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IN the Preface to Volume III 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 help 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 

I 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. xxxi. p. 315, 1917) was published too late to be considered 
in Volume III. 
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. 


Botany School, Cambridge, 
May, 1918. 



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. C. S. 

June 18, 1919. 


GINKGOALES. Pp. 1—60. 


A. Recent 1-8 

B. Fossil 8-60 

i. Petrified wood 8-10 ; ii. Leaves 10-50, 58-60 ; GINK- 

GOITES 10-33 ; Oinkgocladua 33 ; BAIERA 33-50 ; 
iii. Flowers and Seeds 50-58 ; STENORACHIS 5 1 58; 



Pp. 61—75. 

PHYLLUM 69-75. 



93-101; TRICHOPITYS 101-103; SEWARDIA 103-105. 





Fossil Gymnospermous wood (Coniferales) .... 167-172 

Classification op Fossil Coniferous wood . . . 172-176 

I. DADOXYLON (and Araucarioxylon) 176-186; II. CU- 


203; IV. MESEMBRIOXYLON 203-212; V. PARA- 

CEDROXYLON 212, 213; VI. CEDROXYLON 213-219; 


OXYLON 231-234; IX. WOODWORTHIA 234, 235; 
242 ; XIII. ANOMALOXYLON 242, 243 ; XIV. THYL- 

CONIFERALES (continued). Pp. 245—301. 


Araucarineae 245-277 

252-256; ARAUOARITES 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 277-301 
WALCHIA 277-282; SCHIZODENDRON {Telodendron) 
282-286 ; H APALOXYLON 286, 287 ; GOMPHOSTRO- 
BUS 287, 288 ; VOLTZIA 289-295 ; SWEDENBORGI A 
295, 296; Strobilites 296; ULLMANNIA 296-299; 
ALBERTIA 300, 301. 

CONIFERALES (continued). Pp. 302—335. 

Cupressineae 302-335 

31 1-315 ; BRACHYPHYLLUM 315-322, 324-328 ; 
332, 333; CRYPTOMERITES 334, 335; CRYPTO- m 



CONIFERALES (continued). Pp. 336—367. 

Callitrineae 336-346 


Sequoiineae 346-365 

SEQUOIITES 349-351, 352-357; Conites 351,352; GEI 
NITZIA 357-361 ; EUGEINITZIA 361, 362 ; PSEUDO- 

Sciadopittneae 365-36 




CONIFER ALES (continued). Pp. 368—404. 


PHYLLUM 380, 381; PITYOSTROBUS 381-395 

CONIFERALES (continued). Pp. 405—423. 



Strobilites 412, 413; Saxegothopsis 413. 


PHYLLOCLADITES 413, 414; ProtophyllocladmAlb, 416; 
P'll'ip.ocladus 417; Phyttoc&adopsi* 417. 


422 ; CEPHALOTAXOPSIS 422. 423. 







Pp. 424—446. 

LISSYA 426-428; ELATOCLADUS 429-432, 435, 436; 
438; DACTYOLEPIS 438; RARITANIA 438, 439; 
BENSTEDTIA 445, 446. 



INCERTAE SEDIS. Pp. 447—458. 

NAGEIOPSIS 456-458. 


GNETALES. Pp. 459—472. 





FitONTisriECE. Araucaria Bidwillii. 


630, 631. Ginkgo biloba 2, 5 

632. Ginlcgoites obovata ; G. antarctica 12 

633. Ginkgoites digitata var. Huttoni. A. G. Nathorst . . . 15 

634. Ginkgoites digitata. A. G. Nathorst 16 

635. Ginkgoites adiantoides; G pluripartita; G digitata; G. sibirica; 

G. multinervis; G. moltenensis 17 

636. Ginkgoites digitata; Ginkgo biloba 18 

637. Ginkgoites digitata . . . . • 19 

638. Ginkgoites digitata. [Manchester Phil. Lit. Society.] . . 20 

639. Ginkgoites digitata. [Trustees of the British Museum.] . . 20 

640. Ginkgoites digitata var. polaris . . . .. . . 21 

641. Ginkgoites sibirica; Baiera Brauniana. [Royal Society of 

Scotland.] 25 

642. Ginkgoites Obrutschewi. [Geolog. Survey, Russia.] ... 26 

643. Ginkgoites lobata; G. crassipes. [Geolog. Survey, India.] . 28 

644. Ginkgoites adiantoides 31 

645. Baiera Raymondi ; Ginkgoites Geinitzi 38 

646. Baiera paucipartita 39 

647. Baiera spectabilis; B. longifolia 41 

648. Baiera Simmondsi. [Australian Museum, Sydney.] . • 43 

649. 650. Baiera gracilis 45 

651. Baiera gracilis forma Muensteriana. M. S 46 

652. Baiera Phillipsi . 48 

653. Baiera Lindleyana 49 

654. Antholithus sp 52 

655. Antholithus Zeilleri. A. G. Nathorst 52 

656. Stenorachis scanicus 56 

657. Stenorachis lepida. [Geolog. Surveys, India and Russia.] . 57 

658. Eretmophyllum saighanense; E. pubescens .... 59 

659. Ginkgodium Nathorsti 62 

660. Czekanowskia Murrayana. [Trustees of the British Museum.] 64 

661. Czekanowskia microphylla. [Trustees of the British Museum.] 66 

662. Destniophyllum, Solmsi 72 

663. Phoenicopsis speciosa 73 

664. Ginkgopsis Czekanowskii ; Nephropsis integerrima. M. Zalessky 78 
665,666. Psygmophyllum flabellatum 82,83 



667. Psygmopliyllum Kidstoni. T. X. Leslie 85 

668. Psygmophyllum Hoyden*. [Geolog. Survey, India. J . . 86 

669. Psygmophyllum Grasserti 87 

670. Rhipidopsis ginkgoides. M. Zalessky . . . . . 91 

671. 672. Dicranophyllum gallicum 95, 97 

673- Trichopitys heteromorpka 102 

674. Sewardia latifolia 104 

674*. Sequoia sempervirens. A. Henry 106 

675. Phyllocladus trichomanoides ; P. hypophylla; P. alpina . . 107 

676. Podocarpus latijolia 109 

677. Cryptomeria japonica 109 

678. Araucaria excelsa ; A. Rulei. [Royal Society.] . . .110 

679. Araucaria Cookii var. luxurians; A. Cookii; A. Muelleri; 

A. Montana. [Royal Society.] Ill 

680. Araucaria brasiliensis. [Royal Society.] . . . .112 

681. Araucaria Cunninghamii. [Royal Society.] . 112 

682. Agathis Moorei; A. loranthi folia. [Royal Society.] . . 113 

683. Araucaria Cookii; A. imbricaia; A. Bidivillii; A. excelsa. 

[Royal Society.] 114 

684. Microspores, microsporophylls, cones and cone-scales, and seeds 

of various Conifers 116 

685. Saxegothaea conspicua. [New Phytologist.] .... 119 

686. Pinus monophylla; Taxodium mucronatum; T. distichum . 119 

687. Saxegothaea conspicua. [New Phytologist.] .... 120 

688. Tofreya nucifera. [New Phytologist.] 121 

689. Araucaria imbricata. G. R. Wieland 125 

690. Sequoia sempervirens; Abies sp. E. C. Jeffrey . . . 130 

69 1 . Araucaria Bidwillii ; A . Cookii ; Agathis bornensis ; Larix ameri- 

cana. R. B Thomson 132 

692. Saxegothaea conspicua. [New Phytologist.] .... 133 

693. Medullary rays and tracheids of various Conifers . . .137 

694. Sections of various Coniferous leaves 141 

695. Agathis Moorei; A. australis; A. vitiensis; Podocarpus Motley i. 

[Royal Society.] 144 

696. Agathis Moorei; A. australis. [Royal Society.] . . . 144 

697. Araucaria Bidwillii. [Royal Society.] 145 

698. Araucaxia excelsa 146 

699. Cupressus Macnabiav.a. [Camb. Univ. Press.] . . . 147 

700. Libocedrus decurrens. [Camb. Univ. Press.] .... 148 

701. Alhrotaxis cupressoides 150 

702. Sequoia giganlea 152, 153 

703. Actinostrobus pyramidahs; Tetraclinis articulata. W. T. 
Saxton 154 




704. Pinus excelsa 154 

705. Pseudolarix Kaempferi; Abies bracteata; A. concolor; A 

Fraseri; Larix Grijjithi .... 

706. Branches of Abietineae. [Camb. Univ. Press.] 

707. Podocarpus formosensis. R. Dummer 

708. Dacrydium elatum; D. araucarioides 

709. Cephalotaxus Henryi ..... 

710. Drimys Winteri 

711. Veronica Hectori; Callitris calcarata; Veronica cupressoides 

Athrotaxis cupressoides; Crassida lycopodioides 

712. Section of Amethyst Mountain. [Camb. Univ. Press.] . 

713. Dadoxylon (Araucarioxylon) breveradiatum 

714. Dadoxylon (Araucarioxylon) kerguelense .... 

715. Gupressinoxylon sp. ....... 

716. Gupressinoxylon vectense. C. A. Barber 

717. Gupressinoxylon Hortii. [Trustees of the British Museum.] 

718. Gupressinoxylon poly ommatum ; G. Holdenae; G. Koettlitzi 

719. Gupressinoxylon tener 

720. Gupressinoxylon (Taxodioxylon) Taxodii; Gupressinoxylon 

(Taxodioxylon) Sequoianvm, 200 

721. Mesembrioxylonbedfordeyise. [Trustees of the British Museum.] 208 

722. Mesembrioxylon Schwendae ; Mesembrioxylon sp. . . . 209 

723. Gedroxylon transiens ; Gedroxylon blevillense . . . .215 

724. Brachyphyllum eathiense; Gedroxylon Hornei . . . .216 

725. Pityoxylon eiggense 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 Edwardsi. [Trustees of the British Museum.] 233 

730. Xenoxylon phyllocladoides ; X. latiporosum .... 240 

731. Anomaloxylon magnoradialum 242 

732. Dammarites Tolli 248 

733. Dammarites borealis. A. G. Nathorst 249 

734-736. Conites Juddi. [Royal Society of Scotland.] . . 252-254 

737. Araucarites sphaerocarpus. [Camb. Univ. Press.] . . . 257 

738. Araucarites ooliticus. [W. G. Carruthers.] 

739. Araucarites ooliticus. [Trustees of the British Museum.] 

740. Araucarites Milleri; A. Jejfreyi; A. Rogersi; A. Brodiei; 

Haberleinii; A. cutchensis; A. Phillipsi; Araucarites sp. 

741. Araucarites Slernbergii 

742. Elatides Williamsonis ' • . 

743. Elatides curvifolia 

744. Pagiophyllum peregrinum ...... 





Walchia pinijormis ........ 278 

Schizodendron speciosum 283 

Gomphostrobus bifidus 288 

Voltziaheterophylla; V. Liebeana . . . . . 290 

Swedenborgia cryptomer aides ....... 295 

XJllmannia frumentaria ; U. Bronni 297 

Cupressites taxiformis ........ 306 

Cupressinocladus salicornoides ...... 308 

Aihrotaxites Ungeri ; A. lycopodioides 313 

Brachyphyllum expansum . . . . . . .317 

755, 756. Brachyphyllum expansum [Geolog. Survey, India.] 317, 319 

757. Brachyphyllum obesum 322 

Brachyoxylon notabile; Araucariopitys americana; Protodam 
mara speciosa; Brachyphyllum crassum. E. C. Jeffrey 

Taxodites europaev.s 

Moriconia cydotoxon 

Callurites curia 

Callitrites Brongniarti; C. helvetica; C. europaeus 
Frenelopsis Hoheneggeri ; F. ram.osissima 
Sequoia magnifica. [Camb. Univ. Press.] 
Sequoiites concinna. A. G. Nathorst 
Conites oralis. [W. G. Carruthers.] 

Sequoiites Langsdorf.i 

Sequoiites Couttsiae. [Camb. Univ. Press.] 
770. Geinitzia Reichenbachii .... 



771. Sphenolepidium Kurrianum .... 

772, 773. Piiyites Solmsi 

774. Pityites (Pinites) eirensis .... 

775. Pityocladus longifolius. A. G. Nathorst 

776. Pityophyllum longijolium .... 

777. Pityocladus kobukensis [Geolog. Survey, Russia ] 

778. Pityostrobus Dunkeri 

779. Pityostrobus Leckenbyi. [W. G. Carruthers.] . 

780. Pityostrobus Benstedi. [Trustees of the British Museum 

781. Pityostrobus oblongus. [Trustees of the British Museum 

782. Pityostrobus {Pinites) Andraei 

783. 784. Pityostrobus {Pinites) macrocephalus 

785. Pinites Plutonis 

786. Pinus Timleri; Keteleeria Loehri 

787. Picea excelsa. [Camb. Univ. Press.] 

788. Pityospermum Lundgreni. A. G. Nathorst 

789. Pityospermum Nanseni; Pityospermum sp. 






Fig. pace 

791. Prepinus statensis. E. C. Jeffrey . . . . . .401 

792. Podocarpites eocaenica 406 

793. Podocarpites elegans ........ 408 

794. Sequoites Tournali ; Podocarpites incerta .... 408 

795. Podocarpites Campbelli 409 

796. Slachyolaxus elegans. A. G. Nathorst '411 

797. Slrobilites Milleri. [Royal Society of Scotland.] . . . 413 

798. Protophyllocladus subintegrifolius; Brepanolepis rotundifolia ; 

Phyllocladopsis heterophylla 416 

799. Masculostr obits Zeilleri; Masculostrobus sp. [Royal Society of 

Scotland.] 425 

800. Pallisya sphenolepis ........ 428 

801. Elatocladus heterophylla 430 

802. Elatocladus plana. [Geolog. Survey, India.] . . . .431 

803. Reiinosporites indica 433 

804. Elatocladus elegans? E. Schimperi ...... 434 

805. Elatocladus elegans . . . . . . . . 437 

806. Androvettia statensis; Geinitzia Reichenbachii. E. C. Jeffrey . 437 

807. Raritania gracilis 439 

808. Schizolepis Moelleri; S. Braunii; Schizolepis? retrofiexa . . 441 

809. Schizolepidella gracilis 443 

810. Cyparissidium gracih; C. minimum 444 

811. Podozamites lanceolatus. [Trustees of the British Museum.] . 448 

812. Cycadocarpidium Erdmanni; C. Swabii; Podozamites distans . 450 

813. Podozamites lanceolatus? [Geolog. Survey, India.] . . 453 

814. Podozamites Reinii 456 

815. Nageiopsis anglica. [Trustees of the British Museum.] . . 457 

816. Nageiopsis longifoha 458 

817. Gnetum africanum . 463 

818. Wehvitschia mirabilis. A. H. Church ..... 466 


A. Recent. 

In the account of the Coniferae contributed to Die Natiir- 
lichen Pflanzenfamilien 1 the genus Ginkgo, in accordance with the 
prevailing custom, was included in the Taxeae with Taxus, Cephalo- 
taxus, and Torreya. Eichler had previously referred Ginkgo, or 
Salisburia, to a separate family, the Salisburyeae 2 . 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 creation 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 3 . 

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 4 . 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 
has been restored to regions where it survived as late as the 
Tertiary epoch. In habit Ginkgo 5 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 Eichler (89) p. 108. 2 Seward and Gowan (00) B. p. 113. 

3 Engler (97) pp. 19, 341. 4 Elwes and Henry (06) Vol. I. p. 58. 

* For a full account of the genus, see Sprecher (07). 

s. rv 1 


Fig. 630. Leaves of Ginkgo biloba. (Slightly less than half natural size.) 


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 1 and, as Tupper 2 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 3 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, t): 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 4 . A group of large cells with brown contents 
occurs above and below each collateral endarch bundle. The 

1 Seward and Gowan (00) B. PI. ix. fig. 42. 2 Tupper (11) p. 376. 

3 Johnson (14) p. 171. * Sprecher (07) pp. 68—71; Bertrand, C. E. (74). 



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 papillae 1 
(fig. 636, C). The epidermal cell-walls are slightly undulate 2 . 
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 reniformis 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 3 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 4 . Jeffrey and Torrey 5 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. 

2 Bertrand, C. E. (74) PI. iv. figs. 9, 10. 

3 Jeffrey (14) PI. xxm. figs. 7, 8. 4 Starr (10). 
5 Jeffrey and Torrey (16). 




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 












<& & 


Gt. 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, t, transfusion-tracheids. (A, B, after Seward and Gowan; D, after 

the partially modified lamina of a foliage-leaf may bear marginal 

ovules, the lamina being continuous with the collar (fig. 631, D, c) 

it the base of the ovule. In other cases the peduncle may give 

>ff several stalked ovules, as in the specimen described by Fujii 1 

1 Fujii (96); Seward and Gowan (00) B. PL ix. figs. 1—5; Sprecher (07) p. 138. 


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 1 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 2 of short reticulate tracheids as in 
the Palaeozoic seed Stephanospermum 3 . 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 5 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 Riviere (15). Sprecher (07) figs. 120, 147, 148. 

3 See Vol. in. p. 326. 4 Shaw, F. J. F. (08). 5 Fujii (10) p. 216. 



germination of Ginkgo seeds after 45 days' immersion in sea- 
water 1 . The embryo has normally two hypogean cotyledons 
though three are not uncommon. Velenovsky 2 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 (110 /a x 80 /x) spirally 
ciliated antherozoids. After fertilisation the egg-nucleus divides, 
as in some Cycads, until 256 free nuclei are formed 3 , but in Ginkgo 
the subsequent production of walls results in a tissue, called by 
Lyon 4 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 5 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, 
s in the Palaeozoic genus Mesoxylon, which pass up the petiole. 
Annual rings are fairly distinct though, as Nicol 6 recognised, less 
vious 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 7 . 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 


Ewart (08) p. 78. 2 Velenovsky (07) p. 457, fig. 291a. 

Coulter and Chamberlain (03). » Lyon (04). ■ Saxton (10 4 ). 

Nicol (34) A. p. 147. 7 Nakamura (83) p. 25; Fujioka (13) PI. 


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 1 
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 2 . The secondary phloem consists of dis- 
continuous rows of fibres in addition to sieve-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 crvstals 3 . 

B. Fossil. 
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 4 who 
states that the ray-cells of Ginkgo are larger than those in any 
genus of Conifers. It has-been claimed by Felix 5 that Ginkgo is 

1 Jeffrey (12) p. 548. 2 Miiller (90) PL xiv. 

3 For anatomical details, see also Kleeberg (85) ; Essner (86) ; Strasburger (91 ) ; 
Seward and Gowan (00) B.; Penhallow (07); Sprecber (07); Burgerstein (08); 
Tupper(ll). 4 Essner (86). 5 Felix (94). 



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 1 who considers that the wood of Ginkgo — 
though difficult to define precisely in an analytical 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 2 proposed the generic name 
Physematopitys 3 for some Tertiary wood that he believed to possess 
the anatomical characters of Ginkgo biloba. Kraus 4 subsequently 
recognised resin-cells in the wood of Goeppert's type-species, 

hysematopitys 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 Traite de Paleontologie, but mentioned it as a synonym 
of Cupressinoxylon. Beust 5 and Barber 6 among other authors 

dopt the same course. It has more recently been stated by 
Krausel 7 that Goeppert's genus Physematopitys has the characters 
of Protopiceoxylon. Goeppert s 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 9 includes the specimen in Pinus succinifera. 

Schroeter 10 described some wood from beds on the Mackenzie 
river in North Canada, referred to the Miocene period, as Ginkgo sp. 

1 Gothan (06) p. 103. 2 Goeppert (50) p. 242, PL xlix. figs. 1—5. 

3 <t>var)fj.a, 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. 

9 Conwentz (90) A. p. 26. 10 Schroeter (80) p. 32, PL in. fig. 27—29. 


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 1 from beds, probably Eocene in age, 
the Caucasus agrees with Cupressinoxylon in the presence of rows 
of resin-parenchyma in the wood, and the depth of the rays greatb 
exceeds that in Ginkgo biloba. Penhallow 2 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 3 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 hav< 
not been found. 

ii. Leaves. 
GINKGOITES. Gen. nov. 

It has been customary to use the generic name Ginkgo boil 
for the recent species and for fossil leaves from Mesozoic anc 
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 ix. fig. 4. 

2 Penhallow (02) p. 43, Pis. xh., xm. 3 Platen (08) p. 143. 


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 Palaeophyllales 1 , 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 

1 Arber, E. A. N. (12) p. 405. 




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 primigenia 1 should pro- 
bably be referred to the genus Psygmophyllum : like many other 
supposed Palaeozoic species assigned to the Ginkgoales or to 
Ginkgo they afford no satisfactory evidence of affinity to the 
surviving genus. Other examples of leaves from Palaeozoic rocks 
described as species of Ginkgo or Salisburia on inadequate grounds 
are described in the latter part of this chapter. The Rhaetic 
leaves described by Brauns as Cyclopteris crenata and afterwards 
referred by Nathorst, with some doubt, to Ginkgo are described 
in the account of Psygmophyllum 2 .* 

Ginkgoites obovata Nathorst. 

Fig. 632 A shows the form of the specimen from the Rhaetic 
beds of Scania od which Nathorst 3 founded the species Ginkg 


Fig. 632. A. Ginkgoites obovata. B. Ginkgoites antarctica. (Nat. size; A, after 
Nathorst; B, drawn from a specimen in the University Museum, Brisbane.) 

1 Saporta and Marion (85), p. 145, fig. 74. 
3 Nathorst (86) p. 93, PL xx. fig. 5. 

2 See page 88. 


obovata : the obovate lamina is 5-6 cm. long with a maximum 
breadth of 2-5 cm. ; 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. 1 (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 2 (fig. 642, p. 26). 

Ginkgoites Geinitzi Nathorst. 

The leaf from the Rhaetic beds of Scania on which this species 
was founded was originally referred by Nathorst 3 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 Ginkgoites to Baiera. 
This species like many others from Rhaetic rocks is hardly dis- 
tinguishable from some Jurassic types. 

inkgoites antarctica Saporta. 

Under the name Salisburia antarctica Saporta 4 described a 
ingle leaf from Australia believed to be of Lower Lias age, but 
no precise information is given with regard to the locality. Shirley 5 
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 
iven off. This leaf is practically identical with some of the 
mailer, entire, examples on the short shoots of the recent type, 
he rather larger specimen figured by Shirley does not present so 
triking a similarity to these of the existing species. The lack of 
efinite information as to the provenance of the type-specimen 

1 See page 61. 2 Seward (11) p. 46, Pis. in. — vi. 

3 Nathorst (78) B. p. 26, PI. xm. fig. 17. 

4 Saporta and Marion (85) p. 142, fig. 71, A. ; Ratte (88) PI. in. fig. 1 ; Renault 
85) A. PI. ii. fig. 19. 

5 Shirley (98) PI. i. fig. 1. 


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 1829 1 as 
Sphenopteris latij'olia, but under the same name Brongniart 2 had 
a year previously recorded a Carboniferous Fern. In 1830 Bron- 
gniart 3 figured and described another Yorkshire specimen as Cyclo- 
pteris digitata : the generic name Cyclopteris was adopted by Dunker 
and other authors until Heer 4 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 Salisburia 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 5 , 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 Huttoni is retained as a form-designation for the 
more deeply lobed examples included in the species G. digitata (e.g. 

1 Phillips (29) A. PI. vn. fig. 18. See Fontaine in Ward (05) B. p. 121. 

2 Brongniart (28) A. p. 51. 3 Brongniart (28 2 ) A. p. 219, PI. lxi bis, figs. 2, 3. 
4 Heer (81 2 ); (77) i. p. 40. ' 5 Seward (00) B. p. 256. 


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 1 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 

lg. 633. Ginkgoites digitaia 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. 




There is also a further difficulty in regard to terminology: 
the employment of the two generic names Ginkgo or Ginkgoites 
and Baiera reveals a striking lack of 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 GinJcgoites, so 
also the adoption of Ginkgoites or Baiera is to a large extent the 

Fig. 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 Jhemselves afford no sure guide as to precise affinity. The 

South African, Rhaetic, specimen shown in fig. 635, L was originally 

described as Baiera moltenensis 1 but it might equally well be referred 

to Ginkgoites. 

1 Seward (08) B. PI. n. fig. 4. 




The leaves of Ginkgo digitate have a long slender petiole (fig. 634) ; 
the lamina is semiorbicuiar or obcuneate, entire, or more or less 
deeply divided into equal lobes, or irregularly divided into several 

Fig. 635. Ginkgoites. (| nat. size.) 

A. Ginkgoites adiantoides, Tertiary, Island of Mull. 

B. G. pluripartita, Wealden, North Germany (after Schenk). 

C. G. digitate, Jurassic, Japan (after Yokoyama). 

D. G. digitate var. Hutioni, Jurassic, Australia (after Stirling). 

E. G. sibirica, Jurassic, Siberia (after Heer). 

F. G. digiteta, Jurassic, Turkestan. 

G. G. multinervis, Lower Cretaceous, Greenland (after Heer). 
H. G. digitate, Jurassic, Oregon (after Fontaine). 

I. G. digitata, Jurassic, Yorkshire. 

J. G. digitata, Jurassic, Scotland (after Stopes). 

K. G. digitata, Jurassic (or Wealden), Franz Josef Land (after Nathorst). 

L. G. moltenensis, Rhaetic, South Africa. 

M. G. 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 
■5. iv 2 




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 

Fig. 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 1 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 2 . 

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. 2 Seward (11) p. 47, PI. v. fig. 62. 




Fig. 637. Ginkgoites dig; fata. 
(f nat. size.) M. 8. 

occasionally occur in association with the foliage of this and other 
Jurassic species. Male flowers 1 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 2 , 
but with the proviso that it may 
be merely a variety of G. digitata, 
a view that Nathorst 3 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 wifch the leaves of Ginkgo as the soil beneath a living 
Ginkgo tree may be in autumn 4 .' In some specimens from 
the Yorkshire coast the lamina is practically entire as in a leaf 
from Scarborough in the York Museum figured in 1900 5 . 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 6 . 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 

B 1 See page 51. 2 Heer (77) i. p. 44, PI. x. figs. 7—9. 

3 Nathorst (97) p. 15; for a discussion of the age of the Spitsbergen beds, see 
Nathorst (13 2 ). 

I 4 Nathorst (ll 3 ) p. 221. 
5 Seward and Gowan (00) B. PL x. fig. 54. 
6 Seward (00) PL n. fig. 5. 





Fig. 638. Ginkgoites digitata. (Manchester Museum; nat. size. 

Fia. 639. Ginkgoites digitata. (Cirencester Museum; nat. size.] 




acuminata 1 . A deeply divided obcuneate leaf, only 2 — 4 cm. 
broad, is figured by Dr Slopes 2 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 dig i tat a 
var. Huttoni 3 . 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 5 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 from 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 seen 
in impressions of Ginkgoites and due in 
all probability to the shrinkage of the 
mesophyll tissue into transverse bands 6 . 
This form of leaf has been described as 
Gingko polaris Nath. 7 ; it is smaller than most examples of Ginkgoites 

1 Seward (04) B. p. 98. 

2 Stopes (07) PL xxvu. fig. 4. For a map of the Sutherland Coast, see Seward 
il 2 ). 

3 Seward and Bancroft (13) p. 886. 4 Heer (7G) PL lviii. fig. 20. 
5 Heer (77) i. PL x. fig. 2. 
1 For similar transverse wrinkling, see Schenk (71) B. PL xxiv. figs. 7, 8; 

Seward (11) PL xi. fig. 74; Heer (77) ii. PL xi. fig. 1 b. 

7 Newton and Teall (97) PL xxxvin. figs. 1, 2; (98) PL xxix. fig. 3; Nathorst 
<99) PL i. fig. 8 and Thomas (11) PL iv. fig. 8. 

Fig. 640. Ginkgoites digitata 
var. polari*. Franz Josef 
Land. (Museum of the Geo- 
logical Survey, London ; 
nat. size.) M. S. 


digitata and the veins are more crowded. For this type I suggest 
the designation G. digitata var. polaris. Solms-Laubach 1 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 2 . Fig. 635, K 
shows a lobed leaf similar to that represented in fig. 640, described 
by Nathorst 3 from Franz Josef Land and compared by him with 
G. sibirica Heer and G. fabellata 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 4 , 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 5 : 
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 6 . 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 G. digitata var. Huttoni 1 ', 
and Krystofovic 8 has described G. digitata from Jurassic beds in 
Ussuriland at the northern end of the Muravjev-Amurskyj penin- 
sula. Some good specimens are recorded from Bornholm 9 . oi 
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 Huttoni have been figured from Middle Jurassic rocks 
in- Oregon 10 (fig. 635, H). Some particularly large examples are 
named by Fontaine G. Huttoni var. magnifolia but these are not 

1 Solms-Laubach (04) PI. i. fig. 10. 

2 Newton and Teall (97) PI. xli. fig. 10. 

3 Nathorst (99) p. 11, PI. i. figs. 8—19. 4 Ibid. PL t. figs. 20, 21. 
5 Hartz (96) PI. xix. fig. 1. 6 Johnstrup (83). 

7 Nathorst (07) PI. i. fig. 20. 

8 Krystofovic (10) PI. m. fig. 1. 

9 Bartholin (94) p. 96, PI. xn. figs. 1—3; (10) PI. in. figs. 9, 10. 
10 Fontaine in Ward (05) B. Pis. xxx. — xxxn., xlvi. 


specifically distinct from G. digitata. G. digitata is represented in 
Upper Jurassic or Wealden beds in Alaska (Cape Lisburne) 1 . 

Other Localities. 

From Southern Eussia Thomas 2 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 
Eichwald 3 as Cyelopteris incisa from the same district are examples 
of G. digitata. The species is recorded also from Jurassic rocks 
in Turkestan 4 (fig. 635, F), Chinese Dzungaria 5 on the west border 
of Mongolia, from the region to the east of Lake Baikal 6 , and from 
Afghanistan 7 . The incomplete specimen figured by Feistmantel 8 
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 
fghan specimen already quoted. The piece of a leaf figured by 
eistmantel as Ginkgo sp. belongs to a similar leaf, but the venation 
finer and it may be identical with Ginkgo crassipes Feist. 9 
Leaves of the G. digitata type are recorded from Jurassic beds 
in Victoria 10 (fig. 635, D) and from Japan 11 (fig. 635, C). Tuzson 

I has figured a bilobed petiolate leaf from Jurassic rocks in Hungary 
as Ginkgo parvifolia 12 ; 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 

1 Knowlton (14) PL xliv. 

2 Thomas, H. H. (11) p. 73, PI. iv. fig. 7; PI. vtii. fig. 2. 

3 Eichwald (65) PI. iv. fig. 6. 4 Seward (07 2 ) PL vn. 
5 Seward (11) PL in. fig. 40. ° Krasser (05) PL n. fig. 3. 

7 Seward (12) PL iv. fig. 51. 

8 Feistmantel (77 4 ) PL I. fig. 1. 

9 See page 28. 13 Seward (04 2 ) B. fig. 35. 
11 Yokoyama (88) B. PL xin, fig. 2. 13 Tuzson (14) PL xrv. fig. 1. 



Ginkgoites sibirica Heer. 

The specific name sibirica was given by Heer 1 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 G. 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 G. sibirica states that G. Schmidtiana 
is a smaller form of the same species 2 , but Ward 3 points out* that 
as G. Schmidtiana is described on one page and G. 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- 
goites 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 Schmidtiana and G. lepida, are fairly common 
in Jurassic rocks and occur also in Cretaceous floras; they are 
recorded from Kimeridgian beds in Scotland 4 (fig. 641, A), 
Jurassic strata in Siberia, China 5 (described by Yokoyama as 

1 Heer (77) ii. p. 61. PL vn. fig. 6 ; PL ix. fig. 5b; PL xi. ; Heer (82) ii A. p. 16 
Pis. TV., v. 

2 Fontaine in Ward (05) B. p. 125, PL xxxin. 3 Ibid. p. 126 (footnote). 

4 Seward (ll 2 ) p. 679, fig. 9, A. 

5 Yokoyama (06) B. PL vn. ; Krasaer (05) PL n. fig. 5. 




G.flabellata, also the similar leaves figured by Krasser as G. Schmidt- 
iana var. parvifolia), Turkestan 1 , and Western America 2 ; also from 
Upper Jurassic beds in Japan 3 and Franz Josef Land 4 and from 

A B 

FlG. 641. A. OinkgoiUs sibirica. B. Baiera Brauniana; from Upper Jurassic 
(Kimeridgian) bods on the coast of Sutherland. 

Lower Cretaceous beds in Canada 5 . A similar form of Ginkgoites 
is recorded from Jurassic rocks in Victoria 6 . 

It should be added that some Jurassic specimens described as 
species of Baiera, e.g. Baiera Phillipsi 7 Xath. are very near to 
G. sibirica and in such cases the choice of Baiera or Ginkgoites 
is not determined by any satisfactory standard. 


Ginkgo digiiata (Brongn.), Schmalhausen (79) A. p. 33, PI. v. fig. 4 b. 
Gingko sibirica Heer?, Ibid. p. 34, PI. iv. fig. 2 &.] 

The incomplete leaf-fragments from the Altai mountains re- 
erred by Schmalhausen to these species are too incomplete to be 
determined with any degree of certainty. The precise age of the 
beds is uncertain but, as Zeiller 8 has shown, they are probably 
Permian. There is a certain resemblance between the specimen 
referred to Ginkgo digitata and some leaves from Permo-Carboni- 
ferous strata in Kashmir described as Psygmophyllum Hollandi 9 . 

1 Seward (07 2 ) PL vn. 2 Fontaine in Ward (05) B. PL xxxin. 

3 Geyler (77) B. PL xxxi. fig. 6; Yokoyama (89) B. PL xiv. 

4 Newton and Teall (97) PL xxxviu. ; Nathorst (99). 

5 Dawson (85) PL ir. ■ Seward (04 2 ) B. p. 177. 

7 Seward (00) B. p 270. 

8 Zeiller (96) A. 9 Seward (07) p. 59, PL xra. figs. 3—6. 




Ginkgoites whitbiensis (Nathorst). 

This name was proposed by Nathorst 1 for a leaf from the 
Jurassic rocks of the Yorkshire coast in the British Museum similar 
to some of the smaller forms referred to G. digitate but characterised 
by the deltoid form of the lamina, its deep dissection into six 
more or less pointed segments, three on each side of a broad median 
V-shaped sinus, and by the small size (1-5 cm. broad and 2-5 cm. 
deep) of the lamina. It is hardly possible to decide whether this 
and similar small leaves should be regarded as varieties, e.g. G. digi- 
tata var. polaris, or assigned to a distinct species. Fontaine 2 com- 
pares some leaves figured by him from Oregon as Ginkgo sp. with 
Nathorst's species, but they are probably nearer to the examples 
described by Nathorst and others as G. polaris. A small bilobed 
leaf figured by Raciborski 3 from Rhaetic beds near Cracow as 
Ginkgo aff. whitbiensis is more likely to be a young leaf of the 
Fern Hausmannia. 

Fig. 642. Ginkgoites Obnrtscheivi. 

Ginkgoites Obrutschewi Seward. 

This species, named after Prof. Obrutschew who made a col- 
lection of plants from Jurassic rocks in Chinese Dzungaria 4 on 
the western border of Mongolia (lat. 85° N. long. 45° E.), agrees 
fairly closely with some forms of G. digitata. The lamina is deeply 

1 Nathorst (80) A. p. 74; Seward (00) B. p. 261, PI. ix. fig. 8, 

2 Fontaine in Ward (05) B. PI. xxxiv. 

3 Raciborski (91) PI. iv. fig. 13. 4 Seward (11) p. 46, Pis. m.— vn. 


bilobed and each half may be similarly divided : the segments 
are obtuse and not truncate (fig. 642, A) ; the veins are approxi- 
mately 1 mm. apart. The walls of the epidermal cells are straight 
and not sinuous ; the stomata (fig. 642, B) are practically confined 
to the lower surface and are less numerous than in Ginkgo biloba. 
The guard-cells are surrounded by a group of broadly triangular 
cells with papillose, thickly cuticularised, inner Avails, but the 
overarching papillae are rather less prominent than in G. digitala. 
It is interesting to find short secretory tracts at intervals in the 
intercostal regions of this species agreeing with those in the leaves 
of the recent species. A leaf described from Jurassic rocks in 
Amurland as Ginkgo sp., cf. G. Obrutschewi 1 agrees closely with 
the type-specimens. 

Ginkgoties crassipes (Feistmantel). 

This Upper Gondwana species (fig. 643, B) was described from 
the Madras coast as Ginkgo crassipes 2 . An examination of the 
type-specimens enables me to confirm generally the accuracy of 
Feistmantel's figures except in one point, namely the supposed 
presence of a median ridge extending from the petiole through the 
lower third of the lamina, which gives the impression of a midrib : 
this is merely a shallow groove that is clearly accidental. The 
leaves appear to be entire;' the lamina is obcuneate and passes 
into a fairly stout petiole; the veins are occasionally forked and 
approximately 0-5 mm. apart. The piece of lamina described by 
Feistmantel as Ginkgo sp. 3 may be specifically identical with this 

Ginkgoites pluripartita (Schimper). 

This Wealden species was first described by Dunker 4 as 
Cyclopteris digitata Brongn. and Ettingshausen 5 also regarded the 
German specimens as identical with the English Jurassic type. 
Schimper 6 proposed the name Baiera pluripartita because of the 
deeply divided lamina and the comparatively narrow segments : 

1 Seward (12 3 ) PI. i. fig. 9. 

2 Feistmantel (77) p. 197, figs. 6, 7; (79) p. 31, Pis. xv., xvi. 

3 Feistmantel (79) PI. xvi. fig. 12. 

4 Dunker (46) A. p. 9. PI. I. figs. 8, 10; PI. v. figs. 5, 6; PI. VI. fig. 11. 

5 Ettingshausen (52) p. 12 ? PI. iv. fig. 2. 
Schimper (69) A. p. 423. 



while substituting Ginhgoites for Baiera, it is advisable to retain 
Scliimper's specific designation on the ground that the deep division 
of the lamina appears to be the rule in the Wealden leaves whereas 
in G. digitata the leaves are usually much less deeply dissected and 

A . .... 


Fig. 643. Ginkgoites lobata (A) and Ginhgoites crassipes (B). 
(Indian Geological Survey, Calcutta.) 

have broader segments. The largest specimen is one figured by 
Ettingshausen with a lamina 7-5 cm. broad and 4-5 cm. deep ; 
there is generally a deep median sinus and each half of the lamina 
is subdivided into relatively narrow obtuse or truncate obcuneate 
segments (fig. 635, B). Schenk 1 describes the epidermal cells as 
1 Schenk (71) B. p. 212, Pis. xxiv., xxv. fig. 7. 


polygonal with straight walls ; the stomata are surrounded by 
5 — 6 accessory cells as in Ginkgo biloba and they are confined to 
the lower surface. Although there is practically no difference, 
as regards form and venation, between this Wealden species and 
some of the Jurassic leaves referred to G. digitata var. Huttoni the 
distinctive specific name is retained for the reason already men- 

Leaves of a very similar form are figured by Heer as Ginkgo 
iitultinervis 1 (fig. 635, G), from Upper Cretaceous (Cenomanian) 
beds of West Greenland, and Baiera arctica from the Kome beds 
(Urgonian) of West Greenland 2 . 

The specimen from the Atane (Cenomanian) beds of Greenland 
figured by Heer as Ginkgo primordialis 3 appears to be an entire leaf 
with a long petiole 2-5 mm. broad : the original impression in the 
Stockholm Museum, too incomplete to serve as the type of a species, 
shows a very imperfect lamina and a long axis that has probably 
no connexion with the leaf. 

Ginkgoites adiantoides (Unger). 

The Tertiary leaves on which this species was founded were in 
the first instance described as Ginkgo biloba* and, as several writers 
have pointed out, so far as regards form and venation there is no 
good reason for drawing a distinction between the fossils and the 
leaves of the recent species. In a note published in 1913 Depape 5 
definitely adopts the name Ginkgo biloba for Tertiary leaves which 
he regards as specifically identical with Unger's species. In the 
absence of any satisfactory evidence as to the nature of the repro- 
ductive organs and in view of the considerable interval that 
separates the Tertiary and recent plants, it is clearly inadvisable 
to assume specific identity. In adopting the generic designation 
Ginkgoites instead of Ginkgo I am following the custom generally 
recognised of distinguishing fossil forms by a special termination, 
though there is no implication that all species so named are 
generically distinct from the surviving type. In proposing the 

1 Heer (82) ii. B. p. 46, PL vm. figs. 2—4 ; PL ix. fig. 3 b. For an account of the 
stratigraphy and maps of these plant-beds on the coasts of the Xoursoak peninsula, 
see Johnstrup (83) and White and Schuchert (98). 

2 Heer (75) ii. B. p. 37, PL m. fig. 3. 

3 Heer (75) ii. B. p. 100, PL xxvn. fig. 1. 

4 See Gardner (86) p. 99. 5 Depape (13). 


name Salisburia adiantoides for the Miocene leaves from Sengallia 
in North Italy Unger 1 indicated their probable identity with 
Ginkgo biloba L., the generally adopted name for the existing 
species which Smith in 1797 proposed should be called Salisburia 
adiantifolia 2 . The Miocene leaves from Sengallia figured by 
Massalongo and Scarabelli 3 have an entire, irregularly crenulate 
or a more or less deeply bilobed lamina very like that of Ginkgo 
biloba: the specimen named by them S. Procaccini* should also 
be included in G. adiantoides. The resemblance to the recent 
leaves extends to the presence of short secretory tracts between 
the veins, but these were referred by Massalongo to a fungus 
which he named Sclerotites Salisburiae 5 . 

The two specimens reproduced in fig. 644 from the Eocene beds 
in the Isle of Mull illustrate the broad fan-like lamina that varies 
from 5 to 10 cm. in breadth and may be entire, unevenly lobed or 
symmetrically bilobed. The venation is identical with that of the 
living species : the characteristic marginal veins on the lower edge 
of the lamina are clearly seen in fig. 644, A. The preservation 
of these British leaves described by Mr Starkie Gardner 6 is 
exceptionally good; they occur as purple impressions in white 
clay interbedded with basaltic sheets in the cliffs of Ardtun Head 
in the Island of Mull. Though perhaps on the average these 
Eocene leaves from Mull are larger than those of the Maidenhair 
tree some examples of the latter exceed in size any of the fossils. 

Leaves identical with or very similar to the Italian and Scottish 
specimens are recorded from both Tertiary and Upper Cretaceous 
rocks in many parts of the world. Specimens collected by Dr Lyall 
from Tertiary (Miocene or Eocene) beds on Disco Island off the 
West coast of Greenland, lat. N. 70°, were described by Heer 7 as 
Salisburia borealis and he also speaks of them as Salisburia adian- 
toides var. borealis 8 . An examination of the original specimens 
in the Kew Museum and an impression in the Dublin Museum 

1 Unger (45) p. 211 ; (50) A. p. 392. 2 Smith, J E. (1797). 

3 Massalongo and Scarabelli (58) p. 163, PI. I. fig. 1; PI. vi. fig. 18; PI. vii. 
fig. 2; PI. xxxix. fig. 12. 

4 Ibid. p. 165, PI. xxxix. fig. 1. 

5 Ibid. PL i. fig. 1 a. 6 Gardner (86) p. 99, PI. xxi. ; (87) A. 

7 Heer (68) i. p. 95, PI. n. fig. 1 ; PI. xlvii. fig. 4 a, 

8 Ibid. p. 183. 




collected by^ir Leopold McClintock enables me to confirm Gard- 
ner's view as to the identity of S. borealis and S. adiantoides. 
Examples of G. adiantoides were also obtained from Atanekerdluk 1 
on the Noursoak peninsula to the north of Disco Island. Heer 

Ginkgoites adiantoides. From Eocene beds in the Island of Mull. 
(British Museum, V. 1060.) 

figures a very well preserved leaf with a long petiole from Atane- 
kerdluk which he refers to S. adiantoides. In a later account of 
this flora he refers the species to Ginkgo. Heer regarded these 

1 He< 

1 Heer (68) i. p. 57, PL lxxvii. figs. 9—12; PL xvn. fig. 14. 


Greenland beds as Miocene, but it has been suggested that the 
may belong to the Eocene period 1 

Heer 2 also described specimens of Ginkgoites adiantoides from 
beds, assigned to the Miocene period, in Sachalin Island. This 
species has been obtained from the Laramie series, a thick succession 
of brackish water strata deposited on both sides of the Rocky 
Mountains ' extending from Mexico far into British North American 
Territory' and including both Upper Cretaceous and Lower 
Tertiary strata 3 . Leaves described by Lester Ward as Ginkgo 
laramiensis* and regarded by him as intermediate between G. adian- 
toides and G. biloba are indistinguishable by any definite character 
from G. adiantoides. Ward's species is also recorded by Knowlton 5 
from the Montana formation, a series of beds formerly included in 
the Laramie group, in Wyoming. Ginkgoites adiantoides occurs 
in Upper Cretaceous beds in British Columbia and specimens 
described as Salisburia pusilla 6 by Dawson, which I believe t 
belong to this species, were found in the Upper Cretaceous o 
Vancouver Island. From beds of the same age Penhallow 7 ha 
described some wood as Ginkgo jpusilla though it is not clear o 
what grounds it is assigned to the genus Ginkgo. 

It is interesting to find leaves of this type recorded from Lower 
Pliocene beds at Saint Marcel-d'Ardeche in France 8 and from 
Upper Pliocene beds in the Lower Main valley in Germany 9 . 

It is therefore abundantly clear that trees, apparently indis- 
tinguishable as regards the form of the leaves from Ginkgo biloba, 
flourished as recently as the Pliocene period in western Europe 
and in the Eocene period grew as far north as latitude 70°. 

Records of seeds referred to Ginkgo are very meagre and add 
nothing of importance to our knowledge of Tertiary species. Some 
pyritised seeds were described by Ettingshausen and Gardner 10 

1 White, D. and Schuchert (98) p. 367. 

2 Heer (78) v. p. 21, PI. n. figs. 7—10. 

3 Ward (87) p. 15, PL xxx. figs. 5, 6; (85) PL xxxi. pp. 4—6. 

4 Ibid. PL i. fig. 4 ; PL xxxi. fig. 4. 

5 Knowlton (00) p. 31, PL iv. figs. 7—10; PL v. fig. 5. 

6 Dawson (93) p. 56, PL vi. figs. 11 — 14. Heer's species Ginkgo pusilla is 
founded on Jurassic leaves; Heer (77) ii. p. 61. 

7 Penhallow (02) p. 43, Pis. xn., xm. 8 Depape (13). 
9 Engelhardt and Kinkelin (08) p. 196, PL xxm. fig. 18. 

10 Ettingshausen (79) p. 392. 


from the London clay of Sheppey as Salisburia eocenica and 
afterwards figured by Gardner as Ginkgo 2 , eocenica 1 . The 
specimens (11x9 mm.) 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 Ginkgoiies adiantoides from 
the Upper Pliocene of the Frankfurt district 2 , but neither the 
German nor English specimens possess any interest as records of 
Ginkgoiies seeds. 

Ginkgocladus. Ettingshausen. 

An imperfect leaf -like impression described by Ettingshausen 3 
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- 
liensis i ) are of value as botanical records. 

BAIERA. Braun. 

This generic name was first used by Braun 5 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 6 examined Braun's 
Rhaetic specimens and failed to discover any indication of the 
presence of secondary veins. In 1877 Heer 7 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, PI. ix. figs. 31—34. 

2 Engelhardt and Kinkelin (08) p. 196, PI. xxin. figs. 16—18. 

3 Ettingshausen (87) p. 39, PI. vn. fig. 19. 

4 Ibid. (88) p. 103, PI. viii. 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 
linger' s genus Jeanpaulia which has since been discarded. 

I 7 Heer (77) ii. p. 51. 
s. iv .3 


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 
JStachyopitys Preslii which that author afterwards regarded as 
microstrobili of Baiera Muensteriana. Reference is made to 
these and similar fossils in the account of examples of male 
flowers 1 . Seeds have been referred to Baiera 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 2 and other species may be seed 
bearing organs of Baiera, but there is no satisfactory evidence ir 
support of this view. In all probability some of the associated 
seeds belong to Baiera, also some of the microstrobili, e.g. Leuthardt'e 
Swiss specimens described on a later page, but in the present 
state of our knowledge it is preferable to regard these specimei 
as reproductive organs that cannot be assigned with certainty 
any particular species of Baiera or Ginkgoites. 

Leaves assigned to Baiera 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 Baiera 
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. 

2 Fontaine (89) B. pp. 265 etc., Pis. 134 etc. ; Berry (11) p. 372. 



more suitable name for certain forms of leaf is not governed by 
any definite criterion: specimens described as Ginkgo sibirica, 
G. lepida, G. concinna 1 , etc. are indistinguishable from leaves 
referred by authors to Baiera. Similarly such a species as 
B. Lindleyana 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 2 , 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 Baiera 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 as in Pinus. Some specimens described by Salfeld 3 
from the Solenhofen beds of Bavaria as Baiera 1 ! 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. 

I The specimens described by Fontaine and Berry from the 
1 Heer (77) ii. PI. xm. figs. 6—8. 
2 Nathorst (78 2 ) B. PI. xxi. 
3 Salfeld (07) B. p. 195. PI. xx. fig. 3; PL xxi. fig. 1. 
4 Page 105. 


Potomac group in Virginia as Baiera foliosa 1 , 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 Dicranophyllwn : their precise position 
cannot be definitely determined. 

The structure of the cuticles is known in a few species from 
accounts published by Sehenk 2 , Nathorst 3 and Thomas 4 ; 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 5 has drawn attention to the presence in 
B. spectabilis of traces of some secreted substance in the mesophyli 
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 Baiera-like leaves from the Potomac 
beds, originally referred by Fontaine to his genus Baieropsis and 
regarded by him as Ginkgoaceous, illustrates the possibilities of 
error in determinations founded on leaves alone. The fertile 
examples of Baieropsis have been made the type of a new genus 
Schizaeopsis 6 , other species of Fontaine's genus being transfers 
to the genus Acrostichopteris ; they differ from Baiera in thei 
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 Baiera 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 xciv. fig. 13; Berry (11) p. 372, PI. lix. 

2 Schenk (67) A. PI. vi. figs. 1, 2; PL ix. figs. 11—13. 

3 Nathorst (06). 4 Thomas (13 2 ) p. 244, fig. 5. 
5 Nathorst (06) p. 9, fig. 9. 6 Berry (11) p. 214; (ll 2 ). 




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. Baiera 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 Ginkgoale3 with leaves of the type 
that still survives. Baiera is clearly an older form than Ginkgoites ; 
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 1 , 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 2 from Permian beds 
of Mansfeld, a species which Geinitz 3 also recorded, but under the 
generic name Zonarites. Potonie 4 and other authors, following 
Heer, transferred the species to Baiera. Heer's combination, 
B. digitata, had, however, already been used by Schimper 5 for the 
Jurassic species usually called Ginkgoites digitata. 

Leaves of the form represented by B. virginiana may be closely 
allied to Saporta's Permian species Ginkgophyllum (Psygmophyllum) 
Grasserti 6 . 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 

Fontaine and White (80) B. p. 103, PI. xxxvii. figs. 11, 12. 
Brongniart (28 2 ) A. p. 69, PL ix. fig. 1. 
Geinitz (62) PL xxxi. figs. 1, 2. 

Potonie (93) A. p. 237, PL xxxn. fig. 2; PL xxxni. fig. 6. 
5 Schimper (69) A. p. 423. 6 Page 87. 



Schizopteris, e.g. S.Guembeli from Upper Carboniferous and Permian 
strata 1 ,, but other Permian specimens assigned to Schizopteris , 
e.g. S. dichotoma and S. trichomanoides 2 are generically distinct 
and probably belong to Pt endosperms. 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 3 (fig. 645, A^difTers 
but little from B. virginiana ; the narrow cuneate leaf is divided 

Fig. 645. A, Baiera Raymondi. B, Ginkgoites Geinitzi. 
(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 xh. fig. 7 ; Goeppert (64) A. PI. ix. figs. 6, 7. 

2 Zeiller (06) B. PI. i. figs. 7, 8. 3 Renault (88) p. 324, fig. 48. 




portion of the lamina is 5 mm. broad and the ultimate segments 
with obtuse apices have a breadth of 2-5 mm. The venation is 
imperfectly shown in the specimens figured by Renault and Zeiller 1 . 
Baiera 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 2 and Leuthardt 3 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 4 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 

(After Nathorst ; J nat. size ) 

1 Zeiller (06) B. p. 202, PI. xlviii. figs. 1, 2. 

2 Heer (76) A. Pis. xxix., xxx.. xxxvi. 

3 Leuthardt (03) p. 7 r Pis. n.— iv. 

4 Nathorst (86) p. 94 ; Pis. xx- xxn. 


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 paucipartita is described 
by Yokoyama 1 from Rhaetic beds in Japan and an imperfect 
specimen from Rhaetic strata in New Zealand has been assigned 
by Arber 2 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 bilobed 
linear segments. 

Baiera spectabilis Nathorst. 

The leaves of this Rhaetic species from the south of Sweden 3 
reach a length of 25 cm. ; the coriaceous lamina is obcuneate and 
fan-like but narrower than the leaves of such a species as B. Sim- 
mondsi. 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 3 cm. 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) PI. n. fig. 5. 2 Arber, E. A. N. (13) PI. vii. figs. 2, 3. 

, 3 Nathorst (06). i Thomas (13) PI. xxv. figs. 3, 4. 




Fig. 647. A, Baiera spectabilis; B, stoma C, D, epidermal cells and stoma of 
Baiera longifolia. (A, B, after Nathorst A, f n at. size C, D, after Thomas.) 


resembles B. pulchella, a Jurassic species described by Heer 1 from 
East Siberia and by Bartholin 2 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 3 to B. pulchella is probably, as Nathorst 
suggests, a piece of a B. spectabilis leaf. 

Baiera Simmondsi (Shirley). 

The leaves described by Shirley 4 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 5 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 
leaves agree closely with Fontaine's Triassic species, Baiera multi- 
fida 6 , from Virginia : the plant-beds of the Richmond coalfield 
are correlated with the Lunz plant-beds in Austria 7 , 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 8 : 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. 

1 Heer (77) ii. p. 114, PI. xx. fig. 3c; PI. xxn. fig. la; PL xxvm. fig. 3. 

2 Bartholin (94) PL xi. fig. 5. 3 Moller (03) PL iv. fig. 19. 
4 Shirley (98) p. 12, PL n. 5 Ratte (87) PL xvii; (88). 

6 Fontaine (83) B. p. 87, Pis. xlv.— xlvii. 

7 Berry (12). 8 Heer (77) ii. p. 115, PL xxvm. fig. 2 




long and 23 cm. in breadth ; there are nearly 60 ultimate linear 
segments with obtuse apices and, in the smaller subdivisions, 

Fig. 648. Baiera Simmondsi. (Australian Museum, Sydney; * 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. 


I'l I I 1 1 I Ul 

c flora 

Fontaine speaks of the lamina of his species as reaching a length 
of 25 cm. ; both in the method of division and in the form of the 
segments, B. multifield agrees closely with the specimen shown in 
fig. 648. A similar form of leaf is figured by Solms-Laubach 1 
from Rhaetic beds in Chile as Baiera ? Steinmanni, but the lamina 
only is preserved. Schenk's B. taeniata 2 from the Rhaetic flon 
of Franconia is another similar type. 

Baiera stormbergensis Seward. 

The specimens described from the Stormberg series (Rhaetic) 
of South Africa 3 are portions of leaves that must have reached a 
length of 12 cm. or more and a breadth of 10 cm. 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. Schenki 4 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. 

Baiera Muensteriana (Presl). 

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 5 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. 

2 Schenk (67) A. p. 26, PL v. figs. 1—4; PL vi. figs. 1, 2. 

3 Seward (03) B. p. 64, PL viii. fig. 3. 

4 Feistmantel (89) p. 72, PL m. 5 Schenk (67) A. p. 39, PL ix. 




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 1 and several European localities, 
but it is impossible to determine its geographical range 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 

Fig. 649. Baiera gracilis. Type- 
specimen of Bunbury (Bun- 
bury Collection. Botany 
School, Cambridge). 

Fig. 650. Baiera gracilis. (British Museum, 39208.) 

imperfect leaf with a fan-shaped lamina deeply divided into forked 
linear segments 2 : a better example is reproduced in fig. 650. Bean 

1 Schenk (87) B. PI. vm. fig. 44; Zeiller (05) p. 194. 

2 Bunbury (51) A. p. 182, PL xn. fig. 3. 




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 1 Heer 
from the Siberian Jurassic flora is another very similar form; 
also Baiera incurvata Heer 2 from the Lower Cretaceous of Green- 
land, B. angnstiloba Heer 3 , 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 5 , also leaves 
recently referred by Halle 6 to B. australis from the Lower Cretaceous 
plant-beds of Patagonia. Baiera gracilis is recorded from Upper 
Jurassic (or Lower Cretaceous) beds in Alaska 7 , but leaves of this 

Fig. 651. Baiera gracilis forma Mucnsteriana. 
(British Museum, f nat. size.) M. S. 

type are rare in the Jurassic strata of North America. The 
Rhaetic species B. Muensleriana 8 (Presl) described by Schenk from 
Franconia and by other authors is a closely allied type which 

1 Heer (77) ii. PI. xra. figs. 6—8. 2 Ibd. (82) B. PL xm. fig. 6. 

3 Ibd. (78) ii. PL vti. fig. 2; Krasser (05) PL n. fte. 10; Schenk (83) A. PL ltii. 
fig. 1. 

4 Tenison-Woods (83) A„P1. iv. fig. 3. 

5 Seward (04 2 ) B. figs. 36—38. e Ha lle (13) Pis. iv., v. 
7 Fontaine in Ward (05) B. PL xliv. fig. 2. 8 Schenk (67) A. PL ix. 


cannot always be distinguished from B. gracilis. The example 
shown in fig. 651 from the Yorkshire coast has been named B. gra- 
cilis forma Muensleriana to denote its close resemblance to the 
Rhaetic species 1 . B. Guilhaumali 2 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 Ginkgoites sibirica. 

Baiera longifolia (Pomel). 

Pomel 3 described this Jurassic species as Dicropteris longifolia 
and Heer substituted the generic name Baiera i . 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 5 , recorded also by Moller from Bornholm, 
is probably not a distinct type. Thomas 6 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 18 cm. 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 7 records this species from Jurassic rocks 

1 Seward (00) B. p. 264. « Zeiller (03) B. PI. l. figs. 16—19. 

3 Pomel (49) p. 9. 

4 Heer (77) ii. p. 52, PI. vn. figs. 2, 3; PI. vm. ; PI. ix. figs. 1—11; PI. x. 
figs. 6, 7; PI. xv. fig. 116. 

» Ibid. p. 56, PI. x. figs. 1—5; PI. vn. fig. 1; (82) B. PI. m. figs. 4—8; Moller 
(03) PI. v. fig. 3. 

6 Thomas (13) p. 243, PI. xxv. figs. 3, 4. 

7 Krasser (05) p. 18, PI. I. fig. 16. 




in China but the photographic reproduction is unfortunately to 
obscure to afford any indication as to the nature of the specimen 

Baiera Phillipsi Nathorst. 

This Jurassic species 1 (fig. 652) illustrates the absence of any 
definite dividing line between Baeira and Ginkgoites; it agrees 
very closely with G. sibirica and 
with leaves assigned to G. lepida 
and other 'species.' Fig. 652 is 
drawn from Phillips' type- speci- 
men 2 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 Baiera longifolia?. 
Krasser records B. Phillipsi from 
Jurassic strata in Sardinia 4 . This 
species shades into B. gracilis and 
the very similar B. australis McCoy, 
especially resembling some leaves 
included by Halle 5 in the latter 

Baiera Lindleyana (Schimper). 

Leaves of this type were first figured by Lindley and Hutton 6 
as Solenites ? furcata and transferred by Braun to Baiera. 
Schimper 7 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 Baiera gracilis. 

MlQ. 652. Baiera Phillipsi. (York 
Museum; £ nat. size.) M.S. 

1 Nathorst (80) A. p. 76. 

3 Seward (00) B. p. 270. 

5 Halle (13) Pis iv., v. 

6 Lindley and Hutton (37) A. PI. 209. 

2 Phillips (75) A. PI. vit. fie 
4 Krasser (13) p. 5. 

7 Schimper (69) A. p. 683. 





In a former account of this species 1 1 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 
lamina are separated by a smaller 
angle and if cuticular preparations 
are available the stomata afford a 
means of differentiation : in Baiera 
the guard-cells are surrounded by 
a circular group of cells, while in 
Czekanowskia the subsidiary cells 
are longer and narrower, forming a more oblong group. 

Baiera Lindleyana is recorded also from Middle Jurassic rocks 
Chinese 3 Dzungaria and from Upper Jurassic rocks in Scotland 4 . 
Some specimens described by Fontaine 5 from the Black Hills 
Lower Cretaceous) as Czekanowskia nervosa Heer are, as Berry 6 
points out, probably leaves of a Baiera, and I am disposed to refer 
them to B. Lindleyana. 
Baiera Brauniana (Dunker). 

This species 7 , represented by leaves from Wealden and Upper 
urassic 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) PI. iv. fig. 44. 

4 Seward (ll 2 ) PI. v. fig. 105. 

5 Fontaine in Ward (99) B. p. 685, PI. 169, figs. 1, 2. 

6 Berry (11) p. 374. ' Dunker (46) A. p. 11, PL v. fig. 4. 
s. iv 4 

Fig. 653. Baiera Lindleyana. 
(British Museum; A, 39208; B, 
V. 3682.) 


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) 1 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 2 from the Wealden of North Germany. 

Baiera spetsbergensis Nathorst. 

This species, one of the smallest representatives of the genus, 
is described by Nathorst 3 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 and 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 ? eocenica. 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 biloba*. 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 (8 — 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 Ginkgoites. 

1 Seward (ll 2 ) p. 680. 

2 Schenk (71) B. p. 224, PL in. figs. 9—14. 

3 Nathorst (97) p. 53, PI. in. figs. 6—12. 

4 Heer (82) ii. A. p. 16, Pis. iv ., v; (77) ii. p. 57, PI. xi. 


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. xxxviii 1 . 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 2 , 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 microsporophvlls 

our information is scanty and indecisive. Nathorst 3 first suggested 

that some small carbonised bodies from Yorkshire Jurassic beds 

figured by Phillips 4 as 'unknown leaves' are probably fragments 

of male flowers of some species of Ginkgoites. The specimen of 

hich Phillips figured a small portion is shown in fig. 654, B; it 

onsists of a slender axis with several short and partially broken 

ateral branches bearing terminal groups of oblong bodies 4 mm. 

ong and 1 mm. wide, 2 — 4 in each group : these suggest comparison 

ith 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 

mm. long, but in the occurrence of two to four microsporangia 

n a single microsporophyll the resemblance between the fossil 

nd recent form is fairly close 5 . Unfortunately it has not been 

ossible 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 

3e 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. in. p. 502. 2 Heer (82) A. p. 21, PI. ix. 

J Nathorst (80) A. p. 75. 4 Phillips (29) A. ; (75) A. PI. vii. fig. 23. 

5 See page 5. « Seward (00) B. p. 260, fig. 45. 




the axis is 2-3 cm. long and some microsporangia are seen in their 
original position, while others are detached 1 . 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 2 as Antholithus Zeilleri present a 
certain resemblance to these supposed male flowers. One of 

Fig. 654. Antholithus sp. (Sedgwick Museum, Fig. 655. Antholithus Zeilleri. 
Cambridge; A, ca. x I h; B,ca. x 2.) A, drawn (After Nathorst; x 2f.) 

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 — 43 /x 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 

1 Seward and Gowan (00) B. PI. ix. fig. 28. 

2 Nathorst (08) p. 20, PI. iv. 


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 1 in favour of 
regarding Antholiihus Zeilleri as the male organ of the plant 
which bore the fronds known as Lepidopteris Ottonis (Gopp.), 
originally described by Goeppert as Alethopteris 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 Antholiihus 
Zeilleri 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 Antholiihus and the 
latter is only found in beds containing the fronds. Nathorst 
called attention to resemblances between the cuticle of Antholithus 
and that of Baiera leaves, but according to the later investigations 
of Antevs, with which it would appear that Nathorst is in sympathy, 
there is a closer correspondence as regards cuticular structure with 

Fossils regarded by Nathorst as closely allied to his species are 
described by Leuthardt 2 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 4 cm. 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 
Preslii* from the Rhaetic of Franconia which he subsequently 
regarded as the male flower of Baiera Muensteriana Heer, a com- 
parison previously made by Heer 4 . This type consists of an axis 
bearing short lateral appendages terminating in oval bodies opening 

1 Antevs (14). 2 Leuthardt (03) p. 9 ? PL m. 

3 Schenk (67) A. p. 185, PI. xliv. figs. 9—12. 

4 Heer (77) ii. p. 52; Schenk (90) A. p. 261. 



at maturity into 10 — 12 spreading lobes each of which resembles 
in appearance a microsporangium of Antholithus Zeilleri. 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 Ehaetic age, described 
by Shirley 1 from Ipswich, Queensland, as Stachyopilys annularioides 
and S. Simmondsi require further investigation; they may be 
allied, to Stachyopilys Preslii Sch., though neither their morpho- 
logical nature nor systematic position can be settled without fresh 
data. Halle 2 describes some examples of a similar kind from the 
Jurassic beds of Graham Land as Stachyopilys , 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 3 and from the Stormberg (Rhaetic) series of South Africa 
as Stachyopilys sp. 4 

The generic name Ginkgoanthus has been adopted by Nathorst 
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 Stachyopilys Preslii as 
a type apart. As regards the English and Swiss specimens, the 
probability would seem to be that they are the microstrobili o 
some members of the Ginkgoales. 


This generic name 7 was first used by Saporta 8 for Nathorst's 
Zamiostrobus scanicus from Rhaetic and Liassic rocks in Scania 9 

1 Shirley (98) p. 13.. PI. xvin. 2 Halle (13) p. 88, PI. vi. fig. 13. 

3 Geinitz (76) B. p. 12, PI. n. 4 Seward (03) B. p. 66, PI. ix. fig. 2. 

5 Nathorst (99) p. 13.. PL I. figs. 33, 49. 

6 Used by Nathorst in Linnaeus's and not Brongniart's sense; Nathorst (08 
p. 23. 7 <tt€v6s, narrow ; p&xt-s, the backbone. 

8 Saporta (75) A. PL cxvn. ; (79) A. p. 193. 

9 Nathorst (75); (97) p. 20; (02) PL i. pp. 16, 17. 



and for a Liassic species from Belgium, 8. Ponceleti. Nathorst 
subsequently adopted Saporta's genus. I have elsewhere suggested 1 
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 2 (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) 3 . 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 
Xathorst 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 4 and afterwards transferred to Stenorachis, is represented 
by a comparatively slender axis 10 cm. 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. PI. xi. ; (78) ii. PL vi. fig. 8; (82) ii. A. pp. 18, 21, Pis. vi., ix. 
Heer (82) ii. A. p. 21. * Nathorst (75). 



species referred by Nathorst to the Cycadophyta 1 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 


Fig. 656. Stenorackis scanicus. (After Nathorst ; A, nat. size ; B, x 2. 

grounds for such relationship, but the resemblance of this and 
other species of Stenorachis to abnormal seed-bearing shoots o 
Ginkgo biloba may be significant (cf. fig. 631, D, p. 5). 

A similar but rather smaller type was described by Heer 2 
from Upper Jurassic rocks in Spitzbergen as Carpolithes striolatus. 
Nathorst 3 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 4 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 5 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. 

2 Heer (77) i. p. 47, PI. ix. fig. 17. 
4 Shirley (98) p. 16, PI. xx. 

3 Nathorst (97) p. 20, PI. i. fig. 15. 

5 Nathorst (02) p. 17, PI. L figs. 18—21. 




lepida (= Ginkgoites sibirica) and described by him under that 
name 1 . Similar specimens are correlated by Heer with other 
species of Ginkgo leaves, and Ginkgo grandiflora 2 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 z 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 species of Ginkgo. Fig. 657, B 

Fig. 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 4 which is undoubtedly of the same type as Heer's 
European examples. One of Heer's specimens from the Jurassic 
beds of Amurland 5 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 PI. xi. 2 Heer (82) A. p. 18, PI. vi. figs. 1—6. 

3 Ibid. p. 21, PI. ix. 4 Seward (12) p. 23, PL iv. fig. 52. 

5 Seward (12 3 ) p. 28, PI. I. fig. 8. 


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) 1 
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 Krystofovic 2 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 3 should be 
included in Stenorachis, as also Raciborski's Ixostrobus Siemirad- 
zkii* 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 
Stenorachis, 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. 


A genus founded 5 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 6 . Leaves oblanceolate to linear reaching a 
length of 12 cm. and a breadth of 2 cm. ; in the type-species, 

1 Seward (04 2 ) B. p. 179, figs. 39, 40. 

2 Krystofovic (10) PL in. fig. 5. 3 Seward (04 2 ) B. PL xix. 

4 Raciborski (92) PL n. figs. 5—8. 

5 Thomas (13). 6 ipeTfiov. a paddle. 




E. pubescens Thorn., 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 — 1-5 mm. apart, dieho- 
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 maybe 7 cm. long and 
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 
are confined to the subsidiary cells. 

A leaf figured by Ettingshausen 1 
from Wealden rocks as Cyclopteris 
squamata, which Schenk 2 suggests may be a segment of a Gink- 
goites, should probably be included in Eretmophylbnn. 

Fig. 658. A, Eretmophyllum saigha- 
nense. B — D, E. pubescens. D, 
epidermal cells. (A, after Seward ; 
B— D, after Thomas.) 

1 Ettingshausen (52) PI. iv. fig. 1. 

2 Schenk (71) B. p. 213. 



[CH. XL 

Eretmophyllum saighanense (Seward). 

This species (fig. 658, A), originally referred with some misgiving 
to Podozamites 1 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 Eretmo- 
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 2 from Jurassic-Cretaceous rocks of Alaska 
as Nageiopsis longifolia? Font, have little claim to be included in 
that genus 3 and may perhaps be allied to Eretmophyllum. 

1 Seward (12) p. 35, PI. iv. fig. 53. 

2 Fontaine in Ward (05) B. PI. xlv. figs. 1—5. 

3 Berry (10) p. 190. 



GINKGODIUM. Yokoyama. 

Yokoyama 1 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 Baiera. Gink- 
godiurn 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 Eretmophyllum 
(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 Nathorsti Yokoyama. 

The type-species was founded on specimens from strata in 
Japan assigned by Yokoyama 2 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, Pis. ii., in., vm., ix., xn. 




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 1 
records this species from the 
Bathonian series of Kamenka 
in the south of Russia (fig. 659, 
B). The specimens from Alaska 
named by Fontaine 2 Ginkgodium ? 
alaskense agree more closely with 


Heer 3 gave this name (after Fig. 659. Ginkgodium Nathorsti. (Nat. 
Czekanowski who discovered the JJ^A***' Yoko ^ ama; B ' aftei 
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 4 . Heer assigned to this 
genus some seeds associated with the leaves, also what he believed 
to be a male flower 5 , an example of a reproductive shoot of the 
type described on page 57 as Stenorachls. 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, PI. iv. figs. 9—11 ; PI. vra. fig. 3. 

2 Fontaine in Ward (05) B. p. 168, PI. xliv. figs. 3, 4. 

3 Heer (77) ii. p. 65. 4 Cf. Pinus flexilis ; Bot. Mag. Tab. 8467. 
5 Heer (82) ii. PI. vi. fig. 7. 


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 1 
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 2 , and some particularly 
good examples were collected by Prof. Obrutschew from Jurassic 
rocks in the Djair Mountains in Chinese Dzungaria 3 ; these occur 
with carbonised remains of Ginkgoites leaves in papery masses 
similar in the manner of preservation to the Palaeozoic paper 
coal from Russia 4 . The epidermal cells of the Dzungaria Czek- 
anowskia, possibly idt ntical 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 5 , but in the Rhaetic specimens 
from Scania the heavily cuticularised accessory cells are generally 
more numerous. 

It has been suggested by Jeffrey 6 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, Pis. iv., v., Tt, vn. 
4 Vol. i. p. 68. 5 Nathorst (06). 
6 Jeffrey (07) ; Hollick and Jeffrey (09) B. p. 63, PI. vi. figs. 1-3. 




Jurassic floras and a few examples are recorded from Cretaceous 

Czekanowskia Murray ana (Lindley and Hutton). 

On the specimen shown in fig. 660, A Lindley and Hutton 1 
founded the species Solenites Murrayana which they compared 

A B 

Fig. 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 Isoetes 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 

1 Lindley and Hutton (34) A. PI. cxxi. 


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 1 . 
This species was included by Saporta in Jeanpaulia and later 
transferred to Trichopitys, while Zigno 2 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 3 
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 4 of the species 
I went too far in uniting C. Murrayana and C. rigida. In Czeka- 
nowskia Murrayana the leaves, usually about 1 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 5 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 G . 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 leaves in a fascicle, a habit 
ot shown by any typical examples of B. Lindleyana though not 

1 Nathorst (06) PI. n. 

- Zigno (56) A. p. 216. For other references, see Seward (00) B. p. 280. 
3 Phillips (29) A. PL x. fig. 12. i Seward (00) B. p. 279. 

5 Phillips (75) A. p. 200, fig. 9. 6 Seward (00) B. p. 266. 

S. IV =; 




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 
segments are comparativelybroad 
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 Fl( \ J* 1 ; , c ^°."Tt? micr °? h t 

* ° (Phillips). (British Museum No. 39 

leaves are unbranched, and as 283; nat. size.) 

Nathorst 1 suggests the name 

Soleniies 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 01 

the simple or branched habit of the leaves of otherwise simil 


Czekanowskia rigida Heer. 

This species, founded on specimens from Siberian Jurassic 
rocks 2 , 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 3 
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. 

2 Heer (77) ii. p. 70; (78) ii p. 7; (82) A. p. 19. 

3 Nathorst (06). 


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 striatums; 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 Baiera 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 1 
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 
Lindleyana. Berry 2 points out a similarity between Fontaine's 
fossils and Baiera foliosa Font. Heer's type-specimens of C. 
nervosa from the Wealden of Portugal 3 are more like Czekanoivskia 
leaves. The leaves described from Siberia as C. setacea Heer 4 , 
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 

zekanowskia dichotoma Heer and C. capillaris Newberry. 

The branched leaves described under these names 5 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 6 from Middle Cretaceous beds as C. capillaris occur in 
closely packed groups, but no specimens have been discovered 
showing any scale-covered supporting axis. While admitting the 

)robability that these species and C. nervosa 7 from Wealden strata 

1 Fontaine in Ward (99) B. p. 685, PI. clxix. figs. 1, 2. 

2 Berry (11) p. 374. 3 Heer (81) PI. xvn. 

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, PI. vi. figs. 1—3. 

7 Heer (81) PL xvn. 



may be allied to Czekanowskia rigida, such evidence as is available 
points to a maximum development of Czekanowskia in the Jurassic- 


In 1870 Heer 1 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 Feildenia 
for Torellia because of the previous use of the latter name by 
Zoologists. Heer compared Feildenia 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 2 . 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 3 , 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 Feildenia, at best an ill-defined 
genus, can be distinguished from Phoenicopsis. 

Feildenia rigida Heer. 

This species, from Miocene beds of Spitzbergen 4 and Grinnell 
Land 5 , 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) Li. p. 44, PI. vi. figs. 3—12; PI. xvi. fig. 1 b. 

5 Heer (78) i. p. 20, Pis. I., n., vm. 


Feildenia NordensJcoldi Nathorst. 

A species from Upper Jurassic rocks in Spitzbergen founded by 
Nathorst 1 on leaves similar to those of F. rigida but smaller ; the 
lamina is generally 3 — 4 mm. broad and may reach a length of 
4-5 cm. 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. Nordenskoldi are described by Nathorst 
as Feildenia sp. 2 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 was founded by Heer 3 on linear leaves from 
Middle Jurassic strata in Siberia ; the leaves, in extreme cases 20 cm. 
long and varying in breadth from 2 mm. to 2 cm., occur in clusters of 
ix or more and even as many as twenty on very short and relatively 
road axes covered with small scale-leaves. These dwarf-shoots 
ere 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 
laced on the occurrence of the so-called interstitial veins : in 
me species of Phoenicopsis the parallel veins show no trace of 
smaller 'vein' between them, but occasionally in a leaf of the 
me species there are indications of interstitial 'veins.' The 
readth of the lamina is also an uncertain guide : well preserved 
pecimens show that the leaves may reach a considerable length 
nd that the lamina gradually decreases in breadth towards the 
arrow base. Species have been needlessly multiplied particularly 

1 Nathorst (97) p. 56, PI. ra. figs. 16—27. 

2 Nathorst (99) p. 15.. PI. i. figs. 25—30, 32. 3 Heer (77) ii. p. 49. 


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 1 . There is no information available as to 
the epidermal structure of the various types of Phoenicopsis 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 2 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 Cordaites. The Jurassic specimens 
from North Germany on which Salfeld 3 founded the genus Phyl- 
lotenia 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- 
phyllum* 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 Cordaites : 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) Pis. i., n. 3 Salfeld (09) B. p. 26, PI. iv. fig. 3. 
4 Lesquereux (78) p. 322; (80) A. p. 556, PI. 82, fig. 1. 


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 1 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 2 . Similar leaves collected by Nansen are 
described by Nathorst 3 . 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 
3 mm. 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, A). 
The vascular bundles are collateral, enclosed in a sheath of rather 
thick- walled cells : the xylem elements are spiral and scalariform 
and, as Solms states, the occasional preservation of single rows of 
circular bordered pits and the occurrence of lateral sieve-plates 
point to a Gymnospermous affinity. The mesophyll is fairly 
homogeneous and lacunar as shown in fig. 662, A and B : in the 
tangential section (B) the lacunar mesophyll is seen between the 
veins and their associated rows of rectangular cells. The epidermal 
cells are thickly cuticularised and papillose. In one section a few 
stomata were found showing two dark guard-cells, 40 fx long, 
surrounded by six faintly outlined cells (fig. 662, C) agreeing with 
those of Ginkgo and Ginkgoites, except in the absence of any 
overarching papillate subsidiary cells. This may, however, be 

1 Nathorst (07) p. 4. 

2 Newton and Teall (97) ; (98). 3 Nathorst (99). 




due to the section having passed slightly below the level of the 
epidermal surface. The cells of the epidermis are short and have 
straight walls. 

Though we should not be justified in asserting that the leaves 
named Desmiophyllum Solmsi are examples of Phoenicopsis, the 
probability is that they belong to that genus. Phoenicopsis is 

Fig. 662. Desmiophyllum Solmsi. A, transverse sections of leaves ; B. tangentia 
section of the lamina showing veins and mesophyll; C, stomata. (From 
sections in the Museum of the Geological Survey. London.) 

especially characteristic of Jurassic rocks and is best represented 
in the Middle Jurassic series of Siberia. The genus is recorded also 
from Spitzbergen 1 , Franz Josef Land, and Bornholm 2 : a species, 
P. Gunni 3 , has been described from Upper Jurassic rocks in Scot- 
land, the only example of the genus in Britain. Feistmantel 4 

1 Nathorst (97), p. 16. 2 Moller (03) p. 30. 

3 Seward (ll 2 ) p. 681, PI. ix. fig. 35 4 Feistmantel (77) fig. 9 




has figured an imperfect specimen from Upper Gondwana rocks 
in India that may be correctly referred to this genus. Incomplete 
leaves from the Rhaetic beds of Sweden described by Nathorst 1 
as Phoenicopsis cf. speciosa Heer may be examples of the genus, 
but the name DesmiophyUiim would be more appropriate. 

Fig. 663. Phoenicopsis speciosa. (After Heer; f nat. size.) 

*hoenicopsis speciosa Heer. 

In this Siberian Jurassic species 2 the leaves reach a length of 
50 cm. and a breadth of 7—8 mm. ; there are from 15 to 23 veins 

1 Nathorst (86) p. 96, PI. xxv. figs. 25, 26. 

2 Heer (77) ii. p. 112, Pis. xxix., xxx. 


in the lamina and a faintly marked interstitial 'vein/ probably 
the impression of a stereome strand, between each pair of veins. 
In one specimen Heer found 21 leaves in a single cluster but 
usually the number on a single dwarf-shoot is smaller. It was 
the superficial resemblance of a cluster of these leaves (fig. 663) 
to the leaf of some Palms that suggested the name Phoenicopsis. 
The leaves described by Heer as P. latior 1 are not distinguished 
from P. speciosa by any very definite character. Examples of 
detached leaves from Lower Jurassic rocks in Bornholm described 
by Moller 2 as cf. Phoenicopsis latior may equally well be referred 
to Podozamites. 

Phoenicopsis angustifolia Heer. 

The leaves are 4 mm. broad or less and have 6 — 8 veins without 
interstitial veins 3 . This species is recorded from Russia 4 , Siberia, 
China, and the Arctic regions, and leaves of similar type are 
represented by Phoenicopsis media Krasser 5 , which is probably 
merely a form of P. angustifolia 6 ; P. taschkessiensis Krass. from 
China ; also Chinese specimens first described by Potonie 7 withoul 
a specific name and afterwards named by Krasser 8 P. Potoniei. 

The species Phoenicopsis Gunni from Scottish Kimeridge beds 
is a similar type with leaves 3 — 4 mm. broad and 12 cm. long with 
eight veins and indications of interstitial 'veins.' 

? Phoenicopsis elongatus (Morris). 

Morris 9 founded this species on a linear leaf, now in the Britisl 
Museum, from the Jerusalem Basin (Triassic), Tasmania, which he 
referred to Zeugophyllites , a genus founded by Brongniart on a 
specimen from the Lower Gondwana rocks of India but never 
figured. To the same species McCoy 10 referred some broader 
leaves from Permo-Carboniferous strata in New South Wales: 
these were shown by Etheridge 11 and Arber 12 to be distinct from 
Morris's type and the latter author identified them with Noeg- 

1 Heer (77) ii. p. 113, Pis. xxix., xxxi. 2 Moller (03) p. 31. 

3 Heer (77) ii. pp. 51, 113, Pis. i., n„ xxx. 

4 Seward (07 2 ), PL vm. fig. 69. 5 Krasser (00) B. p. 147, PL m. fig. 4. 
« Nathorst (07) p. 7. Seward (11) p. 50. 

7 Potonie (03). 8 Krasser (05) p. 23. 

9 Morris in Strzelecki (45) B. p. 250, PL vi. figs. 5, 5 a. 
i° McCoy (47) B. " Etheridge (93) p. 75. 12 Arber (02 2 ) B. p. 17; (03 2 


gerathiopsis Goepperti (Schmal.) a species that is now recognised 
as identical with Cordaites (Noeggerathiopsis) Hislopi (Bunb.). 
Feistmantel 1 , who reproduces Morris's original figure of Zeugo- 
phyllites elongatus, assigns the Tasmanian plant to Podozamites. 
In 1903 2 I expressed the opinion that this species is a Phoenicopsis 
and figured some specimens from the Stormberg (Rhaetic) series 
of South Africa as Phoenicopsis elongatus. The leaves reach a 
length of more than 16 cm. ; the lamina is gradually tapered to 
an acuminate termination which may be the base, the distal end 
having a bluntly rounded apex. The veins are parallel and simple. 
Similar leaves have been described by Szajnocha and by Kurtz 3 
from Rhaetic rocks in South America. In the absence of specimens 
attached to an axis it is impossible to speak with confidence as to 
the systematic position of the detached leaves, but they bear a 
very close resemblance to some of the European examples of 
Phoenicopsis. The occurrence of these leaves in Australia, South 
Africa, and South America and the Phoenicopsis-\ike leaves recorded 
from India 4 , though not proving the existence of Phoenicopsis in 
the later vegetation of Gondwana Land, afford some evidence of 
its occurrence in the southern floras. 

1 Feistmantel (90) A. p. 150, PI. xxi. fig. 6. 

2 Seward (03) B. p. 67, PL ix. figs. 1, 9, 10. 

3 Szajnocha (88) B. p. 19, PI. n. fig. -4; Kurtz (03). 

4 Feistmantel (77) fig. 9. 



In this section are included several Palaeozoic genera most 
which have been assigned to the Ginkgoales on evidence which n 
most cases is wholly inadequate. 

Probably the oldest specimens referred to the Ginkgoales are 
some imperfect leaves from Middle Devonian rocks in Bohemia 
described by Potonie and Bernard as Barrandeina Dusliana 
(Krejci) 1 , but there is no evidence of affinity to this or to any other 
class of plants. 


This designation has recently been proposed by Zalessky 2 foi 
Schmalhausen' s Permian species Ginkgo cuneata which has n< 
substantial claim to be regarded as generically identical with 
Ginkgo biloba. Zalessky considers that it may belong to the 
Ginkgoales though the available data hardly justify more than a 
suggestion of possible relationship. 

Glottophyllum cuneatum (Schmalhausen). 

A Permian species from the Altai mountains 3 represented by 
obovate spathulate leaf 11 cm. long and 4 cm. broad with a compara- 
tively long ' stalk ' consisting of a narrow portion of the lamina : 
the lamina is traversed by slightly spreading veins about 1 mm. 
apart. In form this leaf resembles the genus Eretmophyllum* (cf. 
fig. 658) and may be a species of that genus, but in the absence 
of any information with regard to the epidermal features it is 
inadvisable to adopt a name implying affinity to the Ginkgoales. 

1 Potonie and Bernard (04) p. 45. 

2 Zalessky (12 2 ) p. 28 (footnote). 

3 Schmalhausen (79) A. p. 34, PI. iv. fig. 5 
* Thomas, H. H. (13). 



ian a 

3V an 


GINKGOPSIS. Zalessky. 

This generic term proposed by Zalessky 1 for Schmalhausen's 
Ginkgo Czekanowskii, but not denned by him, may be adopted for 
certain leaves resembling those of Ginkgo but which there is no 
adequate reason for regarding as the leaves of a member of the 

Ginkgopsis Czekanowskii (Schmalhausen) and Ginkgopsis miniita 
The small leaves, or leaflets, described by Schmalhausen 2 from 
the Permian of the Lower Tunguska river as Ginkgo Czekanowskii 
are more or less orbicular and the lamina, approximately 1-5 cm. 
broad, is divided into several bilobed segments (fig. 664, C). It 
is by no means certain that all the fragments included in this 
species are specifically identical; some bear a close resemblance 
to Ginkgo minuta Nath. 3 from the Rhaetic of Scania, a type which 
he assigns with hesitation to Ginkgo and compares with the leaves 
of Acrostichum peltatum. It is impossible without additional data 
to determine either the systematic position of the specimens 
included in these two species or their morphological nature. 
Through the kindness of Prof. Zalessky I have been able to examine 
photographs of some of the original material and, as shown in 
fig. 664, C, the supposed leaves are borne on a rachis-like axis, 
possibly of some Pteridosperm, a circumstance which though not 
proving that they are leaflets of a compound frond, favours that 


This name is proposed by Zalessky 4 for the leaves described by 

Schmalhausen and Renault respectively as Ginkgo integerrima and 

Ginkgo martensis on the ground that these Permian specimens do 

not afford satisfactory evidence of close affinity to the genus 

'inkgo. The two sets of leaves are probably specifically identical. 

Nephropsis integerrima (Schmalhausen). 

In fig. 664, A, B, are reproduced two of the leaves from the 

1 Zalessky (12 2 ) p. 28 (footnote). 

2 Schmalhausen (79) A. p. 84, PI. xvi. figs. 8—10. 

3 Nathorst (86) p. 93, PI. xin. fig. 112; PI. xx. figs. 14—16. 

4 Zalessky (12 2 ) p. 28 (footnote). 




Permian of the Lower Tunguska on which Schmalhausen founded 
his species Ginkgo integerrima 1 ; the characteristic features are : 
the transversely elongate form of the entire lamina, the spreading, 
dichotomously branched, veins and the short and relatively broad 
stalk-like basal portion of the lamina. As Zeiller 2 says, this 
species is probably identical with Ginkgo martensis 3 from Permian 
beds near Toulon-sur-Arroux (Saone-et-Loire) founded on a single 
leaf 3 cm. broad and 1-6 cm. deep. Zeiller reproduces Renault's 
figure in his volume on the flora of Blanzy and Creusot 4 and state 
that the specimen could not be found : he retains the name Ginkgo 

Fig. 664. C, Ginkgopsis Czekanowskii. A, B, Nephropsis integerrima. (From the 
originals of Schmalhausen's figures, supplied by Prof. Zalessky; nat. size.) 

but with the addition of a question-mark. The Permian leaves 
differ from those of Ginkgo in the absence of marginal veins — though 
this is of secondary importance — in the absence of a true petiole, 
in the form of the lamina and, from the majority of recent Ginkgo 
leaves, in the entire margin. The supposed leaves may be leaflets 
of a compound frond or possibly bracts from some fertile shoot. 

1 Schmalhausen (79) A. p. 85, PI. xvi. figs. 12—15. 

- Zeiller (96) A. p. 475. 

3 Renault (88) p. 232, fig- 47 c. 

* Zeiller (06) B. PI. XLvm. fig. 3. 



This generic name 1 was instituted by Schimper 2 for large 
cuneate leaves (figs. 665 — 667) from Upper Carboniferous and 
Permian rocks that had previously been included in Sternberg's 
genus Noeggerathia : the type-species is Noeggerathia flabellata 
Lind. and Hutt. 3 Psygmophyllum is thus defined by Schimper: 
' Folia pinnatisecta, pinnis erecto-patentibus, e basi valde angustata 
flabelliformis, longitudina liter flabellatim plicatis, plus minus 
profunde pinnatisectis vel margine lobatis seu crenatis; nervis 
pluries dichotomis, erecto-radiantibus.' Among other species 
referred by Schimper to Psygmophyllum are Noeggerathia expansa 
and N. cuneifolia from Permian beds in the Ural mountains 
figured by Kutorga 4 , Brongniart and later authors. In 1878 
Saporta 5 published a note giving the results of an examination 
of Brongniart' s specimens : he expressed the opinion that these 
species are portions of compound fronds comparable with Eremo- 
pteris, differing morphologically from Psygmophyllum flabellatum 
which he regarded as a shoot bearing simple leaves. He also 
pointed out that in the Ural fossils the leaflets have a more or less 
well defined midrib in contrast to the regular flabelliform dicho- 
tomous venation in the leaves of the English species, P. flabelli- 
forme : the latter he assigns to the genus Ginkgophyllum previously 
established for a Permian species, G. Grasserti 6 , consisting of an 
axis bearing spirally disposed cuneate and deeply divided leaves 
(fig. 669) very similar to some of the older Baiera leaves; while 
Schimper's genus Psygmophyllum is applied to the two Russian 
species P. expansum and P. cuneifolium. Saporta included a 
third Ural species in Psygmophyllum, P. santagoulourensis. An 
examination of some specimens of Brongniart' s Noeggerathia 
expansa in the British Museum leads me so far to agree with 
Saporta in the opinion that some of the specimens referred 
to that species are generically distinct from P. flabellatum. 
Schmalhausen's figures of the Ural species, which he refers to 

1 \(/0yfia, a fan. 2 Schimper (70) A. Vol. n. p. 192. 

3 Lindley and Hutton (32) A. Vol. i. Pis. xxvm. — xxix. 

4 Kutorga (44); Brongniart (45) B. PI. E; Schmalhausen (87) Pis. in., iv. 
See also Arber (12) p. 401. 

5 Saporta (78) ; (78 2 ) ; (78 3 ). 6 Saporta (75). 


Psygmophyllum, show a considerable variation in the venation of 
the leaves or leaflets : those represented in his Plate in. figs. 8 — 10 x 
agree with P. fldbellatum while others differ in the presence of a 
midrib (e.g. Schmalhausen's PI. lv. fig. 3). There has been con- 
siderable confusion in regard to the determination of these Russian 
specimens: as Arber says, Kutorga's Cyclopteris gigantea 2 is 
probably a true Psygmophyllum though other specimens subse 
quently referred to P. expansum should not be included in that 
genus. Zalessky 3 in 1912 proposed a new generic name Palamo- 
phyllum for the Russian species but retained Psygmophyllum for 
a Mongolian Permian specimen which he named P. mongolicum: 
this fossil is clearly a portion of a compound frond with leaflets 
like those of some forms of Palaeopteris. In a later paper 4 this 
author assigns Psygmophyllum mongolicum to P alamo fhyllum, but 
on his attention being called by Zeiller to Saporta's note of 1878, 
Zalessky 5 decided to abandon his proposed genus Palamophyllum 
in favour of Psygmophyllum. 

Confusion has also been caused by lack of uniformity in the 
use of the two generic names Psygmophyllum and Ginkgophyllum. 
The type-species of these genera I believe to be generically identical ; 
they agree in the general form of the leaves, the lamina being much 
more deeply divided in the type-species of Ginkgophyllum, also in 
the decurrence of the narrow basal portion of the lamina, and both 
are probably shoots, though the morphological nature of these and 
other types included in Psygmophyllum is by no means clear. 

Arber retains both names: as a matter of convenience he 
restricts Psygmophyllum to leaves that are entire or only slightly 
lobed, e.g. P. fiabellatum (fig. 665), and the more deeply dissected 
leaves such as those of Psygmophyllum Grasserti (fig. 669) he refers 
to Ginkgophyllum. This distinction is, however, purely arbitrary 
and on the analogy of the leaves of Ginkgo biloba it would seem 
preferable to include both deeply divided and more or less entire 
leaves in the same genus. Cambier 6 and Renier prefer the name 
Psygmophyllum to Ginkgophyllum on the ground that in the leaves 

1 Schmalhausen (87). 2 Kutorga (44) PI. n. fig. 7. 

3 Zalessky (12) p. 38, PI. vn. fig. 5. 4 Zalessky (12 2 ) p. 27. 

5 This decision was communicated in a letter (October, 1913). A. C. S. 

6 Cambier and Renier ( 10). 



assigned by some authors to Ginkgophyllum there are no marginal 
veins like those in the lamina of a Ginkgo leaf. This objection, 
though not in itself fundamental, is based on a sound principle, 
namely an objection to the assumption of affinity to Ginkgo implied 
by Ginkgophyllum, an assumption that rests on a superficial 
resemblance unsupported by any evidence of real value. 

The name Psygmophyllum is adopted both for entire and deeply 
divided leaves of larger dimensions than the similar leaflets of 
fronds included in such genera as Palaeopteris and Adiantites. 
Specimens usually occur as detached leaves, but when the leaves 
are attached to an axis the lamina is usually contracted into a 
fairly long, decurrent, basal portion ; there is no true petiole. 

The veins spread from the base of the lamina and are repeatedly 
forked ; they may be very numerous and in some forms occasion- 
ally anastomose, as in P. flabellatum, or much farthei apart, as in 
P. majus Arb. and P. Brownii (Daws.). The genus is at best a 
purely artificial one; we know nothing as to the reproductive 
organs or the anatomical structure, nor is it possible to determine 
in many instances whether the specimens are portions of compound 
fronds or shoots bearing simple leaves. 

Psygmophyllum ranges from Devonian to Permian strata, and 
if the Rhaetic leaves named Psygmophyllum ? crenatum (Nath.) are 
accepted as evidence, the vertical distribution must be extended. 
The genus occurs in England, Ireland, the continent of Europe, 
Spitzbergen, and North America; it is also a member of the 
Permo-Carboniferous floras of South Africa and India. 

Psygmophyllum flabellatum (Lindley and Hutton). 

The name Noeggerathia was given by Lindley and Hutton 1 to 
some specimens from the Newcastle Coal Measures; of the two 
examples figured one consists of a slender supporting axis bearing 
several torn cuneate leaves, which they speak of as part of a com- 
pound frond, and the other is a single leaf or leaflet. It is stated 
by Prof. Lebour and Dr Arber 2 that the original specimens cannot 
be found in the Hutton collection in the Newcastle Museum. In 
Mr Howse's Catalogue 3 a specimen is described as possibly the 
original of the larger example, and some years ago I examined a 

1 Lindley and Hutton (32) A. Pis. xxviii., xxix. 

2 Arber (12) p. 394. 3 Howse (88) A. p. 109. 
S. iv 6 



leaf in the Hutton collection which was believed to be the specimen 
reproduced on Plate 29 of the Fossil Flora. Fig. 665 represents 
an impression in the British Museum 1 from the Newcastle Coal 
Measures; the cuneate and partially torn lamina, 15 cm. long, is 
characterised by the very large number of forked veins, approxi- 
mately three to a breadth of 1 mm. In this and another specimen 2 


Fig. 665. Psygmophyllum flabellatum. (Brit. Mus. No. 40578; 
ca. § nat. size.) M. S. 

it can be seen that the veins occasionally anastomose, but this 
feature is more clearly shown in some better specimens in the 
Sunderland Museum 3 . One of these is represented half natural 
size in fig. 666. The leaves have no true petiole and are attached 
to the axis by narrow decurrent bases. A careful examination of 
the specimen did not enable me to detect any satisfactory evidence 
1 No. 40578. 2 No. 38927. 3 For other figures, see Arber (12). 




of the sheathing nature of the leaf-bases described by Arber, but 
it was on this example that I noticed the anastomosing of the 
veins (fig. 666, A — C), a character not hitherto recorded in the 
genus 1 . The lamina is torn and it is difficult to determine the 
degree of original lobing. Dr Arber believes that such specimens 
as that reproduced in fig. 666 are portions of a herbaceous plant 

Fig. 666. Psygmophyllum flabellatum. A — C, anastomosing veins. 
(From a specimen in the Sunderland Museum; \ nat. size.) M. S. 

and quite distinct from Ginkgo. While agreeing with the latter 
statement I do not regard the shoot-nature as definitely de- 
monstrated, though that is the interpretation usually accepted 
and not improbably correct. The habit appears to be identical 

1 A good specimen in the Manchester Museum also shows anastomosing veins. 



with that of the Permian species named by Saporta Ginkgophyllum 
Grasserti (fig. 669) and his figure 1 of that type suggests a shoot 
rather than part of a compound frond. Some species referred by 
authors to Psygmophyllum are certainly pinnae, while others bear 
a closer resemblance to shoots with simple leaves. Until better 
material is available we cannot determine either the morphological 
nature or the systematic position of the various examples assigned 
to this provisional genus. 

Psygmophyllum Kolderupi Nathorst. 

Nathorst 2 has recently founded this species, naming it after 
Dr Kolderup, on specimens from Devonian strata in West Norway 
consisting of pieces of shoots, or possibly compound fronds, 
bearing spirally disposed fan-like leaves or leaflets on long stalks 
and reaching a breadth of 15 to 30 mm. ; the veins are fine and 
repeatedly forked. The habit appears to be similar to that of 
Psygmophyllum flabellatum, but it is hardly possible to say whether 
we are dealing with fragments of a frond or branches bearin 
simple leaves. 


Psygmophyllum Kidstoni Seward. 

This species, very similar to P. flabellatum, is founded on 
specimens discovered by Mr Leslie in the Permo-Carboniferous 
rocks at Vereeniging, South Africa 3 . The cuneate leaves reach a 
length of 13 cm. and in some cases are deeply divided into two 
truncate lobes (fig. 667). The veins appear to be identical with 
those of P. flabellatum though no definite anastomosing has been 
detected. A photograph recently received of a new specimen 
shows some indication of a few cross veins, but the occasional 
anastomosing of veins should not be regarded as a feature of great 
importance. The axis of this species is broader than any so far 
found in the case of the English species, and the leaves are attached 
by a similar decurrent base. Incomplete leaves similar to P. 
Kidstoni though probably not specifically identical are described 
by Dun 4 from Permo-Carboniferous strata at Sydney as Rhipi- 

1 Saporta (84) PI. 152, fig. 2. 

2 Nathorst (15) p. 25, PL i. figs. 6—11; PL it. figs. 2—5. 

3 Seward (03) B. p. 93, PL xn. ; Arber (05) B. p. 213, fig. 47. 
* Dun (10) PI. 51. 



dopsis ginkgoides var. Suessmilchi ; they are probably referable 
to Psygmophyllum. 

Psygmophyllum Williamsoni Nathorst. 

A species 1 founded on imperfect leaves from the Upper Devonian 
of Spitzbergen agreeing closely in shape and venation with P. 
flabellatum. This is the oldest European example of the genus. 
In answer to an enquiry with regard to the venation Prof. Nathorst 
kindly informed me that he was unable to detect any definite 
traces of anastomosis and that the veins agree in their spacing 
with those of P. flabellatum. 

Fig. 667. Psygmophyllum Kidsloni. From Vereeniging, >S. Africa. 
(^ nat. size.) 

Psygmophyllum majus Arber. 

The large flabellate leaves 2 , often more than 16 cm. long and 
15 cm. broad, representing this Lower Carboniferous or Upper 
Devonian species from Newfoundland, are distinguished from 
P. flabellatum by their broader and less kite-like lamina and by 
the coarser venation. The distal margin is almost entire or 
characterised by broad and shallow lobes : as in P. Williamsoni 
no axis occurs with the leaves. As Arber points out, this species 

1 Nathorst (94) A. PL n. figs. 1, 2; Arber (12) PL xlh. fig. 4. 

2 Arber (12) p. 392, Pis. xlti.— xliv. 




bears some resemblance to Psygmophyllum Brownii originally 
described by Dawson 1 as Cyclopteris Brownii from Upper Devoni 
strata in Maine. 

Psygmophyllum Haydeni Seward. 

In 1905 some incomplete specimens were described from 
Permo-Carboniferous rocks in Kashmir as Psygmophyllum sp. 2 
for which, on the discovery of better material, a specific name was 
proposed 3 . The leaves reach a length of 13 cm. and are charac- 
terised by the division of the lamina into six or more obcuneate 

Fig. 668. Psygmophyllum Haydeni. (Nat. size.) 

segments, the divisions sometimes extending to the base of the 
broad part of the leaf (fig. 668). In the upper part of the lamina 
there are three to four veins per millimetre but lower in the 
lamina the veins are 1 mm. apart. Dr Arber 4 suggests that this 
species should rather be referred to Ginkgophyllum or Rhipidopsis. 

Psygmophyllum Hollandi Seward. 

This less satisfactory species is represented by some imperfect 

leaves from Carboniferous rocks of Kashmir 5 . The lamina is 

1 Dawson (63) PL xvn. fig. 6. 2 Seward and Woodward (05) B. 

3 Seward (07 5 ) B. 4 Arber (12) p. 400 (footnote). 5 Seward (07 5 ) B. 




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 Grasserti (Saporta). 

This Permian species from Lodeve 1 , France, agrees closely with 
P. flabellatum in the size and outline of the leavesas also in their 

Fig. 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 2 . Johnston 3 
describes a species, Ginkgophyllum australe, from ' Mesozoic ' rocks 

1 Saporta (75) p. 1018; (84) PI. 152, fig. 2. 

2 Saporta and Marion (85) p. 144, fig. 73, B; Saporta (82). 

3 Johnston (86) p. 178. 


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 1 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-\ike venation. The leaves agree closely with those of 
P. Grasserti 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 2 as Cyclopteris 
crenata and subsequently figured by Nathorst 3 as Ginkgo ? crenata, 
to some Permo-Carboniferous leaves included in Psygmophyllum 
suggests affinity with that genus. The obovate lamina, approxi- 
mately 12 cm. long, is slightly lobed on the upper margin and 
contracted below into a stalk-like base; the forked veins are 
nearly 3 mm. apart. Nathorst compares the species with Psygmo- 
phyllum flabellatum Lind. and Hutt., but the Rhaetic specimen 
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 4 a 
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 Psygmophyllum than of Ginkgo. 

1 Johnson (14). Since this account was written Pi of. 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). 
4 Arber (12) p. 392, Pis. xlii.— xliv. 


Other records of Psygmophyllum. 

Psygmophyllum primigenium (Saporta). Some leaves dis- 
covered by Grand'Eury in Permian rocks of the Urals were 
described by Saporta 1 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 2 affords a good 
example of a specimen that may be a Psygmophyllum leaf or a 
leaflet of a frond of the Noeggerathia type. Lesquereux 3 considers 
Dawson's specimen to be a pinnule of Palaeopteris. The same 
remark applies to Lesquereux's species Noeggerathia obtusa* from 
the Coal Measures of Pennsylvania included by Arber in Psygmo- 
phyllum ; it is probably a pinnate frond. 

The Russian species P. expansum and P. cuneifolium are dis- 
cussed in the account of the genus. The species Psygmophyllum 
Delvali 5 Camb. and Ren. from the Westphalian of Belgium is now 
admitted to be a leaf of Cordaites. A species described by Sand- 
berger 6 from the Permian of the Black Forest as Ginkgophyllum 
minus has been assigned by Sterzel 7 to Bicranophyllum. A leaf 
figured by Schmalhausen 8 from the Permian of East Russia as 
Baiera gigas is no doubt a Psygmophyllum allied to P. Kidstoni. 
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 9 are, as Zeiller 
points out, pinnules of a frond of the Eremopteris 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, PI. xvn. fig. 6; (71) A. p. 46, PI. xvi. fig. 172. 

3 Lesquereux (80) A. p. 305. 4 Ibid. PI. xlix. figs. 6, 7. 

5 Cambier and Renier (10) PI. vi. fig. 1. 

6 Sandberger (90) p. 101. 7 Sterzel (07) p. 820. 

8 Schmalhausen (87) PI. v. fig. 10. 

9 Schenk (83) A. p. 221, figs. 7, 8; PL xliii. figs. 22—24. 


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 1 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 2 . 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 Rhipidopsis 
as a genus of Gymnosperms of uncertain affinity. Schmalhausen 
attributes to Rhipidopsis some Samaropsis seeds 3 found in associa- 
tion with the leaves, and Kurtz 4 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 d 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 5 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 7 , may reach a 

1 Schmalhausen (79) A. p. 50, Pis. vi., viii. 

2 See p. 86, fig. 668. 

3 Schmalhausen (79) A. PI. viii. fig«. 9—11. 

4 Zeiller (96) A. p. 467. 5 Ibid. p. 471. 

6 Schmalhausen (87) PL in. fig. 10. 

7 Schmalhausen (79) A. p. 50, Pis. vi., viii. 



length including the petiole of 30 cm. 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, 

Fig. 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 


reaches a length of 10 cm. This species has been recorded from 
the Argentine but no figures have been published. It is no 
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 1 . (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 2 states that some 
specimens found by him in Permo-Carboniferous beds in th 
Argentine may belong to this species. 

Rhipidopsis gondwanensis sp. no v. 

The specimens for which this name is proposed were described 
by Feistmantel as Rhipidopsis ginkgoides 3 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 14 cm. 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 4 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 xlvi. A. 

2 Zeiller (96) A. p. 467. 

3 Feistmantel (81) p. 257, PI. n. fig. 1 ; (86) PI. in. A. figs. 1, 2. 

4 Fontaine and White (80) B. p. 99, PI. xxxvm. 





a thickened lower margin extending horizontally a short distance 
on either side of the petiole and presenting the appearance of 
being 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. 


This genus was first described by Grand'Eury 1 who, before the 
publication of the full description of the type-species, suggested 
the substitution of Eotaxites for Dicranophyllum 2 the name finally 
adopted 3 . The genus is 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 
Gymno sperms until we possess fuller information with regard to 
the reproductive organs or the anatomical structure. 

1 Compt. Rend. Vol. lxxx. p. 1021, 1875. 2 dUpavos, two-pointed. 

3 Grand'Eury (77) A. p. 272, Pis. xiv., xxx. 


In habit Dicranophyllum resembles Lepidodendron ; it is i 
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 7 mm., 
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. n. 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 Lepidodendron. 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 Sigillaria. 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. in. 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 




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. ii. p. 35). There is some evidence that the pith 
was discoid as in Cordaites 1 . The microsporophylls are said to 
be borne in small ovoid strobili in the axils of foliage leaves, but 

671. Dkranophyllum 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 2 
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 lxxi. figs. 3, 4. 

2 Zeiller (78) PI. x. figs. 1 a and 3. 


a radially segmented distal portion bearing some imperfectly 
preserved ovoid 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 1 demonstrate the occurrence of small 
oval ovules or seeds, 4x3 mm., 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 galhcum var. Parchemineyi 2 . Some 
leaves of D. striatum 3 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'Eury. 

This species, one of the two described by Grand'Eury in 1877 5 , 
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. PI. lxxi. fig. 5. 

2 Renault (96) A. p. 375. 3 Renault and Zeiller (88) A. p. 632. 

4 Fontaine (89) B. p. 213, PI. xciv. fig. 3; Berry (11) PI. lix. 

5 Grand'Eury (77) A. p. 272, Pis. xiv., xxx. Zeiller (80) A. PI. lxxvi. figs. 1, 2. 



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 1 and elsewhere; it 

occurs also in the Coal Measures of 

Portugal 2 . 

The specimens described by Grand'- 
Eury from Gard as D. tripartitum 3 , which 
I had an opportunity of examining in the 
Ecole des Mines, Paris, are not specific- 
ally distinguishable from D. gallicum. 
A1 j . j £ 7^ 77- Fig. 672. Dicranophyllum qal- 

A large decorticated stem or D. ; qauicum ,. ,».. v%, / 

° J licum. (After Zeiller from 

in the Paris Collection recalls a decor- Renault; § nat. size.) 
ticated stem of Lepidodendron. Some 

'imperfect specimens described by White 4 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- 

I 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 5 as Cyperites ? sp. on 
the ground of the similarity of the lamina to some fragments, 
probably of Lepidodendroid leaves, described by Lindley and 
Hutton 6 as Cyperites bicarinata and subsequently included by 
Heer 7 in his genus Distrigophyllum. In a note to his account of 
Mesozoic plants from Portugal Heer 8 renames the plant Distrigo- 
phyllum lusitanicum and compares it with Dicranophyllum gallicum 
Grand'Eury. De Lima 9 recognised the true nature of the speci- 
mens from the Stephanian of Portugal and published a full 

1 Zeiller (92 2 ) A. p. 96. 2 Lima, de (88) Pis. i., in. 

3 Grand'Eury (90) A. p. 335, PI. vi. figs. 12, 13. 

4 White (99) B. p. 272, PI. xli. fig. 10; PL lxxih. fig. 1. 

5 Gomes (65) p. 32, Pis. i., v. 6 Lindley and Hutton (32) A. PI. XLiir. 
7 Heer (76) A. p. 39. 8 Ibid. (81) p. 11, footnote 1. 
9 Lima, de (88). 

s. iv 7 



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 art 
given off at a much more acute angle. 

Dicranophyllum robustum Zeiller. 

This type 1 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 2 . 

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 Sigillaria. 

Dicranophyllum Richiri Renier 3 . 

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 

Dicranophyllum anglicum Kidston. 

This, the only British species, has recently been described from 
the Westphalian beds of Staffordshire 4 . The crowded leaves, 
3-50 cm. 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, Pis. xiv., xv. 

3 Renier (07) p. 186, PL xvn. figs. 3—7; (10 2 ) PL cxvn. 

4 Kidston (14) p. 170, PL xiv. figs. 3, 3 a. 







of 1-25 mm. 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 1 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 2 are also too fragmentary to be accepted as 
trustworthy evidence of the occurrence of the genus in the southern 

Dicranophyllum striatum Grand'Eury. 

This species like several others is founded on detached leaves 3 , 
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 4 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 5 , 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 6 from the Coal Measures 

1 Schenk (83) A. p. 222, PL JJM. figs. 11, 12. 

2 Ibid. PI. xlii. figs. 17, 18. 

3 Grand'Euiy (77) A. p. 275. Renault and Zeiller (88) A. PI. lxxt. fig. 2. 
1 Renault and Zeiller (88) A. p. 631, PI. lxxi. fig. 1. 

5 Sterzel (07) p. 391, PI. xv. figs. 9—11. 

6 Lesquereux (80) A. pp. 553, 554, Pis. lxxxiti., lxxxvii. 


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 with a frayed 

Dicranophyllum glabrum (Dawson). 

Under this name Dr Stopes 1 has recently described a well 
preserved leaf from the Westphalian series of New Brunswick. 
The specific name was first applied by Dawson 2 to specimens which 
he referred doubtfully to Psilophyton. The type-specimen is 9 cm. 
long and 3 mm. 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 3 

The imperfect specimens described by Dawson 4 from Devonian 
rocks in Queensland as Dicranophyllum australicum and sub- 
sequently figured by Jack and Etheridge 5 consist of a slender 
axis, 3 mm. 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 6 : as shown by drawings reproduced by Potonie 7 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 falcate 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, PI. xvra. fig. 47. 

2 Dawson (62) p. 315: for other references, see Stopes loc. tit. 

3 Renault (96) A. p. 378; (93) A. PL lxxxii. fig. 2. 

4 Dawson (81) A. p. 306, PI. xm. figs. 15, 16. 

5 Jack and Etheridge (92) B. p. 49. 

6 Schenk (90) A. erroneously includes Sigillariostrobus bifidus Geinitz, (73) 
PL in. figs. 5 — 7, in Dicranophyllum. See also Sterzel (93) A. p. 111. 

7 Potonie (93) A. PL xxvm. figs. 1, 2. 


A foliage-shoot described by Renault 1 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. 


Saporta 2 proposed the generic name Trichopitys in 1875 for 
some shoots from the Permian beds of Lodeve 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 3 .' 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 T. 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 4 from Lodeve (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. PI. lxxxii. fig. 1. 

2 Saporta (75) p. 1020. 

3 Ibid. (84) p. 263, PI. cm. fig. 1. 
* Zeiller (00 2 ) B. p. 254, fig. 182. 




Renault 1 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 

Fig. 673. Trichopitys heteromorpha. (After Zeiller; { nat. size.) 

forking of the segments than T. Milleryensis, and a leaf recenl 
figured by Dr Stopes 2 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 3 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 lxxxh. fig. 2 ; (96) A. p. 378. 

2 Stopes (14) p. 79, PI. xviii. fig. 47. 

3 White (99) B. p. 272, PI. XLi/fig. 10; PI. lxxiii. fig. 1. 


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 T. 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 Jeanpaulia, and T. Lindleyana 1 ; 
in the latter species he included the specimens doubtfully assigned 
by Lindley and Hutton to Solenites as Solenites ? furcata 2 . These 
and other Jurassic leaves that are referred by some authors to 
Trichopitys are usually regarded as examples of Baiera 3 ; there are 
no adequate grounds for believing them to be closely related to 
the Permian Trichopitys. 

Zeiller 4 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) PI. clv. figs. 1—9. 2 Lindley and Hutton (37) A. PI. 209. 

3 Seward (00) B. p. 266. 4 Zeiller (ll 2 ) p. 234. 




Sewardia latifolia (Saporta). 

1849. Otozamites latifolia Brongniart, Tableau, p. 106. 

1872. Sphenozamiies latifolius Schimper 1 , Traite, Vol. n. p. 163. 

1875. Cycadorachis armata Saporta, Plant, Jurass. p. 196, PL 117, fig. 1. 

1895. Withamia armata Seward, Wealden Flora, Vol. n. p. 174, PL n. 

figs. 1,2; PL v. fig. 1. 
1900. Sewardia latifolia Zeiller, Elements Paleobot, p. 233. 

This species is represented by woody axes, about 1 cm. in 
breadth reaching a length of 50 cm., from the Wealden beds of 
Sussex, bearing more or less orbicular, entire, leaves or leaf-like 

Fig. 674. Sewardia latifolia. Axis (A) and single leaf (B). (British 
Museum; f nat. size.) 

organs, 6 cm. 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 Cyclopteris 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 2 suggested 

1 See also Saporta (75) A. p. 188, Pis. 112, 113. 

Seward (95) A. p. 175. 


the generic name Acanthoza mites 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 Sphenoza mites 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 — 3J 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 1 that the apparent lamina of the hooked leaves is a 
winged petiole, the hooks representing lateral leaflets. 

Seward ia longifolia (Salfeld). 

This species was described by Salfeld 2 from the Solenhofen 
beds (Upper Jurassic) of Bavaria as Baiera ? longifolia Heer : it 
is founded on branches nearly 30 cm. long bearing large fan- 
shaped deeply divided leaves, or leaf-like organs, in the axils of 
Tecurved 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, Pis. xx.. xxi. 

Fig. 674*. Sequoia sempervirens. Near Crescent City, California. (From 
a photograph by Professor A. Henry.) 



The Coniferales, by far the largest section of the Gymnosperrj 
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 their 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 




problem that has long exercised the ingenuity of investigators. 
The view expressed by Jeffrey 1 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 


Fie 075. A, B, Phyllocladus trkhomanoides. C, P. hypophylla. D, E, 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. 


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 1 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 2 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 leaf is characteristic of some species of 
Podocarpus (fig. 676). The presence of long and short shoots is 
a striking feature of Pinus, Larix, Pseudolarix, Cedrus, and 
Sciadopitys : the short shoot, as Goebel says ' takes no part in 
the construction of the permanent skeleton of the tree 3 .' The 
whorled arrangement of leaves characteristic of several Cupres- 
sineae and the Callitrineae is not a constant feature and, as in 
Lycopodium, both whorled 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 strobili 4 . Proliferous cones are not uncommon in 
some genera : the prolonged axis of the cone of Cryptomeria 
japonica shown in fig. 677 bears 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) PI. 815. 3 Goebel (05) p. 444. 

4 For examples see Sterzel (76); Eichler (82); Worsdell (04); Bartlett (13); 
Shaw, W. R. (96); Robertson (06); Renner (04); Bayer (08); etc. 





transverse dehiscence. The microstrobili of Cedrus are similar 
but longer. In Torreya (fig. 684, D), Taxodium, Widdringtonia, 

Fig. 676. Podocarpus latifolia. (§ nat. size.) M. S. 

Fig. 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 




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 Finns. In Araucaria Muelleri and 
A. Rulei (fig. 678) the microstrobili 
reach a length of 25 cm. : in A. excelsa 
and A. Cookii 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 Cunninghamia 1 , 
Pseudolarix 2 , and Keteleeria are borne 
in umbels, while in Cryptomeria and 
Taxodium 3 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 Microcachrys* the 
wings vary from 2 to 6 (fig. 684, C) 
and in Dacrydium 5 and Podocarpus 6 
(fig. 684, A) there are 2 or 3 small 
bladders. In Taxus, Cephalotaxus, 
Torreya, Sciadopitys, the Cupressineae, 
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 7 , 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 mam- 
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) PI. cm. ' 2 Bot. Mag. Jan. 1908. 

3 Gard. Chron. Nov. 25, 1893, 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). 

Fig. 678. Microstrobilus ot 
Araucaria excelsa (A) and 
Araucaria Rulei (B). (After 
Seward and Ford; A, nat. 
size; B, i nat. size.) 




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 
Bidicillii 1 , similar to some cones of Encephalartos, may be 28 cm. 
in diameter and in some species of Agathis 2 they exceed 11 cm. 


Fig. 679. A, Araucaria Cookii var. luxurians. B, A. Cookii. C, Araucaria Cookii,. 
microstrobilus. D, E, Araucaria Muelleri, part of a microstrobilus and a single 
sporophyll; ps, microsporangia. F, Araucaria Montana, branch. (After 
Seward and Ford.) 

and are 14 cm. 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, I). 

1 Gard. Chron. April 14, 1894, p. 465. 
"2 Seward and Ford (06) B. 




Fig. 680. Araucaria brasiliensis, cone. (£ nat. size; from a specimen 
in the Royal Gardens, Kew.) 

Fig. 681. Araucaria Cunniughamii, cone. 
\\ nat. size.) 

(After Seward and Ford; 




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. Bidivillii broad and woody (fig. 683, B, C). In 
Finns, 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 r 

Fig. 682. A, Agathis Moorei, cone-scales; s, scar of seed. B, Agathis loranihifolia, 
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. Coulteri, 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 Abies, 
Larix, Pseudotsuga the dual nature of the scales is obvious at 
s. iv 8 




maturity, the bract-scale usually extending beyond the edge of 
the seminiferous scale (fig. 705), in Abies bracteata reaching 
length of 5 cm. In Sequoia (fig. 702 B) the cone-scales show no 
outward sign of a double structure, but each scale contains two 

Fig. 683. Cone-scales of Araucaria Cookii (A), A. imbricata (B), A. Bidwillii (C), 
and A. excelsa (D, E) ; I, 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 


represents the lowest term of a series, the upper end of which is 
represented by the scales of Agaihis 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 1 . The scales of such genera as Sciadopitys, Athrotaxis, 
Cryptomeria (fig. 684, S, 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 ao 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 
Arauearia 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 
Athrotaxis (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 2 . 

In a recently published paper on the vascular anatomy of the 
megasporophylls of Conifers by Miss Aase 3 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. A separate origin 
for bract and seminiferous scale-bundles is recorded in several 
Abietineae and in some other Conifers. The origin of the vascular 

1 For references to literature on the morphologj- of cones, see Coulter and 
Chamberlain (10); Worsdell (04) ; Rendle(04); Lotsy(ll); also Celakovsky (82); 
Kramer (85); Bayer (08); Aase (15). 

2 Worsdell (99). 3 Aase (15). 




Fig. 684. A, B, C, microspores of Podocarpus Totara (A). Pinus Laricio (B), Micro- 
cachrys tetragona (C). D, E, microsporophylls of Torreya califomica (D) and Abies 
alba (E). Megasporophylls etc. of Podocarpus spicata (F) ; P. Totara (G),o, ovule ; 
P. neriifolia (H); P. imbricafu (I), o, ovule; Cunninghamia sinensis (K), 
m, membrane ; Cryptomeria (L, M). N, 0, megastrobilus and seed of Athrotaxis 
laxifolia. P, megasporophyll and seed of Dacrydium Balansae; a, epimatium. 
Q, seed of Cupressus sempervirens. R, S, megasporophyll of Sciadopitys ; a, ovu- 
liferous scale ; 6, bract-scale. T, megasporophyll of M icrocachrys ; a, epimatium. 
V, seed of Torreya ; i, ii, integuments ; a, vascular strands. [After Burlingame 
(A), Coulter and Chamberlain (B), Thomson (C), Gard. Chron, (D, N, 0), 
Kirchner, Loew and Schroter (E, Q), Pilger (F, H, P), Gibbs (G, I), Eichler 
(K— M, S), Worsdell (R, T), Oliver (V).] 


supply of the double cone-scale is, however, not constant even in 
the same cone, and in Araucaria Bidwillii 1 each cone-scale is 
supplied by two separate strands from the vascular axis though 
in other species a single bundle enters the cone-scale and divides 
later. It would appear, therefore, that the single or double origin 
of the lower normally orientated bundles and of the upper set of 
inversely orientated bundles is far from constant, and the data 
derived from anatomical study do not afford a satisfactory means 
of determining the morphological nature of the cone-scales. It is 
held by Jeffrey and his school that the Abietineae represent the 
oldest members of the Coniferales and that the Araucarineae are 
a more recent development, the apparently single cone-scale of 
Araucaria and Agathis being derived from the double cone-scales 
of the Abietineae. Some botanists, e.g. Vierhapper 2 , while 
believing that the seminiferous scale of the Abietineae is an organ 
belonging to an axillary shoot subtended by a bract-scale and that 
the cone-scales of other Conifers are also double structures, whether 
or not externally divided, regard the Araucarineae as earlier in 
origin than the Abietineae. If the cone-scales of the Araucarineae, 
to take the extreme type, are in origin double and homologous 
with the obviously double cone-scales of the Abietineae it is more 
logical to regard the Abietineae as the precursors of the Araucari- 
neae. The evidence afforded by fossils in my opinion lends strong 
support to the greater antiquity of the Araucarineae, and I venture 
to believe that no adequate reasons have been given for regarding 
the cone-scales of the Araucarineae as other than simple leaves 
bearing ovules. If, as seems probable, the Coniferales are mono- 
phyletic in origin the cone-scales of the different families are in 
all probability variants of a common type and, in opposition to 
the view which is most in favour, I regard the double cone-scales 
of the Abietineae and the corresponding organs of other Conifers 
which afford evidence of a double structure as derivatives of a 
simple form of sporophyll strictly comparable with the sporophyll 
of a Lycopodium,, the placental outgrowth assuming an increasing 
degree of individuality in the different lines of evolution illustrated 
by various types of strobilus. It is noteworthy that the transition 
from foliage leaves to megasporophylls in the Araucarineae is 
1 Worsdell (99). 2 Vierhapper (10). 


of^en very gradual in contrast to the much more sharply defined 
break between leaves and scales in most of the Abietineae and 
many other Conifers. The cone-scales of the Abietineae are 
recognised as more complex and more recent developments, the 
seminiferous scale being an excessively enlarged placental out- 
growth from a megasporophyll, while in the intermediate types 
such as Sequoia, Cryptomeria, and others the separation between 
the two parts of the cone-scale is much less complete 1 . This 
morphological question is too complex to discuss fully in a 
general summary: students should, however, be warned that 
several botanists do not agree with the opinion that is here 
expressed. It is at least fair to add that the views expressed 
by Prof. Jeffrey and his pupils with regard to the relative positions 
of the Araucarineae and the Abietineae in an evolutionary series 
are stated with an assurance which is misleading to those unfamiliar 
with the nature of the evidence 2 . 

In Saxegothaea (fig. 685), Dacrydiwn (fig. 684, P), and some other 
genera each ovule is surrounded by a cup-like integument (fig. 684, 
T a), formerly called the arillus but recently styled the epimatium 3 ; 
this* is by some authors considered to be the equivalent of the 
seminiferous scale. 

This inadequate account may serve to call attention to a complex 
morphological problem which has an important bearing on questions 
connected with the relative positions of the several genera. It would 
be out of place to enter fully into this difficult subject, but it is one 
that demands careful attention by students of extinct types. 

The number of seeds borne on each scale is an important 
feature in the recognition of genera. In the Abietineae each 
scale usually bears two seeds though it is not uncommon to find 
single-seeded seminiferous scales such as those of Pinus monophylla 
(fig. 686, A — C) in which there is a deep cavity showing that the 
seed was partially embedded in the supporting organ. Such a 
scale might, as a fossil, be easily mistaken for an Araucarian cone- 
scale. In Cunninghamia there are three seeds to each scale (fig. 
684, K) ; in Aihrotaxis and Cryptomeria 3 — 6 ; in Sequoia 5 ; and 

1 Eichler(81). 

2 For a recent discussion on the origin and relationship of the Araucarineae see 
Burlingame's paper (15 2 ) which appeared after this chapter was written. 

3 Pilger (03) p. 16. 




Fig. 685. SaxegotJiaea conspicua. Section of ovule ; n, nucellus ; i, integument ; 
st, stigma-like apex of nucellus; o, epimatium; c, young megaspore. (After 

Fig. 686. A — C, Pinus monophylla, cone-scales with two seeds (A), one seed (B), 
and in distal view (C). D, E, Taxodium mucronatum, cone in surface-view (D) 
and section (E); s, seed. F, Taxodium distichum, scale. (From specimens in 
the British Museum.) M. S. 




in Sciadopitys 7 — 9. Fig. 686, E represents half a cone of Taxodium 
in which the distally expanded woody ends of the scales are tightly 
joined by their edges and form a hard case enclosing as in an 
ovary several angular seeds, the slender stalks being shrivelled 
and inconspicuous. The cones of the Cupressineae and Calli- 
trineae are characterised by a whorled arrangement and a com- 
paratively small number of the scales. In Cupressus the cones 
are oval or spherical and each scale bears 6 — 20 seeds: in the 
Callitrineae the cones are valvular (figs. 703, 762, B — D) and the 
scales vary from 2 to 3 in Callitris and from 
7 to 8 in Widdringionia. The small cones 
of Saxegothaea consist of one-seeded mega- 
sporophylls which become fleshy and par- 
tially concrescent (fig. 687) ; in Juniperus 
the strobilus has the appearance of a berry ; 
in Microcachrys the leaves pass gradually 
into the single-seeded verticillate megaspo- 
rophylls, each with two vascular strands 1 
(fig. 684, T) and an epimatium, a, on one side 
of the ovule; in the ripe cone the mega- 
sporophylls are fleshy but not connate as in \ 
Saxegothaea 2 . In Dacrydium the megaspo- 
rophylls differ but slightly from the foliage 
leaves in some species, e.g. in D. Balansae 
(fig. 684, P) a single leaf at the apex of a 
branch bears an ovule partially covered by 
a hood-like epimatium. In Torreya 3 a very 
short shoot in the axil of a leaf bears two 
bracts and each subtends an ovule and two 
pairs of bracteoles. The seeds of T. cali- 
fornica, which may be 4 cm. long, are enclosed by a thick integument 
differentiated into a sarcotesta and sclerotesta surrounding a 
ruminated endosperm (fig. 688) : there is a ring of vascular bundles 
at the limit of the free part of the integument and this is regarded 
by Oliver 4 as homologous with the tracheal plate at the base of 

Fig. 687. Saxegothaea con- 
spicua. (After Stiles.) 

1 Worsdell (99). 

2 Stiles (08); Noren*(08); Tison (09); Bot. Mag. Tab. 8664 (1916). 

3 Robertson (04) ; (07). 

4 Oliver (02) ; (03). 




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. Cephalotaxus 1 has plum- like seeds 
similar to those of Torreya. In Phyllocladus 2 (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 

Fig. 688. Torreya micifera, 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; mw, wall of megaspore; n, nucellus; ps, pro- 
thallus. (After Oliver.) 

without any megasporophylls. In Podocarpus 3 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). 

3 Pilger(03); Brooks and Stiles (10); Stiles (12); Gibbs(12); Sinnott (13). 


fertile and its apex overtops the ovule, while a second bract is 
sterile. Fig. 684, H shows the strobilus of P. neriifolia 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. Gliick 1 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 
lies in a pocket formed by the enclosing wing, while in Pinus the 
wing embraces the seed like a pair of pincers. In Agaihis (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 2 , 
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 Cuppy 3 to be a primitive 
feature, is recorded in Podocarpus MaJcoyi*. 

The relation between nucellus and integument is less uniform 
in Conifers than in Cycads. In some genera, e.g. Agaihis (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 5 and to a greater degree in Saxegothaea (fig. 685, st) 

1 Gliiek (02) p. 402. 2 Coker (09). 3 Guppy (06). 

* Lloyd (02). 5 Seward and Ford (06) B. 


this protrudes through the micropyle : in Fitzroya patagonica the 
prolongation of the integument as a micropylar tube with a stigma- 
like terminal expansion is particularly striking. There is no 
regular pollen-chamber as in Cycads and Ginkgo, but in Pseudo- 
tsuga 1 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 2 . The archegonia 
of Conifers usually occur at the apex of the proth alius 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 Taxodium 34, in Agaihis 60, 
irregularly distributed on the sides of the prothallus. In Widdring- 
tonia? 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. 


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 
gen era 6 . Arnoldi 7 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 8 and Lawson 9 favours the removal of 

! Taxodium and Cryptomeria to the Cupressineae. Miyake's re- 
searches 10 point to a similar affinity in the case of Cunninghamia. 
The genera Athrotaxis, Fokienia, and Taiivania 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 Ibid. (13). 5 Shaw, W. R. (96); Arnoldi (01). 

6 See also Saxton (13 2 ). 7 Arnoldi (01); Lawson (10); Radais (94). 

8 Coker (03). 9 Lawson (04). 10 Miyake (10). 




used by Masters, has been revived by Saxton 1 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 2 makes Phyllocladus the sole genus 
of Phyllocladoideae, while Miss Robertson 3 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 4 . 

I. ARAUCARINEAE. Agathis, Araucaria. 

II. CUPRESSINEAE. Cupressus, Chamaecyparis, Libo- 
cedrus, Thuya, Juniperus, Fitzroya, Diselma, Thujopsis, Taxodium, 
Glyptostrobus , Cryptomeria, Cunninghamia , Taiwania, Fokienia, 
Athrotaxis, Tetraclinis. 

III. CALLITRINEAE. Callitris, Actinostrobus, Widdring- 


ABIETINEAE. Pinus, Cedrus, Larix, Pseudolarix, 
Picea, Tsuga, Abies, Pseudotsuga, Keteleeria. 

VII. PODOCARPINEAE. Podocarpus, Dacrydium, Micro- 
cachrys, Acmopyle, Pherosphaera, Saxegothaea. 


IX. TAXINEAE. Taxus, 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 5 , 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 role with the Polypodiaceae among the Ferns, 

1 Saxton(10 2 ); (13 2 ). 2 Pilger (03). 3 Robertson (06). 

4 For other views on classification, see Vierhapper (10). 

5 Drude(90); Eftgler(89); Graner(94); Hildebrand (61); Vierhapper (10). 




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 1 , Siam, the Malay region, the Philippines 2 , 
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 nourishes in northern 
Europe and Siberia, Canada and the northern United States, the 
Himalayas, and Japan. Tsuga is more especially a North American 

Fig. 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 

I is characteristic of N.W. America and is recorded from Formosa. 
Pseudolarix is a native of N.E. China and Formosa. Cedrus 3 
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). 

Fox worthy (11). 

3 Hooker, J. D. (62). 



land, New Caledonia, and the Queen Charlotte Islands. The tw 
trees of Araucaria Bidwillii shown in the Frontispiece are survivon 
of a forest on the hills of Queensland. The photograph reproduced 
in fig. 689, for which I am indebted to Dr Wieland 1 , illustrates the 
habit of Araucaria imbricata on the eastern slopes of the Andes 
in South- West Argentina where the trunks reach a diameter o 
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 Calif ornian 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. Libocedrus, 
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; Diselma 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. Taiwania 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. 

SEQUOIINEAE. Sequoia is confined to the Pacific coast of 
North California ; S. sempervirens the species with ' the stronger 
hold upon existence' extends into Oregon, while 8. gigantea forms 
groves in the valleys of the Sierra Nevada. 

PODOCARPINEAE. Podocarpus, one of the more successful 
genera, is essentially a southern type : in Africa it extends from 
1 Wieland (16), p. 224. 


Cape Colony through East Africa to Abyssinia; it occurs in 
S. America from Patagonia to Brazil and replaces Pinus on the 
mountains of Costa Rica 1 ; in the West Indies, Malaya, in the 
Himalayas, China, Japan, Formosa 2 , Tasmania, New Zealand, New 
Caledonia, the Fiji Islands and New Guinea. Dacrydium 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 3 and occurs in 
New Zealand, Tasmania, New Caledonia, New Guinea, and one 
species grows in the Chilean swamps. Saxegothaea 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 Agaihis, Podocarpus, 
and Dacrydium, and his remarks illustrate the importance of 
taking into account p#laeobotanical 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 5 .' 

TAXINEAE. Taxus 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 

Harshberger (11) p. 304. 2 Diimmer (12). 3 Stapf (96). 

4 Groom (16). 5 Guppy (06) p. 300. 




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 T. calif ornica and the Pacific 
separates the latter from the two species in China and Japan 1 . 
Cephalotaxus lives in central China, Japan, and India. 

Anatomical features. 

The anatomy of Conifers, more especially from the point oi 
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 2 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 

The wood of a Conifer consists only of tracheids, with or without 

resin-canals, and xylem-parenchyma and is characterised by 

narrow medullary rays usually one-cell broad. For convenience 

in description it is proposed to speak of the wood of the Conifer 

type as pycnoxylic 3 in distinction to the Cycadean type of 

wood which is styled manoxylic. The presence or absence of 

1 Berry (08 2 ) p. 648. 2 Jeffrey (05) p. 1. 

3 ttvkvSs, compact; ixavos, porous, loose in texture. 


well defined rings of growth should be noted and attention paid 
to the breadth of the late summer ('autumn') wood: Goeppert 1 
considered the breadth of annual rings a character of importance, 
but Kraus 2 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 3 . Though as a rule there is a considerable difference in 
the thickness between the walls of the spring and summer tracheids, 
in Podocarpus Nagi 4 the difference is slight. In some species 
of Araucaria the rings are absent or feebly marked, a fact noticed 
long ago by Nicol 5 , and in other Conifers, e.g., Widdringtonia 
juniperoides and Tetraclinis 6 there may be no definite rings; in 
Libocedrus macrolepis 1 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 8 . 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 9 . 

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 10 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 Nieol (34) A. p. 139. 

6 Conwentz (90) A. p. 33. 

7 Fujioka (13) p. 213. 8 Thomson (13) p. 33; Gothan (07) p. 25. 
9 Seward (92) B; Gothan (08 2 ) ; Antevs(16); (17). 

10 Jeffrey (03) ; (05); etc.; Penhallow (07) pp. 123 et seq. ; Jones (13 2 ). 

IV 9 




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, Pseudolarix, 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 veitical 1 . 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 

#n|it't*'i$» .0 - 


t »•»•(» '; vi 


Fig. 690. 

A, Sequoia semperrirens. 

B, Abies sp. showing traumatic canals in the 
(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. 



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- 
hallow 1 holds, and I believe rightly, that they are not primitive 
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 2 suggests, be correlated with a greater need of protection. 
Kirsch 3 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 4 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 5 , 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 6 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 eithei single or double, 

1 Penhallow (07) p. 150. 2 Gothan (07) p. 40. 

3 Kirsch (11); Thomson (13) p. 38. See also Burlingame (15 2 ). 

4 Conwentz (90) A, p. 45. 5 Groom and Rushton (13). 
1 Chrysler (08) B. p. 204. 





4 *^f*j 




I 3£l 

| 0)3 


Flg. 691. A, Araucaria Bidwillii, tracheids from the cone-axis. B — D, F, Agathis 
bornensis ; B, wood of branch, multiseriate pits and, at the ray, scalarif orm pits ; 
C, tangential section; D, radial section of branch showing transition from 
alternate and opposite to scalarif orm 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.) 




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 1 
(fig. 691, D) and Conwentz 2 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 3 : 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 4 . 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 5 (fig. 692) the pits are as a rule uni- 
seriate and often flattened. Worsdell 6 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 poly st achy a 7 and I have seen similar " 

pits in the wood of Torreya calif ornica and several Fi°- 6y 2. Tracheids 
other conifers other than the Araucarineae. The oi Saxegothaea con- 

spicua. (After 

size of the bordered pits though worthy of notice stiles.) 
is not in itself a feature of much value. As Nicol 
fiist pointed out, in Araucaria they are larger than in Taxus; in 
Pinus they are larger than in Araucaria: Kraus 8 speaks of the 

2 Conwentz (92) p. 35. 

1 Jeffrey (12) PI. vi. fig. 6. 

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). 


Araucarian pits as small (9 — 12-8 /z) in contrast to the broader pits 
(up to 21/x) 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 type 1 . 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 2 , also in some species of 
Cupressus 3 . In Pseudotsuga spiral bands may occur in all the tra- 
cheids of an annual ring. 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 4 . 
The walls of tracheids frequently exhibit well marked spiral 
patterns 5 , 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 Sequoia xylem-parenchyma may be absent though 
it is present in older branches 6 . Such parenchyma occasionally 
occurs in Abietineoiis wood 7 , 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. Kows of parenchyma occur in the wood of 
Abies pectinata 8 , also in Podocarpus and Dacrydium. In Taxodium 9 
the thick horizontal walls of the cells are a characteristic feature. 
Wood-parenchyma is raie in the Araucarineae and, as Penhallow 10 

1 Schenk in Schimper and Schenkt90) A. p. 848. 

2 Bailey (09). 3 Jones (12); (13). 

4 Gothan (05) p. 54; Penhallow (07) p. 41 ; Burgerstein (08) p. 104; Kraus (83) 
p. 103; Nakamura (83). 

5 Kraus (88); Gothan (05). 6 Conwentz (92) p. 35. 
7 Burgerstein (06); Bailey (09). 8 Kny (10). 

9 Schroeter (80) p. 30. 10 Penhallow (04). 


pointed out, tracheids with horizontal patches of resin may be 
mistaken for resiniferous parenchyma; but true parenchyma 
occasionally occurs 1 . 

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 2 , apparently overlooking the fact that 
this term (Sanio's ' Balken ') was used by Miiller 3 for the horizontal 
bars previously described by Winkler 4 on the tracheids of Araucaria 
brasiliensis (fig. 693, I). Groom and Rushton 5 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 6 .' 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 7 
has shown that in the first-year wood of Araucaria and in the cone 
of A. Bidwillii the pits are not always contiguous and rims of 
Sanio may then be present. Moreover, as Thomson 8 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 9 , 
Sanio's rims are not represented 10 . 

The most recent contribution to our knowledge of the rims of 
Sanio is by Mr Sifton 11 who describes them in petioles of Cycas 

• » Jeffrey (12) p. 536. 2 Gerry (10). 

3 Miiller (90). ■ Winkler (72). 

5 Groom and Rushton (13). See also Rushton (16). 

6 Holden (13) p. 252; (13 2 ). 7 Jeffrey (12) PL vi. fig. b 
8 Thomson (13) p. 22. 9 Gothan (10) p. 32. 

10 In a recent paper entitled 'Gliding growth and bars of Sanio' (Grossenbacher, 
Amer. Journ. Bot. vol. i. 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). 


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 tangential 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 1 has given the 
name Abietineous pitting. In the great majority of recent 
genera other than members of the Abietineae the horizontal and 
tangential walls are smooth (fig. 693, L, 0) ; but there are exceptions. 
The ray cells in the cone-scales of Agathis are pitted and species 
of Juniperus 2 , 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. 43, 45, fig. 7; Stopes (15) p. 63. 






| ool ool 


jj°° \oS\ 



0\Ja \<D 

N , U 

&\\ & 



0\\ CPO 


In p 

M O 

Fig. 693. A, Abies Veitchii, medullar}- 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, 0, Pits in medullary-ray cells (radial view) in 
Cedrus atlantica (G) ; Taxodium rtisticli um (H) ; Podocarpus andina (K) ; P. salici- 
folia(L) ; Glyptostrobus (M) ; Sciado])itys (N) ; sp, spring wood ; s, summer wood ; 
Thuya gigantea (0). 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.) 



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 th 
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 cuirent terminology. There is 
a difference of opinion as to the value of medullary-ray pitting a 
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 i 
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 ('Eipore') 
occupies the field in some species of Finns but the same type 
occurs also in Sciadopitys 1 , species of Podocarpus 2 (fig. 693, K, N) 
Microcachrys, Dacrydium, and Phyllocladus 3 . The Araucarinea 
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 4 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) PI. cxxxviii. ; Fujioka (13). 

2 Kleeberg (85). 

3 Schenk in Schimper and Schenk (90) A. p. 855. 

4 Thomson (13) p. 30. 


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 
geneia 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 Cedras, Taxodium, and 
Glyptostrobus, while in Juniperus and several other genera the pits 
are smaller and more numerous 1 . 

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 2 . In a single species 
of Abies the depth varies from 1 to 63 cells 3 . 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, Larix, 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*, in 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 

Penhallow (07). ■ Essner (86); Barber (98). 

Fuiioka (13). 4 Strasburger (91) p. 21. 



occur in Abies 1 and several members of other families. Chrysler 2 
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, Finns Strobus, or, as in other species of Pinus 
('Hard Pines'), their walls are characterised by irregular ingrowths 
or pegs. Wettstein 3 states that in Picea omorica horizontal 
tracheids though common in the main stem do not occur in the 
rays of branches. De Bary 4 recognised tracheids in the rays of 
Sciadopitys ; they have been recorded also in Juniper us, Cupressus, 
Thuja, Sequoia 5 and, as the result of wounding, in Cunninghamia 6 . 

The occuirence 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 7 and Cryptomeria. 
In some cases, e.g., Abies magnificat, 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 9 states that he found none in Podocarpus. 

The structure of the stomata in the leaves of Conifers is fairb 
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 

1 Thompson (12). 2 Chrysler (15). 

3 Wettstein (90) p. 511. 

4 De Bary (84) A. p. 490. See also Tassi (05) quoted by Vierhapper (10). 

5 Gordon, M. (12); Jones (13 2 ). 6 Jeffrey (08). 
7 Kubart (IP) 8 Jeffrey (05). 9 Thomson (13) p. 31. 





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 1 . Caution must be ex- 
ercised in using the number of resin-ducts as a diagnostic character. 
Schroeter 2 draws attention to the occasional absence of ducts in 
Picea excelsa leaves while in others 1 or 2 are present. In most 
leaves theie is some mechanical tissue immediately below the 
epidermis either as scattered fibres or a continuous layer, but in 

i. 694. Leaves in transverse section. A, Cephalotaxus Fortunei. B, Torreya 
myristica; J, transfusion-tissue. C, Araucaria imbricaia; t, transfusion-tissue 
D, Pinus monophylla ; e, endodermis. E, Cryptomeria japonica. F, Cedrus 
Libnni; 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- 
larix and some other genera. The structure, extent, and position 
of short isodiametric tracheids in association with the conducting 

tin important feature. This tissue, the elements of which 

142 COSTIFERALES (recent) [ch. 

are usually termed transfusion-tracheids 1 , is regarded by some 
authors as homologous with the centripetal wood of Cycadean 
leaves (Vol. in. p. 31) and Bernard definitely adopts the term centri- 
petal xylem. Jeffrey 2 and, more recently, Takeda 3 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, t) 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 4 . 

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., Araucaria, 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 5 , 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 2 ). 

3 Takeda (13); see also Thomson (13). 

4 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). 


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 1 has described the northern evergreen Conifeis 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 woo.d-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 2 . 

ARAUCARINEAE. There is a close agreement in the structure of the 
wood in the two members of this family. 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 the 
xylem-elements. In the wood of Agathis robusta 3 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 4 considered by Lignier to possess diagnostic 
importance: Jeffrey 5 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 6 . 

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 5 cm. ; A. macrophylla 17x5 cm.). 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 Naturlichen Pflanzenfamilien the student should consult the Report of the 
Conifer Conference, Journal of the Royal Horticultural Society, Vol. xrv. 1892. 

3 Xoelle (10). 

4 Thiselton-Dyer (OP); Seward and Ford (06) B. 

5 Jeffrey (12) p. 565.. 6 Thomson (13) p. 15. 




Fig. 695. A, Agathis Moorei. B, E, Agathis australis. C, Agathis vitiensis. D, 
Podocarpus Motleyi. (After Seward and Ford; £ nat. size.) 


Fig. 696. Agathis Moorei (A) and Agathis australis (B) cones. (After Seward 
and Ford; A, £ nat. size; B, f nat. size.) 




bear a close superficial resemblance to Cedar cones but the latter are 
relatively narrow and often more or less flattened at the apex 1 . Bommer 2 
calls attention to the resemblance of Agathis cones to those of the Dicotyledon 
Dammaropsis kingiana (Moraceae). 

Araucaria. (Frontispiece; figs. 678 — 681, 689, etc.) The falcate tetra- 
gonal leaves of A. excelsa illustrate one type of leaf that is seen in its smallest 
form in A. Balansae (4 — 5 x 2-5 mm.). In A. Bidwillii (fig. 697) the leaves 
are subsessile and the flat lamina may reach a length of 7 cm. : in A. Housteinii 

Fig. 697. Araucaria Bidwillii. (After Seward and Ford ; nat. size.) 

the ovate-lanceolate leaves may be 10 cm. by 1 cm. Dimorphism 3 in the foliage 
of a single shoot is not uncommon. The striking difference in some species 
between the juvenile and adult foliage is illustrated in fig. 698. In the broad- 
Jeaved species (Colymbea section) resin-canals occur between the veins (fig. 694, 
C) and in the Eutacta section, e.g., A. excelsa, the canals are scattered. Strobili 
and cone-scales of Araucaria are described in an earlier part of this chapter 
(see page 113). 

1 Fliche (96). 

2 Bommer (03) B. PI. x. figs. 164, 165. 

3 Masters (91); Bommer (03) B. PI. v. fig. 23; Siebold (70) PI. cxl. 

s. IV 





CUPRESSINEAE. The absence of resin-canals in the xylem is a feature 
shared by other families; but in the occurrence of xylem-parenchyma in 
different regions of the wood the Cupressineae differ as a rule from the Abie- 
tineae, though this is not a constant distinguishing character. The pits in 
the field vary from 1 to 6 or 8 in some genera, e.g., Taxodium and Glypto- 
strobus 1 : Gothan 2 applies the term Cupressoid to medullary-ray pits character- 

Fig. 698. Araucaria excelsa. A, Mature form. B, Seedling, (f nat. size.) 

ised by an upper and lower border (fig. 693, H) but, as pointed out on a previous 
page, the position of the pore is by no means constant and in Taxodium the 
ray pits are fairly large and almost simple. 

Cupressus and Chamaecyparis. By Beissner 3 and many other authors both 
names are used in a generic sense, though Masters regards Chamaecyparis as 
a subsection of Cupressus. One distinguishing feature is the presence of more 

1 Kleeberg (85). 

2 Gothan (05) p. 47. 

Beissner (91). 




than 2 seeds on each cone-scale of Cupressus. The leaves are whorled, in 
4 ranks, and appressed to the axis (fig. 699). The young foliage of Cupressus, 
Thuya, and other genera (fig. 700), which in the adult state has the form of 
scale-leaves, consists of spreading linear leaves: individuals in which this 
juvenile foliage persists are often spoken of as species of the 'genus ' Retinospora. 
Cones oblong or globular, composed of a few pairs of scales with peltate distal 
ends either smooth or provided with a prominent umbo (fig. 699). 

[G. 699. Cupressus Macnabiana. (From Rendle and the Gardeners' Chronicle.) 

Thuya. Very similar to Cupressus in the habit of its bi- or tri-pinnate 
shoots. The cones of 8 — 10 decussate scales are distinguished from those of 
Cupressus by their elongated, oblong, form and by the upwardly directed 
scales with thickened apices in contrast to the more horizontal and peltate 
scales of Cupressus: there are 2 seeds to each scale, winged (sect. Euthuja) 
or wingless (sect. Biota). 

Libocedrus. Foliage shoots often broader than in Thuya; in L. decurrens 
the appressed, flat, leaves- are characterised by a long decurrent portion and 



in L. Doniana the whorled arrangement is less obvious. There is a large 
canal below the midrib of the leaf as in Thuya and Gupressus. The cones 
(4 — 6 valves) are longer and relatively narrower than in Thuya; seeds un- 

Fig. 700. A, Young plant of Libocedrus decurrens; c, cotyledons; pr, primordial 
leaves; tr, transitional leaves. B, Branch with adult (ad) foliage. (After 
Rendle, from Veitch. ) 

equally winged. Juniperus. The polymorphism of the shoots is especially 
striking; the leaves, 2 — 3 in each whorl, being small and appressed, spinous 
and spreading, or flat and linear. The very narrow leaves of J. Sabina 



differ widely from the broader, flat and sharply pointed leaves of J. drupacea. 
There is a resin-canal near the lower surface 1 or, in some species, e.g., J. oxy- 
cedrus, there may be no canal. The more or less globular or elongated fleshy 
strobilus consists of 1 — 4 whorls of megasporophylls. 

Fitzroya and Diselma. In some examples of F. patagonica the leaves are 
short and crowded and slightly falcate, in others more spreading; while in 
Diselma Archeri 2 the shoots resemble those of some Lycopodiums. The 
leaves are in alternate ternary whorls and there is a single canal below the 
vein. The cones consist of 3 alternate trimerous whorls and the fertile scales 
bear a variable number of winged ovules. Thujopsis. Shoots similar to 
those of Thuya but the decussate leaves are rather larger 3 . Cones narrow, 
consisting of 8 clavate scales with 5 small winged seeds on each scale. 

Taxodium. The slender deciduous shoots, 8 — 10 cm. long, bear 2-ranked 
linear leaves 2 cm. long, acute and lanceolate, also small leaves 10 — 17 mm. 
long and barely 1 mm. broad. Circular depressed branch-scars are a charac- 
teristic feature of the leafless shoots. Globular cones 4 composed of a few 
peltate scales with an irregular crenulate upper margin (fig. 686, D — F) each 
scale bearing 2 slightly winged seeds. The almost leafless spikes of micro- 
strobili are a noteworthy feature. The wood of Taxodium closely resembles 
that of Sequoia sempervirens : the thicker horizontal walls of the xylem- 
parenchyma are quoted by Gothan as a characteristic feature 5 of Taxodium. 
Lingelsheim 6 says that this distinction is not valid; he states that simple pits 
occur in the cross-walls of the xylem-parenchyma of Taxodium but not in the 
cells of Sequoia. Pits are, however, present in the parenchyma of both these 
genera. Glyptostrobus. Similar in habit to Taxodium, but the leaves are not 
2-ranked 7 ; the cones are more pyriform; the seeds are smooth and narrow 
with lateral wings. The comparatively large circular pits on the radial walls 
of the ray cells are characteristic 8 . 

Cryptomeria. Foliage-shoots (fig. 677) as in Araucaria excelsa: a large 
resin-canal occurs below the midrib of the laterally compressed leaves (fig. 694, 
E). The cones have 20 — 30 peltate scales characterised by the deeply cleft 
comb-like upper portion (fig. 684, M). The xylem-parenchyma is said by 
Fujioka to be confined to the region between the spring and summer wood : 
the pits in the field are variable in position and the breadth of the pore may be 
vertical or oblique. 

Cunninghamia. Leaves densely spiral and spreading, narrow, lanceolate, 
acuminate and serrate, 2-5 — 5 cm. long, with a narrow decurrent base: a large 
canal below the midrib. Cones ovoid -globular, 4 cm. long, composed of 
broad thin scales with a serrate edge and a fimbriate membrane on the abaxial 

1 Kirchner, Loew, and Schroter (06) pp. 293, 314. 

2 Hooker, J. D. (60) PL xcviii. 3 Fujioka (13). 

4 Gard. Chron. Nov. 25, 1893, p. 657. 

5 Gothan (06); (09). 6 Lingelsheim (08). 
7 Masters (00). 8 Kleeberg (85). 




side of the 3 winged seeds (fig. 684, K). Branch-scars occur on the stems 1 . 
Taiwania. This genus 2 has the habit of Cryptomeria and cones recalling 
those of Cunninghamia and Tsuga : each scale bears 2 seeds. Fohienia. The 
single species 3 is, in certain respects, intermediate between Cupressus and 
Libocedrus; the cones are globose like those of Chamaecyparis and each 
scale bears two unequally winged seeds; the foliage is nearly identical with 
that of Libocedrus. 

Fig. 701. Athrotaxis cupressoides. 

Athrotaxis. (Figs. 684, N, O; 701.) Leaves short, loosely spreading 
and slightly imbricate {A.*), similar to those of Araucaria excelsa, 

1 Lotsy (11) p. 51. 2 Gard. Chron. Feb. 4, 1911. 

3 A second species has recently been described by Hayata (17) 

4 Baker and Smith (10) p. 305 (and photograph). 


more appressed and smaller, decussate or in close spirals (A. cupressoides) or, 
in A. laxifolia 1 , more like the foliage of Sequoia gigantea. The apex of each 
cone-scale has a sharp point at the distal end. Tetraclinis (fig. 703, B). This 
genus, which occurs in Algeria and Morocco, and is usually placed next to 
Callitris and Widdringtonia, has recently been transferred by Saxton 2 , as the 
result of his work on the life-history of T. articulata, to the Cupressineae : he 
believes that the Callitrineae were derived from the Northern Cupressineae 
through some type resembling Tetraclinis. The flattened foliage-shoots 
resemble Salicornia and bear 4-ranked small leaves; the cones consist of 
4 nearly equal decussate scales, each scale having 2 seeds with unequal wings. 

SCIADOPITINEAE. Sciadopitys. The single species is characterised by 
the long and rigid 'double needle' reaching a length of 12 cm. with a broad 
furrow on the upper surface and a deeper groove on the lower face. Mr 
Boodle 3 has recently described an example of concrescence in needles of Pinus 
Laricio Poir. var. nigricans Pari, which points to the probability that a morpho- 
logical similarity exists between the double needle of Sciadopitys and abnormal, 
fused, leaves of Pinus. Reference is made by Boodle to other possible views 
that have been advanced with regard to the morphological nature of Sciadopitys 
needles. The oblong woody cones, 7x4 cm. 4 are fairly easy to identify by 
the rounded and reflexed upper margins of the scales (fig. 684, S) ; there are 
5 — 15 seeds, with a narrow wing, on each cone-scale. 

SEQUOIINEAE. Sequoia. Attention has already been called to some 
of the anatomical features. The oval pits on the field (2 — 6) have an upper 
and a lower border though the pore is not infrequently obliquely vertical. 
Anatomically the wood of S. sempervirens is considered by Gothan 5 to agree 
more closely with that of Taxodium than with S. gigantea. The leaves of 
S. gigantea (=WeUingtonia) are 3-angled and decurrent, -5 mm. long; those 
on the fertile shoots are broader, shorter and imbricate. S. sempervirens 
bears 2-ranked linear, sessile, but not decurrent leaves with an abruptly 
spinous apex. There is a resin- canal below the midrib and transfusion- 
tracheids form conspicuous lateral groups. The cones of the two species are 
of the same type, but thost of S. gigantea are larger and occasionally reach a 
length of 9—10 cm. (fig. 702). 

CALLITRINEAE. The foliage-shoots, similar to those of some Junipers, 
are characterised by short decurrent scale-like ' leaves in alternate ternary 
or decussate whorls; the ovate or globular cones are composed of a few 
valvate scales. Actinostrobus 6 (fig. 703, A). The very small leaves in ternary 
whorls have a free apex and the slender shoots closely simulate those of some 
species of Veronica and Thamnea depressa (Bruniaceae). The 6 cone-scales 
are oblong, acute, and the base of the cone is invested by 6 rows of crowded 
scales which gradually pass into the foliage -leaves : each scale has 2 — 3 winged 

1 Gard. Chron. Jan. 31, 1891, p. 147. 2 Saxton (13 2 ); (13 3 ). 

3 Boodle (15). 4 Siebold (70) PI. on 

5 Gothan (06); (09). See also Jeffrey (03); Gordon (12). 

6 Saxton (13). 



seeds. Callitris. Similar in habit to Tetraclinis (fig. 703, B) : in some forms 
(C. arborea) the small leaves are closely appressed to the axis ; C. glauca 1 shows 
a considerable range in the form of the leaves, and in C. rhomboidalis the 

Fig. 702 A. Sequoia gigantea. Shoot with cones. (Nat. size.) 

shoots are especially slender. Saxton 2 points out that the tracheids have a 
single row of separate pits and draws attention to the occurrence of horizontal 

1 Baker and Smith (10) p. 118. 

2 Saxton (10 2 ); (10 3 ). The wood of the Callitrineae requires more thorough 




bands above and below each pit. Cones usually spherical, reaching 2-5' cm. 
in length, with 6 smooth or tuberculate valves. In C. Macleayana the cones 
are pyramidal and have 6 — 8 valves 1 . 

Widdringtonia. The adult foliage-shoots bear rather longer leaves in 
decussate pairs 2 ; the twigs are cylindrical and not flat as in Thuya. In 
quickly growing shoots the leaves may be spiral; they have no hypodermal 

%&«* jjr 











1 > '■ 


Fig. 702 B. Sequoia gigantea. An unusually large cone. 
(British Museum; nat. size.) 

layer like that in the leaves of Callitris. Spirally thickened tracheids occasion- 
ally occur in the wood. The cones consist of 4 decussate thick and warty 
valves, each scale bearing 7 — 8 winged seeds. 

ABIETINEAE. Pinus. The needle-like leaves on short shoots are a 
striking feature. In P. silvestris and many other species each short shoot 

1 Baker and Smith (10). 

2 Rendle (96). 



A B 

Fig. 703. A, Actinostrobus pyramidalis. B, Tetraclinis articulate. (After Saxton.) 

Fig. 704. Pinus excelsa, cone and foliage-spur. 
(From a photograph by A. Howard.) 



bears 2 needles plano-convex in section. In P. monophylla (fig. 694, D) each 
shoot has usually a single sharp-pointed cylindrical leaf 4 cm. long. Three- 
needled Pines occur in N. America and the Himalayas but not in Europe. 
In P. Cembra, P. pence (the only European examples), P. Strobus, P. excelsa 
(fig. 704), P. koraiensis, etc., the short shoots bear 5 needles and each is trian- 
gular. Thomson 1 regards the dwarf-shoot of Pinus as a derivative of a longer 
shoot with spirally disposed needles: he has recently described examples of 
dwarf -shoots in recent Pines bearing an abnormally large number of leaves. 
In P. silvestris shoots with 3 leaves are not uncommon : in P. excelsa wounding 
caused the development of as many as 15 needles on a single dwarf-shoot. 
Jeffrey, on the other hand, regards the spur-shoot as a primitive attribute of 
the coniferous stock. The length of the leaf varies considerably and may 
reach 30 cm. ; the margin is generally entire, but in P. Cembra the apical 
portion is finely serrate. The structure of Pine leaves 2 is well known, but 
reference may be made to the twin bundles in hard Pines, a distinction from the 
singlestrand in the soft Pines 3 , thepresenceof a well-defined endodermis( fig. 694, 
D, e) and the infoldings of the chlorenchyma 4 . There are two types of cone, 
that of the Pinaster group in which the distal end of the woody seminiferous 
scale is more or less pyramidal and has a central umbo (cf. fig. 785), prolonged 
in some species, P. ponderosa 5 , P. Jeffrey^, etc., into a short recurved spine 
or, in P. Coulteri, into a large curved spinous process. Another type, illus- 
trated by the Strobus group, is distinguished by the flatter imbricate scales 
with the umbo at the tip of the free rounded margin (fig. 704). The cones of 
the latter type resemble those of Picea. The variation in the size and form of 
cones from the same tree in Pinus excelsa is worthy of notice from a palaeo- 
botanical standpoint 7 . 

Cedrus. The more slender needles of Cedrus, most of which occur in 
clusters on short shoots, are approximately triangular in section (fig. 694, F) 
and have a single bundle and two canals next the lower epidermis. The 
persistent leaf-bases resemble those of Tsuga and Picea (fig. 706, B, D). Canals, 
though not normally present in the wood, are induced by wounding; xylem- 
parenchyma may occur in the late summer-wood and ray-tracheids are present, 
but less prominent than in Pines and some other genera. The deciduous 
nature of the mature cone-scales would lead one to expect their preservation 
as fossils. A characteristic feature of the cones is the presence of radially 
disposed lines normal to the edge of the sporophylls 8 . Larix. The narrow 
linear decurrent leaves, keeled on the lower surface, have a resin-canal close 

1 Thomson (14). 

2 Zang (04) attempts to classify Pine leaves on the basis of anatomy. 

3 For anatomical features of hard and soft Pines, see Holden, R. (13 3 ); Jeffrey 
and Chrysler (06). 

4 Kirchner, Loew, and Schroter (06) figs. 188, 222, etc. 

5 Gard. Ckron. Nov. 15, 1890, pp. 561, 569. 

6 Ibid. March 23, 1889. 7 Bommer (03) B. Pls.n., iv. 
8 Fliche (96) p. 86. 


to the epidermis at each angle. There is a single vascular bundle but n 
thick-walled hypoderm. Bailey 1 notes the occurrence of wood-parenchym 
on the outer face of the summer-wood. The persistent subglobose or more 
elongated cones, reaching a length of 10 cm. (fig. 705, G), are in most species 
characterised by the large size of the bract-scales like those of Abies and 
Pseudotsuga. Bommer calls attention to the superficial resemblance of Larch 
cones to the fruit of Petrophila diversifolia 2 (Proteaceae). Pseudolarix 
resembles Larix in habit (fig. 705, A, B); the male strobili are umbellate; 
the leaves linear lanceolate and up to 7 cm. long. The cones are ovate and 
bear loosely imbricate, pointed deciduous scales; the bract-scale is shorter 
than in Larix. Picea. The short and narrow leaves are tetragonal or more 
or less flat in section : a canal occurs below the vein or there may be two lateral 
resin-ducts. The persistent bases of the leaves form prominent pegs (fig. 706, 
D). The cylindrical or oval cones, reaching 16 cm. in length, consist of leathery, 
concave imbricate scales; the bract-scales are concealed 3 . The stem of a 
variety of P. excelsa (var. tuberculata) A , characterised by large conical tubercles 
of cork, bears a close resemblance to that of Xanihoxylum (Rutaceae) 5 . Tsuga. 
The leaves are flat, decurrent (fig. 706, B), similar to those of Taxus, Abies, and 
some species of Picea ; but there is a single resin-canal below the vein and the 
lamina is petiolate and not sessile as in Sequoia sempervirens. In some cases 
the lamina is finely serrate. The cones closely resemble those of Picea except 
in their smaller size; the bract-scales are usually concealed, but in T. Pat 
toniana 6 they are longer and reach a length of 7-5 cm. 

Pseudotsuga. The leaves resemble those of Tsuga, but there are tw 
lateral canals and the lamina is attached by a narrow base to a slightly pro 
minent leaf-cushion (fig. 706, C). The wood is like that of Larix but the 
tracheids often possess spiral bands, and xylem-parenchyma occasionally 
occurs. Resin-ducts are present in both normal and injured stems. The 
cones, 5 — 10 cm. long, resemble those of Tsuga but differ in the 3-pronged 
conspicuous bract-scale; they are pendulous and the scales are persistent. 
Keteleeria. Shoots similar in habit to those of Abies; leaves flat, a canal at 
each angle. Cones similar to those of Pinus Cembra: microstrobili umbellate. 
Abies. Leaves usually flatter than in Picea and often larger ; the apex may 
be notched though this is not a constant feature 7 . There is no persistent 
leaf- base (fig. 706, A). Two lateral canals 8 are generally present in the leaf, 
but in some species the canals are median 9 . Transfusion- tracheids may form a 
ring round the phloem (A. magnifica). Cones large (in A. nobilis 25 x 10 cm.) 
and in most species the bract-scales are conspicuous (fig. 705, C, F). The cones 

1 Bailey (09). 2 Bommer, toe. cit. PI. ix. figs. 167, 168. 

3 For figures illustrating the range in form of cones, see Kirchner, Loew, and 
Schroter (06) pp. 152, 153. 

4 Schroter (97) figs. 24, 26. 

5 Barber (92). • Gard. Chron. June 3, 1893, p. 659. 

7 Kirchner, Loew, and Schroter (06) fig. 26, p. 89. 

8 Ibid. p. 90. 9 Elwes and Henry (06) p. 717. 






Fig. 705. A, B, Short shoot and cone of Pseudolarix Kaempferi. C, Abies bracteaia, 
showing the long bract-scales. D, E, Abies concolor, cone-scale; b, bract- 
scale; s, seed. F. Abies Fraseri, cone-scale ; 6, bract-scale. G, Larix Griffithi; 
b, bract scale; s, seed. (C — F, from the Gardeners'' Chronicle.) M. S. 

A M B 

Fig. 706. Branches of Abietineae. A, Abies pectinata. B, Tsuga canadensis. 
C, Pseudotsuga Douglasii. D, Picea excelsa. E, Cedrus Libani. F, Larix 
europaea. G. Pseudolarix Kaempferi. (After Rendle from Veitch and after 
Engler and Prantl.) 


of some species with concealed bract-scales, e.g., A. violacea 1 , A. Webbiana 2 , 

A. concolor (fig. 705, D, E), are hardly distinguishable from those of Picea anc 

the larger cones of Tsuga, while those of Abies amabilis resemble Cedar cones. 

PODOCARPINEAE. The wood agrees with that of the Cupressineae 


the bordered pits on the radial walls of the medullary rays in the region o 
the spring-wood are narrow, elliptical, and vertical (fig. 693, K), while in the 
Cupressineae the pore is horizontal (cf. fig. 693, H), a feature that is not 
sufficiently constant to afford a trustworthy criterion. Podocarpus. Leaf- 
lamina linear and short (P. nivalis, etc.) as in Taxus, longer and broader (in P. 
elatus reaching a length of 29 cm. and 2 cm. broad), as in Cephalotaxus (fig 
694, A) and the pinnae of Cycas small appressed and scale-like (P. cupressina 
or ovate and provided with several veins (P. Nagi, 6x2 cm., P. Wallichiana 
13 — 15 x 3-5 cm.). The leaves are opposite in the section Nageia (fig. 707) 
scattered in other species. Tison 4 has shown that in the broad-leaved species 
only one vascular bundle is given off from the stele and this branches in the 
petiole, not in the cortex as in Agathis. The leaf of P. maerophyUa* 
(5 — 6 cm. x 7 mm.) represents a fairly common type: short reticulate tracheids 
occur on each side of the vein and elongated tracheids extend from the midrib to 
the leaf -edge. There are three canals near the lower surface, but in some species 
only one is present. P. formose?isis 6 (fig. 707) has leaves like those of P. Nagi, 
but more rigid, thicker, and smaller; the epidermal cell-walls are thick and 
elongated and 4 — 5 cells surround each stoma (fig. 707, a, b). The presence 
of hypodermal mechanical tissue distinguishes Podocarpus leaves from those 
of Torreya, and another characteristic feature is afforded by the two kinds of 
accessory tracheids in place of the ordinary transfusion-tissue in the great ma- 
jority of leaves. Reference has already been made to the ' cones.' Dacrydium. 
The dimorphism of the foliage-shoots is illustrated in fig. 708. The very 
simple type of megastrobilus is a striking feature (fig. 684, P). Microcachrys. 
Leaves small, appressed, in decussate pairs 7 . As in Dacrydium there is 
a single canal below the vein. Gothan notices the frequent occurrence of a 
single large pit in the field of the rays. The fleshy mulberry-like cone is 
a peculiar feature. Saxegothaea (fig. 687). Lindley 8 , the author of the genus, 
describes this Conifer as having the male flowers of a Podocarp, the cones of 
an Agathis, the fruit of a Juniper, and the habit of Taxus. There is a canal 
below the vein, also lateral transfusion-tracheids but less numerous than in 
Podocarpus. The occurrence of pits in the horizontal walls of the ray cells 
is a character in which Saxegothaea differs from the genera comprised under 
the general term Cupressinoxylon 9 . The cones have already been described. 

1 Gard. Chron. Dec. 27, 1890. 2 Ibid. Oct, 3, 1891. 

3 Gothan (05) p. 47. * Tison (12) PL iv. fig. 2. 

5 Stiles (12). 6 Dummer (12). 

7 Hooker, W. J. Icones Plantarum, 1843. 

8 Veitch (00) p. 158; for figures, see Gard. Chron. June 22, 1889, p. 782. 

9 See p. 186. 






Fig. 707. Podocarjms formosensis. A, Foliage-shoot, a, b, Epidermis with 
stomata. (A, after Dummer; £ nat. size.) 




Acmopyle. This generic name was given to an imperfectly investigated Ne 1 
Caledonian Conifer formerly known as Dacrydium Pancheri. The sessile, 
decurrent, falcate leaves on the lateral branches (1 — 1-6 cm. x 2-5 mm.) are 
a characteristic feature, those on the main axis being small and scale-like 1 . 


A B 

Fig. 708. A, Dacrydium datum. B, Dacrydium araucarioides. (From specimen 
in the British Museum.) 

Pherosphaera. The vegetative shoots resemble those of Microcachrys and 
some Lycopods. The genus is peculiar among the Podocarpineae in the absence 
of an epimatium. Sinnott 2 regards the megastrobili, composed of 2 — 5 one- 
seeded scales, as 'the last step in reduction,' but the difficulty is to discriminate 
between simplicity as a primitive and as a reduction-phenomenon. Phyllo- 
cladus (fig. 675). The cladodes have a central rib and several vascular 

1 Pilger (03) p. 117. Mr Sahni (Emmanuel College, Cambridge) has in prepar- 
ation an account of Acmopyle based on material collected in New Caledonia by 
Mr Compton. 

2 Sinnott (13) p. 73; see also Masters (93) p. 6; Hooker. J. D., Icon. Plant., 1882, 
PL 1383; Baker and Smith (10). 



strands, and stomata occur on both surfaces 1 . The small megastrobili 
consist of spiral or decussate scales, each with one seed. Miss Young 2 mentions 
the occurrence of tracheids with spiral bands : a single large pit occupies the 
field of a ray-cell as in Dacrydium Franklini 3 . The seed in the early stage of 
development shows signs of a ruminated endosperm. 

TAXINEAE. The occurrence of spiral bands in the secondary tracheids 
is a family-character but similar tracheids are not uncommon in other Conifers. 
Taxus. The difference in leaf-form and in the arrangement of the foliage is 
well illustrated by Kirchner and Schroter 4 . The leaves have recurved margins, 
a mucronate acute apex, and a small median vein. Torreya. In habit this 
genus resembles Taxus but the leaves are longer (6 cm. in T. calif ornica) ; 
there is a resin-canal below the vein and two conspicuous stomatal grooves on 
the lower surface with papillose epidermal cells (fig. 694, B) : the midrib is not 
prominent. Cephalotaxus. Leaves linear, more or less falcate, reaching a 
length of 12 cm. in C. Henryi (fig. 709), with a prominent midrib and one canal 

Cephalotaxus Henryi. (British Museum; \ nat. size.) 

(fig. 694, A) ; there are no stomatal grooves. Seeds like those of Torreya in 
size and in the thick fleshy sarcotesta and inner shell ; the endosperm is not 
ruminated. Rothert 5 has described an interesting departure from the usual 
structure in the stem of C. koraina (according to Beissner = C. pedunculata 
var. fastigiata) : a resin-canal occupies the centre of the pith and several short 
tracheids occur internal to the edge of the xylem-cylinder. There do not 
appear to be any anatomical characters apart from the spiral bands of the 
secondary tracheids by which the Taxineae can be distinguished from some 
Cupressineae and other Conifers: xylem-parenchyma is said to be present 
in both spring and summer wood of Cephalotaxus drupacea, while it is unrepre- 
sented in Taxus cuspid ata and Torreya nucifera 6 ; the pits in the field vary 
in number and may be simple or bordered and in the latter case the pore is 
obliquely vertical. 

1 Robertson (06) ; Bernard (04) B. ; Stiles (12). 

2 Young, M. S. (10). 3 Gothan (05) p. 55. 

4 Kirchner, Loew, and Schroter (06) p. 69. 

5 Rothert (99). 6 Fujioka (13) 

S IV 11 




Sources of error in the determination of fossil Conifers. 

The determination of fossil Conifers is one of the most difficult 
tasks of the palaeobotanist. It is comparatively seldom that well- 
preserved cones are found in organic 
connexion with the twigs that bore 
them and the cones rarely exhibit those 
features which are the best guides to 
affinity. Excessive trust in superficial 
similarity has frequently led to the em- 
ployment of generic names suggesting 
relationships which are thoroughly mis- 
leading. In comparing fossil and recent 
forms authors are apt to confine their 
attention to the better-known types, 
forgetting that it is often with the less 
familiar and geographically restricted 
genera that extinct plants are most 
elosely allied. Even the data supplied 
by petrified wood are often insufficient 
to enable the student to do more than 
refer a specimen to some comprehen- 
sive genus based on characters shared 
by several recent genera. Though it 
is as a rule easy to distinguish between 
the wood of a Conifer and that of 
Cycads and Dicotyledons, the agree- 
ment between the xylem elements of 
many Cycads and those of the Arau- 
carineae is sufficiently close to afford 
opportunity for error. The homo- 
geneous structure of the secondary 
wood of some Magnoliaceous genera, 
e.g., Trochodendron and Drimys 1 (fig. 
710), closely simulates that of a Conifer, 
but the medullary rays are approxi- 
mately equal in breadth to the tra- 
cheids and the cells are more elongated 
1 Groppler (94); Solereder (99) p. 34; (08) p. 5 

Fir:. 710. Drimys Winteri. Trans- 
verse section of part of a stem. 
p, pith; /, pericycle fibres; 
c, cambium. 

See also Jeffrey and Cole (16). 




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 
D, Athrotaxis cupressoides , 

B, Callitris calcarata; C, Veronica 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 



arboreum ; Dacrydium Bidwillii 1 , D. Franklini and D. araucarioidei 
(fig. 708, B) recall some species of Lycopodium, and D. cupressinufn 2 , 
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 Agaihis 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 Taxus-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 oi 
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 laxifolium, D.Kirkii, D.elatium (fig.708 ? 
A), Juniperus chinensis, Araucaria excelsa, Callitris glauca, Podo- 
carpus cupressina 3 , 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 taxonomk 
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) Pis. xyiii., xix. 

3 Bennett and Brown (52) PI. x 




The 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 etude n'ouvre des perspec- 
tives plus etendues, mais aucune aussi n'exige plus de reserve et de 
tatonnements 1 .' 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 2 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. (15 2 ). 



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 1 . 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). 


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. Cioss-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 

I 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 
re 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. 

Fossil Gymnospermous wood (Coniferales). 

The earliest attempts to identify petrified wood are summarised 
y Goeppert 1 , Knowlton 2 , and other authors. Luidius (Lhwyd) 3 
t the end of the seventeenth century employed the general desig- 
nation Lithoxylon, and the termination -xylon 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 (89 2 ); Gothan (05). 3 Luidius (1699) A. 


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 1 adopted 
the method of adding the prefix Rhizo- to generic terms for wood 
believed to belong to roots, and Felix 2 and Lignier 3 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 o 
this kind. Barber 4 , in his critical paper on a species of Cupressino 
xylon, calls attention to Strasburger's description of an old mori 
bund stem of Larix with root-like characters and Gothan 5 speaks 
of a branch of Finns 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 6 and 
Witham whose work was rendered possible by methods of section 
cutting first employed, according to Nicol, by a Mr Sanderson 
lapidary. Opinions expressed by Nicol on methods of investi 
gating petrified wood are still pertinent aftei 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, i 
would be requisite to have a thorough knowledge of the structur 
at least of all the different tribes of recent Coniferae; and ye 
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 (07 2 ). 

4 Barber (98). 6 Gothan (05) p. 19. 6 Nicol (34) A. p. 141. 





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- 
menon 1 . The partial obliteration of bordered pits by decay may 
cause them to appear separate though originally in contact (fig. 
475, B, Vol. in. 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 2 . 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 3 . 

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 Antevs 4 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). 

3 For methods, see Jeffrey and Chrysler (06); Hollick and Jeffrey (09) B. 
Gothan (09); Sinnott (09). 

4 Antevs (17). 


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 1 
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 2 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 3 . A modified form of this scheme was pub- 
lished by Schenk 4 and more recently Penhallow 5 , Jeffrey, Lignier, 
Gothan, Dr Stopes, and other authors have considerably extended 
our knowledge. Dr Gothan 6 , 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). 2 Stopes (15) p. 247. See p. 419, Vol. m. 

3 Kraus in Schimper (72) A. p. 363. 

4 Schimper and Schenk (90) A. p. 860. 

5 Penhallow (07). 6 Gothan (05). 




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, 




Fig. 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 


of former ages : the petrified forests in the desert east of Cairo 1 
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 2 ; the succession of Tertiary forests in 2000 ft. 
of volcanic sediment exposed on the sides of Amethyst Mountain 
in the Yellowstone Park 3 (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 fitting on the tracheitis. 
I. DADOXYLON. (Including Araucarioxylon 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. 
Kims 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 fits are separate and circular and, if in two 
or more rows, offosite. Contiguous and more or less flattened fits 
occur Sfasmodically on the tracheids of the wood of the genera included 
in this section. Well-developed rims of Sanio usually occur on the 

1 Unger (59). 

2 A good example of an Arizona tree-trunk is exhibited in the Plant-Gallery of 
the British Museum. See Ward (00 3 ). 

3 Knowlton (99); Holmes (78); Seward (ll 3 ) p. 60, fig. 6. 


II. CUPRESSINOXYLON. (Including Glyptostroboxylon and 


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. 


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 Cupressinoxylon. 

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 
Cupressinoxylon. 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. 


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 


field. In the characters of .the normal wood this genus agrees 
most closely with Cupressinoxylon and Mesembrioxylon. 



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 

Medullary rays uniseriate ; all the walls are pitted ; tracheid 
may be present in the rays though they are usually absent. Ther 
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 Piceoxylon and Pinuxylon. 

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 o 
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. 




Similar to Pityoxylon 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. 



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. 



Similar to Woodworthia in the possession 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. 

XL 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 




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?). 


When uniseriate the pits on the tracheids are usually contiguous 
and flattened ; if biseriate the pits are opposite ; separate pits also 

True resin-canals absent, but canal-like spaces lined with small 
cells occur in some large medullary rays and constitute a charac- 
teristic featuie. 

Medullary rays uniseriate; pits confined to the radial walls; 
2 — 3 circular simple pits in the field. 




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 rilled with tyloses. 

Medullary rays uniseriate; all the walls pitted. 



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 Araucarioxylori]. 

The anatomical characters implied by the expression 'wood of 
the Araucarian type' are enumerated in the Chapter on Eecent 
Conifers, and in Chapter xxxiii. 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 1 . 
The name Araucarites was used by Goeppert 2 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 3 has been widely used, especially for Palaeozoic 
wood having the characters of Araucana or Cordaites. This 
designation leaves open the question of precise systematic position. 
In 1882 Kraus 4 instituted the genus Araucarioxylori, 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 Araucarioxylori for Mesozoic and Tertiary 

1 See p. 248, Vol. m. 2 Goeppert (45). 

3 Endlicher (47). 4 Kraus in Schimper (72) A. p. 370. 


wood has already been criticised x : 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 Araucarioxylon 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: Dadoxylon valdajolense (Moug.) 2 and 
D. Rhodeanum (Goepp.) 3 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 4 
subdivides wood of the Dadoxylon type into several genera in- 
cluding Pycnophyllum, to which is referred Dadoxylon Brandling ii 
(Lind. and Hutt.), Ullmannites applied to the type recently 
named Eristopliyton 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 Araucarioxylon may be added 
after Dadoxylon. Such a species as Dadoxylon permicum Merck. 5 
is one of many examples of a Dadoxylon that cannot be more pre- 
cisely identified. Dadoxylon australe Crie 6 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. Crie's species should not be confused with 

1 See p. 249, Vol. m. See also Gothan (05) p. 14; Potonie (02) p. 289. 

2 Fliche(03); Mougeot (52) A. 

3 Potonie (99) B. p. 294; Gothan (07) p. 17; Tuzson (09) p. 34. 

4 Tuzson (09) p. 17. 5 Mercklin (55) p. 53. 
6 Crie (89) p. 5, Pis. I., in., v. 

s. iv 12 



Dadoxylon australe Arber 1 based on Palaeozoic wood fro 
Australia : the substitution of Dadoxylon for Araucarioxylon, tl 
name used by Crie for his New Caledonian species, necessitates 
a new specific designation for Arber's type, which it is proposed 
to rename Dadoxylon Arberi. From Liassic beds in Yorkshire 
Miss Holden 2 has described a species as Araucarioxylon 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 Artisia 3 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 Xenoxylon* 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. It is, 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 5 
lays greater stress on the nature of the pitting on the walls of 
medullary cells, but Jeffrey 6 has discovered typical Abietineous 
ray cells in the cone-axis of Araucaria and Agathis. Miss Holden 7 
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 Araucarioxylon sp. 8 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 9 that the ray cells of such 

1 Arber (05) B. p. 191, text-figs. 40—43. 

2 Holden, R. (13 2 ) p. 540, PI. xl. fig. 28. 3 See p. 248, Vol. m. 

4 See page 248, Vol. m. 5 Gothan (05). « Jeffrey (12). 

7 Holden (14) (13 2 ) p. 544. See also Sifton (15). 

8 Holden (14) p. 171. 9 Thomson (13). 


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 1 
was instituted by Caspary for specimens distinguished from most 
examples of Dadoxylon by the presence of scattered xylem-paren- 
chyma but, as Gothan 2 points out, a distinctive name is super- 
fluous; the type-species of Araucariopsis, 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 general- 
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. Pis. xiv., xv. 

2 Gothan (10) p. 9. 



they afford with regard to the phylogeny of the Araucarineae an< 
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 let 
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 1 . 

In the following brief descriptions of species of Dadoxylon a fe^ 
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 xxxiii. 
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, ?n addition to Kraus and other authorities, Lignier (07 2 ). 


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 1 , from Franconia and Wiirtemberg, is considered 
by Schimper 2 on the ground of association to be the wood of 
a Voltzia, and Tuzson 3 adopts the generic name Pagiophyllites 
implying relationship with Pagiophyllum ; he includes in D. 
keuperianum the species Araucarioxylon wiirt ember gicum Kr. and 
A. thuringicum Born 4 . 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 5 from Pennsylvania : he records Araucarioxylon vir- 
ginianum, a species described by Knowlton 6 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. austrah (Crie). 

Dadoxylon septentrionale Gothan. 

This species 7 , 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. 2 Schimper (72) A. p. 384. 

3 Tuzson (09), p. 30, fig. 5. 4 Schimper (72) A. p. 384. 

5 Wherry (12). e Knowlton (89 2 ), PL vn. figs. 2—5. 

7 Gothan (10) p. 8, PI. i. figs. 4—8, PI. n. fig. 1. 


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 1 described this species, from Liassic strata in Madagascar, 
under the name Aran car ioxylon : 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 2 
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 (Eichwald), recorded from Moscow 3 , 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 zeelandiae* (Stopes). 

A Cretaceous species from Amuri Bluff, New Zealand 5 , founded 
on a piece of decorticated stem 8 cm. in diameter preserved partly i 
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 6 and other authors, afford particularly good 

1 Fliche (05) p. 350, PI. x. fig. 1. 

2 Lignier (07 2 ) p. 257, PI. xvn. figs. 10—13. 3 Eichwald (68) PI. v. fig. 12. 

4 The specific name is given by Dr Stopes as novae zeelandii. 

5 Stopes (14 2 ) PI. xx. 6 Thomson, R. B. (13). 


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 1 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 Araucarioxylon Ettingshauseni. 
A specimen of wood from Amuri described by Ettingshausen as 
Dammara Oweni 2 appears to be undoubtedly Araucarian. 

Dadoxylon sp. (Holden). 

An interesting type of stem-wood has been described by Miss 
Holden 3 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 Araucarioxylon. 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, Pis. n. figs. 1, 2; vi. figs. 10—12. 

2 Ibid, p. 16, PI. vi. figs. 13—15. 3 Holden, R. (13 2 ). 




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 1 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 2 and in 
Dadoxylon septentrionale Goth. The phloem 
includes well-preserved sieve-tubes and the 
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. 


Fig. 713. Dadoxylon (Ar- 
aucarioxylon ) bre veradi - 
atum, (After Lignier.) 

Among other Cretaceous species are Dadoxylon albianum 3 
(Fliche) from L'Aube; Dadoxylon Dantzii Pot. 4 from beds pro- 
bably of Upper Cretaceous age in East Africa, without definite rings 

1 Lignier (07 2 ) p. 290, PI. xix. 2 See page 179. 

3 Fliche (97) p 8, figs. 2—4. 4 Potonie (02) Pis. i., n. 




of growth and with a single row of contiguous pits on the tracheid 
walls; Dadoxylon virginianum (Knowlton) 1 from the Potomac 
series; Dadoxylon barremianum (Fliche) 2 from the Lower Creta- 
ceous of France; Dadoxylon noveboracense (Holl. and Jeff.) 3 from 
the Middle Cretaceous beds of Staten Island ; Dadoxylon Zuffardii 
Negri 4 from middle Cretaceous rocks in the Gulf of Tripoli ; Dado- 
xylon tankoense (Stopes and Fujii) 5 from Upper Cretaceous beds 
in Japan; Dadoxylon madagascariense (Fliche) 6 from Madagascar. 

Dadoxylon (Araucarioxylon) kerguelense sp. nov. 

In 1881 Goeppert 7 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 8 
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. 

Fig. 714. 

A B 

Dadoxylon (Araucarioxylon) kerguelense. (British Museum.) 

The sections on which this Kerguelen species is founded are in 
the British Museum 9 ; 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 2 ) PL TIL 

3 Hollick and Jeffrey (09) B. p. 58, PL xxi. 

5 Stopes and Fujii (10) PL m. figs. 17, 18. 

6 Fliche (00 2 ) p. 472, fig. 1. 

' Goeppert (81). 8 Gothan (08) p. 13. 

Fliche (00) p. 18. 
Negri (14). 

Sections V. 8388, V. 8390. 




tracheids, contiguous, alternate, and often slightly flattene( 
(fig. 714, B). The medullary rays have usually 5 — 8 elliptica 
pits in the field (fig. 714, A). 

Dadoxylon (Araucarioxylori) pseudoparenchymatosum Gothan. 

A species from Tertiary or possibly Upper Cretaceous rocks 
in Seymour Island (S. lat. 64° 16 'J 1 agreeing closely with the wooc 
of recent Araucarineae. The annual rings are distinct ; there are 
1 — 2 rows of pits on the tracheids 10 — 12 fi in diameter; the 
medullary rays, 2 — 10 cells deep, are usually uniseriate and there 
are several small oblique pits in the field. Cross-bars 2 (Miiller's 
'querbalken') like those described in Araucaria brasiliensis occur 
in some of the tracheids. 

Dadoxylon Doeringii Conwentz 3 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 4 , recorded from 
several localities in the Libyan desert; Dadoxylon Robertianum 
(Schenk) 5 of Tertiary or possibly Cretaceous age from the pro- 
vince of Nagpur, India; Araucarioxylon koreanum (Felix) 6 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 1 '. 


The name Cupressinoxylon 8 or, as written by Kraus, Cupresso- 
xylon 9 , is usually applied to fossil wood exhibiting the following 
features : — Annual rings well defined, often nairow ; 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 i. figs. 12—16. 

2 See page 135. 

4 Unger (59); Schenk (80) p. '3, Pis. i., n. 

5 Schenk (82 2 ). 

7 Gothan (10) p. 23. 

9 Kraus in Schimper (72) A. p. 374. 

Conwentz (85) p. 16. 

Felix (87) p. 518. 
Goeppert (50) p. 196. 


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. Gothan 1 , who uses Cupressinoxylon 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 2 states 
that 60 — 150 rays occur in 1 square millimetre, another feature 
of doubtful value. In some species included in Cupressinoxylon 
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 
4ype 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 2 ) p. 245. 




of the wood; xylem-parenchyma normally abundant and not 
restricted to the end of the year's growth. The photographs 

Fig. 715. Cupressinoxtjlon sp. Lough Neagh, Ireland; 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 1 in Ireland, probably of Upper Tertiary age, illustrate 

1 See vol. i. p. 80. 


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 1 on wood from Lough Neagh, 
the generic name being altered by Kraus to Cupressinoxyhn 2 . 

Kraus 3 enumerates 46 recent species of Conifers with wood of 
the Cupressinoxyhn type and Beust 4 mentions 37 species. As 
used by most authors Cupressinoxyhn 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 Cupressinoxyhn, e.g., Taxodioxylon 
(= Cupressinoxyhn Sequoianum Merck. 5 ), Podocarpoxyhn (= C. 
Hookeri Arb. 6 ), Xenoxyhn (= Cupressinoxyhn Barberi Sew. 7 ): 
these and other species formerly included in Cupressinoxyhn 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 . Hoi den . ] 

Miss Holden 8 has proposed a new generic name, Paracupressi- 
noxylon for wood which agrees with Cupressinoxyhn 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 

I 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). 

5 Mercklin (55) PL xvn. 6 Arber (04); Gothan (08) p. 7. 

• Seward (04) B. p. 60, PL vn. 

8 Holden, R. (13 2 ) p. 537; (14) p. 173. 


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 all important, implies the neglect of other 
characters, 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 Protocedroxylon and Thylloxylon. 

Cupressinoxylon liasinum Lignier. 

This Liassic species 1 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 

1 Lignier (07 2 ) p. 306, PI. xxi. figs. 58—61 ; PI. xxm. fig. 83. 




and rarely bordered, in the field. Kesin-cells are abundant. 
Lignier suggests that if the genus Cwpressinoxylon is subdivided 
this species might be referred to Glyptostroboxylon, though the 
medullary-ray pitting is not consistent with the characters of that 

Cwpressinoxylon vectense Barber. 

Founded on both stem and root wood from the Lower Green- 
sand of the Isle of Wight and described by Barber 1 with a thorough- 

Fio. 716. Cwpressinoxylon 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 2 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) Pis. xxm.— xxiv. 

2 Stopes (15) p. 169, text-fig. 50; also text-figs. 48, 49, and PI. xv. 


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. Kesin-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 irregularl 


Cupressinoxylon McGeei Knowlton. 

This is one of several species from the Potomac lignites included 
by Knowlton 1 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 2 has described some wood of 
Lower Cretaceous age 3 from King Charles Land as Cupressinoxylon, 
cf. C. McGeei, 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 of a 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 4 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. 

1 Knowlton (89 2 ) p. 46, PI. II. fig. 5; PI. in. figs. 1—5. 

2 Gothan (07 2 ) p. 19, fig. 10. 

3 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). 

4 Stopes (15) p. 180, text-figs. 51—53. 




Cupressinoxylon cryptomerioides Stopes. 

A species 1 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 

Fig. 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 2 . The cortex contains large 

1 Stopes (15) p. 186, Pis. xvi., xvn., text-figs. 54, 55. 

2 See page 198. 





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- 

Cupressinoxylon Hortii Stopes. 

This Lower Greensand species 1 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 th( 
majority of fossil woods within narrow limits. 

Cwpressinoxylon 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. Kesin-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 2 

» Stopes (15) p. 194, PL xvm. text-figs. 56—58. 2 Jeffrey (03). 


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 Thomp- 
son 1 . 

Cupressinoxylon 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 2 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-Harms worth Expedition. The species is named after 
Dr Koettlitz, the geologist of the Expedition 3 . 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. It is 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, PI. xlt. fig. 11. 3 Koettlitz (98). 





none on the tangential or horizontal walls 

broad; 2 — 4 small circular or oval (simple?) pits in the field, but 


* ... 


Fig. 718. A, D, Cupressinoxylon polyommatum. B, Phloem showing sieve-plates 
(Franz Josef Land). C, Cupressinoxylon Holdenae. E, Cupressinoxylon 
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 1 was founded was 
obtained from Banksland, from Miocene rocks, N. lat. 74° 40', 

1 Cramer (68) p. 172, Pis. xxxiv., xxxv-, xxxvu. 


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 irregulai 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/x; 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 Merkusii 1 , 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) Taxodii (fig. 720, A, B). 

Cupressinoxylon taxodioides Conwentz. 

Under this name Conwentz 2 describes some wood, probably of 
Pliocene age, from California which he compares with Sequoia 
semper virens. 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 (Glyptostrobusl) neosi- 
biricum 3 , 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 

1 Groom and Rushton (13) p. 484. 2 Conwentz (78) Pis. xin., xiv. 

3 Schmalhausen (90) PI. n. pp. 44 — 49; Gothan (05) p. 50. 




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 Beust 1 from Kerguelen 


[Glyptostroboxylon Conwentz.] 

This generic name was first employed by Conwentz 2 for woo 
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 3 that this species 
should be transferred to Podocarpoxylon, though in the medullary- 
ray pitting it differs from typical representatives of the Podocar- 
pineae. Kraus 4 described the pits in the radial 
walls of the ray cells of Glyptostrobus as large 
and circular and in a Tertiary specimen from 
Niederwollstadt, named Glyptostrobus tener Kr., 
he figures the pits in the field as simple (fig. 
719) ; these are said to be 1 — 8 in number and 
they are arranged in horizontal series, a feature 
characteristic of Taxodium. 

Sections of wood of Glyptostrobus hetero- 
phyllus which I have examined show 2 — 8 pits 
in the field but there is a distinct border and the 
pore is narrow and oblique or in the spring- wood 
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 







O [ 





Fig. 719. Cupressin- 
oxylon tener (Kraus). 
(.After Kraus.) 

Beust (85) PI. iv. 
Gothan (08) p. 9. 

2 Conwentz (85) p. 13. 

4 Kraus (64) p. 195, PI. v. fig. 12. 


that the pits in the region of the spring-wood may be simple. 
The same author 1 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. 


The generic name Taxodioxylon was applied to a Tertiary 
species from Hungary, originally referred by Felix 2 to Rhizotaxo- 
dioxylon, on the ground of a resemblance in structure to the wood 
of the recent genus Taxodium. Schenk 3 , who examined the type- 
specimen, confirmed this comparison. Taxodioxylon has been 
adopted for fossils agreeing with the wood of Taxodium and 
Sequoia sempervirens : Sequoia gigantea, on the other hand, agrees 
more closely with typical species of Cupressinoxylon. Taxodio- 
xylon, 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 

Gothan (05) p. 49. * Felix (84) p. 38. 

Schimper and Schenk (90) A. p. 872. 




horizontal rows is a characteristic feature. The characters of 
Taxodioxylon 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 
Taxodioxylon and this character has been quoted as a trustworthy 





O o o 
o o o 

o c 

j o a 

la o O 1 

O O o 

\° ° A- 

o a o o 

a O o j 


A B C 

Fig. 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 1 . In Glyptostrobus 
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. 

See page 149. 


Cupressinoxylon (Taxodioxylon) Taxodii Gothan. 

In his description of the Tertiary beds at Senftenberg Potonie 1 
identified some of the wood as Taxodium and compared the de- 
posits with those of a Taxodium-swsaonip. Gothan 2 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. 

Cupressinoxylon (Taxodioxylon) Sequoianum Mercklin. 

Gothan 3 recognised this species, originally described by Mercklin 4 
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. Taxodii; 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 
Cupressinoxylon uniradiatum Goepp. 5 from Briihl afterwards 
recorded by Conwentz 6 as Rhizocupressinoxylon from Tertiary beds 
in Silesia. Schmalhausen 7 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 Taxodioxylon type is afforded 
)y T. palustre described by Felix 8 from Tertiary rocks in Hungary 
and recorded also from sub-Oligocene beds in Silesia. Gothan 9 

1 Potonie (96); see also Eberdt (94). 2 Gothan (06) p. 164. 

3 Ibid. p. 165; (09) p. 518. 4 Mercklin (55) p. 65, PL xvn. 

5 Goeppert (50) p. 203, PL xxvu. 6 Conwentz (80) A. p. 25, Pis. iv., v. 

7 Schmalhausen (83) PL xn. 8 Felix (82) p. 278; (84) p. 38 

9 Gothan (10) pp. 40, 43, PL vn. 


assigns to Taxodioxyhn 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 Langsdorfii. 

The name Taxodioxyhn is retained as a section of Cupressino- 
xylon 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. 


Unger 1 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 2 . The 
name, in the form Taxoxylum, was substituted for Taxites em- 
ployed by Goeppert 3 for some species of Tertiary wood. Apart 
from the presence of spiral bands Taxoxylon 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 Taxites scalariformis, was renamed by Unger Taxoxylu 
Goepperti. According to Schenk 4 it is the only species among 
those recorded by Goeppert which should be retained in Taxoxylon, 
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 Taxites Ayckii (after Herr Aycke) of Ter- 
tiary age was retained by Kraus 5 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 6 also 

1 Unger (47) p. 33. 2 See page 134. 

3 Goeppert (40). 4 Schimper and Schenk (90) A. p. 859. 

5 Goeppert (40) p. 188; (50) p. 244; Kraus (64) p. 197. 

6 Lingelsheim (08) p. 27. 


states that there are no true spiral bands in Geoppert's supposed 
Taxoxylon. The Permian species Taxoxylon ginkgoides Ren. 1 and 
Grand' Eury's Upper Carboniferous species T. stephanense 2 are 
probably founded on wood of the Araucarian type in which the 
tracheids show spiral striation. 

An Aptian species Taxoxylon anglicum Stopes 3 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. 


This generic name 4 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 5 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, PI. xix. text -fig. 59. 

4 /j.e<rr)ix.Pptv6s, southern. 5 Stopes (15) p. 210. 


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 

Podocarpoxylon. This name 1 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 iows 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 aie 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 Podocarpoxylon and Cupressino- 
xylon, 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 

Phyllocladoxylon. This name was given by Gothan 2 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 aie 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 3 , but in Phyllocladoxylon the tracheal pits are often 

1 Gothan (05) p. 48 2 Ibid. p. 55; (10) p. 37. 3 See page 238. 


separate and are smaller than in Xenoxylon. The pits on the 
medullary-ray 'cells are also smaller, though it is doubtful if this 
is a constant character. Miss Holden 1 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 araucarioides 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- 
brioxylon (= 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 Phyllocladoxylon, 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 

I Phyllocladoxylon. If the Yorkshire stems are Araucarian so too are 
those from King Charles Land and Seymour Island 2 . In one of 
Miss Holden's species the tracheal pitting is not of the Araucarian 

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. Mesembrioxylon is applied to woods in which the general 

1 Holden, R. (13 2 ) p. 536, PI. xxxix. figs. 7—10. 2 Gothan (07 2 ); (08). 


, but 

features are similar to those associated with Cupressinoxylon, 
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. 

Mesembrioxylon sp. (= ? Podocarpoxylon sp. Gothan). 

This wood from Bathonian rocks in Russian Poland 1 affords 
an example of the difficulty of distinguishing clearly between 
Podocarpoxylon and Glyptostroboxylon : 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 2 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 
piriform, 1 — 2 in the field. The anatomical features described 
hardly afford adequate reasons for assigning the wood to Podo- 
carpoxylon 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 3 as 

1 Gothan (06 2 ) p. 456, fig. 5. 2 Holden, R. (13 2 ) p. 542, PL xl. figs. 31, 32. 

3 Thomas, H. H. (11) p. 80, PL v. figs. 5—7. 



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 /a in diameter; the medullary rays 
have one or rarely two large simple pits in the field. 

Mesembrioxylon woburnense (St opes). 

An Aptian (Lower Greensand) species from Bedfordshire 
founded on two blocks of secondary wood and referred to Podo- 
carpoxylon 1 . 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 2 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- 

I 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 Cedroxylon and 
in some other genera. 
Mesembrioxylon Gothani (Stopes). 
Dr Stopes regards this species 3 , 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, PI. xxi. text-fig. 64. 

3 Ibid. p. 228, text-figs. 65, 66. 




Fig. 721. Mesembrioxylon bedfordense. Radial section showing groups of adjacent 
pits on the tracheids, t; the medullary-ray pits, m, and a few with a border, p; 
n, narrow part of a tracheid. (After Stopes.) 




Mesembrioxylon Schivendae (Kubart). 

This species is recorded from Attersee in Upper Austria and 
though probably of Tertiary age it may be derived from Cretaceous 
strata 1 . 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 — 3 pits (fig. 722, A, B) but occasionally as many as 5 in the field ; 










<;. 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 Podocar poxy Ion, 
from Seymour Island 2 there is no xylem-parenchyma ; the medul- 
lary-ray cells have 1 — 2 elliptical-circular pits in the field and the 
I major axis is oblique. The evidence as to affinity is far from con- 

1 Kubart (ll 2 ) PL m. text-figs. 1 

2 Gothan (08) p. 8, PL I. figs. 9—11 


S. IV 




Mesembrioxylon sp. (Gothan). 

Gothan 1 described some wood as Phyllocladoxylon sp. whic 
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 fi high, and smaller than in Xenoxylon; 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 2 , the bordeied 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- 
xylon are usually more horizontally stretched as in Phyllocladus 
and some species of Pinus. Gothan compares Cupressinoxylon 
Hookeri Arb. 3 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 4 , 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 present day, an 
inference which, though not improbably correct, rests on slender 
evidence in this particular case. 

Mesembrioxylon Mulleri (Schenk). 

This species from Pliocene strata in New South Wales, was 
described by Schenk 5 as Phyllocladus: the pits on the tracheids 

1 Gothan (07 2 ) p. 9, fig. 2; (08) p. 6 (footnote). 

2 Ibid. (08) p. 4, PL i. figs. 4—8. 3 Arber (04). 4 Gothan (08) p. 25. 
5 Schenk in Schimper and Schenk (90) A. p. 873, figs. 424, 425. 


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. 

1 Mesembrioxylon Hooker i (Arber) (= Cwpressinoxylon Hooker i 
A species 1 founded on a splendid specimen of silicified wood 
nearly 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 medullarv 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. 

Arber (04). 

H— 2 


It has been suggested by Gothan 1 that this species should be 
referred to Podocarpoxylon : the pitting of the medullary-ray cells, 
though seldom preserved, seems to differ from the typical Cupress 
inoxylon form. It is therefore referred though with some hesitation 
to Mesembrioxylon. 


Paracedroxylon scituatense Sinnott. The generic name Para- 
cedroxylon 2 was instituted for this Cretaceous species from Scituate 
Massachusetts, in order to indicate its resemblance to Cedroxylon 
as defined byJKraus, 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 o: 
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 tracheid 
have a single row of bordered pits, generally circular. The medul 
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 Cwpressinoxylon and it is not distinguished 
by any well-marked features from Mesembrioxylon. 

Jeffrey 3 has described the axis of a Geinitzia cone from the 

1 Gothan (08) p. 7. 2 Sinnott (09). 3 Jeffrey (11). 


Vtataram formation as exhibiting the features of Parace- 


This generic name was instituted 1 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 2 , 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 Eleoxylon 3 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 4 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 5 speaks of the absence of parenchyma in some species of 
Cedroxylon. Gothan 6 , 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 presence 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 

Kraus in Schimper (72) A. p. 370. 2 Schimper and Schenk (90) A. p. 862. 

Brongniart (49) A. p. 76. 4 Barber (98) p. 332. 

Lignier (07 2 ) p. 245. ■ Gothan (05) p. 45. 


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, Pseudolarix, 
while in wood of the Cwpressinoxylon 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 medullary rays 
per square millimetre in Cwpressinoxylon. 

The characters of Cedroxylon may be briefly summarised a 
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 sometime 
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 1 and 
from Lower Cretaceous beds in King Charles Land 2 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. 

2 Ibid. (07 2 ) p. 26, figs. 14, 15; PL i. fig. 1. 

3 Groom and Rushton (13) PL xxv. figs. 47, 48. 






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. 

Fig. 72.'}. A, 

B, Cedroxylon transiens. 
Gothan; C, 

C, Cedroxylon blevillense. 
after Lignier.) 

(A, B, after 

I Wood of similar type was described by Schroeter 1 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 Hornei Seward and Bancroft. 

An Upper Jurassic species from Helmsdale 2 , Sutherland, a 
locality from which Hugh Miller lecorded 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), PI. I. figs 1—8. 

■ Seward and Bancroft (13) p. 883, text-fig. 5; PL n. figs. 22- 






and uniseriate ; there are 2 — 4 simple or faintly bordered circular 
pits in the field and pits occur also on the tangential and horizontal 



@ o \\ 


! ' 


I ° 



Fig. 724. A, Brachyphyllum eathiense; stoma. B — F, CedroxyJon Hornei. 
(After Seward and Bancroft.) 

Cedroxylon cedroides Gothan. 

In this species from King Charles Land 1 , 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 (07 2 ) p. 23, figs. 11—13. 


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, Pseudolarix, and Tsuga; the 
Abietineous features are relatively more conspicuous than in 
Cedroxylon transiens in which the tracheal pitting is more Arau- 

Cedroxylon maidstonense St opes. 

In this wood 1 , 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 
'ay cells. 

Cedroxylon pottoniense Stopes. 

This species 2 of the same geological age, from Potton in Bedford- 
shire, differs from C. maidstonense in the comparative abundance 

>f xylem-parenchyma : the medullary-ray cells show very clearly 
;he Abietineous type of pitting. 

Cedroxylon blevillense Lignier. 

In this species from the gault of Bleville (Seine-Inferieure) 3 
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 

ire small, numerous, and have an oblique pore (fig. 723, C). The 
tangential walls of the ray cells are sometimes pitted. There is 

10 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 xn. text-figs. 41—43. 2 Ibid. p. 154, text-fig. 44. 

3 Lignier (07 2 ) p. 267, PL xvm. figs. 15—17; PL xxi. fig. 66; PL xxn. fig. 72; 
•1. xxiii. fig. 87. 


A Triassic species described by Wherry 1 as Brachyoxylon penn- 
sylvanicum may, as that author suggested, belong to Cedroxylon: 
the tracheids have 1 — 2 rows of pits, usually separate but sometimes 
alternate and hexagonal as in Cedroxylon transiens and C. Homei ; 
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. Crie 2 describes a 
species, C. auslrale, from the Trias of New Caledonia though 
his figures and descriptions do not afford satisfactory evidence 
in support of this reference. Schenk 3 mentions Cedroxylon 
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 
Mercklin 4 from Jurassic rocks of Russia, Pinites jarassicus, 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 5 as 
Cormovedroxylon jurense from the Braun Jura of Galicia is com- 
pared by him with Pinites jurensis. Fliche 6 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 7 from the Lower Cretaceous of 
Haute Marne and a Cenomanian species C. manekildense Fliche 8 , 
but in neither case are the data adequate. 

C. matsumurae Stopes and Fujii 9 is an Upper Cretaceous 
Japanese species with 1 — 2 rows of tracheid-pits, generally opposite 

1 Wherry (12) PI. iv. 2 Crie (89) Pis. n.— v. 

3 Schimper and Schenk (90) A. p. 871. 

4 Mercklin (55) p. 48, PL vm. figs. G— 10. 5 Felix (82) p. 264. 
,! Fliche (97) p. 7. 7 Ibid. (00) PL n. fig. 1. 

8 Ibid. (96) PL xv. fig. 3. 

9 Stopes and Fujii (10) p. 42, PL i. fig. 10; PL iv. figs. 20—23. 


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 

Among Tertiary species reference may be made to Cedroxylon 
affine Kraus 2 from Sicily, without resin-parenchyma and charac- 
terised by usually two large simple pits in the field ; C. Hoheneggeri 
Felix 3 from the Eocene of Moravia figured by Schenk as from 
Cretaceous strata ; C. Hermanni Sch. 4 , an incompletely described 
species from Assam, probably of Tertiary age. 


Kraus 5 included in this genus some of the species previously 
referred by Goeppert to Pinites ; others he assigned to Cedroxylon. 
Pityoxylon is distinguished from Cwpressinoxylon 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 6 makes 
use of two generic names, Piceoxylon 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 

Stopes and Fujii (10) PI. iv. figs. 24—26. 

Kraus (83). 3 Felix (82) p. 268. 

Schenk (82 2 ) p. 355. 5 Kraus in Schimper (72) A. p. 377. 

Gothan (05) p. 102. 


pitting in the ray cells, and by the occurrence of numerous pits in 
the tangential walls of the summer tracheids. 

Pinuxyhn is used by Gothan in preference to Pinoxylon, the 
name adopted by Knowlton 1 for wood in which there are no resin- 
canals in the medullary rays. In Pinuxyhn 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 2 follows Jeffrey and Chrysler 3 in regarding Gothan' s 
twofold division as unnecessary. 

In the majority of species referred to Pityoxylon 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 4 , who follow previous 
authors in quoting Pinites Conwentzianus Goepp. 5 , 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, Pityoxylon chasense 
Pen. 6 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. Freeh 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. 

5 Goeppert and Stenzel (88) p. 54, Pis. xi., xn. 
• Penhallow (00) p. 76; Thomson and Allin (12). 


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 Dadoxylon 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 1 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 2 Pinites eiggensis 
and afterwards 3 referred by him to the genus Pence 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 Eigg in the Inner Hebrides, 
indley and Hutton 4 and Nicol 5 also gave short descriptions of the 
structure and Miller 6 in the Cruise of the Betsey alludes to a fossil 
trunk as 'an ancient tree of the Oolites.' The wood occurs with 

Ifragmental 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 7 . 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 
n 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. Pis. 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 
irlier conclusions of Sir Archibald Geikie as to the geological history of Eigg and 
;ives references to previous notices of the fossil wood; Seward (ll 2 ) p. 652. 



from the same source as the Dadoxylon wood. Though probably 
Jurassic, a Tertiary source is by no means ruled out. 

Kraus 1 transferred Witham's species to the genus Pityoxylon 
and that name is used by Schroeter 2 and Schenk 3 . The specimens 
on which the following account is based are most of them in the 
British Museum. Annual rings clearly denned, usually 1 — 1*5 mm. 

Fig. 725. Pityoxylon eiggenst; 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 

1 Kraus in Schimper (72) A. p. 378. 2 Schroeter (80) p. 13. 

3 Schimper and Schenk (90) A. pp. 855, 874. 


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 hoiizontal 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 1 . 

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 2 . 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 3 is founded on a specimen of wood obtained by 
Mr Kufford from Wealden beds at Ecclesbourne on the Sussex 

1 I 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, Pis. n. } in. 



I thft 

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 

Fig. 726. Pityoxylon Sewardi. Radial section showing tracheids, tr, in a medullary 
ray; c, 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 


opposite but stellate groups also occur as in Cedroxylon transiens 
Goth. j(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 1 
(Conw.) from the Lower Cretaceous of Sweden. 

Pityoxylon Seviardi Stopes. 

This species 2 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 beais a 
striking resemblance to the recent species Pinus monticola 3 . 

Pityoxylon Benstedi Stopes. 

In this Lower Greensand species 4 from Kent the resin-canals 
often contain tyloses as in P. Nathorsti Conw. and the epithelial 
cells have very thick walls, a feature suggesting comparison with 
the genus Larix. 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 Larix. 

Pityoxylon statenense Jeffrey and Chrysler. 

A species from the Middle Cretaceous of Staten Island 5 found 
in association with the short shoots described by Jeffrey and 

1 Conwentz (92) p. 13, Pis. i. — m., vi., vii. 

2 Stopes (15) p. 95, Pis. iv., v. text-figs. 23, 24. 3 Ibid, text-fig. 25, p. 103. 

4 Ibid. (15) p. 105, Pis. v.— vii. text-figs. 26, 27. 

5 Jeffrey and Chrysler (06). 

s. iv 15 




Hollick as Finns 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 specie 
of Pinus; xylem-parenchyma confined to the periphery of the 

Fig. 727. Pityoxylon Benstedi. Radial section showing the tracheal pitting, tr; 
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 


medullary rays occur, the latter with horizontal resin-ducts. There 
are no ray-tracheids. There is usually one circular or elliptical 
pit in 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 of ray- 
tracheids are regarded by the authors of the species as ancestral 

Pityoxylon protoscleropitys (Holden). 

A Middle Cretaceous species 1 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. Kims 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. 

I The presence of horizontal tracheids in the medullary rays is 

an important character: in Pityoxylon scituatensiformis (Bailey) 2 , 
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 
haracters of a modern hard Pine, yet retaining certain ancestral 

1 Holden (13 3 ). 2 Bailey (11). 




features, as the association of primary and fascicular leaves, the 
latter borne on brachyblasts subtended by a foliar trace.' 

Miss Holden 1 has also described from New Jersey two species 
which she compares with Prepinus, namely Pityoxylon foliosum 
and P. anomalum. 

Pityoxylon Nathorsti (Conwentz). 

Under the name Pinus Nathorsti Conwentz 2 described 
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 lesin-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 3 . The 
medullary aie 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 

Pityoxylon zezoense (Suzuki). 

This Upper Cretaceous Japanese species was described as 
Abiocaulis zezoensis i , 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 3 ). 2 Conwentz (92) p. 13, Pis. I.— in., vi., vn. 

3 Brooks and Sharpies (14). 4 Suzuki (10). 




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 Piceoxylon Pseudotsugae 1 : it is interesting 
as a type of Pityoxylon 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 Piceoxylon. Bailey 2 points out that in 
the absence of spiral bands in the ray-tracheids the fossil species 
resembles Pseudotsuga Douglasii, while in Pseudotsuga macrocarpa 
the tracheids of the rays have spiral bands. 

Fritel and Viguier 3 have described a species from Eocene beds 
in the Paris Basin as Piceoxylon Gotham 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 MacClurii (?) Heer 5 ; 
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. pidchrum. Annual rings well marked; 

1 Gothan (06 3 ). 

3 Fritel and Viguier (11) p. 63. 

4 Cramer (68) p. 171, PL xxxiv 

5 Ibid. Pis. xxxv, xxxvi. 

fig. 1 

2 Bailey (09) p. 54. 
PI. xxxvi figs. 6—8. 



bordered pits large, approximately 25 /a 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. 1 from the Braunkohle of Rheinland, so 
named from its resemblance to the North American Finns Parrya, 
characterised by horizontal tracheids with smooth walls and thin- 
walled epithelial cells ; also Pity oxylon pineoides Kraus 2 a Sicilian 
Tertiary species without ray-tracheids. 

Pity oxylon succinifer (Goeppert). 

This species from the Oligocene amber beds of the Baltic coast 
was first named Pinites succinifer 3 and several years later fully 
described and admirably illustrated as Pinus succinifer*. 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 5 , a feature occasionally seen in recent 
wood. The medullary raj^s 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, PI. i. figs. 1—3. 

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. 


rays are said to consist exclusively of tracheal tissue 1 . 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 2 and Felix 3 , 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, 
Pityoxylon 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 hoiizontal resin-canals is clearly shown. 


Protopiceoxylon exstinctum Gothan. The generic name Proto- 
piceoxylon was proposed for some Lower Cretaceous wood from 
King Charles Land 4 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. exstinctum are also included specimens from Spitz bergen 5 
and some of the material on which Cramer 6 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. PI. ix. fig. 2. 

2 Knowlton (99) p. 763, Pis. cvi. — cviii., cxn. — cxv., cxviii., cxix. 

3 Felix (96) p. 254. 4 Gothan (07 2 ) p. 32, figs. 16, 17; PI. i. figs. 2—6. 


Proiopiceoxylon 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 aie 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 Anomaloxylon 
but in that genus there is no Abietineous pitting on the medullary- 
ray cells; similar canals are described by Jeffrey 1 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 Piceoxylon and 
Pinuxylon 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 

A species from the Black Hills described by Knowlton 2 as 
Pinoxylon dacotense agrees with Protopiceoxylon 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. 

Proiopiceoxylon 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). 2 Knowlton in Ward (00) B. p. 420, PI. clxxix. 





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 

Fig. 728. Protopiceoxylon arcticum. 
(Cambridge Botany School.) 

Fig. 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- 
piceoxylon 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 



Protopiceoxylon Edwardsi Stopes. 

Founded on a branch from the Lower Greensand of Sussex, 
showing 17 annual rings, having the following characters 1 : 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- waMed 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. 


Woodworthia arizonica Jeffrey. This genus 2 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 Araacariopitys. 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 oi 
the lateral walls. 

On the surface of the wood are several small scars and a fe^ 
larger ones, the former representing short shoots subtended by 
leaf -trace ; the shoots are not infrequently branched as they pass 
through the secondary xylem, a feature recorded also in Ginkgo 3 . 
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 in. text-figs. 17—22. 

2 Jeffrey (10 2 ), Pis. xxxi., xxxn. 3 Tupper (11). 


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 Araa- 
cariopitys 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 piobability 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. 

A genus founded by Jeffrey 1 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 Jeffrey. 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 
resent 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) Pis. xxvm — xxx. 


It is suggested by Jeffrey that Araucariopitys may be the stem 
of Heer's genus Czekanowskia 1 , 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 2 . I 
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 €>f an Abietineous alliance, the tendency towards an Arau 
carian pitting of the tracheids being reminiscent of ancestra 
types in which that character was more pronounced. 


Protocedroxylon araucarioides. Gothan. The type-species of 
the genus founded on Upper Jurassic wood 3 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 characteis 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 4 and Miss 
Holden 5 . 

1 See page 63. 2 See also Gothan (10) p. 30. 

3 Gothan (10) p. 27, Pis. v., vi. 4 Jeffrey (12) p. 533, PL i. figs, a, b. 

5 Holden (13 2 ) p. 538, PL xl. figs. 17—21. 



Annual rings well marked ; boidered pits on the radial walls of 
the tracheids in 1 — 3 rows; in the Spitzbergen wood these are 
from 20 to 24 /x 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* 1 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 1 , 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 necessarily 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 lelationship. 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 2 under the generic name Metacedroxylon from Coral- 
lian beds on the Sutherland coast of Scotland, is founded on a piece 


of stem showing 75 rings of growth. There are no resin-canal 
and no xylem-parenchyma ; the tracheids have uniseriate bordere 
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 ; th 
pitting is of the Abietineous type. In the abundance of tylos- 
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 medullaiy 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 Paronai 1 . 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 legard to the relative value 
of tracheal or medullary-ray pitting whether Protocedroxylon 
should be placed neaier to the Araucarineae or to the Abietineae. 
Miss Holden 2 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 Conifer.' 


Gothan 3 instituted this generic name for some Upper Jurassic 
wood, originally described by Cramer 4 from Green Harbour, 

1 Negri (14) p. 340, PL v. figs. 1—6; PL vi. figs. 1—3. 

2 Holden, R. (14) p. 538 3 Gothan (05) p. 38. 
4 Cramer (68) PL xl. ; Schroeter (80) p. 7. 


Spitzbergen, as Pinites latiforosus 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 1 has 
suggested a possible connexion between Elatides and Xenoxylon. 
Xenoxylon would seem to have been widely distributed in later 
Jurassic floras. 

Xenoxylon conchylianum Fliche. 

Fliche 2 refers to Xenoxylon a piece of wood from the Muschel- 
kalk of the Vosges chaiacterised 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. Fliche 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 xxm. figs. 4—5. 



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 1 enabled him to confirm the main points of 
the original description: he regards Ciamer's species Piniles 
pauciporosus as identical with the type-species with which he also 
identifies Araucarioxylon koreanum Felix 2 . Xenoxylon latiporosum 
is characterised by the large size of the pits on the radial walls of 

Fig. 730. A, Xenoxylon phyllocladoides. B, C, Xenoxylon latiporosum. 
(A, after Gothan; B, C, after Cramer.) 

the tracheids, 20 — 40 fx broad and 15 — 20 \x 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, Pis. iv., v. 

Felix (87) PL xxv. fig. I. 


parenchyma but tylose-like cross- walls occur in some of the xylem 

The nature of the pitting led Kraus 1 to include Cramer's 
species in Araucarioxylon and Miss Holden 2 , who records this 
species from the Yorkshire coast, regards the absence of Sanio's 
rims as evidence of Araucarian affinity. The medullary-ray 
pitting is, however, very different from that in recent Araucarineae 
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 3 , 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 4 as Araucarioxylon latiporosum 
(Cram.) and, with some hesitation, Cupressinoxylon Barberi Sew. 5 
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 30 /x to 24 x 36 /x; 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 6 , Yorkshire, and Germany. 

Though similar to Araucaria and Agathis in the flattened con- 
tiguous pits, Xenoxylon 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. 7 and pits of similar form occur in the wood of 
the recent Magnoliaceous plant Drimys Winteri 8 . 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 (06 2 ) p. 454, fig. 4; (10) p. 36, PI. vi. figs. 9, 10, etc. 

4 Conwentz (82) p. 170 

5 Seward (04) B. PL vii.; Holden, R. (14) p. 535. 

6 Gothan (08 2 ) p. 10, figs. 3—9. 

7 Felix (86) A. PL v. fig. 4. 8 Groppler (94) Pis. i., it. 

iv 16 




(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.) 1 , 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 Araucarineac 
and Taxaceae. 


Anomaloxylon magnoradiatum Gothan. Gothan proposed this 
name for some Upper Jurassic wood from Spitz- 
bergen 2 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 
or 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 
rows, not alternate. The medullary rays are 
uniseriate, generally 5 — 7 cells deep, or in places 
forming broad and deep spindle-shaped areas 
either empty or containing a large circular canal- 
like passage. These peculiar rays, as seen in a 


ray showing the 
small cells and a 
resin-canal. (After 

tangential section of the wood, are a conspicuous feature and are 
1 See page 203. 2 Gothan (10) p. 10, PI. L figs. 9— 11 ; PI. n. figs. 2, 3. 


often lined by a layer of small cells (fig. 731). In the presence of 
these large medullary rays Anomaloxylon resembles Thylloxylon, 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 1 
than to any other family Anomaloxylon is a type which cannot 
be assigned to a definite position. 


Thylloxylon irregular -e Gothan. The generic name Thylloxylon 
was given to a single species of Upper Jurassic age from Spitz- 
bergen 2 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 f — 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- 
xylon, 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, PI. vi. figs. 2—8. 




Dr Marie Stopes 1 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 
Araucarioxylon Lindleii (Witham). The genus is a striking ex- 
ample of a combination of Araucarian and Abietineous characters 2 , 
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 
lings 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 ofl 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. 

Planoxylon Lindleii (Witham). 

This Liassic species from Whitby was originally referred by 
Witham 3 to the genus Pence ; subsequently included in Araucario- 
xylon 4 ' it has recently been transferred by Dr Stopes to her new 
genus Planoxylon 5 . 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) 

2 Tr\av do /, 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, Pis. ix., xv. 4 Seward (04) B. p. 56, Pis. vi., vn. 
5 Stopes (16) pp. 118, 120, text figs. 6, 7. 





Shoots, Cone-scales, etc. 

The 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 Coniferales 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, 
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- 
orthy evidence of relationship to existing types. 



PROTODAMMARA. Hollick and Jeffrey. 

The distinctive characters of the recent genus Agathis {Dammar a) 
re briefly described in Chapter XLIII. Fossil records bearing on 
he past history of Agathis are more meagre and more difficult of 

interpretation than those relating to Araucaria. The evidence 
t present available points to the greater antiquity of Araucaria 

at least as regards the type of cone characteristic of that genus. 


On the other hand the type of foliage-shoot represented by existing 
species of Agathis — without taking into account the Palaeozoic 
leaves assigned to Gordaites, 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 1 . 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 Fric and Bayer 2 as Dammaro 
phyllum striatum and D. bohemicum exhibit a close agreement in 
shape and venation with those of some species of Agathis, thoug 
they differ but slightly from some forms of Podozamites, e.g 
P. Reinii Geyl. (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 3 : these, prob- 
ably specifically identical, forms present, as Lesquereux says, a close 
resemblance to Agathis robusta. It is 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 Podozamites is essentially a Rhaetic and Jurassic 
genus. The leaves figured by Saporta 4 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 5 from the Cretaceous of Long Island as 
1 See page 447. 2 Fric and Bayer (01) B. p. 96. 

3 Lesquereux (91) p. 32, PI. t figs. 9—11. 

4 Saporta (94) B. PI. xxxm. fig. 5; PL xxxv. fig 12. 

5 Hollick (12) PJ. 163, figs. 2, 3. 



Podozamites lanceolatus the term Desmiophyllum would be a more 
appropriate generic designation, the name Dammarites being 
adopted for broader foLms. 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 1 points 
out Velenovsky's Tertiary species Podozamites miocenicus may be 
a leaf of Agathis or possibly a Podocarpus. 

Dammarites Bayeri 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 marginatum from the Cenomanian of Greenland but wisely 
excluded by Zeiller 2 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 3 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 disjancta 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. univervis 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, e.g., 
Dammarites albens Presl. 4 from the Quadersandstein of Bohemia 
and D. crassipes Goepp. 5 These two species are united by 

1 Schimper and Schenk (90) A. p. 279. 

2 Zeiller (05 2 ) p. 17, PI. vn. figs. 8—11. 

3 Ettingshausen (87) p. 15, PL I. figs. 20—24. 

4 Sternberg (38) A. PL lii. ; Corda in Reuss (46) B. PL xlviii. ; Goeppert (50) 
p. 237 ; Schimper and Schenk (90) A. p. 279, fig. 292 b 

5 Goeppert (50) PL xlv. fig. 6; Corda in Reuss (46) B. 




Velenovsky under a single type which he calls Krannera mirabilis 1 
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 similai 
to tuberous Cycadean species but it is doubtful if they were pro- 
vided with leaves of the type included in Krannera mirabilis. 
Schmalhausen 2 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 3 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- 
scales obtained from Cretaceous 
strata in Greenland and some 
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 4 , but it was Heer 5 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. microlevis, etc. 

Fig. 732. Dammar ites Tolli. (After 
Schmalhausen; nat. size.) 

1 Velenovsky (85) B. p. 1, Pis. i., iv. 
3 Ibid. (90) p. 14, PI. l. fig. 19. 

2 Schmalhausen (83) p. 313, PL xxxvi. 
4 Hollick (06) p. 38. 5 Heer (82). 


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 1 . The larger forms may con- 
veniently be included in Dammarites borealis Heer and smaller 
forms are illustrated by Protodammara speciosa Holl. and Jeff. 2 
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 
resinous material. Other Greenland ex- Fl °' 7 f- A™"" b °- 

reahs. Cone-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, Museum; nat " size - } 
page 323. Fossils of similar form were described by Heer from 
the same locality as Eucalyptus Geinitzii 3 . Krasser 4 and some 
other writers have retained the generic name Eucalyptus on the 
ground of association with Eucalyptus-like leaves. Hollick speaks 
f scales like D. borealis as ' among the most abundant and charac- 
teristic remains found in the Cretaceous deposits of America and 
IEurope 5 ': 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 6 , 
and anatomical evidence derived from Protodammara supports the 
view that the cone-scales were rich in resinous substance. Both 

1 Hollick (97) PI. xi. figs. 5—8; (06) p. 37; Newberry and Hollick (95) p. 46. 

2 Hollick and Jeffrey (06) p. 199, PI. I. figs. 5—13; PI. n. figs. 1—5. 

3 Heer (82) p. 93. 

4 Krasser (96) B. PI. xvi. fig. 6. 

5 Velenovsky (89) PL I. figs. 28, 29. 

6 Newberry and Hollick (95) p. 47 and see also Berry (07). 


large and small cone-scales are recorded by White 1 , Berry 2 , 
Knowlton, and especially by Hollick from several places on the 
Atlantic Coastal plain. Knowlton's species Dammarites acicu- 
laris 3 is probably identical with D. borealis. As examples of 
smaller forms reference may be made to D. northportensis*, D. 
minor 5 , and the scales now included in Protodammara. Thes 
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 b 
Hollick and Jeffrey afford strong arguments in favour of a 
affinity to Agathis. We have no proof as to the nature of th 
vegetative shoots of the parent-plants. Newberry states that i 
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 Geinitzii, with 
dicotyledonous leaves as evidence of original connexion. 



Protodammara speciosa Hollick and Jeffrey. 

It is probable that these scales 6 are generically identical witl 
the larger forms referred to D. borealis and other species, e.g. 
D. cliffwoodensis 7 , but until anatomical evidence is obtained a 
distinction should be recognised. The type-specimens were found 
at KreiscTierville, 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, E, 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 xlviii. figs. 8—11. 

3 Knowlton (05) PI. xv. figs. 2—5. 

4 Hollick (04) p. 405, PI. lxx. figs. 1, 2. 5 Ibid. (12) PI. n. figs. 35—37. 

6 Hollick and Jeffrey (06); (09) B. p. 46, Pis. iv., x., xiv.— xvi. 

7 Hollick (97) PI, xi. figs 5—8. 


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 1 (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 Protodamtnara 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 Protodamtnara . The American scales occur in association with 
shoots of the type represented by Brachyphyllum macrocarpum 
Newb. 2 (=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 
macrocarp inn 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. 



Conites 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 1 : 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; «, seed. 
B, forma a; s, spaces, probably resin-canals, filled with crystalline materials 
and simulating seeds. C, forma /3; ab, ridge. 

forma a, j3, y, and 8 (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 

1 Seward and Bancroft (13) p. 873, PI. I. figs. 9—12; PI. n. figs. 14—21. 




by Hollick and Jeffrey to Protodammara. The parenchymatous 
tissue of the cone-scales contains several thick-walled idioblasts 
and the 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 

Fig. 735. Conites Juddi, forma a; cone-scale in median section; I, ligule; s, s, 
(?) resinous material; vb, vascular bundles. (x6.) 

is a ligule on the abaxial side of the seed or seeds (fig. 736, B, I). 
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. 
scale from the specimen represented in fig. 734, B shows 






in longitudinal section (fig. 735) a large cavity in the lower part o 
the scale containing at each end a dark patch of some secrete 
substance s, s ; above this is a vascular strand vb, extending into 
the distal end of the scale near which is a ligular outgrowth I, and 
below this is a depression on the upper face of the scale in which 

Fig. 736. Conites Juddi, forma 8. A, Longitudinal section of cone; x, xylem, 
d, seed, e, j, cone-scales. B, Scale in longitudinal section ; c, vascular bundle ; 
a, periderm; b, palisade-tissue; I, 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 


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, x, 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 / (fig. 736, A). 

Cones similar to Conites Juddi are described by Velenovsky 1 
as Fricia 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 Jeffrey 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. 

I 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. PI. hi. fig. 6; PI. vm. fig. 13. 


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 Cunninghamia, 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 sphaerocarpus Carruthers. 

This species (fig. 737), from Inferior Oolite rocks at Bruton 
Somersetshire 1 , 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 Cookii (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 

Araucarites ooliticus (Carruthers). 

This species was originally described by Carruthers as Kaida- 
carpum ooliticum 2 from the Great Oolite of Northamptonshire and 
regarded as an inflorescence of some Pandanaceous plant. Zigno 3 
transferred it to Pandanocarpum. An examination of the type- 
specimen in the Northampton Museum led me to refer the cone to 
Araucarites*. 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 xi.; Seward (04) B. p. 131; (IP) p. 116, fig. 18. 

2 Carruthers (68) p. 156, PI. ix. 3 Zigno (85) p. 3. 

3 Seward (96 2 ) p. 216;- (04) B. p. 135. 




Fig. 737. Arauca rites sphaerocarpus. (British Museum, 41,036; f nat. size.) 

Fig. 738. Arauca rites ooliticus. (After Carruthers ; § nat. size.) 
s. iv 17 




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 

Fig. 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 1 from the Coralline Oolite at Malton, Yorkshire, as 
Araucarites Hudlestoni is probably another example of this species : 
in one of the specimens of A. Hudlestoni 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. ooliticus. 
It is possible that A. ooliticus is specifically identical with A. sfhae- 
rocarfus ; it is at least a closely allied type. 

The specimen figured by Lindley and Hutton 2 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. 

2 Lindley and Hutton (34) A. PI. cxxix. 


Pleydell 1 from the Inferior Oolite of Dorsetshire, A. sphaericus 
(Carr.) 2 , originally referred to Cycadeostrobus, and other species. 

Araucarites pippingfordensis (Ung.) 3 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 4 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 5 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 
overed 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 3 ) p. 105; Seward (04) B. p. 138. 

3 Seward (13) p. 104. 

4 Carruthers (69 2 ) p. 3, PL v. figs. 1—6; Seward (04) B. p. 137, PI. in. fig. 5; 
PI. xii. fig. 2. 5 Fliche (00) p. 11, PI. i. 



Araucarites hespera Wieland. This type is described by Wie- 
land 1 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 lacking 
details, and the photograph support his conclusion as to the 
Araucarian affinity of the specimen. 

The generic name Doliostrobus was instituted by Marion 2 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 Agaihis than Araucaria. Laurent 3 refers a small piece 
of a foliage-shoot from the Aquitanian beds in the Puy-de-D6me to 
Doliostrobus Sternbergii, though there is no information with regard 
to the cones. Gardner 4 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 Araucaria is that in Agaihis the scales 
are persistent, but as pointed out elsewhere 5 cones of Agaihis 
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 2 ) p. 4, fig. 1. 

2 Marion (84). 3 Laurent (12) PL v. fig. 7. 
4 Gardner (86) p. 93, Pis. xxn., xxm. 5 Seward and Ford (06) B. p. 359. 



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 1 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 Araucaria 
supplies confirmatory evidence. 

Pseudo- Araucaria. Fliche. 

The generic name Pseudo- Araucaria was given by Fliche 2 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 Eutacta 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 
he 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, PI. vi. figs. 3—5; PI. vn. figs. 1, 2. 



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 scale 
that can reasonably be referred to the genus Araucarites is 
A. Delafondi founded by Zeiller 1 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 2 mm. 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. Cookii. The 
number of names given to the fossfl scales is but a rough index of 

1 Zeiller (06) B. p. 215, PL l. fig. 1. 




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 Phillipsi Carruthers. 

Carruthers 1 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 

i ' f 

Fig. 740. 1, Araucarites Miller i (Upper Jurassic, Scotland); 2, A. Jeffreyi (Cre- 
taceous, N. America); 3, A. Rogersi (Wealden, S. Africa); 4, 1, A. Brodiei 
(Middle Jurassic, England); 5, A. Haberleinii (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. ) 

cuneate, nearly as long as broad (fig. 740, 9), and in shape similar 
to those of the cone already described as Araucarites ooliticus 

Examples of French Jurassic cone-scales are afforded by 
Araucarites Moreauana Sap. 2 , from Corallian beds near St Mihiel 

1 Carruthers (69 2 ) p. 6, PL n. figs. 7—9; Seward (00) B. p. 285, PI. x. fig. 4. 

2 Saporta (84) p. 425, Pis. clxxxiv., clxxxv. 



and other localities, similar to those of A. Brodiei (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 Eutacta type. Araucarites Falsani Sap. 2 is founded 
on twigs similar to those of Araucaria excelsa and scales character- 
ised by stout terminal spines. Both Araucarites Falsani and A. 
microphylla 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 3 from Solenhofen as Araucarites Haberleinii (fig. 740, 5). 
Salfeld figures some detached scales from the Malm of South- 
West Germany as Araucaria' 1 . 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 5 shows 
several scales still in their natural position. This type bears a 
striking resemblance to seme 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 Raj mahal series reach a breadth of 
5 cm. The scales of A. cutchensis 1 recorded from the Cutch flora 
(fig. 740, 8) and elsewhere are smaller, but in some cases it is 

1 Saporta (84) p. 431, Pis. clxxxvi., clxxxvii. 

2 Ibid. p. 439, Pis. clxxxvi., clxxxvii. 

3 Thiselton-Dyer (72). 

4 Seward (03) B. p. 37, PL vi. figs. 4—7. 

5 Tate (67) p. 147. 6 Feistmantel (77 2 ) p. 186, PI. vm. figs. 9—12 
7 Ibid. (76 2 ) p. 62, Pis. vil— ix., xn. ; (82) PI. m. 




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 1 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 nariow 
linear appendage. As Halle says, the scales closely resemble those 
of A. Brodiei Carr. 

Cone-scales from Jurassic rocks in Victoria 2 , 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. 3 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 4 from Middle Cretaceous 
rocks in North Carolina as Araucarites Jeffreyi (fig. 740, 2). These 
scales are associated with the foliage-shoots referred to Araucarites 
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 5 from Virginia. 


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 (13 2 ) p. 72, PI. vm. figs. 3—10. 

2 Seward (04 2 ) B. p. 181, figs. 42, 43. 

3 Fontaine in Ward (99) B. p. 669, PL clxiii. figs. 1—9. 

4 Berry (08) p. 258, PI. xvi.; (14) p. 20. 

5 Ibid. (11) p. 399, PL lxxvii. fig. 5. 


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 Elatides. 

An example of a Palaeozoic fossil which has been assigned to 
Araucarites on slender grounds is afforded by Araucarites Oldhami 

Araucarites Oldhami Zeiller. 

The specimen from the Lower Gondwana rocks in India to 
which this name is applied 1 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- 
caria 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 2 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 3 is founded 
on fragments of sterile branches bearing elliptical-ovate leaves 
resembling the foliage of Araucaria imbricata and the fossil species 
Araucarites Nathorsti Dus. A very similar type is represented by 
Araucarites bladenensis, described by Berry 4 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 vn. fig. 6. 2 Feistmantel (79 2 ) PI. xin. fig. 6. 

3 Hollick (97) p. 128, PL xn. figs. 3 a, 4. 

4 Berry (08) p. 255, Pis. xn.— xiv.; (14) pp. 19, 105, PL m. figs. 6, 7; PL xix. 
figs. 1, 2. 



veins and imperfectly preserved stomata occur in rows on the 
lower surface. Berry compares the species with Araucaria Tou- 
casi figured by Saporta 1 from Turonian rocks in the South of 
France. Another species founded on a sterile shoot is Araucarites 
Hatcheri described by Wieland 2 from Upper Cretaceous rocks in 

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 Amncaria 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 Sternbergii Goeppert. 

This species was founded on sterile branches, from the rich 
Eocene flora of Haring in the Tyrol 3 , 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. Sternbergii and A. Goepperti Sternb. : the 
latter species was founded by Sternberg 5 on a Tertiary cone from 
Haring in the Tyrol. Ettingshausen 6 subsequently figured 
several good examples of vegetative shoots of this type from 
Haring and described a subglobose cone, figured by Gardner 7 , 
which he refers to the same species: this author also records 
A. Sternbergii from Bilin in Bohemia 8 , but under the generic 
name Sequoia: in his account of the occurrence of the species 
in Carinthia 9 he adopts the designation Araucarites. Ettings- 
hausen figures a single cone-scale from Eocene beds in Styria as 
Araucarites schoeneggensis 10 and compares it to the scales of 

1 Saporta (79) A. p. 198, PL xxvil, 2. 2 Wieland (08 2 ) p. 6, fig. 2. 

3 Goeppert (50) p. 236, PL xliv. fig. 1. 

4 Ibid. PL xliv. fig. 2. 5 Sternberg (38) A. PL xxxix. fig. 4. 
6 Ettingshausen (55) p. 36, Pis. vn., vm. 7 Gardner (86) p. 96. 

8 Ettingshausen (67 2 ) p. 116, PL xin. figs. 3—8. 

9 Ibid. (85). 10 Ibid. (90) PL t. fig. 93. 




A. Sternbergii. The latter species is recorded also by Massalongo 1 
from Eocene rocks in Italy and on imperfect evidence by Heer 2 
from Switzerland. Gardner 3 describes several good specimens of 
vegetative shoots from the Eocene flora of Bournemouth which 
he names Araucarites Goepperti Sternb. though the specific name 
Sternbergii would be more appropriate 
as that designation was first applied ft 
to. similar branches from Haring and Jjj^ 
A. Goepperti 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- 
hamii : with reference to the absence 

of COnes or COne-SCales he quotes the Fig. 741. Araucarites Sternbergii. 
fact, Communicated to him by an (British Museum, V, 523; nat. 

observer in Madeira, that the foliage 81ze ^ 
of A. Cunninghamii 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 


Massalongo (59) Pis. v. — vn. 
Gardner (86) p. 55, PL xi. fig. 1 ; PI. 

2 Heer (55) A. PI. xxi. fig. 5. 
4 Ibid. PL xi. 


included in Araucarites Sternbergii or A. Goepperti to those of 
A. Cunninghamii and other species the probability of generic 
identity is such as to justify the retention of the designation 

Araucarites Haastii (Ettingshausen). 

Ettingshausen 1 described this species as Araucaria Haastii 
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 5 cm. and 2 cm. 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 2 , but the specimen is too 
imperfect to warrant the use of the designation Araucarites. 

Araucarites Nathorsti (Dusen). 

This species, described as Araucaria Nathorsti, is recorded by 
Dusen 3 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 Dusen states, 
in their blunter apices. 

Araucarites imponens (Dusen). 

Nathorst 4 first suggested a reference to Araucaria of the single 

leaf on which this species was founded 5 : 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 n. 2 Ibid. p. 155, PI. I. fig. 18. 

3 Dusen (99) p. 105, PI. xh. 4 Nathorst (04 2 ) B. 

5 Dusen (08) p. 11, PI. I. figs. 16, 17. 





leaves of A. Bidwilli and A. brasiliensis, but the single impressioi 
is hardly sufficient to demonstrate the existence of Araucarites 
in this southern flora (lat. 64° 16' S.). On the other hand th( 
occurrence of wood of the Araucarian type 1 in Seymour Island 
in beds that are either Lower Tertiary or Upper Cretaceous 
supports the conclusion of Nathorst and Dusen. 


Heer 2 proposed this name 3 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, E. ovalis, represented by oval cones 
2-7 cm. long and 6 — 7 mm. broad, E. Brandtiana characterised by 
cylindrical cones, and E. falcata founded on vegetative branches 
very similar to those of E. Williamsonis. Nathorst 4 has included 
these three species in Elatides curvifolia (Dunk.) a Wealdei 
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 vegetativ 
characters of Elatides are those of Araucaria Cunninghamii, 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 wer 
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. Elatides is characteristic of Rhaetic, Jurassic, and 
Wealden floras. 

Elatides Sternbergii (Nilsson). 

A Rhaetic species originally described by Nilsson from Rhaetic 

rocks in the South of Sweden as Abies Sternbergii, subsequently 

1 Gothan(08). 2 Heer (77) ii. pp. 77—79, PL xiv. 3 e\dr V , Fir. 

4 Nathorst (97) pp. 35, 58, Pis. I., n., iv., vi. 





included by Nathorst 1 in Palissya Braunii but afterwards recog- 
nised by him as a distinct species 2 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 4 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. Elatides Sternbergii, 
though similar in the habit of the vegetative shoots to E. 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 E. Williamsonis and E. curvifolia. A cone figured by 
Nathorst 5 is practically identical in external form with one of 
E. Williamsonis illustrated in volume I. of the Jurassic Flora of the 
Yorkshire Coast 6 . 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. 

latides Williamsonis (Brongniart). 
This Jurassic species described by Brongniart 7 as Lycopodites 
Williamsonis, was figured by Phillips 8 as L. uncifolius and by 
Lindley and Hutton 9 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 10 , the name substituted by Heer for Pomel's Pachy- 
fhyllum, 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 2 ) B. p. 28, PL iv. figs. 1—3. 

2 Ibid. (86) p. 107, PL xxiii. figs. 8—12; Pis. xxiv., xxv. 

3 Ibid. (97) p 34; (08). 

4 Schenk (67) A. PL xli. fig. 7. See also Solms-Laubach (91) A. p. 73. 

5 Nathorst (86), PL xxv. fig. 8. 6 Seward (00) B. PL x. fig. 3. 
7 Brongniart (28) A. p. 83. 8 Phillips (29) A. PL vm. fig. 3 
9 Lindley and Hutton (33) A. PL xcin. 10 Seward (00) B. p. 291. 




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 Eutacta species of Araucaria 
and with Cryftomeria. The megastrobili are cylindrical, approxi- 
mately 6 cm. long and 2 cm. in diameter, bearing imbricate, flat, 
scales with narrow pointed distal ends resembling the free portion 

Fig. 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, 
2 cm. 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 1 (fig. 744) (Lind. and Hutt.), but in the 

1 See page 276. 




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 1 first described this Wealden type (fig. 743) from North 
Germany as Lycopodites and it was referred by Ettingshausen 2 to 
Araucarites, the generic name, though probably correctly express- 

Fig. 743. Elatides curvifolia. ( After Nathorst ; nat. size.) 

the position of the fossil Conifer, being used without adequate 
reasons. The identity of the cones discovered by Nathorst 3 in 
he 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 Reichenbacliii* to Elatides curvifolia, as also 
the employment of Dunker' s specific term for Elatides ovalis and 
E. Brandtiana Heer. The cones of E. 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 vn. fig. 9. 

2 Ettingshausen (52) PL n. 

3 Nathorst (97) pp. 35, 58. Pis. i., n., iv., vi. 

4 Heer (75) ii. PL xxxvi. figs. 1—8; PL xxxvn. figs. 1, 2. 

s. iv ]8 


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 E. Williamsonis. On a 
specimen of a cone of this species from Kimeridge strata in 
Scotland 1 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 Araucaria ; 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 2 ; it is 
also recorded from Kimeridgian strata in the North of Scotland. 


Heer 3 instituted this genus in place of Pachyphyllum, previously 
adopted by Pomel 4 for a section of his genus Moreauia, 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 Araacarites and Brachyphyllum. Tuzson 5 
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 6 that Pagiophyllum may most conveni- 

1 Seward (ll 2 ) p. 684, fig. 10. 2 Ibid. (95) A. p. 205; (ll 2 ) p. 685. 

3 Heer (81) p. 11. 4 Pomel (49) p. 352. 

5 Tuzson (09) p. 30. 6 Seward (12) p. 41. 


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 Pagiophyllum 1 ,' 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 2 
afterwards named V. Foettleri by Stur 3 , Pagiophyllum Sandbergi 
by Schenk 4 and figured by Schutze 5 as P. Foettleri, 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 (Dadoxylon) 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 6 
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, PI. vm. 

3 Stur (68) p. 104. 

4 Schimper and Schenk (90) A. pp. 276, 290. 

5 Schutze (01) PL vi. fig. 1. 

6 Zeiller (06) B. p. 219, PL li. 





Pagiophyllum peregrinum (Lindley and Hutton). 

This species, first named by Lindley Araucaria peregrina 1 , was 
founded on material from the Lias 
of Lyme Regis in Dorsetshire 2 . 
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 3 
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 William- 
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 

Fig. 744, 

Pagiophyllum peregrinum. 
(British Museum; nat. size.) 

1 Lindley and Hutton (33) A. PI. lxxxviii. 

2 Seward (04) B. p. 48, PI. v. ;. Saporta (84) p. 383, Pis. 173—176. 

3 Zeiller (06) B. p. 219. 


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. 

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. Sternberg. 

The name Walchia 1 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 Eutacta 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 2 , but it is only in a few specimens 
that seeds are preserved on the cone-scales: in some species, e.g., 
W. frondosa Ren. and W. fertilis Ken. 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. 




Fig. 745. Walchia piniformis. (British Museum; £ nat. size.) 


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 1 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 2 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 3 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 aJcernate bordered polygonal pits, and narrow medullary 
rays. It 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 regibn 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 

1 Bergeron (84). 2 Mougeot (52) A. p. 27.. PI. iv. 3 Fliche (03). 


which Fliche regards as leaf-bases and compares with those on 
Araucarian stems. While admitting the possibility that the wood 
belongs to Gomphostrobas 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 1 and 
Upper Coal Measures 2 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 o 
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 4 . The pinnately branched shoots 

1 Vernon (12) p. 607. 2 Kidston (02) B. 

3 Schlotheim (20) A. p. 415, PI. xxm. fig. la; PL xxv. fig. 1. 

4 £.gr.Stefani(01)p. Ill; Weiss, C. E. (72) p. 179; Heer (76) A. p. 57; Goeppert 
(65) p. 236; Grand'Eury (77) A. p. 514. 




(fig. 745) are characterised by the more or less oblique insertion of 
the slender branchlets and by the comparatively long, narrow, 
falcate, decurrent leaves. The ovoid or cylindrical megastrobili 
terminal on the ultimate shoots bear imbricate ovate-lanceolate 
sporophylls, but their preservation is not such as to throw any 
light on the structure of the seed-bearing organs. Zeiller 1 mentions 
a cone from Lodeve (Permian) 10 cm. long and 1 — 1-2 cm. in 
diameter, but the average length is less than this. A branch 
figured by Potonie 2 from the Permian of Thuringia shows elliptical 
leaf-cushions very like those on the larger stem described by 
Mougeot as Araucarites valdajolensis. Potonie 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 

idston 3 suggests, this may not be a specific character. Renault's 
species W. fertilis* represents a similar form but with smaller 
leaves, and each branchlet ends in a long and narrow strobilus 

hich 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 5 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 

1 Zeiller (06) B. p. 204, PL l. figs. 3, 5; Bergeron (84). 

2 Potonie (93) A. p. 218. 3 Kidston (86) A. p. 15. 

4 Renault (96) A. p. 359; (93) A. PI. lxxx. fig. 2. 

5 Zeiller (92) A. p. 99, PL xv. fig. 3. 



Walchia imbricata Schimper. 

In habit this type 1 closely resembles the foliage-shoots 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 2 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 Kidston from the Upper 
Coal Measures of central England. 

Walchia Schneideri Zeiller. 

This species 3 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. 4 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. Schneideri: 
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. 


Prof. C. E. Weiss 5 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, PI. xlix. figs. 1, 2. 

3 Ibid. p. 206, PI. xlviii. figs. 4, 5. 

4 Eichwald (60) B. PI. xix. fig. 1. 

5 Weiss, C. E. (72) p. 182, Pis. xix., xx. 




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 1 from Russia as species 
of Schizodendron and Angiodendron. 
Potonie, 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 2 , in view of this 
agreement, adopted the older name 
Schizodendron. The latter author 3 
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 
Potonie 4 proved that in bbth cases 
the median groove extends up the 

lower portion of each projecting Fig. 746. Schizodendron specio- 

area from its base and represents sum - (After Potonie; f nat. 
an out-going leaf-trace; he also slze '' 
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 bv Miss Holden 5 . The true nature of 

1 Eichwald (60) B. PI. xvm. 
3 Ibid. (80) p. 263, PL v. 
5 Holden, R. (13) p. 245. 

2 Zeiller (92 2 ) A. p. 102. 
4 Potonie (88). 


the Tylodendron casts is also shown in specimens from the Lower 
Permian of Saxony in the Chemnitz Museum 1 . 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 2 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 Endolepis has in several instances 
been found in connexion with the foliage of Voltzia 3 (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 art 
usually uniseriate. Potonie 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 pseudo whorl 4 , marks 
the position of branches. The pith-cast of a recent Cycad (fig. 398, 
p. 29, Vol. in.) 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 5 has recently described an interesting example 
of Schizodendron which throws some fresh light on structural 
features. The specimen was found in Cheshire but not in situ 
and nothing is known as to its geological age; it consists of a 

1 Sterzel (00). 2 Fliche (10) p. 212, Pis. xix., xx. 

3 Seward (90). See also page 290. 

4 Zeiller (92 2 ) A. PI. xv. fig. 5. 5 Weiss, F. E. (13 2 ). 



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 Potonie : 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 S. Cowardi the presence of secretory canals is a Cycadean 
feature. Bain and Dawson 1 , 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- 

Kri Saxony, and Zeiller, who noticed a similar association 


in French Permian rocks, expressed the opinion that the casts 
belonged to Walchia stems. In this connexion it is noteworthy 
that shoots of Voltzia 1 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 2 , from France, Russia 3 , and Canada while the British 
specimen, though presumably from English rocks, was not found 
in situ. 


Renault 4 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 
dftep. 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); Potonie (88); (93) A. etc. 

3 Schmalhausen (87) PI. vn. fig. 34. 

4 Renault (96) A. p. 360; (93) A. PI. lxxvi. 


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. 


This generic name was instituted by Marion 2 for Permian 
foliage-shoots from Lodeve 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 3 had pre- 
viously described detached sporophylls from the Lower Permian 
of Saxony, of the same type as those on which the genus Go mp h ostro- 
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. Potonie 4 , who 
examined the original fossils in the Dresden Museum — and 1 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 

1873. Sigillariostrobus bifidus, Geinitz, Neues Jahrbuch Min. p. 700, PI. v. 

figs. 5—7. 
1890. Dicranophyllum gallicum, Schenk in Schimper and Schenk, p. 266. 

1890. Gomphostrobus heterophyllus, Marion, Compt. Rend. ex. p. 892. 

1891. Psilotiphyllum bifidum, Potonie, Ber. deutsch. Bot. Ges. Bd. ix. 

p. 256. 

1892. Gomphostrobus bifidus, Zeiller, Bassin Houill. Perm. Brive, p. 101, 

PL xv. fig. 12. 

Gomphostrobus bifidus (Geinitz). 

Potonie 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. ii. p. 26. 2 Marion (90) A. 3 Geinitz (73). 

4 Potonie (93) A. p. 197 Pis. xxvn., xxvm., xxxiii. 





Fig. 747. Gomphostrobus 
bifidus; a, leaf -scar; b. 
scar of sporangium?; 
c, torn piece of tissue. 
(A, B, after Potonie; 
C, Zeiller.) 

with two divergent distal prongs varying considerably in the angle 
of divergence, a variation noticed also by Zeiller. On one example 
(fig. 747, B) Potonie 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- 

The species, represented usually by de- 
tached sporophylls only, is recorded from 
Lodeve, Brive, and other French localities 1 , 
also from Permian localities in Germany. 
It was referred by Schenk to Dicrano- 
phyllum, but in that genus it is the foliage- 
leaves that are forked and there is no reason 
to assume any close relationship between 
the two imperfectly known types. If the scar 
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 2 records the association of Gomphostrobus 
with Dadoxylon wood. Potonie, 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 Walchia 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 3 as Voltzia revealed the 
occurrence of some small distally forked leaves very similar to the 
sporophylls of the European Gomphostrobus. 

1 Zeiller (92 2 ) A. p. 101, PI. xv. fig. 12; (06) B. p. 213, PL l. figs. 6—8. 

2 Sterzel (00) p. 6. 

3 Feistmantel (79 2 ) Pis. xxn. et seq. 




VOLTZIA. Brongniart. 

Brongniart 1 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 Ullmannia: 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 Voltzia 
though without proof of connexion with the vegetative shoots. 
Gothan 2 , who favours a Taxodineous alliance, points out that 
wood associated with Voltzia 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. Liebeana in the British Museum from 

era 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. A recent 
view 3 that Voltzia affords an illustration of a generalised type 
combining Araucarian and Abietineous features is in part based 
on an assumption that the cone-scales are double like those of the 
Abietineae. That the genus is a generalised type is probable, 
but the data are insufficient to warrant any definite statement as 
to which Coniferae are the nearest allies. The range of the genus 
difficult to define : if we include the species V. kenperiana, also 
Heer's genus Leptostrobus, the geological range extends from the 
Permian to Middle Jurassic floras. The typical species are charac- 
teristic of Permian and Lower Triassic rocks. The similarity in 

abit of Walchia, some species of Ullmannia, and Voltzia renders 

1 Brongniart (28) p. 448, Pis. xv.— xvn. 2 Gothan (10) p. 31. 

3 Holden, R. (13). 

s. IV 19 




exceedingly difficult the determination of sterile branches. 
Further, the fact that the specimens of this presumably arborescent 
genus are usually small branch-fragments sets a limit to our know- 
ledge of the external features of the individual plants. 

Voltzia heterophylla Brongniart. 

The examination of numerous specimens from the Bunter 
sandstone of the Vosges led Schimper and Mougeot to include 

Fig. 748. Voltzia. A. B, Voltzia heterophylla (B, medullary cast). C — F, Voltzia 
Liebeana. (A, B, after Seward; C — F after Geinitz.) 

under this name Voltzia brevifolia and V . rigida of Brongniart 1 . 
The Triassic species Voltzia heterophylla (fig. 748, A) is the best 
known representative of the genus: the vegetative shoots agree 
very closely with those of Araucaria excelsa and differ but little 
from shoots of Walchia, some forms of Ullmannia, and Pagiophyl- 

1 Brongniart (28) p. 446; Schimper and Mougeot (44) A. p. 21, Pis. I., vi. — ix. 



lum. Schiitze 1 has given a long list of references to records of 
V. heterophylla, but in making use of such lists it should be remem- 
bered that in the absence of reproductive organs the specific or 
even generic determination of specimens resembling in habit 
Araucaria excelsa is a hopeless task. The heterophylly of Voltzia 
heterophylla is a striking feature : long linear obtuse leaves, 2 — 5 
cm. long, occur in close association with falcate decurrent foliage. 
Small oval strobili characterised by crowded imbricate appendages 
are figured by Schimper and Mougeot as male cones, but in no case 
have any sporangia been detected : similar strobili are also figured 
by Leuthardt 2 from the Keuper of the Basel district. The mega- 
strobili are longer and bear cuneate cone-scales, with 3 — 5 rounded 
lobes on the upper surface, arranged in a lax spiral. The marginal 
lobes of the scales are less deeply separated from one another 
than in V. Liebeana (fig. 748, D — F). We have no satisfactory 
information with regard to the nature or method of attachment 
of the seeds. Saporta 3 figures a cone from Soultz-les-Bains 
showing, as he asserts, the impressions of seeds, but the drawing 
affords no definite evidence as to the relation of cone-scales and 
eeds. Saporta regards the cone-scales as double, each consisting 
of an ovuliferous scale and a bract-scale more or less completely 
fused as in the recent genus Taxodium. The assumption that the 
scales are double rests on a very slender basis, and even in the 
much better preserved specimens of V . Liebeana 'there is nothing 
to indicate that the scale was double 4 .' In a recent paper Miss 
Holden 5 speaks of Voltzia cone-scales as double in terms suggesting 
a well-established fact, though this is by no means the case. 

Blanckenhorn 6 and other authors have described specimens of 
Voltzia heterophylla showing elongated leaf-cushions which they 
compare with similar raised areas on the Permian casts on which 
Weiss founded the genus Tylodendron (Schizodendron 1 ). In the 
latter genus the supposed leaf-cushions are casts of medullary 
rays at the inner edge of the secondary xylem, and an examination 
of Voltzia specimens in the Strassburg Museum 8 convinced me that 

1 Schiitze (01). 2 Leuthardt (03) p. 10, PL iv. 

3 Saporta (84) PL 154, fig. 4. 4 Solms-Laubach (91) A. p. 68. 

5 Holden, R. (13) p. 251. 

6 Blanckenhorn (85) B. PL xx. figs. 17—20. 

7 See page 282. 8 Seward (90). 



the same explanation applies to Voltzia heterophylla. Triassic 
Voltzia casts were referred by Schleiden to a distinct genus Endo- 
lepis: examples figured by Schenk 1 afford a good illustration of 
their close resemblance to Schizodendron. The medullary casts 
of the Triassic genus differ from those of Schizodendron in their 
smaller diameter and in the absence of periodic swellings : narrow 
slits in the elongated areas mark the position of out-going leaf- traces 
(fig. 748, B). A similar though larger form of cast is figured by 
Miss Holden 2 from Coburg and New Brunswick and referred to 
Voltzia coburgensis. Fliche 3 has described a form from the 
Muschelkalk of France with more slender shoots than in most 
examples of V. heterophylla. Specimens from Swiss Triassic beds 
figured by Heer 4 and Leuthardt 5 as examples of this species are 
too incomplete to be identified with certainty. Feistmantel's 
Indian specimens 6 referred to V. heterophylla, which I have re- 
cently examined, from Lower Gondwana strata, show a variation in 
leaf -form suggestive of the European species, but the determination 
is open to question. A supposed cone-scale figured by Feist- 
mantel resembles in outline the lobed scales of V. Liebeana. Some 
very incomplete branches regarded by Feistmantel as pieces of 
Albertia shoots are probably identical with the impressions assigned 
to Voltzia. On a few of the smaller Indian specimens I found 
leaves 5 mm. long divided into two slightly divergent prongs, 
a feature unknown in Voltzia but suggesting Gomphostrobus or 
small leaves of Dicranophyllum. Some small seeds figured by 
Zeiller 7 from the Karharbari beds of India as probably belonging 
to Voltzia cannot be determined with confidence. 

Voltzia walchiaeformis Fliche. 

Fliche 8 gives this name to vegetative shoots from the Bunter 
of the Vosges characterised by a Walchia-like habit of branching, 
the pinnately disposed lateral branches being given off at about 
40° ; the leaves are elliptical, short and broad, more or less appressed 
and less spreading than in Walchia. It is, however, impossible 
without the confirmatory evidence of strobili to distinguish 

1 Schenk (68) p. 80, PI. vi. 2 Holden, R. (13). 

3 Fliche (10) Pis. xvra., xix. 4 Heer (76) A. PI. xxx. 

5 Leuthardt (03) PI. iv. figs. 2—5. 6 Feistmantel (79 2 ) Pis. xxn— xxv. 

7 Zeiller (02) B. PI. vn. fig. 9. 8 Fliche (10) p. 198, PI. xxi. 



clearly between certain forms of Walchia and Voltzia. A very 
similar type is figured by Schiitze 1 as Widdringtonites keuperianus 
Heer 2 from Stuttgart but with no justification for the use of a 
generic name implying relationship with Widdringtonia. 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 Liebeana Geinitz. 

A Permian species 3 , 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 7 cm. 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 3 mm., 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. Liebeana but have slightly 
narrower lobes. 

Voltzia keuperiana (Schimper). 

This specific name 5 was given to fertile shoots characterised 
by long and lax strobili called by many authors V. coburgensis 
Schauroth 6 , a designation first applied to a cast resembling Lygi- 
nodendron (cf. fig. 401, Vol. in. 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. PI. xxx. figs. 4, 5. 

3 Geinitz (80) p. 26, PI. v. 4 Heer (76 2 ). 

5 Schimper (72) A. p. 243, PI. lxxvi. 6 Schauroth (52) p. 540. 



megasporophylls constitute the distinctive features of V. keuper- 
iana: 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- 
phylld. 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 1 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 2 adopted the 
generic name Voltzia and Potonie proposed a new term Voltziopsis 
to be used in a provisional and wide sense for Voltzia keuperiana 
Cheirolepis Escheri Heer, Heer's Leptostrobus* and Nathorst 
Swedenborgia 5 (fig. 749), including species ranging from the Keupe 
to Middle Jurassic strata in contrast to the Lower Triassic and 
Permian range of typical representatives of Voltzia. The species 
Cheirolepis Escheri, included by Potonie 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 Munsteri Schenk 6 . The genus Cheirolepis was in- 
stituted by Schimper 7 as a substitute for Brachyphyllum for the 
Rhaetic species B. Munsteri 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 8 was hardly justifiable in the absence of sporo- 

1 Heer (77) ii. p. 72. 2 Schimper and Schenk (90) A. p. 290, fig. 199. 

3 Potonie (99) B. p. 304. 4 Heer (77) ii. p. 72. 

5 Nathorst (78) B. p. 30, PI. xvi. figs. 6—12. 

6 Schenk (67) A. p. 187. PI. xliii. figs. 1—12. 

7 Schimper (72) A. p. 247. 8 Seward (00) B. p. 294. 






phylls. The species Voltzia recubariensis (Mass.) represented by 
several vegetative shoots, imperfect cones, cone-scales, and seeds 
in the Muschelkalk beds of Kecoaro 1 illustrates the impossibility 
in the case of sterile specimens of drawing any satisfactory line 
between Vollzia 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 7 cm. in length and 1-5 cm. 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 2 includes in Leptostrobus clusters of long 
linear leaves apparently borne on short shoots and resembling the 
needles of Pityites Solmsi Sew., Schizolepis Braunii, and Jeffrey's 
Prepinus. These leaves, described as L. rigida and L. angusti- 
folia, 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 3 . It 
would be difficult to draw any 
generic distinction between Lepto- 

strobus and Voltzia especially V 


This genus 4 is founded on lax 
oval strobili bearing small cone- 
scales with long stalks and a 
single seed (fig. 749, B). In the 
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 

Fig. 749. Swedenborgia cryptomerides. 
A, Strobilus. B, Fertile leaf. (After 
Nathorst; nat. size.) 

1 Schenk (68) Pis. vn. et seq. 
3 Ibid, (77) ii. PI. xtii. fig. 14. 

2 Heer (82) p. 23, Pis. vn., vm. 
4 Nathorst (78) B. p. 30, PI. xvi. 



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 Potonie's 
genus Voltziopsis. 

Strobilites. Strobilites laxus 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 1 , which may be allied to Voltzia, 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 2 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. Ullmannic 
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. Bidwilli. 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 3 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 4 uses 

1 Seward (08) B. p. 101, text-fig. 7; PI. v. fig. 3. 2 Goeppert (50) p. 185. 

3 Solms-Laubach (84) PI. m. fig. 16; Schimper and Schenk (90) A. p. 275, figs. 
190, 191. 4 Tuzson (09) p. 23. 




the generic name Ullmannites for wood having the Araucarian type 
of tracheal pitting which he believes to belong to Ullmannia, but 
under the former genus are included types of wood that are not 
generically identical and afford no evidence of connexion with 
Ullmannia. Ullmannia 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 

Fig. 750. Ullmannia. A — C, Ullmannia frumentaria. D, E, U. Bronni. (A, after 
Geinitz; B, C, after Solras-Laubach ; D, E, after Potonie.) 

biarmica Eich. 1 being represented by both sterile and fertile 
>ranches. Imperfectly preserved impressions from the Permian 

of Durham 2 have been referred to Ullmannia but no satisfactory 
specimens have been discovered in English beds. The fragment 

figured by Lindley and Hutton 3 as Voltzia Phillipsi may belong to 

an Ullmannia. 

1 Schmalhausen (87) PL vi. 

2 Murchison and Harkness (64) p. 154; 

3 Lindley and Hutton (37) A. PL 195. 

Kirby (64). 



Ullmannia Bronni Goeppert. 

The fragments of foliage-shoots on which Goeppert 1 founded thi 
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 Cwpressites 
Ullmanni. 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 Ilmenau (Thuringia) and 
both were described by Brongniart as examples of Fucoides 2 . 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 (cf. fig. 750, C) associated with parenchyma 3 . 
As in the leaves of recent Conifers the transfusion-tissue persis 
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 frumentarius*, is the 
commonest fossil in the Ilmenau mines. The leaves are lanceolate, 

1 Goeppert (50) p. 185, PI. xx. For synonymy, see Solms-Laubach (84); 
Geinitz (80). 2 Brongniart (28) A. PI. ix. 

3 Solms-Laubach (84) PI. ra. figs. 1, 4, 6, 15. 

4 Schlotheim (20) A. PI. xxvu. 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 triangularis 1 , 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 2 . 

Strobilites. Strobilites Bronni (Goeppert). 

Under this provisional name Solms-Laubach 3 described the 
problematical fossils from the Frankenberg copper mines which 
Goeppert believed to be the megastrobili and cone-scales of 
Ullmannia Bronni. 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 4 cm. x 2 cm. of rather smaller scales without radial ribs 
and characterised by a stout, rounded, peripheral border. 

1 Geinitz (80) PL in. figs. 11—15. 

2 Zeiller (06) B. p. 219, PI. l. 

3 Solms-Laubach (84). 


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 1 substituted 
Haidingera. Schimper and Mougeot 2 figure reconstructions of 
both male and female cones and a single cone-scale bearing a 
median seed. Schenk 3 , who examined the original specimens, 
states that the supposed male cone is a young megastrobilus of 
Voltzia, and Solms-Laubach 4 , who also examined the material in 
the Strassburg Museum, considers that Schimper's statement that 
the seed-scale and cones belong to Albertia is ' altogether arbitral 
and unsupported.' There would seem to be no reason for conned 
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- 
tums. The branches described as A. elliptica are not distinguishable 
by any clearly marked feature from A. latifolia. Albertia Braunii 
has larger obovate leaves and A. speciosa has broadly linear leaves 
reaching a length of 4-5 cm. There is a very close resemblance 
between the shoots from the Bunter beds and those of some forms 
of Ullmannia especially U . frumentaria (Schlot.) 6 , and it is doubt- 
ful whether any useful purpose is served by the retention of the 
designation Albertia : the descriptions of the reproductive shool 
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 ai 
no grounds for regarding the specimens usually referred to Albertia 
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 7 to 

1 Endlicher (47) p. 303. 

2 Schimper and Mougeot (44) A. PI. i. 

3 Schenk in Schimper and Schenk (90) A. p. 284. 

4 Solms-Laubach (91) A. p. 75. 

5 Schimper and Mougeot (44) A. Pis. i. — v. 

6 Geinitz (80) PI. in. 

7 Feistmantel (79) p. 29, PI. xxvi. fig. 2. 



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 xxxiii. as Dacloxylon 
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 than is 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, (iii) Such evidence 
as is afforded by Tertiary records, though meagre and often in- 
complete, points to the continued existence of the family in the 
orthern Hemisphere at least in the older Tertiary floras. 



The 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 th 
later Jurassic and Lower Cretaceous floras Conifers agreein 
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. 


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 1 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 2 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. 


Goeppert's term Cupressites 3 is retained for fossils which there 
is reason for associating 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. 


in a wider sense. Bowerbank 1 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 2 in 
this genus afford examples of the inconstancy of leaf-arrangemen 
in a single type and demonstrate the impossibility of drawing an 
definite distinction between this genus and Thuytes as used b 
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 3 , 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, Pis. 202 et seq. 

3 Ibid. PL 214. 


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. 1 
and Biota orientalis Endl. succinea Goepp. 2 from the Oligocene 
amber deposits on the Baltic coast. Schlechtendal 3 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 MacHenryi Baily. 

This name was given 4 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 5 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 6 as Pinites Pritchardi belongs to the species which 
furnished the Antrim specimens : the wood, subsequently referred 
to Cupressinoxylon 1 , 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 14 mm. 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, PI. xv. figs. 199—206. 

2 Ibid. p. 42, PI. xv. figs. 180—198. 

3 SchlechtendaJ (02) Pis. I., n 

4 Baily, W. H. (69) p. 361, PI. xv. fig. 5. $ 

5 Gardner (86) p. 82, PI. xvi. figs. 8, 9; PI. xvm. fig. 1; -PI. xix. 

6 Goeppert (50) p. 220. 7 Kraus in Schimper (72) A. p. 376. 
S. TV 20 


flourished on the western edge of Europe during the period oi 
volcanic activity responsible for the widespread sheets of lava ii 
the North-East of Ireland and the Western Isles of Scotland. 

Cupressites taxiformis Unger. 

This species was founded by Unger 1 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 leave 
are appressed to the axis and free only at the apex, resembling on 
a small scale those of Sequoia 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 2 from 
the Isle of Wight and several specimens are figured by Gardner 3 
from the Middle Bagshot beds of Bournemouth. The material 
from the latter locality consists of sterile shoots with linear Taxus- 
like leaves from 5 to 15 mm. long and occasionally, on the same 
axis, smaller decurrent leaves, though generally the two forms ar 
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 4 : it is characterised by peltate scales 
with a central umbo and a wrinkled surface. 

The evidence in favour of assigning all the _ 

. ° Fig. 751. Cupressites 

sterile shoots to the same species is not con- taxiformis. (Fro 
vincing, though a similar combination is met a specimen in th 
with in the recent species Glyptostrobus hetero- Bj ?^ , Mu 
phyllus. If we confine our attention to the cones 
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 dannot be determined without 
further investigation. 

1 Unger (47) p. 18, PL vm. figs. 1—3; PI. ix. figs. 1—4. 

2 De la Harpe in Bristow (62) PI. v. fig. 2. 

3 Gardner (86) p. 26, Pl.^- figs. 1—13; PL v. figs. 13, 14; PL vn. fig. 8; PL ix. 
figs. 22—26, 28—30. 

4 Ibid. PL ix. fig. 27. 



Another Tertiary representative of the recent genus Cupressus 
is that described from the Oligocene amber beds of East Prussia 
as Cupressus sempervirens L. succinea Goepp. and Menge 1 . This 
species is founded on fragments of sterile shoots with a well pre- 
served male flower showing very clearly the form of the sporophylls. 


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 2 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 3 . 

Cupressinocladus salicornoides (Unger). 

A sterile piece of branch figured by Lindley 4 from Provence as 
Thuya is probably identical with Unger's species recorded by 
Saporta 5 from the same locality. The type-specimens on which 
Unger founded the species Thuites salicornoides (fig. 752) are from 
Eocene beds in Croatia 6 ; 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 1-5 cm. By later authors 
this species, recorded from Styria 7 , Bohemia 8 , Switzerland 9 , the 
Oligocene beds of East Prussia 10 , Italy 11 , the Miocene of France 
and from other localities 12 , is spoken of as Libocedrus, but the few 
examples of cones that have been figured do not exhibit with 

1 Goeppert and Menge (83) A. p. 45, PI. xvi. figs. 218—224. 

2 Heer (82) i. p. 49, PL xxix. figs. 1, 2; PI. XLin. fig. Id. 

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 2 ) p. 42, PL i. fig. 4. 6 Unger (47) p. 11, PL n. 

7 Ettingshausen (70) p. 39; (88) p. 273. 

8 Ibid. (67 2 ) p. 109, PL v. figs. 1—7, 14; Engelhardt (85) PL vm. figs. 27—30. 

9 Heer (55) A. p. 47, PL xxi. fig. 2. 
" Goeppert and Menge (83) A. PL xv. figs. 175—177. 
11 Massalongo (59) p. 153, PL v. figs. 20—23. 12 Marty (08). 



sufficient clearness morphological features that justify the con 
elusion that the specimens agree more closely with Libocedru 
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 

Fig. 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 1 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 2 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 3 also records the species from Aquitanian 
beds in the Puy-de-D6me. 

1 Heer (83) p. 58, Pis. lxx., lxxxvi., lxxxvii. ; Beust (85) p. 40, Pis. in., v. 

2 Gardner (86) p. 25, PI. n. figs. 17—20. 3 Laurent (12) p. 69. 


(ii) Species previously referred to Thujopsis. 

Cupressinocladus massiliensis (Saporta). The small twigs de- 
scribed by Saporta from Provence as Thujopsis massiliensis 1 and 
compared by him with Thuya occidentalis are very similar to those 
named by the same author T. europaea 2 : in neither case is there 
any justification for the use of the generic name Thujopsis. Heer 3 
records T. 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 4 refer some fragments from the Baltic amber beds to 
Thujopsis europaea, but no reproductive organs are figured. 

(iii) 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 Thuites 
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 5 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, PI. i. fig. 6; PI. iv. fig. 2. 

2 Ibid..(65 2 ) PI. i. fig. 5. 

3 Heer (68) i. p. 90, PI. l. fig. 11; (75) iii. p. 7, PI. i. figs. 13—29. 

4 Goeppert and Menge (83) A. PI. xvi. figs. 215—217. 

5 Seward (95) A. p. 209, PI. xx. fig. 6. 


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 1 to Thuites Ehrenswaerdi and T. 
Meriana 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. 2 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 3 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 5 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 6 figure specimens from the lignite 
beds at Kreischerville as J. hypnoides and believed them to be 
identical with those described by Newberry as J. ynacilenta ; they 
also speak of the association of Dammara [Agathis] scales. 

The Tertiary Greenland species J. tertiarius and J. gracilis 7 , 
founded by Heer on sterile branches, are equally unsatisfactory as 

1 Heer (71) iii. p. 38, PI. n. figs. 25, 26; (82) i. PL i. 

2 Goeppert and Menge (83) A. p. 44, PI. xvi. figs. 211—214. 

3 Heer (82) i. p. 47, PI. xliv. fig. 3; PI. xlvi. fig. 18. 

4 Ibid. PI. xxxv. figs. 10, 11. 5 Newberry and Hollick (95) PI. x. fig. 7. 

6 Hollick and Jeffrey (09) B. p. 61, PI. v. figs. 5, 6. 

7 Heer (83) p. 57, Pis. lxx., cit., cvi. 


records of Conifers closely allied to Juniperus, and the same remark 
applies to Juniperites eocenica described by Ettingshausen 1 from 
Haring in the Tyrol. A single male flower figured by Goeppert 
and Menge 2 from the Baltic amber as Juniperus Hartmannianus 
may be correctly referred to that genus though other recent genera 
are not excluded. 

Echinosirobus Schimper. 

Proposed in the first instance by Schimper 3 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, Echinosirobus is dis- 

PhyUostrobus Saporta. 

This generic name was given by Saporta 4 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 5 
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. 

Athrotaxites Unger. 

This name was proposed by Unger 6 for a branched cone-bearing 
shoot from Solenhofen agreeing in vegetative characters with 

1 Ettingshausen (55) PI. v. fig. 6. 

2 Goeppert and Menge (83) A. p. 39, PI. xiv. figs. 156, 157. 

3 Schimper (72) A. p. 330. 4 Saporta (84) p. 635. 
5 Thiselton-Dyer (72). « Unger (49) PL v. figs. 1, 2. 



specimens previously figured by Sternberg as Colder files. The 
specimen in the Munich Museum on which the type-species Athro- 
taxites lycopodioides was founded was examined and re-figured by 
Schimper 1 and by Saporta (fig, 753, C) 2 : the former author 
substituted for Athrotaxites a new genus Echinostrobns 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 Sternbergii. Saporta, who 
believed Unger's type to be intermediate between Athrotaxis 
and Cryptomeria as regards the features of the cones, retained 
Schimper's designation Echinostrobns Sternbergii. Unger 3 in sub- 
sequent accounts of Solenhofen plants extended the application of 
Athrotaxites to sterile shoots, and this course was also followed by 
Thiselton-Dyer 4 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 5 . Most 
of the sterile specimens referred to Athrotaxites by Unger and other 
authors should be transferred to Brachyphyllum. 



Athrotaxopsis Fontaine. 

Fontaine 6 instituted this name for some fertile Coniferous 
shoots from the Potomac formation similar in habit to species in 
eluded by Saporta in his genus Palaeocyparis and to the genu 
Thuites but bearing cones different from those of Athrotaxites 
lycopodioides. Berry 7 subsequently reduced the number of Fon 
taine's species and pointed out that some of his specimens ar< 
indistinguishable from shoots included in Sphenolepidium, a con- 
clusion to which I had been led 8 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 lxxv. fig. 21. 

2 Saporta (84) PL 199. 3 Unger (52); (54 2 ). 
4 Thiselton-Dyer (72). 5 Halle (13) p. 40. 

6 Fontaine (89) B. p. 239. 7 Berry (ll 4 ) 8 Seward (95) A. p. 202. 




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 

Fig. 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 Athrotaxis is a 


survival of a type of Conifer which was widely spread in Jura 

Athrotaxites lycopodioides Unger (= Echinostrobus Sternbergii 
Reference has already been made to the vegetative characters 
of this Upper Jurassic species from Solenhofen 1 . 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 Ungeri Halle. 

This species 2 , 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. 

In his memoir on British Eocene Gymnosperms Gardner 3 
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 4 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, Pis. n.— v. 

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. 


pieces of vegetative organs and in one case a cone to Athrotaxis, 
but the evidence on which the species Athrotaxis (?) subulata 1 is 
founded has little value. 


Brongniart 2 proposed this name for a Jurassic species, Br achy - 
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 3 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 
Brachy phyllum. 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 4 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 5 figured a fertile specimen from the 
Rhaetic rocks of Franconia, which he referred to Brachyphyllum 
speciosum Miinst., and this was re-figured by Schenk 6 as Palissya 
aperta though as others have pointed out the cones are very 
different from those of Palissya: as Nathorst 7 says, they have the 
characters of the genus Elatides, and the same is true of some cones 
figured by Saporta 8 and assigned by him to Brachyphyllum. The 
name Elatides is reserved for specimens characterised by a certain 

1 Gardner (86) p. 43, PI. xi. 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. 

6 Schenk (67) A. PL xlii. figs. 1—13. 7 Nathorst (97) p. 34. 

* Saporta (84) PL 165, fig. 1; PL 167, figs. 2, 3; PL 171, figs. 7—9. 


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 1 
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. 2 and Banc, and a Japanese species 
originally described as Yezonia vulgaris by Drs Stopes and Fujii 3 
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. 869. 

3 Stopes and F-jju (10) p. 23; Jeffrey (10 3 ). 




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 expansion (Sternberg). 

In transferring this widely spread Jurassic species, founded by 
Sternberg on a specimen from the Stonesfield (fig. 754), Oxford- 

Fig. 754. Brachyphyllum expansum. 
(Sedgwick Museum, Cambridge.) 

Fig. 755. Brachyphyllum expansum. 
(Figured by Feistmantel as Echino- 
strobus expansus.) 

shire, as Thuites expansus 1 , 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, PI. xxxvm. 
B. p. 142. 

s. 1, 2. For synonymy see Seward (04) 


Brachyphyllum. Schimper 1 included Sternberg's species in Echino- 
strobus and Saporta 2 adopted the designation Palaeocyparis. The 
small amount of evidence with regard to the structure of the cones 
does not afford an adequate reason for retaining the generic name 

Specimens from Jurassic rocks in India described by Feist- 
mantel 3 (fig. 755) as Echinostrobus expansus, superficially at least 
very similar to the European Thuites expansus, have recently been 
examined by Miss Holden 4 . 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. Munsteri 
and B. affine 5 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 (76 2 ) p. 60, Pis. ix., x. 

4 Holden, R. (15 2 ) p. 221, PI. xi. 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. 




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 

Fig. 756. Brachyphyllum expansuml. (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 


each cone-scale has a funnel-like cavity near one edge and the 
upper side of the cavity is radially ridged 1 . 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 FeistmantePs 
Pachyphyllum heterophyllum 2 from Indian Jurassic beds: on the 
stouter apris 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 3 and both agree 
superficially at least with European examples of Brachyphyllum 

Brachyphyllum mamillare Brongniart. 

This specific name 4 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 5 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. mamillare. 

1 Seward (04) B. PL ix. fig. 4. 

2 Feistmantel (79) PL xi. fig. 4. 3 Ibid. PL xi. fig. 2. 

4 Brongniart (28) A. p. 109. 5 Lindley and Hutton (35) A. PL clxvii. 


Among these are some of the shoots referred by him to Echinostrobus 
expansus 1 and others described as E. rajmahalensis Feist, and 
E. rhombicus 2 . An examination of some of the figured specimens 
referred by Feistmantel to Pachyphyllwn (= 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 3 . 

Zeiller 4 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 5 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 

I 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- 

I 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 6 is represented in the Potomac formation by specimens 
referred by Fontaine 7 to Brachyphyllum crassicaule, and there are 

* Feistmantel (76 2 ) PL ix. figs. 6—9; PL x. figs. 3, 4. 

2 Ibid. (79) PL xii. figs. 2, 10; (82) PL m. fig. 6. 

3 Saporta (84) Pis. 165—172. * ZeiUer (00) p. 3. 
5 Seward (95) A. p. 215, PL xvn. « Heer (81) p. 20, PL xvn. 
7 Fontaine (89) B. p. 221, PL c. fig. 4; PL cix. figs. 1—7. 

s iv 21 


many specimens recorded both from Jurassic and Cretaceous rocki 
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 1 for Portuguese specimens. Apart from 
the absence of thornlike branches this species is Fig. 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 2 . 

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, A). 
Normally there are no resiniferous cells in the xylem though these 
occur in wounded specimens. The medullary rays are said to have 
numerous pits on the radial walls. Jeffrey has described in detail 
the wound-reactions of Brachyoxylon 3 : fig. 758, B represents part 
of a transverse section showing a mass of resiniferous parenchyma 
and a row of resin-canals stretching tangentially from the wounded 
area. Wood exhibiting the same normal and traumatic features is 
mentioned by Jeffrey from Martha's Vineyard and the Potomac 
formation. It is pointed out that Brachyoxylon differs from typical 
Araucarian wood in the frequent occurrence of circular and separate 
bordered pits and in the power of developing traumatic resin-canals. 

1 Saporta (94) B. p. 176, PL xxxi. 

2 Hollick and Jeffrey (09) B. p. 54, Pis. xin., xiv. 3 Jeffrey (06). 




Fig. 758. A, B, Brachyoxylon notabile; A, tracheitis of the secondary xylem; 
B, traumatic resin-canals. C, D, Araucariopitys americana; C, traumatic 
resin-canals; D, section of stem. E, F, Protodammara speciosa, cone-scales 
(x7), see page 250. G, BrachyphyUum crassum. (After Jeffrey.) 



In the combination of the Araucarian and the common type of 
tracheal pitting Brachyoxylon agrees with some other genera of 
Mesozoic woods, e.g. species of Cedroxylon, and in the formation of 
traumatic resin-canals it resembles Abies and other genera of 
Abietineae as also Sequoia. Jeffrey's view is that Brachyoxylon is 
undoubtedly Araucarian though in its wound-reactions it differs 
from the present representatives of the Araucarineae : in this 
respect he considers the genus to hold the same relation to recent 
Araucarineae as Sequoia holds in respect of its power of developing 
resin-canals in response to injury to other allied genera in which 
no such reaction occurs. Admitting the Araucarian arrangement 
of pits on some though by no means on all tracheids, the sum of 
characters hardly warrants the inclusion of Brachyoxylon in the 
Araucarineae : as in several other Mesozoic genera there is in some 
degree a mixture of characters indicative of a generalised type and, 
while Jeffrey sees in this combination evidence of the derivation of 
Araucarian Conifers from an Abietineous ancestry, I venture to 
regard the spasmodic recurrence of the Araucarian type of pitting 
as a partial persistence of characters inherited from an ancient 
Araucarian stock. 

Miss Holden 1 has described some wood from Cliffwood, New 
Jersey, which she refers to Brachyoxylon, differing from that 
described by Jeffrey in the presence of fibres in the secondary 
phloem, a feature associated generally with Cupressineae, Taxo 
dineae, and the Podocarpineae. In the Cliffwood material 
medullary rays are said to have smooth walls, a feature in which 
they differ from those of the Abietineae. 

BRACHYPHYLLUM. Brachyphyllum crassum Lesquereux. 

This name was given by Lesquereux 2 to a large branched 
vegetative shoot from the Dakota group, and for specimens, 
believed by Hollick to be identical with Lesquereux's species, from 
the Amboy clays Newberry proposed the name macrocarpum 3 : this 
specific name was not published and in the Amboy clay mono- 
graph 4 the designation Brachyphyllum crassum is adopted. The 

1 Holden, R. (14) p. 171. 2 Lesquereux (91) p. 32, PL n. fig. 5. 

3 Hollick in Newberry and Hollick (95) p. 51 (footnote). 

4 Newberry and Hollick (95) p. 51, PI. Vii. figs. 1 — 7. 





same name was given by Tenison- Woods 1 to a form of Brachy- 
phyllum from Queensland in 1883, the year in which Lesquereux 
published the name Thuites crassus 2 for the Dakota specimens 
afterwards transferred by him to Brachyphyllum. Neither author 
gave an illustration of the type-specimen and the Australian type 
is still unfigured, but Lesquereux's type is illustrated in the Dakota 
Flora. It would therefore seem reasonable to retain the specific 
name crassum rather than to adopt the designation macrocarpum 
revived by Hollick and Jeffrey. This species was found in the 
Middle Cretaceous beds in Staten Island in a condition which 
enabled Hollick and Jeffrey to supply important information with 
regard to anatomical characters. In their preliminary account 3 of 
the foliage-shoots these authors included in Brachyphyllum macro- 
carpum some wood exhibiting well-defined characters suggestive 
of Araucarian affinities, and Jeffrey 4 in another contribution speaks 
of the wood as that of Brachyphyllum. In a subsequent description 
of the Staten Island material the authors 5 state that they are no 
longer in a position to affirm that the fragments of wood belong 
to Brachyphyllum. This change of view is important as it was from 
the characters of the wood that some of the arguments in favour of 
an Araucarian affinity of the species B. macrocarpum were derived. 
The structural features of the foliage-shoots alone, though in some 
respects agreeing with those characteristic of recent Araucarineae, 
are not known in sufficient completeness to settle definitely the 
precise position of the species. 

The foliage-shoots have triangular, appressed, leaves identical 
in the form of the lamina with that in many Jurassic species of 
Brachyphyllum (fig. 758, G), and in the method of branching as 
also in the shape of the ultimate branchiets the specimens agree 
with typical representatives of the genus. The branches have a 
large pith containing nests of sclerous cells: external to the vascular 
tissue is a ring of resin-canals and a deep-seated periderm, beyond 
which are other canals belonging to the adnate leaves. Strands of 
stereome occur immediately below the epidermis and these are 
responsible for the longitudinal striations which often characterise 



1 Tenison-Woods (83). 2 Lesquereux (83) p. 32. 

3 Hollick and Jeffrey (06) p. 200. * Jeffrey (06). 

Hollick and Jeffrey (09) B. p. 55. 


impressions of Brachyphyllum leaves. In the younger branches the 
vascular tissue consists of separate bundles and a gap is formed on 
the exit of the single leaf-trace: the brace divides in the outer 
cortex into a number of fine strands ' which finally become lost in 
a continuous band of transfusion-tissue' beneath the palisade 
parenchyma 1 . No details are given with regard to the pitting of 
the tracheids or the structure of the medullary rays, but the 
authors state that the phloem showed no indication of the presence 
of any thick-walled fibres. In a later paper Jeffrey 2 states that in 
older stems of Brachyphyllum crassum the pits are flattened by 
mutual contact though in younger branches this feature is often 
not distinguishable. A single specimen is described as probably 
a cone of Brachyphyllum*: this is, however, much smaller than 
any cone previously recorded in connexion with Brachyphyllum 
shoots and the anatomical data do not furnish any proof of its 
morphological nature. 

The species is recorded from the Magothy formation 4 , ClifTwood, 
and from the Raritan formation; Berry also describes a large 
example from the Woodbine formation, Texas, as B. niacrocarpum 
var. formosum 5 . 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 6 . 

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 7 : 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 3 ) p. 770. 

3 Hollick and Jeffrey (09) B. p. 37, PI. ix. figs. 5, 6; PI. xi. fig. 3; PL xiv. fig. 3. 

4 Berry (05) p. 44; (06) p. 168; (ll 3 ) p. 81. 

5 Ibid, (12 3 ) p. 392, PI. xxx. 6 Seward (95) A. p. 218. 
7 Seward and Bancroft (13) p. 869, PI. i. figs. 2—4. 


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 1 and Thompson 2 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 

Brachyphyllum vulgare (Stopes and Fujii). % 

In their account of Upper Cretaceous plants from Hokkaido, 
Japan, Drs Stopes and Fujii 3 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 4 drew attention to the striking 
resemblance between Yezonia and Brachyphyllum crassum and 
fully justified his substitution of Brachyphyllum for the new genus. 
Dr Stopes 5 , 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 3 ) p. 768, PI. lxv. fig. 6. 2 Thompson (12 3 ) PI. vi. fig. 12. 

3 Stopes and Fujii (10) p. 23. 4 Jeffrey (10 3 ). 

6 Stopes (ll 3 ) 



nrr is 

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. 


This generic name, adopted by Endlicher 1 from Unger, is 
employed for fossil species believed to be nearly related bo 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 2 , 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 s , but these are of little or no value as trustworthy 
records. Similarly his species G. intermedius from the Pa toot beds 4 
founded on dimorphic, sterile, shoots affords no substantial evi 
dence of affinity to Taxodium or Glyptostrobus. 

Taxodites europaeus Brongniart. 

This species was first described by Brongniart 5 from Tertiar 
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 6 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. Pis. xvi., xx., xxn. 4 Ibid. (75) ii. PL lii. 

5 Brongniart (33). 6 Heer (55) A. Pis. xix., xx. 




says, it bears a striking resemblance to the existing Chinese type 
Glyptostrobus heterophyllus. Unger 1 describes well 
preserved specimens from Greece and the species 
is recorded, on the evidence of cones as well as 
sterile shoots, from Leoben 2 and other localities in 
Styria 3 , also from Miocene beds in Bohemia 4 . 
Laurent 5 figures examples from Aquitanian beds 
in the Puy-de-D6me and Saporta and Marion 6 
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 
Tertiary beds in Bosnia 7 and Nathorst 8 found the 
species in Arctic Ellesmere Land. The sterile frag- 
ments figured by Goeppert and Menge 9 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 10 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 Taxodiumeocaenicum 
and compared with the Floridan Conifer Taxodium distichum var. 
imbricataria Mett. Gardner points out with reason that specimens 
described by Heer 11 from Miocene beds in Greenland and Alaska 
as Glyptostrobus europaeus and from rocks of the same age in 

Fig. 759. Tdxo- 
dites europaeus. 
(After Heer; 
nat. size.) 

1 Unger (67) PI. I. 
3 Ibid. (90) PL i. 

Ettingshausen (88 2 ) PL ii. 

4 Ibid. (67 2 ) Pis. x., xi.; Velenovsky (81); Unger (52) PL xxxiv. 

5 Laurent (12) Pis. v., vi. 

6 Saporta and Marion (76) Pis. xxiii., xxxni. 

7 Engelhardt (12) Pis. xxxn., xxxvi., xxxvn. 8 Nathorst (15 2 ). 
9 Goeppert and Menge (83) A. PL xvi. 

10 Gardner (86) Pis. in., iv., xxiv. 

11 Heer (68) i. Pis. m., xlv.; (71) iii. Pis. I., m.; (77) i. Pis. XL, xn., xxvi.; (78) 
Pis. ix., xiii.; (83) Pis. lxx., lxxv. 


Spitzbergen as G. Ungeri may be fragments of Conifers mor 
closely allied to Sequoia. 

The fossils originally referred by Lesquereux 1 to Glyptostrob 
gracillimus from the Dakota group were afterwards transferred by 
him to Frenelites Reichii, at a later date removed by Newberry to 
Sequoia and finally described by Jeffrey 2 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 flora 
of Europe. 

Taxodites miocenicum (= Taxodium distichum miocenicum Heer). 

Numerous specimens of branches and some male and female 
flowers have been described by Heer 3 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 Taxodium. The shoots bear di- 
stichous, linear leaves, reaching a length of 2 cm. and 2 — 3 mm. 
broad. The absence of a decurrent base is spoken of as a character 
distinguishing Taxodium from Sequoia Langsdorfii. 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 4 consists of an axis bearing a few oval bud 
3 mm. long which may possibly be groups of microsporophylls, but 
the figures are far from convincing. Nathorst 5 speaks of the 
occurrence of foliage-shoots, flowers, and seeds in the Tertiary 
Taxodium shales of Spitzbergen. Ettingshausen 6 figures shoots 
and flowers from Bilin as Taxodium dubium which are in all 
probability closely allied to the recent species. Goeppert and 

1 Newberry and Hollick (95) PI. ix. 2 Jeffrey (11). 

3 Heer (68) i. Pis. n., xn., xlv. ; (71) Pis. m., iv., etc.; (77) Pis. xni., xxv. ; 
(78) Pis. viii., ix.; (83) Pis. lxx., etc. 

4 Heer (68) i. PI. n.; Schimper and Schenk (90) A. p. 294, fig. 203. 

5 Nathorst (IP) p. 223. 6 Ettingshausen (67 2 ) PI. x. 


Menge 1 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 2 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 3 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 4 and, despite tne 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 5 ; it is represented by deciduous twigs, cones, 
seeds, stumps, and knees 6 : 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 yubariensis Stopes and Fujii. This genus 
is founded on a single detached cone from Upper Cretaceous rocks 
at Hokkaido in Japan 7 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. Konishii; 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 
;he seeds had been shed at the time of f ossilisation : three pro- 

1 Goeppert and Menge (83) A. PI. xvi. figs. 227—229. 

2 Lingelsheim (08) p. 34. 

3 Engelhardt and Kinkelin (08) PI. xxm. figs. 19—21. 

4 Squinabol (92) PI. xvi.; Berry (09) p. 22, fig. 1. 

5 Berry (07 2 ); (09 2 ); (12 4 ); (15). 6 Ibid. (09 2 ), figs. 1, 2. 
7 Stopes and Fujii (10) p. 45 ? PI. v. figs. 27—34. 


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 1 
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, PI. vn. figs. 23—27. 


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 1 , in his reference to the genus in an account 
of the Sezanne 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 cyclotoxon Debey and Ettingshausen. 

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 2 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 3 he de- 
scribed well preserved examples as Moriconia 
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) 4 , Staten 
Island 5 and Block Island. As Hollick points 
out, a large impression included by Heer in Fig.760. Moriconia cy- 
Moriconia should rather be referred to Brachy- dotoxon. (After New - 
phyllum : in the arrangement of the leaves and in e "" y . an , ° 1C ; 

r a ° . nat. size.) 

the form of the short and blunt lateral foliage- 
shoots Moriconia agrees closely with some examples of Brachy- 
phyUum crassum 6 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 median 
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, PI. xxvi. fig. 18. 

3 Ibid. (82) i. p. 49, PL xxxm.; (83) Pis. liii., liv. 

4 Newberry and Hollick (95) p. 55, PI. x. 

5 Hollick (06) PI. in. ; Berry (03) PL xlvih. 6 Berry (06) PL ix. fig. 1. 



The species Cryptomerites divaricatus, for which Bunbury 1 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 2 figured a sterile piece of foliage-shoot from Eocene leaf- 
beds in County Antrim as Sequoia du Noyeri which Gardner 3 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 
Cryptomeria Sternbergii (Goepp.), the specific name being adopted 
because he considered some examples figured by Ettingshausen 4 as 
Araucarites Sternbergii Goepp. to be identical with the Irish fossils, 
though most of the specimens described by authors, including the 
author of the species, as Araucarites Sternbergii 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 Cryptomeria. 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 Sternbergii 

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, Pis. x., xx., xxi. 
4 Ettingshausen (55) PL v. 


specimens figured by Ettingshausen from Monte Promina as Aran- 
carites Sternbergii and some of the impressions from Greenland re- 
ferred by Heer 1 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 Cryplomerites 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. 

Cryptomeriopsis antiqua Stopes and Fujii. The name Crypto- 
meriopsis 2 - 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 3 . 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. antiqua 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- specie^. Prof. Jeffrey 4 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 5 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. PI. n. figs. 1 — 4. 

2 Stopes and Fujii (10) p. 52, PL i. fig. 11; PL vi. figs. 35-41. 

3 Suzuki (10) p. 185. i Jeffrey (10 3 ) p. 771. 5 Stopes (ll 3 ) 



It has already been pointed out that there is good reason for 
treating the three existing genera Callitris, Widdringtonia, and 
Actinoslrobus as members of a distinct family. The genus Tetra- 
clinis, as Saxton 1 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, Tetraclinis 
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, Callitris, 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 
Callitris and Calythrix, 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 Callitris, 
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 (13 2 ); (13 3 ). 


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 1 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 2 
from the London Clay as Cupressites globosus and C. elongatus but 
Gardner 3 , who examined the original specimens, is sceptical as to 
their connexion with the Callitrineae. Ettingshausen 4 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 Widdring- 
tonites keuperianus Heer 5 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 6 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 Bowerbank (40) p. 52, PI. x. 

3 Gardner (86) p. 20. 

4 Ettingshausen (72) p. 164, PL n. 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 


cones were found. Similarly Widdringtoniles gracilis Sap. and 
W. creyensis Sap. from the Corallian and Kimeridgian of France 1 
respectively are founded solely on sterile shoots. The specimen 
figured by Eichwald 2 from Jurassic rocks on the southern border 
of the Caspian sea as Widdringtoniles denliculatus 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 3 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. 

Callitrites Reichii (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 4 as Frenelites Reichii on some branched shoots from Cre- 
taceous rocks in Saxony and afterwards described by Heer 5 from 
the Patoot beds of West Greenland under the generic name 
Widdringtoniles 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 6 in his monograph except two small 
examples which it is suggested may be immature microstrobili. 
Velenovsky 7 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 Aclinostrobus and those described by 
Berry as Widdtingtonites 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 8 from the 

1 Saporta (84) Pis. 201, 202. 2 Eichwald (68) p. 43, PI. iv. fig. 9. 

3 Zeiller (11) PL n. fig. 6. 4 Ettingshausen (67) p. 246, PL I. fig. 10. 

5 Heer (82) i. p. 13, PL lii. figs. 4, 5. 

6 Newberry and Hollick (95) PL vm. 

7 Velenovsky (85) B. p. 27, Pis. vm., x.; (87) figs. 14—16. 

8 Krasser (96) B. p. 126, Pis xiv., xvn. 


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 1 regard the shoots referred by them to Widdringlonia 
Reichii 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 2 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 3 as Taxodium ramosum, but in the absence of cones a 
definite determination of affinity is hardly possible. 

Callilriies subtilis (Heer). 

Founded by Heer 4 on slender twigs bearing spirally disposed, 
appressed, leaves from the Cretaceous beds of Atanekerdluk in 
Greenland and described by Newberry 5 from the Amboy clays. 
Hollick 6 and Berry 7 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 8 . 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. Reichii and the two species have often been confused ; 
also to Cyparissidium minimum as figured by Velenovsky 9 , 
Juniperus macilenta Heer 10 and Widdringionites fascicularis Holl. 11 

1 HoUick and Jeffrey (09) B. p. 29, Pis. v., vin., xx.; Hollick (06) p. 44, PI. iv. 
figs. 6—8. 

2 Berry (ll 3 ) p. 87, PL viii. 

3 Fontaine (89) B. p. 251, Pis. cxxm., cxxiv., etc.; Berry (ll 4 ) p. 302. 

4 Heer (74) B. PI. xxvm. fig. 1. 

5 Newberry and Hollick (95) p. 57, PI. x. figs. 2 — 4. 

6 HoUick (06) p. 45, PI. iv. figs. 2—5. 7 Berry (12 2 ). 

8 Ibid. (14) p. 25, PI. xi. figs. 14—17. 

9 Velenovsky (85) B. PL ix. figs. 6, 7; PI. x. fig. 4. 

10 Heer (75) ii. PL xxvm. fig. lc. 

11 Hollick (06) PL iv. fig. 1. 






A. cone of five valves. 

B, C, two views of a sec- 
tion of a f our- valved cone. 
(British Museum, drawn 
from specimens described 
by Gardner; nat. size.) 

Callitrites curia (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 1 , 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 2 cm. Fig. 761, 
A shows a cone of five valves and B, C, are 
two views of a section of a cone consisting 
of four valves. A similar type described 
by Gardner as Callitris Ettingshauseni 2 , 
also from Sheppey, is represented by glo- 
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 3 is incorrect in recording Callitrites curta from 
the Isle of Wight. 

Callitrites Brongniarti (Endlicher). 

This species, first described by Brongniart 4 as Equisetum 
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 5 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 6 subsequently described good specimens as 

1 Gardner (86) p. 21, PI. ix. figs. 7, 21. 2 Ibid. PI. ix. figs. 1—6. 

3 Ettingshausen (79) p. 392; (80) p. 231. 

4 Brongniart (22) A. p. 329, PI. v. fig. 3; Endlicher (47) p. 274. 

5 Unger (47) p. 22, Pis. VI., vn. 6 Ibid. (67) p. 42, PI. i. figs. 1, 2. 




Callitris Brongniarti from Miocene beds in Euboea, but Saporta 1 
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 2 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. Ettingshausen 3 states that the species is very 
abundant at Haring in the Tyrol: that author describes some 

762. A, A', B, Callitrites Brongniarti. C, Callitrites helvetica. D, Callitrites 
europaea. (A, A', after linger; B, after Saporta; C, after Heer; D, after 
Engelhardt and Kinkelin.) 

sterile shoots from Eocene beds in New South Wales as Callitris 
prisca* which he compares with C. Brongniarti. Well preserved 
shoots are described by Watelet 5 from the Paris Basin. Engel- 
hardt 6 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 7 from the Pliocene beds of the Frankfurt 

1 Saporta (68) p. 316. 

2 Ibid. (62) p. 209, PI. n. fig. 6; PI. m. fig. 1; (65 2 ) p. 39, PI. I. fig. 6. 

3 Ettingshausen (55) p. 34, PI. v. figs. 7—35. 

4 Ibid. (86) p. 95, PI. vm. figs. 3, 4. 

6 Engelhardt (85) p. 314, PI. vhi. fig. 32. 

7 Engelhardt and Kinkelin (08) PI. xxm. fig. 5. 

Watelet (66) A. PI. xxxn. 


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. antiqua (Sap.) from Provence 1 , repre- 
sented by shoots with spiral, sub-opposite or opposite appressed 
scale-leaves and by globular cones with four valves. As Solms- 
Laubach says 2 , the cones agree closely with those of Widdringtonia, 
though it would be difficult to decide between that genus and 

Some good specimens are figured by Heer 3 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 4 as three species of Widdringtonites, 
and in one case, W. legitimus, 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 5 frOm Tertiary beds near Frankfurt are 
founded on unconvincing specimens. Engelhardt and Kinkelin 6 
describe pyramidal cones with 5 — 6 valves 1 — 15 cm. long (fig. 
762, D), which they refer to Frenelites europaeus, from the Upper 
Pliocene 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. 


Schenk 7 instituted this generic name for specimens originally 
described by Ettingshausen 8 from Wealden beds in Silesia as 

1 Saporta (62) PI. n. fig. 7; (02 2 ) PI. m. fig. 3; (65 2 ) PL I. fig. 4; (73) PI. n. 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. 
fi Engelhardt and Kinkelin (08) PI. xxm. fig. 5. 7 Schenk (71) p. 13, PI. i. 
8 Ettingshausen (52) p. 26, PL i. figs. 6, 7. 



Thuiles 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 1 (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 2 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 3 to Frenelopsis 
gracilis Hollick and Jeffrey 4 , 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, PI. xi.; Thompson (12 3 ) PI. v. 

2 Heer (75) ii. p. 73, PL xvra. figs. 5—8; (82) i. p. 7, PI. n. figs. 1—3. 

3 Newberry and Hollick (95) p. 59, PI. xm. figs. 1—3. 

4 Hollick and Jeffrey (09) B. p. 26. . 


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. 

j Frenelopsis Hoheneggeri (Ettingshausen). 
| Frenelopsis occidentalis Heer 1 . 

Though the specimens referred to these two species may 
correctly separated their close agreement in habit poiuts 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. Hoheneggeri 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 ol 
4 — 5 accessory cells overarching the stomatal depression 2 . 

Specimens described from Bohemia by Velenovsky 3 as F. 
bohemica resemble F. Hoheneggeri 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 4 : 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 F. Hoheneggeri (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 5 . 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 xn. figs. 3—7; Saporta (94) B. pp. 139, 199, 214; Pis. 


2 Thompson (12 3 ). 3 Velenovsky (88) figs. 1—3, 10. 

4 Fontaine (89) B. p. 215, Pis. 95—101; Berry (11) p. 422, Pis. lxxi., lxxii. 

5 Berry (10 2 ). 




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 2 
made the type of a new genus Pseudofrenelopsis, but the features 


Fig. 763. A, B, Frenelopsis Hoheneggeri. C, D, F. raniosissima. E, F. occiden- 
talism (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 BecMesia anomala Sew. 3 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, Pis. cxi., cxii., clxviii. 

2 Nathorst (93) p. 52, figs. 6—9. 3 Seward (95) A. p. 179, PI. xiv. 


Further light will no doubt be thrown on the nature of Frene 
lopsis when the results of the investigations of Hollick and JefTre 
are published: ib 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 Callitrinea 
than to any other plants. The genus ranges from Wealden t 
Cenomanian rocks. 


In view of the restricted range of the two surviving species oi 
Sequoia and the peculiarities of the genus, to which expression is 
given by the institution of the family-name Sequoiineae 1 , 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 denned to enable us to discriminate 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- 
bachii affords no real evidence of affinity to the recent genus and 
the same remark applies to specimens included in Heer's genug 
Sphenolepidium and compared by authors with Sequoia. Some of 
the imperfectly preserved Jurassic cones agreeing superficially with 
those of Sequoia may well belong to species of Sequoiineae. 

Though in the majority of instances Jurassic and Cretaceous 
records do not prove the former presence of Sequoia 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 2 . 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. 


genial south on the American continent than in Europe where the 
retreat from Arctic regions ended in extinction. 

Penhallow 1 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 2 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. 
«Jn 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 3 described two species from 
Eocene beds in the North-West Territory as Sequoia Langsdorfii 
and S. Burgessii, both of which were previously described by 
Dawson but assigned by him to different positions. In the wood 

I 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 

I form on the outer face of the summer-wood. The pitting of 
the medullary-ray cells is not described. A peculiar feature in 
S. Burgessii, if the wood is correctly referred to Sequoia, is the 
occurrence of two kinds of medullary rays, uniseriate and fusiform, 
the latter containing resin-canals. No resin-canals occur in the 

I 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 4 Sequoia 
magnifica. A few of the trunks reach a diameter of 6 — 10 ft. 
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, Pis. civ., cv., ex., cxi., cxvu. 




and a height of 30 ft. (fig. 764). The details are imperfectly pre- 
served: a few of 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 

Fig. 764. Petrified tree in the Yellowstone National Park (Sequoia magnified 
Knowlton). (From a photograph kindly supplied by Prof. Knowlton.) 

scattered as in Cupressinoxylon: 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, 


the main Island of Japan, recently described by Yasui 1 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 
Sequoiineae 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 Sequoia. 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 Sequoiites and others which may reasonably be so named. 

SEQUOIITES. Brongniart. 

I Sequoiites problematica (Fliche and Zeiller). 
This species, originally described as Sequoia problematica 2 , 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 3 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 4 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. 

L 1 Yasui (17). 2 Fliche and Zeiller (04). 3 Zeiller (00). 






Sequoiites giganteoides (Stopes). 

This species, under the name Sequoia giganteoides, has recently 
been founded by Dr Stopes 1 on a small petrified fragment of a ver; 

Fig. 765. Sequoiites concinna. Foliage-shoot from the Senonian of Greenland. 
(Stockholm Museum; nat. size.) 

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, PI. ii. text-fig. 16. 


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. 

Sequoiites concinna Heer. 

Heer 1 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 Sphenolejridium 
Sternbergianum from English and other Wealden rocks, as also with 
the shoots of Sequoiites 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 2 from Upper Cretaceous beds in Pike 
County, Arkansas. 

Conites. Conites Gardneri (Carruthers). 

Carruthers 3 described a cone and a piece of vegetative shoot 
from the Gault of Folkestone as Sequoiites Gardneri but neither 
specimen affords any satisfactory evidence of relationship with 
Sequoia. The shoot is of the Pagiophyllum type, and the cone, 
2-5 x 1-5 cm., consists of spirally disposed scales with four-sided 
rhomboidal 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 Sequoiites. 
A specimen figured by Lange 4 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. Pis. li., lii., etc. 

2 Berry (17) p. 172, PL vn. figs. 1—5. 

3 Carruthers (69 2 ) p. 7, PI. I. figs. 7, 8. 

4 Lange (90) PL xxxn. fig. 7. 




Conites ovalis (Carruthers). 

The type-specimen from the Gault of Folkestone 1 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. 

Sequoiites Holsti Conwentz ex Nathorst MS. 
This species 2 , 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 
favourable to Nathorst's adoption of the 
generic name Sequoiites. The species is, 

however, not above suspicion as a record of a Conifer closely 
allied to Sequoia. 

Sequoiites Langsdorfii (Brongniart). 

Brongniart 3 instituted this Tertiary species under the generic 
name Taxites, and Heer 4 in his description of foliage-shoots from 
Miocene beds in Switzerland adopted the designation Sequoia. In 
habit S. Langsdorfii is practically identical with Sequoia sempervirens 
and by many authors it is spoken of as the direct ancestor of the 
recent species. Under this species Schimper 5 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 

Fig. 766. Conites oralis. 
(After Carruthers; nat. 
size. ) 

1 Carruthers (71) p. 3, with text-figure. 

2 Conwentz (92) p. 28, Pis. in., iv., viii. 

3 Brongniart (28) A. p. 108. 
5 Schimper (72) A. p. 216. 

4 Heer (55) A. p. 54, Pis. xx., xxi. 


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. 
Sequoiites Langsdorjii is recorded by Gardner 1 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 2 , from Miocene beds in Greece 3 
where it is represented by foliage-shoots and cones, from Italy 4 , 
Germany, and other European localities. A very similar form, 
originally named by Brongniart 5 Taxites Tournali, is described by 
Gardner 6 from the Middle Bagshot^beds of Bournemouth and with 
it he unites S. Hardti 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- 
nali is recorded by Saporta 7 who figures branches and cones from 
Eocene strata in Provence. Laurent 8 figures fragmentary speci- 
mens, which he refers to S. Langsdorjii, from the Aquitanian series 
in the Puy-de-D6me. 

Sequoiites Langsdorjii is very abundant in Arctic Miocene rocks: 
Nathorst 9 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. - Ettingshausen (57) PL i. fig. 3. 

3 Unger (47) PL u. figs. 17 — 23. Saporta (68) refers S. Langsdorjii as figured by 
Unger to S. Tournali. 

1 Squinabol (92) p. 26, Pis. xv., xvi. 5 Brongniart (28) A. p. 108. 

8 Gardner (86) p. 40. 7 Saporta (65 2 ) PL n. fig. 1. 

8 Laurent (12) p. 65. 9 Nathorst (IP) p. 225. 

s. iv 23 




fine teeth on the edge of the lamina 1 . Similar teeth are stated bv 
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 2 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 3 figures stexile 
shoots from the Sichota- Alin moun- 
tains. Penhallow 4 records the spe- 
cies from British Columbia and *"*' 767 .* 8eq^Lang«hrfii (Fn 

specimens m 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 5 , and the species 
is said to be one of the most abundant and widely distributed 
types in the Yellowstone National Park 6 . 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 2 ) p. 10, PL i. figs. 1—15. 

2 Heer (68) Pis. EL, xx., lv., etc.; (71) Pis. xl., xliii. 
(77) i. Pis. xii., xiii., xxv.; (7*) v. PL I.; (82) i. PL liii. 

3 Palibin (04) PL n. 

4 Penhallow (02) pp. 44, 68; (03) p. 41. 

5 Schmalhausen (90) PL I. figs. 2—11. 

6 Knowlton (99) p. 682. 

etc.; (75) iii. PL n. 


ments of an ancient lake mixed with volcanic ash contain many 
plant and insect remains and Prof. CockerelPs careful investigations 
have led to the discovery of several new types 1 . Staub 2 , 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. Langsdorjii is said to occur in beds of Lower Pliocene 
age in France 3 and a minute cone, only 2 by 1-9 mm. has recently 
been described by Mr and Mrs Reid from Pliocene deposits in 
Holland 4 . 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. 

Sequoiites Couttsiae (Heer). 

This species was founded by Heer 5 as Sequoia 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 6 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 7 

I 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), (08 2 ), (08 3 ). 2 Staub (87) B. PL xix. p. 249. 

3 Depape (13). i Reid, C. and E. M. (15) p. 55, PL i. fig. 13. 

5 Heer (62) p. 1051, Pis. lix.— lxi. 6 Beust (85) PL m. figs. 1—8. 

Reid, C. and E. M. (10) p. 170, PL xv. figs. 23—27. 



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 


Fig. 768. Sequoiites Couttsiae. Twigs A, B, and cone-scales C, D, from Bovey 
Tracey. (Photographs by Mr and Mrs Clement Reid; x 3.) 

a ring of four or occasionally five cells. Gardner 1 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 Sequoia sempervirens while 
the vegetative branches agree with S. gigantea. There are 3 — 5 seeds 
on each scale. Several examples of Sequoiites Couttsiae are figured 
by Heer 2 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, PI. vi. 

2 Heer (68) p. 94, Pis. m., viil, xlv.; (71) Pis. xl., xliii., etc.; (83) PI. lxviii. 


S. Whymperi. This more robust form occurs also in Spitzbergen, 
on the Mackenzie River, and elsewhere. Knowlton 1 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 2 from the Western Territories as Sequoia affinis bear 
a close resemblance to S. Couttsiae. The latter species is recorded 
by Penhallow 3 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 4 , and Saporta 5 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 6 from Eocene beds in South- West 
Russia as S. Couttsiae, though possibly correctly named, are not 
convincing. Palibin 7 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 8 , be examples of 
S. Couttsiae. 

GEINITZIA. Endlicher. 

The name Geinitzia was given by Endlicher 9 to a piece of sterile 
shoot from Lower Cretaceous strata in Saxony previously figured 
by Geinitz 10 as Araucarites Reichenbachii (fig. 769), and in the new 
genus was also included Cryptomeria primaeva Corda 11 . 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 Geinitzia 

I Knowlton (99) B. p. 681. 2 Lesquereux (78) B. PL lxv. 

3 Penhallow (02) p. 50. 4 Bleicher and Fliche (92) p. 382. 

5 Saporta (65*) PI. n. 6 Schmalhausen (83 2 ) Pis. xxxn., XXXVI. 

7 Palibin (01) p. 499. 8 Gardner (86) p. 39. 

9 Endlicher (47) p. 280. 10 Geinitz (42) PI. xxiv. fig. 4. 

II Corda in Reuss (46) B. PI. xliii. figs. 1—11. 




was applied to branches without any recognisable cones. In 1868 
Heer 1 figured specimens from the Kome beds in Greenland which 
he believed to be identical with Araucarites Reichenbachii 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 

Fig. 769. Geinitzia Reichenbachii. 
(After Geinitz,) 

Fig. 770. Geinitzia Reichenbachii. 
(After Heer; nat. size.) 

Sequoia for Araucarites on the ground that some cones in the 
Tubingen Museum from Lower Cretaceous beds in Moravia, 
attached to branches apparently identical with Araucarites Reichen- 
bachii, presented a very close resemblance to those of recent 
Sequoias. The Moravian specimens, which he afterwards figured 2 , 
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 3 from Greenland as Sequoia 

1 Heer (68) PI. xliii. 2 Ibid. (69) PI. i. 

3 Ibid. (75) ii. Pis. xn., xx., xxxiv., etc. 


Reichenbachii 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 1 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 2 . On the other hand the name Sequoia Reichen- 
bachii is applied in some cases to fragments of sterile branches un- 
accompanied with cones 3 : in one instance 4 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 

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, it is 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 5 that the branches from Lower 
Cretaceous or Upper Jurassic rocks in Spitzbergen figured by 
Heer 6 as Sequoia Reichenbachii are examples of Elatides curvifolia 
(Dunk.). The genus Elatides 7 is characterised by cones differing in 
their flatter scales and more elongated form from those usually 
assigned to Sequoia 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 8 serves 

1 Velenovsky (85) B. Pis. vin., ix. 

2 Ward (99) B. Pis. 165, 166; Hollick (06) PI. n. fig. 40; PI. in. figs. 4, 5. 

3 Krasser (96) B. PI. xvn. fig. 14; Schenk (71) PL xxiv. figs. 6, 7. 

4 Lange (90) p. 660. ■ Nathorst (97) p. 35. 

6 Heer (75) ii. Pis. xxxvi.. xxxvu. 7 See page 270. 8 See page 274. 


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 1 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 Reichenbachii 
but longer in form. Subsequently Heer 2 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 3 also gives good drawings of Geinitzia formosa. A well pre 
served cone very like Heer's G. formosa was described by Newberry 4 
from the Amboy clays as Sequoia 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 ii 
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 Jeffrey 5 obtained good cones from th 
Matawan formation apparently identical with Sequoia gradllima 
(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 ). 2 Heer (71 2 ) p. 6, Pis. i., n. 

3 Schimper and Schenk (90) A. p. 299. 

4 Newberry and Hollick (95) p. 50, PL ix. figs. 1—3. 5 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 
Paracedroxylon. 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 1 describe some sterile branches as Geinitzia Reichenbachii 
(fig. 806, D ; page 437) which agree closely with specimens referred by 
authors to Sequoia Reichenbachii 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 Sequoia, 
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 
Brachyoxylon 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 Pseudogeinitzia. 

EUGEINITZIA. Hollick and Jeffrey. 

Eria froxima Hollick and Jeffrey, 
scales on which this species is founded 2 closely resemble 
recent species of Sequoia and Geinitzia gracillima. The 
1 Hollick and Jeffrey (09) B. p. 38. 


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 Geinitzi 

PSEUDOGEINITZIA. Hollick and Jeffrey. 
Pseudogeinitzia sequoiiformis Hollick and Jeffrey. 

A special generic name 1 is given to some four-sided scales on 
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. Reichenbachii, exhibit certain 
Araucarian tendencies and that the cone, Geinitzia gracilliuxi. a> 
also the detached cone-scales, Eugeinitzia and Pseudogeinitzu 
cannot be included in Sequoia. 

Until more is known of the morphological nature of the coin 
described by Heer and other authors as Sequoia Reichenbachii, 
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 Sequoia, much too freely used 
by palaeobotanists, has in some cases 2 been applied to cone-bearing 
branches that are almost certainly identical with Sphenolepiditm/ 
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, PI. x. fig. 14; PI. xxv. fig. 4. 

2 Heer (69) p. 11, PI. i. figs. 10—13 (' Sequoia faMigiata'). 



Heer 1 instituted Sphenolepidium in place of Sphenolepis, pro- 
posed by Schenk 2 for Wealden Coniferous branches, because of the 
previous use of the latter name by Agassiz for a genus of fishes. 
Berry 3 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 Avith 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 BrachyphyUum. The genus is charac- 
teristic of Wealden or Lower Cretaceous strata and is represented 
in several European districts and in North America. 




Sphenolepidium Stembergianum (Dunker). 

This species was originally described by Dunker 1 from North 
Germany as Muscites Stembergianus and by later authors placed 
in Araucarites, Widdringtonites, and other genera 2 . It is impossible 
to determine the specific limits of this species 3 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. Stembergianum . 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. Stembergianum are almost certainly 
examples of Elatides curvifolia. 

Sphenolepidium Kurrianum (Dunker). 

Dunker 4 originally adopted the generic name 
Thuites; later authors preferred BrachyphyJhnn. 
Widdringtonites, Araucarites and other names. 
Fontaine 5 , who records this species from the 
Potomac formation, includes in his genus Athro- 
taxopsis specimens which cannot be distinguished 
by any features of morphological importance 
from Sphenolepidium Kurrianum . The leaves are 
ovate, more or less appressed, agreeing with 
Brachyphyllum or in some examples intermediate 
between the type of foliage assigned to Pagio- 
phyllum and Brachyphyllum. The Wealden speci- 
men reproduced in fig. 771 is placed in Spheno- 
lepidium because of its association with branches, 
identical in habit, bearing cones; if found 

Fig. 771. Spheno- 
lepidium Kurria- 
num. From the 
Wealden of Sus- 
sex. (British Mu- 
seum, V. 2303; 
£ nat. size.) 

as an iso 


1 Dunker (46) A. PL vn. fig. 10. 

2 For references, see Seward (95) A. p. 205; Berry (ll 4 ) p. 293. 

3 Seward (ll 2 ) p. 685. 

4 Dunker (46) A. p. 20. 5 Fontaine (89) B. Pis. cxxxv., etc. 


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 S. 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. 


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 1 for impressions of shoots 
from Russia bearing whorled linear leaves which he believed to be 
closely allied to or generically identical with Sciadopitys. Zeiller 2 , 
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 3 says, they may be compared with several 
recent genera including Sciadopitys, but without anatomical data 
accurate determination is impossible. It is stated by Schenk 4 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 5 describe some single 
leaves preserved in Baltic amber as Sciadopitytes linearis and 
S. 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. - 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, PI. xm. figs. 117—123. 

les oi 



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 1 but the occurrence of such features in other 
recent genera precludes a definite reference to any one type. 


This name has recently been revived by Halle 2 for two species 
Cretaceous leaves from Greenland, one of which, Pintis Crameri 
Heer, though compared by Schenk with the leaves of Sciadopitys, 
was not actually included in Sciadopitytes, while the other is a new 
species, Sciadopitytes Nathorsii. Halle describes these leaves as 
'Conifer-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 Sciadopitytes does not imply more than the 
facts support. The leaves named by Heer 3 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. PI. xliv. figs. 7—18; (75) ii. PI. xxm. pp. 9—15; Halle (15) 
p. 509, PI. xm. figs. 1—13. 




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 Nathorsti 1 , 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 xn. figs. 16—29. 



The 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. xliv : 
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 substantia) 
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 


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. in. p. 298) in the partially decayed 
wood of Antarcticoxylon 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 1 , 

I 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 2 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. 

S. IV 24 


Heer 1 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 2 described a cone as 
Abies Loehri, and this has more recently been identified as Keteleeria 
(fig. 786, C, p. 394) by Engelhardt and Kinkelin 3 on the strength 
of its external resemblance to K . Davidiana. The reference to Abies 4 
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 5 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, 
Larix, 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 Pityites 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. PI. ix. figs. 6—8. 

2 Geyler and Kinkelin (90) p. 16, PI. i. figs. 13—15. 

3 Engelhardt and Kinkelin (08) p. 216, PL xxvi. fig. 7. 

4 Goeppert and Menge (83) A. PL xm. figs. 107—110. 

5 Endlicher (47) p. 283. 


terms suggested by Nathorst and mentioned below. Pityites is, 
however, appropriate for such specimens as those represented in 
figs. 772, 773 which show a direct connexion between cones and 

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 1 , 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 Pityanihus 
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 2 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 3 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 Pityites is 
employed for specimens of a more complete kind. The name 
Abietites has often been used for vegetative shoots and cones 4 
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 E.g. Geinitz (80) p. 12; Fontaine in Ward (05) B. PI. lxviii. figs. 14—17; 
Thomas (11) PI. iv. fig. 16; PI. v. figs. 1, 2. 



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 made 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 o 
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. recen 
Pines and other Conifers. The winged seeds of Agathis differ in th 
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 Abietineou 
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 o 
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 an 
later Mesozoic strata, but we have no means of determining in th 
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 1 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 2 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 3 from the culm of Altendorf as Pinites antecedents, 

1 Renault (93) A. PL xxxn. fig. 1; (96) A. p. 377. 

2 Page 101. 3 Stur (75) A. PI. xiv. fig. 4. 


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 Phoenicofsis, 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 1 
having typical Abietineous features, a fact of importance in relation 
to the widely spread Palaeozoic woods agreeing in essentials with 
the Araucarian type. 

Pitijites 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 2 : 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 Finns Strobus, P. excelsa (fig. 773; cf. fig. 704), Picea 
and Abies', they agree closely with Pityostrobus Carruthersi (Gard.) 
as also with P. Andraei (Coem.) 3 from Lower Cretaceous rocks in 
Belgium and with the smaller cones from the Potomac formation 
described by Fontaine 4 as Abietites ellifticus. The preservation is 
not sufficiently good to show the number 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 Larix 
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. 5 from the Cretaceous beds 
of Kreischerville. Dr Stopes 6 , 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, Pis. xvin., xix. 3 Gardner (86 2 ). 

4 Fontaine (89) B. PL cxxxm. figs. 2—4. 

5 Hollick and Jeffrey (09) B. p. 19. 

6 Stopes (15) p. 157. 




Pityites (Pinites) eirensis sp. nov. 

In his account of petrified material from Franz Josef Lane 
Solms-Laubach 1 describes sections of a Pinus-like leaf from Bell 

Fig. 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, PI. i. fig. 14; PL n. fig. 3. For evidence as to 
geological age, see Newton and Teall (97), (98); Nathorst (99). 




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 Finns 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 

Fig. 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 



possessing them is mentioned in Chapter xliii. The leaf for which 
the specific name eirensis (from Eira harbour) is proposed affords 

Fig. 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. 

xlvih] pityocladtjs 377 


Under this name, used as a subgenus of Pinites, Nathorst 1 
includes branches bearing short shoots similar in habit to those of 
Cedrus and Larix. 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 

Pityocladus Nathorsti Seward. 

In his description of Schizolepis Follini Nath. 2 from Rhaetic 
rocks in Scania, Nathorst includes not only cones with lobed scales 
characteristic of Schizolepis but leaves and branches. Solms- 
Laubach 3 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 Palsjo, 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 4 
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 Follini 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 

1 Nathorst (97) p. 62. 2 Ibid. (78) B. p. 28. 




Pityocladus longifolius (Nathorst). [And Pityophyllum longifolium 
This species was first described by Nathorst 1 from Rhaetic beds 
in Scania as Taxites longifolius and afterwards as ? Cycadites. 


Fig. 776. Pityophgt- 
lum long ifol iio 
(Nat. size.) 

Fig. 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.) 

Moller 2 adopted Nathorst' s generic name Pityophyllum for de- 
tached leaves and that designation should be used for specimens 

1 Nathorst (78 2 ) B. p. 50. « Moller (03) p. 40, PI. vi. 


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 Pityophyttum 
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 Follini. An accurate 
specific delimitation of Pityophyttum 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 Turkestan 1 , South Russia 2 , Oregon 3 , Spitz- 
bergen 4 , and elsewhere. 

Pityocladus Schenki Seward. 

Schenk 5 also refers to Schizolepis several specimens of branches 
and leaves as well as cones which he includes in Schizolepis Braunii 
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 6 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 Braunii. 

Pityocladus kobukensis Seward. 

This species originally described from the Jurassic beds on the 
Kubuk River in Chinese Dzungaria as Pinites 1 is founded on 

ranches bearing short shoots almost identical with P. Schenki. 

hort 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 (07*) B. p. 32. 2 Thomas (11) p. 78, PL vn. figs. 58—61. 

3 Fontaine in Ward B. (05) PL xxxv. 4 Nathorst (97). 

5 Schenk (67) A. PL xliv. figs. 1—4. 6 Ibid. PL xliv. fig. 2. 

7 Seward (11) p. 54, PL iv. figs. 47—51; PL v. fig. 65. 




branches are associated numerous needles, 1 mm. broad and at 
least 5 cm. 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 1 from Li- 
assic strata as Halockloris baruthina 
Ett. A branch with short shoots from 
Jurassic rocks in Amurland described 
as Pinites sp. cf. P. JcobtiJcensis 2 may be 
specifically identical with the Dzungaria 
fossils: an example of the same type 
lent to me by Dr Krystofovic 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) cf. P. kobukensis 3 . 

Similar though smaller specimens Fig. 77 
of Abietineous short shoots are de- 
scribed by Nathorst 4 as Pinites (Pityo- 
cladus) spp. a and b from Upper Jurassic beds in Spitzbergen, and 
compared by him with Pityites Solmsi. 


This name 5 is applied to detached leaves of needle-like form like 
those of recent Pines or to long linear leaves broader and natter 
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 6 and recent 

1 Ettingshausen (52) B. PI. n. fig. 4. 

2 Seward (12 3 ) PL in. 3 Krystofovic (15) PL vi. fig. 9. 

4 Nathorst (97) PL in. figs. 28—30, PL iv. figs. 13, 14, 23. 

5 Ibid. (97) p. 62. 6 See page 374; also Nathorst (97) PL v. figs. 1—10 

Pityocladus kobuken 
sis; b, branch-scar. (After 
Seward: nat. size.) 


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- 
lotaxus 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. Nordenskioldi (Heer) and similar 
forms are characteristic fossils in Jurassic and Cretaceous strata. 

Pityophyllum Lindstromi Nathorst. 

Under this name Nathorst 1 includes leaves described by Heer 
from Cretaceous strata in Greenland as Pinus Quenstedti and 
P. Peter seni, 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 

I 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 2 from Cretaceous and Jurassic rocks. 
Pityophyllum Nordenskioldi 3 (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 4 : 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. 


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, Pis. v., vn.; (99) p. 20. 

2 Ibid. (97) pp. 41, 68, Pis. v., vi. 3 Heer (78) ii. PL n.; Nathorst (97) p. 18. 
4 For references see Moller (03) p. 39; Seward (11) p. 53; Krystofovic (10) 

PL in. fig. 10; Thomas (11) p. 78. 


that Nathorst's term is more appropriate for specimens which do 
not afford evidence 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 1 speaks of this Kimeridge cone from Dorsetshire as 
the oldest example of a Pine-cone. It is represented by a single 
imperfectly preserved specimen, 2x2 cm., 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 2 . 

Pityostrobus strobiformis (Fliche and Zeiller). 

A species, described as Pinites strobiformis 3 , 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 (69 2 ) p. 2, PI. n. fig. 10. 2 Ibid. (71) p. 2. 

3 Fliche and Zeiller (04) p. 802, PI. xix. fig. 6. 


Pityostrobus Sauvagei (Fliche and Zeiller). 

The type-specimen of this species 1 , 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 Dunkeri (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 

IWealden strata as also from other countries. Gardner 2 instituted 
the following species: Pinites Carruthersi, P. valdensis, P. cylin- 
droides 3 , P. pottoniensis 3 , 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. Carruthersi, 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 4 Pinites Dunkeri were originally 
described by Mantell as Abietites Dunkeri*", they reach a length of 
over 33 cm. and have a 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, PI. xix. fig. 7. 

- Gardner (86 2 ). 

3 See also Seward (95) A. p. 193; Stopes (15) pp. 138, 140. 

1 Carruthers (66 2 ) PI. xxi. figs. 1, 2. 6 Seward (95) A. p. 194. 




Fig. 778. Pityostrobus Dunkeri. (British Museum; nat. size.) 




Velenovsky 1 from Lower Cretaceous rocks in Bohemia as Pinus 
longissima, a species recently recorded by Dr Stopes 2 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 Leckenbyi (Carruthers). 

This species was first described by Carruthers 3 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 

Fig. 779. Pityostrobus Leckenbyi. From the Lower Greensand of the 
Isle of Wight. (After Carruthers; £ nat. size.) 

scales agree in external form with those of Cedrus and Dr Stepes 
has recently proposed the generic name Cedrostrobus* in order to 
emphasise this resemblance. Prof. Fliche 5 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 
Leckenbyi. A cone of similar form is also described by Coemans 
from Belgium as Pinus Corneti G and compared by him with Cedrus. 

Velenovsky (85) B. PI. i. figs. 14—17. 
Stopes (15) p. 141, text-fig. 38. 
Carruthers (69 2 ) PI. i. figs. 1—5. 
Fliche (96) p. 200, PI. vm. 

S. IV 

4 Stopes (15) p. 143, text-fig. 39. 
6 Coemans (66) p. 11, PI. v. fig. 3. 





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 Pityostrobus 
as it is by no means certain that Carruthers' type is more closely 
allied to Cedrus than to Abies. 

Dr Stopes includes in Cedrostrobus a second species, Cedrostrobus 
Mantelli 1 , from the Lower Greensand of Kent which Carruthers 
originally named Pinites: she compares with it a Potomac cone 
described by Berry 2 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 3 founded the species Abies 
Benstedi is from the Lower Greensand of Kent. It was subsequently 

Fig. 780. Pityostrobus Benstedi. Tangential section showing, os, ovuliferous scale; 
bs, bract-scale; o, the two ovules on the ovuliferous scale; e, endosperm; u\ wing. 
(After Stopes; x8.) 

described by Carruthers 4 as Pinites and regarded by him as prob- 
ably more nearly allied to Cedrus than to Pinus. Dr Stopes 5 has 
recently made a further examination of the structure of this type 

1 Stopes (15) p. 145, text-fig. 40. 
3 Mantell (46) p. 52, PI. n. fig. 2. 

2 Berry (11) PL lxx. fig. 4. 
4 Carruthers (66 2 ) p. 541. 

Stopes (15) p. 130, text-figs. 32, 33; Pis. x., xi. 


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 1 and afterwards referred by Carruthers 2 
to Pinites, has recently been more fully described by Dr Stopes 3 
under Pinostrobus. The cone, 14 cm. long and nearly 5 cm. in 
diameter, bears overlapping scales with a thickened, curved, distal 
margin 2 cm. broad and, on the exposed surface of the specimen, 
1-3 cm. 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 5 gave an account of a cone from Sidmouth in 
Devonshire, which he referred to Pinites oblongus, though Dr Stopes 6 
expresses a doubt as to the identity of his specimen with that 
described by Lindley. Schimper 7 employs the name Cedrus and 
Goeppert assigns the species to Abietites 8 . It is impossible to 
determine the position of the specimen represented in fig. 781 
among the Abietineae. 

1 Mantell (43) p. 34. 2 Carruthers (66 2 ) p. 541, PL xx. figs. 5, 6. 

3 Stopes (15) p. 123, Pis. x., xi. 4 Lindley and Hutton (35) A. PL 137. 

5 WiUiamson (86). 6 Stopes (15) p. 135. 

7 Schimper (72) A. p. 299. 8 Goeppert (50) p. 207. 





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 

Fig. 781. 

Pityostrobus oblongus. (After Lindley and Hutton, from Stopes; 
nat. size.) 

d Of 

Pityostrobus hexagonus (Carruthers). 

A large cone 15 cm. long and 4 cm. in diameter composed 
stout woody scales with hexagonal apophyses was described by 
Carruthers as Pinites hexagonus from the Gault of the South of 
England 1 ; 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. Quenstedti Heer 2 . 

Pityostrobus (Pinites) Andraei (Coemans). 

The cones of this species (fig. 782), the commonest type in the 
Lower Cretaceous rocks of Hainault 3 , are 10 — 14 cm. long and 

1 Carruthers (71) p. 2, PI. xv. 2 Heer (71 2 ). 

3 Coemans (66) p. 12, PI. iv. %. 4; PI. v. fig. 1. 




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 1 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 ihe 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 Fig 
Abietineous cones are founded deserves careful 
consideration and would probably yield results 
of considerable importance. 

782. Pityostrobus 
(Pinites) Andraei. 
(After Coemans; 
nat. size.^ 

Pityostrobus (Pinites) macrocefhalus (Lindley and Hutton). 

This species, founded on a cone 12 cm. long and 6 cm. in 
diameter, was in the first instance described by Lindley and 
Hutton 2 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 
orata 3 . Endlicher 4 assigned the cones to Zamiostrobus and Miquel 5 

1 Heer (69) p. 13, PI. n. fig. 11. 
3 Ibid. (37) A. PI. ccxxvi A. 

2 Lindley and Hutton (35) A. PI. cxxv. 
Endlicher (40) p. 72. 5 Miquel (42) p. 75. 


proposed the name Z. Henslotvi : their Abietineous nature was first 
recognised by Corda 1 , and Carruthers 2 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. macrocefhalus to Eocene beds at the 
junction of the Woolwich and Thanet beds with the London Clay. 
Both Carruthers and Gardner 3 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 4 . 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. 

2 Carruthers (66 2 ) pp. 536, 540, PI. xxi. 

3 Gardner (86) pp. 63, 65, PI. xiv. 

4 Dutt (16). 




ovule shown in fig. 784, A, the contracted nucellus, n, forms a 
cylindrical column which presents a misleading resemblance to the 

Fig. 783. Pityostrobus (Pinites) macrocephdlus. (After Gardner; nat. size.) 

Fig. 784. Pityostrobus (Pinites) macrocephalus. A, section of an ovule; n, nucellus. 
B, longitudinal section of jpart 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 




surface-features are more like those of the cones of Finns exa 
(cf. fig. 704, p. 154) and similar types. 

Pityostrobus (Pinites) Plutonis (Baily). 

This species was founded by Baily 1 on part of a cone from the 
plant-beds in the basalts of Antrim and described, in greater detail 

Fig. 785. Pityostrobus {Pinites) Plutonis. (After Gardner; nat. size.) 

and from better material from the same locality, by Gardner 2 . 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 -bases. 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) PI. xv. fig. 1. 2 Gardner (86) p. 69, Pis. xv.— xvm. 


same locality is named by Gardner Pinus Bailyi 1 . 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. 2 from Provence, P. Kotschyeana (Ung.), 
originally described by Unger and recorded by Tuzson 3 from 
Hungary, P. transsylvanica Pax 4 , a North American form, which 
the author of the species compares with Pinus Balfouriana, and 
P. prae-montana described by Mogan 5 from Lower Austria. 

Pityostrobus (Pinites) palaeostrobus (Ettingshausen). 

This type originally described by Ettingshausen 6 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 7 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 8 , Germany, France, and elsewhere. A cone of similar 
form is described by Unger 9 from the Oligocene of Kumi as Pinus 
megalopis and it is associated with quinary fascicles. 

Pityostrobus MacClurii (Heer). 

This species described by Heer 10 as Pinus (Abies) MacClurii 
from Miocene beds in Banks Land, lat. 74° 27' N. is represented by 
a specimen in the Dublin Museum. The narrow oval cone, 6x1-5 
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 11 to Pinus Abies L. do not afford satisfactory evidence of 
their generic position. 

1 Gardner (86) p. 73. 2 Saporta (73) p. 94, PL n. 

3 Tuzson (09 2 ) p. 240, Pis. xiv., xv. 4 Pax (07) p. 310. 

« Mogan (03) figs. 1—3. 6 Ettingshausen (55). 

7 Heer (83) PI. lxx. fig. 8; PI. lxxxvii. figs. 5, 6. 8 Staub (85). 
9 Unger (67) PI. xvi. 

10 Heer (68) i. p. 134, PI. xx. figs. 16—18. 

11 Ibid. (71) iii. p. 41, PI. v. figs. 35—49. 




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 Timleri 1 
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 witl 
Pinus Gerardiana from Afghanistan. 

B A 

Fig. 786. A, B, part of a cone and a seed of Pinus Timleri. 
Loehri. See page 370. (After Engelhardt and Kinkelin 


C, Keteleeria 
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 2 identified a small 
cone from the Cromer Forest bed as Pinus silvestris, and from this 
horizon in Sweden Nathorst 3 records the same species ; it is recorded 
also from Pleistocene deposits associated with Elephas primigenius 
in France 4 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 

1 Engelhardt and Kinkelin (08) p. 205, PL xxv. figs. 

2 Reid, C. and E. M. (15) PI. i. figs. 7, 8. 


Nathorst (10) p. 1354. 

Fliche (00). 




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 1 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 2 to Picea excelsa var. alpestris. 
Sernander 3 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 4 and other authors who have™ « D -, D - * t? 

J ■ . J . . Fig. 787. Picea excelsa. From 

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 ei ' na ' hUe '' 
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 5 
from the Patoot (Cretaceous) beds in Greenland as Ophioglossum 
granulalum and afterwards described by Newberry 6 from the 
Amboy clays, has recently been identified by Dr Stopes 7 as a long 
microstrobilus of some Abietineous Conifer, probably a Pinus. 

1 Reid, C. and E. M. (08) PI. 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 3 ); Penhallow (04). 5 Heer (83) PI. lvii. figs. 8. 9. 
6 Newberry and Hollick (95) PI. ix. figs. 11—13. 7 Stopes (ll 4 ) text-figs. 1, 2. 


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. 


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 

Pityospermum Lundgreni Nathorst. 

Nathorst described several winged seeds from the Rhaetic beds 
of Scania as Pinus Lundgreni 1 ] 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 
some imperfect cylindrical cones bearing OLJ 
thin imbricate scales and reaching a length Fig. 788. Pityospermum 
of 3—5 cm. and a diameter of 1-2—2 cm.; Lundgreni. From Stab- 

i i , -i ,i •,■!■, ,i barp in Scania; Rhaetic. 

he also suggested the possibility that some (Sto F ckholm Museum: 
short shoots and long needle-like leaves de- n at. size.) 
scribed as Schizolepis Follini Nath. may 

belong to the plant which bore the cones and seeds. In a later 
account of Schizolepis 2 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 a, 13—17; PL xv. figs. 1—2. 

2 Ibid. (97) p. 38, 

in. long anu 'i im. 

I It 






Abietineous seeds is a valid reason 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 1 as Pinus Timleri (fig. 786, B) bears a 
close resemblance in the form of the wing to some of the Rhaetic 

Pityospermum Nilssoni Nathorst. 

This species, also from the Rhaetic flora of Scania 2 , 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- 
seni, and other Jurassic types from the wings of any recent seeds. 

Pityospermum Nanseni 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 3 . The name Pityospermum Nanseni 
was applied by Nathorst 4 to similar speci- 
mens obtained by Dr Nansen from the same F * G ; 789 ' A ' Pityospermum 

„ , . , . , . -~ Nanseni. B, Pityosper 

region. Other winged seeds from .branz 

Josef Land closely resemble Heer's species 
Pinus Maakiana 5 from Jurassic rocks in 
Siberia. A seed, 1*2 cm. long, from Weal- 
den beds in the South of England is repro- 
Iduced in fig. 789, B 6 : this is possibly a 
distinct species, but the specific determina- 
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) PI. xxv. fig. 4. 

2 Nathorst (78) B. p. 32, PL xv. figs. 17—19. 

3 Newton and Teall (97) PL xxxvm. 

4 Nathorst (99) p. 18, PL n. figs. 12, 13. 

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.) 

Heer (77) ii. PL xvi. fig. 1. 

6 Seward (95) A, p. 198. 


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. 


This generic name has been adopted 1 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, Dacrydium 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 2 : one is shown in fig. 491, G (Vol. in. p. 298); the 
longest axis is 80 /x 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 /x. 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. 
It 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. Khaetic. 
Pityosporites sp. 

Among the spores found by Nathorst 3 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/x and in the 
shape of the bladders it agrees closely with the microspores of 
Picea excelsa*. 

1 Seward (14) p. 23. 2 Ibid. p. 23, PL vm. fig. 45. 

3 Nathorst (08) p. 13, PL n. 

4 Kirchner, Loew, and Schroter (06) p. 151, fig. 68. 




Pityosporites sp. 

In his account of petrified plant-remains from Franz Josef 
Land, probably of Wealden or approximately Wealden age, Graf 
Solms-Laubach 1 mentions the occurrence of well preserved pollen 
with bladders and figures a piece of a cone with flat scales similar 

Fig, 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 — 90 /x; 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 cynarocephala Saporta. Saporta 2 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 3 says, they 
are probably Abietineous. 

1 Solms-Laubach (04) p. 11. 

2 Zvto/j.09, cut up; \ewls, scale. 

3 Zeiller (00) B. p. 278. 

Saporta (65 2 ) p. 55, PI. n. fig. 3. 



C rossotolepis Perroti Fliche. This generic name 1 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 Menziesii 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 Entomolepis 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 Engelmaim 
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 an)' 
recent types. 


PREPINUS. Jeffrey. 

This genus was instituted 2 for lignitic specimens of short sho 
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 Kpoaourds, fringed, tasselled. Fliche (99) p. 474, PI. xn. 

2 Jeffrey (08 2 ) Pis. xm., xiv.; Hollick and Jeffrey (09) B. p. 19, Pis. ix., 





Prepinus statensis Jeffrey. 

The short shoots (fig. 791, B), rather less than 1 cm. long, 
consist of a relatively broad axis bearing on the upper part nume- 
rous 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, x 7; C, x 9£.) 

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 

s. IV 



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 
and 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. 

Jeffrey compares this species with Pinites (--=Pityites) Solmsi 
Sew. 1 (fig. 772) from English Wealden beds and with shoots 
described by Fontaine 2 from the Potomac series under Heer's 
generic name Leptostrobus. 

Prepinus viticitensis Jeffrey. 

This species 3 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 Pityites 
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, Pis. xvtii., xix. See page 373. 

2 Fontaine (89) B. p. 227, Pis. ci.— civ. etc. 3 Jeffrey (10). 


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 1 , quoted by Jeffrey, suggests. 
Jeffrey 2 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 3 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, 
Larix, and Pseudolarix, 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 (JO 2 ) p. 331. 3 Thomson (14). 





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 1 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 

1 Thomson (14), PI. xxn. fig. 10. 



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. Yelenovsky 1 
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 a more comprehensive 
genus such as Taxites 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 2 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. PI. xn. figs. 5—11. 2 Guppy (06) p. 302. 



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 

A B C 

Fig. 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 eocaenica (Unger). 

This species, from Eocene beds in South Styria, was founded 
by Unger 1 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, PI. xxm. figs. 11—16. 


Gardner 1 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 2 records specimens of this type 
from Eocene beds at Haring in the Austrian Tyrol where the 
species is said to be abundant, from Leoben 3 in Styria, Bilin 4 in 
Bohemia, Sagor in Carinthia 5 , and elsewhere. Heer 6 states that 
the species is common in Swiss Eocene deposits and Engelhardt 7 
describes examples from Oligocene rocks in Bohemia. A similar 
or perhaps specifically identical type was figured by Lindley 8 from 
Eocene plant-beds at Aix in Provence as Podocarpus macrophylla, 
but Saporta 9 , who figured additional specimens from the same 
locality, proposed the name Podocarpus Lindley ana on the ground 
that Lindley's designation implies identity with a recent species. 
A leaf referred to P. eocaenica is figured by Massalongo 10 from 
Tertiary beds in Italy. 

Some vegetative shoots originally described by De la Harpe 11 
from Alum Bay as Cupressites elegans are referred by. Gardner 12 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, PI. n. figs. 6—15. 

2 Ettingshausen (55) p. 37, PI. ix. figs. 14, 15. 

3 Ibid. (88 2 ) p. 277. 

4 Ibid. (67 2 ) p. 118, PI. xm. figs. 1, 2. 

5 Ibid. (85) p. 6, PI. xxviii. fig. 12. 

6 Heer (55) A. p. 53, PI. xx. fig. 3. 

7 Engelhardt (85) p. 315, PI. vni. figs. 37, 38. 

8 Lindley in Murchison and Lyell (29) p. 298, fig. A. 

9 Saporta (62) p. 216, PI. n. fig. 7. 

10 Massalongo (59) p. 166, PI. v. fig. 36. 

11 De la Harpe in Bristow (62) p. Ill, PI. v. fig. 3. 

12 Gardner (86) PI. vm. 





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- 
londinensis 1 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 2 as Podo- 
carpus borealis, consisting of small falcate leaves 
and seed-like bodies, are too imperfect to be 
determined with accuracy. Fig. 794, C, C repre- 
sents a type from Bournemouth described as 
Podocarpus incerta which differs from other species 
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 Nageia : 

Fig. 793. Podocar- 
pites elegans. Sup- 
posed fertile shoot. 
(Specimen in the 
British Museum 
figured by Gard- 

A C C B 

Fig. 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 3 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 4 

1 Gardner (86) p. 52, PL ix. figs. 35, 36. 2 Ibid, (87) A. PI. xm. figs. 3—11. 

3 Schmalhausen (83 2 ) PL xxxn. 4 Ettingshausen (86) PL vm. figs. 25—27. 




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 ParJceri 1 , 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 


Fig. 795. Podocarpites Campbell*. 

(After Gardner; nat. size.) 

of Mull as Podocarpus Campbelli 2 (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. Thunbergii. 

1 Ettingshausen (87) PI. i. figs. 12—14. 

2 Gardner (86) p. 97, PI. xxvi. 


Fossils believed to be related to Dacrydium. 

The records of the rocks afford very little information wi 
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 (Stachyotaxus) 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 1 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 2 as Dacrydium densifolium have 
no claim to be accepted as branches of a Podocarpineous Conifer. 
Ettingshausen 3 figures from Eocene beds in Australia and New 
Zealand sterile twigs assigned respectively to Dacrydium cupressi- 
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 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, PI. xlii. figs. 21—26. 

2 Velenovsky (85) B. PI. xn. figs. 1—2. 

3 Ettingshausen (86) PI. vui. figs. 23, 24; (88) PI. I. fig. 19. 

4 Nathorst (86) p. 98. 




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 1 are 
dimorphic ; some of the leaves are appressed and imbricate as in 

\ D 

Fig. 796. Stachyotaxus elegans. A, B, | nat. size; C, slightly enlarged; 
D, x 3£. (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 1 cm. The epidermal cells have straight 
walls and the stomata occur in two rows on the lower surface. 

t^astrobili (fig. 796, B) have the form of spikes about 


5 — 6 cm. 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 Stachyotaxus 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 1 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 2 refers to Stachyotaxus septentrionalis some sterile shoot 3 
from Lower Jurassic, or Rhaetic, beds in East Greenland, and 
Halle 3 draws attention to the superficial resemblance to the 
Swedish type of some vegetative twigs from Graham Land which 
he refers to the genus Elatocladus. 

Strobilites. Strobilites Milleri Seward and Bancroft. 

The specimen on which this species is founded 4 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 5 . 
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 (6x5 mm.) which are no doubt 
seeds : each shows a differentiation into an inner portion surrounded 

1 Gibbs (12) p. 539. 

2 Hartz (96) PL xix. figs. 2, 3. 

3 Halle (13 2 ) p. 83. 

4 Seward and Bancroft (13) p. 882, PL I. fig. 13. 

5 Miller (57) B. p. 493. 




by a flat border and it may be that the latter is the impression 

of a sarcotesta. Another possible interpre- imijjufi iijoi 

tation is that the oval bodies are seeds in 

intimate association with fertile bracts. 

The strobilus .bears a close resemblance 

to Stachyotaxus elegans Nath. 1 from the 

Khaetic of Sweden compared by the author 

of the species with an ovuliferous shoot 

of Podocarpus spicata and Dacrydium 

Franhlini and believed to be allied to the 

recent genus Dacrydium, a view upheld 

by Miss Gibbs 2 in her account of recent 

Podocarps. It is not improbable that 

Strobilites Milleri is more closely allied to 

the Podocarpineae than to any other family 

of Conifers. 

Saxegothopsis Dusen. 

In his account of a Tertiary flora, 
possibly Oligocene, from localities on the 
Magellan straits Dusen 3 describes a single 
leaf as Saxegothopsis fuegianus on the 
ground that it resembles the leaves of Saxe- 
gothaea conspicua. The lamina, rather less 
than 2 cm. long, is linear-lanceolate with a 
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. Dusen admits the lack of satisfactory evidence 
indicating generic identity, but the specimen hardly merits the 
istinction of being made the type of a new genus. 

Fig. 797. Strobilites Milleri. 
(After Seward; Edin- 
burgh Museum ; nat. 





This generic name was given by Heer 4 to a fossil, subsequently 
transferred to a new genus Drepanolepis 5 (fig. 798, C), which affords 
Nathorst (08 2 ) PI. n. figs. 1—27. 2 Gibbs (12) p. 539. 

3 Dusen (99) p. 105, PI. xi. fig. 10. 

4 Heer (75) ii. p. 124, PI. xxxv. figs. 17- 


Nathorst (97) p. 43. 


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 asplenioides 1 . 

Phyllocladites asplenioides (Ettingshausen). 

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 fairlv 
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 

Ettingshausen's species is the only fossil that has come undei 
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 n 2 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 3 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 4 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, PI. vin. figs. 28—31. 

2 Page 543. 3 Berry (03) B. 4 Seward (04) B. p. 31. 


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 1 from 
Dakota beds in Nebraska as Phyllocladus subintegrifolius. Heer 2 
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 8 cm. and a breadth of 2 cm. Heer classes the 
species among plants of uncertain position and compares the 
impressions with the phylloclades of Phyllanthus. The species is 
recorded by Hollick 3 from Martha's Vineyard and other localities, 
by Berry 4 from the Raritan flora, and as Thinnfeldia Lesque- 
reuxiana by Newberry and Hollick 5 from the Amboy clays and 
other Cretaceous floras 6 : 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 7 
has described a specimen from the Great Oolite of Marquise 
(N. France) as Protophyllocladus sp. : this is the first European 

I record for Berry's genus. 
A similar species is represented by Protophyllocladus polymorphus 
(Lesq.) first described by Lesquereux 8 from Vancouver Island as 
Salisburia polymorpha and afterwards transferred by Knowlton 9 
to Thinnfeldia. Another closely allied type is Protophyllocladus 
lobatus Berry 10 from Upper Cretaceous rocks in South Carolina. 
Some of these supposed phylloclades closely resemble flattened 

1 Lesquereux (74) p. 54, PI. i. fig. 12. 

2 Heer (82) i. p. 37, PI. xliv. figs. 9, 10; PL xlvi. figs. 1—12. 

3 Hollick (06) p. 36, PI. v. figs. 1—6. 4 Berry (ll 3 ) p. 98, PL ix. 

5 Newberry and Hollick (95) p. 59, PL XI. 

6 Lesquereux (91) PL n. 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 n. figs. 9—13. 




leaf-like branches from Kreischerville for which Hollick and Jeffrey 
instituted the genus Androvettia 1 (fig. 806, A — C) : some examples 
of that genus are clearly distinct as there are small leaves borne 

Fig. 798. A, B. Protophyllocladus subintegrifolius. C. Drepanolepis rotundi folia. 
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. 


Palaeocladus Ettingshausen. 

This name was applied to a single species, Palaeocladus cunei- 
formis, from New South Wales 1 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 2 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 

I the generic name Nageiopsis than to make use of a designation 
suggesting a relationship which has not been established. 
T AXITE S. Brongniar t. 

This generic name, first proposed by Brongniart 3 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 Taxus, 
Sequoia sempervirens , Cephqlotaxus 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 
:ommon form and the designation Taxites, as generally employed, 

1 Ettingshausen (86) p. 93, PI. vm. figs. 33, 33 a. 

2 Fontaine (89) B. p. 204, PI. lxxxiv. fig. 5, PI. CLXvn. fig. 4; Berry (03) B. 

3 Brongniart (28) A. p. 108. 

s. iv 27 


cannot therefore be interpreted as indicative of affinity to Taxus. 
Taxites 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 1 
has proposed to employ the generic name Elatocladus for sterile 
shoots of the Taxites 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 2 , 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 3 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*. Taxites should be 
retained for fertile branches or reproductive organs which there is 
some good reason for believing to be more closely related to Taxua 
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 5 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 
Sequoiites Langsdorfii and not decurrent. 

The generic names Cephalotaxites and Cephalotaxopsis applied 
by Heer 6 and Fontaine 7 respectively to foliage-shoots resembling 
those of recent species of Cephalotaxus 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, Cephalotaxites, 
like Torreyites and Taxites, is more appropriately restricted to 

1 Halle (13 2 ) p. 83. 2 See page 426. 3 See page 432. 4 See page 410. 

5 Heer (68) p. 95, PI. i. figs. 21—24; PI. xlv. fig. 1. 

6 Ibid. (83), p. 10. 7 Fontaine (89) B. p. 235. 


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 1 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-Tertiary strata. 

The foliage-shoots described as species of Taxites are as a rule 
valueless as records of Taxus, Cephalotaxus, 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 2 and similar examples were described by Heer 3 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 5 , and 
Clement Reid 6 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 7 . 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 8 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. 


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 lx. fig. 19. 

3 Heer (62) p. 1078, PL lxx. figs. 15—23. 

4 C. and E. M. Reid (10) p. 172. 

5 Ibid. p. 171, PL xvi. figs. 42, 43; (08) PL xv. fig. 145. 

6 Reid (99) B. p. 151. 7 Elwes and Henry (06) p. 99. 8 Conwentz (01). 




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 ungrooved 
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 1 , using 
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 distinc; 
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 2 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 3 has described two species, Torreya Dichsoniana and 

1 Berry (08 2 ). 2 Ibid. (14) pp. 107, 123. 

3 Heer (75) ii. p. 70, PL xvm. figs. 1—4; p. 71, PL xvn. figs. 1, 2; (82) i. PL ii. 
fig. 11. 


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 Taxites 
Jeffreyi 1 : there are no adequate grounds for the use of the name 
Torreya. Fontaine's Potomac species Torreya virginica 2 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 3 . 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 5 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 6 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 

ias Sequoia Langsdorfii as fragments of a Torreya and re-name the 
species T. bilinica. 
Some detached leaves and seeds from Upper Pliocene beds in 
the Main Valley (Frankfurt) are referred by Engelhardt and 
Kinkelin 7 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 (ll 2 ) p. 688. 2 Fontaine (89) B. p. 234, PL cix. fig. 8. 

3 Ibid. p. 235, PI. cxiii. fig. 4. 4 Yokoyama (89) p. 230, PI. xxn. figs. 11, 12. 

6 Heer (83) p. 56, PI. lxx, fig. la. 
G Saporta and Marion (76) p. 87, PI. xxn. figs. 6, 7. 
Engelhardt and Kinkelin (08) p. 191, PI. xxm. figs. 6—8. 


VESQUIA. Berfcrand. 

Vesquia tournaisii Bertrand. The name Vesquia, after the 
French botanist Julien Vesque, was given by Bertrand 1 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. 


Fontaine 2 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 3 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 4 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 Pityophyllum. 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. Pis. civ. — cvm. 


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 Cephalotaxus. 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 Cephalotaxus. The presence of depressed 
stomatal regions is a characteristic feature of Torreya and not of 

IHeer 1 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 2 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 Cephalotaxus europaea 3 founded on sterile shoots 
from the Aquitanian of Manosque in the South of France affords 
no convincing evidence of generic identity with Cephalotaxus. 

Seeds from Upper Pliocene beds in the Main valley are referred 
by Engelhardt and Kinkelin 4 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 

1 Heer (83) p. 10, PL liii. fig. 12. ' 2 Berry (10*). 

3 Saporta (93) p. 42, PL v. fig. 4. 

4 Engelhardt and Kinkelin (08) p. 194, PL xxm. fig. 11. 




Trioolepis Leclerei Zeiller. This generic name was proposed bv 
Zeiller for a cone from the Rhaetic flora of Tonkin 1 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 10 cm. 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 fold*; 
and close to the base are slight depressions indicating the former 
presence of seeds 5 mm. long and 1-5 mm. broad. On some oi 
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 Cunninghamia though 
this in itself is probably of comparatively small importance. 


This designation was proposed for fossils which are in all 
probability male strobili of Gymnosperms 2 . 

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 13 cm. long bearing numerous small branches with 

1 Zeiller (03) B. p. 208, PI. l. fig. 15; PI. F, fig. 2. 2 Seward (ll 2 ) p. 686, 

CH. L] 



Fig. 799. Masculostrobus. A, M. Zeilleri, nat. size; a, b, portions enlarged; 
c, microspores from A; d, Masculostrobus sp.; e, spores from d. (British 


spirally disposed sporophylls. A few oval spores (fig. 799, c), 
20 — 30 /x 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 
Vidali 1 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. 

Moller and Halle 2 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 z 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 4 instituted this genus for a type of Conifer previously 
described by Braun 5 from the Khaetic 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. Braunii, was first figured by Goeppert 6 and 
described in more detail by Schenk 7 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 Elatides 8 . Endlicher's designation 

1 Zeiller and Vidal (02) p. 7, PL n. 

2 Moller and Halle (13) p. 36, PI. vi figs. 9—18 

3 Hollick and Jeffrey (09) B. p. 66. 

4 Endlicher (47) p. 306. 5 Braun (43). 6 Goeppert (50) PI. xlviii. 
7 Schenk (67) A. p. 175, PI. xli. 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 1 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). 

Xathorst 2 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 Presl 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 Taxites 
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 (08 2 ). 




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 double 1 . Schenk 2 described 
the cone-scales as bearing 10 — 12 seeds on 
the edge and Saporta 3 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 4 which partially encloses 
the seeds of Dacrydium and other recent 
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. 




Fig. 800. Palissya spkeno- 
lepis. A, B, sporophyll 
in surface-view and in 
section. C, cupules of two 
seeds. (After Nathorst.) 

1 Solms-Laubach (91) A. p. 73. 

2 Schenk in Schimper and Schenk (90) A. p. 336. 

3 Saporta (84) p. 513. 

4 Seepage 118. 

l] elatocladus 429 


This genus was founded with a view to reduce the 'present 
state of intolerable confusion in the classification of the Conifers 1 .' 
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 Taxas, but notwithstanding the 
employment of Taxites by authors who do not intend to convey 
the impression of relationship to Taxus, 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). 
Entocladtis 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 Taxites 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 (13 2 ) p. 82. iXdrri, Pine or Fir; k\&8os, shoot. 




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 (Taxites 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 1 are freely branched and the ultimate branches show a 

Fig. 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 (13 2 ) p. 84 ? PI. vni. text-fig. 18. 




Halle 1 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 2 . 

Elatocladus zamioides (Leckenby ex Bean ms.). 

This type, from the Middle Jurassic of Yorkshire, was described 
by Leckenby 3 as Cycadites zamioides and subsequently transferred 

Fig. 802. 

Elatocladus plana. (Specimens figured by Feistmantel 
as Taxites planus ; nat. size. ) 

to Taxites*. The type-specimen, in the Sedgwick Museum, Cam- 
bridge, consists of a slender axis bearing two-ranked spirally 

1 Halle (13 2 ) p. 86, PI. vni. ~ Stirling (00) PL in. figs. 8, 9. 

Leckenby (64) A. p. 77, PI. vni. fig. L 
4 Seward (00) B. p. 300, PI. x. fig. 5. 


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 1 in Cephalotaxopsis though there is 
no evidence of relationship to the recent Cephalotaxus. Shoots of 
similar habit are figured by Feistmantel 2 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 Taxites, 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 


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 fossil* 
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 3 to separate the Indian 
specimens from Palissya and Taxites 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. Pis. cvi.— cvm. 2 Feistmantel (79) Pis. xin.— xv. 

3 Holden, R. (15 2 ). 

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 is intended. 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 indie us and transferred 
by Feistmantel to Palissya. 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 an v indication of a median -c, OAO D ,. .. . .. . . 

" Fig. 803. Retinosporites indica. A, piece 

astomatic region such as one would f 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 
Presl 1 gave the name Cunninghamites to some sterile shoots 
from Rhaetic and Lower Cretaceous strata on the ground of their 
resemblance to branches of Cunninghamia 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. PI. XLvm. fig. 3; Goeppert (50) PI. xlvii. 
s. iv 28 




branches from the Keuper of Germany which he named C. dubius : 
this species is identified by Saporta 1 with Palissya Braunii, but 
the latter name is now restricted to shoots bearing a particular 
form of strobilus. Nathorst 2 , on the other hand, suggests that 
C. dubius may belong to Elatides, and the same author instituted 
a new generic name Camptophyllum 3 for fragmentary foliage-shoots 

Fig. 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 4 . The name Cunninghamites is given by Oldham and 

1 Saporta (84) p. 511. 2 Nathorst (08 2 ) p. 10. 

3 Ibid. (78) B. PI. xvi. figs. 13—16. 

4 Schenk in Schimper and Schenk (90) A. p. 351, fig. 236. 

l] elatocladus 435 

Morris 1 to Jurassic Indian shoots which Feistmantel 2 afterwards 
transferred to Palissya: these have recently been included by 
Halle 3 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. Velenovsky 4 
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 
Bidwillii or A. brasiliensis. There is no doubt that under Cunning- 
hamites are included branches of many different Conifers. 

Elatocladus elegans (Corda). 

Originally described by Corda 5 from Lower Cretaceous rocks 
in Bohemia as Cunninghamia elegans, this species is recorded from 
many Cretaceous localities; from the Patoot beds in Greenland 6 , 
Moravia 7 , Westphalia 8 , Upper Cretaceous beds in Bulgaria 9 , the 
Amboy clays 10 , Cliff wood, Martha's Vineyard 11 , Georgia, Carolina 12 , 
and other places in North America 13 . The specimen shown in 
fig. 805 from Morayia shows a midrib in a few leaves, while in the 

1 Oldham and Morris (63) PL xxxn. fig. 10. 

2 Feistmantel (76 2 ) p. 55. 

3 Halle (13 2 ) p. 86, PI. vm. 4 Velenovsky (87). 
3 Corda in Reuss (46) B. PL xlix. 
6 Heer (83) PL liii. fig. 1. 7 Ibid. (69) PL i. 

8 Hosius and von der Marck (80) B. PL xxxvu. 

9 Zeiller (05 2 ) PL vh. fig. 14. 10 Newberry and Hollick (95) PL v. 
11 Hollick (06) PL in. fig. 1. 12 Berry (10 4 ) PL xx.; (14) p. 106. 
13 See Hollick (06) for other references. 

b28— 2 


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 1 assigns some branches 
to C. stenophylla but these may be younger forms of C. elegans. 
Similarly C. squamosus 2 , 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 Sequoia and also 
referred by him to Torreya: there is no possibility of deciding the 
precise systematic position of these and similar specimens. Ettings- 
hausen 4 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 5 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. 

2 Heer (71 2 ) PI. I. figs. 5—7; Schimper and Schenk (90) A. p. 282; Berry (03) 
p. 64. 

3 Heer (82) B. PI. xxix. fig. 12. 4 Ettingshausen (72) PL i. fig. 30. 
6 Hollick and Jeffrey (09) B. .p. 22, Pis. ni., vn., viii., xxvm., xxix. 




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 medullarv rays are not described. The data are 

Fig. 806. A — C, Androvettia statenensis. D, Geinitzia 
Reichenbachii. (A, B, x 6; C, x 7; D, x 7; after 
Fig. 805. Elatocladus Hollick and Jeffrey.) 

elegans. (Nat. size; 
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 
1 Berry (14) p. 103, PI. xvm. 


points out, bear a close resemblance to species of Moriconia (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 1 referred to the lower half of the Upper Cretaceous as 
Androvettia carolinensis. 

DACTYOLEPIS. Hollick and Jeffrey. 

Dactyolepis cryptonierioides Hollick and Jeffrey. The generic 
name 2 was instituted for some detached, cuneate, cone-scales from 
the Cretaceous beds at Kreischerville in Staten Island, approxi- 
mately 4 mm. 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 Jeffrey. 

The name Raritania 3 , 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 Baiera 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 4 ) p. 183, PI. xix. 

2 Hollick and Jeffrey (09) B. p. 52, PI. x. figs. 12, 13. 

3 Ibid. (09) B. p. 26, Pis. vi., ix., x., xx. 

4 Newberry and Hollick (95) p. 59 r PI. xn. figs. 1—3 a. 




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' 

Fig. 807. 

A, B, Raritania gracilis; C, Raritanial. 
A, B, x6; C, x 10.) 

(After Hollick and Jeffrey. 

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 Brachyphyllmn 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. 


Braun 1 instituted this genus for a strobilus from Rhaetic 
rocks in Germany which he called Schizolepis liaso-keuperinus, 

1 Braun (47) p. 86. 


characterised by its deeply split cone-scales. Schenk 1 subsequently 
substituted the name S. Braunii 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-scale 
the systematic position must be left an open question. The gen 
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 2 , 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 affinit 
A comparison may also be suggested with the reflexed cone-seal 
of Picea Breweriana. 

Schizolepis Braunii Schenk. 

It has already been painted 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 

1 Schenk (67) A. p. 179. 

2 Nathorst (97) p. 61. 

3 Schenk (67) A. p. 179, PI. xliv.; Schimper and Schenk (90) A. p. 306. 





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 Follini Nathorst. 

In this species from the Rhaetic flora 
of Scania 1 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 2 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 
Raciborski 3 refers a specimen from rocks 
possibly of Rhaetic age in Poland. 

Schizolepis Moelleri Seward. 

Under this name 4 some detached bi- 
lobed scales without seeds were described 
from Jurassic rocks in Turkestan (fig. 808, A) and the South of 
Russia 5 , and Krystofovic has recently discovered a cone of the 
same species in Jurassic strata in Transbaikalia 6 . 

Schizolepis cylindrica Nathorst. 

Founded on a long and narrow, incomplete, strobilus from 
Upper Jurassic rocks in Spitzbergen 7 bearing deeply lobed scales. 
A second species, S. retroflexa 8 (fig. 808, C), assigned with some 

1 Nathorst (78) B. p. 28, Pis. xiv., xv. 

2 Saporta (84) p. 502. 

3 Raciborski (92) p. 354, PL n. figs. 1, 20 d. 

4 Seward (07 2 ) p. 36, PL vn. figs. 64—66. 

5 Thomas (11) p. 79, PL v. fig. 4. 

6 KrystofoviS (15) p. 95, PL vi. figs. 10, 11. 

7 Nathorst (97) p. 39, PL n. figs. 1, 2. 

8 Ibid. p. 60, PL EEL figs. 31, 32; PL vi. figs. 11, 12. 

Fig. 808. Schizolepis. A, Schi- 
zolepis Moelleri. B, Schizo- 
lepis Braunii. C, Schizo- 
lepis(?) retroflexa. (A, after 
Seward; B, after Schenk; 
C, slightly enlarged, after 


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 1 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 Tmesipteris; the relationship of the Spitzbergen fossils 
must be left uncertain pending more evidence. 


Heer described a specimen of Jurassic age from Spitzbergen 
as Phyllocladites rotundifolia 2 which he considered to be closely 
allied to Phyllocladus : an examination of the type-specimen led 
Nathorst 3 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 


Schizolepidella gracilis Halle. The specimens on which this 
genus is founded are from the Hope Bay flora in Graham Land 5 , 
probably of Middle Jurassic age; they consist of slender sterile 
shoots reaching a maximum length of 12 cm. and 2 mm. broad, 
rarely branched and bearing small leaves, 2x1-5 mm., apparently 

1 Saporta (94) B. PI. xxxm. fig. 4. 

2 Heer (75) ii. p. 124, PI. xxxv. figs. 17, 18. 

3 Nathorst (97) p. 43, PI. vi. figs. 24, 25. 

4 Ibid. p. 71, PI. in. figs. 33—37. 5 Halle (13 2 ) p. 90, PI. ix. figs. 18—21. 




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 1 . 



, 1 



A B 

Fig. 809. Schizolepi- 
della gracilis. (After 
Halle; A, enlarged; 

This name was given 2 to foliage-shoots and B, nat.size.) 
cones, from the Urgonian rocks of Greenland, originally described as 
Widdringtonites gracilis 3 . 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- 
\mia. and mentions the striated surface of the fossil cone-scales 
is a distinctive feature, though that may be due, in part at least, 
the state of preservation. The flat form of the cone-scales is a 
character in which Cyparissidium differs from genera such as 
kquoiites, and from the Callitrineae the cones are distinguished by 
he spiral arrangement of the scales. 


See page 439. 
2 Heer (75) ii. PL xvn. fig. 5 b, c; 

Pis. xix. , xx., xxi.; (82) pp. 16, 50, Pis. I., vn. 
3 Ibid. (68) p. 83. 




Cyparissidium is characteristic of Lower Cretaceous strata 
though Nathorst 1 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 2 . 

Fig. 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 20 mm. long and 
11 — 20 mm. broad with scales having a breadth of 12 mm., a 

1 Nathorst (86) p. 103, PI. xxn. 

2 See page 410. 

l] benstedtia 445 

rounded distal border, a rnucronate apex, and a striated dorsal 
face. Velenovsky 1 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 2 
(fig. 810, C). Schenk 3 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 4 is based on insufficient evidence. 


In 1862 Mackie 5 figured a fossil stem from Lower Greensand 
rocks in Kent which Konig afterwards named Dracaena Benstedtii. 
This name was adopted by Morris and Mantell for the supposed 
'Dragon tree.' In 1868 Carruthers 6 expressed the opinion that 
the fossils are more likely to be Pandanaceous stems, while Gardner 7 
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. in. 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 40 cm. and a 
diameter of 15 cm. Smaller specimens with similar surface-cha- 
racters are described by Fliche 9 from Lower Cretaceous beds in 
France as Conifer ocaulon colymbeaeforme and compared by him 
to stems of Araucaria imbricata : other examples closely resembling 

1 Velenovsky (85) B. p. 17, PL vin. 2 Ibid. Pis. ix., x. 

3 Schenk (76) B. p. 167, PL xxix. figs. 10, 11. 
* Hollick (06) p. 46, PL m. fig. 11. 

5 Mackie (62). 6 Carruthers (68) p. 154 (footnote). 

Gardner (86 2 ) p. 201. 8 Seward (96 2 ) p. 216. 9 Fliche (00). 


the French stems are recorded from the Uitenhage (Wealden) 
series of South Africa and the Kimeridge rocks of Sutherland 1 , 
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 2 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 3 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 Benstedtia, at least as regards 
the specimens which afforded anatomical evidence; in the French 
and South African examples no internal structure is preserved. 
Dr Knowlton 4 pointed out that the generic name Benstedtia should 
be preferred to Coniferocaulon on the ground of priority and he 
named the English specimens Benstedtia Benstedtii. Dr Stopes 5 
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 6 . 

1 Seward (03) B. p. 34; (ll 2 ) p. 690. 

2 Bancroft (13) pp. 72, 85. 3 Stopes (11). 
4 Knowlton (11). 5 Stopes (IP). 
6 Stopes (15) p. 159, Pis. xiii., xiv. 





The name Podozamites 1 was instituted for certain species pre- 
viously included in Za mites 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 Podozamites differs in no 
very important respect from Zamites, 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 Podozamites has been 
regarded as Cycadean, but Schenk's discovery of a specimen of 
Podozamites 2 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 Glossozamites 3 . 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 4 
who was strengthened in his view by the characters of the seed- 
bearing sporophylls described by Nathorst 5 and provisionally 
connected by him with Podozamites. Nathorst 6 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. PI. xxxvi. fig. 3. 

3 Ibid. (71) PI. ii. 4 Zeiller (03) B. p. 159. 

5 Nathorst (86) p. 91, PI. xxvi. 6 Ibid. (86) PI. xvi. fig. 10. 



Fig. 811. Podozamites lanceolatus. (Nat. size; British Museum, 39,303.) 


Schuster 1 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 2 , 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 3 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 1902 4 more fully described the type-species 
G. Erdmanni. 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 Erdmanni 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 
interpreted 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, A is 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 (ll 4 ). 2 Seward (00) B. p. 242. 

3 Nathorst (ll 4 ). 4 Ibid. (02) p. 8, PI. I. figs. 5, 6. 

7 29 




rocks in Tonkin 1 where Podozamites also occurs. Heer 2 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. 

Fig. 812. A, B, Cycadocarpidiwm Erdmanni. A, Restoration of fertile shoot. 
B, Single sporophyll with seeds. C, D, Cycadocarpidiwm 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 

1 Nathorst (ll 4 ) p. 5. 2 Heer (77) i. PI. vm. fig. 4. 


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 1 speaks of Cycadocarpidium Erdmannias 
differing from C. Swabii 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. Erdtnanni and the swellings in C. Swabii 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 Podozamites, 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 Nathorsti 2 and separate leaves of Podo- 
zamites ; a similar comparison may be made between the latter genus 
and EretmophyUum, a genus instituted by Thomas 3 and referred 
to the Ginkgoales (cf. figs. 658, 659, pp. 59, 62). There is indeed some 

I resemblance between Cycadocarpidium sporophylls and abnormal 
1 Schuster (114). - Vokoyama (89) B. Pis. vin., ix. 3 Thomas (13). 



seed-bearing leaves of Ginkgo. We have as yet but little to guide 
us in 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 1 to his genus Nageiopsis, 
should not be overlooked from the point of view of their possible 
relationship to Podozamites. 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 2 , 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 Nageiopsis figured by 
Fontaine a zone of crowded scars 3 (fig. 816, s) such as may be seen 
on an Agathis shoot. 

Many of the leaves described as Podozamites are of little value 
as evidence of the occurrence of the genus. In the case of imperfec t 
specimens of detached leaves it is often impossible to distinguish 
between Podozamites, Phoenicopsis, and the leaves of Araucariati 
plants, or pinnae of some species of Zamites. It is therefore net 
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 4 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. angustifolius (Eich.) and P. mar- 
ginatus Heer 5 afford no proof of the presence of Podozamites: 
similarly Velenovsky's species P. miocenica from Bohemia 6 might 
equally well be referred to the genus Dammarites. Well pre- 
served specimens have been described by Zeiller 7 from the 

1 Fontaine (89) B. p. 195. 2 Berry (10). 

3 Fontaine (89) B. PI. lxxvi. fig. 5. 4 Hollick (12) Pis. 162, 163. 

5 Newberry and Hollick (95) PI. xm. figs. 1 — 6. 

6 Velenovsky (81) PL I. figs. 18—20. ^ 7 Zeiller (03) B. 




Rhaetic flora of Tonkin 1 . 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 2 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 _ 
r to Fig. 813. A, Specimen figured 

the stomata, usually with six accessory by Feistmantel as Podo- 
cells, occur in the intercostal regions on 
both surfaces: the characters of the epi- 
dermis are favourable to a relationship 
with the Coniferales and they are not 

inconsistent with the inclusion of the fossils in the genus Phoe- 
nicopsis. Halle 3 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 4 
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. 

Walkom (17) p. 20. 

Feistmantel (82) p. 39, PI. n. figs. 2—5. 
Halle (13). 
4 Fontaine in Ward (05) B. Pis. xxiv., XXV., xliv. Knowlton (14) Pis. v., vi. 


zamites lanceolatus. B, Piece 
of lamina enlarged. (Cal- 
cutta Museum; Geol. Surv., 


Podozamites distans (Presl). 

This Rhaetic species 1 differs very slightly from the Jurassic 
type Podozamites lanceolatus and there has been much confusion 
on the part of authors between the two forms 2 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 20 cm. 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, 04 — 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 w T alls and the stomata, which occur 
on the lower surface, either in rows or scattered, are surrounded by 
small subsidiary cells (fig. 812, E). The leaves are usually rather 
broader in proportion to their length than those of P. lanceolatus 
and the apex is less pointed. Braun 3 instituted two varieties, 
longifolius and latifolius, and to these Schenk 4 added others. Th? 
species is recorded from the Rhaetic of Scania 5 , where it is abundant, 
from Persia 6 , Tonkin, and many other regions: it occurs also in 
Jurassic strata 7 , but on the whole P. distans is a characteristic 
member of Rhaetic floras. 

The Rhaetic species Podozamites Schenki Heer 8 founded on 
Jurassic specimens from Siberia and described by Zeiller 9 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. 10 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. PI. xli. fig. 1. 

2 See Zeiller (03) B. p. 159 for examples of P. distans referred to P. lanceolatus. 

3 Braun (47) p. 85. * Schenk (67) A. Pis. xxxv., xxxvi. 

5 Nathorst (78) B. Pis. xni., XT. 

6 Zeiller (03) p. 193. "' For references, see Zeiller (03) B. p. 159. 
8 Heer (77) ii. p. 45. 9 Zeiller (03) B. PI. xlii. 

10 Lindley and Hutton (36) A. PI. xciv. 


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 
Zamia 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 1 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 
rocks in India as P. lanceolatus (fig. 813) should, I am inclined to 
think, be assigned to Phoenicopsis. 

Podozamites Reinii Geyler; Podozamites stonesfieldensis Seward; 
Podozamites Griesbachi 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. Reinii 2 (fig. 814) the broadly 
oval leaves are attached to a slender axis. P. stonesfieldensis 3 
from the Great Oolite of Stonesfield is probably identical with the 
leaves originally described by Buckman as Naiadea ovata and Lilia 

E n *i\ the leaves are oblong-ovate, approximately 8 x 3-5 cm. ; 
na is rather abruptly contracted at the base and more 

For references see Seward (00) B. p. 242; (07 2 ); (11). 

Geyler (77) B. Pis. xxxra., xxxiv. 

Seward (04) B. p. 121, PL in. fig. 4; PL xi. figs. 1, 2. 




gradually tapered towards the apex; the veins are slightly more 
than 1 mm. apart and converge at each 
end of the lamina. The species re- 
sembles P. lanceolatus var. latifolius 
figured by Schenk 1 from China. 

Podozamites Griesbachi 2 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. Reinii but the venation is still 
coarser in P. stonesfieldensis . A similar 
form of leaf is figured by Velenovsky 3 
from the Lower Cretaceous of Bohemia 
as P. striatus. 

Fig. 814. Podozamites Reinii. 
(After Geyler; f nat. size.) 

Podozamites Kidstoni Etheridge. 

In this type 4 , 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 b and similar leaves are figured by 
Chapman 6 from Jurassic rocks in Victoria, Australia. Etheridge's 
type-specimen is from the Burrum Coal Measures ('? Triassic) ol 
Queensland. This form of leaf agrees closely with some of 
Fontaine's Potomac species of Nageiopsis, e.g. N. obtusa and N. 
heterophylla 1 ', and a similar though not identical form is described 
by Nathorst 8 from the Ehaetic of Sweden as P. ovalis, distinguished 
by its broadly rounded and mucronate apex. 

NAGEIOPSIS. Fontaine. 

This genus was established 9 for vegetative shoots abundantly 
represented in the Potomac flora many of which closely resemble 
Podozamites, though differing in their branched habit and in the 
veins being less convergent in the apical region of the lamina. 


1 Schenk (83) A. PI. xlix. figs. 4 6, 5. 

2 Seward (12) p. 36, PI. iv. fig. 58; PI. vi. fig. 79 

3 Velenovsky (85) B. PI. n. fig. 8. 

4 Jack and Etheridge (92) B. p. 317, PI. xvra 
■ Seward (12) PI. iv. fig. 39. 

7 Fontaine (89) B. Pis. lxxxiv., lxxxv. 

8 Nathorst (78) B. PI. xin. fig. 5. 

figs. 6, 7. 
6 Chapman (09) PL xvni. 

9 Fontaine (89) B. p. 194. 




Fontaine recognised the similarity between Nageiopsis and shoots 
of Podocarpus belonging to the section Nageia: this suggested the 
choice of the generic name. Berry 1 in his revision of Nageiopsis 
transfers some of Fontaine's species to Podozamites ; he also reduces 
the number of the species retained in Nageiopsis on the ground that 
Fontaine attached too much importance to variations in the size 
and form of the leaves. I have elsewhere suggested 2 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 Nageiopsis are hardly distinguishable 
from Zamites Btichianus 3 . 

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. 

Nageiopsis anglica Seward. 

This species 4 founded on the small specimen 
represented in fig. 815 has distichous leaves 
1 — 1-5 cm. long with several parallel veins. 
The English Jurassic type agrees generally 
with Nageiopsis microphylla Font, and N. des- 
crescens Font. :. a similar form is recorded from 
the Wealden beds of Sussex 5 . 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 
5 mm. to 1-3 cm. in breadth; there are 9—12 veins unbranched 

1 Berry (10). 2 Seward (12) p. 33. ■ 3 Berry (11) PI. lxi. 

4 Seward (00) B. p. 288, fig. 51. 5 Ibid. (95) A. p. 211, PI. xn. fig. 3. 

Fig. 815. Nageiopsis an- 
glica. (From a speci- 
men in the Whitby 
Museum; § nat. size.) 



[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, *, 
presumably of bud-scales, at one place on the axis. This species 

Fig. 816. Nageiojisis longifolia. (A, after Fontaine; nat. size.) 

is very abundant in the Potomac beds of Virginia and Maryland 1 
and is recorded by Fontaine from several other localities though 
for the most part on slender evidence. 

Nageiopsis zamioides Fontaine. 

In this species 2 the leaves are ovate-lanceolate and shorter thai 
in N. longifolia; they have a maximum breadth of 2 cm. anc 
reach a length of 7 cm. The example described from English 
Wealden beds as Nageiopsis sp. cf. N. heterophylla agrees closely 
with Fontaine's Potomac species. 

1 Fontaine (89) B. p. 195, Pis. lxxv.— lxxix. ; lxxxv.; Berry (11) p. 384. 

2 Ibid. p. 196, Pis. lxxix. — lxxxi.; Berry (11) p. 386, Pis. Lxn., lxiii. 



I. 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 unnecessary to deal fully with the recent 
types 1 , 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 2 , are 
distinguished by a dichasial system of branching, a character 
foreign to Gymnosperms as a whole though exhibited by the stem 
of Wielandiella, a member of the Bennettitales. There are good 
reasons for believing that pollination is effected by insects 3 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 (06 2 ) p. 274; (09) p. 343; Berridge and Sanday (07) p. 172; Karsten 
(92); Porsch (10). 


Welwitschia, in some species of Gnetum, and occasionally ii 
Ephedra. The seeds are albuminous and the embryos have tw( 
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 1 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° S. 
and has not been found more than 50 miles from the coast. 


Shrubs, in some species with climbing branches, characterisec 
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, e.g. 
Ephedra altissima, the leaves may reach 3 cm. in length and a 
breadth of 1 — 1-5 mm. Monoecious or dioecious; flowers uni- 
sexual; bisexual inflorescences are recorded in E. campylopoda 2 . 
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. altissima, 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 3 , two vascular bundles 
supply the outer envelope (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). 


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 1 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 
Welwitschia. 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 2 found that the seeds of Ephedra distachya germinated 
after 93 days' immersion in sea- water. 

The male flowers 3 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 4 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 5 
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 Wehvitschia. Anatomi- 
cally 6 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) 
- 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 2 ). See also Jeffrey (17) p. 357. 



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 1 . 
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. 


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 2 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 3 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 4 . In appearance 
the antherophore of Gnetum approaches most closely to the stamen 
of an Angiosperm. The female flowers 5 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 mieropylar tube. There are no 

1 Hill and de Fraine (10) p. 329. 2 Karsten (93); Lotsy (11) p. 347, fig. 209. 

3 Caporn (16). 4 See also Pearson (15). 

5 Thoday (Sykes) (11); Lignier and Tison (13). See also Pearson (17). 




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 1 agrees much more 
closely with Wehvitschia than with Ephedra. Attention has been 
called to certain resemblances between the seed of Gnetum and 

Fig. 817. Seed of Gnetum africanum. a, outermost envelope; v, vascular strands; 
b, outer integument; /, flange of micropylar tube; n, nucellar cap; c, inner 
integument; p, endosperm. (After Mrs Thoday.) 

that of Bennettites 2 . 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,/, from the apical region of the inner integument; the inner 

1 Pearson (09); see also Karsten (92); (93 2 ); Lotsy (99). 

2 Berridge (11); (12); Thoday (Sykes) (11). 


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 1 and in transverse section presents a striking similarity to 
sections through the same region of a Bennettites seed (figs. 524, 527, 
Vol. in. 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 2 pointed out that in the vascular bundles of the leaver 
the parenchyma of the medullary rays forms continuous plates, 
gymnospermous character. 


This remarkable genus, discovered by Welwitsch in 1860 anc 
described by Sir Joseph Hooker 3 , presents striking peculiarities in 
the habit of- the vegetative body. A Wehvitschia plant has been 
aptly termed an adult seedling 4 ; the large and squat tuberous 
stem, morphologically the swollen hypocotyl, may be as much 
as 4-5 m. 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. 

2 Strasburger (91) p. 148. For an account of the anatomy of Gnetum, see 
Duthie (12); La Riviere (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 2 ) p. 333. 


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 1 .' 
The venation is parallel and there are numerous cross-connexions, 
some ending blindly in the mesophyll 2 . 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 3 , 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 1 mm. in diameter. There is no embryo-sac but 
the nucellus acts as a nectary, the drop-mechanism of the functional 

1 Pearson (06 2 ) p. 270. 

2 de Bary (84) A. fig. 157; Sykes (10 2 ); Takeda (13 2 ). 

3 Lignier and Tison (12). 





ovule in the female flower being retained in the sterile ovule of the 
male 1 . 

In contrast to the indefinite, spirally disposed, bracts or 





Fig. 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 Cycadeoidea 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). 


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 1 . 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 steiile 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 Gnetum is also a trophophyte. 

For an account of the anatomy of Welwitschia and Gnetum the 
student is referred to original sources. Miss Sykes 2 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 (06 2 ); (09). 

2 Sykes (10); (10 2 ); Worsdell (01 2 ); Pearson (12); Bower (81); (82); Hill and de 
Fraine (10); Boodle and Worsdell (94); Mary R. H. Thomson (16); Henriette 
C. C, La Riviere (16). 



the Cycads to the Angiosperms 1 . It has, for example, been sug- 
gested by Hallier 2 that they are reduced Dicotyledons comparable 
with the Loranthaceae and Myxodendraceae ; while Lignier and 
Tison 3 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 4 in a paper 
on the reproductive organs of Gnetum Gnemon published in 1915, 
and in a later contribution 5 , 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 6 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-Angiosperrn 
alliance, there must be one, I think, but at present I cannot bring 
myself to believe that it is direct 7 .' 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 2 ) : additional references to literature are given at the end of this 
paper. See also Caporn (16). 

5 Pearson (17). 6 Thompson, W. P. (16). 7 Seward (17) p. ix. 


of the true position of this puzzling group which, despite the 
lack of palaeobotanical evidence, is probably a survival from a 
remote past. 

II. Gnetales (Fossil). 

Arber and Parkin 1 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 Gnetales 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. 
Keference has already been made 2 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 3 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, PI. xxvi. 


accepted as trustworthy records. The pieces of vegetative branches 
and the paired nuts described by Heer 1 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 2 Ephedrites Johnianus and similar 
specimens referred by Goeppert from the same Oligocene beds to 
Ephedra Mengeana have been identified by Conwentz 3 as fragments 
of flowering shoots of a Loranthaceous genus, Patzea. Engelhardt 4 
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 accspted 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 Welwitschui 
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 Welwilschia 5 ., though in some cases possibly 
justified by actual relationship, cannot be considered to have any 
importance unless supported by additional evidence. The seecs 
named by Renault Gnetopsis and subsequently investigated by 
Oliver and Salisbury 6 are now recognised as types closely allied 
to Lagenostoma and other Pteridosperm seeds from Carboniferous 

In their monograph of the Pliocene Floras of the Dutch- 
Prussian Border Mr and Mrs Clement Reid figure under the name 
Gnetum scandens var. robustum 1 a piece of axis 8 mm. long and 
4 mm. 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, Pis. xiv., xv. 

2 Goeppert and Berendt (45) A. Pis. iv., v.; Goeppert and Menge (83) A. PI. xvi. 

3 Conwentz (86) pp. 136, 138, PI. xm. figs. 8—20. 

4 Engelhardt (91) p. 647. 5 Seward (04) B. pp. 19, 20. 
■ Oliver and Salisbury (11) p. 34. 

7 Reid, C. and E. M. (15) p. 55, PI. xx. fig. 27. 


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 Gnetum 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 1 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, ProteaepkyUum and referred others to Ficus, Sapindopsis 
etc. A revision of the Patuxent fossils by Berry 2 has led to a 
considerable simplification in nomenclature and to the conclusion 
that some at least of these Lower Potomac leaves are Gnetalean. 

I A comparison of some of Fontaine's figures of Ficus virginiensis, 
species of Ficophyllum, ProteaepkyUum, 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- 
teaephyUum ovatum Font. 3 It is possible that a careful study of 

1 Fontaine (89) B. pp. 281 et seq. See also Seward (14 2 ). 

2 Berry (11) pp. 64, 148, 499, etc. 

3 Fontaine (89) B. Pis. 139, 141, 144, 145, etc. 


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 iv. 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. 

(Volumes III. and IV.) 

[With a few exceptions this list does not include books and papers 
given in the Bibliographies in Volumes I. and II.] 

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 Generale de Botanique' (Paris); lists 
given by Arber in the 'Progressus Rei Botanicae' (Leiden), vol. I. 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,' I. and n. (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 Riviere. ( 15) On the ribbing of the 

seeds of Ginkgo. Annals of Botany, vol. xxix. p. 591. 
Andersson, J. G. ( 10) Die Ver&nderungen 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. li. No. 7. 
(14 2 ) The Swedish Species of Ptilozamites Nath. Ibid. Bd. li. 

No. 10. 
(16) Das Fehlen resp. Vorkommen der Jahresringe in Palao- und 

Mesozoischen Holzern und das klimatische Zeugnis dieser Erschei- 

nungen. Geolog. For en. Stockholm Forhand. Bd. xxxviii. 
— (17) Die Jahresringe der Holzgewachse und die Bedeutung der- 

selben als klimatischer Indikator. Progressus Rei Botanicae, p. 285. 



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. xxviii. 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. XL pt. iv. p. 281. 

(03) On the roots of Medullosa anglica. Ann. Bot. vol. xvu 

p. 425. 

■ (03 2 ) Discussion on Dr Kurtz's paper (1903). Quart. Jour n. Cf 

Soc. vol. lix. p. 26. 

(04) Cupressinoxylon Hookeri sp. nov. a large silicified tree froi 

Tasmania. Geol. Mag. [v], vol. i. p. 7. 

•■ (05) On some new species of Lagenostoma, a type of Pterido- 

spermous seed from the Coal Measures. Proc. B. Soc. vol. lxxi. B, 
p. 245. 

(07) On Triassic species of the genera Zamites and Pterophyttum, 

types of fronds belonging to the Cycadophyta. Trans. Linn. SocS 
vol. vn. pt. vii. p. 109. 

(08) On a new Pteridosperm possessing the Sphenopteris type of 

foliage. Ann. Bot. vol. xxn. 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 2 ) On the affinities of the Triassic plant Yuccites vogesiacas 

Schimp. and Moug. Geol. Mag. [v], vol. vi. 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. v 
p. 391. 

(12 2 ) Fossil Plants from the Kent Coalfield. Geol. Mag. [i 

vol. ix. 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. B. Soc* 
vol. lxxxvi. p. 344. 

(13 2 ) 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. xxviii. p. 81. 

(14 2 ) 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 Angksperms. 
Journ. Linn. Soc. vol. xxxviii. p. 29. 

(08) Studies on the Evolution of the Angiosperms. Ann. Bot. 

vol. xxn. p. 489. 

Arnoldi, W. (01) Beitrage zur Morphologie einiger Gymnospermen. 
Bull. Nat. Moscow, No. 4, 1900. 



Bailey, I. W. (09) The structure of the wood in the Pineae. Bot. Gaz. 

vol. XLVin. 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. Geol. Soc. 

vol. xxv. p. 357. 
Bain, F. and Sir W. Dawson. (85) Notes on the Geology and Fossil 

Flora of Prince Edward Island. Canadian Bee. Set. vol. i. ( 1884-85) 

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 africanum, a new Medullosean stem. Trans. Linn. Soc. 

vol. viii. pt. ii. p. 69. 

( 14) Pteridosperm Anatomy and its relation to that of the Cycads. 

New Phyt. vol. xin. p. 41. 

Barber, C. A. (92) On the nature and development of the corky excres- 
cences on stems of Zanthoxylum. Ann. Bot. vol. VI. p. 155. 

(98) Cupressionoxylon vectense; a fossil Conifer from the Lower 

Greensand of Shanklin, in the Isle of Wight. Ann. Bot. vol. xn. 
p. 329 

Bartholin, C. T. (94) Nogle i den bornholmske Juraformation fore- 
kommende Planteforsteninger. Bot. Tidskrift (Copenhagen), Bd. 
xix. p. 87. 

(10) Planteforsteninger fra Holster hus paa Bornholm. Danmarks 

Geol. Unders. n. Raek. No. 24. 

Bartlett, A. W. (13) Note on the occurrence of an abnormal bispo- 

rangiate strobilus of Larix europaea DC. Ann. Bot. vol. xxvii. p. 575. 
Bassler, H. (16) A Cycadophyte from the North American Coal 

Measures. Amer. Journ. Sci. vol. xlii. p. 21. 
Bayer, A. (08) Zur Deutung der weiblichen Bliiten der Cupressineen 

nebst Bemerkungen iiber Cryptomeria. Beiheft Bot. Cent. Bd. xxiii. 

Abt. i. p. 27. 
Beissner, L. (91) Handbuch der Nadelholzkunde. Berlin. 
Bennett, J. J. and R. Brown. (52) Plantae Javanicae rariores. London, 

Benson, Margaret. (08) On the contents of the pollen-chamber of a 
specimen of Lagenostoma ovoides. Bot. Gaz. vol. lv. p. 409. 

(12) Cordaites 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. xxiii. p. 623. 


Berger, R. (48) De fructibus et seminibus ex formatione lithan 

Diss. Inaug. Vratislaviae. 
Bergeron, J. (84) Note sur les strobiles du Walchia piniformis. Bull. 

soc. geol. France [3], Tome xn. p. 583. 
Berridge, E. M. (11) On some 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. xxvi. 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. in. No. 9, p. 45. 

(05) Additions to the fossil Flora from Cliff wood. Bull. Torrey 

Bot. Club, vol. xxxn. p. 43. 

(06) Contributions to the Mesozoic Flora of the Atlantic Coas 

Plain. I. Ibid. vol. xxxin. p. 33. 

(07) Coastal Plain Amber. Torreya, vol. vn. p. 4. 

(07 2 ) 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 2 ) 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. xvii. p. 19. 

(09 2 ) Pleistocene Swamp deposits in Virginia. Amer. Nat. 

vol. xliii. p. 432. 

(10) A revision of the Fossil Plants of the genus Nageiopsis of 

Fontaine. Proc. U. S. Nat. Mus. vol. xxxvni. p. 185. 

(10 2 ) The epidermal characters of Frenelopsis ramosissimus. Bot. 

Gaz. vol. l. p. 305. 

(10 3 ) Additions to the Pleistocene Flora of New Jersey. Torreya, 

vol. x. p. 261. 

(10 4 ) Contributions to the Mesozoic Flora of the Atlantic Coastal 

Plain. V. North Carolina. Bull. Torrey Bot. Club, vol. xxxvn. 
p. 181. 

(11) The Lower Cretaceous deposits of Maryland. (Berry, Clark, 

and Bibbin.) Maryland Geol. Surv. 

• (ll 2 ) A Lower Cretaceous species of Schizaeaceae from Eastern 

North America. Ann. Bot. vol. xxv. p. 193. 
(II 3 ) The Flora of the Raritan Formation. Geol. Surv. New 

Jersey, Bull. 3. 

(II 4 ) A Revision of several genera of Gymnospermous plants from 

the Potomac group in Maryland and Virginia. Proc. U. S. Nat. 
Mus. vol. xl. p. 289. 

(12) The age of the plant-bearing shales of the Richmond coal- 
field. Amer. Journ. Sci. vol. xxxiv. p. 224. 




Berry, E. W. (12 2 ) Notes on the genus Widdringtoniles. Bull. Torr. Bot. 
Club, vol. xxxix. p. 341. 

(12 3 ) Contributions to the Mesozoic Flora of the Atlantic Coastal 

Plain. VII. Texas. Bull. Torrey Bot. Club, vol. xxxix. p. 387. 

(12 4 ) 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. Sure. Professional papers, No. 84. 

(15) The Mississippi River Bluffs at Columbus and Hickman, 

Kentucky, and their Fossil Flora. Proc. U. S. Nat. Mus. vol. xlviii. 
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. xliv. p. 167. 

Bertrand, C. E. (74) Anatomie comparee des tiges et des feuilles chez 
les Gnetacees et les Coniferes. Ann. Sci. nat. [v], vol. xx. p. 5. 

(83) Note sur le genre Vesquia, Taxinee fossile du Terrain 

Aachenien de Tournai. Bull. soc. bot. France [3], Tome v. p. 293. 

(89) Les Poroxylons vegetaux fossiles de l'epoque houilliere. 

Ann. soc. Belg. de Microscopie, Tome xm. Fasc. i. 

(98) Remarques sur la structure des grains de pollen de Cordaites. 

Assoc. Franc, pour Vavanc. des Sci. Nantes (1898), p. 436. 

(07) Les caracteristiques du genre Rhabdocarpus d'apres les 

preparations de la collection B. Renault. Bull. soc. bot. [4], Tome vu, 
p. 654. 

(07 2 ) Les caracteristiques du genre Diplotesta de Brongniart. 

Bull. soc. bot. France [4], tome vu. p. 388. 

(07 3 ) Les caracteristiques du genre Leptocaryon de Brongniart. 

Ibid. p. 452. 

(07 4 ) Les caracteristiques du genre Taxospermum de Brongniart. 

Ibid. p. 213. 

(07 5 ) Remarques sur le Taxospermum angulosum. Compt. rend. 

d'assoc. Frang. pour Vavanc. des sci. (Reims, 1907), p. 410. 

(08) Les caracteristiques du Cycadinocarpus angustodunensis de 

B. Renault. Bull. soc. bot. France [4], tome vni. p. 326. 

(08 2 ) Les caracteristiques du genre Cardiccarpus d'apres les graines 

silicifiees etudiees par Ad. Brongniart et B. Renault. Ibid. p. 391. 

(08 3 ) La specification 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. 

(11) 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. n. p. 243. 
Bertrand, P. (08) Sur les stipes de Clepsydropsis. Compt. Rend. Nov. 16, 


Bertrand, P. (11) Structure des stipes d' Asterochlaena 
Mem. soc. geol. Nord, Tome vn. i. 

(13) Les Fructifications de Neuropteridees recueillies dans le 

. terrain houiller du Nord de la France. Ann. soc. geol. Nord, Tome 

xlii. p. 113. 

( 14) l£tat actuel de nos Connaissances sur les Genres ■ Cladoxylon ' 

et 'Steloxylon.' Compt. Rend. de. Vaxsoc. franc, pour V Avancement 
des Sciences (Havre, 1914), p. 446. 

Beust, F. (85) Untersuchung iiber fossile Holzer aus Gronland. Neve 

Denksch. allqem. Schweiz. Ges. gesammt. Naturwiss. Bd. xxix. 
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 a l'etude des Terrains Tertiaires 

d' Alsace. Bull. soc. geol. France [3], Tome xx. p. 375. 
Bodenbender, W. (96) Beobachtungen iiber Devon- und Gondwana 

Schichten in der Argentinischen Republik. Zeitsch. Deutsch. geol. 

Ges. Bd. xlviii. p. 743. 
(02) Contribucion al Conocimiento de la Precordillera de San Juan 

de Mendoza. Bol. Acad. Nac. Cienc. Cordoba, vol. xvn. p. 203. 
Boodle, L. A. (15) Concrescent and solitary Foliage leaves in Pinus. 

New Phyt. vol. xiv. 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. Bol. vol. vin. p. 231. 
Boulay. (79) Recherches de paleontologie vegetale dans le terrain 

houiller du Nord de la France. Ann. soc. sclent. Bruxelles, ann. iv. 

pt, 2, 1880. 

(87) Notice sur la Flore tertiaire des environs de Privas ( Ardiche), 

Bull. soc. hot. France, Tome xxxiv. p. 227. 

(88) Notice sur les Plantes fossiles des gres tertiaires de Saint - 

Saturnin (Marne et Loire). J own. Bot. Ann. 2, p. 921. 

Bower, F. 0. (81) On the germination and histology of the seedlings of 
Welwitschia mirabilis. Quart. Journ. Micr. soc. vol. xxi. pp. 15, 571. 

(82) The germination and embryogeny of Gnetum Gnemon. Ibid. 

vol. xxii. [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. 

Bowerbank, J. S. (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. A Lad. Wiss. Berlin, p. 289. 

(75 2 ) Die Diagnosen dreier im Jahre 1873 von G. Wallis in Neu 

Granada entdeckter Cycadeen. Ibid. p. 376. 


Braun, C. F. W. (43) Beitrage zur Petrefactenkunde Bayreuth (Graf zu 
Miinster), Heft vi. Bayreuth. 

(47) Die Fossile Gewachse aus den Grenzschiehten zwischen dem 

Lias und Keuper des neu aufgefundenen Pflanzenlagers in dem 
Steinbruche von Veitlahm bei Culmbach. Flora, p. 81. 

(49) Beitrage zur Urgeschichte der Pflanzen. VI. Welfrichia 

cine neue Gattung fossiler Rhizantheen. Progr. Hi. Jahresber. 
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p. 47. 
Brenchley, Winifred E. (13) On Branching specimens of Lyginodendron 

Oldkamium Will. Journ. Linn. Soc. vol. xli. p. 349. 
Bristow, H. W. (62) The Geology of the Isle of Wight. Mem. GeoL 

■ Surv. Great Britain. 
Brongniart, A. (25) Observations sur les vegetaux fossiles renformes 

dans les gres de Hoer en Scanie. Ann. Sci. nat. vol. iv. p. 200. 

(28) Essai d'une Flore du gres bigarre. Ann. Sci. nat. vol. v. 

p. 435. 

(33) Notice sur une Conifere fossile du terrain d'eau douce de 

Tile d'lliodroma, Ann. Sci. nat. tome xxx. p. 175. 

(74) Etudes sur les graihes fossiles trouvees a l'etat silicifie dans 

le terrain Houiller de Saint ^tienne. Ann. Sci. nat. tome xx. [5], 

p. 234. 

(81) Recherches sur les graines fossiles silicifiees. Paris. 

Bronn, H. G. (58) Beitrage zur triassischen Schiefer von Raibl. Neues 

Jakrb. Min. p. 129. 
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Buchman, J. (45) Outline of the Geology of the neighbourhood of 

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in the Oolite quarries of the Isle of Portland. Trans. Geol. Soc. [2], 

vol. ii. p. 395. 
(37) Geology and Mineralogy considered with reference to Natural 

Theology. London. 
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und A. G. Nathorst. Cent. Min. Geol. ; Paleont. p. 442. 
Burgestein, A. (06) Zur Holzanatomie der Tanne, Fichte und Larche. 

Ber. deutsch. Bot. Ges. Bd. xxiv. Heft vi. p. 295. 

(08) Vergleichende Anatomie des Holzes der Koniferen. Wiesner- 

Festschrift, Wien. 

Burlingame, L. (08) The staminate cone and male gametophyte of 
Podocarpus. Bot. Gaz. vol. xlvi. p. 161. 


Burlingame, L. (13) The Morphology of Araucaria brasilie?isis. Bot. 
Gaz, vol. lv. p. 97. 

(15) The Morphology of Araucaria brasiliensis. Ibid. vol. lix. 

p. 1. 

(15 2 ) The Origin and Relationships of the Araucarians. Ibid. 

vol. 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. xli. ix. p. 1. 
Caldwell, 0. W. (07) Microcycas calocoma. Bot. Gaz. vol. xliv. p. 118. 
Caldwell, 0. W. and C. F. Baker. (07) The identity of Microcycas 

calocoma, Ibid. vol. xliii. p. 130. 
Cambier, R. et A. Renier. ( 10) Psygmophyllum Delvali n. sp. du Terrain 

houiller de Charleroi. Ann. soc. geol. Belg. Tome n. p. 23. (Mem. 

in 4to. ) 
Capellini, G. and Conte E. Solms-Laubach. (92) I Tronchi di Bennettitee 

dei Musei Italiani. Mem. Reale Acad, 8ci. Istit. Bologna [5], 

torn. ii. p. 161. 
Caporn, A. St. C. (16) A note on the male inflorescence of a species of 

Gnelum from Singapore. Ann. Bolus Herb. vol. n. pt. i. 
Carano, E (04) Contribuzione alia conoscenza della Morfologia e dello 

sviluppo del fascio vascolare delle foglie delle Cicadacee. Ann, di 

Bot. vol. i. p. 109 (Rome). 
Carpentier, A. (11) Sur quelques fructifications et inflorescences du 

Westphalien du Nord de la France. Rev. Gen. Bot, tome xxiii. p. 1 . 

(13) Contribution a l'etude du Carbonifere du Nord de la France. 

Mem. soc. geol. du Nord, tome vn. ii. p. 1. 

Carruthers, W. (66) On Araucarian cones from the Secondary beds 
Britain. Geol, Mag. vol. in. p. 249. 

(66 2 ) On some fossil Coniferous fruits. Ibid. vol. in. p. 534. 

(67) On Cycadeoidea Yatesii sp. nov. a fossil Cycadean stem from 

the Potton sands, Bedfordshire. Ibid. vol. iv. p. 199. 

(67 2 ) On Gymnospermous Fruits from the Secondary rocks of 

Britain. Journ. Bot. vol. v. p. 1. 

(67 3 ) On some Cycadean Fruits from the Secondary rocks 

Britain. Geol. Mag. vol. I v. 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. vi. p. 1. 

(69 2 ) On some undescribed Coniferous fruits from the Secondary 

rocks of Britain. Ibid. vol. vi. 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. vni. p. 1. 

(77) Description of a new species of Araucarites from the Coralline 

Oolite of Malton. Quart. Journ. Geol. Soc. vol. xxxin. p. 402. 




Carruthers, W. (93) On Cycas Taiwaniana, sp. nov. and C. Seemanni 
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) Einige fossile Holzer Preussens. 
K. Preuss. Geol. LandesanstaU, Bd. ix. Heft iii. p. 113. 

Celakovsky, L. (82) Zur Kritik der Ansichten von den Fruchtschuppe 
der Abietineen. Abh. K. bbhm. Ges. Wiss. Prag [vi], Bd n. 

Chamberlain, C. J. (06) The ovule and female gametophyte of Dioon. 
Bot. Gaz. vol. xlii. p. 321. 

(09) Dioon spinulosum. Ibid. vol. xlviit. p. 401. 

(10) See Coulter and Chamberlain. 

(10 2 ) Fertilization and embryogeny in Dioon edule. Bot. Gaz. 

vol. l. p. 415. 

(11) The adult Cycad Trunk. Ibid. vol. lii. p. 81. 

(12) Morphology of Ceratozamia. Ibid. vol. liii. p. 1. 

(12 2 ) A round-the-world Botanical Excursion. Pop. Sci. Monthly, 

vol. lxxxi. p. 417. 

(12 3 ) 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. 

Chapman, F. (09) Jurassic Plant-remains from Gippsland, Pt. ii. Bee. 

Geol. Surv. Victoria, vol. in. 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 Wehvitschia 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. in. No. 3. 

(08) Description of Tertiary Plants. TI. Amer. Journ. Sci. 

vol. xxvi [4], p. 537. 

(08 2 ). The Fossil Flora of Florissant, Colorado. Bull. Amer. Mus. 

Nat. Hist. vol. xxiv. p. 71. 

(08 3 ) Florissant; a Miocene Pompeii. Pop. Sci. Monthly (Aug. 

1908, p. 112). 

Coemans, E. (66) Description de la flore fossile du premier etage 
du terrain Cretace du Hainaut. Mem. Acad. R. Belg., tome 


Coker, W. C. (03) On the gametophyte and embryo of Taxodium. Bot. 
Gaz. vol. xxxvr. p. 1. 

(09) Vitality of Pine seeds and the delayed opening of cones. 

Amer. Nat. vol. xliii. p. 677. 

Compter, G. (94) Die fossile Flora des untern Keupers von Ostthuringen. 
Zeitsch. fur Naturwiss. Leipzig, Bd. lxvii. p. 205. 

(03) Cycadeenfruchte aus der Lettenkohle von Apolda. Zeitsch. 

filr Naturwiss. Stuttgart, Bd. lxxv. p. 171. 

Compton, R. H. (08) See South and Compton. 

s. iv 31 


Conwentz, H. (78) Ueber ein tertiares Vorkommen Cypressenartigei 
Holzer bei Calistoga in Californien. Neues Jahrb. Min. p. 800. 

(82) Fossile Holzer aus der Sammlung der Konig. geol. Landes- 

anstalt zu Berlin. Jahrb. K. preuss. geol. Land. Bergakad. Berlin 
fiir das Jahr 1881, p. 144. 

(85) Sobre algunos arboles fosilcs del Rio Negro. Bol. Acad. Nac. 

Cienc. Cordoba, torn. vn. p. 575. 

(86) Die Angiospermen des Bernsteins. Danzig. 

(89) Ueber Thyllen und Thyllen-ahnliche Bildungen, vornehmlich 

im Holze der Bernsteinbaume. Ber. Deutsch. Bot. Ges. Bd. vn. 
p. (34). 

(92) Untersuchungen liber fossile Holzer Schwedens. K. Svensk. 

Vetenskapsakad. Hand. Bd. xxiv. No. 13. 

(01) The past history of the Yew in Great Britain and Ireland. 

Rep. list Meeting Brit. Assoc. (Glasgow), p. 839. 

Coulter, J. M. and C. J. Chamberlain. (03) The Embryogeny of Zam ia. 

Bot. Gaz. vol. xxxv. p. 184. 

(10) Morphology of Gymnosperms. Chicago. 

Cramer, C. (68) Fossile Holzer der Arctischen Zone. Heer's Foss. Flor. 

Arct. vol. i. p. 167. 
Cri£, L. (89) Beitrage zur Kenntniss der fossilen Flora einiger Inseln des 

Sudpacifischen und Indischen Oceans. Pal. Abhand. (Dames and 

Kayser) (N.F.), Bd. i. Heft ii. 
Daguillon, A. (90) Recherches morphologiques sur les feuilles des 

Coniferes. Rev. Gen. 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. n. p. 132. 

(62) On the Flora of the Devonian period in North- east en 

America. Ibid. vol. xvm. p. 296. 

(63) Further Observations on the Devonian Plants of Mf 

Gaspe, and New York. Ibid. vol. xix. p. 458. 

(63 2 ) Synopsis of the Flora of the Carboniferous Period in Not 

Scotia. The Canadian Nat. and Geologist, vol. vtii. p. 431. 

(81) Notes on New Erian (Devonian) plants. Quart. Journ. Geol. 

Soc. vol. xxxvu. p. 299. 

(85) On the Mesozoic Floras of the Rocky Mountains Region of 

Canada. Trans. R. Soc. Canada, sect. iv. p. 1. 

(90) On new plants from the Erian and Carboniferous, and on the 

characters and affinities of Palaeozoic Gymnosperms. Canadian 
Rec. Sci. vol. iv. p. 1. 

(93) On new species of Cretaceous plants from Vancouver Island. 

Trans. R. Soc. Canada, sect. iv. p. 53. 

Dawson, Sir J. W. and D. P. Penhallow. (91) Note on the specimens of 
Fossil wood from the Erian (Devonian) of New York and Kentucky. 
The Canadian Rec. Sci. vol. iv. p. 242. 

Depape, Gr. (13) Sur la presence du Ginkgo biloba L. dans le Pliocene 
'nferieur de Saint-Marcel-d'Ardeche. Compt. Rend. vol. 157, p. 957. 


Depape, G. and A. Carpentier. (13) Presence des genus Gnetopsis B. 
Ren. and R. Zeill. et Urnatopteris Kidst. dans le Westphalien du 
Xord de la France. Ann. soc. geol. du Nord, tome xlii. p. 294. 

(15) Sur quelques graines et fructifications du Westphalien du 

Xord de la France. Rev. Gen. Bot. vol. xxvn. p. -321. 

Dorety, Helen A. (08) The embryo of Ceratozamia, a physiological study. 
Bot. Gaz. vol. xlv. p. 412. 

(08 2 ) The seedling of Ceratozamia. Ibid. vol. xlvi. p. 205. 

(09) The extrafascicular cambium of Ceratozamia. Ibid. vol. xlvii. 

p. 149. 

(09 2 ) 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. 
Douville\ H. et R. Zeiller. (08) Sur le terrain houiller du Sud Oranais. 

Compt. Rend, tome cxlvi. p. 732. 
Drude, 0. (90) Handbuch der Pflanzengeographie. Stuttgart. 
Diimmer, R. (12) Podocarpus jormosensis. 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. xliv. p. 615. 
Duns, J. (72) On Cardiocarpon. Proc. R. Soc. Edinburgh, p. 692. 
Dusanek, F. (13) Spaltoffnungen der Cycadaceen. (Abstract in the 

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Dusen, P. (99) Uber die Tertiare Flora der Magellans-Lander. Wiss. 

Ergeb. Schwed. Exped. nach den Magellansldndern unter Leitung von 

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(08) tjber die Tertiare Flora der Seymour-Insel. Wiss. Ergeb. 

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Duthie, Augusta V. (12) Anatomy of Gnetum africanum. Ann. Bot. 

vol. xxvi. p. 593. 
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Eames, A. J. (13) The Morphology of Agathis australis. Ann. Bot. 

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Senftenberg. Jahrb. K. Preuss. Geol. Land. Bergakad. Berlin, 

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ber. K. Akad. Wiss. Berlin, p. 1020. 

(82) Uber Bildungabweichungen bei Fichtenzapfen. Sitzber. K. 

Akad. Wiss. Berlin, p. 40. 

-s (89) Gymnospermae. Engler and Prantl ; Die Natilrlichen 

Pflanzenfa?nilien, Teil ii. 
Eichwald, E. (53-68) Lethaea rossica. Stuttgart. 
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Elwes, H. J. (12) The Flora of Formosa. Gard. Chron. July 
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Elwes, H. J. and A. Henry. (06) The trees of Great Britain and Ireland, 
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(47) Synopsis Coniferarum. Sangalli. 

Engelhardt, H. (85) Die Tertiarflora des Jesuitengra bens bei Kundratitz 
in Nordbohmen. Nov. Act. K. Leop.-Car. Deutsch. Akad. Natur- 
forsch. Bd. xlviii. No. 3, p. 299. 

(91) Uber Tertiarpflanzen 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. xn. 
p. 593. 

Engelhardt, H. and F. Kinkelin. (08) Oberpliocene Flora und Fauna 
des Unter-Maintales. Abh. Senck. Naturforsch. Ges. Bd. xxix. 
Heft iii. p. 151. 

Engler, A. (89) Engler and Prantl; Die Naturlichen Pflanzenfamilien, 
Teil ii. Abt, i. 

(95) Die Pflanzenwelt Ost Afrikas, etc. Th. C. Berlin. 

(97) Engler and Prantl; Die Naturlichen Pflanzenfamilien. 

Nachtrag zu Teilen ii-iv. 

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. Bee. Geol. Surv. Neiv South Wales, 

vol. in. 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. \Y . 

Haidinger, vol. iv. p. 95. 

(52) Beitrag zur naheren Kenntniss der Flora der Wealdenperiode. 

Abh. K.k. geol. Beichs. Wien, Bd. i. Abth. iii. No. 2, p. 1. 

(52 2 ) Die Steinkohlenflora von Stradonitz. Ibid. Bd. I. Abth. iii. 

No. 4. 

(55) Die Tertiare Flora von Haring in Tirol. Ibid. Bd. n. Abth. ii. 

No. 2. 

(57) Die Fossile Flora von Koflach in Steiermark. Jahrb. K.k. 

geol. Beichs. Bd. viii. p. 738. 

(58) Beitrage zur Kenntniss der fossilen Flora von Sotzka in 

Untersteiermark. Sitz. K. Akad. Wiss. Wien, Bd. xxviit. p. 471. • 

(67) Die Kreideflora von Niederschoena in Sachsen. Ibid. Bd. lv. 

Abth. i. p. 235. 

(67 2 ) Die fossile Flora des Tertiar-Bechens von Bilin. Denksch, 

Wiss. Akad. Wien, Bd. xxvi. p. 79. 


Ettingshausen, C. von. (70) Beitrag zur Kenntniss der Tertiarflora 
Steiermarks. Sitz. K. Akad. Wien, Bel. lx. Abth. i. p. 17. 

(72) Die Fossile Flora von Sagor in Krain. Denksch. Wiss. Akad. 

Wien, Bd. xxxn. p. 32. 

(78) Beitrag zur Erforschung der Phylogenie der Pflanzenarten. 

Ibid. Bd. xxxviii. p. 65. 

(79) Report on Phyto-Palaeontological Investigations of the 

Fossil Flora of Sheppey. Proc. R. Soc. vol. xxix. 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. l. p. 1. 

(86) Beitrage zur Kenntniss der Tertiarflora Australiens. Ibid. 

Bd. liii. p. 81. 

(87) Beitrage zur Kenntniss der Fossile Flora Xeuseelands. 

Denksch. K. Akad. Wiss. Wien, Bd. liii. p. 3. 

(88) Contributions to the Tertiary Flora of Australia. Mem. geol. 

Surv.N.S.W. Pal.Xo.2. 

(88 2 ) Die Fossile Flora von Leoben in Steiermark. Denksch. K. 

Akad. Wiss. Wien, Bd. liv. Abth. i. p. 261. 

' (90) Die Fossile Flora von Schoenegg bei Wies in Steiermark. 

Ibid. Bd. lvii. p. 61. 
Ewart, A. J. (08) On the longevity of seeds. Proc. R. Soc. Victoria, 

vol. xxi. [X.S.], 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. xxn. p. 289. 

(76) Xotes on the age of some Fossil Floras in India. Rec. Geol. 

Surv. Ind. vol. ix. pt. iv. p. 115. 

(76 2 ) Jurassic (Oolitic) Flora of Kach. Fossil Flora of the Gond- 

wana System, pt. i. vol. n. 1880. 

(77) Xotes on Fossil Floras in India. Rec. Geol. Surv. Ind. 

vol. x. pt. iv. p. 196. 

(77 2 ) Jurassic (Liassic) Flora of the Rajmahal group in the 

Rajmahal hills. Foss. Flor. Gond. Sijst. vol. I. pt. ii. 

(77 3 ) Jurassic (Liassic) Flora of the Rajmahal group from Golapili, 

near Ellore, S. Godaveri. Ibid. pt. iii. 

(77 4 ) Flora of the Jabalpur group (Upper Gondwanas) in the 

Son-Xarbada region. Ibid. vol. n. pt. ii. 

(77 5 ) Ueber die Gattung WiUiamsonia Carr. in Indien. Palaeon- 

tolog. Beit. Palaeonlograph. Suppl. in. Lief. iii. 

(79) Upper Gondwana Flora of the outliers on the Madras coast. 

Foss. Flor. Gond. Si/st. vol. I. pt, iv. 

(79 2 ) The Flora of the Talchir-Karharbari beds. Ibid, vol. in. 

pt, i. 

(80) The Flora of the Damuda-Panchet Divisions. Ibid. vol. in. 

pt. ii. 


Feistmantel, 0. (80 2 ) Note on the fossil genera Noeggerathia Sternbt 

Noeggerathiopsis Feist, and Rhiptozamites Schmal. Rec. Geol. Surr. 
India, vol. xin. pt. i. p. 61. 

(80 3 ) Further notes on the correlation of the Gondwana Flora 

with other Floras. Rec. Qeol. 8urv. Ind. vol. xin. pt. iii. p. 190. 

(81) Palaeontological notes from the Hazaribagh and Lohardagga 

Districts. Ibid. vol. xiv. pt. iii. p. 241. 

(81 2 ) The Flora of the Talchir-Karharbari beds. Foss. Flor. 

Gond. Syst. vol. in. pt. i. 

(81 3 ) The Flora of the Damuda-Panchet Divisions. Ibid. vol. in. 

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. vol. iv. pt. ii. 

— — (89) Ubersichtliche Darstellung der geologisch-palaeontologischer 
Verhaltnisse Siid-Afrikas. Th. i. Abh. K. bohm. Ges. Wis8, [vii], 
Bd. in. 
Felix, J. (82) Beitrage zur Kenntniss fossiler Coniferen-Holzer. Engler's 
Jahrb. Bd. in. p. 260. 

(84) Die Holzopale Ungarns in PalaeophytoJogischen Hinsicht. 

Mitt. Jahrb. K. TJng. geol. Anst. Bd. vn. 

(87) Untersuchungen iiber fossile Holzer. Zeitsch. Deutsch. geol. 

Ges. p. 517. 

(94) Untersuchungen iiber fossile Holzer. Ibid. Heft I. p. 79. 

(96) Untersuchungen iiber fossile Holzer. Ibid. Heft n. p. 249. 

Fiedler, H. (57) Die Fossile Friichte der Steinkohlen-formation. Acad. 

Caes. Leop. Nov. Acta, Bd. xxvi. p. 239. 
Fliche, P. (96) tftude sur la flore fossile de l'Argonne (Albien-Ceno- 
manien). Bidl. soc. sci. Nancy. 

(97) Note sur les nodules et bois mineralises trouves a St Parres- 

les-vaudes (Aube) dans les gres verts infracretaces. Mem. soc. 
Acad, de V Aube, tome lx. 

(99) Note sur quelques fossiles vegetaux de l'Oligocene dans les 

Alpes Francaises. Bull. soc. geol. France [3], tome xxvit. p. 466. 

(00) Contribution a la Flore fossile de la Haute-Marne ( Infra - 

cretacee). Bull. soc. sci. Nancy. 

(00 2 ) Note sur un bois fossile de Madagascar. Bull. soc. geol. 

France [3], tome xxvin. p. 470. 

(03) Note sur des bois silicifies Permiens de la vallee de Celles 


(05) Note sur des bois fossiles de Madagascar. Bull. soc. geol. 

France [4], tome v. p. 346. 

(10) Flore Fossile du Trias en Lorraine et Franche-Comte avec 

des considerations finales par M. R. Zeiller. Paris. 

Fliche, P. and Bleicher. (82) ^tude sur la flore de l'oolithe inferieure 
aux environs de Nancy. Bull. soc. sci. Nancy. 



Fliche, P. and R. Zeiller. (04) Note sur une florule Portlandienne des 

environs de Boulogne-sur-mer. Bull. soc. geol. 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. xvt. p. 261. 
Foxworthy, F. W. (11) Philippine Gynmosperms. Philipp. Journ. Sci. 

(C) Botany, vol. vi. Xo. 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. Ibid. vol. xxviii. p. 251. 

Fritel, P. H. and R. Viguier. (11) Etude anatomique de deux bois 

Eocenes. Ann. sci. not. [9], tome xiv. p. 63. 
Fujii, K. (96) On the different views hitherto proposed regarding the 

morphology of the flowers of Ginkgo biloba. Bot. Mag. Tokyo, 

vol. x. Xo. 109, p. 13. 

( 10) Some remarks on the Cretaceous Fossil Flora and the causes 

of extinction. Bot. Mag. Tokyo, vol. xxiv. Xo. 284, p. 197. 

Fnjioka, M. (13) Studien iiber den anatomischen Bau des Holzes der 
japanischen Xadelbaumc. Journ. Coll. Agric. Imp. Univ. Tokyo, 
vol. iv. Xo. 4, p. 201. 

Gardner, J. S. (86) A monograph of the British Eocene Flora, vol. n. 
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 in. Dresden und Leipzig. 

(62) Dyas oder die Zechstein -formation und das Rothliegende, 

Heft n. Leipzig. 

(63) Uber zwei neue 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. Ibid. p. 1. 

(80) Xachtrage zur Dyas I. Miltheil. 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) Oberpliocan Flora aus den Baugraben 

des Klarbeckens bei Xiederrad etc. Abh. SencJc. Naturforsch. Ges- 

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Gibbs, L. S. (12) On the Development of the female strobilus in Podo* 

carpus. Ann. Bot. voL xxvi. p. 515. 


t. Grossh. 

Gliick, H. (02) Eine fossile Fichte aus dem Neckertahl. Mitt. 
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Goc, M. J. le. (14) Observations on the centripetal and centrifugal 
xylems in the petioles of Cycads. Ann. Bot. vol. xxviii. 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 Holzer, 
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(41) Uber den Bernstein.... Uebersicht den Arbeiten und Veran- 

derungen der Schles. Ges. fur Vaterland. Kultur. Brest ax. 

(41 2 ) Taxites scalariformis, eine neue Art fossilen Holzes. Arch, fiir 

Min., Geog., etc. Bd. xv. p. 727. 

(44) Ueber die fossilen Cycadeen iiberhaupt, mit Rucksicht auf 

die in Schlesien vorkommenden Arten. Uebersicht Arbeit, und 
Verand. 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. 

(45 2 ) Description des vegetaux fossiles recueillis par M. P. de 

Tchihatcheff en Siberie. Voyage scientifique dans V Altai oriental, 
p. 379. Paris. 

(46) Ueber die fossile Flora der mittleren Juraschichten in 

Oberschlesien. Uebers. Arbeit, und Verand. Schles. Ges. vat. Kidtur 
im Jahre 1845, Breslau, 1846, p. 139. 

(47) Zur Flora des Quadersandsteins in Schlesien. Nachtrag. Nov. 

Act. Ac. Caes. Leop.-Car. vol. xxn. p. 355. 

(50) Monographic der fossilen Coniferen. N aturuerkundige Ver- 

hand. Holland. Maatschap. Wettenschappen Haarlem. Leiden. 

(52) Fossile Flora des Ubergangsgebirges. Nova Acta Caes. 

Leop. -Carol. Nat. Cur. Bd. xxn. (supplement). 

(53) Ueber die gegenwartigen Verhaltnisse der Palaontologie in 

Schlesien, so wie liber fossile Cycadeen. Denksch. Schles. Ges. fiir 
Vaterland. Kultur. 

(65) Die fossile Flora der Permischen Formation. Palaeont. 

Bd. ii. p. 1. 

(65 2 ) Uber die fossile Kreideflora und ihre Leitpflanzen. Zeit. 

Deutsch. geol. Ges. Bd. xvii. p. 638. 

(66) Beitrage zur Kenntniss fossilen Cycadeen. Neues Jahrb. 

Min. p. 129. 

(80) Beitrage zur Pathologie und Morphologic fossiler Stamme. 

Palaeontol. [N.F.], Bd. vm. iii. p. 131. 

(81) Revision meiner Arbeiten uber die Stamme der fossilen 

Coniferen, insbesondere der Araucariten, und uber die Descendenz- 
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Goeppert, H. R. and G. Stenzel. (81) Die Medulloseae. Palaeontol. 
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(88) Nachtrage zur Kenntniss der Coniferenholzer der Palaeo- 

zoischen Formationen. Abh. K. Preuss. Akad. Wiss. Berlin. 


Gomes, B. A. (65) Vegetaes Fosseis. Flora fossil do Terrens Carbonifero 
das visinhancas dio Porto, serra do Bussaio, e moinho d'ordem 
proximo a alcacer do Sal. Comm. geol. Portugal. Lisbon. 

Gordon, Marjorie. (12) Ray-traeheids in Sequoia sempervirens. New 
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Carboniferous rocks of Pettycur (Fife). Proc. Camb. Phil. Soc. 
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(12) On Rhetinangium Arbcri, a new genus of Cycadofilices from 

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pt. iv. p. 813. 

Gothan, W. (05) Zur Anatomie lebender und fossiler Gymnospermer- 
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(06) Die fossilen Coniferenholzer von Senftenberg. Abh. K. 

Preuss. Geol. Landesanst. Bergakad. [N.F.], Heft xlvi. p. 155. 

(06 2 ) Fossile Holzer aus dem Bathonien von Russisch-Polen. 

Verhand. K. Russ. Min. Ges. zu St Petersburg [ii], Bd. xliv. Lief. i. 
p. 435. 

(06 3 ) Piceoxylon Pseudotsugae als fossiles Holz. Potonie's Abbild. 

und Beschreib. Foss. Pflanz. Lief. iv. 80. 

(07) Uber die Wandlungen der Hoftiipfelung bei den Gymnosper- 

men im Laufe der geologischen Epochen und ihre physiologische 
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— — (07 2 ) Die Fossile .Holzer von Konig Karls Land. K. Svensk. 
VetensJeapsakad. Hand. Bd. xlii. No. 10, p. 1. 

(08) Die Fossile Holzer von der Seymour- und Snow Hill-Insel. 

Wiss. Ergeb. Schwedis. Sudpolar-Exped. 1901-03, Bd. in. Lief. viii. 

(08 2 ) Die Frage der Klima-differenzirung im Jura und in der 

Kreideformation im Lichte palaobotanischen Tatsachen. Jahrb. 
K. Preuss. Geol. Landes. fur 1908, Bd. xxix. Th. ii. Heft 2, p. 220. 

(09) Uber Braunkohlenholzer des rheinischen Tertiars. Jahrb. 

K. Preuss. geol. Land. Bd. xxx. Teil i. Heft 3, p. 516. 

■- (10) Die Fossile Holzreste von Spitzbergen. K. Svensk. Veten- 

skapsakad. Hand. Bd. xlv. No. viii. 

(11) Uber einige Permo-Carbonische Pflanzen von der unteien 

Tunguska (Sibirien). Zeitsch. Deutsch. Geol. Ges. Bd. lxiii. Heft 4, 
p. 418. 

(13) Die oberschlesische Steinkohlennora. Teil I. K. Preuss. 

geol. Landes. [N.F.], Heft lxxv. 

Gourlie, W. (44) Notice of the Fossil Plants in the Glasgow Museum. 

Proc. Phil. Soc. Glasgow, vol. I. 1844, p. 105. 
Grand'Eury, C. (00) Sur les tiges debout, les souches et racines de 

Cordaites. Compt. Rend, tome cxxx. (April 30). 

(04) Sur les graines des Neuropteridees. Compt. Rend, tome 

cxxxix. p. 23. 

(04 2 ) Sur les graines des Neuropteridees. Ibid. p. 782. 


Grand'Eury, C. (05) Sur les Bhabdocarpus, les graines et l'evolutio: 
Cordaitees. Ibid, tome cxl. p. 995. 

(05 2 ) Sur les graines de Sphenopteris^ sur 1'attributlon des Co- 

donospermum et sur 1' extreme variete des 'graines de fougeres.' 
Ibid. p. 812. 

(13) Recherches g6obotaniques sur les forets et sols fossiles et sur 

la vegetation et la flore houilleres, en deux parties et dix livraisons. 
Pt. i. Livr. ii., Paris et Liege. 

Graner, F. (94) Die geographische Verbreitung der Holzarten. I. Die 
Coniferen. Forstwissenschaft. Centralblalt, Berlin (August). 

Griffith, W. (59) Remarks on Gnetum. Trans. Linn. Soc. vol. xxn. 
pt. iv. p. 299. 

Groom, P. (10) Remarks on the Oecology of Coniferae. Ann. Bot. 
vol. xxjv. p. 241. 

(16) A Note on the Vegetative Anatomy of Pterosphaera 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. xli. p. 457. 
Groppler, R. (94) Vergleichende Anatomie des Holzes der Magnoliaceen. 

Biblioth. Bot. Bd. vi. Heft 31. Stuttgart. 
Grossenbacher, J. G. (15) Medullary spots and their cause. Bull. Torr. 

Bot. Club, vol. xlii. p. 227. 
Guppy, H. B. (06) Observations of a Naturalist in the Pacific between 

1896 and 1899. London. 
Gutbier, A. von. (49) Die Versteinerungen des Zechsteingebirges und 

Rothhegenden 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 for Bot. Upsala, Bd. ix. 

No. 14. 

(12) On the occurrence of Dictyozamites in South America. 

Palaeobot. Zeitsch. Bd. I. Heft i. p. 40. 

(13) Some Mesozoic plant-bearing deposits in Patagonia and 

Tierra del Fuego and their Floras. K. Svensk. Vetenskapsakad. 
Hand. Bd. li. No. 3. 

(13 2 ) The Mesozoic Flora of Graham Land. Wiss. Ergeb. Schwed. 

siidpolar. Exped. 1901-03, Bd. m. Lief. 14, p. 1. 

See Moller, H. J. and T. G. Halle (13). 

(15) Some xerophytic leaf -structures in Mesozoic plants. Geol. 

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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 Eigg. Quart. 
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(08) The Geology of the small Isles of Inverness-shire. Mem. 

Geol. Surv. Scotland. 

Harpe, P. de la. (62) See Bristow, H. W. 


Harshberger, J. W. (98) Water- storage and conduction in Senecio 

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rania), vol. n. Xo. 1. 
(11) Phytogeographic Survey of X. America. (Die Veget. der 

Erde; Engler and Drude, xui. Leipzig.) 
Hartz, N. (96) Planteforsteninger fra Cap Stewart i 0stgronland. 

Meddel. om Grfinland, xix. Copenhagen. 
Hayata, A. (06) On Taiwania, a new genus of Conif erae from the Island 

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( 10) Botanical Survey by the Govt, of Formosa. Congr. Int. Bot. 

(Bruxelles), p. 59. 

(17) Some Conifers from Tonkin and Yunnan. Bot. Magazine, 

vol. xxxi. p. 113. 

Heer, 0. (62) On the Fossil Flora of Bovey Tracey. Phil, Trans. R. Soc. 
vol. clii. p. 1039. 

(68) i. Die in Xordgronland, auf der Melville-Insel, im Banks- 
land, an Mackenzie, im Island und in Spitzbergen entdeckten 
fossilen Pflanzen. Flor. Foss. Arct. vol. I. Zurich. 

(69) Beitrage zur Kreide-Flora. I. Flora von Moletein in 

Mahren. Neue Denksch. Allgem. Schiveiz. Oes. gesammt. Naturwiss. 
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(71) iii. Die Miocene Flora und Fauna Spitzbergens. Flor. 

Foss. Arct. vol. n. 

(71 2 ) Beitrage zur Kreide-Flora. II. Kreide Flora von Quedlinburg. 

Neue Denksch. Allgem. Schweiz. Ges. gesammt. Naturwiss. Bd. xxiv. 

(75) ii. Die Kreide-Flora der arctischen Zone. Flor. Foss. Arct. 

vol. in. 

(75) iii. Xachtrage zur Miocenen Flora Gronlands. Ibid. 

(76) Flora Fossilis Helvetiae. Zurich. 

(76 2 ) Uber Permische Pflanzen von Funkkirchen in Ungarn. 

Mitt. Jahrb. K. Ung. Geol. Anst. Bd. v. 

(77) i. Beitrage zur fossilen Flora Spitzbergens. Flor. Foss. 

Arct. 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 Beitrage zur fossilen Flora Sibiriens und des Amurlandes. 


(78) v. Beitrage zur Miocenen Flora von Sachalin. Ibid. 

(81) Contributions a la Flore du Portugal. Sect. Trav. Geol. Port. 


(81 2 ) Zur Geschichte der Ginkgo-artigen Baume. Engler's Bot. 

Jahrb. Bd. I. p. 1. 

(82) i. Flora fossilis Gronlandica. Flor. Foss. Arct. vol. vr. 

(83) Flora fossilis Gronlandica. Ibid. vol. VII. 


Helmhacher, R. (71) Sitzber. d. K. Bohm. Ges. Wiss. p. 81. 

Henry, A. (06) See Elwes and Henry. 

Herzfeld, S. (10) Die Entwicklungsgeschichte der weiblichen Bliite von 

Cryptomeria japonica Don. Ein Beitrag zur Deutung der Frucht- 

schuppe der Coniferen. Sitzber. Akad. Wiss. Wien, Bd. cxix. 

Abt. i. p. 807. 
Hick, T. (95) On Kaloxylon Hookeri Will, and Lyginodendron oldhamium 

Will. Mem. Proc. Manchester Lit. Phil. Soc. [4], vol. ix. p. 109. 
Hilderhrand, F. (61) Die Verbreitung der Coniferen. Rhein. und 

Westphal. Verhand. Bd. xvin. p. 199. 
Hill, T. Gr. and E. de Fraine. (10) On the seedling structure of Gymno- 

sperms. IV. Ann. Bot. vol. xxiv. p. 319. 
Hirase, S. (98) Etudes sur la fecundation et l'embryogenie du Ginkgo 

biloba. Journ. Coll. Sci. Imp. Univ. Tokyo, vol. xn. p. 103. 
Hohlke, F. (02) Ueber die Harzbehalter und die Harzbildung bei den 

Polypodiaceen und einigen Phanerogamen. Beiheft Bot. Cent. 

Bd. xi. p. 8. 
Holden, H. S. (10) ■ Note on a wounded Myeloxylon. New Phyt. vol. ix. 

p. 253. 
Holden, Ruth. (13) Some fossil plants from Eastern Canada. Ann. Bot. 

vol. xxvii. p. 243. 

(13 2 ) Contributions to the anatomy of Mesozoic Conifers. No. 1. 

Jurassic Coniferous wood from Yorkshire. Ann. Bot. vol. xxvii. 
p. 533. 

(13 3 ) Cretaceous Pityoxyla from Cliff wood, New Jersey. Proc. 

Amer. Acad. Arts and Sci. vol. xvi. p. 609. 

(14) Contributions to the anatomy of Mesozoic Conifers. II. 

Cretaceous Lignites from Cliffwood, N. Jersey. Bot. Gaz. vol. lviii. 
p. 168. 

(14 2 ) On the relation between Cycadites and Pseudocycas. New 

Phyt. vol. xin. p. 334. 

(15) A Jurassic wood from Scotland. Ibid. vol. xiv. p. 205. 

(15 2 ) 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. Sci. vol. xvi. p. 124. 

(04) Additions to the Palaeontology of the Cretaceous formation on 

Long Island. No. II. Bull. N. Y. Bot. Gard, vol. in. No. 11, p. 403. 

(06) The Cretaceous Flora of southern N. Y. and New England. 

U. S. Geol. Surv. Mon. vol. l. 

(06 2 ) Systematic Palaeontology of the Pleistocene deposits of 

Maryland. Contributions from the New York Bot. Gard. No. 85. 

(12) Additions to the Paleobotany of the Cretaceous formation 

on Long Island. Bull. N. Y. Bot. Gard. vol. vni. No. 28, p. 154. 

Hollick, A. and E. C. Jeffrey. (06) Affinities of certain Cretaceous plant- 
remains commonly referred to the genera Dammara and Brachy- 
phyllum. Amer. Nat. vol. XL. p. 189. 


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, n. p. 47. 

Hooker, J. D. (52) Dacrydium laxifolium. Icones Plant, vol. v. pi. 815. 

(60) Flora Tasmanica. London. 

(62) On the Cedars of Lebanon, Taurus, Algeria, and India. 

Nat. Hist. Rev. p. 11. 

(63) On Welwitschia, a new genus of Gnetaceae. Trans. Linn. 

Soc. vol. xxiv. p. 1. 

Hooker, J. D. and E. W. Binney. (55) On the structure of certain 
limestone nodules enclosed in seams of bituminous coal, with a 
description of some Trigonocarpons contained in them . Phil. Trans. 
R. Soc. vol. CXLIX. 

Horich, 0. (06) Potonie's Abbildungen und Beschreibungen fossilen 
Pflanzen-Reste. Lief. iv. 69, 70. 

Howse, R. (88) A catalogue of Fossil Plants from the Hutton collection. 
Nat. Hist. Trans. Northumberland, Durham, and Newcastle-upon- 
Tyne, vol. x. 

Hutchinson, A. H. (14) The male gametophyte of Abies. Bot. Gaz. 
vol. ivn. p. 148. 

(15) On the male gametophyte of Picea canadensis. Ibid. 

vol. lix. p. 287. 

Jeffrey, E. C. (03) The comparative anatomy and phylogeny of the 
Conifers. I. The genus Sequoia. Mem. Boston Soc. Nat. Hist. 
vol. v, Xo. 10, p. 441. 

(04) A fossil Sequoia from the Sierra Nevada. Bot. Gaz. vol. 

xxxvhi. p. 321. 

(05) The comparative anatomy and phylogeny of the Conifers. 

The Abietineae. Mem. Boston Soc. Nat. Hist. vol. vr. No. 1. 

(06) The wound Reactions of Brachyphyllum. Ann. Bot. vol. xx. 

p. 383. 

(07) Araucariopitys, a new genus of Araucarians. Bot. Gaz. 

vol. xliv. p. 435. 

■ (08) Traumatic ray-tracheids in Cunninghamia sinensis. Ann. 

Bot. vol. xxn. p. 593. 
(08 2 ) On the structure of the leaf in Cretaceous Pines. Ibid. 

vol. xxn. p. 207. 

(10) A new Prepinus from Martha's Vineyard. Proc. Boston Soc. 

nat. Hist. vol. xxxiv. No. 10, p. 333. 

( 10 2 ) A new Araucarian genus from the Triassic. Ibid. vol. xxxiv. 

No. 9, p. 325. 

(10 3 ) On the affinities of Yezonia. Ann. Bot. vol. xxiv. p. 769. 

(11) The affinities of Geinitzia gracillima. Bot. Gaz. vol. l. 

p. 21. 

(12) The History, Comparative Anatomy and Evolution of the 

Araucarioxylon type. Proc. Amer. Acad. Arts Sci. vol. XLvm. 
No. 13.. p. 532. 


Jeffrey, E. C. (14) Spore-conditions in hybrids and the. mutation hypo- 
thesis of de Vries. Bot. Gaz. vol. LVin. p. 322. 

(17) The Anatomy of Woody Plants. Chicago 1917. 

Jeffrey, E. C. and M. A. Chrysler. (06) On Cretaceous Pityoxyla. Bot. 
Gaz. vol. XLn. p. 1. 

(06 2 ) The Lignites of Brandon. Contrib. from the Phanerogamic 

Labt. of Harvard Univ. No. vi. 

(07) The microgametophyte of the Podocarpineae. Amer. Nat. 

vol. xli. 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. Lxn. p. 281. 

Johnson, T. (11) A seed-bearing Irish Pteridosperm, Crossotheca 
Honinghausi Kidst. Sci. Proc. B. Dublin Soc. vol. xiii. p. 1. 

(12) Heterangium hibernicum sp. nov. a seed-bearing Heterangium 

from Co. Cork. Ibid. vol. xm. 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. B. Soc. 

Tasmania for 1886, p. 160. 
Johnstnip, M. F. (83) Recherches sur les fossiles appartenant aux 

formations Cretacee et Miocene, sur la cote occidentale du Gr0nland. 

Medd. om Grfinland, 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 2 ) 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 xi. p. 195. 

(93 2 ) Zur Entwickelungsgeschichte der Gattung Gnetum. Cohn's 

Beit. Biol. Pflanz. vr. p. 337. 

Kershaw, E. M. (09) The structure and development of the ovule of 
Myrica Gale. Ann. Bot. vol. xxiii. p. 353. 

(12) Structure and development of the ovule of Bowenia spec- 

tabilis. Ibid. vol. xxvi. p. 625. 

Kidst on, 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. veil p. 47. 


Kidston, R. (90) The Yorkshire Carboniferous Flora. Trans. Yorks. 
Nat. Union, pt. xiv. 

(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. cxcvn. p. 1. 

(04 2 ) On the Fructification of Neuropteris heterophylla Brongn. 

Proc. R. Soc. vol. lxxii. p. 487. 

(04 3 ) Some Fossil Plants collected by Mr A. Sinclair from the 

Ayrshire coalfield. Kilmarnock Glenfield Ramblers Soc. Annals 
(1901-04), No. iv. Kilmarnock. 

(05) Preliminary Note on the occurrence of Microsporangia in 

organic connection with the Foliage of Lyginodendron. Proc. R. 
Soc. vol. lxxvt. p. 358. 

(11) Les V6getaux houillers recueillis dans le Hainaut Beige. 

Mem. Mus. Roy. d'hist. nat. Belg. tome iv. 

(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. l. 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. xlvtii. pt. ii. p. 263. 
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(00) Studi