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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
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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
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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
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‘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
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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
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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.
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(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.
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characters and affinities of Palaeozoic Gymnosperms. Canadian
Rec. Sci. vol. iv. p. 1,
—— (93) On new species of Cretaceous plants from Vancouver Island.
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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
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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.
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Eichler, A. W. (81) Uber die weiblichen Bliithen der Coniferen. Monats-
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—— (82) Uber Bildungabweichungen bei Fichtenzapfen. Sitzber. K.
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31—2
484 LIST OF WORKS
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Endlicher, §. (40) Genera plantarum secundum ordines naturales
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—“... (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.
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Engelhardt, H. and F. Kinkelin. (08) Oberpliocene Flora und Fauna
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Engler, A. (89) Engler and Prantl; Die Natiirlichen Pflanzenfamilien,
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— (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.
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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
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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.
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—— (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
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Gordon, Marjorie. (12) Ray-tracheids in Sequoia sempervirens. New
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Carboniferous rocks of Pettycur (Fife). Proc. Camb. Phil. Soc.
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—— (12) On Rhetinangiwm Arberi, a new genus of Cycadofilices from
the Calciferous sandstone series. Trans. R. Soc. Edinb. vol. xuvit. ~
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Gothan, (05) Zur Anatomie lebender und fossiler Gymnospermer-
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—— (06) Die fossilen Coniferenhélzer von Senftenberg. Abh. K.
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—— (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
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(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.
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—— (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. ,
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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.
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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. ;
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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.
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—— (12) On the occurrence of Dictyozamites in South America.
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—— (18) Some Mesozoic plant-bearing deposits in Patagonia and
Tierra del Fuego and their Floras. K. Svensk. Vetenskapsakad.
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—— (13?) The Mesozoic Flora of Graham Land. Wiss. Ergeb. Schwed.
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— See Moller, H. J. and T. G. Halle (13).
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Hallier, H. (05) Provisional scheme of the Natural (Phylogenetic)
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Harker, A. (06) The Geological structure of the Sgurr of Figg. Quart.
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—— (08) The Geology of the small Isles of Inverness-shire. Mem.
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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
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Hartz, N. (96) Planteforsteninger fra Cap Stewart i @stgrgnland.
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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,
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vol. ci. p. 1039.
—— (68) i. Die in Nordgronland, auf der Melville- Insel, im Banks-
land, an Mackenzie, im Island und in Spitzbergen entdeckten
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— (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.
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—— (71*) Beitrage zur Kreide-Flora. II. Kreide tiers van (ueitiniore,
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—— (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
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Hick, T. (95) On Kalorylon Hookeri Will. and Lyginodendron oldhamium
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Hilderbrand, F. (61) Die Verbreitung der Coniferen. Rhein. und
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Héhlke, F. (02) Ueber die Harzbehalter und die Harzbildung bei den
Polypodiaceen und einigen Phanerogamen. Beiheft Bot. Cent.
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Holden, H.S. (10) Note on a wounded Myeloxylon. New Phyt. vol. 1x.
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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.
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ae (14) Contributions to the anatomy of Mesozoic Conifers. II.
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p. 168.
—— (14?) On the relation between Cycadites and Pseudocycas. New
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—— (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.
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on Long Island. Bull. N. Y. Bot. Gard. vol. vitt. No. 28, p. 154.
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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.
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Hooker, J. D. and E. W. Binney. (55). On the structure of certain
limestone nodules enclosed in seams of bituminous coal, with a
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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.
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Hutchinson, A. H. (14) The male gametophyte of Abies. Bot. Gaz.
vol. tv. p. 148.
— (15) On the male gametophyte of Picea canadensis. Ibid.
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— (04) A fossil Sequoia from the Sierra Nevada. Bot. Gaz. vol.
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— (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.
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—— (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
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494 ; LIST OF WORKS
Jeffrey, E. 0. (14) Spore-conditions in hybrids and the mutation hypo-
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—— (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-
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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-
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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.
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— (05) The geology of the Perry basin in South-eastern Maine, with
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Schwefelgruben Siciliens. Abdh. Naturf. Ges. Halle, Bd. xvi. p. 79.
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Kriusel, R. (13) Beitrage zur Kenntniss der Holzer aus der Schlesischen
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Krystofovié, A. (10) Jurassic Plants from Ussuriland. Mem. Com.
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Ostrau-Karwiner Kohlenbecken. Sitzber. K. Akad. Wiss. Wien,
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— (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.
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Kutorga,S. (42) Beitrag zur Palaeontologie Russlands. Verhand. Russ.-
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Lang, W. H. (97) Studies in the Development and Morphology of
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—— (00) Studies in the Development and Morphology of Cycadean
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—— (09) The gametophytes and embryo of Pseudotsuga Douglasii.
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— (04) Notes complémentaires sur la structure du Bennettites
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—— (07) Sur un moule litigieux de Williamsonia gigas (Ta. and H.)
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(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
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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
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Lignier, O. (13%) Différenciation des tissus dans le Bourgeon végétatif
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Lignier, 0. et A. Tison. (11) Les Gnétales sont des Angiospermes
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—— (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-
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Lindley, J. See Murchison and Lyell (29). — _
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— (80) Nagra anmarkningar om Williamsonia, Carruthers. Ibid. No. 9.
—— (81) Berittelse, afgifven till Kongl. Vetenskaps-Akad. Ofver.
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—— (86) Om Floran i Skanes kolférande Bildningar. I. Floran
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—— (89) Sur la présence du Genre Dictyozamites, Old. dans les Couches
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—— (93) Beitrige zur Geologie und Palaeontologie der Republik
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—— (97) Zur Mesozoischen Flora Spitzbergens. K. Svensk. Veten-
skapsakad. Hand. Bd. xxx. No. 1, p. 5.
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502 LIST OF WORKS
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—— (06) Om nagra Ginkgovaxter fran Kolgrufvorna vid Stabbarp
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—— (07%) Paldobotanische Mitteilungen. 2. Die Kutikula der
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— (07) Jbid. I. Pseudocycas, eine neue Cycadophytengattung aus
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—— (08) Uber die Untersuchungen kutinisierter fossiler Pflanzen-
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Bd. xiut. No. 6, p. 3. i
—— (08%) Paldobot. Mitt. 7. Uber Palissya, Stachyotaxus and Palaeo-
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—— (09) Paldobot. Mitt. 8. Uber Williamsonia, Wielandia, Cycado-
cephalus und Welirichia. Ibid. Bd. xuv. No. 4.
—— (092) Uber die Gattung Nilssonia Brongn. mit besonderer Beriick-
sichtung Schwedischen Arten. K. Svensk. Vetenskapsakad. Hand.
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—— (10) Excursion C3. Dépéts fossiliféres (plantes) quaternaires de
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— (11) Palaobot. Mitt. 9. Neue Beitrage zur Kenntniss der William-
sonia-Bliiten. A. Svensk. Vetenskapsakad. Hand. Bd. xtvt. No. 4.
— (11°) Bemerkungen iiber Weltrichia Fr. Braun. Arkiv Bot.
(K. Svensk. Vetenskapsakad. Stockholm), Bd. u. No. 7, p. 1.
-——— (11%) Fossil floras of the Arctic Regions as evidence of geological
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— (1l*) Paldobot. Mitt. 10. Uber die Gattung Cycadocarpidium
Nath. nebst einigen Bemerkungen iiber Podozamites. K. Svensk.
Vetenskapsakad. Hand. Bd. xuvt. No. 8.
—— (12) Die Mikrosporophylle von Wielandiella. Arkiv Bot. Stock-
holm, Bd. xu. No. 6, p. 1.
—— (12°) Paldobot. Mitt. 11. Zur Kenntniss der Cycadocephalus
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— (13) How are the names Williamsonia and Wielandiella to be
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—— (73) Report of the Geological Survey of Ohio, vol. 1.
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Newton, E. T. and J.J. H. Teall. (97) Notes on a collection of rocks
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—— (98) Additional notes on rocks and fossils from Franz Josef Land.
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_» Verholzung der Nadeln einiger Coniferen. Pringsheim’s Jahrb.
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506 LIST OF WORKS
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—— (68) Ibid. tome 1x. p. 5.
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508 LIST OF WORKS
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—— (71) Die Fossilen Pflanzen der Wernsdorfer Schichten in der
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—— (82) Ueber Medullosa elegans. Engler’s Jahrb. Bd. ut. p. 156.
—— (82?) Die von dem Gebriidern Schagintweit in Indien gesammelten
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Schenk, A. (89) Ueber Medullosa Cotta. Abh. K. Stichs. Ges. Wiss.
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Ablagerungen im Osten des Europaischen Russlands. Mém.
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510 LIST OF WORKS
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—— (ll) The Evolution of Plants. London.
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Scott, D. H. and E.C. Jeffrey. (14) On Fossil Plants showing structure,
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Scott, D. H. and A. J. Maslen. (07) The structure of the Palaeozoic
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Sellards, E.H. (03) Codonotheca, a new type of spore-bearing organ from
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— (96) A newspecies of Conifer, Pinites Ruffordi, from the English
Wealden formation. Journ. Linn. Soc. vol. xxx. p. 417.
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Camb. Phil. Soc. vol. 1x. p. 340.
—— (97°) A contribution to our knowledge of Lyginodendron. Ann.
Bot. vol. xi. p. 65.
—— (97%) On the association of Sigillaria and Glossopteris in 8. Africa.
Quart. Journ. Geol. Soc. vol. Lui. p. 315.
—— (974) On the leaves of Bennettites. Proc. Camb. Phil. Soc. vol. rx.
p. 273.
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—— (03) On the occurrence of Dictyozamites in England, with Remarks
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—— (06) Notes on Cycads. Proc. Camb. Phil. Soc. vol. xt. pt. v. p. 299.
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REFERRED TO IN THE TEXT 511
Seward, A. C. (07?) Jurassic plants from Caucasia and Turkestan. Mém.
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—— (10) Article ‘““Gymnosperms,” Encyclop. Brit. edit. 11, vol. xm.
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—— (11) Jurassic plants from Chinese Dzungaria. Mém. com. geol.
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—— (11?) The Jurassic Flora of Sutherland. Trans. R. Soc. Edinb.
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— (11%) Links with the Past in the Plant world. Cambridge.
— (l1l*) The Jurassic Flora of Yorkshire. The Naturalist, January,
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— (12) Mesozoic plants from Afghanistan and Afghan-Turkestan.
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— (12?) A petrified Williamsonia from Scotland. Phil. Trans. R.
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—— (123) Jurassic plants from Amurland. Mém. com. geol. St Péters-
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512 LIST OF WORKS
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—— (00) Gruppe verkieselter Araucariten Stamme. Bericht Natur-
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— (07) Die Karbon- und Rotliegendfloren im Grossherzogtum Baden.
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516 LIST OF WORKS
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518 LIST OF WORKS
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—— (02) Notes on living Cycads. I. On the Zamias of Florida.
Ibid. vol. xtm. p. 331. ;
—— (06) American Fossil Cycads. Washington.
—- (08) Historic Fossil Cycads. Amer. Journ. Sci. vol. XXv. p. 93.
— (08%) Two new Araucarias from the Western Cretaceous. Geol.
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—— (09) The Williamsonias of the Mixteca Alta. Bot. Gaz. vol. XLVI.
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— (11) On the Williamsonia Tribe. Amer. Journ. Sct. vol. XXXII.
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—— (11°) A study of some American Fossil Cycads. Pt. v. Ibid.
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— (12) Ibid. pt. vi. On the smaller flower-buds of Cycadeoidea.
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—— (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
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Williamson, W. C. (40) On the Distribution of Fossil remains on the
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REFERRED TO IN THE TEXT 519
Williamson, W. C. (51) On the structure and affinities of the plants
known as Sternbergiae. Manchester Lit. Phil. Soc. [2], vol. 1x.
p. 340.
—— (69) On the structure and affinities of some exogens from the
Coal Measures. Monthly Micros. Journ. vol. 1. p. 66.
— (70) Contributions towards the history of Zamia gigas L. and H.
Trans. Linn. Soc. vol. XXVI. p. 663.
—— (72) Notice of further researches among the plants of the Coal
Measures. Proc. R. Soc. vol. xx. p. 435.
—— (72?) On the structure of the Dicotyledons of the Coal Measures.
Rep. 41st meeting (Edinburgh) of the Brit. Assoc. p. 111.
—— (74) On the Organisation of the Fossil Plants of the Coal Measures
VI. Phil. Trans. R. Soc. vol. CLxX11. p. 675.
—— (76) On some fossil seeds from the Lower Carboniferous beds of
Lancashire. 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.
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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
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