generally true that in the ray cells of Abietineae these walls are pitted while in the Araucarineae they are unpitted, and it is equally true that the predominance of alternate and contiguous pits on the tracheids is evidence of Arau- carian affinity. Though generally absent from Araucarian wood xylem-parenchyma occasionally occurs in stems otherwise iden- tical with the usual Araucarian type. The genus Araucariopsis+ was instituted by Caspary for specimens distinguished from most examples of Dadoxylon by the presence of scattered xylem-paren- chyma but, as Gothan? points out, a distinctive name is super- fluous; the type-species of Araucartopsis, A. macractis, should be transferred to Dadoxylon; in the possession of xylem-parenchyma it agrees with Dadoxylon septentrionale Goth. from Spitzbergen. The importance attached by Jeffrey and other American authors to the presence of Sanio’s rims on the tracheal walls is, I venture to think, greatly overestimated. The determination of fossil wood is to a large extent a question of relative values. There is clear evidence, and-it would be surprising were it not so, that in several extinct types there are combinations of character pointing to less sharply defined boundaries than those which delimit existing families and genera. It is in the conclusions drawn from genexal- | ised types that authors differ. An outstanding fact is the pre- dominance in Palaeozoic stems of the Araucarian form of tracheal pitting which is unquestionably a much older type than that characteristic of the Abietineae. The following definition is based on specimens agreeing in the sum of their characters with recent Araucarineae, but there are various genera described by Jeffrey, Miss Holden, and other authors and believed by them to be more or less closely allied to the Araucarineae which are not provided for in the definition. These genera are treated separately as generalised types and the decision as to the nature of the evidence 1 Caspary and Triebel (89) p. 81. Pls. xrv., xv. 2 Gothan (10) p. 9. 12—2 180 CONIFERALES [cH. they afford with regard to the phylogeny of the Araucarineae and. the Abietineae must be left to the individual student. Annual rings occasionally well marked but not infrequently absent or indistinct. Tracheids with uniseriate and more or less flattened bordered pits or with two or more rows of alternate polygonal pits on the radial walls. The alternate disposition, even if unaccompanied by flattening and the polygonal contour of the pits, is a Dadoxylon feature if it is the dominant arrangement ‘and not a sporadic occurrence. Bordered pits occasionally occur on the tangential walls but they are smaller and comparatively rare. Rims of Sanio absent. Xylem-parenchyma usually absent or feebly developed; resiniferous tracheids occasionally present. Medullary rays uniseriate, very rarely double, homogeneous, 1—30 or as many as 50 cells deep; walls comparatively thin and without pits on the horizontal and vertical walls; the radial walls may show 1—15 small pits, the oblique pore being occasionally enclosed in a feebly developed border. In view of the entire absence of pits on the ray cells of at least some recent Araucarias the structure of the ray cells in fossil stems requires careful revision}. In the following brief descriptions of species of Dadozylon a few examples are chosen to illustrate the wide geological and geo- graphical range of fossil wood of this type, but it must be remem- bered that in many cases no positive statement can be made with regard to the nature of the parent-plant beyond the facts afforded by the anatomical characters of the stem. Evidence bearing on the geological age of the Araucarineae is discussed in the course of the description of genera founded on vegetative shoots and reproductive organs. Species of Dadoxylon from Carboniferous and Permian strata have already been described in Chapter xxXxXIII. as more probably referable to Cordaites or at least to the Cordai- tales, and it is not by any means impossible that some of the Dadoxylons recorded from Triassic or even higher strata may belong to Cordaitalean species rather than to the Araucarineae. | The evidence afforded by petrified wood in conjunction with that derived from vegetative remains lends probability to the view that Araucarian plants existed at least as early as the later Palaeozoic 1 See, in addition to Kraus and other authorities, Lignier (072). XLIv] . DADOXYLON : 181 age. The lack of satisfactory knowledge with regard to the mor- phology of the reproductive organs of such genera as Walchia, Voltzia, and other plants closely resembling living Araucarias in the habit of their foliage-shoots precludes any definite statement as to the precise degree of relationship between these and other types and existing Araucarineae, though it is certain that the Araucarineae were at least foreshadowed before the close of the Palaeozoic era. Dadoxylon keuperianum (Goepp.). This species!, from Franconia and Wiirtemberg, is considered by Schimper? on the ground of association to be the wood of a Voltzia, and Tuzson® adopts the generic name Pagiophyllites implying relationship with Pagiophyllum; he includes in D. keupervanum the species Araucarioxylon wiirtembergicum Kr. and A. thuringicum Born*. The tracheids of D. keuperianum have one or more rows of contiguous and more or less flattened pits; the medullary rays are uniseriate and 2—50 cells in depth; Tuzson figures 2—4 circular simple pits in the field. Though possibly belonging to Voltzia or Pagiophyllum this wood is best retained in Dadoxylon. Other Triassic Dadoxylons are described by Wherry® from Pennsylvania: he records Araucariorylon vir- ginianum, a species described by Knowlton® from Potomac beds, and A, vanartsdaleni: in both forms the tracheal pits are com- pressed and alternate, the rings of growth indistinct, and the medullary-ray cells are said to have no pits. Reference has already been made to a Triassic species from New Caledonia, D. australe (Crié). Dadoxylon septentrionale Gothan. \“ This species’, founded on material believed by Gothan to be ‘Triassic in age, has the following characters :—Annual rings often distinct macroscopically but microscopically showing little con- trast between spring- and summer-wood; bordered pits in a single row and separate or polygonal and in two alternate rows; they are often arranged in stellate clusters as in some species of Cedroxylon. 1 Goeppert (81) p. 42. ® Schimper (72) A. p. 384. 3 Tuzson (09), p. 30, fig. 5. 4 Schimper (72) A. p. 384. 5 Wherry (12). 8 Knowlton (892), Pl. vi. figs. 2—5. —7 Gothan (10) p. 8, Pl. t figs. 4—8, PL. 1. fig. 1. 182 : CONIFERALES [CH. Medullary rays uniseriate, reaching a depth of 30 cells, 2—4. elliptical oblique pits in the field; wood-parenchyma, often with dark contents, is not uncommon. Gothan emphasises the abund- ance of xylem-parenchyma as a character in which this species differs from typical Dadoxylons. Dadoxylon mahajambjense (Fliche). Fliche? described this species, from Liassic strata in Madagascar, under the name Aravcariorylon: the radial walls of the tracheids have two rows of contiguous and alternate pits; the medullary - rays are uniseriate, usually 8—16 cells deep, and small circular pits occur on the radial walls. Dadoxylon divescence (Lignier). An Oxfordian species from Normandy characterised by leaf- traces larger than those of Araucaria imbricata and, as Lignier? states, suggesting leaves comparable in size with those of Cordaites. The tracheids have 1—4 rows of pits and the medullary rays are 8—11 cells in depth. Dadoxylon argillicola (Kichwald), recorded from Moscow’, is another example of a similar type of wood, and many other in- stances might be quoted in illustration of the wide distribution of vegetative organs in Jurassic beds agreeing anatomically with the Araucarineae. Dadoxylon (Araucarioxylon) novae zeclandiae* (Stopes). A Cretaceous species from Amuri Bluff, New Zealand, founded ona piece of decorticated stem 8 cm. in diameter preserved partly in silica and in part in a calcareous medium. The small pith, not more than 1 mm. in diameter, is imperfectly petrified: the tracheids of the secondary xylem, which shows well-marked annual rings, have biseriate, alternate, hexagonal, pits; there is no xylem-paren- chyma. An interesting feature is the occurrence of tracheids on each side of the medullary rays with thicker walls and containing discs of resin: these resiniferous tracheids, similar to those de- scribed by Thomson® and other authors, afford particularly good Fliche (05) p. 350, Pl. x. fig. 1. Lignier (07?) p. 257, Pl. xvm. figs. 10—13. 3 Eichwald (68) Pl. v. fig. 12. The specific name is given by Dr Stopes as novae zcelandit. Stopes (14°) Pl. xx. 8 Thomson, R. B. (13). 1 2 4 5 XLIV] DADOXYLON 183 examples of this type of element. The uniseriate medullary rays are usually 3—4 cells deep and are described as having 5—6 bordered pits with oblique pores in the field. In view of Thomson’s conclusion with regard to the absence of pits on the ray cells in recent Araucarineae and considering the form of the pits as shown in Dr Stopes’ drawing it may be doubted whether the pits actually belong to the walls of the ray cells. Dr Stopes comments on the scarcity of Araucarian remains recorded from New Zealand and adds a brief account of some Tertiary wood described by Ettingshausen! as Araucaria Haasti, a species founded on foliage-shoots which, without any adequate reason, it is surmised belong to the petrified wood. The latter is poorly preserved: Dr Stopes examined the type-specimen and found that the bordered pits on the tracheids are circular and not compressed; she expresses some doubt as to its Araucarian affinity, but renames the species Araucariorylon Ettingshausent. A specimen of wood from Amuri described by Ettingshausen as Dammara Oweni® appears to be undoubtedly Araucarian. Dadozxylon sp. (Holden). An interesting type of stem-wood has been described by Miss Holden? as Araucarioxylon sp. from the Cretaceous lignites of Cliftwood, New Jersey, which shows Araucarian characters in com- bination with certain anatomical features not usually associated with Araucarioxzylon. The tracheids for the most part have alternate compressed pits; the medullary rays consist of thin- walled cells and there is no xylem-parenchyma. Near the inner edge of the wood the tracheids are characterised by bordered pits in opposite pairs and rims of Sanio occur between adjacent pairs. The specimen is said to supply the only missing link in the chain - of evidence pointing to the derivation of the Araucarineae from the Abietineae. Opposite pits are figured by Miss Holden in tra- cheids from the cone-axis of Araucaria Bidwillii and, as stated else- where, Sanio’s rims are not unknown in Araucarian wood. I have adopted the name Dadoxylon because the characters as a whole are consistent with that designation though it might be contended that a new name is desirable to indicate the occurrence of unusual 1 Ettingshausen (87) p. 154, Pls. rz. figs. 1, 2; v1. figs. 1O—12. 2 Ibid. p. 16, Pl. vi. figs. 13—15. 3 Holden, R. (137). % ya io { | 184 CONIFERALES [cH. features. The occurrence of opposite pits and rims of Sanio in the younger portion of the xylem is regarded as evidence in support of the view that the stem, while in the main Araucarian, exhibits features indicative of the origin of the Araucarineae from the Abietineae. The presence of opposite pits in wood in which the normal arrangement is alternate is not surprising if it is admitted that the Coniferous pitting is derived from an earlier scalariform type. Even in stems in which the alternate or opposite pitting is well established it is not very uncommon to find occasional departures from the normal pattern. This Cretaceous stem is one of many generalised types, and the arguments based on the admixture of characters in favour of the greater antiquity of the Abietineae do not present any insuperable difficulty to the opposite view namely that the Araucarineae preceded the other families of the Coniferales. Dadoxylon (Araucarioxylon) breveradiatum (Lignier). This species from the Cenomanian of Normandy? affords one of the few examples of the preservation of phloem, cortex, and pith. Lignier adopted the generic name Arauca- riocaulon: he describes the tracheids as having 1—3 rows of pits, crowded and alter- nate but not flattened. The medullary rays are very short, 1—3 and rarely 4 cells in depth; there are said to be 8—15 pits in the field (fig. 713). Resiniferous parenchyma is abundant as in the wood referred by Caspary to a special genus Araucariopsis® and in Fye.713. Dadoxylon (Ar- Dadoxylon septentrionale Goth. The phloem — @wearioxylon)breveradi- includes well-preserved sieve-tubes and the a ence, cortex is characterised by numerous sclerites, an Araucarian feature. This species appears to be an aberrant type the position of which is by no means clear. Among other Cretaceous species are Dadorylon albianum® (Fiche) from L’Aube; Dadoxylon Dantzii Pot.4 from beds pro- bably of Upper Cretaceous age in East Africa, without definite rings 1 Lignier (07?) p. 290, PL. xrx. 2 See page 179. 3 Fliche (97) p 8, figs. 2—4. 4 Potonié (02) Pls. 1, 12. XLIV] DADOXYLON 185 of growth and with a single row of contiguous pits on the tracheid walls; Dadozylon virginianum (Knowlton)! from the Potomac series; Dadoxylon barremianum (Fliche)? from the Lower Creta- ceous of France; Dadoxylon noveboracense (Holl. and Jeff.)* from the Middle Cretaceous beds of Staten Island; Dadoxylon Zuffardic Negri* from middle Cretaceous rocks in the Gulf of Tripoli; Dado- xylon tankoense (Stopes and Fujii)® from Upper Cretaceous beds in Japan; Dadoxylon madagascariense (Fliche)® from Madagascar. Dadoxylon (Araucarioxylon) kerguelense sp. nov. In 1881 Goeppert? mentioned some wood received from Kergue- len Island from Baron von Schleinitz, probably of Tertiary age, under the name Araucarites Schleinitzi et Hookeri, but, as Gothan® points out, it is not clear whether he refers to one or two species, and as there are no figures or full description Goeppert’s designation cannot stand. * 4 a * * * 3 * an ue * AED SREB Sy Py Fie. 714. Dadoxylon (Araucarioxylon) kerguelense. (British Museum.) The sections on which this Kerguelen species is founded are in the British Museum®; they show the following characters :— Annual rings narrow, often 15—20 tracheids broad, the summer wood being frequently represented by only two rows of elements. There are 1 or 2 rows of bordered pits on the radial walls of the 1 Knowlton (89?) Pl. vu. 2 Fliche (00) p. 18. 3 Hollick and Jeffrey (09) B. p. 58, Pl. xxr. 4 Negri (14). 5 Stopes and Fujii (10) Pl. ut. figs. 17, 18. 6 Fliche (002) p. 472, fig. 1.. 7 Goeppert (81). 8 Gothan (08) p. 13. ® Sections V. 8388, V. 8390. 186 CONIFERALES [cH. tracheids, contiguous, alternate, and often slightly flattened (fig. 714, B). The medullary rays have usually 5—8 elliptical pits in the field (fig. 714, A). Dadoxylon (Araucuriorylon) pseudoparenchymatosum Gothan. A species from Tertiary or possibly Upper Cretaceous rocks in Seymour Island (S. lat. 64° 16’)! agreeing closely with the wood of recent Araucarineae. The annual rings are distinct; there are 1—2 rows of pits on the tracheids 10—12y in diametér; the medullary rays, 2—10 cells deep, are usually uniseriate and there are several small oblique pits in the field. Cross-bars? (Miiller’s ‘querbalken’) like those described in Araucaria brasiliensis occur in some of the tracheids. Dadoxylon Doeringit Conwentz is a Patagonian species of Sub- Oligocene age characterised by distinct annual rings; rays up to 40 cells in depth, with 1—2 pits in the field. Among other Ter- tiary species are Dadoxylon aegyptiacum Unger*, recorded from several localities in the Libyan desert; Dadozxylon Robertianum (Schenk)> of Tertiary or possibly Cretaceous age from the pro- vince of Nagpur, India; Araucariorylon koreanum (Felix)® from Korea, characterised by the occurrence of a single row of con- tinuous pits on the tracheids, is referred by Gothan to the genus Xenoxylon and regarded as identical with X. latiporosum’. II. CUPRESSINOXYLON. Goeppert. The name Cupressinoxylon® or, as written by Kraus, Cupresso- xylon®, is usually applied to fossil wood exhibiting the following features :—Annual rings well defined, often narrow; vertical rows of parenchyma, often containing resin and recognisable by their dark contents even in transverse section (fig. 715, A), scattered through the spring- and summer-wood. Bordered pits on the tracheids usually separate and circular and if in more than one row opposite; medullary-ray cells generally characterised by the presence of several small pits in the field. Used in this sense 1 Gothan (08) p. 10, Pl. 1. figs. 12—16. 2 See page 135. 3 Conwentz (85) p. 16. 4 Unger (59); Schenk (80) p. 3, Pls. 1., 1. 5 Schenk (822). ® Felix (87) p. 518. ? Gothan (10) p. 23. * Goeppert (50) p. 196. ® Kraus in Schimper (72) A. p. 374. XLIV] CUPRESSINOXYLON 187 Cupressinoxylon denotes wood similar to Cedroxylon except in the greater abundance of xylem-parenchyma and its occurrence in the spring- as well as in the summer-wood. The medullary rays afford another distinction which according to some authors is more trustworthy than the presence or distribution of the xylem- parenchyma. Gothan1, who uses Cupressinorylon in a more restricted sense, lays stress on the pitting of the medullary cells as a distinctive feature: the pitting is confined to the radial walls, or in other words there is no Abietineous pitting in Cupressinoxylon in the stricter sense. The medullary-ray pits have a broadly elliptical pore which is more or less horizontal at least in the spring-tracheids—the Cupressoid type in contrast to the Podo- carpoid type in which in the spring-wood the pore is narrower and more vertical, though in some Podocarps the bordered pits are replaced by large simple pits: in the summer-wood the difference between the Cupressoid and Podocarpoid type disappears. In Cupressinoxylon the medullary rays are uniseriate and not very deep though the depth is a variable character. Lignier? states that 60—150 rays occur in 1 square millimetre, another feature _ of doubtful value. In some species included in Cupressinorylon the pitting of the tracheids is partly Araucarian as it also is in certain types of Cedroxylon. The presence of Sanio’s rims, though not mentioned by many authors, is regarded by Jeffrey and some other American botanists as an important character to be expected in all Coniferous wood other than that of the Araucarineae, It is clear that unusually good preservation is essential for the recognition of such features of the medullary-ray cells as Gothan includes in his definition of the genus; unless the tissues are well preserved the generic separation of Coniferous types except within very wide limits is impossible. The name Cupressinoxylon may conveniently be restricted to wood having | the usual type of tracheal pitting though pits of the Araucarian type may occur locally, with medullary rays in which the pitting | is confined to the radial walls and generally with several fairly | small and apparently more or less definitely bordered -pits in the | field, the pores in the spring-wood being elliptical and more or | less horizontal. Resin-canals absent except in wounded regions 1 Gothan (05) p. 39. 2 Lignier (07?) p. 245. ! 188 CONIFERALES [cH. of the wood; xylem-parenchyma normally abundant and not restricted to the end of the year’s growth. The photographs Fic. 715. Cupressinoxylon sp. Lough Neagh, Treland; p, xylem-parenchyma. (British Museum; A. V. 8248, B. V. 8253, C. V. 8257.) reproduced in fig. 715 of sections of a piece of petrified wood from Lough Neagh?in Ireland, probably of Upper Tertiary age, illustrate 1 See vol. 1. p. 80. XLIV] CUPRESSINOXYLON 189 very clearly the appearance of xylem- or resin-parenchyma as seen in transverse, radial and tangential section. The presence of dark contents renders these cells conspicuous in transverse section (A) and both contents and cross-walls are seen in the longi- tudinal sections (B,C). In fig. 715, B, the opposite and scattered bordered pits are shown on the radial walls of the tracheids with an occasional tendency to an alternate arrangement. Goeppert founded his species Pinites Pritchardti} on wood from Lough Neagh,- the generic name being altered by Kraus to Cupressinoxylon?. Kraus® enumerates 46 recent species of Conifers with wood of the Cupressinoxylon type and Beust* mentions 37 species. As used by most authors Cupressinoxylon includes members of the Cupressineae (in the wide sense as used on page 124), Podocar- pineae, Abies Webbiana, and some other recent genera. As already stated Gothan has essayed the difficult task of defining the genus in such a way as to restrict the wood so named to a smaller number of recent Conifers, recognising as distinct genera certain species previously included in Cupressinoxylon, e.g., Taxodioxylon (= Cupressinoxylon Sequoianum Merck.*}, Podocarpoxylon (= C. Hookert Arb.°), Xenoxylon (= Cupressinoxylon Barberi Sew.’): these and other species formerly included in Cupressinoxylon are distributed among genera not always well defined but instituted with a view to increase the value of fossil species considered in - relationship to recent: types. [Paracupressinoxylon. Holden.] Miss Holden® has proposed a new generic name, Paracupressi- noxylon for wood which agrees with Cupressinoxylon in having xylem-parenchyma scattered through the year’s growth but differing in the Abietineous pitting of the medullary-ray cells and in the absence of Sanio’s rims. In Paracupressinoxylon are in- cluded two species, P. cedroides and P. cupressoides, both Jurassic and the latter also Cretaceous in age, which appear to differ too much from one another to be referred to one genus. Both species 1 Goeppert (50) p. 220. 2 Kraus in Schimper and Schenk (72) A. p. 376. 3 Kraus (64). 4 Beust (85). . 3 Mercklin (55) Pl. xvi. © 6 Arber (04); Gothan (08) p. 7. 7 Seward (04) B. p. 60, Pl. vit. 8 Holden, R. (132) p. 5387; (14) p. 173. 190 . CONIFERALES [cH. are regarded by the author of the genus as undoubted repre- sentatives of the Araucarineae, a determination that is hardly con- sistent with the affinity implied by the generic name. The species P. cupressoides, from Yorkshire and the Cretaceous lignites of New Jersey, is characterised by the restriction of pits to the radial wall of the, medullary-ray cells, the absence of Sanio’s rims, the presence of scattered xylem-parenchyma, and by the occurrence . both of scattered and crowded pits on the tracheal walls. More- over in this species the phloem shows an alternation of hard and soft elements. The affinity suggested by these features would seem to be to Cupressinoxylon. On the other hand, Paracupres- sinoxylon cedroides, founded on material from the Yorkshire coast, is characterised by the Abietineous pitting of the medullary-ray cells, an admixture of scattered and compressed bordered pits on the radial walls of the tracheids, scattered xylem-parenchyma, no alternation of hard and soft bast, and by the absence of any sclerous cells in the pith. This species also illustrates the occur- rence of resin-canals in wounded regions of the wood. The refer- ence of both these species to the Araucarineae, chiefly because of the absence of Sanio’s rims, though consistent with the principle that this character is ail important, implies the neglect of other chaiacters, more especially the nature of the medullary-ray pitting, which in the case of recent Conifers are unquestionably of taxo- nomic importance. The species P. cedroides should not, in my opinion, be included with P. cupressoides in one genus; it is pro- bably more closely allied to the Abietineae than to any other family. It should, however, be remembered that pitting of the horizontal and tangential walls of medullary-ray cells is a feature that is not confined to the Abietineae; it occurs also in some recent Junipers and the extinct genera Protocedrozylon and Thylloxylon. Cupressinoxylon Nasinum Lignier. This Liassic species! from Orne, France, is founded on the wood of a pentarch and hexarch root: the bordered pits on the radial walls of the tracheids are usually in one row, occasionally in two opposite series; smaller pits occur on the tangential walls; medul- lary rays 1—5 cells deep with a few ovoid-oblong pits, often simple Lignier (07?) p. 306, Pl. xxz. figs. 58—61; Pl. xxu. fig. 83. XLIv] CUPRESSINOXYLON 191 and rarely bordered, in the field. Resin-cells are abundant. Lignier suggests that if the genus Cupressinozylon is subdivided this species might be referred to Glyptostroborylon, though the medullary-ray pitting is not consistent with the characters of that genus. Cupressinoxylon vectense Barber. Founded on both stem and root wood from the Lower Green- sand of the Isle of Wight and described by Barber! with a thorough- Fic. 716. Cupressinoxylon vectense. Transverse section of branch-wood showing a compound ring. (After Barber.) ness of detail rarely met with in accounts of petrified wood. Dr Stopes? has recently re-described this species, adding a figure of the medullary-ray pitting. Annual rings well marked, often illustrat- ing the occurrence of compound rings of growth (fig. 716) which are discussed in detail in the original account; bordered pits in a 1 Barber (98) Pls. xx11.—xxIv. , 2 Stopes (15) p. 169, text-fig. 50; also text-figs. 48, 49, and Pl. xv. 192 CONIFERALES [CH. single row, rarely double in the roots, free and circular in branches, often contiguous and compressed in roots ; tangential pits common. Medullary rays usually uniseriate, 1—16 cells deep, pits confined to the radial walls, usually 1 but sometimes 2—4 oval and oblique pits in the field. Resin-canals absent; resin-parenchyma in vertical rows, abundant and scattered. The pith consists of pitted parenchymatous cells separated by intercellular spaces ; in the roots the rows of tracheids pass directly into the cells of the pith; in the branches they terminate in small groups of cells irregularly arranged. Cupressinoxylon McGee Knowlton. This is one of several species from the Potomac lignites included by Knowlton? in Cupressinoxylon. The annual rings are well marked ; the tracheids have 2—3 rows of opposite and circular pits on the radial walls and small bordered pits are abundant on the tangential walls. The uniseriate medullary rays, 2—49 cells deep, have 1—2 oval apparently simple pits in the field and resin- parenchyma is abundant. Gothan? has described some wood of Lower Cretaceous age® from King Charles Land as Cupressinoxylon, cf. C. McGeet, agreeing with Knowlton’s type in the medullary-ray pitting; there are 2—4 simple pits in the field, elliptical and hori- zontal; an indication of a border was seen in some of the pits in the region of the summer-wood, but the general absence ofa border, if an original feature, is a difference between this wood and that of recent genera included in Cupressinoxylon. There is no Abie- tineous pitting on the ray cells. The species Cupressinoxylon luccombense described by Dr Stopes? from the Lower Greensand of the Isle of Wight closely resembles C. vectense, but it has stone-cells in the pith, the tracheids are larger and there are usually 3—4 pits in the field in place of 1 or sometimes 2—3 in C. vectense; moreover in the latter species the pits of the ray cells are more uniform in size. * Knowlton (89?) p. 46, Pl. 1. fig. 5; Pl. rz. figs. 1—5. 2 Gothan (07%) p. 19, fig. 10. $ Gothan speaks of the King Charles Land fossils as Jurassic, but the beds have since been shown to belong to the Cretaceous system. See Burckhardt (11). + Stopes (15) p. 180, text-figs. 51—53. XLIV] CUPRESSINOXYLON 193 Cupressinoxylon cryptomerioides Stopes. A species! founded on a small branch from the Lower Green- sand of Kent showing the following features:—the primary xylem, composed of spiral and scalariform elements, also tracheids Fic. 717. Cupressinoxylon Hortii. Radial section showing the pits of the medullary rays, mp; the pits on the tracheids, and the wood-parenchyma, rp. (After Stopes.) with small circular pits, forms projecting wedges in the perimedul- lary region; there are no resin-canals and resiniferous parenchyma is abundant; the medullary rays, generally 2—3 cells in depth, have two circular pits in the field, a feature regarded by Kraus as indicative of affinity to Glyptostrobus?. The cortex contains large 1 Stopes (15) p. 186, Pls. xvi., xvur., text-figs. 54, 55. 2 See page 198. 194 CONIFERALES [CH. resin-canals. Dr Stopes draws attention to certain features, particularly the pitting of the medullary-ray cells and the structure of the cortex, in which this species resembles the genus Crypto- meria. Cupressinorylon Hortii Stopes. This Lower Greensand species! from Bedfordshire is distin- ~ guished from other types by the very numerous medullary rays, often separated from one another by a single row of tracheids, which are both uniseriate and multiseriate, the same ray exhibiting both forms at different heights. The rays attain a depth of 80 cells and there is generally one large oval or circular pit in the field (fig. 717) though two are occasionally present. This species is hardly a typical Cupressinoxylon and Dr Stopes points out that the large single pit in the field is suggestive of Podocarpoxylon: it affords another illustration of the impossibility of identifying the majority of fossil woods within narrow limits. Cupressinoxylon Holdenae sp. nov. A species of Eocene age from the London Clay of Faversham in the Cambridge Botany School collection characterised by well- -defined annual rings and the presence in some but not all the bands of summer-wood of resin-canals (fig. 718, C) which vary in size, some of the larger being formed by the confluence of smaller adjacent canals. The presence of rather thick-walled, pitted, cells lining the canals or in close association with them is a characteristic feature. Tyloses occur in some of the canals. The large number of canals in each row suggests their development in response to traumatic stimuli. The bordered pits occur in single, separate, rows or in double and opposite rows with rims of Sanio occasionally preserved. In a few places the pits of a single row are in contact and slightly flattened. Resin-parenchyma occurs in vertical rows in both spring- and summer-wood. Medullary rays uniseriate, reach- ing occasionally 30 cells in depth; the tangential and horizontal walls are unpitted and on the radial walls the pits are preserved only in a few places; there appear to be 2—4 fairly large simple pits in the field. The crowded series of canals (fig. 718, C) are identical with the traumatic ducts described in Sequoia sempervirens? 1 Stopes (15) p. 194, Pl. xv. text-figs. 56—58, ® Jeffrey (03). XLIV] CUPRESSINOXYLON 195 and Abies (cf. fig. 690, B, p. 130). The absence of Abietineous pitting in the ray cells, the distribution of the canals, and the presence of scattered rows of xylem-parenchyma are features indicating affinity to Sequoia sempervirens. The preservation is not sufficiently good to warrant any definite statement with regard to the pits on the radial walls of the ray cells: the absence of a border is in contrast to the pits in Sequoia, but the apparent lack of a border may be due to imperfect preservation or to decay. Miss Holden, who carefully examined the sections, called my atten- tion to the occasional occurrence of obscure and narrow cells of unequal breadth on the edge of some medullary rays bearing a resemblance to the ghost-like ray-tracheids described by Thome: sont. Cupressinozylon Koettlitzi sp. nov. Silicified wood is by no means uncommon in the Franz Josef Archipelago and several specimens have been found in talus-heaps - and in basaltic lavas. The age may be Upper Jurassic, Cretaceous, or possibly Tertiary. A radial longitudinal section of a piece of wood is figured, though not named, by Newton and Teall? and without a full description. The following account is based on sections cut from the same block in the possession of the Geological Survey, which was collected at Northbrook Island by members of the Jackson-Harmsworth Expedition. The species is named after Dr Koettlitz, the geologist of the Expedition’. Annual rings narrow and distinct: there are no resin-canals and no clearly preserved xylem-parenchyma, though in a few places there are indications of what appear to be elongated cells containing a dark substance. Itis noteworthy that in some recent Cupressineae resin- parenchyma is not invariably present. The bordered pits on the radial walls of the tracheids are variable in their arrangement; they occur in single rows (fig. 718, E), contiguous and sometimes slightly flattened or more or less widely scattered, also in double rows with an opposite or occasionally. an alternate disposition. There are a few pits on the tangential walls of the tracheids and rims of Sanio are seen in places on the radial faces. The medullary rays are 1—25 cells deep, uniseriate and. very rarely two-cells 1 Thompson (12). 2 Newton and Teall-(97) p. 508, Pl. xx. fig. ll. 3 Koettlitz (98). 13—2 196 _ CONIFERALES [CH. broad; 2—4 small circular or oval (simple?) pits in the field, but none on the tangential or horizontal walls. ) Yor Be c| Fic. 718. A, D, Cupressinoxylon polyommatum. B, Phloem showing sieve-plates (Franz Josef Land). C, Cupressinoxylon Holdenae. E, Cupressinoxylon Koett- litzi. (A, D, Dublin Museum; B, E, Geological Survey Museum, Jermyn Street; C, Cambridge Botany School.) Cupressinoxylon polyommatum Cramer. The petrified wood on which this species! was founded was obtained from Banksland, from Miocene rocks, N. lat. 74° 40’, 1 Cramer (68) p. 172, Pls. XXXIV., XXXV., XXXVII. XLIv] CUPRESSINOXYLON 197 long. 122° W., during the voyage of H.M.S. Investigator in 1851. Through the kindness of Prof. Johnson I was able to have sections cut from Dr Cramer’s specimen in the Dublin Museum. Annual rings clearly marked; summer-wood narrow. The most striking feature is the irregular distribution and unusually large number of small bordered pits on the radial walls of the tracheids. The pits vary in size and have an average diameter of 13-77; they are usually separate but occasionally those of a single row are in contact. There are frequently as many as 3—4 rows (fig. 718, A) and occasionally 5 opposite and separate pits, a feature suggesting comparison with Pinus Merkusit1, but in that species the pits of the three opposite rows are in contact. Rims of Sanio’are occasionally present. Rows of narrow parenchyma occur in different regions of the wood. Medullary rays uniseriate, 1—16 cells deep, with 2—4 simple large oval pits in the field (fig. 718, D); there are no pits on the other walls of the ray cells. The pitting in the field is very similar to that in Taxodium and: Cupressinoxylon (Taxodioxylon) Taxodia (fig. 720, A, B). Cupressinoxylon taxodioides Conwentz. Under this name Conwentz? describes some wood, tohabiy of Pliocene age, from California which he compares with Sequoia sempervirens: He speaks of one stem 22 metres long and with a maximum diameter of 3—4 m.; the bordered pits are in 1—2 rows on the radial walls and small pits occur on the tangential walls; the medullary rays are usually two-cells broad and have 3—4 generally elliptical pits in the field apparently simple and arranged in a horizontal row. The rays are usually 15—20 cells deep but may reach a depth of 56 cells. Resin-parenchyma occurs in vertical rows but it is not stated whether it is confined to any definite region of the wood. Schmalhausen’s species Cupressinoxylon (Glyptostrobus?) neost- biricum3, characterised by medullary rays 13—20 or even 40—48 cells deep, and 1—2 circular or oval pits in the field, though com- pared by him with Glyptostrobus, cannot safely be regarded as more nearly allied to that genus than to some other members of the Cupressineae. 1 Groom and Rushton (13) p. 484. 2 Conwentz (78) Pls. xmt., xiv. 3 Schmalhausen (90) Pl. u. pp. 44—49; Gothan (05) p. 50. 198 CONIFERALES [cH. Many of the species included by authors are not described in sufficient detail to satisfy modern requirements with regard to the structure of the medullary rays and other characters. From the point of view of geographical distribution reference may be made to Cupressinoxylon antarcticum described by Beust1 from Kerguelen Land. [Glyptostroboxylon Conwentz. ] This generic name was first employed by Conwentz® for wood from sub-Oligocene beds in Argentina which he described as Glyptostroboxylon Goepperti: no figures accompany the description and it is hardly possible to determine with accuracy the precise affinity of the specimen. The annual rings are said to be distinct, the pits on the tracheids uniseriate and contiguous; resin-paren- chyma occurs, and the medullary-ray cells have large circular pits on the radial walls. It is suggested by Gothan® that this species should be transferred to Podocarpozylon, though in the medullary- “ray pitting it differs from typical representatives of the Podocar- pineae. Kraus* described the pits in the radial walls of the ray cells of Glyptostrobus as large (©) S and circular and in a Tertiary specimen from Niederwéllstadt, named Glyptostrobus tener Kr., te) (=) he figures the pits in the field as simple (fig. = 719); these are said to be 1—8 in number and |{a|| OO}O° they are arranged in horizontal series, a feature oor St characteristic of Taxodium. rs) ra} ; ool§8 Sections of wood of Glyptostrobus hetero- ||2 OO} phyllus which I have examined show 2—8 pits in the field but there is a distinct border and the "0-719. Cupressin- i i i . oxylon tener (Kraus). pore is narrow and oblique or in the spring-wood (After Kraus.) the pore may be broader and almost horizontal. Xylem-parenchyma is scattered through the wood and the thick transverse walls are pitted. Gothan believes that the medullary-ray pitting of Glyptostrobus is distinguishable from the Cupressoid type by the increase in the breadth of the pore as the ray cells are followed from the summer- to the spring-wood and by the fact 1 Beust (85) Pl. rv. 2 Conwentz (85) p. 13. 3 Gothan (08) p. 9. 4 Kraus (64) p. 195, Pl. v. fig. 12. XLIv] CUPRESSINOXYLON 199 that the pits in the region of the spring-wood may be simple. The same author! points out that this type of medullary-ray pitting occurs in Cunninghamia. The impression produced by an examination of the recent Glyptostrobus is that this so-called Glyptostroboid pitting is not a sufficiently well-defined ‘type to serve as a trustworthy diagnostic character. The medullary-ray pitting in the region of the summer-wood is similar to that in some species of Podocarpoxylon (= Mesembrioxylon), while in the spring- wood the pits in the field are rather of the type associated with Cupressinoxylon and the scattered xylem-parenchyma is another characteristic of the latter genus. There would seem to be no adequate grounds for regarding the two fossil species referred to Glyptostroboxylon as more nearly related to Glyptostrobus than to certain other recent genera. The retention of the name Glypto- stroboxylon is inadvisable in that it implies an affinity which is not supported by satisfactory evidence. CUPRESSINOXYLON, sub-genus TAXODIOXYLON Felix. The generic name Tazodiorylon was applied to a Tertiary species from Hungary, originally referred by Felix? to Rhizotaxo- _ dioxylon, on the ground of a resemblance in structure to the wood of the recent genus Taxodium. Schenk®, who examined the type- specimen, confirmed this comparison. Tazxodioxylon has . been adopted for fossils agreeing with the wood of Tazodiwm and Sequoia sempervirens: Sequoia gigantea, on the other hand, agrees more closely with typical species of Cupressinoxylon. Taxodio- zylon, while similar in most respects to Cupressinoxylon, is said to differ in the medullary-ray pitting, the pits in the field being almost simple and elliptical with their long axis horizontal in con- trast to the more definitely bordered pits of the Cupressoid type. This distinction is, however, not entirely satisfactory: in the wood of Taxodium the pits in the field are rather large and, though often simple, they occasionally present the appearance of pits with a well-developed border and the pore may be almost vertical or horizontal. In the recent species, as in some fossil examples, the tendency of the pits to arrange themselves in one or two 1 Gothan (05) p. 49. 2 Felix (84) p. 38. ? Schimper and Schenk (90) A. p. 872. 200 CONIFERALES [CH. horizontal rows is a characteristic feature. The characters of Taxodiorylon may be summarised as follows:—Annual rings distinct; bordered pits on the radial walls of the tracheids in 1—4 rows, circular and separate and, if in two or three rows, opposite ; rims of Sanio present. Medullary-ray cells pitted only on the radial walls; pits in the field, 2—8 in number, often arranged in horizontal rows (fig. 720, A), sometimes fairly large, simple, or bordered and horizontally elliptical. The thick walls of the xylem- parenchyma (fig. 720, B) are characteristic of Taxodium and Tazodioxylon and this character has been quoted as a trustworthy ‘S) OD Cl EO I] @ Col] © 4 eee in 0) [jeS SUE = © [a&pfooo SeS|l SS_|] @ on met Ss) (ssl (0) tooo] P=! ae q | 6 f prams ao 9 9 ees 8 | T aa A B Cc Fic. 720. A, B, Cupressinoxylon (Taxodioxylon) Taxodii. C, Cupressinoxylon (Taxo- dioxylon) Sequoianum. (A, B, after Gothan; C, after Mercklin.) distinction between Taxodium and Sequoia sempervirens though, as already pointed out, this is not a safe test?. In Gilyptostrobus heterophyllus the transverse walls of the xylem-parenchyma are also thick and pitted and this tissue in some species of Cupressus exhibits precisely similar features. It is in certain inconstant features that Taxodioxylon differs from Cupressinoxylon. In several instances the occurrence of wood referred to Taxodioxylon in beds containing impressions of foliage-shoots like those of Taxodium and Sequoia sempervirens strengthens the conclusions based on anatomical characters. 1 See page 149. XLIV] CUPRESSINOXYLON 201 Cupressinoxylon (Taxodioxylon) Taxodii Gothan. In his description of the Tertiary beds at Senftenberg Potonié! identified some of the wood as Tazodium and compared the de- posits with those of a Taxodium-swamp. Gothan® has given a fuller description of the wood, which is characterised by xylem- parenchyma with thick transverse walls (fig. 720, B) and fairly large elliptical pits in the field; in the region of the spring-wood the medullary-ray pits have a horizontal pore and in the late wood the pore is said to be vertical. The bordered pits on the vertical walls of the xylem-parenchyma have a vertical pore in the summer- wood but it is horizontal in the spring-wood. Cupressinozylon (Taxodioxylon) Sequotanum Mercklin. Gothan? recognised thisspecies, originally described by Mercklin? from an unknown locality, in Tertiary beds in Germany associated with foliage-shoots agreeing with Sequoia sempervirens. The walls of the xylem-parenchyma are thinner than in C. Tazxodii; there are 2—7 apparently simple pits in the field (fig. 720, C) in one or two horizontal rows. There may be three rows of opposite pits on the radial walls of the tracheids. It may be that C’. Sequoianum is the wood of a Sequoia. A closely allied species is represented by Cupressinozylon uniradiatum Goepp.? from Briihl afterwards recorded by Conwentz® as Rhizocupressinoxylon from Tertiary beds in Silesia. Schmalhausen’ has described some interesting speci- mens of wood from Tertiary beds in Russia as Cupressinoxylon Sequoianum characterised by the possession of elliptical simple pits in the ray cells (from 2 to 6 in the field) of the transversely elongated form characteristic of Taxodioxylon. Xylem-paren- chyma is fairly abundant and the tracheids are peculiar in having three or rarely four bordered pits on the walls as in some recent Pines. Another example of wood of the Tazodioxylon type is afforded by T. palustre described by Felix® from Tertiary rocks in Hungary and recorded also from sub-Oligocene beds in Silesia. Gothan® 1 Potonié (96); see also Eberdt (94). 2 Gothan (06) p. 164. 3 [bid. p. 165; (09) p. 518. 4 Mercklin (55) p. 65, Pl. xvi. 5 Goeppert (50) p. 203, Pl. xxvu. ® Conwentz (80) A. p. 25, Pls. 1v., v. 7 Schmalhausen (83) Pl. x11. 8 Felix (82) p. 278; (84) p. 38 ® Gothan (10) pp. 40, 43, PI. vir. 202 CONIFERALES [cH. assigns to Taaodioxylon two specimens from Tertiary strata in Spitzbergen and it is probable that the wood may belong to plants which bore the twigs described by authors as Taxodium distichum and Sequoia Langsdorfit. The name Taxodiozylon is retained as a section of Cupressino- zylon and not as a separate genus on the ground that the characters on which it is based do not appear to be sufficiently distinctive or constant to warrant its recognition as a well-defined generic type. II. TAXOXYLON. Unger. Unger! gave this name to fossil wood characterised by the presence of spiral bands in the secondary tracheids, a feature especially associated with the recent genera Taxus, Torreya, and Cephalotaxus, but by no means unknown in other Conifers®. The name, in the form Tazorylum, was substituted for Tazxites em- ployed by Goeppert*® for some species of Tertiary wood. Apart from the presence of spiral bands Tazxoxylon agrees with Cupres- sinoxylon, though according to Lignier the medullary rays are deeper in the latter genus. Taxoxylon scalariforme (Goeppert). This Tertiary species, originally described by Goeppert from Hungary as Tavites scalariformis, was renamed by Unger Tazorylum Goepperti. According to Schenk‘ it is the only species among those recorded by Goeppert which should be retained in Taxozylon, the spiral pattern of the tracheids being due to the presence of true bands and not, as in the other species, the result of enzyme action on the wood which produces a spiral striation closely simulating spiral bands. The bordered pits on the tracheids are circular and separate; the medullary rays are uniseriate and from 1 to 10 cells in depth; there are no resin-canals. Goeppert’s species Taaxites Ayckii (after Herr Aycke) of Ter- tiary age was retained by Kraus® as an example of that genus but afterwards transferred to Cupressinoxylon on the ground of the absence of true spiral bands in the tracheids. Lingelsheim® also 1 Unger (47) p. 33. 2 See page 134. 3 Goeppert (40). 4 Schimper and Schenk (90) A. p. 859. > Goeppert (40) p. 188; (50) p. 244; Kraus (64) p. 197. ® Lingelsheim (08) p. 27. XLIv | MESEMBRIOXYLON . 203 states that there are no true spiral bands in Geoppert’s supposed Taxoxylon. The Permian species Taxoxylon ginkgoides Ren.t and Grand’Eury’s Upper Carboniferous species J. stephanense? are probably founded on wood of the Araucarian type in which the tracheids show spiral striation. An Aptian species Taxoxylon anglicum Stopes is referred to that genus because of the occurrence of a spiral marking on the tracheids which the author of the species believes to indicate the presence of true spiral bands in the wood of recent Taxaceae, and because of the groups of 3—4 bordered pits in the fields of the medullary rays. There are no resin-canals in the wood ; the tracheids have a single row of circular pits on the radial walls and occasional rims of Sanio are preserved. There are 1—6 pits in the field and the presence of a border is regarded by Dr Stopes as an argument in favour of the tracheal nature of some of the medullary-ray elements, though the appearance of the cells does not afford any substantial ground for interpreting them as other than parenchymatous elements. An examination of the type-specimens convinced me that the spiral markings on the tracheids are not true bands like those of recent Taxineous wood and the pitting of the medullary-ray cells is in itself by no means a trustworthy criterion. There are, I venture to think, no good reasons for referring this wood to the genus Taxoxylon. IV. MESEMBRIOXYLON. Gen. nov. This generic name* is proposed for fossil wood exhibiting certain features associated with several recent genera which have a southern distribution. It is intended to replace Gothan’s two genera Podocarpoxylon and Phyllocladoxylon, types differing from one another in features which, as Dr Stopes® points out, are too inconstant to justify the retention of both designations. Moreover the use of Gothan’s names implies affinities to recent genera which there are no adequate reasons for assuming. In this instance, as ‘in many others, the anatomical characters do not enable us to 1 Renault (85) p. 163. 2 Grand’Eury (90) A. p. 317; Gothan (05) p. 68. “3 Stopes (15) p. 204, Pl. x1x. text-fig. 59. + 4 peonuBpwvds, southern. 5 Stopes (15) p. 210. 204 . CONIFERALES - [oH: assign fossil species to a position within the Coniferales sufficiently definite to be denoted by the use of a name implying close rela- tionship to a particular genus as distinct from a group of allied types. Podocarpoxylon. This name! has been applied to wood agreeing in structure with recent species of Podocarpus and Dacrydium more closely than with other Conifers. As generally understood the genus stands for wood without resin-canals, possessing xylem- parenchyma not necessarily confined to a particular region of the year’s growth. In Podocarpoxylon aparenchymatosum Goth. xylem-parenchyma is absent. The bordered pits on the tracheids are in 1—2 rows and, if in two series, the pits are opposite or sub- opposite (Podocarpoxylon Schwendae Kub.); rims of Sanio are present. There is no Abietineous pitting in the ray cells; the pits in the field are typically Podocarpoid, that is there are few in the field and these appear to be bordered and characterised by an elliptical or linear pore which is oblique or more or less vertical. The medullary-ray pitting next the summer-wood does not afford a satisfactory means of separating Podocarpozylon and Cupressino- zylon, but in the region of the spring-elements the Podocarpoid type is a distinguishing feature of Podocarpoxylon, though as stated in the account of recent Conifers the position of the pore is by no means a constant character. On the other hand, the pits in the field may be large and simple as in Sciadopitys, some species of Podocarpus, in Phyllocladoxylon, Xenoxylon and some other genera. Phyllocladozylon. This name was given by Gothan? to wood similar to Podocarpoxylon but differing chiefly in the occurrence of large, simple pits in the field [‘Eiporen’], a feature shared with Sciadopitys and some species of Podocarpus, by Microcachrys, Dacrydium, and Pherosphaera. The tracheids have 1—2 rows of bordered pits on the radial walls, scattered and circular, but not infrequently contiguous and flattened, and if in two rows they may be alternate. There are no clear indications of Sanio’s rims in the specimens figured by Gothan and Schenk. There are no resin- canals and no xylem-parenchyma. Phyllocladoxylon agrees closely with Xenoxylon®, but in Phyllocladoxylon the tracheal pits are often 1 Gothan (05) p. 48 2 Ibid. p. 55; (10) p. 37. 3 Sée page 238. XLIV] MESEMBRIOXYLON ; 205 separate and are smaller than in Xenozylon. The pits on the medullary-ray cells are also smaller, though it is doubtful if this is a constant character. Miss Holden? has instituted a new generic name, Paraphyllocladoxylon, for two specimens of wood from Jurassic rocks on the Yorkshire coast which do not appear to differ from Mesembrioxylon in any respect calling for generic recognition. In Paraphyllocladoxylon eboracense, from the Oolite of Scarborough, the tracheids have usually scattered and circular pits on the radial walls and pits are also abundant on the tangential walls: in Paraphyllocladoxylon araucartoides the pits on the radial walls are always closely compressed and flattened. There is no Abietineous pitting and there may be one or occasionally two large simple pores in the field like those in Xenoxylon and Mesem- briozylon (= Phyllocladoxylon of Gothan) but smaller than those of Xenoxylon. Xylem-parenchyma is absent, but some tracheids have apparent cross-walls that are believed to be resin-plates. Miss Holden recognises the close resemblance of her species to Gothan’s Phyllocladexylon, but a new name is employed on the ground that the absence of Sanio’s rims shows that the wood of the Yorkshire plants is Araucarian. The absence of Sanio’s rims cannot be confidently regarded as an original feature and, assuming - this negative character to be a real one, it does not differentiate the specimens from those described by Gothan; Gothan’s figures afford no evidence of the presence of Sanio’s rims in his species of Phyllocladoxylon. If the Yorkshire stems are Araucarian so too are those from King Charles Land and Seymour Island?. In one of Miss Holden’s species the tracheal pitting is not of the Araucarian type, while in the other it is Araucarian ; the pitting of the medullary rays is opposed to an affinity to any recent Araucarian Conifer. Both of the Yorkshire species are therefore transferred to Mesem- brioxylon: their anatomical characters indicate that they are generalised types which cannot legitimately be included in any family based solely on existing Conifers. While recognising that it is not always easy to draw a definite distinction between Xenoxylon and Mesembrioxylon the two names may be conveniently retained, the former being used in a much more restricted sense than the latter. Mesembriozylon is applied to woods in which the general 1 Holden, R. (13?) p. 536, Pl. xxxrx. figs. 7—10. 2 Gothan (077); (08). 206 ; CONIFERALES [CH. _ features are similar to those associated with Cupressinozylon, but the xylem-parenchyma may not be always present and the medul- lary-ray cells have one or two large simple pits, or two or more smaller bordered pits, in the field, the pore being rather vertical than horizontal. Mesembrioxylon undoubtedly includes species which if additional data were available would be assigned to ‘distinct genera. Apart from the probability that anatomical characters were even less restricted in their range through different types in former periods than they are in existing genera, the impossibility of discriminating between certain closely allied recent Conifers points to the advisability of employing designations for fossil woods in a wide sense and thus avoiding the danger of misleading students in search of material on which to base con- clusions with regard to the relative antiquity of existing genera. Mesembriozylon sp. (=? Podocarpoxylon sp. Gothan). This wood from Bathonian rocks in Russian Poland? affords an example of the difficulty of distinguishing clearly between Podocarpoxrylon and Glyptostroborylon: the bordered pits on the radial walls of the tracheids are separate or contiguous and slightly flattened; xylem-parenchyma occasionally occurs; there are usually two fairly large circular simple pits in the field (fig. 722, C). The systematic position of this wood cannot be regarded as — well established. Miss Holden? has recently described two specimens from the Jurassic beds on the Yorkshire coast as Podocarpoxylon sp. but the evidence in support of affinity to the Podocarpineae is not by any means conclusive. In one specimen there is no xylem- parenchyma and in the other parenchyma occurs at the end of the year’s growth. The pits on the medullary rays are described as piciform, 1—2 in the field. The anatomical features described hardly afford adequate reasons for assigning the wood to Podo- carpozylon rather than to Cupressinoxylon. ? Mesembrioxylon sp. (Thomas). A specimen of imperfectly preserved wood from Jurassic rocks in the Izium district, in South Russia, described by Thomas? as 1 Gothan (06?) p. 456, fig. 5. * Holden, R. (13?) p. 542, Pl. xx. figs. 31, 32. 3 Thomas, H. H. (11) p. 80, Pl. v. figs. 5—7. XLIv | MESEMBRIOXYLON 207 Phyllocladoxylon sp. may perhaps be included in Mesembrioxylon though the preservation is hardly sufficiently good to admit of accurate determination. The bordered pits on the tracheids are circular and separate, about 15 in diameter; the medullary rays have one or rarely two large simple pits in the field. Mesembrioxylon woburnense (Stopes). An Aptian (Lower Greensand) species from Bedfordshire founded on two blocks of secondary wood and referred to Podo- carpoxylon1, The tracheids have 1—2 rows of bordered pits, the pits in two series being opposite; Sanio’s rims are present. Resin- parenchyma is abundant all through the wood; the medullary rays are for the most part 3 cells deep but vary from 1 to 25; there is one large circular or oval pit, or sometimes two, in the field, and a narrow border is occasionally preserved. This species is near to M. Schwendae but there are fewer pits in the field in the English type. Mesembrioxylon bedfordense (Stopes). This Aptian species” is especially characterised by the arrange- ment of the bordered pits on the radial walls of the tracheids; the pits are uniseriate and occur in chains of 3—10, the border being flattened above and below by contact (fig. 721, t): the narrower parts of the xylem-elements are often without pits. Xylem- parenchyma is scattered through the wood and the medullary-ray cells have an oval or nearly circular large pit, sometimes with a border (fig. 721, m, p), in the field. The contiguous pits constitute an Araucarian feature though similar pits occur in Cedrozylon and in some other genera. Mesembrioxylon Gothani (Stopes). Dr Stopes regards this species®, from the Aptian of the Isle of Wight, as highly suggestive of the genus Phyllocladus. The medullary rays are generally 2—4 cells deep and there are 1—2 large oval simple pits in the field. Xylem-parenchyma is sparsely scattered through the wood, and stone-cells occur in the pith. 1 Stopes (15) p. 211, Pl. xx. text-figs. 60—63. 2 Ibid. p. 223, Pl. xx1. text-fig. 64. 3 Ibid. p. 228, text-figs. 65, 66. 208 CONIFERALES [CH. 1‘ (csccscmamarces Fic. 721. Mesembrioxylon bedfordense. Radial section showing groups of adjacent pits on the tracheids, 4; the medullary-ray pits, m, and a few with a border, p; n, narrow part of a tracheid. (After Stopes.) XLIV] MESEMBRIOXYLON 209 Mesembrioxylon Schwendae (Kubart). This species is recorded from Attersee in Upper Austria and though probably of Tertiary age it may be derived from Cretaceous strata. Xylem-parenchyma is present; the bordered pits on the radial walls of the tracheids are in 1—2 rows, usually separate but if contiguous not flattened; if in 2 rows opposite or sub-opposite ; the medullary rays reach a depth of 13 cells; there are generally’ 1—8 pits (fig. 722, A, B) but occasionally as many as 5 in the field; a 1 Or ©) B ed VO |, OG : O09 | . ee = eS ll | Wl (exe) ‘A Go Cc “5 So Fic. 722. A, B, Mesembrioxylon Schwendae. C, Mesembrioxylon sp. (A, B, after : Kubart; C, after Gothan.) they are bordered and the pore is obliquely vertical, though this feature is inconstant and in some places the pore is circular or the bordered pit may be replaced by a large simple pit. Sclerous cells occur in the pith. Mesembrioxylon aparenchymatosum (Gothan). In this Tertiary species, included by Gothan in Podocarpoxylon, from Seymour Island? there is no xylem-parenchyma; the medul- lary-ray cells have 1—2 elliptical-circular pits in the field and the major axis is oblique. The evidence as to affinity is far from con- clusive. 1 Kubart (11%) PL. m1. text-figs. 1—10. 2 Gothan (08) p. 8, PL. 1. figs. 9—11. 14 210 CONIFERALES [cH. Mesembrioxylon sp. (Gothan). Gothan! described some wood as Phyllocladorylon sp. which he originally stated to have been derived from King Charles Land, but Nathorst pointed out that it came from Scoresby Sound in East Greenland, N. lat. 70° 50’. The pits on the tracheids are in 1—2 rows, about 16 high, and smaller than in Xenozylon; they are scattered or contiguous and flattened, but more often separate; the medullary rays are composed of a small number of cells and there are 1—2 simple pores in the field. Mesembrioxylon antarcticum (Gothan). In this Tertiary species from Seymour Island, included by Gothan in Phyllocladoxylon?, the bordered tracheal pits are generally uniseriate and separate; the pitting of the medullary-ray cells is like that in Mesembrioxylon sp. (fig. 722, C). In the account of this wood Gothan points out that the similar pits on the walls of the medullary rays of Sciadopitys are smaller than in the fossil type; in contrast to the tendency towards a vertical elongation of the ray pits in some recent Podocarpineae those of Mesembrio- zylon are usually more horizontally stretched as in Phyllocladus and some species of Pinus. Gothan compares Cupressinoxylon Hookeri Arb.? with Mesembrioxylon antarcticum, but the former is distinguished by the smaller pits on the ray cells and by the occurrence of xylem-parenchyma. It is noteworthy that, as Gothan states’, the only recent South American Conifer possessing large simple pits in the medullary-ray cells is Podocarpus andina, while on the other hand there are several Australian genera agreeing closely with the Seymour Island species in the character of the medullary-ray pitting: from this it is concluded that in Tertiary times there was a closer connexion between the. South American and Australian regions than at the preseht day, an inference which, though not improbably correct, rests on slender evidence in this particular case. Mesembrioxylon Miillert (Schenk). This species from Pliocene strata in New South Wales, was described by Schenk’ as Phyllocladus: the pits on the tracheids 1 Gothan (07?) p. 9, fig. 2; (08) p. 6 (footnote). 2 Ibid. (08) p. 4, Pl. 1. figs. 4—8. 3 Arber (04). 4 Gothan (08) p. 25 ® Schenk in Schimper and Schenk (90) A. p. 873, figs. 424, 425. XLIV | MESEMBRIOXYLON 211 are represented by Schenk as widely separated and the single pits in the field are narrower and more oblique than in other examples of the genus. The impression made by these species, formerly referred to Phyllocladoxylon and distinguished from one another by no very well-defined characters, is that they agree with certain recent Podocarpineae and with Sciadopitys more closely than with any other recent Conifers; but in the absence of any definite evidence with regard to foliage or reproductive organs it is im- possible to select any one existing genus as the modern representa-- tive of the Arctic and South American fossil species. If the absence of Sanio’s rims is accepted as a criterion of affinity, some species of Mesembrioxylon would be included in the list of types allied to the Araucarineae, but even assuming that the preservation of the wood is such as to admit of their recognition, were they present, their absence does not nullify the evidence afforded by the tracheal and medullary-ray pitting. 2? Mesembrioxylon Hookeri (Arber) (= Cupressinoxylon Hookeri Arber). A species! founded on a splendid specimen of silicified wood neatly 9 ft. long and with a diameter of 3 ft. from Tasmania exhibited in the Geological Department of the British Museum. The stem was discovered early in the nineteenth century in Ter- tiary basaltic lava on the Macquarie plains. Dr Arber quotes Sir Joseph Hooker who gives an interesting account of the method of preservation of the decorticated wood. The annual rings are well marked and narrow; the tracheids have usually a single row of circular and scattered bordered pits on their radial walls and smaller pits are abundant on the tangential walls. Sanio’s rims are clearly shown on some of the tracheids. The medullary rays are generally uniseriate and in exceptional cases reach a depth of over 20 cells. Arber speaks of the occurrence of a small simple pit on the radial wall of the ray cells; an examination of the sec- tions in the British Museum showed that for the most part the pitting on the ray cells is not preserved but in some places a single, fairly large, simple pit occurs in the field. Resin-parenchyma is present in both spring- and summer-wood. 1 Arber (04). 14—2 212 CONIFERALES [cH. It has been suggested by Gothan! that this species should be referred to Podocarpoxrylon: the pitting of the medullary-ray cells, though seldom preserved, seems to differ from the typical Cupress- inozylon form. It is therefore referred though with some hesitation to Mesembrioxylon. =~ V. PARACEDROXYLON. Sinnott. Paracedroxylon scituatense Sinnott. The generic name Para- cedroxylon? was instituted for this Cretaceous species from Scituate, Massachusetts, in order to indicate its resemblance to Cedroxylon as defined by Kraus, more especially as regards the pitting of the tracheids and the absence of resin-canals and xylem-parenchyma in the normal wood, and to show that in some features it differs from that genus, namely in the absence of Sanio’s rims and in the smooth and thin unpitted horizontal walls of the medullary-ray cells. In typical species of Cedroxylon xylem-parenchyma is present and the tracheal pitting alone is not a distinguishing feature. The annual rings are broad and not well defined; resini- ferous parenchyma is restricted to wounded regions. Groups of thin-walled cells, which it is suggested may be abortive resin- canals, though there is no evidence that this is the case, and thick- walled parenchyma occur in the wounded tissue. The tracheids have a single row of bordered pits, generally circular. The medui- lary rays, 2—12 or more cells in depth, are pitted only on the radial walls and there are 4—-6.circular pits with an oblique slit-like pore in the field. The occurrence of bands of much thickened and pitted parenchyma is regarded as evidence of wounding: canal-like spaces occur in the traumatic tissue. Moreover in the affected regions the medullary-ray cells often show pitting on their hori- zontal and tangential walls. Sinnott regards the absence of Sanio’s rims as indicative of Araucarian affinity while the traumatic phenomena are interpreted as Abietineous characters. The genus rests on a slender basis: except for the absence of xylem-parenchyma the normal wood differs very slightly from Cupressinozylon and it is not distinguished by any well-marked features from Mesembrioxylon. Jeffrey? has described the axis of a Geinitzia cone from the 1 Gothan (08) p. 7. ? Sinnott (09). 3 Jeffrey (11). XLIV] CEDROXYLON 213 Mataram formation as exhibiting the features of - Parace- droxylon. VI. CEDROXYLON. Kraus. This generic name was instituted! for fossil wood agreeing with Cupressinoxylon in the arrangement of the pits on the tracheids and in the absence of resin-canals, but differing in the scarcity or absence of xylem-parenchyma. As defined by Schenk?, Cedroxylon stands for fossil wood agreeing generally with that of recent species of Cedrus, Abies, and Tsuga with or without tracheids in the medullary rays. Brongniart’s genus Hleoxylon? is included by Schenk as a synonym of Cedroxylon. The chief distinguishing character of Cedroxylon as compared with Cupressinoxylon, as used by some authors, is the more restricted occurrence of xylem- parenchyma; in Cedroxylon it is confined to the end of each year’s wood whereas in Cupressinoxylon the parenchyma is not so limited in its distribution. A closer examination of different types of wood included in Cedroxylon. shows that the xylem-parenchyma is an unsafe guide: Barber* states that he found more xylem- parenchyma per square millimetre in Cedrus wood than in Crypto- meria (a genus included in the general term Cupressinoxylon) and Lignier® speaks of the absence of parenchyma in some species of Cedroxylon. Gothan®, who has discussed the distinctive features of these and other genera in considerable detail, points out that in Abies Webbiana xylem-parenchyma is abundant as in Cupress- inoxylon, while in some Cupressineae the parenchyma is so scarce that it is often difficult to discover. It is clear that a test based on the preserice or distribution of xylem-parenchyma is unsatis- factory; the application of such a test would lead to the inclusion of both Abietineous and Cupressineous genera in one generic type. In typical cases the distribution of xylem-parenchyma is none the less a useful character, but Conifers with parenchyma scattered through the year’s growth are not confined to types usually in- cluded in the comprehensive genus Cupressinoxylon: many Podo- carps and some other genera not members of the Cupressineae possess abundant parenchyma in the wood. The structure of the 1 Kraus in Schimper (72) A. p. 370. 2 Schimper and Schenk (90) A. p. 862. 3 Brongniart (49) A. p. 76. 4 Barber (98) p. 332. 5 Lignier (077) p. 245. 5 Gothan (05) p. 45. 214 CONIFERALES [CH. medullary-ray cells affords an important distinguishing feature. In Cedroxylon they are characterised by the occurrence of pits on all the walls as in Abies, Tsuga, Cedrus, Larix, Picea, Pseudolariz, while in wood of the Cupressinoxylon type there is no Abietineous pitting but only pits on the radial walls. Another distinguishing feature, mentioned by Lignier,—whether important or not is open to question—is based on the greater number of sy rays per square millimetre in Cupressinozylon. The characters of Cedroxylon may be briefly Faeiieed as follows :—Annual rings well marked; bordered pits on the radial walls of the tracheids usually circular and separate and if in more than one row, opposite, but in some species the Araucarian type of pitting also occurs (fig. 723), the pits being contiguous and alternate or sometimes arranged in stellate clusters. Xylem-parenchyma typically confined to the end of an annual ring, but sometimes absent; medullary rays generally uniseriate and composed ex- clusively of parenchyma though horizontal tracheids may occur; pits on all the walls of medullary-ray cells as in the Abietineae; on the radial walls there may be 1—6 apparently simple circular pits in the field. There are no resin-canals except as the result of injury. Cedroxylon transiens Gothan. This species from Upper Jurassic rocks in Spitzbergen! and from Lower Cretaceous beds in King Charles Land? illustrates the admixture of opposite, separate, and contiguous, alternate, pits in the same wood (fig. 723, A, B); stellate groups also occur (fig. 723, A), an arrangement occasionally seen in some recent Pines, e.g., Pinus Merkusii?. The Araucarioid type of pitting is charac- teristic of the spring-wood where it is associated with the more usual opposite and separate arrangement. Smaller separate pits occur on the tangential walls of some of the tracheids. Resin-canals are absent, though a solitary example is recorded by Gothan. The medullary rays have several small pits in the field, also pits on the tangential and horizontal walls; the xylem-parenchyma is confined to the end of the year’s wood ; the cross-walls are pitted (fig. 723, B). 1 Gothan (10) p. 38; Pl. vi. figs. 11—13. ° Ibid. (07°) p. 26, figs. 14,15; Pl. r. fig. 1. 8 Groom and Rushton (13) Pl. xxv. figs. 47, 48. XLIV | CEDROXYLON 215 Gothan makes no mention of Sanio’s rims. The Abietineous features predominate over the Araucarian, the latter being limited to the local occurrence of polygonal and alternate bordered pits. Fic. 723. A, B, Cedroxylon transiens. C, Cedroxylon blevillense. (A, B, after Gothan; C, after Lignier.) Wood of similar type was described by Schroeter! from King Charles Land as Pinus (Larix) Johnseni: resin-canals, possibly due to wounding, occur in the summer-wood. The medullary rays are 1—18 cells deep and there are 1—3 simple circular pits in-the field; all the walls of the ray cells are pitted. As in Cedroxylon transiens the Araucarian type of pitting is represented on some of the tracheids. Cedroxylon Hornet Seward and Bancroft. An Upper Jurassic species from Helmsdale?, Sutherland, a locality from which Hugh Miller recorded numerous specimens of fossil wood which is still abundant on the beach immediately north of Helmsdale. The annual rings are well defined: the bordered pits are usually in a single row on the radial walls of the tracheids, occasionally in contact and flattened; double rows of opposite pits are not uncommon. Xylem-parenchyma is confined to the late wood. Medullary rays, 1—26 cells deep, generally 8—12, 1 Schroeter (80), Pl. 1. figs 1—8. 2 Seward and Bancroft (13) p. 883, text-fig. 5; Pl. 11. figs. 22—25. 216 CONIFERALES [cH. and uniseriate; there are 2—4 simple or faintly bordered circular pits in the field and pits occur also on the tangential and horizontal walls. ® © © @ Fig. 724. A, Brachyphyllum eathiense; stoma. B—F, Cedroxylon Hornei. (After Seward and Bancroft.) Cedroxylon cedroides Gothan. In this species from King Charles Land}, probably Lower Cretaceous, the pitting of the tracheids is in the main of the usual Coniferous type and not Araucarioid, though in places the pits are 1 Gothan (072) p. 23, figs. 11—13. XLIV] CEDROXYLON 217 alternate and form stellate groups. The medullary rays, reaching a depth of 30 cells, show very clearly the Abietineous pitting and there are 4—5 simple circular pits in the field. The occasional occurrence of single large pores in the field would seem to be due, at least in part, to the destructive action of fungi. Xylem- parenchyma occurs in the summer-wood. The structure agrees with that of the wood of Cedrus, Pseudolariz, and Tsuga; the Abietineous features are relatively more conspicuous than in Cedroxylon transiens in which the tracheal pitting is more Arau- carian. Cedroxylon maidstonense Stopes. In this wood!, from the Lower Greensand of Kent, the rings of growth are well marked; the bordered pits on the radial walls of the tracheids are usually uniseriate and Sanio’s rims are dis- tinctly preserved. Xylem-parenchyma is absent or very rare and there are no resin-canals. The medullary rays are seldom deeper than 10 cells; there are 4—6, or occasionally more, oval or circular pits in the field and some have a slit-like pore and are bordered; pits are clearly shown on the tangential walls of the ray cells, Cedroxylon pottoniense Stopes. This species? of the same geological age, from Potton in Bedford- shire, differs from C. maidstonense in the comparative abundance of xylem-parenchyma: the medullary-ray cells show very clearly the Abietineous type of pitting. Cedroxylon blevillense Lignier. In this species from the gault of Bléville (Seine-Inférieure)* the tracheal pits are usually uniseriate and separate but if in two rows they may be either opposite or alternate. The pits in the field are small, numerous, and have an oblique pore (fig. 723, C). The tangential walls of the ray cells are sometimes pitted. There is no resiniferous parenchyma. The characters afford another illus- tration of the impossibility of drawing any clearly defined line between Cedroxylon and allied generic types. 1 Stopes (15) p. 149, Pl. xu. text-figs. 41—43. 2 Tbid. p. 154, text-fig. 44. 3 Lignier (07?) p. 267, Pl. xvu. figs. 15—17; Pl. xxr. fig. 66; Pl. xxir. fig. 72; Pl. xxiu. fig. 87. 218 CONIFERALES [CcH. A Triassic species described by Wherry! as Brachyoxylon penn- sylvanicum may, as that author suggested, belong to Cedrozylon: the tracheids have 1—2 rows of pits, usually separate but sometimes alternate and hexagonal as in Cedroxylon transiens and C. Hornet; no description is given of the medullary-ray pitting or of any xylem-parenchyma. There are no adequate grounds for referring this Triassic wood to Cedroxylon. Several species of wood from Triassic and higher horizons have been assigned to Cedroxylon, but in many cases the descriptions fall short of modern standards and accurate determination is impossible. Crié? describes a species, C. australe, from the Trias of New Caledonia though his figures and descriptions do not afford satisfactory evidence in support of this reference. Schenk? mentions Cedrorylon pertinax (Goepp.) as the oldest representative of the genus and speaks of it as Rhaetic, while Gothan refers the species to a Juras- sic horizon. A species founded by Goeppert and described by Mercklin4 from Jurassic rocks of Russia, Pinites jurassicus, may be a Cedroxylon: the bordered pits are usually separate and opposite but sometimes in contiguous groups. Mercklin states that small thick-walled cells, often with dark contents, occur at the outer limit of each ring. A specimen described by Felix® as Cormocedroxylon jurense from the Braun Jura of Galicia is com- pared by him with Pinites jurensis. Fliche® records, though with- out complete diagnoses, some French Lower Cretaceous species: the tracheids of Cedroxylon reticulatum Sap., from the Albian of L’ Aube, are characterised by pits which are usually separate but may be contiguous and flattened. Cones closely resembling those of Cedrus occur in the same beds. This author gives partial descrip- tions of C. barremianum Fliche? from the Lower Cretaceous of Haute Marne and a Cenomanian species C. manekildense Fliche®, but in neither case are the data adequate. C. matsumurae Stopes and Fujii® is an Upper Cretaceous Japanese species with 1—2 rows of tracheid-pits, generally opposite 1 Wherry (12) Pl. rv. 2 Crié (89) Pls. 1.—v. 3 Schimper and Schenk (90) A. p. 871. 4 Mercklin (55) p. 48, Pl. vin. figs, 6—10. 5 Felix (82) p. 264. 5 Fliche (97) p. 7. ? Ibid. (00) Pl. 1. fig. 1. 8 Ibid. (96) Pl. xv. fig. 3. 9 Stopes and Fujii (10) p. 42, PL 1. fig. 10; PL. iv. figs. 20—23. XLIV] PITYOXYLON 219 but sometimes alternate though not contiguous. The medullary rays, 5—12 or rarely 20 cells deep, are imperfectly preserved. Another species, C. Yendoi St.1 and Fuj. from the same locality is also founded on material that is insufficient for accurate deter- mination. Sporadically occurring resin-ducts are regarded as traumatic. Among Tertiary species reference may be made to Cedroxylon affine Kraus* from Sicily, without resin-parenchyma and charac- terised by usually two large simple pits in the field; C. Hoheneggert Felix? from the Eocene of Moravia figured by Schenk as from Cretaceous strata; C. Hermanni Sch.*, an incompletely described species from Assam, probably of Tertiary age. VII. PITYOXYLON. Kraus. Kraus® included in this genus some of the species previously referred by Goeppert to Pinites; others he assigned to Cedroxylon. Pityoxylon is distinguished from Cupressinoxylon and Cedroxylon by the normal occurrence of resin-canals in the wood and by the presence of horizontal tracheids in some of the medullary rays. Within the limits of the genus the following differences occur in the characters of the medullary rays and the resin-canals :—the walls of the ray-tracheids are smooth or provided with dentate ingrowths; the pits on the medullary-ray cells are large and simple or smaller and apparently bordered, and there may be one or several pits in the field; the parenchyma of the resin-canals has thin or thick walls. As generally employed Pityoxylon includes species exhibiting anatomical features met with in Pinus, Picea, Larix, Pseudotsuga, and some other Abietineae. Gothan® makes * use of two generic names, Piceorylon and Pinuxylon, to denote the possibility of more precise comparison with recent types than is - implied by Kraus’s more comprehensive term. Piceoxylon is characterised by thick-walled epithelial cells lining the resin-canals, by small pits in the ray cells, spiral tracheids in the summer-wood, the absence of teeth in the ray-tracheids, clearly marked Abietineous 1 Stopes and Fujii (10) Pl. rv. figs, 24—26. 2 Kraus (83). 3 Felix (82) p. 268. 4 Schenk (82?) p. 355. 5 Kraus in Schimper (72) A. p. 377. ® Gothan (05) p. 102. 220 CONIFERALES [cH. pitting in the ray cells, and by the occurrence of numerous pits in the tangential walls of the summer tracheids. Pinuzylon is used by Gothan in preference to Pinoxylon, the name adopted by Knowlton! for wood in which there are no resin- canals in the medullary rays. In Pinuaylon the walls of the epithelial cells are thin, rarely thick; the medullary rays have large simple pits in the spring-wood; there are no spiral bands in the tracheids. The horizontal tracheids have smooth or dentate walls and the Abietineous pitting is much reduced. The distinctions on which these two genera are based are thus not very clearly defined and it is only in particularly well-preserved material that the two generic types can be recognised with certainty. Dr Stopes? follows Jeffrey and Chrysler? in regarding Gothan’s twofold division as unnecessary. In the majority of species referred to Pityorylon the published information is insufficient for a sub-division in Gothan’s sense and as a rule the generic name stands for wood of an Abietineous type which cannot be assigned with confidence to any one recent genus. The question of the antiquity of the Abietineae has been confused by the too liberal use of the term Pinites by Goeppert and some other authors for stems which have no claim to be placed in the genus Pityoxylon. Jeffrey and Chrysler*, who follow previous authors in quoting Pinites Conwentzianus Goepp.®, described as a Carboniferous species from Waldenburgh, as evidence of a Palaeo- zoic Pinus-like wood, state that the species receives ‘full confir- mation from the description of a similar type, Pityorylon chasense Pen.§ from the Permian of Kansas.’ Goeppert and Stenzel state that Pinites Conwentzianus was found on a rubbish-heap (‘Halde’), but Goeppert apparently entertained no doubt as to its Carboni- ferous age. Through the courtesy of Prof. Frech of Breslau I was able to examine the original sections and convinced myself that the wood is Abietineous: the rings of growth are well defined ; horizontal tracheids occur in some of the rays and the tracheal pits, 1—2 rows, are widely separated, though occasionally the pits 1 Knowlton in Ward (00) B. p. 420. 2 Stopes (15). 3 Jeffrey and Chrysler (06). + Ibid..p. 13. > Goeppert and Stenzel (88) p. 54, Pls. x1., x11. 5 Penhallow (00) p. 76; Thomson and Allin (12). “XLIV] - PITYOXYLON 221 of a single row are in contact and slightly flattened. Feeling sceptical as to the Carboniferous age of the wood I wrote to Dr Conwentz who confirmed my doubts with regard to the value of the evidence as to the geological horizon. Thomson and Allin have shown that Penhallow’s Pityoxylon cannot be accepted as trust- worthy evidence of the occurrence of a Palaeozoic Abietineous type. Pityoxylon chasense is not an Abietineous species; it is founded on Dadozylon wood devoid of annual rings and without resin-canals traversing the medullary rays. The fragments of wood from the Muschelkalk of Recoaro figured by Schleiden and Schenk! as Pinites Goeppertianus afford no evidence of Abietineous affinity beyond the occurrence of separate bordered pits on the walls of the tracheids. Pityoxylon eiggense (Witham). The petrified wood first named by Witham? Pinites eiggensis and afterwards$ referred by him to the genus Peuce was originally recorded by Macculoch in 1814 from below the massive and pre- cipitous ridge of pitchstone which forms a striking feature above the basaltic lavas of the Sgurr of Higg in the Inner Hebrides. Lindley and Hutton‘ and Nicol® also gave short descriptions of the structure and Miller® in the Cruzse of the Betsey alludes to.a fossil trunk as ‘an ancient tree of the Oolites.’ The wood occurs with fragmental sedimentary rocks below the pitchstone and not actually in situ; Mr Harker’s thorough examination of the island led him to the conclusion that the wood and associated rock- fragments are derived from Jurassic (Oxfordian) strata and were carried up by volcanic agency’. Mr Harker tells me that he has never seen the Pityoxylon with any undoubted matrix adherent; it occurs with wood of a different type (Dadoxylon) which is em- bedded in a white sandstone agreeing exactly with the Great Estuarine Sandstone of Eigg in which similar wood has been found in place. It is, however, possible that Pityoxylon did not come 1 Schenk (68) Pl. v. figs. 4—7. 2 Witham (31). 3 Ibid. (33) A. Pls. xiv., xv. 4 Lindley and Hutton (33) A. Pl. xxx. 5 Nicol (34) A. p. 154. 6 Miller (58) p. 37. 7 Harker (06) p. 55; (08) p. 52. In these memoirs Mr Harker discusses the -earlier conclusions of Sir Archibald Geikie as to the geological history of Eigg and gives references to previous notices of the fossil wood; Seward (11°) p. 652. 222 CONIFERALES [cH. from the same source as the Dadoxylon wood. Though probably Jurassic, a Tertiary source is by no means ruled out. Kraus! transferred Witham’s species to the genus Pityoxylon and that name is used by Schroeter? and Schenk’. The specimens on which the following account is based are most of them in the British Museum. Annual rings clearly defined, usually 1—1-5 mm. ° . 2 e x q , A ? P B 3 ve aie os Cc D Fie. 725. Pityoxylon eiggense; c, resin-canal in a fusiform medullary ray. (British Museum, 51427, 51641, 51727.) broad; the reduction in diameter of the summer-tracheids extends over several rows, the transition being much more gradual than in some types of Coniferous wood. A characteristic feature is the occurrence of more or less circular patches where the tracheids have been destroyed with the exception of a single tracheid or a ? Schroeter (80) p. 13. Kraus in Schimper (72) A. p. 378. * Schimper and Schenk (90) A. pp. 855, 874. XLIV] PITYOXYLON 223 small group in the centre of a clear crystalline matrix. Some of these patches simulate resin-canals, a fact which led Schenk to deny the existence of true canals. All stages of decay are shown, _from the partial obliteration of a circular group of tracheids to the destruction of the group, one central element being left, or to the formation of a canal-like cavity (fig. 725, A). It is often difficult to decide whether a clear space in the wood is a canal or the result of post-mortem changes, but there is no doubt as to the occurrence of some true secretory canals in different regions of the wood. There is very little parenchyma accompanying the canals. The medullary.rays are of two kinds, uniseriate, 1—13 cells in depth, though usually about 6 cells in depth, and lenticular rays with a central canal (fig. 725, C, c) identical with those in a modern Pine. Fig. 725, D shows part of a vertical canal with some parenchymatous lining in continuity with a hotizontal canal in a broad medullary ray. Several small pits occur on the tangential and horizontal walls of the ray cells, and the radial walls, which are less clearly preserved, occasionally show 1—3 elliptical pits. In radial section the upper and lower cells of a medullary ray are often distinguished by their less uniform breadth and resemble in this respect ray tracheids. A careful examination of sections revealed the existence of bordered pits in the tangential walls of these elements and confirmed their tracheal nature?. The bordered pits are generally single and sparsely scattered on the radial walls of the tracheids; occasionally the pits are in contact and a few double and opposite rows occur. An interesting feature is the occurrence in some tracheids of a biconcave patch of some brown substance agreeing closely with resinous deposits described by Penhallow®. If this species is from Jurassic strata its close resemblance to recent types of Pinus is a fact of consider- able interest. Pityoxylon Ruffordi Seward. This species? is founded on a specimen of wood obtained by Mr Rufford from Wealden beds at Ecclesbourne on the Sussex 1 T am indebted to Miss Ruth Holden for calling my attention to this feature and for other assistance in the examination of the sections. 2 Penhallow (04) p. 526. 3 Seward (95) A. p. 199; (96) p. 417, Pls. m1., IIL. 224 CONIFERALES [CH. coast: the sections on which the description is based are in the British Museum. Annual rings well marked, varying in breadth from 1 to 3 mm.; resin-canals are abundant both in the spring- and Fic. 726. Pityorylon Sewardi. Radial section showing tracheids, tr, in a medullary ray; ¢, parenchyma of the ray with simple pits. (After Stopes.) autumn-wood and horizontal canals occur in the lenticular medul- lary rays: some of the canals are occupied by large rounded cells like tyloses. The bordered pits on the radial walls of the tracheids form single or double rows; in the latter case the pits are generally XLIV] PITYOXYLON 225. opposite but stellate groups also occur as in Cedroxylon transiens Goth. (fig. 723, A) and several other species. The uniseriate medullary rays reach a depth of 30 cells: there are usually 2—4 oval or circular pits in the field. A similar type of stem is represented by Pityoxylon Nathorsti+ (Conw.) from the Lower Cretaceous of Sweden. Pityoxylon Sewardi Stopes. This species? is founded on a petrified branch, not less than 18 cm. in diameter, from the Lower Greensand of Kent. It exhibits the usual features characteristic of the genus; the wood contains horizontal and vertical canals with thin-walled epithelial cells. The medullary rays are larger and more abundant than in most Coni- ferous woods and horizontal tracheids (fig. 726, tr) occur inter- spersed with the parenchymatous cells, c, as well as on the upper and lower margins, an arrangement in which the fossil bears a striking resemblance to the recent species Pinus monticola*. Pityoxylon Benstedi Stopes. In this Lower Greensand species* from Kent the resin-canals often contain tyloses as in P. Nathorstt Conw. and the epithelial cells have very thick walls, a feature suggesting comparison with the genus Lariz. The medullary rays show well-marked Abie- tineous pitting (fig. 727, a) and ray-tracheids (rt) occur. Rims of Sanio are shown in fig. 727 between the circular bordered pits, tr. The difference between the tracheal and ordinary parenchy- matous elements of the rays, as represented in fig. 727 from a drawing by Dr Marie Stopes, is not very clearly defined and in the upper ray shown in the figure part of a tracheid is seen abutting laterally on parenchymatous ray cells, the only difference between them being in the form of the pits, a criterion which is largely dependent for its value on the state of preservation. Dr Stopes is inclined to regard this species as most nearly allied to Lariz. Pityoxylon statenense Jefirey and Chrysler. - A species from the Middle Cretaceous of Staten Island® found in association with the short shoots described by Jeffrey and 1 Conwentz (92) p. 13, Pls. 1.—111., vi., VIr. 2 Stopes (15) p. 95, Pls. 1v., v. text-figs. 23, 24. 3 [bid. text-fig. 25, p. 103. 4 Ibid. (15) p. 105, Pls. v.—vit. text-figs. 26, 27. 5 Jeffrey and Chrysler (06). Ss. IV 15 226 CONIFERALES [cH. Hollick as Pinus triphylla, etc. The anatomical features are as follows :—annual rings narrow, not clearly marked owing to the walls of the summer-tracheids being thinner than in recent species of Pinus; xylem-parenchyma confined to the periphery of the (aha @ 1 I L981 § = f ‘S) © = -62W rt. © r-) bane | spo S OL PEM IOC | ote Bsle he = Fic. 727. Pityoxylon Benstedi. Radial section showing the tracheal pitting, ir; rt, ray-tracheids; p, ray cells with large pits; a, typical Abietineous pitting of end-walls of medullary-ray cells. (After Stopes.) resin-canals which occur in any part of the wood and are often filled with tyloses. The tracheids have a single row of pits not contiguous or flattened; the pits on the tangential walls are confined to the summer-tracheids; both linear and fusiform XLIV] PITYOXYLON 227 medullary rays occur, the latter with horizontal resin-ducts. There are no ray-tracheids. There is usually one circular or elliptical pitin the field. A second species from the same locality, Pityoxylon scituatense, differs only in some unimportant features from P. staten- ense. These fossils differ from recent Pines as also from Picea, Pseudotsuga, and Larix in the absence of ray-tracheids. In the restriction of bordered pits to the tangential walls of the tracheids of the summer-wood they agree with the soft Pines, but though this character is generally lacking in hard Pines, Jeffrey and Chrys- ler point out that in some haid Pines without pits on the tangential walls of the tracheids of vegetative shoots the summer elements of the cones have tangential pits. The occurrence of bordered pits on the tangential walls of the late wood and the absence ofray- tracheids are regarded by the authors of the species as ancestral features. Pityoxylon protoscleropitys (Holden). A Middle Cretaceous species! from New Jersey, referred by Miss Holden to Pinus, showing the following features :—annual rings well developed; linear and fusiform medullary rays, hori- zontal and vertical resin-canals, bordered pits uniseriate and scat- tered on the radial walls of the tracheids; none on the tangential walls. Rims of Sanio are present. There are 1—2 pits in the field with a lenticular pore and circular border; the other walls of the ray cells are abundantly pitted. Ray-tracheids occur on the margins of the medullary rays and rarely interspersed with the parenchyma; their walls are denticulate as in recent hard Pines. The presence of horizontal tracheids in the medullary rays is an important character: in Pityoxylon scituatensiformis (Bailey)?, another Middle Cretaceous species, ray-tracheids are present but they have smooth walls and are not met with in the first 10—15 rings of wood, whereas in P. protoscleropitys they occur even in the wood of the first year. In this connexion the presence of ray- tracheids in Pityoxylon eiggense is noteworthy at least if that species is from a Jurassic source. Pityoxylon protoscleropitys is considered by Miss Holden to be ‘probably the earliest form with all the characters of a modern hard Pine, yet retaining certain ancestral 1 Holden (134). * Bailey (11). 15—2 228 CONIFERALES . [ CH. features, as the association of primary and fascicular leaves, the latter borne on brachyblasts subtended by a foliar trace.’ Miss Holden? has also described from New Jersey two species which she compares with Prepinus, namely Pityorylon foliosum and P. anomalum. Pityoxylon Nathorsti (Conwentz). Under the name Pinus Nathorsti Conwentz? described in considerable detail specimens of petrified wood from the Senonian Holma sandstone of Sweden, also a cone and two detached needles. The rings of growth agree with those of stems and older branches in the gradual increase in the thickness of the tracheid-walls in passing from the spring to the late summer elements. The bor- dered pits on the radial walls of the tracheids are uniseriate and separate and none were found on the tangential walls. Both horizontal and vertical 1esin-canals occur, several of them with well-preserved tyloses; the epithelial cells are thin-walled and unpitted as in Pinus silvestris. It may be that the abundance of tyloses is connected with the presence of fungal mycelia as in wood of Hevea stems recently described by Mr Brooks?. The medullary a1e uniseriate, generally 5—7 cells deep, also fusiform and with resin-canals: the preservation is not sufficiently good to admit of any definite statement as to the occurrence of horizontal tracheids. Pityoxylon zezoense (Suzuki). This Upper Cretaceous Japanese species was described as Abtocaulis zezoensis*, but in view of the occurrence of features suggesting comparison with Pinus as well as with Abies the more comprehensive generic name is preferable. In the presence of pits on the tangential and horizontal walls of the medullary-ray cells and in the pitting of the tracheids this wood conforms to the Abietineous type. There are no ray-tracheids: the pitting on the radial walls of the ray cells agrees in part with that in Abies and in some of the cells there are large circular pits like those of Pinus. Normal resin-canals are present in the second ring only, others being interpreted as traumatic. Xylem-parenchyma is 1 Holden (13%). ’ 2 Conwentz (92) p. 13, Pls. 1.—u11., vz, vu. 3 Brooks and Sharples (14). 4 Suzuki (10). va XLIV] PITYOXYLON 229. sparsely distributed. There would seem to be little difference of importance between this species and wood referred to Cedroxylon. The number of Cretaceous examples of Pityoxylon might be considerably extended: for an account of French species reference should be made to Lignier and to Fliche. Pityoxylon Pseudotsugae (Gothan). Gothan described this species, from South Nevada and prob- ably of Tertiary age, as Piceorylon Pseudotsugae!: it is interesting as a type of Pityorylon agreeing closely with the recent genus Pseudotsuga in the presence of spiral bands in the tracheids. There are both vertical and horizontal resin-canals and the ray cells have Abietineous pitting. Xylem-parenchyma occurs next the summer- wood and the epithelial cells have thick walls, features in agree- ment with Gothan’s genus Piceorylon. Bailey? points out that in the absence of spiral bands in the ray-tracheids the fossil species resembles Pseudotsuga Douglasti, while in Pseudotsuga macrocarpa the tracheids of the rays have spiral bands. Fritel and Viguier® have described a species from Eocene beds in the Paris Basin as Piceorylon Gothani in which some of the xylem- tracheids have spiral bands. Pityoxylon pulchrum (Cramer). A Tertiary species* originally described from material collected by Sir Leopold MacClintock in Banksland as Cupressinoxylon pulchrum. A piece of wood in the Dublin Museum labelled ‘from Ballast Bay, Baring Island, given by Sir L. MacClintock’ agrees very closely with Cramer’s type-specimen, and as the resemblance extends to most of the anatomical characters, I believe it to be the material on which C. pulchrum was founded. The chief difference is that the Dublin wood has resin-canals as in the specimen described by Cramer as Pinus MacClurit (?) Heer’; in C. pulchrum no resin-ducts are recorded. On the other hand in the sum of its characters the Dublin specimen agrees much more closely with C. pulchrum. Annual rings well marked; 1 Gothan (06%). 2 Bailey (09) p. 54. 3 Fritel and Viguier (11) p. 63. * Cramer (68) p. 171, Pl. xxxrv. fig. 1; Pl. xxxvi figs. 6—8. 5 Ibid. Pls. xxxv, XXXVI. 230 CONIFERALES _ [CH bordered pits large, approximately 25y in diameter, in 1—2 opposite rows, sometimes in contact and slightly flattened. Medullary rays uniseriate, 1—14 cells deep, also fusiform rays containing a horizontal canal; 2 or 3 large oval pits occur on the radial walls of the ray cells and in a few cases pits on the tangential walls. Ray-tracheids with bordered pits occur on the edges of the medullary rays. : Among other Tertiary species reference may be made to Pity- oxylon parryoides Goth. from the Braunkohle of Rheinland, so named from its resemblance to the North American Pinus Parrya, characterised by horizontal tracheids with smooth walls and thin- walled epithelial cells; also Pityoxylon pineoides Kraus? a Sicilian Tertiary species without ray-tracheids. Pityoxylon succinifer (Goeppert). This species from the Oligocene amber beds of the Balti¢ coast was first named Pinites succinifer® and several years later fully described and admirably illustrated as Pinus succinefer*. It affords a striking illustration of the possibilities of amber as a petrifying agent and shows several features of anatomical interest. The roots are represented by pieces of wood in a pathological state: the tracheids have 1—3 rows of pits on their radial walls and some of them contain tyloses; the walls of the ray-tracheids have dentate ingrowths. The stem and branch wood is more complete. Sieve-tubes and sieve-plates are exceptionally well preserved and both cortex and pith tissues are represented. The tracheids have 1—2 rows of separate pits; a spiral sculpturing on the walls of the tracheids was mistaken by Menge for the spiral bands characteristic of the Taxineae and he named the species Taxoxylum electrochyton. Conwentz describes tyloses in the tracheids, also a crescentic patch of parenchyma in the wood passing into a mass of resin®, a feature occasionally seen in recent wood. The medullary rays have 1—4 pits in the field; both ray- tracheids and horizontal resin-canals occur and in some cases 1 Gothan (09) p. 523, figs. 3—5. 2 Kraus (83) p. 83, Pl. 1. figs. 1—3. ® Goeppert (41) p. 39; Goeppert and Berendt (45) A. p. 61. 4 Conwentz (90) A. p. 26, with numerous plates. 5 Ibid. (89); (90) A. p. 48; cf. Hollick and Jeffrey (09) B. Pl. xxi. fig. 4. > XLIvV] PROTOPICEOXYLON 231 rays are said to consist exclusively of tracheal tissue. Pine needles and cones have been obtained from the amber beds. The Fossil forests of the Yellowstone Park include examples of Pityoxylon trees some of which have been described by Knowl- ton? and Felix®, but unfortunately the anatomical details are not as a rule well preserved. The most striking exposure ‘of the Tertiary (probably Miocene) trees is on the slopes of Amethyst mountain (fig. 712), where a succession of forests is represented throughout the 2000 ft. of strata. Felix describes a species, Pityozylon fallax, chiefly interesting from the point of view of a comparison between the stem and root wood of the same tree: the elements of the root are in general larger than those in the stem. Knowlton gives an account of P. Aldersoni and P. ame- thystinum, species which may be identical: the pits on the tracheids and medullary rays are seldom preserved, but the occurrence of both vertical and horizontal resin-canals is clearly shown. VIII. PROTOPICEOXYLON. Gothan. Protopiceoxylon exstinctum Gothan. The generic name Proto- piceoxzylon was proposed for some Lower Cretaceous wood from King Charles Land? possessing Abietineous characters, intermediate between Cedroxylon and Pityoxylon in having only vertical resin- canals, at least in uninjured wood. The anatomical features of the type-specimen are complicated by the occurrence of additional resin-canals in wounded portions of the stem. It is difficult to determine the precise extent of the traumatic influences, but the presence of callus-wood healing a wound leaves no doubt as to the correctness of Gothan’s conclusion that certain features are abnormal and due to the effects of wounding. In the species P. eaxstinctum are also included specimens from Spitzbergen® and some of the material on which Cramer® founded his species Pinites cavernosus: the later specific name is not retained on the ground that the original diagnosis is incorrect and it was only after examining sections of the type-specimen that Gothan recog- nised the true nature of Cramer’s species. 1 Conwentz (90) A. Pl. 1x. fig. 2. 2 Knowlton (99) p. 763, Pls. cvI.—cviil., CXII.—CXV., CXVIII., CXIX. 3 Felix (96) p. 254. 4 Gothan (07?) p. 32, figs. 16,17; PI. 1. figs. 2—6. 5 Gothan (10) p. 15, Pl. m1. figs. 5—8; Pl. in. figs. 1—4, 6—8. ° Cramer (68). 232 CONIFERALES [cH. Protopiceoxylon exstinctum shows the following characters :— annual rings well marked; vertical resin-canals occur in the wood but there are no canals in the medullary rays except a few of unusually large diameter in wounded areas; there is no xylem- parenchyma apart from the resin-canals. Tracheids with 1—2 rows of bordered pits on the radial walls, separate and circular, also contiguous and flattened, opposite, or sometimes alternate: in the occurrence of the Araucarian type of pitting on some tracheids this species agrees with several types of Mesozoic wood. Medullary rays uniseriate, characterised by well-developed Abie- tineous pitting; on the radial walls there are 2—4 circular and bordered pits in the field. No undoubted ray-tracheids were noticed ; numerous small pits occur on the horizontal walls of paren- chymatous cells associated with the resin-canals. The pith consists of parenchyma with thin sclerenchymatous diaphragms. The horizontal canals, presumably traumatic, in some of the medullary rays resemble in their large size those in Anomalozxylon but in that genus there is no Abietineous pitting on the medullary- ray cells; similar canals are described by Jeffrey+ in wounded wood of Cedrus and other Conifers. In the occurrence of vertical canals only in the normal wood Protopiceoxylon is intermediate between Cedroxylon, which has no canals, and Piceorylon and Pinuzylon of Gothan (= Pityoxylon of Kraus), the fossil represen- tatives of such recent genera as Larix, Picea, and Pinus, in which both vertical and horizontal ducts occur. Gothan holds, and probably with good reason, that vertical canals preceded those in the medullary rays and regards the fossil species as a primitive type. A species from the Black Hills described by Knowlton? as Pinoxylon dacotense agrees with Protopiceorylon in having only vertical canals, but it is not clear whether they are normal or traumatic: Piceoxylon would seem to be the more appropriate designation for Knowlton’s species. Protopiceoxylon articum sp. nov. This species is founded on a specimen from Cape Flora, Franz Josef Land, probably Oxfordian in age. Annual rings are distinct 1 Jeffrey (03); (05). * Knowlton in Ward (00) B. p. 420, Pl. cuxxrx. XLIV] PROTOPICEOXYLON 233 and narrow; several oval or circular spaces are conspicuous in transverse section, some being true canals and others the result of decay. There are 2—3 opposite rows of bordered pits on the radial walls of the tracheids. Partially destroyed rows of resin- parenchyma occur which probably belong to secretory canals. The uniseriate and comparatively deep medullary rays, 20—30 cells, are characterised by rather thick and pitted horizontal and i aN Fic. 728. Protopiceorylon arcticum. (Cambridge Botany School.) Fic. 729. Protopiceoxylon Ed- wardsi. Longitudinal view of the thick-walled, pitted, epithelial cells of the resin- canals, (After Stopes.) vertical walls (fig. 728); 4—5 small simple pits occur on a few of the cells and on the upper and lower edges of some of the rays are empty elements of unequal breadth which in all probability are ray-tracheids. The wood agrees in the presence of vertical canals only and in the structure of the medullary rays with Proto- piceorylon exstinctum Goth. In Gothan’s species there are 2—4 bordered pits in the field, but the absence of a border in the Franz Josef Land wood may be a consequence of imperfect preservation. , 234 CONIFERALES [cH. Protopiceoxylon Edwardsi Stopes. Founded on a branch from the Lower Greensand of Sussex, showing 17 annual rings, having the following characters!: a large pith nearly 3 mm. in diameter composed of parenchyma without stone-cells; tracheids with usually one row of circular bordered pits; vertical canals in the summer-wood and associated with a small amount of resiniferous parenchyma; medullary rays uniseriate, with Abietineous pitting and 2—4 more or less circular pits in the field. The small size of the resin-canals is a character- istic feature, also their thick-walled pitted epithelial cells (fig. 729). The species differs from Gothan’s P. exstinctum in the smaller diameter of the canals, the absence of traumatic horizontal canals, and in the greater number of the vertical secretory passages. IX. WOODWORTHIA. Jeffrey. Woodworthia arizonica Jeffrey. This genus? is founded on specimens from the Triassic petrified forest of Arizona character- ised by the occurrence of short shoots in the secondary wood com- parable with those in the stem of Araucariopitys. In the type- specimen the annual rings are not very clearly defined: the pitting on the tracheids is definitely Araucarian. The medullary rays are uniseriate, 2—9 cells deep: they appear to have pits only on the lateral walls. On the surface of the wood are several small scars and a few larger ones, the former representing short shoots subtended by a leaf-trace; the shoots are not infrequently branched as they pass through the secondary xylem, a feature recorded also in Ginkgo?. Jeffrey describes the short shoots as having a limited existence and disappearing in the wood at a comparatively short distance from the pith; they have no rings of growth, a character associated with short-lived leaf-spurs in recent species but a feature in which they differ from those of Ginkgo. The leaf-traces subtending the short shoots, in contrast to those of Araucaria, are not persistent throughout the secondary wood. Jeffrey regards this fact as an argument against the view that the persistence of the traces in Araucaria is a primitive character; but it is worthy of note that 1 Stopes (15) p. 81, Pl. m1. text-figs. 17—22. * Jeffrey (107), Pls. XXXI., XXXII. 3 Tupper (11). XLIV] ARAUCARIOPITYS © 235 the leaves accompanying the shoots of Woodworthia are not strictly comparable with those of the foliar organs of recent Araucarias which have no short shoots in their axils. Jeffrey regards the short shoot as a primitive attribute of the coniferous stock and its occurrence in the stems of Woodworthia and Arau: cartopitys is held to be evidence in support of the interpretation of the seminiferous scales of Abietineous genera as metamorphosed short shoots, an interpretation which is open to question. The presence of short shoots is not a monopoly of the Abietineae and their presence in a stem may be regarded as a point of contact with Ginkgo as well as with Abietineous plants. Attention is called _ elsewhere to the probability that foliar spurs like those of Pinus are specialised forms of ordinary shoots. However we may interpret the characters exhibited by Woodworthia, the genus is an interesting example of an extinct type illustrating the combina- tion with Araucarian characters of a morphological feature that is no longer represented in the Araucarineae. X. ARAUCARIOPITYS. Jeffrey. A genus founded by Jeffrey? on a stem from the Middle Cre- taceous beds of Staten Island, New York, showing on its decorti- cated surface scars of short shoots and in the structure of the wood both Abietineous and Araucarian features. Araucariopitys americana Jefirey. The bordered pits on the radial walls of the tracheids are often contiguous and flattened though in places separate and circular, usually arranged as a single row. All the walls of the ray cells are pitted as in the Abietineae. The large number of vertical resin-canals (fig. 758, C, D, page 323) in a single tangential row is regarded as evidence of traumatic origin. This conclusion is based on the fact that when canals are present in wood that is known to have been wounded they occur in crowded tangentially arranged rows in contrast to their sparser distribution in the normal wood. There are no canals in the uninjured tissues of Araucariopitys. Diaphragms of sclerous cells occur in the pith as in some recent Abietineae. The short shoots are shown in tangential section of the stem and stated to be accom- panied by a subtending leaf. 1 Jeffrey (07) Pls. xxvi1.—xxx. 236 CONIFERALES [cH. It is suggested by Jeffrey that Araucariopitys may be the stem of Heer’s genus Czekanowskia', but there is no proof of a connexion and evidence afforded by Czekanowskia favours an alliance with the Ginkgoales. Araucariopitys is regarded by Jeffrey as ‘unquestion- ably Araucarian’ though ‘nearer in structure to the Abietineae than any other known Araucarian genus living or extinct.’ The chief Araucarian feature would appear to be the occurrence of flattened tracheal pits, but Jeffrey points out that not only are the pits in a single row and sometimes separate, but even when in double rows the pits may be opposite. The other characters, e.g., the pitting of the medullary-ray cells and the presence of resin-canals, are more Abietineous than Araucarian?. In view of the occasional occurrence in Abietineous genera of con- tiguous and alternate pits on the tracheids it is open to doubt whether there are adequate grounds for assuming a definite Araucarian affinity. Araucariopitys is one of several genera described by Jeffrey and other American authors exhibiting features shared by recent Araucarineae and Abietineae which are claimed as evidence of the greater antiquity of the Abietineous type. In this genus the balance of evidence would seem to be in favour of an Abietineous alliance, the tendency towards an Arau- carian pitting of the tracheids being reminiscent of ancestral types in which that character was more pronounced. XI. PROTOCEDROXYLON. Gothan. Protocedroxylon araucarioides. Gothan. The type-species of the genus founded on Upper Jurassic wood? from the Esmarks Glacier, Spitzbergen, is one of the most striking examples of a group of generalised types from Upper Jurassic strata especially from the Arctic regions. The generic name emphasises the Abie- tineous characters while the specific term gives expression to the presence of Araucarian features. The following account is based chiefly on the description by Gothan, and a few additional facts are taken from an account of some specimens from Liassic and Oolitic rocks on the Yorkshire coast by Prof. Jeffrey? and Miss Holden?®. 1 See page 63. 2 See also Gothan (10) p. 30. 3 Gothan (10) p. 27, Pls. v., vr. 4 Jeffrey (12) p. 533, Pl. 1. figs. a, b. 5 Holden (13?) p. 538, Pl. xu. figs. 17—21. XLIV] PROTOCEDROXYLON 237 Annual rings well marked; bordered pits on the radial walls of the tracheids in 1—3 rows; in the Spitzbergen wood these are from 20 to 24 in height, dimensions larger than in recent Arau- carineae, and in the type-specimen the pits are always contiguous, more or less flattened and alternate—that is Araucarian; in the English specimens the pits when in a single row are often separate and circular but equally often contiguous. Jeffrey points out that the alternate pits when in more than one series are less crowded than in Araucarian wood. There are no Sanio’s rims. The medullary rays are uniseriate and characterised by Abietin- eous pitting on the horizontal and tangential walls; on the radial walls there are 1—3 circular, apparently unbordered, pits in the field. Xylem-parenchyma is practically absent. An interesting feature is the abundance both in the Spitzbergen and Yorkshire material of tyloses in many of the tracheids, a feature occasionally met with in recent Conifers? as in some other fossil species. The pitting of the tracheids in the type-specimen may be described as exclusively Araucarian, but in the English specimens separate pits also occur though on the whole the Araucarian type is domi- nant. The pitting of the medullary rays is on the other hand definitely Abietineous. The American authors, particularly Miss Holden}, consider that the absence of Sanio’s rims suffices to tip the balance on the Araucarian side. On most of the tracheids the crowding of the pits precludes the occurrence of Sanio’s rims and in other cases their absence is not necessazily an original feature. Abietineous pitting is recorded by Jeffrey in the cone- axis of an Agathis and it has also been found in Araucaria; but in the Araucarineae it is very exceptional: its occurrence as a constant feature in Protocedroxylon may be regarded as an indica- tion of Abietineous t1elationship. No substantial assistance is afforded by impressions in Spitzbergen rocks: the abundance of Elatides is consistent with the occurrence of Araucarian wood, but impressions of Abietineous Conifers afford at least as strong an argument in favour of the occurrence of Abietineous wood. Protocedroxylon scoticum (Holden). This species, described by Miss Holden? under the generic name Metacedrozylon from Coral- lian beds on the Sutherland coast of Scotland, is founded on a piece 1 See page 178. 2 Holden, R. (15). 238 CONIFERALES (cH. of stem showing 75 rings of growth. There are no resin-canals and no xylem-parenchyma; the tracheids have uniseriate bordered pits on the radial walls only and they are almost invariably com- pressed by mutual contact; the presence of a torus is a feature characteristic of the Abietineae and not of the Araucarineae. Bars of Sanio are present but there are no rims of Sanio. Tyloses are abundant in the tracheids. The medullary rays are 2—20 cells deep, generally uniseriate though occasionally biseriate; the pitting is of the Abietineous type. In the abundance of tyloses and in other characters the wood resembles Protocedroxylon araucarioides Goth. a species transferred by Miss Holden to Metacedroxylon, but it differs in the absence of tangential tracheal pits and in the occurrence of biseriate medullary rays. Some fossil wood, which is not very well preserved, from Middle Cretaceous rocks near Iefren in the Gulf of Tripoli is de- scribed by Negri as Protocedroxylon Paronat+. The tracheal pits where biseriate are often alternate and compressed; rims of Sanio are absent: the latter feature, deduced from negative evidence, and considering the state of preservation, is surely of little value. The presence of Araucarian pitting on the tracheids in several, Jurassic species is far from surprising in view of the prevalence of that type of pitting in Palaeozoic stems; moreover an ad- mixture of characters is a natural result of progressive develop- ment. It is a matter of opinion with 1egard to the relative value of tracheal or medullary-ray pitting whether Protocedroxylon should be placed nearer to the Araucarineae or to the Abietineae. Miss Holden? discards the name Protocedroxylon for Metacedroxylon on the ground that the former implies Abietineous affinity, a fine shade of difference that hardly gives adequate expression to her conclusion that ‘Metacedroxylon araucarioides cannot be other than an Araucarian Conffer.’ XII. XENOXYLON. Gothan. Gothan? instituted this generic name for some Upper Jurassic wood, originally described by Cramer? from Green Harbour, 1 Negri (14) p. 340, Pl. v. figs. 1—6; PI. vi. figs. 1—3. 2 Holden, R. (14) p. 538 ® Gothan (05) p. 38. 1 Cramer (68) Pl. xt.; Schroeter (80) p. 7. XLIV] XENOXYLON 239 Spitzbergen, as Pinites latiporosus in order to give expression to the combination of distinctive features in both the tracheal and medullary-ray pitting. The most striking characteristics are the very large size of the bordered pits of the tracheids, the occasional (X. phyllocladoides) or constant (X. latiporosum) occurrence of contiguous and vertically flattened and transversely elongated pits on the radial walls, the absence of pits on the transverse and tangential walls of the medullary-ray ‘cells, and the presence of large simple pores on the lateral walls. There is no definite evidence as to the nature of the foliage, though Nathorst! has suggested a possible connexion between Elatides and Xenozylon. Xenozylon would seem to have been widely distributed in later Jurassic floras. Xenoxylon conchylianum Fliche. Fliche? refers to Xenoxylon a piece of wood fai the Muschel- kalk of the Vosges characterised by (i) the occurrence of uniseriate bordered pits compressed above and below and occupying the whole breadth of the tracheids and (ii) a single elliptical pit in the field. The regularity of the pores in the medullary-ray cells and the fact that they are most clearly shown where the preservation is best favour the conclusion that they are an original feature. The medullary rays are usually from 5 to 10 cells in depth. There are » no resin-canals and no xylem-parenchyma. Fiche states that the pits on the tracheids are rather less flattened than in X. lati- porosum and they occupy a greater breadth of tracheal wall. The photographs accompanying the description are unfortunately too small to show the important characters. The annual rings are faintly marked and the summer elements are confined to 4—5 rows. Despite the resemblance between this Triassic species and those previously described it is by no means certain that Fliche’s species is generically identical with the younger types. Large simple pores occur in the medullary-ray cells of recent Conifers belonging to different families, and it is not uncommon to find the bordered pits on the radial walls of tracheids in contact and slightly flattened in wood normally characterised by circular and separate pits.. In 1 Nathorst (97) p. 42. 2 Fliche (10) p. 232, Pl. xxru. figs. 4—5. 240 CONIFERALES [cH. this connexion it is noteworthy that Fliche states that the pits in his wood are occasionally circular. Xenoxylon latiporosum (Cramer). Gothan’s examination of the specimens on which Cramer founded this species! enabled him to confirm the main points of the original description: he regards Ciamer’s species Pinites pauctporosus as identical with the type-species with which he also identifies Araucarioxylon koreanum Felix?. Xenoxylon latiporosum is characterised by the large size of the pits on the radial walls of Slo 5 Yo Jef] CD) () S TST ESTs Ss eS TOD ONION Mou 2 AIS] |S E (| | IS (oe) ed |© . A Lap) See EO aes QO) OTe LOT OTS Sigeite C 2) OVO Fic. 730. A, Xenozxylon phyllocladoides. B,C, Xenoxylon latiporosum. (A, after Gothan; B, C, after Cramer.) the tracheids, 20—40p broad and 15—20w high, their vertically flattened form (fig. 730, B) and their occurrence in one or two con- tiguous rows, the pits of double rows being generally opposite. The medullary rays are uniseriate, reaching 17 cells in depth, character- ised by the narrow form of the cells, the absence of pits on the horizontal and vertical walls and by the presence of large simple pores on the lateral walls, usually one pore in the field (fig. 730, C) which it almost fills, or occasionally two; there is no xylem- 1 Gothan (10) p. 23, Pls. rv., v. > Felix (87) Pl. xxv. fig. 1. XLIV] ‘XENOXYLON 241 parenchyma but tylose-like cross-walls occur in some of the xylem elements. The nature of the pitting led Kraus? to include Cramer’s species in Araucarioxylon and Miss Holden?, who records this species from the Yorkshire coast, regards the absence of Sanio’s tims as evidence of Araucarian affinity. The medullary-ray pitting is, however, very different from that in recent Araucarineaé and the absence of Sanio’s rims may well be a natural consequence of the crowded arrangement of the tracheal pits. Xenoxylon phyllocladoides Gothan. This species, founded on material from the Bathonian of Russian Poland$, differs from X. latiporosum in the not infrequent occurrence of separate and circular pits on the tracheids: in it are included specimens from Liassic rocks at Gallberges near Salzgitter in Germany described by Conwentz* as Araucarioxylon latuporosum (Cram.) and, with some hesitation, Cupressinoxylon Barberi Sew.® from the Yorkshire coast. The tracheal pits are uniseriate, flattened or separate and circular (fig. 730, A), or in two rows, generally though not invariably opposite; they vary in size from 22 x 30p to 24 x 36; the medullary rays are generally less than 10 cells deep and in pitting agree with those of the type-species. This species is recorded from Poland, Spitzbergen, King Charles Land§, Yorkshire, and Germany. Though similar to Araucaria and Agathis in the flattened con- tiguous pits, Xenozylon differs in the elliptical form of the border and pore, also in the occurrence of separate and circular pits and in the occurrence of opposite pairs. In the form of the pits on the tracheids Xenoxylon resembles the Palaeozoic species Dadoxylon _protopityoides Fel.” and pits of similar form occur in the wood of the recent Magnoliaceous plant Drimys Winteri®. From the Abietineae the genus is distinguished by the restriction of the medullary-ray pitting to the radial walls, though the large pores 1 Kraus in Schimper (72) A. p. 384. 2 Holden, R. (14) p. 536, Pl. xxxix. figs. 5, 6. 3 Gothan (062) p. 454, fig. 4; (10) p. 36, Pl. vi. figs. 9, 10, ete. 4 Conwentz (82) p. 170 5 Seward (04) B. Pl. vi.; Holden, R. (14) p. 535. 5 Gothan (087) p. 10, figs. 3—9. 7? Felix (86) A. Pl. v. fig. 4. 8 Groppler (94) Pls. 1., 1. ' $. IV 16 242 CONIFERALES [cH. (fig. 730, C) in the field resemble those of some Pines and other Abietineae as also those of Sciadopitys (fig. 693, N): in Xenoxylon there are no resin-canals and no xylem-parenchyma. Gothan considers that while differing in the sum of its characters from any other type of Conifer, Xenoxylon shows most resemblance to Gothan’s genus Phyllocladoxylon (= Mesembrioxylon Sew.), a genus including fossil species which suggest affinity not only with Phyl- locladus but with other members of the Taxaceae. In his account of Xenoxylon phyllocladoides from King Charles Land Gothan describes instructive examples of the effect of the action of fungal hyphae on the structure of tissues. The genus may be described as a generalised type exhibiting features shared by the Araucarineae and Taxaceae. XII. ANOMALOXYLON. Gothan. Anomalozylon magnoradiatum Gothan. Gothan proposed this name for some Upper Jurassic wood from Spitz- _ bergen? which cannot be definitely assigned to a family-position: its most striking feature is the occurrence of large spindle-shaped medullary rays containing a large ‘canal,’ or spaces lined by a single layer of cells (fig. 731). The rings of growth are well marked, the summer-wood being composed of a very few rows of tracheids in abrupt juxtaposition to the larger spring elements, a character associated with roots. There are no vertical resin-canals and no regular o7 typical horizontal canals. Xylem-parenchyma is rare or absent. The bordered pits on the radial walls of the tracheids are in 1—2 rows, separate and circular or, more frequently, con- tiguous and more or less flattened but, if in two Fae se ne rows, not alternate. The medullary rays are yin magnoradi- uniserlate, generally 5—7 cells deep, or in places atwm. Medullary forming broad and deep spindle-shaped areas Y showing the : eon : small cells and a either empty or containing a large circular canal- eain canal. (After like passage. These peculiar rays, as seen in a Gothan.) tangential section of the wood, are a conspicuous feature and are 4 See page 203. * Gothan (10) p. 10, Pl. 1. figs. 9—11; Pl. m. figs. 2, 3. XLIV] THYLLOXYLON 243 often lined by a layer of small cells (fig. 731). In the presence of these large medullary rays Anomaloaylon resembles Thyllozylon, but in the latter genus the rays are smaller and more uniform in size. Gothan discusses the nature of these medullary rays and inclines to the view that they agree more closely with abnormal or traumatic formations in certain Conifers than with any normal structures. There are no pits on the horizontal or tangential walls of the ray cells and there are 2—3 simple circular pits in the field. The general impression gained from an examination of Gothan’s photographs is that no true canals occur, and that the peculiar medullary rays owe their form to partial decay. of abnormal patches of parenchyma possibly produced as the result of wounding. Though on the whole nearer in structure to the Taxodineae? than to any other family Anomaloxylon is a type which cannot be assigned to a definite position. XIV. THYLLOXYLON. Gothan. Thyllocylon irregulare Gothan. The generic name Thylloxylon was given to a single species of Upper Jurassic age from Spitz- bergen? on account of the occurrence of tiillen-like parenchyma in horizontal canal-like spaces in some of the larger medullary rays. The wood is characterised by separate bordered pits in the summer tracheids and 1—2 rows of alternate contiguous, Arau- carioid, pits on the spring elements; xylem-parenchyma occurs only at the end of the year’s growth. The medullary cells have Abietineous pitting and there are 2—3 small circular, apparently simple, pits in the field, or occasionally only one in the region of the late wood. The rays are uniseriate or 2—3 cells broad and some medullary rays closely resemble those of certain Abietineae possessing horizontal resin-canals; but in Thylloxylon there are no true canals. The central parenchyma of some of the broad rays is replaced by a canal-like space and these spaces are often filled with spherical tiillen-like tissue, a feature shared with Anomalo- aylon, but in that genus there is no Abietineous pitting on the medullary-ray cells. There are no vertical resin-canals. 1 That is Athrotaxis, Cryptomeria, Sequoia, Taxodium, and Sciadopitys, genera which are now (see page 126) assigned to different families. 2 Gothan (10) p. 34, Pl. vi. figs. 2—8. 16—2 244 CONIFERALES [CH. XLIV XV. PLANOXYLON. Stopes. Dr Marie Stopes? has recently instituted this generic name for a piece of Coniferous wood from Middle (or Upper?) Cretaceous rocks in New Zealand and in it she also includes the Liassic species Araucariozylon Lindleii (Witham). The genus is a striking ex- ample of a combination of Araucarian and Abietineous characters?, and, as Dr Stopes points out, it resembles in this respect Cedroxylon transiens Goth. and other generalised types. Planoxylon Hectori Stopes. The type-specimen, from Amuri Bluff, New Zealand, is part of a stem 150 years old or more. The rings of growth are well marked; the tracheids have 1—3 rows of alternate and hexagonal bordered pits on the radial walls and there may be a single row of separate pits on the elements at the end of an annual ring. The medullary rays are nearly always uniseriate, 1—24 cells deep but usually from 3 to 9 cells in depth; all the walls of the ray cells are pitted and there are 1—2 vertical rows of three pits in the field in the neighbourhood of the spring tracheids and generally a single vertical pair in the region of the late wood. Xylem-parenchyma appears to occur only between the spring tracheids and the latest formed wood of the previous year. Like many other fossil stems this species indicates the existence of Conifers with typical Araucarian pitting on the tracheids and equally well defined Abietineous pitting on the medullary-ray cells. It is especially interesting as showing the presence in the southern hemisphere of a type very similar to Cedroxylon transiens and other species recorded from high northern latitudes. Planozylon Lindlew (Witham). This Liassic species from Whitby was originally referred by Witham to the genus Peuce; subsequently included in Araucario- zylon* it has recently been transferred by Dr Stopes to her new genus Planozylon®. The pitting of the tracheids is essentially Araucarian; there are 1—3 rows of alternate hexagonal pits on the radial walls, but the pitting of the medullary-ray cells, as Dr Stopes has shown, is typically Abietineous. 1 Stopes (16) ° mAavdouat, to wander; suggesting that ‘the forms comprising the genus were moving from one position to another in a systematic sense.’ 3 Witham (33) A. p. 58, Pls. 1x., xv. 4 Seward (04) B. p. 56, Pls. v1., vu. 5 Stopes (16) pp. 118, 120, text-figs. 6, 7. CHAPTER XLV. CONIFERALES. SHooTs, CoNE-SCALES, ETC. TuE majority of the vegetative and fertile shoots, cone-scales, seeds, etc., selected in illustration of the past history and geographi- cal distribution of the Coniferales are described under the different families enumerated on page 124. Under each family are included not only specimens which, with a fair amount of confidence, can be assigned to a family-position but also genera of doubtful affinity which it has been contended afford evidence of greater or less value in favour of an alliance with the family under which they are described. The inclusion of certain genera in a chapter or section devoted to a particular family does not necessarily mean that they show clear evidence of relationship to that family: many of the genera might with equal propriety be relegated to Chapter L, which is devoted to Conzferales incertae sedis. On the other hand some genera included in that category would by other authors be given a place in the Araucarineae or some other family. I have endeavoured to state the different views expressed by authors with regard to the affinity of imperfectly known genera, but in many instances the available data do not afford any trust- worthy evidence of relationship to existing types. ARAUCARINEAE. DAMMARITES. Presl. PROTODAMMARA. Hollick and Jeffrey. The distinctive characters of the recent genus Agathis (Dammara) are briefly described in Chapter XLIII. Fossil records bearing on the past history of Agathis are more meagre and more difficult of interpretation than those relating to Araucaria. The evidence at present available points to the greater antiquity of Araucaria at least as regards the type of cone characteristic of that genus. 246 ARAUCARINEAE [CH. On the other hand the type of foliage-shoot represented by existing species of Agathis—without taking into account the Palaeozoic leaves assigned to Cordaites, some species of which bear a close superficial resemblance to those of certain examples of the recent genus—is widely represented in Rhaetic and Jurassic floras by Podozamites!. There is, however, no proof that Podozamites was nearly related to Agathis, and, indeed, such information as we have with regard to the reproductive organs of that genus does not point to any very close Araucarian affinity. Fossil wood gives no help towards a distinction between the two members of the Araucarineae nor do impressions of vegetative shoots materially aid us. . Palaeobotanical literature contains a few records of leaves referred to Dammara or Dammarites but in no case is there any conclusive evidence of generic identity of the fossils with the recent genus. Leaves from Lower Cretaceous rocks in Bohemia described by Velenovsky and by Fri¢ and Bayer? as Dammaro- phyllum striatum and D. bohemicum exhibit a close agreement in shape and venation with those of some species of Agathis, though they differ but slightly from some forms of Podozamites, e.g., P. Reinii Gey). (fig. 814, p. 456). Other leaves that may belong to plants similar to Agathis are represented by Dammarites caudatus and D. emarginatus Lesq. from the Dakota series®: these, prob- ably specifically identical, forms present, as Lesquereux says, a close resemblance to Agathis robusta. Itis impossible without additional data to determine the true position of these and similar leaves though it is permissible to regard them as possible examples of the foliage of Conifers closely allied to Agathis. Similarly, some detached leaves from Cretaceous and Tertiary strata referred to Podozamites may well be more akin to Agathis especially in view of the fact that Podozamiates is essentially a Rhaetic and Jurassic genus. The leaves figured by Saporta* from Lower Cretaceous beds in Portugal as Podozamites ellipsoideus agree closely with those of Agathis. In the case of separate linear leaves like those described by Hollick® from the Cretaceous of Long Island as 1 See page 447. 2 Fric and Bayer (01) B. p. 96. 3 Lesquereux (91) p. 32, Pl. 1 figs. 9—11. 4 Saporta (94) B. Pl. xxxm. fig. 5; Pl. xxxv. fig 12. 5 Hollick (12) PJ. 163, figs. 2, 3. XLV] DAMMARITES 247 Podozamites lanceolatus the term Desmiophyllum would be a more appropriate generic designation, the name Dammarites being adopted for broader forms. This distinction is purely arbitrary and it must be admitted that there is no substantial justification for the use of a generic name implying affinity with Agathis. Unless there are adequate grounds for assuming generic identity of detached Tertiary and Cretaceous leaves with Podozamites it is inadvisable to make use of that designation. As Schenk?! points out Velenovsky’s Tertiary species Podozamites miocenicus may be a leaf of Agathis or possibly a Podocarpus. Dammarites Bayer Zeiller. This name was given to some oval-lanceolate leaves from Upper Cretaceous beds in Bulgaria varying in length from 10 to 12 cm. and from 15 to 30 mm. broad agreeing closely with Heer’s Podo- zamites marginatus from the Cenomanian of Greenland but wisely excluded by Zeiller? from that genus, though on grounds which are no longer cogent if the interpretation of Podozamites impressions as shoots and not pinnate leaves is accepted. Ettingshausen? records two species of Dammarites from Tertiary rocks in New Zealand: Dammarites Oweni includes in addition to leaves a cone-scale, the impression of a cone, and some petrified wood of the Araucarian type. There is no proof that these disjuncta membra belong to the same plant though it is not improbable that they are parts of a Conifer closely allied to Agathis. Ettingshausen’s second species D. wnivervis is founded on a leaf and a supposed cone-scale of doubtful value. The data furnished by leaves alone are of little value. In addition to the cone described from New Zealand by Ettingshausen other examples are recorded as species of Dammarites but without any satisfactory evidence of affinity to the recent genus, ¢.g., Dammarites albens Presl.4 from the Quadersandstein of Bohemia and D. crassipes Goepp.® These two species are united by 1 Schimper and Schenk (90) A. p. 279. 2 Zeiller (05?) p. 17, Pl. vu. figs. 8—11. 3 Ettingshausen (87) p. 15, Pl. 1. figs. 20—24. 4 Sternberg (38) A. Pl. ru1.; Corda in Reuss (46) B. Pl. xtvu.; Goeppert (50) p. 237; Schimper and Schenk (90) A. p. 279, fig. 292 b. 5 Goeppert (50) Pl. xtv. fig. 6; Corda in Reuss (46) B. 248 ARAUCARINEAE [cH. Velenovsky under a single type which he calls Krannera mirabilis? from a name suggested by Corda: additional examples superior in preservation to those previously figured are illustrated in Velenovsky’s memoir on the Bohemian Cretaceous Gymnosperms. Velenovsky regards the supposed cones as stems bearing crowded woody scales which originally had long Cordaites-like leaves at- tached to a transverse ridge just internal to the thickened distal ends: there appears to be no absolute proof in support of this connexion between scales and foliage-leaves, but one specimen figured shows portions of leaf-like organs attached to two of the scales, though these may be petiolar and not pieces of laminae. Reference is made elsewhere to the Krannera leaves. It is probable that, as Velenovsky believes, the supposed cones are stems similar to tuberous Cycadean species but it is doubtful if they were pro- vided with leaves of the type included in Krannera mirabilis. Schmalhausen? figures a Tertiary cone from Russia as Dammara Armaschewskii which in the form of the distal ends of the scales resembles Araucaria brasiliensis and species of Pinus, but some detached scales agree closely in shape and in the possession of a single seed with those of Agathis. Small detached cone-scales of Tertiary age, described by Schmalhausen® as Dammara Tolli from the New Siberian Islands, (fig. 732), may be allied to Agathis ; they agree generally with those of D. borealis and other western types. It is, however, from detached cone- Beales Obtammed “ront: Urebatee ns 4 ano, iaemmutun Tau (At strata in Greenland and some Schmalhausen; nat. size.) European localities but especially from the Eastern United States that the most promising in- formation has been gained. Hitchcock first recorded these scales from Martha’s Vineyard and spoke of them as ‘seed-vessels’ of some Coniferous plants?, but it was Heer> who compared them with the cone-scales of Agathis. The latter author described several examples from Cenomanian strata in West Greenland as Dammara borealis, D. microlepis, etc. 1 Velenovsky (85) B. p. 1, Pls. 1.,1v. |? Schmalhausen (83) p. 313, PL xxxv1. 3 Ibid. (90) p. 14, Pl. 1. fig. 19. 4 Hollick (06) p. 38. 5 Heer (82). XLV] DAMMARITES 249 Dammarites borealis (Heer). Though it is clearly impossible to define with any precision the limits of species based on detached scales varying considerably in size and shape, several types have been recorded, particularly from different localities on the Atlantic Coastal plain of North America’. The larger forms may con- veniently be, included in Dammarites borealis Heer and smaller forms are illustrated by Protodammara speciosa Holl. and Jeff.” It is probable that these two types are generically identical, but the name Protodammara implies the presence of certain structural features while Heer’s species is founded on casts or impressions. A specimen of the latter species from Greenland is shown in fig. 733, the scale is 22 mm. broad and is characterised by several parallel lines, either vascular bundles or resin-canals, and the white patches represent some exuded — f : - Fia. 733. Dammarites bo- resinous material. Other Greenland ex- ““)) 7.” Gone-scale from amples are more elongated basally and are —Igdlokungnak, West identical in shape with the smaller scales Greenland. (Stockholm from Staten Island seen in fig. 758, E, F, Mupsisray ABU Eiee) page 323. Fossils of similar form were described by Heer from the same locality as Eucalyptus Geinitzii?. Krasser* and some other writers have retained the generic name Eucalyptus on the ground of association with Eucalyptus-like leaves. Hollick speaks of scales like D. borealis as ‘among the most abundant and charac- teristic remains found in the Cretaceous deposits of America and Europe®’: he adds that the name Dammarites is chosen for the sake of convenience rather than from a conviction that it represents their true generic relationship. Newberry in describing this type from the Amboy clays states that some of the scales have grooves, corresponding to the dark lines in fig. 733, filled with amber®, and anatomical evidence derived from Protodammara supports the view that the cone-scales were rich in resinous substance. Both 1 Hollick (97) PL. x1. figs. 5—8; (06) p. 37; Newberry and Hollick (95) p. 46. 2 Hollick and Jeffrey (06) p. 199, PJ. 1. figs. 5—13; PI. 1. figs. 1—5. 3 Heer (82) p. 93. 4 Krasser (96) B. Pl. xvi. fig. 6. 5 Velenovsky (89) Pl. 1. figs. 28, 29. 6 Newberry and Hollick (95) p. 47 and see also Berry (07). 250 ARAUCARINEAE [cH. large and small cone-scales are recorded by White1, Berry?, Knowlton, and especially by Hollick from several places on the Atlantic Coastal plain. Knowlton’s species Dammarites acicu- laris? is probably identical with D. borealis. As examples of smaller forms reference may be made to D. northportensis*, D.. minor®, and the scales now included in Protodammara. These numerous scales occur as detached specimens and without seeds, but their resemblance to the cone-scales of Agathis and the ana- tomical features exhibited by the lignitic examples described by Hollick and Jeffrey afford strong arguments in favour of an affinity to Agathis. We have no proof as to the nature of the vegetative shoots of the parent-plants. Newberry states that in the Amboy clays the scales occur in association with shoots like Heer’s Juniperus macilenta, in some cases, apparently, attached. On the other hand Krasser considered the association of the speci- mens from Moravia, which he called Eucalyptus Geinitzw, with dicotyledonous leaves as evidence of original connexion. Protodammara speciosa Hollick and Jeffrey. It is probable that these scales® are generically identical with the larger forms referred to D. borealis and other species, ¢.9., D. cliffwoodensis’, but until anatomical evidence is obtained a distinction should be recognised. The type-specimens were found at Kreischerville, Staten Island, New Jersey, in Middle Cretaceous beds, and the species is thus defined:—kite-shaped cone-scales from 4 to 6 mm. long by 4—6 mm. broad above, abruptly narrowed from about the middle to the base (fig. 758, EK, F, page 323), rounded, incurved, and apiculate above; resin-ducts five or more, extending down the lower surface of the limb; seed-scars three in number, crescentically arranged above the middle and approxi- mately in the broadest part of the scale, with the central one higher up than the laterals. Near the base of a scale there is a single vascular bundle with the xylem uppermost: at a higher level a single strand with reversed orientation is given off and the original 1 White (90) p. 97. 2 Berry (03) Pl. xiv. figs. 8—11. 3 Knowlton (05) Pl. xv. figs. 2—5. F 4 Hollick (04) p. 405, Pl. uxx. figs. 1, 2. 5 Ibid. (12) Pl. w. figs. 35—37. ® Hollick and Jeffrey (06); (09) B. p. 46, Pls. 1v., x., XIV.—xvI. 7 Hollick (97) Pl, x1. figs 5—8. XLV] PROTODAMMARA 251 bundle divides into three. In the lower portion of the scale there are seven resin-canals and above these is a band of transfusion- tracheids surrounding and connecting the vascular bundles. In median longitudinal section a scale shows a terminal spinous process similar to that in the scales of Conites Juddi} (fig. 734); on the adaxial side of this the scale is swollen and internal to the swollen part is a small pit marking the position of the middle of the three seeds. The upper surface of the scales is covered with periderm and stone-cells occur in the ground-tissue. In anatomical characters Protodammara resembles the scales of recent Arauca- rineae more closely than those of any other Conifers, and in the absence of a definite ligule and in the relation of the seeds to the scale the fossil scales are similar to those of Agathis. In the description of a Scotch Upper Jurassic cone, Conites Juddi, attention is called to a close resemblance in anatomical features to Protodammara. The American scales occur in association with shoots of the type represented by Brachyphyllum macrocarpum — Newb.? (=B. crassum) (fig. 758, G), an association noted also in other localities than Kreischerville. The structure of these shoots is described under the genus Brachyphyllum®, but as regards the scales the important point is that if this association means original connexion, the habit of the parent-plant was wholly different from that of any recent Agathis or Araucaria. Brachyphyllum macrocarpum is regarded by Hollick and Jeffrey as certainly Araucarian. Wood of the Araucarian type is also found in associa- tion with the Protodammara scales and the Brachyphyllum shoots. Considering the cone-scales by themselves, their position would seem to be next to Agathis though they differ in bearing three seeds in place of the single seed in the recent genus: the number of seeds borne on the larger scales such as D. borealis is not known. Protodammara affords an interesting illustration of the co-existence of characters now characteristic of the Araucarineae with others no longer exhibited by members of that family: assuming a con- nexion between Brachyphyllum macrocarpum and the cone-scales, the habit of the vegetative shoots furnishes a further illustration of a wider range in the morphological features of fossil Conifers allied to existing Araucarineae. 1 See page 252. 2 Hollick and Jeffrey (09) B. p. 33. 3 See page 322. 252 ARAUCARINEAE fou. Contes Juddi Seward and Bancroft. This name was given to partially petrified cones of Upper Jurassic age collected by Hugh Miller on the North-east coast of Scotland!: the fossils though differing in size and to some extent in form are included under one specific term but distinguished as Fig. 734. Conites Juddi. A, forma y; ab, space where a section was cut; s, seed. B, forma a; s, spaces, probably resin-canals, filled with crystalline materials and simulating seeds. C, forma 8; ab, ridge. forma a, B, y, and 6 (fig. 734). The type-specimens are in the Royal Scottish Museum, Edinburgh. Spirally disposed thick scales are attached by a comparatively narrow base to a thick axis and the individual scales agree closely in shape with those described as Dammarites borealis and with smaller forms referred ? Seward and Bancroft (13) p. 873, Pl. 1. figs. 9—12; Pl. 1. figs. 14—21. XLV] CONITES 253 by Hollick and Jeffrey to Protodammara. The parenchymatous tissue of the cone-scales contains several thick-walled idioblasts and thé resin-canals and spaces form a conspicuous feature. A series of vascular bundles runs radially through the scale, but no evidence has been obtained of the occurrence of a double set of vascular strands like those in Protodammara. The seeds—the number of which, whether one or more, cannot be determined— lie in a depression near the proximal end of the scales and there Fic. 735. Conites Juddi, forma a; cone-scale in median section; 1, ligule; s, s, (2) resinous material; vb, vascular bundles. ( x 6.) is a ligule on the abaxial side of the seed or seeds (fig. 736, B, 1). The uppermost scales on the two cones shown in fig. 734, B and C, illustrate the striking similarity to such detached scales as those of Dammarites borealis: the raised patches, s, s, simulating seeds, are formed by a crystalline substance filling cavities in the scales and probably corresponding to the resin-ducts which form a charac- teristic feature in the scales of Dammarites and Protodammara. A cone-scale from the specimen represented in fig. 734, B shows 254 ARAUCARINEAE — [ou. in longitudinal section (fig. 735) a large cavity in the lower part of the scale containing at each end a dark patch of some secreted — substance s, s; above this is a vascular strand vb, extending into the distal end of the scale near which is a ligular outgrowth J, and below this is a depression on the upper face of the scale in which Fic. 736. Conites Juddi, forma 6. A, Longitudinal section of cone; 2, xylem, d, seed, e, f, cone-scales. B, Scale in longitudinal section; c, vascular bundle; a, periderm; 5b, palisade-tissue; J, ligule; s, seed. (After Seward and Bancroft.) a seed was originally situated. The cones shown in fig. 734, B, C, have lost their seeds and indicate a persistent habit in contrast to the cones of Agathis and, presumably, the cones which possessed scales like Dammarites borealis. The cone seen in fig. 734, A, is probably younger; the scales are more crowded and in one of XLV) CONITES 255 them is the cast of a seed, s. One of the scales, 1-7 cm. long, of this cone is represented in section in fig. 736, B: a vascular bundle, c, runs through the length of the tissues towards the blunt spinous distal end above which is a prominent hump and next to this a ligule, /, close to the depression in which the seed, s, was situated. Below the vascular bundle, c, a band of periderm surrounds a central area of decayed tissue, a, a. Next the lower surface at b are a few layers of palisade cells, a characteristic feature. The specimen shown in section in fig. 736, A, is described as forma 8: the secondary xylem, z, of the axis is not sufficiently well pre- served to throw any light on the nature of the tracheal pitting. The large (white) cavities at first sight suggesting seeds near the axis are bounded by periderm and correspond to the partially destroyed tissue in fig. 736, B: a delicate structure, the nature of which could not be determined, occurs in the cavity d, fig. 736, A. In the scale e several smaller cavities are seen near the upper face above the vascular strands and below the latter is a larger cavity ; similar cavities are shown in the scale f (fig. 736, A). Cones similar to Conites Juddi are described by Velenovsky1 as Friéia nobilis and Sequoia fastigiata, both from Lower Cretaceous strata, but it is impossible to say whether the resemblance has any significance. In several anatomical characters the scales of Conites Juddi resemble those of Protodammara described by Hollick and Jefirey from Kreischerville. The only indication of pits on the xylem tracheids in the Scottish cones was seen in the scale shown in fig. 736, B: the pits are for the most part uniseriate but occasionally contiguous though generally not actually in contact. Large idioblasts and resin-cavities occur in both the Scottish and American cones, but in the former the occurrence of a ligule is a distinguishing feature in which they agree with cone-scales of recent Araucarias. In Araucaria the seeds are embedded in the substance of the scales while in the fossil species they are situated in a depression on the upper face, a feature in which Conites Juddi agrees more closely with the cones of Agathis. In the sporophylls of Conites Juddi, which anatomically are close to those of recent Araucarineae, characters occur which are now shared between Araucaria and Agathis. The apparently small size of the seeds 1 Velenovsky (85) B. Pl. 11. fig. 6; Pl. virr. fig. 13. 256 ARAUCARINEAE [CH. and their relation to the ligular outgrowth, as well as the occurrence of separate bordered pits on the tracheids suggest comparison with the recent genus Cunningham, though the structure of the scales is more akin to that of Araucarian sporophylls. The com- bination of features which are now distributed among different genera is to be expected in extinct types belonging to evolutionary stages anterior to the divergence of characters along independent lines. The main conclusion is that the affinities are Araucarian though the morphological characters are such as to indicate a com- bination of features no longer found in a single genus. Cones exhibiting a close resemblance to those of Araucaria. ARAUCARITES. Presl. Araucarites sphaerocarpus Carruthers. This species (fig. 737), from Inferior Oolite rocks at Bruton, Somersetshiret, affords a good example of a large Araucarian cone 13 cm. in diameter very similar in form to some recent species (cf. fig. 680 and fig. 681). The rhomboidal scales, 2 cm. broad at the distal end, are laterally winged as in Araucaria Cookui (fig. 638, A) and bear a single seed embedded in the middle of the upper surface: on the exposed distal ends is a transverse groove and on some of the more complete examples a short rounded umbo is seen below the groove; in some scales a. transverse row of pits marks the position of vascular bundles just below the transverse depression. Araucarites ooliticus (Carruthers). This species was originally described by Carruthers as Kazda- carpum ooliticum? from the Great Oolite of Northamptonshire and regarded as an inflorescence of some Pandanaceous plant. Zigno® transferred it to Pandanocarpum. An examination of the type- specimen in the Northampton Museum led me to refer the cone to Araucaritest. The type-specimen (fig. 738) is a portion of a cone 9 cm. long consisting of a stout central axis covered with spirally disposed deep pits bounded by a crystalline reticulum; the pits 1 Carruthers (66) Pl. x1.; Seward (04) B. p. 131; (113) p. 116, fig. 18. 2 Carruthers (68) p. 156, Pl. rx. 3 Zigno (85) p. 3. 3 Seward (96?) p. 216; (04) B. p. 135. XLV] ARAUCARITES 257 Fig. 737. Araucarites sphaerocarpus. (British Museum, 41,036; 3 nat. size.) yt: Fic. 738. Araucarites ooliticus. (After Carruthers; 3 nat. size.) 8. IV 17 258 ARAUCARINEAE [CH. being cavities in the proximal portion of the scales in which the seeds were embedded. Numerous imbricate scales are attached laterally to the central region and partially hidden in the matrix. The scales are approximately 1-7 cm. broad and slightly winged. The single seed on each scale, the general form of the cone, the shape of the individual scales, and the occurrence of sterile scales at the base of the axis are features in which the fossil is practically identical with recent forms. Fig. 739 shows a piece of a smaller Fic. 739. Araucarites ooliticus. A, Scale in surface-view showing the projecting end of a seed. B, Scale seen from the proximal end showing the seed-cavity. C, part of cone. (Northampton Museum; nat. size.) cone (in the Northampton Museum), of the same type; this specimen shows the appearance of the scales in end-view (C), in surface-view (A), and as seen from the proximal end with the seed-cavity (B). In fig. 739, A, the base of a seed is seen projecting from the middle of the laterally expanded scale. An oblong-ovate cone described by Carruthers! from the Coralline Oolite at Malton, Yorkshire, as Araucarites Hudlestoni is probably another example of this species: in one of the specimens of A. Hudlestons in the York Museum a broad central region is occupied by a mass of pisolite to which numerous cone-scales are attached. The scales are shown in section and in several of them there is a single seed lying in a cavity occupying the proximal end of the scale precisely as in A. ooltticus. It is possible that A. ooliticus is specifically identical with A. sphae- rocarpus; it is at least a closely allied type. The specimen figured by Lindley and Hutton? as Strobilites Bucklandi appears to be indistinguishable from A. ooliticus. Similar cones are illustrated by Araucarites Cleminshawi Mansell- 1 Carruthers (77); Seward (04) B. p. 133. ? Lindley and Hutton (34) A. Pl. cxxrx. XLV] ARAUCARITES ; 259 Pleydell* from the Inferior Oolite of Dorsetshire, A. sphaericus (Carr.)?, originally referred to Cycadeostrobus, and other species. Araucarites pippingfordensis (Ung.)® is a Wealden species first described by Fitton and named by Unger Zamiostrobus pipping- fordensis but recognised by Carruthers as a cone closely allied to A. sphaerocarpus. Detached scales which may belong to this species have recently been figured from the Wealden beds on the Sussex coast. The cone described as Araucarites (Conites) sp. from the same locality is possibly identical with A. pippingfordensis. Araucarites Brodiei Carruthers. The specimens from the Stonesfield slate* on which this species was founded are rather larger than the scales of A. Phillipsi: the scale shown in fig. 740, 4, is 3 cm. long and 1-7 cm. broad; a raised edge just beyond the single seed no doubt corresponds to the so- called ligule on an Araucarian scale and the distal spinous process is another feature shared with recent types. Araucarites (Sarcostrobus) Paulini (Fliche). A specimen described by Fliche® from Lower Cretaceous rocks in the Haute-Marne, France, is made the type of a new genus Sarcostrobus on the ground that the seeds are not so completely covered by the tissues of the scales as in recent Araucarias. The elliptical cone is 5-5 cm. long and 3-8 cm. in diameter; in form, in the stout axis, and in the shape of the single-seeded cone-scales it closely resembles the megastrobili of Araucaria excelsa and Jurassic species such as Araucarites ooliticus (Carr.) (fig. 738): the small seeds are sunk in a cavity at the proximal end of the - scale, but Fliche states that they are not covered on their upper side by the substance of the scale. He is no doubt correct in assigning the cone to the Araucarineae, but the slight differences between the relation of seeds to scales referred to by Fliche do not appear to be sufficiently important to justify the creation of a distinctive generic name; moreover the preseivation of the speci- mens renders accurate description of details very difficult. 1 Mansell-Pleydell (85). 2 Carruthers (67%) p. 105; Seward (04) B. p. 138. 3 Seward (13) p. 104. : 4 Carruthers (692) p. 3, Pl. v. figs. 1—6; Seward (04) B. p. 137, Pl. 1. fig. 5; Pl. x11. fig. 2. 5 Fliche (00) p. 11, PL 1. 17—2 260 ARAUCARINEAE [cH. Araucarites hespera Wieland. This type is described by Wie- land! from a specimen obtained from Upper Cretaceous rocks in South Dakota consisting of half an eroded cone bearing scales with small seeds. Wieland’s description, though brief and facking details, and the photograph support his conclusion as to the Araucarian affinity of the specimen. The generic name Doliostrobus was instituted by Marion? for specimens of foliage-shoots from Oligocene beds in the South of France agreeing with Araucarites Sternbergii, on the ground that the reproductive organs exhibit features more like those charac- teristic of Agathis than Araucaria. Laurent? refers a small piece of a foliage-shoot from the Aquitanian beds in the Puy-de-Déme to Doliostrobus Sternbergvi, though there is no information with regard to the cones. Gardner? describes branches from the Bembridge marls in the Isle of Wight as Doliostrobus Sternbergii (Goepp.) and accepts Marion’s conclusion as to the intermediate character of the genus: the foliage-shoots, though rather more slender than those from Bournemouth referred by him to Araucaria Goepperti, are exactly similar in habit and cannot be distinguished by any feature of importance. Gardner reproduces a drawing communicated by Marion of a foliage-shoot bearing a terminal cone-axis from which the scales have fallen, also several detached cone-scales, agreeing closely in size and shape with scales of Araucaria excelsa and other recent species, and an impression described as a seed with a lateral wing. The supposed seed has, however, a terminal wing and moreover it is as large as the detached scales: it is permissible to suggest that it may be an imperfectly preserved cone-scale, but without examining the actual specimen any definite assertion would be hazardous. Gardner states that a reason for comparing the scales with those of an Araucarta is that in Agathis the scales are persistent, but as pointed out elsewhere’ cones of Agathis very readily fall to pieces and the scales easily become detached from the axis. Having regard to the nature of the sterile shoots, the form of the cone, as shown in a drawing published by Gardner of a specimen sent to him by Ettingshausen from Haring, and an 1 Wieland (08°) p. 4, fig. 1. 2 Marion (84). 3 Laurent (12) Pl. v. fig. 7. 4 Gardner (86) p. 93, Pls. xxm., Xx. 5 Seward and Ford (06) B. p. 359. XLV] ' PSEUDO-ARAUCARIA 261 unconvincing specimen of a winged seed figured from Marion's ' drawing, there would seem to be no valid reason for drawing a distinction between Doliostrobus and Araucarites or for regarding Marion’s and Gardner’s fossils as intermediate between Araucaria and Agathis. Attention has been called on a previous page! to the danger of placing too much confidence in the resemblance of foliage-shoots of fossil specimens to those of recent types, but in this case the presence of cones and scales like those of Arawcaria supplies confirmatory evidence. Pseudo-Araucaria. Fliche., The generic name Pseudo-Araucaria was given by Fliche? to several cones from the Lower Cretaceous beds of the Argonne which he described under three specific names, Pseudo-Araucaria Loppinetti, P. major, P. Lamberti. Externally they are similar to those of some recent Araucarias and in shape agree with cones of Cedrus: a stout axis bears deciduous scales with two seeds, the seeds of each pair being separated from one another by a median ridge of the cone-scale which covers them laterally. The seeds appear to bear a relation to the scale similar to that between the single seed and the cone-scale of an Araucaria. The cone-scales are slightly expanded laterally as in the Hutacta section of the recent genus. Fliche’s descriptions are unfortunately inadequately illustrated and it is difficult to obtain a very clear impression of the structural features. The most interesting peculiarity of these cones is the occurrence of two seeds in each cone-scale agreeing in their position on the sporophyll with the single seed of Araucaria: the author of the genus regards it as a type intermediate between the Abietineae and the Araucarineae. Araucarian cone-scales. The question of the lower geological limit of cones or cone- scales of the Araucarian type is one which cannot be settled with any certainty: there are many examples of vegetative organs very similar in habit to Araucaria excelsa and allied species recorded from Triassic, Permian, and to a less extent from Upper Carboni- ferous strata, also others which agree in the broader form of the 1 See page 162. 2 Fliche (96) p. 70, Pl. vi. figs. 3—5; Pl. vir. figs. 1, 2. 262 _ ARAUCARINEAE [CH. leaves with Araucaria Bidwilli and A. imbricata; but the majority of these shoots are referred to such genera as Voltzia, Walchia, Albertia, and Ullmannia. It is pointed out in the description of these genera that there are reasons for believing them to have Araucarian affinities, though there is no definite evidence that any of them bore cones exhibiting the same order of resemblance to those of recent Araucarineae as is the case with Jurassic and Cretaceous types. Araucarites Delafondi Zeiller. One of the very few Palaeozoic species of seed-bearing scales that can reasonably be referred to the genus Araucarites is A, Delafondi founded by Zeiller} on some detached scales from Permian beds at Charmoy; the scales are broadly triangular 10—12 mm. long and 8—10 mm. broad, the base is cuneate and truncate, the apical margin is rounded and has a small median depression instead of the usual spine. In the middle of the scale is a shallow depression which contained a single seed 8—10 mm. long and 2mm. broad. As Zeiller says, there is no absolute certainty as to the affinity of this species but the scales are un- questionably very similar to those of Mesozoic and recent species of Araucarites and Araucaria. It is suggested that the vegetative shoots of Ullmannia frumentaria (fig. 750) from the same beds may belong to the plant which bore cones with scales of A. Delafondi. The occurrence of widely distributed Jurassic cone-scales, bearing a single seed and agreeing very closely in their shape and size, as also in the laterally expanded borders and in many cases in the presence of a distal spinous process, with those of recent species of Araucaria especially those belonging to the section Eutacta, bears striking testimony to the former extended geo- graphical distribution of Araucarian plants. It has been pointed out in a previous chapter that a single seed occasionally occurs on the seminiferous scales of recent Pine cones (fig. 686, B), but in the scales now under consideration the occurrence of a single seed is a constant feature and moreover the form of the scales is identical with that of such species as Araucaria excelsa and A. Cookit. The number of names given to the fossil scales is but a rough index of } Zeiller (06) B. p. 215, Pl. x. fig. 1. XLV] ARAUCARITES 263 the number of actual species: it is obviously impossible to decide with any assurance how much value should be attached to differ- ences in size or to slight variations in form, but the main point is that cones and cone-scales of the Araucarian type are among the most familiar Jurassic fossils. The following selected examples are chosen in illustration of this statement and reference to others will be found in some of the sources quoted in the footnotes. Araucarites Phillipst Carruthers. Carruthers! described this species from the Middle Jurassic rocks on the Yorkshire coast: the type-specimen is in the Leckenby collection in the Sedgwick Museum, Cambridge. The scales are NGA ew _ Fic. 740. 1, Araucarites Milleri (Upper Jurassic, Scotland); 2, A. Jeffreyi (Cre- taceous, N. America); 3, A. Rogersi (Wealden, 8. Africa); 4, 7, A. Brodiet (Middle Jurassic, England); 5, A. Héberleinii (Middle Jurassic, Germany); 6, Araucarites sp. (Middle Jurassic, Australia); 8, A. cutchensis (Middle Jurassic, India); 9, A. Phillipsi (Middle Jurassic, England). (Slightly reduced; 1, 3, 4, 6, 7, 9, after Seward; 2, after Berry; 5, after Thiselton- Dyer; 8 after Feistmantel.) ee cuneate, nearly as long as broad (fig. 740, 9), and in shape similar to those of the cone already described as Araucarites ooliticus -(Carr.). Examples of French Jurassic cone-scales are afforded by Avraucarites Moreauana Sap.%, from Corallian beds near St Mihiel 1 Carruthers (692) p. 6, Pl. 1. figs. 7—9; Seward (00) B. p. 285, Pl. x. fig. 4. 2 Saporta (84) p. 425, Pls. cLXXXIV., CLXXXV. 264 ARAUCARINEAE [CH. and other localities, similar to those of A. Brodie (fig. 740, 4, 7) but reaching a length of 4-5 cm., kite-shaped and provided with a terminal spine; also Araucarites microphylla Sap.1 represented by foliage-shoots and cone-scales; the shoots bear linear-lanceolate leaves similar to those of Araucaria Bidwilli but smaller, and the ~ scales are of the Hutacta type. Araucarites Falsani Sap.” is founded on twigs similar to those of Araucaria excelsa and scales character- ised by stout terminal spines. Both Araucarites Falsani arid A. nucrophylla are from Kimeridgian strata in Ain. German cone-scales hardly distinguishable from some of the British and French examples are described by Sir William Thisel- ton-Dyer® from Solenhofen as Araucarites Haberleinit (fig. 740, 5). Salfeld figures some detached scales from the Malm of South- West Germany as Araucaria? which are undoubtedly Araucarian cone-scales agreeing closely with A. Milleri from Scotland (fig. 740, 1). Araucarites Rogersi Seward. The scales of this species (fig. 740, 3)4 from the Uitenhage (Wealden) series of Cape Colony reach a length of 3 cm. and the straight distal margin bears a median spine; there is no indication of a ligule. A specimen in the British Museum collected by Atherstone and referred to by Tate> shows several scales still in their natural position. This type bears a striking resemblance to some of the Indian specimens described by Feistmantel and is similar to the North American Neocomian species A. wyomingensis Font. Araucarites macropterus Feistmantel and A. cutchensis Feist. Several examples of typical Araucarian cone-scales are figured by Feistmantel from Upper Gondwana rocks in India. The scales described as A. macropterus® are distinguished by their large size; specimens from the Rajmahal series reach a breadth of 5em. The scales of A. cutchensis? recorded from the Cutch flora (fig. 740, 8) and elsewhere are smaller, but in some cases it is 1 Saporta (84) p. 431, Pls. CcLXXXVI., CLXXXVII. 2 Ibid. p. 439, Pls. cLXXXVI., CLXXXVII. 3 Thiselton-Dyer (72). 4 Seward (03) B. p. 37, Pl. vr. figs. 4—7. ° Tate (67) p. 147. 6 Feistmantel (77°) p. 186, Pl, virt. figs. 9—12 * Ibid. (76*) p. 62, Pls. vil.—tx., xit.; (82) Pl. m1. XLV] ARAUCARITES 265 impossible to draw any sharp line between the two species; they agree very closely with both British and French Jurassic types. Araucarites cutchensis Feist. is recorded by Halle! from the Upper Jurassic flora of Graham Land. The scales exhibit a con- siderable range in size and shape and more than one type may be represented. They are always more or less cuneate and have a narrow truncate base; some of them show broad lateral wing-like extensions; the distal end is nearly truncate and bears a narlow linear appendage. As Halle says, the scales closely resemble those of A. Brodiet Carr. Cone-scales from Jurassic rocks in Victoria?, Australia, de- scribed as Araucarites sp., A and B, demonstrate the occurrence of cones with scales almost identical with A. Phillipsi and other European forms. There is a comparative scarcity of Araucarian “cone-scales in Jurassic and Cretaceous strata in North America but some examples are recorded. Araucarites wyomingensis Font.? from the Lower Cretaceous of the Black Hills is represented by broadly cuneate scales 1-2 cm. long and with a maximum breadth of 9 mm. and a broad beak at the apex bearing seeds 4—5 mm. long. Larger cone-scales are described by Berry* from Middle Cretaceous rocks in North Carolina as Araucarites Jeffreyi (fig. 740, 2). These scales are associated with the foliage-shoots referred to Araucarttes bladensis and the two may belong to one plant. The same author also figures a specimen from the Upper Potomac series as Arau- carites patapscoensis® from Virginia. Foliage-shoots. There are numerous examples of foliage-shoots among Mesozoic, and to a less extent Palaeozoic, strata which bear a striking resem- blance to branches of recent species of Araucaria, especially species of the Eutacta section, but in many cases confirmatory evidence such as would be afforded by reproductive shoots is lacking. The practice adopted by some authors of referring 1 Halle (132) p. 72, Pl. vim. figs. 3—10. 2 Seward (04?) B. p. 181, figs. 42, 43. 3 Fontaine in Ward (99) B. p. 669, Pl. cixur. figs. 1—9. 4 Berry (08) p. 258, Pl. xvi.; (14) p. 20. 5 Ibid. (11) p. 399, Pl. uxxvit. fig. 5. 266 ; - ARAUCARINEAE [on. impressions of vegetative branches to the genus Araucarites solely on the ground of similarity in habit and leaf-form to the recent genus is not in accordance with sound principles, though in some instances the implied relationship may be a reality. Pending more satisfactory evidence many of the sterile Araucaria-like shoots are referred to Pagiophyllum, while branches of similar habit bearing oval cones are included in the genus Hlatides. An example of a Palaeozoic fossil which has been assigned to Araucarites on slender grounds is afforded by Araucarites Oldhami Zeiller. Araucarites Oldhami Zeiller. The specimen from the Lower Gondwana rocks in India to which this name is applied! consists of an axis bearing spirally disposed lanceolate-acuminate leaves reaching 4:5 cm. in length, slightly contracted at the base and longitudinally striated: portions of the axis show rhomboidal and feebly convex areas separated by narrow scars where the laminae have been broken off. As Zeiller says, the resemblance of the shoot to a branch of Arau- carta imbricata is very close, but considering the age of the beds and the absence of any Araucarian cone-scales from rocks at this horizon in India it is questionable whether it is wise to adopt the name Araucarites. It is not unlikely that a small specimen figured by Feistmantel® from the Karharbari coal-field as possibly a Fern rhizome is a portion of a leafless axis of Zeiller’s species. Araucarites ovatus Hollick. This species from Cretaceous strata in New Jersey? is founded on fragments of sterile branches bearing elliptical-ovate leaves resembling the foliage of Araucaria imbricata and the fossil species Araucarites Nathorstt Dus. A very similar type is represented by Araucarites bladenensis, described by Berry4 as Araucaria bladen- ensis, from the Upper Cretaceous rocks of Carolina and Alabama: the leaves are decurrent, ovate-lanceolate, about 1-6 by 0-8 cm. with a cuspidate apex and rounded base; there are 14—16 parallel 1 Zeiller (02) B. p. 36, Pl. vi. fig. 6. 2 Feistmantel (79?) Pl. xu. fig. 6. 3 Hollick (97) p. 128, Pl. xm. figs. 3 a, 4. 4 Berry (08) p. 255, Pls. x11.—xrv.; (14) pp. 19, 105, Pl. m1. figs. 6, 7; Pl. xrx. figs. 1, 2. XLV] ARAUCARITES 267 veins and imperfectly preserved stomata occur in rows on the lower surface. Berry compares the species with Araucaria Tou- casi figured by Saporta! from Turonian rocks in the South of France. Another species founded on a sterile shoot is Araucarites Hatchert described by Wieland? from Upper Cretaceous rocks in Wyoming. These and other examples that might be quoted, though re- ferred to the Araucarineae on evidence that cannot be considered conclusive, are probably correctly determined; the comparison with Araucaria Bidwilli and A. imbricata suggested by the striking resemblance of the leaves is supported by the occurrence of Arau- carian cone-scales in some of the localities. Araucarites Sternbergit Goeppert. This species was founded on sterile branches, from the rich Eocene flora of Haring in the Tyrol, practically identical in habit with foliage-shoots of Araucaria excelsa and other recent species. From the same locality Goeppert* figured an imperfectly pre- served cone approximately 6 cm. long and 3 cm. in diameter characterised by imbricate, spirally disposed scales with reflexed apices which he compares to a male cone of Araucaria imbricata incorrectly spoken of as A. excelsa: Goeppert suggests the possible specific identity of A. Sternbergui and A. Goepperti Sternb.: the latter species was founded by Sternberg® on a Tertiary cone from Haring in the Tyrol. Ettingshausen® subsequently figured several good examples of vegetative shoots of this type from Haring and described a subglobose cone, figured by Gardner’, which he refers to the same species: this author also records A, Sternbergii from Bilin in Bohemia’, but under the generic name Sequoia: in his account of the occurrence of the species in Carinthia® he adopts the designation Araucarites. Ettings- hausen figures a single cone-scale from Eocene beds in Styria as Araucarites schoeneggensis and compares it to the scales of 1 Saporta (79) A. p. 198, Pl. xxvu., 2. 2 Wieland (08%) p. 6, fig. 2. 3 Goeppert (50) p. 236, PI. xxiv. fig. 1. 4 Ibid. Pl. xxiv. fig. 2. 5 Sternberg (38) A. PI. xxx1x. fig. 4. 6 Ettingshausen (55) p. 36, Pls. vu., vim. ? Gardner (86) p. 96. 8 Ettingshausen (672) p. 116, Pl. xu. figs. 3—8. ® Ibid. (85). 10 Tbid. (90) Pl. t. fig. 93. 268 ARAUCARINEAE [CH. A. Sternbergii. The latter species is recorded also by Massalongo? from Eocene rocks in Italy and on imperfect evidence by Heer? from Switzerland. Gardner? describes several good specimens of vegetative shoots from the Eocene flora of Bournemouth which he names Araucarites Goeppert: Sternb. though the specific name Sternbergit would be more appropriate as that designation was first applied to similar branches from Haring and A. Goeppertt was founded on a de- . tached cone. Two small pieces of larger specimens in the British Mu- seum from Bournemouth are repre- sented in fig. 741 in illustration of the very close resemblance of the leaves to those of recent species. Gardner draws attention to the similarity of some of the fossil examples to deci- duous shoots of Araucaria Cunning- hamw: with reference to the absence of cones or cone-scales he quotes the we. 741. Araucarites Sternbergii. fact, communicated to him by an (British Museum, V, 523; nat. observer in Madeira, that the foliage °*) of A. Cunninghamit requires two or three days to sink while mature seeds do not begin to sink before the fifth or sixth day, so that in moving water shoots and seeds would necessarily be deposited separately. Some of the fragments of branches described by Gardner as Athrotaxis (%) subulata* may well belong to Araucarites. It must be admitted that in the case of the English specimens, as in many others, the use of the generic name Araucarites is based on the evidence of vegetative branches only, but Gardner correctly states that in the shoots of similar habit referred to Cryptomeria the leaves are straighter, and moreover the presence on some of the shoots of the latter of persistent cones like those of Cryptomeria japonica constitutes a clear distinction. Having regard to the very striking resemblance of the widely spread Tertiary specimens 1 Massalongo (59) Pls. v.—v1. 2 Heer (55) A. Pl. xx. fig. 5. 3 Gardner (86) p. 55, Pl. x1. fig. 1; Pl. xm. 4 Ibid. Pl. xt. XLv] ARAUCARITES - 269 included in Araucarites Sternbergii or A. Goeppertt to those of A. Cunninghamii and other species the probability of generic identity is such as to justify the retention of the designation Araucarites. Araucarites Haastii (Ettingshausen). Ettingshausen! described this species as Araucaria Haastit from beds at Shag Point, New Zealand, believed to be of Eocene age; it is represented by sterile branches bearing crowded ovate- lanceolate, acuminate, leaves apparently of leathery texture reaching a length of 5cm. and 2cm. or less in breadth. As Ettingshausen says, they agree very closely with the leaves of Araucaria imbricata but like those of A. Nathorsti Dus. they have a less spinous apex than in the recent species. Some petrified wood from Malvern Hills in New Zealand is referred by Ettings- hausen to the same species but without any evidence of con- nexion between the wood and the foliage-shoots. The same author describes a branch similar in habit to Araucaria excelsa, from Shag Point, as Araucaria Danai, but the specimen is too imperfect to warrant the use of the designation Araucarites. Araucarites Nathorsti (Dusén). This species, described as Araucaria Nathorsti, is recorded by Dusén? from Punta Arenas on the Magellan Straits: the age of the beds is believed to be Oligocene though the precise horizon has not been determined. The material consists of fragments of foliage-shoots bearing short and relatively broad leaves of leathery texture, varying from linear to ovate; they agree closely with the leaves of Araucaria imbricata, differing chiefly, as Dusén states, in their blunter apices. Araucarites imponens (Dusén). Nathorst‘ first suggested a reference to Araucaria of the single leaf on which this species was founded®: it was collected in a marine volcanic tuff in Seymour Island and is probably of Lower Tertiary age. The leaf is linear, 6 cm. long, and tapers gradually towards an incomplete apex; it agrees in form and size with 1 Ettingshausen (87) p. 154, Pl. 1. 2 Ibid. p. 155, PI. 1. fig. 18. 3 Dusén (99) p. 105, Pl. xn. 4 Nathorst (04?) B. 5 Dusén (08) p. 11, Pl. 1. figs. 16, 17. 270 ARAUCARINEAE (cu: leaves of A. Bidwilli and A. brasiliensis, but the single impression is hardly sufficient to demonstrate the existence of Araucarites in this southern flora (lat. 64° 16’ S.). On the other hand the occurrence of wood of the Araucarian type? in Seymour Island in beds that are either Lower Tertiary or Upper Cretaceous supports the conclusion of Nathorst and Dusén. ELATIDES. Heer. Heer? proposed this name? for some Jurassic Coniferous remains from Siberia characterised by spirally disposed falcate leaves (figs. 742, 743) and cones similar externally to those of Picea, Abies and other Abietineae. The genus is based primarily on the form of the cones and cone-scales. In the new genus were included three species, H. ovalis, represented by oval cones 2-7 cm. long and 6--7 mm. broad, #. Brandtiana characterised by cylindrical cones, and £. falcata founded on vegetative branches very similar to those of £. Williamsonis. Nathorst* has included these three species in Elatides curvifolia (Dunk.) a Wealden species abundantly represented in the plant-beds of Spitzbergen (fig. 743). In the absence of cones it is impossible to draw any satisfactory distinction between foliage-shoots belonging to Elatides and those referred by authors to Sequoia, Pagiophyllum, and other genera. It is therefore only in cases where cones are present that the designation Elatides is ‘admissible. The vegetative characters of Elatides are those of Araucaria Cunninghamit, A. excelsa, and allied species while the cones consist of flat imbricate scales with narrower and more or less pointed or spinous distal ends. There is some reason to believe that the cone-scales were _ monospermic but the evidence is not conclusive and rests on a single species. The data are insufficient to fix definitely the posi- tion of the genus, though it is in all probability a member of the Araucarineae. Slatides is characteristic of Rhaetic, Jurassic, and Wealden floras. Elatides Sternbergit (Nilsson). A Rhaetic species originally described by Nilsson from Rhaetic rocks in the South of Sweden as Abies Sternbergii, subsequently 1 Gothan (08). * Heer (77) ii. pp. 77—79, PL. x1v. 3 édary, Fir. + Nathorst (97) pp. 35, 58, Pls. 1, m., Iv., VI. XLV] ELATIDES 271 included by Nathorst! in Palissya Brawnii but afterwards recog- nised by him as a distinct species? and recently transferred to Elatides®. An examination of specimens in the Stockholm Museum leads me to agree with the substitution of the designation Elatides. Nathorst has also pointed out that some of the cones from the Rhaetic of Franconia referred by Schenk! to Palissya Braunii are of the Elatides type and distinct from cones of Palis- sya which are characterised by their more open habit and by other more important morphological features. Hlatides Sternbergii, though similar in the habit of the vegetative shoots to £. William- sonis from Jurassic strata, differs in the narrower and straighter leaves which may reach a length of 2—3 cm. and are either straight or slightly curved in contrast to the stouter and strongly falcate leaves of EZ. Willsamsonis and E. curvifolia. A cone figured by Nathorst® is practically identical in external form with one of E. Willamsonis illustrated in volume I. of the Jurassic Flora of the’ Yorkshire Coast®. We have no knowledge of the structure of the reproductive shoots and no evidence other than the habit of the foliage-shoots with regard to systematic position: it is, however, probable that this Rhaetic species is closely allied to the later Jurassic and Wealden types. Elatides Williamsonis (Brongniart). This Jurassic species described by Brongniart? as Lycopodites Wilkamsonis, was figured by Philhips® as L. uneifolius and by Lindley and Hutton® under Brongniart’s name. The specimens figured by the English authors are in the York and Manchester Museums respectively. Schimper transferred the species to Pachy- phyllum and it. has usually been assigned to that genus or to Pagiophyllum”, the name substituted by Heer for Pomel’s Pachy- phyllum, a designation now reserved for sterile shoots and there- fore inapplicable to the present species which possesses cones of the Elatides type. The vegetative shoots are monopodially 1 Nathorst (78?) B. p. 28, Pl. rv. figs. 1—3. 2 Ibid. (86) p. 107, Pl. xxii. figs. 8—12; Pls. xxiv., xxv. 3 Ibid. (97)-p 34; (08). 4 Schenk (67) A. Pl. xxi. fig. 7. See also Solms-Laubach (91) A. p. 73. 5 Nathorst (86), Pl. xxv. fig. 8. 8 Seward (00) B. Pl. x. fig. 3. ? Brongniart (28) A. p. 83. 8 Phillips (29) A. Pl. vu. fig. 3 ® Lindley and Hutton (33) A. Pl. xcr. 10 Seward (00) B. p. 291. 272 ARAUCARINEAE [CH. branched, the smaller branches being given off at an acute angle; the leaves are crowded, fleshy, tetragonal and falcate (fig. 742), agreeing closely with the foliage of the Hutacta species of Araucaria and with Cryptomeria. The megastrobili are cylindrical, approxi- mately 6 cm. long and 2cm. in diameter, bearing imbricate, flat, scales with narrow pointed distal ends resembling the free portion Ul Fic. 742. Elatides Williamsonis. (British Museum; nat. size.) of the foliage-leaves. No specimens have been described showing seeds attached to the scales. The more slender microstrobili, 2em. long, bear sporophylls at right angles to the axis with tri- angular upturned distal ends characterised by a median keel. In the vegetative shoots this species closely resembles the Liassic Pagiophyllum peregrinum? (fig. 744) (Lind. and Hutt.), but in the 1 See page 276. XLV] ELATIDES 273 absence of cones the latter species is retained in Pagiophyllum. From the Wealden species E. curvifolia, E. Williamsonis differs in its stouter and more crowded leaves though the differences are slight both in the cones and vegetative shoots. In all probability this species is represented in several Jurassic floras, but unless cones are present specimens should be referred to Pagiophyllum. Elatides curvifolia (Dunker). Dunker? first described this Wealden type (fig. 743) from North Germany as Lycopodites and it was referred by Ettingshausen? to Araucarites, the generic name, though probably correctly express- Fic. 743. Elatides curvifolia. (After Nathorst; nat. size.) ing the position of the fossil Conifer, being used without adequate reasons. The identity of the cones discovered by Nathorst? in the Wealden or Upper Jurassic beds of Spitzbergen with those on which Heer founded the genus Elatides led to the adoption of that generic term. Nathorst’s discovery of several fertile branches justifies his reference of Heer’s specimens from Spitzbergen described as Sequoia Reichenbachii* to Elatides curvifolia, as also the employment of Dunker’s specific term for Llatides ovalis and E. Brandtiana Heer. The cones of H. curvifolia are cylindrical or oval and it is suggested by Nathorst that these forms might be regarded as varieties, the oval form being spoken of as var. ovalis 1 Dunker (46) A. p. 20, Pl. vm. fig. 9. ? Ettingshausen (52) Pl. nm. 3 Nathorst (97) pp. 35, 58, Pls. 1., 0., Iv., VI. 4 Heer (75) ii, Pl. xxxvi. figs. 1—8; Pl. xxxvit. figs. 1, 2. 274 ARAUCARINEAE [CH. after Heer’s specific name and the cylindrical cones being distin- guished as var. Brandtiana, but it is doubtful whether the retention of these varietal names is advisable. The cone-scales have pointed apices and agree closely with those of #. Willkamsonis. On a specimen of a cone of this species from Kimeridge strata in Scotland! one scale afforded evidence of the occurrence of a single seed as in Araucaria. Nathorst regards some smaller cones on a branch from Spitzbergen as microstrobili and suggests that longi- tudinal striae on the sporophylls may represent long microspor- angia like those of Araucarza; but the preservation is too imperfect to demonstrate the nature of the specimen. The vegetative branches bear falcate leaves rather more slender and as a rule less crowded than in E. Williamsonis: on older branches from which the free part of the lamina has fallen there are leaf-bases or in some cases an oval leaf-scar. This type is characteristic of Wealden strata in Spitzbergen, North Germany, and other European localities; it is no doubt represented by some of the impressions of branches assigned to Sphenolepidium Sternbergianum?; it is also recorded from Kimeridgian strata in the North of Scotland. PAGIOPHYLLUM. Heer. Heer? instituted this genus in place of Pachyphyllum, previously adopted by Pomel‘ for a section of his genus Moreawia, on the ground that the latter name had been applied to a member of the Orchidaceae. Some of the species referred to Pagiophyllum have also been included in Araucarites and Brachyphyllum. Tuzson® instituted a new genus Pagiophyllites for petrified Mesozoic wood having Araucarian features, the type-species being P. keuperianus (Goepp.), but no evidence is furnished in support of a connexion of this wood with foliage-shoots of Pagiophyllum. Schimper and Saporta include in their diagnosis of the genus both vegetative and reproductive shoots and consider Pagiophyllum to be allied to Agathis, Cunninghamia, and Araucaria. Certain species have in recent years been transferred to Elatides because of the occur- rence of cones conforming to Heer’s genus. It has been suggested® that Pagiophyllum may most conveni- 1 Seward (112) p. 684, fig. 10. 2 [bid. (95) A. p. 205; (112) p. 685. 3 Heer (81) p. 11. 4 Pomel (49) p. 352. 5 Tuzson (09) p. 30. ® Seward (12) p. 41. XLV] ; PAGIOPHYLLUM 275 ently be reserved for vegetative branches of Conifers (fig. 744) possessing foliage like that of Araucaria excelsa and allied species, which in the absence of cones cannot safely be referred to Elatides or other genera based, in part at least, on strobilar characters. Pagiophyllum is essentially an artificial genus: as Solms-Laubach says, ‘it is only in accordance with old custom to distinguish the Ullmanniae of the Zechstein from Pagiophyllum1,’ and it is equally difficult to draw any clearly defined line between this genus and some forms included by authors in Brachyphyllum.. A Triassic species from Raibl originally referred to Voltzia heterophylla® afterwards named V. Foettleri by Stur®, Pagiophyllum Sandbergi by Schenk‘ and figured by Schiitze® as P. Foettlert, has the habit of a Brachyphyllum. This is one of many examples of sterile shoots illustrating the arbitrary use of generic names for coni- ferous remains which afford no definite evidence of their systematic position, The Araucarian habit is in itself of little value as ‘evidence of affinity, but the abundance of petrified wood with ' Araucarian features (Dadozylon) in strata yielding Pagiophyllum shoots suggests an Araucarian alliance, and the fact that some Pagiophyllum shoots bear Elatides cones affording indications of Araucarian characters points in the same direction. It cannot be assumed that all Pagiophyllum shoots bore similar cones, and it is mainly on this account that the employment of Pagiophyllum as a provisional designation is recommended. Pagiophyllum is widely distributed in Jurassic strata and extends into Cretaceous and Tertiary rocks: it occurs also in pre-Jurassic floras and has recently been described by Zeiller® from the Permian of France. It should be recognised that this extended use of the name is not in accordance with general practice, but it is adopted on the ground that, as in recent Conifers so in the case of extinct types, similarity in the habit of vegetative branches does not necessarily imply close relationship as regards the more important characters of the reproductive shoots. 1 Solms-Laubach (91) A. p. 77. 2 Bronn (58) p. 135, Pl. vi. 3 Stur (68) p. 104. . 4 Schimper and Schenk (90) A. pp. 276, 290. 5 Schiitze (01) Pl. v1. fig. 1. 6 Zeiller (06) B. p. 219, Pl. x1. 18—2 276 ARAUCARINEAE , [CH. Pagiophyllum peregrinum (Lindley and Hutton). This species, first named by Lindley Araucaria peregrina’, was founded on material from the Lias of Lyme Regis in Dorsetshire?. It is possible that the generic name chosen by Lindley correctly expresses the position of the species, but decisive evidence is lacking. Vegetative shoots bear crowded imbricate, spirally dis- posed, leaves tetragonal in section, broadly triangular, sometimes fal- cate and more or less appressed to the stem in the lower portion of the lamina (fig. 744). There is a distinct dorsal keel and occasion- ally rows of papillae are visible on the lamina; the apex is obtuse or acute. The leaves vary consi- derably in size and shape. Zeiller® describes the cuticle of the dorsal and ventral surfaces of some leaves on Permian specimens from Blanzy: the stomata occur in longitudinal rows on the lower face only, the guard-cells being usually at right-angles to the long axis of the leaf. In habit this species agrees closely with Elatides Wailliam- sonis, a Middle Jurassic type; it occurs in Jurassic rocks of Eng- land, France, Germany, Italy, and elsewhere, the oldest recorded ex- amples being those described by Fic. 744. Pagiophyllum peregrinum. (British Museum; nat. size.) 1 Lindley and Hutton (33) A. Pl. yxxxvit. * Seward (04) B. p. 48, Pl. v.; Saporta (84) p. 383, Pls. 173—176. 3 Zeiller (06) B. p. 219. XLV] WALCHIA 277 Zeiller from the Permian of France which he refers to Pagio- phyllum in preference to Ullmannia, pointing out that the leaves are relatively longer and less appressed to the axis than in the shoots known as U. Bronni Goepp. (fig. 750, D, E). Triassic specimens from North Italy in the Bologna Museum named Pagiophyllum Rotzoanum appear to be indistinguishable from the English species, It is unnecessary to describe other examples of the genus as the Pagiophyllum type is illustrated by many Mesozoic and Tertiary species referred by authors to Sequoia, Geinitzia, Elatides, Sphenolepidium, and other genera. The important point is that in place of generic names connoting definite forms of cone, the designation Pagiophyllum should be adopted for all foliage-shoots of a certain habit which afford no satisfactory evidence as to the nature of the reproductive shoots. PataEozoic CONIFERS EXHIBITING CERTAIN FEATURES SsUG- GESTIVE OF ARAUCARIAN AFFINITY BUT WHICH CANNOT BE DEFINITELY ASSIGNED TO THAT OR TO ANY OTHER FAMILY oF CONIFERALES ON THE EVIDENCE AT PRESENT AVAILABLE, WALCHIA. Sternberg. The name Walchia! is applied to foliage-shoots, occasionally bearing terminal cones, from Permian and to a less extent Upper Carboniferous rocks, which present a striking agreement in habit with branches of Araucaria excelsa and other recent species of the section Hutacta of Araucaria. Information with regard to repro- ductive shoots is very incomplete and we have little more than circumstantial evidence as to the anatomical features of the stem. In many cases the ultimate branches bear terminal cones similar to the megastrobili of Elatides?, but it is only in a few specimens that seeds are preserved on the cone-scales: in some species, ¢.g., W. frondosa Ren. and W. fertilis Ren. the fertile shoots appear to be of a distinct type though the evidence is not wholly satisfactory. It is probable, as several authors have suggested, that the species included in Walchia, were our information fuller, would be referred to more than one generic type. The resemblance of branches of Walchia to the foliage-shoots of Lepidodendron, especially in the 1 Sternberg (26) A. p. xxii. 2 See page 272. 278 CONIFERALES INCERTAE SEDIS [CH. Fig. 745. Walchia piniformis. (British Museum; } nat. size.) XLv] WALCHIA 279 case of specimens too small to show the characteristic branching- habit, has led to confusion between the two genera. It is often very difficult to draw a definite line between Walchia and Ullmannia, and in the absence of sporophylls the genus Gomphostrobus may be easily confused with species of Walchia. Foliage-shoots characterised by a pinnate arrangement of the ultimate branches (fig. 745) attached at right-angles or obliquely to an axis of higher order. Leaves spirally disposed, crowded and imbricate, short and ovate or linear and spreading, usually tetra- gonal and more or less falcate and decurrent. The dimorphism of branches and differences due to age or position on the tree render a satisfactory delimitation of species almost impossible though a few fairly well defined types can be recognised with reasonable certainty. . As Bergeron! says, in the absence of strobili the separation of species represented only by sterile shoots is hardly possible. Further reference is made to the features exhibited by reproductive shoots in the appended account of a few selected types. Information with regard to the anatomical characters of Walchia is very scanty and is based on evidence afforded by the association of foliage-shoots and petrified wood-or on inferences drawn from unconvincing considerations. Among specimens which may belong to this genus one of the more interesting is that on which Mougeot? founded the species Araucarites valdajolensis. The type-specimen, from the Permian of Val d’Ajol in the Vosges, has been refigured and critically discussed by Fliche? though no complete investigation of its structure has been made. The cylindrical piece of stem, 9 cm. in diameter, has a large pith and a -broad zone of secondary wood composed of tracheids, with two rows of alcernate bordered polygonal pits, and narrow medullary rays. 4; seems clear from Mougeot’s brief account and from the description of other specimens by Fliche that the anatomical features are Araucarian though we have no information as to the structure of the inner edge of the xylem, a region of special importance as regards comparison with other types possessing a similar Araucarian pitting on the tracheids. The surface of Mougeot’s specimen is characterised by numerous spirally dis- posed, elliptical projections 5—7 mm. long and 3—4 mm. wide 2 Bergeron (84). 2 Mougeot (52) A. p. 27, PI. rv. 3 Fliche (03). 280 CONIFERALES INCERTAE SEDIS [CH. which Fliche regards as leaf-bases and compares with those on Araucarian stems. While admitting the possibility that the wood belongs to Gomphostrobus or some Cordaitean species Fliche con- siders Walchia the most likely genus. The comparative closeness of the leaf-bases would seem to be a difficulty: in stems as large as the type-specimen of W. valdajolensis one would expect to find the leaf-bases more widely separated and tangentially stretched. It is by no means unlikely that the supposed surface-features may belong to a deeper zone of the cortex of a partially decorticated stem; but in any case they do not suggest a stem of Cordaites or Mesoxylon. It is impossible to assign the species with confidence to Walchia though Fliche may be correct in his opinion as to the likelihood of that being its true position. The generic name Araucarites implies a degree of affinity which has not been estab- lished and the designation Dadoxylon would be more in keeping with the facts. Walchia is especially characteristic of Permian floras though it has been shown to occur in the Stephanian of several countries. In Britain Walchia is recorded from a very few Permian! and Upper Coal Measures? localities. We cannot speak with confidence as to the position of the genus: the striking resemblance in the system of branching and in the foliage-shoots to certain species of Araucaria at once suggests a possible affinity to the Araucarineae, and this slender basis of comparison receives support from the occurrence in a few instances of single seeds on the upper face of sporophylls and from the Araucarian type of pitting in wood associated with Walchia branches. It may fairly be said that although proof is lacking there is a strong presumption in favour of regarding this Permo-Carboniferous genus as more nearly allied to the Araucarineae than to any other family of Gymnosperms. Walchia piniformis (Schlotheim). This, the commonest species, was originally described by Schlotheim as Lycopodiolithes piniformis?: it occurs in both Permian and Stephanian strata’. The pinnately branched shoots 1 Vernon (12) p. 607. 2 Kidston (02) B. ® Schlotheim (20) A. p. 415, Pl. xxut. fig. la; Pl. xxv. fig. 1, 4 E.g. Stefani (01) p. 111; Weiss, C. E. (72) p. 179; Heer (76) A. p.57; Goeppert (65) p. 236; Grand’Eury (77) A. p. 514. XLV] ' -WALCHTA 281 (fig. 745) are characterised by the more or less oblique insertion of the slender branchlets and by the comparatively long, narrow, faleate, decurrent leaves. The ovoid or cylindrical megastrobili terminal on the ultimate shoots bear imbricate ovate-lanceolate sporophylls, but their preservation is not suck as to throw any light on the structure of the seed-bearing organs. Zeiller! mentions a cone from Lodéve (Permian) 10cm. long and 1—1-2 cm. in diameter, but the average length is less than this. A branch figured by Potonié? from the Permian of Thuringia shows elliptical leaf-cushions very like those on the larger stem described by Mougeot as Araucarites valdajolensis. Potonié has drawn atten- tion to the difficulty of distinguishing small specimens of this species from W. filiciformis and W. linearifolia, and some forms described as W. imbricata and W. hypnoides are by no means clearly distinguished from W. piniformis. In W. filiciformis (Schloth.) the leaves are characterised by the downward curve of the lamina near the base though this in itself is hardly a decisive criterion. In W. linearifolia Goepp. the leaves are rather more delicate and less falcate, while in W. imbricata they are usually shorter, relatively broader, and more strongly imbricate and incurved. W. hypnoides (Brongn.) is a smaller form though, as Kidston® suggests, this may not be a specific character. Renault’s species W. fertilist represents a similar form but with smaller leaves, and each branchlet ends in a long and narrow strobilus which affords no indication of the nature of the sporophylls. Walchia filiciformis (Schlotheim). This species, recorded from Permian and Stephanian beds, and differing but slightly from W. piniformis is important as supplying more satisfactory evidence as to the nature of the megastrobili. Zeiller> has described a fertile specimen from the Permian of Brive in which the ovate-lanceolate cone-scales (sporophylls) bear on their upper concave face single ovoid seeds, 7—8 mm. long. The strobili appear to be lax in the arrangement of the monospermic sporophylls. 1 Zeiller (06) B. p. 204, Pl. x. figs. 3,5; Bergeron (84). 2 Potonié (93) A. p. 218. 3 Kidston (86) A. p. 15. 4 Renault (96) A. p. 359; (93) A. Pl. pxxx. fig. 2. 5 Zeiller (92) A. p. 99, Pl. xv. fig. 3. 282 : CONIFERALES INCERTAE SEDIS [CH. Walchia imbricata Schimper. In habit this type! closely resembles the foliage- as of Araucaria Rulei on a smaller scale, the crowded leaves being strongly incurved and imbricate: it is readily distinguishable from Ullmannia BRonni Goepp. (fig. 750, D, E). A good example is figured by Zeiller? in which the branches are unusually large, 8 mm.—1-2 cm. in diameter: the species occurs in Permian and Stephanian strata and is recorded by oo from the Upper Coal Measures of central England. Walchia Schneidert Zeiller. This species? from Charmoy is characterised by the long filiform leaves, -6 mm. broad and 1-2 cm. long, usually straight and decurrent. The ultimate branches, some of which bear compara- tively long and slender cones, are oblique and alternate as in W. piniformis. W. foliosa Eich.* from the Permian of Russia is a similar form but with less delicate leaves. Walchia frondosa Renault. A species from the Permian of Autun having slender leaves rather shorter and more falcate than those of W. Schnevderi: some of the branches bear a terminal globular bud superficially resembling the ovuliferous shoot of Taxus. It is, however, not improbable that the buds are purely vegetative like those figured by Bergeron on a specimen of W. piniformis. SCHIZODENDRON. Hichwald. (TYLODENDRON Weiss.) Prof. C. E. Weiss® instituted the generic name Tylodendron for casts from Upper Carboniferous and Permian strata in Germany which he described as branches of a Conifer with spirally arranged rhomboidal raised areas or pulvini each of which has a median slit in its apical portion (fig. 746). The elongate, narrow, raised areas (cf. the medullary cast of a Voltzia, which shows precisely similar areas, represented in fig. 748) were regarded as casts of leaf- cushions and the slit was interpreted as a resin-canal. 1 Schimper (72) A. p. 239. 2 Zeiller (06) B. p. 211, Pl. xurx. figs. 1, 2. 3 Ibid. p. 206, Pl. xivmt. figs. 4, 5. 4 Eichwald (60) B. Pl. xrx. fig. 1. 5 Weiss, C. E. (72) p. 182, Pls. xrx., xx. XLv] SCHIZODENDRON 283 Schizodendron speciosum (Weiss). One of the casts figured by Weiss and assigned by him to this species has a length of 70 cm. and at intervals of about 30 cm. shows periodic swellings where it assumes a barrel-shaped form. Pieces of wood attached to some of the casts were investigated by Dippel who found that they agreed anatomically with Araucarian stems. Weiss considered his specimens to be generically identical with casts figured by Eichwald! from Russia as species of Schizodendron and Angiodendron. Potonié, while uncertain as to the close agreement with some of Eich- wald’s fossils, regarded Tylodendron as identical with Eichwald’s Schizo- dendron, and Zeiller?, in view of this agreement, adopted the older name Schizodendron. The latter author? formerly believed Tylodendron, as described by Weiss, to be- distin- guished from Schizodendron by the apical occurrence of the slit on the so-called leaf-cushion in contrast to the basal slit in Schizodendron, but Potonié! proved that in both cases the median groove extends up the lower portion of each projecting Fic. 746. Schizodendron specio- area from its base and represents ‘wm. (After Potonié; ¢ nat. an out-going leaf-trace; he also ©) demonstrated that Schizodendron is a pith-cast, the tapered areas being the inner ends of medullary rays. In some specimens the casts afford some indication of a discoid pith. The relation between the wood and the pith-casts is also very clearly shown in a section of a petrified stem of Permian age from Prince Edward Island described by Miss Holden®. The true nature of —S LESS ‘1 Eichwald (60) B. Pl. xvm. 2 Zeiller (92?) A. p. 102. 3 Ibid. (80) p. 263, Pl. v. 4 Potonié (88). 5 Holden, R. (13) p. 245. 284 CONIFERALES INCERTAE SEDIS [CH. the Tylodendron casts is also shown in specimens from the Lower Permian of Saxony in the Chemnitz Museum! Casts similar to those described as Tylodendron and Schizodendron were recorded by Schleiden in 1846 and referred to a new genus Endolepis: he believed them to be casts of the pith-cavity of some Dicotyledonous stem. Examples of Endolepis have been described by Schenk and more recently by Fliche? who discusses the history of the genus and on the ground of priority adopts Schleiden’s name in prefer- ence to Schizodendron. It is, however, preferable to retain Schizodendron for the larger casts with periodic swellings. The smaller type represented by EHndolepis has in several instances been found in connexion with the foliage of Volézia3 (fig. 748) and it is questionable if a special designation is needed. In the form of the raised areas on the surface of the cast Schizodendron and Endolepis appear to be identical: while suggesting the advisability of retaining the former name I recognise that the Permian and Triassic casts may belong to stems which are closely allied or even generically identical. The structure of the wood of Schizodendron speciosum is of the Araucarian type; the tracheids have 1—3 rows of contiguous and alternate pits on the radial walls and the medullary rays are usually uniseriate. Potonié compares the pith-casts of the Palaeo- zoic stems with those of recent species of Araucaria and Agathis: the pith of the recent species is much smaller but in both fossil and recent medullary casts there are periodic swellings where the presence of scars, sometimes in a whorl or pseudowhorl*, marks the position of branches. The pith-cast of a recent Cycad (fig. 398, p. 29, Vol. 111.) bears a general resemblance to Schizodendron: in Araucaria the medullary rays are narrower and so produce narrower raised areas on a pith-cast. Schizodendron Cowardi (F. E. Weiss). Prof. F. E. Weiss® has recently described an interesting example of Schizodendron which throws some fresh light on structural features. The specimen was found in Cheshire but not i situ and nothing is known as to its geological age; it consists of a 1 Sterzel (00). ® Fliche (10) p. 212, Pls. x1x., xx. 3 Seward (90). See also page 299. : + Zeiller (92?) A. Pl. xv. fig. 5. 5 Weiss, F. E. (13). XLV] _ SCHIZODENDRON 285 petrified barrel-shaped piece of pith with portions of the inner edge of the xylem-cylinder. The surface-features agree with those of S. speciosum, each rhombic area being divided for a third of its length by a median groove. The pith is composed of thin-walled parenchyma with several secretory canals in the outer region; patches of xylem are preserved in the depressions between the lozenge-shaped areas showing the same anatomical characters as those described by Potonié: internal to the secondary xylem are very small groups of tracheids separated by 1—2 rows of parenchyma from the secondary elements, which pursue a sinuous longitudinal course. These tracheal strands are, as Weiss points out, at least superficially comparable with the primary xylem of such a type as Pitys antiqua. The innermost elements of the secondary xylem are usually scalariform and these pass gradually into tracheids with two alternate rows of bordered pits often slightly polygonal. The leaf-traces are formed of two endarch strands which coalesce as they pass downwards and eventually merge laterally with the secondary xylem. Medullary casts with the external features of Schizodendron might well belong to stems which are not identical in anatomical characters, and from casts alone all that can be inferred is the presence in the vascular cylinder of medullary rays with fairly broad inner faces separated by prominent wedges of tracheids, also the spiral disposition of leaves each supplied with a single vascular bundle given off from the lower angle of the xylem- meshes. In the case of Schizodendron Cowardi the presence of small strands of primary xylem suggests comparison with such a genus as Pitys or Mesopitys, while in the other examples there is no indication of any xylem internal to the main cylinder. The characters of the secondary xylem point to an Araucarian or Cordaitean affinity and the pith agrees with that of Araucaria, though in 8. Cowardi the presence of secretory canals is a Cycadean feature. Bain and Dawson}, though they did not correctly inter- pret the surface-characters of Schizodendron, referred to it as representing decorticated branches of the Conifer Walchia. Sterzel recorded the association of Schizodendron with Walchia foliage- shoots in Saxony, and Zeiller, who noticed a similar association 1 Bain and Dawson (85). 286 CONIFERALES INCERTAE SEDIS cH. in French Permian rocks, expressed the opinion that the casts belonged to Walchia stems. In this connexion it is noteworthy that shoots of Voltzia! also possess medullary casts (fig. 748, A, B) with the superficial features of Schizodendron. Though we have no proof of a connexion between casts and leaf-bearing branches, it is probable that some forms of Schizodendron represent the pith- casts of Walchia: if this view is correct it affords another argument in favour of connecting Walchia with the Araucarineae, but how close the connexion is cannot be definitely settled without further evidence as to the reproductive shoots. Schizodendron, though not confined to Permian rocks, is most abundant in beds of that age; it is recorded from several localities in Germany?, from France, Russia’, and Canada while the British specimen, though presumably from English rocks, was not found m situ. HAPALOXYLON. Renault. Renault? instituted this genus for a cylindrical stem 2 cm. in diameter from the Permian of Autun characterised especially by the parenchymatous structure of the secondary xylem. The type-species, Hapaloxylon Rochei, resembles Araucarites valda- jolensis, a Permian species founded by Mougeot, in its spirally disposed leaf-scars each with an elongated groove marking the position of the leaf-trace. The solid parenchymatous pith is surrounded by a narrow zone of 2—3 layers of tracheids with a single row of bordered pits which Renault speaks of as primary xylem: this forms the inner edge of a broad cylinder of homogene- ous parenchyma traversed by uniseriate medullary rays 1—3 cells deep. The secondary-xylem elements are rectangular 7—8 times as long as broad and without pits. Beyond the cambium is a broad zone of secondary phloem consisting of a regular alternation of well preserved sieve-tubes with lateral sieve-plates and paren- chyma. The cortex contains some secretory sacs and is bounded by periderm. The inference drawn by Renault is that the leaves were small like those of Walchia and each had a single vein. There is, 1 Seward (90). 2 Weiss, C. E. (72); (74); Potonié (88); (93) A. etc. 3 Schmalhausen (87) Pl. viz. fig. 34. + Renault (96) A. p. 360; (93) A. Pl. yxxvi. XLV] “GOMPHOSTROBUS 287 however, no definite evidence as to the nature of the foliage: the stem structure represents a type previously unrecorded among Gymnosperms, but comparable with the structure of the stem of Aeschynomene a recent genus of the Leguminosae. GOMPHOSTROBUS!. Marion. This generic name was instituted by Marion? for Permian foliage-shoots from Lodéve bearing vegetative leaves similar to those of Walchia and Araucaria excelsa but distinguished by the bifurcate form (fig. 747) of the relatively long sporophylls borne in a crowded cluster on the apical region of the axis. Marion referred Gomphostrobus to the Coniferales. Geinitz? had pre- viously described detached sporophylls from the Lower Permian of Saxony, of the same type as those on which the genus Gomphostro- bus was founded, as Sigillariostrobus bifidus. The Saxon specimens are represented in the drawings published by Geinitz as distally- forked scales bearing a single seed at the base. Potonié*, who examined the original fossils in the Dresden Museum—and I am able to confirm his view—states that there are no undoubted seeds but only a faintly outlined area near the proximal end of each scale which no doubt marks the position of a seed or sporan gium. 1873. Sigillariostrobus bifidus, Geinitz, Neues Jahrbuch Min. p. 700, Pl. v. figs. 5—7. 1890. Deseanophathan gallicum, Schenk in Schimper and Schenk, p. 266. 1890. Gomphostrobus heterophyllus, Marion, Compt. Rend. cx. p. 892. 1891. Psilotiphyllum bifidum, Potonié, Ber. deutsch. Bot. Ges. Bd. rx. p. 256. ye 1892. Gomphostrobus bifidus, Zeiller, Bassin Houill. Perm. Brive, p. 101, _Pl. xv. fig. 12. Gomphostrobus bifidus (Geinitz). Potonié in his account of the genus and type-species repro- duces Marion’s original drawings showing a Walchia-like axis with short falcate leaves bearing crowded linear and distally forked ~ sporophylls reaching a length of 8 cm. in the apical region. The sporophylls, which are bent to one side, giving the impression of wind-blown foliage, consist of a simple lamina 8-5 to 25 mm. long, 1 See vol. 1. p. 26. 2 Marion (90) A. 3 Geinitz (73). 4 Potonié (93) A. p. 197 Pls. xxvu., XXVIIL, XXXII. 288 CONIFERALES INCERTAE SEDIS [cH. with two divergent distal prongs varying considerably in the angle of divergence, a variation noticed also by Zeiller. On one example (fig. 747, B) Potonié records the occurrence of two scars; a lower scar, a, representing the attachment of the lamina and a second scar, b, which he attributes to a sporan- gium. The species, represented usually by de- : bp C tached sporophylls only, is recorded from Lodéve, Brive, and other French localities}, A wt c also from Permian localities in Germany. a B It was referred by Schenk to Dicrano- Fic. 747. Gomphostrobus phyllum, but in that genus it is the foliage- —#fidus; a, leaf-scar; b. leaves that are forked and there isnoreason ““" © sporangium?; eaves that are forked and t is n son a, tots pede OF Heews, to assume any close relationship between (A, B, after Potonié; . the two imperfectly known types. If the scar C, Zeiller.) at the base of the sporophylls marks the position of a seed a com- parison with the Araucarineae is suggested, and in this connexion it is noteworthy that Sterzel? records the association of Gonvphostrobus with Dadoxylon wood. Potonié, who at first overlooked Marion’s paper, proposed the name Psilotiphyllum to give expression to his opinion that the Permian plant is a Palaeozoic member of the Psilotales, a conclusion based on insufficient evidence. We have no definite information with regard to the nature of the organ borne on the sporophylls. The same author compares the sporo- phylls of Gomphostrobus with the leaves of Sphenophyllum though the verticillate disposition of the leaves of the latter genus is a well- defined difference. It would seem, as Zeiller says, that Gompho- strobus is probably allied to Walchza though its position cannot be precisely determined without further data. A recent examination of some specimens from Lower Gondwana rocks in India described by Feistmantel? as Voltzia revealed the occurrence of some small distally forked leaves very similar to the sporophylls of the Huropean Gomphostrobus. 1 Zeiller (927) A. p. 101, Pl. xv. fig. 12; (06) B. p. 213, PL. x. figs. 6—8. 2 Sterzel (00) p. 6. 3 Feistmantel (79?) Pls. xxI1. et seq. XLV] VOLTZIA 289 VOLTZIA. Brongniart. Brongniart! instituted this genus for foliage-shoots from the Bunter sandstones of the Vosges, the name being chosen in com- memoration of Voltz; he compared the branches with those of Araucaria excelsa but added that the cone-scales bore three ovules. The leaves show considerable variation even on the same axis, a feature shared with Walchia and Ulimannia: the megastrobili are characterised by a lax disposition and the fan-like, lobed or crenulate form of the megasporophylls, which in the best preserved type, V. Liebeana (fig. 748, C—-F), bear three ovate seeds on the upper surface: Many authors compare the Triassic genus with members of the Taxodineae, e.g., Cryptomeria, and the Araucari- neae: wood of the Araucarian type has been referred to Volizia though without proof of connexion with the vegetative shoots. Gothan?, who favours a Taxodineous alliance, points out that wood associated with Volizia has Araucarian pitting on the tracheids, though he adds that the occurrence of typical Araucarian pitting in stems possessing other characters foreign to the recent Araucarineae justifies the conclusion that the presence of alternate polygonal pits on the tracheids is not necessarily proof of Arau- carian affinity. An examination of some carbonised fragments attached to cone-scales of V. Lvebeana in the British Museum from Gera revealed the occurrence of uniseriate pits both separate and in contact with one another. It is probable that Voltzia is related to the Araucarineae though in what degree is uncertain. Leuthardt (03) Pl. rv. figs. 2—5. § Feistmantel (792) Pls. xx11.—xxv. ? Zeiller (02) B. Pl. vit. fig. 9. 8 Fliche (10) p. 198, Pl. xx1. XLV] VOLTZIA 293 clearly between certain forms of Walchia and Voltzia. A very similar type is figured by Schiitze! as Widdringtonites keuperianus Heer* from Stuttgart but with no justification for the use of a generic name implying relationship with Wvddringtonia. The fragments of foliage-shoots on which Heer founded this species are too small and of too common a type to be referred to a genus implying any definite position in the Coniferales. Voltzia Inebeana Geinitz. A Permian species®, characteristic of the Zechstein copper- ~ bearing beds of Gera and other localities, represented by foliage- shoots (fig. 748, C), well preserved cone-scales, and strobili. The vegetative branches closely resemble those of V. heterophylla and Ullmannia selaginoides: there is the same inconstancy in leaf-form as in the Bunter species. The strobili are also similar to those of V. heterophylla: the largest example figured by Geinitz is 2-5 cm. in diameter and 7cm. long. The cone-scales (fig. 748, D—F) have five lobes, deeper than in. V. heterophylla, and the central lobe is longer than the others. The occurrence of three seeds is a characteristic feature; these are ovate, 5 x 3mm., and have a narrow marginal wing. From their close association with strobili Geinitz identified some spherical seeds, formerly described by him as Cyclocarpon eiselianum, as those of V. Liebeana. This author figures an imperfectly preserved strobilus as a male catkin; it resembles those of V. heterophylla, but no microspores have been discovered. Heer describes some foliage-shoots and detached lobed cone-scales from the Permian of Hungary as V. hungarica!: the scales are similar to those of V. Lrebeana but have slightly narrower lobes. Voltzia keuperiana (Schimper). This specific name® was given to fertile shoots characterised by long and lax strobili called by many authors V. coburgensis Schauroth®, a designation first applied to a cast resembling Lygi- nodendron (cf. fig. 401, Vol. 11. p. 37) from the Keuper of Coburg and having no proved connexion with Voltzia. The strobili and 1 Schiitze (01) Pl. x. 2 Heer (65) A. fig. 31; (76) A. Pl. xxx. figs. 4, 5. 3 Geinitz (80) p. 26, Pl. v. 4 Heer (76%). 5 Schimper (72) A. p. 243, Pl. Lxxvi. 6 Schauroth (52) p. 540. 294 CONIFERALES INCERTAE SEDIS [cH. megasporophylls constitute the distinctive features of V. kewper- tana: Schimper figures two strobili approximately 18 cm. long characterised by fan-shaped scales; the lamina has a fairly long stalk gradually passing into a broad rounded distal portion with a crenulate edge, the sinuses between the numerous crenulations being continued as grooves over the face of the expanded portion of the scale. No information is available as to the seeds. The similarity in the general plan of the strobili, apart from the clearly marked distinguishing feature of the megasporophylls, points to a generic affinity between this species and V. Liebeana and V. hetero- phylla. Schimper states that the strobili of V. keuperiana occur in groups in contrast to the solitary cones of other types, and in view of this distinction and the form of the cone-scales he employed the generic name Glyptolepis for which Heer substituted Glypto- lepidium+ on the ground of the previous use of Glyptolepis for a fossil fish. Schimper refers to this species the wood named by Goeppert Araucarites keuperianus (= Dadoxylon keuperianum) but there is no proof of actual connexion. Schenk? adopted the generic name Voltzia and Potonié proposed a new term Voltziopsis® to be used in a provisional and wide sense for Voltzia kewperiana, Cheirolepis Eschert Heer, Heer’s Leptostrobus* and Nathorst’s Swedenborgia® (fig. 749), including species ranging from the Keuper to Middle Jurassic strata in contrast to the Lower Triassic and Permian range of typical representatives of Voltzia. The species Cheirolepis Escheri, included by Potonié in his genus Voltziopsis, was founded by Heer on an imperfectly preserved scale from the Lower Lias of Switzerland resembling the lobed cone-scales of Cheirolepis Miinstert Schenk®. The genus Cheirolepis was in- stituted by Schimper’ as a substitute for Brachyphyllum for the Rhaetic species B. Miinstert Schenk, the new name being chosen because of the digitate margin of the cone-scales which are said to bear single seeds. My former employment of Schimper’s generic term for sterile branches originally named by Phillips Brachy- phyllum setosum® was hardly justifiable in the absence of sporo- 1 Heer (77) ii. p. 72. ® Schimper and Schenk (90) A. p. 290, fig. 199. 3 Potonié (99) B. p. 304. 4 Heer (77) ii. p. 72. 5 Nathorst (78) B. p. 30, Pl. xvi. figs. 6—12. ® Schenk (67) A. p. 187, Pl. xxm. figs. 1—12. 7 Schimper (72) A. p. 247. 8 Seward (00) B. p. 294. XLV] VOLTZIA; SWEDENBORGIA 295 phylls. The species Volizia recubariensis (Mass.) represented by several vegetative shoots, imperfect cones, cone-scales, and seeds in the Muschelkalk beds of Recoaro! illustrates the impossibility in the case of sterile specimens of drawing any satisfactory line between Volizia and Pagiophyllum. Heer instituted the genus Leptostrobus for strobili from Jurassic strata in Siberia agreeing closely in habit and in the form of the megasporophylls with those of Voltzia. The strobili, referred by Heer to three species, do not exhibit any well-marked specific differences; the longest example, L. crassipes, is Tem. in length and 1-5cm. broad: the scales, 7—8 mm. broad, are entire at the distal margin or more or less lobed and in some specimens the scales are hardly distinguishable from those of V. heterophylla. Heer states that two-winged seeds are borne on some of the scales though the evidence is not clear. In a later account Heer? includes in Leptostrobus clusters of long linear leaves apparently borne on short shoots and resembling the needles of Pityites Solmst Sew., Schizolepis Braunit, and Jefirey’s Prepinus. These leaves, described as L. rigida and L. angusti- fola, though in close association with strobili are not actually connected with them; they differ considerably from the short, triangular, imbricate leaves shown in one of Heer’s figures imme- diately below the fertile portion of an axis of L. crassipes®. It would be difficult to draw any / generic distinction between Lepto- strobus and Voltzia especially V. ff keuperiana. B SWEDENBORGIA. Nathorst. This genus? is founded on lax : A oval strobili bearing small cone- scales with long stalks and a Fia. 749. Swedenborgia cryptomerides. single seed (fig. 749, B). In the cA peancnir als 7 leaf. (After type-species, S. cryptomerides, the cones reach 7 cm. in length and the scales are divided distally into 4 or 5 acute digitate lobes. Nathorst compares the strobili 1 Schenk (68) Pls. vit. et seq. 2 Heer (82) p. 23, Pls. vii., vit. 3 Jbid, (77) ii. Pl. x1. fig. 14. 4 Nathorst (78) B. p. 30, Pl. xvz. 296 CONIFERALES INCERTAE SEDIS [cH. with those of Cryptomeria and Voltzia but mentions the presence of a single seed as an important distinguishing feature. To unite Swedenborgia with Voltzia would be misleading, and there is no valid reason for replacing Nathorst’s term by Potonié’s genus Voliziopsis. Strobilites. Sirobilites lacus Seward. This name was applied to a lax strobilus, 39 cm. long and 1-3 cm. broad at the base, from Rhaetic beds on the Orange River, South Africa!, which may be allied to Voltzea, though in the absence of ‘seeds its position cannot be determined. The sporophylls consist of a lamina with a rounded distal edge and a radially folded surface attached by a short horizontal stalk resembling the seed-bearing scales of Voltzia heterophylla, V. coburgensis, and to some extent ~ Heer’s Jurassic Leptostrobus. ULLMANNIA. Goeppert. Goeppert? in his description of Ullmannia refers to the extensive literature on the fossils from the Permian copper mines of Frank- enberg on which the genus was founded: the most complete of these earlier accounts is that of Ullmann. In habit similar to Walchia, Ullmannia is represented by various forms of foliage- shoots and impressions of buds and cones, but the data are in- sufficient to settle its position in the Coniferales. Ullmannia Bronni (fig. 750, D), the type-species, is practically identical in leaf-form and habit with the Mesozoic genus Pagiophyllum, while the species U. frumentaria (fig. 750, A) agrees closely with such recent Conifers as Araucaria excelsa and A. Bidwill. The branches bear spirally disposed crowded leaves with a median vein and numerous longitudinal striations on the lamina. The association of impressions of foliage-shoots with wood having the Araucarian type of pitting? affords contributory evidence, though by no means proof, of Araucarian affinity. In the absence of any definite information as to the structure of the reproductive shoots Ull- mannia must be left for the present as a Conifer which cannot be assigned with certainty to a systematic position. Tuzson? uses + Seward (08) B. p. 101, text-fig. 7; Pl. v. fig. 3. 2 Goeppert (50) p. 185. * Solms-Laubach (84) Pl. m1. fig. 16; Schimper and Schenk (90) A. p. 275, figs. 190, 191. 4 Tuzson (09) p. 23. XLV] ULLMANNIA 297 the generic name Ullmannites for wood having the Araucarian type of tracheal pitting which he believes to belong to Ullmanmia, but under the former genus are included types of wood that are not generically identical and afford no evidence of connexion with Ullmannia. Ulimannia is essentially a Permian genus especially characteristic of the copper-bearing rocks of Frankenberg in Hessen and Ilmenau in Thuringia; it is recorded also from France and by Eichwald and Schmalhausen from Russia, the species Fic. 750. Ullmannia. A—C, Ullmannia frumentaria. D, E, U. Bronni. (A, after Geinitz; B, C, after Solms-Laubach; D, E, after Potonié.) U. biarmica Eich.1 being represented by both sterile and fertile branches. Imperfectly preserved impressions from the Permian of Durham? have been referred to Ullmannia but no satisfactory specimens have been discovered in English beds. The fragment figured by Lindley and Hutton® as Voltzia Phillipst may belong to an Ullmannia. 1 Schmalhausen (87) Pl. v1. 2 Murchison and Harkness (64) p. 154; Kirby (64). 3 Lindley and Hutton (37) A. Pl. 195. 298 CONIFERALES INCERTAE SEDIS [cH. Ulimannia Bronni Goeppert. The fragments of foliage-shoots on which Goeppert! founded this species were described by some earlier authors as the Frankenberg ears of corn and by Schlotheim as Poacites phalaroides. Bronn, who first identified the fossils as Coniferous, named them Cupressites Ulimanni. We know nothing of the structure of the reproductive shoots, and the cones referred by Goeppert to this species have since been assigned to the genus Strobilites. In habit and leaf-form Ullmannia Bronni is indistinguishable from certain Triassic and Ju- rassic foliage-shoots referred by most authors to Pagiophyllum. The leaves are imbricate (fig. 750, D, E); the lamina is oval or broadly linear, elliptical, and characterised by longitudinal striations. Some imperfectly preserved leaves examined by Solms-Laubach showed clear indications of the presence of a midrib with lateral groups of transfusion-tracheids, as in U. selaginoides and U. frumentaria. Ullmannia selaginoides (Brongniart). This and the following species, U. frumentaria, are founded on vegetative shoots from the Permian of IJmenau (Thuringia) and both were described by Brongniart as examples of Fucoides?. The leaves of U. selaginoides are longer than those of U. Bronni, linear and almost uniform in breadth, elliptical in section. There is a single vascular bundle accompanied by wings of reticulate trans- fusion-tracheids (ef. fig. 750, C) associated with parenchyma®. As in the leaves of recent Conifers the transfusion-tissue persists in the apical region of the lamina. There are 1—2 rows of hypo- dermal fibres below the epidermis with sunken stomata and the mesophyll consists largely of palisade-cells (cf. fig. 750, B). The stele of the shoot has a large pith with nests of dark cells enclosed by a cylinder of secondary xylem consisting of tracheids having a single row of separate circular pits on the radial wall and uniseriate medullary rays 1—6 cells deep. Ullmannia frumentaria (Schlotheim). This species, originally named Carpolithes frumentarius4, is the commonest fossil in the IImenau mines. The leaves are lanceolate, 1 Goeppert (50) p. 185, Pl. xx. For synonymy, see Solms-Laubach (84); Geinitz (80). ? Brongniart (28) A. Pl. 1x. 3 Solms-Laubach (84) Pl. 11. figs. 1, +, 6, 15. 4 Schlotheim (20) A. Pl. xxvii. fig. 1 For figures, see Geinitz (80); etc. XLV] ULLMANNIA; STROBILITES 299 acute, decurrent, and more or less falcate and like those of other species characterised by longitudinal striae on the dorsal face (fig. 750, A). Well-preserved impressions of this species in the Dresden Museum bear a close resemblance to shoots of Araucaria Bidwilli and A. brasiliensis. Some specimens show laterally attached oval cones, but it is not clear if these are reproductive shoots or vegetative buds and nothing is known as to the nature of the sporophylls. The seeds, Cardiocarpus triangularis1, referred by Geinitz to this species on the ground of association, cannot be safely assigned to Ullmannia. In leaf-structure (fig. 750, B, C) U. frumentaria closely resembles U. selaginoides but the hypo- dermal fibres form oval strands instead of 1—2 layers. The species has also been recorded from the Permian of France’. Strobilites. Strobilites Bronni (Goeppert). Under this provisional name Solms-Laubach® described the problematical fossils from the Frankenberg copper mines which Goeppert believed to be the megastrobili and cone-scales of Ullmannia Bronm. In the absence of any satisfactory evidence of connexion with the vegetative shoots on which U. Bronni was founded it is better to follow Solms-Laubach in the adoption of the non-committal name Strobilites. These star-stones (‘Sterngrau- pen’) of the miners consist of more or less circular bodies bearing some resemblance to the peltate cone-scales of Cupressus; they occur either singly or in cone-like groups. No seeds have been found attached to the scales nor is there any proof that they were borne by a Conifer. The larger scales, 15—25 mm. in diameter, are characterised by 8—12 radial ridges and a central depression (? umbo), and to the under surface is attached a cylindrical stalk usually in the centre but occasionally excentric. The scales are sometimes found in almost spherical clusters and another form described by Solms-Laubach consists of a cylindrical cone-like aggregate 4m. x 2 cm. of rather smaller scales without radial ribs and characterised by a stout, rounded, peripheral border. 1 Geinitz (80) Pl. m1. figs. 11—15. 2 Zeiller (06) B. p. 219, Pl. x. 3 Solms-Laubach (84). 300 CONIFERALES INCERTAE SEDIS [cH. ALBERTIA. Schimper. This generic name was given by Schimper to vegetative branches from the Bunter of the Vosges agreeing in habit with shoots of some species of Agathis but differing in the broader insertion of the lamina. For Albertia Endlicher! substituted Haidingera. Schimper and Mougeot? figure reconstructions of both male and female cones and a single cone-scale bearing a median seed. Schenk?, who examined the original specimens, states that the supposed male cone is a young megastrobilus of Voltzia, and Solms-Laubach4, who also examined the material in the Strassburg Museum, considers that Schimper’s statement that the seed-scale and cones belong to Alvertia is ‘altogether arbitrary and unsupported.’ There would seem to be no reason‘for connect- ing the cones figured by Schimper and Mougeot with the shoots referred by those authors to four species of Albertia®. Albertia lati- folia is founded on branches bearing fairly large (2-5 x 1 cm.), obovate, slightly decurrent leaves with numerous longitudinal stria- tions. The branches described as A. ellvptica are not distinguishable by any clearly marked feature from A. latifolia. Albertia Braunw has larger obovate leaves and A. speciosa has broadly linear leaves reaching a length of 45cm. There is a very close resemblance between the shoots from the Bunter beds and those of some forms of Ullmannia especially U. frumentaria (Schlot.)®, and it is doubt- ful whether any useful purpose is served by the retention of the designation Albertia: the descriptions of the reproductive shoots are misleading and rest on no substantial basis and the sterile branches exhibit no characters by which they can be generically separated from Ullmannia. The important point is that there are no grounds for regarding the specimens usually referred to Albertra as Araucarian other than the uncertain and untrustworthy evidence afforded by a similarity to Agathis. The fragmentary impression from the Karharbari beds of India assigned by Feistmantel’ to 1 Endlicher (47) p. 303. 2 Schimper and Mougeot (44) A. Pl. 1. 3 Schenk in Schimper and Schenk (90) A. p. 284. 4 Solms-Laubach (91) A. p. 75. 5 Schimper and Mougeot (44) A. Pls. 1.—v. ® Geinitz (80) Pl. m1. * Feistmantel (79) p. 29, Pl. xxvt. fig. 2. XLV | ARAUCARINEAE 301 Albertia is too imperfect to be determined ; it may be identical with those referred by the same author to Voltzia. The abundance and wide distribution of wood with Araucarian features in Palaeozoic rocks though, for reasons already stated, not admissible as proof of the occurrence of members of the Araucarineae, at least shows the great antiquity and predominance of the Araucarian type. There can be no reasonable doubt that much of the wood described in Chapter xxxu1. as Dadoxylon belonged to Araucarian plants, more especially the examples furnished by Mesozoic and Tertiary strata. In considering the past history of the family the evidence of the wood must be taken into account. The conclusions drawn from a survey of the fossil records are: (i) the type represented by Araucaria is older than that now illus- trated by Agathis. In other words Araucaria possesses features, especially those associated with the megastrobili, which extend farther back without departing far from the existing type thanis the case with Agathis. (ii) The Araucarineae foreshadowed in the later Carboniferous and earlier Permian periods were in all probability established as a family in Rhaetic times, and in the Jurassic and earlier Cretaceous periods the Araucarineae were almost cosmo- politan and represented by numerous forms. (ili) Such evidence as is afforded by Tertiary records, though meagre and often in- complete, points to the continued existence of the family in the Northern Hemisphere at least in the older Tertiary floras. CHAPTER XLVI. CUPRESSINEAE. Tue published records of fossil Conifers would seem to justify the conclusion that the Cupressineae were widely distributed and represented by a wealth of genera during the latter part of the Mesozoic era particularly in the later Jurassic floras, but on closer inspection of the material a student, having any familiarity with the external features of recent genera, cannot fail to recognise the wholly inadequate nature of the data on which the systematic determinations are based. It is undoubtedly true that in the later Jurassic and Lower Cretaceous floras Conifers agreeing generally in habit and in the possession of appressed imbricate leaves with such genera as Cupressus, Chamaecyparis, and Thuya were among the most characteristic types: some have the leaves in decussate pairs with an occasional tendency to a spiral phyllo- taxis while others possess leaves of the same form but spirally disposed. Almost all are sterile and when cones are present the form and arrangement of the scales often suggest comparison with recent types other than the Cupressineae. In the Chapter on recent Conifers attention is called to the inconstancy of leaf- arrangement in certain species and to the close resemblance between vegetative shoots of plants belonging to different families. Fossil coniferous branches referred by authors to the Cupressineae afford a striking illustration of the insufficiency of the evidence on which sterile impressions have been named. This statement, though primarily concerned with Mesozoic records, applies also to many Tertiary species. The records of the rocks clearly show that European Tertiary floras contained a considerable number of Cupressineous types that are now confined to other regions, but a critical examination of the older fossils leads to the conclusion that in very many cases accurate determination of the affinities of sterile branches, superficially resembling existing members of the Cupressineae, is impossible without additional information. * CH. XLVI] THUYTES; CUPRESSINOCLADUS 303 In the absence of well-preserved cones or anatomical data it is possible that a comparative examination of cuticular membranes might furnish useful results. It is, however, only rarely that such information can be obtained and the only safe course to follow is to use, with greater freedom than has generally been the practice, provisional generic names which do not imply affinities to recent genera. Provisional names that have reference only to vegetative features should be superseded by designations denoting characters of greater taxonomic significance when the necessary information is available. As a preliminary to the description of a few selected types it may be useful to consider the sense in which some generic names have been employed and at the same time to state whether the retention of certain names or their use in a modified sense is advisable. Thuytes Brongniart. Brongniart! instituted this term for ‘Branches like those of Thuya; fruit unknown.’ The name, in the form Thuyites, had been used a few years previously by Sternberg and it was adopted by Unger? as Thuites for both sterile branches and cones similar to those of the recent genus. Thuytes has been widely used for , vegetative branches agreeing generally in habit with those of Thuya or Cupressus and some other Cupressineae, but with the exception of some Tertiary species the designation has reference in: nearly all cases to the form and arrangement of the leaves. The employment of Thuytes in this wide sense is open to criticism on the ground that in accordance with the usual practice specimens so named would be considered to be more nearly related to Thuya than to any other genus. It is therefore proposed to adopt the generic designation Thuites only for such specimens as afford evidence of close affinity to the recent genus and to discontinue its use for sterile shoots which suggest comparison not only with Thuya but also with Cupressus, Libocedrus and other genera. CUPRESSINOCLADUS. Gen. nov. . Goeppert’s term Cupressites® is retained for fossils which there is reason for associatirig with Cupressus and should not be employed 1 Brongniart (28) A. p. 109; Seward (04) B. p. 140. 2 Unger (50) A. p. 346. - 3 Goeppert (50) p. 183. 304 CUPRESSINEAE [CH. in a wider sense. Bowerbank! adopted the form Cupressinites for some fossil cones from the London Clay in order to avoid the implication of affinity only to Cupressus which is suggested by Cupressites. This generic name would be convenient for Cupres- sineous branches had it not been restricted in the first instance to cones: to avoid the revival of a term and its employment in a new sense it is proposed to adopt the name Cupressinocladus for vege- tative shoots agreeing in the habit of branching and in the predomi- nance of a decussate arrangement of appressed leaves with recent Cupressineae such as Cupressus, Thuya, Libocedrus and similar types. When cones are present which throw any light on generic affinity some other term should be adopted. It will, however, be found in practice that the choice of the most appropriate name is exceedingly difficult; and no sharp line can be drawn between. certain specimens which conform in part to Cupressinocladus and in part to the characters of Brachyphyllum. Palaeocyparis Saporta. The published illustrations of Conifers included by Saporta? in this genus afford examples of the inconstancy of leaf-arrangement in a single type and demonstrate the impossibility of drawing any definite distinction between this genus and Thuytes as used by Saporta. With one exception all the specimens from Jurassic rocks referred by Saporta to his genus are sterile and in habit agree with several recent genera of the Cupressineae particularly Cupressus, Chamaecyparis, and Thuya. In Palaeocyparis are in- cluded species previously referred to Echinostrobus, Thuytes, and Athrotaxites, The branching is in one plane; the leaves are stated to be usually though not invariably decussate, more or less tri- angular, appressed, and imbricate, rarely free at the apex of the lamina. The supposed cone described in the case of Palaeocyparis elegans, a species from Upper Jurassic beds in France, is only 7 by 9 mm. and it is not clear whether it is a true cone or a vegetative bud or perhaps a male flower. The genus is practically founded on vegetative characters only. An objection to the retention of Saporta’s term is that several of the specimens may legitimately be included in a previously established genus Brachyphyllum. 1 Bowerbank (40) p. 51. 2 Saporta (84) p. 574, Pls. 202 et seq. 3 Ibid. Pl. 214. XLVI] THUITES; CUPRESSITES 305 THUITES. Brongniart emend. As stated on a previous page it is proposed to limit this name to fossils affording good evidence of close affinity to the recent genus Thuya. Among the few examples that appear to fulfil this con- dition are those described as Thuya occidentalis L. succinea Goepp.* and Biota orientalis Endl. succinea Goepp.” from the Oligocene amber deposits on the Baltic coast. Schlechtendal? records specimens from Oligocene beds at Weimar, consisting of shoots with opposite pairs of leaves, on which a resin-canal is seen below the apex, and the remains of male flowers, which he names Thuya occidentalis var. thuringica. CUPRESSITES. Goeppert. Cupressites MacHenryt Baily. This name was given* to a piece of shoot of Cupressus-like habit from the Eocene beds of Antrim. Baily’s figure, which Gardner says is inaccurate, shows the small scale-leaves as spiral, but the specimens subsequently described by Gardner® from the same locality leave no doubt as to the decussate arrangement of the foliage. Gardner adopts the name Cupressus Pritchardi on the assumption that the fossil wood from Lough Neagh described by Goeppert® as Pinites Pritchardi belongs to the species which furnished the Antrim specimens: the wood, subsequently referred to Cupressinoxylon’, affords a typical example of that genus as is clearly shown by the photographs reproduced in fig. 715 (p. 188), but there is no proof of any connexion between it and the branches from the Antrim leaf-beds. The shoots are characterised by their slender pinnately arranged branchlets with small decussate leaves, and the cones, about 14mm. long, bear 10 hexagonal scales of the Cupressus type very similar to those of Cupressites taxiformis. No seeds were found attached to the cone-scales. The abundance of specimens in the Irish beds indicates that this Cupressineous species was a common tree in the forests which 1 Goeppert and Menge (83) A. p. 43, Pl. xv, figs. 199—206. 2 Ibid. p. 42, Pl. xv. figs. 180—198. 3 Schlechtenda] (02) Pls. 1., 11. 4 Baily, W..H. (69) p. 361, Pl. xv. fig. 5. 5 Gardner (86) p. 82, Pl. xvi. figs. 8,9; Pl. xvii. fig. 1; Pl. xxx. § Goeppert (50) p. 220. 7 Kraus in Schimper (72) A. p. 376. s. IV 20 306 CUPRESSINEAE [CH. flourished on the western edge of Europe during the period of volcanic activity responsible for the widespread sheets of lava in the North-East of Ireland and the Western Isles of Scotland. Cupressites taxiformis Unger. This species was founded by Unger! on sterile and fertile shoots from Eocene beds in the Tyrol. Many of the sterile branches are similar in the form of the linear leaves to Taxus, but other leaves are appressed to the axis and free only at the apex, resembling on a small scale those of Sequowa gigantea. The cones consist of polygonal, peltate, scales probably verticillate and superficially similar to the strobili of Cupressus: they are borne on shoots with scale-like leaves. The species is recorded by De la Harpe? from the Isle of Wight and several specimens are figured by Gardner? from the Middle Bagshot beds of Bournemouth. The material from the latter locality consists of sterile shoots with linear Tazus- like leaves from 5 to 15 mm. long and occasionally, on the same axis, smaller decurrent leaves, though generally the two forms are found on different twigs. The cones occur only in connexion with the shoots bearing small ap- pressed leaves: the specimen shown in fig. 751 has been re-drawn from one of Gardner’s figured specimens?: it is characterised by peltate scales with a central umbo and a wrinkled surface. The evidence in favour of assigning all the 5.4. 751. Curmesilies sterile shoots to the same species is not con- — jaziformis. (From vincing, though a similar combination is met a specimen in the with in the recent species Glyptostrobus hetero- Baers ee phyllus. If we confine our attention to the cones avs they may reasonably be retained in the genus Cupressites and regarded as evidence of the existence in Western Europe in the Eocene period of a type closely allied to the genus Cupressus. The position of the sterile shoots cannot be determined without further investigation. 1 Unger (47) p. 18, Pl. vim. figs. 1—3; Pl. rx. figs. 1—4. 2 De la Harpe in Bristow (62) Pl. v. fig. 2. 3 Gardner (86) p. 26, Pl. 1. figs. 1—13; Pl. v. figs. 13, 14; Pl. vu. fig. 8; Pl. rx. figs. 22—26, 28—30. 4 Ibid, Pl. 1x. fig. 27. XLVI] CUPRESSINOCLADUS 307 Another Tertiary representative of the recent genus Cupressus is that described from the Oligocene amber beds of Hast Prussia as Cupressus sempervirens L. succinea Goepp. and Menge!. This species is founded on fragments of sterile shoots with a well pre- served male flower showing very clearly the form of the sporophylls. CUPRESSINOCLADUS. Gen. nov. The following examples of Cupressineous shoots that do not afford satisfactory evidence of relationship to any particular recent genus are given in illustration of the desirability of employ- ing such a non-committal generic term as Cupressinocladus. (i) Species previously referred to Libocedrus. The Cretaceous specimens from the Atane beds of West Green- land described by Heer? as Libocedrus cretacea are unaccompanied by any cones and may equally well be compared with species of Thuya: specimens from the Amboy clays, believed to be identical with Heer’s, are assigned to the latter genus by Newberry?. Cupressinocladus salicornoides (Unger). A sterile piece of branch figured by Lindley* from Provence as Thuya is probably identical with Unger’s species recorded by Saporta® from the same locality. The type-specimens on which Unger founded the species Thuites salicornoides (fig. 752) are from Eocene beds in Croatia®; they do not bear any mature cones and cannot be assigned with confidence to any recent genus. The flattened shoots bear appressed leaves in decussate pairs and the decurrent lamina may reach a length of 15cm. By later authors this species, recorded from Styria’, Bohemia®, Switzerland®, the Oligocene beds of East Prussia’, Italy, the Miocene of France and from other localities, is spoken of as Lebocedrus, but the few examples of cones that have been figured do not exhibit with 1 Goeppert and Menge (83) A. p. 45, Pl. xv1. figs. 218—224. 2 Heer (82) i. p. 49, Pl. xxrx. figs. 1, 2; Pl. xxi. fig. 1d. 3 Newberry and Hollick (95) p. 53, Pl. x. figs. 1, la. 4 In Murchison and Lyell (29) p. 298, fig. B. 5 Saporta (65?) p. 42, Pl. 1. fig. 4. 8 Unger (47) p. 11, Pl. 11. ? Ettingshausen (70) p. 39; (88) p. 273. 8 Ibid. (672) p: 109, Pl. v. figs. 1—7, 14; Engelhardt (85) Pl. vit. figs. 27—30. ® Heer (55) A. p. 47, Pl. xx1. fig. 2. : 1) Goeppert and Menge (83) A. PL. xv. figs. 175—177. 11 Massalongo (59) p. 153, PI. v. figs. 20—23. 12 Marty (08). 20-2 308 CUPRESSINEAE [CH. sufficient clearness morphological features that justify the con- clusion that the specimens agree more closely with Libocedrus than with Thuya or Thujopsis. The present discontinuous dis- tribution of Libocedrus (page 126) is favourable to the view that it was formerly much more widely spread, but despite the very Fic. 752. Cupressinocladus salicornoides. (After Unger; nat. size.) close resemblance between the sterile shoots of the Tertiary Conifer to those of some existing species of the genus it would be unwise to adopt the designation Libocedrus or Libocedrites. Heer? described fragments from Miocene strata in Greenland as Libocedrus Sabiniana including a supposed cone-scale, which is too indistinct to be determined. To this species Beust refers some petrified wood from the same locality. The sterile specimens recorded by Gardner? from the Woolwich beds of Kent as Libo- cedrus adpressa, though similar to L. salicornoides, are no doubt specifically distinct: they are valueless as evidence of the existence of Libocedrus. Laurent? also records the species from Aquitanian beds in the Puy-de-Déme. 1 Heer (83) p. 58, Pls. LXX., LXXXVI., LXxxvul.; Beust (85) p. 40, Pls. o1., v. 2 Gardner (86) p. 25, Pl. 1. figs. 17—20. 3 Laurent (12) p. 69. XLVI] CUPRESSINOCLADUS . 309 (ii) Species previously referred to Thujopsis. Cupressinocladus massiliensis (Saporta). The small twigs de- scribed by Saporta from Provence as Thujopsis massiliensis! and compared by him with Thuya occidentalis are very similar to those named by the same author 7’. ewropaea®: in neither case is there any justification for the use of the generic name Thujopsis. Heer? records 7’, europaea from Miocene beds in Greenland and in a later account adopts the name Biota orientalis on the ground of the occurrence of imperfect cones and elongated cone-scales comparable with those of Biota (Thuya), but the figured specimens are too indistinct to warrant the employment of the generic term Thuites in the more restricted sense advocated above. Goeppert and Menge* refer some fragments from the Baltic amber beds to Thujopsis europaea, but no reproductive organs are figured. (ili) Species previously referred to Thuya or Thuites. Several Jurassic and Lower Cretaceous specimens formerly included in Thuites are now transferred to Brachyphyllum on the ground that the choice of one or other of these names has fre- quently been determined by characters that are both inconstant and of little morphological importance. There are, however, several examples of Coniferous shoots from Mesozoic and Tertiary strata that are clearly distinguished from such types as Thuttes expansa (= Brachyphyllum expansum), in which the verticillate arrangement of the leaves is not a well marked or constant feature, by the very regular disposition of appressed leaves in decussate pairs as in recent species of Libocedrus and some other Cupres- sineae: for this form of shoot the generic name Cupressinocladus is now adopted. : Cupressinocladus valdensis Seward. A species described as Thuites valdensis® from a single specimen from Wealden beds on the Sussex coast, characterised by decussate appressed leaves with a comparatively long basal portion con- crescent with the axis of the branch and a free short triangular 1 Saporta (65) p. 72, Pl. 1. fig. 6; Pl. rv. fig. 2. 2 [bid (65%) Pl. 1. fig. 5. , 3 Heer (68) i. p. 90, Pl. x. fig. 11; (75) iii. p. 7, Pl. 1. figs. 13—29. 4 Goeppert and Menge (83) A. Pl. xvr. figs. 215—217. 5 Seward (95) A. p. 209, Pl. xx. fig. 6. 310 CUPRESSINEAE _‘([cH. apex, agreeing closely with some forms of Libocedrus and with the younger branches of Frenelopsis. Though accurate determination of the position of such specimens is impossible, they afford evidence of the fairly widespread occurrence of Conifers in Mesozoic and Tertiary strata exhibiting a striking resemblance in habit to recent Cupressineous genera. Vegetative branches from Miocene beds in Spitzbergen and Greenland referred by Heer! to Thuttes Ehrenswaerdi and T. Mervana respectively afford examples of specimens which would be more appropriately included in the genus Cupressinocladus. Similarly the fragments described by Goeppert and Menge as Thuya Mengeana Goepp.? afford no convincing evidence of generic identity with the recent genus. (iv) Species previously referred to Juniperus or Juniperites. Cupressinocladus hypnoides (Heer). The slender sterile branches from the Lower Cretaceous beds on the West of Greenland described as Juniperus hypnoides® afford no substantial evidence of relationship to Juniperus rather than to some other member of the Cupressineae or Callitrineae. Specimens from the same locality which may be specifically identical with Juniperus hypnoides are described by Heer as J. macilenta*. The leaves are very small and occur on the slender axes in opposite pairs. Newberry records J. macilenta® from the Amboy clays and states that cone-scales of Dammara are associated with the foliage-shoots, though Hollick in a note to the description says that he was unable to find any such scales with the vegetative branches. Hollick and Jeffrey® figure specimens from the lignite beds at Kreischerville as J. hypnoides and believed them to be identical with those described by Newberry as J. macilenta; they also speak of the association of Dammara [Agathis] scales. The Tertiary Greenland species J. tertiarius and J. gracilis’, founded by Heer on sterile branches, are equally unsatisfactory as 1 Heer (71) iii. p. 38, Pl. 1. figs. 25, 26; (82)i. Pl. 1. 2 Goeppert and Menge (83) A. p. 44, Pl. xvr. figs. 211—214. 3 Heer (82) i. p. 47, Pl. xiv. fig. 3; Pl. xiv. fig. 18. 4 Ibid. Pl. xxxv. figs. 10, 11. ® Newberry and Hollick (95) Pl. x. fig. 7. 6 Hollick and Jeffrey (09) B. p. 61, Pl. v. figs. 5, 6. 7 Heer (83) p. 57, Pls. LXx., CIT., CVI. XLVI] ECHINOSTROBUS, ETC. 311 records of Conifers closely allied to Juniperus, and the same remark applies to Juniperites eocenica described by Ettingshausen! from Haring in the Tyrol. A single male flower figured by Goeppert and Menge? from the Baltic amber as Juniperus Hartmannianus may be correctly referred to that genus though other recent genera are not excluded. Echinostrobus Schimper. Proposed in the first instance by Schimper? for Unger’s Athro- taxites lycopodioides, this term was adopted for several sterile shoots such as those named by Brongniart Thuytes expansus, characterised by the possession of decussate leaves like those of Thuya and Cupressus with others agreeing more closely with Brachyphyllum. As the name has reference to the spinous nature of the cone-scales, and as it is now agreed that Unger’s earlier name Athrotaxites may be appropriately employed, Echinostrobus is dis- carded. Phyllostrobus Saporta. , This generic name was given by Saporta* to an Upper Jurassic fertile shoot with whorled leaves of the Thuites form bearing a single cone compared with those of Libocedrus. The impression conveyed by Saporta’s figures is that the preservation of the cone is too imperfect to warrant the institution of a new generic term. Condylites Thiselton-Dyer. This name applied to specimens from the Solenhofen slates® has reference to the elbow-like insertion of lateral branches: the foliage is like that of Brachyphyllum, and the cones, which are imperfectly preserved, are compared with those of Thuya. As in the case of Saporta’s Phyllostrobus the cones are too obscure to admit of any satisfactory description. Athrotacxites Unger. This name was proposed by Unger for a branched cone-bearing shoot from Solenhofen agreeing in vegetative characters with 1 Ettingshausen (55) Pl. v. fig. 6. 2 Goeppert and Menge (83) A. p. 39, Pl. xiv. figs. 156, 157. 3 Schimper (72) A. p. 330. ‘4 Saporta (84) p. 635. 5 Thiselton-Dyer (72). - 8 Unger (49) Pl. v. figs. 1, 2. 312 CUPRESSINEAE [CH. specimens previously figured by Sternberg as Caulerpites. The specimen in the Munich Museum on which the type-species Athro- taxites lycopodioides was founded was examined and re-figured by Schimper! and by Saporta (fig. 753, C)?: the former author substituted for Athrotaxites a new genus Echinostrobus in order to avoid the implication of relationship with Athrotaxis which he was not prepared to accept, and without adequate reason altered Unger’s specific name lycopodioides to Sternberg. Saporta, who believed Unger’s type to be intermediate between Athrotaxis and Cryptomeria as regards the features of the cones, retained Schimper’s designation Echinostrobus Sternbergit. Unger? in sub- sequent accounts of Solenhofen plants extended the application of Athrotarites to sterile shoots, and this course was also followed by Thiselton-Dyer? who expressed agreement with Unger as regards the resemblance of the fossil cones to those of the recent genus. In accordance with the principle advocated on a previous page it is suggested that Unger’s generic name should be retained only for specimens which afford evidence, other than mere resemblance of foliage-shoots, of affinity to the recent genus Athrotaxis: this use of Unger’s term has recently been adopted by Halle®. Most of the sterile specimens referred to Athrotaxites by Unger and other authors should be transferred to Brachyphyllum. Athrotaxopsis Fontaine. Fontaine® instituted this name for some fertile Coniferous shoots from the Potomac formation similar in habit to species in- cluded by Saporta in his genus Palaeocyparis and to the genus Thu:tes but bearing cones different from those of Athrotaxites lycopodioides. Berry’ subsequently reduced the number of Fon- taine’s species and pointed out that some of his specimens are indistinguishable from shoots included in Sphenolepidium, a con- clusion to which I had been led® by a comparison of Wealden specimens of Sphenolepidium Kurrianum with Fontaine's figures. Some of the Potomac examples agree in vegetative characters with the genus Brachyphyllum. In the absence of any substantial 1 Schimper (74) A. Pl. uxxv. fig. 21. 2 Saporta (84) Pl. 199. 3 Unger (52); (54°). 4 Thiselton-Dyer (72). 5 Halle (13) p. 40. 6 Fontaine (89) B. p. 239. ? Berry (11%). 8 Seward (95) A. p. 202. XLYI] ATHROTAXITES 313 grounds for assuming any direct relationship between the Potomac Conifers and Athrotaxis, and in view of the fact that the American specimens can be accommodated in previously instituted genera, there are no good reasons for retaining Fontaine’s name. ATHROTAXITES. Unger emend. The Tasmanian genus Athrotaxis (page 150) is one of the existing Conifers of which our knowledge of morphological features is very Fic. 753. A, B, D, Athrotaxites Ungeri. C, Athrotaxites lycopodioides. (A, B, D, after Halle; A, nat. size. C, after Saporta; nat. size.) incomplete: its restricted range and the striking resemblance of the vegetative characters to those of many Jurassic species in- cluded in Brachyphyllum suggest antiquity, but palaeobotanical records have furnished very little evidence of value in support of this conclusion. It is however probable that like several other plants now confined to the southern hemisphere Athrotazis is a 314 CUPRESSINEAE [cH. survival of a type of Conifer which was widely spread in Jurassic floras. Athrotaxites lycopodioides Unger (= Echinostrobus Sternbergu Schimp.). Reference has already been made to the vegetative characters | of this Upper Jurassic species from Solenhofent. In the crowded imbricate leaves and in the blunt stiff branches it agrees very - closely with Athrotaxis cupressoides: the globular cones (fig. 753, C), though incompletely preserved, also exhibit in the comparatively small number of cone-scales and their thick spinous distal ends a distinct similarity to those of the recent genus. Nothing is known of the seeds. Athrotaxites Ungert Halle. This species”, founded on fertile specimens from the San Martin flora of Patagonia, probably of Upper Jurassic or Wealden age, is practically identical with Unger’s type: the branches are more slender and the globular cones, 10—15 mm. in diameter, appear to be of the same type. The cone-scales have a cuneate base and a thick spathulate distal end prolonged into a short pointed apex (fig. 753, A, B); the thickening of the scales close behind the apex recalls the form characteristic of recent. cones (cf. fig. 684, N, p. 116). As Halle says, the vegetative features (fig. 753, D) of this and the preceding type are those of Brachyphyllum. Jn his memoir on British Eocene Gymnosperms Gardner? referred to Athrotaxis some of the foliage-shoots and cones from Bovey Tracey in Devonshire which had previously been included in Sequoia Couttsiae Heer: the reasons for the change of genus are by no means adequate. Mr and Mrs Clement Reid¢ in their recent investigation of the Bovey Tracey material, which they refer to an Upper Oligocene age, made a careful examination of numerous Sequoia fragments including a comparison of fossil cuticular membranes with the epidermis of both Sequoia and Athrotaxis leaves: they were unable to discover any evidence of the presence of representatives of the latter genus. Gardner also assigns some 1 See page 312; Unger (49). 2 Halle (13) p. 40, Pls. m.—yv. 3 Gardner (86) p. 90, Pl. vi. figs. 1—9; Pl. x. figs. 6—9. 4 Reid, C. and E. M. Reid (10) p. 171. XLVI] . - | BRACHYPHYLLUM 315 pieces of vegetative organs and in one case a cone to Athrotaxis, but the evidence on which the species Athrotaxis (?) subulata} is founded has little value. BRACHYPHYLLUM. Brongniart. Brongniart? proposed this name for a Jurassic species, Brachy- phyllum mamillare, founded on sterile branches characterised by pinnate branching in one plane and spirally disposed appressed leaves with a thick lamina of triangular, conical, or hexagonal form. He afterwards? extended the term to other Jurassic species and called attention to the striking resemblance of the fossil shoots to those of Athrotaxis. The photograph of Athrotaxis cupressoides shown in fig. 701 (p. 150) affords a very good idea of the habit of Brachyphyllum. Specimens in which the pinnate ramification is a conspicuous feature are more like shoots of Thuya or Cupressus, and on the smaller branches the leaves may assume a decussate arrange- ment. On older branches the leaves are often hexagonal and more or less convex, while on the branchlets they are more triangular or conical and are free at the apex, which in some forms is bent out- wards from the axis (fig. 756). In nearly all cases specimens referred to Brachyphyllum are sterile and, except in examples where the preservation of the cones is too imperfect to afford any evidence of morphological characters, it is suggested that the generic name should be reserved for sterile branches and regarded as purely provisional. As Saporta‘ points out in his account of the genus, considerable confusion has been caused by attempts to assign species to. several positions in the Coniferales on wholly insufficient evidence. Unger® figured a fertile specimen from the Rhaetic rocks of Franconia, which he referred to Brachyphyllum speciosum Miinst., and this was re-figured by Schenk® as Palissya aperta though as others have pointed out the cones are very different from those of Palissya: as Nathorst’ says, they have the characters of the genus Hlatides, and the same is true of some cones figured by Saporta® and assigned by him to Brachyphyllum. The name Elatides is reserved for specimens characterised by a certain 1 Gardner (86) p. 43, Pl. x1. 2 Brongniart (28) A. p. 109. 3 Ibid. (49) A. p. 69. 4 Saporta (84) p. 310. 5 Unger (49) Pl. v. figs. 3, 4. 8 Schenk (67) A. Pl. xxi. figs. 1—13. 7 Nathorst (97) p. 34. 8 Saporta (84) Pl. 165, fig. 1; Pl. 167, figs. 2, 3; Pl. 171, figs. 7—9. 316 CUPRESSINEAE [cH. type of cone (fig. 742) and although typical examples of the genus bear leaves differing in their greater freedom from the axis from the more concrescent foliage of Brachyphyllum, it is not possible in all cases to draw a definite line between the two forms of shoot. Until a few years ago nothing was known as to the anatomical features of Brachyphyllum but the researches of Hollick and Jeffrey! have partially made good this deficiency : these authors investigated the structure of Brachyphyllum macrocarpum Newb. (= B. crassum) (fig. 758, G) from the Cretaceous beds of Kreischerville and demon- strated a close resemblance in some characters to recent Araucarias. They fully recognise that it would be unsafe to assume the presence of similar anatomical features in other species, though it is reason- able to expect the occurrence of such characters in many species not yet found as petrifications. Other examples of shoots of the Brachyphyllum habit furnishing information with regard to anatomy are B. eathiense Sew.? and Banc. and a Japanese species originally described as Yezonia vulgaris by Drs Stopes and Fujii’ and since transferred to Brachyphyllum. As the great majority of specimens referred to Brachyphyllum give no anatomical infor- mation the generic name is usually applied to fossils exhibiting only external features; it is a form-genus. The introduction of anatomical characters, based on the examination of a very small number of examples, into a general definition might seriously mislead students with regard to the affinities of species known only as impressions. On the other hand as some species of Brachy- ° phyllum exhibit anatomical features of diagnostic value the definition of the genus may be extended, in certain cases only, to include the information furnished by such examples as those de- scribed on pages 322—328. The inclusion of some species, e.g. the well-known type usually referred to Thuites, T. expansus, in Brachyphyllum is a change which may be regarded as retrograde, but an examination of specimens of that type shows the impossibility of recognising any constant verticillate disposition of the leaves such as would justify the adoption of Cupressinocladus or some other name implying 1 Hollick and Jeffrey (09) B. p. 33. 2 Seward and Bancroft (13) p. &69. 3 Stopes and Fujii (10) p. 23; Jeffrey (10%). XLVI] BRACHYPHYLLUM 317 affinity to recent Cupressineae. By slightly extending the use of Brachyphyllum we avoid the danger of giving a false impression of affinity and lighten the task of dealing with material which is of secondary botanical importance. Brachyphyllum expansum (Sternberg). In transferring this widely spread Jurassic species, founded by Sternberg on a specimen from the Stonesfield (fig. 754), Oxford- Fic. 754. Brachyphyllum expansum. Fic. 755. Brachyphyllum expansum. (Sedgwick Museum, Cambridge.) (Figured by Feistmantel as Echino- strobus expansus.) shire, as Thuites expansus1, from Thuites to Brachyphyllum the application of the latter name is extended to include Coniferous shoots in which the decussate arrangement of the leaves is more apparent than in the majority of species usually referred to 1 Sternberg (23) A. p. 38, Pl. xxxvut. figs. 1, 2. For synonymy see Seward (04) & Pp & B. p. 142. 318 CUPRESSINEAE : [cH. Brachyphyllum. Schimper} included Sternberg’s species in Echino- strobus and Saporta? adopted the designation Palacocyparis. The small amount of evidence with regard to the structure of the cones does not afford an adequate reason for retaining the generic name Thuites. Specimens from Jurassic rocks in India described by Feist- mantel? (fig. 755) as Echinostrobus expansus, superficially at least very similar to the European Thuites expansus, have recently been examined by Miss Holden*. The epidermal cells of the small decussate leaves are irregular in shape; the stomata are scattered but there is an astomatic area down the centre of the lamina. The stomata are sunk and have four accessory cells. Miss Holden points out that the epidermal features of this Indian type are different from those of Brachyphyllum macrocarpum, B. Miinsteri and B. affine® in which rows of stomata alternate with strands of sclerenchyma: this difference is legitimately used as an argument in favour of retaining the generic name Thuites rather than em- ploying Brachyphyllum. It is, however, as a rule impossible to obtain any information with regard to the cuticular features, and from the external characters of impressions of foliage-shoots we cannot draw any satisfactory line between specimens referred to Brachyphyllum and Thuites. Miss Holden’s work affords an illustration of the possibility of employing epidermal features as a means of separating shoots which in habit appear to belong to one generic type. So far as I know we have no data with regard to the epidermal structure of the European Thuites expansus and we cannot therefore say whether the Indian species are identical or not with those included in the same species from other regions, In habit Brachyphyllum expansum agrees with B. mamillare and other types as also with recent species of Thuya and Cupressus: in some examples the branchlets are crowded and in others the ramification is much more open; the small appressed leaves are broadly triangular or longer and relatively narrower than in such 1 Schimper (72) A. p. 333. 2 Saporta (84) p. 600, Pl. 209. 3 Feistmantel (762) p. 69, Pls. 1x., x. 4 Holden, R. (15%) p. 221, Pl. x1. figs. 2, 5, 6. The specimens examined were kindly sent to the Cambridge Botany School by the Director of the Indian Geo- logical Survey. 5 Hollick and Jeffrey (09) B.; Schenk (67) A. XLVI] BRACHYPHYLLUM 319 species as B. mamillare or B. crassum: the apical portion of the lamina is free and may be slightly falcate. In a few cases globular Fic. 756. - Brachyphyllum expansum?. (The original of Feistmantel’s Pachyphyllum heterophyllum. Calcutta Museum, Geol. Surv. India; nat. size.) cones occur on the foliage-shoots characterised by spirally disposed scales: in a specimen from the Stonesfield slate described in 1904 320 CUPRESSINEAE [cH. each cone-scale has a funnel-like cavity near one edge and the upper side of the cavity is radially ridged’. No seeds have been found in connexion with the cones. The male flowers are longer and narrower and consist of numerous sporophylls attached at right-angles and expanded distally into a peltate lamina. The specimen reproduced in fig. 756 is the original of Feistmantel’s Pachyphyllum heterophyllum? from Indian Jurassic beds: on the stouter axis there are spirally disposed triangular leaf-bases while on the smaller branches the leaf-lamina is preserved and appears to be thick, sub-falcate, and tetragonal. This specimen is in my opinion indistinguishable from that shown in fig. 755, which Feistmantel figures as Echinostrobus expansus*? and both agree superficially at least with European examples of Brachyphyllum exrpansum. Brachyphyllum manallare Brongniart. This specific name* has been applied to specimens from many Jurassic localities and it might well be extended to others regarded by authors as distinct species. An accurate specific determination of the numerous Brachyphyllum shoots is indeed hopeless without other characters than those afforded by impressions and casts. In habit the species resembles Athrotaxis cupressoides: the branches are given off at a fairly wide angle; the leaves are small, fleshy, and more or less triangular with a median dorsal keel and usually spirally disposed. There has been some confusion between this species and Sternberg’s Thuites expansus: the specimen from the Yorkshire coast figured by Lindley and Hutton® under the latter name, now in the Manchester Museum, is undoubtedly identical with Brongniart’s species. There is a considerable difference in the degree of freedom of the upper part of the lamina from the axis; in some specimens the leaf is almost entirely concrescent with the axis and in others the leaves are more open and attached only by the basal part of the lamina. Feistmantel figures several specimens of Brachyphyllum from Indian Jurassic localities under different names, many of which appear to be indistinguishable superficially from B. mamullare. 1 Seward (04) B. Pl. 1x. fig. 4. 2 Feistmantel (79) Pl. x1. fig. 4. 4 Brongniart (28) A. p. 109. 3 Tbid. Pl. x1. fig. 2. 5 Lindley and Hutton (35) A. Pl. chxvi. XLVI] BRACHYPHYLLUM 321 Among these are some of the shoots referred by him to Echinostrobus expansus! and others described as FE. rajmahalensis Feist. and E. rhombicus?. An examination of some of the figured specimens referred by Feistmantel to Pachyphyllum (= Pagiophyllum) peri- grinum (Lind. and Hutt.) leads me to include them at least pro- visionally in B. mamillare. The generic distinction between the form-genera Brachyphyllum and Pagiophyllum is by no means always clearly marked. Among many European examples of the Brachyphyllum mamil- lare form of Conifer, reference may. be made to the illustrations by Saporta of the French Jurassic specimens referred to Brachy- phyllum Moreauanum Brongn., B. nepos Sap. and a form with more slender branches, B. gracile®. Zeiller* records specimens of foliage-shoots with cones super- ficially resembling those of. Sequoia from Lower Jurassic beds in Madagascar which he assigns to Brachyphyllum and compares with B. nepos. Brachyphyllum spinosum Seward. A Wealden species® founded on several well preserved specimens from the coast of Sussex characterised by the possession of short, thorn-like, lateral branches clothed with fleshy leaves with a longitudinally striated lamina of the usual Brachyphyllum type. Two or three of these spinous shoots occur at the same level on the parent-axis. The stouter branches are covered with spirally dis- posed polygonal leaf-bases, while on the more slender branches the broad and short leaves assume a more or less regular decussate disposition. In leaf-form and branching-habit this species agrees closely with several other examples of the genus, but the spinous shoots are a distinctive feature. Brachyphyllum obesum Heer. This species originally described from Lower Cretaceous strata in Portugal® is represented in the Potomac formation by specimens referred by Fontaine’ to Brachyphyllum crassicaule, and there are 1 Feistmantel (76?) Pl. 1x. figs. 6—9; Pl. x. figs. 3, 4. 2 Tbid. (79) Pl. xu. figs. 2, 10> (82) Pl. m1. fig. 6. 2 Saporta (84) Pls. 165—172. - 4 Zeiller (00) p. 3. 5 Seward (95) A. p. 215, Pl. xvi. 8 Heer (81) p. 20, Pl. xvu. 7 Fontaine (89) B. p. 221, Pl. c. fig. 4; Pl. crx. figs. 1—7. s Iv 21 322 CUPRESSINEAE [CH. many specimens recorded both from Jurassic and Cretaceous rocks which differ in no important features from Heer’s type (fig. 757). An examination of branches of the recent species Cupressus Lawsoniana shows a considerable difference in the form of ramifi- cation depending on the development of nume- rous or few lateral shoots, and such differences afford an argument against the use of distinctive names such as B. obesiforme and others adopted by Saporta! for Portuguese specimens. Apart from the absence of thornlike branches this species is Fic. 757. Brachy- hardly distinguishable from B. spinosum. phyllum obesum. (After Heer; nat. BRACHYOXYLON. Hollick and Jeffrey. size. ) This generic name was proposed for pieces of wood from the Middle Cretaceous beds in Staten Island originally regarded as that of the plant which bore the foliage-shoots described from the same locality by Hollick and Jeffrey as Brachyophyllum macrocarpum, but as the result of further study it was recognised that lack of proof of any connexion between wood and shoots necessitated a new genus’. : Brachyoxylon notabile Hollick and Jeffrey. The tracheids of the xylem have separate pits usually in a single row, but they are occasionally flattened and very rarely there are two alternate rows of polygonal pits (fig. 758, . It is practically impossible to distinguish the present species so far as external features are concerned from such species as B. obesum Heer, B. crassicaule Font. and others®. Brachyphyllum eathiense Seward and Bancroft. The type-specimen of this species was originally figured by Miller as an ‘imbricated stem’ from Upper Jurassic rocks in the North of Scotland’: it consists of a branched shoot bearing in places some broadly triangular imbricate leaves with longitudinal ridges on the surface of the lamina. The pith includes some scattered thick-walled elements: no information of importance was obtained as to the structure of the vascular tissue of the stele. The short fleshy leaves have a well-protected epidermis succeeded by palisade-tissue and groups of hypodermal fibres while the rest of the mesophyll consists of parenchyma with secretory sacs and 1 For figures, see Hollick and Jeffrey (09) B. 2 Jeffrey (10°) p. 770. ® Hollick and Jeffrey (09) B. p. 37, Pl. rx. figs. 5, 6; Pl. x1. fig. 3; Pl. xiv. fig. 3. 4 Berry (05) p. 44; (06) p. 168; (115) p. 81. 5 Ibid. (123) p. 392, Pl. xxx. 6 Seward (95) A. p. 218. ? Seward and Bancroft (13) p. 869, Pl. 1. figs. 2—4. XLVI] BRACHYPHYLLUM 327 portions of leaf-traces. The most striking feature is the occurrence of reticulately pitted, isodiametric tracheids, closely resembling those in recent Araucarian leaves and in Brachyphyllum crassum. Stomata were found on some of the leaves agreeing in the possession of four accessory cells (fig. 724, A, page 216) with those described by Jeffrey? and Thompson? in B. crassum. Though comparable with Araucarian leaves in the structure and distribution of the trans- fusion-tissue and in the branching leaf-traces, there is hardly sufficient evidence to warrant any positive statement with regard to the relationship to recent genera of the American and Scottish species. Brachyphyllum vulgare (Stopes and Fujii). In their account of Upper Cretaceous plants from Hokkaido, Japan, Drs Stopes and Fujii? instituted a new genus Yezonia for some petrified shoots which they suggested should be placed in a special family of Gymnosperms. The specimens of foliage-shoots they described as Yezonia vulgaris and for a cone, which they con:. sider may belong to the vegetative branches, the generic name Yezostrobus was proposed. The slender foliage-shoots bear appressed ' leaves, apparently spirally disposed, agreeing closely with those of recent Cupressineae in their form and relation to the axis; but in the absence of impressions their surface-features cannot be clearly determined. Anatomically the shoots agree very closely with Brachyphyllum crassum: the pith contains groups of sclerous cells; the leaf-traces branch repeatedly in the base of the leaf, and trans- fusion-tissue is abundant in the mesophyll. The secondary xylem shows uniseriate separate pits on the tracheids, and the medullary rays are 1—2 cells in depth. Jeffrey* drew attention to the striking resemblance between Yezonia and Brachyphyllum crassum and fully justified his substitution of Brachyphyllum for the new genus. Dr Stopes*, while agreeing with this conclusion, points out that evidence furnished by fructifications can alone settle the question of generic identity; she states that the supposed cone attributed by Hollick and Jeffrey to Brachyphyllum differs widely from Yezostrobus which may be the cone of the Japanese species. The 1 Jeffrey (10°) p. 768, Pl. Lxv. fig. 6. 2 Thompson (12%) Pl. vz. fig. 12. 3 Stopes and Fujii (10) p. 23. 4 Jeffrey (10%). 5 Stopes (11°) 328 *CUPRESSINEAE [CH. American cone may, however, be a vegetative bud, and nothing is known as to its seed-bearing appendages. Yezostrobus has not been proved to have any connexion with the foliage-shoots of Brachy- phyllum. Disregarding the two cones, there can be no doubt as to the very close similarity between the American and Japanese shoots. TAXODITES. Unger. This generic name, adopted by Endlicher? from Unger, i is employed for fossil species believed to be nearly related to the recent genera Taxodium and Glyptostrobus. Reference is made in the chapter on Coniferous woods to supposed examples of these genera. The separation of the two recent Conifers is based on features which cannot be applied to fossil impressions and even in the case of the existing types Beissner?, following Bentham and Hooker, does not accept the Far Eastern species referred by Endlicher to Glyptostrobus as representatives of a distinct genus, but includes them in Taxodium. Heer describes some fragments of shoots from the Lower Cretaceous of Greenland as Glyptostrobus groenlandicus®, but these are of little or no value as trustworthy records. Similarly his species G. intermedius from the Patoot beds* founded on dimorphic, sterile, shoots affords no substantial evi- dence of affinity to Taxodium or Glyptostrobus. Tazxodites europaeus Brongniart. This species was first described by Brongniart® from Tertiary beds in Greece. The branches bear leaves varying in size and form, some being appressed and triangular while others are more elongate and freer from the axis: the oval or globular cones, with a maximum diameter of 15 mm.., consist of 18—20 scales agreeing in their rounded crenulate edges and radially grooved surface with those of Glypto- strobus. Brongniart states that this species occurs also in Germany, Bohemia, and at Oeningen. Heer® figures good examples of vege- tative shoots and cones as Glyptostrobus europaeus from Oeningen ; the scale-leaves are decurrent and the oval cones have semi- circular scales with 6—8 grooves (fig. 759). This is described as one of the commonest fossils in the Swiss deposits and, as Heer 1 Endlicher (47) p. 278. 2 Beissner (91) p. 148. 3 Heer (75) ii. Pls. XVI, XX., XXII. 4 Ibid. (75) ii. Pl. uo. 5 Brongniart (33). 6 Heer (55) A. Pls. xIx., xx. XLVI] TAXODITES 329 says, it bears a striking resemblance to the existing Chinese type Glyptostrobus heterophylius. Unger! describes well preserved specimens from Greece and the species is recorded, on the evidence of cones as well as sterile shoots, from Leoben? and other localities in Styria’, also from Miocene beds in Bohemia‘. Laurent® figures examples from Aquitanian beds in the Puy-de-Déme and Saporta and Marion® refer to Glyptostrobus europaeus fragments of sterile branches and an imperfect cone from Pliocene beds in the Province of Ain and mention the occurrence of the same type in Pliocene strata in the valley of the Arno. Vegetative shoots are recorded from yg. 759, Taxo- Tertiary beds in Bosnia’ and Nathorst® found the —dites_ europaeus. species in Arctic Ellesmere Land. The sterile frag- (After. Heers nat. size.) ments figured by Goeppert and Menge® from the : Oligocene beds on the Baltic coast, though possibly correctly determined, afford no proof of affinity to the genus Glypto- strobus. Some very good specimens from Eocene beds at Reading are described by Gardner! as examples of this species but the cones are immature and do not furnish convincing evidence of close relationship to the recent genus. The same remark applies to specimens figured by this author from Bournemouth. Specimens from the latter locality, characterised by their long slender branches with spirally disposed leaves having long decurrent bases and pro- jecting apices, are referred to a distinct species Taxodium eocaenicum and compared with the Floridan Conifer Taxodium distichum var. imbricataria Mett. Gardner points out with reason that specimens described by Heer! from Miocene beds in Greenland and Alaska as Glyptostrobus europaeus and from rocks of the same age in 1 Unger (67) Pl. 1. 2 Ettingshausen (88) Pl. 1. 3 bid. (90) Pl. 1. 4 Ibid. (672) Pls. x., x1.; Velenovsky (81); Unger (52) Pl. xxxtv. 5 Laurent (12) Pls. v., vz. 6 Saporta and Marion (76) Pls. xx1ir., XXXVII. 7 Engelhardt (12) Pls. xxxu., XXXVI., XXXVI. 8 Nathorst (15). ® Goeppert and Menge (83) A. PL. xv1. 10 Gardner (86) Pls. m1., IV., XXIV. 11 Heer (68) i. Pls. 1., xtv.; (71) iii. Pls. 1, 11.; (77) i. Pls. X1., XU1., XXVI.; (78) Pls. rx., xmt.; (83) Pls. uxx., LXxv. 330 CUPRESSINEAE [CH. Spitzbergen as G. Ungeri may be fragments of Conifers more closely allied to Sequoza. The fossils originally referred by Lesquereux! to Glyptostrobus gracillimus from the Dakota group were afterwards transferred by him to Frenelites Reichti, at a later date removed by Newberry to Sequoia and finally described by Jeffrey? as Geinitzia gracillima. Neglecting records based on sterile specimens only it is clear that Conifers closely allied to Taxodium and especially to Glypto- strobus heterophyllus of China were abundant in the Tertiary floras of Europe. : Taxodites miocenicum (= Taxodium distichum miocenicum Heer). Numerous specimens of branches and some male and female flowers have been described by Heer? from Miocene beds in Green- land, Spitzbergen, Grinnell Land, Northern Siberia, and Sachalin Island as Taxodium distichum miocenicum. Though in certain cases the material is too imperfect to determine with accuracy, some of the fossils bear a striking resemblance to Taxodium distichum both as regards vegetative features and cones. In this species Heer includes specimens originally named by Sternberg Phyllites dubius and afterwards transferred to Tazxodium. The shoots bear di- stichous, linear leaves, reaching a length of 2cm. and 2—3 mm. broad. The absence of a decurrent base is spoken of as a character distinguishing Taxodium {from Sequoia Langsdorfix, Specimens from Grinnell Land now in the Dublin Museum described by Heer as Taxodium distichum miocenicum bear leaves contracted at the base but not decurrent. A specimen from Grinnell Land said to be a male inflorescence‘ consists of an axis bearing a few oval buds 3 mm. long which may possibly be groups of microsporophylls, but the figures are far from convincing. Nathorst® speaks of the occurrence of foliage-shoots, flowers, and seeds in the Tertiary Taxodium shales of Spitzbergen. Ettingshausen® figures shoots and flowers from Bilin as Taxodiwm dubiwm which are in all probability closely allied to the recent species. Goeppert and 1 Newberry and Hollick (95) Pl. 1x. 2 Jeffrey (11). 3 Heer (68) i. Pls. m., x1, XLV.; (71) Pls. 1., Iv., etc.; (77) Pls. x1m., xxv.; (78) Pls. vit, Ix.; (83) Pls. Lxx., ete. 4 Heer (68) i. Pl. u.; Schimper and Schenk (90) A. p. 294, fig. 203. 5 Nathorst (11°) p. 223. ® Kttingshausen (672) Pl. x. XLVI] CUNNINGHAMIOSTROBUS 331 Menge! refer some detached leaves from the Baltic amber to Taxodium distichum and a cone is referred to Taxodites Beckianus but without any clear evidence of affinity to the recent genus. Lingelsheim? records some wood from Tertiary rocks in Silesia which he refers to Taxodium- and at the same locality he found masses of pollen some of which he assigns to that genus. Engel- hardt and Kinkelin® describe cones of the Taxodium type as Taxodium distichum var. pliocenicum from the Frankfurt basin. Heer’s species is also recorded from several other Tertiary | floras* and, despite the fragmentary nature of the material, there is good reason for regarding the evidence as an indication of the widespread occurrence of a Conifer in Tertiary Europe closely related to the Swamp Cypress of North ‘America. The species is stated to be abundant in Pleistocene beds in North America from New Jersey to Maryland, Virginia and elsewhere on the eastern side of the continent®; it is represented by deciduous twigs, cones, seeds, stumps, and ISnees®: its occurrence points to the existence of Cypress swamps over a wide area, also to the migration of the existing species towards the south. CUNNINGHAMIOSTROBUS. Stopes and Fujii. Cunninghamiostrobus yubariensis Stopes and Fujii. This genus is founded on a single detached cone from Upper Cretaceous rocks at Hokkaido in Japan’ which in size and form agrees with cones of Cunninghamia, and the anatomical features of the cone-scales support this comparison. The cone, 2 x 3 cm., is intermediate in size between those of Cunninghamia sinensis (cf. fig. 684, K) and | C. Konishit; the scales being more like those of C. sinensis; they are 9—10 mm. across and characterised by the presence of a median pad of tissue projecting slightly from the upper surface presumably close to the attachment of the ovules, but no ovules or seeds were found and the open habit of the cone indicates that the seeds had been shed at the time of fossilisation: three pro- 1 Goeppert and Menge (83) A. Pl. xvi. figs. 227—229. 2 Lingelsheim (08) p. 34. 3 Engelhardt and Kinkelin (08) PJ. xx1m. figs. 19—21. 4 Squinabol (92) Pl. xv1.; Berry (09) p. 22, fig. 1. 5 Berry (072); (092); (124); (15). 8 Ibid. (097), figs. 1, 2. 7 Stopes and Fujii (10) p. 45, Pl. v. figs. 27—34. 332 CUPRESSINEAE (cH. tuberances on a scale at the apex of the cone may represent aborted ovules though the nature of these is problematical. Near the base of a cone-scale there is a single transversely elongated vascular bundle which subdivides higher in the scale into a series of normally orientated vascular strands, and in one scale a much smaller bundle, probably an ovular trace, was found immediately above the main strand. The parenchymatous ground-tissue contains a few sclerous elements and several resin-canals, the larger ducts forming a series across the scale, and near the base a single large canal occurs below the broad vascular bundle as in Cunninghamia. Groups of trans- fusion-tracheids occur between the vascular strands. The preservation of the tissues of the cone-axis is not good enough to throw any light on the question of affinity and it.is from the morphology of the seedless scales that any conclusions must be drawn. The cone-scales show no indication of a division into the two organs characteristic of the Abietineae nor is there any evidence of a ligular outgrowth like that of an Araucarian scale. The resemblances in both form and anatomical characters to the sporophylls of Cunninghamia exhibited by the fossil cone appear to be such as to justify the employment of a generic name implying close relationship. Cunninghamites Presl. The employment of this name by many authors for sterile branches (e.g. fig. 805) superficially resembling foliage-shoots of Cunninghamia sinensis suggests an affinity which is not supported by any substantial evidence and while in some cases the fossils may belong to plants closely allied to the recent genus, there is no definite justification for assuming such alliance. The Lower Cretaceous species of Cunninghamites and similar forms are there- fore relegated to Halle’s genus Elatocladus. MORICONIA. Debey and Ettingshausen. This generic name was applied by Debey and Ettingshausen? to some obscure impressions from the Cretaceous beds of Aix-la- Chapelle which they described as portions of a plant ‘incertae sedis’; the specimens superficially resemble the pinnae of a fern with broadly linear pinnules, but the occurrence of curved lines 1 Debey and Ettingshausen (55) B. p. 239, Pl. vi. figs. 23—27. XLVI] MORICONIA 333 at right-angles to the long axis of the pinnules (fig. 760) suggested that some at least of the vein-like markings might be the boun- daries of small scale-like leaves similar to those of Libocedrus and other Conifers. Saporta?, in his reference to the genus in an account of the Sézanne flora, assigns Moriconia to the Cupressineae, a ' determination in accordance with the habit of the foliage-shoots, though in the absence of reproductive organs it is impossible to fix its position more precisely. The characteristic features are the pinnate branching, the flattened form of the branches, and the geometrically regular decussate short and broad leaves. The genus is recorded only from Lower and Middle Cretaceous rocks. Moriconia cyclotozon Debey and Kttingshausen. This, the type-species, is recorded from Cretaceous rocks at Aix-la-Chapelle, from the West coast of Greenland and the Atlantic coastal plain. Heer? figured an imperfectly preserved specimen from Disco as Pecopteris kudlistensis in which an indication is given of the occurrence of the actual leaves, but some years later? he de- scribed well preserved examples as Moricomia cyclotoxon, and, as the result of an inspection of drawings supplied by Debey, identified them ' with the type-species. The same type is re- corded from the Amboy clays (fig. 760)*, Staten Island® and Block Island. As Hollick points out, a large impression included by Heer in Fie.760. Moriconiacy- Moriconia should rather be referred to Brachy- ¢lotoxon. (After New- phyllum: in the arrangement of the leaves and in aa mone the form of the short and blunt lateral foliage- shoots Moriconia agrees closely with some examples of Brachy- phyllum crassum® in which the leaves appear to be regularly de- cussate. The leaves of Moriconia are wholly appressed and the upper edge of the lamina is rounded and almost truncate; a médian line, possibly due to the presence of a dorsal keel, runs down the middle of the exposed broad surface of the shoots. 1 Saporta (68) A. p. 301. 2 Heer (75) ii. p. 97, Pl. xxvi. fig. 18. 3 Ibid, (82) i. p. 49, Pl. xxxi11.; (83) Pls. Lu., Liv. 4 Newberry and Hollick (95) p. 55, Pl. x. 5 Hollick (06) Pl. m.; Berry (03) Pl. xiv. ® Berry (06) Pl. 1x. fig. 1. 334 CUPRESSINEAE _ [cH. CRYPTOMERITES. Bunbury. The species Cryptomerites divaricatus, for which Bunbury? pro- posed this generic name, is more probably Araucarian than a type allied to Cryptomeria: the choice of the term was suggested solely by vegetative characters and Bunbury recognised that these agreed with species of Araucaria as well as with Cryptomeria. The designa- tion Cryptomerites should be restricted to fossils which there is good reason for believing to be allied to the recent genus. Cryptomerites du Noyeri (Baily). Baily? figured a sterile piece of foliage-shoot from Eocene leaf- beds in County Antrim as Sequoia du Noyeri which Gardner? sub- sequently stated to be identical with specimens obtained from the same locality bearing cones similar to those of Cryptomeria. Gardner described the Irish specimens and others from Mull as Cryplomeria Sternbergii (Goepp.), the specific name being adopted because he considered some examples figured by Ettingshausen4 as Araucarites Sternbergit Goepp. to be identical with the Ivish fossils, though most of the specimens described by authors, including the author of the species, as Araucarites Sternbergit are believed to be identical with Araucarites Goepperti Sternb. In these circumstances it seems desirable to employ the specific name du Noyeri used by Baily. Gardner’s material consists of foliage-shoots agreeing in their spirally disposed leaves, 4—7 mm. in length and falcate in form, with branches of some species of Araucaria, Dacrydium, and Cryptomeria japonica. The occurrence of associated cones, in some cases attached to the vegetative shoots, affords fairly good evidence in support of comparison with Crypiomeria. The sub-globose cones, 15—20 mm. in diameter, consist of a comparatively small number of scales attached by a narrow base and gradually widening towards the distal edge which is deeply fringed. The general appearance of the cones, especially those from Glenarm in Antrim, is similar to those of Cryptomeria (cf. fig. 684, M) and taking into account the characters of the sterile branches the assumption of affinity to that genus appears to be well founded, though actual proof of close relationship is lacking. Gardner includes in Cryptomeria Sternbergit 1 Bunbury (51) A. p. 190, Pl. xm. fig. 4; Seward (00) B. p. 287. 2 Baily (69) Pl. xv. fig. 4. 3 Gardner (86) p. 85, Pls. X., XX., XXI. 4 Ettingshausen (55) Pl. v. XLVI] CRYPTOMERIOPSIS 335 specimens figured by Ettingshausen from Monte Promina as Arau- carites Sternbergui and some of the impressions from Greenland re- ferred by Heer! to Sequoia Sternbergi. The Miocene fragments figured by Heer afford no evidence of affinity other than that of leaf-form, and the use of the term Cryptomerites should therefore be avoided. If the Eocene plant is correctly regarded as closely allied to Cryptomeria it supplies another striking illustration of the change in the geographical distribution of Conifers since the early part of the Tertiary period. CRYPTOMERIOPSIS. Stopes and Fujii. Cryptomertopsis antiqua Stopes and Fujii. The name Crypto- meriopsis? was proposed for some petrified twigs from Upper Cretaceous beds in Japan resembling in habit and structural features the recent Conifer Cryptomeria japonica. The xylem of the axis consists of tracheids with uniseriate separate, circular, bordered pits; there are no resin-canals and no xylem-parenchyma,; the presence of the latter tissue is recorded by Suzuki in a second Japanese species C. mesozoica®. The medullary rays are usually one-cell deep in the type-species and there are a few (1—3 in C. mesozoica) oval pits in the field. In C. antigua the phloem is said to consist of soft tissue only, but fibres occur in C. mesozoica. An undivided leaf-trace supplies each leaf. The four-sided leaves are characterised by the presence of three canals, a large central canal below the vascular bundle and two lateral ducts; the vascular bundle is accompanied by well-developed lateral groups of trans- fusion-tracheids. The leaves of C’. mesozoica differ in a few details from those of the type-species. Prof. Jeffrey* maintains that _ Cryptomeriopsis is generically identical with Geinitzia as described by Hollick and Jeffrey from Staten Island and should be included in the Araucarineae. Dr Stopes® adheres to the view that the Japanese fossils are closely allied to Cryptomeria and afford no evidence of affinity to Araucaria: the structure of the xylem shows no Araucarian features in the pitting of the tracheids and, while accurate determination of systematic position must depend upon the evidence of reproductive shoots, the evidence of the vegetative shoots favours comparison with Cryptomeria rather than with Araucaria. 1 Heer (75) iii. Pl. 11. figs. 1—4. 2 Stopes and Fujii (10) p. 52, Pl. 1. fig. 11; Pl. vi. figs. 35—41. 3 Suzuki (10) p. 185. 4 Jeffrey (10%) p. 771. 5 Stopes (11%). CHAPTER XLVII. CALLITRINEAE. It has already been pointed out that there is good reason for treating the three existing genera Callitris, Widdringtonia, and Actinostrobus as members of a distinct family. The genus Tetra- clinis, as Saxton! has shown, while agreeing with the Callitrineae in certain features, exhibits a closer resemblance in its gametophyte to the Cupressineae and is regarded as a type connecting the two families Cupressineae and Callitrineae. So far as external characters are concerned, and these are the features from which the palaeo- botanist is compelled to draw such conclusions as he can, Tetrachinis falls into line with the Callitrineae. The discontinuous distribution of the recent species of these four genera suggests antiquity and a former more extended range. Palaeobotanical literature contains numerous records of Widdringtonia, Calittris, Frenela or Frenelites based in many cases on sterile shoots and sometimes on cones and seeds more or less closely resembling those of recent forms. The generic name Frenela has now been discarded in favour of Callitris: it was proposed by Miquel in 1826 to avoid confusion between Callitvis and Calythriz, the latter being the name of a Myrtaceous genus. An inspection of the published figures of supposed fossil representatives of the Callitrineae shows that the name Widdring- tonia or Widdringtonites has sometimes been applied to fertile shoots with cones differing in the number of the valves from those of recent species and more closely resembling the cones of Calliris, Tetraclinis, or Actinostrobus. Moreover the number of valves in recent cones, though usually constant, is not invariably the same and in imperfectly preserved specimens it is often difficult to differentiate satisfactorily between the four genera. In the case of many sterile shoots preserved as impressions it is practically im- possible to distinguish clearly between those of the Callitrineae and 1 Saxton (132); (134). CH. XLVIT] CALLITRITES. 337 slender branches of Juniperus, Thuya, and other Cupressineae. Even when cones are preserved there is some danger of confusion with fruits of certain Dicotyledons, e.g. Lagerstroemia macrocarpa (Lythraceae). In view, of the difficulties of precise determination the most convenient course is to adopt the generic name Callitrites in a comprehensive sense without as a rule attempting to assign the fossil to one of the existing genera of the Callitrineae. CALLITRITES. Endlicher. Endlicher! employed the generic names Widdringtonites, Calli-_ trites, Frenelites, and Actinostrobites, but the material seldom justifies such discrimination. The name Actinostrobites was pro- posed in the first instance for some cones described by Bowerbank? from the London Clay as Cupressites globosus and C. elongatus but Gardner®, who examined the original specimens, is sceptical as to their connexion with the Callitrineae. Ettingshausen* described a small cone from Miocene beds in Carinthia as Actinostrobus mio- cenica on the ground that there appear to be traces of scales at the base of each of the six small linear valves of the cone. The specimen is too imperfect to be determined with any accuracy. It is im- possible to express any considered opinion with regard to the validity of the numerous Tertiary records of Callitris and Widdring- tonia without access to the actual material, though many of the illustrations lend strong support to the identification of the speci- mens as examples of some Callitrineous type. Despite the imper- fection of many of the records there can be no doubt as to the former occurrence of representatives of the Callitrineae in Tertiary floras in Europe. The pinnately branched sterile shoots referred to Weddring- lonites keuperianus Heer® from the Trias of Switzerland and Germany bear a close resemblance to some forms of Walchia and there is no sound reason for assigning the species to the Calli- trineae. Saporta® described fragments of branches from the Lower Lias of France as examples of Heer’s type, but in this case also no 1 Endlicher (47) p. 271. 2 Bowerhank (40) p. 52, PI. x. 3 Gardner (86) p. 20. 4 Ettingshausen (72) p. 164, PI. 11. figs. 9—-12. 5 Heer (65) A. p. 52, fig. 31; Schiitze (01) Pl. x.; Schenk in Schimper and Schenk (90) A. p. 311. 6 Saporta (84) Pl. 201, fig. 1. s IV 22 338 ; CALLITRINEAE [CH. cones were found. Similarly Widdringtonites gracilis Sap. and W. creyensis Sap. from the Corallian and Kimeridgian of France? respectively are founded solely on sterile shoots. The specimen figured by Eichwald? from Jurassic rocks on the southern border of the Caspian sea as Widdringtonites denticulatus has the habit of an Araucaria and the supposed cone, which may be some foreign body not actually attached, affords no evidence of affinity to the Callitrineae. Zeiller? describes a small fragment from Liassic beds in the Commune of Cherveux bearing small rhomboidal decussate leaves similar to Widdringtonites liassinus (Kurr) as figured by Salfeld and to W. keuperianus, but the material affords no definite indication of relationship to the Callitrineae. Cailitrites Reichit (Ettingshausen). This species, recorded from several Cretaceous localities in the Eastern United States and elsewhere, is in many cases represented only by slender sterile shoots and its position among the Coniferae is by no means clearly established. It was founded by Ettings- hausen* as Frenelites Reichvi on some branched shoots from Cre- taceous rocks in Saxony and afterwards described by Heer® from the Patoot beds of West Greenland under the generic name Widdringtonites though without satisfactory evidence in support of relationship to Widdringtonia. This species is one of the com- monest Conifers in the Amboy clays of New Jersey, but no cones are figured by Newberry® in his monograph except two small examples which it is suggested may be immature microstrobili. Velenovsky’ figures sterile branches from the Perucer beds of Bohemia and an ovate cone, 13 mm. long, with four valves, which resembles a small cone of Actinostrobus and those described by Berry as Widdringtonites subtilis. Some of the twigs bear terminal elliptical bodies regarded as male flowers. The leaves of this species are usually spiral and, with the exception of the apex, closely appressed. Callitrites Reichii is also recorded by Krasser§ from the 1 Saporta (84) Pls. 201, 202. 2 Kichwald (68) p. 43, Pl. rv. fig. 9. 3 Zeiller (11) Pl. 1. fig. 6. 4 Kttingshausen (67) p. 246, Pl. 1. fig. 10. 5 Heer (82) i. p. 13, Pl. iru. figs. 4, 5. 5 Newberry and Hollick (95) Pl. vit. 7 Velenovsky (85) B. p. 27, Pls. vut., x.; (87) figs. 14—16. 8 Krasser (96) B. p. 126, Pls xiv., xv. XLVII] CALLITRITES © ; 339 Cenomanian of Moravia where it is represented by both sterile and fertile shoots; the cones are quadrivalvate. It occurs in the Middle Cretaceous of Staten Island though without any cones: Hollick and Jeffrey! regard the shoots referred by them to Widdringtonia Reich as Araucarian on the ground that the bordered pits on the tracheal walls are usually contiguous. There is, however, no sub- stantial reason for assigning these vegetative organs to the Arau- carineae though the structure of the wood shows an Araucarian tendency. Berry? records the species from the Cenomanian Raritan formation of New Jersey and he expresses the opinion that the species is closely allied to some Potomac specimens described by Fontaine? as Taxodium ramosum, but in the absence of cones a definite determination of affinity is hardly possible. Callitrites subtilis (Heer). Founded by Heer‘ on slender twigs bearing spirally disposed, appressed, leaves from the Cretaceous beds of Atanekerdluk in .Greenland and described by Newberry® from the Amboy clays. - Hollick® and Berry’ have also recorded the species from Cretaceous strata in different parts of the Eastern United States and the latter author figures examples from Upper Cretaceous beds in South Carolina’.+ The epidermal cells are regularly rectangular and the stomata are surrounded by 5—6 accessory cells. Berry figures conical cones, 7—9 mm. long by 4—5 mm. in diameter, composed of four thick scales differing somewhat in shape from the cones of recent species. The sterile shoots of this species bear a close resem- blance to C. Reichti and the two species have often been confused; also to Cyparissidium minimum as figured by Velenovsky®, Juniperus macilenta Heer! and Widdringtonites fascicularis Holl. 1 Hollick and Jeffrey (09) B. p. 29, Pls. v., vim., xx.; Hollick (06) p. 44, PL. 1v. figs. 6—8. 2 Berry (11%) p. 87, Pl. vit. 3 Fontaine (89) B. p. 251, Pls. cxxui1., CXxIVv., etc.; Berry (11*) p. 302. 4 Heer (74) B. Pl. xxvim. fig. 1. 5 Newberry and Hollick (95) p. 57, Pl. x. figs. 2—4. 5 Hollick (06) p. 45, Pl. rv. figs. 2—5. 7 Berry (12). 8 Ibid. (14) p. 25, Pl. x1. figs. 14—17. 9 Velenovsky (85) B. Pl. rx. figs. 6, 7; Pl. x. fig. 4. 10 Heer (75) ii. Pl. xxvuit. fig. le. 4 Hollick (06) Pl. rv. fig. 1. 22—2 340 CALLITRINEAE [CH. Callitrites curta (Bowerbank). Bowerbank referred several pyritised cones from the London Clay of the Island of Sheppey to Cupres- sites and some of them he compared with species of Callitris. Gardner}, as the result of an examination of Bowerbank’s type- specimens, reduced the number of species and adopted the name Callitris. The speci- mens, in the British Museum, assigned to Callitrites curta are conical cones composed of 4, 5, or rarely 6 thick and woody valves which are sometimes unequal in size: the largest has a diameter of 2cm. Fig. 761, nn A shows a cone of five valves and B, C, are a Ae: TRDIRCREES ‘OH : : : , cone of five valves. two views of a section of a cone consisting _B, C, two views of a sec- of four valves. A similar type described tion of afour-valvedcone. by Gardner as Callitris Ettingshauseni?, aaa ace sae oike f rom specimens described also from Sheppey, is represented by glo- by Gardner; nat. size.) bular cones 12—15 mm. in diameter and composed of 6—8 scales. These two species are probably correctly referred to the Callitrineae though the pyritised cones are the only portions of the plant preserved in the Sheppey clay. Gardner states that Ettingshausen® is incorrect in recording Callitrites curta from the Isle of Wight. Callitrites Brongniarti (Endlicher). This species, first described by Brongniart* as Hquisetum brachyodon from the Paris Basin, is recorded from many European localities, in some cases represented only by sterile shoots but frequently also by cones and small winged seeds. Some well pre- served specimens are figured by Unger® from the Tyrol (fig. 762, A, A’) under the name Thuyites callitrina characterised by regularly whorled leaves, apparently four at each node, with a relatively long and narrow appressed lamina and a small free apex and by - valvate cones. Unger® subsequently described good specimens as 1 Gardner (86) p. 21, Pl. 1x. figs. 7, 21. ? Ibid. Pl. rx. figs. 1—6. 3 Ettingshausen (79) p. 392; (80) p. 231. 4 Brongniart (22) A. p. 329, Pl. v. fig. 3; Endlicher (47) p. 274. 5 Unger (47) p. 22, Pls. vi., viz. 8 Ibid. (67) p. 42, PL. 1. figs. 1, 2. XLVII] CALLITRITES 341 Callitris Brongniartt from Miocene beds in Euboea, but Saporta! considers these impressions to be more closely allied to Widdring- tonia and renames them Widdringtonia kumensis. Good examples of quadrivalvate cones (fig. 762, B) are figured by Saporta? from the Eocene beds of Aix and Armissan in Provence, showing in some cases two outer broader valves and two internal laterally com- pressed valves. Httingshausen® states that the species is very abundant at Haring in the Tyrol: that author describes some ves SSS =D RAL vas Stes SY eS Fig. 762. A, A’, B, Callitrites Brongniarti. C, Callitrites helvetica. D, Callitrites europaea. (A, A’, after Unger; B, after Saporta; C, after Heer; D, after Engelhardt and Kinkelin.) sterile shoots from Eocene beds in New South Wales as Callatris prisca* which he compares with C. Brongniarti. Well preserved shoots are described by Watelet® from the Paris Basin. Engel- hardt® records the species from Oligocene beds in Bohemia but on the inadequate evidence of a winged seed; it is recorded also by Engelhardt and Kinkelin’ from the Pliocene beds of the Frankfurt 1 Saporta (68) p. 316. 2 Ibid. (62) p. 209, Pl. m. fig. 6; Pl. mm. fig. 1; (65%) p. 39, Pl. 1. fig. 6. 3 Ettingshausen (55) p. 34, Pl. v. figs. 7—35.. 4 Ibid. (86) p. 95, P]. vim. figs. 3, 4. 5 Watelet (66) A. Pl. xxx. ® Engelhardt (85) p. 314, Pl. vat. fig. 32. 7 Engelhardt and Kinkelin (08) Pl. xxmu. fig. 5. 342 ‘ CALLITRINEAE [CH. district though on slender grounds. It may be, as Masters suggested, that some at least of the Tertiary specimens included in Callitrites Brongniarti are more closely allied to the existing genus Tetraclinis than to Callitris. ; Among other species that may be included in Callitrites are C. brachyphylla (Sap.) and C. antigua (Sap.) from Provencel, repre- sented by shoots with spiral, sub-opposite or opposite appressed scale-leaves and by globular cones with four valves. As Solms- Laubach says®, the cones agree closely with those of Widdringtonia, though it would be difficult to decide between that genus and Tetractinas. Some good specimens are figured by Heer? from the Oeningen beds as Widdringtonia helvetica, now transferred to Callitrites, con- sisting of branched filiform foliage-shoots with small appressed leaves and cones with four valves (fig. 762, C). Fragments of branches with small appressed leaves in opposite pairs from the Oligocene amber beds of the Baltic coast are de- scribed by Goeppert and Menge? as three species of Widdringtonites, and in one case, W. legitamus, the species is founded on a cone 6 mm. long and 2-5 mm. broad which is not above suspicion as a record of a Callitrineous strobilus. Frenela europaea and F. Ewal- dana described by Ludwig® from Tertiary beds near Frankfurt are founded on unconvincing specimens. Engelhardt and Kinkelin® describe pyramidal cones with 5—6 valves 1—1-5 cm. long (fig. 762, D), which they refer to Frenelites europaeus, from the Upper Phocene beds of the Lower Main valley. Many other similar instances might be quoted, but on the other hand there is ample evidence of the presence in the earlier Tertiary floras in Europe of Conifers agreeing both in vegetative and reproductive shoots with existing species now confined to Africa and Australia. FRENELOPSIS. Schenk. Schenk? instituted this generic name for specimens originally described by Ettingshausen® from Wealden beds in Silesia as 1 Saporta (62) Pl. 1. fig. 7; (62?) Pl. mn. fig. 3; (652) Pl. 1. fig. 4; (73) Pl. m1. fig. 1. 2 Solms-Laubach (91) A. p. 60. 3 Heer (55) A. p. 48, Pl. xvi. figs. 2—18. 4 Goeppert and Menge (83) A. p. 39. 5 Ludwig (59) A. pp. 69, 136. 6 Engelhardt and Kinkelin (08) PL. xxim. fig. 5. * Schenk (71) p. 13, Pl. 1. 8 Ettingshausen (52) p. 26, Pl. 1. figs. 6, 7. XLVII] FRENELOPSIS 343 Thuites Hoheneggeri on the ground that the external features of ° the vegetative shoots indicate an affinity to the recent genus Frenela (= Callitris) rather than to Thuya or Cupressus. The: resemblance to Callitris was recognised by Ettingshausen. The most striking features of Frenelopsis are the comparatively long internodes of the jointed stems and branches (fig. 763, A), the occurrence of appressed leaves in opposite pairs or four in a verticil, - concrescent with the whole internodal surface and projecting slightly above each nodal line as small broadly triangular scales, . the presence of longitudinal lines of small dots on the internodal regions due to rows of stomata characterised by 4—5 accessory cells surrounding the depressed guard-cells! (fig. 763, D, E). The smaller branches closely resemble those of species of Cupressineae (fig. 763, C) or Callitrineae in leaf-form and branching, but older branches from which the leaves have partially or wholly disappeared often differ considerably from the younger foliage-shoots and by themselves afford little or no indication of their true nature. Further details are given in the description of representative species. Frenelopsis is characteristic of Wealden or higher horizons in the Lower Cretaceous series; it occurs in Silesia, Bohemia, Portugal and the South of France and in some North American localities, particularly in the Potomac formation. Heer? records the species from Lower Cretaceous rocks in West Greenland but some of the original specimens which I had an opportunity of examining in the Stockholm Museum afforded no satisfactory. evidence of their systematic position. Though assigned by Heer to the Gnetales, Frenelopsis is usually regarded as a Conifer agreeing with Callitris more closely than with any other existing genus. In their descrip- tion of some fossil shoots referred by Newberry® to Frenelopsis gracilis Hollick and Jeffrey, who institute a new genus Raritania for this species, state that they have reason to believe that some American specimens correctly assigned to Frenelopsis are examples of Gnetalean plants. Nothing is known of any reproductive organs, but such information as we have with regard to the habit of the vegetative shoots and the structure of the stomata would seem to 1 Zeiller (82) A. p. 231, Pl. x1.; Thompson (12%) Pl. v. 2 Heer (75) ii. p. 73, Pl. xvut. ‘figs, 5—8; (82) i. p. 7, Pl. a. fige. 1—3. 3 Newberry and Hollick (95) p. 59, Pl. x11. figs. 1—3. 4 Hollick and Jeffrey (09) B. p. 26. 344 CALLITRINEAE [cH. ‘ be in favour of including this Lower Cretaceous genus among the Coniferales and regarding it as probably allied to the Callitrineae. ‘But the data are insufficient to form the basis of any definite state- ment as to the position of the genus. | ieee Hoheneggeri pEchingphimseny: Frenelopsis occidentalis Heert. Though the specimens referred to these two species may be correctly separated their close agreement in habit points to a single type so far at least as concerns the characters as a whole. The specimen represented in fig. 763, A, B, originally described by Zeiller from the province of Gard, illustrates the method of branch- ing and the form of the leaves borne in whorls of four. Schenk, in his account of Lower Cretaceous material of F. Hoheneggert from Wernsdorf in the Carpathians, states that the leaves are in decussate pairs or sometimes in verticils of four. The epidermal cells have straight walls and a thick cuticle; the stomata form longitudinal rows on the internodes and are characterised by the presence of 4—5 accessory cells overarching the stomatal depression?. Specimens described from Bohemia by Velenovsky? as F. bohemica resemble F. H oheneggeri both in habit and in the structure of the stomata. Frenelopsis ramosissima Fontaine. This species is represented by numerous well preserved speci- mens in the Potomac formation?: some of the stems have a diameter of 5 cm. and lateral branches are given off in whorls of 3-—5; there are three leaves at each node with broadly triangular apices and concrescent decurrent bases as in Ff’. Hohenegger? (fig. 763, C). The stomata are arranged in longitudinal rows and agree in the posses- sion of a rosette of accessory cells (fig. 763, D) with F. Hoheneggeri: several of the epidermal cells are provided with short spinous pro- cesses®, This species is represented by specimens showing clearly the cupressoid habit of the smaller foliage-shoots (fig. 763, C). 1 Heer (81) p. 21, Pl. x1 figs. 3—7; Saporta (94) B. pp. 139, 199, 214; Pls. XXXVI, XXXVI. 2 Thompson (123), 3 Velenovsky (88) figs. 1—3, 10. 4 Fontaine (89) B. p. 215, Pls. 95—101; Berry (11) p. 422, Pls. LxXx1., LXXIL. 5 Berry (10?). XLVIT] FRENELOPSIS 345 In another Potomac species, F. parceramosa Font.1, there appears to be a single leaf at each node: this form resembles some specimens from Lower Cretaceous rocks in Mexico which Nathorst? made the type of a new genus Pseudofrenelopsis, but the features Fic. 763. A, B, Frenelopsis Hoheneggeri. C, D, F. ramosissima. 3, P. occiden- talis. (A, B, after Zeiller; C, D, after Berry; E, after Thompson.) are hardly sufficiently well exhibited to throw much light on the nature of the shoots. It is possible that some Wealden branches from English strata described as Becklesia anomala Sew.? may belong to some species of Frenelopsis, but the absence of any leaves or nodal marks precludes their inclusion in this genus. 1 Fontaine (89) B. p. 218, Pls. cx1., CXII., CLXVII. 2 Nathorst (93) p. 52, figs. 6—9. 3 Seward (95) A. p. 179, Pl. xiv. 346 SEQUOIINEAE [cH. Further light will no doubt be thrown on the nature of Frene- lopsis when the results of the investigations of Hollick and Jeffrey are published: it may be that Heer was correct in his attribution of the Portuguese specimens to the Gnetales though in the habit of the branching, especially in F. ramosissima, and in the structure of the stomata there is a closer resemblance to recent Callitrineae than to any other plants. The genus ranges from Wealden to Cenomanian rocks. SEQUOIINEAE. In view of the restricted range of the two surviving species of Sequoia and the peculiarities of the genus, to which expression is given by the institution of the family-name Sequoiineae!, the question of geological antiquity and past distribution assumes a special interest. Reference has already been made to fossil wood presenting features now found in Sequoia, but it is very doubtful if the anatomical characters of the recent species are sufficiently well defined to enable us to discriminatt between the wood of Sequoia and certain other Conifers. Many of the impressions of vegetative shoots and cones described as Sequoia from Jurassic and especially Lower Cretaceous strata do not bear a close scrutiny. The widely spread species often referred to as Sequoia Reichen- bachit affords no real evidence of affinity to the recent genus and the same remark applies to specimens included in Heer’s genus Sphenolepidium and compared by authors with Sequoia. Some of the imperfectly preserved Jurassic cones agreeing superficially with those of Seguova may well belong to species of Sequoiineae. Though in the majority of instances Jurassic and Cretaceous records do not prove the former presence of Sequova or a closely allied type, some of them afford justification for the belief that the American trees are survivals from at least the later floras of the Mesozoic era. On the other hand Tertiary strata in many parts of the world supply clear evidence of the wide distribution of Sequoia or some nearly related Conifers in Europe and elsewhere. The inference suggested is that the recent species survive in California because of the greater possibilities of migration towards the more 1 See page 151. 2 Mr E. W. Berry (16) has recently published a sketch-map illustrating the world- wide distribution of fossils referred to Sequoia. ra XLVIT] SEQUOIINEAE 347 genial south on the American continent than in Europe where the retreat from Arctic regions ended in extinction. Penhallow! records some petrified wood from Cretaceous strata in Alberta which he names Sequoia albertensis and regards as very similar to the wood of Sequoia sempervirens, but the evidence in favour of a reference to the existing genus is inconclusive. Resin- cells are scattered through the wood; the medullary rays have 1—2 bordered pits in the field, the broadly elliptical pore being generally diagonal to the cell-axis. Tertiary wood from different localities in North America is referred to Sequoia on evidence that is far from conclusive. Prof. Jeffrey? described a particularly well preserved piece of stem from the Miocene auriferous gravels of the Sierra Nevada, near the home of Sequoia gigantea, as Sequoia Penhallowi; though I am informed that he is now inclined to refer the wood to the Abietineae. In his account attention is called to certain features, e.g. the pitting on the end-walls of the medullary-ray cells, the scarcity of xylem- parenchyma, and the presence of vertical and horizontal resin- canals, believed to be traumatic, which are certainly suggestive of abietineous affinity. Prof. Penhallow? described two species from Eocene beds in the North-West Territory as Sequoia Langsdorfir and S. Burgessii, both of which were previously described by Dawson but assigned by him to different positions. In the wood believed to belong to the plant which bore the well-known twigs recorded by many authors as S. Langsdorfii resin-cells are numerous and scattered and resin-canals are present only in a rudimentary form on the outer face of the summer-wood. The pitting of the medullary-ray cells is not described. A peculiar feature in S. Burgessti, if the wood is correctly referred to Sequova, is the occurrence of two kinds of medullary rays, uniseriate and fusiform, the latter containing resin-canals. No resin-canals occur in the wood. Attention has been called (p. 171, fig. 712) to the abundance of petrified stems in the Lower Tertiary deposits in the Yellow- stone Park: some of these are named by Mr Knowlton* Sequova magnifica. A few of the trunks reach a diameter of 6—10 it. 1 Penhallow (08) p. 83, figs. 1—6. 2 Jeffrey (04). 3 Penhallow (03) pp. 41—46, figs. 2—8. 4 Knowlton (99) p. 761, Pls. civ., cv., CX., CXL, CXVI. 348 SEQUOIINEAE (cH. and a height of 30 ft. (fig. 764). The details are imperfectly pre- served: a few ot the tracheids show traces of single and double rows of small bordered pits, but no pits are shown on the walls of the medullary-ray cells. Resin-parenchyma is abundant and Fic. 764. Petrified tree in the Yellowstone National Park (Sequoia magnifica Knowlton). (From a photograph kindly supplied by Prof. Knowlton.) scattered as in Cupressinozylon: it is doubtful whether the wood of Sequoia can be distinguished from that of some other genera included in the genus Cupressinoxylon. Specimens of wood from the Tertiary coal-field of Aichi-Gifu in the middle region of Hondo, XLVIT] SEQUOLITES 349 the main Island of Japan, recently described by Yasui! as Sequoia hondoensis has the following characters: narrow annual rings, tracheal pits usually uniseriate though often biseriate and opposite on the broader tracheids, rims of Sanio present, medullary-ray cells with oval bordered pits on the lateral walls but unpitted else- where,-resin-cells scattered through the spring- and summer-wood, resin-canals present which are believed to be traumatic. The occurrence of this wood according to the author of the species ‘completes in an interesting way the evidence for the existence’ of Sequoia ‘in Cenozoic times throughout temperate regions of the whole northern hemisphere.’ While it is probable that the Sequolineae were very widely spread in the Tertiary period it is open to question if the anatomical evidence is sufficiently clear to justify the reference of the Japanese wood to Seguota. The chief reason for the adoption of that generic name is the occurrence of resin-canals similar to the traumatic ducts in the recent species. The following descriptions include fossils which cannot be re- ferred to Sequowites and others which may reasonably be so named. SEQUOIITES. Brongniart. Sequovites problematica (Fliche and Zeiller). This species, originally described as Sequoia problematica?, is founded on a small elliptical cone from Upper Jurassic rocks in the Boulogne district: in the form of the scales, which show a ridge extending from the edges of the distal surface to a central de- pression in the middle of the cone-scales, the fossil suggests affinity to the recent genus. Zeiller? also records a cone from Jurassic strata in Madagascar associated with branches of the Brachyphyllum type which he says presents all the characters of Sequoia. It must, how- ever, be admitted that in both these cases close relationship to Sequoia has not been demonstrated. Under the name Sequoia minor Velenovsky * describes specimens from the Lower Cretaceous strata of Bohemia consisting of foliage- shoots with small imbricate linear-lanceolate leaves and a small terminal, spherical, cone the sporophylls of which have rhomboidal distal ends and a central umbo: but as in most fossils referred to Sequoia the evidence of generic affinity is inadequate. 1 Yasui (17). 2 Fliche and Zeiller (04). 3 Zeiller (00). 4 Velenovsky (87) p. 638, figs. 11, 12. 350 SEQUOLIINEAE [cH. Sequovites giganteoides (Stopes). This species, under the name Sequoia giganteoides, has recently been founded by Dr Stopes? on a small petrified fragment of a very Fic. 765. Sequovites concinna. Foliage-shoot from the Senonian of Greenland. (Stockholm Museum; nat. size.) a slender foliage-shoot from the Lower Greensand of Luccomb Chine in the Isle of Wight. The pith contains stone-cells, and a single, 1 Stopes (15) p. 70, Pl. 11. text-fig. 16. XLVI] SEQUOLITES 351 undivided leaf-trace enters each decurrent leaf-base; there is a central large resin-canal in the leaves and a considerable develop- ment of transfusion-tracheids on each side of the secretory passage. Palisade-cells are a conspicuous feature and one or two layers of hypoderm fibres occur next the lower epidermis. The author of the species points out the close resemblance between the fossil and the leaves and shoot-axis of Sequoia gigantea. Sequovites concinna Heer. Heer! described several specimens of foliage-shoots and cones from the Patoot beds in West Greenland as Sequoia concinna, the commonest Conifer in these rich Lower Cretaceous strata. The form of the sparsely branched shoots with their long and slender branchlets and straight or slightly curved, decurrent, acuminate, leaves (fig. 765) agree closely with those known as Sphenolepidium Sternbergianum from English and other Wealden rocks, as also with the shoots of Sequovites Couttsiae. The oval cones, 23 x 20 mm., consist of a few scales with 5—6 angled thick distal ends on which there is a median transverse line and a central scar. This species, represented by sterile shoots and cones, has recently . been recorded by Berry? from Upper Cretaceous beds in Pike County, Arkansas. Conites. Conites Gardnert (Carruthers). Carruthers* described a cone and a piece of vegetative shoot from the Gault of Folkestone as Sequovites Gardnert but neither specimen affords any satisfactory evidence of relationship with Sequoia. The shoot is of the Pagiophyllum type, and the cone, 2-5 x 15em., consists of spirally disposed scales with four-sided thomboidal distal ends. There is no information with regard to the seeds: the data being wholly insufficient to serve as a criterion of affinity, the generic name Conites is substituted for Sequozites. A specimen figured by Lange* from the Aachen Sands as Carpolithes hemlocinus Schloth. and compared by him to a Sequoia cone agrees closely with the English species. 1 Heer (83) p. 13, Pls. Li., LIL., etc. 2 Berry (17) p. 172, Pl. vit. figs. 1—5. 3 Carruthers (69?) p. 7, Pl. 1. figs. 7, 8. 4 Lange (90) Pl. xxxu1. fig. 7. 352 SEQUOIINEAE [cH. Conites ovalis (Carruthers). The type-specimen from the Gault of Folkestone? is an oval cone 6 cm. long and about 2-5 cm. in diameter; the scales are cuneate and the exposed ends transversely elongated and hexagonal (fig. 766). It bears a close resemblance to Geinitzia gracillima, but in the absence of any details with regard to anatomical features or seeds the non-com- mittal name Conites is employed. Sequotites Holsti Conwentz ex Nathorst MS. This species?, from the Holma sand- stone (Senonian) of Sweden, is founded on fragments of foliage-shoots covered with spirally disposed, appressed, broadly tri- angular leaves. The specimens are not well enough preserved to show in detail the ana- tomical features, but Conwentz considers such characters as he was able to recognise tas Gueeiiiee ae favourable to Nathorst’s adoption of the gize.) SAS generic name Sequoiites. The species is, however, not above suspicion as a record of a Conifer closely allied to Sequoia. Fic. 766. Conites ovalis. Sequoutes Langsdorfii (Brongniart). Brongniart? instituted this Tertiary species under the generic name Tazites, and Heer? in his description of foliage-shoots from Miocene beds in Switzerland adopted the designation Sequoia. In habit S. Langsdorfii is practically identical with Sequova sempervirens and by many authors it is spoken of as the direct ancestor of the recent species. Under this species Schimper? includes a fairly long list of synonyms—species referred to Taxites, Taxodium, Cupres- sites, and other genera—which serves to emphasise the fact that impressions of sterile branches with distichous, linear, leaves cannot 1 Carruthers (71) p. 3, with text-figure. 2 Conwentz (92) p. 28, Pls. 11., Iv., VIIt. 3 Brongniart (28) A. p. 108. 1 Heer (55) A. p. 54, Pls. xx., XXI. 5 Schimper (72) A. p. 216. XLv11| SEQUOIITES 353 in many cases be identified with Sequoia with absolute certainty. Differences are pointed out by authors in their description of species between the leaves of such recent forms as Taxus baccata and Sequoia sempervirens, but an examination of actual specimens re- veals the inadequacy of such fine distinctions as are sometimes quoted. Our knowledge of the cones is confined to external characters and these afford a more substantial basis than the foliage-shoots on which to form an opinion with regard to the striking similarity between the Tertiary and existing species. Sequovites Langsdov fii is recorded by Gardner? from the Eocene beds in Mull, but the identification rests on sterile branches bearing linear-lanceolate decurrent leaves 6—9 mm. long; the main axis of a branched specimen bears scale-like leaves appressed to the stem except at the distal end of the lamina and, as in the recent species, scale-leaves occur at the base of each lateral shoot. The species is recorded also from Styria?, from Miocene beds in Greece® where it is represented by foliage-shoots and cones, from Italy‘, Germany, and other European localities. A very similar form, originally named by Brongniart® Taxites Tournali, is described by Gardner® from the Middle Bagshot beds of Bournemouth and with it he unites S. Hardt: Heer founded on material from Bovey Tracey. S. Tournali is characterised by the association of distichous (fig. 794, A, B, p. 408) and smaller appressed leaves like those of Sequoia gigantea, an association also met with in S. sempervirens. S. Tour- nalt is recorded by Saporta’ who figures branches and cones from Eocene strata in Provence. Laurent® figures fragmentary speci- mens, which he refers to S. Langsdorfii, from the Aquitanian series in the Puy-de-Dome. Sequovites Langsdorfii is very abundant in Arctic Miocene rocks: Nathorst® speaks of numerous branches in Tertiary clays in Elles- mere Land in a remarkable state of preservation so that they could be washed out and isolated like dried specimens in a herbarium. A peculiarity of the Ellesmere specimens is the occurrence of very 1 Gardner (86) p. 41, PI. x. fig. 1. 2 Ettingshausen (57) Pl. 1. fig. 3. 3 Unger (47) Pl. 11. figs. 17—23. Saporta (68) refers S. Langsdorfit as figured by Unger to 8. Tournali. 4 Squinabol (92) p. 26, Pls. xv., XvI. 5 Brongniart (28) A. p. 108. 6 Gardner (86) p. 40. 7 Saporta (65°) Pl. m. fig. 1. § Laurent (12) p. 65. 9 Nathorst (11%) p. 225. Ss. IV 23 354 SEQUOIINEAE ‘{CH. fine teeth on the edge of the lamina!. Similar teeth are stated by Nathorst to have been seen in one or two examples of Sequoia sempervirens, and it suggested that the papillae, which are a normal feature of the recent species, were more strongly developed in the Tertiary type. Heer? records the species from Miocene beds in Greenland and states that it is one of the commonest Conifers in Disco Island, from the Mackenzie River, Alaska, Spitzbergen, and Sachalin Island. The fragments reproduced in fig. 767 were col- lected in Disco Island and are now in the Dublin Museum with other fossils described by Heer; the long linear leaves, A, are decurrent and in some cases the lamina shows fine transverse striations: the smaller leaves shown in fig. 767, B, are referred by Heer to a distinct species S. brevifolia, but there is no - important difference between the two forms. Palibin® figures sterile shoots from the Sichota-Alin moun- tains. Penhallow? records the spe- cies from British Columbia and 9-767. Sequotites Langsdorfi. (From specimens in the Dublin Museum other localities, and to the same described by Heer.) type he assigns some petrified wood from the Queen Charlotte Islands though without any real evidence of connexion. Twigs and cones are described by Schmalhausen from Tertiary strata in the New Siberian Islands®, and the species is said to be one of the most abundant and widely distributed types in the Yellowstone National Park®. Remains of more than one species of Sequoia are recorded from Florissant, Colorado, which has recently been described as a Miocene Pompeii: the sedi- 1 Nathorst (15?) p. 10, Pl. 1. figs. 1—15. 2 Heer (68) Pls. 1., xx., Lv., ete.; (71) Pls. xL., xum1., etc.; (75) iii. Pl. o.; (77) i. Pls. xm, xu., xxv.; (78) v. Pl. 1.3 (82) i. Pl. uo. 3 Palibin (04) PL. 11. 1 Penhallow (02) pp. 44, 68; (03) p. 41. 5 Schmalhausen (90) Pl. 1. fgs, 2—11. ® Knowlton (99) p. 682. XLVII] SEQUOIITES 355 . ments of an ancient lake mixed with volcanic ash contain many plant and insect remains and Prof. Cockerell’s careful investigations have led to the discovery of several new types!. Staub2, who records the species from Aquitanian beds in Hungary, gives a list of references to other authors. Making allowances for doubtful identifications based on sterile branches there remain a sufficient number of authentic records to demonstrate the wide range of this species and allied forms in Europe and the Arctic regions during the Eocene and Miocene periods. S. Langsdorfii is said to occur in beds of Lower Pliocene age in France® and a minute cone, only 2 by 1-9 mm. has recently been described by Mr and Mrs Reid from Pliocene deposits in Holland?. The Dutch specimen is referred to Sequoia with some hesitation and it is suggested it may be an immature cone of an undescribed species, which possibly marks the last appearance of the genus in Europe. - Sequotites Couttsiae (Heer). This species was founded by Heer® as Sequova Couttsiae on material from Oligocene beds which form a basin-shaped depres- sion in the granitic rocks of Dartmoor in Devonshire. The material consists of foliage-shoots (fig. 768, A, B), similar in habit to those of the recent species Sequoia gigantea, and globose or sub-globose cones with peltate scales and winged compressed seeds like those of Sequoia sempervirens. Several seeds are said to occur on each cone- scale. Beust® examined wood from Bovey Tracey in which he found tracheids with separate bordered pits and resiniferous xylem- parenchyma as in the recent species. Mr and Mrs Clement Reid’? have recently investigated the Bovey Tracey material and their conclusion is that ‘Sequoia Couttsiae is a true Sequoia and close to the living Sequoia sempervirens and S. gigantea.’ They give the following description of the cones: ‘Broadly oval and abruptly narrowed into the stalk, or somewhat cordate; at the base are a few small recurved wedge-shaped barren scales, the lower ones having their stalks strongly reflexed, the middle ones with stalks 1 Cockerell (06), (08), (082), (083). 2 Staub (87) B. Pl. xry. p. 249. 3 Depape (13). 4 Reid, C. and E. M. (15) p. 55, PI. 1. fig. 13. 5 Heer (62) p. 1051, Pls. nrx.—1x1. 6 Beust (85) Pl. my. figs. 1—8. 7 Reid, C. and E. M. (10) p. 170, Pl. xv. figs. 23—27. 23—2 356 SEQUOIINEAE [cH. at right-angles to the axis; at the apex is a rosette of a few almost sessile barren scales; the arrangement of the scales is distinctly spiral. It is not easy to count the number of the scales, as none of the cones we found are perfect. There would seem to be 20—24 fully developed scales, besides a few undeveloped round the apex and base.’ The scales vary in shape (fig. 768, C, D) and are cuneate or umbrella-shaped, the rugose distal ends have lines radiating from a central umbo; the winged seeds are pendant beneath the thick involuted margin and on the upper surface of one scale five seeds were found. Preparations of the cuticular membrane of the leaves showed irregularly scattered stomata, each surrounded by Fic. 768. Sequoiites Couttsiae. Twigs A, B, and cone-scales C, D, from Bovey Tracey. (Photographs by Mr and Mrs Clement Reid; x3.) a ring of four or occasionally five cells. Gardner! has also described specimens from Bovey Tracey and Hampshire characterised by imbricate keeled decurrent leaves with a free, divergent or falcate, apex and in older branches by more obtuse appressed leaves. The cones in size and form resemble those of Sequova sempervirens while the vegetative branches agree with S. gigantea. There are 3—5 seeds on each scale. Several examples of Sequowtes Couttsiae are figured by Heer? from Miocene beds in West Greenland and he speaks of the species as the commonest Conifer in Disco Island. Gardner points out that the northern form has larger cones and stouter foliage- shoots than the British type and proposes for it a new specific name 1 Gardner (86) p. 36, Pl. vz. ? Heer (68) p. 94, Pls. m1., vii., XLV.; (71) Pls. xu., XLq1., ete.; (83) Pl. yxvmt. XLVII] GEINITZIA 357 S. Whympert. This more robust form occurs also in Spitzbergen, on the Mackenzie River, and elsewhere. Knowlton! records S. Couttsiae along with other species from beds probably of Miocene age in the Yellowstone Park: he assigns to this type specimens described by Lesquereux from Colorado as Glyptostrobus Ungeri and others from the Fort Union Group referred by Newberry to Glyptostrobus europaeus. The cone-bearing branches figured by Lesquereux* from the Western Territories as Sequoia affinis bear a close resemblance to S. Couttsiae. The latter species is recorded by Penhallow’ from the Eocene beds on the Deer River in Canada (lat. 51° and 54° N.). The same or a closely allied type is recorded from Miocene beds in Alsace*, and Saporta® describes very good examples of S. Couttsiae from the Eocene beds at Armissan in Provence. According to Gardner the material referred by Saporta to Heer’s species includes at least two other species. Specimens described by Schmalhausen® from Eocene beds in South-West Russia as S. Couttsiae, though possibly correctly named, are not convincing. Palibin? records this species from Oligocene beds at Molotytchi in the Fatej district, Russia, and discusses the geological age of the strata from which Schmalhausen’s plants were obtained. Ettingshausen’s specimens from Bilin in Bohemia assigned by him to Taxodium dubium may, as Gardner says®, be examples of S. Couttsiae. GEINITZIA. Endlicher. The name Getniizia was given by Endlicher® to a piece of sterile shoot from Lower Cretaceous strata in Saxony previously figured by Geinitz! as Araucarites Reichenbachw (fig. 769), and in the new genus was also included Cryptomeria primaeva Corda". Both these species were referred by Endlicher to Geinitzia cretacea. Corda’s species was founded on several foliage-shoots from Lower Cre- taceous rocks in Bohemia with the habit of Araucaria excelsa and in one or two instances bearing what appear to be terminal buds described by Corda as small cones. In the first instance Geinatzia 1 Knowlton (99) B. p. 681. 2 Lesquereux (78) B. Pl. uxv. 3 Penhallow (02) p. 50. 4 Bleicher and Fliche (92) p. 382. 5 Saporta (65?) Pl. m1. 6 Schmalhausen (83°) Pls. XXXII, XXXVI. 7 Palibin (01) p. 499. 8 Gardner (86) p. 39. ® Endlicher (47) p. 280. 10 Geinitz (42) Pl. xxtv. fig. 4. “4 Corda in Reuss (46) B. Pl. xii. figs. 1—11. 358 SEQUOIINEAE [cH. was applied to branches without any recognisable cones. In 1868 Heer! figured specimens from the Kome beds in Greenland which he believed to be identical with Araucarites Reichenbachtt Gein: though the foliage-shoots bear shorter leaves than those on the type-specimen of Geinitz: Heer states that he was able to examine the type-specimen and assured himself of the specific identity of the German and Greenland specimens; he substituted the generic name Fic. 769. Geinitzia Reichenbachii. Fic. 770. Geinitzia Reichenbachii. (After Geinitz.) ; (After Heer; nat. size.) Sequoia for Araucarites on the ground.that some cones in the Tiibingen Museum from Lower Cretaceous beds in Moravia, attached to branches apparently identical with Araucarites Reichen- bachit, presented a very close. resemblance to those of recent. Sequoias. The Moravian specimens, which he afterwards figured?, are oval and the cone-scales have distally expanded distal ends (fig. 770) like those of Sequoia, but no evidence was obtained as to the number of seeds. Additional examples of vegetative shoots and cones were described by Heer? from Greenland as Sequoia 1 Heer (68) Pl. xuit. 2 Ibid. (69) Pl, 1. 3 Ibid. (75) ii. Pls. x11., XX., XXXIV., ete. XLVI] GEINITZIA 359 * Reichenbachw and this species is recorded by authors from many Lower Cretaceous localities, but in no case is any conclusive evi- dence brought forward in support of the assumed generic identity with Sequoia. Specimens of Sequoia Reichenbachii with foliage and cones are figured by Velenovsky! from Bohemia showing clearly the characteristic peltate cone-scales, and similar examples though with rather larger cones are described from Lower Cretaceous strata in North America®. On the other hand the name Sequoia Reichen- bachw is applied in some cases to fragments of sterile branches un- accompanied with cones: in one instance* evidence was obtained of the occurrence of separate circular bordered pits on the tracheids of some vegetative branches from the Cretaceous beds of Aix-la- Chapelle. It is impossible to say whether such shoots bore cones like those of Geinitzia or Elatides; some at least belong to Elatides curvifolia. Two conclusions are suggested by an examination of the records so far quoted: the use of the generic name Sequoia is not based on any solid foundation and, secondly, itis unsafe to assume that fragments of sterile branches bearing falcate leaves similar to those on fertile shoots referred to S. Reichenbachii belong to that species. The common occurrence of Mesozoic specimens agreeing more or less closely with Araucaria excelsa, while demonstrating the abundance of that form of vegetative shoot, by no means proves the equally wide occurrence of one specific type. It has, for example, been shown by Nathorst® that the branches from Lower Cretaceous or Upper Jurassic rocks in Spitzbergen figured by Heer® as Sequoia Reichenbachit are examples of Elatides curvifolia (Dunk.). The genus Hlatides’ is characterised by cones differing in their flatter scales and more elongated form from those usually assigned to Sequova though the foliage-shoots are of the same type. It is therefore advisable to adopt some provisional generic term for sterile shoots resembling in habit those of Araucaria excelsa and which in the absence of cones cannot be safely assigned to a genus founded on the cone-characters. The name Pagiophyllum® serves 1 Velenovsky (85) B. Pls. vim., 1x. 2 Ward (99) B. Pls. 165, 166; Hollick (06) Pl. 11. fig. 40; Pl. 1. figs. 4, 5. 3 Krasser (96) B. Pl. xvu. fig. 14; Schenk (71) Pl. xx1v. figs. 6, 7. 4 Lange (90) p. 660. 5 Nathorst (97) p. 35. ® Heer (75) ii. Pls. XXXVI., XXXVII. 7 See page 270. 8 See page 274. 360 SEQUOIINEAE [CH. this purpose and it should be applied to sterile branches of the Araucaria type which cannot reasonably be referred to Elatides, Geinitzia, or other genera connoting certain types of fertile shoot. It has, however, been pointed out that in the first instance Geinitzia was applied to sterile shoots, but later this designation came to be associated with cones of elongate-oval form bearing peltate scales. In 1852 Unger! applied Geinitzia to a specimen from Neustadt consisting of a slender piece of foliage-shoot and an imperfectly preserved cone similar to the cones of Heer’s Sequoia Reichenbachit but longer in form. Subsequently Heer? described under the name Geinitzia formosa shoots and cones from Lower Cretaceous strata at Quedlinburg: the cones are similar in form to that figured by Unger and bear cone-scales with polygonal distal ends having a central umbo and radially disposed lines on the exposed surface. Schenk also gives good drawings of Geinitzia formosa. A well pre- served cone very like Heer’s @. formosa was described by Newberry* from the Amboy clays as Sequota gracillima, the specific name having been previously used by Lesquereux for sterile branches from Dakota in conjunction with the generic name Glyptostrobus. Newberry adopted Lesquereux’s specific term because he found in the Dakota beds cones like that from the Amboy clays associated with the branches described by Lesquereux. Newberry’s cone is practically identical with that of Heer’s Geinitzia formosa, but it is _noteworthy that the former is borne on a slender branch having small appressed leaves in place of the more spreading falcate leaves of Heer’s species. This difference in the foliage is of secondary importance in comparison with the close resemblance between the cones. Subsequently Jefirey® obtained good cones from the Matawan formation apparently identical with Sequoia gracillima (Lesq.) as figured by Newberry and he was able to investigate the anatomical features. The pith of the cone-axis contains groups of sclerous cells: the phloem differs from that of Sequoia in the absence of fibres, while the secondary wood has no resin-cells—another difference from Sequoia: the tracheal pits are circular and in no case contiguous and there are no rims of Sanio. The latter feature 1 Unger (52). 2 Heer (71°) p. 6, Pls. 1, 1. 3 Schimper and Schenk (90) A. p. 299. + Newberry and Hollick (95) p. 50, Pl. rx. figs. 1—3. ° Jeffrey (11). XLVII] GEINITZIA; EUGEINITZIA 361 is regarded by Jeffrey as an essential character of the Araucarineae and the absence of any Abietineous pitting in the medullary-ray cells is another Araucarian feature. The conclusion drawn by Jeffrey is that despite the absence of Araucarian pitting on the tracheids the anatomical details point to an Araucarian relationship, the wood of the cone-axis having the characters of Sinnott’s genus Paracedrozylon. No information was obtained with regard to the seeds. Jeffrey's examination of the cone shows, as he says, that it does not agree structurally with the cones of recent Sequoias, but the reference to the Araucarineae rests on a slender basis. In their account of the Kreischerville plants Hollick and Jeffrey! describe some sterile branches as Geinitzia Reichenbach (fig. 806, D ; page 437) which agree closely with specimens referred by authors to Sequoia Reichenbachwi though they might equally well be identified with Elatides curvifolia (Dunk.). For such sterile twigs the name Pagiophyllum would be preferable. The pith of the Kreischer- ville shoots contains groups of sclerous cells; the leaf-bases show in transverse section three resin-canals and these are enclosed by the transfusion-tissue which accompanies the vascular bundle. In its distribution the transfusion-tissue differs from that in Sequovra, which is confined to the flanks of the vascular strand, and agrees with the corresponding tissue in Araucarian leaves. There is no ' xylem-parenchyma and the tracheids have 1—2 rows of bordered pits, in contact or sometimes separate and if in two rows alternate. The wood agrees with that described by Hollick and Jeffrey as Brachyorylon and shows a decided Araucarian affinity. In the absence of cones attached to the shoots it is not possible to settle definitely the systematic position of the specimens. A fact in favour of identifying the branches with Sequoia (or more appro- priately Geinitzia) Reichenbachii is the occurrence in the same beds of detached cone-scales very similar to those of G. gracillima, which are referred to two new genera, Eugeinitzia and Pseudogeimitza. EUGEINITZIA. Hollick and Jeffrey. Eugeinitzia proxima Hollick and Jeftrey. The scales on which this species is founded? closely resemble those of recent species of Sequoia and Geinitzia gracillima, The 1 Hollick and Jeffrey (09) B. p. 38. 2 Ibid. p. 43, Pl. x. fig. 10; Pl. xxv. figs. 1—3. 362 SEQUOILNEAE [CH. vascular bundles were found to be arranged round the margin of the peltate portion of the scales and completely surrounded by transfusion tissue, ‘a feature of marked contrast to the scale- bundles in Sequoia and at the same time one which indicates a strong affinity with the Araucarineae.’ The mature scales afforded no indication of the number or place of attachment of the seeds, but an immature cone lent support to the view that each scale bore four ovules on the peduncle near the cone-axis. Hollick and Jeffrey regard the scales as Araucarian and think it probable that they were connected with the twigs named by them Geinitzia Reichenbachit. PSEUDOGEINITZIA. Hollick and Jeffrey. Pseudogeinitzia sequoiiformis Hollick and Jeffrey. A special generic name? is given to some four-sided scales on the ground that they not only differ in their tetragonal form from the hexagonal scales of Eugeinitzia but probably belonged to a smaller cone, As in the former type the vascular bundles are enclosed by transfusion-tracheids. The investigations of the American botanists show that the sterile branches, G. Reichenbach, exhibit certain Araucarian tendencies and that the cone, Geinitzia gracillima, as also the detached cone-scales, Hugeinitzia and Pseudogeinitza, cannot be included in Sequoia. Until more is known of the morphological nature of the cones described by Heer and other authors as Sequoia Reichenbachit, S. ambigua, etc., their relationship to existing Conifers cannot be settled; but meanwhile it would seem convenient to include both the smaller oval cones and the longer forms represented by G. gracillima in the same genus Geinitzia, applying the name to cones having spirally arranged scales with peltate distal ends superficially resembling those of Sequoia. The name Sequoza, much too freely used by palaeobotanists, has in some cases” been applied to cone-bearing branches that are almost certainly identical with Sphenolepidium Kurrianum (Dunk.). On the other hand for sterile foliage-shoots unconnected with cones the non-committal name Pagiophyllum, is suggested on the ground that foliage-shoots alone cannot be more precisely determined. 1 Hollick and Jeffrey (09) B. p. 45, Pl. x. fig. 14; Pl. xxv. fig. 4. 2 Heer (69) p. 11, Pl. 1. figs. 10—13 (‘Sequoia fastigiata’). XLVII] _ SPHENOLEPIDIUM 363 SPHENOLEPIDIUM. Heer. | Heer! instituted Sphenolepidium in place of Sphenolepis, pro- posed by Schenk? for Wealden Coniferous branches, because of the previous use of the latter name by Agassiz for a genus of fishes. Berry* has recently reverted to the original form Sphenolepis on . the ground that its employment by zoologists is not a serious objection. Schenk’s definition of his genus includes both vegetative organs and cones, but it is desirable that the name Sphenolepidium should be restricted to fertile specimens or at least to specimens which can with reasonable certainty be connected with cone- bearing examples. The habit of the foliage-shoots of the two best- known Wealden and Lower Cretaceous species, S. Sternbergianum and S. Kurrianum, is of the type which leads authors to employ such generic terms as Sequoia, Athrotaxites or Athrotaxopsis, Widdringtonites, Glyptostrobus, Araucarites, and Cyparissidium, but in the absence of cones it is impossible to feel confidence in any attempts to distribute such sterile specimens among genera which are characterised not only by a certain form of foliage-shoot but also by a particular type of cone. The generic name Sphenolepidium should be retained only for specimens with small, more or less globose, cones possessing spirally disposed cone-scales, cuneate, relatively broad and fairly thick. The cones are much smaller and have relatively broader and thicker scales than those of Elatides though there is no essential difference in the vegetative characters of the two genera. No cones have been described throwing any light on the affinity of the genus and like many others it must be left for the present in the category of Coniferae incertae sedis. The leaves are spirally disposed on the comparatively slender branches and are either ovate, triangular, and free only in the acuminate region, or longer and more spreading and falcate; the latter type agrees with Pagiophyllum while some forms bearing Sphenolepidium cones are rather of the Brachyphyl- lum type. Many of the specimens recorded as Sphenolepidium afford no evidence as to the nature of the cones and should be assigned to Pagiophyllum or Brachyphyllum. The genus is charac- teristic of Wealden or Lower Cretaceous strata and is represented ‘in several European districts and in North America. } Heer (81) p. 19. 2 Schenk (71) B. p. 243. 3 Berry (114) p. 290. 364 SEQUOLINEAE _ (cH. Sphenolepidium Sternbergianum (Dunker). This species was originally described by Dunker+ from North Germany as Muscites Sternbergianus and by later authors placed in Araucarites, Widdringtonites, and other genera”. It is impossible to determine the specific limits of this species? and S. Kurrianum (fig. 771): the cones exhibit no well-defined dis- tinguishing characters and the chief distinction is the more spreading foliage of the Araucarian or Pagiophyllum type of S. Sternbergianum. As Berry suggests, this species—described from the Potomac formation and elsewhere in North America and from several European localities— is probably represented in the Lower Cretaceous flora of Greenland under such names as Glypto- strobus groenlandicus Heer and Sequoia fastigiata. Some of the English and German fossils attri- buted to S. Sternbergianum are almost certainly examples of Elatides curvifolia. Sphenolepidium Kurrianum (Dunker). Dunker? originally adopted the generic name Thuites; later authors preferred Brachyphyllum, Widdringtonites, Araucarites and other names. Fontaine>, who records this species from the a Potomac formation, includes in his genus Athro- i taxopsis specimens which cannot be distinguished ie by any features of morphological importance is from Sphenolepidium Kurrianum. The leaves are a ovate, more or less appressed, agreeing with fe. 771. Spheno- Brachyphyllum or in some examples intermediate —‘epidium Kurria- - hie. + f doll : a te Paee num. From the etween the type of foliage assigned to Pagio- Wealden of Sus. phyllum and Brachyphyllum. The Wealden speci- sex. (British Mu- men reproduced in fig. 771 is placed in Spheno- seu, V. 2303; lepidium because of its association with branches, *™*" “7° ) identical in habit, bearing cones; if found as an isolated 1 Dunker (46) A. Pl. viz. fig. 10. 2 For references, see Seward (95) A. p. 205; Berry (114) p. 293. 3 Seward (11°) p. 685. ' 4 Dunker (46) A. p. 20. 5 Fontaine (89) B. Pls. oxxxv., ete. J XLVIT] SCIADOPITINEAE 365 fossil it would be referred to Brachyphyllum. The figured specimen shows the variable form and size of the leaves and there is good reason to believe that the plants represented by the frag- ments included in one or other species of Sphenolepidium were characterised by a considerable range in the habit of the foliage- shoots, a fact which renders of little importance the separation into 8. Kurrianum and S. Sternbergianum based on the form of the leaves in detached branches. The small cones borne terminally on slender branches resemble superficially the cones of Athrotaxis, but no facts are available as to the structure of the cone-scales and there is no evidence on which to found an opinion as to the position of the genus. SCIADOPITINEAE. Though several fossil plants have been compared with the existing species Sciadopitys verticillata, in no case is there any con- clusive evidence of the occurrence of this type of Conifer. Schmal- hausen founded the genus Cyclopitys! for impressions of shoots from Russia bearing whorled linear leaves which he believed to be closely allied to or generically identical with Sciadopitys. Zeiller?, who brought forward strong arguments for assigning the strata regarded by Schmalhausen as Jurassic to the Permian period, considers Cyclopitys to be an Equisetaceous plant. Detached linear leaves similar to those of Cyclopitys are abundant in many Jurassic floras and, as Nathorst® says, they may be compared with several recent genera including Sciadopitys, but without anatomical data accurate determination is impossible. It is stated by Schenk* that the Cretaceous leaves described by Heer as Pinus Crameri agree in their epidermal features with the foliage of Sciadopitys, but in this as in other cases generic identity or even close relationship has not been demonstrated. Goeppert and Menge® describe some single leaves preserved in Baltic amber as Sciadopitytes linearis and 8. glaucescens; they speak of the leaves as having a single vein on the upper face and two veins on the lower surface though it is not clear what morphological feature is represented by the ‘veins.’ 1 Schmalhausen (79) A. p. 39. 2 Zeiller (96) A. p. 477. 3 Nathorst (97) p. 19. 4 Schimper and Schenk (90) A. p. 293 5 Goeppert and Menge (83) A. p. 36, Pl. xr. figs. 117—123. 366 SCIADOPITINEAE [CH. Schenk states that these Oligocene leaves are Dicotyledonous and not the leaves of a Conifer.’ Specimens of fossil wood have been described exhibiting certain features, especially the pitting of the medullary-ray cells, similar to those of Sciadopitys! but the occurrence of such features in other recent genera precludes a definite reference to any one type. SCIADOPITYTES. Goeppert. This name has recently been revived by Halle? for two species of Cretaceous leaves from Greenland, one of which, Pinus Crameri Heer, though compared by Schenk with the leaves of Sciadopitys, was not actually included in Sciadopitytes, while the other is a new species, Scradopitytes Nathorsti. Halle describes these leaves as ‘Coniter-like in habit, with a dorsal groove which is protected by elongated papillae and whose epidermal tissue differs from that of the rest of the leaf through a non-seriate arrangement of the cells and the occurrence of stomata.’ The outstanding feature of the leaves of Sciadopitys is the double nature of the lamina and the morphological peculiarities which have led to its recognition as a phylloclade; but, as Halle admits, there is no evidence that the fossils are other than ordinary simple leaves. The interesting characters described by Halle amply justify the use of a generic name separating the leaves from those known only as impressions, without any structural features pre- served, and referred to Pityophyllum. It is, however, open to ques- tion whether the name Sctadopitytes does not imply more than the facts support. The leaves named by Heer? Pinus Crameri are about 12 mm. long and 2:5 mm. broad: the apex is bluntly rounded and the base is slightly widened. Halle points out that there is evidence that the leaves were cylindrical. The carbonised leaves of this species form thick masses in the shale and excellent pre- parations of the cuticle can be obtained. Halle considerably extends Schenk’s account of the epidermal characters. The apparent midrib is a groove and there is no indication of a true median vein. Rather large stomata are crowded in the groove and 1 See page 138. 2 Halle (15) p. 508. 3 Heer (68) i. Pl. xxiv. figs. 7—18; (75) ii. Pl. xx. pp. 9—15; Halle (15) p. 509, Pl. xu. figs. 1—13. XLVII] SCIADOPITYTES 367 are surrounded by somewhat tangentially elongated cells, the other cells between the stomata being much smaller. The cells on the sloping sides of the groove bear cylindrical papillae. On the whole the structure recalls that of the recent Sciadopitys though as Halle shows there are certain differences. The second species, Sciadopitytes Nathorsti1, was discovered by Nathorst in the Middle beds of Atanekerdluk in West Greenland. The leaves are at least 40 mm. long and about 1 mm. broad: the stomata are confined to the groove as in S. Crameri and numerous papillae are borne on the borders of the median depression. The stomata are not so crowded as in S. Crameri and differ less in size from the other epidermal cells. These two species, though exhibit- ing some similarity to Sciadopitys, can hardly be assumed to belong to plants more closely allied to the recent Japanese Conifer than to other existing forms. The occurrence of the characters described by Halle may be recognised by adding the name Sciadopitytes after the non-committal term Pityophyllum. 1 Halle (15) p. 512, Pl. xm. figs. 16—29. CHAPTER XLVIII. ABIETINEAE. TuE relative antiquity of the different families of the Coniferales is a question which every student of the geological history of the group desires to answer. Reference has already been made to the different views that are held with regard to the phylogenetic relations of the Araucarineae and the Abietineae: conclusions on this subject are based partly on the morphological characters ex- hibited by recent types and in part on palaeobotanical data. The evidence afforded by petrified wood is briefly dealt with in Ch. xuiv: this shows that the features associated with modern Abietineae do not stretch as far back into the past as is the case with the type represented by the wood of the Araucarineae. The evidence derived from a study of impressions of foliage-shoots and cones as well as the meagre data supplied by petrified cones is less easy to interpret because of the greater imperfection of the records. The southern distribution of the Araucarineae predisposes the student in favour of a southern origin, while the essentially northern range of the Abietineae suggests that this family had its birth north of the equator. But conclusions based on such considerations require confirmation from other kinds of evidence. In the Jurassic- Wealden period the Araucarineae were well represented in the northern hemisphere and the impression gained from a survey of Jurassic records is that the Araucarineae shared with other types an almost world-wide distribution. It is much easier for a palaeobotanist to form an opinion as to the period of maximum development and vigour of a given set of plants than to discover a substantial foundation on which to rest a view as to the first appearance or the original home of the earliest representatives of the family-type. It is, perhaps, significant that the Araucarineae are represented in the Jurassic floras of Graham Land on the edge of Antarctica, Australia, and India. The Abietineae, on the other hand, do not CH. XLVIIT] ABIETINEAE 369 bulk largely in Mesozoic floras before the closing stages of the _ Jurassic period and more especially in the earlier days of the Cretaceous era. The abundance of Abietineous cones in Lower Cretaceous strata, a period later than that in which the Arau- carineae are abundantly preserved in plant-bearing deposits, at least points to a later maximum development of the Abietineae, and such data as we have seem to favour a northern rather than a southern origin. Winged seeds, hardly.distinguishable from those of modern Pines (fig. 788, p. 396), from Rhaetic beds in the South of Sweden, foliage-shoots from beds of the same age exhibiting features now associated with the Abietineae, demand serious consideration in connexion with the antiquity of the family, though it can hardly be maintained that they furnish proof of the existence in Rhaetic and Liassic floras of true Abietineae. The occurrence of a winged pollen-grain (fig. 491, G; Vol. 11. p. 298) in the partially decayed wood of Antarcticozylon might be urged as a plea for a southern origin of the family, but an extended bladder-like exine is not a monopoly of the microspores of the Abietineae. The following types selected in illustration of the fossil records of the Abietineae show how difficult it is in many cases to deter- mine the precise position within the family to which cones or foliage-shoots should be assigned. Palaeobotanical literature con- tains many species referred to Abies or Abietites, Cedrus, and other genera, but it is usually impossible from the available data to carry identification so far. A few examples may be quoted: certain Lower Cretaceous cones bear a very close resemblance to those of Cedrus, but an examination of some of the less familiar cones of existing species of Abies and Picea shows that the reasons for connecting the fossils with Cedrus are not entirely satisfactory. The fossil wood described under Cedroxylon does not denote that the parent-plants were more closely allied to Cedrus than to some other genera of the same family. Boulay? has described some seeds from Miocene beds in France as Cedrus vivariensis which he unhesitatingly regards as generically identical with those of recent Cedars, and there is no reason to doubt the correctness of this conclusion. Cone-scales bearing two seeds from Miocene beds in Spitzbergen described by 1 See page 385. 2 Boulay (87) p. 235. 370 ABIETINEAE [cH. Heer! as Pinus (Cedrus) Lopalini may belong to a true cedar cone, but the evidence is hardly convincing. It is not too much to say that even Tertiary records of Conifers seldom enable us to dis- criminate between individual genera. In the absence of anatomical data the needle-like leaves scattered through Mesozoic and Tertiary strata cannot be identified with reasonable certainty. From Upper: Pliocene beds in Germany Geyler and Kinkelin? described a cone as Abies Loehri, and this has more recently been identified as Keteleeria (fig. 786, C, p. 394) by Engelhardt and Kinkelin® on the strength of its external resemblance to K. Davidiana. The reference to Abies# of some leaves enclosed in the Baltic amber affords an example of the assistance afforded by characters recognisable in well preserved material, and it is probable that a fuller knowledge of the epidermal characters of recent Conifer leaves may supply a useful aid to more precise identification. PITYITES. Gen. nov. Endlicher® employed the name Pinites for leaves, male flowers, and cones considered to be closely allied either to recent species of Pinus or to some other genus of the Abietineae, such as Abies, Lariz, or Picea. Many authors have adopted the generic name Pinus in cases where the evidence appears to them sufficiently strong to indicate identity with the existing genus, but it is only cones and foliage-shoots from Tertiary and Pleistocene beds that can as a rule be definitely assigned to such a position. It may, perhaps, be carrying consistency too far to restrict Endlicher’s designation to such specimens as there is good reason for connecting with the recent genus Pinus; but the more restricted use of Pinites has the merit of being less likely to mislead the student and, chiefly on that account, I propose to adopt the genus Prtyites for Abietineous fossils which cannot with confidence be referred to a more precise position. In practice this designation will not often be employed as in most cases cones and vegetative organs occur as separate fossils and are most conveniently described under the 1 Heer (78) i. Pl. 1x. figs. 6—8. 2 Geyler and Kinkelin (90) p. 16, Pl. 1. figs. 13—15. 3 Engelhardt and Kinkelin (08) p. 216, Pl. xxv. fig. 7. 4 Goeppert and Menge (83) A. Pl. xm. figs. 107—110. 5 Endlicher (47) p. 283. : XLVI] PITYITES 371 terms suggested by Nathorst and mentioned below. Putyites is, however, appropriate for such specimens as those represented in figs. 772, 773 which show a direct connexion between cones and foliage-shoots. - Goeppert adopted Pinites for fossil wood in a wide sense, but it has long been the custom to describe petrified wood agreeing structurally with recent Pines and other members of the Abietineae under Kraus’s term Pityoxylon. Nathorst!, with a view to greater convenience, proposed certain subgeneric names as qualifying epithets indicating the nature of the fossils but not implying a direct connexion with Pinus: he adopted the names Pityanthus for male flowers suggesting alliance with those of some Abietineous genus, Pityostrobus for cones, Pityolepis for cone-scales, Pityosper- mum for seeds, Pityocladus for vegetative shoots, and Pityophyllum for detached leaves. To these the name Pityosporites? has recently been added. ; The generic or rather subgeneric term Pityophyllum is apt to mislead the student if used in conjunction with Pinites: the leaves so named, as Nathorst admits, are in many instances almost - certainly derived from plants which do not belong to the Abie- tineae. Under Pityophyllum are included both needle-like leaves which are probably Abietineous with others having a broader lamina (fig. 776) and much more likely to be connected with such genera as Cephalotaxus, Torreya, or Podocarpus. The term Pityosporites® is proposed for microspores provided with wings similar to those of Pinus and other members of the Abietineae, though in this case also relationship with another family, namely the Podocarpineae, is not excluded. These terms whether used as subgeneric titles or as generic designations serve a useful purpose for disjuncta membra, while the name Pitytes is employed for specimens of a more complete kind. The name Abietites has often been used for vegetative shoots and cones* which there is no adequate reason for assigning to a position nearer to Abies than to other genera of the same family: it is desirable to restrict the term to fossils which afford evidence of 1 Nathorst (97) p. 62; (99) p. 16. ~ 2 Seward (14) p. 23. 3 Ibid. 4 Eg. Geinitz (80) p. 12; Fontaine in Ward (05) B. Pl. uxviut. figs. 14—17; Thomas (11) Pl. rv. fig. 16; Pl. v. figs. 1, 2. 24—2 372 ABIETINEAE [cH.” affinity to the recent genus. Similarly, names such as Laricites Cedrites and others implying a more precise determination than is suggested by Pityites may conveniently be used either as sub- generic or generic terms. In the account of recent. Conifers allusion is finds: to the views held by students of fossil plants with regard to the relative position of the Abietineae and the Araucarineae in a chronological sequence. The types selected for description are intended to serve as guides to those who wish to draw conclusions from the geological records, but so long as we have to trust chiefly to impressions without the more certain guidance of anatomical data the inferences drawn cannot be regarded as other than provisional. The evidence of fossil seeds is difficult to interpret, as its value depends on the amount. of importance to be attached to the occurrence of speci- mens closely resembling in the form of the wing the seeds of recent Pines and other Conifers. The winged seeds of Agathis differ in the shape of the membranous appendage from those of Abietineous species, and the oldest winged seeds attributed to the Abietineae, from Rhaetic rocks, exhibit a closer agreement with the Abietineous type. On the other hand it is questionable whether the form of a wing constitutes a safe criterion of affinity. A similar difficulty is presented by ‘winged’ pollen-grains: a bladder-like extension of the exine though usually associated with the Abietineae is a character which is not confined to that family. Foliage-shoots like those of recent Abietineae are recorded from Rhaetic rocks and later Mesozoic strata, but we have no means of determining in the case of the oldest examples whether their superficial resemblance to branches of Cedrus and other genera has a phylogenetic signifi- cance. The generic name Pinites is applied by Renault? to a slender branch from Permian rocks in France bearing spirally disposed filiform leaves 3 cm. long apparently borne singly and directly on the main axis, not on short shoots. It is elsewhere? suggested that this specimen, Pinites permiensis, may belong to a plant allied to Dicranophyllum: there is certainly no adequate reason for the employment of the generic term Pinites. Similarly an impression figured by Stur? from the culm of Altendorf as Pinites antecedens, 1 Renault (93) A. Pl. xxxu. fig. 1; (96) A. p. 377. 2 Page 101. 3 Stur (75) A. Pl. xiv. fig. 4. XLVIII] PITYITES 373 which I was able to examine in the Vienna collection, is too frag- mentary to be determined. The occurrence of linear leaves in fascicles is in itself no real evidence of Abietineous affinity: the clustered leaves of Czekanowskia and Phoenicopsis, especially the former, though essentially similar in habit to the foliage-shoots of some Abietineae are generally believed to belong to plants of another class. The evidence furnished by petrified wood has already been considered: the important point is that there is no satisfactory case of the occurrence of fossil wood of Palaeozoic age! having typical Abietineous features, a fact of importance in relation to the widely spread Palaeozoic woods agreeing in essentials with the Araucarian type. Pityites Solmsi Seward. This name was proposed for some cones attached to foliage- shoots as well as detached cones and vegetative branches from Wealden rocks on the coast of Sussex?: the type-specimens form part of the rich. Rufford collection in the British Museum. The branches are covered with the elongated persistent bases of scale- leaves and in the axils of these are borne numerous long needles (fig. 772). The cones are oblong and bear broad, rounded, scales like those of Pinus Strobus, P. excelsa (fig. 773; of. fig. 704), Picea and Abies; they agree closely with Pityostrobus Carruthersi (Gard.) as also with P. Andraei (Coem.)® from Lower Cretaceous rocks in Belgium and with the smaller cones from the Potomac formation described by Fontaine* as Abietites ellipticus. The preservation is not sufficiently good to show the nimber of leaves in each foliage- spur: the needles may have been borne in dense clusters as in Cedrus. In general habit the species resembles Cedrus and Lari though the greater length of the needles is more in accordance with recent species of Pinus. Shoots similar to those of this species are represented by Prepinus statensis Jeff. from the Cretaceous beds of Kreischerville. Dr Stopes®, following the example of Berry, refers this species to Abietites. - 1 See page 220, also Thomson and Allin (12). + 2 Seward (95) A. p. 196, Pls. xvumr., X1X. 3 Gardner (867). 4 Fontaine (89) B. Pl. cxxxiit. figs. 2—4. : 5 Hollick and Jeffrey (09) B. p. 19, 8 Stopes (15) p. 157. 374 ABIETINEAE [CcH. Pityites (Pinites) evrensis sp. nov. In his account of petrified material from Franz Josef Land Solms-Laubach! describes sections of a Pinus-like leaf from Bell | | Fic. 772. Pityites Solmsi. (British Museum, V. 2169; nat. size.) Island (Eira harbour) probably of the same geological age as the plant-beds of Cape Stephen, which is believed to be Upper Jurassic or Lower Cretaceous. Through the kindness of the Director of the 1 Solms-Laubach (04) p. 12, Pl. 1. fig. 14; Pl. m. fig. 3. For evidence as to geological age, see Newton and Teall (97), (98); Nathorst (99). XLVUI] PITYITES 375 Geological Survey I have been able to examine the sections in the Jermyn Street Museum. Graf Solms-Laubach describes the leaves as oval in section, the upper face strongly convex and the lower almost flat as in two-needled Pines, but as shown in fig. 774, A the leaves may be approximately cylindrical (1 mm. in diameter), like those of Pinus monophylla or the leaves of Cedrus. There is a single vein accompanied by some radially disposed transfusion-tracheids, the whole being enclosed in a single layer of rather thick-walled Fic. 773. Pityites Solmsi. (British Museum, V. 2146; nat. size.) cells. There is no distinct division of the bundle into two halves but there are indications of the presence of a broad median medul- lary ray. The mesophyll-cells have prominent infoldings precisely as in recent Pines, Cedrus, and some other Abietineae (fig. 774, B; cf. fig. 694): the epidermis has a thick cuticle and below it are 1—2 layers of small thick-walled elements. Solms-Laubach speaks of two resin-canals, one at each side of the lamina, but I was unable to distinguish any undoubted canals in the leaf shown in fig. 774. The occasional absence of canals in Abietineous leaves normally 376 ABIETINEAE [oH. possessing them is mentioned in Chapter xt. The leaf for which the specific name eirensis (from Eira harbour) is proposed affords Fic. 774. Pityites (Pinites) eirensis. A, section of leaf. B, mesophyll enlarged. (Museum of the Geological Survey.) an interesting example of an Abietineous type, in all probability of Upper Jurassic age, exhibiting a remarkable resemblance to certain recent species especially Pinus monophylla. XLvur] PITYOCLADUS 377 PITYOCLADUS. Nathorst. Under this name, used as a subgenus of Pinites, Nathorst? includes branches bearing short shoots similar in habit to those of ‘Cedrus and Lariw. Branches of this type, bearing leaves and cones, are illustrated by Pityites Solmsi (figs. 772, 773), but in some cases such vegetative shoots occur as detached fossils and it is to them that Nathorst’s term may conveniently be applied. The striking resemblance of the fossil specimens to shoots of Cedrus and Larix and the frequent association or attachment of needle-like leaves afford strong grounds for assigning the branch-fragments to the Abietineae. Pityocladus Nathorsti Seward. In his description of Schizolepis Follini Nath.? from Rhaetic rocks in Scania, Nathorst includes not only cones with lobed scales characteristic of Schizolepis but leaves and branches. Solms- Laubach® expressed the opinion that we know nothing of the foliage of Schizolepis ‘for there is nothing to make it even probable that the numerous needles which lie one above another in the beds at Palsjé, any more than the branches beset with needle-bearing shoots which Schenk has referred to this genus, have any connexion with Schizolepis.’ In a later account of Schizolepis, Nathorst* suggests the advisability of separating the leaves and branches. from the Schizolepis cones, though as he says the association of the .two sets of organs in more than one locality may be significant. It is, therefore, preferable to assign the vegetative organs to Pityocladus, at the same time keeping in mind the possibility of an original connexion with the cones described under the generic name Schizolepis. In order to avoid confusion I have removed the branches and associated leaves from Schizolepis Follina to a dis- tinct species Pityocladus Nathorsti. The specimens figured by Nathorst consist of (i) a fairly stout axis bearing a smaller lateral shoot like that on which the leaf-clusters of Cedrus and Larix are borne; (ii) separate short shoots characterised by zones of small scars alternating with smooth areas; (iii) numerous crowded linear leaves. 1 Nathorst (97) p. 62. 2 Ibid. (78) B. p. 28. . 3 Solms-Laubach (91) A. p. 70. . 4 Nathorst (97) p. 38. 378 ABIETINEAE _ (CH. Pityocladus longifolius (Nathorst). [And Pityophyllum longifolium Nathorst. | This species was first described by Nathorst! from Rhaetic beds in Seania as Taxites longifolius and afterwards as ? Cycadites. Fic. 776. Pityophyl- lum longifolium. (Nat. size.) Fia. 775. Pityocladus longifolius. Leaves attached to a short shoot covered with scales. (Nat. size: a piece of lamina enlarged to show the fine trans- verse striations. From a specimen from Scania in the Stockholm Museum.) Méller? adopted Nathorst’s generic name Pityophyllum for de- tached leaves and that designation should be used for specimens 1 Nathorst (787) B. p. 50. 2 Moller (03) p. 40, Pl. vz. XLvuT] PITYOCLADUS 379 which afford no evidence as to the nature of the axis. The specimen from the Stockholm Collection represented in fig. 775, from the Rhaetic rocks of Scania, is especially interesting as affording one of the few examples of leaves of the type known as Pityophyllum longifolium attached to an axis covered with short scales. The lamina varies from 1 to 5 mm. in breadth and may be broader: as seen in the enlarged portion the lamina is transversely wrinkled, an appearance characteristic of most forms of the broader Pityo- phyllum leaves and probably produced by contraction on drying. There is little difference between this species and the leaves figured by Nathorst from Scania as Schizolepis Follint. An accurate specific delimitation of Pityophyllum leaves is hopeless. Detached leaves (fig. 776) similar to those shown in fig. 775 are recorded by Moller from Lower Jurassic beds in Bornholm, by other authors from Jurassic strata in Turkestan1, South Russia”, Oregon, Spitz- bergen*, and elsewhere. Pityocladus Schenkt Seward. Schenk also refers to Schizolepis several specimens of branches and leaves ag well as cones which he includes in Schizolepis Braunit Schenk. The larger branches bear leaf-cushions and short lateral shoots with scale-leaves at the base, and in some examples tufts of needles occur on the short shoots. A specimen described by Braun as Isoetes pumilus is identified by Schenk in his monograph of the Rhaetic Flora® as a leaf-bearing shoot like that of a recent Pine. The Swedish specimen represented in fig. 775, also of Rhaetic age, is similar to those included by Schenk in Schizolepis Braunit. Pityocladus kobukensis Seward. This species originally described from the Jurassic beds on the Kubuk River in Chinese Dzungaria as Pinites? is founded on branches bearing short shoots almost identical with P. Schenkv. Short shoots not more than 1 cm. long are borne spirally on a thicker axis and covered with small leaf-scars (fig. 777) exactly as in the corresponding shoots of Cedrus or Larix. With the 1 Seward (074) B. p. 32. 2 Thomas (11) p. 78, Pl. vu. figs. 58—61. 3 Fontaine in Ward B. (05) Pl. xxxv. 4 Nathorst (97). 5 Schenk (67) A. Pl. xtiv. figs. 1—4. 6 Ibid. Pl. xiv. fig. 2. 7 Seward (11) p. 54, Pl. rv. figs. 47—51; Pl. v. fig. 65. 380 ABIETINEAE — [CH. branches are associated numerous needles, 1 mm. broad and at least 5cm. long, sometimes covering the whole surface of the rock. The specimens agree closely with the foliage-shoots of Pityites Solmsi: similar examples are described by Ettingshausen! from Li- assic strata as Halochloris baruthina Ett. A branch with short shoots from Jurassic rocks in Amurland described as Pinites sp. of. P. kobukensis® may be specifically identical with the Dzungaria fossils: an example of the same type lent to me by Dr Krystofovie from Jurassic beds of Amurland shows a forked lateral foliage-shoot. This author has recently described a specimen from Jurassic rocks in Transbaikalia as Pin- ites (Pityophyllum) ef. P. kobukensis?®. Similar though smaller specimens Fic. 777. Pityocladus kobuken- of Abietineous short shoots are de- ame eran Lae scribed by Nathorst‘4 as Pinites (Pityo- , cladus) spp. a and b from Upper Jurassic beds in Eee and compared by him with Pityites Solmst. PITYOPHYLLUM. Nathorst. This name is applied to detached leaves of needle-like form like those of recent Pines or to long linear leaves broader and flatter than the needles of Pinus. Some of the specimens referred ‘to this genus are very similar to the leaves of Keteleeria. In a few cases (fig. 775) the leaves are still attached to a short shoot but usually they occur as detached specimens (fig. 776). The genus is met with in Rhaetic strata but is specially abundant in Jurassic floras and persists through Cretaceous and Tertiary rocks. The leaves generally described under this generic term are broader and flatter than such leaves as those of Pityites Solmsi® and recent 1 Ettingshausen (52) B. PI. m1. fig. 4. 2 Seward (12%) Pl. mr. 3 Krystofovié (15) Pl. vz. fig. 9. 4 Nathorst (97) Pl. 11. figs. 28—30, Pl. rv. figs. 13, 14, 23. 5 Ibid. (97) p. 62. 8 See page 374; also Nathorst (97) Pl. v. figs. 1—10 XLVI] PITYOPHYLLUM; PITYOSTROBUS 381 Pines, and the presence of a fine transverse wrinkling on the lamina is a characteristic feature. Pityophyllum, if employed for both the narrower and broader forms, includes specimens which in all pro- bability belong to Conifers of more than one family: some are certainly Abietineous but the flatter and broader forms bear a closer resemblance to leaves of some species of Podocarpus, Cepha- lotazus or Torreya. Nathorst, who instituted the generic name Pityophyllum, recognises that many of the specimens so named have no real botanical value. Detached leaves of the type included in this comprehensive genus are of little interest; but it is note- worthy that such species as P. Nordenskidldi (Heer) and similar forms are characteristic fossils in Jurassic and Cretaceous strata. Pityophyllum Lindstrémi Nathorst. Under this name Nathorst! includes leaves described by Heer from Cretaceous strata in Greenland as Pinus Quenstedti and P. Peterseni, also specimens from Upper Jurassic beds in Spitz- bergen. The leaves reach a length of at least 8 cm. and are 1—2 mm. broad; the lamina tapers gradually towards the base and is more abruptly narrowed in the apical region; there is a prominent mid- rib on one side and sometimes indications of two finer marginal ‘veins,’ also other longitudinal striations which may mark the position of rows of stomata. These leaves are broader than those of Pityites Solmsi and narrower than very similar specimens described by Heer, Nathorst, and other authors as Pinus or Pityophyllum Staratschini? from Cretaceous and Jurassic rocks. Pityophyllum Nordenskiéldi? (Heer) from rocks of the same age is another similar form having a tendency to a slightly sickle-shaped and transversely wrinkled lamina (cf. fig. 776). Pityophyllum is abundantly represented in Jurassic Floras*: the specimens are, however, of very little interest to the botanist as it is impossible to assign them to a family position in the Coniferales. PITYOSTROBUS. Nathorst. This name is used in preference to Feistmantel’s genus Pino- strobus, recently resuscitated by Dr Marie Stopes, on the ground 1 Nathorst (97) pp. 40, 67, Pls. v., vi1.; (99) p. 20. 2 Ibid. (97) pp. 41, 68, Pls. v., vi. 3 Heer (78) ii. Pl. 11.; Nathorst (97) p. 18. 4 For references see Moller (03) p. 39; Seward (11) p. 53; Krystofovic (10) Pl. mu. fig. 10; Thomas (11) p. 78. . 382 ABLETINEAE ’ (CH. that Nathorst’s term is more appropriate for specimens which do not afford evidenee of closer affinity to Pinus than to other genera of the Abietineae. In cases where the specimens may reasonably be regarded as more nearly allied to Pinus than to any other genus the designation Pinites may be added. There can be no question of the abundance of Abietineous Conifers in Tertiary floras and it is equally true that cones of the Pityostrobus type are widely spread in Lower Cretaceous strata especially in Europe. The evidence furnished by cones clearly points to the existence in Upper Jurassic floras of Conifers closely resembling in the general form of their strobili recent members of the Abietineae. The wide distribution of cone-scales and cones of the Araucarian type in Middle Jurassic floras is in striking con- trast to the scarcity of cones of the Abietineous form in rocks older than the uppermost Jurassic and Lower Cretaceous series. Pityostrobus dejectus (Carruthers). Carruthers! speaks of this Kimeridge cone from Dorsetshire as the oldest example of a Pine-cone. It is represented by a single imperfectly preserved specimen, 2 x 2cm., of globular form with partially destroyed broad and thin cone-scales: though it super- ficially resembles some recent Abietineous cones there is scarcely enough evidence to warrant its inclusion in the Abietineae. The cone was first described as Pinites depressus but owing to the previous use of that specific name by Coemans it was re-named P. dejectus*. Pityostrobus strobiformis (Fliche and Zeiller). A species, described as Pinites strobiformis*®, from Portlandian rocks near Boulogne founded on a single incomplete cone similar in form and in the possession of apparently flat, imbricate, scales to Pinus excelsa (cf. fig. 704, p. 154). The surface-features are not shown on the weathered specimen and there is no definite informa- tion with regard to the number or position of the seeds, but as the authors of the species state the narrow elongate and slightly curved form of the cone, which was probably about 17 cm. long, affords a valid reason for comparison with recent Pines. 1 Carruthers (692) p. 2, Pl. 1. fig. 10. 2 Ibid. (71) p. 2. 3 Fliche and Zeiller (04) p. 802, Pl. x1rx. fig. 6. XLVIT] PITYOSTROBUS. 383 Pityostrobus Sauvagei (Fliche and Zeiller). The type-specimen of this species!, from the Portlandian of Boulogne, is an ovoid cone 4-5 cm. long characterised by distally expanded scales and resembling the small cones of Pinus Laricio. In the absence of further data precise identification is not possible though the fossil is probably correctly regarded as an Abietineous cone of the Pinus Pinaster type. Having regard to the fact that in this and the preceding species the determination is based solely on external form no very definite statement is admissible as to systematic position, but such evidence as there is favours the view that in these two cones we have Jurassic representatives of two sections of the genus Pinus. Pityostrobus Dunkert (Carruthers). Several detached cones bearing imbricate scales, broad and flat like those of Picea, some species of Abies in which the ovuliferous scales are longer than the bract-scales, and certain species of Pinus characterised by flat scales instead of the woody scales of the Pinus silvestris and P. Pinaster type have been described from British Wealden strata as also from other countries. Gardner? instituted the following species: Pinites Carruthersi, P. valdensis, P. cylin- - droides®, P. pottoniensis®, but an examination of the type-specimens shows that the distinctive features are not sufficiently well marked to warrant so many specific names. The Lower Greensand specimen from Potton, P. cylindroides, is water-worn and the shape of the imperfect scales is not the original form; it may possibly be identical with P. valdensis, P. Carrutherst, and P. pottoniensis, and there are no important features in which these forms differ from the longer cones of Pityostrobus Dunkeri. The cones from Brook in the Isle of Wight named by Carruthers* Pinites Dunkert were originally described by Mantell as Abietites Dunkeri*®; they reach a length of over 33 cm. and havea breadth of 3 cm., they are elongate-oval and relatively narrow and the long scales are attached to a slender axis (fig. 778). The seeds, apparently two on each scale, are oval and compressed. Cones of similar form and length are described by 1 Fliche and Zeiller (04) p. 804, Pl. xrx. fig. 7. 2 Gardner (867). . 3 See also Seward (95) A. p. 193; Stopes (15) pp. 138, 140. 4 Carruthers (662) Pl. xx1. figs. 1, 2. 5 Seward (95) A. p. 194. 384 ABIETINEAE _ [CH Fig. 778. Pityostrobus Dunkeri. (British Museum; nat. size.) XLVI] PITYOSTROBUS 385 Velenovsky! from Lower Cretaceous rocks in Bohemia as Pinus longissima, a species recently recorded by Dr Stopes? from the Lower Greensand of England. Though in the absence of foliage-shoots cones of this type cannot be assigned with certainty to any one recent genus, their great length suggests comparison with those of Pinus Lambertiana and P. excelsa rather than with cones of recent species of Picea. Pityostrobus Leckenbyt (Carruthers). This species was first described by Carruthers? from a specimen in the Leckenby Collection, Cambridge, from the Lower Greensand of the Isle of Wight. It is 10 cm. long and 5 cm. in diameter; the Fic. 779. Pityostrobus Leckenbyi. From the Lower Greensand of the Isle of Wight. (After Carruthers; 4 nat. size.) scales agree in external form with those of Cedrus and Dr Stopes has recently proposed the generic name Cedrostrobus* in order to emphasise this resemblance. Prof. Fliche® described a cone from the Argonne as Cedrus oblonga which he believed to be identical specifically with Abies oblonga of Lindley and Hutton, but Dr Stopes gives Fliche’s name as a synonym of Cedrostrobus Leckenbyt. A cone of similar form is also described by Coemans from Belgium as Pinus Corneti® and compared by him with Cedrus. 1 Velenovsky (85) B. Pl. 1. figs. 14—17. ' * Stopes (15) p. 141, text-fig. 38. 3 Carruthers (69?) Pl. 1. figs. 1—5. 4 Stopes (15) p. 143, text-fig. 39. 5 Fliche (96) p. 200, Pl. vit. 8 Coemans (66) p. 11, Pl. v. fig. 3. Ss. IV . 25 386 ABIETINEAE (cH. Though superficially very like a cone of Cedrus (fig. 779), Pityo- strobus Leckenbyi also strongly resembles some species of Picea and Abies in which the bract-scales do not project beyond the semini- ferous scales. I have adopted the non-committal term Pztyostrobus as it is by no means certain that Carruthers’ type is more closely allied to Cedrus than to Abves. Dr Stopes includes in Cedrostrobus a second species, Cedrostrobus Mantelli1, from the Lower Greensand of Kent which Carruthers originally named Pinites: she compares with it a Potomac cone described by Berry? as Cedrus Leei. But these species do not afford any proof of close relationship to the recent genus Cedrus. It is probable that some of the numerous cones found in Lower Cre- taceous rocks belong to trees having the characters of Cedrus, though in the absence of more decisive evidence than has so far been furnished it would seem preferable to retain the wider desig- nation Pityostrobus. Pityostrobus Benstedi (Mantell). The small oval cone on which Mantell? founded the species Abies Benstedi is from the Lower Greensand of Kent. It was subsequently Fic. 780. Pityostrobus Benstedi. Tangential section showing, os, ovuliferous scale; bs, bract-scale; 0, the two ovules on the ovuliferous scale; e, endosperm; w, wing. (After Stopes; x 8.) described by Carruthers! as Pinites and regarded by him as prob- ably more nearly allied to Cedrus than to Pinus. Dr Stopes® has recently made a further examination of the structure of this type 1 Stopes (15) p. 145, text-fig. 40. 2 Berry (11) Pl. uxx. fig. 4. 3 Mantell (46) p. 52, Pl. m1. fig. 2. 4 Carruthers (66?) p. 541. 5 Stopes (15) p. 130, text-figs. 32, 33; Pls. x., x1. XLVI] PITYOSTROBUS 387 and. believes it to be more closely related to Abies. The cone-scales, as seen in fig. 780, show their double nature, and on the semini- ferous scale are two ovules provided with wings, w. The ovules are immature and there is a small space in the middle of the endo- sperm (fig. 780, e). Pityostrobus (Pinites) sussexiensis (Mantell). This Aptian (Lower Greensand) species, originally named by Mantell Zamia sussexiensis! and afterwards referred by Carruthers? to Pinites, has recently been more fully described by Dr Stopes® under Pinostrobus. The cone, 14 cm. long and nearly 5cm. in diameter, bears overlapping scales with a thickened, curved, distal margin 2 cm. broad and, on the exposed surface of the specimen, 13cm. deep. The seeds, two on each scale, have a corrugated stone-layer in the testa and bear massive, broad wings. A section through the middle of a scale shows an irregularly scattered double set of variously orientated vascular bundles and resin-canals. The ‘species closely resembles Pinus excelsa and P. Strobus and is con- sidered by Dr Stopes to occupy a position between these two types. The inference to be drawn from this and several other cones from Lower Cretaceous strata is that Abietineous cones having more or less flat scales as seen on the surface were more abundant in Europe in the early Cretaceous forests than those in which the distal ends of the scales are rhomboidal as in Pinus Pinaster. Pityostrobus oblongus (Lindley and Hutton). The type-specimen, a water-worn cone from Dorsetshire, pre- sumably from Lower Greensand rocks, was described by Lindley and Hutton as Abies oblonga*: it was assigned by some authors to Pinites. Williamson® gave an account of a cone from Sidmouth in Devonshire, which he referred to Pinites oblongus, though Dr Stopes® expresses a doubt as to the identity of his specimen with that described by Lindley. Schimper? employs the name Cedrus and Goeppert assigns the species to Abietites*. It is impossible to determine the position of the specimen represented in fig. 781 among the Abietineae. 1 Mantell (43) p. 34. 2 Carruthers (662) p. 541, Pl. xx. figs. 5, 6. 3 Stopes (15) p. 123, Pls. x., xr. 4 Lindley and Hutton (35) A. Pl. 137. 3 Williamson (86). 5 Stopes (15) p. 135. ? Schimper (72) A. p. 299. 8 Goeppert (50) p. 207. 25—2 388 : ABIETINEAE . [cH. The type-specimen bears a resemblance to P. Leckenbyi (fig. 779); the scales are broad and thin at the distal end and the axis is relatively slender. The French specimens from Lower Cretaceous rocks referred by Fliche to this species as Cedrus oblonga are con- sidered by Dr Stopes to be specifically identical with Pityostrobus Leckenbyt. Fic. 781. Pityostrobus oblongus. (After Lindley and Hutton, from Stopes; nat. size.) Pityostrobus hexagonus (Carruthers). A large cone 15 cm. long and 4 cm. in diameter composed of stout woody scales with hexagonal apophyses was described by Carruthers as Pinites hexagonus from the Gault of the South of England’; it agrees externally with recent cones of the Pinaster type but the distal ends of the scales are almost flat and nothing is known of the internal structure. The species may be compared with the Lower Cretaceous species P. Quenstedtt Heer?. Pityostrobus (Pinites) Andraei (Coemans). The cones of this species (fig. 782), the commonest type in the Lower Cretaceous rocks of Hainault?, are 10—14 cm. long and 1 Carruthers (71) p. 2, Pl. xv. 2 Heer (71°). * Coemans (66) p. 12, Pl. rv. fig. 4; Pl. v. fig. 1. XLVII] PITYOSTROBUS 389 2-25 cm. in diameter. The cone-scales are compared with those of Pinus excelsa, but the distal ends are stouter than in the recent species and more like those of P. Pinaster. Heer! compares P. Andraei with his Pinus Quenstedti from Moravia in which the scales have thick apophyses with a central umbo. The needles of the Moravian species are 20 cm, long and appear to be either 3 or 5 in a fascicle. It is impossible within the limits of a general text-book to discuss the bearings of the nume- rous Tertiary records of Abietineous cones, many of them undoubtedly borne by species of Pinus. A few examples only are mentioned primarily in order to draw the attention of students to the importance of making a critical examination of Tertiary and Pleistocene Coni- fers. The neglect of Tertiary plants is largely due to the unscientific treatment by authors of detached leaves of Angiosperms which in many instances are referred to recent genera on wholly inadequate grounds, but the more trustworthy nature of the material on which species of Fic. 782. Pityostrobus Abietineous cones are founded deserves careful = (Pinites) Andraei. consideration and would probably yield results ee of considerable importance. Pityostrobus (Pinites) macrocephalus (Lindley and Hutton). This species, founded on a cone 12cm. long and 6cm. in diameter, was in the first instance described by Lindley and Hutton? from an account furnished by Prof. Henslow and named Zamia macrocephala; it was found near Dover and believed to be derived from the ‘Greensand formation.’ A second specimen from Faversham in Kent was described by the same authors as Zamia ovata’, Endlicher4 assigned the cones to Zamiostrobus and Miquel® 1 Heer (69) p. 13, PL. 1. fig. 11. 2 Lindley and Hutton (35) A. Pl. cxxv. 3 [bid. (37) A. Pl. coxxvi A. 4 Endlicher (40) p. 72. 5 Miquel (42) p. 75. 390 ABIETINEAE [CH. proposed the name Z. Henslowi: their Abietineous nature was first recognised by Corda}, and Carruthers? subsequently gave some account of the internal structure and employed the generic name Pinites. The discovery of additional specimens in situ enabled Carruthers to assign P. macrocephalus to Eocene beds at the junction of the Woolwich and Thanet beds with the London Clay. Both Carruthers and Gardner? retain both specific names, but an -examination of the specimens convinces me that there are no differences worthy of specific recognition. The following brief account is based on an examination of sections in the British Museum and in part on notes supplied by Mr Dutt of Queens’ College, Cambridge, who is preparing a fuller account of the material? The cones are ovoid-cylindrical and obtuse; the weathered surface (fig. 783) shows slightly convex polygonal areas without any trace of a central umbo. The axis is slender in com- parison with that of most recent species of Pinus; the stele includes a fairly large pith of thick-walled cells surrounded by a vascular cylinder in which foliar gaps are formed by the exit of the double sporophyll-traces. It is noteworthy that no resin-canals occur in the xylem. A ring of large resin-canals lined with thin-walled epithelial cells occurs outside the phloem. The cone-scales: are given off almost at right-angles and then bend sharply upwards and become slightly broader near the surface of the cone (fig. 784, B). In one section a portion of a subtending bract-scale was recognised. The seminiferous scales are composed of thick-walled cells and contain idioblasts like those in Araucarian leaves, also resin-canals: two ovules occur in a depression near the base of the scales. The sporophyll-trace divides in the scale into several bundles, and in places there are indications of a second series of inversely orientated strands. The comparatively large ovules, nearly 1 cm. long, are attached by a short stalk, and in places the remains of.a wing can be seen. Although the integument is thick and lignified and the micropyle closed there are no embryos and no indication of archegonia in the partially preserved nucellar tissue. In the 1 Corda in Reuss (46) B. * Carruthers (66?) pp. 536, 540, Pl. xx. 3 Gardner (86) pp. 63, 65, Pl. xrv. 4 Dutt (16). XLVIII] PITYOSTROBUS 391 ovule shown in fig. 784, A, the contracted nucellus, n, forms a cylindrical column which presents a misleading resemblance to the mn i a nee yy oy H q ND ey RON Fic. 783. Pityostrobus (Pinites) macrocephalus, (After Gardner; nat. size.) Fig. 784. Pityostrobus (Pinites) macrocephalus. A, section of an ovule; n, nucellus. B, longitudinal section of part of the cone. (From sections in the British Museum; B, slightly reduced.) prothallus tent-pole of Ginkgo ovules: at its blunt apex are two winged pollen-grains. Prothallus-tissue is also represented. Carruthers compared the species with Pinus Pinaster but the 392 ABIETINEAE (CH. surface-features are more like those of the cones of Pinus excelsa (of. fig. 704, p. 154) and similar types. Pityostrobus (Pinites) Plutonis (Baily). This species was founded by Baily1 on part of a cone from the plant-beds in the basalts of Antrim and described, in greater detail Fic. 785. Pityostrobus (Pinites) Plutonis. (After Gardner; nat. size.) and from better material from the same locality, by Gardner®. The cones are 7—9 cm. long and 2—3 cm. in diameter, characterised by woody scales with sub-hexagonal apophyses with a central umbo and a rounded upper margin; there are two seeds with long and narrow wings on each scale. In one case (fig. 785) three cones are attached in an erect position to a branch covered with persistent leaf-bages. The foliage-leaves were borne in pairs and reached a length of 10—15 cm. A similar type with shorter needles from the 1 Baily (69) Pl. xv. fig. 1. 2 Gardner (86) p. 69, Pls. xv.—xvum1. XLVIIT] PITYOSTROBUS 393 same locality is named by Gardner Pinus Bailyi1, These Irish specimens agree in the cones and foliage-spurs with such recent species as Pinus halepensis and P. Pinaster, but the apparently erect position of the cones of the fossil type is a distinctive feature. Tertiary cones similar to Pityostrobus Plutonis are illustrated by Pinus robustifolia Sap.? from Provence, P. Kotschyeana (Ung.), _ originally described by Unger and recorded by Tuzson® from Hungary, P. transsylvanica Pax*, a North American form, which the author of the species compares with Pinus Balfouriana, and _P. prae-montana described by Mogan® from Lower Austria. Pityostrobus (Pinites) palaeostrobus (Kittingshausen). This type originally described by Ettingshausen® from Haring in the Tyrol is recorded from many Tertiary localities. The cones are ovate sub-cylindrical with scales of the Pinus Strobus form and the needles are borne in fascicles of five. Heer? refers to this species © some thin and long needles from the Miocene of Greenland, but it is not clear that the needles are in fives. The species is recorded from Hungary®, Germany, France, and elsewhere. A cone of similar form is described by Unger® from the Oligocene of Kumi as Pinus megalopis and it is associated with quinary fascicles. Pityostrobus MacClurw (Heer). This species described by Heer! as Pinus (Abies) MacClurat from Miocene beds in Banks Land, lat. 74° 27’ N. is represented by a specimen in the Dublin Museum. The narrow oval cone, 6 x 1:5 cm., consists of imbricate scales with the upper margin rounded or irregularly truncate: some of the scales show indications of a pair of seeds. Though similar to cones of Picea, the fossil cannot be definitely assigned to any recent genus. Similarly, Miocene speci- mens of cones, scales, and leaves from Spitzbergen referred by Heer!! to Pinus Abies L. do not afford satisfactory evidence of their generic position. 1 Gardner (86) p. 73. _ # Saporta (73) p. 94, Pl. 1. 3 Tuzson (092) p. 240, Pls. xtv., xv. 4 Pax (07) p. 310. 5 Mogan (08) figs. 1—3. ” § Ettingshausen (55). 7 Heer (83) Pl. uxx. fig. 8; Pl. uxxxvuz. figs. 5, 6. 8 Staub (85). ® Unger (67) Pl. xvi. 10 Heer (68) i. p. 134, Pl. xx. figs. 16—18. 11 Ibid. (71) iii. p. 41, Pl. v. figs. 35—49. 394 ABIETINEAE [cH. While most of the Tertiary species of Pityostrobus agree closely with recent types some exhibit more or less striking peculiarities. A species described by Engelhardt and Kinkelin as Pinus Timleri4 from Pliocene beds near Frankfurt is founded on pieces of large cones characterised by cone-scales with a conical distal end having 3 to 5 flat surfaces (fig. 786, A). The authors compare it with Pinus Gerardiana from Afghanistan. Fic. 786. A, B, part of a cone and a seed of Pinus Timleri. OC, Keteleeria Loehri. See page 370. (After Engelhardt and Kinkelin; nat. size.) The material obtained from Pleistocene beds is often well enough preserved to afford trustworthy data with regard to the later geo- logical history of different genera. Clement Reid? identified a small cone from the Cromer Forest bed as Pinus silvestris, and from this horizon in Sweden Nathorst* records the same species; it is recorded also from Pleistocene deposits associated with Elephas primigenius in France‘ -and similar evidence has been obtained from Switzer- land, Germany, Denmark, and other regions. At a later date the former range of Pinus silvestris and other types is illustrated by the Pa 1 Engelhardt and Kinkelin (08) p. 205, Pl. xxv. figs. 1—4. 2 Reid, C. and E. M. (15) Pl. x. figs. 7, 8. 3 Nathorst (10) p. 1354. 4 Fliche (00). XLVI] PITYANTHUS 395 evidence of submerged forests and, as we ascend the scale, the records become more legible and the prehistoric merges into the historic era. A cone apparently identical with the Spruce Fir (Picea excelsa) found in the Pre- glacial beds on the Norfolk! coast (fig. 787) is a relic of the flora which existed in England when the Rhine after receiving ‘many large tributaries—now separate rivers—seems to have flowed across the present bed of the North Sea.’ The same species is recorded from Pliocene beds on the Dutch-Prussian frontier, also from the valleys of the Main and Neckar, the speci- mens from the latter locality being referred by Gliick? to Picea excelsa var. alpestris. Sernander? has discussed the past history of Picea in Scandinavia and quotes records of the occurrence of the genus in other parts of Europe. Similar instances of the wider range of Abietineous genera are given by Berry* and other authors who have p27 piven ee described Pleistocene plants in North Pre-glacial beds at Mun- America. From the facts at present avail- desley, Norfolk. (After able it would seem that Pinus and allied Beis nat. size.) genera were more abundantly represented in the Tertiary and Post- Tertiary floras in Europe than in American strata of the same age. PITYANTHUS. Nathorst. Pityanthus granulatus (Heer). This species, described by Heer® from the Patoot (Cretaceous) beds in Greenland as Ophioglossum granulatum and afterwards described by Newberry® from the Amboy clays, has recently been identified by Dr Stopes’ as a long microstrobilus of some Abietineous Conifer, probably a Pinus. 1 Reid, C. and E. M. (08) Pl. xv. fig. 147. 2 Gliick (02). 3 Sernander (93). See also Andersson (10) and W. B. Wright (14) for excellent summaries of Pleistocene history. 4 Berry (07); (10°); Penhallow (04). 5 Heer (83) Pl. ivi. figs. 8.9. ® Newberry and Hollick (95) Pl. 1x. figs. 1I—13. 7 Stopes (11*) text-figs. 1, 2. 396 ABIETINEAE [CH. This author examined the American specimen, which she regards as a fertile shoot of a three-needled Pine: the strobilus is 35 mm. long and from it winged pollen-grains were isolated. In its unusual length the strobilus resembles the male flowers of Pinus australis from Florida. PITYOSPERMUM. Nathorst. The few specimens chosen for description afford examples of some of the oldest records of fossils, agreeing in the form of the wing with recent Abietineous seeds and, as far as I know, none have been discovered in strata below the Rhaetic. From Tertiary rocks numerous winged seeds are recorded, but these are of no special interest and they are usually accompanied with foliage-shoots, cones, or other fossils which afford more trustworthy data as to relationship. Pityospermum Lundgreni Nathorst. Nathorst described several winged seeds from the Rhaetic beds of Scania as Pinus Lundgreni}; they are 9-—11 mm. long and 4 mm. broad, the actual seeds being 3—4 mm. in length. Two examples from Stabbarp in the Stockholm Museum are represented in fig. \ 788. To the same species Nathorst referred y some imperfect cylindrical cones bearing ——_ thin imbricate scales and reaching a length Fic. 788. Pityospermum of 3—5 cm. and a diameter of 1:2—2cm.; Lundgreni. From Stab- ee barp in Scania; Rhaetic. he also suggested the possibility that some (Stockholm, Musser, short shoots and long needle-like leaves de- nat. size.) scribed as Schizolepis Follint Nath. may belong to the plant which bore the cones and seeds. In a later account of Schizolepis? he expressed the opinion that in the absence of any proof of actual connexion the leaves and short shoots should be separated from Schizolepis and included in Pinites. These leaf-fascicles are described under the name Pityo- phyllum and the seeds, which occur as separate fossils, are alone included in Pityospermum Lundgreni. The striking resemblance of the seeds to those from Franz Josef Land (fig. 789) and recent 1 Nathorst (78) B, p. 31, Pl. xiv. figs. 9 @, 13—17; Pl. xv. figs. 1—2. 2 Ibid. (97) p. 38. XLVIIZ] PITYOSPERMUM 397 Abietineous seeds is a valid redson for suggesting the inclusion of the Rhaetic specimens in the Abietineae, though it would be going too far to conclude that the seeds were borne on cones generically identical with or even closely related to those of any existing representative of the family. A Pliocene seed figured by Engelhardt and Kinkelin! as Pinus Timleri (fig. 786, B) bears a close resemblance in the form of the wing to some of the Rhaetic specimens. Pityospermum Nilssom Nathorst. This species, also from the Rhaetic flora of Scania?, is character- ised by the much longer wing (2-7 cm.) which in size and form differs much more widely than Pityospermum Lundgreni, P. Nan- sent, and other Jurassic types from the wings of any recent seeds. Pityospermum Nansen Nathorst. The seed shown in fig. 789, A, 11 mm. long, is drawn from a specimen in the Museum of the Geological Survey (Jermyn Street) collected by Dr Koettlitz in Franz Josef Land and of Upper Jurassic or Wealden age: ‘this and other seeds are figured by Newton and Teall?, The name Pityospermum Nanseni was applied by Nathorst* to similar speci- mens obtained by Dr Nansen from the same region. Other winged seeds from Franz Josef Land closely resemble Heer’s species Pinus Maakiana® from Jurassic rocks in Siberia. A seed, 1-2 cm. long, from Weal- den beds in the South of England is repro- duced in fig. 789, B®: this is possibly a distinct species, but the specific determina- Fic. 789. A, Pityospermum Nanseni. B, Pityosper- mum sp. (A, drawn from a specimen in the Museum of the Geological Survey figured by Newton and Teall; B, from a speci- men, V. 2323, in the British Museum from Wealden rocks.) tion of separate seeds of this form is of little value unless the differences are well marked. The important point is the striking resemblance between such seeds as those shown in figs. 788, 789 1 Engelhardt and Kinkelin (08) Pl. xxv. fig. 4. 2 Nathorst (78) B. p. 32, Pl. xv. figs. 17—19. 3 Newton and Teall (97) Pl. xxXVIII. 4 Nathorst (99) p. 18, Pl. u. figs. 12, 13. 5 Heer (77) ii. Pl. xvi. fig. 1. 8 Seward (95) A, p. 198. 398 ABIETINEAE [cH. and seeds of recent Pines and other Abietineae. It is not possible to determine the precise generic affinity of seeds of this type, but their practical identity with recent Abietineous seeds warrants their reference to that family. ; PITYOSPORITES. Seward. This generic name has been adopted! for spores, provided. with bladder-like extensions of the exine, agreeing in size and form with those of recent Abietineous genera. Winged pollen occur also in the Podocarpineae, but the fossil examples so far recorded are much more like the microspores of Abietineous genera than those of Podocarpus, Dacrydiwm and Microcachrys. Pityosporites antarcticus Seward. In the course of examining sections of wood collected by Mr Priestley on the Priestley Glacier (approximately lat. 74° S.) I noticed two small microspores in the siliceous matrix of the partially decayed stem?: one is shown in fig. 491, G (Vol. 111. p. 298); the longest axis is 80 and the central part bears two bladders charac- terised by a fine surface-reticulation similar to that on recent spores. A microspore of Pinus silvestris has a length of 75. It is very unlikely that the spores have any connexion with the stem in which they are preserved; they bear a much closer resemblance to the microspores of Abietineous genera than to the spores of the Podo- carpineae: the probability is that the Antarctic specimens belong to some Abietineous Conifer though this cannot be definitely stated. lt is probable that the upper part of the Beacon Sandstone, from which the boulder containing the fossil is believed to have been derived, is not older than Lower Mesozoic, e.g. Rhaetic. Pityosporites sp. Among the spores found by Nathorst? in Liassic clay from Hor in Scania were several winged microspores, one of which is repro- duced in fig. 790, C from a photograph kindly supplied by Prof. Nathorst. The length of the spore is about 100 and in the - shape of the bladders it agrees closely with the microspores of Picea excelsa?. 1 Seward (14) p. 23. i ? Ibid. p. 23, Pl. vii. fig. 45. 3 Nathorst (08) p. 13, Pl. 1. 4 Kirchner, Loew, and Schréter (06) p. 151, fig. 68. XLVIII] ENTOMOLEPIS 399 Pityosporites sp. In his account of petrified plant-remains from Franz Josef Land, probably of Wealden or approximately Wealden age, Graf Solms-Laubach! mentions the occurrence of well preserved pollen with bladders and figures a piece of a cone with flat scales similar Fic. 790. A, B, Pityosporites sp. from Franz Josef Land. C, Pityosporites sp. from Scania. (A, B, from specimens in the Museum of the Geological Survey ; C, after Nathorst. ) to that of a Picea. The drawings reproduced in fig. 790, A, B were made from specimens found in sections of the material examined by Solms-Laubach in the Geological Survey collection. The longest diameter is 70—90y; in form and size the microspores resemble those of recent Pines. The apparently broad wall shown in fig. 790, A is the result of the obliquity of the section. ENTOMOLEPIS. Saporta. Entomolepis cynarocephala Saporta. Saporta? instituted this generic name for some cones from the Oligocene plant-beds of Provence 8—10 cm. long, ovate-elliptical, and composed of spirally arranged coriaceous scales not thickened at the apex but prolonged beyond the imbricate broad portion into a long recurved, acuminate and fimbriate, spinous process. No seeds have been found and there is no evidence as to internal structure. Saporta considers the cones to belong to some extinct type and, as Zeiller® says, they are probably Abietineous. 1 Solms-Laubach (04) p. 11. 2 évrouos, cut up; Aeris, scale. Saporta (65?) p. 55, Pl. . fig. 3. 3 Zeiller (00) B. p. 278. 400 ABIETINEAE [CH. CROSSOTOLEPIS. Fliche. Crossotolepis Perroti Fliche. This generic name! was proposed for an imperfectly preserved cone, from Oligocene beds near Embrun in the French Alps, of elongate-cylindrical form, 13-3 cm. long and 3-5 cm. in diameter, characterised by the fimbriate edge of the imbricate, highly inclined, scales, which bear two seeds. The cone agrees closely with several recent Abietineous types especially with Picea Menziesit and other species of Picea, but is distinguished by the deeply fimbriate upper margin of the thin cone-scales; it is difficult to determine how far this feature is the result of secondary causes: Fliche believes it to be an original character comparable with that which led Saporta to found the genus Hntomolepis for an Oligocene cone from Armissan in Provence. It is not certain whether the seeds are winged. Fliche is no doubt correct in his conclusion that Crossotolepis is an Abietineous cone closely allied to Picea and probably related to Entomolepis. He refers the two Oligocene genera to the Abietineae; they differ from any recent forms in the greater dissection of the distal edges of the seed-bearing scales, which in this respect are comparable with the more feebly lobed scales of the cones of Picea Engelmann and other species. Our knowledge of both genera is, however, meagre and all that can be said is that the type-specimens afford some evidence of the former occurrence of some Tertiary Abietineous Conifers distinguished by the distally dissected scales from any recent types. PREPINUS. Jeffrey. This genus was instituted? for lignitic specimens of short shoots and leaves from Middle Cretaceous beds on Staten Island, N.Y. characterised by the large and indefinite number of leaves borne on a single short shoot, the presence of a basal sheath of scale-’ leaves, and by certain anatomical features, particularly the mesarch structure of the single leaf-bundle, the occurrence of a complex system of transfusion tissue, and other features. 1 xpocawrés, fringed, tasselled. Fliche (99) p. 474, Pl. xm. 2 Jeffrey (087) Pls. xm, x1v.; Hollick and Jeffrey (09) B. p. 19, Pls. 1x, XXI.—XXIv. XLVIII] PREPINUS 401 Prepinus statensis Jeffrey. The short shoots (fig. 791, B), rather less than lem. long, consist of a relatively broad axis bearing on the upper part nume- tous spirally disposed truncate portions of leaves, in some cases Fig. 791. Prepinus statensis. A, Transverse section of a detached leaf believed to belong to P. statensis. B. Short shoot showing the basal portions of needles and, below, the scars of scale-leaves. C. Transverse section through part of a leaf-fascicle. (After Jeffrey; A, x 30; B, x7; C, x94.) more than 20 on a single shoot. Scars of scale-leaves are represented by rhombic areas near the base. There is no evidence as to the nature of the branches on which the foliage-spurs were pro- duced. The leaves are polygonal in section (fig. 791, A, C): there are two marginal resin-canals and a single median vascular bundle as in some recent Pines. There is a considerable development of Ss. IV . 26 402 ABIETINEAE é [cH. sclerous tissue in the ground-tissue of the leaf and a complete absence of mesophyll with infolded walls like that of recent and some fossil Pines (cf. fig. 774). The phloem is represented by a crescentic space in the leaf shown in fig. 791, A. The dark zone surrounding the bundle consists of thick-walled and relatively long transfusion-tracheids and external to these is a broader sheath of short transfusion-tracheids, but there is no endodermal layer aud no admixture of parenchyma with the tracheids. The xylem is composed partly of centripetal and in part of centrifugal elements : the centrifugal xylem forms an uninterrupted arc next the phloem, and between this and the transfusion-tissue on the lower side of the bundle seen in fig. 791, A the centripetal xylem is represented by radial rows of tracheids separated by spaces. Spiral proto- xylem elements occur between the two groups of metaxylem. It is interesting to find similar transfusion-tissue in some leaves of true Pines described by Jeffrey from the same beds, but their bundles are double and composed of centrifugal xylem only as in modern species.. The pith of the axis of a Prepinus shoot contains nests of sclerous cells: the leaf-traces pass through the cortex as single bundles, and the single ring of wood contains a row of resin-canals blocked by tyloses. The tracheids have uniseriate bordered pits which are often contiguous and separated by Sanio’s rims. Jefirey compares this species with Pinites (=Pityites) Solmsi Sew.1 (fig. 772) from English Wealden beds and with shoots described by Fontaine? from the Potomac series under Heer’s generic name Leptostrobus. Prepinus viticitensis Jefirey. This species*? was founded on specimens of short shoots from the Lower Cretaceous clays of Gay Head, Martha’s Vineyard, Massachusetts, which are considered to be closely allied to Potyites Solmsi. The wood of the axis, representing a single year’s growth, contains two series of resin-canals and, as in the type-species, the canals of the leaves are in continuity with those in the cortex of the shoot-axis, whereas in recent Pines the leaf-canals end blindly. 1 Seward (95) A. p. 196, Pls. xvmu., x1x. See page 373. 2 Fontaine (89) B. p. 227, Pls. c1.—crv. etc. 3 Jeffrey (10). XLVIII] PREPINUS 403 As the choice of the name Prepinus implies, Jeffrey regards the type of foliage-shoot represented by these species as the direct ancestor of the leaf-spurs of recent Pines. The short shoots of Prepinus are smaller than those of existing species of Pinus, but in the numerous and spirally arranged leaves they resemble those of Cedrus and Larix. Anatomically the fossil leaves differ widely from any Abietineous types, and were it not for the occurrence of true Pine needles in association with Prepinus, which to some extent bridge the gap between Prepinus and Pinus, one might be sceptical with regard to the close affinity of Prepinus to recent Pines. Jeffrey compares the structure of the leaf of P. statensis with that of some leaves of Cordaites, but the agreement is probably not so close as Dr Stopes’ description!, quoted by Jeffrey, suggests. Jeffrey? regards the short shoots of Pinus and other Abietineae as a primitive attribute of the Coniferous stock and as one of several reasons for believing the Abietineae to be the oldest tribe of Conifers. Prof. Thomson? has recently discussed the value of the evidence based on the short shoots of Pinus and Prepinus and comes to the conclusion that the foliage-spurs of Pinus are specialised shoots and do not belong to the category of: primitive forms. In Cedrus, Larix, and Pseudolarix the leaves are spirally disposed on the short shoots, while in Pinus they are fewer and cyclic. The frequent occurrence of more than the normal number of leaves on the foliage-spurs of Pinus has already been mentioned: in healthy plants supernumerary foliage-leaves are not uncommon and an increase in the number of needles is also induced by wounding. The spirally arranged scale-leaves below the whorled leaves on a short shoot of Pinus are homologous with the scale- leaves on ordinary branches, and on seedling Pines they are re- placed by the primordial leaves: transitional forms occur between these three forms of leaf. The persistent short shoots of Cedrus, Lariz, and Pseudolariz, as also of Ginkgo, are regarded as the more primitive condition as compared with the deciduous nature of the cyclic foliage-shoots of Pinus. Thomson notes that short shoots of Pinus may proliferate like those of Cedrus and Larix. He concludes that ancestrally ‘the leaves of the Pines were spirally arranged on ordinary branches and that the spur is derived from 1 Stopes (03). 2 Jeffrey (10?) p. 331. 3 Thomson (14). 26—2 404 ABIETINEAE (CH. XLVIII this condition.’ The fossil shoots from the Potomac series de- scribed as Leptostrobus longifolius differ in the larger number of the needles from modern Pines and resemble abnormal short shoots of Pinus excelsa! produced by wounding. The short foliage-shoot of Prepinus furnishes a more completely known example of a branch bearing spirally disposed leaves. In view of the palaeonto- logical evidence and of the facts obtained from a study of recent Pines it would seem that, as Thomson holds, the present form of the Pine spur is the result of specialisation and not a primitive feature. 1 Thomson (14), Pl. xxm. fig. 10. CHAPTER XLIX. PODOCARPINEAE. THE data on which to base any conclusions as to the antiquity or former distribution of the genus Podocarpus or of Conifers believed to be closely allied to recent Podocarps are unfortunately derived from records which in the majority of cases are far from satisfactory and consist mainly of detached leaves. Velenovsky? refers some linear leaves from the Perucer series of Bohemia to Podocarpus (P. cretacea), but they exhibit no distinctive characters. Some of the numerous leaves described as species of Podocarpus are in all probability correctly regarded as Tertiary representatives of the recent genus, but it is often impossible to state with any confidence that detached leaves should be referred to Podocarpus, or preferably to Podocarpites, rather than to 2 more comprehensive genus such as Faxites or Elatocladus. In spite of the fragmentary nature of the evidence and the fact that no undoubted example of a Podocarpus fertile shoot has been discovered, an examination of the published records leads to the conclusion that in Tertiary floras, particularly in those of Eocene age, species closely allied to existing Podocarps were abundant in Europe, a conclusion that is especially interesting from the point of view of the present geographical distribution of the Podocarpineae. A brief account of some of the better known examples of Tertiary species of Podocarpus, which are transferred to the genus Podocarpites in accordance with the practice usually adopted in the case of fossil species, may serve to illustrate the nature of the material and the wide range of the specimens. Dr Guppy? in his very suggestive remarks on the present dis- tribution and means of dispersal of Podocarpus writes: ‘If we assign a home in the high latitudes of the northern hemisphere to a genus that was well represented in Europe in the Tertiary period, a movement of migration southward would explain most of the difficulties in the present distribution. The great vertical range 1 Velenovsky (85) B. Pl. xu. figs. 5—11. 2 Guppy (06) p. 302. 406 PODOCARPINEAE [cH. of some of the species leads us to attribute a corresponding power of adaptation to the genus in respect of widely different climates.... - With such a capacity for adaptation, migration of the genus would be rendered easy over the globe.’ The geological history of the Podocarpineae is unfortunately very fragmentary but such data as are available lend support to the view that there was ‘a centre of diffusion in the extreme north,’ the present distribution of the B Cc Fic. 792. Podocarpites eocaenica. (A, after Heer; B, C, after Gardner; nat. size.) family being as in the case of the Araucarineae the result of migration from other parts of the world where the plants are now represented only in the floras of a bygone age. PODOCARPITES. Andrae. Podocarpites eocaenica (Unger). This species, from Eocene beds in South Styria, was founded’ by Unger? on detached sub-falcate leaves 3—12 cm. long (fig. 792, A) with a short petiole or a more or less sessile lamina. 1 Unger (51) p. 158, Pl. xxi. figs. 11—16, XLIx] PODOCARPITES 407 Gardner? has described examples of this species from the Middle Bagshot beds of Bournemouth, the Lower Bagshot of Alum Bay in the Isle of Wight, and from other British localities, A good impression from Bournemouth is reproduced in fig. 792, B: the leaf is 7 cm. long and 3 mm. broad, the apex is sharply pointed and there is no petiole. Fig. 792, C shows a leaf of slightly different form which may belong to the species. No reproductive organs have been found. Ettingshausen? records specimens of this type from Eocene beds at Haring in the Austrian Tyrol where the species is said to be abundant, from Leoben® in Styria, Bilin in Bohemia, Sagor in Carinthia®, and elsewhere. Heer® states that the species is common in Swiss Eocene deposits and Engelhardt? describes examples from Oligocene rocks in Bohemia. A similar or perhaps specifically identical type was figured by Lindley® from * Kocene plant-beds at Aix in Provence as Podocarpus macrophylla, but Saporta®, who figured additional specimens from the same locality, proposed the name Podocarpus Lindleyana on the ground that Lindley’s designation implies identity with a recent species. A leaf referred to P. eocaenica is figured by Massalongo!® from Tertiary beds in Italy. Some vegetative shoots originally described by De la Harpe! from Alum Bay as Cupressites elegans are referred by Gardner?” to Podocarpus and this determination derives support from the variety in the foliage illustrated by his specimens: in some branches the linear leaves are two-ranked while in others the leaves are spirally disposed and three-sided, a diversity met with in recent species. The imperfectly preserved fragment reproduced in fig. 793 from a careful drawing by Miss Woodward of the actual specimen is figured by Gardner as a fertile branch bearing a single seed with 1 Gardner (86) p. 48, Pl. 1. figs. 6—15. 2 Ettingshausen (55) p. 37, Pl. 1x. figs. 14, 15. 3 Ibid. (88?) p. 277. 4 Ibid. (67?) p. 118, Pl. xin. figs. 1, 2. 3 Ibid. (85) p. 6, Pl. xxvui. fig. 12. 6 Heer (55) A. p. 53, Pl. xx. fig. 3. 7 Engelhardt (85) p. 315, Pl. vit. figs. 37, 38. 8 Lindley in Murchison and Lyell (29) p. 298, fig. A. 9 Saporta (62) p. 216, Pl. 1. fig. 7. 10 Massalongo (59) p. 166, Pl. v. fig. 36. 1 De la Harpe in Bristow (62) p. 111, Pl. v. fig. 3. 72 Gardner (86) Pl. vi. 408 PODOCARPINEAE [cH. a fleshy base as in certain existing species, but the details are too indistinct to afford any proof of affinity to Podo- carpus. Gardner also describes a globose wrinkled seed, 16 mm. in diameter, as ? Podocarpus argillae- fr londinensist from the London Clay which bears a close resemblance to the seeds of Podocarpus elata. The specimens from Eocene beds in the Island of Mull described by Gardner? as Podo- 5, 793, Podocar- carpus borealis, consisting of small falcate leaves ites elegans. Sup- and seed-like bodies, are too imperfect to be posed fertile shoot. determined with accuracy. Fig. 794, C, C’ repre- oe Ne British Museum sents a type from Bournemouth described as figured by Gard- Podocar pus incerta which differs fromother species 2°) in the absence of a definite midrib; the linear-lanceolate coriaceous leaves, reaching a length of 3.cm., are decurrent and appear to have’ several parallel veins, a feature characteristic of the section Nageta: A Cc Cc’ B Fic. 794. A, B, Sequoiites Tournali (see p. 353). C, C’, Podocarpites incerta. (From — specimens in the British Museum described by J. Starkie Gardner; A, B, V. 524; C, V. 522; nat. size.) it is, however, doubtful whether this species should be included in a genus implying affinity to Podocarpus. Some leaves figured by Schmalhausen® from Oligocene strata in Russia as species of Podocarpus are too imperfect to afford any trustworthy evidence as to the occurrence of the genus Podocarpites. Ettingshausen* 1 Gardner (86) p. 52, Pl. 1x. figs. 35, 36. 2 Ibid. (87) A. Pl. xm. figs. 3—11. 3 Schmalhausen (83?) Pl. xxxII. 4 Ettingshausen (86) Pl. vm. figs. 25—27. XLIX] _ PODOCARPITES 409 describes as Podocarpus prae-cupressina foliage-shoots and a very imperfectly preserved seed from Eocene rocks in New South Wales: the same author records Podocarpus Parkerit, a doubtful species, from New Zealand. Podocarpites Campbelli Gardner. Shoots bearing leaves similar to those of P. eocaenica are described by Gardner from the Eocene plant-beds in the basalts EM Fic. 795. Podocarpites Campbelli. (After Gardner; nat. size.) of Mull as Podocarpus Campbells? (fig. 795). The linear acuminate straight or slightly curved leaves with a contracted decurrent base are about 7 cm. long and possess a well-defined midrib. Gardner states that the late Prof. Oliver regarded the fossils as the branches of some Podocarp; they are compared with Podocarpus falcata of the Cape and Tropical Africa and with the South African species P. Thunbergiw. 1 Ettingshausen (87) Pl. 1. figs. 12—14. 2 Gardner (86) p. 97, Pl. XxvI. 410 PODOCARPINEAE [cH. Fossils believed to be related to DACRYDIUM. The records of the rocks afford very little information with regard to the past history of Conifers allied to the recent genus Dacrydium. The marked dimorphism of the foliage-shoots (fig. 708, p- 160), their close resemblance to branches of some other Conifers, as also to Lycopodiaceous plants and some of the larger Mosses, are serious difficulties in the way of recognising representatives of this genus among impressions of vegetative branches. It is inter- esting to find that the most promising piece of evidence of the occurrence of a fossil type (Stachyotarus) allied to Dacrydium is furnished by a Rhaetic flora, a fact pointing to a high antiquity of the plan of reproductive shoot characteristic of existing species. Schenk! compares with Dacrydium some obscure and small fragments from the Coal Measures of China which he described as Conchophyllum Richthofeni, but there are no substantial grounds for such comparison. The specimens consist of pieces of slender axes bearing spirally disposed bracts or small leaves showing at the base of the ovate-oblong lamina a slight depression from which a seed may have fallen. The Lower Cretaceous foliage-shoots from Bohemia described by Velenovsky? as Dacrydium densifolium have no claim to be accepted as branches of a Podocarpineous Conifer. Ettingshausen® figures from Eocene beds in Australia and New Zealand sterile twigs assigned respectively to Dacrydium cupresst- noides and D. prae-cupressinum: in neither case is there any evi- dence as to the nature of the reproductive organs, and the form of the foliage-shoots might with equal probability be interpreted as evidence of other Conifers or of some Lycopodiaceous plant. STACHYOTAXUS. Nathorst. Stachyotaxus elegans Nathorst. The genus Stachyotaxus* was instituted for some Rhaetic specimens from Scania originally named by Agardh Sargassum septentrionale and Caulerpa septentrionalis; the former was re- named by Nathorst Carpolithes septentrionalis and the latter 1 Schenk (83) A. p. 223, Pl. xxi. figs. 21—26. 2 Velenovsky (85) B. Pl. xm. figs. 1—2. 3 Ettingshausen (86) Pl. vim. figs. 23, 24; (88) Pl. 1. fig. 19. 4 Nathorst (86) p. 98. XLIX] STACHYOTAXUS 411 Cyparissidium septentrionale. The discovery of additional material led Nathorst to transfer some of the specimens to a new genus Stachyotaxus. The foliage-shoots of Stachyotaxus elegans! are dimorphic; some of the leaves are appressed and imbricate as in Fic. 796. Stachyotaxus elegans. A, B, % nat. size; C, slightly enlarged; D, x3}. . (After Nathorst.) Cyparissidium and some other Conifers while others are linear and distichous (fig. 796, A), sessile and decurrent, with a lamina reaching a length of about lem. The epidermal cells have straight walls and the stomata occur in two rows on the lower surface. The megastrobili (fig. 796,B) have the form of spikes about 1 Nathorst (082) p. 11, Pls. 1, 1m. 412 PODOCARPINEAE [CH. 5—6cem. long; the fairly stout axis bears sporophylls, approxi- mately at right-angles, consisting of a short and relatively thick stalk expanded into a triangular scale bearing two ovate seeds, 3—3-5 mm. long, each being enclosed basally in a cupule (fig. 796, C, D): the distal end of each sporophyll forms an upturned acumin- ate apex. The cuticle of the seed-coat shows that the latter consisted of thick-walled cells, and within the testa Nathorst found the remains of a much more delicate membrane, possibly repre- senting the nucellus. Nathorst compares the sporophylls with those of recent Dacrydiums, though in Stachyotarus there are normally two seeds on each sporophyll and not one as in the recent genus, a difference possibly of no great importance. In habit the fertile shoots of the fossil type are comparable with those of Podocarpus spicata. Miss Gibbs! in her account of recent Podo- carpineae expresses agreement with Nathorst’s view that Stachyo- taxus is probably a member of that family. Nathorst describes a second Swedish species but from a slightly lower horizon in the Rhaetic series. This species, Stachyotaxus elegans, is characterised by longer and stouter megastrobili reaching at least a length of 12 cm. and by longer linear leaves 10 cm. long. Hartz? refers to Stachyotarus septentrionalis some sterile shoots from Lower Jurassic, or Rhaetic, beds in East Greenland, and Halle? draws attention to the superficial resemblance to the Swedish type of some vegetative twigs from Graham Land which he refers to the genus Hlatocladus. Strobilites. Strobilites Mulleri Seward and Bancroft. The specimen on which this species is founded* was obtained by Hugh Miller from Upper Jurassic beds on the North-East coast of Scotland and inaccurately figured in the Testimony of the Rocks®. Fig. 797 is from a careful drawing by Mr T. A. Brock of the original specimen in the Edinburgh Museum. A slender axis bears nume- rous spirally disposed oval bodies (6 x 5 mm.) which are no doubt seeds: each shows a differentiation into an inner portion surrounded 1 Gibbs (12) p. 539. ? Hartz (96) Pl. x1x. figs. 2, 3. 3 Halle (13?) p. 83. 4 Seward and Bancroft (13) p. 882, Pl. 1. fig. 13. 5 Miller (57) B. p. 493. XLIX] PHYLLOCLADITES 413 by a flat border and it may be that the latter is the impression of a sarcotesta. Another possible interpre- tation is that the oval bodies are seeds in intimate association with fertile bracts. The strobilus bears a close resemblance to Stachyotazus elegans Nath. from the Rhaetic of Sweden compared by the author of the species with an ovuliferous shoot of Podocarpus spicata and Dacrydium Franklin and believed to be allied to the recent genus Dacrydium, a view upheld by Miss Gibbs? in her account of recent Podocarps. It is not improbable that Strobilites Millert is more closely allied to the Podocarpineae than to any other family of Conifers. Saxegothopsis Dusén. In his account of a Tertiary flora, possibly Oligocene, from localities on the Magellan straits Dusén? describes a single leaf as Sazxegothopsis fuegianus on the : ground that it resembles the leaves of Saxe- p97. sirobilites ead gothaea conspicua. The lamina, rather less‘ (After Seward; Edin- than 2cm. long, is linear-lanceolate witha burgh Museum; nat. spinous apex and a short stalk; no veins “*) are. shown in the drawing. There is no substantial reason for regarding this solitary fossil as a fragment of a Conifer allied to Saxegothaea. Dusén admits the lack of satisfactory evidence indicating generic identity, but the specimen hardly merits the distinction of being made the type of a new genus. PHYLLOCLADINEAE. PHYLLOCLADITES. Heer. This generic name was given by Heer‘ to a fossil, subsequently transferred to a new genus Drepanolepis® (fig. 798, C), which affords 1 Nathorst (08) Pl. m. figs. 1—27. 2 Gibbs (12) p. 539. 3 Dusén (99) p. 105, Pl. x1. fig. 10. 4 Heer (75) ii. p. 124, Pl. xxxv. figs. 17—21. 5 Nathorst (97) p. 43. 414 PHYLLOCLADINEAE [CH. . no real evidence of a relationship to the recent genus Phyllocladus, but Heer’s term may be retained for a species described by Ettingshausen as Phyllocladus aspleniordes?. Phyllocladites asplenioides (Kttingshausen). This Tertiary species from New South Wales presents a close resemblance to Phyllocladus (fig. 675, p. 107) and is probably an Eocene representative of the genus. The specimens consist of fairly stout axes bearing cuneate and irregularly lobed leaf-like organs, in some cases apparently subtended by small scales—a circum- stance which justifies Ettingshausen’s suggestion that the lateral members are phylloclades. In one case a seed occurs at the base of a phylloclade. The phylloclades are practically identical with some forms of Thinnfeldia, particularly with American examples referred by Berry to his genus Protophyllocladus; the laminae of the Australian species agree both in form and venation with those of the Cretaceous American impressions, but in the latter there is no good reason for interpreting the leaf-like organs as flattened branches. Ettingshausen’s species is the only fossil that has come under my notice that has any substantial claim to be considered a satis- factory record of the recent genus Phyllocladus. In the account of Thinnfeldia in Volume 11? reference is made to the resemblance of some impressions included in that genus to the phylloclades of Phyllocladus, a resemblance which led Ettingshausen to assign the type-species of Thinnfeldia to the Coniferae. Berry? considers that Ettingshausen’s comparison with Phyllocladus, though not applicable to Jurassic and other of the older species of Thinnfeldia, is valid in respect of certain Middle and Upper Cretaceous forms for which he instituted the genus Protophyllocladus. Attention has previously been called? to the inadequacy of the evidence in support of the conclusion implied by the adoption of the name Proto- phyllocladus. The specimens for which this name was instituted consist of comparatively large coriaceous leaf-like impressions, linear or ovate-lanceolate with an entire, undulate, or crenulate margin, provided with a short petiole prolonged as a stout midrib from which numerous simple veins are given off at an acute angle. In 1 Ettingshausen (86) p. 94, Pl. vir. figs. 283—31. 2 Page 543. 3 Berry (03) B. 4 Seward (04) B, p. 31. XLIX] PROTOPHYLLOCLADUS 415 venation and to a large extent in shape the fossils conform to the characters of Thinnfeldia. Protophyllocladus subintegrifolius (Lesquereux). This species was originally described by Lesquereux! from Dakota beds in Nebraska as Phyllocladus subintegrifolius. Heer? described similar or possibly identical specimens from the Atane beds of Greenland as Thinnfeldia Lesquereuxiana and included Lesquereux’s name as a synonym: these Greenland specimens reach a length of 8cm. and a breadth of 2cm. Heer classes the species among plants of uncertain position and compares the impressions with the phylloclades of Phyllanthus. The species is recorded by Hollick® from Martha’s Vineyard and other localities, by Berry? from the Raritan flora, and as Thinnfeldia Lesque- reuxiana by Newberry and Hollick® from the Amboy clays and other Cretaceous floras®: most of the examples are detached leaves (or ? phylloclades), linear, spathulate, or ovate with an entire or toothed margin while a few are branched (fig. 798, A, B). In no single case is there any evidence in favour of regarding the speci- mens as phylloclades rather than leaves. Until additional facts are obtained it would seem preferable either to retain the generic name Thinnfeldia used by several authors or to adopt some title which does not suggest a relationship to any recent genus. Zeiller? has described a specimen from the Great Oolite of Marquise (N. France) as Protophyllocladus sp.: this is the first European record for Berry’s genus. A similar species is represented by Protophyllocladus polymorphus (Lesq.) first described by Lesquereux® from Vancouver Island as Salisburia polymorpha and afterwards transferred by Knowlton® to Thinnfeldia. Another closely allied type is Protophyllocladus lobatus Berry! from Upper Cretaceous rocks in South Carolina. Some of these supposed phylloclades closely resemble flattened 1 Lesquereux (74) p. 54, Pl. 1. fig. 12. 2 Heer (82) i. p. 37, Pl. xxiv. figs. 9, 10; Pl. xiv. figs. 1—12. P 3 Hollick (06) p. 36, Pl. v. figs. 1—6. 4 Berry (11°) p. 98, Pl. rx. 5 Newberry and Hollick (95) p. 59, Pl. x1. 6 Lesquereux (91) Pl. 1. figs. 1—3. 7 Zeiller (12) p. 13. 8 Lesquereux (78) B. p. 84, Pl. Lx. figs. 40, 41. 9 Knowlton (93) p. 47, Pl. v. figs. 1—4; Berry (03) B. p. 442. 10 Berry (03) B.; (14) p. 17, PL. m. figs. 9—13. 416 " PHYLLOCLADINEAE [cH. leaf-like branches from Kreischerville for which Hollick and Jeffrey instituted the genus Androvettia! (fig. 806, A—C): some examples of that genus are clearly distinct as there are small leaves borne Fic. 798. A, B. Protophyllocladus subintegrifolius. C. Drepanolepis rotundifolia. D. Phyllocladopsis heterophylla. (A, B, after Newberry and Hollick; C, after Nathorst; D, after Fontaine. Nat. size.) : on the flat phylloclade, but other specimens are hardly distinguish- able from Protophyllocladus. Though it would be foolish to deny that the marked resemblance as regards form and venation between certain species of Protophyllocladus or Thinnfeldia and Phyllocladus may be significant, it is true that no satisfactory evidence has been produced in support of actual affinity to the recent genus. 1 See page 436. XLIX] TAXITES 417 Palaeocladus Ettingshausen. This name was applied to a single species, Palaeocladus cunei- _ formis, from New South Wales! founded on a specimen described as a linear cuneiform phylloclade with a median rib from which a few simple veins are given off at an acute angle, each passing up the middle of a lateral tooth; the teeth or serrations are regarded as lateral phylloclades concrescent with one another and with the main flattened axis. A few small scales occur at the base of the compound phylloclade. Ettingshausen’s ingenious interpretation, though possibly correct, rests on too slender a basis to justify the assumption of any affinity to Phyllocladus. Phyllocladopsis Fontaine. This generic term was proposed for some branched foliage- shoots from the Potomac formation? characterised by small broadly ovate leaves with spreading veins. The type-species Phylloclad- opsis heterophylla (fig. 798, D), as Fontaine and Berry state, bears a close resemblance to some forms of Nageiopsis and there can be little doubt as to the foliar nature of the appendages, which afford no evidence of morphological affinity to the phylloclades of a Phyllocladus. The species must be left for the present as a plant of uncertain position: it would seem more appropriate to adopt the generic name Nageiopsis than to make use of a designation suggesting a relationship which has not been established. TAXINEAE. TAXITES. Brongniart. This generic name, first proposed by Brongniart® for some Tertiary and one Jurassic species, has been widely used for vege- tative shoots bearing spirally disposed and distichously placed linear leaves (e.g. fig. 802) resembling in habit those of Tazus, Sequoia sempervirens, Cephalotaxus and some other recent Conifers. In the absence of anatomical characters or reproductive organs it is impossible to determine the precise position of shoots of this common form and, the designation Tazites, as generally employed, 1 Ettingshausen (86) p. 93, Pl. vt. figs. 33, 33a. 2 Fontaine (89) B. p. 204, Pl. uxxxrv. fig. 5, Pl. cuxvu. fig. 4; Berry (03) B. 3 Brongniart (28) A. p. 108. 8s. IV 27 418 TAXINEAE [cH. cannot therefore be interpreted as indicative of affinity to Taxus. Tazites has also been applied to fossil wood believed to belong to Conifers allied to Taxus or other members of the Taxeae, but this name has been superseded by Taxoxylon. Taxites is misleading in its implication of relationship as the specimens so named afford no proof of their systematic position within the Coniferales. Halle? has proposed to employ the generic name Elatocladus for sterile shoots of the Taaxites type, the latter name being restricted to fossils which there are good grounds for assigning to the Taxeae. Among other genera to which authors have referred vegetative shoots superficially similar to Taxus is Palissya?, a genus founded in part on a definite type of reproductive shoot very different from that of Taxus. The investigation of the epidermal structure of some specimens of Jurassic age placed by Feistmantel in Palissya has led Miss Holden to institute a new genus Retinosporites® based on characters recognisable only in the case of fossils which retain their cuticular features. Another example of foliage-shoots of the Taxites habit, which are shown by their fertile shoots to have no connexion with the Taxeae, is Stachyotaxus*. Tazxites should be retained for fertile branches or reproductive organs which there is some good reason for believing to be more closely related to Taxus than to any other genus. So far as I am aware, none of the specimens described as species of Taxites supply sufficient justifica- tion for being so designated. Fragments of sterile shoots from the Miocene beds of Western Greenland described by Heer® as Taxites Olriki and now in the Dublin Museum afford no proof of close affinity to Taxus: the leaves are rather larger than those of Sequotites Langsdorfii and not decurrent. The generic names Cephalotaxites and Cephalotaxopsis applied by Heer® and Fontaine’ respectively to foliage-shoots resembling those of recent species of Cephalotarus suggest an affinity that is not supported by data based on reproductive shoots. While such a designation as Cephalotaxopsis may be used without a definite implication of relationship to the recent genus, Cephalotazites, like Torreyites and Tazxites, is more appropriately restricted to 1 Halle (13) p. 83. * See page 426. 3 See page 432. 4 See page 410. > Heer (68) p. 95, Pl. 1. figs. 21—24; Pl. xxv. fig. 1. ® Ibid. (83), p. LU. * Fontaine (89) B. p. 235. XLIX] TORREYITES 419 specimens which there is good reason for connecting with Torreya and Taxus respectively. Among the numerous specimens of wood from Mesozoic and Tertiary beds referred by authors to Taxoxylon! there are very few which show true spiral bands on the secondary-xylem elements: there are-no undoubted examples of the Taxineous type of stem | from pre-Tertiarv strata. The foliage-shoots described as species of Tamites are as a rule valueless as records of Taxus, Cephalotavus, or Torreya. Some small seeds very like those of Taxus baccata are described by Ludwig from the Oligocene lignites of West Germany as Taxus margaritifera® and similar examples were described by Heer® as Carpolithes nitens from the lignites of Bovey Tracey: the Bovey seeds have recently been named by Mr and Mrs Reid Taxus (?) nitens*, The latter authors refer some globose mucronate seeds from the preglacial deposits of Norfolk to Taxus baccata®, and Clement Reid® records the same species from preglacial beds in ’ Suffolk, from interglacial beds in Suffolk and from the peat below sea-level in the Thames valley. The genus Taxus ranges through parts of Europe, Asia, North America, Algeria, and occurs sporadi- cally on the mountains of Sumatra, Celebes and the Philippines; it is noteworthy that there is little difference between the several species which are probably mere geographical forms’. The Yew is still wild in parts of Sussex, Hampshire, and Wiltshire, in a few localities in Scotland, but still rarer in Ireland. It is clear from the Pleistocene records that Taxus was formerly much more widely spread. Dr Conwentz® has shown that many places in Germany and the British Isles derive their names from the Yew, and the same author found that several prehistoric wooden articles in the Dublin Museum are made from Yew wood. TORREYITES. The evidence on which several fossil Coniferous branches from Cretaceous and Tertiary rocks have been referred to Torreya is in 1 See page 202. 2 Ludwig (61) p. 73, Pl. ux. fig. 19. 3 Heer (62) p. 1078, Pl. uxx. figs. 15—23. 4C. and E. M. Reid (10) p. 172. 5 Ibid. p. 171, Pl. xvz. figs. 42, 43; (08) Pl. xv. fig. 145. 6 Reid (99) B. p. 151. 7 Elwes and Henry (06) p. 99. 8 Conwentz (01). 27—2 420 TAXINEAE [cH. most cases unconvincing and with one exception no facts as to epidermal characters are available. Leaves of recent species of Torreya (fig. 694, B, p. 141) are characterised by two well marked stomatal grooves on the lower surface, and another feature is the absence of a prominent midrib: the leaves of Cephalotaxus, similar in form and size to those of Torreya, differ in the flat wngrooved — lower surface and the prominence of the midrib on the upper surface of the lamina. We have no information with regard to any fossil seeds of the Torreya type, a type to which reference is made in the account of fossil Palaeozoic seeds. The present distribution of Torreya-suggests that it was formerly more widely spread, but the data at present available do not admit of any very satisfactory statement of its past history. Torreyites carolianus (Berry). Berry described this species as Tumion carolianum}, using the unfamiliar generic name which has been substituted by purists in nomenclature for Torreya. The material from Middle Cretaceous rocks in North Carolina consists of twigs with spirally disposed flat linear-lanceolate leaves 2-5—3 cm. long and with a maximum breadth of 3 mm., gradually tapering towards the slender apex and slightly contracted at the decurrent base. There is no distinct midrib, but in the proximal part of the lamina a more opaque band indicates the position of the vascular tissue: on either side of the middle line is a band in which the stomata are scattered; the long axis of the guard-cells tends to be at right-angles to the length of the leaf as in recent species and the fossil stomata gener- ally resemble those of existing types. Some less satisfactory specimens from Upper Cretaceous beds in Georgia are doubtfully referred by Berry? to this species. In view of the characters of the vegetative fragments from Carolina it seems reasonable to adopt the generic name Torreyites. Berry has published a map showing the distribution of Cretaceous representatives of Torreya, but it is questionable whether the nature of the records constitutes a solid foundation. Heer® has described two species, Torreya Dicksoniana and 1 Berry (08?). ® Ibid. (14) pp. 107, 123. 3 Heer (75) ii. p. 70, Pl. xva. figs. 1—4; p. 71, Pl. xvu. figs. 1, 2; (82) i. Pl. x. fig. 11. XLIX | TORREYITES 421 T. parvifolia, from the Lower Cretaceous beds of Greenland: the leaves of the former exhibit a close agreement with those of the recent species though the evidence in support of generic identity is far from decisive. The second species differs in the smaller leaves: an examination of one of the figured specimens in the Stockholm Museum led me to the conclusion that it may be identical with an Upper Jurassic form from Scotland described as Tazites Jeffreyi1: there are no adequate grounds for the use of the name. Torreya. Fontaine’s Potomac species Torreya virginica? is founded on a piece of shoot bearing long linear leaves with no obvious midrib but with two strong lines between the middle and the edges of the lamina which suggest stomatal grooves. This specimen is of less value than the type-specimen of Torreyites carolinianus because of the lack of information with regard to the stomata. A second species of very little botanical value is described from the same formation as Torreya falcata’. Yokoyama’s Upper Jurassic or Wealden species Torreya venusta* from Japan has no claim to be included among records of Torreya. The Miocene species from Greenland, Torreya borealis, described by Heer® is founded on sterile twigs with broad linear leaves which afford no definite indication of relationship to the recent genus. Some fragments from Pliocene beds near Lyon described by Saporta and Marion® as Torreya nucifera var. brevifolia show two stomatal grooves and may be correctly identified. These authors regard the specimens described by Ettingshausen from Bilin in Bohemia as Sequoia Langsdorfi as fragments of a Torreya and re-name the species 7’. bilinica. Some detached leaves and seeds from Upper Pliocene beds in the Main Valley (Frankfurt) are referred by Engelhardt and Kinkelin? to Torreya nucifera fossilis: the seeds bear a close resemblance to those of the existing species. While there is fairly good evidence from Tertiary localities of the comparatively recent occurrence of Torreya in Europe the records cannot be regarded as conclusive. 1 Seward (11?) p. 688. 2 Fontaine (89) B. p. 234, Pl. crx. fig. 8. Ibid. p. 235, Pl. oxr. fig. 4. 4 Yokoyama (89) p. 230, Pl. xxu. figs. 11, 12. Heer (83) p. 56, Pl. Lxx, fig. 7a. Saporta and Marion (76) p. 87, Pl. xxu. figs. 6, 7. Engelhardt and Kinkelin (08) p. 191, Pl. xxi. figs. 6—8. ao ow to 422 TAXINEAE [cH. VESQUIA. Bertrand. Vesquiu tournaisii Bertrand. The name Vesquia, after the French botanist Julien Vesque, was given by Bertrand? to seeds from Lower Cretaceous strata at Tournai which he described as intermediate in certain respects between Taxus and Torreya; they are two or three times as large as the seeds of Taxus and about one-third the size of those of Torreya. The ligneous shell is ribbed and on each side at the base is a large orifice marking the position of a vascular bundle which is continued through the length of the marginal ribs. The seeds are elliptical in transverse section and prolonged apically into a micropylar beak agreeing structurally with the micropyle of Taxus and Torreya. Bertrand also found anatomical features in the shell similar to those of the recent genera. In the absence of illustrations it is difficult to follow the description in detail, but the facts appear to favour Bertrand’s conclusions with regard to the affinities of the fossil species. CEPHALOTAXOPSIS. Fontaine. Fontaine? gave this name to specimens of vegetative shoots, abundant in the Patuxent formation in the Potomac group, closely resembling in habit recent species of Cephalotaxus: the characters of the genus have been revised by Berry? who adds some particulars as to the structure of the epidermis. Fontaine’s four species are reduced by Berry to two. No seeds have been found attached to the branches, but the American authors consider that some associated seeds may belong to the genus. Cephalotaxopsis magnifolia Fontaine. With this species Berry* includes C. ramosa Font. The branches are fairly robust and in some cases bear lateral shoots in whorls or pseudo-whorls; the leaves are distichous, linear-lanceolate, rather abruptly rounded at the base and tapering gradually to a mucronate apex, with an average length of 4—5 cm. and a breadth of 3—4 mm. Groups of bud-scale scars occasionally occur at the base of an ultimate shoot. The thick lamina may be transversely wrinkled as in Petyophyllum. There is a distinct midrib and a short distance 1 Bertrand (83). 2 Fontaine (89) B. p. 235. 3 Berry (11) p. 374. 4 Ibid. p. 377, Pl. Lx. fig. 1; Fontaine loc. cit. Pls. crv.—cvmi. XLIx] CEPHALOTAXOPSIS 423 on either side of it is a stomatal groove in which stomata are irregularly scattered; the orientation of the guard-cells though not constant tends to be parallel to the long axis of the leaf. The epidermal cells are thick-walled and quadrangular or hexagonal and arranged in regular rows. Berry states that the stomatal grooves are a prominent feature and that there is some evidence of the occurrence of woolly hairs, characters suggestive of Torreya rather than Cephalotazus. Branches from the Potomac beds with shorter leaves are referred to Cephalotaxopsis brevifolia in which is included C. microphylla Font. Berry points out that the _ photograph of C. brevifolia which he gives serves to ‘emphasize the idealisation and inaccuracy of the former figures of this plant.’ Such evidence as is available favours the comparison of these species with recent Taxineae, but the structure of the leaves of C. magnifolia, so far as it is indicated in the epidermal preparations described by Berry, would seem to be in favour of a closer affinity to Torreya than to Cephalotarus. The presence of depressed stomatal regions is a characteristic feature of Torreya and not of Cephalotazus. Heer described a specimen from the Lower Cretaceous beds of Greenland as Cephalotaxites insignis consisting of a small piece of foliage-shoot with, apparently attached to it, an oval seed (18 x 13 mm.) in the form of a mould. Without examining the type-specimen it would be rash to accept the determination as correct. Berry? has described some seeds from Mid-Cretaceous rocks in Carolina as Cephalotaxospermum carolinianum, approxi- mately 10 by 8 mm. in size, ovoid acuminate, and resembling the seeds of Cephalotaxus, but no anatomical features are preserved. Saporta’s species Cephalotaxzus europaea® founded on sterile shoots from the Aquitanian of Manosque in the South of France affords: no convincing evidence of generic identity with Cephalotazus. Seeds from Upper Pliocene beds in the Main valley are referred by Engelhardt and Kinkelin4 to three species of Cephalotaxus, the type-species being C. francofurtana. They bear a close resemblance to the recent seeds but this is hardly sufficient to prove their generic identity. 1 Heer (83) p. 10, Pl. xu. fig. 12. 2 Berry (101). 3 Saporta (93) p. 42, Pl. v. fig. 4. 4 Engelhardt and Kinkelin (08) p. 194, Pl. xxmu. fig. 11. CHAPTER L., CONIFERALES INCERTAE SEDIS. TRIOOLEPIS. Zeiller. Trioolepis Leclerei Zeiller. This generic name was proposed by Zeiller for a cone from the Rhaetic flora of Tonkin! which he at first placed in the comprehensive genus Conites. In general appearance the specimen resembles a cone of a Picea; it is elongate- oval and incomplete, more than 10cm. long and about 3 cm. in diameter. The impression shows numerous spirally disposed, imbricate, scales apparently thin, oval-linear in form, 12—15 mm. long and 6—7 mm. broad, suddenly contracted to an obtuse apex; the surface is marked by more or less distinct longitudinal folds and ‘close to the base are slight depressions indicating the former presence of seeds 5mm. long and 15mm. broad. On some of the scales there is a faint curved trilobed line in the upper third of the ventral face which, it is suggested, may possibly mark the limit of an ovuliferous scale fused to a subtending bract-scale, but there is no substantial ground for any conclusion as to the morpho- logical nature of the cone-scales. Zeiller states that there is no decisive evidence with regard to the systematic position of this fossil: the presence of three seeds recalls Cunninghama though this in itself is probably of comparatively small importance. MASCULOSTROBUS. Seward. This designation was proposed for fossils which are in all probability male strobili of Gymnosperms?. Masculostrobus Zeilleri Seward. This, the type-species, was discovered by the late Dr Gunn in Kimeridgian strata on the North-East of Scotland: it consists of a slender axis 13cm. long bearing numerous small branches with * Zeiller (03) B. p. 208, Pl. u. fig. 15; Pl. F, fig. 2. 2 Seward (112) p. 686. CH. L] MASCULOSTROBUS 425 Fic. 799. Masculostrobus. A, M. Zeilleri, nat. size; a, b, portions enlarged; c, microspores from A; d, Masculostrobus sp.; e, spores from d. (British Museum.) 426 CONIFERALES INCERTAE SEDIS [cH. spirally disposed sporophylls. A few oval spores (fig. 799, c), 20—30 in diameter, were obtained from some of the sporophylls. The specimen is probably an inflorescence of male flowers of a Conifer; it bears some resemblance to Zeiller’s Pseudoasterophyllites Vidal! from the Kimeridgian of Spain, but in that type the leaves are in whorls and the shoot appears to be sterile. The smaller example shown in fig. 799, d, e, may be part of a microstrobilus of Elatides curvifolia, but in the absence of any connexion with that species it should be retained in Masculostrobus. Méller and Halle? in their account of a flora from South-East Scania, probably of Wealden age, describe some fragments of microstrobili which they assign to Masculostrobus. The Cretaceous specimens from Kreischerville described by Hollick and Jeffrey as Strobilites microsporophorus® may be transferred to Masculostrobus. This species, represented by portions of small strobili a few milli- metres long, is characterised by sporophylls in which the leaf- traces are enclosed by transfusion-tracheids, a feature suggesting comparison with modern Araucarineae, and the spores have two bladder-like wings. It may well be, as the authors of this type suggest, that the extension of the exine of the microspores, now restricted to the Abietineae and Podocarpineae, may be an ancient character and formerly more widely spread among the Coniferales. PALISSYA. Endlicher. Endlicher? instituted this genus for a type of Conifer previously described by Braun® from the Rhaetic flora of Franconia as Cunninghamites sphenolepis, characterised by distichous, linear, leaves and a megastrobilus composed of loosely imbricate scales. The name Palissya was selected to commemorate Bernard Palissy. The type-species, P. Brawnii, was first figured by Goeppert® and described in more detail by Schenk’ who, as Nathorst points out, included two distinct types of reproductive shoot under the name Palissya: one of them is retained in that genus while the other agrees with cones referred to Elatedes®’. Endlicher’s designation 1 Zeiller and Vidal (02) p. 7, Pl. a. 2 Moller and Halle (13) p. 36, Pl. v1 figs. 9—18 3 Hollick and Jeffrey (09) B. p. 66. + Endlicher (47) p. 306. 5 Braun (43). ° Goeppert (50) Pl. xivmt. 7 Schenk (67) A. p. 175, Pl. x1. figs. 2—14. 8 Ibid. fig. 7. L] , PALISSYA 427 has been employed by several authors for Mesozoic shoots with linear leaves unaccompanied by reproductive organs and having therefore no claim to be assigned to a genus characterised by a well defined type of strobilus. Further reference to the mis- application of Palissya is made in the account of some Indian specimens recently assigned to Retinosporites. In the absence of any evidence of the occurrence of strobili, shoots similar in habit to Palissya should be described under Halle’s generic name Elatocladus. An impression of a shoot from Triassic beds in Bucks County, Pennsylvania, recently described by Wherry! as Palissya longifolia, would be more appropriately referred to Elato- cladus as it affords no indication of the nature of the fertile branches. Palissya sphenolepis (Braun). ‘ _ Nathorst? in his recent and able account of this species reverts to the older specific name on the ground that Endlicher’s name P. Braunii is not in accordance with the laws of priority. It is possible though not certain that some specimens described under different names by Pres] prior to the publication of Braun’s account of Cunninghamites sphenolepis may belong to this species. Schenk considerably extended Braun’s description and was the first to publish figures of ripe cones showing certain morphological features of the seed-bearing scales which authors have differently interpreted. Palissya sphenolepis is a Rhaetic species recorded from Franconia and Scania, possibly more closely allied to the genus Cunninghamia than to any other existing Conifer; but, as Nathorst suggests, it may belong to an extinct section of Gymnosperms. The foliage- shoots bear spirally disposed, two-ranked, leaves of the Tamites form; the lamina is narrow and linear with a median vein, decurrent on the axis as a persistent leaf-cushion. The epidermal cells have straight walls and the stomata, confined to the lower surface, occur in two rows. Palissya cannot be identified with any degree of certainty in the absence of well-preserved strobili. The mega-_ strobili are cylindrical and relatively narrow; in an immature con-- dition they closely resemble those of Elatides, the surface being formed of the lanceolate, imbricate, distal ends of crowded cone- scales. It is the older strobili with elongated internodes that 1 Wherry (16). 2 Nathorst (087). 428 CONIFERALES INCERTAE SEDIS (cH. constitute the most striking feature of the genus: the cone-scales (sporophylls) are entire, elongate-lanceolate (fig. 800) with an acuminate apex, and each bears 5—6 pairs of seeds characterised by a cup-like basal investment or cupule (fig. 800,C). The sporophylls have a strongly developed keel on the lower surface and a less distinct median rib between the two rows of seeds on the upper face (fig. 800, A, B). There is no evidence to support the view that the cone-scales are doublet. Schenk? described the cone-scales as bearing 10—12 seeds on the edge and Saporta? believed the seeds to be lobes of a seminiferous scale, each lobe supporting one seed. Nathorst’s in- vestigation of Scanian material has thrown a welcome light on the nature of the mega- strobili as interpreted by previous authors. The bodies described by Schenk as seeds are projecting spherical casts of cup-like organs which originally embraced the lower portions of the seeds. The morphological nature of the cupule cannot be determined, but as Nathorst suggests it may correspond to the epimatium?* which partially encloses the seeds of Dacrydiwm and other recent Fig. 800. Palissya spheno- lepis. A, B, sporophyll in surface-view and in section. C, cupules of two seeds. (After Nathorst.) Conifers. Nathorst compares the cone-scales of Palissya with those of the genus Cunninghamia (fig. 684, K, p. 116) in which each sporo- phyll bears three seeds on the adaxial side of a membranous out- growth stretched across the scale. The resemblance would seem to be closer than Nathorst suspects as the membrane in Cunning- hamia does not arise as a continuous strip of tissue but as three separate ligule-like pieces, one on the abaxial side of each seed. 1 Solms-Laubach (91) A. p. 73. 2 Schenk in Schimper and Schenk (90) A. p. 336. 3 Saporta (84) p. 513. 4 See page 118. L] ELATOCLADUS 429 ELATOCLADUS. Halle. This genus was founded with a view to reduce the ‘present state of intolerable confusion in the classification of the Conifers?.’ Halle expresses the opinion, with which most botanists cannot but agree, that sterile shoots of Conifers should not be described under generic names which imply the possession of a certain type of cone. The occurrence of foliage-shoots of similar or even identical habit in certain recent genera possessing distinct types of repro- ductive shoots serves to emphasise the unscientific character of the too common practice of assigning fossils to genera distinguished by a particular form of cone even though the specimens in question afford no evidence of the nature of the fertile branches. The generic name Taxites has been widely employed for dorsiventral branch-fragments bearing linear leaves with a pseudo-distichous arrangement as in the recent Taxus, but notwithstanding the employment of Taxites by authors who do not intend to convey the impression of relationship to Tazxus, it is preferable to reserve Taxites for specimens which there are reasonable grounds for believing to be related to the Taxeae. Halle insists that it is undesirable to use one name for dorsiventral shoots and another for shoots with spirally disposed leaves, as fossil forms are known, e.g. Stachyotaxus elegans, Nathorst, in which both types: occur on the same plant. Similar cases of dimorphism are well illustrated by Dacrydium and other recent Conifers (cf. fig. 708, p. 160). Entocladus is proposed for sterile coniferous branches of the radial or dorsiventral type, ‘which do not show any characters that permit them to be included in one of the genera instituted for more peculiar forms.’ The name serves a useful purpose for sterile shoots which it has been the custom to include in Tazites and for types such as Elatocladus heterophylla Halle, which bear both distichous, linear leaves and crowded scale-like leaves similar to those of Brachy- phyllum and some forms referred to Pagiophyllum. It is, however, desirable to retain Brachyphyllum and Pagiophyllum for sterile shoots exhibiting no well marked dimorphism and bearing fleshy appressed leaves and four-sided falcate leaves respectively. Used 1 Halle (132) p. 82. éAdrq, Pine or Fir; «Addos, shoot. 430 CONIFERALES INCERTAE SEDIS _ [oH. in this narrower sense Elatocladus is more likely to serve the object which the author had in view. It is noteworthy that in some specimens of Elatocladus (Tasites spp.) the leaves have a transversely wrinkled lamina, a feature usually associated with the detached linear leaves assigned to the genus Pityophyllum. Elatocladus heterophylla Halle. The shoots of this species from the Jurassic flora of Graham Land! are freely branched and the ultimate branches show a Fic. 801. Elatocladus heterophylla. (After Halle; nat. size.) tendency towards a distichous arrangement. The radially sym- metrical branches bear short and thick, acute, more or less appressed leaves or the lamina may be slightly falcate, while the dorsiventral shoots have narrow linear acute leaves in two ranks (fig. 801). A faintly marked midrib is present in both forms of leaf. Halle states that the shorter leaves differ from those of Brachyphyllum in being rather longer and less closely appressed to the axis, though this is a difference of secondary importance. The main interest of the species is its dimorphism. In Elatocladus is also included the Indian species, recorded by 1 Halle (132) p. 84, Pl. va. text-fig. 18. L] ELATOCLADUS 431 Halle? from Graham Land, originally described by Oldham and Morris as Cunninghamites confertus and subsequently removed by Feistmantel to Palissya: the leaves are distichous with a sessile and decurrent lamina attached at a wide angle. The apex of the leaves is obtuse and a midrib is present. In Elatocladus conferta Halle in- cludes the Australian form Palissya australis as figured by Stirling?. Elatocladus zamioides (Leckenby ex Bean ms.). This type, from the Middle Jurassic of Yorkshire, was described by Leckenby? as Cycadites zamioides. and subsequently transferred » fae th irs Fic. 802. Elatocladus plana. (Specimens figured by Feistmantel as Tazites planus; nat. size.) to Taxitest. The type-specimen, in the Sedgwick Museum, Cam- bridge, consists of a slender axis bearing two-ranked spirally 1 Halle (132) p. 86, Pl. vit. 2 Stirling (00) Pl. mu. figs. 8, 9. 3 Leckenby (64) A. p. 77, Pl. vit. fig. 1. 4 Seward (00) B. p. 300, Pl. x. fig. 5. 432 CONIFERALES INCERTAE SEDIS [cH. attached linear leaves slightly more than 2 cm. long and 1-8 mm, broad, basally contracted, with an acute apex. It closely resembles shoots of recent Taxeae and Sequoia sempervirens, also some Potomac species included by Fontaine? in Cephalotaxopsis though there is no evidence of relationship to the recent Cephalotarus. Shoots of similar habit are figured by Feistmantel2 from Indian Jurassic beds as Taxites planus but the lamina is not contracted at the base. Two of Feistmantel’s figured specimens are reproduced in fig. 802. These afford good examples of fossil branches which it has been the custom to refer to Taxctes, but without information with regard to the epidermal characters it is impossible to determine their affinities. The form of the leaf-bases agrees with that shown in fig. 803 and it is probable that Taxites planus may be another example of Miss Holden’s genus Retinosporites, though in the absence of anatomical data Elatocladus is the more appropriate designation. RETINOSPORITES. Holden. Feistmantel used the name Palissya for some Indian Jurassic vegetative coniferous shoots which afford no evidence of affinity to that genus as represented by P. Braunii. Some of his fossils may be identical with the British species Taxites zamioides, now assigned to Elatocladus, while the examples described by Feist- mantel as Palissya sp. and Palissya indica have been transferred to a new genus Retinosporites. The Indian impressions afford no evidence of a midrib; the upper epidermis consists of cells with straight walls and there are no stomata, while on the lower face of the lamina stomata are irregularly scattered, the long axis of the guard-cells being more or less parallel to the margin of the leaf. The absence of a midrib, at least so far as regards-impressions and cuticular preparations, led Miss Holden? to separate the Indian specimens from Palissya and Tazites as vegetative shoots included by authors in genera having leaves with a distinct median vein and in which the stomata are in rows on the lower surface. The generic name Retinosporites, spelt by Miss Holden Retinosporitis, is proposed on the ground that the only flat-leaved Conifers among 1 Fontaine (89) B. Pls. cv1.—cv1u. 2 Feistmantel (79) Pls. xm.—xv. 3 Holden, R. (15%), L] CUNNINGHAMITES 433 those examined showing similar epidermal characters were certain seedlings of the Retinospora type. Miss Holden recognises that _ Retinosporites suggests relationship to such recent Cupressineous species as are included under the genus Retinospora, but she states that no such implication isintended. The new designation, though not very happily chosen, may be retained for shoots with linear leaves (fig. 803) without a midrib and having the stomatal features described in Miss Holden’s account of R. indica. Retinosporites indica (Oldham and Morris). An Indian species originally described by Oldham and Morris as Taxites indicus and transferred by Feistmantel to Palessya.' The leaves are linear and decurrent (fig. 803, A) and without a midrib. The epidermal cells have straight walls and the stomata, though oc- casionally present on the upper surface, are scattered on the lower epidermis as in the Retinospora foliage of Thuya or Juniperus, without any indication of a median A . B . : Fic. 803. Retinosporites indica. A, piece astomatic region such as one would of shoot; B, stoma.. (After R. Holden.) expect in leaves possessing a mid-- rib. There are generally six accessory cells and the guard-cells are sunk below the level of the epidermis (fig. 803, B). Sterile foliage-shoots formerly referred to the genus Cunninghamites. Pres]! gave the name Cunninghamites to some sterile shoots from Rhaetic and Lower Cretaceous strata on the ground of their resemblance to branches of Cunninghamaa and both his specimens and the large number, particularly from Cretaceous beds, referred by authors to Presl’s genus afford no real evidence of affinity to the recent Conifer. The type-species Cunninghamites oxycedrus, from Lower Cretaceous rocks in Saxony, is probably identical with Corda’s species Cunninghamia elegans (fig. 804) from Lower Creta- * ceous beds in Bohemia. Presl also included in Cunninghamites 1 Sternberg (38) A. Pl. xivum. fig. 3; Goeppert (50) Pl. xiv. 8. IV 28 434 CONIFERALES INCERTAE SEDIS [CH. branches from the Keuper of Germany which he named C. dubius: this species is identified by Saporta? with Palissya Braunu, but the latter name is now restricted to shoots bearing a particular form of strobilus. Nathorst?, on the other hand, suggests that C. dubius may belong to Elatides, and the same author instituted a new generic name Camptophyllum? for fragmentary foliage-shoots Fic. 804. A, Elatocladus elegans? B, Elatocladus Schimperi. (A, after Velenovsky; B, after Nathorst. Nat. size.) from the Rhaetic of Scania which he named C. Schimperi (fig. 804, B): these bear a close resemblance to C. elegans; the linear- lanceolate leaves are 15—20 mm. long and the lamina is recurved, probably as the result of drying. It is impossible to determine the position of this type and it may conveniently be transferred to Elatocladus. Some of Nathorst’s specimens are also figured by Schenk’. The name Cunninghamites is given by Oldham and 1 Saporta (84) p. 511. ? Nathorst (08?) p. 10. 8 Ibid. (78) B. Pl. xv. figs. 13—16. 4 Schenk in Schimper and Schenk (90) A. p. 351, fig. 236. L] ELATOCLADUS 435 Morris! to Jurassic Indian shoots which Feistmantel? afterwards transferred to Palissya: these have recently been included by Halle® in Elatocladus as E. conferta and recorded by him from the rich Antarctic flora of Graham Land. In no case have we any definite information with regard to the cones borne by the Cunninghamites type of shoot. Velenovsky4 figures some Lower Cretaceous cone-scales from Bohemia, which he refers to Cunninghamia, resembling in shape the scales named by Hollick and Jeffrey Protodammara, but the resemblance may be only superficial. The vegetative branches assigned by authors to Cunninghamites have linear-lanceolate leaves usually showing a distinct midrib and often other parallel lines on the lamina which are probably due to hypodermal fibres. The leaves may reach a length of 6 cm. and are 1—4 mm. broad; the edge is entire and finely serrate as in Cunninghamia sinensis. A characteristic feature is the occurrence of persistent decurrent leaf-bases on the branches which in some specimens that have lost the free portion of the lamina present a close similarity to Brachyphyllum. Some of the examples of Cunninghamites may well be shoots of a plant allied to Araucaria Bidwillit or A. brasiliensis. There is no doubt that under Cunning- hamates are included branches of many different Conifers. Elatocladus elegans (Corda). Originally described by Corda® from Lower Cretaceous rocks in Bohemia as Cunninghamua elegans, this species is recorded from many Cretaceous localities; from the Patoot beds in Greenland®, Moravia’, Westphalia®, Upper Cretaceous beds in Bulgaria®, the Amboy clays”, Cliffwood, Martha’s Vineyard", Georgia, Carolina”, and other places in North America®. The specimen shown in fig. 805 from Moravia shows a midrib in a few leaves, while in the 1 Oldham and Morris (63) Pl. xxxu. fig. 10. 2 Feistmantel (767) p. 55. 3 Halle (13?) p. 86, Pl. vir. 4 Velenovsky (87). 5 Corda in Reuss (46) B. Pl. xz1rx. S Heer (83) Pl. um. fig. 1. ? Ibid. (69) Pl. 1. 8 Hosius and von der Marck (80) B. Pl. xxxvu. ® Zeiller (05?) Pl. vu. fig. 14. 10 Newberry and Hollick (95) Pl. v. 1. Hollick (06) Pl. mz. fig. 1. 12 Berry (104) Pl. xx.; (14) p. 106. 13 See Hollick (06) for other references. 28—2 436 CONIFERALES INCERTAE SEDIS (CH. Bohemia specimen represented in fig. 804 the midrib is more obvious and the leaf-bases have a more regular form. The branch- ing is sparse and not pinnate. Velenovsky? assigns some branches to C. stenophylla but these may be younger forms of C. elegans. Similarly C. squamosus®, as figured by Heer and other authors, affords no satisfactory evidence of specific difference from C. elegans. Impressions from the Atane beds of Greenland described by Heer? as C. borealis have been compared by Schenk with Sequoza and also referred by him to Torreya: there is no possibility of deciding the precise systematic position of these and similar specimens. Ettings- hausen* has described as Cunninghamites miocenica fragments of shoots from Sagor in Carinthia bearing linear leaves with a finely serrate edge. ANDROVETTIA. Hollick and Jeffrey. This genus was instituted® for Cretaceous fossils from Staten Island superficially resembling Fern leaves with a pinnate venation and an irregularly lobed or incised margin. The leaf-like fragments are, however, stem-structures bearing minute scale-like leaves attached to the edges and surface. In habit these phylloclades agree with Phyllocladus, but on anatomical grounds the authors of the genus regard it as Araucarian though the evidence is far from convincing. Androvettia statenensis Hollick and Jeffrey. Some of the specimens show no indication of their phylloclade- nature and, as impressions, would be identified as Fern pinnules or referred to Thinnfeldia. Others, after bleaching in chlorine- water, showed a fairly stout vascular axis giving off simple or forked branches at an acute angle; small decurrent leaves free only at the apex occur on the margins of the shoots (fig. 806, A, B). In a few cases the phylloclades bear short axillary branches with immature cones, possibly microstrobili. There are three vascular cylinders in the section reproduced in fig. 806, C, and in the narrow wings of the ‘lamina’ there are the 1 Velenovsky (85) B. p. 15. * Heer (71*) Pl. 1. figs. 5—7; Schimper and Schenk (90) A. p. 282; Berry (03) p. 64. ® Heer (82) B, Pl. xxrx. fig. 12. 4 Ettingshausen (72) Pl. 1. fig. 30. ° Hollick and Jeffrey (09) B. p. 22, Pls. m., viz, VII., XXVIUI., XXIX. . L] ANDROVETTIA 437 traces of two or three leaves. Several stomata occur on the sur- face of the phylloclade, each surrounded by 4—5 accessory cells. Sclerotic cells are present in the pith. The secondary xylem is of the coniferous type and the uniseriate bordered pits on the tracheids may be either separate and circular or flattened by contact. No resin-cells, such as occur in the wood of Phyllocladus, were recog- — nised. The medullary rays are not described. The data are Fic. 806. A—C, Androvettia statenensis. D, Geinitzia Reichenbachii. (A, B, x 6; C, x7; D, x 7; after Fic. 805. Hlatocladus Hollick and Jeffrey.) elegans. | (Nat. size; ae after Heer.) hardly sufficient to justify the inclusion of this type in the Araucarineae: the occasional flattening of the tracheal pits and the absence of resin-cells are not fatal to an alliance to Phyllocladus. The precise position of the genus within the Coniferales must for the present be left in doubt. Androvettia elegans Berry. This species, from the Upper Cretaceous of Georgia, is repre-~ sented by dorsiventral fern-like vegetative shoots which, as Berry? 1 Berry (14) p. 103, Pl. xvut. 438 CONIFERALES INCERTAE SEDIS [CH.. . points out, bear a close resemblance to species of Moricoma (cf. fig. 760). The branches are distichously arranged and the opposite lateral leaves are stout, falcate, and decurrent while those on the upper and lower faces are represented by scales on the middle line of the phylloclades. The same author describes specimens from beds in North Carolina! referred to the lower half of the Upper Cretaceous as Androvettia carolinensis. DACTYOLEPIS. Hollick and Jeffrey. Dactyolepis cryptomerioides Hollick and Jeffrey. The generic name? was instituted for some detached, cuneate, cone-scales fr6m the Cretaceous beds at Kreischerville in Staten Island, approxi- mately 4mm. long, composed of an upper and a lower segment. The upper portion is divided distally into as many as seven irregular short finger-like processes and the lower part is entire. Each of the processes possesses a single vascular bundle ‘completely surrounded by a cordon of transfusion-tissue, thus betraying its Araucarineous relationship.’ The scales which are without seeds are compared with those of Voltzia. There is, however, no proof that Voltzia had double scales. The view that Dactyolepis is Araucarian may fairly be said to rest on an insufficient basis. RARITANIA. Hollick and Jefirey. The name Raritania®, after the Raritan formation, was given to some Cretaceous fossils from Kreischerville identical with New Jersey specimens described by Newberry as Frenelopsis gracilis* on the ground that they belong to a type distinct from Frenelopsis as generally understood. . Raritania gracilis (Newberry). The specimens so named con- sist of slender, dichotomously branched, axes bearing minute leaves resembling Psilotum triquetrum and in the form of the branching the leaves of Batera Lindleyana (Schimp.). The dis- | tinguishing feature is the occurrence of the prickle-like leaves (fig. 807, B) invisible to the unaided eye (fig. 807, A). A small 1 Berry (10%) p. 183, PL. xrx. * Hollick and Jeffrey (09) B. p. 52, Pl. x. figs. 12, 13. 3 Ibid. (09) B. p. 26, Pls. vi., 1x., x., Xx. * Newberry and Hollick (95) p. 59. Pl. xm. figs. 1—3 a. L] RARITANIA; SCHIZOLEPIS 439 imperfectly preserved cone was found on a peduncle having leaves similar to those on the vegetative twigs. Some fragmentary lignitic branches (fig. 807, C) associated with the impressions showed the anatomical characters of a Conifer; but Hollick and Jeffrey, though believing that the fragments ‘almost certainly’ / \ 7 ft WW 8 & Fic. 807. A, B, Raritania gracilis; C, Raritania?. (After Hollick and Jeffrey. A, B, x 6; C, x 10.) belong to Raritania, admit that there is no proof of their identity with the dichotomously branched impressions. The secondary xylem of the cylindrical stele of the twigs, one of which is shown in fig. 807, C, agrees with that of Brachyphyllum macrocarpum Newb. The uniseriate bordered pits are occasionally flattened; the cortex, confluent with the decurrent leaves, contains sclerotic cells and each leaf has a resin-canal subtending the leaf-trace. The genus is referred to the Araucarineae on the evidence of the occurrence of flattened pits at the ends of some of the tracheids, a conclusion difficult to accept without considerable reservation. SCHIZOLEPIS. Braun. Braun! instituted this genus for a strobilus from Rhaetic rocks in Germany which he called Schizolepis liaso-keuperinus, 1 Braun (47) p. 86. 440 CONIFERALES INCERTAE SEDIS [cH. characterised by its deeply split cone-scales. Schenk? subsequently substituted the name S. Brawnit and included under that designa- tion Braun’s Isoetites pumilus, a species founded on a foliage-shoot, also some other similar vegetative branches believed to belong to the plant which bore the cones. As here used, the term Schizolepis is restricted to cones and cone-scales since there is no definite evidence as to the nature of the foliage-shoots connected with the strobili. Schizolepis cannot be referred on any satis- factory grounds to a definite position among the Coniferales: it is possibly an extinct type allied to recent Abietineae, but until more is known with regard to the morphology of the cone-scales the systematic position must be left an open question. The genus is represented by strobili from Rhaetic beds in Franconia, Scania, and Poland; detached scales from Middle Jurassic floras are also included in Schizolepis (fig. 808), and Nathorst has described incomplete strobili from Upper Jurassic or Wealden strata in Spitzbergen. Attention has been called to a resemblance between Schizolepis scales and the fertile leaves of Tmesipteris?, but there is no reason for regarding this as indicative of relationship. More than one author has compared the bilobed cone-scales of Schizolepis with the 3—5-lobed scales of Voltzia and Cheirolepis though this comparison rests on a feature which in itself is no proof of affinity. A comparison may also be suggested with the reflexed cone-scales of Picea Breweriana. Schizolepis Braunit Schenk. It has already been pointed out that under this name Schenk® included both cones and vegetative shoots though he recognised the lack of any decisive evidence of common parentage. While agreeing with Nathorst that the association with Schizolepis strobili of similar vegetative shoots both in Germany and Sweden may be more than accidental, in the present state of our knowledge it is preferable to refer the leaves and branches to Pityophyllum or Pityocladus. In the younger strobili the bilobed scales are more or less pressed against the axis and in older examples they are more spreading: each scale has two lanceolate lobes and is attached l Schenk (67) A. p. 179. ? Nathorst (97) p. 61. * Schenk (67) A. p. 179, Pl. xx1v.; Schimper and Schenk (90) A. p. 306. L] SCHIZOLEPIS 441 by a narrow stalk-like basal portion (fig. 808, B). Schenk states that there are-two anatropous seeds to each scale, but it is not clear if the actual seeds are present. Schizolepis Follint Nathorst. In this species from the Rhaetic flora of Scania! the bilobed scales are sessile and broader than in S. Braunii. In some specimens there appear to be two seeds near the base of a scale. Saporta? regards the cone-scales of the type-species and S. Follini as double structures, the trans- verse line shown in Schenk’s figures below the forking of the scale being the limit of the bract-scale, while the thinner distal lobed part represents the seminiferous scale; an interpretation which rests on very slender evidence. To this species Fre.808, Schizolepis. A, Sehé- Raciborski? refers a specimen from rocks zolepis Moelleri. B, Schizo- possibly of Rhaetic age in Poland. lepis Braunit. C, Schizo- . . ; lepis(?) retroflexa. (A, after Schizolepis Moellers Seward. Seward; B, after Schenk; Under this name* some detached bi- eee enlarged, “alter lobed scales without seeds were described from Jurassic rocks in Turkestan (fig. 808, A) and the South of Russia®, and Krystofovié has recently discovered a cone of the same species in Jurassic strata in Transbaikalia®. Schizolepis cylindrica Nathorst. Founded on a long and narrow, incomplete, strobilus from Upper Jurassic rocks in Spitzbergen’ bearing deeply lobed scales. A second species, S. retroflera® (fig. 808, C), assigned with some 1 Nathorst (78) B. p. 28, Pls. xIv., xv. 2 Saporta (84) p. 502. 3 Raciborski (92) p. 354, Pl. m. figs. 1, 20d. 4 Seward (072) p. 36, Pl. viz. figs. 64—66. 5 Thomas (11) p. 79, Pl. v. fig. 4. & Krystofovité (15) p. 95, Pl. v1. figs. 10, 11. 7 Nathorst (97) p. 39, Pl. 1. figs. 1, 2. 8 Ibid. p. 60, Pl. m1. figs. 31, 32; Pl. vz. figs. 11, 12. 442 , CONIFERALES INCERTAE SEDIS [cH. doubt to Schizolepis, is characterised by pendulous stalked scales: in neither of these Spitzbergen forms are there any seeds on the strobili. Nathorst quotes a species from Portugal described by Saporta? as Palaeolepis bicornuta as being possibly allied to Schizolepis, but there are not sufficient grounds for assuming any close affinity. He also draws attention to the resemblance of the bilobed leaves or scales of Schizolepis(?) retroflexa to the fertile leaves of T'mesipteris; the relationship of the Spitzbergen fossils must be left uncertain pending more evidence. DREPANOLEPIS. Nathorst. Heer described a specimen of Jurassic age from Spitzbergen as Phyllocladites rotundifolia® which he considered to be closely allied to Phyllocladus: an examination of the type-specimen led Nathorst? to institute a new generic name Drepanolepis. As Nathorst’s revised description and more accurate drawing show, there are no substantial grounds for assuming any relationship between the fossil and Phyllocladus. Drepanolepis rotundifolia consists of a fairly stout axis bearing spirally disposed thick, falcate scales each of which bore a seed, or possibly a sporangium, near the base (fig. 798, C). A similar type is described by Nathorst as Drepanolepis angustior* characterised by the narrower form of the scales and a broader axis. Both species may be described as strobili of open habit with single-seeded sporophylls: it is impossible to determine the systematic position of the genus, though as Nathorst says it is probably a type of fertile Gymnospermous shoot. There is no reason for comparing the specimens with Phyllocladus. SCHIZOLEPIDELLA. Halle. Schizolepidella gracilis Halle. The specimens on which this genus is founded are from the Hope Bay flora in Graham Land, probably of Middle Jurassic age; they consist of slender sterile shoots reaching a maximum length of 12cm. and 2 mm. broad, rarely branched and bearing small leaves, 2 x 1-5 mm., apparently } Saporta (94) B. Pl. xxxumm. fig. 4. * Heer (75) ii. p. 124, Pl. xxxv. figs. 17, 18. 3 Nathorst (97) p. 43, Pl. vi. figs. 24, 25. 4 Ibid. p. 71, Pl. m1. figs. 33—37. ® Halle (13?) p. 90, Pl. 1x. figs. 18—21. L} CYPARISSIDIUM 443 spirally disposed (fig. 809). The lamina is rounded, ovate or obovate, and always bilobed at the broad apex: no veins were detected. As Halle says, it is im- possible to determine the affinities of the frag- ments but he thinks they may belong to pen- dulous branches of a Conifer. Attention is called to a resemblance to some Hepaticae, and a pos- sible relationship to: Lycopodium or the Psilotales may also be suggested. The choice of the generic name is not intended to imply anything more than a superficial similarity between the leaves and the bilobed strobilar appendages of Schizo- lepis}. : Fic. 809. Schizolepi- CYPARISSIDIUM. Heer. _ della gracilis. (After Halle; A, enlarged ; This name was given? to foliage-shoots and_B, nat. size.) cones, from the Urgonian rocks of Greenland, originally described as Widdringtonites gracilis?. The smaller sterile branches are indistin- guishable from specimens referred by authors to Widdringtonites while the larger examples might be included in Brachyphyllum. The leaves are small, appressed, and imbricate, similar to those of some recent Cupressineae and Callitrineae but spirally disposed and not verticillate (fig. 810); the shoots agree also with Microcachrys and other recent Conifers. The cones are composed of a small number of flat scales (fig. 810, B) too imperfectly preserved to afford any definite evidence as to the affinities of the genus. Heer states that a detached cone-scale shows the impression of a single seed, but the material is insufficient to form the basis of a comparison with the Araucarineae; he points out a resemblance to Cunning- hamia, and mentions the striated surface of the fossil cone-scales as a distinctive feature, though that may be due, in part at least, to the state of preservation. The flat form of the cone-scales is a character in which Cyparissidiwm differs from genera such as Sequotites, and from the Callitrineae the cones are distinguished by the spiral arrangement of the scales. 1 See page 439. 2 Heer (75) ii. Pl. xv. fig. 5 6, c; Pls. x1x., XX., XXI.; (82) pp. 16, 50, Pls. 1., viz, xXXvm@M. 3 Ibid. (68) p. 83. 444 CONIFERALES INCERTAE SEDIS [cH. Cyparissidium is characteristic of Lower Cretaceous strata though Nathorst! has recorded a species, C. Nilssonianum, from Rhaetic rocks in Scania with cones having scales more pointed and lanceolate than those of Heer’s species. A second Rhaetic species, C. septentrionale, has been transferred by Nathorst to the genus Stachyotaxus?. Fic. 810. Cyparissidium. A, B, Cyparissidium gracile. C, C. minimum. (A, B, after Heer; C, after Velenovsky; nat. size.) Cyparissidium gracile Heer. The type-species (fig. 810, A, B) is one of the most abundant plants in the Kome beds at Pattorfik in West Greenland and it occurs also in the Atane beds. The cones are 26 mm. long and 11—20 mm. broad with scales having a breadth of 12mm., a 1 Nathorst (86) p. 103, Pl. xxi. 2 See page 410. L] BENSTEDTIA 445 rounded distal border, a mucronate apex, and a striated dorsal face. Velenovsky! records this species from Lower Cretaceous rocks in Bohemia but without the essential evidence of cones: the same author describes a smaller type from Bohemia as C. minimum? (fig. 810, C). Schenk® figures some fragments from Lower Creta- ceous rocks in the Tyrol as Cyparissidium cretaceum, but in the absence of cones there is no sufficient reason for their inclusion in Cyparissidium. Hollick’s record of C. gracile from the Cretaceous of Block Island? is based on insufficient evidence. BENSTEDTIA. Seward. In 1862 Mackie® figured a fossil stem from Lower Greensand rocks in Kent which Konig afterwards named Dracaena Benstedtit. This name was adopted by Morris and Mantell for the supposed ‘Dragon tree.’ In 1868 Carruthers® expressed the opinion that the fossils are more likely to be Pandanaceous stems, while Gardner’ spoke of a possible Cycadean affinity. An examination of Mackie’s specimens and others in the British Museum led me to suggest a comparison with stems of recent species of Cycads, particularly stems of Zamia which do not retain the armour of leaf-bases (fig. 381 B, vol. 111. p. 5) characteristic of most Cycadales, and to institute a new generic name Benstedtia® in preference to a designa- tion implying an improbable relationship. The genus is defined as follows: stems characterised by irregular and interrupted grooves and broader ridges running transversely, with occasional small elliptical protuberances irregularly disposed on the surface. There are no distinct leaf-scars but branch-scars occasionally occur; the upward convergence of the transverse wrinklings indi- cates bifurcation in some specimens. The English examples reach a length of over 40cm. and a diameter of 15cm. Smaller specimens with similar surface-cha- racters are described by Fliche® from Lower Cretaceous beds in France as Coniferocaulon colymbeaeforme and compared by him to stems of Araucaria imbricata: other examples closely resembling 1 Velenovsky (85) B. p. 17, Pl. vm. 2 Ibid. Pls. Ix., x. 3 Schenk (76) B. p. 167, Pl. xxix. figs. 10, 11. 4 Hollick (06) p. 46, Pl. m. fig. 11. 5 Mackie (62). ® Carruthers (68) p. 154 (footnote). 7 Gardner (86%) p. 201. ° Seward (967) p. 216. 9 Fliche (00). 446 CONIFERALES INCERTAE SEDIS [cH. L the French stems are recorded from the Uitenhage (Wealden) series of South Africa and the Kimeridge rocks of Sutherland}, Scotland, in the former case as Benstedtia sp., and in the latter as Coniferocaulon colymbeaeforme because of the presence of a small pith more suggestive of a Conifer than of a Cycadean branch. A large specimen from Jurassic rocks in India is described by Miss Bancroft? as Coniferocaulon sp.; this agrees very closely in surface- features with the casts from Kent, but an examination of transverse and longitudinal sections demonstrated that the apparent surface is not the actual surface, and such anatomical data as it was possible to obtain clearly indicated the Coniferous nature of the wood, conclusions in agreement with those reached by Dr Stopes in the case of some English specimens. The reference of these stems to Cycads or Conifers was based entirely on surface-characters and it was recognised that no definite conclusion was possible without anatomical confirmation. Dr Marie Stopes® succeeded in obtaining preparations of tracheids from a Lower Greensand cast showing uniseriate and separate bordered pits of the Abietineous type thus disproving a Cycadean affinity. This discovery led to the substitution of Coniferocaulon for Benstedtra, at least as regards the specimens which afforded anatomical evidence; in the French and South African examples no internal structure is preserved. Dr Knowlton* pointed out that the generic name Benstedtia should be preferred to Contferocaulon on the ground of priority and he named the English specimens Benstedtia Benstedtw. Dr Stopes replied to this criticism by asking why the Dragon tree, which is merely a partially decorticated piece of badly: preserved Coniferous wood, should have a name. Specimens exhibiting distinctive surface-features, whether complete or decorticated and even if they are in some cases at least portions of Coniferous stems, are none the less entitled to some recognition as a matter of convenience. Some excellent illustrations of Benstedtia casts are given by Dr Stopes in her recently published Catalogue of Lower Greensand Plants®. Seward (03) B. p. 34; (112) p. 690. Bancroft (13) pp. 72, 85. 3 Stopes (11). Knowlton (11). 5 Stopes (11%). Stopes (15) p. 159, Pls. xu, xiv. eof pe CHAPTER, LI. PODOZAMITES AND NAGEIOPSIS; GENERA INCERTAE SEDIS. PODOZAMITES Braun, and CYCADOCARPIDIUM Nathorst. The name Podozamutes+ was instituted for certain species pre- viously included in Zamutes characterised by the possession of distant alternate pinnae with a contracted base and veins slightly spreading in the proximal part of the lamina but for the most part approxi- mately parallel. As defined by Braun Podozamutes differs in no very important respect from Zamuites, and the latter name is retained by Schenk for Z. distans Presl in preference to Podozamites applied to that species by Braun. By most authors Podozamuites has been - regarded as Cycadean, but Schenk’s discovery of a specimen of Podozamites? in the Rhaetic beds of Franconia showing a cluster of small scale-leaves at the base of the axis led him to suggest a possible affinity to Agathis as an alternative to the generally accepted view that the appendages are leaflets or pinnae homo- logous with those of a pinnate Cycadean frond. In a later paper Schenk included in Podozamites some undoubted pinnate fronds on which Schimper founded the genus Glossozametes?. Schenk was, however, influenced in his preference for a Cycadean alliance by the structure of the epidermal cells (fig. 812, E) which have straight walls, and the same consideration weighed with Zeiller* who was strengthened in his view by the characters of the seed- bearing sporophylls described by Nathorst® and provisionally connected by him with Podozamites. Nathorst® described a speci- men from the Rhaetic strata of Scania agreeing in the presence of basal scale-leaves with that figured by Schenk, and more recently 1 Braun (43) p. 36. 2 Schenk (67) A. Pl. xxxvz. fig. 3. 3 Jbid. (71) Pl. o. 4 Zeiller (03) B. p. 159. ® Nathorst (86) p. 91, Pl. xxvr. 6 Ibid. (86) Pl. xvi. fig. 10. 448 PODOZAMITES [cH. Fra. 811. Podozamites lanceolatus. (Nat. size; British Museum, 39,303.) Li] CYCADOCARPIDIUM 449 Schuster! has published photographs of some examples of Podo- zamites distans in which the base of the axis is invested by small imbricate scales and in connexion with it are two other clusters of similar scales, probably unexpanded buds. In 1900 I expressed the opinion that Podozamites is probably a Conifer, the supposed pinnate fronds (fig. 811) being foliage-shoots like those of recent species of Agathis. The most important recent contribution to our knowledge of Podozamites is due to Nathorst: in 1911 he published additional facts? with regard to some seed-bearing organs from the Rhaetic of Scania for which he proposed the generic name Cycado- carpidium in 1886 and in 19024 more fully described the type-species C. Erdmann. Until the publication of Nathorst’s more recent paper Cycadocarpidium was known only as detached sporophylls found in beds containing Podozamites leaves. The following de- scription is abridged from Nathorst’s account. Cycadocarpidium Erdmannc is represented by ovate sporophylls consisting of a sterile portion 9 mm. long and at most 6 mm. broad with 4—5 simple veins, tapering to a short and slender pedicel on each side of which is an oval seed (fig. 812, A—-D) with an obliquely placed triangular lamina compared by Nathorst with a cupule and interpretéd by Schuster as a leaflet. A specimen figured by Nathorst shows several] sporophylls attached to a common axis, and supports his view that the seed-bearing organs were borne as imbricate carpellary scales. Fig. 812, Ais drawn from Nathorst’s restoration of a cone-like cluster of sporophylls. Another type of sporophyll, Cycadocarpidium Swabii, is distinguished by the larger dimensions of the lamina, 4-1 cm. long and 16 mm. broad, with 10 veins: in this type the two small seeds are apparently without any appen- dages (fig. 812, C, D). A third species, C. redivivum, is founded on small detached leaves and bud-like clusters previously as- signed to Podozamites distans. These are now recognised as small Cycadocarpidium sporophylls. There is a strong probability that Cycadocarpidium was borne on a Podozamites shoot ; in form and venation the sterile lamina of the sporophylls agrees with the leaves of Podozamites and the two organs are constantly associated in the Scanian beds. Zeiller records C. Erdmanni in Rhaetic 1 Schuster (114). 2 Seward (00) B. p. 242. 3 Nathorst (11+). 4 Ibid. (02) p. 8, Pl. 1. figs. 5, 6. 8s. IV 29 450 GENERA INCERTAE SEDIS — [CH. rocks in Tonkin! where Podozamites also occurs. Heer? in his account of some impressions of Podozamites from Spitzbergen figured a seed in close association with what he believed to be a carpellary leaf like that of a Cycad and suggested a connexion between the seed and Podozamites: this supposed connexion has, however, little to support it. Fic. 812. A, B, Cycadocarpidium Erdmanni. A, Restoration of fertile shoot. B, Single sporophyll with seeds. C, D, Cycadocarpidium Swabii. (After Nathorst.) E, Podozamites distans. Epidermal cells. (After Schenk.) Nathorst’s view of the sporophylls is that each is a single fertile leaf bearing two ovules comparable morphologically with the megasporophylls of Dioon, but, as he points out, the terminal portion of the sporophyll of Cycadocarpidiwm is much more leaf- like than the sterile distal end of the megasporophylls of Dioon, Encephalartos Zamia. He considers that the evidence afforded * Nathorst (11*) p. 5. 2 Heer (77) i. Pl. vim. fig. 4. LI] CYCADOCARPIDIUM 451 by these seed-bearing organs favours a Cycadean alliance: on the other hand he has satisfied himself that some Podozamites specimens are shoots with spiral linear leaves like those of Agathis. It is in many cases very difficult to say whether the axis of a Podozamites bears the leaves in two ranks or spirally. Nathorst speaks of some examples in which the leaves are not spiral and suggests the existence of two kinds of branch some with spiral and some with two-ranked leaves as in certain Conifers. But if this is the case one can hardly imagine that the two-ranked arrangement is not due to the twisting of the leaves of shoots with spirally disposed foliage. In a recent contribution to the systematic position of Podozamites Schuster! speaks of Cycadocarpidium Erdmanni as differing from C. Swabit in the presence of two ‘rudimentary leaflets’ (the triangular lamina shown in fig. 812, B) which in the latter species are represented only by two small swellings at the upper ends of the seeds: he regards C. Erdmanni as the more primitive type. The lamina is homologised with the cover-scale or bract of the double cone-scale of the Abietineae; he compares the two leaflets of C. Hrdmanni and the swellings in C. Swabit with abnormal seminiferous scales of an Abietineous cone. Schuster’s view is that Podozamites distans is a primitive Conifer evolved from the base of a Cycadofilicinean line which gave rise to the Ginkgoales, a supposition based on a very slender foundation. Nathorst regards Podozamites as an intermediate type related both to Cycads and Conifers; he does not, however, overlook the fact that the sporophylls of Cycadocarpidium may be compared with those of some Conifers even though their resemblance to Cycadean sporophylls would seem to be closer. Additional data are needed before we can settle the position of Podozumites, but such informa- tion as we have may be said to point to the conclusion that it is nearer to the Conifers or the Ginkgoales than to any other group of Gymnosperms. Nathorst calls attention to a similarity between Yokoyama’s Ginkgodium Nathorstc? and separate leaves of Podo- zanutes ; a similar comparison may be made between the latter genus and Eretmophyllum, a genus instituted by Thomas? and referred to the Ginkgoales (ef. figs. 658, 659, pp.59,62). There is indeed some resemblance between C'ycadocarpidiwm sporophylls and abnormal 1 Schuster (114). 2 Yokoyama (89) B. Pls. vim, rx. 3 Thomas (13). 29—2 452 GENERA INCERTAE SEDIS [CH. seed-bearing leaves of Ginkgo. We have as yet but little to guide usin our attempts to trace the ancestry of that remarkable survival Ginkgo biloba, and it is highly probable that, if more satisfactory records of older members of the Ginkgoales were available, we should be able considerably to extend the range in morphological characters which in the present representative of the group is comparatively restricted. The numerous leaf-bearing axes, many of them branched, referred by Fontaine! to his genus Nagetopsis, should not be overlooked from the point of view of their possible relationship to Podozamates. The branching habit of these Potomac specimens is no bar to an affinity to Podozamites if examples of the genus are no longer to be interpreted in terms of a Cycadean frond. Berry?, in a recent revision of Fontaine’s genus, refers some species to Podozamites which he still regards as Cycadean. It is interesting to find on a specimen of Nagewopsis figured by Fontaine a zone of crowded scars? (fig. 816, s) such as may be seen on an Agathis shoot. Many of the leaves described as Podozamuites are of little value as evidence of the occurrence of the genus. In the case of imperfect specimens of detached leaves it is often impossible to distinguish between Podozamites, Phoenicopsis, and the leaves of Araucarian plants, or pinnae of some species of Zamites. It is therefore not possible to state with confidence the geological range of the genus. Undoubted examples of Podozamites are essentially Rhaetic and Jurassic fossils, and there can be no doubt as to the abundance and wide geographical range of the genus in both these periods. Such leaves as those recently figured by Hollick* from Cretaceous beds of Long Island as Podozamites lanceolatus certainly agree closely in form with that species, but they are all detached speci- mens: the fragmentary leaves from the Middle Cretaceous beds of the Amboy clays described as P. angusttfolius (Eich.) and P. mar- ginatus Heer® afford no proof of the presence of Podozamites: similarly Velenovsky’s species P. miocenica from Bohemia® might equally well be referred to the genus Dammarites. Well pre- served specimens have been described by Zeiller? from the 1 Fontaine (89) B. p. 195. ? Berry (10). 3 Fontaine (89) B. Pl. Lxxvi. fig. 5. 1 Hollick (12) Pls. 162, 163. 5 Newberry and Hollick (95) Pl. xm. figs. 1—6. 5 Velenovsky (81) Pl. 1. figs. 18—20. * Zeiller (03) B. Lt] PODOZAMITES 453 Rhaetic flora of Tonkin!. There are few satisfactory records of the genus from the southern hemisphere, and we have no actual proof of its existence in India, though Feistmantel? refers to Podozamites detached leaves, which, as an examination of the original specimens shows, may have been borne on Podozamites shoots, but they may also be examples of Phoenicopsis. One of the leaves figured by Feistmantel from the ° Jabalpur group as Podozamites lanceolatus is reproduced in fig. 813 from a drawing recently made from the actual fossil: the lamina shows several fine parallel stria- tions between the more clearly marked veins. Miss Holden, who examined the carbonised cuticles of some of the Indian leaves lent to me by the Director of the Indian Geological Survey, found that the epidermal cells have straight walls and 5 13 A Bastien teed the stomata, usually with six accessory py Feistmantel as Podo- cells, occur in the intercostal regions on Soman ait ga both surfaces: the characters of the epi- a a caw. dermis are favourable to a relationship — Jndia.) with the Coniferales and they are not inconsistent with the inclusion of the fossils in the genus Phoe- nicopsis. Halle? has recently described some imperfect leaves from Patagonia as probably Podozamites, but as he pertinently says the evidence is not enough to establish the correctness of the determination. Some of the leaves from the Potomac beds in- cluded by Fontaine in Podozamites are of little value as authentic records of the genus, but there is still considerable doubt as to the relationship between this genus and Nageiopsis which was very abundant in the Potomac flora. The leaves figured by Fontaine* from the Jurassic of Oregon and from Alaska are also not above suspicion as records of Podozamites, though there is no doubt that the genus was represented in some of the North American floras. 1 Walkom (17) p. 20. 2 Feistmantel (82) p. 39, Pl. 1. figs. 2—5. 3 Halle (13). : 4 Fontaine in Ward (05) B. Pls. xxtv., xxv., XLIv. Knowlton (14) Pls. v., vi. 454 GENERA INCERTAE SEDIS (cH. Podozamites distans (Presl). This Rhaetic species! differs very slightly from the Jurassic type Podozamites lanceolatus and there has been much confusion on the part of authors between the two forms? which, indeed, cannot always be clearly distinguished. P. distans is often repre- sented only by detached leaves but in some specimens the shoot reaches a length of 20cm. The slender axis bears distant, lanceolate or ovate-lanceolate leaves, sometimes slightly falcate with a rounded or obtusely pointed apex and gradually contracted at the base which, as Zeiller says, may assume the form of a very short pedicel. The leaves may be 4—7 cm. long and 5—14 mm. broad; the veins, 0:-4—0-7 mm. apart, are dichotomously branched in the proximal portion of the lamina but elsewhere parallel and simple, except that they slightly converge at the apex. The epidermal cells have straight walls and the stomata, which occur on the lower surface, either in rows or scattered, are surrounded by small subsidiary cells (fig. 812, H). The leaves are usually rather broader in proportion to.their length than those of P. lanceolatus and the apex is less pointed. Braun? instituted two varieties, longifolius and latifolius, and to these Schenk# added others. The species is recorded from the Rhaetic of Scania®, where it is abundant, from Persia®, Tonkin, and many other regions: it occurs also in Jurassic strata’, but on the whole P. distans is a characteristic member of Rhaetic floras. The Rhaetic species Podozamites Schenki Heer® founded on Jurassic specimens from Siberia and described by Zeiller? and Nathorst from Tonkin, Persia, and Sweden is distinguished from P. distans by the smaller shoots and the more acuminate leaves. Podozamites lanceolatus (Lindley and Hutton). The type-specimen of Zamia lanceolata Lind. and Hutt.!° in the Manchester Museum from the Middle Jurassic beds of Yorkshire consists of a slender axis bearing scattered and distant linear- 1 Sternberg (38) A. Pl. xxr. fig. 1. ? See Zeiller (03) B. p. 159 for examples of P. distans referred to P. lanceolatus. 3 Braun (47) p. 85. 4 Schenk (67) A. Pls. xxxv., XXXVI. 5 Nathorst (78) B. Pls. xm, xv. 8 Zeiller (05) p. 193. * For references, see Zeiller (03) B. p. 159. 8 Heer (77) ii. p. 45. - ® Zeiller (03) B. Pl. xia. ke ° Lindley and Hutton (36) A. Pl. xctv. Li] rs PODOZAMITES 455 lanceolate leaves up to 7 cm. long and 7 mm. broad; the lamina has a tapered acuminate apex and a less gradually though not abruptly contracted base. The leaves differ from the leaflets of Zama which they superficially resemble in their less abruptly contracted proximal end. In habit a shoot of P. lanceolatus (fig. 811) very closely resembles the fronds of Zamia media. Some of the leaves in the type-specimen appear to be laterally attached, while others appear to be given off from the upper surface. The leaves of this as of other species are frequently found detached. The variability in the form of the leaves has led to the employment of several varietal names, and if not used too freely the addition of some descriptive term to the specific name may often serve a useful purpose. Fig. 811 represents a good example of the species from the Yorkshire coast. The method of attachment of the leaves is not always clear, but their irregular distribution and the slender axis are features more in accordance with a foliage-shoot than a pinnate frond. Podozamites lanceolatus is a widely dis- tributed Jurassic species! recorded from many European localities extending to North Siberia and Spitzbergen as well as from North America, Turkestan, Afghanistan, Japan, China, and elsewhere. The specimens figured by Feistmantel from Upper Gondwana tocks in India as P. lanceolatus (fig. 813) should, I am inclined to think, be assigned to Phoenicopsis. Podozamites Reinit Geyler; Podozamites stonesfieldensis Seward; Podozamites Griesbacht Seward. These species from Jurassic strata serve as examples of a broader type of the genus represented in the last two species by detached leaves only. In these as in many other cases one cannot feel absolute confidence as to the correctness of the determination. In some of the Japanese examples of P. Reini? (fig. 814) the broadly oval leaves are attached to a slender axis. P. stonesfieldensis® from the Great Oolite of Stonesfield is probably identical with the leaves originally described by Buckman as Navadea ovata and Liha lanceolata: the leaves are oblong-ovate, approximately 8 x 3-5 cm.; the lamina is rather abruptly contracted at the base and more 1 For references see Seward (00) B. p. 242; (07°); (11). 2 Geyler (77) B. Pls. xxxmI., XXXIv. 3 Seward (04) B. p. 121, Pl. m1. fig. 4; Pl. x1. figs. 1, 2. 456 GENERA INCERTAE SEDIS’ [cH. gradually tapered towards the apex; the veins are slightly more than 1mm. apart and converge at each end of the lamina. The species re- - sembles P. lanceolatus var. latifolius figured by Schenk! from China. Podozamites Griesbachi? is a Jurassic species from Afghanistan similar in the shape and size of the leaves to the other two species; it has a coarser venation than P. Reiniz but the venation is still coarser in P. stonesfieldensis. A similar form of leaf is figured by Velenovsky? from the Lower Cretaceous of Bohemia 5. ¢14 podozamites Reinii. as P. striatus. (After Geyler; 3 nat. size.) Podozamites Kidstoni Etheridge. In this type’, usually represented by detached leaves, the lamina is smaller than in other species, short and broad with a blunt apex and abruptly contracted at the base. The species is recorded from Afghanistan> and similar leaves are figured by Chapman® from Jurassic rocks in Victoria, Australia. Etheridge’s type-specimen is from the Burrum Coal Measures (? Triassic) of Queensland. This form of leaf agrees closely with some of Fontaine’s Potomac species of Nageiopsis, e.g. N. obtusa and N. heterophylla’, and a similar though not identical form is described by Nathorst® from the Rhaetic of Sweden as P. ovalis, distinguished by its broadly rounded and mucronate apex. NAGEIOPSIS. Fontaine. This genus was established® for vegetative shoots abundantly represented in the Potomac flora many of which closely resemble Podozamates, though differing in their branched habit and in the veins being less convergent in the apical region of the lamina. 1 Schenk (83) A. Pl. xurx. figs. 4b, 5. ° Seward (12) p. 36, Pl. rv. fig. 58; Pl. vi. fig. 79. > Velenovsky (85) B. Pl. m. fig. 8. 4 Jack and Etheridge (92) B. p. 317, Pl. xvmu. figs. 6, 7. 5 Seward (12) Pl. rv. fig. 39. 6 Chapman (09) Pl. xv. * Fontaine (89) B. Pls. uxxxIv., LXxXxv. 8 Nathorst (78) B. Pl. xu. fig. 5. ® Fontaine (89) B. p. 194. LI] NAGEIOPSIS 457 Fontaine recognised the similarity between Nagetopsis and shoots of Podocarpus belonging to the section Nageia: this suggested the choice of the generic name. Berry! in his revision of Nagevopsis transfers some of Fontaine’s species to Podozamites; he also reduces the number of the species retained in Nagevopsis on the ground that Fontaine attached too much importance to variations in the size and form of the leaves. I have elsewhere suggested? that some of the shoots referred to Nageiopsis may be Araucarian, as in habit they closely resemble Araucaria Bidwilli and Agathis. Until reproductive organs are discovered it is impossible to speak with confidence with regard to the position of the genus. It may be closely allied to Podozamites or, as Fontaine believed, it may be related to Podocarpus. It should be noted that some of the speci- mens included by Fontaine in Nagetopsis are hardly distinguishable from Zamites Buchianus®. Trees or shrubs characterised by irregularly branched foliage- shoots bearing leaves usually in two ranks but spirally attached; the leaves exhibit a wide range in size and shape, long and linear or lanceolate, acute or subacute, more or less abruptly contracted at the proximal end and attached by a very short stalk; there are several parallel veins dichoto- : mously branched near the base of the lamina. -¢gmms Nageiopsis anglica Seward. This species founded on the small specimen represented in fig. 815 has distichous leaves — 1—1-5 cm. long with several parallel veins. py. 815, Nageiopsis an- The English Jurassic type agrees generally — glica. (From a speci- with Nageiopsis microphylla Font. and N. des- men in the Whitby ae ae 4 Museum; # nat. size.) crescens Font.: a similar form is recorded from the Wealden beds of Sussex’. Though satisfactory evidence of affinity is lacking it is permissible to suggest an Araucarian affinity. Nageiopsis longifolia Fontaine. The linear-lanceolate leaves reach a length of 8—20 cm. and 5mm. to 1-3cm. in breadth; there are 9--12 veins unbranched 1 Berry (10). 2 Seward (12) p. 33. 3 Berry (11) Pl. uxt. 4 Seward (00) B. p. 288, fig. 51. 5 Ibid. (95) A. p. 211, Pl. x11. fig. 3. 458 GENERA INCERTAE SEDIS [CH. LI except at the base (fig. 816); the lamina is abruptly narrowed and attached by a short and slightly twisted stalk. Though apparently inserted laterally the leaves are in all probability spirally disposed. In one of Fontaine’s figures there is a group of small scars, fig. 816, s, - presumably of bud-scales, at one place on the axis. This species Fic. 816. Nageiopsis longifolia. (A, after Fontaine; nat. size.) is very abundant in the Potomac beds of Virginia and Maryland! and is recorded by Fontaine from several other localities though for the most part on slender evidence. Nageiopsis zamioides Fontaine. In this species? the leaves are ovate-lanceolate and shorter than in N. longifolia; they have a maximum breadth of 2cm. and reach a length of 7cm. The example described from English Wealden beds as Nagetopsis sp. cf. N. heterophylla agrees closely with Fontaine’s Potomac species. 1 Fontaine (89) B. p. 195, Pls. uxxxv.—LxxIx.; Lxxxv.; Berry (11) p. 384. . 2 Ibid. p. 196, Pls. Lxx1x.—.Lxxx1.; Berry (11) p. 386, Pls. uxu., Lx. CHAPTER LI. GNETALES. IT. Recent. In this group of Gymnosperms are included three genera, Ephedra, Gnetum, and Welwitschia. They differ widely from one another in vegetative features, and Ephedra, the most primitive, is distinguished by certain important peculiarities of the repro- ductive organs. EPHEDROIDEAE. Ephedra. GNETOIDEAE. Gnetum, Welwitschia. Having regard to our exceedingly meagre knowledge of fossil representatives, it is urinecessary to deal fully with the recent types!, but the members of this aberrant section of seed-plants exhibit morphological characters of interest from the point of view of comparison with the Bennettitales and the Angiosperms. Though in external appearance the three genera are poles asunder, they have in common certain features both in the vegetative and reproductive organs which differentiate them from all other Gymnosperms and connect them more closely than the Cycads or Conifers with the Angiosperms. The leaves are opposite; the secondary xylem contains vessels in addition to tracheids; the male and female flowers are characterised by the possession of one or two envelopes in addition to the usual single integument; the inflorescences, occasionally though not as a rule bisporangiate*, are distinguished by a dichasial system of branching, a character foreign to Gymnosperms as a whole though exhibited by the stem of Wielandiella, a mémber of the Bennettitales. There are good reasons for believing that pollination is effected by insects? in 1 For a general account of the group, with illustrations, the student is referred to Wettstein (11) and Lotsy (11), or to Coulter and Chamberlain (10). 2 Land (04); Berridge and Sanday (07) p. 127; Lotsy (11) p. 293. 3 Pearson (062) p. 274; (09) p. 343; Berridge and Sanday (07) p. 172; Karsten (92); Porsch (10). ae 460 GNETALES [cH. Welwitschia, in some species of Gnetum, and occasionally in Ephedra. The seeds are albuminous and the embryos have two cotyledons. Archegonia are produced in the female prothallus of ‘ Ephedra while in Gnetum and Welwitschia these organs are repre- sented by single cells as in the Angiosperms or by nuclei. Ephedra has a wide distribution in the warm temperate regions of the northern hemisphere: in America it occurs on both sides of the equator and from the Mediterranean region it reaches to Brittany in the west and North Africa in the south. Gnetum extends both east and west in the tropics: Gnetum scandens is a widely spread Asiatic species, and the genus occurs in Angola and in some other parts of Africa. Welwitschia is confined to a littoral strip of desert in extra-tropical South Africa from 14° S. to 23° 8. and has not been found more than 50 miles from the coast. Ephedra. Shrubs, in some species with climbing branches, characterised by an Equisetum-like habit of the younger shoots which form long jointed and slightly fluted branches bearing whorls of two or sometimes three, scaly, concrescent leaves. In rare cases, ¢€.g. Ephedra altissima, the leaves may reach 3cm. in length and a breadth of 1--1-5mm. Monoecious or dioecious; flowers uni- sexual; bisexual inflorescences are recorded in E. campylopoda?. The female flowers occur in strobili on a dichasially branched inflorescence; each strobilus consists of three pairs of bracts, in some species the bracts are more numerous. There is generally a single ovule in E. alftssima, but in most species there are two or as many as six ovules in a single strobilus. The ovules are enclosed by two envelopes regarded by some authors as a perianth and an integument and by others as two integuments. In E. distachya, as described by Mrs Thoday and Miss Berridge®, two vascular bundles supply the outer énvelope (outer integument) one running up each angle of the flattened side of the flower. The thin inner integument becomes free from the nucellus at a distance of two-thirds its length and projects beyond the outer envelope as a long style-like micropylar tube. A ring of bundles runs a short distance up the inner integument but ends low down in a mass of transfusion- 1 Stapf (89). 2 Wettstein (11) p. 417. 3 Thoday (Sykes) and Berridge (12). Lit] | EPHEDRA 461 tracheids. Attention is drawn to the resemblance of the outer integument to the integument of Bennettites, and the single ovule of Ephedra is considered to be the representative of the whole ovulate strobilus of Bennettites of which it is a much reduced derivative. There is a deep pollen-chamber at the apex of the nucellus! and there are 2—8 long-necked archegonia at the summit of the prothallus (endosperm). In its female prothallus and sexual apparatus Ephedra differs considerably from Gnetum and Welurtschia. The second envelope forms the hard shell of the seed which is enclosed by bracts either in the form of membranous wings (sect. Alatae) or as a red or yellow flesh (sect. Pseudobaccatae). Ewart? found that the seeds of Ephedra distachya germinated after 93 days’ immersion in sea-water. The male flowers? occur also in strobili on dichasial inflorescences, a single flower occurs in the axil of each of the fertile bracts. A flower consists of a short axis bearing a pair of membranous appendages and the flower-axis is prolonged as a simple or bifid stalk bearing bilocular synangia, 2—6 according to the species. In some cases the central stalk or antherophore of the flower is flattened and laminar* instead of the usual cylindrical form: it has been interpreted both as an axial and a foliar structure, but the latter interpretation is probably correct. Arber and Parkin® regard the antherophore as having been formed from two fused members, and this view is adopted by Mrs Thoday and Miss Berridge. On the basis of this interpretation the microsporophylls of Ephedra, represented by the antherophore, are considered to be homologous with the disc of sporophylls of a Benettitean flower and with the stamens in a male flower of Welwitschia. Anatomi- cally® Ephedra exhibits a closer agreement with the Conifers and in some respects with the Dicotyledons than with recent Cycads or the Bennettitales. The presence of vessels in the secondary xylem is an Angiospermous feature though in structure they differ from the Angiospermous type; the pitting of the tracheids is in the main Abietineous but the occurrence of compressed pits 1 Land (04); for other references, see Lignier and Tison (12); also Sigrianski (13) 2 Ewart (08). 3 Thibout (96) gives a good account of the male flowers of the Gnetales. 4 Thoday and Berridge (12) p. 970. 5 Arber and Parkin (08). 6 Thompson (12). See also Jeffrey (17) p. 357. 462 GNETALES [cH. furnishes a point of contact with the Araucarineae; rims of Sanio occur and xylem-parenchyma is abundant; the medullary rays are multiseriate as in Dicotyledons. The bast on the other hand is essentially gymnospermous. The occasional occurrence of spiral bands in the tracheids and the presence of lignified trabeculae in the xylem-elements are other Coniferous traits. The leaf-trace is double, a feature met with in Agathis as well as in recent Cycads but not in the Bennettitales. The anatomy of seedlings affords further indications of resemblance to Araucaria and the Podocarps!}. It would seem, then, that the case for a relationship between the Gnetales and the Bennettitales founded on the facts of floral morphology does not derive support from the anatomical features of the most primitive genus of the group. Gnetum. Small trees or climbers with long and slender stems; the inter- nodes, sometimes reaching a length of 15 cm., bear pairs of ovate- oblong or lanceolate-acuminate leaves 11—18 cm. long by 4—7 cm. broad. The leaves? agree in form and venation with those of many Dicotyledons and could not be distinguished from them in a fossil state. The epidermal cells have undulate walls. The flowers are in spikes; at each node two fused bracts form a cupular structure in the axil of which the male or female flowers are borne on an annular swelling. The male flowers? are in 3—5 whorls: each consists of an envelope of two coherent leaves enclosing a central column, as in Ephedra, which bears at the apex one, two, or rarely four unilocular sporangia or reduced synangia. The antherophore eventually elongates and pushes the anthers through an aperture at the summit of the floral envelope*. In appearance the antherophore of Gnetum approaches most closely to the stamen of an Angiosperm. The female flowers® occur in a single series, 5—8 in a whorl; each consists of an ovule surrounded by three envelopes; the outermost is coloured and succulent, the middle envelope or outer integument is differentiated after fertilisation into an inner sclerotesta and an outer sarcotesta, while the inner- most covering is prolonged as a micropylar tube. There are no 1 Hill and de Fraine (10) p. 329. > Karsten (93); Lotsy (11) p. 347, fig. 209. 3 Caporn (16). + See also Pearson (15). ° Thoday (Sykes) (11); Lignier and Tison (13). See also Pearson (17). Lit] GNETUM 463 archegonia: in some species the megaspore contains numerous free ‘nuclei all of which are potentially sexual; after fertilisation a sterile nutritive tissue, or endosperm, is formed in the lower part of the spore. In Gnetum Gnemon the endosperm is often formed before fertilisation. In the great reduction of the female apparatus and in the nature of the endosperm Gnetum! agrees much more closely with Welwitschia than with Ephedra. Attention has been called to certain resemblances between the seed of Gnetum and Fic. 817. Seed of Gnetum africanum. a, outermost envelope; v, vascular strands; b, outer integument; f, flange of micropylar tube; 7, nucellar cap; c, inner integument; p, endosperm. (After Mrs Thoday.) that of Bennettites?. For convenience of comparison a diagram- matic section of a seed of Gnetum africanum is reproduced in - fig. 817. The outermost envelope, a, forms a green succulent covering free to the apex; the middle envelope, 6 (outer integument), is free from the other envelopes except at the apex where it is locked to the inner integument by the downward growth of a flange, f, from the apical region of the inner integument; the inner 1 Pearson (09); see also Karsten (92); (93%); Lotsy (99). 2 Berridge (11); (12); Thoday (Sykes) (11). 464 GNETALES [CH. integument, c, is united to the nucellus for about two-thirds of the length of the seed-body. The nucellus consists of a few layers of cells and at the apex forms a nucellar cap, n, the cells of which are lignified; this cap is supported by a short tent-pole produced from the summit of the endosperm. A ring of vascular bundles enters the base of the seed and forms three series, the two outer pass up to the tip of both the two outer coverings, v, v,:and the inner series extends up the inner integument as far as the level where nucellus and integument part company. In the seed shown in fig. 817 the micropyle is closed and the tissue in the closed region of the canal is lignified and dark. Among other features in which this seed agrees with that of Bennettites Morierei is the inner zone of the outer integument, composed of a palisade and a fibrous layer; the fibrous layer becomes five-angled in the upper part of the seed? and in transverse section presents a striking similarity to sections through the same region of a Bennettites seed (figs. 524, 527, Vol. 11. pp. 397, 402). The oval fleshy seeds are able to germinate after lying some months in sea-water. There is a fairly close resemblance between Gnetum and Ephedra as regards anatomical: characters, but some species of Gnetum (sect. Thoa) are character- ised by the formation of successive cambial cylinders as in Cycas. Strasburger? pointed out that in the vascular bundles of the leaves the parenchyma of the medullary rays forms continuous plates, a gymnospermous character. Welwitschia. This remarkable genus, discovered by Welwitsch in 1860 and described by Sir Joseph Hooker, presents striking peculiarities in | the habit of the vegetative body. A Welwitschia plant has been aptly termed an adult seedling’; the large and squat tuberous stem, morphologically the swollen hypocotyl, may be as much as 45m. in girth. The seedling has two cotyledons and an ex- ceptionally long radicle: at an early stage a pair of isobilateral leaves is produced at right-angles to the cotyledons and these 1 Thoday (Sykes) (11) p. 1116, text-fig. 11. * Strasburger (91) p. 148. For an account of the anatomy of Gnetum, see Duthie (12); La Riviére (16); Thomson, M. R. H. (16). 3 Hooker, J. D. (63). For figures of Welwitschia, see also Gard. Chron. Jan. 22, p. 49, 1910. _ 4 Sykes (10?) p. 333. . Lit] WELWITSCHIA 465 persist as the only leaves throughout the long life of the plant, attaining a length of 5m. The tough lamina is torn into strips by the wind and ‘the extraordinary appearance of the shapeless mass of coiled and twisted leaf-ribands standing out in bold relief from the sharp glistening dead landscape passes description!.’ The venation is parallel and there are numerous cross-connexions, some ending blindly in the mesophyll?. Welwitschia is dioecious and the flowers are borne in inflorescences with a dichasial branch- system produced from pits on the crown of the stem; the female inflorescences, which are larger than the male, reach a length of 30 cm. and bear cones about 7 cm. long. The female flowers occur singly in the axils of bracts which form four orthostichies giving a four-angled form to the cones. Each flower may produce two small leaf-rudiments®, but the flower proper consists of an ovule with two envelopes; the outer, called by Hooker the perianth, is considerably extended tangentially and in. the ripe seed forms a wing-like appendage producing an appearance almost identical with that of some Samaropsis seeds. .The inner integument is prolonged upwards like a long and slender hollow bristle for a distance of 4—5 mm. beyond the upper edge of the subtending bract. The inner envelope has no vascular supply. The secretion of sugar in the micropylar tube attracts the pollinating insect Odontopus sexpunctulatus. The staminate cones are smaller and the subtending bracts connate. The outer envelope of the flower is formed of two membranous segments without vascular bundles which may be styled lateral prophylls of the axillary shoot: internal to these are two fused members forming a sac-like investment with free rounded lobes also without a vascular supply. Within these perianth-segments is the staminal tube bearing six free stamens each supplied with a vascular bundle and bearing a terminal trilocular synangium (fig. 818). The centre of the flower is occupied by a pyriform ovule surrounded by a thin integument continued as a slightly kinked stylar tube terminating in a flat stigmatic disc 1mm. in diameter. There is no embryo-sac but the nucellus acts as a nectary, the drop-mechanism of the functional 1 Pearson (06?) p. 270. 2 de Bary (84) A. fig. 157; Sykes (107); Takeda (13%). 3 Lignier and Tison (12). 466 ' G@NETALES [cH. ovule in the female flower being retained in the sterile ovule of the male}. In contrast to the indefinite, spirally disposed, bracts or Fic. 818. Welwitschia mirabilis. Staminate flower (sectional elevation), sub- tending bract and the two missing stamens indicated by dotted lines. (From a drawing kindly supplied by Dr A. H. Church.) perianth of Cycadecidea flowers Welwitschia has only two connate segments, and the staminal disc of Welwitschia is considerably 1 This account is based on the excellent description with illustrations by Dr Church (14). LT] WELWITSCHIA 467 reduced; the gynoecium consists of only one functional ovule in- stead of an indefinite number as in Cycadeoidea. Church regards the resemblance between the flowers of these two genera as an instance of parallel development, which does not imply relationship. He thinks there is ‘no indication whatever of any relation to the car- pellary flowers of the Angiosperms.’ Hooker’s account of the ovule has recently been considerably extended by the important researches of Pearson}. The megaspore consists of two regions, an upper fertile and a lower sterile portion; each is composed of ‘cells’ with more than one nucleus; some of the ‘cells’ of the fertile region grow upwards as tubes into the nucellar cone where pollen-tubes are encountered and fertilisation ensues. The nuclei in each ‘cell’ of the sterile region fuse and uninucleate cells are produced; this tissue now grows considerably in size and cell-divisions occur resulting in the formation of an endosperm. Pearson regards the free nuclei that are in the embryo-sac at the time of septation into the multinucleate ‘cells’ as all alike, and all potential gametes. It follows, therefore, that the endosperm formed in the lower portion of the sac is the product of fusion of sexual nuclei; it is not a gametophyte or a sporophyte and Pearson proposes for it the new term trophophyte, ‘a bye-product resulting from the fusion of potentially sexual nuclei and functioning in the same manner as the prothallus of the lower seed-plants.’ More recent work by this author confirms his opinion that the endosperm of Ginetum is also a trophophyte. For an account of the anatomy of Welwitschia and Gnetum the student is referred to original sources. Miss Sykes? called atten- tion to certain interesting characters,—the occurrence of reticulately pitted protoxylem elements in the stem, the arrangement of separate and not contiguous bordered pits in 1—2 rows on the tracheids, and to the presence of concentric steles and inversely orientated bands of vascular tissue in the stem and inflorescences closely resembling Medullosean features. In certain respects the Gnetales are closer than the Conifers or 1 Pearson (062); (09). 2 Sykes (10); (102); Worsdell (012); Pearson (12); Bower (81); (82); Hill and de Fraine (10); Boodle and Worsdell (94); Mary R. H. Thomson (16); Henriette C. C, La Riviére (16). 30—2 468 GNETALES [cH. the Cycads to the Angiosperms!. It has, for example, been sug- gested by Hallier? that they are reduced Dicotyledons comparable with the Loranthaceae and Myxodendraceae; while Lignier and Tison? regard them as a group of Angiosperms nearest to the Amentales. The question of relationship between the Gnetales and the Angiosperms, especially the difficult problems connected with the endosperm, was fully considered by Pearson in a paper on the reproductive organs of Gnetum Gnemon published in 1915, and in a later contribution®, published after his death, the morpho- logical problems are reviewed in the light of more recent work. The same subject is dealt with by Prof. Thompson® in a recent paper in which he calls attention to the form of the inflorescence, the arrangement of the parts of the flowers, the presence of an ovary with a style, the germination of the microspores at some distance from the nucellus, as evidence of affinity to the Angiosperms, and concludes that the ancestors of the Angiosperms were ‘not far removed from the genus Gnetum.’ On the other hand some botanists prefer to regard the Gnetales as a blindly-ending branch of Gymno- sperms with no direct relationship to the Flowering plants. Difficult as it is to believe that plants so different, when the sum of characters is considered, as the Gnetales and the Bennettitales are off-shoots of a common stock, it would be rash to assume that such resemblances as have been emphasised by Miss Sykes and other authors have no phylogenetic value. At the time of his death (November, 1916) Professor Pearson ‘was engaged upon a volume on the Gnetales: in April, 1916, he wrote, ‘A large part of the book on the Gnetales is written, though it will need some revision....As to the Gnetalean-Angiosperm alliance, there must be one, I think, but at present I cannot bring myself to believe that it is direct’.” Had Pearson been able to complete his work it is certain that a statement of his most recent conclusions would have enabled botanists to obtain a clearer view 1 Arber and Parkin (07); (08); references to other authors will be found in these papers. See also Lignier and Tison (12); Lignier (03); Lignier and Tison (11). 2 Hallier (05) p. 153. 3 Lignier and Tison (11). * Pearson (15*): additional references to literature are given at the end of this paper. See also Caporn (16). 5 Pearson (17). & Thompson, W. P. (16). ? Seward (17) p. ix. LIT] FOSSIL GNETALES 469 of the true position of this puzzling group which, despite the lack of palaeobotanical evidence, is probably a survival from a remote past. II. Gwetazes (Fossit). Arber and Parkin! and other authors have called attention to the lack of any trustworthy records of Gnetalean plants in the sedimentary strata of different periods.. Several specimens have been described either as generically identical with Ephedra or as probable representatives of the two other members of the group, but while some are incorrectly determined others are too im- perfect to be accepted as evidence. In view of the morphological features characteristic of the present members of the Gneétales and the geographical distribution of the species of Ephedra, Gnetum, and Welwitschia, it would seem safe to conclude that the absence of fossil forms is not explicable on the hypothesis of a recent origin of the group, but is rather the result of the imperfection of the geological record and of the difficulty of distinguishing between fragmentary remains of Gnetalean genera and vegetative or repro- ductive organs of similar external form belonging to other plants. Reference has already been made” to certain characters shared by the seeds of Gnetum and Bennettites and in spite of the great and obvious differences separating the Gnetales and Cycadales it would seem probable that the striking similarity between the seeds of Gnetum and those of the Bennettitales has some phylo- genetic significance. But even granting a phylogenetic significance to the evidence brought forward by Mrs Thoday and other authors, we have still to admit that an indication of some former connexion between the Gnetales and the Bennettitalean line is rather the shadow of evidence with regard to the geological history of the Gnetales and not a substantial contribution to our knowledge of the antiquity of this section of the Gymnosperms. The specimens described by Unger? from Eocene beds in Styria as Ephedrites sotzkianus, though very similar to those of Ephedra fragilis with which they are compared, are too fragmentary to be 1 Arber and Parkin (08) p. 507. 2 See page 463. 3 Unger (51) p. 159, Pl. xxv1. 470 GNETALES) [cH.. accepted as trustworthy records. The pieces of vegetative branches and the paired nuts described by Heer! from Jurassic strata in Siberia as Ephedrites antiquus are of no botanical value. Portions of inflorescences preserved in amber from the Baltic coast and named by Goeppert and Berendt? Ephedrites Johnianus and similar specimens referred by Goeppert from the same Oligocene beds to Ephedra Mengeana have been identified by Conwentz? as fragments of flowering shoots of a Loranthaceous genus, Patzea. Engelhardt* refers some slender branches from Tertiary beds in Chile to Ephedra but they, like most of the specimens recorded as fossil represen- tatives of the genus, are too incomplete to be accepted as evidence. In the absence of anatomical data or of well preserved flowers it would be exceedingly difficult to recognise impressions of vegetative shoots of Ephedra and to distinguish them from Dicotyledonous twigs of similar habit. Similarly the torn lamina of a Welwitschia leaf bears too close a resemblance to other linear parallel-veined leaves to be recognisable unless the preservation is such as to show traces of the characteristic venation mentioned in the account of -the recent genus. Comparisons between some fossil seeds and the winged seeds of Welwitschia®, though in some cases possibly justified by actual relationship, cannot be considered to have any ‘importance unless supported by additional evidence. The seeds named by Renault Gnetopsis and subsequently investigated by Oliver and Salisbury® are now recognised as types closely allied to Lagenostoma and other Pteridosperm seeds from Carboniferous rocks. In their monograph of the Phocene Floras of the Dutch- Prussian Border Mr and Mrs Clement Reid figure under the name Gnetum scandens var. robustum’ a piece of axis 8 mm. long and 4mm. broad showing eight nodes bearing crowded scars of some deciduous appendages. The authors speak of the specimen as ‘a portion of a male inflorescence of a Gnetum...so close to that of the living G. scandens that we cannot separate it.’ If their 1 Heer (77) ii. p. 82, Pls. xiv., xv. 2 Goeppert and Berendt (45) A. Pls. 1v., v.; Goeppert and Menge (83) A. Pl. xvi. 3 Conwentz (86) pp. 136, 138, Pl. xm. figs. 8—20. : 4 Engelhardt (91) p. 647. - 5 Seward (04) B. pp. 19, 20. 7 Oliver and Salisbury (11) p. 34. Reid, C, and E. M. (15) p. 55, Pl. xx. fig. 27. Lit]. FOSSIL GNETALES 471 identification is correct—and though the evidence is hardly con- clusive the resemblance between the fragment from Renver and an inflorescence axis of Gnetum is undoubtedly striking—it points to the occurrence in a Pliocene European flora of a genus that is now mainly tropical and which had not so far been recognised with any certainty in a fossil state. The striking resemblance of Gnetwm leaves to those of some Dicotyledons is an obvious difficulty in the way of the identification of impressions. It is among the oldest examples of supposed Dicotyledons that search should be made for possible representatives of the genus Gnetum. Among the earliest records of Angiosperms are those described by Fontaine! from the Patuxent series of the Potomac formation which rests on Palaeozoic crystalline rocks and contains the remains of a flora that is clearly Jurassic or Wealden in its general facies; but with Jurassic Gymnosperms and Ferns are associated some Dicotyledon-like leaves of ovate and linear form for some of which Fontaine instituted the genera Rogersia, Fico- phyllum, Proteaephyllum and referred others to Ficus, Sapindopsis etc. A revision of the Patuxent fossils by Berry? has led to a considerable simplification in nomenclature and to the conclusion that some at least of these Lower Potomac leaves are Gnetalean. A comparison of some of Fontaine’s figures of Ficus virginiensis, species of Ficophyllum, Proteaephyllum, and Rogersia with a leaf of Gnetum Gnemon reveals a very close agreement, as regards form and venation, consistent with Berry’s suggestion. It is by no means unlikely that these forerunners of the Dicotyledonous type that occur as foreign elements in a typical Jurassic flora, without an admixture of undoubted Angiosperms like those which occupy an important position in the upper beds of the Potomac formation, may belong to plants more closely allied to Gnetum than to any Angiosperm. Attention is especially called to the following species as revised by Berry and illustrated in Fontaine’s monograph: Ficophyllum oblongifolium (Font.), Rogersia longifolia Font., Pro- teaephyllum ovatum Font.? It is possible that a careful study of 1 Fontaine (89) B. pp. 281 ef sey. See also Seward (14?). 2 Berry (11) pp. 64, 148, 499, ete. 3 Fontaine (89) B. Pls. 139, 141, 144, 145, ete. 472 GNETALES [CH. LII the venation-characters of these and other fossil leaves may lead to the discovery of criteria which may enable us to separate the leaves of Gnetum from similar Dicotyledonous foliage. It is with a keen sense of the incompleteness of my task that Volume tv. is concluded without any attempt to deal with the abundant if, in very many cases, undecipherable records of Angio- sperms. The omission of this branch of Palaeobotany in what purports to be a general text-book calls for a word of explanation. A mere summary of conclusions so far published with regard to the geological history of Flowering plants would not yield results commensurate with the labour involved. What is needed is a critical examination, as far as possible, of the actual specimens and a careful scrutiny of the evidence on which determinations are based. It is undoubtedly the fact that a large number of leaf-impressions are practically valueless as trustworthy data, and I venture to think that it is only with the cooperation of trained systematists that any satisfactory estimate can be formed as to the value of the fragmentary documents preserved in Cretaceous and Tertiary strata. It is preferable to omit, at least for the pre- sent, this part of the subject than for the sake of completeness— in a treatise that is very far from complete in its treatment of the groups that have been considered—to essay a task for which the - author recognises that he is very inadequately equipped. LIST OF WORKS REFERRED TO IN THE TEXT. (Votumes III. anp IV.) [With a few exceptions this list does not include books and papers given in the Bibliographies in Volumes I. and IT.] ra The following are some of the Bibliographies which students will find useful for additional references:—Geological Literature added to the Geological Society’s Library, published from time to time by the Society ; Prof. Zeiller’s lists in the ‘Revue Générale de Botanique’ (Paris); lists given by Arber in the ‘ Progressus Rei Botanicae’ (Leiden), vol. 1. Heft i. p. 218, 1907; Jongman’s ‘ Die Palaeobotanische Literatur’ (Jena), 1910-13; also the International Catalogue of Scientific Literature (Botany and Geology). For the Literature dealing with Cretaceous plants the student should refer to Dr Marie Stopes’ ‘Cretaceous Flora,’ 1. and 1. (British Museum Catalogues, 1913, 1915). The dates of books published in parts given in the footnotes to this volume are as a rule those of the concluding part. For the dates of separate parts of books relating to Palaeozoic floras the student is referred to Prof. Zeiller’s valuable list at the end of the ‘Flore Fossile du Bassin Houiller de Valenciennes.’ Useful bibliographies of the writings of Saporta, Heer, and Ettingshausen have been compiled by Zeiller (96), Malloizel and Zeiller (N.D.), and Krasser (97). Aase, Hannah C. (15) Vascular anatomy of the megasporophylls of Conifers. Botanical Gazette, vol. Lx. p. 277. Affourtit, M. F. A. and H.C. C. La Riviére. (15) On the ribbing of the seeds of Ginkgo. Annals of Botany, vol. xxix. p. 591. Andersson, J.G. (10) Die Veranderungen des Klimas seit dem Maximum der letzten Eiszeit. (Collection of papers published by the Int. Geol. Congress; edited by J. G. Andersson.) Stockholm. Andrews, E.B, (75) Descriptions of Fossil Plants from the Coal Measures of Ohio. Geol. Surv. Ohio. Antevs, E. (14) Lepidopteris Ottonis (Gopp.) Schimp. and Antholithus Zeilleri Nath. K. Svensk. Vetenskapsakad. Hand. Bd. ut. No. 7. —— (142) The Swedish Species of Ptilozamites Nath. Ibid. Bd. 11. No. 10. : . —— (16) Das Fehlen resp. Vorkommen der Jahresringe in Paldo- und ’ Mesozoischen Hélzern und das klimatische Zeugnis dieser Erschei- nungen. Geolog. Féren. Stockholm Forhand. Bd. xxxvitt. — (17) Die Jahresringe der Holzgewachse und die Bedeutung der- . gelben als klimatischer Indikator. Progressus Rei Botanicae, p. 285. 474 LIST OF WORKS Arber, Agnes. (See also Robertson, A.) (10) On the structure of the Palaeozoic seed Mitrospermum compressum (Will.). Ann. Bot. vol. xxiv. p. 491. (14) A note on Trigonocarpus. Ibid. vol. xxv. p. 195. - Arber, E. A. Newell. (02) Notes on the Binney collection of Coal-Measure Plants. Pt. iii. The type-specimens of Lyginodendron oldhamium (Binney). Proc. Camb. Phil. Soc. vol. x1. pt. iv. p. 281. (03) On the roots of Meduilosa anglica. Ann. Bot. vol. Xvit-. p. 425. (03?) Discussion on Dr Kurtz’s paper (1903). Quart. Journ. Geol. Soc. vol. LIx. p. 26. (04) Cupressinoxylon Hookeri sp. nov. a large silicified tree from Tasmania. Geol. Mag. [v], vol. 1. p. 7. (05) On some new species of Lagenostoma, a type of Pterido- spermous seed from the Coal Measures. Proc. R. Soc. vol. txxi. B, p. 245. (07) On Triassic species of the genera Zamites and Pterophyllum, types of fronds belonging to the Cycadophyta. Trans. Linn. Soc. vol. vit. pt. vii. p. 109. (08) On a new Pteridosperm possessing the Sphenopteris type of foliage. Ann. Bot. vol. xxtt. p. 57. (09) On the Fossil Plants of the Waldershare and Fredville series of the Kent Coalfield. Quart. Journ. Geol. Soc. vol. Lxv. p. 21. (09?) On the affinities of the Triassic plant Yuccites vogesiacus Schimp. and Moug. Geol. Mag. [v], vol. vz. p. 11. (12) On Psygmophyllum majus sp. nov. from the Lower Carboni- ferous rocks of Newfoundland, together with a Revision of the genus and Remarks on its affinities. Trans. Linn. Soc. vol. vit. p. 391. (12?) Fossil Plants from the Kent Coalfield. Geol. Mag. [v], vol. 1x. p. 97. (13) A preliminary note on the Fossil] Plants of the Mount Potts beds, New Zealand, collected by Mr D. G. Lillie, Biologist to Capt. Scott’s Antarctic Expedition in the ‘Terra Nova.” Proc. R. Soc. vol. LXxxvi. p. 344. (13?) The structure of Dadoxylon Kayi. Quart. Journ. Geol. Soc. vol. LXIx. p. 454. (14) A Revision of the Seed impressions of the British Coal Measures. Ann. Bot. vol. xxvut. p. 81. (14?) On the Fossil Flora of the Kent Coalfield. Quart. Journ. Geol. Soc. vol. LXx. p. 54. Arber, E. A. Newell and J. Parkin. (07) On the origin of Angiosperms. Journ, Linn. Soc. vol. XXxXvutt. p. 29. (08) Studies on the Evolution of the Angiosperms. Ann. Bot. vol. xxi p. 489. Arnoldi, W. (01) Beitrige zur Morphologie einiger Gymnospermen. Bull. Nat. Moscow, No. 4, 1900. REFERRED TO IN THE TEXT 475 Bailey, I. W. (09) The structure of the wood in the Pineae. Bot. Gaz. vol. XLVI. p. 47. —- (11) A Cretaceous Pityoxylon with marginal tracheids. Ann. Bot. vol. xxv. p. 315. Baily, W. H. (69) Notice of Plant-remains from Beds interstratified with the Basalts in the county of Antrim. Quart. Journ. (eol. Soc. vol. xxv. p. 357. - Bain, F. and Sir W. Dawson. (85) Notes on the Geology and Fossil Flora of Prince Edward Island. Canadian Rec. Sci. vol. 1. (1884-83) p. 154. Baker, R. T. and H. C. Smith. (10) Research on the Pines of Australia. Dpt. Public Instruction, Tech. Educ. Ser. No. 16. Sydney. Bancroft, Nellie. (13) On some Indian Jurassic Gymnosperms and Rhexoxylon africanu UM, A. DEW Medullosean sean stem. Trans. Linn. Soc. - vol. Vit. pt. i. p. 69. —— (14) Pteridosperm Anatomy and its relation to that of the Cycads. _ New Phyt. vol. xu. p. 41. Barber, C. A. (92) On the nature and development of the corky excres- cences on stems of Zanthoxylum. Ann. Bot. vol. v1. p. 155. —— (98) Cupressionorylon vectense; a fossil Conifer from the Lower Greensand of Shanklin, in the Isle of Wight. Ann. Bot. vol. xm. - p. 329 Bartholin, C. T. (94) Nogle i den bornholmske Juraformation fore- kommende Planteforsteninger. Bot. Tidskrift (Copenhagen), Bd. XIX. p. 87. —— (10) _Planteforsteninger fra Holsterhus paa Bornholm. Danmarks Geol. Unders. 1. Raek. No. 24. Bartlett, A. W. (13) Note on the occurrence of an abnormal bispo- rangiate strobilus of Larix europaea DC. Ann. Bot. vol. xxvm. p. 575. Bassler, H. (16) A Cycadophyte from the North American Coal Measures. Amer. Journ, Sci. vol. xt. p. 21. Bayer, A. (08) Zur Deutung der weiblichen Bliiten der Cupressineen nebst Bemerkungen iiber Cryptomeria. Beiheft Bot. Cent. Bd. xxi. Abt. 1. p. 27. Beissner, L. (91) Handbuch der Nadelholzkunde. Berlin. Bennett, J. J.andR. Brown. (52) Plantae Javanicae rariores. London, 1838-52. Benson, Margaret. (08) On the contents of the pollen-chamber of a specimen of Lagenostoma ovoides. Bot. Gaz. vol. Lv. p. 409. — (12) Cordaittes Felicis, sp. nov., a Cordaitean leaf from the Coal Measures of England. Ann. Bot. vol. xxvi. p. 202. —— (14) Sphaerostoma ovale (Conostoma ovale et intermedianum, Williamson), a Lower Carboniferous ovule from Pettycur, Fife- shire, Scotland. Trans. R. Soc. Edinburgh, vol. L. pt. i. No. i. p. 1. Benson, M. and E. J. Welsford. (09) The morphology of the ovule and female flower of Juglans regia and of a few allied genera. Ann. Bot. vol. xxuI. p. 623. 476 LIST OF WORKS Berger, R. (48) De fructibus et seminibus ex formatione lithanthracum. Diss. Inaug. Vratislaviae. Bergeron, J. (84) Note sur les strobiles du Walchia piniformis. Bull. soc. géol. France [3], Tome xu. p. 583. Berridge, BE. M. (11) Onsome points of resemblance between Gnetalean and Bennettitean seeds. New Phyt. vol. x. p. 140. —— (12) The structure of the female strobilus in Gnetum Gnemon. Ann. Bot. vol. xxv. p. 987. Berridge, Emily M. and Elizabeth Sanday. (07) Oogenesis and embryo- geny in Ephedra distachya. New Phyt. vol. vi. p. 127.. Berry, E. W. (03) The Flora of the Matawan Formation. Bull. New York Bot. Gard. vol. 11. No. 9, p. 45. ; —— (05) Additions to the fossil Flora from Cliffwood. Bull. Torrey Bot. Club, vol. xxxi1. p. 43. — (06) Contributions to the Mesozoic Flora of the Atlantic Coastal Plain. I. /bid. vol. xxxut. p. 33. —— (07) Coastal Plain Amber. Torreya, vol. vu. p. 4. —— (07%) Contributions to the Pleistocene Flora of North Carolina. Journ. Geol. vol. xv. No. 4, p. 338. — (08) Some Araucarian remains from the Atlantic Coastal Plain. Bull. Torrey Bot. Club, vol. xxxv. p. 249. —— (08?) A Mid-Cretaceous species of Torreya. Amer. Journ. Sci. vol. xxv. p. 382. ; —— (09) A Miocene Flora from the Virginian Coastal Plain. Journ. Geol. vol. Xvi. p. 19. —— (09%) Pleistocene Swamp deposits in Virginia. Amer. Nat. vol. xLur. p. 432. — (10) A revision of the Fossi] Plants of the genus Nageiopsis of Fontaine. Proc. U. S. Nat. Mus. vol. xxxvit. p. 185. —— (10°) The epidermal characters of Frenelopsis ramosissimus. Bot. Gaz. vol. L. p. 305. —— (10°) Additions to the Pleistocene Flora of New Jersey. Torreya, vol. x. p. 261. — (104) Contributions to the Mesozoic Flora of the Atlantic Coastal Plain. V. North Carolina. Bull. Torrey Bot. Club, vol. xXxxvut. p. 181. — (ll) The Lower Cretaceous deposits of Maryland. (Berry, Clark, and Bibbin.) Maryland Geol. Surv. g —— (11?) A Lower Cretaceous species of Schizaeaceae from Eastern North Ameiica. Ann. Bot. vol. xxv. p. 193. —— (115) The Flora of the Raritan Formation. Geol. Surv. New Jersey, Bull. 3. : — (1l*) A Revision of several genera of Gymnospermous plants from the Potomac group in Maryland and Virginia. Proc. U. 8. Nat. Mus. vol. xu. p. 289. —— (12) The age of the plant-bearing shales of the Richmond coal- field. Amer. Journ. Sci. vol. XXXIV. p. 224. REFERRED TO IN THE TEXT 477 Berry, E. W. (12?) Notes on the genus Widdringtonites. Bull. Torr. Bot. Club, vol. Xxxrx. p. 341. —— (128) Contributions to the Mesozoic Flora of the Atlantic Coastal Plain. VII. Texas. Bull. Torrey Bot. Club, vol. xxxtx. p. 387. —— (124) Pleistocene plants from the Blue Ridge in Virginia. Amer. Journ. Sci. vol. XXXIV. p. 218. —— (14) The Upper Cretaceous and Eocene floras of South Carolina and Georgia. U.S. Geol. Surv. Professional papers, No. 84. —— (15) The Mississippi River Bluffs at Columbus and Hickman, Kentucky, and their Fossil Flora. Proc. U. S. Nat. Mus. vol. xuvit. p. 293. —— (16) The Geological history of Gymnosperms. The Plant World, vol. XIx. p. 27. ; = —— (17) Contributions to the Mesozoic Flora of the Atlantic Coastal Plain, XII. Arkansas. Bull. Torr. Bot. Club, vol. xut1v. p. 167. Bertrand, C. E. (74) Anatomie comparée des tiges et des feuilles chez les Gnétacées et les Coniféres. Ann. Sct. nat. [v], vol. xx. p. 5. —— (83) Note sur le genre Vesquia, Taxinée fossile du Terrain: Aachénien de Tournai. Bull. soc. bot. France [3], Tome v. p. 293. —— (89) Les Poroxylons végétaux fossiles de l’époque houilliére. Ann. soc. Belg. de Microscopie, Tome xt. Fase. i. —— (98) Remarques sur la structure des grains de pollen de Cordaites. Assoc. Frang. pour Pavanc. des Sci. Nantes (1898), p. 436. —— (07) Les caractéristiques du genre Rhabdocarpus d’aprés les préparations de la collection B. Renault. Bull. soc. bot. [4], Tome vu, p. 654. —-- (072) Les caractéristiques du genre Diplotesta de Brongniart. Bull. soc. bot. France [4], tome vu. p. 388. «= —— (07%) Les caractéristiques du genre Leptocaryon de Brongniart. », Ibid. p. 452. ‘i —— (074) Les caractéristiques du genre Taxospermum de Brongniart. Ibid. p. 213. —— (075) Remarques sur le Tarospermum angulosum. Compt. rend. @ assoc. Frang. pour Vavanc. des sci. (Reims, 1907), p. 410. — (08) Les caractéristiques du Cycadinocarpus angustodunensis de B. Renault. Bull. soc. bot. France [4], tome vit. p. 326. —— (08%) Lescaractéristiques du genre Cardiccarpus d’aprés les graines . silicifiées étudiées par Ad. Brongniart et B. Renault. Ibid. p. 391. —— (08%) Laspécification des Cardiocarpus de la collection B. Renault. _ Ibid. p. 454. —— (09) Sur le genre Compsotesta de Ad. Brongniart. Ann. Jard. Bot. Buit. [2], suppl. 3. —— (ll) Le Bourgeon femelle des Cordaites. Nancy. Bertrand, C. E. et B. Renault. (82) Recherches sur les Poroxylons. Arch. bot. du Nord de la France, vol. u. p. 243. Bertrand, P. (08) Surles stipes de Clepsydropsis. Compt. Rend. Nov. 16, 1908. 478 LIST OF WORKS - Bertrand, P. (11) Structure des stipes d’Asterochlaena laxa Sterzel. Meém. soc. géol. Nord, Tome vu. i. — (13) Les Fructifications de Neuroptéridées recueillies dans le terrain houiller du Nord de la France. Ann. soc. géol. Nord, Tome xu. p. 113. —— (14) Etat actuel de nos Connaissances sur les Genres ‘Cladoxylon’ et ‘Steloxylon.’ Compt. Rend. de lassoc. frang. pour ? Avancement des Sciences (Havre, 1914), p. 446. Beust, F. (85) Untersuchung iiber fossile Holzer aus Grénland. Neue Denisch. allgem. Schweiz. Ges. gesammt. Naturwiss. Bd. xx1x. Binney, E. W. (66) On Fossil wood in calcareous nodules found in the upper foot coal near Oldham. Proc. Lit. Phil. Soc. Manchester, vol. v. p. 113. Bleicher and Fliche. (92) Contribution & l’étude des Terrains Tertiaires @ Alsace. Bull. soc. géol. France [3], Tome xx. p. 375. Bodenbender, W. (96) Beobachtungen iiber Devon- und Gondwana Schichten in der Argentinischen Republik. Zeitsch. Deutsch. geol. Ges. Bd. xivit. p. 743. -—— (02) Contribucion al Conocimiento de la Bescon ilera de San Juan de Mendoza. Bol. Acad. Nac. Cienc. Cordoba, vol. xvu. p. 203. Boodle, L. A. (15) Concrescent and solitary Foliage-leaves in Pinus. New Phyt. vol. xtv. p. 19. Boodle, L. A. and W. C. Worsdell. (94) On the comparative anatomy of the Casuarineae, with special reference to the Gnetaceae and Cupuliferae. Ann. Bot. vol. vit. p. 231. Boulay. (79) Recherches de paléontologie végétale dans le terrain houiller du Nord de la France. Ann. soc. scient. Bruxelles, ann. iv. pt. 2, 1880. —— (87) Notice sur la Flore tertiaire des environs de Privas (Ardiche), Bull. soc. bot. France, Tome xxxiv. p. 227. —— (88) Notice sur les Plantes fossiles des grés tertiaires de Saint- Saturnin (Marne et Loire). Journ. Bot. Ann. 2, p. 921. Bower, F.O. (81) On the germination and histology of the seedlings of Welwitschia mirabilis. Quart. Journ. Micr. soc. vol. Xx1. pp. 15, 571. —— (82) The germination and embryogeny of Gnetum Gnemon. Ibid. vol. xxi. [N.S.], p. 277. —— (84) On the structure of Rhynchopetalum montanum. Journ. Lin. Soc. vol. xx. p. 440. , —— (12) Studies in the Phylogeny of the Filicales. II. Lophosira, and its relation to the Cyatheoideae and other Ferns. Ann. Bot. vol. XXvI. p. 269. As Bowerbank, J. 8. (40) History of the Fossil fruits and seeds of the London Clay. London. Braun, A. (75) Die Frage nach der Gymnospermie der Cycadeen. Monatsber. K. Preuss. Akad. Wiss. Berlin, p. 289. —— (75°) Die Diagnosen dreier im Jahre 1873 von G. Wallis in Neu Granada entdeckter Cycadeen. Ibid. p. 376. REFERRED TO IN THE TEXT 479. Braun, C.F.W. (43) Beitrige zur Petrefactenkunde Bayreuth (Graf zu Miinster), Heft vi. Bayreuth. — (47) Die Fossile Gewichse aus den Grenzschichten zwischen dem Lias und Keuper des neu aufgefundenen Pflanzenlagers in dem Steinbriiche von Veitlahm hei Culmbach. Flora, p. 81. —— (49) Beitrage zur Urgeschichte der Pflanzen. VI. Weltrichia eine neue Gattung fossiler Rhizantheen. Progr. iit. Jahresber. K. Kreis-Landwirthsch. und Gewerbschule zu Bayreuth. Brauns, D. (66) Der Sandstein bei Seinstedt unweit des Fallsteins und die in ihm vorkommenden Pflanzenreste. Paleont. Bd. 1x. p. 47. Brenchley, Winifred E. (13), On Branching specimens of Lyginodendron Oldhamium Will. Journ. Linn. Soc. vol. xii. p. 349. Bristow, H. W. (62) The Geology of the Isle of Wight. Mem. Geol. Surv. Great Britain. Brongniart, A. (25) Observations sur les végétaux fossiles renformés dans les grés de Hoer en Scanie. Ann. Sci. nat. vol. tv. p. 200. —— (28) Essai d’une Flore du grés bigarré. Ann. Sci. nat. vol. v. “3 p. 435. : —— (33) Notice sur une Conifére fossile du terrain d’eau douce de Vile d’Iliodroma. Ann. Sci. nat. tome xxx. p. 175. —— (74) Etudes sur les graines fossiles trouvées & I’état silicifié dans le terrain Hoviller de Saint Etienne. Ann. Sci. nat. tome xx. [5], . 234. — *(8l) Recherches sur les graines fossiles silicifiées. Paris. Bronn, H.G. (58) Beitrage zur triassischen Schiefer von Raibl. Neues Jahrb. Min. p. 129. Brooks, F. T.and A, Sharples. (14) Pink disease. Bull. No. 21, Depart. Agric. Fed. Malay States. Brooks, F. T. and W. Stiles. (10) The structure of Podocarpus spinulosus (Smith) R. Br. Ann. Bot. vol. xx1v. p. 305. Buchman, J. (45) Outline of the Geology of the neighbourhood of Cheltenham (in collaboration with R. I. Murchison and H. E, Strickland). London. Buckland, W. (28) On the Cycadecideae, a Family of Fossil Plants found in the Oolite quarries of the Isle of Portland. Trans. Geol. Soc. [2], vol. 1. p. 395. —— (37) Geology and Mineralogy considered with reference to Natural Theology. London. Burckhardt, C. (11) Bemerkungen zu einigen Arbeiten von W. Gothan und A. G. Nathorst. Cent. Min. Geol.; Paleont. p. 442. Burgestein, A. (06) Zur Holzanatomie der Tanne, Fichte und Liarche. Ber. deutsch. Bot. Ges. Bd. xx1v. Heft vi. p. 295. —— (08) Vergleichende Anatomie des Holzes der Koniferen. Wveesner- Festschrift, Wien. Burlingame, L. (08) The staminate cone and male gametophyte of Podocarpus. Bot. Gaz. vol. xuvi. p. 161. 480 LIST OF WORKS Burlingame, L. (13) The Morphology of Araucaria brasiliensis. Bot. Gaz. vol. Lv. p. 97. (15) The Morphology of Araucaria brasiliensis. Ibid. vol. LIx- 7. p. 1 (152) The Origin and Relationships of the Araucarians. Ibid. vo]. Lx. p. 1. . Butterworth, J. (97) Some further investigations of Fossil seeds of the genus Lagenostoma (Williamson) from the Lower-Coal Measures, Oldham. Mem. Proc. Manchester Lit. and Phil. Soc. vol. X11. ix. p. 1. Caldwell, 0. W. (07) Microcycas calocoma. Bot. Gaz. vol. xxiv. p. 118. Caldwell, O. W. and C. F. Baker. (07) The identity of Microcycas calocoma. Ibid. vol. xm. p. 130. Cambier, R. et A. Renier. (10) Psygmophyllum Delvali n. sp. du Terrain houiller de Charleroi. © Ann. soc. géol. Belg. Tome 1. p. 23. (Mém. in 4to.) Capellini, G. and Conte E.Solms-Laubach. (92) I Tronchi di Bennettitee dei Musei Italiani’ Mem. Reale Acad. Sci. Istit. Bologna [5], tom. I. p. 161. Caporn, A. St. C. (16) A note on the male inflorescence of a species of Gne/um from Singapore. Ann. Bolus Herb. vol. 1. pt. i. Carano, H (04) Contribuzione alla conoscenza della Morfologia e dello sviluppo del fascio vascolare delle foglie delle Cicadacee. Ann. di Bot. vol. 1. p. 109 (Rome). Carpentier, A. (11) Sur quelques fructifications et inflorescences du Westphalien du Nord dela France. Rev. Gén. Bot. tome xxl. p. 1. —— (13) Contribution & étude du Carbonifére du Nord de la France. Meém. soc. géol. du Nord, tome vit. ii. p. 1. Carruthers, W. (66) On Araucarian cones from the Secondary beds of Britain. Geol. Mag. vol. mt. p. 249. —— (66?) On some fossil Coniferous fruits. Ibid. vol. m1. p. 534. —— (67) On Cycadeoidea Yatesii sp. nov. a fossil Cycadean stem from the Potton sands, Bedfordshire. Ibid. vol. rv. p. 199. —— (67?) On Gymnospermous Fruits from the Secondary rocks of Britain. Journ. Bot. vol. v. p. 1. —— (67%) On some Cycadean Fruits from the Secondary rocks of Britain. Geol. Mag. vol. 1v. p. 101. —— (68) British Fossil Pandanaceae. Ibid. vol. v. p. 153. —— (69) On Beania, a new genus of Cycadean Fruit from the Yorkshire Oolite. Ibid. vol. v1. p. 1. —— (69°) On some undescribed Coniferous fruits from the Secondary rocks of Britain. Ibid. vol. v1. p. 1. —— (70) On Fossil Cycadean stems from the Secondary rocks of Britain. Trans. Linn. Soc. vol. xxvi. p. 675. —— (71) On two undescribed Coniferous fruits from the Secondary rocks of Britain. Geol. Mag. vol. vt. p. 1. —— (77) Description of a new species of Araucarites from the Coralline Oolite of Malton. Quart. Journ. Geol. Soc. vol. xxxt1. p. 402. REFERRED TO IN THE TEXT 481 Carruthers, W. (93) On Cycas Taiwaniana, sp. nov, and C. Seemannt ; R. Br. Journ. Bot. vol. xxxi. p. 1. Carter, M. Geraldine. (11) A Reconsideration of the origin of Trans- fusion-tissue. Ann. Bot. vol. xxv. p. 975. Caspary, R. and R. Triebel. (89) Linige fossile Holzer ses : K. Preuss..Geol. Landesanstalt, Bd. 1x. Heft iii. p. 113. Celakovsky, L. (82) Zur Kritik der Ansichten von den Fruchtschuppe der Abietineen. Abh. K. béhm. Ges. Wiss. Prag [vi], Bd 11. Chamberlain, C. J. (06) The ovule and female gametophyte of Dioon. . Bot. Gaz. vol. xuu. p. 321. —— (09) Dioon spinulosum. Ibid. vol. xvi. p. 401. — (10). See Coulter and Chamberlain. — (10?) Fertilization and embryogeny in Dioon edule. Bot. Gaz. vol. L. p. 415. — (1l) The adult Cycad Trunk. Jbid. vol. Liz. p. 81. —-- (12) Morphology of Ceratozamia. Ibid. vol. Lu. p. 1. —— (12?) Around-the-world Botanical Excursion. Pop. Sci. Monthly, vol. LXxx1. p. 417. ‘ —— (128) Two species of Bowenia. Bot. Gaz. vol. Liv. p. 419. — (13) Macrozamia Moorei, a connecting link between living and fossil Cycads. Bot. Gaz. vol. Lv. p. 141. és Chapman, F. (09) Jurassic Plant-remains from Gippsland, Pt. ii. Ree. Geol. Surv. Victoria, vol. 11. pt. i. p. 103. Chrysler, M. A. (15) The Medullary rays of Cedrus. Bot. Gaz. vol. LIx. p. 387. Church, A. H. (14) On the Floral Morphology of Welwitschia mirabilis (Hooker). Phil. Trans. R. Soc. vol. 205, p. 115. ; Cockerell, T. D. A. (06) The Fossil Flora and Fauna of the Florissant (Colorado) shales. Univ. Colorado Series, vol. m1. No. 3. —— (08) Description of Tertiary Plants. TI. Amer. Journ. Sct. vol. xxvi [4], p. 537. —— (087). The Fossil Flora of Florissant, Capias: Bull. Amer. Mus. Nat. Hist. vol. xxiv. p. 71. _——. (08°) Florissant; a Miocene Pompeii. Pop. Sci. Monthly (Aug. 1908, p. 112). Coemans, E. (66) Description de la flore fossile du premier étage du terrain Crétacé du Hainaut. Mém. Acad. R. Belg., tome XXXVI. Coker, W. C. (03) On the gametophyte and embryo of Tazodium. Bot. Gaz. vol. XxxvI. p. 1. — (09) Vitality of Pine seeds and the delayed opening of cones. _ Amer. Nat. vol. xu. p. 677. Compter, G. (94) Die fossile Flora des untern Keupers von Ostthiiringen. Zeitsch. fiir Naturwiss. Leipzig, Bd. uxvu. p. 205. —— (03) Cycadeenfriichte aus der Lettenkohle von ee Zeitsch. fiir Naturwiss. Stuttgart, Bd. uxxv. p. 171. Compton, R. H. (08) See South and Compton. s. IV 31 482 LIST OF WORKS Conwentz, H. (78) Ueber ein tertiires Vorkommen Cypressenartiger Holzer bei Calistoga in Californien. Neues Jahrb. Min. p. 800. — (82) Fossile Holzer aus der Sammlung der K6nig. geol. Landes- anstalt zu Berlin. Jahrb. K. preuss. geol. Land. Bergakad. Berlin fir das Jahr 1881, p. 144. ; : — (85) Sobre algunos arboles fosiles del Rio Negro. Bol. Acad. Nac. Ciene. Cordoba, tom. vi. p. 575. — (86) Die Angiospermen des Bernsteins. Danzig. —— (89) Ueber Thyllen und Thyllen-ahnliche Bildungen, vornehmlich im Hélze der Bernsteinbiume. Ber. Deutsch. Bot. Ges. Bd. vit. p. (34). —— (92) Untersuchungen iiber fossile Hélzer Schwedens. K. Svensk. Vetenskapsakad. Hand. Bd. xxiv. No. 13. —— (01) The past history of the Yew in Great Britain and Ireland. Rep. 71st Meeting Brit. Assoc. (Glasgow), p. 839. Coulter, J.M. and C.J. Chamberlain. (03) The Embryogeny of Zamia. Bot. Gaz. vol. xxxv. p. 184. —— (10) Morphology of Gymnosperms. Chicago. Cramer, C. (68) Fossile Holzer der Arctischen Zone. Heer’s Foss. Flor. Aret, vol. I. p. 167. ae af Crié, L. (89) Beitrige zur Kenntniss der fossilen Flora einiger Inseln des Siidpacifischen und Indischen Oceans. Pal. Abhand. (Dames and Kayser) (N.F.), Bd. 1. Heft ii. Daguillon, A. (90) Recherches morphologiques sur les feuilles des Coniféres. Rev. Gén. Bot. tome 11, p. 154. Dawson, J.W. (46) Notices of some Fossils found in the Coal Formation of Nova Scotia. Quart. Journ. Geol. Soc. vol. 11. p. 132. —— (62) On the Flora of the Devonian period in North-eastern America, Jbid. vol. xvut. p. 296. —— (63) Further Observations on the Devonian Plants of Maine, Gaspé, and New York. Ibid. vol. xrx. p. 458. -—— (63°) Synopsis of the Flora of the Carboniferous Period in Nova Scotia. Phe Canadian Nat: and Geologist, vol. vu. p. 431. —— (81) Notes on New Erian (Devonian) plants. Quart. Journ. Geol. Soc. vol. XXXv1I. p. 299. —— (85) On the Mesozoic Floras of the Rocky Mountains Region of Canada. Trans. R. Soc. Canada, sect. tv. p. 1. —— (90) On new plants from the Erian and Carboniferous, and on the characters and affinities of Palaeozoic Gymnosperms. Canadian Rec. Sci. vol. iv. p. 1, —— (93) On new species of Cretaceous plants from Vancouver Island. Trans. R. Soc. Canada, sect. iv. p. 53. Dawson, Sir J. W. and D, P. Penhallow. (91) Note on the specimens of Fossil wood from the Erian (Devonian) of New York and Kentucky. The Canadian Rec. Sci. vol. 1v. p. 242. Depape, G. (13) Sur la présence du Ginkgo biloba L. dans le Pliocéne ‘nférieur de Saint-Marcel-d’Ardéche. Compt. Rend. vol. 157, p. 957. REFERRED TO IN THE TEXT 483 Depape, G. and A. Carpentier. (13) Présence des genus Gnetopsis B. Ren. and R. Zeill. et Urnatopteris Kidst. dans le Westphalien du Nord de la France. Ann. soc. géol. du Nord, tome xuu. p. 294. — (15) Sur quelques graines et fructifications du Westphalien du Nord de la France. Rev. Gén. Bot. vol. xxvm. p. 321. Dorety, Helen A. (08) The embryo of Ceratozamia, a physiological study. Bot. Gaz. vol. xiv. p. 412. — (08?) The secdling of Ceratozamia. Ibid. vol. xivr. p. 205. —— (09) Theextrafascicular cambium of Ceratozamia. Ibid. vol. XLVI. p. 149. — (09?) Vascular anatomy of the seedling of Microcycas Calocoma. Ibid. p. 139. Dorrien-Smith, A.A. (11) A Botanizing expedition to West Australia. Journ. R. Hort. Soc. vol. XXXVI. p. 285. Douvillé, H. et R. Zeiller. (08) Sur le terrain houiller du Sud Oranais. Compt. Rend. tome CXLVI. p. 732. Drude, O. (90) Handbuch der Pflanzengeographie. Stuitgart. Diimmer, R. (12) Podocarpus formosensis. Gard. Chron. Oct. 19, p. 295. Dun, W. S. (10) Notes on some Fossil plants from the roof of the coal seam in the Sydney Harbour Colliery. Journ. Proc. R. Soc. New South Wales, vol. xt1v. p. 615. Duns, J. (72) On Cardiocarpon. Proc. R. Soc. Edinburgh, p. 692. DuSdnek, F. (13) Spaltdffnungen der Cycadaceen. (Abstract in the Bot. Cent. Bd. cxxv. p. 340.) Dusén, P, (99) Uher die Tertidre Flora der Magellans-Lander. Wiss. Ergeb. Schwed. Exped. nach den Mageillanslandern unter Leitung von O. Nordenskjéld, Bd. 1. No. iv. p. 87. —— (08) Uber die Tertiire Flora der Seymour-Insel. Wiss. Ergeb. Schwed. siidpolar-Exped. 1901-03, Bd. m1. Lief. iii. p. 1. Duthie, Augusta V. (12) Anatomy of Gnetum africanum. Ann. Bot. vol. xxvi. p. 593. Dutt, C. P. (16) Pityostrobus macrocephalus, L. and H. A Tertiary Cone showing Ovular structures. Ann. Bot. vol. xxx. p. 529. Eames, A. J. (13) The Morphology of Agathis australis. Ann. Bot. vol. xxvu. p. 1. Eberdt, O. (94) Die Braunkohlen Ablagerungen in der Gegend von Senftenberg. Jahrb. K. Preuss. Geol. Land. Bergakad. Berlin, Bd. xrv. p. 212. Eichler, A. W. (81) Uber die weiblichen Bliithen der Coniferen. Monats- ber. K. Akad. Wiss. Berlin, p. 1020. —— (82) Uber Bildungabweichungen bei Fichtenzapfen. Sitzber. K. Akad. Wiss. Berlin, p. 40. —— (89) Gymnospermae. Engler and Prantl; Die Natiirlichen Pflanzenfamilien, Teil ii. Hichwald, E. (53-68) Lethaea rossica. Stuttgart. Elkins, Marion G. and G. R. Wieland. (14) Cordaitean wood from the Indiana Black Shale. Amer. Journ. Sci. vol. xxxvu. p. 65. 31—2 484 LIST OF WORKS Elwes, H. J. (12) The Flora of Formosa. Gard. Chron. July 13, 1912, . 25. Elwes, H. J.and A. Henry. (06) The trees of Great Britain and Ireland, vol.1. Edinburgh. Endlicher, §. (40) Genera plantarum secundum ordines naturales disposita. Vindobonae, 1836-40. —“... (47) Synopsis Coniferarum. Sangaili. Engelhardt, H. (85) Die Tertidrflora des Jesuitengrabens bei Kundratitz in Nordbéhmen. Nov. Act. K. Leop.-Car. Deutsch. Akad. Natur- forsch. Bd. xiv. No. 3, p. 299. —— (91) Uber Tertiadrpflanzen von Chile. Abh. Senckenberg. naturforsch. Ges. p. 629. —— (12) Weiterer Beitrag zur Kenntniss der fossilen Tertiarflora Bosniens. Wiss. Mitt. aus Bosnien und der Herzegowina, Bd. xu. p. 593. Engelhardt, H. and F. Kinkelin. (08) Oberpliocene Flora und Fauna des Unter-Maintales. Abh. Senck. Naturforsch. Ges. Bd. xx1x. Heft iii. p. 151. Engler, A. (89) Engler and Prantl; Die Natiirlichen Pflanzenfamilien, ; Teil u. Abt. i. — (95) Die Pflanzenwelt Ost Afrikas, etc. Th. C. Berlin. —— (97) Engler and Prantl; Die Natiirlichen Pflanzenfamilien. P Nachtrag zu Teilen 1-1Vv. ‘Essner, B. (86) Ueber den diagnostischen Werth der Anzahl und Hohe der Markstrahlen bei den Coniferen. Abh. naturforsch. Ges. Halle, Bd. xvi. p. 1. Etheridge, R. (93) On the occurrence of a plant allied to Schizoneura in the Hawkesbury Sandstone. Rec. Geol. Surv. New South Wales, vol. UI. pt. iii, p. 74. Ettingshausen, C.von. (51) Ueber einige neue und interessante Taenio- pteris Arten aus den Sammlungen des Kais. Hof Mineralien Cabinetes und der K.k. geol. Reichsanstalt. Naturwiss. Abh. W. Haidinger, vol. Iv. p. 95. —— (52) Beitrag zur naheren Kenntniss der Flora der Wealdenperiode. Abh, K.k. geol. Reichs. Wien, Bd. 1. Abth. iii. No. 2, p. 1. —— (52?) Die Steinkohlenflora von Stradonitz. Jbid. Bd. 1. Abth. iii. No. 4. —— (55) Die Tertiare Flora von Haring in Tirol. Jbid. Bd. a. Abth. ii. No. 2. — (57) Die Fossile Flora von Kéflach in Steiermark. Jahrb. K.k. geol. Reichs. Bd. vit. p. 738. —— (58) Beitraége zur Kenntniss der fossilen Flora von Sotzka in Untersteiermark. Sttz. K. Akad. Wiss. Wien, Bd. xxvut. p. 471. —— (67) Die Kreideflora von Niederschoena in Sachsen. Jbid. Bd. Ly. Abth. i. p. 235. —— (67°) Die fossile Flora des Tertiaér-Bechens von Bilin. Denksch. Wiss. Akad. Wien, Bd. xxv1. p. 79. REFERRED TO IN THE TEXT 485 Ettingshausen, C. von. (70) Beitrag zur Kenntniss der Tertidrflora Steiermarks. itz. K. Akad. Wien, Bd. ux. Abth. i. p. 17. —— (72) Die Fossile Flora von Sagor in Krain. Denksch. Wiss. Akad. Wien, Bd. xxxu. p. 32. —— (78) Beitrag zur Erforschung der Phylogenie der Pflanzenarten. Ibid. Bd. xxxvut. p. 65. —— (79) Report on Phyto-Palaeontological Investigations of ‘the Fossil] Flora of Sheppey. Proc. R. Soc. vol. xxtx. p. 388. —— (80) Report on Phyto-Palaeontological Investigations of the Fossil Flora of Alum Bay. Ibid. vol. xxx. p. 228. — (85) Die Fossile Flora von Sagor in Krain. Denksch. Wiss. Akad. Wien, Bd. t. p. 1. —— (86) Beitrage zur Kenntniss der Tertiarflora Australiens. Ibid. Bd. ii. p. 81. —— (87) Beitrage zur Kenntniss der Fossile Flora Neuseelands. Denksch. K. Akad. Wiss. Wien, Bd. uu. p. 3. —— (88) Contributions to the Tertiary Flora of Australia. Mem. geol. Surv. N.S.W. Pal. No. 2. : —— (88?) Die Fossile Flora von Leoben in Steiermark. Denksch. K. Akad. Wiss. Wien, Bd. tiv. Abth. 1. p. 261. — (90) Die Fossile Flora von Schoenegg bei Wies in Steiermark. Ibid. Bd. tvu. p. 61. Ewart, A. J. (08) On the longevity of seeds. Proc. R. Soc. Victoria, vol. xxr. [N.S.1, pt. i. p. 1. Feistmantel, 0. (72) Beitrag zur Kenntniss der Ausdehnung des sogen- nanten Wyraner Gasschiefers und seiner Flora. Jahrb. K.k. geol. Reichs. Wien, Bd. xxi. p. 289. , —— (76) Notes on the age of some Fossi] Floras in India. Rec. Geol. Surv. Ind. vol. 1x. pt. iv. p. 115. —— (76?) Jurassic (Oolitic) Flora of Kach. Fossil Flora of the Gond- wana System, pt. i. vol. 11. 1880. —— (77) Notes on Fossil Floras in India. Rec. Geol. Surv. Ind. vol. x. pt. iv. p. 196. —— (77°) Jurassic (Liassic) Flora of the Rajmahal group in the Rajmahal hills. Foss. Flor. Gond. Syst. vol. 1. pt. ii. —— (77°) Jurassic (Liassic) Flora of the Rajmahal group from Golapili, near Ellore, 8. Godaveri. bid. pt. iii. —— (774) Flora of the Jabalpur group (Upper Gondwanas) in the Son-Narbada region. Jhid. vol. 11. pt. ii. —— (77°) Ueber die Gattung Williamsonia Carr. in Indien. Palaeon- tolog. Beit. Palaeontograph. Suppl. 1. Lief. iii. —— (79) Upper Gondwana Flora of the outliers on the Madras coast. Foss. Flor. Gond. Syst. vol. 1. pt. iv. —— (79%) The Flora of the Talchir-Karharbari beds. Ibid. vol. ut. pt. i. —— (80) The Flora of the Damuda-Panchet Divisions. Jbid. vol. 11. pt. ii. 486 _ LIST OF WORKS Feistmantel, 0. (802) Note on the fossil genera Noeggerathia Sternberg, Noeggerathiopsis Feist. and Rhiptozamites Schmal. Rec. Geol. Surv. India, vol. x1. pt. i. p. 61. ; —— (80%) Further notes on the correlation of the Gondwana Flora with other Floras. Rec. Geol. Surv. Ind. vol. x1. pt. iii. p. 190. —— (81) Palaeontological notes from the Haz4rib4gh and Lohardagga Districts. Ibid. vol. xiv. pt. iii. p. 241. —— (812) The Flora of the Talchir-Karharbari beds. Foss. Flor. Gond. Syst. vol. 111. pt. i. —— (81%) The Flora of the Damuda-Panchet Divisions. Jbid. vol. m1. pt. iii. — (82) The Fossil Flora of the South Rewah Gondwana Basin. Ibid. vol. Iv. pt. i. —— (86) The Fossil Flora of some of the Coalfields in Western Bengal. Ibid. vo]. Iv. pt. ii. —— (89) Ubersichtliche Darstellung der geologisch-palaeontologischer Verhaltnisse Siid-Afrikas. Th. i. Abs. K. bohm. Ges. Wiss. [vii], Bd. 11. Felix, J. (82) Beitrage zur Kenntniss fossiler Coniferen-Hélzer. Engler’s Jahrb. Bd. m1. p. 260. —— (84) Die Holzopale Ungarns in Palaeophytologischen Hinsicht. Mitt. Jahrb. K. Ung. geol. Anst. Bd. vu. — (87) Untersuchungen iiber fossile Hélzer. Zeitsch. Deutsch. geol. Ges. p. 517. — (94) Untersuchungen iiber fossile Hélzer. Ibid. Heft 1. p. 79. — (96) Untersuchungen iiber fossile Hélzer. Ibid. Heft 1. p. 249. Fiedler, H. (57) Die Fossile Friichte der Steinkohlen-formation. Acad. Caes. Leop. Nov. Acta, Bd. xxv. p. 239. Fliche, P. (96) Etude sur la flore fossile de Argonne (Albien-Ceno- manien). Bull. soc. sci. Nancy. —— (97) Note sur les nodules et bois minéralisés trouvés & St Parres- ; les-vaudes (Aube) dans les grés verts infracrétacés. Mem. soc. Acad. de l Aube, tome Lx. —— (99) Note sur quelques fossiles végétaux de lOligocéne dans les Alpes Frangaises. Bull. soc. géol. France [3], tome xxvit. p. 466. —— (00) Contribution 4 la Flore fossile de la Haute-Marne (Infra- crétacée). Bull. soc. sci. Nancy. —— (00?) Note sur un bois fossile de Madagascar. Bull. soc. géol. France [3], tome xxvii. p. 470. —— (03) Note sur des bois silicifiés Permiens de la vallée de Celles ai (Vosges). (05) Note sur des bois fossiles de Madagascar. Bull. soc. géol. ae [4], tome v. p. 346. — (10) Flore Fossile du Trias en Lorraine et Franche-Comté avec des considérations finales par M. R. Zeiller. Paris. Fliche, P. and Bleicher. (82) Etude sur la flore de l’oolithe inférieure aux environs de Nancy. Bull. soc. sci. Nancy. REFERRED TO IN THE TEXT 487 Fliche, P. and R. Zeiller. (04) Note sur une florule Portlandienne des environs de Boulogne-sur-mer. Bull. soc. géol. France [4], tome Iv. p. 787. Fontaine, W. M. (93) Notes on some Fossil plants from the Trinity Division of the Comanche series of Texas. Proc. U. S. Nat. Mus. vol. Xvi. p. 261. Foxworthy, F.W. (11) Philippine Gymnosperms. Philipp. Journ. Sci. (C) Botany, vol. v1. No. 3, p. 149. Fraine, E. de. (12) On the structure and affinities of Sutcliffia, in the light of a newly discovered specimen. Ann. Bot. vol. XXvI. p. 1031. —— (14) On Medullosa centrofilis, a new species of Medullosa from the Lower Coal Measures. [bid. vol. xxvitt. p. 251. Fritel, P. H. and R. Viguier. (11) Etude anatomique de deux bois ; Kocénes. Ann. sci. nat. [9], tome xrv. p. 63. Fujii, K. (96) On the different views hitherto proposed regarding the morphology of the flowers of Ginkgo biloba. Bot. Mag. Tokyo, vo]. x. No. 109, p. 13. — (10) Some remarks on the Cretaceous Fossi! Flora and the causes of extinction. Bot. Mag. Tokyo, vol. xxiv. No. 284, p. 197. Fujioka, M. (13) Studien itiber den anatomischen Bau des Hélzes der japanischen Nadelbiume. Journ. Coll. Agric. Imp. Univ. Tokyo, vol. 1v. No. 4, p. 201. NGardner, J. 8. (86) A monograph of the British Eocene Flora, vol. 1. Gymnospermae. Palaeont. Soc. London. —— (86) Second Report of the Committee appointed for the purpose of reporting on the Fossil Plants of the Tertiary and Secondary beds of the United Kingdom. Rep. of the 56th Meeting Brit. Assoc. Geinitz, H. B. (42) Charakteristik der Schichten und Petrefacten des Sachsischen Kreidegebirges. Heft 11. Dresden und Leipzig. — (62) Dyas oder die Zechstein-formation und das Rothliegende, Heft mu. Leipzig. —— (63) Uberzweineue Dyadische Pflanzen. Neues Jahrb. Min. p. 525. —— (71) Uber fossile Pflanzen aus der Steinkohlen-Formation am Altai. Leipzig. —— (73) Versteinerungen aus dem Brandschiefer der unteren Dyas von Weissig bei Pillnitz in Sachsen. Neues Jahrb. Min. p. 681. —— (75) Uber neue Aufschliisse im Brandschiefer der unteren Dyas von Weissig bei Pillnitz in Sachsen. Jbid. p. 1. —— (80) Nachtrage zur Dyas I. Muittheil. aus dem K. Min.-geol. und Praehist. Mus. Dresden, Heft 3. Gerry, E. (10) The distribution of the Bars of Sanio in the Coniferales. Ann. Bot. vol. xxiv. p. 119. Geyler, T. and F. Kinkelin. (90) Oberpliocin Flora aus den Baugraben des Klarbeckens bei Niederrad etc. Abh. Senck. Naturforsch. Ges. Bd. xv. p. 1. Gibbs, L. S. (12) On the Development of the female strobilus in Podo- carpus. Ann. Bot. vol. xxvi. p. 515. 488 LIST OF WORKS Gliick, H. (02) Eine fossile Fichte aus dem Neckertahl. Mitt. Grossh. Bad. geolog. Landesanst. Bd. 1v. Heft iv. p. 399. Goc, M. J. le. (14) Observations on the centripetal and centrifugal xylems in the petioles of Cycads. Ann. Bot. vol. xxvui. p. 183. Goebel, K. (05) Organography of Plants. Pt. ii. Oxford. Goeppert, H.R. (40) Uber die neulichst im Basalttuff des hohen Seelbach- kopfes bei Siegen entdeckten bituminosen und versteinerten Hélzer, s0 wie iiber die der Braunkohlenformation tiberhaupt. Arch. fiir Min., Geog., etc. (Karsten und von Decken), Bd. xiv. p. 182. —— (41) Uber den Bernstein....Vebersicht den Arbeiten und Verdn- derungen der Schles. Ges. fiir Vaterlind. Kultur. Breslau. —— (412) Tazites scalariformis, eine neue Art fossilen Holzes. Arch. fiir Min., Geog., etc. Bd. xv. p. 727. —— (44) Ueber die fossilen Cycadeen tiberhaupt, mit Riicksicht auf die in Schlesien vorkommenden Arten. Uebersicht Arbeit. und Verdnd. Schlesisch. Ges. vat. Kult. 1843, p. 114. Breslau. —— (45) F. Wimmer’s Flora von Schlesien nebst einer Uebersicht der Fossilen Flora Schlesiens von H. R. Goeppert. Breslau. —— (452) Description des végétaux fossiles recueillis par M. P. de Tchihatcheff en Sibérie. Voyage scientifique dans [Altai oriental, p. 379. Paris. —— (46) Ueber die fossile Flora der mittleren Jurasckichten in Oberschlesien. Uebers. Arbeit. und Verind. Schles. Ges. vat. Kultur im Jahre 1845, Breslau, 1846, p. 139. —— (47) Zur Flora des Quadersandsteins in Schlesien. Nachtrag. Nov. Act. Ac. Caes. Leop.-Car. vol. xxit. p. 355. —— (50) Monographie der fossilen Coniferen. Naturwerkundige Ver- hand. Holland. Maatschap. Wettenschappen Haarlem. Leiden. —— (52) Fossile Flora des Ubergangsgebirges. Nova Acta Caes. Leop.-Carol. Nat. Cur. Bd. xxi. (supplement). —— (53) Ueber die gegenwartigen Verhaltnisse der Palaontologie in Schlesien, so wie iiber fossile Cycadeen. Denksch. Schles. Ges. fir Vaterland. Kultur. -—— (65) Die fossile Flora der Permischen Formation. Palaeont. Bd. 1. p. 1. —— (65%) Uber die fossile Kreideflora und ihre Leitpflanzen. Zeit. Deutsch. geol. Ges. Bd. xvir. p. 638. — (66) Beitrige zur Kenntniss fossilen Cycadeen. Neues Jahrb. Min. p. 129. — (80) Beitrage zur Pathologie und Morphologie fossiler Stimme. Palaeontol. [N.F.], Bd. vu. ii. p. 131. —— (81) Revision meiner Arbeiten iiber die Stamme der fossilen Coniferen, insbesondere der Araucariten, und iiber die Descendenz- lehre. Bot. Cent. Bd. v., v1. p. 378. Goeppert, H. R. and G. Stenzel. (81) Die Medulloseae. Palaeontol. (N.F.], Bd. vor. p. 113. — (88) Nachtrige zur Kenntniss der Coniferenhélzer der Palaeo- zoischen Formationen. Abhk. K. Preuss. Akad. Wiss. Berlin. REFERRED TO IN THE TEXT 489 Gomes, B. A. (65) Vegetaes Fosseis. Flora fossil do Terrens Carbonifero das visinhangas dio Porto, serra do Bussaio, e moinho d’ordem ; proximo a alcacer do Sal. Comm. geol. Portugal. Lisbon. Gordon, Marjorie. (12) Ray-tracheids in Sequoia sempervirens. New Phyt. voi. xt. p. 1. Gordon, W. T. (10) On a new species of Physostoma from the Lower Carboniferous rocks of Pettycur (Fife). Proc. Camb. Phil. Soc. vol. xv. pt. v. p. 395. —— (12) On Rhetinangiwm Arberi, a new genus of Cycadofilices from the Calciferous sandstone series. Trans. R. Soc. Edinb. vol. xuvit. ~ JZ pt. a p- 813. Gothan, (05) Zur Anatomie lebender und fossiler Gymnospermer- mee Abh. K. Preuss. geol. Landes. [N.F.], Heft xrrv. p. 1. —— (06) Die fossilen Coniferenhélzer von Senftenberg. Abh. K. Preuss. Geol. Landesanst. Bergakad. [N.F.], Heft xtv1. p. 155. —— (06?) Fossile Hélzer aus dem Bathonien von Russisch-Polen. Verhand. K. Russ. Min. Ges. zu St Petersburg [ii], Bd. xi1v. Lief. i. p. 435. —— (06°) Piceorylon Pseudotsugae als fossiles Holz. Potonié’s Abbild. und Beschreib. Foss. Pflanz. Lief. 1v. 80. —— (07) Uber die Wandlungen der Hoftiipfelung bei den Gymnosper- men im Laufe der geologischen Epochen und ihre physiologische ra Bedeutung. Stiz. Ges. Naturforsch. Freunde, No. 2, p. 13. (07?) Die Fossile Holzer von Konig Karls Land. K. Svensk. Vetenskapsakad. Hand. Bd. xuix. No. 10, p. 1. — (08) Die Fossile Hélzer von der Seymour- und Snow Hill-Insel. Wiss. Ergeb. Schwedis. Siidpolar-Exped. 1901-03, Bd. ut. Lief. viii. Stockholm. —— (08?) Die Frage der Klima-differenzirung im Jura und in deg Kreideformation im Lichte paléobotanischen Tatsachen. Jahrb. K. Preuss. Geol. Landes. fiir 1908, Bd. xxix. Th. ii. Heft 2, p. 220. , -—— (09) Uber Braunkohlenhélzer des rheinischen Tertiadrs. Jahrb. K. Preuss. geol. Land. Bd. xxx. Teil i. Heft 3, p. 516. —— (10) Die Fossile Holzreste von Spitzbergen. K. Svensk. Veten- skapsakad. Hand. Bd. xiv. No. viii. —— (11) Uber einige Permo-Carbonische Pflanzen von der unteren Tunguska (Sibirien). Zeitsch. Deutsch. Geol. Ges. Bd. x1. Heft 4, p. 418. —— (13) Die oberschlesische Steinkohlenflora. Teil 1 K. Preuss. geol. Landes, [N.F.], Heft txxv. Gourlie, W. (44) Notice of the Fossil Plants in the Glasgow Museum. Proc. Phil. Soc. Glasgow, vol. 1. 1844, p. 105. Grand’Eury, C. (00) Sur les tiges debout, les souches et racines de Cordaites. Compt. Rend. tome cxxx. (April 30). —— (04) Sur les graines des Neuroptéridées. Compt. Rend. tome OXXXIX. p. 23. ——- (04?) Sur les graines des Neuroptéridées. Ibid. p. 782. 490. | LIST OF WORKS Grand’Eury, C. (05) Sur les Rhabdocarpus, les graines et Pévolution des Cordaitées. did. tome CXL. p. 995. i —— (052) Sur les graines de Sphenopteris, sur l’attribution des Co- donospermum et sur V'extréme variété des ‘graines de fougéres.’ Ibid. p. 812. : —— (13) Recherches géobotaniques sur les foréts et sols fossiles et sur la végétation et la flore houilléres, en deux parties et dix livraisons. Pt. 1. Livr. ii., Paris et [nége. Graner, F. (94) Die geographische Verbreitung der Holzarten. I. Die Coniferen. Forstwissenschaft. Centralblatt, Berlin (August). Griffith, W. (59) Remarks on Gnetum. Trans. Linn. Soc. vol. Xxt1. pt. iv. p. 299. Groom, P. (10) Remarks on the Oecology of Coniferae. Ann. Bot. vol. xxiv. p. 241. —— (16) A Note onthe Vegetative Anatomy of Plerosphaera Fitzgeraldi F.v.M. Annals Bot. vol. xxx. p. 311. Groom, P. and W. Rushton. (13) The structure of the wood of East Indian species of Pinus. . Journ. Linn. Soc. vol. xut. p. 457. Groppler,R. (94) Vergleichende Anatomie des Hélzes der Magnoliaceen. 2 Biblioth. Bot. Bd. v1. Heft 31. Stuttgart. ; Grossenbacher, J.G. (15) Medullary spots and their cause. Bull. Torr. Bot. Club, vol. xii. p. 227. Guppy, H. B. (06) Observations of a Naturalist in the Pacific between 1896 and 1899. London. Gutbier, A. von. (49) Die Versteinerungen des Zechsteingebirges und Rothliegenden oder des Permischen Systems in Sachsen. Dresden and Leipzig. Halle, T. G. (10) A Gymnosperm with Cordaitean-like leaves from the Rhaetic beds of Scania. Arkiv fér Bot. Upsala, Bd. rx. No. 14. —— (12) On the occurrence of Dictyozamites in South America. Palaeobot. Zeitsch. Bd. 1. Heft i. p. 40. —— (18) Some Mesozoic plant-bearing deposits in Patagonia and Tierra del Fuego and their Floras. K. Svensk. Vetenskapsakad. Hand. Bd. ur. No. 3. —— (13?) The Mesozoic Flora of Graham Land. Wiss. Ergeb. Schwed. stidpolar. Exped. 1901-03, Bd. un. Lief. 14, p. 1. — See Moller, H. J. and T. G. Halle (13). —— (15) Some xerophytic Jeaf-structures in Mesozoic plants. Geol. Foren. Stockholm Férhand. Bd. xxxvu. H. v. p. 493. Hallier, H. (05) Provisional scheme of the Natural (Phylogenetic) system of Flowering Plants. New Phyt. vol. tv. p. 151. Harker, A. (06) The Geological structure of the Sgurr of Figg. Quart. Journ. Geol. Soc. vol. Lx. p. 40. —— (08) The Geology of the small Isles of Inverness-shire. Mem. yeol. Surv. Scotland. Harpe, P. de la. (62) See Bristow, H. W. REFERRED TO IN THE TEXT 491 Harshberger, J. W. (98) Water-storage and conduction in Senecio praecox DC. from Mexico. Contrib. Bot. Labt. (Univ. Pennsyl- vania), vol. 1. No. 1. —— (11) Phytogeographic Survey of N. America. (Die Veget. der Erde; Engler and Drude, x1. Leipzig.) . Hartz, N. (96) Planteforsteninger fra Cap Stewart i @stgrgnland. Meddel. om Grdnland, x1x. Copenhagen. Hayata, A. (06) On Taiwania, a new genus of Coniferae from the Island of Formosa. Journ. Linn. Soc. vol. xxxvit. p. 330. —— (07) On Taiwania and its affinity to other genera. Bot. Mag. (Tokyo), vol. xxi. p. 21. : — (10) Botanicakg§urvey by the Govt. of Formosa. Congr. Int. Bot. (Bruxelles), p. 59. —— (17) Some Conifers from Tonkin and Yunnan. Bot. Mageine, vol. xxxr. p. 113. Heer, 0. (62) On the Fossil Flora of Bovey Tracey. Phil. Trans. R. Soc. vol. ci. p. 1039. —— (68) i. Die in Nordgronland, auf der Melville- Insel, im Banks- land, an Mackenzie, im Island und in Spitzbergen entdeckten fossilen Pflanzen. Flor. Foss. Arct. vol. 1. Ziirich, — (69) Beitrage zur Kreide-Flora. J. Flora von Moletein in Mahrén. Neue Denksch. Allgem. Schweiz. Ges. gesammt. Naturwiss. Bd. xxit. — (71) iii. Die Miocene Flora und Fauna Spitzbergens. Flor. eu Arct. vol. II. —— (71*) Beitrage zur Kreide-Flora. II. Kreide tiers van (ueitiniore, Neue Denksch. Aligem. Schweiz. Ges. ge t. Naturwiss. Bd. xxiv. —— (75) ii. Die Kreide-Flora der anchisehity Zone. Flor. Foss. Arct. vol. m1. —— (75) iii. Nachtrige zur Miocenen Flora Grénlands. Ibid. —— (76) Flora Fossilis Helvetiae. Ziirich. —— (76?) Uber Permische Pflanzen von Fiinkkirchen in Ungarn. Mitt. Jahrb. K. Ung. Geol. Anst. Bd. v. —— (77) i. Beitrage zur fossilen Flora Spitzbergens. Flor. Foss. Aret. vol. Iv. — (77) ii. Beitrage zur Jura-Flora Ost Sibiriens und des Amur- landes. Ibid. —— (78) i. Die Miocene Flora des Grinnell-Lands. Ibid. vol. v. —— (78) ii. Beitrige zur fossilen Flora Sibiriens und des Amurlandes. Ibid. — (78) v. Beitrage zur Miocenen Flora von Sachalin. Ibid. — (81) Contributions 4 la Flore du Portugal. Sect. Trav. Geol. Port. (Lisbon). — (81?) Zur Geschichte der Ginkgo-artigen Baume. Engler’s Bot. Jahrb, Bd. 1. p. 1. —— (82) i. Flora fossilis Grénlandica. Flor. Foss. Arct. vol. v1. , — (83) Flora fossilis Groénlandica. Ibid. vol. vit. 492 LIST OF WORKS Helmhacher, R. (71) Sitzber. d. K. Bohm. Ges. Wiss. p. 81. Henry, A. (06) See Elwes and Henry. Herzfeld, S. (10) Die Entwicklungsgeschichte der weiblichen Bliite von Cryptomeria japonica Don. Ein Beitrag zur Deutung der Frucht- schuppe der Coniferen. Sitzber. Akad. Wiss. Wien, Bd. cxrx. Abt. i. p. 807. Hick, T. (95) On Kalorylon Hookeri Will. and Lyginodendron oldhamium Will. Mem. Proc. Manchester Lit. Phil. Soc. [4], vol. 1x. p. 109. Hilderbrand, F. (61) Die Verbreitung der Coniferen. Rhein. und Westphal. Verhand. Bd. xvi. p. 199. Hill, T. G. and E. de Fraine. (10) On the seedling structure of Gymno- sperms. IV. Ann. Bot. vol. xxiv. p. 319. Hirase, 8. (98) Etudes sur la fécundation et l’embryogenie du Ginkgo biloba. Journ. Coll. Sci. Imp. Univ. Tokyo, vol. xu. p. 1038. Héhlke, F. (02) Ueber die Harzbehalter und die Harzbildung bei den Polypodiaceen und einigen Phanerogamen. Beiheft Bot. Cent. Bd. x1. p. 8. Holden, H.S. (10) Note on a wounded Myeloxylon. New Phyt. vol. 1x. p. 253. Holden, Ruth. (13) Some fossil plants from Eastern Canada. Ann. Bot. Fi vol. xxvir. p. 243. (13?) Contributions to the anatomy of Mesozoic Conifers. No. 1. oe Coniferous wood from Yorkshire. Ann. Bot. vol. xxvii. p. 533. —— (138%) Cretaceous Pityoxyla from Cliffwood, New Jersey. Proc. Amer. Acad. Arts and Sci. vol. xvi. p. 609. ae (14) Contributions to the anatomy of Mesozoic Conifers. II. Cretaceous Lignites from Cliffwood, N. Jersey. Bot. Gaz. vol. Lv1. p. 168. —— (14?) On the relation between Cycadites and Pseudocycas. New Phyt. vol. xu. p. 334. —— (15) A Jurassic wood from Scotland. Ibid. vol. xiv. p. 205. — (15%) On the cuticles of some Indian Conifers. Bot. Gaz. vol. Lx. p. 215. Hollick, A. (97) The Cretaceous clay marl exposed at Cliffwood, N. J. Trans. N. Y. Acad. Sct. vol. xvi. p. 124. —— (04) Additions to the Palaeontology of the Cretaceous formation on Long Island. No. II. Bull. N. Y. Bot. Gard. vol. ut. No. 11, p. 403. —— (06) The Cretaceous Flora of southern N. Y. and New England. U. 8. Geol. Surv. Mon. vol. L. —— (06°) Systematic Palaeontology of the Pleistocene deposits of Maryland. Contributions from the New York Bot. Gard. No. 85. —— (12) Additions to the Palaeobotany of the Cretaceous formation on Long Island. Bull. N. Y. Bot. Gard. vol. vitt. No. 28, p. 154. Hollick, A. and E. C. Jeffrey. (06) Affinities of certain Cretaceous plant- remains commonly referred to the genera Dammara and Brachy- phyllum. Amer. Nat. vol. xu. p. 189. REFERRED TO IN THE TEXT 493 Holmes, W. H. (78) Fossil Forests of the Volcanic Tertiary formations of the Yellowstone National Park. Ann. Rep. Geol. Geogr. Surv. U.S.A. pt. m1. p. 47. Hooker, J.D. (52) Dacrydium lazifolium. Icones Plant. vol. v. pl. 815. —— (60) Flora Tasmanica. London. — (62) On the Cedars of Lebanon, Taurus, Algeria, and India. Nat. Hist. Rev. p. 11. —— (63) On Welwitschia, a new genus of Gnetaceae. Trans. Linn. Soc. vol. xxiv. p. l. Hooker, J. D. and E. W. Binney. (55). On the structure of certain limestone nodules enclosed in seams of bituminous coal, with a description of some Trigonocarpons contained inthem. Phil. Trans. R. Soc. vol. CXLIX. Horich, 0. (06) Potonié’s Abbildungen und Beschreibungen fossilen Pflanzen-Reste. Lief. iv. 69, 70. Howse, R. (88) A catalogue of Fossil Plants from the Hutton collection. Nat. Hist. Trans. Northumberland, Durham, and Newcastle-upon- Tyne, vol. x. Hutchinson, A. H. (14) The male gametophyte of Abies. Bot. Gaz. vol. tv. p. 148. — (15) On the male gametophyte of Picea canadensis. Ibid. vol. Lx. p. 287. ; Jeffrey, E. C. (03) The comparative anatomy and phylogeny of the Conifers. I. The genus Sequoia. Mem. Boston Soc. Nat. Hist. . vol. v. No. 10, p. 441. — (04) A fossil Sequoia from the Sierra Nevada. Bot. Gaz. vol. XXXVI. p. 321. — (05) The comparative anatomy and phylogeny of the Conifers. The Abietineae. Mem. Boston Soc. Nat. Hist. vol. v1. No. 1. — (06) The wound Reactions of Brachyphyllum. Ann. Bot. vol. xx. p. 383. — (07) Araucariopitys, a new genus of Araucarians. Bot. Gaz. vol. xiv. p. 435. ; — (08) Traumatic ray-tracheids in Cunninghamia sinensis. Ann. Bot. vol. xx. p. 593. —— (08?) On the structure of the leaf in Cretaceous Pines. Jbid. vol. xxu. p. 207. —— (10) Anew Prepinus from Martha’s Vineyard. Proc. Boston Soc. nat. Hist. vol. xxxiv. No. 10, p. 333. — (10?) Anew Araucarian genus from the Triassic. hid. vol. xxxtv. No. 9, p. 325. —— (10%) On the affinities of Yezonia. * Ann. Bot. vol. xxtv. p. 769. — (1l) The affinities of Geinitzia gracilima. Bot. Gaz. vol. t. p. 21. —— (12) The History, Comparative Anatomy and Evolution of the Araucarioxylon type. Proc. Amer. Acad. Arts Sci. vol. xuvut. No. 13, p. 532. 494 ; LIST OF WORKS Jeffrey, E. 0. (14) Spore-conditions in hybrids and the mutation hypo- thesis of de Vries. Bot. Gaz. vol. Lvm. p. 322. —— (17) The Anatomy of Woody Plants. Chicago 1917. Jeffrey, E. C. and M. A. Chrysler. (06) On Cretaceous Pityoxyla. Bot. Gaz. vol. xuu. p. 1. —— (062) The Lignites of Brandon. Contrib. from the Phanerogamic Labt. of Harvard Univ. No. vi. —— (07) The microgametophyte of the Podocarpineae. Amer. Nat. vol. xur. No. 486, p. 355. Jeffrey, E. C. and Ruth D. Cole. (16) Experimental Investigations on the genus Drimys. Ann. Bot. vol. xxx. p. 359. Jeffrey, E. C. and R. E. Torrey. (16) Ginkgo and the microsporangial mechanisms of the seed plants. Bot. Gaz. vol. Lx. p. 281. Johnson, T. (11) A seed-bearing Irish Pteridosperm, Crossotheca Héninghausi Kidst. Sci. Proc. R. Dublin Soc. vol. xm. p. 1. —— (12) Heterangium hibernicum sp. nov. a seed-bearing Heterangium from Co. Cork. Ibid. vol. x11. No, 20. —— (14) Ginkgophyllum kiltorkense sp. nov. Ibid. vol. xiv. p. 169. —— (17) Spermolithus devonicus, Gen. et sp. nov., and other Pterido- sperms from the Upper Devonian beds at Kiltoscan, Co. Kilkenny. Ibid. vol. xv. p. 245. Johnston, R. H. (86) Fresh contributions to our knowledge of the Plants of Mesozoic age in Tasmania. Papers and Proc. R. Soc. Tasmania for 1886, p. 160. Johnstrup, M. F. (83) Recherches sur les fossiles appartenant aux formations Crétacée et Miocéne, sur la céte occidentale du Grgnland. Medd. om Grgnland, vol. v. ; Jones, W.S. (12) The structure of the Timbers of some common genera of Coniferous trees. Quart. Journ. Forestry, April. —— (13) The minute structure of the wood of Cupressus macrocarpa. Ibid. — (13?) Ray-tracheids in Sequoia sempervirens and their pathological character. Lampeter. Karsten, G. (92) Beitrag zur Entwickelungsgeschichte einiger Gnetum Arten. Bot, Zeit. p. 205. — (93) Untersuchungen iiber die Gattung Gnetum. I. Ann. Jard. Bot. Buitenzorg, tome x1. p. 195. — (93%) Zur Entwickelungsgeschichte der Gattung Gnetum. Cohn’s Beit. Biol. Pflanz. v1. p. 337. Kershaw, E.M. (09) The structure and development of the ovule of Myrica Gale. Ann. Bot. vol. xxim. p. 353. — (12) Structure and development of the ovule of Bowenia spec- tabilis. Ibid. vol. xxvi. p. 625. Kidston, R. (84) On a new species of Schiitzia from the Calciferous sandstone of Scotland. Ann. Mag. Nat. Hist. vol. xm. p. 77. — (86) Notes on some fossil plants collected by Mr R. Dunlop, Airdrie, from the Lanarkshire coal-field. Trans. Geol. Soc. Glasgow, vol. vim. p. 47. REFERRED TO IN THE TEXT 495 Kidston, R. (90) The Yorkshire Carboniferous Flora. Trans. Yorks. Nat. Union, pt. Xv. —— (92) Notes on some fossil plants from the Lancashire Coal Measures. Trans. Manchester Geol. Soc. pt. xiii. vol. XxI. — (04) On the Fructification of Neuropteris heterophylla Brongn. Phil. Trans. R. Soc. vol. oxevu. p. 1. —— (04?) On the Fructification of Neuropteris heterophylla Brongn. Proc. R. Soc. vol. pxxu. p. 487. — (04°) Some Fossil Plants collected by Mr A. Sinclair from the Ayrshire coalfield. Kilmarnock Glenfield Ramblers Soc. Annals (1901-04), No. iv. Adlmarnock. —— (05) Preliminary Note on the occurrence of Microsporangia in organic connection with the Foliage of Lyginodendron. Proc. R. - Soc. vol. LXXvI. p. 358. ‘ — (11) Les Végétaux houillers recueillis dans le Hainaut Belge. Mém. Mus. Roy. @hist. nat. Belg. tome tv. —— (14) On the Fossil Flora of the Staffordshire coalfields. Pt. iii. The Fossil Flora of the Westphalian series of the S. Staffs. coalfield. Trans. R. Soc. Edinb. vol. u. pt. i. p. 73. Kidston, R. and D. T. Gwynne-Vaughan. (12) On the Carboniferous Flora of Berwickshire. Pt. i. Stenomyelon tuedianum Kidst. Trans. R. Soc. Edinb. vol. xvii. pt. ii. p. 263. | Kidston, R. and W. J. Jongmans. (11) Sur la Fructification de Neuro- pteris obliqua Brongn. Arch. Neerl. sci. exact. nat. [11. B], tomer. p. 25. Kirby, J. W. (64) On some remains of Fishes and Plants from the ‘Upper limestone’ of the Permian series of Durham. Quart. Journ. Geol. Soc. vol. xx. p. 349. Kirchner, 0., E. T. Loew, and C. Schréter. (06) Die Coniferen und Gnetaceen Mitteleuropas. Stutigart. Kirk, T. (89) The Forest Flora of New Zealand. Wellington. Kirsch, 8. (11) The Origin and Development of resin-canals in the Coniferae, with special reference to the Development of Thyloses and their correlation with the Thylosed strands of the Pteridophytes. Trans. R. Soc. Canada, sect. iv. p. 43. Kisch, Mabel H. (13) The Physiological Anatomy of the periderm of fossil Lycopodiales. Ann. Bot. vol. xxvir. p. 281. Kleeberg, A. (85) Die Markstrahlen der Coniferen. (Inaug. Diss.) Bot. Zeit. Bd. xii. Klein, L. (81) Bau und Verzweigung einiger dorsiventral gebaute Polypodiaceen. Nov. Act. K. Leop. Car. Deutsch. Akad. Natur- forsch. Bd. xia. No. 7, p. 335. Knowlton, F. H. (89) Description of the fossil woods and lignites from Arkansas. Ann. Rep. Geol. Surv. Arkansas, vol. 11. p. 249. —— (89?) Fossil wood and Lignite of the Potomac formation. Bull. U. 8. Geol. Surv. No. 56. — (90) A Revision of the genus Araucarioxylon of Kraus, with the compiled descriptions and partial synonymy of fhe species. Proc. U. S. Mus. vol. xu. p. 601. 496 LIST OF WORKS Knowlton, F. H. (93) The Laramie and the overlying Livingstone formation in Montana. Report on the Flora. Bull. U. S. Geol. Surv. No. 105. —— (99) Fossil Flora of the Yellowstone National Park. Monographs. xxxu. U.S. Geol. Surv. pt. ii. chap. XIv. —— (00) Floraof the Montana formation. Bull. U.S. Geol. Surv. No. 163. — (05) The geology of the Perry basin in South-eastern Maine, with a chapter on the fossil plants. U. S. Geol. Surv. Prof. Papers, No. 35. é —— (ll) The correct technical name for the Dragon tree of the Kentish Rag. Geol. Mag. [v], vol. vir. p. 467. — (14) The Jurassic Flora of Cape Lisburne, Alaska. U.S. Geol. Surv. Prof. Papers, No. 85-D, p. 39. Kny, L. (10) Uber die Verteilung des Holzparenchyma bei Abies pectinata DC. Ann. Jard. Bot. Buit. [2], Suppl. ur: p. 645. Koettlitz, R. (98) Observations on the geology of Franz Josef Land. Quart. Journ. Geol. Soc. vol. tiv. p. 620. Kosmovsky, C. (92) Quelques mots sur les couches 4 végétaux fossiles dans la Russie orientale et en Sibérie. Bull. soc. Imp. Nat. Moscou [N.S.], tome v. p. 170. Kramer, A. (85) Beitrage zur Kenntniss der Entwickelungsgeschichte und des anatomischen Baues der Fruchtblatter der Cupressineen und der Placenten der Abietineen. Flora, xu. p. 519. Krasser, F. (91) Uber die fossile Flora der rhatischen Schichten Persiens. Sitzber. K. Akad. Wiss. Wien, Bd. c. Abth. i. p. 413. — (97): Constantine Freiherr von Ettingshausen. Oéesterr. bot. Zeitsch. Nos. 9 and 10. — (03) Konstantin von Ettingshausen’s Studien itiber die fossile Flora von Ouricanga in Brasilien. Sitzber. K. Akad. Wiss. Wien, Bd. cxm. Abt. i. p. 852. — (05) Fossile Pflanzen aus Transbaikalien der Mongolei und Mandschurei. Denksch. K. Akad. Wiss. Wien, Bd. Lxxviml. p. 589. —— (09) Zur Kenntniss der fossilen Flora der Lunzer Schichten. Jahrb. K.k. geol. Reichs. Wien, Bd. tix, Heft i. p. 101. —— (12) Williamsonia in Sardinien. Sitzber. K. Akad. Wien, Bd. oxxr. Abt. i. p. 944. —— (13) Die fossile Flora der Williamsonien bergenden Juraschichten von Sardinien. Jbid. Bd. cxxu. Kraus, G. (64) Mikroskopische Untersuchungen iiber der Bau lebenden Nadelhélzer. Wiirzb. Naturwiss. Zeitsch. Bd. v. p. 144. —— (66) Uber den Bau der Cycadeenfiedern. Prings. Jahrb. Bd. 1v. sp. 805. —— (88) Beitrige zur Kenntniss fossiler Holzer. I. Holzer aus den Schwefelgruben Siciliens. Abdh. Naturf. Ges. Halle, Bd. xvi. p. 79. —— (92) Beitrage zur Kenntniss fossiler Holzer. Abh. Naturforsch. Ges. Halle, Bd. xvu. p. 67. — (96) Physiologisches aus den Tropen. Ann. Jard. Bot. Buit. vol. x1. p. 217. REFERRED TO IN THE TEXT 497 Kriusel, R. (13) Beitrage zur Kenntniss der Holzer aus der Schlesischen Braunkohle. Teil1. (Inaug. Diss. Breslau.) Bot. Cent. Bd. cxxut. p. 123. Krystofovié, A. (10) Jurassic Plants from Ussuriland. Mem. Com. Geol. [N.8.], Livr. 56. —— (15) Plant remains from Jurassic lake-deposits of Transbaikalia. Mém. Soc. Imp. Russe Mineralog. [2], 1. Kubart, B. (08) Pflanzenversteinerungen enthaltende Knollen aus dem Ostrau-Karwiner Kohlenbecken. Sitzber. K. Akad. Wiss. Wien, Bd. cxvu. Abt. i. p. 573. — (11) Corda’s sphaerosiderite aus dem Steinkohlenbecken Radnitz- Béaz in Béhmen nebst Bemerkungen iitber Chorionopteris gleicheni- oides Corda. Ibid. Bd. cxx. Abt. i. p. 1035. —— (11°) Podocarpoxylon Schwendae, ein fossiles Holz von Altersee (Oberésterreich). Osterr. bot. Zeitsch. No. 5, p. 161. —— (14) Uber die Cycadofilicineen Heterangium und Lyginodendron aus dem Ostrauer Kohlenbecken. Osterr. bot. Zeitsch. No. 1. ii. p. 8. Kurtz, F. (03) Remarks upon Mr E. A. Arber’s communication on the Clarke collection of Fossil Plants from New South Wales. Quart. Journ. Geol. Soc. vol. LIX. p. 25. Kutorga,S. (42) Beitrag zur Palaeontologie Russlands. Verhand. Russ.- Kais. Mineral. Ges. St Petersburg. —— (44) Zweiter Beitrag zur Palaeontologie Russlands. Ibid. p. 62. Land, W.J.G. (04) Spermatogenesis and Oogenesis in Ephedra trifurca. Bot. Gaz. vol. Xxxvut. p. 1. Lang, W. H. (97) Studies in the Development and Morphology of Cycadean sporangia. I. The microsporangia of Stangeria para- doxa. Ann. Bot. vol. xt. p. 421.- —— (00) Studies in the Development and Morphology of Cycadean sporangia. II. The ovule of Stangeria paradoxa. Ibid. vol. xiv. p. 281. Lange, T. (90) Beitraige zur Kenntniss der Flora des Aachener Sandes. Zeitsch. Deutsch. geol. Ges. Bd. XLIt. p. 658. Laurent, L. (12) Flore fossile des Schistes de Manat (Puy-de-Déme). Ann. Mus. @hist. nat. Marseille (Geol.), tome XIv. p. 3. Lawson, A. A. (04) The gametophytes, fertilization, and embryo of Cryptomeria japonica. Ann. Bot. vol. xvi. p. 417. —— (09) The gametophytes and embryo of Pseudotsuga Douglasii. Ibid. vol. xxi. p. 163. —— (10) The gametophytes and embryo of Sciadopitys verticillata. Ibid. vol. xxiv. p. 403. Lebour, G. A. (77) Itustrations of Fossil Plants; being an autotype reproduction of selected drawings. London. Lesquereux, L. (74) Contributions to the Fossil Flora of the Western Territories, Pt. 1. The Cretaceous Flora. Rep. U. 8. Geol. Surv. Territ. vol. VI. s. IV 32 498 LIST OF WORKS Lesquereux, L. (78) Onthe Cordaites and their related generic divisions, in the Carboniferous formation of the United States. Proc. Amer. Phil. Soc. (Philadelphia), vol. xviz. p. 315. — (83) Contributions to the Fossil Flora of the Western Territories. Pt. iii. The Cretaceous and Tertiary Floras. U. S. Geol. Surv. Territ. vol. M1. -—— (91) The Flora of the Dakota group. Monographs U. S. Geol. Surv. vol. Xvu. Leuthardt, F. (03) Die Keuper Flora von Neuewelt bei Basel. Abh. Schweiz. palaeont. Ges. Bd. xxx. p. 1. Lignier, O. (92) La nervation taenioptéridée de folioles de C'ycas et le tissu de transfusion. Bull. soc. Linn. Normandie [4], vol. v1. fasc. 1. -—— (94) La nervation des Cycadées est dichotomique. Assoc. Franc. pour Vavancement de sci. (Caen). — (94?) Végétaux Fossiles de Normandie. Structure et affinités du Bennettites Morierei S. and M. sp. Mém. soc. Linn. Normand. tome Xvul. p. l. —— (95) Végétaux fossiles de Normandie. II. Contributions 4 la flore liassique de Ste Honorine-la-Guillaume (Orne). bid. vol. xvi. —— (01) Ibid. TI. Etude anatomique du Cycadeoidea micromyela Mor. Ibid. vol. xx. p. 331. -—— (08) Le fruit du Williamsonia gigas Carr. Mém. soc. Linn. Normand. vol. xxt. p. 19. —— (03%) La Fleur des Gnétacées est-celle intermédiaire entre celle des Gymnospermes et celle des Angiospermes? Bull. soc. Iinn. Normand. [5], vol. va. p. 55. — (04) Notes complémentaires sur la structure du Bennettites aforierei 8S. and M. Bull. soc. Linn. Normand. [5], vol. vai. p. 3. —— (06) Radiculttes reticulatus, radicelle fossile de Séquoinée. Bull. soc. bot. France, tome V1. [iii.], p. 193. —— (07) Sur un moule litigieux de Williamsonia gigas (Ta. and H.) Jf Carr. Ibid. [6], vol. 1. (07?) Végétaux fossiles de Normandie. IV. Bois divers (sér. 1). Mém. soc. Linn. Normand. vol]. xx1. p. 239. —— (09) Le Bennettites Morieret (S. and M.) Lignier ne serait-il pas C@origine infracrétacée? Bull. soc. Linn. Normand. [6], vol. 1. p. 214. — (11) Le Bennettites Morierei (S. and M.) Lign. se reproduisait probablement par parthénogénése. Bull. soc. bot. France [4], tome XI. p. 224. . ; —— (11°) Les “Radiculites reticulatus” Lign. soit probablement des radicelles de Cordaitales. Assoc. Franc. Avanc. Sct. xu. (Dijon), p. 509. [See also Lignier (06).] — (12) Stomates des écailles interséminales chez le Bennettites Jorierei (S. and M.). Jbid. tome xm. p. 425. -—— (13) Végétaux fossiles de Normandie. VII. Contributions a la Flore Jurassique. d/ém. soc. Linn. Normand. vol. xxtv. p. 69. REFERRED TO IN THE TEXT 499 Lignier, O. (13%) Différenciation des tissus dans le Bourgeon végétatif du Cordaites lingulatus B. Ren. Ann. sci. nat. [7], vol. xvIt. p. 233. Lignier, 0. et A. Tison. (11) Les Gnétales sont des Angiospermes apétales. Compt. Rend. Jan. 23. --— (12) Les Gnétales, leurs fleurs et leur position systématique. Ann. scr. nat. [N.S.]. —— (13) L’ovule tritégumenté des Gnetum est probablement un axe @inflorescence. Bull. soc. bot. France [4], tome xm. p. 64. —— (137) Un nouveau Sporange Séminiforme, Mittagia semini- formis, gen. et sp. nov. Mém. Soc. Linn. Normandie, tome xxiv. p. 49. Lima, W. de. (88) Flora Fossil de Portugal. Monographia do Genero Dicranophyllum. Comm. dos Trab. geol. Portugal. Lindley, J. See Murchison and Lyell (29). — _ Lingelsheim, A. (08) Uher die Braunkohlenhdlzer von Saarau. Jahres. Ber. Schles. Ges. Vaterland. Cultur. Bd. Lxxxv. Lloyd, F. E. (02) Vivipary in Podocarpus. Torreya, 1. p. 113. Lomax, J. (02) Onsome features in relation to Lyginodendron oldhamium. Ann. Bot. vol. xvi. p. 601. Lotsy, J. (99) Contributions to the life-history of the genus Gnetum. Ann. Jard. Bot. Buit. [2], vol. 1 p. 46. —— (09) Vortrage itber botanischeStammesgeschichte. Bd.m. Jena. —— (11) Ibid. Bd. m. Ludwig, R. (61) Fossile Pflanzen aus der Altesten Abtheilung der Rheinisch-Wetterauer Tertiér-Formation. Palaeontograph. Bd. vu. p. 39. — (69) Fossile Pflanzenreste aus den paldolithischen Formation der Umgegend von Dillenburg, Bidenhopf und Friedberg und aus den Saalfeldischen. Jbid. Bd. xvu. p. 105. Lyon, H. L. (04) The Embryogény of Ginkgo. Minnesota Bot. Stud. XXII. p. 275. McBride, T. H. (93) A new Cycad. Amer. Geologist, vol. x11. p. 248. Mackie, S. J. (62) The Dragon Tree of the Kentish Rag. Geologist, vol. v. p. 401. McLean, R.C. (12) Two Fossil prothallifrom the Lower Coal Measures. New Phyt. vol. x1. p. 305. McNab, W. R. (70) On the structure of a Lignite from the Old Red Sandstone. Trans. Bot. Soc. Edinburgh, vol. x. p. 312. Mahlert, A. (85) Beitrage zur Kenntniss der Anatomie der Laubblatter der Coniferen mit besonderer Berichsichtigung des Spaltéffnungs- Apparates. Bot. Cent. Bd. xxiv. p. 54. Malloizel, G. and R. Zeiller. (N.D.) Bibliographie et tables icono- graphiques (O. Heer). Stockholm. Mansell-Pleydell, J. C. (85) Notes on a cone from the Inferior Oolite Beds of Sherborne. Proc. Dorset Nat. Hist. Antig. Field Club, . vol. v. p. 141. 32—2 500 LIST OF WORKS Mantell, G. (27) Illustrations of the Geology of Sussex. London. ; —— (43) Description of some fossil plants from the chalk formation of the south-east of England. Proc. Geol. Soc. vol. Iv. Pp. 34. — (46) Description of some Fossil Fruits from the Chalk formation of the south-east of England. Quart. Journ. Geol. Soc. vol. m. p. 51. Marion, A. F. (84) Sur les caractéres d’une Conifére tertiaire, voisine des Dammareées (Doliostrobus Sternbergii). Compt. Rend. vol. xctv. p.821. Marsh, A.S. (14) Notes on the Anatomy of Stangeria paradoxa. New Phyt. vol. xu. p. 18. Marty, P. (08) Sur la Flore fossile de Lugarde (Cantal). Compt. Rend. vol. oxLvm. p. 395. Maslen, A.J. (10) See Scott and Maslen. — (ll) The structure of Mesoxylon Sutcliffi (Scott). Ann. Bot. vol. xxv. p. 38]. Massalongo, A. (59) Studii sulla Flora Fossile e geologia stratigrafica del Senigalliese. Verona. Massalongo, A. and G. Scarabelli. (58) Studii sulla Flora Fossile e geologia stratigrafica del Senigalliese. Verona. Masters, M.T. (91) Review of some points in the comparative mor- phology, anatomy, and life-history of the Coniferae. Journ. Linn. Soc, vol. XXvui. p. 236. —— (93) Notes on the genera of Taxaceae and Coniferae. Ibid. vol. xxx. p. 1. —— (00) Tazxodium and Glyptstrobus. Journ. Bot. (February, 1900). Matte, H. (04) Recherches sur l’appareil libéro-ligneux des Cycadées. Caen. —— (08) Sur le développement morphologique et anatomique des Cycadacées. Mém. soc. Linn. Normand. tome XXIII. Matthew, G. F. (10) Revision of the Flora of the Little River group. Trans. R. Soc. Canada, sect. iv. vol. 11. [3], p. 77. Mercklin, C. E. von. (55) Palaeodendrologicon Rossicum. St Petersburg. Miller, H. (58) The Cruise of the Betsey. Edinburgh. Miquel, F. A. W. (42) Monographia Cycadearum. — (47) Collectanea nova ad Cycadearum cognitionem. Linnaea, Bd. xix. p. 411. —— (51) Over de Rangschikking der fossiele Cycadeae. Tijdsch. Wiss. Nat. Wet. vol. Iv. p. 205. — (69) On the sexual organs of the Cycadaceae. Journ. Bot. vol. vit. p. 64. Mirande, M. (05) Recherches sur le développement et l’anatomie de Cassythacées. Ann. Sci. nat. [ix], vol. 1 p. 181. Miyake, K. (06) Uber die Spermatozoiden von Cycas revoluta. Ber. Deutsch. bot. Ges. Bd. xxiv. p. 78. —— (10) The Development of the gametophytes and embryogeny in Cunninghamia sinensis. Betheft Bot. Cent. Bd. xxxvu. Abt. i. Heft 1. Mogan, L. (03) Untersuchungen iiber eine fossile Konifere. Sitzber. K. Akad. Wiss. Wien, Bd. cxm. Abt. i. p. 829. REFERRED TO IN THE TEXT 501 Mohl, H. von. (62) Hinige anatomische und physiologische Bemerkungen iiber das Holz der Baumwurzeln. Bot. Zeit. p. 225. Moller, H. (03) Bidrag till Bornholms Fossila Flora (Rhat och Lias). Gymnospermer. K. Svensk. Vetenskapsakad. Hand. Bd. xxxvt. No. vi. Miller, H. J. and T.G. Halle. (13) The Fossil Flora of the Coal-bearing deposits of south-eastern Scania. Arkiv Bot. (Stockholm), Bd. x11. No. 7. - ; Moriére, J. (69) Note sur deux végétaux trouvés dans le département du Calvados. Mém. soc. Linn. Normand. vol. xv. Morris, J. (40) Memoir to illustrate a Geological map of Cutch (Grant, C. W.). Trans. Geol. Soc. [2], vol. v. pt. ii. p. 289. —— (41) Remarks upon the Recent and Fossil Cycadeae. Ann. Mag. Nat. Hist. vol. vu. p. 110. - — (54) A Catalogue of British Fossils. London. Miller, C. (90) Ueher die Balken in den Holzelementen der Coniferen. Ber. Deutsch. bot. Ges. Bd. vat. p. 17. Murchison, Sir R. and R. Harkness. (64) On the Permian rocks of the North-west of England, and their extension into Scotland. Quart. Journ. Geol. Soc. vol. xx. p. 144. Murchison, Sir R. and C. Lyell. (29) On the Tertiary Freshwater formations of Aix, in Provence, including the coal-field of Fuveau. Description of plants by J. Lindley. Edinburgh New Phil. Journ. . 287. arsine. Y. (83) Ueber den anatomischen Bau des Holzes der wichtigsten Japanischen Coniferen. Unters. aus dem Forstbot. Instit. zu Miinchen. ut. Berlin. Nathorst, A.G. (75) Om en Cycadékotte fran den ratiska formationens lager vid TinkarpiSkane. Ofver. K. Vetenskapsakad. Férh. No. 10. — (78) Om Ginkgo crenota Brauns sp. frin sandstenen vid Senstedt naira Braunschweg. Ibid. No. 3. — (80) Nagra anmarkningar om Williamsonia, Carruthers. Ibid. No. 9. —— (81) Berittelse, afgifven till Kongl. Vetenskaps-Akad. Ofver. K. Vetenskapsakad. Férhand. No. 1. —— (86) Om Floran i Skanes kolférande Bildningar. I. Floran vid Bjuf. Sver. geol. Unters. ser. C, Nos. 27, 33, 85 (1878-86). —— (88) Nya anmarkningar om Williamsonia. Ofver. K. Vetenskaps- akad. Férh. No. 6. —— (89) Sur la présence du Genre Dictyozamites, Old. dans les Couches Jurassiquesde Bornholm. Overs. K. Dansk. Vidensk. Selsk. Férhandl. p. 96. i ‘ —— (93) Beitrige zur Geologie und Palaeontologie der Republik Mexico (Felix und Link), Th. u. Heft i. Leipzig. —— (97) Zur Mesozoischen Flora Spitzbergens. K. Svensk. Veten- skapsakad. Hand. Bd. xxx. No. 1, p. 5. —— (97%) Nachtragliche Bemerkungen iiber die Mesozoische Flora Spitzbergens. Ofvers. K. Vet.-Akad. Forhand. No. 8. 502 LIST OF WORKS Nathorst, A.G. (99) The Norwegian North Polar Expedition 1893-96. Scientific Results, edit. F. Nansen. III. Fossil Plants from Franz Josef Land. London and Christiania. —— (02) Beitrage zur Kenntniss einiger Mesozoischen Cycadophyten. K. Svensk. Vetenskapsakad. Hand. Bd. xxxvi. No. 4. —— (06) Om nagra Ginkgovaxter fran Kolgrufvorna vid Stabbarp i Skane. Lunds Univ. 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Féren. Stockholm Férh. Bd. xxxv. H. vi. p. 361. —— (13?) Die Pflanzenfiihrenden Horizonte innerhalb der Grenz- schichten des Jura und der Kreide Spitzbergens. Geol. Féren. Stockholm Férh, Bd. xxxv. H. iv. p. 273. REFERRED TO IN THE TEXT 503 Nathorst, A. G. (14) Zur Fossilen Flora der Polarlinder. Teil 1. Lief. iv. Nachtrage zur Palaozoischen Flora Spitzbergens. Stockholm. —— (15) Zur Devonflora des westlichen Norwegens. Bergens Mus. Aarbog, No. 7.° —— (15?) Tertiére Pflanzenreste aus Ellesmere-Land. Rep. second Norwegian Arct. Exped. in the “Fram” 1898-02, No. 35. Kristiania. Negri, G. (14), Sopra alcuni legni fossili del Gebel Tripolitano. Boll. Soc. geol. Ital. vol. Xxx. p. 321. Nestler, A. (95) Ein Beitrag zur Anatomie der Cycadeenfiedern. Prings- heim’s Jahrb. Bd. xxvu. p. 341. Newberry, J.S. (54) New Fossil Plants from Ohio. Annals of Science; including the Transactions of the American Association for the Advance- ment of Science, vol. 1. 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XXv. ae Oliver, F. W. and D. H. Scott. (03) On Lagenostoma Lomaxi, the seed of Lyginodendron. Proc. R. Soc. vol. LXxI. p. 477. — (04) On the structure of the Palaeozoic seed Lagenostoma Lomazxi, with a statement of the evidence upon which it is referred to Lygino- dendron. Phil. Trans. R. Soc. vol. cxcvu. p. 193. Osborne, T. G. B. (09) The lateral roots of Amyelon radicans Will. and their Mycorhiza. Ann. Bot. vol. xx. p. 603. Palibin, J. (01) Quelques données relatives aux débris végétaux contenus dans les sables blancs et les grés quartzeux de la Russie méridionale. Bull. Com. Geol. tome xx. p. 447. —— (04) Pflanzenreste vom Sichota-Alin Gebirge. Verhand. K. Russ. Mineral. Ges. Bd. xu1t. Lief. i. p. 31. Patrick, J.S. (44) On the Fossil vegetables of the Sandstone of Ayrshire. Ann. Mag. Nat. Hist. vol. xu. p. 283. Pavolini, A. F. (09) La Stangeria paradora Th. Moore. Nuov. Giorn. Bot. Italiano [N.S.], vol. xv1. p. 335. Firenze. Pax, F. (07) Beitrage zur fossilen Flora der Karpathen. 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Africa, vol. 111. p. 259. Reid, Clement and Fleanor M. Reid. (08) On the Preglacial Flora of Britain. Journ. Linn. Soc. vol. xxxvim. p. 206. —— (10) The Lignites of Bovey Tracey. Phil. Trans. R. Soc. vol. cct. . 161. — ne The Pliocene Floras of the Dutch-Prussian border. Meded.. Rijksopsporing Delfstoffen, No. 6. The Hague. Renault, B. (79) Sur un nouveau groupe de tiges fossiles silicifiées de Pépoque houilliére. Compt. Rend. tome LXXXvut. p. 35. —— (80) Sur une nouvelle espéce de Poroxylon. Compt. Rend. tome xcr. p. 860. —— (80%) Cours de Botanique fossile. Tome 1. Paris. —— (83) Cours de Botanique fossile. Tome m1. — (85) Cours de Botanique fossile. Tome rv. —— (87) Note sur le Clathropodium MMorieri. Bull. soc. Linn. Nor- mand. [4], vol. 1. p. 3. — (88) Les Plantes fossiles. Paris. —— (89) Sur un nouveau genre fossile de tige cycadéenne. Compt. Rend. vol. crx. p. 1173. —— (96) Note sur le genre Métacordaite. Soc. @hist. nat. d Autun. Renault, B. and R. Zeiller. 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(78) Sur le nouveau groupe Paléozoique des Dolero- phyllées. bid. vol. LXxxvu. p. 393. —— (82) Sur quelques types de végétaux récemment observés a l’état fossile. Ibid. vol. xLiv. p. 922. : —— (84) Paléontologie Francaise. Plantes Jurassiques, tome II. — (91) Ibid. tome tv. Types Proangiospermiques. —— (93) Revue des travaux de Paléontologie végétale. Rev. Gén. tome v. p. l. Saporta, G.de and A. F. Marion. (76) Recherches sur les végétaux fossiles de Meximieux. Arch. Mus. @hist. nat. de Lyon. Lyon. —— (78) Révision de la flore Heersienne de Gelinden. Mém. cour. et Meém. sav. érang. tome xu. (Acad. roy. sci. etc. Belg.). —— (85) L’Evolution du Régne végétal. Tomes. Les Phanérogames. Paris. . Saxton, W. T. (10) Contributions to the life-history of Widdringtonia cupressoides. Bot. Gaz. vol. u. p. 31. — (10%) Contributions to the life-history of Callitris. Ann. Boi. vol. xxiv. p. 557. —— (10%) Notes on the anatomy of Widdringtonia and Callitris. S. African Journ. Sci. p. 282. — (104) The development of the embryo of Encephalartos. Bot. Gaz. vol. XLIx. p. 13. — (12) Note onan abnormal prothallus of Pinus maritima L. Ann. Bot. vol. xxvi. p. 943. —— (13) Contributions to the life-history of Actinostrobus pyramidalis Miq. Ann. Bot. vol. xxvit. p. 321. —— (13?) The classification of Conifers. New Phyt. vol. x1. p. 242. — (J3*) Contributions to the life-history of Tetraclinis articulata, Masters, with some notes on the Phylogeny of the Cupressoideae and the Callitroideae. Ann. Bot. vol. xxvu. p. 577. Schauroth, C. von. (52) Herr von Schauroth an Herrn Beyrich. Zeit. . Deutsch. geol. Ges. Bd. Iv. p. 538. Schenk, A. (67) Ueber die Flora der schwarzen Schiefer von Raibl. Wiirzburg. Naturwiss. Zeitsch. vol. v1. p. 10. —— (68) Ueber die Pflanzenreste des Muschelkalkes von Recoaro. Benecke’s Geog.-Pal. Beit. Bd. 1. Miinchen. —— (68°) Beitrige zur Flora der Vorwelt. Palaeontol. Bd. xvi. p. 218. —— (71) Die Fossilen Pflanzen der Wernsdorfer Schichten in der Nordkarpathen. Jbid. Bd. xrx. p. 1. —— (80) Ueber fossile Hélzer aus der Libyschen Wiiste. Bot. Zeit. Bd. xxxvut. p. 657. —— (82) Ueber Medullosa elegans. Engler’s Jahrb. Bd. ut. p. 156. —— (82?) Die von dem Gebriidern Schagintweit in Indien gesammelten fossilen Holzer. Engler’s Bot. Jahrb. Bd. m1. p. 353. —— (83) Fossile Hélzer der libyschen Wiiste (Die Libysche Waste, Bd. u.). REFERRED TO IN THE TEXT 509 Schenk, A. (89) Ueber Medullosa Cotta. Abh. K. Stichs. Ges. Wiss. Bd. xv. p. 523, Schlechtendal, D. von. (02) Thuja occidentalis thuringiaca. Zeitsch. Naturwiss. Stuttgart. Bd. uxxv. p. 33. Schmalhausen, J. (83) Die Pflanzenreste der Steinkohlenformation am éstlichen Abhange des Ural Gebirges. Mém. Acad. 8. Pétersbourg, vol. xxx1. No. 13. —— (83?) Beitrage zur Tertiir Flora siid-west Russlands. Paleontol. Abh. (Dames and Kayser), Bd. 1. Heft iv. p. 285. —— (87) Die Pflanzenreste der Artinskischen und Permischen Ablagerungen im Osten des Europaischen Russlands. Mém. Com. géoi. St Pétersbourg, vol. 1. No. iv. p. 1. —— (90) Wissenschaftliche Resultate der von der Akad. der Wiss. zur Erforschung des Janalandes und der Neusibirischen Inseln in den Jahren 1885, 1886 ausgesandten Expedition. Abt.u. Tertidre Pflanzen der Insel Neusibirien. Mém. Pacad. Imp. Sci. St Péters- bourg [vii], vol. xxxvit. No. 5. Schneider, W. (13) Vergleichend-morphologische Untersuchungen iiber die Kurztriebe einiger Arten von Pinus. Flora[N.F.], Bd. v. p. 385. Schroeter, C. (80) Untersuchung iiber fossile Hélzer aus der arctischen Zone. (Heer’s Flor. Foss. Arct. vol. v1. 1882.) —— (97) Ueber die Vielgestaltigkeit der Fichte (Picea excelsa T.ink.). Vierteljahr. Naturforsch. Ges. Ziirich. Jahrg. xu. p. 125. Schuster, J. (11) Ueber Goeppert’s Rawmeria im Zwinger zu Dresden. Sitzber. K. Bayer. Akad. 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Bot. vol. xxvi. p. 1077. —— (12°) The structure of the stomata of certain Cretaceous Conifers. Bot. Gaz. vol. y1v. p. 63. —— (16) The Morphology and Affinities of Gnetum. Amer. Journ. Bot. vol. m1. p. 135. Thomson, R. B. (07) The Araucarineae, a Protosiphonogamic method of Fertilisation. Science [N.8.], vol. xxv. p. 272. —— .(09) Onthe pollen of Microcachrys tetragona. Bot. Gaz. vol. XLVI. p. 26. Gees (13) On the comparative anatomy and affinities of the Arau- carineae. Phil. Trans. R. Soc. vol. cctv. p. 1. 33—2 516 LIST OF WORKS Thomson, R. B. (14) The spur-shoot of the Pines. Bot. Gaz. vol. LVI. p. 362. = Thomson, R. B. and A. E. Allin. (12) Do the Ahietineae extend to the Carboniferous? Bot. Gaz. vol. Li. p. 339. ; Thomson, Mary R. H. (16) A note on the wood of Gnetum Gnemon. Ann. Bolus Herb. vol. 1. pt. ii. p. 81. Tison, A. (09) Sur le Saxegothaea conspicua Lind. Mém. soc. Linn. Normand. vol. xx1u. p. 139. — (12) Sur la persistance de la nervation dichotomique chez Jes Coniféres. Bull. soc. Linn. Normandie [vi], vol. 1v. p. 30. —— (12°) See Lignier and Tison. Tupper, W. W. (11) Notes on Ginkgo biloba. Bot. Gaz. vol. LI. p. 374. — Tuzson, J. (09) Monographie der fossilen Pflanzenreste der Balatonsee- gegend. Result. der Wiss. Erforsch. des Balatonsees, Bd. 1. Teil i. Budapest. — (09?) Beitrige zur fossilen Flora Ungarns. Mutt. Jahrb. K. Ung. Geol. Reichs. Ba. xxi. Heft viii. p. 233. ; —— (14) Beitrage zur fossilen Flora Ungarns. Mitt. Jahrb. K. Ungarisch. Geol. Reichs. Bd. xx1. Heft viii. Unger, F. (45) Synopsis Plantarum Fossilium. Leipzig. —— (47) Chloris Protogaea. Leipzig. —— (49) Einige interessante Pflanzenabdriicke aus der K. Petre- factensammlung in Miinchen. Bot. Zeit. p. 345. — (51) Die Fossile Flora von Sotzka. Denksch. K. Akad. Wiss. Wien, Bd. m. Abt. ii. p. 131. —— (52) Ueber einige fossile Pflanzen aus den lithographischen Schiefer von Solenhofen. Palaeontograph. Bd. wu. p. 251. — (52?) Iconographia plantarum fossilium. Abbildungen und Beschrei- bungen fossiler Pflanzen. Denksch. K. Akad. Wiss. Wien, Bd. rv. p. 73. —— (54) Zur Flora des Cypridinenschiefers. Sitzber. K. Akad. Wiss. Wien, Bd. xt. p. 595. — (54%) Jurassische Pflanzenreste. Palaeontograph. Bd. Iv. p. 39. — (59) Der versteinerte Wald bei Cairo. Sitzber. K. Akad. Wiss. Wien, Bd. xxx. p. 209. : —— (67) Die fossile Flora von Kumi. Denksch. K. Akad. Wiss. Wien, Bd. xxvii. p. 27. Veitch’s Manual of the Conifers. (00) London. Velenovsky, J. (81) Die Flora aus den Ausgebrannten Tertiadren Letten von Vrsovic bei Laun. Abh. béhm. Ges. [vi], Bd. 1. —— (87) Neue Beitrage zur Kenntniss der Pflanzen des bohmischen Cenomans. Sitzber. K. bohm. Ges. Wiss. Jahrg. 1886, p. 633. — (88) Ueber einige neue Pflanzenformen der béhmischen Kreide- formation. Jhid. Jahrg. 1887. —— (89) Kvétena Ceského Cenomanu. Abs. K. bohm. Ges. Wiss. (vii], Bd. or. —— (07) Vergleichende Morphologie der Pflanzen. Teil ii. Prag. REFERRED TO IN THE TEXT 517 Vernon, R. D. 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XXv. p. 93. — (08%) Two new Araucarias from the Western Cretaceous. Geol. Surv. South Dakota. —— (09) The Williamsonias of the Mixteca Alta. Bot. Gaz. vol. XLVI. p. 427. 3 — (11) On the Williamsonia Tribe. Amer. Journ. Sct. vol. XXXII. p. 433. ; —— (11°) A study of some American Fossil Cycads. Pt. v. Ibid. vol. xxx. p. 133. — (12) Ibid. pt. vi. On the smaller flower-buds of Cycadeoidea. Ibid. vol. xxx. p. 73. —— (13) The Liassic Flora of the Mixteca Alta of Mexico, its com- position, age, and source. bid. vol. xxxvi. p. 251. —— (14) A study of some American Fossil Cycads. Pt. vii. Furthe notes on disk structure. Jbid. vol. xxxvum. p. 117. —— (16) American Fossil Cycads. Vol. 1. Taxonomy. Washington. Wiesner, J. (03) Die Rohstoffe des Pflanzenreiches. Bd. u. Leipzig. Wild, G. (00) On new and interesting features in Trigonocarpus olivae- formis. Trans. Manchester Geol. Soc. vol. Xxv1. pt. xv. p. 484. Williamson, W. C. 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Rep. 45th meeting (Bristol) of the Brit. Assoc. p. 159. —— (86) On the morphology of Pinites oblongus. Mem. Proc. Man- chester Lit. Phil. Soc. vol. x. [3], p. 189. —— (87) On the Organisation of the Fossil Plants of the Coal Measures, XI. Phil. Trans. R. Soc. vol. 178. ' —— (90) Ibid. XVII. Ibid. vol. cLXxx1. p. 89. Wiliamson, W. ©. and D. H. Scott. (94) The root of Lyginodendron oldhamium Will. Proc. R. Soc, vol. vi. p. 128. —— (95) Further observations on the Organisation of the Fossil Plants of the Coal Measures. III. Lyginodendron and Heterangium. Phil. Trans. R. Soc. vol. CLXXXVI. p. 703. Wills, Lucy. (14) Plant cuticles from the Coal Measures. Geol. Alag. [6], vol. 1. p. 385. : Wills, L. T. (10) The fossiliferous Lower Keuper rocks of Worcester- shire. Proc. Geol. Assoc. vol. xx. p. 249. Winkler, ©. (72) Zur Anatomie von Araucaria brasiliensis. Bot. Zeit. Jahrg. xxx. p. 581. Witham, H. (30) On the vegétable fossils from Lennel Braes near Coldstream, upon the banks of the Tweed in Berwickshire. Phil. Mag. vol. vi. p. 16. — (31) Observations on fossil-vegetables accompanied by representa- tions of their internal structure as seen through the microscope. Edinb. Journ. Sci. vol. v. p. 183. Worsdell, W. C. (96) The anatomy of the stem of Macrozamia com- pared with that of other genera of Cycadeae. Ann. Bot. vol. x. p. 601. —— (97) On transfusion-tissue, its origin and function in the leaves of , Gymnospermous plants. Trans. Linn. Soc. vol. v. [2], p. 301. —— (98) The vascular structure of the sporophylls of the Cycadaceae. Ann. Bot. vol. xii. p. 203. —— (98?) The comparative anatomy of certain genera of the Cyca- daceae. Journ. Linn. Soc. vol. xxx. p. 437. —— (99) Observations on the vascular system of the female flowers of Conifers. Ann. Bot. vol. xu. p. 527. 520 LIST OF WORKS Worsdell, W. C. (00) The comparative anatomy of certain species of Encephalartos. Trans. Linn. Soc. vol. Vv. pt. xiv. p- 445. ' —— (002) The affinities of the Mesozoic fossil Bennettites Gibsonianus Carr. Ann. Bot. vol. xiv. p. 717. —— (01) Contributions to the comparative anatomy of the Cycadaceae. Trans. Linn. Soc. vol. vi. pt. ii. p. 109. —— (012) The vascular structure of the flowers of the Gnetaceae. Ann. Bot. vol. xv. p. 766. —— (04) The structure and morphology of the ovule. Ann. Bot. vol. xvii. p. 57. —— (05) Fasciation, its meaning and origin. New Phyt. vol. Iv. . 55. — 06) The structure and origin of the Cycadaceae. Ann. Bot. vol. xx. p. 129, ; Wright, W. B. (14) The Quaternary Ice Age. London. Yabe, H. (13) Mesozoische Pflanzen von Omoto. Sci. Rep. of the Tokoku Imp. Univ. (ser. ii. Geol.), Bd. 1. Heft iv. p. 57. Yasui, K. (17) A Fossil Wood of Sequoia from the Tertiary of Japan. Ann. Bot. vol. xxxi. p. 101. Yates, J. (55) Notice of Zamia gigas. Proc. Yorks. Phil. Soc. vol. 1. p. 37. Yokoyama, M. (94) Mesozoic plants from Kozuke, Kii, Awa, and Tosa. Journ. Coll. Sct. Imp. Univ. Japan, vol. vii. pt. iil. p. 201. —— (05) Mesozoic plants from Nagato and Bitchu. Ibid. vol. xx. Young, J. (69) Trans. Nat. Hist. Soc. Glasgow, vol. 1. pl. 1v. (No text.) —— (76) Catalogue of the Western Scottish fossils. Glasgow. Young, Mary S. (07) The male gametophyte of Dacrydium. Bot. Gaz. vol. xiv. p. 189. —— (10) The morphology of the Podocarpineae. Ibid. vol. L. p. 81. Zalessky, M. (05) Uber Friichte aus den Unter Carbon-Ablagerungen des Mstabeckens in Nord Russland. Bull. acad. Imp. sci. St. Péters- bourg, tome xxi. No. iii. p. 113. —— (05?) Notiz tiber die obercarbonische Flora des Steinkohlenreviers von Jantai in der siidlichen Mandshurei. Verhand. K. Russ. Min. Ges. St. Pétersbourg [2], Bd. xir. p. 385. —— (09) Communication préliminaire sur un nouveau Dadoxylon provenant du Dévonien supérieur du bassin du Donetz. Bull. acad. Imp. sei. St. Pétersbourg. —— (10) On the discovery of the calcareous concretions known as coal balls in one of the seams of the Carboniferous strata of the Donetz basin. Ibid. p. 477. —— (ll) Etude sur l’anatomie du Dadoxylon Tchihatcheffi Goepp. sp. Meém. com. géol. St. Pétersb. [N.S.], Livr. 68, p. 1. — (11) Note préliminaire sur le Coenoxylon Scotti, nov. gen. et sp. Etudes Paléobotaniques, pt.1. St. Petersburg. —— (12) Sur le Cordaites aequalis Goepp. sp. de Sibérie et sur son identité avec la Noeggerathiopsis Hislopi Bunb. sp. de la Flore du Gondwana. Mém. com. géol. St. Pétersb. [N.8.], Livr. 86. REFERRED TO IN THE TEXT 521 Zalessky, M. (12?) On the impressions of plants from the coal- bearing deposits of SudZenka, Siberia. Bull. soc. Natural. Orel. pt. Iv. —— (18) Flore Gondwanienne du Bassin de la Pétchora. I. Riviére Adzva. Bull. soc. Oural. @amis des Sci. Nat. & Ekatérinebourg, vol. XXXII. Zang, W. (04) Die Anatomie der Kiefernadel und ihre Verwendung zur systematischen Gliederung der Gattung Pinus. (Diss. Inaug.) Giessen. Zeiller, R. (78) Sur une nouvelle espéce de Dicranophyllum. Bull. soc. géol. France [3], vol. vi. p. 611. —— (80) Note sur quelques plantes fossiles du terrain permien de la Corréze. Ibid. vol. vit. [3], p. 196. — (96) Le Marquis G. de Saporta, sa vie et ses travaux. Bull. soc. géol. France [3], vol. xxiv. p. 197. —— (00) Sur les végétaux fossiles recueillis par M. Villiaume dans les gites charbonneux du Nord-ouest de Madagascar. Compt. Rend. Jun. 5. —— (02) Nouvelles observations sur la-flore fossile du bassin de Kousnetzk (Sibérie). Ibid. tome cxxxiv. p. 887. —— (05) Sur les plantes Rhétiennes de la Perse recueillies par M. J. de Morgan. Bull. soc. géol. France [4], tome v. p. 190. —— (05°) Sur quelques empreintes végétales de la formation char- konneuse supracrétacce des Balkans. Ann. Mines. — (08) See Douvillé and Zeiller. —— (11) Note sur quelques végétaux in‘raliasiques des environs de Niort. Bull. soc. géol. France [4], tome xt. p. 321. — (11°) Sur une flore triasique découverte a Madagascar. Compt. Rend. vol. cum. p. 230. - —— (12) Sur quelques végétaux fossiles de la Grande Oolite de Mar- quise. Bull. soc. acad. Boulogne-sur-mer, tome Ix. p. 5. —— (N.D.) See Malloizel and Zeiller. Zeiller, R. and P. Fliche. (04) Découverte de strobiles de Sequoia et de Pin dans le Portlandien des environs de Boulogne-sur-mer. Compt. Rend. tome cxxxvit. p. 1020. Zeiller, R. and L. M. Vidal. (02) Sobre algunas impresiones vegetales del Kimeridgense de Santa Marfa de Meya. Mem. Real Acad. Cienc. y Artes Barcelona, vol. tv. No. 26. Zigno, A. de. (53) Découverte d’une flore Jurassique analogue a celle de Scarborough dans les couches oolitiques des Alpes Vénitiennes. Bull. soc. géol. France [2], vol. x. p. 268. —— (73-85) Flora fossilis formationis Oolithicae. Vol. m. Padua, 1873-85. Zopf, W. (92) Ueber einige niedere Algenpilze (Phycomyceten) und eine neue Methode ihre Keime aus dem Wasser zu isoliren. Abh. Natur- forsch. Gcs. Halle, Bd. xvu. p. 79. INDEX (VoLtumes I.—IV.) Fossil Genera Abietites 1v. 369 Abiocaulis rv. 228 Acanthocarpus rr. 305 Acetabularia 1. 166 alcicularia 1. 166 «lcrostichites 11. 340 elctinostrobites Iv. 337 Adiantites 1. 376; m1. 173 Aetheotesta m1. 331 Albertia tv. 262, 300 Alethopteris 1. 573 aAlgites 1. 148, 204 Alloiopteris u. 470, 579; ur. 86, 109 Amphibennettites 11. 418 Amyelon mm. 261 Anachoropteris 1. 462 Anarthrocanna 1. 283 Androstrobus tr. 505 Androvettia Iv. 436 Aneimites 11. 346; ut. 173 Angiopteridium m1. 485 . Ankyropteris 1. 450 Annularia 1. 336 Anomaloxylon tv. 175, 242 Anomopteris 1. 329 Anomozamites m1. 548 Antarcticoxylon m1. 296 Antholithus rv. 51 Anthrophyopsis m1. 578 Aphlebia 1. 525 Araucariopitys Iv. 175, 234 Araucariopsis Iv. 179 Araucarioxylon m1. 248; 1v. 176 Araucarites Iv. 256 Arberia tr. 516; ut. 352 Archaeocalamites 1. 383 Archaeopitys 11. 290 Archaeopteris 111. 560 alrchaeosigillaria 1. 268 alrctopodium rt. 200 Arthrodendron 1. 326 Arthropitys 1. 325 alrlisia 11. 246 Aspidiaria m. 124 Asplenites 11. 580 Asterochlaena u. 448; m1. 205 Asterophyllites 1. 332 Asteropteris m1. 203 Asterotheca 1. 398 -Astromyelon 1. 342 Athrotaxites tv. 311 Athrotaxopsis tv. 312 Aulacopteris 1. 567; 1. 106 Bacillus 1. 135 Bactryllium 1, 154 Baiera tv. 33 Baieropsis iv. 36 Bambusium m1. 277 Barrandeina rv. 76 Beania ur. 502; rv. 51, 55 Becklesia tv. 345 Bennettites ut. 367; rv. 463 Bensonites 11. 469 Benstedtia rv. 445 Bergeria 11. 126 Bernouillia 1. 409 Blastolepis 11. 460 Blechnoxylon 1. 509 Boroviczia 11. 360 Bothrodendron 1. 248 Bothrostrobus 1. 262 Botrychioxylon m1. 459 Botryoconus 11. 264 Botryopteris 1. 436 Bowmannites 1. 401 Brachyoxylon tv. 322 Brachyphyllum rv. 311 Brittsia 1. 464 Brukmannia 1. 350 Buchlandia wu. 478 Bulbopodium ut. 368, 479 Calamites 1. 295 Calamitina 1. 367 Calamocladus 3. 332 Calamodendron 1. 327 Calamophyllites 1. 371 Calamopitys 1. 301; 11. 190; rv. 177 Calamopteris 111. 190 INDEX Calamostachys 1. 350 Calamosyriz u1. 190 Callipterrdium m1. 560 Callipteris u1. 557; 111. 109 Callitrites Iv. 337 Callixylon 11. 291 Calymmatotheca 11. 407; m1. 50 Camptopteris 11. 389 Cardiocarpon 11. 264 Cardiocarpus 11. 224, 333 Cardiopteris 1. 523 Carolopteris 11. 360 Carpolithes 111, 353, 497; Iv. 298, 351 Carpolithus 111. 363, 497; Iv. 34, 410, 419 Caulerpa t. 142, 158 Caulerpites 1. 142, 158 Caulopteris 11. 421 Cedrites Iv. 372 Cedrostrobus Iv. 385 Cedroxylon tv. 174, 212 Cephalotaxites Iv. 418 Cephalotaxopsis 1v. 422 Cephalotaxospermum Iv. 423 Cephalotheca 11. 537 Ceratozamia m1. 508 Chara 1. 223 Cheirolepis 1 84; Iv. 294 Cheirostrobus 1. 413; 1. 7 Chiropteris 11. 431 Chondrites 1. 144 Chorionopteris 1. 476 Cingularia 1. 364 Cladophlebis 1. 343 Cladosporites 1. 220 Cladoxylon m1. 200 Clathraria 1. 203; m1. 480 Clathropodium m1. 479 Clathropteris 11. 385 Clepsydropsis u. 448; m1. 204 Coccospheres 1. 120 Codonotheca nt. 124 Codonospermum m1. 330 Coelotrochium 1. 176 Coenoxylon tt. 252, 293 Colpospermum ui. 362 Colpoxylon m1. 109, 142 Colymbetes 11. 491 Compsotesta 111. 347 Condylites 111. 467; 1v. 311 Confervites 1. 177 Coniferocaulon Iv. 445 Comopteris 11. 367 Conites tv. 252, 351, 424 .- Conostoma 11. 79, 312 Cordaianthus tn. 264° Cordaicarpon it. 334 Cordaicarpus 111. 220, 334 Cordaicladus 111. 220 Cordaioxylon 111. 220 523 Cordaispermum tt. 335 Cordattes 1. 219 Cordaixylon m1. 223 Cornucarpus 11. 337 Corynepteris 11. 469; m1. 511 Coscinodiscus 1. 153 Crematopteris 11. 523 Crossochorda 1, 148 Crossotheca 1. 396; 11. 52 Crossotolepis tv. 400 Crossozamia 111. 500 Cruziana 1. 144 Cryptomeriopsis Iv. 335 Cryptomerites Iv. 335 Ctenidopsis 111. 580 Ctenis m1. 578 Ctenophyllum ut. 528, 585 Ctenopsis m1. 585 Ctenopteris 11. 548; 111. 511 Ctenozamites 1. 548 Cunninghamiostrobus tv. 331 Cunninghamites Iv. 433 Cupressinites Iv. 304 Cupressinocladus Iv. 303, 307 Cupressinoxylon Iv. 173, 186 Cupressites Iv. 305 Cycadella 111. 374, 417 Cycadeoidea 111. 367 Cycadeomyelon 111. 490 Cycadeorachis 111. 591 Cycadeospermum ut. 496 Cycadeostrobus 11. 503 Cycadinocarpus 111. 333, 341 Cycadites 111. 558 Cycadocarpidium Iv. 449 Cycadocephalus 111. 473 Cycadolepis m1. 494 Cycadopteris 11. 544 Cycadorachis 11. 590 Cycadospadix 111. 500 Cycadoxylon 1m. 185 Cyclocarpon 111. 335, 340 Cyclopitys Iv. 365 Cyclopteris 11. 571; 111. 130, 496 Cyclospermum i. 349 Cyclostigma m1. 251 Cyclozamia 111. 368 Cylindropodium 1. 368 Cymbella 1. 153 Cymopolia 1. 169 Cyparissidiwm Iv. 443 Czekanowskia 1v. 62 Dacrydium iv. 410 Dactylopora 1. 175. Dactyloporella 1. 176 Dactylotheca 1. 404 Dactyolepis tv. 438 Dadoxylon tt. 248, 289; 1v. 172, 176 Dammara tv. 248 524 Dammarites tv. 245 Dammarophyllum tv. 246 Danaeites 1. 398 Danaeopsis 1. 407 Dasyporella 1. 176 Decagonocarpus 11. 357 Dendrophycus 1. 146 Derbeyella 11. 516 Desmiophyllum m1. 236, 278; rv. 69 Dichopteris 1. 550 Dicranophyllum iv. 93 Dictyophyllum 11. 380 Dictyopteris 11. 571; ur. 546 Dictyothalamus 11. 127 Dictyozamites tr. 546 Dioonites m1. 509 Diplolabis 11. 446 Diplopora 1. 173 Diplopterotesta 111: 360 Diplotesta m1. 265 Diplotmema wu. 532; 11. 77 Discopteris 11. 403 Dolerophyllum ur. 130 Doliostrobus tv. 260 Dory-Cordaites m1, 224 Dory-Cycadolepis 111. 494 Drepanolepis tv. 442 Echinostipes 111. 368 Echinostrobus tv. 311 Edraxylon rt. 47 Elatides tv. 270 Elatocladus iv. 429 Eleoxylon tv. 213 Encephalartopsis 111. 508 Endolepis m1. 278 Entolepis Iv. 292 Entomolepis tv. 399 Eophyton 1. 144 Ephedrites tv. 469 LEiquisetites 1. 257 Eremopteris 11. 169 Eretmophyllum tv. 58 Eriotesta wt. 319, 332 Eristophyton m1. 194 Etapteris 11. 435, 465 Eucalamites 1. 376 Lu-Cordaites 111. 580 Eugeinitzia iv. 361 Eury-Cycadolepis, m1. 495 Euryphyllum ui. 238 Favularian. 198 Feildenia tv. 69, 69 Feistmantelia ut. 514 Fittonia m1. 487 Flicheia 1. 448 Frenelites tv. 336 Frenelopsis tv. 342 Fricia ut. 505; 1v. 255 INDEX Fucoides 1. 142 Gallionella 1. 163 Gangamopteris 11. 512; m1. 258 Gaudrya 1. 332 Geinttzia Iv. 357 Gigantopteris 1. 557 Ginkgo (see Ginkgoites) Ginkgoanthus rv. 54 Ginkgocladus iv. 33 Ginkgodium tv. 60 Ginkgoites tv. 10 Ginkgophyllum iv. 86 Ginkgopsis Iv. 77 Girvanella 1. 124 Gleichenites m1. 351 Glenopteris 1. 538 Gloioconis 1. 130 Glossopteris 11. 496; 11. 140 Glossozamites III. 530; Iv. 447 Glottophyllum Iv. 76 Glyptostrobites Iv. 357 Glyptostroboxylon Iv. 173, 198 Gnetopsis 11. 317; Iv. 470 Gomphonema i. 153 Gomphostrobus 11. 253; Iv. ann Goniolina 1. 176 Grammatopteris 11. 434 Gyroporeila 1. 175 . Halimeda 1. 164 Haliserites 11. 27 Halonia a. 135 Hapalopteris 11. 406 Hapaloxylon Iv. 286 Hausmannia 11. 390 Hawlea 11. 398 Heterangium 11. 70, 179 Hexagonocarpus ut. 116, 356 Hexapterospermum i. 321 Hierogramma ut. 200 Holcospermum ut. 361 Hostinella 1. 28 Hymenophyllites 11. 363 Isoetes 1. 66 Isoetites 11. 67; rv. 440 Isoetopsis 11. 68 Jeanpaulia Iv. 33 Jordania 111. 336 Juniperites rv. 310 Kalymma ut. 194 Kidstonia 1. 325 Kiltorkensia tv. 88 Klukia 1. 347 Knorria 11, 124 Krammera 11. 277; 1v. 248 INDEX Laccopieris 11. 355 Lagenospermum ut. 64, 316 Lagenostoma wt. 55, 307 Laricites tv. 372 Lepidocarpon 1. 271 Lepidocaryon ut. 346 Lepidodendron 1. 93 Lepidophloios 11. 104 Lepidophyllum 1. 181 Lepidopteris 1v. 53 Lepidostrobus Iv. 181 Leptrostrobus Iv. 294 Lesleya 11. 517 Libocedrites Iv. 308 Linatopleris a. 544 Linopteris 11. 572; 11. 86, 111 Lithothamnion 1. 185 Lomatopteris 11. 544 Lonchopteris 11. 494, 576 Lycopodites 11. 76; 1v. 273 Lycostrobus 11. 88; 11. 506 Lyginodendron 111. 36 (see also Lygino- — pleris) Lyginopteris u1. 35 Lygodium 1. 350 Macrostachya 1. 362 Macrotaeniopteris 11. 486 Malacotesta 11. 363 Mantellia 111. 367 Marattiopsis m1. 407 Marchantites 1. 233 Mariopteris u. 532; m1. 111 Marsilidium u. 474 Masculostrobus 111. 505; Iv. 424 Matonidium u. 361 Medullosa m1. 86 Megalopteris 1. 509 Megalorhachis 11. 204 Megalospermum m1. 326 Megaloxylon m1. 175 Megaphyton a. 422 Mesembrioxylon tv. 173, 203 ' Mesopitys m1. 295 Mesostrobus 11. 195 Mesoxylon m1. 214, 270 Metacedroxylon tv. 175, 238 Metaclepsydropsis 1. 447 Metacordaites 111. 260 Miadesmia 11. 275 Micrococcus 1. 136 Microdictyon 11. 369 Microspermum v1. 365 Mitrospermum m1. 345 Mittagra m1. 63 Mixoneura 11. 555 Moriconia iv. 332 Mucor 1. 213 Muscites 1. 238 Musocarpus ut. 361 525 Myeloxylon 11. 89; 105 Myxomycetes 1. 205 Nageiopsis iv. 456 Nathorstia 1. 410 Nematophycus 1. 192 Nephropsis v.77 * Neuropteridium u. 519 Neuropteris 1. 565; 111. 56, 109, 113 Neuropterocarpus m1. 116 Neuropteromedullosa 11. 156 Neurospermum 111. 116 Neurotheca 11. 110 Nitssonia m1. 511, 566 Nilssoniopteris 111. 569 Niponophyllum 1. 282 Noeggerathia 11. 428; 111. 589; 1v. 79 Noeggerathianthus 111. 264 Noeggerathiopsis u1. 220, 238 Odontopteris 11. 552; 111. 86 Oldhamia 1. 145 Oligocarpia 11. 352 Omphalophloios 11. 264 Oncopteris 11. 372 Onoclea 11. 380 Onychiopsis 1. 377 Oochytrium 1. 216 Ophioglossites 11. 428 Osmunda 11. 339 Osmundites 11. 331 Otozamites 111. 460, 537 Ottokaria 11. 498; m1, 124, 139 Ovulites 1. 161 Pachypteris 11. 550 Pachytesta 1. 574; m1. 115, 323 Pachytheca 1. 202 Pagiophyllites 1v. 177 Pagiophyllum rv. 181, 270 Palaeachyla 1. 127 Pataeocladus iv. 417 Palaeocyparis 1v. 304 Palaeohepatica 1. 234 Palaeolepis tv. 442 Palaeomyces 1. 222 Palaeoperone 1. 215 Palaeopitys 111. 210 (footnote) Palaeoporella 1. 176 Palaeopteris 11. 560 Palaeostachya 1. 350, 357 Palaeoxylon 111. 100 Palamophyllum tv. 80 Palissya tv. 315 Palmatopteris 11. 532 Paracalamostachys 1. 361 Paracedroxylon tv. 173, 212 Paracupressinoxylon Iv. 189 Parapecopteris 11. 398 Paraphyllocladoxylon tv. 205 526 INDEX Parapitys m1. 294 Protophyllocladus tv. 414 Pecopteris 1. 576; m1. 166 Protopiceoxylon tv: 176, 231 Pecopteromedultosa m1. 156 Protopitys 11. 207 Pelourdea m1. 277 Protopteris 1. 370 Penicillus ¥. 161 Protosalvinia a. 476 Peridium 1.117 | Psaronius u. 412; m1. 88, 168 Peronosporites 1. 218 Pseudo-Araucaria tv. 261 Phoenicopsis m1. 236; tv. 63, 452 Pseudoasterophyllites 1v. 426 Phycodes 1. 149 Pseudobornia u. 8 Phycopsis 1. 192 Pseudoctenis 11. 584 Phylladoderma ut. 244 Pseudocycas 11. 558 Phyllocladites rv. 413 Pseudofrenelopsis 1v. 345 Phyllocladopsis tv. 417 Pseudogeinitzia 1v. 361 Phyllocladoxylon rv. 203 Psilophyton 1. 26 Phyllostrobus tv. 311 Psilotiphyllum 1v. 288 Phyllotaenia 1. 280 Psilotites 1. 24 Phyllotheca 1. 281 Psygmophyllum tv. 34, 79 Physematopitys tv. 9 Pteridotheca 11. 325; un. 54 Physostoma 111. 62, 309 Plerispermostrobus um. 66 Piceoxylon tv. 229 Pterophyllum m1. 548 Pila 1. 180 Pterospermum 1. 357 Pinakodendron u. 264 Ptilophyilum mm. 379, 512 Pinites m1. 35, etc.; 1v. 101, 387 Ptilozamites m. 546; ut. 511 Pinostrobus Iv. 387 Ptychocarpus 1. 397 Pinoxylon Iv. 220 Ptychopteris 11. 422 Pinus Iv. 216, ete. Ptychotesta m1. 321 Pinuxylon tv. 1li4 Ptychoxylon ww. 187 Pityanthus wv. 371, 395 Pychnophyllum ut. 223 Pityites Iv. 370 Pyxidicula t. 154 Pityocladus iv. 377 Pityolepis Iv. 371 Rachiopteris . 436, 448; m1. 47 Pityophyllum iv. 378 Radiculites m1. 217 Pityospermum iv. 396 Radiospermum ut. 64 Pityosporites Iv. 398 Raritania 1v. 438 Pityostrobus tv. 371 Raumeria 1. 409 Pityoxylon ut. 251; rv. 219 Reinschia 1. 106 Pitys U1. 285 Renauiltia u. 406 Plagiozamites m1. 589 Retinosporites Iv. 418, 433, Planoxylon tv. 176, 244 Rhabdocarpus 111. 90, 341 Platylepis m1. 369 Rhabdoporella 1. 176 Platypterygiwm 1. 555 Rhabdospermum ur. 116, 339 Platypterospermum 111. 362 Rhabdospheres 1. = Pleuromera u1. 68 Rhacophyllum w. 52 Poa-Cordaites 11. 224 Rhacophyton a. 537. Podocarpites tv. 405 Rhacopteris 1. 427 Podocarpoxylon Iv. 173, 204 Rhetinangium ut. 181 Podocar pus tv. 408, £11 Rhexoxylon m1. 146 Podozamites 1v. 447 Rhipidopsis tv. 90 Poecilitostachys 1. 91 Rhiptozamites wu, 238 Polylophospermum 1. 329 Rhizocordaites u1. 224 Polyporus 1. 221 Rhizocupressoxylon tv. 201 Polypterocarpus Wt. 357 Rhizodendron a. 375 Polypterospermum 1, 323 RLhizomopteris 0. 381 Poroxvylon m1. 83, 214 Rhizotaxodioxylon iv. 199 Potontea ur. 111 Rhodea tr. 364 Prepinus tv. 400 Rhynchogonium m1. 358 _ Protaeophyllum rv. 471 Rhytidodendron 1, 251 Protoannularia 1. 338 Rhytidolepis 1. 198 Protocedroxylon iv. 236 Rogersia tv. 471 Protodammara tv. 244 Rosellinites 1. 209 INDEX P 527 Ruffordia u. 350 Sagenopteris 11. 477 Salvinia, 11. 476 ’ Samaropsis 11. 168, 265, 333; tv. 90 465 Sapindopsis iv. 471 Saportaea Iv. 92 Sarcostrobus tv. 259 Sarcotaxus 1. 265 Saxegothopsis tv. 413 Schizaeopsis iv. 36 Schizodendron 111. 250; tv. 282 Schizolepidella tv. 442 Schizolepis Iv. 377, 439 Schizoneura 1. 254, 291 Schizopodium m1. 369 Schizopteris m1. 464, 525; Iv. 38 Schizospermum im. 319 Schizostachys 11. 464 Schizoxylon 111. 200 Schiitzia m1. 126 Sciadopitytes Iv. 366 Scleropteris 1. 552 Scolecopteris 11. 401 Scuto-Cordaites 111. 237 Selaginellites u. 74, 85 Senftenbergia u. 347; 1. 63 Sequoia 1v. 347 Sequoiites Iv. 349 Sewardia m. 589; tv. 103 Sigiilaria u. 69, 196 Sigillariophyllum 1. 200 . Sigillariopsis 1. 213 Sigillariostrobus 1. 215 Siphonema 1. 160 Solenites rv. 66 Solenopora 1. 189 Sorocladus 11. 110 Speirocarpus 11. 409 Spencerites 11. 192 Sphaerites 1. 209 Sphaerocodium 1. 160 Sphaerostoma tt. 78, 316 Sphenolepidium tv. 277, 362 Sphenolepis iv. 363 Sphenophyllostachys 1. 400; 11. 1 - Sphenophyllum 1. 387; 1. 1 Sphenopleris 1. 529; m1. 49, 109 Sphenothallus 1. 148 Sphenozamites 11. 587 Spirophyton 1. 144 Spiropteris 1. 579 Sporocarpon 111. 309 Stachannularia 5. 340, 361 Stachyopitys Iv. 54 Stachyopteris 1v. 54 Stachyotaxus tv. 410 Staphylopteris a. 531 Stauropteris 11. 465 Steloxylon ut. 163 Stenomyelon 11. 183 Stenorachis ut. 503; tv. 51, 54 Stephanospermum 111. 302, 326 Stigmaria 11. 226 Stigmariopsis 11. 233 oe m1. 124, 141; Iv. 296, 299, Sturiella 1. 324 Stylocalamites 1. 367, 374 Subzamites m1. 521 Suicliffia 11. 87, 149 Swedenborgia tv. 295 Sycidium 1. 173 Syncardium m. 200 Synedra 1. 153 ~ Taeniopteris 11. 485; m1. 1, 467 Taxites tv. 202, 417 Taxodioxylon rv. 173, 199 Taxodites 1v. 328 Taxospermum 1. 265, 347 Taxoxylon iv. 173, 203 Telangium u. 532; m1. 54 Thamnocladus 1. 27 Thamnopteris 1. 329 Thaumatopteris 11. 385 Thinnfeldia u. 537; tv. 414 Thuites tv. 303 Thuya tv. 305 Thylloxylon iv. 176, 243 Thysanotesta 111. 304 Titanophyllum U1, 283 Todeopsis 11. 324 Todites 1. 339 Torellia 1v. 68 Torreya iv. 421 Torreyites Iv. 418 Tracheotheca 11. 437 Trichopitys tv. 48, 101 Trigonocarpus ut. 61, 115, 319 Triletes 11. 192 Trioolepis Iv. 424 Triploporella 1. 177 Triplosporites 11. 190 Tripterospermum I. 321 Trizygia 1. 411 Tubicaulis 11. 434 Tylodendron 111. 250; Iv. 282 Tysonia IIL. 385 Ullmannia 1v. 262, 296 Ulimannites ut. 199; tv. 179 Ulodendron u. 128 Ulospermum 11. 497 Urnatopteris 11. 396, 407 Urophlyctites 11. 247 Uteria 1. 177 Vectia ut. 419; tv. 170 528 Vermiporella 1. 172 Vertebraria 11. 501 Vesquia tv. 422 Volkelia 111. 208 Volkmannia 1. 350, 491 Voltzia rv. 181, 282 Voltziopsis 1v. 294 Walchia tv. 277, 286 Wardia m1. 172 Weichselia | 494 Weltrichia 1. 475 Whittleseya 111. 128 Widdringtonia tv. 339 Widdringtonites 1v. 336 Wielandia ot. 463 Wielandiella m1. 454, 463 Williamsonia m1. 372, 421, 531 Williamsoniella ut. 434, 467 Withamia tv. 103 Woodwardites 11. 377 INDEX Woodworthia tv. 133 Xenophyton 11. 158 Xenopteris 11. 555 Xenoxylon m. 213; rv. 175, Yatesia 11. 386 Yezonia Iv. 327 Yezostrobus tv. 327 Yuccites m1. 277 Zalesskya u. 326; m1. 180 Zamiophyllum m1. 529 Zamiopsis 1. 512 Zamiostrobus m. 503 Zamites 111. 281, 529 Zeilleria u. 407 Zeugophyllites 1v. 74 Zonatrichites 1. 129 Zygopteris 11. 418, 443 Zygosporites 1. 220 238 INDEX (VotumE IV.) Aase, H. C. 115, 142 Abies 108, 122-139 (passim), 156, 195, 213, 270, 324, 369-371, 383, 387 A. alba 116 A. balsamae 137 A. bracteata 114, 157 it. concolor 157, 158 al. Fraseri 157 A. homolepis 137 aA. Loehri 390 A, magnifica 140, 156 A. oblonga 385, 387 A? pectinata 134, 157 A. Veitchit 137 al. violacea 158 A. Webbiana 158, 189, 213 Abietineae 124, 125, 153-158, 368-404 Abietites 369-373, 387 A. Dunkeri 383 A. ellipticus 373 Abiocaulis zezoense 228 Acanthozamites 105 - Acmopyle 127, 160 -lcrostichopteris 36 lctinostrobites 337 ctinostrobus 123-126, 151, 336, 338 wl. miocenica 337 A. pyramidalis 154 Aeschynomene 287 Agardh, J. G. 410 Agathis 111-125 (passim), 133, 136, 140, 143, 145, 164, 178, 237, 245- 261 (passim), 300, 447, 451 A. australis 144 al. bornensis 132 A. loranthifolia 113, 143 A. macrophylla 143 al. Moorei 113, 144 al. robusta 143 A. speciosa 300 A. vitiensis 144 Albertia 262, 292, 300, 301 A. Braunit 300 A. elliptica 300 A. latifolia 300 A. speciosa 300 Alethopteris Ottonis 53 Ss. IV Anacardium 122 Androvettia 416, 436-438 A. carolinensis 438 A, elegans 437 lL. slatenensis 436, 437 Angiodendron 283 Anomaloxylon 175, 232, 242, 243 A, magnoradiatum 242, 243 Antevs, E. 53, 169 Antholithus 53 A. Schmidtianus 51, 57, 58 A. Zeillert 52-54 Araucaria 140-142; passim A. Balansae 145 A. Bidwillit 111, 114, 117, 126, 132, 135, 145, 183, 262-270 (passim), 296, 299, 435, 457 A. brasiliensis 112, 137, 186, 270, 435 aA. Cookii 111, 113, 114, 132, 256, 262 — var. luxurians 111 A. Cunninghamii 112, 268-270 A. Danai 269 at. excelsa 113, 114, 145, 164, 259-296 (passim), 359 al. Goepperti 260 A. Haasti 183 A. Housteinit 145 A. imbricata 125, 126, 141, 262-269 (passim), 445 A. montana 111 A. Muellert 110, 111 A. Nathorsti 269 al. peregrina 276 A. Rulet 110, 282 A. Toucasi 267 Araucarineae 124, 125, 143, 145, 245- 277 Araucariopitys 63, 175, 234-236 A. americana 235, 323 Araucariopsis 179, 184 Araucarioxylon 172, 176-186, 244 A. koreanum 186, 240 A. latiporosum 241 A, Lindleti 244 al. thuringicum 181 A. vanartsdaleni 181 al. virginianum 181, 185 34 530 A, wiirtembergicum 181 Araucarites 176, 177, 256-270, 280, 364 A. bladenensis 265, 266 . Brodiei 259, 263-265 . Cleminshawi 258 . cutchensis 263-265 . Delafondi 262 . Falsani 264 Goepperti 267-269, 334 Haastii 269 Haberieinii 263, 264 Hatcheri 267 hespera 260 Hudlestoni 258 imponens 269 . Jeffreyi 263, 265 keuperianus 294 macroplerus 264 . microphylla 264 . Millert 263, 264 Moreauana 263 . Nathorsti 266, 269 Oldhami 266 . ooliticus 256-259, 263 ovatus 266 patapscoensis 265 Phillipsi 259, 263, 265 pippingfordensis 259 Reichenbachii 357, 358 . Rogersi 263, 264 Schleiniizi et Hookeri 185 . schoeneggensis 267 . sphaericus 259 . sphaerocarpus 256-258 . Sternbergii 260, 334, 335 A, valdajolensis 279, 281, 286 A. wyomingensis 264, 265 Araucarites (Conites) sp. 259 A. (Sarcostrobus) Paulini 259 Arber, E. A. N. 12, 40, 74, 80-89 (passim), 178, 21] Arber, E. A. N. and J. Parkin 461, 469 Arnoldi, W. 123 Artisia 178 Atherstone, W. G. H. 264 Athrotaxis 115, 118, 123, 126, 150, 312 -315, 365 A. cupressoides 151, 163, 314, 320 A, laxifolia 116, 151 A. selaginoides 150 A, subulata 268, 315 Athrotaxites 304, 311-315, 363 A. lycopodioides 311-314 A, Ungeri 313, 314 Athrotaxopsis 312, 313, 363 BER DRDEREERE RARE RERRE BERR REDE RRR Baccharis scolopendra 164 Baiera 11, 12, 16, 33-50, 87, 101, 103 B. angustiloba 46 B. arctica 29 B. australis 46, 48 B. bidens 46 B. Brauniana 25, 49, 50 B. Czekanowskiana 47 B. delicatula 46 B. dichotoma 44 B. digitata 37, 38 B. foliosa 36, 96 — forma Muensteriana 46, 47 Guilhaumali 47 incurvata 46 Lindleyana 35, 47-50, 65, 67, 438 longifolia 35, 40, 41, 44, 47, 48, 105 microphylla 49, 65 moltenensis 16 Muensteriana 34, 38, 44, 45, 53 mullifida 39, 42, 44 palmata 42 paucipartita 35, 39, 40, 66 Phillipsi 25, 48 pluripartita 27 pulchelia 42 Raymondi 38 Schenkii 44. Simmondsi 35, 38, 40-43, 47 spectabilis 35, 36, 40-42, 44 spetsbergensis 50 Steinmanni 44 stormbergensis 44 . taeniata 44 . virginiana 37, 38 Baieropsis 36 Bailey, I. W. 156, 229 Baily, W. H. 334, 392 Bancroft, N. 446 Barber, C. A. 9, 163, 191, 213 Barrandeina Dusliana 76 Bartholin, C. T. 42 Bary, A. de 140 Beania 51, 55 B. gracilis 56 B. geminata 56 Becklesia anomala 345 Beissner, L. 161, 328 Bennettites 461, 463, 469 B. Morieri 464 Benstedtia 445, 446 B. Benstedti 446 Bergeron, J. 279, 282 Berry, E. W. passim be Bo Bo BB yy by bs sb bs bs By ba bbb * Bertrand, C. E. 422 Beust, F. 9, 189, 198, 308, 355 Biota orientalis 309 B. succinea 305 Blanckenhorn, M. 291 Bommer, C. 145, 162 Boodle, L. A. 151 INDEX Bowerbank, J. 8. 337 Brachyoxylon 322-324, 361 B. notabile 322, 323 B. pennsylvanicum 218 — - Brachyphyllum 251, 274, 275, 304, 324 -333 (passim), 364, 365, 429, 430, 435, 443 . crassicaule 321, 326 crassum 316, 319, 323-327, 333 . eathiense 216, 316, 326, 327 . expansum 309, 317-320 . gracile 321 . macrocarpum 251, 316, 322, 325, 439 — var. formosum 326 mamillare 315, 318-320 Joreauanum 321 Muensteri 294 nepos 321 obesiforme 322 obesum 321, 322, 326 setosum 294 speciosum 315 . spinosum 321 . vulgare 327 Braun, C. F. W. 34, 44, 48, 426-429 Brauns, D. 12, 88 Brongniart, A. passim Brooks, F. T. 228 Buckman, J. 19, 455 Bunbury, C. F. 46 by by by tu by by bo bo bo by bo bo be be be Calamopitys 177 Callitrineae 124, 126, 151-153, 336- 346 Callitris 120, 151, 152, 336, 337, 342 C. arborea 152 C. calcarata 163 C. glauca 152, 164 C. Macleayana 153 C. rhomboidalis 152 Callitrites 337-342 . antiqua 342 . brachyphylla 342 Brongniarti 340-342 curta 340 Eitingshauseni 340 . europaea 341 . helvetica 341 » prisca 341 . Reichit 338 C. subtilis 339 Cambier, R. and A. Renier 80 Camptophyllum 434 C. Schimperi 434 Cardiocarpus triangularis 299 Carpolithes frumentarius 298 C. hemlocinus 351 C. nitens 419 C. septentrionalis 410 C. striolatus 56 aagaagaaga 531 Carpolithus ternatus 34 Carruthers, W. 51, 256-263, 351, 385, 386, 390 Caspary, R. 179, 184 Caulerpa septentrionalis 410 Caulerpites 312 Cedrites 372 Cedrostrobus 385 C. Leckenbyi 385 C. Mantelli 386 Cedrozylon 174, 181, 187, 207, 212- 219, 229, 231, 232, 324, 369 . affine 219 . australe 218 barremianum 218 . blevillense 217, 218 . cedroides 216 Hermanni 219 Hoheneggeri 219 Hornet 215-218 maidstonense 217 manekildense 218 matsumurae 218 pottoniense 217 . reticulata 218 . transionis 214-218, 225 - Yendoi 219 edrus 108, 122, 125, 130, 139-141, 155, 213-217, 261, 369-386 (passim), 403 C. atlantica 137 C. Leet 386 C. Libani 141, 157 C. oblonga 385-388 C. vivariensis 369 Cephalotaxites 418 C. insignis 423 Cephalotaxopsis 418, 422, 423, 432 C. brevifolia 423 ~ C. magnifolia 422, 423 C. microphylla 423 C. ramosa 422 Cephalotaxospermum carolinianum 423 Cephalotaxus 121, 128, 161, 164, 169, 202, 417-423, 432 C. drupacea 161 2 C. europaea 423 C. Fortunei 141 C. francofurtana 423 C. Henry 161 C. koraina 161 : Chamaecyparis 126, 146, 302, 304 Cheirolepis 294, 440 C. Escheri 294 C. Muensteri 294 Chrysler, M. A. 131, 140 Cockerell, T. D. A. 355 Coemans, E. 385 Coker, W. C. 123 Conchophyllum Richthofeni 410 34—2 oaagagagagsasaaa 5382 Condylites 311 Coniferales (fossil) 165-446 — (recent) 106-164 Coniferocaulon colymbeaeforme 446 Conites 351, 424 C. Gardnert 351 C. Juddi 251-256 C. ovalis 352 Conwentz, H. passim Corda, A. J. 248, 390 Cordaioxylon 177 Cordaites 70, 176, 178, 180, 280, 403 C. (Noeggerathiopsis) Hislopi 75 Cramer, C. 197, 238 Crassula lycopodioides 163 Crié, L. 177, 178, 218 Crossotolepis 400 ~C. Perrott 400 Cryptomeria 108-110, 115, 118, 140, 141, 149, 194, 213, 268, 272, 289, 296, 312, 334, 335 C. japonica 108, 109, 268, 334, 335 — var. spiralis 164 C. primaeva 357 C. Sternbergit 334 Cryptomeriopsis 335 C. antiqua 335 C. mesozoica 335 Cryptomerites 335 C. divaricatus 334 C. du Noyert 334 Cunninghamia 110, 115, 118, 122, 123, 140, 149, 150, 199, 256, 331, 332, 424, 428, 433-436, 443 C. elegans 433, 435, 436 C. Konishii 331 C, sinensis 116, 331, 332, 362, 435 C. squamosus 436 C. stenophylla 436 Cunninghamiostrobus 331, 332 C. yubariensis 331 Cunninghamites 433-436 C. borealis 436 C. confertus 431 C. dubias 434 C. miocenica 436 C. oxycedrus 433 C. Sphenolepis 426 Cupressineae 124, 126, 146-151, 302- 335 Cupressinites 304 Cupressinocladus 303, 304, 307-311, 445, C, hypnoides 310 C. massiliensis 309 C. salicornoides 307, 308 C. valdensis 309 Cupressinorylon 9, 158, 173, 186-202, 204, 206, 212, 213, 305, 348 INDEX . antarclicum 198 . Barberi 189, 241 cryptomerioides 193 Holdenae 194, 196 Hookeri 189, 210, 211 Hortit 193, 194 Koettlitzi 195, 196 liasinum 190 luccombense 193 McGeei 192 polyommatum 196, 197 pulchrum 229 Sequoianum 189 Taxodii 201 taxodioides 197 tener 198 uniradiatum 201 . vectense 191, 192 . (Glyptostroboxylon) neosibiricum 197 (T'axodioxylon) Sequoianum 200, 201 C. (Taxodioxylon) Taxodii 197, 201 Cupressites 303-307 C. elegans 407 | C. elongatus 337 C. globosus 337 C. MacHenryi 305 C. taxiformis 305, 306 2 Cupressus 120, 126, 134-148 (passim), 299-318 (passim), 340, 343 C. Lawsoniana 322 C. MacNabiana 147 C. Pritchard 305 C. sempervirens 116 C. sempervirens succinea 307 Cycadeoidea 466, 467 Cycadeostrobus 259 Cycadites 378 C. zamioides 431 Cycadocarpidium 447, 449-451 C. Erdmann 449-451 C. redivivum 449 C. Swabii 449, 450 Cycadorachis armata 104 Cycas 7, 164 C. revoluta 135, 136 Cyclopitys 365 Cyclopteris 89 C. crenata 12, 88 C. digitata 14, 27 C. incisa 23 C. squamata 59 Cyparissidium 363, 443-445 C. cretaceum 445 C. gracile 444, 445 C. minimum 339, 444, 445 C. Nilssonianwm 444 C. septentrionale 411, 444 Cyperites 97 C. bicarinata 97 an oaasaaaasaagasaasaa INDEX Czekanowskia 35, 40, 62-68, 236, 373 . capillaris 67 dichotoma 67 . microphylla 49, 65, 66 . Murrayana 64-67 . nervosa 49, 67 . rigida 63, 65-68 . selacea 67 aaaeaaag Dacrydium 110, 120, 127, 134, 138, 158, 160, 204, 334, 398, 410-413, 428 . araucarioides 160, 164 Balansae 116 Bidwitlit 164 cupressinoides 410 cupressinum 140, 164 densifolium 410 elatior 160, 164 Franklini 161, 164 . Kirkii 164 . laxifolium 108, 164 Pancheri 160 . prae-cupressinum 410 Dactyolepis 438 D. eryptomerioides 438 SEPESESESSESE Dadoxylon 172, 176-186, 221, 222, 280, 288, 301 . aegyptiacum 186 . albianum 184 Arberi 178 argillicola 182 australe 177, 178, 18] © barremianum 185 Brandlingii 177 Dantzii 184 divescense 182 Doeringit 186 keuperianum 181, 294 madagascariense 185 mahajambjense 182 noveboracense 185 permicum 177 protopityoides 241 Rhodeanum 177 Rovertianum 186 saxonicum 177 septentrionale. 179-188 tankoense 185 valdajolense 177 virginianum 185 . Zuffardii 185 x (Araucarioxylon) breviradiatum 184 D. (Araucarioxylon) kerguelense 185, 186 D. (Araucarioxylon) novae zeelandiae 182 PHSSSSSSSSSSOESSSSSSSSHSh D. (Arducarioxylon) pseudoparenchy- matosum 186 Dammara 246, 310 (see Agathis) D. Armaschewskii 248 533 D. microlepis 248 D. Oweni 183 D. Tolli 248 Dammarites 245-250, 452 . acicularis 250 albens 247 Bayeri 247 . borealis 248-253, 255 caudatus 246 cliffwoodensis 250 crassipes 247 emarginatus 246 minor 250 northportensis 250 Oweni 247 Tolli 248 . uninervis 247 Dammarophyllum bohemicum 246 D. striatum 246 Dammaropsis Kingiana 145 Dawson, J. W. 89, 100 Debey, M. H. and C. von Ettings- hausen 332 Dehaasia media 164 Depape, G. 29 Desmiophyllum 69-73, 89, 247, 288 D. gracile 70 D. Solmsi 71, 72 Dicranophyllum 36, 93-103, 292, 372 . anglicum 98 . angustifolium 99 . australicum 100 Beneckianum 94, 98, 99 dichotomum 100 dimorphum 100 gallicum var. Parchemineyi 96 glabrum 100, 102 latifolium 287 longifolium 98, 99 lusitanicum 97 Richiri 98 . robustum 95, 98 . striatum 96, 99 . Sternbergii 260 . tripartitum 97 Dicropteris longifolia 47 Dioncophyllum Tholloni 105 Dioon 450 Dippel, L. 283 Diselma 149 - Distrigophyllum lusitanicum 97 Dracaena Benstedti 445 Drepanolepis 413, 442 D. angustior 442 D. rotundifolia 416, 442 Drimys Winteri 162, 241 Dun, W. 8. 84 Dunker, W. 14, 273, 364 Dusén, P. 269, 270, 413 Dutt, C. P. 390 SSSSSSSSSssyd SSSSSSSSSsSsysyssy 534 Echinostrobus 304, 311-315 EF. expansus 318, 320, 321 E. rajmahalensis 321 E. rhombicus 321 E. Sternbergii 312, 314 Eichler, A. W. 1 Eichwald, KE. 23, 283, 338 Elatides 237, 239, 266, 270-275, 315, 359, 360, 363, 426 E. Brandtiana 270, 273 E£. curvifolia 270-274, 359, 361, 364, 426 — var. Brandtiana 274 — var. ovalis 273 E. falcata 270 E. ovalis 270, 273 E, Sternbergit 270, 271 E, Williamsonis 270-276 ELlatocladus 332, 405, 412, 418, 429-435 E. conferta 431, 435 Ei, elegans 435, 437 E. heterophylla 429-431 E. plana 431 : Ei. zamioides 431 Eleoxylon 213 Encephalartos 7, 450 Endlicher, 8. 176, 300, 328, 357, 370, 389, 426 Engejhardt, H. and F. Kinkelin 331, 341, 370, 394, 421, 423 Engler, A. 1 Entolepis 292 Entomolepis 399, 400 E. cynarocephala 399 Eotaxites 93 Ephedra 459-462 E. altissima 460 FE. campylopoda 460 Ff. distachya 460, 461 E. fragilis 469 FE. Mengeana 470 Ephedrites antiquus 470 E. Johnianus 470 FE. Sotzkianus 469 Equisetum brachyodon 340 Eretmophyllum 58-61, 76, 451 E. pubescens 59 E. saighanense 59, 60 E. whitbiense 59 Eristophyton Beinertianum 177 Essner, B. 8, 9 Etheridge, R. 74 Ettingshansen, C. von passim Eucalyptus 249 BE. Geinitzii 249, 250 Bugeinitzia 361, 362 HR. proxima 361; 362 Ewart, A. J. 461 Feilden, Capt. 68 INDEX Feildenia. 60, 68, 69 F. Nordenskidldi 69 F. rigida 68 Feistmantel, O. passim Felix, J. 8, 10, 168, 190, 199, 201, 218, 231 Ficophyllum 471 Ficus 471 F. virginiensis 471 Fitton, W. H. 259 Fitzroya 122, 126, 136, 149 F. patagonica 123 Flabellaria wiminea 65 Fliche, P. 182, 218, 239, 259, 261, 279, 280, 292, 385 Fokienia 122, 123, 126, 150 Fontaine, W. M. passim Fontaine, W. M. and I. C. White 92 Frenela 336, 342 F. europaea 342 F, Bwaldana 342 Frenelites 336, 337 F. europaeus 342 F. Reichti 330, 378 Frenelopsis 310, 342-346 F. bohemica 344 F. gracilis 343, 438 F. Hoheneggeri 344, 345 F. occidentalis 344, 345 F. parceramosa 345 F. ramosissima 344-346 Fri¢, A. and E. Bayer 246 Fricia nobilis 255 Fucoides 298 FF, digitatus 37 Fujii, K. 6 Fujioka, M. 149 Gardner, J. 8. passim Geinitz, H. B. 37, 287, 299, 357 Geinitzia 212, 277, 357-362 G. cretacea 357 G. formosa 360 G. gracillima 330, 352, 361 G. Reichenbachti 358, 361, 362, 437 Geyler, T. and F. Kinkelin 370 Gibbs, L. S. 412 Ginkgo 1-11, 98, 94, 234, 391, 403 (see also Ginkgoites) G. adiantoides 10, 11, 17 G. antarctica 43 G. biloba 1-14, 18, 19, 29-32, 50, 56, 58, 76, 80 . concinna 35, 46 G. crassipes 23 G. crenata 88 G. cuneata 76 G. Czekanowskii 77 G. digitata 25, 50 G. eocenica 33, 50 R INDEX 535 G. flabellata 22, 25 G. grandiflora 57 G. Hermelini 22 G. Huttoni 14, 22 — var. magnifolia 22, 23 G. integerrima 77, 78 G. integriuscula 19 Jaccardi 21 . laramiensis 32 lepida 24, 35, 46 lobata 23 martensis 77, 78 minuta 77 multinervis 29 Obrutschewi 13 parvifolia 23 pluripartita 24, 27, 28 . polaris 21, 26 — var. pygmaea 22 G. primordialis 29 G. pusilla 10, 32 G. Schmidtiana 24 — var. parvifolia 25 G, sibirica 22-25, 35, 50 G. Simmondsi 42 G. whitbiensis 26 Ginkgoales 1-60 Ginkgoanthus 54 Ginkgocladus 33 G. australiensis 33 Ginkgodium 60-62 alaskense 62 G. Nathorsti 13, 61, 62, 451 page 10-33 G. antarctica 13 DAADAQAAARS _ G. adiantoides 29-33 G. antarctica 12-14 G. crassipes 27 G. digitata 14-23, 26, 28, 34, 52 — var. Huttont 15, 17, 21, 22, 29 —- var. polaris 21, 22, 26 G. Geinitzi 13, 38 G. lobata 28 G. moltenensis 17 G, multinervis 17 G, Obrutschewi 26, 27 G. obovata 12, 13 G, pluripartita 17 G. sibirica 24, 25, 47, 51, 57 G. whitbiensis 26 Ginkgophyllum 11, 36, 80, 81, 86, 87 . australe 87 G. Grasserti 30, 37, 84, 88 G. kamenkianum 87 G. minus 89 Ginkgopsis 77 G. Czekanowskii 77, 78 G. minuta 77 Glossozamites 447 Glottophyllum 76 G. cuneatum 76 Gliick, H. 395 Glyptostrobites Ungeri 357 ee aeonaen 173, 191, 198, 199, G. Goepperti 198 Glyptostrobus 126, 137, 139, 146, 149, 193, 197-199, 328, 360, 363 . europaeus 328, 329, 357 . gracillima 330 groenlandicus 328, 364 heterophyllus 198, 200, 306, 329, 330 intermedius 328 tener 198 G. Ungeri 357 Gnetales 459-472 Gnetopsis 470 Gnetum 459, 462-464 G. africanum 463 G. Gnemon 463, 471 — var. scandens 470 G. scandens 460, 470 Goebel, K. 108 Goeppert, H. R. passim Goeppert, H. R. and A. Menge 309, 311, 329, 330, 342, 365 Goeppert, H. R. and G. Stengel 220 Gomes, B. A. 97 Gomphostrobus 100, 279, 280, 287-292 G. bifidus 287, 288 G. heterophyllus 287 Gothan, W. passim Grand’Eury, C. 12, 89, 93 Groom, P. 143 Groom, P. and W. Rushton 135 Gunn, M. 424 Guppy, H. B. 127, 405 AAPARAR Haidingera 390 Halle, T. G. passim Hallier, H. 468 Halochloris baruthina 380 Hapaloxylon 286, 287 H. Rochei 286 Harker, A. 221 Harpe, P. de la 306, 407 Hartz, N. 32, 412 Hausmannia 26 Heer, O. passim Henslow, G. 389 Hirase, S. 1, 6 Holden, R. passim Hollick, A. 246, 250, 310, 333, 339, 452 Hollick, A. and E. C. Jeffrey passim Hooker, J. D. 464, 465 Tsoetes 64, 65 I. pumilus 379 Isoetites pumilus 440 Txostrobus Siemiradzkii 58 536 Jack, R. L. and R. Etheridge 100 Jeanpaulia 33, 65, 103 J. Lindleyana 103 J. Muensteriana 44 J. palmata 42 Jeffrey, E. C. passim Jeffrey, E. C. and M. A. Chrysler, 220, 227 Jeffrey, E. C. and R. E. Torrey, 4 Johnson, T. 3, 197 Johnston, R. H. 87, 88 Juniperites 310 J. eocenica 311 Juniperus 134, 136, 140, 148, 310, 311, 337 . chinensis 164 . drupacea 149 . gracilis 310 . Hartmannianus 311 . hypnoides 310 macilenta 250, 310, 339 . oxycedrus 149 . tertiarius 310 . virginiana 137 BUN Kaidacarpon ooliticum 256 Keteleeria 110, 122, 156, 370, 380 K. Davidiana 370 K. Loehri 394 Kidston, R. 99, 281, 282 Kiltorkensia 88 K. devonica 88 Kirchner, O. and C. Schroeter 16] Kirsch, 8. 131 Knowlton, F. H. passim Koetlitz, R. 21, 195, 397 Krammera mirabilis 248 Krasser, F. 25, 47, 48, 70, 74, 250, 338 Kraus, G. passim Krausel, R. 9 Krystofovié, A. 22, 58, 380, 441 Kurtz, F. 75, 90, 92 Kutorga, 8. 79 Lagenostoma 6 Lagerstroemia macrocarpa 337 Lange, T. 351 Laricites 372 Larix, 108-140 (passim), 155, 156, 214, 219, 225, 227, 232, 370, 373, 377, 379, 403 L. americana 132 DL. europaea 157 L. Griffitht 157 L. leptolepis 134 Laurent, L. 260, 308, 329, 353 Lawson, A. A. 123 Lebour, G. A. 81 Leckenby, J. 431 Lepidopteris 53 INDEX L. Ottonis 53 Leptostrobus 289, 294-296, 402 L. angustifolia 295 L. crassipes 295 L. longifolius 404 L. rigida 295 Leslie, T. N. 84 Lesquereux, L. 70, 89, 99, 324, 330, 357, 360, 415 Leuthardt, F. 39, 53, 291, 292 Lhwyd, E. 167 Libocedrites 308 Libocedrus 122, 126, 147, 150, 303-311, 333 . adpressa 308 . cretacea 307 . decurrens 136, 147 . Doniana 148 . macrolepis 129 . Sabiniana 308 Lignier, O. passim Lignier, O. and A. Tison 468 Lilia lanceolata 455 Lima, W. de 97 Lindley, J. 158, 276, 307, 387 Lindley and Hutton passim Lindsaya reniformis 4 Lingelsheim, A. 149, 202, 331 Lithoxylon 167 Ludwig, R. 342 Lycopodiolithes piniformis 280 Lycopodites 273 L. uncifolius 271 L. Williamsonis 271 Lycopodium arboreum 163 L. tetragonum 164 Lyginopteris 166 Lyon, H. L. 7 eiuieisiuis! McClintock, Sir L. 31, 229 McCoy, Sir F. 74 Macrozamia heteromera 94 Macculoch, J. 221 Manoxylic (wood) 128 Mantell, G. 383, 387 Marion, A. F. 260, 287 Masculostrobus 424426 M, Zeilleri 424426 Massalongo, A. 268, 407 Masters, M. T. 124, 342 Menge, A. 230 Mercklin, C. E. von 201, 218 Mesembrioxylon 138, 173, 199, 203- 212 : M. antarcticum 210 AL. aparenchymatosum 209 M. bedfordense 207, 208 M. Gothani 207 M. Hookeri 211 M. Milleri 210 INDEX M. Schwendae 207, 209 M. woburnense 207 Mesoxylon 7, 280 Metacedroxylon 175, 237, 238 M. araucarioides 238 Microcachrys 110, 116, 127, 158, 204, 398, 443 MM. tetragona 116 Miller, H. 215, 221, 252, 326, 412 Miquel, F. A. W. 336, 389 Miyake, K. 123 Mogan, L. 393 Moller, H. 42, 47, 74, 378, 379 Moller, H. and T. G. Halle 426 Moreauia 274 Moriconia 332, 333, 436 M. cyclotoron 333 Mougeot, A. 279 Miiller, C. 135 Muscites Sternbergianus 364 Nageia 457 Nageiopsis 417, 452, 453, 456-458 N. anglica 457 NV. decrescens 457 NV. heterophylla 456 N. longifolia 60, 457, 458 N. microphylla 457 N. obtusa 456 N. zamioides 458 Naiadea ovata 455 Nathorst, A. G. passim Nephropsis 77, 78 N. integerrima 77, 78 Newberry, J. 8. passim Newton, E. T. and J. J. H. Teall 22, 71, 195, 397 Nicol, W. 7, 129, 133, 168, 221 Nilssonia 51 Noeggerathia 19 N. cuneifolia 79 N. expansa 79 N. flabellata 79 N. gilboensis 89 N. obtusa 89 Noeggerathiopsis Coepneili 74 Obrutschew, V. 63 Odontopus sexpunctulatus 465 Oldham, T. and J. Morris 431, 434 Oliver, D. 409 Ophioglossum granulatum 395 Otozamites latifolia 104 Pachyphyllum 271 P. heterophyllum 319, 320 Pagiophyllites 177, 274 P. keuperianus 177, 274 Pagiophyllum 181, ‘265, 270-277, 291, 295-298, 321, 359, 429 537 P. Foettleri 275 P. peregrinum 274-277, 321 P. Rotzoanum 277 P. Sandbergi 275 P. Williamsonis 271 Palaeocladus 417 P. cuneiformis 417 Palaeocyparis 304, 318 P. elegans 304 Palaeolepis bicornuta 442 Palaeophyllales 11 Palamophyllum 80 Palibin, J. 354, 357 Palissya 315, 418, 426-428, 431, 435 P. aperta 315 P. australis 431 P. Braunii 271, 426, 427, 432, 434 P. longifolia 427 P. Sphenolepis 427, 428 Pandanocarpum 256 Paracedroxylon 173, 212, 213 P. scituatense 212 Paracupressinoxylon 189 P. cedroides 189, 190 P. cupressoides 189, 190 Paraphyllocladoxylon 173, 205 P. araucarioides 205 P. eboracense 205 Patzea 470 Pearson, H. H. W. 467, 468 Pecopteris kudlistensis 333 Penhallow, D. P. 32, 134, 170, 223, 347, 354, 357 Petrophila diversifolic. 156 Peuce 221, 244 Pherosphaera 127, 160, 204 Phillips, J. 14, 49, 51, 65, 271 Phoenicopsis 35, 40,.57, 62, 68-75, 373, 452-455 . angustifolia 74 . elongatus 74, 75 Gunni 72, 74 . latior 74 . media 74 . Potoniet 74 . speciosa 73 . taschkessiensis 74 Phyllanthus 415 Phyllocladineae 124, 413-417 Phyllocladites 413, 414 P. asplenioides 414 P. rotundifolia 442 Phyllocladopsis 417 P. heterophylla 416, 417 Phyllocladoxylon 173, 203-207, 210, 211, 242 Phyllocladus 121, 124, 127, 134, 138, 207, 210, 414-417, 436-442 P. alpina 107 P. asplenioides 414 hyytetytytytyy 538 P. hypophylla 107 P. subintegrifolius 415 P. trichomanoides 107 Phyllostrobus 311 Physematopitys 9 P. excellens 10 P. Goepperti 10 P. salisburioides 9 P. succinea 9 Picea 113, 122, 125, 129, 139, 155, 156, 214, 219, 227, 232, 270, 370, 373, 383, 385, 386, 393 P. Browniana 440 P. Engelmanni 400 P. excelsa 141, 157, 395-398 — var. alpestris 395 P. Menziesti 400 P. omorica 140 Piceoxylon 174, 219, 232 P. Gothani 229 P. Pseudotsugae 229 © Pilger, R. 124 Pilularia 64 Pinites 101, 219, 220, 370, 371, 373, 382 . antecedens 372 Carrutherst 383 cavernosus 231 Conwentzianus 220 cylindroides 383 depressus 382 Dunkeri 383 eiggensis 221 Goeppertianus 22] hexagonus 388 jurassicus 218 jurensis 218 latiporosus 239 oblongus 387 pauciporosus 240 permiensis 101, 372 pottoniensis 383 Pritchardti 189, 305 strobiformis 382 . succinifer 230 . valdensis 383 Pinostrobus 387 Pinoxylon 220 P. dacotense 232 Pinus 108, 110, 123-140 (passim), 153, 210, 223, 228, 232, 382, 403 Abies 393 . australis 396 Bailyi 393 Balfouriana 393 canariensis 139 Cembra 113, 155, 156 . Corneti 385 . Coultert 113, 155 . Crameri 365, 366 bo Aya AU AS Pe Phy Po he She to he Poh ote BND INDEX P. excelsa 154, 155, 378, 382, 385, 389, 392, 404 P. excelsa var. tuberculata 156 P. Gerardiana 394 P. halepensis 393 P. Jeffreyi 155 P. Lambertiana 108, 385 P. Laricio 116, 383 — var. nigricans 151 . longissima 385 . Lundgrent 396 Maakiana 397 MacClurii 229 Merkusii 133, 197, 214 monophylla 118, 119, 155, 375, 376 monticola 225 Nathorsti 228 Parrya 230 Peterseni 381 Peuce 155 Pinaster 383, 387-393 ponderosa 155 Quenstedti 381, 388 robustifolia 393 . silvestris 113, 137, 153, 394, 398 Strobus 140, 155, 173, 387, 393 . succinifer 9 . Timlert 394, 397 triphylla 226 Pinus (Abies) MacClurii 393 P. (Cedrus) Lopalini 370 P. (Larix) Johnseni 215 Pinuaylon 174, 219, 220, 232 Pityanthus 371, 395, 396 P. Dunkeri 383, 384 P. granulatus 395 Pityites 370-376 P. Solmsi 295, 373-381, 402 P. (Pinites) eirensis 374-376 Pityocladus 371, 377-380 P. kobukensis 379, 380 P. longifolius 378 P. Nathorsti 377 P. Schenki 379 Pityolepis 371 Pityophyllum 366, 371, 379-381, 396, 430 P. Lindstrémi 381 P. longifolium 378, 379 P. Nordenskidldi 381 P. Staratschini 381 Pityospermum 371, 396-398 P. Lundgreni 396, 397 P. Nanseni 397, 398 P. Nilssoni 397 Pityosporites 371, 398, 399 P. antarcticus 398 Pityostrobus 371, 381-395 P. Andraei 373, 388, 389 P. Benstedti 386 TUNED UTD TUTTO NT INDEX 539 P. Carruthersi 373 P. elatus 158 P. dejectus 382 P. falcata 409 P. hexagonus 388° P. formosensis 158, 159 P. Kotschyeana 393 P, imbricata 116 P. Leckenbyi 385-388 P. incerta 408 P. MacClurii 393 P. latifolia 109 P. megalopis 393 P. Lindleyana 407 P. oblongus 387, 388 P. macrophylla 158, 407 P. prae-montana 393 P. Makoyi 122 P. Sauvagei 383 P. Motleyi 144 P. strobiformis 382 P. Nageva 129 P. transsylvanica 393 P, Nagi 122, 158 Pityostrobus (Pinites) macrocephalus P. neriifolia 140 389-391 P. nivalis 158 P. (Pinites) palaeostrobus 393 P. Parkeri 409 P. (Pinites) sussexiensis 387 P. polystachya 133 Pityoxylon 170, 174, 219-232 P. praccupressina 409 P. Aldersoni 231 P. salicifolia 137 P. amethystensis 231 P. spicata 116, 122, 412, 413 P. anomalum 228 P. Thunbergiit 409 P. Benstedi 225, 226 P. Totara 116, 121 P. Chasense 220, 221 Podozamites 60, 70, 71, 74, 246, 447- P. eiggense 169, 221-223 456 P. fallax 231 P. angustifolium 452 P. foliosum 228 P. distans 449, 454 P. Nathorsti 225, 228 P. ellipsoideus 246 P. parryoides 239 P. Griesbachi 455, 456 P. pineoides 230 P. imbricata 164 P. protoscleropitys 227 P. Kidstoni 456 P. Pseudotsugae 229 P. lanceolatus 247, 448, 452-455 P. pulchrum 229 — var. latifolius 456 P. Ruffordi 223, 224 P. latifolius 454 P. scituatense 227 P. longifolius 454 P. scituatensiforme 227 P. marginatus 247, 452 P. Sewardi 224, 225 P. miocenicus 247, 452 P. statenense 227 P. ovalis 456 P. succinifer 230 P. Reinii 246, 455, 456 P. zezoense 228 P. Schenki 454 Planoxylon 176, 244 P. stonesfieldensis 455 P. Hectori 244 P. striatus 456 P. Lindleti 244 Pomel, A. 47, 274 Platen, P. 10 Poacordaites 70 Podocarpineae 124, 126, 127, 405-413 Podocarpites 405 P. eocaenica 406, 407 Podocarpoxylon. 173, 189, 194, 198, P. Pe 199, 203, 204, 206, 209, 219 aparenchymatosum 204 Schwendae 204, 207 Podocarpus 110, 121, 126, 127, 134, TODD BTS 139, 141, 158, 164, 204, 247, 398, 405, 408 . andina 137, 210 argillaelondinensis 408 Blumei 164 . Campbelli 409 . cretacea 405 . cupressina 158, 164 . dacrydioides 162 -Potonié, H.'37, 201, 281, 283, 287, 288, 294 Potonié, H. and C. Bernard 76 Prepinus 228, 295, 400-404 P. statensis 373, 401-403 P. viticitensis 402 Presl, C. B. 433 Priestley, B. 398 Protaeophyllum 471 Protocedroxylon 175, 236-238, 360 P. araucarioides 236 P. Paronai 238 P. scoticum 237, 238 Protodammara 244-251, 253, 254, 435 P. speciosa 249, 250, 323 Protophyllocladus 414 P. lobatus 415 P. polymorpha 415 P. subintegrifolius 415, 416 540 Protopiceoxylon 9, 176, 231-234 P. arcticum 232, 233 : P. Edwardsi 233, 234 P. exstinctum 231-234 Pseudo-araucaria 261 P. Lamberti 261 P. Loppinetti 261 Pseudoasterophyllites Vidali 426 Pseudofrenelopsis 345 Pseudogeinitzia 361, 362 P. sequoiiformis 362 Pseudolarix 108, 110, 123, 125, 129, 139, 139, 141, 156, 214, 217, 403 P. Kaempferi 157 Pseudotsuga 113, 122, 128, 125, 134, 139, 156, 174, 227, 229 » P. Douglasii 108, 157, 229 P. macrocarpa 229 Psilophyton 100 Psilotiphyllum 288 P. bifidum 287 Psilotum triquetrum 438 Psygmophyllum 11, 12, 34-37, 79-90, 177 angustilobum 89 Brownii 81, 86 . crenatum 81, 88 cuneifolium 79, 89 . Delvali 89 - . expansum 79, 89, 90 flabellatum 79-84, 87-89 Grasserti 80, 87, 88 Haydeni 86, 90 Hollandi 25, 86, 87 Kidstoni 84, 85, 89 kiltorkense 88 Kolderupi 84 majus 81, 85, 86, 88 mongolicum 80 primigenium 89 santagoulourensis 79 . Williamsoni 85 Pterozonium (Gymnogramme) forme 4 UP AY Pa Soa Pa Ay hy Pah ho Ph ho hy Phy reni- Raciborski, M. 26 Radais, M. 142 Raritania 438, 439 R. gracilis 438, 439 Ratte, F. 42 Reid, C. 394 Reid, C. and E. M. Reid 314, 355, 419. 470 Renault, B. 77, 101, 286, 372 Retinospora 147 Retinosporites 418, 427, 432, 433 R. indica 433 Rhipidopsis 11, 86, 90-92 R. densinervis 92 R. ginkgoides 90-92 INDEX R. ginkgoides var. Siissmilchi 85 R. gondwanensis 92 Rhizocupressinoxylon 201 Rhizotaxodioxylon 199 Robertson, A..124 Rogersia 471 R longifolia 471 Rufford, P. J. 223_ Salfeld, H. 35, 70, 105, 338 Salisburia 1, 12 S. adiantifolia 1 S. adiantoides 30, 31 — var. borealis 30 S. antarctica 13 S. borealis 30, 31 S. eocenica 33 ‘8. polymorpha 415 S. primigenia 12 S. Procaccini 30 S. pusilla 32 Salisbury, E. J. 470 Samaropsis 90, 92, 465 Sandberger, F. von 89 Sanderson, — 168 Sanio’s rims 8, 135, 136, 179, 187 Sapindopsis 471 Saporta, G. de passim Saporta, G. de and A. I’. Marion 329, 421 Saportaea 92, 93 S. grandifolia 93 Sarcostrobus 259 _ Sargassum sepientrionale 410 Saxegothaea 127, 158, 413 S. conspicua 119, 120, 133, 413 Saxegothopsis 413 S. fuegianus 413 Saxton, W. T. 7, 123, 124, 151, 336 Schauroth, C. von 293 Schenk, A. passim Schimper, W. P. passim Schimper, W. P. and A. Mougeot 290, 291, 300 Schizaeopsis 36 Schizodendron 282-286, 291, 292 S. Cowardi 284, 285 S. speciosum 283 Schizolepidella 442, 443 S. gracilis 442, 443 Schizolepis 377, 439-443 S. Braunii 295, 329, 440, 441 S. cylindrica 441 S. Follini 377, 379, 396, 441 ~ S. liaso-keuperianus 439 S. Moelleri 441 S. retroflera 441, 442 Schizopteris 38, 46 S. dichotoma 38 S. Giimbeli 38 INDEX S. trichomanoides 38 Schlechtendal, D. von 305 Schleiden, M. J. 292 Schleiden and Schenk 221 Schmalhausen, J. passim ~ Schroeter, C. 9, 215, 222 Schuster, J. 449, 451 Schiitze, E. 275, 291, 293 Sciadopitineae 124, 151, 365 Sciadopitys 108, 110, 115, 120, 123, 127, 137-141, 151, 204, 210, 242 S. glaucescens 365 | S. linearis 365 S. verticillata 365 Sciadopitytes 366, 367 S. Crameri 367 S. Nathorsti 366, 367 Sclerotites Salisburiae 30 Scolopendrium nigripes 4 Scott, D. H. 177 Sequoia 114, 118, 123, 126, 131, 134, 138, 140, 151, 270, 277, 321, 324, 330, 346, 349, 362 S. affinis 357 S. albertensis 347 S. ambigua 362 . brevifolia 354 . Burgessit 347 . concinna 351 . Couttsiae 314, 355 . du Noyeri 334 . fastigiata 255, 364 . gigantea 126, 131, 151-153, 199, 306, 347, 353-356 . gracillima 360 . hondoensis 349 S. magnifica 347, 348 . minor 349 . Penhallowi 347 S. problematica 347 S. Reichenbachit 273, 346, 358-361 S. sempervirens 106, 127, 130, 149, 151, 194-201, 347, 352-356, 417, 432 S. Whymperi 357 Sequoiineae 124, 126, 151, 346-365 Sequoiites 349-357, 443 S. concinna 350, 351 . Couttsiae 351, 355-357 . Gardneri 351 . giganteoides 350 Hardti 353 . Holsti 352 . Langsdorfii 352-355, 418 . problematica 349 . Tournali 353, 408 Sewardia 35, 103-105 N. latifolia 35, 104 S. longifolia 103-105 Shirley, J. 13, 42, 54, 55 Sifton, H. B. 135, 136 - R RRRRRHRR mh RR Rnnintninints 541 Sigillariostrobus bifidus 287 Sinnott, E. W. 160, 212 Smith, J. E.30 | Solenites 66 S. furcata 48, 103 S. Murrayana 64, 65 Solms-Laubach, Graf zu passim Sphaerococcites Mvensterianus 44 Sphenolepidium 277, 312, 346, 363-365 S. Kurrianum 312, 362-366 S. Sternbergianum 274, 351, 363-365 Sphenolepis 363 S. rhaetica 54 Sphenophyllum 288 Sphenopteris latifolia 14 S. longifolia 48 Sphenozamites latifolius 104 Stachyopitys annularioides 54 "|S. Preslii 34, 53, 54 Stachyotaxus 410-413, 418 S. elegans 410-413 S. septentrionalis 410, 412, 429 Stachypteris Simmondsi 54 Staub, M. 355 Stenorachis 51, 54-62 S. lepida 56-58 S. Ponceleti 55 S. scanicus 55, 56 S. Solmsi 56 Stenzel, G. 89, 288 Stephanospermum 6 Sternberg, C. von 14, 267, 308, 312 Stirling, J. 431 Stopes, M. C. passim Stopes, M. C. and K. Fujii 316, 327 Strasburger, E. 464 Strobilites 296, 299 S. Bronni 299 S. Bucklandi 258 S. laxus 296 S. microsporophorus 426 S. Millert 412, 413 Stur, D. 275, 372 Suzuki, Y. 335 Swedenborgia 294-296 Sykes, M. G. 467, 468 Szajnocha, L. 75 Taiwania 123, 126 Takeda, H. 142 Tate, R. 264 Taxineae 124, 127, 161, 417-423 Taxites 202, 352, 405, 417-419, 429 . duckit 202 . indicus 433 . Jeffreyr 421 . longifolius 378 . Olriki 418 . planus 431, 432 . scalariformis 202 ASH 542 T. Tournala 353 T. zamioides 43 Taxodioxylon 173, 199-202 T. palustre 201 Taxodites 328-331 T. Beckianus 331 T. europaeus 328, 329 T. miocenicum 330 Taxodium 126, 146, 149, 151, 197-201, 291, 328, 330 T. distichum 119, 137, 201, 202, 330, 331 — var. imbricataria 329 — var. miocenicum 330 — var. pliocenicum 331 T. dubium 330, 357 T.. eocaenicum 329 T. mucronatum 108, 119 T. ramosum 339 Taxoxylon 173, 202, 203, 418, 419 T. anglicum 203 T. ginkgoides 203 T. scalariforme 202 T. stephanense 203 Taxoxylum 202 T. electrochyton 230 T. Goepperti 202 Taxus 11), 121, 124, 127, 129, 133, 141, 161, 202, 417-419 T. baccata 419 T. cuspidata 161 T. margaritifera 419 T. nitens 419 _ Tenison Woods, J. E. 325 Tetraclinis 126, 129, 336, 342 T. articulata 151, 154 Thamnea depressa 151 Thinnfeldia 414-416, 436 T. Lesquereuxiana 415 Thiselton-Dyer, Sir W. T. 264, 312 Thoday, M. G. 469 Thoday, M. G. and E. M. Berridge 460, 461 Thomas, H. H. 23, 36, 47, 49, 60, 66, 206, 451 Thompson, W. P. 195, 327 Thomson, R. B. 133, 135, 140, 155, 178, 182, 183, 403, 404 Thuites 303-305, 309, 311 . callitrina 340 . crassus 325 . Ehrenswaerdi 310 . expansus 309, 311, 316-320 . Hoheneggeri 343 . meriana 310 . valdensis 309 Thujopsis 149, 308, 309 T. europaea 309 Thuya 126, 139, 140, 147, 148, 302- 318, 337 fe R= Bm be a= Bae | INDEX T. europaea 309 T. gigantea 137 T. Mengeana 310 T. occidentalis 164, 309 — var. succinea 305 — var. thuringica 305 Thylloxylon 176, 190, 243 T. irregulare 243 Tison, A. 158 Tmesipteris 440, 442 Torellia 68 Torreya 109, 110, 120, 121, 128, 140, 141, 161, 169, 202, 419-423, 436 T. bilinica 421 T. borealis 421 fT. californica 116, 120, 128, 161 T. Dicksoniana 420 T. falcata 421 T. myristica 141 T. nucifera 121, 137, 140, 161, 164 — var. brevifolia 421 T. nucifera fossilis 421 T. parvifolia 421 T. taxifolia 128 T. venusta 421 T. virginica 421 Torreyites 418-420 T. carolianus 420 Trichomanes reniforme 4 Trichopitys 48, 65, 97, 101-103 T.. heterophylla 101-103 T. laciniata 103 T. Milleryensis 100, 102 Trioolepis 424 T. Lecleret 424 Trochodendron 162 Tsuga 125, 130, 140, 150, 156, 158, 213, 214, 217 Tumion carolianum 420 Tupper, W. W. 3 Tuzson, J. 23, 177, 181, 274, 393 Tylodendron 282 Ullmannia 262, 277, 279, 289, 290, 296-299 U. biarmica 297 U. Bronni 277, 282, 296-298 U. frumentaria 262, 296-300 U. selaginoides 298, 299 Ullmannites 177, 297 Unger, F. passim Vectia 170 Velenovsky, J. passim Veronica 151 V. cupressoides 163 V. Hectori 163 Vesquia 422 V. tournalit 422 Vierhapper, F, 117 INDEX Volizia 181, 262, 282-295, 301, 438, 440 ; . brevifolia 290 . coburgensis 293, 296 . Poettleri 275 . heterophylla 275, 290, 296 keuperiana 289, 293, 295 Liebeana 289-294 . Phillips: 297 . recubariensis 295 rigida 290 . walchiaeformis 292, 293 Voltziopsis 294, 296 SNNNNNNNNS Walchia 177, 181, 262, 277-282, 286— 296, 337 W. fertilig 277, 281 W. filiciformis 281 W. foliosa 282 W. frondosa 277, 282 W. hypnoides 281 W. imbricata 281, 282 W. linearifolia 281 W. piniformis 278-282 W. Schneiderit 282 W. valdajolensis 280 Ward, L. F. 24, 32 Weiss, C. E. 282, 283 Wellingtonia 157 Welwitschia 459-468 W. mirabilis 466 Wettstein, R. V. 140 Wherry, E. T. 181, 218, 427 White, D. 97, 102, 250 Widdringtonia 109, 120, 123-126, 151, 153, 336-338, 342 W. juniperoides 129 W. Reichii 339 Widdringtonites 336-338, 363, 364, 443 W. creyensis 338 W. denticulatus 338 W. fascicularis 339 W. gracilis 338, 443 W. helveticus 342 W. keuperianus 293, 337, 338 543 W. legitimus 342 W. liassinus 338 W. subtilis 338 Wieland, G. R. 260, 267 Williamson, W. C. 387 Winkler, C. 135 Withamia 103 W. armata 104 Woodworthia 3, 168, 175, 234, 235 W. arizonica 234, 235 Worsdell, W. C. 133 Xanthoxylon 156 Xenoxylon 168, 175, 178, 189, 204, 205, 238-242 X. condylianum 239 X. latiporosum 186, 239, 240, 241 X. phyllocladoides 239-242 Yasui, K. 349 Yezonia 327 Y. vulgaris $16, 327 Yezostrobus 327, 328 Yokoyama, M. 24, 40, 61 Young, M. S. 161 Zalessky, M. 58, 76, 77, 80, 177 Zamia 7, 445, 450 Z. media 445 Z. ovata 389 Z. sussexiensis 387 Zamiostrobus 389 Z. Henslowi 390 Z. pippingfordensis 259 Z. scanicus 54, 55 Zamites 447, 452 Z. Buchianus 457 Z. distans 447 Z. lanceolata 454 Zeiller, R. passim Zeugophyllites 74 Z. elongatus 75 Zigno, A. de 65, 256 Zonarites 37 CAMBRIDGE: PRINTED BY J. B. PEACE, M.A., AT THE UNIVERSITY PRESS f ; ieee ih ite vite