PRESENTED 


Che Trustees 


OF 


THE BRITISH MUSEUM. 


CATALOGUE 


OF THE 


MESOZOIC PLANTS 


IN THE 


DEPARTMENT OF GEOLOGY 


PARRY 2: 


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in 2007 with funding from 
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http://www. archive.org/details/catalogueofmesozO2brituoft 


CATALOGU E 


OF THE 


MESOZOIC PLANTS 


IN THE 


DEPARTMENT OF GEOLOGY 


BeLEISh MUSEUM 


(NATURAL HISTORY). 


\ / 


THE WEALDEN FLORA. 


Part IIl.—GYMNOSPERM 2. 
PLATES I1.-XX. 


BY 


ASC) SHWARD, M.A. F.G.S. 


UNIVERSITY LECTURER IN BOTANY, CAMBRIDGE. 


LONDON : 
PRINTED BY ORDER OF THE TRUSTEES. 


SOLD BY 
LONGMANS AND CO., 39, PATERNOSTER ROW. 
B. QUARITCH, 15, PICCADILLY. DULAU AND CO., 37, SOHO SQUARE, W. 
KEGAN PAUL AND CO., CHARING CROSS ROAD, W.C. 
AND AT THE 
BRITISH MUSEUM (NATURAL HISTORY), CROMWELL ROAD, S.W. 


1896. 


rs 


HERTFORD : 
PRINTED BY STEPHEN AUSTIN AND SONS. 


PREFACE. 


Tue first part of this Catalogue of the Wealden Plants 
contained figures and descriptions of the Algz, Characee, 
Equisetine, and Filicine; the present volume is devoted 
to the Cycadez and the Conifer. 

In the Author’s conclusions he gives a summary 
(pp. 233-241) of the Wealden flora comprised in these pages, 
from which it appears that the Thallophyta are repre- 
sented by 2 sp.; the Charophyta by 1 sp.; the Bryophyta 
by 1 sp.; the Equisetine by 3 sp.; the Filicine by 23 sp. ; 
the Cycadez by 24 sp.; the Conifers by 17 sp.; uncertain 
forms, 5 species: total, 76 species. 

Mr. Seward considers that “the general characters of the 
vegetation certainly seem to point to a tropical climate, 
and there can be little doubt that the temperature was 
considerably higher than the Wealden districts enjoy at the 
present day” (p. 239). He further adds that, “ Looking 
at the Wealden plants collectively, we notice a very striking 


agreement with the flora of the underlying Jurassic strata, 


vi PREFACE. 


and it would be difficult to point to any well-marked 
or essential difference between the plant-life of the two 
periods. The evidence of palaobotany certainly favours 
the inclusion of the Wealden rocks in the Jurassic series.” 
Mr. Arthur Smith Woodward informs me that the fishes of 
the Wealden beds bear testimony to the same Jurassic alliance. 
We are thus led to conclude that whereas the palzeonto- 
logical evidence, derived from the more purely marine 
deposits, would induce us to place the Wealden beds with 
the overlying and newer Cretaceous series—the peculiar 
estuary, or lake conditions, of these mostly fresh-water 
deposits, full of remains of terrestrial organisms, both 
of plants and animals, would, by their close relationship 
with the underlying and older Purbecks and Oolites, fix 
a Jurassic date to this ancient land surface upon which 


the Wealden flora once flourished. 


HENRY WOODWARD. 


GroLocicaL DEPARTMENT, 
British Museum (Naturat History), 
CromweEtt Roan, 8.W. 


November 16th, 1895. 


AUTHORS PREFACE. 


In the present volume the same method of treatment has 
been followed as in Part I. 

My thanks are again due to Mr. George Murray, 
Mr. Carruthers, and to the Assistants of the Geo- 
logical and Botanical departments generally; also to 
Mr. Rufford and Mr. C. Davies Sherborn. 

To the Director of the Royal Gardens, Kew, I am 
indebted for the facilities afforded me of repeatedly 
examining the exceptionally large collection of cycadean 
plants in the Kew Herbarium. 

I wish also to gratefully acknowledge communica- 
tions from the late Marquis of Saporta, Sir William 
Dawson, Prof. Nathorst, Prof. Lester Ward, and others; 


and to express my thanks to Miss Gertrude Woodward 


vill AUTHOK’S PREFACE. 


for the great care and artistic skill with which she has 
executed the lithographic plates. 
I am indebted to Mr. Gepp, of the Botanical Department, 


for the negative from which Plate VIII. has been printed. 


A. CSE NES: 


CAMBRIDGE, 
November 16th, 1896. 


NOTES. 


The names of authors in the footnotes, when followed by 
a number in brackets, or without a number, will be found in 
the list of works at the end of the present volume (Part II.) ; 
those followed by A. in brackets will be found in the biblio- 


graphy of Part I, 


The great majority of specimens described in Part II. are 
from Ecclesbourne and Fairlight, near Hastings, and form part 
of the Rufford Colleen 

In addition to these, there are a few fossils from the Beckles 
Collection, and from the collections of Mantell, Dawson, and 


others. 


ERRATA. 


P. 65. For Otozamites Klipsteinti var. superba, read Otozamites Klipsteinit 
var. superbus. 


P. 68. For Otozamites Klipsteinii var. longifolia, read Otozamites Klipsteinii 
var. longifolius. 


P. 89. For V. 2742, read V. 2748. 
Plate II. Figs. 1 and 2. For Saportaia, read Withamia. 


LIST OF FIGURES IN THE TEXT. 


PAGE 
1. COycadites Romert, Schenk . 9.0.05 2 .-.: - 28 
2 2 Dioonites Dunkerianus (Gopp:) = =. «+ - « ' 46 
3. ilssonia Schaumburgensis (Dunk.) . . . . . . . 55 
4, Otozamites Goppertianus (Dunk.). . . .... - 74 
PemePCLTIDEL COLUSA Te) > os 5 hal Beieco shea otek eh eee scr 89 
6. Anomozamites Lyellianus (Dunk.). . . . . .. - 93 
ie Carpolithes (Cycadacen)) . os ee, ss 108 
8. Bennettites (Williamsonia) Carruthersi, sp.nov. . . 161 


Ore Vatesias Morrisdis atts 2. ys) teeth, es 2 «| LES 


Group SPERMAPHYTA (PHANEROGAMIA). 


In Engler and Prantl’s invaluable work, Die natiirlichen Pflanzen- 
Jamilien,' we find certain innovations as regards the classification 
of plants: conspicuous among the changes suggested are the terms 
Embryophyta zoidiogama and Embryophyta siphonogama, the former 
being applied to the Bryophyta and Pteridophyta, and in the latter 
are included the Phanerogamia. 

The researches of Hofmeister, and the more recent investigations 
of Strasburger and others, have brought to light a multitude of 
facts, by which we have been led to a more exact knowledge as to 
the natural affinities between the several plant groups. Develop- 
mental study, and our more accurate perception of the homologies 
existing between the different families, have tended to emphasize 
the points of contact between the various divisions of the vegetable 
kingdom. Any system of classification is to be welcomed which 
best enables us to give expression to recognized leading charac- 
teristics, and at the same time to bring out in a concise phraseology 
the differences and resemblances between class and class. Engler’s 
new terms, if not used to supersede the older and widely known 
designations, may at least be recognized as marking a definite 
advance towards a better understanding of phylogenetic problems. 

In dealing with fossil plants we have constantly to face the 
difficulties of classification. With some writers there is a tendency 
to strain the known points of resemblance between living and 
extinct forms, and to include both in one family or sub-class; in 
other cases, needless isolation may be given to fossil genera by 
separating them from existing types. Undoubtedly the most 
natural plan is to endeavour as far as possible to fit together the 
representatives of Paleeozoic, Mesozoic, and Cainozoic genera, with 
present day plants, in a common scheme of classification. It is 
obviously impossible in the vast majority of fossil specimens, 
to discover anything of those characters on which a modern 


aS ANevIL ai, Fae, We 


2 CYCADACER. 


classification of plants is based; but we have to discriminate as 
best we can between valueless and important taxonomic features, 
and to accept within legitimate limits the assistance of evidence 
founded on analogy. To exclude fossil plants from a classification 
based on living types would be at once thoroughly unscientific and 
unnatural. Recent botany and the botany of past ages have too 
often been treated from different standpoints, and the great aim of 
paleobotanical study has thus been entirely lost sight of. The 
more we recognize the fact that plant-life, with its innumerable 
problems awaiting solution, is not confined within the limits of one 
age in the history of the earth, the sooner ought we to attain to 
a natural system of classification. 

The more important characters of the Spermaphyta (Embryophyta 
stphonogama) may be thus briefly summarized :— 

In the great majority of cases the body of the plant is differ- 
entiated into root, stem, and leaves. The embryo is formed as 
the result of fertilization, by means of a pollen-grain tube, of an 
ege-cell enclosed in a macrospore ; the fertilized egg-cell develops 
into an embryo, which more or less completely fills up the 
macrospore and macrosporangium. The seed may or may not 
be enclosed in an ovary. The gametophyte (sexual or oophore 
generation) is considerably reduced, and the sporophyte (asexual 
or sporophore generation) has become much more conspicuous than 
in the Pleridophyta. 


Class GYMNOSPERM Zi. 


Seeds naked, not enclosed in an ovary. Fertilization of the 
ege-cell by means of a pollen-tube. Vegetative structures capable 
of secondary growth in thickness. 


Order CYCADACEA. 


Stem rarely branched, leaves large and generally pinnate. In 
the recent genera flowers always dicecious, and without a perianth. 
The Order Cycadacee, like the Marattiacee among ferns, affords 
an instance of a series of plants of which few survive at the 
present day, but which was abundantly represented in the vege- 


CYCADACER. 5 
4 
tation of former periods. The recent cycads are usually divided 
into nine genera and two families: the Cycade@, including one 
genus, Cycas; and the Zamiee, with the genera Zamia, Cerato- 
samia, Macrozamia, Dioon, Encephalartos, Stangeria, Bowenia, and 
Microcycas.* 

None of the living cyeads occur outside tropical or subtropical 
regions. In Tertiary times the family does not appear to have 
had a wide distribution, nor to have been represented by many 
genera; possibly, however, a closer acquaintance with extra- 
European Tertiary strata may bring to light a greater number 
of cycads from these beds than are at present known. In the 
Mesozoic period cycads occupied a prominent position, and had 
an extended geographical range. The Jurassic strata afford 
abundant evidence that cycadean plants reached their maximum 
development in that era; less numerous in the Triassic vegetation, 
the Cycadacee dwindle down to a few representatives in the 
Permian and Carboniferous floras. 

Before giving a summary of the earlier geological history of this 
exceedingly interesting section of the Gymnosperme, we may take 
note of some of the difficulties which beset any attempt to trace the 
geological history of cycadean piants. As in the case of ferns, and 
indeed of all fossil plants, so here again we have to deal in nearly 
every instance with detached and isolated specimens of stems, 
fronds, flowers, and seeds. The fronds are often abundant enough, 
and their preservation frequently good; but the characters which 
are made use of in generic and specific determinations are such as 
preclude any certain conclusions as to precise botanical affinity. 
The nature of cycadean flowers, and their manner of occurrence 
on the plant, separated as they are from the sterile fronds, present 
an obstacle to exact determination. On the other hand, the fronds 
alone afford, in many instances, convenient data on which to found 
a provisional classification ; their form and general habit of growth 
are fairly uniform, and they do not present the same striking 
variation in leaf form which constitutes one of the many difficulties 
associated with the fronds of fossil ferns. Among recent cycads 
we have a phyllopodium exhibiting, in the majority of species, 
certain distinct and easily recognized characters; usually a pinnate 
structure, with stout and more or less closely set segments 


1 Engler and Prantl, Teil ii. p. 6. See also De Candolle. 


4 CYCADACER. 


traversed either by a single midrib or by a number of equal and 
parallel veins. There are, however, certain variations from the 
familiar cycadean type, even in some of the living genera. In 
the South African genus Stangeria,’ originally described in 1835 
as a fern, the pinne possess a fern-like venation, forming a 
strongly marked contrast to the usual Zamia or Cycas type. 
Writing of this plant in 1854, Smith? notes that the peculiar 
character of the leaf segments renders untenable the criterion of 
venation, usually relied upon in discriminating between fossil ferns 
and eycads. Among fossil leaves there are various genera which 
have been assigned to cycads or ferns according to the preference 
of different authors. The well-known genus Wilssonia has been 
placed by Schenk and others among the /ilicine, but it is usually 
referred to as an extinct member of the Cycadacee; the widely 
distributed Zeniopteris has been assigned to both ferns and cycads, 
but it is generally regarded as a genus of fossil ferns. The genus 
Dictyozamites* and numerous others might be cited as examples 
of doubtful forms which cannot with any certainty be assigned 
either to the Pleridophyta or Gymnosperme. 

In a recent work on the Coal-Measures of Gard, Grand’Eury ‘* 
includes certain leaf forms in the class of gymnosperms, but by 
other writers these have usually been described as ferns. To settle 
such doubtful cases as these, Bornemann® undertook a minute 
comparative examination of the epidermal cells of recent ferns 
and cycads, and found what he considered fairly safe guides 
in the rectangular or wavy outlines of the epidermal cells of 
the leaves of these two sets of plants. Schenk*® has followed 
Bornemann’s example in making use of this anatomical character 
in the case of carbonized epidermal tissues of doubtful fossil 
leaves, but the fern-like wavy walls in the epidermal cells of 
Stangeria leaves preclude any trustworthy reliance on such a 
method of separating ferns and cycads. 


‘ Hooker, Bot. Mag. Pl. 5121, vol. xv. [8] 1859. Reference given to 
Kunze, ete. 

2 Smith, p. 88. 

3 Nathorst (1). 

* Grand’ Eury (1), p. 301. 

5 Bornemann. 

® Schenk (A. 1), Flor. foss. Grenz. Keup. Lias. 


CYCADACE. 5 


As a general rule, the fronds of recent cycads are simply 
pinnate; but in the Australian genus Bowenda,’ with its bipinnate 
leaves, we have an exception to this rule; and it is by no means 
improbable that this character may have been shared by many 
extinct genera. The late Dr. Stur, of Vienna, expressed his 
belief that the well-known Coal-Measure fossils Vewropteris and 
Alethopteris should be included in the list of Paleozoic cycads, 
and this opinion was partly founded on the resemblance of the 
Carboniferous fronds to the branched leaf of Bowenia. The absence 
of any clearly proved fructification in these so-called ferns has 
been referred to by Stur and others in favour of a cycadean 
relationship. Kidston? has recently recorded the occurrence of 
a fertile Neuropteris frond, but the facts he publishes cannot be 
regarded as finally settling the position of these genera. He 
figures a terminal portion of a specimen ‘‘ending in a number of 
dichotomous branchlets, the ultimate divisions being about 8 mm. 
long, and bearing the fruit at their summits.’ Unfortunately the 
very small pinnules associated with this fragment do not furnish 
all the evidence one could desire as to the real nature of the 
specimens. 

Another aberrant form of a recent frond is afforded by the 
Australian cycad Maecrozamia heteromera, Moore,? in which the 
pinne are in some varieties of the species repeatedly forked, 
reminding one to some extent of the Mesozoic species of Baiera. 
Specimens of Macrozamia heteromera, var. Narrabri, and var. glauca, 
in the Royal Gardens, Kew, show very clearly this striking and 
unusual character in cycadean fronds. (Pl. XIII. Figs. 1 and 2.) 

A further variation in the form of cycadean leaves is seen in 
such species as Zamia Skinnert, Warscew, Z. picta (=Z. muricata, 
Willd.),* Z Wallisii, A. Gr., etc.: the pinne of these forms reach 
an unusually large size, and differ in shape from those of most 
members of the family. A single pinna of Z Wallisic in the Kew 
Herbarium measures 87 X 13 cm.; the lamina is traversed by 
a few prominent and forked veins, and exhibits another peculiarity 
in the possession of a short petiole. If we have to rely on leaves 


1 Hooker, Bot. Mag. Pl. 5398, vol. xix. [3] 1863. 
? Kidston (1), p. 150, pl. viii. fig. 7. 

3 Moore, p. 122. 

4 De Candolle, p. 541. 


6 CYCADACEX. 


alone we must necessarily expect to fall into error, but it is 
important not to bind ourselves too closely to the more common 
forms of cycadean fronds in endeavouring to determine the leaves 
of extinct species. Seeing that the existing genera of cycads are 
obviously but a few remnants of a once vigorous and numerous 
family, we should not neglect the less known and more aberrant 
forms of fronds in our comparisons of fossil and recent specimens. 
We are accustomed to include in the Cycadacee a large number 
of Jurassic and Lower Cretaceous fronds which possess some more 
or less close external resemblance to those of living species. That 
such determinations are correct we have no absolute proof, but 
can only trust to the distinctly cyeadean form whieh the leaves 
present. It is possible that among such Mesozoic genera there 
are included some which should rather come under the head of 
Bennettitee, a group of plants nearly allied to the true ecycads, 
but which possess certain peculiarities of structure of sufficient 
importance to exclude them from the Cycadacee as at present 
defined. Silicified stems from the Upper Jurassic and Lower 
Cretaceous rocks of England, France, Italy, America, and other 
places, agree in anatomical structure with the stems of recent 
eycads, but in organic connection with some of these fossil forms 
there has been found a special type of inflorescence, showing a more 
highly organized and specialized structure than is afforded by the 
flowers of existing Cycadee or Zamiee. Our knowledge of the 
vegetative and reproductive structures of Bennettites is mainly 
due to the researches of Carruthers,! Solms-Laubach,? and more 
recently Lignier.* The Bennettitee inflorescence presents certain 
points of contact with the Conifer, and the characters it possesses 
in common with and distinct from those of ecycadean flowers 
suggest that “‘the Bennettitee are posterior to the Cycadacea, at 
least as regards the reproductive structures.”” As Lignier has said 
in his recent paper, we may perhaps regard the Bennettitee as 
a family which has been derived with the cycads from common 
ancestors. We have still to learn what forms of frond were 
possessed by these stems. Carruthers‘ speaks of a ‘‘ remarkable 


Carruthers (1). 
Solms-Laubach (1 and 2). 


Lignier. (For abstract of this paper see Nature, October 18, 1894, p. 594.) 
Loc. cit. p. 697 (footnote). 


CYCADACER. a 


cycadean leaf” from the Lower Greensand, which he suggests may 
possibly represent a frond of Bennettites; the specimen referred to 
is not in organic connection, nor in any close association, with 
a stem, and therefore no satisfactory conclusion can be drawn as 
to its real nature. As yet we can only reply to the question as to 
what was the precise form of Bennettites leaves by mere guesses, 
founded on no surer basis than a vague suspicion of probability. 
The leaf-scars on the surface of the stems suggest a frond of 
eycadean habit; and in all probability many of the Mesozoic leaves 
which we are accustomed to connect with true cycadean stems 
should be referred to Bennettites. To include all cycad-like fronds 
in the Cycadacee as defined for existing species, would almost 
certainly result in assigning many fossil leaves to a wrong position. 
Possibly the better plan would be to assign such fossil fronds as 
may reasonably be referred to cycadean plants, to some more 
comprehensive Natural Order than that of the Cycadacee. 

This brings us to the question of intermediate forms, and the 
association of cycadean structure with several of these synthetic 
types lends an increased interest to the past history of cycads, and 
at the same time enhances the difficulty of systematic treatment. 
The Upper Carboniferous genus Dfyeloxylon (Stenzelia, Goppert, 
Uyelopteris, Renault), found in England, France, and Germany, 
has been assigned by several writers to the Filicing, and placed 
in the Marattiacee or Ophioglossacee; others prefer to include 
it with the cycads. The structure of the vascular bundles of 
Myeloxylon petioles! is in some respects typical of recent cycads ; 
the spiral protoxylem elements being on that side of the xylem 
facing the phleam. The bundles are collateral in form, and often 
accompanied by mechanical or stereome elements. The fundamental 
tissue contains numerous secretory canals, and in some cases strands 
of stereome. One of the most readily recognized features is the 
hypodermal tissue, made up of alternating bands of thick walled 
fibres and thin parenchymatous cells. Occasionally the petiolar 
axis is found to be branched, and small Pecopteris-like pinnules 
have been observed attached to a slender Myeloxylon midrib. This 
discovery by Renault of pinnules in connection with I]yeloxylon 


1 Seward (1). References given to other papers; see also Zeiller (1), p. 290, 
pl. xxvii. fig. 1. 


8 CYCADACER. 


appears to be confirmed by some specimens in the Binney Collection * 
of Coal-Measure plants. In one instance this form of petiole has 
been found inserted on a stem of IMedullosa Leuckarti, Gopp. and 
Stenz., a plant with distinctly cycadean characteristics. Probably 
we may regard Myeloxylon as a synthetic or intermediate form 
exhibiting cycadean and fern characters, but more nearly allied to 
existing Cycadee than to the Lvlicine. In the Coal-Measure genus 
Lyginodendron,® originally described in detail by Williamson in 
1873, we have another important link in the chain of cycadean 
phylogeny. A revision of the English specimens of this plant, and 
an examination of fresh material by Williamson and Scott, has 
brought into greater prominence the clearly defined cycadean 
features exhibited by the Lyginodendron stems. It has recently 
been shown by these observers that Williamson’s genus Aaloxylon 
represents the root of ZLyginodendron, and we have previously 
learned that Rachiopteris aspera, Will., with its sphenopteroid 
pinnules, is a branch of the same plant.’ This is, again, an instance 
of cycadean and pteridophytic characters combined in a synthetic 
genus. The presence of secondary vascular tissue in Lyginodendron 
lends additional interest to this instance of fern-cycad alliance. 
In speaking of the occurrence of diploxyloid structure in this 
genus, Bertrand and Renault‘ regard the existence of such a type 
of vascular bundle in the petioles of recent cycads as a remnant 
of an ancestral structure. 

The same diploxyloid arrangement occurs on an extended scale 
in the Permo-Carboniferous genus Pororylon,® and must be looked 
upon as an important aid in any attempt to trace the lines of 
development of the Cycadacee. Renault has founded the genus 
Cycadoxylon® on a fragment of a silicified branch from Autun, 
in which the structure of the wood and fundamental tissue bears 
a distinct resemblance to a young ecycadean stem. He suggests 
that this type may find its true position between cycads and 


‘ Now in the Woodwardian Museum, Cambridge. 

* Williamson (1, part iv.). The name was proposed by Gourlie in 1843. 
(Williamson, p. 393.) See also Solms-Laubach (A.), Fossil Botany, p. 38. 

8 Williamson (1, pt. vi.), p. 684; also (1, pt. xiii.), p. 298. 

ai) poet 

° Bertrand and Renault (2). 

§ Renault (1), p. 283. 


CYCADACER. 9 


Cordattee. Unger’s genus Cordaites,! with its large parallel 
veined leaves and tall woody stem, affords another example of 
the occurrence of cycadean structures in association with 
anatomical features suggestive of another set of plants; in this 
case it is with the Conifere that cycadean characters appear to 
be combined. In the Mesozoic floras we have Carruthers’ genus 
Bennettites, to which reference has already been made, with its 
combination of cycadean and coniferous characters. Another 
and less accurately known plant, Willamsonia,® offers a difficult 
problem to the palzobotanist; but here, again, we have probably 
to deal with a synthetic type closely allied to Bennettites. 

Enough has been said to show the promising character of 
the study of the geological history of cycads, and we may not 
unreasonably entertain the hope, that we are within a measurable 
distance of deciphering some of the earlier chapters in the records 
of cycadean development. 

Before considering the questions of terminology and the details 
of generic and specific determination of fossil cycadean fronds, we 
may briefly pass in review the recorded facts as to the past history 
of the Cycadacee, and especially such as have reference to the 
representatives of this order in Paleozoic times. In 1868 
Carruthers* expressed the opinion that ‘‘no satisfactory evidence 
exists of the occurrence of Cycadee in any Paleozoic formation.” 
It is true that the facts we at present possess do not allow us 
to affirm that the Paleozoic strata contain examples of plants 
which exhibit typical cyeadean structure, and of such a kind as 
to warrant their inclusion in the Cycadacee as at present defined. 

It has already been shown that certain typical features of cycad 
structure are met with in various Permo-Carboniferous genera, 
but these are associated with other morphological characters 
which are unknown among recent representatives of this class 
of gymnosperms. It would, indeed, be a matter of surprise if 
we found in Paleozoic strata a perfectly typical cycadean genus. 
In the case of Jurassic plants we speak unhesitatingly of cycad 
leaves, although we cannot as a rule support such assertions with 
facts of anatomical details or floral structure If external resem- 


1 Renault (1), p. 323. 
2 See Bennettites. 


3 (1), p. 676. 


10 CYCADACER. 


blance of leaf form is to be trusted at all, we must admit the 
existence in Upper Paleozoic rocks of a few fossil fronds, which 
have as much claim as those from Jurassic strata to be classed 
among the Cycadacee. In reviewing the evidence in favour of 
Paleozoic cycads, we may for convenience sake consider Permian 
and Carboniferous specimens together. 

In 1848 Gutbier! figured and described a Rothliegende plant 
from Rheinsdorf, near Zwickau, which he designated Pterophyllum 
Cotteanum. The figure reminds one to some extent of Ctenis 
falcate, L. and H., but the pinne show no trace of any anasto- 
mosing venation; the specimen cannot well be excluded from the 
provisional cycadean genus Pterophyllum. Eichwald? has figured 
a portion of a frond from the Carboniferous rocks of Konznetzk 
in the Altai Hills, under the name of Pterophyllum inflecum ; this 
also seems to conform to the recognized characters of Pterophyllum. 
Carruthers * has referred to some stems described by Eichwald from 
Russian Permian rocks, but is of opinion that they cannot be 
accepted as satisfactory examples of Paleozoic cycads; the same 
author also calls attention to the specimens described by Presl and 
Guillard as cycadean stems, and shows that they have no claim to 
be placed among fossil eycads. Schmalhausen‘* has more recently 
figured a stem fragment from the Permian of Kargala in Orenburg, 
which he refers to Schimper’s species Clathraria strigata, but 
regards the specimen as a stem of Cordaites lancifolius, Schmalh. 
In 1864 Sandberger® recorded a species of Pterophyllum, P. 
blechniodes, from the Upper Coal-Measures of Holzplatze, near 
Oppenau; the specimen seems to have been reasonably placed 
among cycadean fronds. Gdppert,® in 1843, described what he 
considered to be the oldest known cyecadean frond; this imperfect 
fragment from Konigshiitte, in Silesia, he named Pterophyllum 
gonorrachis. Two other specimens were recorded by the same 
author from Paleozoic strata as Cycadites gyrosus and Cycadites 
taxodinus ;’ the former is a small and imperfect specimen which 


— 


A.) Verstein. Roth. Sachsen, p. 21, pl. viii. fig. 7. 


1 
Vols tap ola, ol artes ae ‘ 
* (1), p. 675. 

4 (1), p. 37, pl. v. figs. 4 and 5. 

5 (1), p. 34, pl. ii. figs, 1-4. 

S5(1), Pp. 00,apl. a. digeens 

7 (2), p. 181, pl. ii. figs. 1-34, 


CYCADACEZ. 11 


it is hardly possible to definitely refer to either eycads or ferns; 
the latter specimen, from the Culm beds, is more distinct, but still 
by no means a satisfactory proof of the existence of a cycadean 
species in the Culm flora. Solms-Laubach! considers that Goppert 
was probably justified in referring the last-named species to the 
Cycadee. An examination of the type specimens in the Breslau 
Museum of these two species of Cycadites led me to regard 
C. gyrosus as too imperfect for identification, and suggested the 
possibility that C. taxodinus might perhaps be regarded as a 
fragment of a coniferous branch. If the evidence for Carboniferous 
cycads rested simply on Goppert’s specimens it would be of little 
value; but there have been many more perfect examples recorded 
from this formation. From the Permo-Carboniferous rocks of 
France we have several records of cycadean fronds. ‘ihe genus 
Pterophylium has been discovered in the Upper Carboniferous beds 
of Montchanin (Saone-et-Loire), and the fragment is figured by 
Saporta and Marion as Pterophyllum Grand’ Euryanum, Sap. et 
Mar.’; the form of the pinne and their manner of attachment to the 
rachis support this determination. Another species is recorded by 
Renault, under the name of Sphenozamites Rochei,* from the Permian 
of Autun; the figure of this plant, given by Saporta and Marion,‘ 
suggests a strong likeness to Noeggerathia, and it may be that if, 
as some believe, the latter genus must be assigned to the /ileine, 
the same position should be given to Renault’s species. Moeggera- 
thia may be left for the present as one of those doubtful forms 
which cannot be definitely assigned to any clearly defined position. 
From the Commentry coal-field, from which so many interesting 
additions have been made by Renault and Zeiller to the Coal- 
Measures flora, we have several new species of cycadean leaves. 
Zamites carbonarius, Ren. and Zeill.,° is the name given to the 
largest of a set of frond fragments from a particular locality in 
this coal-field; the type specimen consists of a portion of a stiff 
rachis bearing a few alternately placed oval pinne, and the form 
of the segments is not unlike that of Noeggerathia. In addition to 


Fossil Botany, p. 86. 

Saporta and Marion, vol. i. p. 109. 

Renault (2). 

Loe. cit. p. 109. 

Flor. Commentry, p. 614, pl. lxvii. fig. 7. See also Renault and Zeiller (1). 


ao Fr Oo ND 


(0 CYCADACER. 


this species, the same authors institute five other specific names? 
for isolated pinne which do not appear to afford any distinct 
indication of specific difference. Potonié,? in his recent work on 
the Permian flora of Thiringen, includes all these five species 
under Zamites carbonarius, and an examination of the figured 
pinne certainly lends support to this view. Zeiller* has defended 
Renault’s determination, on the ground that there are certain 
differences in the venation and form of the pinne which are 
hardly consistent with the suggested inclusion under a single 
species; he is, however, willing to admit that possibly Zamites 
regularis may be identical with Z. Planchardi. Whatever may be 
the specific value of these Commentry specimens, Zeiller regards 
them as undoubtedly fragments of the same generic form, and the 
discovery of more perfect specimens leads him to found a new 
genus, Plagiozamites, as more suitable for their reception than 
Zamites. In speaking of the resemblance between Plagiozamites 
and Noeggerathia, Zeiller expresses an opinion in favour of 
including the latter genus among cycads, using the term cycads * 
in a wide sense. This opinion is partly based on the close 
similarity between Noeggerathia and Plagiozamites on the one 
hand, and on the marked resemblance between the latter genus 
and Zamites on the other. The form of the fronds certainly 
favours this view, but such reasoning from external resemblance 
cannot be accepted as conclusive when we are dealing with cycads 
and ferns. In all these cases we must be prepared to find a 
combination of pteridophytic and cycadean characters, and if we 
were in possession of the facts of anatomical structure, we should 
possibly be quite unable to decide definitely for one or other of 
these two groups of plants. | 
The Commentry flora has furnished an exceedingly fine specimen 
of the genus Pterophyllum*’—P. Fayoli, Ren. and Zeill. This 
example is unusually large and well preserved, and there can be 
little or no hesitation in accepting it as a Paleozoic cycadean 
frond, having an equally strong claim to be described as such as 


1 Flor. Commentry, pp. 615-617, pl. Ixvii. figs. 8-19. 
31), p. 200: 

2 (2) ip. aLia. 

4 Ibid. p. 179. 

® Renault and Zeiller (2), p. 619, pl. lxviii. fig. 1. 


CYCADACEX. 13 


the Mesozoic representatives of the same genus. The genus itself 
is merely a provisional one, and rests on external characters of 
vegetative structures, but the cycadean habit is sufficiently obvious 
to lend confidence to the generally accepted botanical position 
assigned to this and other cycad-like leaves. Portions of gigantic 
leaves are figured by Renault and Zeiller from the Commentry 
coal-field under the generic name Zttanophyllum,! and it is 
suggested that possibly these may belong to Calpoxylon stems, 
which have been referred on anatomical grounds to the Cycadacee, 
but these and many other leaf forms must remain in the list of 
plante incerte sedis until additional facts are available. Renault 
has recently described another species of Permian ecycad, Ptero- 
phyllum Combrayi,? which shows a fairly close resemblance to 
P. Jaegeri, Brong. Enough has been said to show that in Permo- 
Carboniferous times there existed certain forms of leaf structures, 
which must be assigned with the numerous Mesozoic fronds to the 
provisional genera of extinct cycads. The large number of seeds 
from this geological horizon, with their well-preserved structure 
and variety of external form, are naturally a source of difficulty 
as regards systematic position. There are distinct indications of 
eycadean affinity in many of the silicified gymnospermous seeds ; 
some belong, no doubt, to Cordaites, whilst others may be more 
correctly placed in the Conifere. The seeds of the recent genus 
Ginkgo show some points of contact with those of cycads, and 
among the seeds of Paleozoic plants it would not surprise us 
to find cycadean and coniferous characteristics represented in the 
same species. We cannot well do more than speak of these doubtful 
fossils as examples of Paleozoic gymnospermous seeds, many of 
which distinctly resemble the seeds of recent cycads. Grand’Eury® 
includes many such fossils in the family Moeggerattiacee, a sub- 
section of gymnosperms; the choice of this name is not a very 
happy one, seeing that we know so little as to the actual position 
of Sternberg’s genus Woeggerathia. 

Ascending the geologic series from the Permian to the Upper 
Jurassic strata, we find a gradual increase in the number and 
variety of cycadean fronds, and in the Wealden vegetation the 


1 Renault and Zeiller, p. 622, pl. Ixix. 
= (3)5 Us G72. 
3 (1), p. 301. 


14 CYCADACER. 


Cycadacee were represented by many large and striking species. 
Further reference will be made to the Lower Cretaceous cycads in 
the general review of the Wealden flora at the end of this volume. 
Throughout the Cretaceous and Tertiary series we have evidence 
of a decline in the relative importance and numerical proportion 
of the Cycadacee. It has been suggested that possibly the paucity 
of species may in some measure be explained by our very imperfect 
acquaintance with tropical and subtropical Cretaceous and Tertiary 
plant-bearing strata;’ it may be that the rocks of these eras were 
deposited under climatal conditions which were not favourable to 
a rich development of cycads. Heer? has described various frond 
fragments from Tertiary beds which are not particularly satis- 
factory as records of cycadean species. The two species Wilssonia 
Serotina, Heer, and V. pygmea, Heer, from the Miocene flora of 
Sachalin Island, are both founded on fragments which may possibly 
belong to that doubtful genus in which they have been placed. 
From the Upper Fresh-water Molasse of Schaffhausen, the same 
author describes a structureless stem as Cycadites Escheri, Heer ;* 
the appearance of the scale-covered surface lends some support to 
this determination, but the specimen is too imperfect to be of any 
particular importance. Heer figures a fragment of a frond from 
Lausanne under the name Zamites (Dioon?) tertiarius,’ founded 
on a poor and fragmentary specimen. Three species of Tertiary 
cycads are figured by Saporta and Marion in their ? Evolution 
du régne végétal:* one of these is assigned to Zamiostrobus—Z. 
Saportanus, Schimp., and may possibly be rightly described as 
a cycadean cone, but its precise nature cannot be definitely 
ascertained. The other two species, Zamites epibius, Sap., and 
Encephalartos Gorceixianus, Sap., are most probably true cycads. 
Ettingshausen’s New Zealand specimen, described as Zamites sp. ? 
cannot be accepted as trustworthy evidence of a Tertiary cycad.° 
From Australia the same author records Anomozamites Muelleri,® 
Ett., a species based on small fragments of what may be a 


1 Solms-Laubach, p. 85. 

2 Flor. foss. Arct. vol. v. (Flor. Sachalin), pp. 19 and 21, pl. ii. figs. 1-6. 
3 (A.) Fl. Tert. Helvet. p. 46, pl. xv. and pl. xvi. fig. 1. 

4 Les Phanérogames, vol. i. p. 116. 

5 Kttingshausen (1), p. 13, pl. i. fig. 10. 

6 (2), p. 9, pl. viii. figs. 19-22. 


CYCADACER. 15 


cycadean leaf. Another possible Tertiary cycad is described by 
Ettingshausen from the Miocene beds of Leoben; to this the 
name Ceratozamia Hoffmanni, Ett., has been assigned.’ The single 
imperfect pinna which is figured by the author of the species, 
does not afford sufficient evidence that it belongs to this particular 
recent genus. Granting its cycadean nature, and even this entails 
a considerable amount of faith, there is surely no reason why the 
fragment should not be referred to some other genus than the one 
chosen; one might suggest ‘‘ cycadean pinna?”’ as a more fitting 
term than C. Hoffmanni. Goppert’s Tertiary species of a 
Greenland cycad, Zamites arcticus,* is founded on a fairly good 
specimen, and certainly appears to be correctly included among 
the Cycadacee. These few examples of fragments described by 
various writers as cycadean fronds, sufficiently demonstrate the 
meagre relics of this order of gymnosperms in Tertiary rocks. 

In his Monograph on the Jurassic cycads, Saporta*® has given 
a useful and critical summary of the history of the literature on 
fossil Cycadace@, to which is added a series of definitions of the 
chief characters by which the several genera of fronds may be 
recognized. Certain suggested emendations of some of these 
diagnoses will be found under the head of the respective genera 
in the descriptive part of this Catalogue. 

Without following the gradual additions to our knowledge of 
fossil cycadean fronds during the last sixty or seventy years, 
or attempting to discuss the numerous classifications proposed by 
various writers, it may serve a useful purpose to draw attention 
to some of the difficulties and possible sources of error associated 
with the investigation of the past history of cycads. 

The characters generally made use of in the separation of 
distinct genera of fossil cycadean leaves may be enumerated as 
follows: (i.) The method of attachment of the pinne to the 
rachis, and whether persistent or deciduous. (i.) The nature of 
the base of the pinne, auriculate or gradually tapered, etc., the 
presence or absence of a distinct basal callosity. (iii.) The pinna 
apex, whether truncate, acuminate, ete. (iv.) Venation. (v.) The 
angle of insertion of the pinne on the rachis; the alternate 


1 (3), p. 272, pl. iil. fig. 10. 
2 Goppert (2), p. 134, pl. ii. figs: 9 and 10. 
3 (A. 2), Pal. Frang. [2] vol. ii. 1875, pp. 26-45. 


16 CYCADACER. 


or opposite arrangement of the pinne. (vi.) The form of the 
epidermal cell-walls. (vii.) Presence of spines on the segment 
margin. In addition to these more detailed characters, the form 
of the frond as a whole, whether simple, pinnate, or bipinnate, 
and the shape of the individual pinne, long, narrow, broadly 
oval, etc., are important characters to be kept in view. 

In Géppert’s valuable paper on fossil cycads,' the wholesome 
warning is given that to define generic characters within such 
narrow limits as are often adopted, results in an unnecessary 
multiplication of genera, and tends to confusion and to increase 
the difficulties of determination. Allusion has already been made 
to the numerous leaves, the affinities of which cannot be definitely 
settled until further data are forthcoming. As regards the genus 
Nilssonia, some writers have argued for its inclusion among ferns, 
but others prefer to consider it an unusual form of cycadean 
frond. Zeniopteris, Neuropteris, Noeggerathia, and a host of other 
leaves must for the present be left in a somewhat doubtful 
position. The genus Stangerites, instituted by Bornemann,? has 
been used by a few authors as a convenient term for certain 
Teniopteris-like leaves, but the name seems unnecessary, and 
may be ranked among those misleading titles which suggest a 
relationship to a living genus which is not supported by facts 
of any taxonomic value. Saporta, in speaking of this genus, 
remarks that the author of the term Stangerites ‘‘a ajouté a ce 
qui s’était fait avant lui une confusion réellement inextricable et 
périssé de difficultés la synonymie des principales espéces, decrites 
d’aprés leurs feuilles seulement.’ * 

In the recent species Stangeria paradoxa (Moore), it is worthy 
of note that we have pinnz with entire margins, and others with 
deeply cut lobes extending to the midrib; some of the deeply 
divided lamine suggest in a slight measure a WVilssonia form of 
leaf. In a small plant of Cycas circinalis, L., in the Royal 
Gardens, Kew, I noticed an abnormal form of leaf structure at 
the base of a young frond, suggesting another example of an 
approach to the Wilssonia type of leaf. Instead of the ordinary 
uninerved and separate pinne characteristic of Cycas, this 


(CN) jo Ta 
2 p. 58, misspelt ‘‘ Strangerites, nov. gen.”’ 
3 Loe. cit. p. 39. 


CYCADACER. Ly 


particular specimen showed a lamina on either side of the basal 
part of the rachis, having the appearance of several pinne fused 
together laterally, the position of each segment being indicated 
by a strong vein. 

In every classification which is based on artificial characters, 
and which gives us provisional genera, there must necessarily 
be inconsistencies, and in all probability plants possessing no close 
relationship will often be included in the same genus. Among 
fossil ferns this is especially the case; as regards cycads, although 
not perhaps to an equal extent, there are the same difficulties 
to be encountered owing to the isolated and fragmentary nature 
of the specimens on which determinations are based. It may, 
perhaps, be possible to add to the convenience of classification, 
or to minimise the danger of conveying wrong impressions by 
ill-chosen names, by adopting some more admittedly provisional 
classification than is at present employed. An attempt to modify 
our present system, which is too often inadequate and unsatis- 
factory, will be more appropriately undertaken after the Wealden 
and Jurassic genera have been subjected to a detailed treatment. 
For the present, attention may be drawn to some of the obstacles 
in the way of accurate determination of fossil fronds, 

As regards the manner of attachment of pinne to the rachis: 
among recent genera there are some in which the pinne are 
readily detached from the rachis by a well-marked line of articu- 
lation ; e.g. in such forms as Zamia furfuracea, Ait., with broad 
oval pinnee, and other species of the same genus. In species of 
Encephalartos, Ceratozamia, Dioon, etc., there are distinct and 
sharply defined scars left on the axis of the frond on the fall 
of the pinne ; in others, again, the pinne are persistent. Among 
fossil forms, the rachis scars and detached pinne with clearly cut 
bases evidently point to a deciduous habit; but it is often a matter 
of great difficulty to decide definitely as to the existence of such 
a character, and it is quite unsafe to trust to a feature of this 
kind as an essential character in generic classification. It is by 
no means easy in some cases to distinguish the true auriculate 
base of a pinna, from a cordate form produced by the crushing 
and flattening of a thick and leathery segment. Bornemann has 
called attention to this possible source of error, and points to 
the absence of any true auriculate base in the pinne of recent 
fronds. In examining herbaria specimens of some Encephalartos 


C 


18 CYCADACER. 


fronds, such as £. Caffer, Miq., and other species with broad 
stout pinne, one frequently notices that the basal portions of 
the segments have been depressed in such a way as to present 
in surface view the appearance of a distinct auriculate base. 
In some of the examples of Otozamites Klipsteinit (Dunk.) 
var. superba, described in the present volume, this has probably 
been the case; but, thanks to the large number of excellent 
specimens in the Rufford Collection, it is perfectly clear that 
the pinne of this striking plant possessed auriculate bases. 
The absence or presence of a callosity is often a question of 
considerable uncertainty among fossil leaves, and the existence 
of a basal thickening, often none too distinct in the segments of 
recent species, can only be satisfactorily made out in exceedingly 
well-preserved specimens. In some cases there is a distinct 
wrinkling of the coaly surface layer in the position where a 
callosity would naturally occur, and this may no doubt have 
sometimes resulted from a callosity in the living pinna, but in 
others the same appearance may be due to mere bending of the 
frond segments in the process of fossilization. 

It has been shown by more than one writer how easily the 
manner of attachment of the pinne to the rachis may be obscured 
by the frond being seen from its under side. In the case of Dioon 
a view of the upper face of the leaf would lead one to refer it 
to such a genus as Dioonites; but if the lower surface were 
exposed to view Pterophyllum would be the most appropriate 
genus. In a species like IJuacrozamia Denisoni, Moor and Meull., 
in which the pinne are attached along a median line on the 
upper face of the rachis, the same pinne seen from below are 
apparently inserted laterally on the axis, and show no signs of 
decurrent bases. Braun’s figures of Zamites (= Otozamites) brevi- 
folius, Braun,' as seen from above and below, bring out very 
clearly the striking contrast between the two views; the same 
kind of difference is well shown in Feistmantel’s figures of 
Ptilophyllum acutifolium var. maximum, from the Rajmahal Hills 
of India.” 

The comparative breadth of the pinna base is a character which 
varies considerably according to the position of the segment on 


1 Pl. xiii. figs. 138-16. 
2 Feistmantel, Pal. Ind. pt. ii. pl. xl. 


CYCADACER, 19 


the rachis, whether towards the tip or the lower part of the leaf, 
or according to the age of the frond. The terminal pinne are 
often strongly decurrent at the base, whilst the lower segments 
have a uniform width; a young frond of Cycas media, Br., 
shows pinne with no indication of tapering towards the rachis, 
but the older and broader segments are distinctly narrowed. 

Stress is often laid on the form of the pinna apex, whether 
truncate, acute, etc. In the typical form of Pterophyllum the 
pinne have truncated apices, but specimens are occasionally 
referred to this common provisional genus in which the apices of 
the segments are clearly not truncate. Bornemann defines the 
genus as possessing pinnee which may be either straight at the tip 
or obliquely truncate, and this wider definition is probably the 
most satisfactory. In such a specimen as that of Ofozamites 
Géppertianus (Dunk.), figured in Pl. I. Fig. 2, some of the 
pinne are more or less truncate at the tip, and others regularly 
acuminate. In the examples of Zamites Buchianus (Ett.) in the 
British Museum Collection, the variation in the apical terminations 
of the pinne has proved a difficulty, some specimens having 
gradually tapering segments, and others showing obtusely ter- 
minated apices, but the occurrence of some intermediate forms 
throws doubt on the value of such a feature as a leading specific 
characteristic.'_ In dried fronds of Cycas revoluta, Thunb., it is not 
uncommonly found that in many of the pinne the pointed spiny 
apex has been replaced by a rounded termination, with a slight 
median depression at the end of the single vein. As a rule, 
however, the pinne of recent fronds maintain a fairly uniform 
mode of termination in the same species. The venation is not 
always readily made out even in fairly good specimens; the thick 
coriaceous pinne of some recent species, with their indistinct veins, 
prepare us for a similar difficulty im dealing with fossil leaves. 
It is well known that the lower surface of a pinna often shows 
very distinct venation, while the veins on the upper surface are 
quite obscure. In Cycas we have a convenient venation character, 
which is taken as the essential feature of the fossil genus Cycadites; 
but in this case, as we shall see later in describing the genus, 
frequent mistakes have been made in the determination of speci- 
mens, which apparently rest on such a readily recognized character 


HPAL) JOU 


20 CYCADACER. 


as the presence or absence of a midrib. Schenk has pointed out 
Dunker’s error with regard to the supposed Wealden species of 
Cycadites, C. Morrisianus, Dunk., and a careful examination of the 
English material confirms Schenk’s correction. In some recent 
species of Cycas the midrib is by no means obvious on the upper 
surface of the pinne; ¢.g. in a dried specimen of Cycas Cairnsiana 
(Muell.), the upper convex surface of a pinna presents an appear- 
ance suggestive of a few parallel veins, no doubt due to wrinkling, 
rather than of a single midrib. In C. Beddomei, Dyer,’ the 
margins of the pinne are strongly revolute, and a cast of the 
lower surface of a pinna would show too longitudinal ridges 
separated by a distinct groove, the latter being formed by the 
projecting central vein. On the other hand, the tendency to 
a revolute margin in the long, narrow, linear pinne of other 
genera than Cycas, often leads to an appearance which might 
easily be mistaken for a stout midrib in fossil specimens of such 
a leaf. The under surface of the pinne of Encephalartos 
Ghellinckit, Lem, (Pl. XIII. Fig. 3), Zamia angustifolia, Jacq., ete., 
shows a narrow median groove separating the revolute edges of the 
narrow segments, and this same folding might readily give rise to 
a midrib-like character in the segments of fossil fronds. In a few 
exceptional cases there is an anastomosis of the veins in cycadean 
leaves; among fossil fronds Lindley and Hutton established the 
genus Ctenis, for ‘all leaves having the general character of 
Cycadegz, but with veins connected by forks or transverse bars.” * 
As regards living genera, some authors refer to Bowenia and 
Stangeria as having anastomosing veins, but the occurrence of 
anastomosis in the segments of the former genus is denied by 
Engler* and others. The proximity and number of the veins in 
a pinna are characters of no little value in the separation of specific 
forms, but the difficulty of eliminating the effects of fossilization 
and the different appearances presented by the upper and lower 
faces, render it difficult to arrive at any very trustworthy con- 
clusion as to venation characters. In speaking of cycadean venation, 
Bornemann‘ suggests that the characteristic veins of Zama have 


1 Dyer (1). 

2 (A.) Foss. Flor. vol. ii. p. 108. 
3 Engler and Prantl, p. 9. 

4 Loc. cit. p. 39. 


CYCADACER. Ds 


usually been overlooked as a means of identification. The in- 
clination of pinne to the rachis, and their alternate or opposite 
disposition are characters which have been used as the basis of 
specific determination, but such features as these are likely to prove 
misleading unless used with great caution. In one part of a frond 
the pinne may be distinctly opposite, and in another alternate. 
The same kind of variation in the angle of insertion of a segment 
to the rachis, is readily seen in the large fronds of such recent 
species as Ceratozamia mexicana, Brong., Macrozamia Macleay’, Miq., 
and many others; also among fossils in the larger specimens of 
Zamites Buchianus (Ett.), ete. A comparison of the young and 
old fronds of many cycads reveals the same striking difference as 
regards the inclination of the pinne. The open or closely set 
arrangement of pinnz is another misleading character; e.g. in an 
old frond of Encephalartos longifolius, Lehm., the pinnee are for 
the most part in contact with one another, but the young frond 
presents a distinctly open habit, with the pinne much more 
openly arranged. In Ofozamites Goppertianus (Dunk.) there is 
the same difference in this respect between the upper and lower 
portions of the same specimen, e.g. Pl. I. Figs. 1 and 2. 

The form of the epidermal cells is a character of doubtful 
value, and at the same time one which can only be made use of 
under favourable conditions of fossilization. The custom of asso- 
ciating spiny margins with the pinnee of Encephalartos has led an 
American writer to adopt this feature as the leading characteristic 
of his genus Hncephalartopsis.1 Fontaine has founded this new 
genus on some very fragmentary and imperfect pinne with spinous 
margins and anastomosing veins. None of the figured fragments 
afford any clue as to the nature of the pinna base, or as to the 
manner of insertion on the rachis. The material is hopelessly 
inadequate for the institution of a new genus. The fact of the 
fragments possessing anastomosing veins deters Fontaine from 
including them in the recent genus Encephalartos; as it is, he 
prefers to institute a new term, and to consider the species as 
probably a “prototype”? of the recent genus. It is true one is 
accustomed to associate spiny pinne with species of Encephalartos, 
but there are several forms of that genus in which no indication 
of such a character is found; and on the other hand, spiny pinne 


1 Fontaine (A. 2), Potomac Flora, p. 174. 


De) CYCADACER. 


are met with in Dioon edule, Lind., and to a certain extent in 
Zamia Lindeni, etc. Newberry’ has doubtfully referred a small 
portion of a frond from the Rheetic beds of Honduras to the genus 
Encephalartos, but expresses his hesitation as to the true position 
of the specimen by adding a query to the generic name. It is 
suggested by Newberry that the Miocene cycad named by Saporta 
Encephalartos Gorceixianus, does not correspond so closely with any 
living member of the genus as does the Honduras specimen; he 
adds: ‘This correspondence in the form of the pinnules is so 
close that I felt warranted in placing our fossil provisionally in the 
genus Encephalartos. The fructification will of course be necessary 
for a demonstration of generic identity, and has not yet been 
obtained.”” In Lesquereux’ posthumous monograph on the Dakota 
flora, there is a fragment figured and described as a new species, 
under the name Encephalartos eretaceus, Lesq.”; but this is another 
example of what we may regard as the utterly unwarrantable use 
of a recent generic name, and the institution of a new species on 
absolutely insufficient data. It does not seem to have been generally 
recognized that the living species of Lncephalartos present a great 
variety of leaf form, from the long and narrow pinne of such 
species as EZ. Ghellinckii, Lem. (Pl. XIII. Fig. 3), and £. 
eycadifolius, Lehm. (Pl. XIII. Fig. 6), through #. Lehmanna, 
Lehm., etc., to Z£. Caffer, Mig., and £. horridus, Lehm. There 
is a very striking difference between the young and old fronds 
of £. cycadifolius: in the former the pinne are much more oblique 
to the rachis, and have not assumed the stiff and straight character 
which is so pronounced in the latter. Many of the Mesozoic 
cycadean fronds present a striking similarity to LHncephalartos 
leaves, but it would be exceedingly rash to apply the name of 
the recent genus to even the best of these fronds, and still more 
unwise to make use of it for the merest fragments of isolated 
pinne. 

It will be most convenient to consider the Wealden specimens 
referred to the Cycadacee under the headings Frondes and Trunet ; 
and also to describe such seeds and reproductive structures as may 
possibly be included among cycadean fossils. Unfortunately the 
isolated mode of occurrence of leaves, stems, and seeds does not 


1 (1), p. 346, fig. 5. 
2 Lesquereux (A. 3), p. 29, pl. i. fig. 12. 


CYCADITES. 23 


allow, in the great majority of cases, of any certain conclusions 
as to the relation of the detached members one to another. 


FRONDES. 


Cycadites Rémeri, Schenk. 
Cycadites Saporte, sp. nov. 
Dioonites Dunkerianus (Gopp.). 
Dioonites Bronguiarti (Mant.). 
Vilssonia Schaumburgensis (Dunk.). 
Otozamites Klipsteinii (Dunk.). 
Otozamites Klipsteinii (Dunk.), var. superba mihi. 
Otozamites Klipsteinii (Dunk.), var. longifolia mihi. 
Otozamites sp., cf. O. Klipsteinit (Dunk.). 
Otozamites sp., cl. O. Reibeiroanus, Heer. 
Otozamites Goppertianus (Dunk.). 
Zamites Buchianus (Ett.). 
Zamites Oarruthersi, sp. DOV. 
Specimens of doubtful position. 
Anomozamites Lyellianus (Dunk.). 


Genus CYCADITES, Sternberg. 


[Flor. Vorwelt, iv. p. xxxii. 1825.] 


Sternberg proposed this name in 1825 for three fossil plants 
from the Lower Cretaceous of Hor in Scania, and one from Radnitz 
in Bohemia. He defined the genus as follows: ‘ Folia pinnati- 
fida seu pinnata, nervis validis simplicibus e rhachi horizontaliter 
exeuntibus.” 

Sternberg’s species Cycadites Nilssont had been previously 
figured by Nilsson in 1820,' but he left the plant unnamed; 
this species is now included in the genus JVilssonza. Another 
of Sternberg’s species, C. linearis, is no doubt, as Presl first 
suggested,” a fragment of some fossil stem. Cycadites palmatus, 
Sternb., from Radnitz, is probably a fragment of Cordaites, and 
C. samiefolius suggests a coniferous twig. In 1824 Nilsson? 
figured a portion of a leaf from the Quadersandstein of Hor, with 


1 Nilsson (1), pl. iv. fig. 3. 
2 Sternberg (A.), Flor. Vorwelt, fasc. vii. p. 194. 
3 (2), p. 148, pl. ii. dis. figs. 4 and 6. 


24 CYCADITES, 


uninerved and apparently palmately-arranged segments; this he 
described as probably a Filicite. Brongniart! afterwards referred 
Nilsson’s plant to Cycadites, on account of the resemblance of 
the leaf segments to the pinne of the recent genus Cycas. As 
Schenk? has pointed out, Nilsson’s figure in all probability 
represents an Aralia leaf, and the fossil is certainly not a species 
of Cycadites. In Brongniart’s Prodrome*® we have the following 
definition of the genus Cycadites :— 

‘*Feuilles pinnées, a pinnules linéaires, enti¢res adhérentes 
par toute leur base, traversGes par une seule nervure moyenne, 
€paisse ; point de nervures secondaires.”’ 

He regards the single-veined linear pinne as the important 
feature, and in spite of the fact that the first specimen to be 
included under this generic name was incorrectly determined, 
this definition of Cycadites has been generally adhered to. 

Schimper, Saporta, and other authors have, in the main, 
adopted Brongniart’s diagnosis. We may perhaps most con- 
veniently define Cycadites as follows :— 

Frond pinnate, pinne alternate or opposite, linear, lanceolate, 
entire, with a single median vein; attached to the rachis by 
the entire base, the lower margin of which may be slightly 
decurrent on the frond axis, or slightly narrowed towards the 
point of attachment. 

It is better to confine our definition to the frond characters, 
and thus frame it in such a manner that it practically includes 
those fossil fronds which have a cyeadean habit, and resemble 
the recent Cycas in the possession of uninerved segments. In 
several cases Cycadites fronds have been found in close associa- 
tion with characteristic Cycas-like carpellary leaves; but in the 
majority of specimens we have only sterile fronds, and it is 
better, therefore, to have some definition which enables us to 
give such leaves a place in a convenient genus, which does not 
depend upon special characters of fertile leaves. 

The genus Cycadites, as detined by most writers since the 
days of Brongniart, possesses easily recognized characters, and 
ought not to present any very serious difficulty in the way of 


ne (CAzE2) sper 9oe 
2 (A. 1), Fl. foss. Grenz. Keup. p. 158. 
3p. 98. 


bo 
Or 


CYCADITES. 


generic determination. When we come to examine the various 
plant fragments which have been figured as representatives of 
the genus at different geological horizons, it becomes apparent 
that the mere acceptance of a list of Cycadites species as an index 
of the past history of the genus would undoubtedly lead us into 
error. In any case it would be rash to maintain that a record 
of even the most perfectly preserved specimens of the Cycadites 
type of frond, affords an epitome of the geological history of the 
genus Cycas. The occurrence of fossil carpellary leaves very 
similar to, or practically identical with, those of Cycas, lends 
confirmation to the position assigned to many of the Cycadites 
fronds; but as regards other species we can only express the 
opinion that they are parts of a plant which closely resembles 
in habit, and probably in structure, the living genus. It has 
already been pointed out that the pinne of Cycas circinalis, L., 
may occasionally be united laterally and assume a form suggestive, 
in some degree, of Nilssonia or Pterophyllum. No great weight 
can be attached to this single instance of such lateral fusion, but 
it is worth noting as having a possible connection with some 
of the fossil leaf forms which present little resemblance to 
recent fronds. Saporta has called attention to the similarity 
between some Cycadites species and Vilssonia, and one of Heer’s 
species, C. Dicksoni+ from the Cretaceous of Greenland, seems 
to possess pinne which are either in contact with one another, 
or actually united by the margins. 

Berger figured a fragment as Cycadites alatus, Berg.,? and 
compared it with Wilssonia brevis, Brong., the same plant being 
afterwards renamed by Goppert Nilssonia Bergert.§ As regards 
the first record of Cycadites in Paleozoic rocks, it cannot be said 
that there is any very decided evidence of the occurrence of this 
genus, but Goppert’s C. taxodinus is by no means such a doubtful 
representative of the genus as several of the species described 
from newer beds. Gippert’s C. gyrosus may perhaps be a portion 
of a young frond with its pinne circinately rolled, but it is not 
enough to establish the existence of Cycadites in Carboniferous 
times. Sterzel has recently figured an imperfectly preserved 


1 Heer (A. 3.), Fl. foss. Arct. vol. iii. pt. ii. p. 99, pls. xxvii. and xxviii. 
2 Berger, p. 22, pl. iii. figs. 5 and 6. 
8 (1), p. 141. 


26 CYCADITES. 


impression from the Middle Rothliegende of Possendorf, Saxony,' 
which he speaks of as ‘‘ Cycadites ? or Walchia sp.,” but does 
not consider it sufficiently distinct to allow of accurate identi- 
fication. The figure entirely justifies Sterzel’s doubtful attitude. 
In the Mesozoic beds Cycadites fronds become more abundant; 
a list of most of the species has been given by Solms-Laubach.? 

A few of the so-called Cycadites species call for special mention. 
The Jurassic specimens described by Leckenby, from Cloughton, 
as Cycadites zamioides* are probably, as Nathorst suggests, 
fragments of a conifer. The type specimen of Leckenby’s 
species in the Woodwardian Museum, Cambridge, bears a label 
on which Nathorst has written, ‘‘A conifer of the genus 
Palissya”; and Richards,{ who examined the specimens a few 
years ago, adopts this view. In the case of some small indistinct 
impressions, it is often very difficult to decide between a twig of 
a conifer with its spirally arranged leaves extended in one 
plane, and a small cycadean frond with its uninerved pinne 
inserted on the two sides of a rachis. A branch of Cephalotarus 
Fortunet, Hook, might very easily be mistaken for Cycadites if 
found in a fossil state with the details of structure imperfectly 
preserved. Heer has described several species of Cycadites from 
Arctic localities, but the figures do not inspire confidence in his 
determinations. Cycadites Dicksoni® may very probably be a true 
Cycadites; C. sibiricus, Heer,’ and C. gramineus, Heer,’ from 
the Jurassic rocks of Siberia, are both founded on the merest 
fragments of single pinne, and cannot be taken as trustworthy 
records. The institution of species on such minute fragments 
as the figures represent, is to be greatly deplored; the result 
can only be either to mislead those who are willing to accept 
all fossil species described by well-known authors, or to deter 
the more sceptical from attaching any importance to fossil plant 


1 Sterzel, p. 140, pl. xii. fig. 12. 

* (A.), Fossil Botany, p. 86. 

3 Leckenby (A.), Quart. Journ. Geol. Soc. vol. xx. 1864, p. 77, pl. viii. fig. 1. 

eal) sapence 

5 Heer, Fl. foss. Arct. vol. iii. pt. ii.. p. 97, pls. xxvii. and xxviii.; and 
vol. vi. pl. xiv. fig. 10. 

§ Tbid. vol. v. pt. ii. p. 16, pl. iv. fig. 1. 

7 Ibid. fig. 2, ete. 


CYCADITES. 27 


determinations which do not rest on other characters than those 
of external form. Another species from the same beds, C. ? 
planicosta, Heer,! is founded on imperfect pinne, but in this 
case Heer definitely admits the doubtful value of the name. 
The specimen described by the same authority from the Tertiary 
beds of Schaffhausen as Cycadites Escheri* is very likely a 
cycadean stem, but the genus Cycadites has been restricted to 
fossil fronds, and Heer’s stem fragment should be referred to 
some other genus, in order to avoid the confusion likely to arise 
from using the name in a more comprehensive sense. Dawson 
has described some fronds from the Middle Cretaceous of the 
Rocky Mountains, which he names Cycadites Unjuga,* and 
compares with Heer’s Cycadites Dicksoni. The two figures of 
the Canadian specimens do not appear to agree as regards the 
Cycadites form of pinna: in Fig. 2 each pinna appears to have 
several parallel veins, and the general habit seems different 
from that in Fig. 2b; if Fig. 2 be an accurate representation 
of the specimen, and the vein-like lines are not the draughts- 
man’s shading, it could hardly be accepted as a true Cycadites. 
Feistmantel figures a fragment under the name of Cycadites 
constrictus, Feist., and speaks of a midrib in the basally con- 
stricted pinne ; the figure does not show any distinct midrib, 
and leaves one in doubt as to the wisdom of choosing the genus 
Cycadites. 


1.—Cycadites Romeri, Schenk. 
[Fig. 1.] 


1871. Cycadites Rimeri, Schenk, Paleontographica, vol. xix. p. 229, pl. xxxii. 
figs. 1, la. 
1874. “Cycadites Rémeri, Schimper, Trait. pal. vég. vol. iii. p. 552. 


Type. Portion of a frond. Berlin Museum. 
The following definition is given by Schenk for this species :—° 


1 Heer, FI. foss. Arct. vol. iv. pt. ii. pl. iv. fig. 16. 

2 Heer (A. 1), Fl. Tert. Helvet. p. 46, pl. xv. 

8 Dawson (1), p. 20, pl. i. fig. 2. 

4 Feistmantel, Gond. Flor. vol. i. pt. iv. p. 25, pl. vii. fig. 10. 
5 Schenk (A. 2), Paleontographica, vol. xix. p. 229. 


28 CYCADITES. 


‘“‘Folia petiolata pinnata, petiolus validus, segmenta linearia 
patentia integra alterna basi dilatata breviter decurrente sessilia, 
83 cent. longa, 2 mm. lata, uninervia, nervus medianus validus.” 

He refers to the recent species Cycas Siamensis, Miq., as most 
nearly allied to the fossil frond. This is the only example of 
Cycadites recognized by Schenk among the North German 
Wealden plants; the specimens referred by Dunker to that 
genus being without the characteristic single vein in the pinne. 
The English specimens, for which the name Cycadites Saportea, 
sp. noy., is proposed, differ from the present species in their 
narrower and more approximately disposed pinne, inclined 
almost at right angles to the rachis. The scanty material in 
the Rufford Collection referred to Cycadites Rémeri enables us, 
however, to add one or two points to the original diagnosis by 
Schenk :— 

Frond pinnate, rachis strong, pinne linear and narrow, 
obliquely and laterally attracted to the rachis, entire, alternate, 
with slightly broadened and somewhat decurrent base, single 
median vein, apices acuminate and terminating in a sharp point. 


Fie. 1.—Cycadites Rémeri, Schenk (Y. 2738). Slightly enlarged. 


V. 2788. Fig. 1. 

An imperfect specimen, showing several partially preserved 
pinne, the largest of which has a length of 83 cm., as in 
Schenk’s specimen, and a breadth of 3mm. 

In the type specimen of C. Rémert the pinne are broken at 
the apices, but in the English example the sharply acuminate 
tips are clearly preserved, and correspond closely with those 
in (C. Saporte, Cyeas revoluta, Thunb., ete. Each pinna is 
traversed by a median groove, which must probably be regarded 
as the midrib seen from the under side, but it should be noticed 
that there are in some of the pinne slight variations in the 
breadth of the groove, and it occasionally departs somewhat 


CYCADITES. 29 


from a strictly median course. The appearance, indeed, is such 
as to suggest a folding over of the pinne margins. It has 
already been pointed out in the introductory remarks on fossil 
eycadean fronds, how the pinne of such recent species as 
Encephalartos Ghellinckii, Lem., may become folded over until 
a narrow median groove is left in the middle of the lower 
surface of the segment, representing the line of separation of 
the recurved edges (Pl. XIII. Fig. 3). On the other hand, we 
may have a similar curling over in the pinne of a true Cycas; 
but in the present specimen the narrow line is for the most part 
perfectly median and of uniform breadth, and cannot well be 
attributed to any other cause than the presence of a central vein. 
At one corner of the specimen there are three pinne, which 
clearly demonstrate a folding over of the margins, but this is 
in itself no proof of the absence of a single vein. These pinna 
fragments are in oblique contact with what appears to be a 
portion of the rachis, and if we may regard the two as 
organically connected, the segments exhibit the same characters 
as regards the form and attachment of the base as Schenk has 
described in the German examples. LEcclesbourne. ufford Coll. 


2.—Cycadites Saporte, sp. nov. 
fe i Bie, 7s Pl VE. Bie, 64 Pie VL. Bie. 2.) 


Type. Large and well-preserved fronds. British Museum. 

The difficulty of recognizing the essential character of Cycadites 
in the pinne of fossil fronds has made itself felt in no small 
degree in dealing with the present series of specimens. The 
figures and descriptions given by Romer and Dunker of the 
Cycadites-like leaves have to be viewed in the light of Schenk’s 
more recent statements, based on an examination of Dunker’s 
type specimens. If we leave out of consideration those portions 
of cycadean fronds which are figured by Dunker as Cycadites 


1 Schenk (A. 2), Paleeontographica, vol. xix. p. 233. 


30 CYCADITES. 


Morrisianus, Dunk.,! and which present a close agreement in habit 
with the English specimens, we have only C. Rémeri, Schenk, 
among Wealden fronds with which to compare C. Saporte. The 
differences between C. dmeri and the present species are, I 
believe, too well marked to admit of a single specific designation. 

In view of the exceptionally large size of the Ecclesbourne 
fronds, and the satisfactory manner of preservation, it is better to 
adopt a new specific term, and I have ventured to identify the 
name of the Marquis of Saporta with this new form of Lower 
Cretaceous cycad. 

Since the above was written the Marquis of Saporta’s promised 
Monograph on the Flore fossile du Portugal has been published. 
A review of this valuable contribution to Mesozoic paleobotany 
is given in the latter part of the present volume. Among the 
very few remains of cycadean fronds described by Saporta, one 
form of Cycadites appears under the name of C. tenuisectus,? Sap., 
and the figures of the frond fragments show a very distinct resem- 
blance to the English specimens which I have referred to the new 
species, C. Saporte. Possibly the Portuguese and British plants 
should be placed in one species, but for the present at least, 
there are certain differences to be noticed which hardly justify 
this adoption of Saporta’s specific name. In the English fronds 
the pinnz are somewhat stouter, the tips more sharply acuminate, 
and the general habit of the leaf appears to be rather stiffer than 
in C. tenuisectus. 

Frond pinnate, linear, of uniform breadth; rachis broad and 
flattened, marked with obliquely placed lines, terminating proxi- 
mally in a broadened and swollen base. Pinne of uniform breadth, 
alternate or subopposite, attached to the upper surface of the 
rachis, and inserted at right angles or slightly oblique to 
the frond axis, the bases of the two rows of the pinne almost 
in contact; average length of the pinne 6-7 cm., and 1-15mm. 
in breadth; bases slightly broadened and contiguous, apices of 
the long linear pinne terminating in a sharp point; single median 
vein in each segment. 

In 1839 Romer instituted the species Cycadites Brongniarti 


1 Dunker (A. 2), Wealdenbildung, p. 16, pl. vi. fig. 1. 
2 (1), p. 171, pl. xxxii. figs. 1-4 and 6. 


CYCADITES. 31 


for a specimen from the North German Wealden beds, and thus 
defined it': ‘‘C. foliis pinnatis sublinearibus, pinnis numerosis 
linearibus approximatis apice obtusiusculis medio costatis basi sub 
dilatatis.”’ 

He speaks of the pinne as possessing a strong midrib, and his 
figure shows this character very clearly. Dunker has refigured 
Romer’s original specimen, and here again the pinne appear 
to have a distinct median vein; he points out that Mantell’s 
Cycadites Brongniarti* should be placed in the genus Nilssonia, as 
it does not conform to the accepted definition of Cycadites. This 
Tilgate fossil* is now referred to as Dioonites Brongniarti (Mant.). 

In 1852 Ettingshausen* obtained a portion of a cycadean 
frond from near Teschen, in Silesia, and referred it to Romer’s 
species, but at the same time expressing the opinion that it 
represented a form intermediate between C. Brongniarti, Rom., 
and C. Morrisianus, Dunk. ‘This is certainly not the same species 
as Romer’s type, and should, as Schenk suggests, be placed in 
another species; he speaks of it as C. Heerti, Schenk, and 
expresses the opinion that possibly C. Brongniarti, Rom., may 
be simply a partially developed frond of C. Dorrisianus.’ Sub- 
sequently the same author includes both C. Brongniartc and 
C. Morrisianus as synonyms of Dioonites Dunkerianus (Gopp.). 
He states that the type specimen of C. Morrisianus, Dunk., shows 
no indication of a midrib, and must therefore be referred to Ptero- 
phyllum or Dioonites instead of to Cycadites. There is the same 
absence of a median vein, according to Schenk, in the segments 
of C. Brongniarti, Rém., and this must, therefore, be also excluded 
from the genus Cycadites.6 It is not quite clear if Schenk is here 
speaking of Rémer’s original specimen; if he refers to the figured 
specimen as it appears in the illustrations of Romer and Dunker, 
the figures are certainly at variance with Schenk’s description. 
There is the same apparent contradiction between figure and 


1 Romer, F. A. (A.), Verstein. Ool. Geb. p. 9, pl. xvii. fig. 1. 

2 Dunker, Joc. cit. p. 16, pl. ii. fig. 4. 

8 Mantell (A. 4), Geol. S.E. England, p. 238. 

4 (A. 4), Abh. k.-k. geol. Reichs. vol. i. Abth. iii. No. 2, 1852, p. 20, 
ol, a, ieee 

5 (A. 3), Paleontographica, vol. xix. p. 7. 

6 (A. 2), p. 233. 


32 CYCADITES. 


diagnosis in Dvoonites abietinus' (Godpp.), as represented in 
pl. xxxvil. fig. 1 of Schenk’s. monograph. Schimper retains 
Romer’s species, and unites with it Pterophyllum Dunkerianum, 
Gopp., as figured by Dunker,? but in this case the figure shows 
very clearly that the venation is not of the Cycadites type. In 
the face of Schenk’s statements, we cannot, then, accept any of 
Dunker’s figures of what he describes as species of Cycadites 
as really examples of that genus; and, as Saporta*® points out, the 
only representatives of Cycadites so far known for beds of approxi- 
mately Wealden age are C. Réimert, Schenk, and C. Heerii, 
Schenk. In the English specimens referred to the new species 
C. Saporte the preservation is fortunately good, and leaves no 
doubt as to the existence of a true midrib in the pinne. 
C. Roimeri agrees to some extent with this species, but differs 
in its broader pinne and their disposition on the frond axis. The 
specimens of C. Saporte are unusually large, and hence enable 
us to obtain a good idea as to the general habit of the frond; 
if it were not for this fact one might be inclined to include them 
under Schenk’s species. The plant described by Braun from the 
Jurassic sandstones of Steinstedt as C. rectangularis* differs in 
its shorter and broader pinne, and in the fact that they are more 
distinctly at right angles to the rachis. Some of Saporta’s figures 
of what he regards as C. rectangularis, are much more like 
C. Saporte than the type specimen figured by Braun; cf. especi- 
ally pl. xiii. figs. 1 and 3 of the Flore Jurassique. Saporta 
includes C. pectinatus, Berg., as a synonym of Braun’s species, 
and adopts the term rectangularis in preference to the older name 
pectinatus, because of the use of the latter term by Lindley and 
Hutton in connection with the genus Zamites;° perhaps hardly 
a sufficiently sound argument to overrule the priority of Berger’s 
term. 

Berger’s small fragment as figured in his pl. iii. fig. 4, 


1 (A. 2), p. 284. Géoppert, and not Miquel, appears to be author of the two 
specific names abietinus and Dunkerianus; Schenk refers both these species 
to Miquel. 

2 Schimper (A.), Trait. pal. vég. vol. ii. p. 180. 

S Loc. cit. p. 72. 

4 Braun (A.), Paleontographica, vol. ix. p. 56, pl. xiv. fig. 7. 

5 Saporta, loc. cit. p. 70. 


CYCADITES. 33 


evidently belongs to a plant of very similar habit to that of 
C. Saporte; it differs mainly in the greater breadth of the 
pinne, so far at least as it is possible to judge from Berger’s 
figure. 

Another species which bears a still closer resemblance to 
C. Saporte, is C. Rajmahalensis, Old., described by Oldham, 
from India! The figures and description of this plant are in 
close agreement with the Ecclesbourne Wealden species ; the seg- 
ments of the Indian frond seem to be rather more closely arranged 
and somewhat shorter than in the English form. It would, 
however, be somewhat unwise to refer the Wealden specimens 
to Oldham’s species, considering the geological age of the two 
plants, and the less perfect preservation of Cycadites Rajmahalensis. 
In this, as in many other cases of fossil plants, we have to speak 
cautiously as to the relationship of individual members of different 
floras, and must trust rather to the comparative study of the floras 
as a whole, than to the apparent identity of isolated elements. 

Trautschold’s specimen of Cycadites acinaciformis, Traut.,? is 
similar to the present species, but probably not identical with it. 
Schenk suggests that the Russian species is probably identical with 
Pecopteris decipiens, Traut., and must be placed with the ferns. 
It is difficult to speak with much confidence as to the nature 
of the specimen figured as C. acinaciformis, but it certainly bears 
a strong likeness to the cycadean genus.’ 


V. 2777. Pl. VIII. Fig. 2 (+ natural size). 

This exceptionally fine specimen shows one frond 60cm. in 
length, and a second 38cm. long, the latter being inclined to 
the former in such a way as to suggest but little displacement 
from their original position of growth on the parent stem. The 
larger frond, as represented on a small scale in the photograph, 
shows a striking uniformity in the length and breadth of the 
numerous closely set pinne. In several of the segments the 
sharp apices are clearly preserved. Judging by other specimens, 
in which the pinne are longer and more nearly at right angles 


1 Oldham and Morris (A.), Foss. Fl. Gond. p. 15, pl. viii. 

* Trautschold (A. 3), Nouv. Mém. Soc. Nat. Moscou, vol. xiii. 1876, p. 34, 
plese fies 1: 

° Schenk (A. 2), Palwontographica, vol. xix. p. 261. 


34 CYCADITES. 


to the rachis, it is probable that this large example may 
represent a frond not quite fully developed. The stout rachis, 
about 1:4 cm. in breadth, and especially that of the smaller 
frond, shows numerous obliquely running longitudinal lines. 
The contiguous and slightly broadened bases of the pinne are 
very distinct on portions of the larger frond. In some places 
the segments, adhering together by their contiguous bases, have 
been torn en masse from the axis of the leaf. 

The stout and distinct median vein is well marked throughout. 
Near Hastings. Rufford Coll. 


V. 2797. Pl. VI. Figs 5 and 5a. 

Frond 13:5 cm. in length; rachis lem. broad. The long and 
contiguous pinne are attached at right angles to one edge of 
the flattened axes. Midrib distinctly preserved, as in Fig. 5a; 
also the sharply acuminate tips of the segments. Longest pinna 
llcm. ‘The general appearance of this specimen is indicative 


of an older frond than V. 2777. Ecclesbourne. Rufford Coll. 


V. 2124. Pl. III. Fig. 7. 

24 cm. long. At the two extreme ends only one row of 
pinne has been preserved, and the impression of the broad rachis 
is shown on the surface of the rock. In other places the two 
rows of pinne are almost in contact, as in the portion repre- | 
sented in Pl. IV. Fig. 5, and there is very little of the rachis 
visible between the bases of the two sets of pinne. The 
arrangement and general appearance of the segments bear a 
marked resemblance to Dioonites Dunkerianus (Gopp.). Ececles- 
bourne. Rufford Coll. 


V. 1069. A more terminal portion of a frond. Pinne smaller 
and more obliquely inclined to the rachis than in most of the 
other specimens. Cy. the terminal portion with the specimen 
of Dioonites Dunkerianus (V. 2823) figured in Pl. Il. Fig. 8. 
Ecclesbourne. Presented by P. Rufford, Esq., 1885. 


V. 2124. Narrow pinne attached to one side of the flat rachis. 


V. 2124. Several portions of fronds. In one there appears 
to be the broad and thick basal termination of the petiole fairly 


DIOONITES. 30 


clearly preserved. The midrib and pointed apex well seen in 
several of the pinne. LEcclesbourne. Rufford Coll. 


V. 2924. 25cm. long. A single row of pinne attached to 
one margin of the broad flat rachis; midrib distinct. Eccles- 
bourne. Rufford Coll. 


Genus DIOONITES, Miquel. 


[Tijdsch Wis. Nat. Wet. vol. iv. 1851, p. 205.] 


In dealing with such fronds as those figured by Dunker as 
Cycadites Morrisianus, Dunk., and C. Brongniarti, Rom., and 
afterwards described by Schenk as species of Pterophyllum or 
Dioonites, we have to face the difficulty of deciding upon the most 
suitable generic term. The fronds in the Rufford Collection show 
well-marked characters, and leave no doubt as to the form and 
manner of attachment of the pinne; we have long, narrow, linear 
and parallel-veined segments, with acutely pointed tips attached 
by broad and non-auriculate bases to the upper surface of the 
rachis. Must these be included in Pterophyllum or Dioonites, or 
do the generally received definitions of these genera not admit 
of the application of either name to the Wealden fronds? Let 
us briefly summarize some of the various definitions of these 
genera, and note how far they coincide with the characteristic 
features of the present series of specimens. Pterophyllum was 
defined by Brongniart in 1828' as a genus characterized by— 
‘‘Feuilles pinnées, 2 pinnules d’une largeur a peu prés égale, 
s'ins¢rant sur le pétiole par toute la largeur de leur base, tronquées 
au sommet; nervures fines, égales, simple, peu marquées, toutes 
paralléles.”” He speaks of the truncate apices of the pinne as 
an essential character, but does not insist on a lateral or surface 
insertion on the frond axis. Pterophyllum Jaegeri, Brong., is spoken 
of as one of the species of this genus, and in this instance the 
attachment of the segments is apparently lateral. In the Zubleau? 


1 (A. 2), Prodrome, p. 95. 
2 (A. 4), p. 68. 


36 DIOONITES. 


Brongniart points out that subsequent writers have applied his 
generic name to plants which do not conform to the original 
definition. He considers the essential characters to be (i.) a slight 
union of the bases of the pinne; (ii.) the quadrilateral, oblong, 
or linear form of the segments; (iii.) truncately terminated 
segments; and (iv.) the presence of fine parallel veins not convergent 
at the apex. Morris! speaks of Pterophyllum as including plants 
with pinnate fronds and sublinear pinne, inserted by the whole 
base, with the apices truncate or sometimes acute, etc. Miquel? 
keeps closely to Brongniart’s original definition. Goppert* adopts 
a wider definition, and includes in this genus plants with obtusely 
and acutely terminated pinne, ete. Bornemann,*‘ in 1856, defined 
the genus as follows: ‘Frond pinnate or deeply pinnatisect, 
pinne approximate, and with the whole base attached to the 
rachis, short, broad, quadrate, or elongate, straight at the tip 
or obliquely truncate, horizontal or oblique to the rachis; veins 
parallel.” 

Leckenby® assigns the name Pterophyllum to the species 
P. medianum, Leck., with its Milssonia-like lamina, which is 
apparently not attached to the side of the frond axis. Schenk, 
in his Fossele Flora der Grenzschichten® . . . , adopts a very com- 
prehensive definition, and defines the pinne of Pterophyllum as 
distichous, elongate, or adherent, narrow or broad, apex acute or 
truncate; but he says nothing as to the manner of attachment 
to the rachis. He includes P. mmerustans, Gopp., and P. Braunia, 
Gopp., in the same genus. Heer’ prefers the genus Zamites, used 
in an unusually wide sense, for such fronds as his Z. borealis, 
Heer, Z. acutipennis, Heer, etc., which resemble in general form 
the leaves of the Wealden species originally described by Dunker 
as Cycadites Morrisianus. Schimper separates the fronds with 
irregularly pinnatifid leaves from the true Pterophyllum type, and 
institutes for their reception the genus Anomozamites.® In the 


4 p. 58. 

5 Loe. cit. p. 77, pl. viii. fig. 2. 

6 (A. 1), Fl. foss. Grenz. Keup. Lias, p. 163. 
7 Fl. foss. Arct. vol. iii. p. 66, pl. xv. 

8 Trait. pal. vég. vol. ii. p. 140. 


DIOONITES. 37 


genus Pterophyllum he includes fronds with pinne vertically 
attached to the side of the rachis, and having truncate apices. 
His genus Ctenophyllum,' which includes certain forms often 
referred to Pterophyllum, is defined as follows: ‘‘ Folia linealia, 
gracilia; foliolis lateri rachis superiori oblique adfixis, sepius 
oppositis, linealibus, obtusis, basi retro folium infrapositum de- 
fleuntibus, coriaceis, tenuiter et parallele nervosis.”’ 

Saporta departs somewhat from the definitions given by other 
authors,” and restricts Pterophyllum to fronds with pinne attached 
to the side of the rachis, and which are distinct one from another, 
not fused laterally at the base, and having truncate apices. 
Feistmantel,? on the other hand, in speaking of Zamites proximus, 
Feist., points to the separate pinne, which are not connected 
at the base, as a feature inconsistent with the inclusion of the 
plant in the genus Pterophyllum. Nathorst‘ has described 
certain plants from Bjuf as possibly species of Pterophyllum ; 
but to express the absence of perfectly satisfactory evidence, he 
prefixes a query to the generic name. More recently, this 
author has called attention to the lateral insertion of the pinne 
as an essential character of Pterophyllum, and a convenient 
distinguishing feature from MWilssonia.® In Zittel’s Handbuch,® 
the lateral attachment of the pinne, which may or may not be 
distinct at the base, and their rounded or truncate apices are given 
as important generic marks. It is suggested that possibly such 
a frond as Pterophyllum Dunkerianum, Gopp., ought not to be 
included in the genus Pterophyllum, because of the insertion of 
the segments on the upper surface of the frond axis. Solms- 
Laubach’ refers to P. Jaegert, Brong., as an example of one 
form of Pterophyllum frond, and in another place® calls attention 
to the ilssonia-like form of some species of the same genus, 
which agree with other examples of Pterophyllum in the lateral 
insertion of the leaf lamina. 


1 Trait. pal. vég. vol. ii. p. 127. 

2 Loc. cit. p. 43. 

3 Foss. Fl. Gond. vol. i. ser. ii. 2, p. 115. 

4 (A. 1), pt. ii. pp. 69-72. 

5 (A. 3), Denkschr. k. Ak. Wiss. math.-nat. Cl. vol. lvii. p. 6. 
6 (A.), p. 224. 

7 Fossil Botany, p. 88. 

8 Ibid. p. 139. 


38 DIOONITES. 


On the whole, then, the characters generally insisted on seem to 
be the lateral attachment of the pinne to the rachis, and by 
many, but by no means all authors, the truncately terminated 
segments. The confusion which has arisen from constant altera- 
tions by various writers, and from the not uncommon practice 
of including certain fronds in a particular genus, in spite of 
obvious discrepancies between the specimens and the generic 
diagnosis, is sufficiently obvious if we glance at some of the 
better known Pterophyllum species as figured by different authors. 
We have such forms as P. inconstans, Gopp., P. Dunkerianum, 
Gipp., P. Jaegeri, Brong., P. Braunit, Gopp., etc., included in the 
same genus. It is true that in examining fossil fronds we are 
often unable to decide as to the actual manner of attachment of 
the pinnz, and are thus driven to leave the specimen as doubtful, 
or to decide as best we may in the face of difficulties inseparable 
from the determination of isolated leaf fragments. We cannot 
always be sure whether we have the frond preserved with its 
lower or upper side uppermost. It is, however, clear that we 
cannot consistently make use of Brongniart’s genus for such 
specimens as those before us. 

The genus Dioonites of Miquel has been adopted by some 
authors for these narrowly segmented Wealden fronds. This 
again is a generic name which has been made to do duty for 
forms of leaves, which it is difficult to regard as correctly included 
in the same genus, even if the genus be admittedly a provisional 
and artificial one. Miquel is responsible for the proposal of this 
name, and for the following definition:1 ‘Frondes pinnate, 
rigide, crasse. Foliola densa patentissima suprema nune sub- 
imbricata, lanceolata, vel lineari-lanceolata, recta vel subfalcata, 
acuta vel acutiuscula, basi tota latitudine inserta, inferne 
retrorsum subdecurrentia, nervis cum margine parallelis equa- 
libus subtus distinctioribus (cum  sulculis stomatiferis alter- 
nantibus).” He included under this name- several species 
previously described as examples of Pterophyllum and other 
genera. Bornemann adopts Miquel’s genus and extends its use 
to some additional species, but does not make any important 
alteration in the original diagnosis. Schimper retains the term 


(2) spade 


DIOONITES. 39 


Dioonites, and gives the essential characters as follows:' ‘‘ Folia 
pinnata, pinnis pro more angustis, lanceolatis, acutis, obliquis, 
tota latitudine insertis, basique leniter pro- et decurrentibus, 
nervis parallelis.” In Zittel’s Handbuch*® the genus is quoted, 
and Pterophyllum Buchianum, Ett., and P. Brongnarti, Schenk, 
are given as two typical species. The former of these has since 
been transferred by Nathorst to a new genus, Zamiophyllum,* on 
the ground that the pinne are slightly narrowed towards the 
base. Saporta repeats the character of Dioonites* as given by 
previous writers, and figures D. Brongniartt as a typical example; 
but the species referred to by this writer at the end of his 
definition as the typical form of the genus is D. Kurrii, Schimp. 
In Fontaine’s Potomac Flora we find numerous forms included 
under Miquel’s genus, but it must be noted that this author, 
while giving what he refers to as Schimper’s definition of the 
genus, speaks of the pinne as ‘‘sometimes expanded at base so 
as to extend up and down the rachis.” ° ‘This is an important 
alteration, as Schimper describes the pinne as distinctly 
decurrent, and it is this characteristic which is repeated by the 
majority of writers as one of the essential generic features. In 
his definition of Dioonites Buchianus (Ett.) Fontaine refers to 
the pinnez as slightly narrowed at the base, but does not regard 
this character as opposed to the adoption of Miquel’s genus. The 
attachment of the pinne by the whole of a more or less 
decurrent base appears to be the chief characteristic generally 
insisted on. In several definitions of Droonttes no mention is 
made of the place of attachment of the frond segments, whether 
on the surface or sides of the rachis; in several of the figured 
specimens referred to this genus the pinne are inserted laterally. 
Some authors have emphasized the fact that the segments must 
be attached to the upper surface of the rachis, as in D. Brongniarit. 
This position of the pinne affords one point of difference from 
Pterophyllum, and in the decurrent and separate leaves we have 
other features characteristic of Doonites. Nathorst, in discussing 


1 (A.), Trait. pal. vég. vol. ii. p. 128. 
2p. 223. 

3 (A. 3), p. 46. 

4 (A. 2), vol. ii. p. 44. 

5 


40 DIOONITES. 


the generic characters of the plant, to which he applies the name 
Zamiophyllum Buchianum (Ett.), refers to Miquel’s genus Dioonites 
as characterized by the attachment of the pinne to the upper 
surface of the rachis, and by the insertion of the segments almost 
at right angles to the axis; he says nothing as to the decurrent 
bases of the pinne. If we accept this definition, and depart 
from the usually accepted feature of a decurrent pinna base, we 
may well include the Wealden plants under this genus. It is 
certainly not an easy matter to draw a definite line between 
pinne attached to the rachis by the entire base, which is not 
decurrent, and those which are similarly attached, but with their 
bases more or less decurrent. In the English examples of the 
species D. Dunkerianus (Gopp), the pinne towards the upper 
end of the frond are distinctly decurrent, but those occupying 
a lower position cannot be described as possessing decurrent bases. 
Cf. Pl. II. Fig. 3, and Pl. IIT. Fig. 6. There are two other 
genera to which reference should be made, which to a certain 
extent agree in their definitions with such fronds as D. Dunkert- 
anus, etc., viz. Ctenophyllum and Ptilophyilum. The former 
genus was instituted by Schimper! to include certain forms of 
fronds which do not in all essentials comply with the definitions 
of Otozamites on the one hand, and Dvioonites on the other, 
Pterophyllum pecten, L. and H., being taken as the type species. 
The author of the genus afterwards somewhat modified his original 
diagnosis, and pointed out that Pterophyllum Braunianum, Gopp., 
had been erroneously described as a species of Ctenophyllum. The 
genus Ptilophyllum, proposed by Morris in 1840? for certain Indian 
fronds, can with difficulty be distinguished from Ctenophyllum. 
It is thus defined :— . 

‘Fronds pinnate; pinne linear, closely approximated, more or 
less elongate; base variable in form, oblique, round, imbricate, 
sometimes auricled in the upper and sometimes in the lower part. 
Veins slender, equal, parallel.” > 

Goppert long ago expressed the opinion that Morris’ term 
was a needless addition to the list of cycadean genera* It 


1 Toe. cit. vol. il. p. 148. 
2 Morris (2), p. 21. 

3 Morris (1), p. 116. 

au (1) apswlelige 


DIOONITES. 41 


has been found useful by several writers as a convenient name 
to apply to Indian fronds, but as at present used it does not 
appear to be wholly satisfactory. The genus Péilophyllum seems 
to haye been almost confined to Asiatic fronds, and the locality 
of a specimen has probably had too great a share in the 
selection of Ptilophyllum in preference to Ctenophyllum as the 
most suitable name. Nathorst! figures and describes a leaf 
fragment from Japan as Péilophyllum cf. cutchense, Morr., but 
it would seem practically impossible to separate such a form as 
this from some English Jurassic fronds usually placed in the 
genus Ctenophyllum. 

On the whole perhaps the better course is to retain, at least 
for the present, the name Dioonites as the most suitable generic 
designation for the Wealden species D. Dunkerianus (Gopp.). 
We must slightly modify the definition of the genus, and no 
longer insist on the decurrent pinna base as an essential charac- 
teristic. The implied relationship to the recent Dioon is the 
least satisfactory feature of Dioonttes, but possibly we shall be 
able, on a future occasion, to suggest some further alteration in 
the existing nomenclature of fossil cycadean fronds. We may 
define this genus, using the term Dvoonites in a wide and 
provisional sense, as follows :— 

Frond pinnate, pinne at right angles, or more or less obliquely 
inclined to the rachis, attached to the upper surface of the frond 
axis, bases separate, may or may not be decurrent, not narrowed 
towards the point of attachment, apices acuminate, straight or 
slightly truncate, veins parallel. 

In dealing with Dvioonites, as with many other genera, we 
may easily fall into the error of excluding or including certain 
forms owing to our imperfect knowledge as to the manner of 
attachment of the pinne; but it is obviously impossible to 
devise a perfectly satisfactory system, so long as we are limited 
by the exigencies of fossilization and the imperfection of the 
frond fragments. 


1 Nathorst (A. 3), p. 52, pl. iv. fig. 8. 


42 DIOONITES. 


Dioonites Dunkerianus (Géppert). 
[Pl. IL. Fig: 35 Pl. II. Fig. 6.] 


1843.  Nilssonia pecten, Dunker, Progr. p. 7. 
Pterophyllum Dunkerianum, Goppert, Foss. Cycad. p. 52. 
1846. Pterophyllum Dunkerianum, Dunker, Wealdenbildung, p. 14, pl. ii. 
fig. 3, pl. vi. fig. 4. 
Cycadites Morrisianus, Dunker, ibid. p. 16, pl. vii. fig. 1. 
? Pterophyllum abietinum, Dunker, ibid. p. 15, pl. vii. fig. 2. 
1848. Pterophyllum Dunkerianum, Bronn, Index pal. nomencl. p. 1055. 
Cycadites Morrisianus, Bronn, ibid. p. 371. 
1849. Zamites Dunkerianus, Brongniart, Tableau, p. 107. 
Cycadites Morrisianus, Brongniart, ibid. p. 107. 
1850. Pterophyllum Dunkerianum, Unger, Gen. spec. plant. foss. p. 290. 
Cycadites Morrisianus, Unger, ibid. p. 280. 
1851. Dioonites Dunkerianus, Miquel, Rangschik. foss. Cycad. p. 212. 
? Dioonites abietinus, Miquel, ibid. p. 206. 
1852. Pterophyllum Dunkerianum, Ettingshausen, Abh. k.—k. geol. Reichs. 
vol. i. Abth. iii. No. 2, p. 20. 
1856. Dioonites Dunkerianus, Bornemann, Organ. Rest. Lettenkohl. p. 56. 
Cycadites Morrisianus, Bornemann, 7bid. p. 51. 
1869. Dioonites Dunkerianus, Schimper, Trait. pal. vég. vol. u. p. 150. 
Cycadites Morrisianus, Schimper, ibid. p. 180. 
1871. Dioonites Dunkerianus, Schenk, Paleeontographica, vol. xix. p. 232, 
pl. xxxvi. figs. 1-5. 
Cycadites Morrisianus, Schenk, ibid. p. 233. 
? Dioonites abietinus, Schenk, ibid. p. 234, pl. xxxvii. fig. 
1874. Dioonites Dunkerianus, Schimper, loc. cit. vol. iii. p. 550. 


_ 


Type. Portions of fronds. Berlin Museum. 

Gippert! thus defines the species: ‘‘ Pt. fronde pinnata, pinnis 
crassiusculis alternis lineari-acicularibus elongatis pectinato- 
patentissimis subremotis eque distantibus 4-5 nervis subacutis.” 
Dunker originally named this plant Nilssonia pecten, but an 
inspection of drawings received from Dunker led Goppert to dissent 
from the original designation. Dunker, in his Wealdenbildung, 
makes one or two slight alterations in Gdppert’s diagnosis; he 
speaks of the venation as ‘‘nervis 3-4 instinctis,” and adds “‘rhachi 
crassa compressa.”’? The specimen represented in Dunker’s pl. v. 


1 (1), p. 184. 
0. ar) ee 


DIOONITES. 43 


fig. 3 shows the pinnee apparently attached rather to the side than 
to the middle of the upper surface of the rachis; the bases are 
slightly swollen, and the apices pointed. In pl. vi. fig. 4 of the 
same author part of the broad rachis is shown, and the approximate 
and narrowly linear pinne are inserted at right angles to the frond 
axis. The specimen figured by Dunker and named by Goppert 
Pterophyllum abietinum, bears such a strong resemblance to Dioonites 
Dunkerianus as seen from the under side of the frond, that I have 
ventured to insert this species as a possible synonym. Schenk has 
previously called attention to this resemblance, and suggests that 
possibly the similarity may amount to specific identity ; Schenk’s 
figure shows a distinct midrib in the pinne, but this must be an 
error in the sketch or some deceptive appearance in the fossil. 
The specimen referred to Dvuoonites abietinus, by Hosius and 
von Marck, is probably a fragment of Zamites Buchianus (Ktt.).' 
Schenk adopts Miquel’s generic term Deoonttes, which the latter 
author proposed in 1851 for this and other species of Pterophyllum. 
In speaking of the genus Cycadites, reference was made to Schenk’s 
substitution of Dioonites or Pterophyllum for Cycadites, in the case 
of certain specimens previously assigned by Dunker to the latter 
genus; an examination of the type specimens having convinced 
Schenk of the absence of a single median vein in the leaf segments, 
and thereforé of the erroneous adoption of the same Cycadites. 
Schenk’s figure 1, pl. xxxvi.* shows a portion of one side of 
a frond with closely placed long and narrow pinnee, which in their 
manner of attachment suggest a spirally twisted frond axis, such 
as we have in the recent cycad Dacrozamia spiralis, Miq.; but this 
may well be an accident of fossilization. Fig. 5 of Schenk shows 
the same kind of rachis as in Dunker’s fig. 4, pl. vi. The figures 
of the epidermal cells given by this writer show a distinctly 
undulating outline in the walls, and the presence of numerous 
stomata. 

It does not seem quite clear whether Schenk has correctly 
included C. Brongniarti, Rom., as a synonym of the present 
species; * he speaks of Roémer’s species as probably the upper 


1 (A. 1), Paleontographica, vol. xxvi. p. 218, pl. xliv. fig. 199. 
2 (A. 2), Paleontographica, vol. xix. 
3 Loe. cit. p. 233. 


44 DIOONITES. 


portion of a frond of Dioonites Dunkerianus. The evidence of the 
figures of Romer and Dunker does not, however, sufficiently 
support this view to justify our following Schenk’s example 
without having examined the type specimen. In the case of 
Cycadites Morrisianus, Dunk., there can be little doubt that the 
specimens referred by Dunker to Cycadites must be transferred 
to the genus Dioonttes. 

It should be pointed out that Ettingshausen had previously 
suggested the specific identity of Cycadites Morrisianus, Dunk., 
and Pterophyllum Dunkerianum, Gopp.; he considered it possible 
that P. Géppertianum ought to be included with these two species.’ 

The specimen figured by Ettingshausen as Cycadites Brongniarti, 
Rom.,? has since been placed by Schenk in a new species— 
Cycadites Heeri.* 

Among the Ecclesbourne specimens there are several good 
examples which must be included in Goppert’s species. At first 
sight many of them would be referred to Cycadites, and the general 
habit of the frond shows a striking resemblance to that of Cycadites 
Saporte, sp. nov., but a closer examination demonstrates that no 
true midrib can be detected, and that the ridge in some of the 
pinne which closely simulates such a central vein, is merely 
the strongly marked convexity of the upper surface of the 
leaf segments. Among recent cycads the genus Lncephalartos 
affords examples of fronds in which the general habit is strikingly 
similar to that of Dioonttes Dunkerianus: E. Ghellinckii, Lem., as 
shown in Pl. XIII. Figs. 38-5, possesses pinne of about the same 
size, and with a very similar mode of attachment, at least as 
regards their almost horizontal position, but in the lateral insertion 
to the rachis the segments of the recent species differ from those 
of the fossil frond. In F. Ghellinckid the convex upper surface 
of the pinne presents a very similar appearance to that in the 
Wealden frond segments, and the sharply acuminate tips of the 
pinne is practically identical in the two cases. In Pl. XIII. 
Fig. 3 a portion of a frond of this species is represented, natural 
size; in Fig. 4 the median groove on the under side of a single 


TA(AGy4) Sip ezile 
2 Ettingshausen, loc. cit. p. 20, pl. i. fig. 9. 
3 Schenk (A. 3), Paleontographica, vol. xix. p. 7, pl. iii. fig. 4. 


DIOONITES. 45 


pinna is clearly shown; and in Fig. 5 a section of a pinna 
illustrates the strongly revolute form of the margins. Lemaire’s 
figure of this species is very poor, and gives an imperfect idea 
of the habit of the leaf." 

Another species of this recent genus, ZL. cycadvfolius, Lehm. 
(Pl. XIII. Fig. 6), also illustrates a point of contact between 
existing and extinct fronds; it differs from #. Ghellineki in its 
somewhat broader pinne. It would, however, tend to a mis- 
conception of the true nature of the Wealden fronds, if the 
generic term Lxcephalartos were adopted on the strength of the 
striking similarity as regards the character of the fronds; we 
unfortunately know nothing as to the flowers and stems of 
Dioonites Dunkerianus. We may adopt a slightly emended form 
of Schenk’s definition for the present species :— 

Frond pinnate, rachis strong, pinne approximate, thick, linear, 
entire, alternate or subopposite; 2-3mm. broad at the widest 
part, with a length of 1lcm. or more, gradually but slightly 
narrowed towards the distal ends; the two rows of pinne 
attached close together to the upper surface of the rachis; the 
lower margin of the basal end of the pinne either slightly 
decurrent, especially towards the tip of the frond, or somewhat 
broadened and bluntly rounded; towards the apex of the frond 
the segments are obliquely inclined, and in the lower portion 
almost at right angles, to the axis. Veins usually indistinct, 
5-6 parallel equal veins in each pinna. 


W.. 3218; Pls Ill. Fig...6; 

23cm. in length. The upper surface of the pinne strongly 
convex; pinne slightly and gradually tapered towards a pointed 
apex. The arrangement of the segments and their somewhat 
broadened bases are very similar to those in Cycadites Saporte. 
Schenk speaks of the pinne as 4-4} cm. long, but his figure 
represents some with a length of 8 or 9cm. In the present 
specimen the longest pinna has a length of 11lcm., and this does 
not include the actual apex. In nearly all the segments it is 
impossible to make out the venation, but in one or two cases 
the parallel veins are visible. Ecclesbourne. Rufford Coll. 


1 Lemaire, pl. plxyvii. 


46 DIOONITES,. 


V. 2823. Pl. II. Fig. 3. 

Cf. V. 1069. Cycadites Saporte. Small specimen, evidently 
close to the frond apex. The pinne are much more oblique 
and decurrent than in the previous example. LEcclesbourne. 


hufford Coll. 


V. 2821. Fig. 2 

Imperfectly preserved piece of rachis with portions of pinne 
on one side; some of the pinne have well-marked venation and 
acute tips. The figure shows some of the more perfect apices. 
The pinne have a breadth of about 2mm., and each is traversed 
by numerous veins, in some of the segments as many as ten may 
be counted. A comparison of this specimen with V. 3218 (Pl. IT. 
Fig. 6) shows some fairly striking differences, and it is not 
improbable that we have to deal with two specific forms; in 
V. 2821 the veins are more numerous, and the pinne are shorter 
and proportionately broader than in the other examples referred 
to this species. Ecclesbourne. Rufford Coll. 


sh A 


Fia. 2.—? Dioonites Dunkerianus (Gopp.). 


Distal terminations of pinne (V. 2821). 


V. 2124c. Broad and flat rachis very like that of Cycadites 
Saporte. On one side the long and narrow pinne are fairly well 
preserved, showing occasional signs of venation and a strong 
convex upper surface. Ecclesbourne. Rufford Coll. 


V. 2127. Probably a portion of a frond near the apex. The 
broad bases of the pinne and their manner of attachment to the 
rachis clearly seen. Ecclesbourne. Rufford Coll. 


V. 2361. Probably a fragment of this species; broad pinne. 
Ecclesbourne. Rufford Coli. 


DIOONITES. 47 


V. 2822. Here the pinne are more oblique to the axis of the 
frond, and the lower edges of the bases more decurrent, as in the 
terminal fragment shown in Pl. Il. Fig. 3 (V. 2823). Eccles- 
bourne. Rufford Coll. 


V. 2824. Similar terminal portion to preceding specimen. 
Ecclesbourne. Rufford Coll. 


V. 2916. Two specimens. Rachis 16cm. long, apparently 
twisted, showing in the lower portion two alternate or sub- 
opposite pinne attached to its upper surface; in the upper part 
the segments are separated by 6 mm. of rachis, suggesting a view 
of the under side of the frond. Pinne slightly convex, presenting 
the appearance of a bread midrib. Ecclesbourne. Rufford Coll. 


Dioonites Brongniarti (Mant.). 


1833. Cycadites Brongniarti, Mantell, Geol. S.E. England, p. 238. 

1841. Pterophyllum Brongniarti, Morris, Annals, p. 119. 

1842. Hisingera Mantellii, Miquel, Mon. Cycad. p. 62. 

1844. Nilssonia Brongniarti, Géppert, Foss. Cycad. p. 57. 

1848. Cycadites Brongniarti, Bronn, Index pal. nomencl. p. 371. 

1849. Zamites Brongniarti, Brongniart, Tableau, p. 107. 

1850. Nilssonia Brongniarti, Unger, Gen. spec. plant. foss. p. 295. 

1851. Nilssonia Brongniarti, Broun and Rimer, Leth. geog. vol. ii. p. 61, 
pl. xxviii. fig. 14. 

1852. Nilssonia Brongniarti, Ettingshausen, Abh. k.-k. geol. Reichs. vol. i. 
Abth. ii. No. 2, p. 238. 

1854. Pterophyllum Brongniarti, Morris, Brit. foss. p. 19. 

1856. Nilssonia Brongniarti, Bornemann, Organ. Rest. Lettenkohl. p. 59. 

1871. Dioonites Brongniarti, Schenk, Paleontographica, vol. xix. p. 236, 
pl. xxxii. fig. 2. 

1874. Dioonites Brongniarti, Schimper, Trait. pal. vég. vol. iii. p. 551. 

1875. Pterophyllum Brongniarti, Topley, Weald. p. 409. 

1881. Dioonites Brongniarti, Renault, Cours. bot. foss. vol. i. p. 51, pl. iv. 
figs. 18 and 14. 

1889. Dioonites Kotoeit, Yokoyama, Journ. Coll. Sci. Japan, vol. iii. p. 44, 
pl. vil. fig. 1, and pl. xiv. fig. 14. 


48 DIOONITES. 


Type. Imperfect fragment of frond. 

In 1883 Mantell described a badly preserved frond fragment 
from the Tilgate beds of Sussex, which he named Cycadites Brong- 
niarti, using the term Cycadites rather as a general designation 
indicative of cycadean affinity, and not in accordance with the 
narrow sense in which Brongniart defined the genus. Morris sub- 
stituted Pterophyllum for Cycadites, and Dunker called attention 
to the plant figured by Romer as C. Brongniarti, which should not 
be confounded with Mantell’s type described under the same name; 
the latter he suggested should be referred to Nilssonia. Goppert 
adopts Wilssonia as the generic term, and Miquel, Ettingshausen, 
and others follow his example. Schenk, on the other hand, 
points out certain discrepancies between the characters of Mantell’s 
species, as further illustrated by subsequently described examples 
of the same type from the Wealden of North Germany, and the 
genus WVilssonia; he substitutes Miquel’s term Dvéoonites for 
Dunker’s Nilssonia.t Schenk’s specimen is in a better state of 
preservation than the English example, and shows more clearly 
the manner of attachment of the pinne. This species differs from 
Otozamites Goppertianus (Dunk.), in the absence of an auriculate 
base to the segments, in its coarser veins, and in the segments 
being more nearly at right angles to the axis of the frond. We 
may adopt Schenk’s definition: ‘‘ Folia pinnata, segmenta e basi 
latiore apicem versus attenuata acuminata lineari-lanceolata integra 
approximata alterna vel opposita, in petioli latere antico sessilia, 
3 mm. usque ad 2°5 cm. longa, 3°5-5 mm. lata, superiora breviora, 
summa brevissima ovata, superiora oblique patentia, media paten- 
tissima, nervi tenues quinque vel sex tenues eequales paralleli.” 

The plant figured by Leckenby? as Pterophyllum angustifolium, 
Leck., from the Oolite of Gristhorpe, shows a marked similarity 
in general appearance to Dioonites Brongniarti. 

Yokoyama’s Japanese species, D. otoei,3 may probably be 
included as a synonym of Mantell’s plant; the former author 
mentions the greater number of veins in the pinne of his plant 
as a distinguishing feature from D. Brongniarti, and speaks of the 


1 Schenk (A. 2), p. 34. 
2 (A.), pl. vii. fig. 3. 
3 Yokoyama (A. 2), p. 44. 


DIOONITES. 49 


latter form as having 5-6 veins in each leaf segment, whereas in 
D. Kotoei there are 7-14. In Schenk’s figure of Mantell’s species 
there are eight or nine veins shown, and in the solitary specimen 
in the Rufford Collection there appear to be at least eight veins. 
The greater length of the segments is another point referred to by 
Yokoyama as a specific character of his plant; but it is difficult 
on comparing the published figures of the two species to detect any 
distinct difference in this respect. On the whole, I am unable to 
discover any sufficient difference between the two plants to warrant 
the retention of Yokoyama’s specific name. Ptilophyllum oligo- 
neurum, Ten.-Woods,! also agrees closely with the English species. 
In speaking of Pterophyllum Richtnofent, Schenk, from China, 
Schenk? suggests that probably some of the fragments so named 
may be identical with Dioonites Brongniarti; there is certainly 
a close correspondence between the two forms, but perhaps 
hardly a sufficiently strong resemblance to justify the inclusion 
of the Chinese specimens in the synonomy of Mantell’s species. 
The plant figured by Schenk * as Pterophyllum equale, Brong., from 
Persia, resembles Dioonites Brongniarti. In Schenk’s specimen 
the pinne appear to be inserted on the upper surface of the 
rachis, and not laterally as the generic term Pterophyllum implies. 


V. 2748. 26cm. long. The alternately disposed pinne are 
attached to the middle of the upper face of the frond axis. 
Venation clearly marked. The lower margin of the pinne is 
curved gradually upwards, cutting off the veins obliquely, and 
the upper margin is practically horizontal. The segments are 
somewhat less than those in Mantell’s figure, but there can be 
little doubt as to the specific identity of the specimens. Lccles- 
bourne. Rufford Coll. 


1 Jack and Etheridge (A.), pl. xviii. fig. 11. 

2 Richthofen (A.), China, vol. iv. p. 247, pl. xlvii. fig. 7, and pl. xviii. 
figs. 5, 6, and 8. 

3 Schenk (A. 7), Bibl. bot. vi. pl. v. fig. 23. 


50 NILSSONIA. 


Genus NILSSONTA, Brongniart. 
[Ann. Sci. Nat. vol. iv. 1825. p. 200.] 


In 1820 Nilsson? described and figured certain plant remains 
from Hor, a small village north of Lund in Scania, and regarded 
them as probably fern fronds, but he made no attempt to define 
them specifically. In 1825 Brongniart refigured and described 
some of Nilsson’s specimens under the following specific names: 
Nilssonia elongata, N. brevis, N. (2?) equalis, Pterophyllum magus, 
and P. minus. It was on one of Nilsson’s specimens that 
Brongniart founded his genus Milssonia, and also Pterophyllum ; 
the two Hor species referred by Brongniart to this latter genus 
have since been transferred by Nathorst* to Schimper’s genus 
Anomozamites. We have the first complete diagnosis of WVilssonca 
in the Prodrome,? where it is thus defined: ‘‘ Feuilles pinnées ; 
pinnules rapprochées, oblongues, plus ou moins _alongées, 
arrondies au sommet, adhérentes au rachis par toute la largeur 
de leur base, & nervures ‘paralléles dont quelques-unes sont 
beaucoup plus marquées.” In his later work,‘ Brongniart retains 
this name and speaks of Vilssonia as closely allied to Pterophyllum. 
Miquel ® substituted a new generic term, Hisingera, for some of 
the species of Wlssonia, and as an example of the new genus he 
cites Cycadites Brongniarti, Mant. Goppert® accepts Brongniart’s 
genus in its wide sense, and does not suggest the institution of 
any sub-genera. In 1856 Bornemann adopted the following 
definition of Milssonia:’ ‘‘ Frondes coriacese, pinnate, vernatione 
circinate, foliola contigua continue tota latitudine inserta, patentia, 
abbreviata, basi passim coherentia, apice obtusa vel truncata, 
nervis parallelis arcuatis apice confluentibus nonnullis validi- 
oribus.’’ This writer points out the difficulty of recognizing the 


1 Nilsson, p. 108. 


p. 90. 
(A. 4), Tableau, p. 63. 


NILSSONIA. ol 


two different kinds of veins in the leaf divisions. Schenk, in 
his Flora der Grenzschichten' includes Nilssonia among the ferns, 
and refers to certain specimens in which the leaves show 
numerous round projecting structures between the veins, and 
which he regards as sporangia or sori; and it is on the strength 
of these appearances, suggestive of fern fructification, that the 
genus is excluded from the Cycadacee. Schenk speaks of the 
veins as equal and simple, and refers to the epidermal cells as 
having the straight walls characteristic of cycads. No great 
importance should be attached to any argument based on the 
form of the cell walls, as Schenk himself has admitted; but 
the fructification is a much more important feature. Saporta? 
places Milssonia in the Cycadaceez, and considers that Schenk 
was probably deceived by certain leaf parasites, which might 
well present an appearance closely simulating fern sori. Nathorst ® 
follows a similar course, and speaks of our ignorance as to 
the actual nature of Schenk’s sori, seeing that no traces of 
structure have been preserved; he suggests stomata and parasitic 
fungi as two possible explanations of these sorus-like appearances. 
Solms-Laubach* does not accept the proposed explanation as 
satisfactory, and inclines to follow Schenk in classing MWilssonia 
among the ferns, on the strength of the sorus-like bodies on the 
leaf lamina. Nathorst draws special attention to the insertion 
of the leaf segments on the upper surface of the rachis as an 
essential character of the genus; he speaks of the veins as equal 
and simple. Various authors have spoken of two kinds of veins 
in the leaves of Mlssonia, stouter and finer veins, but Nathorst 
remarks that Schenk has recognized his mistake with regard to 
the supposed two sets of veins; he mistook folds in the leaf 
lamina for well-marked simple veins.° Schimper,® in the first 
volume of the Trait. pal. vég., classes Nilssonia with the 
Filicine, and accepts Schenk’s interpretation of the apparently 


1 
( 

2 (A. 2), Pal. Franc. vol. ii. p. 41. 

3 (A. 2), Foss. Fl. Schwedens, p. 20. 

* Fossil Botany, p. 139. 

5 Nathorst (A. 2), p. 18. 

6 p. 488. 


oe NILSSONTA. 


fertile specimens ; but in a later work! by this author we find 
Nilssonia placed close to Pterophyllum in the Cycadacee. 

Without following in further detail the various descriptions 
or definitions of this genus, we may thus sum up the chief 
characters by which the species may best be recognized :— 

Frond coriaceous, the lamina more or less deeply pinnatifid, 
the lines of division generally extending almost to the rachis; 
segments attached to the upper surface of the axis by the 
entire base, contiguous, usually broad and truncate, but varying 
considerably in size and shape; apices obtuse or truncate. Veins 
simple and equal. 

In connection with Mlssonia, which may best be considered 
as a genus of doubtful affinity, but probably cycadean, there are 
three other genera of which some mention must be made,— 
Anomozamites, Ptilozamites, and Pterophyllum. As regards the 
last, Nathorst has on several occasions emphasized the distinct 
difference as regards the manner of attachment of the leaf 
segments; in Pterophyllum they are inserted laterally on the 
rachis; in WVilssonia, as in Dioonites, they are attached to the 
upper face of the leaf axis.2 The genus Anomozamites was 
instituted by Schimper® for certain species of Pterophyllum with 
irregularly pinnatifid leaves, and this term has been generally 
adopted; the veins are described as simple and parallel, and the 
segments as laterally attached. Nathorst, however, has instituted 
a genus, Ptilozamites,* in which are included pinnate and bipinnate 
fronds, which in habit correspond fairly closely with Anomozamites, 
but differ in the possession of forked veins which dichotomize 
at the base, and occasionally branch a second time before reaching 
the margin of the leaf segment. The plant originally described 
by Leckenby as Ctenis Leckenbyi,® the specific name having been 
suggested by Bean, shows very clearly the characters of the 
venation and the branching habit of the frond; this must now 
be included in Nathorst’s Pt/ozamites® as suggested by the author 


1 (A.), Zittel, Handbuch, p. 225. 

2 Nathorst (2), p. 61, and (A. 38), Denkschr. k. Ak. Wiss. vol. lvii. p. 46. 
3 (A.) Trait pal. vég. vol. i. p. 140. 

4 (A. 1), Flor. Hoganas och Helsingborg, p. 21. 

5 (A.), Quart. Journ. Geol. Soc. vol. xx. p. 78, pl. x. fig. 1. 

6 Nathorst (A. 1), Flor. Héganis, p. 21. 


NILSSONIA. 53 


of the genus himself. An inspection of Nathorst’s figures of 
Anomozamites and Ptilozamites species suggests a difficulty in 
certain cases in deciding between the two genera. In such a form 
as Anomozamites gracilis, Nath.,! we have well-marked branching 
in some of the veins, and a close approximation in general 
appearance to other species included in Ptlozamites, cf. e.g. 
P. Heert, Nath.,? and 4. minor * (Brong.). 

Fontaine‘ figures a few small fragments of leaves from the 
Potomae flora under the name Anomozamites, but there is hardly 
enough material to justify even a generic determination, and still 
less to warrant the institution of two new species. 


Nilssonia Schaumburgensis (Dunk.). 
[Figs. 3a, b, and ¢.] 


1843. Pterophyllum Schaumburgense, Dunker, Progr. p. 6. 

1844. Pterophyllum Schaumburgense, Goppert, Foss. Cycad. p. 54. 

1846. Pterophyllum Schaumburgense, Dunker, Wealdenbildung, p. 15, pl. i. 
fig. 7; pl. ii: fig. 1; pl. vi. figs. 5-10. 

1848. Pterophyllum Schaumburgense, Broun, Index pal. nomencl. p. 1056. 

1849. Pterophyllum Schaumburgense, Brongniart, Tableau, p. 107. 

1850. Pterophyllum Schaumburgense, Unger, Gen. spec. plant. foss. p. 292. 

1851. Pterophyllum Schaumburgense, Miquel, Rangschik. foss. Cycad. p. 218. 

1852. Pterophyllum Schaumburgense, Kttingshausen, Abh. k.—k. geol. Reichs. 
vol. i. Abth. ii. No. 2, p. 22. 

1856. Pterophyllum Schaumburgense, Bornemann, Organ. Rest. Lettenkohl. 
p. 58. 

1869. Anomozamites Schawmburgensis, Schimper, Trait. pal. vég. vol. ii. p. 141. 

1871. Anomozamites Schaumburgensis, Schenk, Paleontographica, vol. xix. 
p- 231, pl. xxxiii. 

1883. Pterophyllum Schaumburgense, Peyton, Quart. Journ. Geol. Soc. vol. 
xxxix. Proc. p. 3. 

1890. Nilssonia cf. Schaumburgensis, Nathorst, Denkschr. k. Ak. Wiss. 
math.-nat. Cl. vol. lvii. pp. 45, 49, and 53, pl. i. figs. 6-9. 

1894. Nilssonia Schaumburgensis, Yokoyama, Journ. Coll. Sci. Japan, vol. vii. 
pt. ui. p. 227, pl. xx. figs. 12 and 14; pl. xxi. fig. 14; pl. xxii. 
figs. 5-7. 


1 (A. 1), Flor. Bjuf. p. 65, pl. xy. fig. 15. 

2 Ibid. p. 60, pl. xii. figs. 1 and 7. 

3 Ibid. p. 66, pl. xiv. figs. 5-7, and pl. xviii. fig. 4. 
4 (A. 2), Potomac Flora, p. 167, pl. xxx. 


54 NILSSONIA. 


Type. Several specimens of leaves. 

Dunker defined the species as follows :— 

‘‘Pterophyllum fronde pinnate vel, rarissime quidem, profunde 
pinnatifida, pinnis alternis approximatis sub-obliquis irregularibus, 
oblongo-ovatis, vel quadratis vel rotundatis, infimis subdecurrenti- 
bus, nervis crebris tenuibus instructris, rhachi (suptereti ?) 
longitudine striata.” 

The specimens figured by Dunker from the North German 
Wealden beds show a considerable variation in size and form; this 
variable character is also well brought out in the later and more 
perfect figures in Schenk’s monograph. Such a specimen as that 
represented by Dunker in pl. i. fig. 7' must probably be regarded 
as a leaf seen from the under side, thus showing a Pterophyllum- 
like appearance. Schenk draws attention to the apparent lateral 
attachment of the segments in some of the specimens which are 
seen from the under surface, but notes that there can be no doubt 
as to their actual insertion on the upper surface of the axis. 
Schimper includes this variable Wealden species in his genus 
Anomozamites, and Schenk accepts this determination. Peyton has 
previously recorded the species in the English Wealden beds, but 
no detailed descriptions or figures accompany his note. Nathorst 
records from Japan specimens of what is most probably the same 
species as the English and North German forms; he refers to 
a previous paper? in which he pointed out the true Wilssonia 
character of Dunker’s species, the segments being attached to the 
upper face of the rachis, and not laterally as in Pterophyllum or 
Anomozamites. A leaf fragment closely resembling the present 
species is figured by Schenk* from Persia as Anomozamites minor, 
Schimp. In Yokoyama’s recent contribution on Mesozoic plants 
from Kozuke, etc., several specimens are referred to this species 
on the authority of Nathorst; the figures suggest a lateral attach- 
ment of the unequal segments, but possibly the leaves are shown 
with the under side uppermost. Nathorst’s figures of this species 
from Japan represent typical Wi/ssonia fronds. 


(A. 2), Wealdenbildung. 
(2), p. 82. 
7 


1 
? (2) 
3 (A. 7), Bibl. bot. vi. pl. v. fig. 21. 


NILSSONIA. 55 


V. 2171.** Figs. 8a and 8. The two figured specimens are 
examples of the narrower form of the species: in @ the almost 
entire lamina resembles a small example of Zeniopteris; in 6 
the truncate segments are well shown; and in both cases the 
venation is distinct. Both specimens are represented twice the 
natural size. Rufford Coll. 


V. 2171a. Fig. 3c. Broader specimen, 1:1 em. in breadth. The 
median groove on the upper surface and the veins are very 
distinct. This example serves as a connecting link between the 
larger forms of the plant as described by Schenk, and the smaller 
English specimens. 


Nilssonia Schaumburgensis (Dunk.). 
Fig. 3a and bd (V. 2171**). 


Fie. 3¢ (V. 21714). } (Twice nat. size.) 


V. 2171. Several specimens. The variation in the size and 
division of the lamina is well illustrated in these examples. 
Generally speaking, Schenk’s figures represent leaves with more 
regular lobes than are found in the English specimens. In some 
cases the segments are numerous and very narrow, in others the 
lamina is almost entire. 


V. 21714. A specimen with the lamina entire for a length of 
4-2 cm. 


V. 2171c. This specimen of one of the narrower forms of the 
species shows a depression at the distal end of the lamina, and 
presents a very similar appearance to that of Zeniopterts Beyrichit 
(Schenk) as figured by Schenk. Ecclesbourne. Rufford Coll. 


56 OTOZAMITES. 


V. 2172. This specimen was erroneously included under Tenio- 
pteris Beyrichii in Vol. I. (p. 126). 


V. 2234a. 5mm. broad. In a length of 42cm. there are 
about 23 divisions in the lamina; in the same length of V. 21714, 
the lamina shows no divisions. At one end the segments gradually 
decrease in size until they almost disappear. Ecclesbourne. 

Rufford Coll. 


Other specimens: V. 716. Hastings. Dawson Coll. V. 1436. 
Ecclesbourne. Presented by P. Rufford, Esq., 1886. V. 2284. 
Ecclesbourne. ufford Coll. 


Genus OTOZAMITES, Braun. 
[Miinster, Beitrag. Petrefact. Heft vi. 1843, p. 36.] 


The name Otozamites, instituted by Braun in 1843, was defined 
by him as follows: Leaves pinnate, pinne alternate and ap- 
proximate, auriculate, and attached by a portion of the base; 
veins radiating from the point of attachment to the margins of 
the segments. 

This author includes Zamites falcatus (Sternb.), Z. Bucklandi 
(Brong.), and Z. brevifolius (Braun, etc.) as species of his new 
genus; Z. fulcatus was first figured by Sternberg! as Odontopteris 
Jaleata, and O. Bucklandi was described by Brongniart in 1825 as 
Filicites Bueklandi var. Britannica*; both of these species were 
assigned by Morris * to his genus Pételophyllum. Brongniart adopts 
Braun’s generic name, and points out that Ofopteris, Lindley and 
Hutton, corresponds to Otozamites of Braun. This genus affords 
another example of confusion in nomenclature arising from a 
difference of opinion as to the botanical relationship of the fossil 


1 (A.), Flor. Vorwelt, pl. xxiii. fig. 1, fase. 5 and 6, p. 78. 
2 Brongniart (4), p. 422. 
Sl) ip. dle 


OTOZAMITES. 57 


fronds. Brongniart! says that Otozamites Bucklandi, Brong., was 
figured by De la Beche? as a fern from the Lias of Axminster, 
and by Lindley and Hutton as O¢opteris obtusa,> the typical 
species of the genus. The same writer suggests the advisability 
of distinguishing certain leaf forms under another genus, Spheno- 
zamites, of which the chief characteristic is the absence of an 
auriculate base in the pinne; this genus has come into general 
use, and serves a useful purpose as a convenient provisional 
term. Bornemann‘ has suggested that probably some of the 
plants referred to Ofozamites are without true auriculate pinne, 
the apparently eared form being merely a result of pressure on 
the upper surface of the thick pinne. He removes some of 
Braun’s species from the genus, and speaks of Otozamites brevifolius 
(Sternb.) and O. gramineus (Morr.) (=Zamites gramineus, Morris) 
as typical species. The following is Bornemann’s emended version 
of Braun’s diagnosis:° ‘‘ Leaves pinnate; pinne approximate, 
alternate, or subopposite, lanceolate, pointed or more or less 
blunt, auriculate at the base, and attached to the rachis only by 
the lower part, the upper corner of the auriculate base prolonged 
and partly covering the rachis. Veins radiate from the point of 
attachment towards the margin of the pinne, and are for the 
most part dichotomous.”’ This definition appears to be on the 
whole satisfactory, but Bornemann unfortunately errs in describing 
the pinne as attached to the rachis by the lower portion; the 
manner of insertion of some auriculate pinne cannot correctly be 
described according to his definition. Schenk has discussed at 
some length the botanical position of the genus Otopteris in his 
Flora der Grenzschichten;® he draws attention to a specimen of 
which the segments exhibit a peculiar marginal structure, 
suggestive of a Pteris-like fertile leaf. The structure of the 
epidermal cells is also referred to in support of the inclusion of 
this genus among the Filicine; but in his later writings Schenk 
speaks of Ofozamites as a member of the Cycadacea. 


1 Tableau, p. 61. 

2 Pl. vii. fig. 2. 

3 (A.), Foss. Flor. pl. exxviii. 
4p. 49. 

epi o2. 

5 p. 186. 


58 OTOZAMITES. 


Schimper, in the first volume of his Zract. pal. vég,' adopts 
the genus Otopteris, L. and H., but afterwards (vol. i1.)? accepts 
Braun’s generic name Otozamites ; he institutes a sub-genus Rhombo- 
zamites for Otozamites Beanti and other species, and makes use 
of one of Pomel’s terms, Cyclozamites, for Otozamites Bunburyanus 
and other forms. This subdivision seems quite unnecessary, and 
tends rather to confusion than to useful classification. Saporta® 
retains Otozamites Bucklandi (Brong.) as the type of the genus, 
but in his diagnosis mention is made of certain features which 
set rather narrow limits to the generic characters; the basal 
callosity of the pinne and the auriculate upper angle of the base, 
are features which do not always appear in fronds which must 
be referred on general grounds to the genus Ofozamites. It is 
true we frequently find that the upper lobe of the pinna base 
is more decidedly auriculate than the lower, but this is not 
universal. Saporta’s figures of some of the species of Ofozamites 
show this quite clearly; e¢.g., O. Reglei, Sap.,t O. Brongniartn, 
Schimp.,® ete. In describing the characteristics of the various 
examples of the genus, Saporta points out the numerous varia- 
tions from the normal type. In discussing the geological history 
of the genus, this author refers to the absence of Ofvzamites from 
Wealden and Neocomian strata; since these words were written 
several examples of Wealden forms have been discovered, and 
the material acquired in recent years shows that the small plant 
figured by Dunker as Cyclopteris Klipsteinii® is most probably a 
species of Otozamites. The groups into which Saporta divides 
this genus have been adopted by Schimper in Zittel’s Handbuch.’ 
The recognition of certain typical species as representatives of 
different forms of a genus may in some cases be a convenient 
aid to classification, but there is always the danger of unduly 
emphasizing slight and unimportant differences for the sake of 
such purely arbitrary grouping. In the case of a genus such as 


1 p. 483. 

Za pemlayis 

3p. 45. 

Jnl @bh< 

BSP levenie 

6 (A. 2), Wealdenbildung, pl. ix. figs. 6 and 7. 
7p. 221. 


OTOZAMITES. 59 


Otozamites we know very little indeed as to its exact botanical 
position, and for the present, at least, it will probably be better 
not to bind ourselves to any of these subdivisions of the genus. 
We may adopt a definition of Otozamites very similar to that 
previously quoted from Bornemann, but which gives a more definite 
expression to the variable character of the numerous forms and 
fronds included in Braun’s genus :— 

Frond pinnate; pinne attached to the upper surface of the 
rachis by a portion of the auriculate base, base more or less 
distinctly auriculate, the upper lobe often more prominent than 
the lower; segments may be approximate, imbricate, or distinct. 
Veins numerous and branched, radiating from the point of 
attachment and cut off obliquely by the margin of the pinna; in 
the longer and narrower form of pinna the veins are practically 
parallel to the edges of the segment; pinne vary from long, 
narrow, and linear-lanceolate, with acute tips, to broadly oval 
or almost orbicular in form, with bluntly rounded apices. 

Solms-Laubach,! after speaking of the flabelliform venation of 
Otozamites, goes on to say that in some forms of this genus the 
veins of the pinne conform much less distinctly to that type than 
in others. If we examine the narrower and longer forms of 
Otozamites pinne, the veins become more or less parallel to the 
segment margins soon after leaving the auriculate basal portion. 
In the longer segments of Otozamites gramineus and other species 
this is the case, so that the flabelliform character of the venation 
cannot by any means be relied upon as a constant and easily 
recognized characteristic of the genus. It is often a difficult 
matter to decide whether the pinne bases are actually auriculate ; 
in dealing with some fronds we find it almost impossible to 
draw a well-marked line between Otozamites, Ctenophyllum, and 
Ptilophylium, ete. 


1 Fossil Botany, p. 89. 


60 OTOZAMITES. 


Otozamites Klipsteinii (Dunk.). 


[Pl. I. Figs. 3 and 4; Pl. VII.] 


1846. Cyclopteris Klipsteinii, Dunker, Wealdenbildung, p. 11, pl. ix. figs. 
6 and 7. 

1848. Cyclopteris Klipsteinii, Bronn, Index pal. nomencl. p. 377. 

1849. Adiantites? Klipsteinii, Brongniart, Tableau, p. 107. 

1850. Cyclopteris Klipsteinii, Unger, Gen. spec. plant. foss. p. 95. 

1869. Aneimidium Klipsteinii, Schimper, Trait. pal. vég. vol. i. p. 486. 

1871. Aneimidium Klipsteinii, Schenk, Paleontographica, vol. xix. p. 218, 
pl. xxxi. fig. 6. 


Type. Detached pinna and fragment of frond. 

Dunker defines the species as follows: ‘‘Cyclopteris fronde 
pinnata, pinnulis alternis sessilibus? ovato-oblongis sequalibus, 
nervis creberrimis flabellatis tenerrimis.’” He points out that 
the veins are exceedingly delicate, and apparently dichotomous 
in the upper portions, the venation shown in his figures being 
coarser than it actually is in the specimens. Ettingshausen 
figures four detached leaflets which he describes as intermediate 
in character between Cyclopteris Mantelli and C. Klipsteinii; in 
his drawings there appears to be a distinct suggestion of a 
midrib, but nothing is said in the definition of the species as 
to the existence of a median vein. It has already been pointed 
out (Wealden Catalogue, vol. i. p. 131) that these leaflets are 
certainly not typical examples of Sagenopteris Mantelli (Dunk.), 
as Ettingshausen erroneously states. It will be better to leave 
them out of consideration as doubtful fragments. 

Schenk figures a single pinna of this species, which shows 
very clearly an auriculate base and dichotomously spreading 
veins. The Rufford Collection contains a large number of well- 
preserved fronds with pinne of various sizes, and which in some 
cases are clearly identical with the species figured by Dunker 
and Schenk as Cyelopteris or Aneimidium Klipsteinti. The 
number and variety of the specimens present a difficulty as 
regards specific determination. A casual inspection of the fronds 
with large and broadly oval segments would probably lead one to 
institute a new specific name for their reception, but on carefully 
examining and comparing all the examples, it appears to be 
impossible to determine definite specific limitations. The frond 


OTOZAMITES. 61 


figured in Pl. I. Fig. 3 must certainly be referred to Dunker’s 
species; the pinne agree closely with that figured by Schenk, 
and with the inferior figure by Dunker. We have a pinnate 
frond, with segments attached by an auriculate base; the venation 
agrees with that of Otozamites, and the general character of the 
leaf points to that genus as the most convenient designation for 
the specimens. Another specimen, Pl. II. Fig. 4, possesses 
pinne of the same form as those in V. 2286, but differs in its 
much stouter rachis and in the imbricate disposition of the 
segments. It is difficult to speak confidently as to the relation 
of the several specimens one to another, but probably we have 
in Pl. IL. Fig. 4 the lower portion of a young frond of the same 
form of which Pl. I. Fig. 3 represents a terminal fragment. 
Leaving a more detailed notice of these specimens until later, 
we must turn to the extremely fine examples of the larger 
fronds, such as those represented in Pl. VII. Fig. 9, etc. It is 
possible there may be two or three species represented by this 
splendid series of fronds, which I have referred to the genus 
Otozamites; in some the pinne are longer and narrower than 
in others, and in some we have a shorter and broader form of 
segment. On the whole, in the absence of any constant and 
well-marked differences consistent with separate species, I prefer 
to include nearly all these various forms under one species, 
Otozamites Klipstemit (Dunk.), and resort to descriptive terms 
for the designation of one or two varieties. We may thus 
define this comprehensive species: Frond pinnate, rachis fairly 
stout, tapering to a slender axis in the terminal portion; pinne 
attached to the upper surface of the rachis, in the young frond 
probably imbricate, with a more or less well-marked auriculate 
base; veins numerous, radiating from the point of attachment 
towards the margin of the pinne; apices obtuse, pinne alternate 
or subopposite, in the mature fronds almost at right angles to 
the axis, or more or less obliquely inclined. 

Before describing the individual specimens referred to this 
species and its varieties, we may notice some other forms of the 
genus Otozamites, and other plants with which the present example 
may be compared. Otozamites Beanit (L. and H.)! corresponds 


1 Fossil Flora, pl. xliv. 


62 OTOZAMITES. 


with O. Klipsteinit in having auriculate pinne, broader and stouter 
than the majority of species of the genus. In describing the type 
specimen, Lindley and Hutton ask if the plant may be a pinnated 
leaf of the Cycadeoidee, but reply with a decided negative; in the 
second volume of the Fossi] Flora the generic name Otopteris is 
substituted by the authors of the species for Cyclopteris, under 
which the plant was originally described. In the Leckenby Collec- 
tion in the Woodwardian Museum, Cambridge, there are several 
very fine specimens of Ofozamites Beanii, which in some instances 
show a terminal portion with small pinne very similar to such 
an apical tip as is represented in Pl. I. Fig. 3. In the Yorkshire 
plant the species are, however, very distinctly auriculate, and 
have the upper portion of the base more prominently lobed than 
is the case in O. Klipsteinii (Dunk.). In one of Leckenby’s 
specimens, which possesses a rachis 43°5 cm. long, the largest 
pinna has a length of 3 cm. and a breadth of 1-9 cm., the smallest 
measuring lem. by 7mm. Some of Zigno’s figures of his species 
Otozamites Molinianus) present a striking likeness to O. Beanit, 
and ought most likely to be referred to that species. Kurr’s figure 
of what he calls Zamites Mandelslohi* agrees very closely with 
the specimen represented in Pl. 1V. Fig. 4; the genus Otozamites 
has been rightly substituted by Schimper for Kurr’s species. 
The larger pinne of O. Alipsteinit show a certain resemblance 
to those of some few previously described plants. but in no case 
does the similarity appear sufficiently pronounced to justify a 
reference to the same species. In the third volume of the 
Gondwana Flora of India,’ Feistmantel figures some isolated 
pinne under the name Glossozamites Stoliczkanus, Feist., and 
describes them as probably constituting the largest representatives 
of Schimper’s genus. The form of these large segments is not 
at all unlike that of the Wealden pinne, but in the latter case 
the auriculate base favours the adoption of the genus Otozamites. 
The plant described many years ago by Goppert, from a much 
lower geological horizon, under the name of Cyelopteris frondosa * 
(Gopp.), may be compared, as regards the form of its leaf segments, 


1 (A.), Flor. foss. Oolit. vol. ii. pl. xxxv. 

oP). igen. 

3 Vol. iii. pt. i. pl. xx. figs. 4 and 6. 

4 Schimper (A.), Trait. pal. veg. vol. i. p. 453, pl. xxxv. 


OTOZAMITES. 63 


with the larger specimens of the present species. Schimper places 
this Culm species in his genus Cardiopteris, and describes it as an 
unusually large-leaved neuropteroid fern. The venation of the 
large pinne of the Wealden fronds suggests that of certain fern 
pinnules; and it must be borne in mind in dealing with such leaves 
as those of a neuropteroid or otozamitean type, that we are 
unable to speak dogmatically as to the botanical position of the 
species. 

Among the Jurassic plants figured by Saporta’ there occur 
a few examples of fronds with leaf segments comparable to those 
of O. Klipsteinii; e.g., O. decorus, Sap., and O. lagotis, Brong. 
A detached segment named by Saporta Sphenozamites Brongniartt, 
agrees fairly closely with some of the larger pinne of the English 
Wealden specimens, but the French fragments are too small to allow 
of any precise comparison, and hardly worthy of a special specific 
name. The frond fragments described by Hosius and yon Marck as 
Pterophyllum blechniforme* have a certain resemblance to some of 
the English specimens, but are clearly not specifically identical; the 
generic name Otozamites would seem to be more applicable to this 
species of Hosius and von Marck than that of Pterophyllum. The 
pinnee of Fontaine’s species Zamites tenuinervis* may be compared 
with some of the longer segments of Oftozamites Klipsteinii, but 
there is not sufficient evidence to justify the inclusion of the 
Potomac and English forms in the same species. 

Finally, we have a somewhat similar form of leaf in some 
examples of the Palzozoic genus Cordaites* recently figured by 
Grand’ Eury from the Coal-field of Gard. Whilst drawing attention 
to some of the plants previously described, in which a greater or 
less resemblance may be traced to the present species, it must 
be definitely stated, that we are without any satisfactory evidence 
as to the exact position in the plant kingdom to which these large- 
leaved forms should be referred. Having regard to the general 
habit of the fossil fronds, the apparently stiff nature of the pinne, 
etc., the cycads appear to be the more likely plants with which to 


WV Pips 

2 (A. 1), Paleontographica, vol. xxvi. pl. xliv. fig. 197. 
8 Potomac Flora, pl. xvii. fig. 1, ete. 

4 Grand’ Eury, pl. vi. figs. 14 and 16. 


64 OTOZAMITES. 


compare Otozamites Klipsteini. Among recent cycadean species 
we do not meet with fronds possessing pinne with a well-defined 
auriculate base, but we are not without instances of fronds which 
in other respects bear a decided resemblance to the Wealden plant. 
Zamia purpuracea, Ait., may be cited as one recent species with 
large pinnee comparable to those of the Wealden plant; the young 
pinne of this living cycad are very similar in form to the more 
terminal segments of Otozamites Beanii. In this species of Zamia, 
as also in Z. pygmea, Sims, and in some other forms, etc., the 
pinne are distinctly narrowed towards the point of attachment 
to the rachis, from which they are readily detached, leaving a 
well-marked scar. The recent fronds correspond rather more 
closely with such fossil forms as Podozamites Reinit, Geyl., etc., 
described by Geyler! and others. In Ofozamites latifolius (Phill.)* 
we have another large-leaved form which may to some extent 
be compared with the present species: the specimens of Phillips’ 
species are imperfectly preserved, and do not give any decided 
indication of an auriculate base; the venation appears to agree 
fairly closely with that of Nathorst’s genus Ptilozamites, and the 
prominence of the veins reminds one of some of the large and 
boldly veined segments of such recent species as Zamia picta, 
Z. Skinneri, Warsz., ete. 


V. 2336. Pl. I. Fig. 8. This terminal portion of a frond 
must no doubt be referred to the same species in which the 
detached segments figured by Dunker and Schenk have been 
included. In Dunker’s specimens there is not the same distinctly 
auriculated base as in this example, but this may be put down 
to less perfect preservation and possibly inaccurate drawing ;. in 
Schenk’s leaflet of the same species the auriculate base is distinct, 
and agrees exactly with that in the English frond. The veins 
appear to be rather fewer and farther apart in this terminal 
portion than in the pinne of the larger fronds. Largest pinna 
2em. by 9mm. LEcclesbourne. Rufford Coll. 


V. 2170a. Two detached pinne ; the venation finer, and exactly 
like that in Schenk’s example. Ecclesbourne. Rufford Coll. 


1 (A.), Paleontographica, vol. xxiv. pl. xxxiv. 
2 Phillips, p. 171, fig. 6. 


OTOZAMITES. 65 


O. Klipsteinii (Dunk.), var. superba mihi. 


V. 27450. Pl. I. Fig. 4. 

Rachis 12°5 cm. in length. Largest pinna 4:2 cm. by 1°6 em.; 
smallest 3°8 by 1:2. The largest pinne of V. 2336 has a length 
of 2 cm. and a breadth of 9mm. Some of the pinne are very 
clearly preserved, and show excellent venation. The rachis 
appears to be slender, as in V. 2336. The upper pinne have 
very slightly auriculate bases, which are attached by their central. 
portion to the surface of the frond axis. In the lower pinne the 
base is more distinctly lobed. Cf. Pterophyllum oblongifolium, 
Kurr [ = Glossozamites oblongifolius (Kurr) ].! There is not, I am 
inclined to think, sufficient proof of any important difference 
between V. 2336 (Pl. I. Fig. 3) and V. 2745a (PI. I. Fig. 4) 
to warrant a specific separation; but as we shall find a gradual 
transition from such specimens as V. 2745a to the fronds with 
much larger pinne, e.g. Pl. VII., it will be convenient to institute 
a variety of Dunker’s species under the name superba. Eccles- 
bourne. Rufford Coll. 


V. 2170, Pl. VII. Fig. 9. It may be, as suggested in the 
introductory account of the species, that we have two or three 
species included under O. A lipsteinii (Dunk.), but thanks to the 
numerous and well-preserved specimens it is possible to examine 
an unusually fine series, and so escape to some extent from the 
dangers of fragmentary and imperfect portions of fronds, which 
often lead to unnecessary multiplication of specific names. In 
the present instance we can trace a gradual transition from the 
short and broad segments, such as those represented in Pl. VII. 
Fig. 5 (V. 2745), to the large forms such as Pl. VII. Figs. 1 
and 6 (V. 2122). The leaves with the broader and more or 
less imbricating obliquely set pinnze come very near to O. Beant, 
and those with more separate, longer, and narrower segments 
closely resemble the pinne figured by Fontaine as Zamites 
tenuinervis, Font. In fragments of young fronds or in the lower 
part of older fronds the pinnze may have a length of 2 cm. and 
a breadth of 1-4cem., in the larger segments 8:2 x 2:'4cem. In 
this specimen (V. 2170) the rachis is 23cm. in length; it has 
left a hollow mould in the rock, roofed over by the basal ends 


Sy iturr plead. fee Oo: 


66 OTOZAMITES. 


of the pinne; on one side there are nine pinnw, and on the 
other seven. These show the broad, obtusely terminated form of 
the segments very clearly ; they are attached to the upper surface 
of the rachis by the central portion of the distinctly auriculate 
base. Most of the pinne overlap, the upper edge of each pro- 
jecting over the lower margin of the pinna next above it; very 
similar in habit to some of the larger specimens of O. Beanii in 
the Leckenby Collection. On the under side of the same slab 
there occur portions of pinne of another frond, and a section of 
a rachis mould, about 6mm. broad and 8mm. in depth; also 
a specimen of Cycadites Saporte, sp. nov. Ecclesbourne. 


Rufford Coll. 


V. 2122a. Pl. VII. Fig. 2. Rachis imperfectly preserved ; 
portions of six pinne, the uppermost having the same form as 
the largest of V. 2745 (Pl. I. Fig. 4), the lowest and largest, 
6-4 x 3 em. (Pl. VII. Fig. 2), showing a slight lobe in the middle 
of the upper margin. In some of the segments the point of 
attachment is well shown, and the numerous fine spreading 
veins are well marked in the carbonaceous surface layer. 


V. 21260. Pl. VII. Figs. 4 and 8. Frond 28cm. long.; 11 
pinne on one side. Hollow round mould of rachis, about 4-5 mm. 
in diameter. Short and broad overlapping pinne, 4°8 xX 2°5 cm., 
and very like those of V. 2740a, V. 2740, etc. The upper and 
smaller pinne, as shown in the figure, are narrower and longer 
and not overlapping; these agree exactly with the segments in 
V. 27450 (Pl. I. Fig. 4). Uppermost pinne about 3 cm. x 1°8 cm.; 
a segment separated from this by 11 cm. and attached to about the 
middle of the specimen, measures 4°5 X 2°5cm. The two larger 
pinnee (Pl. VII. Fig. 8) agree very closely with those of V. 2170 
(Pl. VII. Fig. 9); they have a strongly convex upper surface, 
and show a sharp bending down of the lamina near the point 
of insertion to the rachis. This bending of the pinna base is 
exactly similar to the appearance frequently presented by the 
stiff leathery pinne of recent species of Lncephalartos, to which 
allusion has been made in the introductory remarks on fossil 
Cycadacee. 

V. 2126. 32cm. long. Rachis about 4mm. broad, in the form 
of a round hollow mould. Compare the largest pinna of this 
specimen with the smallest of V. 2122. 


OTOZAMITES. 67 


V. 2740. Pl. VII. Fig. 7. Specimen 37cm. long; 13 pinne 
on one side. The marked variation in length of pinne at the 
upper and lower end of the rachis is well seen. Several segments 
show the manner of insertion on the frond axis; venation fairly 
distinct. The pinne present a strongly convex surface, and are 
slightly inclined to the horizontal plane of the rachis, as in 
Weichselia Mantelli (Brong.), (Vol. i. p. 116). At the upper end 
of the axis the pinne bases are 2°7 cm. apart, towards the lower 
end 2—2°5cm. apart. The upper pinne are practically identical 
with those of V. 2170 (Pl. VIL. Fig. 9), but the shorter, broader, 
and closer pinne of V, 2170 are somewhat broader and closer than 


any in V. 2740. 


V. 27402. 30cm. long. The venation exceedingly well marked. 
Rachis 5 mm. broad, flattened. Pinne vary considerably in size 
at the two ends of the frond, 4°8 x 2°5 to 3°55 x 2:lem. The 
auriculate base of many of the segments clearly shown. For the 
most part the segments overlap one another, oblique to the rachis, 
and apparently of a thick and leathery nature. Cf. O. Beanii. 
Ecclesbourne. Rufford Coll. 


V. 2912. Pl. VII. Fig. 3. Rachis 33cm. in length. Lowest 
pinna 5°7 X 2°7cm.; uppermost 6°5 x 38. Venation in excellent 
preservation. Apices bluntly rounded; the lower edge of the base 
appears to be more distinctly lobed than the upper. Smaller 
pinne like the larger ones in V. 2740, and the larger like those 
of V. 2912, ete. 


V. 2912. This specimen shows six pinne attached to the rachis, 
three on each side; pinne 7 X 2°7cm. broad, do not overlap, 
inclined to the horizontal plane of the rock surface. Rachis 
striated longitudinally, and shows cross lines at intervals, but 
the latter are no doubt simply minute transversely running cracks 
in a mineral substance. Cf. V. 2090, V. 2122, etc. Ecclesbourne. 

Rufford Coll. 


V. 2745. Pl. VII. Fig. 5. Rachis 12 cm. long, with six pinne. 
Pinne short and broad. Compare this specimen with V. 2836 
(Pl. I. Fig. 3) and V. 27450 (Pl. I. Fig. 4). Largest pinne 
2°8 xX 1-6 em., lowest and smallest 1°8%1:4 cm. Venation distinct. 
Cf. upper end of V. 2740c. 


68 OTOZAMITES. 


V. 27454. 12°5 cm. long. Seven pinne on one side: largest 
4-216 cm., smallest 3°8 1:2 em. Rachis appears to be 2°5 mm. 
Venation good, upper pinne slightly auriculate, in the lower seg- 
ments the ariculate base is more distinct. Cf. Pterophyllum 
oblongifolium, Kurr. Kcclesbourne. Rufford Coll. 


V. 2090. Part of a rachis and four pinne. Pinne long, not 
overlapping, about 3 cm. apart, of similar form to those of V. 2912, 
Eeclesbourne. Presented by P. Rufford, Esq., 1889. 


V. 2364. Portion of a single pinna; compare with some of the 
lower pinne in V. 2740a; broad and short, 5 x 3°8cm. Very 
different in appearance from V. 2364, but probably the same 
species; in V. 2122 we have the same kind of divergence in the 
form of the segments as between the two specimens V. 2364 
and V. 23640. Ecclesbourne. Rufford Coll. 


O. Klipsteinii var. longifolia, mihi. 


V. 2122. Pl. VII. Figs. 1 and 6. MRachis 43 em. long; 15 
pinne on one side. The pinne in this specimen are of the 
longer and narrower form. The smallest and lowest segment 
measures 7°5 X 2°6cm.; the uppermost and longest 8-4 X 2 cm. 
The lowest pinna connects such an example as this with V. 2912, 
V. 2090, V. 2740, etc. Pinne separate, bases in the lower part 
of the frond 8:5cm. apart, the two uppermost separated by 
25cm. Venation not quite so distinctly seen as in some other 


specimens; the lower pinne show very clearly the auriculate 
form of base. 


V. 2122). This specimen, 20cm. in length, is very imperfectly 
preserved. The pinne are of the longer and narrower type, 
and agree fairly well with those of Zamites tenuinervis, Font., 
but the distinctly auriculate form of the base and the manner 
of insertion on the rachis seem to sufficiently distinguish the 
specimen from the Potomac species, and to connect it with the 
other forms of Otozamites Klipsteinit var. superba. Possibly it 
would be better to designate such forms as V. 2122, V. 21228, 
V. 212380, etce., O. Alipsteinii var. longifolia. Between this 
specimen and V. 2122 there is the closest similarity. Eccles- 
bourne. Rufford Coll. 


OTOZAMITES. 69 


V. 21230. Here again we have a specimen, imperfectly 
preserved, which it is difficult to assign to a perfectly satisfactory 
position. The pinne are long and narrow, and agree with those 
of V. 21224, etc.; but on the other hand this form is nearer to 
that of Zamites tenuinervis, Font. The bases of the pinne are 
less distinctly auriculate, and in some there is a wrinkling of the 
leaf substance near the point of attachment to the frond axis. 
Cf. Encephalartos Lehmanni. Ecclesbourne. Ruffurd Coll. 


Other specimens referred to this species: V. 1069, V. 2126, 
V. 2364, V. 29124, V. 3160 (a good example of O. AVipsteinii 
var. longifolia). 


Otozamites sp. Cf. O. Klipsteinii (Dunk.). 


V. 2734. Pl. II. Fig. 4. 

In Pl. II. Fig. 4 is represented one of two specimens, of which 
one is the counterpart of the other. The rachis has a length of 
12°5cm., and bears at its upper end six imbricate and obliquely 
attached pinne. The base of the rachis is swollen, and shows 
a clean cut surface by which it was attached to the parent stem ; 
just above the base there are traces of filiform appendages, possibly 
scale or hair structures, such as occur on the lower portions of many 
recent cycadean fronds. The bases of the alternately placed pinne 
fit in between one another, and are attached to the upper surface 
of the rachis. Saporta! has figured a very similar specimen as 
Otozamites sp., in which the base of the petiole shows two fairly 
large leaf-like or stipular(?) structures. The French Jurassic 
specimen closely resembles O. Beanii (L. and H.), but it is no 
doubt better to follow Saporta, and retain it as a fragment too 
small to be accurately determined. Another somewhat analogous 
form is figured by the same author as Otozamites marginatus*; but 
in dealing with portions of such partially developed fronds, any 
attempt to assign them to specific forms founded on nature fronds 
must be attended with no little difficulty and risk. Possibly this 


1 (A. 2), Pal. Frane. vol. ii. pl. vi. figs. 3 and 4. 
2 Ibid. pl. cix. fig. 1. 


70 OTOZAMITES. 


specimen (V. 2784) may be the basal portion of a young partially 
expanded frond of the same species (O. Klipsteiniv), of which 
V. 2336 (Pl. I. Fig. 3) is the apical termination of a young frond, 
but it would hardly be wise to do more than offer the suggestion. 
In V. 2336 the rachis appears to be very slender; here, on the other 
hand, it is distinctly strong and broad: how far this constitutes an 
important difference, and how far it is due to one specimen 
being the terminal portion and the other the basal portion, or to 
differences in the manner of preservation, is difficult to decide. 
The individual pinne are very similar to, and possibly identical 
with, the isolated segment figured by Dunker' and Schenk? and 
now classed with V. 2336. Ecclesbourne. Rufford Coll. 


Otozamites sp. Cf. O. Reibeiroanus, Heer. 


V. 2926. This small specimen shows ? pinne attached to the 
upper surface of a rachis. It may belong to the same species 
as V. 2784 (Pl. Il. Fig. 4), but of this there is no obvious 
proof. It agrees very closely with Heer’s Portuguese species 
O. Reibeiroanus The upper lobes of the pinne are more 
prominent than the lower. Ecclesbourne. Rufford Coll. 


Otozamites Goppertianus (Dunk.). 


[Pl. I. Figs. 1 and 2; Fig. 4.] 


1846. Pterophyllum Géppertianum, Dunker, Wealdenbildung, p. 14, lewis 
fig. 6. 

1848. Pterophyllum Géppertianum, Bronn, Index pal. nomencl. p. 1053. 

1849. Zamites Géppertianus, Brongniart, Tableau, p. 107. 

1850. Pterophyllum Géppertianum, Unger, Gen. spec. plant. foss. p. 290. 

1851. Dioonites Géppertianus, Miquel, Tijdsch. Wis. Nat. Wet. iv. p. 7. 

1852. Pterophyllum Géppertianum, Ettingshausen, Abh. k.-k. geol. Reichs. 
vol. i. Abth. iii. No. 2, p. 21. 

1856. Dioonites Gippertianus, Bornemann, Organ. Rest. Lettenkohl. p. 56. 

1870. Dioonites Géppertianus, Schimper, Trait. pal. vég. vol. ii. p. 161. 

1871. Dioonites Géppertianus, Schenk, Palwontographica, vol. xix. p. 230, 
pl. xxxiv. figs. 3 and 4. 


1 Wealdenbildung, pl. ix. figs. 6 and 7. 
2 Paleontographica, vol. xix. pl. xxxi. fig. 6. 
8 Heer (A. 6), Secc. Trab. Geol. Portugal, 1881, pl. ix. figs. 1-9. 


OTOZAMITES. al 


Type. Specimen of frond. Berlin Museum. 

Dunker thus defined the species in 1846: ‘‘ Pterophyllum 
fronde impari-pinnata, pinnis oppositis subrectis elongatis lneari- 
bus acuminatis subremotis, eque distantibus angulo acuto adnatis, 
nervis obsoletis quinis vel senis, rhachi crassa subtereti levi.” 

He speaks of each pinna having five or six equal and thin 
veins, and shows this venation in the slightly magnified portion 
represented in fig. 5a, pl. ii. Dunker’s figures certainly appear 
to justify his choice of the genus Pterophyllum as regards the 
manner of attachment and distal terminations of the segments. 
Reference has already been made, in speaking of the various 
sources of error connected with fossil cycads, to the very different 
appearance presented by a frond when viewed from the upper 
and lower surface. In the present instance it is quite possible, 
and, indeed, considering all the facts before us, I believe very 
probable, that we have in Dunker’s fig. 5 a view of the under 
surface of the frond. The rachis is seen to project considerably 
above the level of the pinne, and the latter are either attached 
by broad bases to its margin, or pass underneath and are united 
to the face of the rachis which is pressed against the rock 
surface. The venation, as Dunker describes and figures it, does 
not accurately correspond with that of the English specimens 
which I have ventured to refer to this species. If, however, 
Dunker’s drawing represents an imperfect fragment seen from 
below, it is very likely that we should find a somewhat different 
appearance presented by the veins to that which is seen in the 
more perfect pinne of the English fronds. If specimen V. 2360 
(Pl. I. Fig. 2) be compared with Dunker’s pl. i. fig. 5, the 
striking resemblance between them in the form and arrangement 
of the pinne cannot fail to be noticed. The English specimen 
seen from the under side would present an appearance very similar 
to that of Dunker’s frond. Turning to Schenk’s account of the 
same species, we find he follows Miquel in adopting the generic 
name Dioonites in preference to Pterophyllum. Schenk includes 
in this species the specimen figured by Dunker as Pecopteris 
linearis, which the former regards as a badly preserved fragment 
of Dioonites Goppertianus ; this does not seem probable, so far as 
the figures enable us to form an opinion. This author extends 
the original diagnosis of the species, and points out that the pinne 
are attached to the upper surface of the frond axis; his figure 


~I 
bo 


OTOZAMITES. 


shows a median groove in the upper surface of the rachis between 
the two rows of segments. The veins, apparently not very distinctly 
shown, are described as parallel, uniform, and delicate. Schenk’s 
fig. 8, pl. xxxiv. probably represents, as in Dunker’s fig. 5, 
the under surface of a frond, but in fig. 4 of Schenk’s plate we 
have a small piece of leaf seen from above, and here the median 
groove and manner of insertion of the segments are indistinctly 
seen. A comparison of V. 2123 (Pl. I. Fig. 1) with Schenk’s 
figs. 4 and 4a reveals a fairly close resemblance; the corre- 
spondence in the general form of the fronds and pinnz suggests 
the same species. In the German specimens we have only small 
and imperfect fragments, but in the Sussex examples the pre- 
servation is particularly good, and the details well marked. The 
pinne of the English specimens do not in every case show 
a distinctly auriculate base; it is only here and there that this 
feature can be seen. Considering the difference in the manner 
of preservation in the two sets of specimens, it is not much to 
be wondered at if no trace of the lobed base can be detected in 
the more imperfect specimens. Another important point is the 
probability, that Schenk’s fig. 4 is the only specimen from the 
German Wealden in which we have a view of the upper surface 
of the rachis. 

A plant from the Lower Cretaceous rocks of Portugal, originally 
figured by Morris! as Zamites gramineus var. munde, and after- 
wards by Heer? as Otozamites angustifolius, shows a certain amount 
of resemblance to the present forms; compare especially Heer, 
pl. ix. fig. 10 and fig. 3d. It must be admitted that the 
English specimens have a broader rachis than is apparent in 
Dunker’s figures, but this may easily be due to the different 
manner of fossilization, and cannot be relied upon as an essential 
difference, considering the nature of the material. Some of 
Saporta’s figures of Otozamites latior, Sap., resemble in general 
characters the Wealden species, but differ in some points of detail 
which sufficiently separate the two forms.’ A comparison of 
Saporta’s pl. xcvil. figs. 1 and 38 with the present specimens 
shows very clearly the different appearance presented by an upper 


1 (8), pl. xxvi. fig. 7. 
2 Heer, loc. cit. pl. ix. 
8 Saporta, pls. xevii. and xcyiil. 


~I 
co 


OTOZAMITES. 


and lower view of the same plant. I am inclined to regard some 
of the examples referred by Saporta to O. datior as identical with 
Heer’s O. angustvfolius, Heer; cf. Saporta, pl. xevii. fig. 2, and 
Heer, pl. ix. fig. 12, ete. The plant figured by Bartholni,’ as 
Otozamites latior, closely resembles the present species. A some- 
what similar habit is also seen in Leckenby’s Otopteris lanceolata: 
the type specimen of this species appears to be identical with 
some other examples which have been assigned to Ctenophyllum 
pectinata; the base of the pinne may be very slightly auriculate, 
but the specimens do not afford satisfactory evidence of this. 

It will be seen from the two specimens (V. 2123 and V. 2360) 
figured in Pl. I. Figs. 1 and 2, that the frond presents a very 
different appearance in the lower and upper portions. The graceful 
and tapering pinne, with their slightly but distinctly auriculate 
base (Fig. 4), and the grooved broad rachis are perhaps the most 
striking specific characteristics. The inclusion of these specimens 
in Dunker’s species may, perhaps, be an error of judgment, but 
I have endeavoured to show on what grounds this course has been 
taken. The institution of a new species would have been in some 
respects more satisfactory than defining afresh an old species 
founded on specimens much less perfect than those now before us, 
but, having recognized the strong probability that the apparent 
differences between Dunker’s type and the English examples is 
due to accidents of fossilization, the original name has been 
retained. We may thus define Otozamites Giéppertianus :— 

Frond pinnate, deciduous; rachis broad, with a surface marked by 
irregular longitudinal lines; pinne alternate, articulate, attached 
by a slightly auriculate base to the middle of the upper surface 
of the rachis; towards the base of the frond the pinne are very 
narrow and short, and farther apart than those attached to the 
middle and upper portion of the frond axis; in certain parts of 
a frond the pinne are approximate and almost imbricate. Lower 
pinne very obliquely attached to the rachis, the upper gradually 
becoming inclined at a greater angle to the axis in passing towards 
the apex of the frond. Pinne linear-lanceolate, very gradually 
tapering from the base to the apex, occasionally somewhat falcate, 
apices acute and slightly symmetrical, the lower margin of a 
segment being sometimes decidedly curved in an upward direction, 


1 Bartholni, pl. iii. fig. 1. 


74 OTOZAMITES. 


the upper edge in some cases almost straight. Narrow elliptical 
scars left on the surface of the rachis where the pinne have 
fallen off. Veins numerous, radiating outwards from the centre 
of the base, but parallel in the greater part of the length of 
each pinne. 


V. 2360. Pl. I. Fig. 2. Rachis 35:°5cm. long; exceedingly 
well preserved at the base, which shows by its clearly cut outline 
the deciduous habit of the frond. Surface wrinklings or irregular 
striations very distinctly marked on the rachis. The figure 
represents only a portion of the specimen; the upper part, not 
shown in the drawing, is practically identical with the portion of 
V. 21238 figured in Pl. I. Fig. 1. The specimen as a whole shows 
very clearly the great contrast between the loosely arranged, very 
narrow, and short basal pinnee; the somewhat more approximate 
and much longer, gradually tapering, and slightly faleate pinne 
in the middle of the frond; and the stiffer, more approximate, 
and broader pinnee towards the apex. ‘The lowest pinna is about 
1 mm. broad, the broadest about 5mm. Some of the pinne have 
a distinctly auriculate base. Cf. Dunker, pl. u. fig. 5. Eccles- 
bourne. Rufford Coll. 


Fic. 4.—Otozamites Goppertianus (Dunk.), (V. 2741). Enlarged. 


V. 2741. Fig. 4. Rachis 18cm. in length. Part of the lower 
portion of a frond corresponding to V. 2360 (Pl. I. Fig. 2). The 
gradual increase in the breadth of the pinne in passing from the 
lower to the upper part of the frond is well marked. Auriculate 
bases very distinct, and venation fairly clear. Ecclesbourne. 


Rufford Coll. 


ZAMITES. UE 


V. 2123. Pl. I. Fig. 1. Rachis 10cm. long; the pinne more 
closely arranged, and resemble on a rather larger scale Schenk’s 
figures of this species. The auriculate character of the bases of 
the segments distinctly indicated, also the median groove in the 
axis of the frond. Numerous fine veins distinct. Cf. the upper 
unfigured portion of V. 2360. 

There is an imperfect specimen of this species in the Museum 
of Practical Geology, Jermyn Street, in which the auriculate 
base and venation characters are fairly well shown. 


Genus ZAMITES, Brongniart. 
[Prodrome, 1828, p. 94.] 


In describing some Japanese examples of the Wealden cyca- 
dean frond named by Ettingshausen' Pterophyllum Buchianum, 
Nathorst? draws attention to certain characters in the pinne 
which, he considers, exclude the species from Pterophyllum or 
Divonites, in which it has generally been placed by other writers. 
He finds that the segments are somewhat narrower towards the 
point of insertion, and apparently laterally attached to the frond 
axis, thus differing in essential features from the above-named 
genera. The splendid series of specimens in the Rufford Collection 
confirms Nathorst’s remarks with regard to the narrowing of the 
piunne towards the point of attachment to the rachis, but on the 
other hand, the English examples show a manner of attachment 
of the pinne more in accordance with the genus Zamuites, the 
pinne being inserted on the upper surface of the rachis, and 
apparently along two more or less distinct lines. The new genus 
Zamiophyllum, proposed by Nathorst for this species as a substitute 
for Pterophyllum or Dioonites, on the ground that the characters 
do not conform in all essential respects with any existing genera, 
cannot well be retained in the light of the more recent facts 
afforded by the large series of English specimens. The genera 
Dioonites and Pterophyllum are quite unsuitable for this species, 


1 (A. 4), Abh. k.-k. geol. Reichs. vol. i. 1852, p. 21, pl. i. fig. 1. 
2 (A. 3), Denkschr. k. Ak. Wiss. vol. lvii. p. 46. 


76 ZAMITES. 


and it would seem that Brongniart’s genus Zamites corresponds 
most closely in general characters with these large Wealden fronds. 

A brief review of some of the numerous definitions proposed 
for Zamites may suffice to emphasize the generally accepted 
characters, justifying at the same time Saporta’s pertinent 
remarks as to the vague sense in which the name has frequently 
been applied. In 1828 Brongniart' proposed Zamites for a few 
fossil fronds showing some points of difference from his other 
genus Zamia; he mentions Z echii, Brong., as an example 
of the former, and describes the pinne as: ‘‘Se recouvraut 
mutuellement et passant sur le pétiole commun; nervures 
divergentes arquées, souvent bifurquées.” 

The name Zamia is afterwards given up as likely to prove 
misleading as regards the relationship of fossil and recent 
leaves, and Zamites is described as characterized by the possession 
of entire pinne, not truncate at the apex, and not decurrent, 
but slightly constricted at the base. Brongniart includes Braun’s 
two genera Podozamites and Pterozamites as two subsections 
of Zamites. Goppert® uses Zamites in a wide sense, and notes 
the resemblance of some species to the genus Hncephalartos 
among recent cycads. ‘The similarity between various fossil 
fronds and species of this living genus has not been sufficiently 
recognized by most palzobotanical authors. Goppert’s definition 
does not restrict Zamites to fronds with basally constricted 
pinne, but includes those with a swollen and auriculate 
base. Pomel,? true to his unfortunate habit of founding new 
genera, proposes Crossozamia for certain species of Zamites, but 
the name has not come into general use. Bornemann* describes 
Zamites as comprising species with a greater or less resemblance 
to the recent Zamias, and defines the genus as follows: ‘‘ Frond 
pinnate, leathery, pinne approximate or distant, from ovate to 
slender and linear in form, contracted at the base, entire or toothed 
on the upper margin, blunt at the top. Veins of uniform size, 
clearly seen on both sides of the pinne, dichotomous. Epidermal 
cells having the same structure as in recent Zamias.” Schenk® 


1 Prodrome, p. 94. 

(1), p. 122. 

Pomel, p. 342. 

Bornemann, p. 54. 

(A. 1), Fl. foss. Grenz. Keup. Lias, p. 158. 


ao -§ © WN 


— 


ZAMITES. af 


gives a diagnosis of Zamites in which the veins in the pinne are 
described as equal and parallel, and dichotomously converging 
towards the apex. He thus includes such forms as correspond 
to Braun’s genus Podozamites. In Zittel’s Handbuch’ we find a 
closer approximation to the definition of Zwmites as generally 
accepted: the pinne are described as attached to the upper surface 
of the rachis, and possessing a basal callosity, rounded or some- 
what contracted at the base; the median veins parallel, and the 
outer veins diverging towards the apex of the pinna. 

Solms-Laubach? notes the occurrence in the Zamite of pinnee 
which articulate with and separate from the frond axis, pointed 
at the apex, abruptly rounded at the base, and attached obliquely 
to the rachis, which they overlap and cover. Solms pertinently 
remarks that isolated segments of species of Zamites may easily 
be mistaken for portions of other fossil genera. Before attempting 
to modify in any way the definitions of Zamites, it may be well 
to consider some of those special features which have been 
referred to by several writers, and to see how far such details 
of frond structure are likely to serve as trustworthy guides. As 
regards the basal callosity usually included in definitions of this 
genus, we must acknowledge the great difficulty to be experi- 
enced in deciding definitely as to its existence in many fossil 
leaves. In the process of fossilization the pinne of a cycadean 
frond are often flattened down against the rock, and closely 
adpressed to the surface of the rachis, and thus there may be 
produced transverse wrinklings just above the point of attachment 
of the segments; such wrinklings may easily suggest in a fossil 
specimen the original existence of a basal callosity. The venation 
may prove useful in determining certain species, but it is not 
very often that fossil specimens are sufficiently well preserved to 
admit of a complete diagnosis of the venation character. If, for 
example, we had neither the basal nor the apical portion of a 
pinna, it would be practically impossible to discriminate between 
some of the long narrow Otozamites pinne, and those of Zamites 
and other genera. 

Nathorst’s recently proposed genus Zamiophyllum, to which 
reference has already been made, was founded partly to include 


1 Zittel (A.), p. 218. 
* Fossil Botany, p. 88. 


78 ZAMITES. 


fronds of which the obliquely inclined pinne have a somewhat 
narrowed base. This decrease in breadth towards the point of 
attachment of a pinna is characteristic of numerous species of 
Zamia, Ceratozamia, and Macrozamia, and might well be included 
in a more extended definition of Zamites. The basal contraction 
of the segments is, as Nathorst points out, directly opposed to 
the accepted definitions of Dioonites and Pterophylium; but there 
seems no good reason to regard such a feature as at variance 
with the genus Zamites. This name Zamites was proposed by 
Brongniart at a time when only two genera of living cycads 
had been recognized, Cycas and Zamia; and the present definition 
of the genus stamps it rather as a comprehensive and provisional 
designation for certain frond characters which are now shared 
by various members of the recent Zamice. If we retain Zamites 
as usually defined, or in a slightly modified form, it must be 
regarded merely as a convenient term to be applied to certain 
fossil fronds in which some of the characteristics of Zama, 
Macrozamia, Ceratozamia, or even Encephalartos may be repre- 
sented. In the recent Zama the pinne are articulated to the 
rachis, and in many forms are readily detached, leaving a distinct 
circular or elliptical scar; this same character is also met with 
in other genera, such as Ceratozamia, Encephalartos, etc. The 
basal callosity often referred to as one of the important charac- 
teristics of Zamites is best seen in some forms of the genus 
Macrozamia, and the manner of insertion and position of the 
pinne on the rachis in species of Zumites find a parallel living 
in species of Encephalartos, Ceratozamia, Macrozamia, and Zamia. 

The following general definition of Zamites may serve to 
indicate those characters which are most readily recognized in 
fossil fronds :— 

Frond pinnate, pinnz more or less obliquely inclined to 
the rachis and attached to the upper surface, apices acuminate 
and tapering, or obtusely rounded, the base may be abruptly 
rounded and marked with a eallosity near the point of attach- 
ment, or the pinne may be slightly and gradually narrowed 
towards the base, margins entire; veins parallel, but slightly 
divergent in the apical portion of each pinna. 

Such a definition is perhaps suggestive of a genus with 
characters expressed in too general terms, and not sufficiently 
limited, but a more complete examination of the different types 


- 
ZAMITES. 19 


of fossil leaves may possibly lead to the institution of other genera 
with more narrowly defined characters. 


Zamites Buchianus'’ (Ett.). 
Bele Te Bios 1-6: PL TV. 3 Pl. Vilieac] 


1847. Pterophyllum saxonicum, Goppert, Nova Acta Ac. Czs. Teop.-Car. 
vol. xxii. p. 362, pl. xxxviii. fig. 13. 
1852. Pterophyllum Buchianum, Ettingshausen, Abh. k.-k. geol. Reichs. vol. i. 
Abth. in. No. 2, p. 21, pl. i. fig. 1. 
1856. ? Dioonites Buchianus, Bornemann, Organ. Rest. Lettenkohl. p. 57. 
1867. Pterophyllum saxonicum, Httingshausen, Sitz. k. Ak. Wiss. Wien. 
vol. lv. Abth. i. p. 11, pl. i. figs. 11 and 12. 
1870. Dioonites Buchianus, Schimper, Trait. pal. vég. vol. ii. p. 149. 
Dioonites saxonicus, Schimper, ibid. p. 211. 
1871. Zamites Gopperti, Schenk, Paleontographica, vol. xix. p. 11, pl. iii. 
fig. 6. 
1879-80. ? Dioonites abietinus, Hosius and von Marck, Paleontographica, 
vol. xxvi. p. 213, pl. xliv. fig. 199. 
1889. Dioonites Buchianus, Fontaine, Potomac Flora, p. 182, pls. lxviii.—Ixxiv. 
Dioonites Buchianus var. angustifolius, Fontaine, ibid. p. 185, pls. 
Ixvii., Ixviii., and Ixxi. 
Dioonites Buchianus var. obtusifolius, Fontaine, ibid. p. 184, pl. clxviii. 
fig. 3. 
1890. Zamiophyllum Buchianum, Nathorst, Denkschr. k. Ak. Wiss. Wien. 
vol. lvii. pp. 46 and 49, pl. ii. figs. 1 and 2; pl. iii. ; pl. v. fig. 2. 
Zamiophylium Naumanni, Nathorst, ibid. p. 47, pl. v. fig. 1. 
1894. Zamiophyllum Buchianum, Yokoyama, Journ. Coll. Sci. Japan, vol. vii. 
pt. iii. p. 223, pl. xx. fig. 1; pl. xxiii. fig. 6; pl. xxvii. fig. 5a, 4; 
pl. xxviii. figs. 1 and 2. 
Zamiophyllum Buchianum var. angustifolia, Yokoyama, loc. cit. p. 224, 
pl. xxii. fig. 4; pl. xxv. fig. 5; pl. xxviii. figs. 8 and 9. 
Zamiophyllum Naumanni, ibid. p. 225, pl. xxii. fig. 3; pl. xxvi. 


Type. Portion of frond. Collection of Herr Hohenegger, Teschen.? 
Ettingshausen defined the species as follows: ‘P. fronde pinnata, 
pinnis circa 1-2dm. longis, 4-7 mm. latis, alternis, linearibus, 
subremotis, subangulo acuto adnatis, nervis creberrimis, tenuissimis 
instructis; rhachide crassiuscula.”’ 

The type specimen is described as the middle portion of a frond 


1 Possibly Zamites Milleri, Zigno, may prove to be identical with this species 
(Zigno, Flor. foss. Oolit. vol. ii. p. 40, pl. xxx. fig. 6; Hugh Miller, Testimony 
of the Rocks, p. 434, fig. 136). 

2 Ettingshausen (A. 4), p. 32. 


80 ZAMITES. 


at least 8-4 ft. in length, and compared with Pterophyllum 
Humboldti, Dunk.; the latter species, however, appears to be a 
typical Pterophyllum species, and quite distinct generically from 
Ettingshausen’s type. Schenk reproduces the figure given by 
Ettingshausen, and points out the fact that none of the pinna 
apices are shown in the specimens. Very probably we may regard 
Zamites Gépperti, Schenk, as Z Buchianus (Ett.), seen from the 
lower surface; Schenk himself compared the former species with 
Pterophyllum saxonicum, Reich.,! from Niederschonen, and there 
seems good reason to follow Fontaine in including P. saxonicum, 
as figured by Ettingshausen, as synonymous with Z. Buchianum. 
Hosius and von Marck? have figured a small fragment of a frond, 
which they refer to P. saxonicum, but the specimen is too im- 
perfect to admit of accurate identification. The same authors 
refer another specimen of cycadean frond to Dvoonites abietinus, 
which shows a distinct resemblance to the smaller forms of Zamites 
Buchianus (ef. Hosius and Marck, pl. xliv. fig. 199, and Pl. III. 
Fig. 1 of the present volume), but perhaps the similarity is hardly 
sufficiently well marked to warrant the inclusion of the fossil in 
the synonym list of the present species, without the addition of 
a query. Some writers have preferred Dioonites to Pterophyllum 
for Ettingshausen’s species; Schimper and Fontaine both adopt 
the former name. The Potomac flora has yielded numerous 
examples of fronds which Fontaine refers to D. Buchianus; he 
speaks of the species as ‘‘one of the most widely diffused and 
characteristic fossils of the Potomac flora.” * This author institutes 
two varieties of Ettingshausen’s species—D. Buchianus, var. 
obtusifolius and var. angustifolius. It might perhaps be advisable to 
adopt Fontaine’s terms, and apply them to certain forms of the 
species represented in the numerous examples from the Wealden 
of Ecclesbourne, but there is the usual difficulty to be faced in 
drawing lines between one form and another. In looking at some 
specimens we find the pinna apices are very distinctly acutely 
tapered, and closely correspond with Fontaine’s D. Buchianus var. 
angustifolius ; but in such a frond as V. 21787, although on the 
whole the pinne are tapered, yet some of the segments terminate 


1 Ettingshausen (A. 8), Sitz. k. Ak. Wiss. Wien. vol. lv. Abth. i. pl. i. 
figs. 11 and 12. 

2 (A. 1), Paleontographica, vol. xxvi. p. 213, pl. xliv. fig. 198. 

3 Potomac Flora, p. 182. 


ee, 


ZAMITES. 81 


more obtusely, and lead us to specimens, such as V. 2227, in 
which we have no longer the form characterized by gradually 
tapered and pointed pinne. A similar variation in the apices and 
in the breadth of the pinnz is pointed out by Yokoyama! in the 
Japanese examples. The frond figured by Goppert as Pterophyllum 
Carnallianum* shows a certain resemblance to some of the forms 
of the present species. Heer* has compared the present species 
with Zamia globuliferus, Heer, from the Kome beds of Greenland. 
Reference has already been made to Nathorst’s substitution of the 
new genus Zamiophyllum for Miquel’s name Dioonites. He points 
out that the species is from Urgonian strata, and not of Wealden 
age as Ettingshausen erroneously states. In referring to the basally 
narrowed pinne in the Japanese exampies, Nathorst notes that 
the character is not apparent in the European specimens; we still 
find, however, in the examples described from the Rufford Collection 
abundant proof of this narrowing of the segments. The largest 
pinna mentioned by this author has a length of 260mm. and 
a breadth of 4-6mm.; in a pinna which is 3°5 mm. broad there 
are 12 veins, in one with a breadth of 6mm. 17 veins, and 22 
veins in pinne 8mm. broad. Nathorst compares Z. Buchianus 
with the recent cycad Zamia media, Jacq. In describing a second 
Japanese form, which he refers to another species, 7 Maumannt, 
he expresses the opinion that it may possibly represent the lower 
part of a frond of Z Buchianus*; it is compared with Zamites 
Schenkii, Schimp. (=Z. Gépperti, Schenk), from the Wernsdorf 
beds. Yokoyama also figures under Nathorst’s species Z. Vaumannt 
(pl. xxi. fig. 3, and pl. xxvi.) a portion of a frond with pinne 
having a breadth up to 20mm., but suggests that it may be 
specifically identical with Z. Buchianus. 

The English examples clearly demonstrate that the pinne are 
attached to the surface of the rachis, and not laterally as Nathorst 
inferred from his less perfect material. Z. Naumanni, Nath., 
is in all probability identical with Z Buchianus; the specimens 
of the former figured by Nathorst seem in some cases, ¢.g. pl. ili. 
and pl. v. fig. 1, to show a surface attachment of the pinne. 


1 Yokoyama, p. 2238. 

2 (1) pled tie. 4. 

3 (A. 3), FI. foss. Arct. vol. vi. pt. i. p. 12, pl. i. 
* (A. 8), Denkschr. k. Ak. Wiss. vol. lvii. p. 47. 


82 ZAMITES. 


In the specimen represented in pl. v. fig. 1 of Nathorst, there 
appear to be two elliptical scars about the middle of the frond 
fragment; these agree very closely with similar scars in several 
of the Rufford examples, and represent the points of detachment 
of pinne. The abundant examples in the National Collection 
enable us to extend and modify the existing definition of the 
species. Among recent species Ceratozamia mexicana, Brong., 
Zamia cycadifolia, Jacq., Macrozamia Macleayt, Miq., and Encepha- 
lartos Lehmanni, Lehm., may be mentioned as possessing fronds 
very similar in form to those of Z. Buchianus (Ett.). 

We may define the species as follows :— 

Frond large, pinnate; rachis longitudinally striated, with a 
fairly broad median groove on the upper surface. Pinne 
alternate, opposite or subopposite, varying in length from 38-4 cm. 
to 20cm., and in breadth from 1:5 to 2mm., attached obliquely to 
the rachis, and slightly thickened and somewhat broadened at the 
actual point of insertion; separated from the rachis by a distinct 
absciss line, leaving an elliptical scar; generally narrowed towards 
the base, but in the narrower pinne this reduction in breadth 
decreases and is not nearly so evident; usually inclined at about 
45° to the rachis, they may be almost at right angles to the frond 
axis, and in the case of young fronds and the apical portions 
of larger ones, the pinne are attached at a much more acute angle. 
The distance between the pinne varies considerably in different 
parts of the same leaf, and in leaves of different ages; apices 
generally tapering to a point, or more or less obtusely rounded. 
Veins numerous, parallel, and as a rule not very prominent. 

V. 2120. Pl. VIII.’ Fig. 1 and Pl. III. Fig. 5. This splendid 
specimen has a length of 77°5cm.; broken at one point, and 
somewhat displaced laterally. Breadth of pinnee varies from 2 to 
9mm. Attachment of the pinne not shown in this example. 
The broader and lower pinne appear to taper towards the rachis 
more than in many specimens. ‘Tips of some of the pinne clearly 
seen, as in Pl. III. Fig. 5; these agree very closely with the 


apices of pinne in V. 21238 (Pl. III. Fig. 3). Certain parts of » 


this frond agree exactly with V. 2925, etc. The apical portion 
appears to be identical with Dvoonttes Buchianus var. angustifolia, 
as figured by Fontaine in his pl. lxx. fig. 2.2, The lower portion 


1 Represented in Pl. VIII. § nat. size. 
* Potomac Flora. 


— ee ee 


ZAMITES. 83 


corresponds to that of V. 2123a, etc. Veins numerous, but not 
well marked. Towards the upper end of the rachis there are 
elliptical scars marking the original places of insertion of detached 
segments. If V. 2120 and V. 2898 be compared, the most striking 
difference seems to be the less tapered and narrowed bases of the 
pinne in the latter specimen. Ecclesbourne. Rufford Coll. 


V. 2227. Pl. III. Fig. 4. Large specimen, with unusually 
broad pinne; length of rachis 388cm., width 1:2cm. The 
oblique attachment of the segments distinctly shown, also the 
nature of the base and the position of the pinne on the axis. 
Surface of the rachis marked with longitudinal strize; about 
1mm. from each margin of the rachis there is a distinct longi- 
tudinally running line. Pinne long, narrow, and slightly curved ; 
one row attached obliquely by tapering bases to the upper surface 
of the frond axis, alternate in position. In the lower part of the 
frond the pinne have a length of about 19cm., and a breadth 
of 1:3cm.; their distal ends are bluntly acuminate, as shown 
in Pl. III. Fig. 4. Very little, if any, indication of a basal 
callosity on the segments. There is only a slight difference in 
size between the upper and lower pinne. Cf. V. 2820. Eccles- 
bourne. Rufford Coll. 


V. 2820. A very similar specimen. Rachis 29cm. long; the 
longest pinna between 17 and 18cm. in length and 1 cm. broad. 
Towards the upper end of the frond the pinne are about 10 or 
11cm. long; the apices are not shown, and therefore the actual 
length can only be approximately measured; breadth of these 
pinne about 8mm. LEcclesbourne. Rufford Coll. 


V. 2125a. Rachis 21cm. long. The upper portion closely 
resembles Dioonites abietinus as figured by Hosius and von Marck. 
Ecclesbourne. Rufford Coll. 


V. 2128. Portion of a rachis of a large frond; the bases of 
detached pinne well preserved, also traces of scars on the rachis. 
Ecclesbourne. Rufford Coll. 


V. 2363. Pl. III. Fig. 2. The chief feature in this specimen 
is the very distinctly and gradually tapering terminations of the 
few pinne which are preserved. Compare the apex of one of 
these pinne, with those of the two shown in Pl. II. Fig. 4 


84 ZAMITES. 


(V. 2120), and the less pointed form of V. 2128¢ (PI. IIT. Fig. 3). 
Ecclesbourne. Rufford Coll. 


V. 2128c. Pl. III. Fig. 3. At the upper end of the specimen 
there is part of a rachis with five pinne attached; the rest of the 
slab shows several imperfect pinne with gradually tapered tips; 
the tips seem to be intermediate in form between those of 
Ve 2227) (Pla ie. 4) sand ~V. 2363 (Pl. LLL. aie 22): 
V. 2123, etc. Ecclesbourne. Rufford Coll. 


V. 2125). Pl. IV. Fig. 1. A young frond, 25cm. long; 
cf. V. 21230, etc. Pinne approximate, very gradually tapering 
distally. The pinne are broadest in the middle of the specimen, 
and narrower towards either end; at the lower end the pinne 
are narrower and less closely arranged, agreeing with those of 
V. 2262 (Pl. Ill. Fig. 1). Compare also V. 2898 and Fontaine’s 
pl. Ixxi. fig. 2. Ecclesbourne. Rufford Coll. 


V. 2125c¢. Pl. IV. Fig. 2. Rachis 33cm. long.  Pinne 
alternate and opposite, rachis depressed, the points of attachment 
of segments clearly shown, apical portions not preserved ; broadest 
pinna 7mm., narrowest 5mm. Venation distinct. Cf. V. 21238a, 
V. 21252, etc. Ecclesbourne. Rufford Coil. 


V. 2128a. Pl. IV. Figs. 4 and 5. 35cm. long. Rachis well 
shown, the longitudinal irregular striations distinct; these pro- 
bably indicate strands of hypodermal sclerenchymatous tissue. 
Tips of pinne not seen, but bases very distinct ; in Figs. 4 and 5 
the surface and oblique insertion is easily recognized, also in some 
pinne the very slightly broadened base. In the lower part of 
the frond the pinne are nearly at right angles to the rachis, but 
more oblique towards the upper end. Veins numerous and fine, 
and in many pinne clearly shown. LKeclesbourne. LRufford Coll. 


V. 2925. Pl. IV. Fig. 3. Specimen very similar to V. 2125a; 
but pinne farther apart, and in this respect identical with the 
lower portion of V. 2123a. In addition to the larger example, 
there is a very small fragment on one side of the slab, which is 
evidently the tip of a frond (Pl. 1V. Fig. 3); in this the segments 
are closer together and more oblique to the rachis than in 
V. 21254, and more like V. 21284 and V. 21254. Ecclesbourne. 

Rufford Coll. 


ZAMITES. 85 


V. 2262. (Pl. III. Fig. 1.) This agrees fairly closely with 
Dioonites abietinus (Gopp.) as figured by Hosius and von Marck?; 
probably their specimen is not a true example of Gippert’s 
species. Small piece of rachis with long and narrow pinne ; mode 
of insertion well seen. Cf. 21254; breadth of pinne about the 
same, but in the present specimen the segments are farther apart 
and more spreading; this difference, however, only applies to the 
upper three-fourths of 21254; in the lower fourth it is practically 
identical with V. 2262. This and some other specimens appear to 
be identical with D. Buchianus var. angustifolia as figured in the 
Potomac Flora, and with some of Yokoyama’s specimens, e.g. 
pl. xxii. fig. 4, pl. xxv. fig. 5, and pl. xxviii. figs. 8 and 9.? 
Cf. V. 2898, V. 1069, and the upper part of 2125a. Longest 
pimna 7°5cm., and 3mm. broad. LEcclesbourne. Rufford Coll. 


V. 720. Two specimens. Fragments of a young frond; pinne 
very obliquely set and approximate. Hastings. Dawson Coll. 


V. 1069. Small specimen, probably the lower part of a frond, 
the pinne being much narrower in the lower than in the upper 
portion ; manner of attachment of segments distinct. Of. V. 2262, 
V. 21250, V. 21230, etc. Ecclesbourne. Rufford Coll. 


V. 2123. Very imperfect example. Bases of several pinne 
show clearly the manner of attachment to the rachis; at the base 
of some of the pinn a wrinkled appearance suggests the existence 
of a callosity. There is not quite the same gradual narrowing of 
the pinne towards the frond axis as in V. 2227 and many other 
specimens. Pinn long, and tapered to an acuminate tip, as in 


V. 2363 (Pl. III. Fig. 2), ete. 


V. 2123. Rachis about 27 em. long; pinne 4 mm. broad, alternate 
and approximate. Cf. V. 2898. The lower part of this specimen 
appears to be identical with the upper portion of V. 2125, also with 
V. 720. Evidently a young frond. 


V. 2123d. Part of a single pinne, showing sharply pointed apex; 
venation not distinct. Ecclesbourne. Rufford Coll. 


1 (A. 1), Paleontographica, vol. xxvi. pl. xliv. fig. 199. 
* Yokoyama, Joc. cit. 


86 ZAMITES. 


V. 2125. Rachis 19cm. in length. Longest pinna 12cm., 
gradually tapering towards the tip, which is not present. The 
long, narrow, and tapering segments of this specimen are exactly 
the same as those in the smaller examples, V. 2262 and V. 1069. 
Cf. Fontaine,! pl. lxxiv. figs. 1-8, ete. Ecclesbourne. 

Rufford Coll. 


V. 28732. Broad rachis, 1:2 mm. at the lowest end, but obviously 
flattened. Groove in the middle, and on each side of the groove 
there are two elliptical scars marking the places to which segments 
were attached; some of the segments appear to have a basal 
callosity ; pinnz opposite or subopposite. Cf. Nathorst’s figure 
(Pl. V. Fig. 1a)? of the rachis of what he calls Z. Naumann. 
Ecclesbourne. Rufford Coll. 


V. 2698. Very large specimen, but the details not well 
preserved. Rachis 43cm. long; pinne of the gradually tapered 
form, as in V. 2363, etc. The upper portion the same as V. 2898, 
V. 21254, etc. Breadth of uppermost pinna 1:5-2mm.; that of 
the lowest lem. This appears to be a frond seen from the lower 
surface, the pinne being, therefore, apparently laterally attached 
to the rachis. Kcclesbourne. Rufford Coll. 


V. 2898. A fine example of what is probably a partially 
expanded frond. Rachis about 20cm. long. Some pinne about 
14cm. in length and 3-4mm. broad; towards the tip the pinne 
are crowded and imbricate, with a width of about 3mm.; towards 
the base 6mm. broad. Sussex. Beckles Colt, 

Cf. the tip of this and V. 2120, V. 21253, etc. V. 3153, 
V. 3154, V. 3156, and V. 3157. Other specimens of this species. 
Ecclesbourne. Rtufford Coll. 


Zamites Carruthersi, sp. nov. 
[Pl. VI. Figs. 2-4.] 


Type. Portions of fronds. British Museum. 

Frond pinnate; rachis longitudinally striate; pinne alternate, 
attached by an oblique base to the outer part of the upper surface 
of the axis, almost at right angles to the rachis in a fully 


1 Fontaine, Potomac Flora. 
2 Denkschr. k. Ak. Wiss. vol. lvii. 


ZAMITES. 87 


developed frond, linear or linear-elliptical, somewhat abruptly 
narrowed towards the base, but slightly broadened at the actual 
base of attachment, separate from the rachis by a well-marked 
articulation. Veins divergent from the base, but for the most 
part parallel, and diverging at the tip of the segments, which is 
bluntly rounded. 

In the examination of the fossil fronds in the National 
Collection, and the comparison of them with the leaves of 
existing species, I have often been struck with the close corre- 
spondence between certain forms of Wealden fronds and species 
of the genus Encephalartos. An inspection of a thoroughly 
representative collection of the different forms of Encephalartos 
fronds, such as are to be found in the exceptionally good 
collection of living and herbarium specimens at the Royal 
Gardens, Kew, tends to very considerably widen one’s conception 
as to the characters of this recent genus of cycads. ‘The 
species generally met with in collections are those in which 
the pinne are broad and more or less spiny; but the genus 
includes various other forms with pinne of quite a different 
form, which often bear a striking resemblance to various fossil 
fronds. As examples of the diversity of leaf form to be met 
with in this genus, the few following species may be cited as 
representatives of some of the forms assumed: LZ. Caffer, Miq., 
with its stout oval pimne, with or without marginal teeth, 
E. pungens, Lehm., with long, narrow, and acuminately terminated 
segments, /. cycadifolius, Lehm., and E#. Ghellinckii, Lem. (Pl. 
XIII. Figs. 8-5), characterized by the long and narrow pinne, 
which form a striking contrast to the broader and better known 
segments of £. horridus, Lehm., and other species. Some of the 
species, e.g. EL. Lehmanni, Lehm., £. cycadifolius, Lehm., and others 
have a close resemblance to some species of Ceratozamia and Zamia. 

In the introductory remarks mention was made of the institution 
by Fontaine of a new genus, Hncephalartopsis, for certain isolated 
pinne from the Potomac beds of North America; and such a genus 
might serve a useful purpose if founded on more satisfactory 
material, but as at present defined it can have but little value. 
Possibly the institution of such a genus, with a wide and modified 
definition, might prove a valuable addition to our list of fossil 
genera, but for the present it will perhaps be better to fall back 
on the old and comprehensive Zamites. I have ventured to 


88 ZAMITES. 


connect the name of Mr. Carruthers with the present species of 
cycadean frond, as a slight recognition of his valuable contributions 
to our knowledge of the fossil Cycadacee. 

Some of the large detached pinne figured by Fontaine as 
examples of his new species Zamites tenuinervis, agree fairly closely 
with those of the present form, but in the Potomac plant the 
venation appears to be coarser, and the bases of the segments 
usually ‘‘abruptly subcordate” ; in the pinna shown in Fontaine’s 
pl. lxxvi. fig. 7 the base seems much more like that in the English 
specimen. The paucity and imperfect character of the Potomac 
material, and the differences already alluded to, hardly warrant 
the adoption of Fontaine’s name for the English forms. There is a 
close resemblance between the present specimens and some of those 
referred to Z. Buchianus, e.g. V. 2123, but in the latter species 
the longer and more gradually tapering piune are sufficiently 
characteristic to distinguish the two forms. Among recent cycads, 
Encephalartos longifolius, Lehm., is one of those which resemble 
very closely in habit the fronds of Z Carruthersi. As examples 
of other fossil fronds to be compared with this species, Zamites 
affinis, Schenk,’ and Palgozamia recta, Tate,” may be mentioned. 


Vi 2lesd-. Pl Vi ie 4. 

In this specimen the manner of attachment of the pinne is 
clearly shown; the line of separation being particularly distinct 
at the base of the middle pinna of the portion of frond represented 
in Fig. 4, Pl. VI. Rachis at least lem. broad, and marked with 
fine longitudinal lines. Venation very distinct, as in V. 2128c. 
Only a portion of the specimen shown in the figure; rachis 
21cm. long. LEcclesbourne. Rufford Coll. 


V. 21286: Pl. Yi. ‘Figs: 2 and 3. 

The pinne are very like those with blunt apices which have 
been included in Z. Buchianus (e.g. V. 2227), but in the present 
specimen the base and manner of attachment of the pinne con- 
stitute the special features. The form of base clearly seen in 
Fig. 8, and the blunt apex with the slightly divergent vems in 
Fig. 2. Rachis in this specimen 13 cm. in length, with portions 
of nine pinnee on one side. Eeclesbourne. Rufford Coll. 


1 Paleontographica, vol. xix. pl. iii. fig. 6. 
2 Tate (A.), Quart. Journ. Geol. Soc. vol. xxiii. pl. y. 


ZAMITES. 89 


The specimens referred to as Otozamites Klipsteindi var. longi- 
folia should be compared with Zamites Carruthersi; e.g. V. 2128a, 
V. 2122, and V. 21224; but in those and similar specimens the 
pinne have a more or less distinctly auriculate base. V. 212380. 
Single pinna. Of. V. 2123d. Ecclesbourne. Rufford Coll. 


Specimens of Doubtful Position. 


V. 2742. Pl. VI. Fig. 1. This may perhaps be the terminal 
portion of a frond of Z. Carruthersi, or possibly of Otozamites 
Klipsteinit var. longifolia, but it is very difficult to feel any great 
confidence in placing it in such a position. Ecclesbourne. 


Rufford Coll. 


a 
AN) 


\\ WY, if 
\\ W/ 


Fig. 5.—? Zamites sp. (V. 2744). 2 nat. size. 


V. 2744. Fig. 5. This specimen and V. 2748a suggest portions 
of a frond very similar to Ctends falcata, L. and H., but no 
indication of anastomosing veins has been detected in the pinne 
of these Wealden examples. The portion of frond shown has 
a length of 13°5cm.; one of the broadest pinne is 7mm. in 
breadth, and is traversed by about ten veins (Fig. 5a). To some 
extent the specimen reminds one, as regards general habit, of 


90 ANOMOZAMITES, 


Zamites Buchianus, but the venation and decurrent pinne are 
distinctive features in the former, and the pinne are more 
oblique than in Kttingshausen’s species. Ecclesbourne. 

Rufford Coll. 


V. 2275. Another terminal piece of frond, agreeing closely 
with V. 2748, and very possibly the same species, but the 
specimen shows no details, being merely a brown stain on the 
surface of a coarse grit. It is possible that these two specimens 
may belong to the terminal portion of a frond of which the 
older and larger segments are shown in Pl. VII. Figs. 1, 4, and 6 


(V. 2122, V. 21262). Ecclesbourne. Rufford Coll. 


V. 27482. In some respects not unlike Nathorst’s figure of 
Zamiophyllum Naumanni, a species of Japanese frond now referred 
to Zamites Buchianus. The present specimen is in all probability 
part of a frond seen from the under side. Cf. Ctenophyllum latv- 
folium, Font., a plant which Fontaine’ refers for no very obvious 
reason to Schimper’s genus Ctenophyllum. V. 3188. Fragment of 
the same form. Ecclesbourne. Rufford Coll. 


Genus ANOMOZAMITES, Schimper. 


[Trait. pal. vég. vol. ii. 1870, p. 140.] 


In discussing the genus WVilssonia mention was made of. 


Schimper’s genus Anomozamites, which he instituted for certain 
Pterophyllum-like leaves possessing the following characteristics : 
‘Folia speciosa, mediocria, vel parva, elongata-oblonga vel 
elongato-linealia, pinnatisecta, hic illic (juniora) integra, nervis 
rhachi perpendicularibus, parallelis, simplicibus vel e basi dicho- 
tomis; pinnis inequalibus, rectangulis, membranaceis vel tenui- 
coriaceis.”’ 

No mention is made in this definition of the place of insertion 
of the segments, whether lateral or on the upper surface of the 
leaf axis. Nathorst has since given special prominence to the 
manner of attachment of the segments as the chief distinguishing 
character between the present genus and WVilssonia;* the same 
author has also instituted a new genus for the reception of 


1 Potomac Flora, p. 176, pl. Ixvii. figs. 2 and 3. 
2 See ante, p. 62. 


ee 


" 
a ae 


ANOMOZAMITES. 91 


Anomozamites-like leaves with dichotomizing veins. We must, 
then, somewhat modify Schimper’s original diagnosis, and the 
following may be adopted as a rough definition of this provisional 
genus Anomozamites; it is a slightly altered version of that in 
Zittel’s Handbuch.) 

Frond comparatively small, linear or tongue-shaped, and usually 
divided into segments which present a more or less obvious 
difference in size, separate or confluent at the base, attached 
laterally to the rachis, and never entirely covering the upper 
face of the frond axis; the segments bluntly rounded or truncate 
distally; veins simple and parallel, generally at right angles to 
the rachis. 

The examples of this genus possess, as a rule, a characteristic 
habit which marks them off from the pinnate fronds of Ptero- 
phyllum with their equal and longer segments. It is difficult in 
some cases to distinguish between the genera Anomozamites and 
Nilssonia. In the former the segments are sometimes attached 
to the rachis in such a manner as to suggest the surface insertion 
of Wilssonia, but there is always some part of the frond axis 
exposed to view, whereas in JWVilssonda the lamina appears to be 
continuous from one side to the other. It is not easy in the 
present instance to decide whether the genus Pterophyllum or 
Anomozamites is the more suitable; both are purely provisional 
genera, and it is not a matter of very great importance which 
term is adopted. 


Anomozamites Lyellianus (Dunk.). 
[Fig. 6.] 
1846. Pterophyllum Lyellianum, Dunker, Wealdenbildung, p. 14, pl. vi. figs. 
1 and 2. 
1848. Pterophyllum Lyellianum, Bronn, Index. pal. nomencl. p. 1056. 
1849. Zamites Lyellianus, Brongniart, Tableau, p. 107. 
1850. Pterophyllum Lyellianum, Unger, Gen. spec. plant. foss. p. 290. 
1851. Dioonites Lyellianus, Miquel, Tijdsch. Wis. nat. Wet. iv. p. 205. 
1852. Pterophyllum Lyellianum, Ettingshausen, Abh. k.—k. geol. Reichs. 
vol. i. Abth. iii. No. 2, p. 22. 
1856. Dioonites Lyellianus, Bornemann, Organ. Rest. Lettenkohl. p. 56. 
1870. Pterophyllum Lyellianum, Schimper, Trait. pal. vég. vol. ii. p. 137. 
1871. Pterophyllum Lyellianum, Schenk, Paleontographica, vol. xix. p. 230 
pl. xxxiv. figs. 1 and 2. 


’ 


ISpip224- 


92 ANOMOZAMITFS. 


Type. Uarge specimens of frond. Berlin Museum. 

Dunker thus defines the species in his Vealdenbildung :— 

“‘Pterophyllum fronde pinnata, pectiniformi, pinnis oppositis 
linearibus que distantibus, approximatis basi fere confluentibus, 
apice obtusis, angulo recto adnatis, nervis iv. vel v. tenerrimis, 
rhachi plana subsuleata.”’ 

The figure of Dunker’s type specimen is very much more sug- 
gestive of the genus Pterophyllum, than are the drawings given by 
Schenk, or that of the solitary specimen in the Rufford Collection. 
Probably the original specimen is part of a frond seen from the 
under surface, thus showing a particularly prominent rachis. It 
may be that the English fossil should be placed in a new species, 
but the apparent differences which distinguish it from Dunker’s 
specimen may be merely such as are the result of a more fully 
developed condition of frond in the latter case. Possibly Zamites 
e@qualis, Dunk.,! should be included in this species. Schenk’s 
figure agrees more closely with the English specimen: this author 
speaks of the segment as being attached to the upper surface of 
the rachis, and not laterally inserted ; if this were really the case, 
the genus Pterophyllum as usually defined would be inapplicable. 
Schenk’s example does not show the rachis sufficiently clearly 
to definitely settle this point, but in all probability, as in our 
specimen, there is a narrow line of axis separating the two rows 
of segments. A close inspection of Schenk’s figure enables us 
to detect certain slight differences in the breadth of the pinne, 
similar to those in the Ecclesbourne specimen. It must be 
remarked, however, that there is very little difference in the 
breadth of the several segments. There is a striking agreement 
as regards general appearance and arrangement of the segments, 
between the Wealden specimen and the Jurassic species Anomo- 
zamites Nilssont. Lindley and Hutton’ figured this plant as 
Pterophyllum Nilssoni (Phill.): from their figure it is not easy to 
decide between ilssonia or Anomozamites as the most suitable 
genus, but an examination of several specimens of this species 
from the Yorkshire coast, shows very clearly the characteristics 
of the latter genus. There are several examples of this form in 


1 Wealdenbildung, pl. vi. fig. 3. 
2 Lindley and Hutton (A.), Fossil Flora, vol. i. pl. lxvii. fig. 2. 


ANOMOZAMITES. 93 


the Leckenby Collection, which have been examined by Nathorst 
and referred by him to Anomozamites. 

In addition to the single specimen of Anomozamites Lyellianus 
in the British Museum, there is a somewhat larger example in the 
Museum of Practical Geology, Jermyn Street. In this latter 
specimen the pinne are broader and have a more open arrange- 
ment. The lateral attachment is very clearly shown, and the four 
or five veins in each segment are distinctly marked. Here and 
there may be noticed slight differences in the breadth of the 
segments, which are arranged alternately towards the upper and 
lower ends of the specimen, but in a few cases the pinne are 
opposite. The Jermyn Street specimen is from the Wealden of 
Ore near Hastings. 


Fia. 6.—Anomozamites Lyellianus (V. 3251). Nat. size. 


V. 3251. Fig. 6. 

Probably a young leaf, showing clearly a gradual diminution 
in the length of the segments towards either end of the rachis. 
Manner of attachment and venations of the pinne clearly pre- 
served; each segment appears to have four or five simple, 
parallel, and distinctly marked veins as shown in Fig. 64. The 


94 CYCADOLEPIS. 


segments appear to be much thicker than in the similar J urassic 
form A. Nilssoni, and their breadth is much more uniform than in 
the latter species. 

A species of a somewhat analogous habit has recently been 
described by Fontaine under the name of Zamites Montanensis from 
the Montana Coal-field.t_ Ecclesbourne. Rufford Coll. 


Genus CYCADOLEPIS, Saporta. 
[Pal. Frang. sér. ii. végétaux, vol. ii. 1875, p. 200.) 


In 1875 Saporta proposed the term Cycadolepis as a convenient 
generic designation for detached bud scales of cycadean fronds. 
He defined it as follows: ‘‘Squame coriacew basi dilatate loco 
insertionis crassee facie interiori plus minusve concave nudeque, 
facie autem dorsali convexiuscule, sursum elongate lanceolato- 
acuminate, extus ad utrumque latus tomento piloso donate.” 

A small number of fronds have been recorded in which larger 
or smaller basal scaly structures are preserved ; as a few examples 
of such, we have Zamites gigas, Morr., as figured by Saporta,? 
Podozamites distans, Presl,? Otozamites sp.,* Podozamites lanceolatus 
minor, Schenk,® etc. Among recent cycads we have, in addition 
to the ordinary pinnate fronds, various forms of smaller scale 
leaves; the latter are particularly well seen in Cycas, where an 
old stem shows a clearly marked alternation of the persistent 
basal portions of fronds alternating with the bases of scale leaves. 
These scale leaves are true leaf structures, in which the green 
assimilating portion of the phyllopodium has not been developed. 


1 (A. 3), Proc. U.S. Nat. Mus. vol. xv. p. 494, pl. Ixxxiv. fig. 4. Fontaine 
omits to mention that Dawson has described a fragment of cycadean frond 
from the Kootanie series of the Rocky Mountains as a new species, Zamites 
Montana, a name dangerously near to Z. Montanensis (see Dawson, Trans. Roy. 
Soc. Canada, 1885, section iv. p. 7, pl. i. fig. 6). 

2 Loc. cit. pl. Ixxxi. fig. 1. 

3 Ibid. pl. lxxvi. fig. 2. 

4 Ibid. pl. Ixxvi. figs. 3 and 4 [see also Zigno (A.), Flor. foss. Oolit. vol. ii. 

pls. xxxy. and xxxvi.]. 
5 Nathorst (A. 1), Flor. Bjuf, i. pl. xvi. fig. 10. 


CYCADOLEPIS, 95 


In the genus Ceratozamia the expanded base of a frond shows 
two lateral stipule-like appendages, and these are also found in 
the same position on the margin of the smaller scale leaves! In 
some forms of Macrozamia, the surface of an old stem is entirely 
enclosed in a thick armour of large persistent petiole bases without 
any accompanying scales. Many recent cycadean stems appear 
to be covered by numerous scale-like structures of identical form; 
it is by no means an easy task in many cases to distinguish 
between the bases of true fronds and those of scale leaves, even 
where both forms of leaf are present. The variation in form 
and size exhibited by the scale leaves of recent species, sufficiently 
demonstrates the futility of attempting any exact generic or 
specific discrimination in the case of the isolated fossil examples. 
It is true we have in Dioon and Cycas fairly characteristic lanceo- 
late scales, often clothed in a dense woolly covering; but a close 
inspection of a tall stem of the latter genus reveals a marked 
difference in the scale leaves towards the apex of the stem, and 
those in the older portions, where there are only the persistent 
broad bases adhering to the plant stem. In dealing with fossil 
scale leaves it will probably be wise to extend the definitions of 
Saporta’s genus and to include in it not merely the ‘elongate 
lanceolate-acuminate”’ forms of bud scales, but also other forms 
of true scale leaves, as well as those structures which may be 
regarded as the persistent bases of petioles. This genus, used 
in a much more comprehensive sense, should afford a convenient 
means of grouping together those detached leaf structures, which 
frequently cannot be definitely referred to any particular genus 
or species. Such isolated plant members, in themselves, perhaps, 
of little value, are worthy of record as contributions to the 
material from which to build up a more complete history of 
fossil cycadean plants. Some of the numerous scales in the 
Rufford Collection may, indeed, be referred to certain forms of 
stems, and no doubt as our material is increased others may 
be recognized as portions of some well-defined genus or species 
of cycad. 

‘In the male and female flowers of some recent species the 
detached scales bear a close resemblance to the sterile leaf 
structures of the stem; it will be well, therefore, to include 


1 Engler and Prantl, Cycadace@, p. 7. 


96 CYCADOLEPIS. 


in the genus Cycadolepis such scale leaves as afford no clear procf 
of their carpellary or antheriferous nature. As an example of such 
a resemblance, it may be noted that some of the smaller examples 
of the Ecclesbourne scales present a distinct agreement in shape 
and size with the detached carpellary scales of such a form as 
Macrozamia Dyeri. Similarly the isolated leaves of the ‘ bulbils”’} 
of Cycas and other genera, erroneously compared by some with 
the inflorescences known as Bennettites and Williamsonia, should 
be included under this comprehensive genus; also the narrow 
lanceolate-acuminate and short broad leaves of the two latter 
genera. 

The only forms included by Saporta in Cycadolepis, e.g. C. 
villosa and C. hirta,? are narrow leaf-like structures similar to the 
scales of Dioon or Cycas. Fliche and Bleicher* have adopted 
Saporta’s name for a very imperfect fragment which seems to 
be practically indeterminable. We may, perhaps, as a matter of 
convenience, and to avoid the obvious danger and inexpediency 
of instituting several more or less meaningless specific names, 
arrange the various detached scales under two main heads, 
basing the distinction of the two sections on the general form 
of the scales. 


Cycadolepis. 


Scale-like leaf structures of cycadean plants, varying consider- 
ably in form and including detached petiolar bases, bud scales, 
etc., also isolated carpellary or antheriferous scales which exhibit 
no trace of ovules or pollen-sacs. 


1.—Cycadolepis (Dory*-Cycadolepis). Scales of a more or 
less linear-lanceolate form, broadest at the base and 
tapering gradually towards the apex. 

2.—C. (Eury?’-Cycadolepis). Broadly oval or orbicular scales, 
with the broadest portion frequently nearer the distal 
than proximal end; thick and fibrous structures. 


Miquel (1), p. 7, pl. ii. figs. I and J. 

Saporta, doc. cit. pp. 201, 202, pl. exiv. figs. 4-6. 

(A.), Bull. Soe. Sci. Nancy [2], vol. v. figs. 9-11, p. 76. 
ddpu =spear. 

evpvs = broad. 


oO - co we ~ 


CYCADOLEPIS. 97 


1.—Dory -Cycadolepis. 


The forms included in this section of Cycadolepis are practically 
such as Saporta describes in his diagnosis of the genus. Saporta’s 
examples are regarded as distinct species, but it is surely un- 
necessary to institute elaborate specific definitions for such isolated 
structures, and especially as the so-called species bear a distinct 
resemblance to one another. Both forms are from the Lower 
Kimmeridgian of the province Ain. C. villosa is compared with 
the scales of Stangeria, and C. hirta with those of Cycas and 
Dioon. It is by no means unlikely that both may belong to 
Williamsonia; similar scales figured by Fontaine in a specimen 
of Williamsonia virginiensis, Fout.,! from the Potomac beds, and 
some of the Wealden Williamsonia scales both present a strong 
likeness to the French specimens, and suggest the possibility 
of generic identity. A specimen in the Rufford Collection 
(V. 2830), consisting of a collection of narrow acuminate hairy 
scales, which is probably part of a cycadean stem, shows some 
scales very similar to C. hirta, Sap. Feistmantel has figured a 
specimen of what he calls C. pilosa? from the Gondwana flora 
of India which agrees very closely with Saporta’s examples, 
also with the leaves of W. virginiensis, Font. The same author 
figures another form as Cycadolepis,3 which may perhaps be 
regarded as a cycadean scale. The specimen figured by Nathorst 
as Cycadospadix integer angustior, Nath.,* suggests a form which 
might be included in the present section of Dory-Cycadolepis. 


V. 2802. Single leaf; linear-lanceolate in form; 7-5 cm. long, 
9mm. broad, delicate hairs on each margin. Appears to be 
identical with Saporta’s “species”? C. villosa. Compare V. 2129, 
in which there are several similar scales aggregated together. 
Very likely this specimen may be a detached leaf of Williamsonia. 
Further reference will be made to this form of scale in the 
descriptions of cycadean stems and some of the specimens of 
Bennettites (Williamsonia). Ecclesbourne. Rufford Coll. 


1 Potomac Flora, pl. exxxiii. figs. 6 and 7. 
? Foss. Fl. Gond. vol. ii. pl. vii. fig. 5. 

3 Ibid. vol. i. pt. iv. pl. xiv. figs. 10-12. 
* Nathorst, Joc. cit. pl. xviii. fig. 6. 


98 CYCADOLEPIS. 


V. 2129*. A scale or possibly petiole base with numerous 
hair-like appendages; similar to V. 2802. Cf. also Cycadolepis 
hirta, Sap., and the Otozamites petiole of Pl. II. Fig. 4 of the 
present volume. V. 2927. Part of another woolly or hairy scale. 
Ecclesbourne. Rufford Coll. 


2.—Eury-Cycadolepis. 


In this section are included several detached scales varying in 
shape from an almost orbicular or somewhat pentagonal form, such 
as V. 2699, represented in Pl. VI. Fig. 6, to the larger and longer 
type as shown in Pl. V. Fig. 2. So far as I have been able to 
discover, these forms have not been previously figured; there 
cannot be much doubt as to their original connection with some 
form of cycadean stem, and indeed some of the specimens are 
identical with the stout curved scales on such stems as those of 
Fittonia. In some cases the scales occur in very close associa- 
tion with stems, but in none are they found actually in place. 
For the present, at any rate, it is better to describe some of 
the more characteristic forms, and to include them all under 
Cycadolepis, suggesting at the same time the very probable 
and indeed almost certain identity of some with the scales of 
Fittonia and other forms of stems. Hosius and von Marck have 
described a specimen from the Gault of Ahaus (Westphalia), which 
they regard as probably made up of a few large petiole bases 
belonging to some form of cycadean stem. One of these 
*‘petiole bases” has a length of 1lem., a breadth of 4:7 ¢m., 
and is 4cm. in thickness. The generic name Jfegalozamia is 
proposed for this doubtful fossil, and the following definition 
is given by these authors: ‘‘ Rhachidum basibus inerassatis 
carnosis falciformibus, costis quatuor longitudinalibus preditis ; 
costis marginalibus acutioribus, costa et dorsali et ventrali 
obtusiori.” + Structures such as this diagnosis describes would 
be legitimately included in the genus Cycadolepis, used in the 
more comprehensive sense as suggested above. 


1 (A. 1), Paleontographica, vol. xxvi. p. 203, pl. xiii. figs. 181-183. 


CYCADOLEPIS. 99 


Vi. 2920) PIS Veerias 2: 

The figure of this large example shows very well the general 
appearance of the longer forms; the surface shown in the drawing 
is strongly convex, and at the distal end somewhat suddenly 
incurved. Some of the dark curved lines seen in the figure are 
irregularly placed grooves suggesting the tracks of some small 
animal, which has slightly eaten into the hard fibrous substance 
of the scale. Very similar markings or grooves have been noticed 
by Grand’Eury' on a leaf of Cordaites, and described by him as 
‘galeries d’insecte.’’ Other lines and striations on the convex 
surface of the scale are probably due to a wrinkling of the leaf 
substance. The large petiole bases on an old stem of Macrozamia 
Douglasi, Hill, in the Botanical Department of the British Museum, 
bear a striking resemblance to this form of Cycadolepis. In the 
recent scales there is the same tendency to terminate in the 
pointed angular fashion as seen in the figured specimen, and 
in both there is a distinct narrowing towards the base of attach- 
ment. The convex under surface of the recent scales is covered 
with a thick down of hairs, and there is a similar wrinkled 


appearance to that of many of the fossil examples. Ecclesbourne. 
Rufford Coll. 


V. 2699. Pl. VI. Figs. 6 and 6a. 

This specimen is a good example of the stouter and more 
pentagonal form of scale, the distal edge is strongly recurved as 
seen in Fig. 6a, and the narrower basal end shows the surface 
of attachment. A comparison of this form of scale with those 
on the stems of Fittonia squamata, Carr.,? Bucklandia sp., Fittonta 
igauxi, Sap.,* ete., shows a very close agreement in size and 
shape. Ecclesbourne. Rufford Coil. 


V. 2799. Another very large specimen, similar to V. 2929 (Pl. V. 
Fig. 2), 13-5em. in length, 7-5em. wide at the broadest part. 
The convex surface is marked in places by reticulated lines and 
wrinklings; towards the distal end the surface is curved gradually 


Ip. 338, ph. xxi, fies 7. 

* Carruthers (1), pl. lvi. fig. 1. (The original specimen is in the Museum 
of Practical Geology, Jermyn Street.) 

3 Saporta, doc. cit. pl. exxvii. 


100 CYCADOLEPIS. 


inwards. At the base there is a well-defined semicircular area 
bounded by a distinct line; this is probably an attachment scar. 
Very similar to MMacrozamia Douglasi, Hill. Another specimen 
with the same registered number has a similar form, and shows 
surface wrinklings. Ecclesbourne. Rufford Coll. 


V. 2131. Several specimens: in some the curvature of the 
distal end is very pronounced. 


V. 2181a. Smaller scales, about 5cm. by 4cm., closely 
resembling those of /ittonia and Bucklandia; ef., e.g., Carruthers,* 
pl. lvi. fig. 1, and Saporta,? pl. lvii. fig. 1. 


V. 2131+. Scale of medium size; shows similarly convex 
surface and recurved apical portion, also distinct basal scar. 
Ecclesbourne. Rufford Coll. 


V. 26992. Part of a very large scale; reticulately marked 
surface. 


V. 26995. 6:5 cm. long, and about 5:5 cm. broad; here the 
narrower basal end is bent sharply back, the opposite end shows 
a well-defined angular margin. 


V. 2699c. 7:5cm. in length; similar in form to the large 
scale. 


V. 2799. Shows the same kind of attachment surface at the 
base; the surface is marked by numerous dots and irregular lines 
suggestive of insect ravages. Ecclesbourne. Rufford Coll. 


V. 2749. Small scale showing distinct rectulate markings on 
the surface. On the same piece of rock there is an impression 
of a cycadean stem showing what appear to be the outlines of 
petiole bases; possibly this may be a badly preserved piece of 
a stem, to which the smaller scales were originally attached. 
Ececlesbourne. Rufford Coll, 


1 (1), Carruthers. 
2 (A. 2), Pal. Franc. vol. ii. 


CARPOLITHES. 101 


V. 2132. Similar specimen, but larger, and the stem impression 
more distinct. Ececlesbourne. Rufford Coll. 


V. 2913. Part of a well-preserved scale; shows very clearly 
the sharp angular contour; the general appearance is very similar 
to that of a large recent cycadean scale. Cf. Macrozamia, sp. 
Ecclesbourne. Rufford Coll. 


Other specimens of similar Eury-Cycadolepis species: V. 2184, 
piece of Sphenopteris Fontaine, Sew., on the same rock; V. 2286, 
V. 2301, V. 2699¢, V. 2828, V. 2929. Ecclesbourne. 

Rufford Coll. 


V. 2800 and V. 2733. These specimens present rather a 
different appearance to that of most of the larger scales; this 
may, however, be due to folding over of the edges, of which 
there is distinct evidence. Some of the specimens of Cycadolepis 
are by no means unlike certain monocotyledous spathes, but there 
can be little doubt as to their cycadean nature. Ecclesbourne. 

Rufford Coll. 


Genus CARPOLITHES, Sternberg. 
[Flor. Vorwelt, Fasc. iv. p. xl. 1823.] 


Fossil seeds are abundant in rocks of various ages, and in some 
cases their excellent preservation enables us to study in detail the 
structure of both testa and nucellus, and to refer them, with 
a considerable degree of certainty, to a particular class, family, 
or genus of plants. The superb illustrations in Brongniart’s 
posthumous work Recherches sur les graines fossiles silicifiées' 
demonstrate in a striking manner, the excellent preservation of 
isolated gymnospermous seeds under certain favourable con- 
ditions; but in spite of the perfection of the mineralized tissues, 
it is scarcely ever possible to assign the detached seeds to their 
respective plants. In Mesozoic rocks seeds are by no means 


1 Paris, 1881. 


102 CARPOLITHES. 


uncommon, but their preservation is usually imperfect, and not 
such as to throw any appreciable light on their exact botanical 
position. The Wealden strata of England have as yet been 
searched in vain, for any satisfactory indications of the existence 
of angiospermous plants in the flora of that period, and this fact 
leads us to the assumption that most probably the Wealden seeds 
are either coniferous or cycadaceous. There are, however, the 
detached tubers of Hyuisetites Burchardti, Dunk., and £. Yokoyama, 
Sew., which may easily be mistaken for seeds. In the first part 
of this Catalogue! doubt was expressed as to the nature of the 
oval bodies described as seeds by Stokes and Webb, and Mantell, 
and by some authors referred to Lyuisetites. 

The name Carpolithes was proposed by Sternberg as a convenient 
and comprehensive genus for ‘‘ Fructus seminavi mono- vel dicoty- 
ledonea, solitaria, structura interna plane obliterata.” This term 
has been adopted by many authors as a designation for isolated 
seeds of doubtful position, and its use is in most cases more 
appropriate than any term indicative of some special class or group 
of plants. In 1849 Pomel proposed Ulospermum,* as a generic name 
for fossil ‘‘ fruits’? resembling those of the recent Cycadacea, but 
this term, like many others suggested by the same writer, has 
not been generally accepted. Schimper instituted the genus 
Cycadinocarpus,’ for ‘‘Semina subglobosa, ovata vel oblonga, 
quoad magnitudinem valde variantia, nune parvula, nunc majora 
volumenque Castanez attingentia; epidermide plus minus crassa 
instructa, levia, haud raro compressione mutua angulosa, epi- 
dermide destituta solida, lignea, sublevia, striata, costata vel 
reticulata, basi insertionis cicatrice lata notata, apice minute 
apiculata.”” In 1875 Saporta substituted Cycadospermum, as a 
more fitting name for detached cycadean seeds than Schimper’s 
genus Cycadinocarpus, on the grounds of an implied misconception 
of the exact morphological nature of the seeds of cycads. The 
genus is thus defined: ‘‘Semina e carpophyllis distracta post 
maturationem in strata pervagata nune majora nune plus minusve 
parvula, plerumque ovata ovatoque-oblonga haud raro compressione 


1 Vol. i. pp. 27, 28. 
2 Pomel, p. 16. 
3 Trait. pal. vég. vol. ii. p. 208. 


CARPOLITHES. 103 


mutua angulosa extus levia vel longitudinaliter striata costataque, 
basi semper rotundiore insertionis cicatrice notata apice autem plus 
minusve attenuata.’ ! 

As regards the difference between fossil specimens of cycadean 
and coniferous seeds, it would seem that we cannot trust to any 
convenient method of distinguishing, in all cases, between the 
two groups of plants. The large oval, or almost spherical seeds 
of certain cycads may generally be distinguished from the typical 
furms of coniferous seeds, but in the latter group we have such 
forms as Ginkgo, Cephalotaxus, and others, in which the size 
approaches more closely to that of the cycadean ovule, than to 
the smaller seeds of such conifers as Larix, Pinus, and many 
others. In many Conifere the presence of a membranous wing 
affords a ready means of identification, at least as regards their 
separation from Cycadacee, but the seeds of many conifers are 
without any winged appendage, and even in the case of winged 
seeds, the thin membrane might readily become detached before 
the seed had been permanently enclosed in a mass of sediment. 
Another obvious source of difficulty, worth referring to in this 
connection, is the very great difference in size exhibited by the 
seeds of the same plant at different stages of growth. The ripe 
seeds of such a genus as Cycas, preserved with the wrinkled 
reddish-brown outer coat intact, present a very different 
appearance from those in which this coat has become detached, 
thus exposing the perfectly smooth inner coat; and a still greater 
contrast is afforded by the more spherical kernel (nucellus), with 
its surface traversed by branching grooves marking the position 
of vascular bundles.2, On the whole, it would seem advisable to 
follow the example of Schenk in his Flora der Grenzschichten,? 
and make use of the old term Carpolithes for gymnospermous 
Mesozoic seeds. In certain cases the character of the seeds, or 
their frequent juxtaposition with cycadean fronds, may enable 
us to speak of them as cycadean with reasonable certainty ; 
when such is the case it will be well to give expression to our 
more accurate knowledge, either by adopting Saporta’s genus, 


' Saporta (A. 2), Pal. Fran. vol. ii. p. 235. 
® This is well shown in the seeds of Cycas circinalis. 
3 Pl. xxxiii. figs. 5-9. 


104 CARPOLITHES. . 


or, possibly the better plan, by adding the word cycadean or 
Cycadaceeé as a descriptive epithet to Carpolithes. If some such 
course as this were generally followed, there would be less cause 
for the not altogether unwarranted criticisms, which students of 
recent plants are in the habit of passing on the misplaced 
dogmatism of palzobotanists. Our records of fossil plants ought 
surely to be sufficiently trustworthy, to be made use of by 
botanists in compiling statistics of the geological history of 
any class or family of plants. It must be admitted that to 
attempt a history of plant development or distribution in the 
various epochs of the earth’s history, by simply accepting as 
reliable data the examples of fossil plants, or fragments of 
plants, described under the names of existing genera, or desig- 
nated by terms plainly suggestive of botanical affinity, would 
lead the too trustful student into hopeless error. Occasionally 
a fossil seed may exhibit some definite and characteristic form, 
for which some special specific term might be added, but in 
the majority of cases where the individual differences are merely 
those of size or slight variation in shape the use of specific 
terms is to be deprecated. In Fontaine’s Potomac Flora several 
seeds are recorded as species of Carpolithes, the genus being 
used in this instance for the ‘‘nut-like seeds of conifers.” ? 
Under the genus Cycadinocarpus the same author places ‘‘ various 
horny seeds which resemble those of cycadean plants more than 
those of conifers.’”’? It is admitted by Fontaine that the correct 
placing of these seeds is impossible; his species are founded in 
some cases on very slight differences in size and shape, and can 
have but little taxonomic value. Saporta* has instituted various 
species for the French Jurassic seeds referred to gymnosperms; 
some of these show fairly well-marked characteristic features, 
but in others it would be difficult to justify the adoption of 
specific designations. In a recent paper by Dawson, several 
gymnospermous seeds are wisely grouped together as examples 
of Carpolithes.‘ 


1p. 264. 

2 p. 270. 

3 Loe. cit. pp. 238-245. 
4 Dawson (2), p. 90. 


CARPOLITHES. 105 


Under certain circumstances, as suggested by Solms-Laubach 
in speaking of Cycadites (Cycas) Steenstrupi, Heer,’ it may 
be legitimate to refer seeds and fruits to certain species of plants, 
even in the absence of any actual proof of organic connection ; 
but it can only be in exceptional cases where the association 
of fronds and seeds renders such a course admissible. As an 
example of what appears to be an instance of a supposed con- 
nection, not sufficiently supported by facts, we may cite Heer’s 
Zamites globuliferus,? where a frond occurs in association with 
seeds, 


Carpolithes. 
Seeds of doubtful botanical position. 


Carpolithes (Cycadacee). 


Fic. 7.—(V. 2130*.) Carpolithes (Cycadacee). 
a. and b. Nat. size. c. Portion of 6 slightly magnified. 


V. 2130" Piet 7, 

This is a particularly well-preserved example of what must be 
regarded as a cycadean seed. It is impossible to refer it to any 
particular genus, but in all probability it belongs to some other 


1 Fossil Botany, p. 86. 
2 Fl. foss. Arct. vol. vi. pt. i. pl. iv. figs. 1-7, etc. 


106 CARPOLITHES. 


form than Cycadites, if we assume that genus to have possessed 
seeds similar to those of the recent Cycas. Species of Macrozamia 
possess ovules closely resembling the present specimen.. ‘The 
seed has a length of 1:8em., and is l-lem. broad. The mould 
from which the kernel (Fig. 7b) is readily removed, is lined with 
a thin structure probably representing the integumentary portion 
of the testa (Fig. 7a); between this and the matrix there is 
a layer of coaly substance. The kernel may probably be regarded 
as a cast of the nucellus, with the impressions of the branched 
vascular bundles clearly seen on its surface. The fossil figured 
by Stokes and Webb as Carpolithus Mantelli,) shows in the 
enlarged drawing similar branched markings on the surface, 
suggestive of vascular strands. It may be that Mantell’s speci- 
men should be retained under its original genus and _ not 
transferred to quisetites, but it is difficult to speak with any 
certainty, at all events in the absence of the type specimen. 
Eeclesbourne. Rufford Coll. 


V. 2129, V. 2131, V. 2699. Large flattened and more or 
less spherical bodies showing coaly substance on the exposed 
surface; in V. 2131 the seed (?) is 4cem. in breadth. It is 
possible that some of them may be scales and not true seeds, 
but their general appearance is not unlike that of some recent 
cycadean ovules, e.g. species of Cycas. Ecclesbourne. 


Rufford Coll. 


Cf. V. 2180", V. 2236. Small specimen of a badly preserved 
seed. 


V. 28272. Seed with pointed apex, not unlike Cycadeospermum 
obovatum, Font.2 V. 2256. Cast of nucellus with remains of 
testa, similar to V. 2827a. V. 2700*. Cast and mould of 
imperfect specimen. Ecclesbourne. Rufford Coll. 


V. 3312. Probably a seed of some cycadean plant; it appears 
to have split partially open along the longest diameter. Eccles- 
bourne. Rufford Coll. 


1 (A.), pls. xlvi. and xlvii. 
2 Potomac Flora, pl. exxxv. fig. 13. 


CARPOLITHES. 107 


Carpolithes. 


V. 2184. Part of a seed-like body. There are several small 
circular holes on the surface of this and a few other specimens, 
filled up with a fine brown dusty material, suggesting the 
borings of some small animal. Similar examples are afforded by 


V. 2739, V. 2918. Kcclesbourne. Rufford Coll. 


The following specimens may also be included under Carpo- 
lithes: V. 21380, V. 2130a, V. 21303, V. 2165, V. 2700, 
V. 2739*, V. 2826, and V. 28270. 


38369. Possibly an imperfect seed, but indeterminable. 


Seed-like Bodies of Doubtful Position. 
Cf. OOLITHES, sp., CARRUTHERS.! 


Pl. IX. Fig. 5 (V. 27962). An oval body partially covered 
with a chitinous-like coat; the dark brown and brittle 
substance which occurs over part of the specimen, suggests some 
resemblance to the dried reddish coat of a Cycas seed. The 
central part does not show any signs of a nucellus or seed 
structure ; it consists of an irregularly indented projecting portion 
of the rock. The external skin exhibits no cellular structure 
under the microscope. 

Compare Oolithes sphericus, Carr.; the figures given by 
Carruthers of this species, present a striking resemblance to 
the present specimen and other similar forms in the Rufford 
Collection. Buckman had previously identified these Stonesfield 
slate bodies as reptilian eggs, and Carruthers’ examination of 
the same material leads him to accept this determination. It 
is probable that whatever position be assigned to the Jurassic 
fossils, it may with equal force be accepted for many of the 
Wealden seed-like bodies. 


V. 2796. In this specimen the outer brown skin has been 
removed. 


V. 2818. Smooth brown skin present. Ecclesbourne. 
Rufford Coll. 


1 (2), p. 447, pl. xix. 


108 FLORES. 


V. 2825. Flattened and subspherical body, with a hard shiny 
and dark brown skin more or less deeply indented. Cf. 
Carruther’s figures of Oolithes, sp. 


V. 2828. Specimens showing a similar brown skin, enclosing 
a smooth central kernel. 


V. 2817. Small specimen with smooth surface, showing at 
the two opposite ends of a diameter a number of very small 
rounded prominences; these are just visible, as small dots, to 
the naked eye. 


V. 2817a. Small body, 5mm. long, with smooth brown coat, 
similar to V. 2796a, ete. V. 2165, fragment. Ecclesbourne. 
Rufford Coll. 


FLORES. 


In Solms-Laubach’s Fossil Botany,’ we have a concise and 
eritical reswmé of the various male and female cycadean flowers 
described in paleeobotanical literature prior to 1887. It will 
be seen from this account, that our knowledge of the floral 
structures of fossil cycadean plants is extremely meagre. In 
the carpophylls of the recent Cycas, we have a well-marked 
and peculiar form of female flower which is readily distinguished 
from the cone-like collection of carpophylls met with in other 
genera; occasionally these Cycas forms of flowers have been 
found in close association with the sterile fronds of Cycadites, 
and justify the conclusion that both structures formed parts of 
the same plant. In other cases, however, we are less fortunate 
in the records of staminal or carpellary leaves, and there must 
be considerable hesitation in accepting several of the examples 
which have been described as true cycadean flowers. 

It will be convenient to adopt Schimper’s genus Androstrobus 
in speaking of a few Wealden specimens, of what appear to be 
male flowers of some genus of cycadean plant. 


1 Solms-Laubach (A.), p. 89. 


ANDROSTROBUS. 109 


Genus ANDROSTROBUS, Schimper. 


[ Trait. pal. vég. vol. ii. p. 199, 1870.] 


Schimper has thus defined the genus: ‘‘Amenta cycadeacea 
antherifera, cylindrica, e squamis imbricatis, latere postico 
antheras sessiles ferentibus efformata.”’ 

A sufficient definition of Androstrobus, would be to speak of 
the genus as a convenient term to apply to such fossils as 
resemble more or less closely the male flowers of recent cycads, 
and which appear to belong to the Cycadacee. 

The genus was founded on a specimen originally described by 
Saporta from the Upper Bathonian of Etrochey as 4. samioides, 
but afterwards renamed A. Balduini' after the discoverer of the 
specimen, the latter specific term being considered more suitable 
as not suggesting such a definite resemblance to a particular form 
of cyead. Saporta’s figures show the outline of several pollen- 
sacs between the spirally arranged staminal leaves, attached 
apparently in the same manner as in recent species. This 
author describes another and more imperfect specimen of a male 
cone, under the name Androstrobus (Zamiostrobus) Guerangert ;* 
the same specimen having been previously referred to by 
Brongniart* as an undoubted example of a male cycadean 
flower. Saporta compares this fossil with the genus Dioon, but, 
as Solms-Laubach * has suggested, there seems to be but slender 
grounds for such a comparison. The specimens described by 
Nathorst® and Heer® respectively as Androstrobus borealis and 
A. Sibiricus, are far from satisfactory, and cannot be accepted 
as entirely trustworthy records of this genus. In addition to 


1 Pal. Franc. vol. ii. p. 209, pl. exv. figs. 1 and 2. 

2 Ibid. p. 37, pl. lxxviil. figs. 1-3. 

3 Tableau, p. 64. ' 

4 Loc. cit. p. 90. 

5 (A. 1), Flor. Héganis, p. 49, pl. ii. figs. 12 and 13, and pl. ii. (Helsingborg) 
figs. 15 and 16. 

6 Fl. foss. Arct. vol. iv. pt. ii. p. 47, pl. iv. figs. 14 and 16. 


110 ANDROSTROBUS. 


the genus Frigia of Velenovsky,! reference may also be made 
to Zamites familiaris (Cord.)? from the Lower Quader of Bohemia, 
which Corda and Carruthers* regard as an example of a male 
cone; the figures of this form lend support to such an opinion. 
The specimen described by Carruthers in his monograph on 
cycadean stems as the ‘‘antheriferous cone of Bucklandia’’* does 
not seem to me to afford any distinct evidence in favour of such 
a determination. 


Androstrobus Nathorsti, sp. nov. 


Pl. IX. Figs. 1-4. 

Type. Specimens of imperfect flowers and detached staminal 
leaves. British Museum. 

It is difficult to give a definition of this species which shall 
be in any sense complete. The following may serve to indicate 
the most obvious features of this somewhat unusual form of 
cycadean cone. 

Axis fairly stout, bearing spirally disposed and more or 
less triangular staminal scales; in section the scales have a 
hexagonal outline, in side view they show a broad base in close 
contact with the axis of the flower, and measure about 1—1:5 cm. 
in length, gradually tapered towards the apex, which is pointed 
or slightly rounded. On some of the staminal leaves there are 
rows of regularly placed angular depressions, probably repre- 
senting the impressions of pollen-sacs, borne towards the basal 
or proximal end of each scale. 


V. 2701. Pl. IX. Figs. 3 and 4. 

This specimen shows several fairly well-preserved scales of 
a male flower, length about 6°5cm., breadth 4em. The central 
axis is not very clearly seen, but there are indications here and 
there of the points of attachment of the sporophylls. The axis 


1 (A. 1), Gym. Bohm. Kreid. p. 8, pl. iii. 
2 Corda in Reuss (A.), Verstein. bohm. Kreid. p. 86, pl. xlix. figs. 10 and 11. 


Sac) Deo: 
4 (1), pl. liv. fig. 6. 


ANDROSTROBUS. 111 


appears to have been about lem. in breadth, but it is difficult 
to estimate the dimensions with any accuracy. The surface of 
the scales is of a brown colour; the longest measures 15cm. in 
a direction at right angles to the floral axis; the surface is con- 
siderably wrinkled and bears obvious traces of having been folded 
and crushed. Towards one end of the specimen the basal part 
of a scale is seen in surface view, and on it are clearly preserved 
what are taken to be the outlines of pollen-sacs (Figs. 3 and 4). 
These are in the form of small depressed areas radiating from 
the proximal portion of the scale surface; each depression is 
bounded by a straight basal wall, and two slightly diverging 
lateral walls, with two apical walls inclined to one another at 
an angle of about 35°; from the apex there is a slight median 
ridge passing to the basal wall. In the upper row there are 
about 14 of these pollen-sac impressions, and below these there 
are the remains of a lower set of similar structures. Traces of 
the pollen-sacs occur on some of the other scales, but less clearly 
preserved. The striking regularity with which these impressions 
are arranged, is much more marked than in the pollen-sacs of 
recent cycads. On the lower surface of a staminal leaf of Dioon 
or Encephalartos, we find on the removal of the pollen-sacs a 
fairly distinct reticulate marking, but of much less regularity than 
in the fossil. The angular outline of the sacs in the present 
specimens may be due, to some extent, to the mutual pressure 
of more or less oval structures, such as we have in the pollen- 
sacs of recent male flowers. 

The tapered free ends of the scales are somewhat similar to 
the narrowed apices of the staminal leaves of species of Ence- 
phalartos; eg. the male flower of #. Altensteinu, Lehm, E£. 
pungens, Lehm., ete. Cf. Androstrobus Balduini, Sap., pl. exv. 
figs. 2a and 26.1 Ecclesbourne. Rufford Coll. 


V. 2810; PIL, Vig. 1. 

7cm. in length. This specimen shows several fairly well- 
preserved scales somewhat closely set on a central axis which 
is narrower than that in V. 2701. Lach staminal leaf presents 
a triangular outline, with a more or less distinct median ridge 


1 Loe. cit. 


112 ANDROSTROBUS. 


extending across the middle; if this be clearly seen in the scales 
on the right-hand side of the axis in Pl. IX. Fig. 1, towards 
the upper margin of some of the scales there are clear indica- 
tions of another projecting angle, e.g. the third from the bottom 
on the right of the axis in Fig. 1. There is a close similarity 
in form between the detached scale figured by Carruthers as 
Araucarites Phillipsti, Carr.,' and those of the present specimens. 
The median line, as seen in the scales of Fig. 1, should be com- 
pared with the prominent lateral angles seen in the end view 
of the scales in Pl. IX. Fig. 2. In one place on the surface 
of the argillaceous matrix, there were some fairly distinct 
impressions of pollen-sacs. LEcclesbourne. Rufford Coll. 


Wee2oll sPlvLX! Pie.2: 

This shows several staminal leaves in end view, some being 
apparently in place and retaining the spiral arrangement. There 
can be little doubt as to the identity of these with V. 2810 and 
V. 2701, the different appearance in the present example being 
due to the fact that here we have a view of the end, and in the 
previous specimen, a view of the flattened sides of the scales. 
The exposed ends show a central depression, and a distinct 
hexagonal outline. The shape and general appearance of the 
scales remind one of the staminal leaves in Zamia, sp.; but in 
the present specimen we are presumably looking at the basal, and 
not the distal ends of the scales; the specimens V. 2810 and 
V. 2701 show the much greater width of the base than the 
apex. It may be, however, that in Fig. 2 (V. 2811) the 
apices have been depressed, and we have a view of the apical 
rather than the basal parts of the scales. 

Cf. Androstrobus Guerangert, Brong.? Ecclesbourne. 


Rufford Coll. 


V. 2236. A smaller specimen than V. 2701 and V. 2810; 
the remains of a central axis with a few well-preserved scales 
attached. No trace of pollen-sacs. Ecclesbourne. ufford Coll. 


1 Carruthers (5), pl. ii. fig. 8. 
2 Loe. cit. pl. lxxviii. fig. 1. 


CONITES. 1138 


Genus CONITES, Sternberg. 
[Flor. Vorwelt, fase. iii. p. 36, 1823.] 


Several writers have called attention to the close resemblance 
between the cones of certain Conifere@ and those of some species 
of cycads. In attempting to determine the true nature of a fossil 
cone, of which the internal structure is either very imperfect or 
entirely wanting, we are met by the great difficulty of clearly 
discriminating between the female flowers of these two groups of 
plants. Carruthers’ has mentioned certain distinctive characters 
of cycadean cones which, he considers, should enable us to 
distinguish them from the corresponding structures of conifers, 
but the main differences which he notes are such as can only 
be recognized by the help of internal structure; he writes: 
‘“‘Any difficulty in determining the affinity of a cone by its 
external characters can easily be solved, as to whether it is 
coniferous, cycadean, or proteaceous, by a transverse section, 
which would show, if the structure is even a little preserved, 
the form of the scale and the position of the seed.”* Unless 
the structure is fairly well preserved there is often no little 
difficulty in deciding in favour of one or other of the two 
orders of plants, Conifere and Cycadacee. 

In view of the generally recognized difficulty of clearly 
separating the cones of these plants, and of distinguishing some 
cones from small cycadean stems, there must be a certain amount 
of hesitation in choosing the most suitable generic term for cone- 
like fossils of doubtful affinity. 

Endlicher* proposed the name Zamiostrobus for a cone originally 
figured by Lindley and Hutton as Zamia macrocephala*; but 
Carruthers has since shown that the original reference of this 
fossil to the Cycadacee cannot be accepted, and it is now known 


1 Carruthers (4), p. 536. 
2 Thid. p. 536. 

3p. 72 (No. 707). 

4 Fossil Flora, pl. exxxvi. 


114 CONITES. 


as Pinites macrocephala (L. and H.).1 Owing to the erroneous 
inclusion of this specimen in the genus Zamiostrobus, and the 
unwarranted application of the name to cones which are clearly 
not cycadean, Carruthers proposed the generic name Cycadeostrobus 
as a more suitable designation for what are ‘‘ supposed to be fruits 
of Cycadee.” In speaking of the cones figured by Carruthers 
under this genus, Solms-Laubach? reasonably suggests that possibly 
several of the fossils may be either small stems or true cones. The 
only certain cone he considers to be that figured as Cycadeostrobus 
Brunonis, Carr., but this, he adds, ‘‘looks more like a cone of 
Araucaria than of Cycadee@.”’ Having had an opportunity of 
examining Carruthers’ type specimens, I must confess to a con- 
siderable amount of scepticism in accepting them as well-authenti- 
cated examples of cycadean flowers. 

In cases where it seems impossible to express oneself with any 
degree of certainty as to whether a specimen is a small stem or 
cone, the better plan is probably to give expression to the doubtful 
affinity by leaving the fossil unnamed, or by prefixing a query to 
any name which it may have already received. The practice of 
replacing some of the older and more indefinite names of the older 
paleobotanists by newer terms more expressive of definite botanical 
affinity, has not always marked an advance in accurate knowledge. 
Such a name as Conites does not, indeed, convey any particular 
information to the mind of botanists as to the nature of the 
fossils so designated, but, on the other hand, Zamiostrobus or 
Cycadeostrobus both definitely suggest either the male or female 
flowers of some form of cycad. 

In the first volume of this Catalogue the term Algites® was 
proposed as a useful generic designation for doubtful forms of 
fossil Alge, in preference to the more committal and frequently 
misleading names often made use of. Although such a course 
as this is, in one sense, rather retrogressive than progressive, 
yet it would at all events minimise the chances of possible error 
if we adopted the old name Conztes for several of the cones 
previously referred to the Cycadacee on what appears to be too 


1 Carruthers (4), p. 538. 
2 Loe. ctt. p. 92. 
3 Seward (2), p. 2. 


CONITES. 115 


often insufficient evidence. If we have distinct cycadean cones 
before us, the name Cycadeostrobus would seem a suitable term 
to apply to them. As in the case of the genus Carpolithes, we 
may always give expression to any bias towards one or other 
group of plants, by adding the word Cycadee or Conifereé as 
qualifying epithets to the more comprehensive generic name. 

I would suggest, then, the revival of the old genus Conites* 
as a convenient generic name for cones of doubtful botanical 
affinity. 


Conites elegans (Carr.). 


1867. Cycadeostrobus elegans, Carruthers, Journ. Bot. vol. v. p. 7, pl. Ivii. 
ia Ge 

1867. Cycadeostrobus ovatus, Carruthers, Joc. cit. p. 6, pl. lvii. fig. 1. 

1870. Zamiostrobus elegans, Schimper, Trait. pal. vég. vol. i. p. 203. 
Zamiostrobus ovatus, Schimper, loc. cit. p. 203. 

1871. Zamiostrobus elegans, Schenk, Paleeontographica, vol. xix. p. 228. 
Zamiostrobus ovatus, Schenk, loc. cit. p. 228. 

1889. Cycadeostrobus elegans, Bristow, Geol. I. Wight, p. 248. 
Cycadeostrobus ovatus, Bristow, loc. cit. p. 248. 


Type. Pyritized specimens, British Museum. 

After an examination of the type specimens of Cycadeostrobus 
elegans and C. ovatus which Carruthers has described, I have 
ventured to include both examples under one specific name. The 
pyritized specimens do not appear to present any distinctive 
characters which can be regarded as of specific value. Unfor- 
tunately the preservation is not such as to enable us to prove 
either cycadean or coniferous relationship. Carruthers speaks of 
Cycadeostrobus elegans as an ‘‘ovoid cone, truncate below; scales 
nearly as deep as they are wide,’”’? and of C. oratus as an ‘‘ ovate 
cone ; scales somewhat broader than deep.’’* In the absence of 
structural characters it is impossible to give any more complete 
diagnosis. 


1 The genus Strodilites was suggested in 1840 by Schimper and Mougeot for 
certain cones from the Triassic beds of the Vosges. 

4 (Gp 1De te 

3 Ibid. p. 6. 


116 TRUNCLI. 


40962. Journ. Bot. vol. v. 1867, pl. lvii. fig. 9. One of the 
two specimens is the type of Carruthers’ species, Cycadeostrobus 
elegans. In the better specimen, as shown in the figure, 
the pyritized cone has been more or less compressed ; at 
the base there is a central depression or scar of attachment of 
a peduncle. The surface view of the scales suggests a wearing 
down of their distal ends. The second example is less perfect 
than the type specimen. Brook Point. Lady Hastings Coll. 


V. 2543. Two specimens, very friable. Cf. Carruthers’ figure 
of Cycadeostrobus ovatus, Carr. Possibly Cycadeostrobus truncatus, 
Carr.,! might also be included as a synonym of the present 
species. Brook. Presented by A. Dendy, Esq., 1888. 


V. 63. Imperfect pyritized specimens. These and V. 2548 are 
somewhat larger than the cone represented by Carruthers in Journ. 
Bot. vol. v. pl. lvii. fig. 9; but they are probably examples of the 
same species. Brook. Purchased 1882. 


V. 2853. Portion of a flattened cone; apparently the same as 
40962. Sussex. Beckles Coll. 


V. 385. Very imperfect pyritized specimens. Brook. 
Presented by C. Westendarp, Esq., 1884. 


TRUNCI. 


In the introductory remarks? on cycadean fronds it was 
suggested that the use of some more general term than that of 
Cycadacee, might prove advantageous in dealing with the remains 
of extinct cycad-like leaves. The chief reason for such a proposal 
is to be found in the character of the floral structures of the 
genus, for which Carruthers instituted the name of Bennettites. 
This plant, as we have already shown, cannot well be included 
in the class Cycadacee as at present defined for recent species; 


1 (3), p. 6, pl. lvii. fig. 3. 
Seles 


TRUNCI, 117 


the same necessity for a more comprehensive class designation is 
equally apparent in the case of cycad-like stems. It would be 
impossible to so far extend the present limits of the Cycadacee, as 
to incorporate under that term all the fossil stem structures in 
which characteristic features of cycadean anatomical structure 
have been recognized; but we must in any case clearly under- 
stand that such stems as Sennettites and others, although very 
closely related to recent cycads in histological details, are, 
however, separated from living forms by certain peculiarities 
in the morphology of their reproductive organs. The above 
heading, therefore, of Zrunei does not exclude such stems as 
are known to be associated with a bennettitean form of floral 
structure; it must be taken in a more comprehensive sense than 
merely including stem structures which agree in all essential 
features with living members of the Cycadee@ or Zamiee. 

The study of cycadean stems has been raised to considerable 
importance by the fact of the preservation, in several: instances, 
of more or less perfect internal structure in fossil specimens. 
As with fronds, so here again we are debarred from any complete 
diagnoses of many fossil stems by the isolated occurrence of the 
leaves and their supporting axes. We must for the present 
restrict ourselves to an investigation of facts as regards the 
anatomy of stem structures; and, as in Pennettites, of the 
accompanying floral shoots. 

The early records of so-called cycadean stems in Paleozoic 
rocks have already been referred to. It is often a matter of 
some considerable difficulty to confidently identify a structureless 
cast or impression of a cycadean trunk ; the imperfectly preserved 
stems of some forms of Sigillaria, Lepidodendron, or Lepidofloyos 
may simulate fairly closely the characteristic appearance of 
cycadean stems. In Grand’Eury’s recent monograph on the 
Coal-field of Gard there is a figure of Lepidofloyos laricinus, 
Sternb.,1 which may be reasonably compared to a stem of 
a cycadean plant, bearing lateral appendages suggestive of a 
bennettitean inflorescence. The tree fern genus Protopteris, with 
its leaf-trace bundle scars imperfectly shown or apparently 
absent, may be mistaken for a cycadean axis with its prominent 


1 Grand’ Eury (1), pl. vi. fig. 17. 


118 TRUNCI. 


petiole bases. An example of such resemblance is afforded by 
a specimen figured by Hosius and von Marck as probably Proto- 
pteris punctata, Sternb. ; the plant represented in their plate xliii. 
fig. 1861 might well be described as an imperfect cycadean stem. 
Again, it is almost impossible in some cases to decide with 
certainty between an imperfect cone and a small cycadean stem. 
The fossil described by Lesquereux from Colorado as Zamiostrobus 
mirabilis,? is obviously a badly preserved stem with basal portions 
of petioles. An examination of such a stem as that of the living 
species of Cycas, is sufficient to demonstrate the difficulties 
attending our attempts to separate into specific forms fragments 
of imperfect stems. The upper part of an old Cycas stem with 
its bud scales still in place, presents a very different appearance to 
the lower portion of the same axis, from which the scale leaves and 
petiole bases have become detached, leaving clean-cut rhomboidal 
scars. As a general rule we have a fairly easy task in identifying 
fossils as cycadean stems. The frond scars and scale leaves which 
clothe the woody axis afford a convenient distinguishing feature ; 
but, on the other hand, it is important to keep in view the existence 
of other forms of stems among recent cycads, in which the well- 
known covering of leaf bases is absent. In such plants as Zamia 
Loddigesti, Miq., and Z. Skinneri, Warsz., the peculiar branched 
stem, with its transversely elongated wrinklings and small knob- 
like protuberances, presents a totally different aspect to the trunks 
of Cycas, Encephalartos, Dioon, and others. It may be noted 
in this connection that the Lower Greensand fossil which Konig 
named Dracena Benstedtii,? and of which the National Collection 
contains several examples from the Kentish Rag of Maidstone, 
and a few recently added by Mr. Rufford from the Ecclesbourne 
Wealden Beds, shows a striking resemblance to the stems of the 
above-named forms of Zamia. The fossils have at all events no 
claim to a generic name implying a monocotyledonous affinity. 

We cannot here undertake a descriptive account of the 
morphology of recent cycadean stems; but for information on 
this head, reference may be made to the treatment of these 


' (A. 1), Paleeontographica, vol. xxvi. 
 Lesquereux (1), p. 70, pl. Ixiii. fig. 1. 
3 Morris (A), Brit. Foss. p. 8. 


TRUNCI. 119 


plants in Engler and Prantl’s Die natiirlichen pflanzenfamilien.' 
Saporta? and Renault? have also given some account of the 
living cyeads, and further details may be found in the writings 
of Brongniart,* Miquel,> Richard,® Karsten,’ Carruthers,® Solms- 
Laubacb, and others. 

By far the greater number of known fossil stems have been 
found in Lower Cretaceous and Jurassic strata, and it is with 
these Mesozoic examples that we are at present concerned. In 
Brongniart’s Prodrome® there is recorded but one example of 
a cycadean stem; this is the plant described by Buckland from 
the Portland dirt-bed, and for which the French author suggested 
the name of Mantellia. The common Clathraria Lyelli, Mant., 
is included by Brongniart among the Monocotyledons. In the 
Tableau we find several additions to the list of cycadean stems, 
and among them is the interesting genus J/edullosa of Cotta; 
this Paleeozoic plant has been subjected to a detailed investigation 
by Goppert and Stenzel," Schenk,” and others, and we may 
probably regard it as an extinct type of cycadean structure, 
using the term cycadean in a wide sense. In 1828 Buckland’ 
figured and described some large specimens of silicified stems 
from the Isle of Portland, and, with the concurrence of Robert 
Brown, instituted a new family, Cycadeoidee, for their reception. 
Buckland fully recognized the close resemblance between these 
‘‘netrified birds’ nests’? and the stems of certain cycads; but 
a new family name was proposed on account of some peculiarity 
as regards the position and size of the rings of wood. In a 


1 Teil. ii. Abth. i. p. 6. 2 Pal. Frang. vol. ii. 
3 (A. 4), Cours bot. foss. vol. i. p. 33. 


a (2). 5 (1) and (3). 

6 Richard. 7 Karsten. 

8 Carruthers (1). See also Solms-Laubach (8). 
2) 0s SR 2) Ws BY). 


11 Goppert and Stenzel.—In speaking of the Medullosee, Solms-Laubach 
remarks (Fossil Botany, p. 100) that ‘‘that in their anatomical structure they 
show many points of resemblance to the Cycadee, though they depart from them, 
according to the most recent investigations, in some important particulars.”’ 

12 Schenk. 13 Buckland (1). 


120 TRUNCI. 


later work,’ this author has given some further description of 
the Portland fossils, and discusses the question of terminology ; 
the genus Cycadites being regarded by Brown as preferable to 
Cycadeoidea, and the name of Mantellia, proposed by Brongniart, 
is thought to be unsuitable, having been already used by Parkinson 
for a genus of fossil Zoophytes. At the present day? it is un- 
fortunately not always held that the use of a particular name 
by palzeozoologists, is a fatal objection to the adoption of the same 
for a fossil plant. Among the new figures added by Buckland to 
those given in his earlier paper, we find some drawings of longi- 
tudinal sections of petioles and axillary buds; the latter have since 
been fully described by Carruthers as the inflorescence of 
Bennettites. In 1870 an important monograph appeared by the 
latter author on Jossil cycadean stems from the Secondary rocks 
of Britain;? the memoir contains full reference to earlier records 
of cycadean stems, and includes figures and descriptions of the 
following new genera — Yatesia, Fittonia, Walliamsonia, and 
Bennettites. Five years later, several additions were made by 
Saporta* to our knowledge of the stems of fossil cycads; he 
founded the genera Bolbopodium, Cylindropodium, Platylepis, 
Clathropodium, and Cycadeomyelon. The numerous terms added or 
substituted for those previously proposed by Carruthers have 
involved the terminology of cycadean stem structures in some 
confusion. In more recent years we have a valuable contribution 
from Solms-Laubach and Capellini® on the examples of bennettitean 
stems preserved in Italian museums. These authors limit the use 
of the term Bennettites to a single species, B. Gibsonianus, Carr., 
and in a still later preliminary paper by Lester Ward,® Carruthers’ 
genus is absorbed into the more comprehensive Cycadeoidea. We 
may look for an important monograph at an early date by Lester 
Ward and Knowlton on the exceedingly fine series of American 
eycadean stems. In Dana’s Manual of Geology’ mention is 


1 (2), p. 453. 

* See Quart. Journ. Geol. Soc. vol. 1. 1894, p. 435. 
3 Carruthers (1). 

4 Pal. Frang. vol. ii. p. 245. 

5 Capellini. 

6 Ward (1), p. 78. 

7p. 472. 


BUCKLANDIA. 121 


made of some large stumps of cycads having been found near 
Baltimore, Maryland, and their age is spoken of by Tyson as 
probably Upper Jurassic. Fontaine’s Potomac Flora’ contains 
a few photographs of these Maryland stems, and a splendid 
specimen has lately been received by the Botanical Department 
of the British Museum. The few facts we so far possess as to 
these American stems lead us to expect a descriptive monograph 
of exceptional interest.* 

The material so far collected from Upper Jurassic and Lower 
Cretaceous strata has already yielded valuable information with 
regard to the anatomy of the vegetative, and in some instances 
of the reproductive, structures of Mesozoic cycadean plants. To 
further extend our knowledge of these various fossil species, a more 
intimate acquaintance with the several types of recent cycads 
is much to be desired; and, as Solms-Laubach* points out, we 
possess no detailed and modern account of the large tuberous 
stems long ago described by Buckland from the dirt-beds of 
Portland. 


Genus BUCKLANDIA, Presl. 


[Sternberg, Flor. Vorwelt, fase. iv. p. xxxiil. 18205.] 


This genus was instituted by Presl for a plant discovered by 
Mantell in the Wealden of Tilgate; the same fossil had been 
previously referred by Stokes and Webb to Clathraria,* a term 
proposed by Brongniart® in 1822 for certain forms of sigillarian 
stems. Mantell® was the first to give a description of these Tilgate 
plants, but he proposed no name for them, merely pointing out 
a probable affinity with the Euphorbiacee, or possibly with the 
arborescent ferns. Carruthers pays a tribute to the ‘‘ remarkable 
discrimination”? with which Presl recognized the cycadean nature 


1 Fontaine (A. 2), pls. clxxiv.—clxxx. 

2 MacBride. 

3 Fossil Botany, p. 99. 

4 Stokes and Webb (A.), Trans. Geol. Soc. [2] vol. i. p. 421. 
5 (3), p. 209. 

6 Mantell (A. 3), Ilust. Geol. Sussex, p. 42. 

7 Loc. cit. p. 682. 


122 BUCKLANDIA. 


of the English fossils. In the Zableau! Brongniart includes Mantell’s 
plant in the Ziliacee, noting at the same time its resemblance to 
the stem of a cycad. Schimper? retains Clathraria for plants of 
the type of Clathraria anomala, Stokes and Webb (C. Lyelli, 
Mant.); and Saporta,? who follows Carruthers in preferring the 
name Bucklandia to Clathraria, speaks of the plant figured by 
Schenk as O. Lyelli as probably a species of Carruthers’ genus 
Littonia. Nathorst, in his Floran vid Bjuf,t on the other hand, 
includes under Clathraria two new species, but in the latter part 
of the same work he substitutes Bucklandia® for Stokes and 
Webb’s genus. Nathorst’s specimens are imperfect fragments 
of stems with alternating series of narrower and crowded leaf- 
scars, and broader and more openly arranged leat bases; he 
compares them with the stem of Cycas. 

The separation of such conventional genera as Bucklandia and 
those proposed by Saporta, is often a matter of great difficulty, 
and so long as we have only imperfect external or internal casts 
to deal with, there must always be a certain amount of doubt 
as to the existence of true generic and specific differences. 
Carruthers thus defines the genus Buchklandia:— 

‘“Trunk cylindrical, sometimes bifurcating, reticulate, with the 
scars of the bases of the leaves, which are arranged in alternating 
series of large and small scars, the large being placed on swellings 
and the small on constrictions of the stems. Andrcoecium a cone (?), 
gyncecium a terminal crown of leaves bearing seeds on their some- 
what altered margins.’ ® 

The so-called male cone referred to in this definition was 
discovered in the same series of strata as those im which 
Bucklandia occurs; it is assigned to this genus on the strength 
of its occurrence in the same beds, and on account of a resemblance 
which its scales present to the sporophylls of a male flower of 
the recent species of Cycas. In the absence of more satisfactory 
evidence than is afforded by this single imperfect specimen, the 
nature of which does not appear to be by any means established, 
we are not in a position to include the male flower in a definition 
of Presl’s genus. ‘The alternating swellings and constrictions 


1 Brongniart (A. 4), p. 91. 4 Nathorst (A. 1), p. 77. 
2 Trait. pal. vég. vol. ii. p. 182. 5 Thid. p. 124. 
3 Loe. cit. p. 307. § Loc. cit. p. 682. 


BUCKLANDIA. 123 


of the stem, and the slight difference in the form and size of 
the leaf-scars, led Carruthers to draw a close parallel between 
Bucklandia and Cycas, and to infer the nature of the female 
flowers. He writes: “If the interpretation I have given of 
the stem of Bucklandia be correct, and if there be good reason, 
from a morphological point of view, for connecting with it the 
seeds and male cone found in the same beds, we have a plant 
which, in these known particulars, cannot be separated generically 
from Cycas.’* In addition to the original species, Bucklandia 
anomala (S. and W.), Carruthers proposes a second specific name 
for certain Wealden stems which he considers to be distinct from 
Stokes and Webb’s type. An examination of the British Museum 
material does not appear to favour this separation into two 
distinct forms, and I have ventured to incorporate both of the 
species into B. anomala. We cannot hope to separate such 
imperfect and structureless specimens into specific forms of any 
real value, seeing what marked variations in surface features we 
must expect to find in examples of cycadean stems clothed with 
a number of more or less decayed leaf bases. It may be noted 
that the upper portion of the specimen of Bucklandia anomala 
figured by Carruthers in pl. liv. fig. 1, shows a close approximation 
in form to that of some forms of the genus /ttonva. 


Bucklandia anomala (Stokes and Webb). 


1824. Clathraria anomala, Stokes and Webb, Trans. Geol. Soc. [2] vol. i. 
p- 422, pls. xlv., xlvi. fig. 8; pl. xlvii. fig. iv. 

1827. Clathraria Lyelli, Mantell, Illust. Geol. Sussex, p. 52, pl. i. fig. 2; 
pl. ii. figs. 4 and 6. 

1828. Clathraria Lyelli, Brongniart, Prodrome, p. 200. 

1833. Olathraria Lyelli, Mantell, Geol. S.E. England, p. 233, pl. i. figs.i., ii, 
and vi. 

1844. Clathraria Lyelli, Mantell, Medals, vol. i. p. 182, fig. 44. 

1847. Clathraria Lyelli,? Mantell, Geol. Excurs. I. Wight, p. 292. 

1848. Clathraria Lyelli, Broun, Index pal. nomencl. 


1 This suggested resemblance to Cycas does not appear to me very close ; 
a comparison of the fossil stem and its pith cast with the stem and pith cast 
of Macrozamia, sp., reveals a striking similarity. 

2 Carruthers, doc. cit. p. 686. 

3 The specimen figured by Mantell (p. 293), Abth. i. p. 305, as Clathraria 
Lyellt is a waterworn fragment of bennettites. 


124 BUCKLANDIA. 


1850. Clathraria Lyelli, Unger, Gen. spec. plant. foss. p. 314. 
1851. Clathraria Lyelli, Ettingshausen, Abh. k.-k. geol. Reichs. vol. i. Abth. 
iii. No. 2, p. 26. 
1851-52. Clathraria Lyelli, Bronn, Leth. geog. vol. ii. p. 63, pl. xxviii. fig. 7. 
1854. Clathraria Lyelli, Morris, Brit. Foss. p. 6. 
1870. Bucklandia anomala, Carruthers, Trans. Linn. Soc. vol. xxvi. p. 686, 
pl. liv. figs. 1-3. 
Bucklandia Mantelli, Carruthers, ibid. p. 686, pl. liy. fig. iv. 
1870. Clathraria Lyelli, Schimper, Trait. pal. vég. vol. ii. p. 182. 
1871. Clathraria Lyelli, Schenk, Paleontographica, vol. xix. p. 227, pl. xxx. 
fig. 7. 
1874. Clathraria Lyelli, Schimper, Trait. pal. vég. vol. iii. p. 553. 
Clathraria Mantelli, Schimper, ibid. p. 543. 
1875. Clathraria Lyelli, Topley, Weald, p. 409. 
1889. Clathraria Lyelli, Bristow, Geol. I. Wight, p. 258. 


Type. Pith casts and structureless casts of the cortical surface 
of stems. British Museum. 

In 1822 Mantell gave a brief description of some fossil stems 
from Tilgate Forest for which he proposed no name, but suggested 
that they might be allied to the Huphorbiacee, or possibly to certain 
arborescent ferns. Two years later Stokes and Webb proposed 
to include these fossils in the genus Clathraria, and gave them 
the name of C. anomala; the surface-markings suggested to them 
a resemblance to the recent Zamia and Cyeas. In the Geology 
of the South-East of England Mantell claims priority for his 
name of Clathraria Lyelli; he notes the occurrence of imperfect 
leaf fragments in association with the stems, and speaks of them 
as linear-lanceolate in shape. Having mentioned such plants as 
he considers most closely allied to the Tilgate fossils, Mantell 
adds: ‘‘the impressions of the petioles on the bark bear a great 
resemblance to those on the stems of Cycas revoluta and 
C. circinalis.” In a later work! the same writer speaks of the 
axis, roots, leaves, and probably fruit of Clathraria as having been 
discovered in close association or connection with one another. 
He gives a woodcut showing some of the long linear-lanceolate 
leaves attached to the stem, and remarks that impressions of such 
yucca-like leaves have often come under his notice. It is 
unfortunate that none of the specimens of Clathraria afford any 
evidence whatever as to the form of the leaves as described by 


1 Mantell, Medals, vol. i. p. 182. 


BUCKLANDIA. 125 


Mantell. In the younger stems Mantell notes that the internal 
axis (pith cast) cannot be distinguished from the outer cortical 
portion. 

Carruthers! gives the following detailed definition of Buchklandia 
anomala: ‘*Sears of the leaves subrhomboidal, the lateral angles 
more or less truncate, inferior angle acute, the superior obtuse or 
somewhat rounded. The surface of the scar in some specimens 
marked with a triradiate ridge. The smaller scars oblong, with 
blunt lateral angles, obtuse inferior, and slightly rounded, almost 
straight superior angle; the scars equal in breadth to the larger 
ones, but not nearly so deep; the cicatrix on the upper margin. 
The bases of the leaves are set somewhat obliquely on the stem, 
their upper margin following the direction of the right-hand 
spiral. Each series of leaves occupies a considerable length of the 
stem. The phyllotaxy is represented by the fraction -°;.”’ 

The new species, Bucklandia Mantelli, is thus described: 
‘Scars of the leaves rhomboidal, the lateral angles acute, the 
inferior and superior angles obtuse, the latter somewhat rounded. 
The small scars equal in breadth to the large ones, and increasing 
in depth from the bottom of the constriction upwards. Each 
swelling of the stem bearing three or four series of leaves, the 
constricted portion much longer, and crowded with the smaller 
scars, forming twelve or fourteen vertical series. The base of the 
leaves set horizontally on the stem. The phyllotaxy is represented 
by the fraction 3°.” It will be noticed that the more acute 
lateral angles of the leaf-scars and certain other slight differences, 
constitute the chief distinctive features of Bucklandia Mantelli, 
Carr. The close agreement between these two forms will be 
pointed out in the following descriptive notes on the British 
Museum specimens :— 


In the original figures of Bucklandia anomala a specimen is 
shown with the cortical portion of the stem separated by some 
coaly substance—no doubt the carbonized remains of the wood— 
from a central sandstone cast; the latter, with its ‘‘ interrupted 
longitudinal ridges,” being a cast of the pith cavity, and the ridges 
the impressions of spaces in the xylem cylinder which were 


1 Loc. cit. p. 686. 
2 Ibid. p. 686. 


126 BUCKLANDIA. 


originally occupied by the cells of the primary medullary rays. 
In speaking of the numerous specimens of these pith casts, 
Carruthers admits the impossibility of referring them with any 
certainty to their respective species of bucklandian stems, and 
suggests that probably they may belong to three or four distinct 
forms of the genus. Saporta’ has instituted a comprehensive 
genus, Cycadeomyelon, with the following definition: ‘‘ Medulla 
centralis primum substantie cellularis disperditione evanida, dein 
sedimento cylindrum lignosum intus vacuum cumulorite substitula 
et tune post ligni circumfusi abolitionem cylindrum plenum plus 
minusve compressum fasciculorum meatuumque impressionibus 
superficialiter notatum efformans.”’ 

In cases where we cannot be certain as to the relation between 
casts of the pith cavity and those of the cortical surface of a stem, 
it will be convenient to make use of Saporta’s genus as a useful 
designation to express the absence of sufficient data for any 
more accurate identification. In some specimens we have the 
clathrarian pith cast enclosed in a bucklandian cortex, but in 
most cases the internal and external casts have been separated. 
Although very probably, as Carruthers suggests, the detached 
pith casts belong to more than one species of Bucklandia, yet 
the very striking resemblance between those internal casts, which 
are still surrounded by the cortical surface, and the isolated 
specimens is sufficient reason for the inclusion of such forms 
under the present genus. Hosius and von der Marck* have 
described an Aptien fossil stem as Clathraria (?) galtiana, and 
compare it with Clathraria Lyelli as figured by Schenk*; it is, 
however, probably not identical with the English type, and 
should perhaps be referred to the genus /ittonza. 

8262. Figured by Stokes and Webb, Trans. Geol. Soc. [2] 
vol. i. pl. xlv. fig. 1. Mantell, Illust. Geol. Sussex, pl. 11. fig. 1.4 
Carruthers, Trans. Linn. Soc. vol. xxvi. pl. liv. fig. 3. 

From one end of the specimen the cast of the hollow pith 
projects beyond the encasing wood and cortex; it shows the 


1 Loc. cit. p. 381. 

2 (A. 1), Paleeontographica, vol. xxvi. pl. xlii. fig. 180. 

3 (A. 2), Paleontographica, vol. xix. pl. xxx. fig. 7. 

4 Mantell explains the repetition of Stokes and Webb’s plates in his ‘* Tllus- 
trations of the Geology of Sussex’’ in a note on page 52 of that work. 


BUCKLANDIA. 2A 


characteristic ridges of Clathraria. The petiole scars have 
rounded or truncate lateral angles, but those towards the lower 
part of the specimen have their lateral angles more acute; this 
is indicated in Carruthers’ figure on the left of the pith cast. 
Probably specifically identical with specimen V. 3309, figured 
by Carruthers as Bucklandia Mantelli. Tilgate Forest. 


8358. Figured by Stokes and Webb, Joc. ecié. pl. xlv. fig. 2. 
Mantell, loc. cit. pl. 11. fig. 2. 

The figure of this specimen does not do full justice to the 
details shown on the stem surface. If some of the scars be 
compared with the lower petiole bases of V. 3308 it will be found 
to be a matter of some difficulty, not to say impossible, to point 
to any distinct difference between the two forms, the former of 
which Carruthers speaks of as Bucklandia Mantelli, and the latter 
as B. anomala. 

A second smaller specimen in the form of a slightly com- 
pressed hollow cast of the bark with weathered petiole bases, and 
on the upper portion bearing scars of scale leaves. Very similar 
to Mantell’s pl. iii. fig. 4a. As shown in Mantell’s figure, there 
is a distinct projecting ridge above the petiole scar, and separated 
by a depression from the main part of the petiole; this is pro- 
bably due to the irregular or unequal weathering of scleren- 
chymatous and parenchymatous tissue. Cf. a stem of Cycas. 
Tilgate. 


V. 3308. Figured by Mantell, Geol. S.E. England, pl. i. fig. 2, 
and by the same author in Illust. Geol. Sussex, pl. i. fig. 2. 

Here again the figures do not do justice to the specimen. In 
this stem the petiole scars are clearly preserved and larger than 
in the other examples of the same species; they are not so much 
obscured, as in many specimens, by the irregular surface ridge 
and projections which Carruthers regards as the remains of a 
gummy exudation on the original plant stem. Some of the more 
prominent scales resemble the form which occurs in close associa- 
tion with V. 2749, and the lower portions of some of the larger 
scales suggest a reticulate marking like that in V. 2749 and in 
several of the isolated scales. The lateral angles of the scars are 
for the most part rounded or obtuse, but in some the angles 


128 BUCKLANDIA. 


are much more acute. The form of many of the leaf base scars 
points to a partially decayed petiole rather than a clean-cut 
surface of a persistent corky base. Cf. Carruthers, loc. cit. 
pl. liv. fig. 1; ef. also Fittonia. Tilgate Forest. 


V. 3309. Figured by Carruthers, pl. liv. fig. 4, as Bucklandia 
Mantelli. 

In this specimen the two kinds of scars are clearly shown, 
also the increased diameter of the axis where the large scars 
occur. The lateral angles of the scale scars are more obviously 


truncate than those of the petiole bases. Cf. 46644. Cuckfield. 


V. 3310. Figured by Carruthers, pl. liv. fig. 2. 

In describing this specimen Carruthers speaks of a ‘ triradiate 
ridge’? on the surface of the scars, but I am unable to recognize 
any such character; the markings are probably the result of 
some secondary changes and cannot be regarded as an original 
character. 


V. 713. This specimen shows the clathrarian pith cast, as in 
8262, but much more clearly. The flattened internal cast has 
a length of 45cm., and exhibits the usual surface features 
characteristic of Clathraria Lyelli. Part of it projects beyond 
the surrounding cortical cast, but it is enclosed to some extent 
by the remains of leaf bases, and between the cortical shell and 
the pith there is a space about 5mm. in width, here and there 
filled with coal; this no doubt represents the portion of the stem 
originally occupied by woody tissue. The pith case is flattened 
and shows alternations of broader and narrower portions; the 
surface markings, in the form of narrower and tapered ridges, 
do not appear to be disposed with any regularity. The large 
leaf bases are approximately 3 cm. in depth and 4:3 cm. in breadth. 
Some of the leaf bases have projecting upper surfaces, as in 
V. 3308, suggesting partially decayed petioles, but in other parts 
of the stem the scars are much flatter and more lke the clean- 
cut bases of detached fronds. 

V. 7182. 23cm. in length. In the broader and lower part 
of the specimen the petiole bases are fairly large, and have much 
the same form as in V. 718. 


BUCKLANDIA. 129 


V. 713d. Poor specimen. On one side an irregularly marked 
surface is exposed, which probably represents the impression of 
the external portion of the wood. Hastings. Dawson Coll. 


V. 213824. Imperfect impression of a stem. Shows some resem- 
blance to V. 2182, but differs in having two well-marked forms 
of scars, and in the absence of ramenta. Part of a scale in 
close connection with the stem, identical with V. 2699, ete. 
Mr. Rufford informs me that he has frequently found large 
scales, such as V. 2799, in association with this form of stem; 
the base of V. 2799 might well have been in attachment with 
scars like those of the present specimen. 


? V. 2182c. A small specimen showing petiole scars: this may 
be an example of Bucklandia, but it also closely resembles the 
genus Fittonia, and its precise nature must be a matter of un- 
certainty. Ecclesbourne. Rufford Coll. 


V. 2749. Small piece of a stem with two or three fairly distinct 
petiole scars, and some narrow and longer scars to which scales 
were probably attached. A single detached scale, with the 
reticulate surface markings, occurs in close association with the 
stem. Cf. Carruthers, loc. cit. pl. liv. fig. 4 (Bucklandia Mantelli). 


V. 2749a. In the lower part of the stem the petiole scars are 
shown with a flat surface, and higher up there appear to be 
casts of the lower portions of the petioles; the latter bear a 
strong resemblance to some of the smaller examples of scales. 
This, like many other specimens, shows a general resemblance 
to Lttonia. Ecclesbourne. Rufford Coll. 


8372. Small specimen showing petiole scars. Tilgate Forest. 
46644. A portion of a small stem with the scars characteristic 
of Bucklandia. Tilgate Forest. Bowerbank Coll. 


V. 3311. Species showing narrow scale scars. Cf. Carruthers, 
loc. cit. pl. liv. fig. 4. 


K 


130 BUCKLANDIA. 


Cycadeomyelon (Bucklandia), sp. 


V. 2332. Probably a portion of a pith cast; but the larger 
medullary ray impressions suggest some other stem than that 
with which the ordinary Clathraria Lyelli is occasionally found. 


V. 2804. Considerably flattened sandstone cast. The form of 
the medullary ray impression and of the leaf-trace bundle is more 
distinctly shown than in most specimens. Schenk’s figure also 
shows this feature clearly. The form of the medullary ray casts 
in Weiss’ genus Tylodendron is very similar to that of the cor- 
responding structures in the present specimen. Good figures of 
Tylodendron will be found in a paper on that genus by Potonié.! 
Cf. also similar casts in Voltzva. 


V. 28040. Similar specimen, but less flattened. Part of a scale 
at one end of the cast. Ecclesbourne. Rufford Coll. 


8268, 12333. Portions of pith casts; probably of the lower 
portion of the stem axis. Tuilgate Forest. 


8264. Figured by Mantell, Illust. Geol. Sussex, pl. i.: 93cm. 
in length; at broadest part the diameter is 13cm. The details 
of surface characters not very clearly preserved. ‘Tilgate Forest. 

Mantell Coll. 


8269. 79cm. long, diameter 13cm. A large branched specimen, 
showing at the summit an apparently dichotomous bifurcation. In 
addition to the large branches, there are the scars of six or seven 
smaller lateral branches, about 2cm. in length; the smaller 
sears are all on one side of the specimen and arranged in a 
fairly regular line. Cf. Goppert’s figure of Cycas revoluta with 
the numerous large and small branches.” Tilgate Forest. 

Mantell Coll. 


8274. 78cm. long, 7°5cm. in diameter. The breadth varies, 
but there is no regular alternation of narrower and thicker 
portions. Tilgate Forest. Mantell Coll. 


1 Potonié (2). 
* Goppert (3), pl. ix. fig. 3. 


FITTONIA. 131 


V. 3307. A long specimen with distinct constrictions at fairly 
regular intervals; the surface projections are rather broader than 
in many examples, and resemble those in V. 2332; they show 
a considerable variation in size, some being identical with the 
usual clathrarian form of medullary ray cast, and others larger 
and more prominent. 


V. 3306. Here again there is a marked variation in the size of 
the ray impressions. A well-defined branch scar. 


V. 713¢c. Hastings. Dawson Coll. 
V. 1880. Near Hastings. Dawson Coll. 
V. 2249. Ecclesbourne. Rufford Coll. 


8272. Large specimen. 


Genus FITTONTA, Carruthers. 


[Trans. Linn. Soc. vol. xxvi. 1870, p. 690.] 


This generic name was proposed by Carruthers for an unusually 
perfect specimen of a structureless cycadean stem, which was 
probably obtained from the Wealden of Brook, in the Isle of 
Wight. Mantell first figured and described this specimen as 
Clathraria Lyelli in his Geological Excursions round the Isle of 
Wight*; a larger and more complete figure appears in Carruthers’ 
Monograph. The genus is thus defined :— 

‘“‘Trunk short, obovate ; woody axis slender, enlarging upwards ; 
cortical layer large. Scales and bases of the petioles large, 
imbricated, at first reflexed, then ascending.” 

Carruthers notes the absence of any fruit or foliage which 
can be referred to this form of stem, but he considers the 
resemblance to Encephalartos sufficiently distinct to justify him 
assigning Fvttonia to a position near to the living genus. Saporta 
adopts the name, and points out the resemblance between Jttonia 
and Bucklandia; he calls attention, however, to certain points 


1 p. 297. 


132 FITTONIA. 


of difference, and considers that Mantell’s specimen has been 
correctly made the type of a new genus. In Bucklandia the 
stem appears to have been frequently branched, but no such 
habit is indicated in the specimens of /%ttonia; in the latter 
genus the increase in size both of the petiole and scale-leaf 
bases, and of the lower portion of the petioles as well as the 
scale leaves, with the subsequent disarticulation of the upper 
part of the frond axis, constitute constant and characteristic 
features. The pith is large and surrounded by a narrow zone 
of wood. The scale leaves, associated here and there with the 
bases of fronds, are distinguished from the latter by their 
thinner distal margins, and the absence of any distinct surface 
of articulation. As Saporta remarks, there is a striking re- 
semblance between some forms of /ttonia and Bucklandia; this 
similarity has already been noted in the descriptions of some 
of the Museum examples of the latter genus, and it is also 
clearly seen in Carruthers’ type specimen in the Jermyn Street 
Museum of Practical Geology. It may be that the two genera 
are not really distinct, but merely represent different forms of 
preservation of very similar, if not identical, plants. In describing 
the single English specimen of JS%ttonia, Mantell quotes the 
opinion of Brongniart,’ to whom a drawing of the fossil was 
sent, that it 1s probably the upper portion of a clathrarian 
stem with persistent petioles. It may be more convenient, with 
the evidence at present available, to retain both generic names, 
and to make use of F’ttonia as a useful designation for a certain 
form of cycadean trunk. 


Fittonia Ruffordi, sp. nov. 
[Pl. IX. Fig. 6.] 


Type. Large impressions of stems. British Museum. 

The lower portion of the petioles persistent, showing a well- 
marked surface from which the upper part of the frond has 
been detached. The persistent and swollen bases are regularly 
disposed in the stem surface, and apparently without any 
alternation of petiole scars and scale-leaf scars. 


1 Mantell (A. 7), Geol. Excurs. I. Wight, p. 298. 


FITTONIA. 133 


V. 2238. Pl. IX. Fig. 6. 

18cm. in length, 10cm. broad. An impression of part of the 
lateral surface of a cycadean stem, covered with a carbonaceous 
layer. The petiole bases are shown with unusual definition, and 
the form of the surface of articulation is particularly clearly 
marked. The swollen portion or cushion below the petiole scar 
presents a fairly close resemblance to some forms of Sigillaria 
Brardii, Brong., with its similarly situated leaf-scar, which 
agrees closely in shape with that of Fittonia. The petiole base 
or cushion has a length of 1:‘7cm. The form of the leaf-scar 
is practically identical with that of the cleanly-cut end of the 
frond of Otozamites Goppertianus as figured in Pl. I. Fig. 2. 
Cf. Fittonia squamata, Carr.,1 and F. insignis, Sap.*; the present 
species differs from these in the smaller size of the leaf base and 
in their much more regular and uniform arrangement on the 
stem. Ecclesbourne. ° Rufford Coll. 


V. 2121. 77cm. long, 8-9 cm. broad. Here again the surface 
of the stem has been carbonized, and the petiole bases present 
a similar appearance to those in V. 22388, but are rather less 
perfectly preserved. The comparatively long and narrow form of 
the leaf base is clearly shown. LEcclesbourne. Rufford Coll. 


V. 2244. 35cm. long. Probably this may be referred to 
Fittonia Ruffordi, but the petiole bases are much less clearly 
preserved. In all the specimens the characteristic feature is the 
ereat length as compared with the breadth of the petiole bases. 
Ecclesbourne. Rufford Coll. 


?V. 3181. Possibly a specimen of this species. Ecclesbourne. 
Rufford Coll. 


1 Carruthers (1), pl. lvi. 
2 Saporta, Pal. Franc. vol. ii. pl. cxxv. figs. 1-3. 


134 BENNETTITES. 


Genus BENNETTITES, Carruthers. 
[Trans. Linn. Soc. vol. xxvi. 1870, p. 681.] 


In 1855 the President of the Linnean Society of that year, 
Robert Brown, exhibited a new form of cycadean stem which 
had been found by Saxby at Bonchurch, in the Isle of Wight.’ 
The name Cycadites Saxbyanus was suggested by Brown for this 
new fossil, of which the two most striking features were the 
elliptical outline of the stem as seen in transverse section, and 
the presence of a bud in the axil of each leaf. Fifteen years 
after the discovery of this stem, Carruthers” published a full and 
scientific description of several examples of the same species. 
As a result of careful investigation of the morphology of these 
specimens, Carruthers instituted the generic name Bennettites 
for the new form of stem, which he found could not be classed 
under any of the recognized subdivisions of the Order Cycadacee. 
After speaking of the elliptical form of the axis and the presence 
of buds in the axils of many of the leaves, Carruthers proceeds 
to describe the anatomical structure of the stem, and calls special 
attention to the leaf-traces as affording another characteristic 
feature of the genus. ‘‘In all the known members of the Order 
(Cycadacee),’* says Carruthers, ‘‘the leaf-traces arise in the 
interior of the cylinder of wood, as bundles of small size, and, 
passing through the meshes of the ligneous cylinder, and then 
through the cortical parenchyma, as small distinct bundles, after 
running for a short distance, at least in some genera, in a 
horizontal direction parallel to the periphery of the stem, they 
pass in the petiole of the leaf.”” In ennettites, however, ‘the 
vascular tissue for each leaf springs from the woody cylinder in 
a single large compact bundle, which as it passes outwards breaks 
up into the different bundles required for the service of the leaf.” 
This peculiar behaviour of the leaf-traces is also referred to 
by Solms-Laubach* as an important distinctive character of 


1 Brown, p. 180. 
oa (LP 

3 Lbid. 

& (2), p. 422. 


BENNETTITES. 135 


Carruthers’ genus. The structure of the petiole bases and the 
ramenta-like scales has been fully described by Carruthers and 
others. In the axils of some of the petiole bases which surround 
the woody axis of the stem, there occur the lateral branches 
which constitute the most important and interesting feature of 
the genus. ‘These organs,” writes Carruthers, ‘‘are not 
properly buds, for although they do not appear to have pushed 
themselves beyond the surfaces of the permanent bases of the 
leaves, they are fully developed organs, and differ from the 
secondary axes of Jfantellia, which are generally broken off 
beyond the surfaces of the permanent bases of the petioles, and 
show there a woody cylinder agreeing in structure with the 
principal axis of the plant. The secondary axis consists of a 
very short and slender stem, bearing a number of simple lnear 
acuminate leaves. These are the only foliar organs hitherto 
found connected with these fossils.’ Each of the axillary 
branches terminates in a subpyriform enlargement, bearing seeds ; 
this terminal portion with its seeds are fully described and 
illustrated by Carruthers and Solms-Laubach. The former 
author, in summing up the affinities of the genus, expresses 
the opinion that ‘‘it must be considered to hold the same 
relation to the other Cycadee, that Taxus, with its succulent, 
cup-shaped pericarp does to the cone-bearing Conifere.”’* 

In his Evnleitung in die Paliéophytologie Graf Solms 
confirms many of the characteristics of Lennettites as described 
by Carruthers, and by a careful examination of the English 
material he is able to settle certain doubtful points, and to carry 
a stage further our knowledge of this interesting type.® <A 
more detailed account was afterwards published by Solms in 
the Botanische Zeitung,* and the article subsequently appeared 
in English in the fifth volume of the Annals of Botany.’ 
Without entering at length into a histological description of. 
Bennettites, we may call attention to some of the more striking 
and characteristic features. The structure and course of the 


1 Carruthers, oc. cit. p. 697. 
2 Tbid. p. 698. 

3 Solms-Laubach (A.), p. 94. 
4 Ibid. (1). 

© bid, (2). 


136 BENNETTITES. 


leaf-trace as described by Carruthers, and the analogy which 
he noted between the trellis-work form of the vascular cylinder 
and the bundle system of a fern stem are confirmed by Solms; 
he adds, however, that a ‘closer examination will doubtless 
disclose a greater affinity with the course of the vascular 
bundles in many Conzfere.”! The pith and cortex are traversed 
by numerous and large gum-canals, and the stems are enclosed 
in the armour of leaf bases characteristic of recent cycads. 
Between the petioles of Bennettites and those of living cycads 
there is the closest agreement; the bases of the leaves are 
separated from one another by a felt of ramenta-like outgrowths 
from the petiole surface. Occasionally the substance of the petiole 
has rotted away, leaving deep cavities occupying the meshes in 
a network of intervening projecting ridges. In Bennettites 
Gibsonianus the structure of the lateral branches bearing the 
remarkable form of fructification is very perfectly preserved, 
and it is evident that we have to deal with a plant differing 
in many important respects from the present types of cycads. 
Referring to the cycadean character of the stem and leaf-stalk 
of Bennettites, Solms remarks: ‘‘ We arrive at the surprising 
result that all the Jurassic and Neocomian stems which are 
termed Cycas-stems, so far as anything is known of their 
structure, belong to Bennettitee, and that not a single one of 
them has been proved to be a genuine stem of Cycadee. This 
further shows how precarious is the identification of fossil remains 
when it rests on superficial characters only.”* The form of the 
lateral fructification branches may be summarized as follows: 
They occasionally arise exactly over a leaf base, but in some 
cases have been pushed somewhat to the side; as a rule it 
appears impossible to ‘‘determine the precise relative position 
of the two kinds of organs.”? The flower-bearing shoots are, 
at all events, not terminal structures, as in recent cycads. Each 
fertile branch is made up of several short internodes, and bears 
spirally-arranged lanceolate acuminate bracts; in structure the 
bracts agree for the most part with the larger leaf bases of the 


1 Solms-Laubach (2), p. 422. 
2 Ibid. p. 424. 
3 p. 431. 


BENNETTITES. Sy/ 


main stem. The apex of the fertile shoots has the form of a 
hemispherical parenchymatous cushion, from which are given off 
a number of closely crowded stalks united into a club-shaped 
group; between these stalks or seed-bearing cords there occur 
smaller structures, the so-called interstitial organs. External to 
these is a comparatively broad parenchymatous band of tissue, 
which arises from a lower level on the cushion than the 
cords and interstitial organs. Surrounding this homogeneous 
peripheral tissue we have several lanceolate bracts encircling 
the entire fructification. Towards the upper surface of the 
spadix, and near its periphery, there are numerous seeds situated 
just internal to the homogeneous tissue, which becomes rather 
more strongly developed towards the apex of the fructification. 
Each seed lies in a flask-shaped pit, and is borne on a long 
stalk or cord; in some seeds the structure of the embryo with 
its radicle, plumule, and two cotyledons, may be clearly 
recognized. Solms suggests that the homogeneous tissue which 
overtops the spadix and contains the seeds, has been formed 
by the union of interstitial organs. A surface view of the 
terminal portion of the fructification would present a number 
of areole, probably raised to the form of pyramidal projections, 
and between these areole or distal ends of interstitial organs, 
there would be seen the narrow openings leading to the flask- 
shaped seed cavities. 

In his description of Bennettites, Solms compares its fructifica- 
tion with that described by Saporta and Marion as Williamsonia 
Morieret from the Oxfordian of Vaches-Noires. This French 
specimen, with its histological details preserved in unusual 
perfection, has recently been thoroughly examined and described 
by Prof. Lignier, of Caen. His description clearly shows that 
Saporta’s and Marion’s species must be referred to Bennettites, 
and is closely allied to the English species of that genus.! In 
the main Lignier’s description agrees with those of Carruthers 
and Solms; there are, however, one or two points in which 
the French author’s account is slightly at variance with that 
of the former writers. Bennettites Morierei has the form of 
a detached ovoid fossil, with a length of 55mm. and a breadth 


1 Lignier. See also Seward (3) for a short abstract of Lignier’s paper. 


138 BENNETTITES, 


of 35mm. At the base a fractured surface reveals the existence 
of a slightly convex receptacle, from which is given off a compact 
cluster of long peduncles, each of which bears at its apex an 
oval seed. The seed-bearing peduncles are surrounded by several 
involucral bracts closely applied to the surface of the fruit. 
Numerous thin lamelle occur in association with the seminiferous 
peduncles; to these Lignier applies the term interseminal scales. 
The seeds are arranged side by side close to the upper surface 
of the mass of peduncles and interseminal scales; the latter 
passing between and beyond the seeds, and their swollen distal 
ends forming a protective covering to the blunt hemispherical apex 
of the fruit. In surface view, the upper part of the specimen 
appears to be made up of a large number of small projecting 
areas with polygonal bases and rounded summits. Here and 
there the projections arrange themselves in the form of rosettes 
round a small central cavity, marking the position of a seed. 

As in Bennettites Gibsonianus the fruit is covered by involucral 
bracts, but we have the interesting suggestion of Lignier that 
these were not simply linear acuminate in form, but that the 
portion of the bracts preserved is merely the petiolar part of a 
leaf structure of which the pinnate or flabellate lamina has been 
detached. This inference is drawn from a detailed examination 
of the vascular strands traversing each bract. After comparing 
and contrasting Bennettitee with the Cycadacee and Conifere, 
Lignier concludes that they represent ‘‘a family which has been 
derived with the cycads from common ancestors, but not from the 
cycads themselves. Of these common ancestors the two families 
have preserved the form of the trunk, the structure of certain 
tissues, the foliar origin of the ovule, ete. But whilst the cycads 
have retained a grouping of carpophylls on a single axis, and 
have acquired special characters, such as the complication of the 
leaf-trace and the lateral position of the ovules, the Bennettitee 
have retained the simple leaf-trace and have acquired a terminal 
position of the ovules, the reduction of the fertile axes to single 
carpophylls, the grouping of these fertile reduced axes, and the 
modification of the neighbouring leaves.”’! 

In a recent paper on fossil cycadean trunks from North 


1 Lignier, loc. cit. p. 73. 


BENNETTITES. 139 


America, Lester Ward! transfers Carruthers’ species of Bennettites 
to Buckland’s genus Cycadeotdea; he considers the fact of the 
fruit being known in a single species, B. Gibsonianus, is not a 
sufficient reason for the retention of the genus. Solms-Laubach * 
had previously restricted Carruthers’ term to the species in 
which the characters of the fructification are known, viz. 
Bennettites Gibsonianus; this use of the term is probably the 
most convenient, and it would seem much better to retain 
Carruthers’ name for stems bearing the bennettitean inflorescence 
than to include these under such a comprehensive and purely 
provisional genus as Cycadeovdea. 


Bennettites (Cycadeoidea) Saxbyanus (Brown). 


1851. Cycadites Saxbyanus, Brown, Proc. Linn. Soe. vol. ii. p. 130. 

1854. Cycadeoidea Saxbyana, Morris, Brit. Foss. p. 7. 

1870. Bennettites Saxbyanus, Carruthers, Trans. Linn. Soc. vol. xxvi. p. 698. 
1874. Bennettites Saxbyanus, Schimper, Trait. pal. vég. vol. i. p. 558. 
1878. Bennettites Saxbyanus, Carruthers, in Dixon’s Geol. Sussex, p. 281. 
1894. Cycadeoidea Saxbyana, Ward, Biol. Soc. Washington, vol. ix. p. 80. 


Type. Stems showing internal structure. British and Oxford 
Museums. 

The specific name of this plant was proposed by Robert Brown 
in honour of Mr. Saxby, who found the first specimen near 
Bonchurch, in the Isle of Wight. The following definition of 
the species was given by Carruthers :— 

“Trunk elliptical, with large medulla, and thin, much inter- 
rupted woody cylinder, vascular bundles passing upwards and 
outwards and breaking up into two rows of small bundles, which 
are parallel to the superior and inferior surfaces of the petiole ; 
section of petiole subtriangular.” 

In accordance with the narrower use of the genus Lennettites, 
as suggested by Solms and others, the present species ought to be 
transferred to Buckland’s Cycadeoidea, but for the present it may 


PWrardy (i) pais. 
? Capellini and Solms-Laubach, p. 29. 


140 BENNETTITES. 


be better to retain Carruthers’ generic name, and by the addition 
of Cycadeoidea to indicate our incomplete knowledge of the 
fructification. Cycadeoidea is employed, therefore, in the wider 
sense of the term, including stems with a circular as well as an 
elliptical transverse section. 


46628 (V. 32331). Figured by Carruthers, Trans. Linn. Soc. 
vol. xxvi. 1870, pl. lvil. fig. 3, as Bennettites Saxbyanus. 

Length 21cm.; at one end the specimen shows the transverse 
and polished section as represented in Carruthers’ figure; at the 
opposite end the stem has been considerably worn. The varying 
degrees of wearing in this specimen show remarkably well the 
striking differences in appearance presented by the several surfaces 
of cortical or woody tissue exposed to view. Carruthers calls 
attention to the striking similarity ‘‘between these fossil stems 
and the caudex of a tree fern’’*; the U-shaped petiolar bundles 
shown in section on part of the specimen present a distinct 
resemblance to the corresponding leaf-traces in fern stems. Brook 
Point, Isle of Wight. Saxby Coll. 


V. 3234. Figured by Carruthers, Joc. eit. pl. lvii. fig. 4. 

The surface of the specimen shows fairly good sections of 
petioles with numerous vascular bundles; the portion represented 
in Carruthers’ figure shows the inner face of the wood, and a 
radial section through the cortex in which the course of the 
leaf-trace bundles is clearly seen. I am unable to detect any 
difference between the present specimen and that bearing the 
number 88360, which has been labelled by Carruthers Bennettites 
Gibsonianus. In his definition of B. Saxbyanus, Carruthers speaks 
of the petioles as subtriangular in section, whilst those of 
B. Gibsonianus are described as subquadrangular. Each of the 
above specimens shows both forms of petioles, and in other respects 
there appear to be no real differences. 


V. 3235. Figured (in part) by Carruthers, Joc. cit. pl. vii. fig. 7. 
The figure shows very clearly the form of the petiole bundles 
as seen in a tangential section of the cortex. The resemblance 


1 A second number recently added (1895). 
? Carruthers, Joc. cit. p. 696. 


BENNETTITES. 141 


between these bundles and the leaf-traces of Protopteris punctata, 
Sternb., is fairly close. Other portions of this specimen demon- 
strate very clearly the mode of origin and course of the petiole 
bundles. The tangential section of the wood, close to the inner 
(pith) face, presents the same appearance as figured by Carruthers 
(pl. lvii. fig. 6). 


V. 3236. Figured by Carruthers, loc. edt. pl. Ivii. figs. 1 and 2. 

The figured portion is the smaller half of a fine specimen in the 
Museum Collection. The surface is made up of triangular cavities 
which constitute the meshes in a network of projecting ridges of 
interpetiolar ramenta. A few inflorescences, or rather the cavities 
originally occupied by inflorescences, are seen in surface view, 
and also in the longitudinally cut face represented in Carruthers’ 
figure. The woolly or finely fibrous ramenta appear to be identical 
with the corresponding interpetiolar structures in some specimens 
in the Rufford Collection from Ecclesbourne (cf. V. 3177, V. 2349, 
and V. 2182). There is a distinct similarity between the slightly 
raised base forming the floor of one of the inflorescence cavities, 
and the rounded receptacle seen in some of the Ecclesbourne 
examples of isolated inflorescences (Bennettites [| Williamsonia] 
Carrutherst), e.g. V. 21296, ete. Brook Point. 


38363. A waterworn specimen showing petiole base projecting 
somewhat beyond the ramental network. Cf. V. 3234 (pl. lvu. 
fig. 4) and 38360. Solms’ refers to this and some other specimens 
as no doubt examples of Bennettitee, but without traces of fructifi- 
cation. An old label on the specimen gives Tilgate as the locality ; 
this has been altered to Brook. Brook. Mantell Coil. 


cf. Bennettites Saxbyanus. 


V. 2132. Part of a large stem, resembling the specimen of 
Bennettites Saxbyanus figured by Carruthers in pl. lvii. fig. 3. 
The external surface shows numerous imperfectly preserved leaf 


1 Solms-Laubach (2), p. 426. 


142 BENNETTITES. 


bases. Between the petioles there appears to have been a fairly 
wide space occupied by ramenta, A side view of the specimen 
shows the inward prolongation of some of the petioles, and 
demonstrates that the so-called bases as seen on the surface 
simply represent the level in the leaf armour to which dis- 
organization has extended. The inner face of the block is probably 
the outer surface of the wood. No fructification seen. LEccles- 
bourne. Rufford Coll. 


Bennettites Gibsonianus, Carr. 


1851. Clathraria Lyellii, Mantell, Petrifactions, p. 46. 

1854. Clathraria Lyellii, Mantell, Medals, vol. i. (edit. 2), p. 163. 

1854. Clathraria Lyellii, Mantell, Geol. Excurs. I. Wight, p. 214. 

1870. Bennettites Gibsonianus, Carruthers, Trans. Linn. Soc. vol. xxvi. p. 700. 

1874. Bennettites Gibsonianus, Schimper, Trait. pal. vég. vol. ili. p. 559. 

1878. Bennettites Gibsonianus, Carruthers, in Dixon’s Geol. Sussex, p. 281. 

1890. Bennettites Gibsonianus, Solms-Laubach, Bot. Zeit. 1890, p. 789, pls. 
ix. and x. 

1891. Bennettites Gibsonianus, Solms-Laubach, Annals Bot. vol. v. p. 419, 
pls. xxv. and xxvi. 

1894. Bennettites Gibsonianus, Lignier, Mém. Soc. Linn. Normandie, vol. 
Xvili. p. 76. 

1894. Cycadeoidea Gibsoni, Ward, Biol. Soc. Washington, vol. ix. p. 80. 


Type. Portions of a large block showing internal structures. 
British Museum and Kew. 

The specimens on which Carruthers founded this species are 
recorded by him as of Lower Greensand age, and Solms-Laubach 
has since confirmed this determination.1 The examples of 2. 
Saxbyanus are regarded as Wealden, and were obtained from Brook 
Point, in the Isle of Wight; 2B. Gcbsonianus was found in Luccomb 
Chine, Isle of Wight, by Mr. Gibson. Some of the specimens 
which Carruthers has referred to the present species are apparently 
from Brook, and from the same beds as B. Saxbyanus. 

Carruthers defines the species as follows :— 

“Trunk compressed, elliptical, with small medulla, and a thick 
subcontinuous woody cylinder; vascular bundles passing almost 


1 Solms-Laubach (2), p. 429. 


BENNETTITES. 143 


directly outwards and breaking up into a double series of small 
bundles, which are parallel to the superior and inferior surfaces of 
the petiole, except that a loop is sent down from the upper series 
into the centre of the petiole. The section of the petiole is sub- 
quadrangular.” 

Lignier has recently extended this definition to include the 
characteristic features of the inflorescence ; he makes the following 
addition to the above diagnosis ' :— 

“ Fruit haut de 3 centim. 4. Bractées involucrales lancéolées, 


acuminées, dépourvues de limbe. Poils écailleux peu larges, 
souvent ventrus, 4 cellules allongées perpendiculairement aux 
faces. Graines longues de 3 A 4 millim., larges de 1:2 a 2°5, non 
anguleuses (ou & peine anguleuses), dépourvues d’épiderme 
rayonnant. Assise réticulée scléreuse et formée de cellules qui 
ont 60 de longeur sur 50 de large.” 

The following specimens are included here as somewhat doubtful 
examples of Bennettites Gibsonianus :— 

38360. Figured by Mantell as Clathraria Lyellit, Medals of 
Creation, vol. i. 1854 (edit. 2), p. 163; Petrifactions and their 
Teachings, 1851, p. 46; Geological Excursions round the Isle of 
Wight, 1854, p. 214. 

The old label on this specimen gives Tilgate Forest as the 
locality, but Carruthers has substituted Brook Point. The pre- 
servation of this waterworn example is very different from that 
of the larger blocks of Bennettites. The surface, as figured by 
Mantell, shows the characteristic ramental network with the 
meshes occupied by petiole bases. Towards the right-hand upper 
corner of the specimen there appears to be an inflorescence shown 
on the waterworn surface. In the absence of any well-defined 
inflorescence, and in view of the striking similarity to, if not 
identity with, V. 3234 (labelled by Carruthers B. Saxbyanus), it 
will be better to regard the exact position of this specimen as 
somewhat doubtful. Solms? quotes Mantell’s description of this 
fossil, and says he believes he recognizes the specimen in the 
British Museum (Geological Department) Collection as one on which 
the word ‘‘ Brook” is written in ink on the upper surface. There 


Lignier (1), p. 76. 
Loc. cit. p. 420. 


1 
2 


144 BENNETTITES. 


can be no doubt as to the original of Mantell’s figure; the fact 
of its being figured is recorded on the label, but there is no word 
‘‘Brook”’ written on it. No doubt Solms is referring to some 
other specimen; in another place the same author speaks of 
Mantell’s figure as a very good representation of a specimen in 
the Geological Department. 


38361 and 88362. Solms' refers to this specimen (cut into two 
pieces, bearing the above registered numbers) as being without 
fructification. On the smaller piece there appears to be inflores- 
cences shown in transverse and longitudinal section. The label on 
one piece bears the name Clathraria Lyellii. Possibly this specimen 
should be referred to Bennettites Saxbyanus. 


V. 82382. Another waterworn specimen cut into three pieces, 
one transverse slice and two larger portions. It would seem 
impossible to definitely refer this imperfect example to one or 
other species of Bennettites. 


Bennettites (Cycadeoidea), sp. 
[PLAX] 


V. 3177. The chief interest of this specimen is in the numerous 
casts of inflorescences which occur on the stem. The surface 
features cannot be accurately made out; the portions shown in 
Pl. XV. Fig. 1 exhibit the conical cavities originally occupied 
by the fertile axes, and the impressions of petioles and ramental 
tissue. The whole surface presents a somewhat waterworn 
appearance, and instead of showing a surface view of the petiole 
bases in the form of rhomboidal sections, it has the form of 
a worn-down surface with an oblique view of petiole casts and 
ramental tissue. 

In longitudinal section the fertile axes have the form of cavities 
narrowed towards the distal end; these cavities were no doubt 
originally occupied by the fleshy terminations of inflorescences. 
The wall of such a cavity (Pl. XV. Fig. 3) shows clearly preserved 


1 Loe. cit. p. 426. 


BENNETTITES. 145 


casts of the bases of crowded involucral bracts. The bracts them- 
selves are represented partly by cavities and partly by carbonaceous 
matter; between the bracts the woolly or hair-like ramenta 
are distinctly shown, and these, as frequently happens in such 
tissue in cycadean stems, have been permeated by a mineralizing 
solution, and so preserved. The bracts appear to be somewhat 
expanded distally (Pl. XV. Fig. 2), as described by Lignier 
in Bennettites Moriérei (Sap. and Mar.).!' In Pl. XV. Fig 3 an 
inflorescence cast is shown, of which the surface is covered by 
a fine and well-marked reticulation; in the upper portion of the 
cavity this reticulum has the form represented in Fig. 5, a fairly 
regular network formed by thin plates projecting from the surface, 
and some of the meshes are partially filled by round or oval 
bodies suggesting very small seeds; the meshes thus filled are 
less angular than those without seed-like bodies. In Fig. 4 is 
shown a wax cast of the inflorescence cavity of Fig. 3, and in 
Fig. 6 the small dot-like depressions on the surface of the 
cast correspond to the small bodies in the meshes of Fig. 5. 
Towards the base of Fig. 4, the reticulum is rather larger, and 
there are no dots in the more basal portion, owing to the absence 
of any ‘‘seeds”’ in this part of the inflorescence (Fig. 7). In the 
middle of one side of this reticulately marked cavity there is a 
narrow longitudinal ridge, probably representing a median groove 
in the inflorescence. The portion of inflorescence figured in Pl. X. 
Fig. 4, should be compared with the present specimen. Further 
reference is made to this stem in the description of the detached 
specimens referred to a new species, Bennettites Carrutherst. Cf. 
Pl. X. Figs. 1 and 4, and Saporta’s figures of Williamsonia gigas, 
Carr., in the Pal. Frang., vol. iv. pl. xiii. fig. 2, and pl. xiv. 
figs. 1 and 2. Keclesbourne. Rufford Coll. 


V. 2896. Imperfectly preserved stem, 26cm. in length, and 
20cm. in broadest part; leaf-stalks and ramenta shown, but no 
inflorescence. Beckles Coll. 


V. 2816. Smaller specimen; no actual stem surface visible. 
The woolly ramenta suggest a connection with V. 2182 and 
B. Saxbyanus (Brown). Ecclesbourne. Rufford Coll. 


1 Lignier (1), p. 26. 


146 BENNETTITES. 


Bennettites (Williamsonia). 
FLORES. 


In the Rufford collection of Wealden plants from the neighbour- 
hood of Hastings, there are several specimens which must be 
assigned to the same position as the well-known Jurassic William- 
sonia. For reasons which are stated more fully below, I have 
referred these Wealden fossils to the genus Bennettites, and am 
led to regard them as portions of the inflorescence of that plant. 
Hitherto typical Williamsonias have not been recorded from any 
Wealden or Lower Cretaceous rocks in England ; the importance 
of the discovery is considerably increased by the fact that the 
specimens appear to throw some new light upon the nature and 
affinity of this anomalous form of inflorescence. Before describing 
the individual fossils in detail, it may be convenient to give a 
short summary of our present position with regard to the opinions 
of palzobotanists on the nature of Williamsonia. 

In A Geological Survey of the Yorkshire Coast, by Young and 
Bird, published in 1822, there is a figure of a specimen from 
the ironstone of Saltwick, which is spoken of as resembling the 
head of an artichoke (Cynara integrifolia), with the ‘covering 
or calyx consisting of numerous lanceolate and striated leaves.’’! 
Another figure in this work represents ‘‘a petrified nut of a 
singular kind”’; these two fossils are examples of what was 
subsequently named Williamsonia.? 

Part of a cycadean frond from the ‘‘ Oolitic rocks of Scarborough” 
is figured by Lindley and Hutton, and named by them Zamia gigas*; 
a few years later Williamson notes the occurrence with this form 
of frond of ‘‘a remarkable fossil, apparently connected with the 
fructification of a Cycas.’* In 1849 Yates® draws attention 
to the identity of Zamia gigas, L. and H., Zamia Mantelli, Brong., 
and Cycadites lanceolatus, Phill. He recognizes the difficulty of 
connecting the leaves and stem with the peculiar form of in- 
florescence associated with them, and while favouring the view 


1 Young and Bird, pl. ii. fig. 6, p. 183. 
2 Thid. pl. iii. fig. 7, p. 186. 

3 Fossil Flora, vol. ii. pl. elxy. 

4 Williamson (1), p. 230. 

5 Yates. 


BENNETTITES. 147 


that Zamia gigas and the fructification are parts of the same 
plant, admits the absence of any actual proof. In the same year 
Mantell! figures a specimen of Williamsonia in his Medals of 
Creation, as the fruit of Zamites lanceolata, and describes the 
lanceolate involucral bracts as concealing the seeds of the in- 
florescence. In a short communication to the Yorkshire Philo- 
sophical Society, Williamson gives a vertical section of a restored 
inflorescence, and represents the ovoid body of the fructification 
as terminating in a cone.” Brongniart calls attention in his 
Tableau,? to the similarity between the Scarborough fossils and 
a specimen described by Buckland‘ from the Inferior Oolite of 
Charmouth, Dorset, under the name of Podocarya. This com- 
parison is one which has, I believe, been justified by recent 
investigations, and it is highly probable that Buckland’s specimen 
is a particularly well-preserved bennettitean inflorescence. It is 
to be hoped that this valuable specimen may be rediscovered,’ 
and subjected to a careful examination. 

The name Podocarya was chosen by Buckland for this Oolitic 
fossil, on the suggestion of Robert Brown. It is described as 
chiefly resembling the inflorescence of the recent genus Pandanus, 
and by most subsequent writers it is included in the Pandanacee. 
Saporta,® in his volume on Types proangiospermiques, reproduces 
Buckland’s figures, and substitutes Williamsonia for Podocarya 
as the generic name; Unger’ had previously named the species 
after Buckland. 

In a specimen figured by Leckenby in 1864° we have leaves 
of Paleozamia pecten (Lind.) in close association with a small 
form of Williamsonia, which Nathorst afterwards referred to 
the new species Walliamsonia Leckenbyi. Carruthers,’ writing 


1 Mantell (1), p. 161. 

2 Williamson (2), p. 47. 

p. 88. 

Buckland (2), vol. i. p. 466, pl. Ixxxiv. 

It is said to be in the Oxford Geological Museum, but cannot be found. 
Pal. Frang. vol. iv. p. 127, pl. cexxxviil. figs. 1-3, and pl. cexxxix. fig. 1. 
Unger (A. 2), Gen. spec. plant. foss. p. 327. 

8 Leckenby (A.), Quart. Journ. Geol. Soc. vol. xx. p. 77, pl. ix. fig. 4a. 
_This specimen is in the Woodwardian (Geological) Museum, Cambridge. It 
agrees closely with Williamson’s ‘‘carpellary disk’’ of Williamsonia gigas, 
Carr., except in its smaller size. 

° Carruthers (3). This paper was also printed in the Geol. Mag. vol. iv. 1867. 


a2 Oo BP w 


148 BENNETTITES. 


in 1867, refers to the opinions of Yates and Williamson, and 
adds: ‘‘I have examined numerous specimens of this fossil 
(Williamsonia gigas) in the British Museum, but have been 
unable to determine anything satisfactorily in regard to the 
precise structure of this anomalous fruit. It presents so many 
peculiarities unknown in the fruit of any modern cycad, that 
for the present at least, and notwithstanding its Zamia-like leaves, 
I must consider it a doubtful cycad.” 

In the volume of the Linnean Society’s Transactions for 1870 
we have the first exhaustive treatment of this Oolitic fossil. 
From his intimate knowledge, both of the fossils and their 
manner of occurrence in the rocks of the Yorkshire coast, 
Williamson was peculiarly fitted to attack this difficult problem. 
Williamson describes and figures what he regards as the stem of 
Zamites gigas, L. and H.; the surface is made up of broad 
lozenge-shaped areas, and is compared with the trunk of a recent 
Cycas. One example is referred to as ‘obviously the apex 
of a stem with portions of seven or eight diverging fronds.” 
The fronds are next described, also certain structures spoken 
of as the ‘‘squamous peduncles”? of the fructification. The 
ereater part of the paper is, however, devoted to a detailed 
examination of the ‘‘organs of fructification.”” Surrounding the 
ovoid inflorescence we have a number of linear bracts constituting 
an involucrum ; usually these involucral leaves have been broken 
off towards the base, and immediately below the broken ends 
there is exposed an annular area of ‘‘radiating cells.” In 
pl. li. fig. 7, Williamson represents a specimen in which the 
bracts have been completely preserved and are continued to the 
base of the fructification. The central part of the whole structure 
is occupied by a pyriform cavity which contracts apically, and 
then expands into a funnel-shaped appendage. In one of the 
figured specimens the form of this central cavity is clearly seen, 
and from its being filled with carbonaceous matter, it is assumed 
that the axis was originally a solid structure. A very similar 
appearance is presented by one of Rufford’s recently discovered 
specimens.” The ring of radiating cells seen at the base of most 
of the examples is considered to be the lowest margin of a layer 


1 Williamson (8). 
lite ek, yeh we 


BENNETTITES. 149 


of cells which ‘formed a cortical layer, arranged vertically upon 
and extending over the entire surface of the pyriform axis.” 
In another type of specimen figured by Williamson, we have 
a disk-shaped structure with a central depression, and split up 
peripherally into several short and narrow segments; this is 
named the ‘‘carpellary disk,” and towards the apex of each of 
these rays there are said to be two small pits on the lower 
surface. These pits are regarded as the places of attachment 
of ovules; but it is now generally agreed that there is no good 
evidence for the existence of well-defined depressions which 
could be described as marking the position of seeds. The general 
conclusion arrived at, and expressed by a restoration of the whole 
plant, is that the two forms of fructification are probably the 
male and female organs of Zamites gigas; the commoner ovoid 
fossil being the male flower, with a pyriform axis originally 
invested by a deciduous antheriferous tissue, and the female 
flower being represented by the much less abundant ‘carpellary 
disk,”” with the ovules inserted in pits towards the tips of the 
star-like rays. 

As a convenient provisional name for these anomalous structures 
Carruthers instituted the genus Williamsonia, and placed it in 
a new tribe—Williamsonee. The genus is thus defined :! ‘Stem 
cylindrical, elongated, marked with the equal-sized, tumid, 
rhomboidal scars of the fallen leaves. eaves ovate-lanceolate 
or linear acuminate, segments numerous, attached to the rachis 
by the central portion, with small free margins; veins numerous, 
parallel, at the base slightly diverging into the free margins. 
Flowers terminal, stamens surrounding a fleshy axis, ovules borne 
on the upper surface of an orbicular laciniate spadix.”’ 

Carruthers thus expresses himself with reference to Williamson’s 
work: ‘‘ He has introduced a clearer apprehension of the different 
forms of the supposed organs of reproduction, by the suggestion | 
that the two kinds represent the different sexes, and by the 
discovery of a seed-bearing spadix.”* In the second volume 
of his Plantes Jurassiques, Saporta considers the problem of 
Williamsonia,? and denies the existence of any satisfactory grounds 


1 Carruthers (1), p. 691. 
2 Thid. p. 692. 
3 Saporta, Pal. Frane. vol. ii. p. 53. 


150 BENNETTITES. 


for the supposed connection between the fronds of Zamites gigas, 
the cycadean stem, and the floral structures ; the last-mentioned he 
prefers to regard as some primitive form of a monocotyledonous 
inflorescence, probably a pandanaceous type, analogous to Yucevtes, 
Podocarya, Eolirion, ete. 

In the Palgontologica Indica there have been described by 
Oldham and Morris,' and afterwards by Feistmantel,? various 
specimens of Wiiliamsonia. The latter author figures certain 
eycad-like stems found in association with the Wailliamsonia 
fossils as belonging to the plant which bore the Williamsonia 
inflorescence. Further additions to our knowledge of the 
distribution of this fructification were made by Nathorst in 1880,? 
and he put forward the opinion that Wélliamsonia should be 
placed with the Balanophoree. This idea he afterwards abandoned, 
and in a later paper* upholds the view that Williamsonia was 
the inflorescence of a plant bearing cycadean fronds. He gives 
a restoration of Anomozamites minor (Brong.) bearing in the 
forks of a branched stem large flowers of the type Williamsonia 
angustifolia, Nath. The restored species presents an appearance 
certainly more suggestive of some extinct form of plant than of 
anything at present in existence. Nathorst associates his other 
species, W. Leckenbyi, with Anomozamites Lindleyanus, Schimp., 
and connects W. gigas, Carr., with Zamites gigas, L. and H., 
adding that possibly Braun’s Weltrichia may be regarded as the 
inflorescence of Otozamites. In another place® the same writer 
refers to the likelihood of a connection or identity of Bennettites 
and Williamsonia. 

In Zigno’s second volume of the Flora fossilis formationis 
Oolithice,’ certain specimens from Italian rocks are described 
under the new generic name of Blastolepis; the figures of the 
two species of this genus, B. ofozamitis and B. falcata, Zig., are 
strikingly suggestive of Carruther’s genus. There can be little 
doubt that these specimens should be assigned to the genus 
Williamsonia. 


1 Oldham and Morris (A), p. 32, pl. xxxii. fig. 12. 

* Feistmantel (2). 3 Nathorst (3). 4 Ibid. (4). 

° Since this was written, I have had an opportunity of examining Nathorst’s 
specimens, and can bear testimony to the accuracy of his description. 

6 Tbid. (1), p. 97. 

7 Zigno (1), p. 178, pl. xlii. figs. 9-11. 


BENNETTITES. 151 


In a recent volume of the Plantes Jurassiques, Saporta’ gives 
a full account of the Williamsonia question, and discusses at 
length such views as have been advanced as to the nature of the 
genus. Several of the Yates’ specimens, now in the Paris 
Museum (Jardin des Plantes), and many others are figured in 
Saporta’s monograph. Some of these figures were originally 
drawn for Brongniart,* whose intention it was to publish a 
memoir on the subject. In addition to this very important 
contribution by Saporta, reference should be made to a paper 
by him in conjunction with Marion in the Comptes Rendus for 
1881,* and also to the account by the same authors in DL’ Evolution 
du regne végétal* The genus Williamsonia, with Yuccites, 
Goniolina, Weltrichia, and others, is included in the special class 
of ‘‘ Types proangiospermiques.”’ ‘‘Ces types sont appelés par 
nous des Proangiospermes, parce, qu’ayant en réalité précédé dans 
Vordre des temps les Angiospermes véritables et ne pouvant étre 
classés méthodiquement parmi ces derniéres, ils se distinguent 
pourtant trés nettement de tous les végétaux passés en revue 
jusqwici, et quwils s’écartent a la fois et des Cryptogames et 
des Gymnospermes, n’ayant d’ailleurs de points de contact 
appréciables ni avec les Cycadées, ni avec les Salisburiées, encore 
moins avec les Coniféres.”* As an introduction to the examination 
of the genus, Saporta writes:® ‘‘Avec les Williamsonia nous 
abordons un des problémes les plus difficiles, un des sujets les 
plus controversés, mais aussi les plus curieux, peut-étre méme 
le plus remarquable de tous ceux que nous offre l’ensemble des 
plantes jurassiques.”” He agrees with Brongniart that Williamsonia 
is probably generically identical with Buckland’s Podocarya. 
Some of Williamson’s conclusions he does not accept; the 
‘‘carpellary disk” of that author, Saporta regards as a terminal 
expansion of the male spadix. One important point to note 
in reference to Saporta’s conclusions is his interpretation of two 
forms of specimens; that figured by Williamson in his pl. lii. 


1 Pal. France. vol. iv. 1891. 

2 Ibid. p. 90 (footnote). 

3 Saporta and Marion (1). 

4 Ibid. (2), Les Phanérogames, vol. i. p. 235. 
§ Saporta, Pal. Frang. vol. iv. p. 63. 

§ Ibid. p. 89. 


152 BENNETTITES. 


figs. 8-61 is described as the male inflorescence, and another form, 
of which several figures are given in the Paléontologie Francaise, 
is regarded as the female inflorescence. Like Williamson, Saporta 
sees in the fibrous layer at the base of the common form of 
Williamsonia an antheriferous tissue; the pyriform central axis 
he describes as expanding distally into a large structure like that 
figured by the English writer as the carpellary disk; this 
terminal expansion appears to have been readily separated by 
a natural surface of disarticulation from the rest of the axis. 
In the whole inflorescence, according to Saporta and Marion, 
we have a male involucre surrounding a conical axis with its 
base enclosed in a circular zone marked by radiating strie; the 
external edge of this zone being occupied by a number of small 
compartments of an irregular hexagonal form, which seem to 
correspond to pollen-sacs. The basal zone represents the sterile 
and persistent portion of an androphore, which, when complete, 
covered the whole of the conical receptacle with a layer of 
staminal appendages. In the female inflorescence the bracts are 
somewhat shorter; the centre was occupied by a more or less 
globular axis, having its surface marked out into a number 
of compartments arranged in the form of facettes grouped in 
rosettes; the general appearance of the whole structure being 
very similar to that presented by Buckland’s Podocarya. The 
ovules were situated in subcortical cavities, which communicated 
with the surface by small openings; the latter appearing as the 
central points of groups of comparatively small meshes of the 
superficial reticulum. In 1869 Moriére described a_ petrified 
fruit from the Oxfordian beds of Vaches-Noires (Calvador),? and 
this specimen is regarded by Saporta and Marion as throwing 
considerable light on the nature of Williamsonia. The recent 
examination of this specimen by Lignier* has already been 
alluded to; his work has afforded us very important data with 
regard to the connection between Bennettites and Wailliamsonia. 

Saporta draws attention to a close resemblance between the 
terminal infundibuliform expansion of Williamsonia and the fossil 


1 Williamson (8). 
2 Moriére, pl. ii. fig. 4. 
3 Lignier (1). 


BENNETTITES. 153 


dealt with many years ago by Braun.’ Braun’s genus Weltrichia 
was instituted for a specimen discovered in the neighbourhood 
of Baireuth, and referred to as a new genus of the Rhizanthee. 
Schenk ? expresses himself as very sceptical as to the correctness 
of Braun’s description of this fossil, and does not consider it of 
any scientific value. 

In discussing the systematic position of Williamsonia and 
other genera, Schenk very reasonably calls attention to the 
absence of any satisfactory evidence in favour of a pandanaceous 
alliance; he suggests that possibly Bennettites and Walliamsonia 
should be classed together, but refers to the absence of histological 
characters as a serious obstacle to any decided conclusion. In 
Zittel’s Handbuch, Schenk® refers to Williamsonia as the female 
inflorescence of Bennettites, and refers to Nathorst’s and Solms’ 
work in support of this opinion. Solms-Laubach‘ agrees with 
Nathorst and Saporta & Marion as to the absence of trustworthy 
evidence of the connection between Zamia gigas and Williamsonia. 
After referring to Moriére’s important specimen, he concludes: ° 
‘‘T have no doubt, therefore, that this specimen belongs to 
Bennettites, but in saying this I have no intention of prejudging 
the question of its relation to Williamsonia ; for it is still possible 
that further discoveries may show the fructifications of Lennettites 
and Williamsonia both belong to similar stems resembling the 
stems of Cycadacea, and confirm the opinion of Williamson and 
Carruthers. But until the truth of these conjectures is ascertained, 
we must be content to leave the relationship of Wialliamsonia 
undetermined.” In another place, Solms* draws attention to the 
probability that we have long been familiar with the male 
inflorescence of Bennettites in the fossil known as Williamsonia. 

In his account of Bennettites etrusca, Cap. and Solms,’ Solms 
figures and describes certain boat-shaped sacs which he regards 
as pollen grains. He goes on to say that, if his interpretation 
of these structures be correct, the inflorescence of ennettites 


1 Braun (1). 

2 Schenk (A. 8), p. 190. 

3 Zittel (A.), Handbuch, p. 805. 

4 Fossil Botany, p. 370. 

5 Ibid. p. 372. 

6 Note on Bennettites in Saporta’s Pal. Franc. vol. iv. p. 303. 
7 Capellini and Solms, p. 202, pl. v. figs. 7 and 8. 


154 BENNETTITES. — 


apparently bore anthers and pollen grains in its early stage of 
development, and afterwards assumed the female condition. The 
figures of isolated ‘‘ pollen grains” are not thoroughly convincing ; 
but if Solms’ opinion be correct, we have a most important 
addition to our knowledge of this interesting genus. 

My examination of the Wealden examples of Williamsonia 
leads me to support the view that this problematical fossil is 
generically identical with Bennettites, and so far as our evidence 
goes, we are, I believe, justified in regarding the former genus 
as a form of inflorescence of the same type as that which has 
been found in organic union with bennettitean stems. This 
opinion is chiefly based on the Wealden forms, but if the suggested 
relationship or identity of these with Bennettites be admitted, we 
have a very strong case for including the Jurassic species in the 
same category. A critical discussion of the Oolite specimens must 
be deferred until the French material has been studied; such a 
question will be best dealt with in a later volume devoted to 
the Jurassic flora. 

As regards the question of male and female inflorescences, I 
am unable to recognize any sexual difference in the various 
examples from the Wealden beds, and there does not seem to 
be any good reason for regarding the so-called male Williamsonias 
among the Jurassic specimens, as in any way proved to be of 
that nature. In comparing Williamsonia with Bennettites we have 
to rely entirely on the female inflorescence of the latter plant, 
and it would seem that so far as our present evidence goes, we 
have more reason for speaking of Williamsonia as the female 
inflorescence. As to the nature of the male inflorescence we 
are still without any very satisfactory evidence. 

The following records have been made of Williamsonia, showing 
a fairly wide distribution; but probably some of these species 
cannot well be retained as trustworthy examples of the genus. 


ENGLAND. Williamsonia gigas, Carr. Inferior Oolite. 
W. Leckenbyi, Nath. “ 33 
W. Bucklandi (Ung.). > * 
FRANCE. W. Pougneti, Sap. Lower Lias. 
W. Moriérei, Sap. Oxfordian. 
W. pictaviensis, Sap. a 
W. Zeilleri, Sap. Kimmeridgian. 
W. Gagnierei, Sap. Portlandian and Purbeckian. 


Portueat. W. minima, Sap. Lower Cretaceous. 


BENNETTITES. 155 
Swepen (Scanta). W. angustifolia, Nath. Rheetic. 
Bornuoim. W. Forchhammeri, Nath. Jurassic. 
GREENLAND. JW. cretacea, Heer. Cenomanian. 
Irany. W. (Blastolepis) otozamitis, Zig. Inferior Oolite. 
W. (Blastolepis) falcata, Zig. 33 35 
W. (Blastolepis) acuminata, Zig. 4 He 
CANADA. W. recentior, Daws. Middle Cretaceous. 
Williamsonia ? Neocomian. 
AMERICA. W. virginiensis, Font. Potomac. 
W. elocata, Lesq. Dakota. 
W.? Riesrvi, Hollick Cretaceous. 
Inpia.! W. Blanfordi, Feist. Kach (Umia beds; Up. Oolite). 


W. cf. gigas, Carr. 


Rajmahal (Lias). 


W. microps, Feist. 4 55 


The second English species, W. Leckenby7, is founded on a small 
rayed disk like the large “‘carpellary disk” of Williamson, and 
is regarded by Saporta as a lobed terminal expansion and not an 
involucre as suggested by Nathorst. Saporta considers it possible 
that the forms referred to this species may be simply a ‘‘ morpho- 
logical variation”? of W. gigas. 

Buckland’s Podocarya is transferred by Saporta to Williamsonia,* 
and it is highly probable that this plant is a bennettitean in- 
florescence. 

The French species W. Pougneti, Sap.,* is founded on an 
imperfect specimen which does not admit of any exact determina- 
tion, and hardly justifies the institution of a new species. W. 
Zeilleri, Sap.,° is also founded on a very poor specimen, and cannot 
be diagnosed with any exactness. W. minima, Sap.,° recently 
described from Portugal, is represented by an imperfect cluster 
of small bracts, and cannot be accepted as a satisfactory record 
of the genus. 

Saporta has pointed out that Heer’s species, W. cretacea, from 
Greenland, shows many points .of divergence from the typical 


1 A fossil figured by Sharpe as Asterophyllites ? from South Africa resembles 
an involucre of Williamsonia ; but Hooker’s description of the specimen is more 
suggestive of such a genus as Schizonewra (Sharpe, Trans. Geol. Soc. vol. vil. 
[2], p. 227). Iam indebted to Prof. Rupert Jones for calling my attention to 
Sharpe’s figure. 

* Saporta, Pal. Franc. vol. iv. p. 167. 3 Ibid. p. 127. 

4 Ibid. p. 124, pl. ccxxxvii. ; see also Saporta and Marion (2), vol. i. p. 234. 

5 Ibid. p. 181, pl. cexxxiv. fig. 3. 

6 Saporta (1), p. 108, pl. xix. fig. 9, 


156 BENNETTITES. 


form;* it may possibly be an inflorescence of some cycad-like 
stem, but is far from satisfactory.? One of the forms figured by 
Zigno*® (Blastolepis acuminata) is named by Saportat W. Jtalica. 
Dawson’s W. recentior is an exceedingly poor specimen and 
hardly worthy of any name. 

It is difficult in the case of some of the above forsila to 
precisely define their geological horizon. The Bornholm species, 
W. Forchhammeri, Nath.,> is from beds which have recently been 
shown by Bartholin to contain 46 species of plants, of which 
25 are regarded as Rheetic, and about 15 as indicating a Lower 
Oolitic age. 

The fragment described by Dawson® as W. recentior is from 
the Canadian Middle Creek series, which is compared with the 
Patoot series of Greenland and the Dakota group of the Western 
United States. 

The unnamed Williamsonia is from the Kootanie formation, 
which is correlated with the Neocomian of Europe.? 

Fontaine’s Potomac series includes rocks differing somewhat 
widely in age, Jurassic and Cretaceous strata being incorporated 
in one formation. 

Williamsonia? Riesii is probably a true Williamsonia with 
numerous and unusually narrow bracts; the author of the species 
compares it with some composite flower. 

As regards the Indian beds, it has long been a difficult problem 
to determine their exact geological position; the Rajmahal beds 
are spoken of by Oldham,® in the recent edition of the Manual 
of Indian Geology, as Liassic, the Kach (Cutch) or Umia beds 
being correlated with the Upper Oolite, but the precise horizon 
can only be approximately stated. 


1 Saporta, Pal. Franc. vol. iv. p. 118. 

2 Heer, Flor. foss. Arct. vol. vi. p. 59, pl. xii. fig. 1; pl. xiii. fig. 9. 
[Since writing the above I have examined Heer’s type specimen of W. eretacea 
in the Geological Museum in Copenhagen, and find the impression is very 
indistinct and unsatisfactory. | 

3 Zigno, loc. cit. p. 173, pl. xlii. fig. 10. 

* Saporta, loc. cit. p. 180, pl. ecli. fig. 3; and pl. eclii. fig. 4. 

5 Nathorst (4). See also Bartholin, p. 112. 

6 Dawson (A. 5), p. 12, pl. iv. fig. 1. 

7 Dawson (2), p. 87. 

8 Medlicott and Blanford, p. 207. 


BENNETTITES. Vow 


Bennettites (Williamsonia) Carruthersi, sp. nov. 
REIS xX andshl. XI. Fig.)8. | 


Type. Specimens in the British Museum from the Fairlight 
clays, Fairlight, near Hastings. 

As a matter of convenience, a new specific name is adopted for 
several specimens in the Rufford Collection. Without a fuller 
knowledge of their anatomical structure it is impossible to give 
an exact specific definition, but the general characters may be 
briefly expressed as follows :— 

Inflorescence ovoid, surrounded by numerous linear bracts, 
enclosing a central axis (from which seed-bearing peduncles and 
interseminal structures were given off); between the involucral 
bracts and the periphery of the spadix there was a regular 
reticulum of projecting ingrowths marking out the surface of 
the inflorescence into small areolations. 

In dealing with detached inflorescences or portions of such 
structures, it is impossible to clearly discriminate between different 
specific forms, as distinct from portions of the same species or 
the same inflorescence in different stages of development. The 
most important features in the following specimens are those 
which serve to connect them, on the one hand with the typical 
Bennettites, and on the other with the Jurassic Williamsonia. 
Some of the examples of these Wealden forms differ from the 
majority in haying short and broad bracts at the base of the 
inflorescence; these we may speak of as B. (Williamsonia) 
Carruthersi var. latifolius. I have ventured to associate the name 
of Mr. Carruthers with the present species; it is to his work 
that we are primarily indebted for our knowledge of Bennettvtes. 


V. S177. Ele Vien. 1, Wa, and 18: 

This specimen is one of the most important of those to be 
described. In general form and appearance it is very similar to 
- Bennettites Moriéret as figured by Lignier,! except that in the 
present specimen the linear bracts are distinctly shown, about 


1 Lignier (1), pl. v. figs. 55 and 56. 


158 BENNETTITES. 


twelve in number, and, as occurs so commonly in the Jurassic 
Williamsonias, they are broken off basally, leaving an annular 
area immediately surrounding the base. This annular area shows 
very distinctly numerous parallel longitudinal striations; these I 
regard as corresponding to the so-called ‘‘antheriferous tissue” of 
some writers in Williamsonia gigas, Carr. At the base we have 
a well-defined rim surrounding a central short and conical cavity 
(Fig. 16); a similar form of axis occupied by carbonaceous matter 
occurs in the specimen represented in Fig. 8. Compare also Saporta, 
pl. xxvi. fig. 3. Length of V. 3177 6cm., breadth 38-5cm. In 
Fig. 1a the truncated base of one of the bracts is represented as seen 
from below; this shows a number of regularly placed projections 
from the face of a bract, extending from the latter to the internal 
fibrous structures. In some of the figures of the inflorescence 
of Bennettites Gibsonianus, Carr., given by Solms-Laubach,! there 
are similar internal projections represented in the inner face of 
the ‘‘outer layer” of the fructification. In describing the 
structure of the inflorescence, Solms writes: ‘‘ Not unfrequently 
sharp and tolerably deep indentations penetrate from without into 
this homogeneous external layer; these indentations are covered 
with the epidermis, and probably answer to the cross-sections of 
a superficial areolation of the entire fructification ; they are par- 
ticularly well and clearly seen near the base of the spadix in 
fig. xil. of pl. xxv.” The same indentations are seen in Solms’ 
pl. xxv. fig. 8, fig. 10, and fig. 11. These ingrowths are, I 
believe, the structures seen in our Fig. la, Pl. X. In the longi- 
tudinal section of Cycadeoidea etrusca, Cap. and Solms,’ figured 
by Capellini and Solms, pl. iv. fig. 1, we have the structures 
clearly represented, and again, on a larger scale, by Carruthers * 
in B. Gibsonianus (pl. lx. fig. 3). In his description of the latter 
species, Solms speaks of the interstitial organs as becoming much 
more numerous towards the periphery of the spadix (pl. xxv. 
figs. 8 and 11); Lignier also refers to this character, and in a 
figure of the peripheral region of the fructification (p. 31, fig. 2), 
shows the superficial bracts on the outside, and internal to these 
much smaller interseminal bracts and atrophied seed-bearing 


1 Solms-Laubach (2), pl. xxv. figs. 8 and 10-12. 
2 Capellini. 
3 Carruthers (1). 


BENNETTITES. 159 


peduncles. The fine longitudinal lines seen on the surface of 
the specimen figured by Lignier on pl. v. figs. 55 and 56, and 
described by him as the peduncles, bear a very close resemblance 
to the fine lines traversing the basal annular area in our figure 
(Pl. X. Fig. 1), and to the so-called antheriferous tissue of 
Williamsonia gigas. It is, I believe, the interseminal structures 
or possibly atrophied peduncles that are seen in such specimens of 
Bennettites Carrutherst as are represented in Pl. X. Fig. 1, Pl. X. 
Fig. 2, Pl. X. Fig. 3, and Pl. XI. Fig. 4. The central conical 
cavity was originally occupied by a fleshy axis, on which were 
borne the seed-bearing peduncles. Ecclesbourne. Rufford Coll. 


V. 3201. Pl. X. Fig. 4. 

In this specimen we have the outer part of the basal portion of 
a larger form, or older inflorescence. In the centre there is a 
somewhat oval area with an uneven surface 2-3cm. in diameter, 
surrounded by a series of slightly raised structures; external to 
this is a concave rim, with its surface marked by a well-defined 
reticulum of projecting ridges. The appearance of this saucer-like 
rim suggests that there were originally numerous narrow bracts in 
close contact with it; the outline of these being indicated by the 
shallow depressions and intervening ridges as shown in the figure. 
This specimen approximately corresponds to the basal portion of 
V. 3177 (Pl. X. Fig. 1), the external margin of the reticulately 
marked rim coinciding with the truncated bases of the involucral 
bracts of such examples as V. 3177 (Pl. X. Fig. 1) and V. 2129 
(Pl. X. Fig. 2). The network is probably formed by the same 
regular indentations which are seen in Fig. la, and described by 
Solms' as forming a regular areolation over the surface of the 
spadix. Between the reticulately marked rim and the central 
boss there would be the continuous external layer of the peripheral 

zone. Cf. Solms, pl. xxv. figs. 4 and 7-12. Ecclesbourne. 
Rufford Coll. 


V. 32025 Pie Fie. 2. 

Here again we have the characteristic Williamsonia gigas base, 
somewhat larger than in V. 3177 (Pl. X. Fig 1). The linear bracts 
are unevenly broken, exposing the numerous fine, interseminal, 


1 Loc. cit. p. 437. 


160 BENNETTITES. 


fibre-like structures. Probably in this specimen the whole in- 
florescence is partially expanded, and is not so conical in form 
as the more completely closed example shown in Pl. X. Fig. 1. 
Cf. W. gigas as figured by Williamson (pl. lii. fig. 3), W. Gagnieret 
as figured by Saporta’ (pl. xxvi. figs. 1-3), ete. Ecclesbourne. 

Rufford Coll. 


V. 21295. PIX. Fig. 3. 

The remains of a fructification similar to but rather smaller 
than V. 3177 (Pl. X. Fig. 1). Occupying the centre is a well- 
defined hemispherical boss, 8 mm. in diameter, differing from that 
in Pl. X. Fig. 16 in its more spherical form; its surface is covered 
with small punctations. At the base of this boss there appears to 
be a small projecting rim, and external to this an almost vertical 
involucre of narrow bracts. Resting on the inner face of these 
bracts is a fibrous material, consisting probably of the same 
slender structures as those seen in V. 8202 (Pl. X. Fig. 2), and 
like them being the small interseminal scales, or peduncles, 
which make up the peripheral portion of the spadix. Compare 
B. Gibsonianus, Carr.: in that species the central boss is more 
cushion-like in form. At a distance of about two-thirds from 
the base the bracts are bent outwards, as shown in the figure. 
Ecclesbourne. Rufford Coll. 


We. 2129¢., Pl. X. Fig. 5. 

A star-like cluster of broader bracts, closely resembling such 
a form as Williamsonia (Blastolepis) acuminata, Zig. (= W. Ltalica, 
Sap.).2. The bracts are arranged in a close spiral, and do not 
form a true whorl. In some there are a number of fine ‘hairs 
obliquely attached to the margin; these are shown on the 
lower margin of the middle left-hand bract in the figure. The 
hairy margin corresponds with that in Cycadolepis villosa, Sap.,° 
and our V. 2802,‘ ete. ‘This specimen probably represents a series 
of bracts which surrounded the base of an inflorescence. LEccles- 
bourne. Rufford Coll. 


1 Pal. Franc. vol. iv. 

2 Saporta, Pal. Frang. vol. iv. pl. ecli. fig. 3. 
3 Ibid. vol. ii. pl. exiy. fig. 4. 

05 Bf 


BENNETTITES. 161 


V. 2793. Pl. XI. Fig. 1. 

The bracts in this specimen are of large size, and less complete. 
In the centre we have the remains of the base of the axis to 
which seed-bearing peduncles were originally attached: ef. V.3201 
(Pl. X. Fig. 4). On one side of the central boss a portion of the 
fibrous annular ring is preserved, showing the impressions of some 
of the peripheral structures of the spadix; external to this we 


have traces of a reticulate structure resembling that of V. 3201 
(Pl. X. Fig. 4). Ecclesbourne. Rufford Coll. 


V. 2129¢. Pl. XI. Fig. 2. 

Portions of large and spreading bracts surrounding a slightly 
irregular conical cavity; on the inner face of this there are 
numerous fine reticulations. Probably of the same type as 
V. 3177 (Pl. X. Fig. 1), but much less complete. Ecclesbourne. 

Rufford Coll. 


Fic. 8.—Bennettites (Williamsonia) Carruthersi, sp. nov. (nat. size). 
Specimen in the possession of Mr. Rufford. 


This specimen (Fig. 8) shows the clearly outlined central 
structure, of conical form, similar to the smaller conical cavity 


M 


162 BENNETTITES. 


in V. 3177 (Pl. X. Fig. 1). The impression immediately below 
the inflorescence is probably not in organic connection with it. 
Preservation imperfect. Fairlight. 


V. 2913. Involucral bracts, surrounding a central boss. 
Towards the base of one of the bracts, and on the surface of the 
rock, there are some traces of the reticulate structure usually 
met with in this position. Ecclesbourne. Rufford Coll. 


V. 3172. Fairly broad involucral leaves surrounding a small 
central cavity. Cf. V. 3177 (Pl. X. Fig. 1). Ecclesbourne. 
Rufford Coll. 


V. 29132. Small specimen showing a portion of the annular 
zone,' with narrow irregular projecting ridges towards its inner 
margin, passing towards the outer edge into distinct reticulations : 
of. V. 3201 (Pl. X. Fig. 4), etc. In the centre is a slightly 
raised boss, separated by a circular groove from the annular 
zone; in the circular groove was probably situated the ‘“‘con- 
tinuous external layer of the peripheral zone,” as figured and 
described by Solms.? Ecclesbourne. Rufford Coll. 


V. 2793a and V. 2793. Very similar to V. 2129c (Pl. X. 
Fig. 5). 


V. 2801. This specimen shows a central portion like that 
in V. 3201 (Pl. X. Fig. 4); surrounding this there are irregular 
radiating lines gradually passing towards the periphery into 
a regular reticulation. Ecclesbourne. Rufford Coll. 


V. 29134. A circle of narrow bracts surrounding a central 


area. ,, Cf. Pl. XI. Fig. 1 (VW. 2798). 


1 There is a specimen very similar to this in the Leckenby Coll. (Cambridge). 
2 Solms (2), pl. xxv. 


BENNETTITES. 163 


V. 21299. Two specimens with an involucre of broader 
bracts surrounding a central boss. It is difficult to decide how 
far the breadth of the bracts may constitute a specific difference. 
These examples are somewhat intermediate between the specimens 
referred to the variety Jatifolius, and the narrow leaved forms. 


Cf. V. 27985 and V. 2129/, ete. 


V. 2254. Small portion of the annular zone. LEcclesbourne. 
Rufford Colt. 


V. 2129. Small specimens showing a central cavity bordered 
by an annular zone of fibrous structures. 


Bennettites (Williamsonia) Carruthersi, sp. nov., 
var. latifolius mihi. 


V. 2129¢. Pl. XI. Fig. 4. 


Compare with a figure of Williamsonia gigas given by Oldham 
and Morris. In the centre is a depressed boss, as in V. 2793 
(Pl. XI. Fig. 1), V. 3201 (Pl. X. Fig. 4), surrounded by a narrow 
ring, and external to this a rim of fibrous structures, about 1:3 cm. 
broad. At a lower level than the fine radiating fibrous structures 
(peripheral interseminal scales or abortive peduncles), we find here 
and there an impression of a reticulately marked surface; this 
reticulum is probably identical with that in V. 3201 (Pl. X. 
Fig. 4), and in V. 8177 (Pl. X. Fig. 1). External to the annular 
ring are the blunt and rounded tips of a few broad bracts. The 
specimen consists of the detached basal portion of an inflorescence 
seen from the inside. Ecclesbourne. Rufford Coll. 


V. 2129/. Pl. XI. Fig. 3. 


Very similar to V. 2129 (Pl. XI. Fig. 4); an involucre of short 
and broad bracts like those of the preceding specimen, but in this 
case seen from the outside. In places where portions of the bracts 
have been removed, a fine reticulate structure is seen on the rock- 
surface. In the centre of many of the polygonal meshes a slight 
magnification reveals the presence of a small black dot of coaly 


164 BENNETTITES. 


substance. A mesh with such a central dot presents some resem- 
blance to the section of a peduncle as figured by Lignier,' the 
dot representing the central vascular bundle. Ecclesbourne. 


Rufford Coll. 


V. 2129a. Very similar to V. 2129/. 


P? Bennettites. 


V. 2304, V. 3174, V. 3176. Indistinct remains of carbonized 
hairy bracts; may possibly belong to Bennettites. Also V. 2830 
and V. 2816. Ecclesbourne. Rufford Coll. 


V. 3163 and V. 3164. A short axis, about 4cm. in breadth, 
terminating in partially expanded bracts. The bracts and hairy 
ramenta closely resemble those of V. 3177 (p. 144), and suggest 
the same plant. May possibly be an axis which bore a 
Williamsonia inflorescence. Kcclesbourne. Rufford Coll. 


V. 2305, V. 2306. May be portions of similar stems, but cannot 
be accurately determined. Ecclesbourne. Rufford Coli. 


V. 2349. A mass of hairy or woolly structures evidently 
arranged round some axis; pressed together as so many thin 
lamine. Very similar to the hairy scale leaves, ete., in the stem 
previously described (V. 3177, p. 144). Ecclesbourne. 

Rufford Coll. 


V. 2930. Woolly scales or masses of ramental structures. 
Keclesbourne. Rufford Coll. 


1 Lignier (1), pl. v. 


YATESIA. 165 


Genus YATESIA, Carruthers. 
[Trans. Linn. Soc. vol. xxvi. 1870, p. 687.] 


This generic name was proposed by Carruthers for a form of 
cycadean stem originally described as Cycadeotdea Yatesi’, Carr.4 
The genus is thus defined :— 

‘Trunk cylindrical, of uniform thickness, and covered with 
the short persistent bases of the petioles; scars of the aborted 
leaves scattered among those of the true leaves. Androecium 
unknown; gyncecium forming a cone, each carpophyll of which 
bears two reflexed ovules.” 

The characters of the floral structures are based on certain 
cones of somewhat doubtful affinity, which have not been found 
in actual connection with Yatesia stems, and cannot, therefore, 
be regarded as of much value. In some of the specimens referred 
to this genus, the general arrangement of the tissues, and to 
some extent the histological structure, have been preserved, but 
the latter are but very imperfectly known. If we examine the 
figures of stems included in Yates¢a, we shall find it a difficult 
task to distinguish some of the examples figured by Carruthers 
from stems referred to the genus Bucklandia. The form of the 
leaf-scars in Bucklandia Milleriana, Carr.,? agrees closely with 
that in the Yatesta stems; and in Y. Joassiana, Carr.,? there 
appear to be distinct indications of the transverse constrictions 
in the stem, as in many bucklandian stems; the form of the 
leaf bases is also very similar in the two genera. Questions as to 
differences in age, stages of growth, and the manner of preservation 
of the stems, render the discrimination and exact limitation of . 
generic types exceedingly difficult, or even impossible; my im- 
pression is, that at all events in some of Carruthers’ species there 
are no satisfactory grounds for the application of two generic 
names. It does not seem possible to draw any distinct line of 


1 Carruthers (6). 
? Carruthers (1), pl. lv. fig. 1. 
3 Jbid. pl. lv. fig. 8. 


166 YATESIA. 


separation between some of the smaller stems of Bucklandia and 
the yatesian form of trunk. Schimper includes Yatesia Joassiana, 
Carr.,! in the genus Clathraria, and expresses surprise that it 
has been referred by Carruthers to another genus. 

The specimens of Yatesia DMorrisii, Carr., described by the 
author of the species, were obtained from the Lower Greensand 
beds of Potton and Leighton Buzzard. One of the examples in 
the National Collection is described as being from the ‘‘ Wealden” 
beds of Leighton Buzzard. The exact age of fossils from these 
beds must be a matter of some uncertainty, owing to the fact 
of many of them being clearly derived forms; it may be noted, 
however, that some of the plant fossils from Potton appear to be 
specifically identical with Wealden types. 

Among the genera instituted by Saporta, that of Cylindropodium 
includes some forms of stems which bear a striking resemblance 
to Carruthers’ species of Yatesva. 


Yatesia Morrisii, Carr. 


1867. Cycadeoidea Morrisii, Carruthers, Geol. Mag. vol. iv. p. 199. 

1870. Yatesia Morrisit, Carruthers, Trans. Linn. Soc. vol. xxvi. p. 688, 
pl. lv. figs. 3-6. 

1874. Yatesia Morrisii, Schimper, Trait. pal. vég. vol. iii. p. 55d. 


Type. Stem with internal structure imperfectly preserved. 
Royal Agricultural College, Cirencester. 

The following is Carruthers’ definition of this species :— 

‘Stem cylindrical, covered with the bases of the petioles, which 
are rhomboidal in form, and terminate in a tumid boss, the apex 
of which is directed upwards. The cellular axis is very large. 
The pith has disappeared, except in one specimen, where sufficient 
of it remains to show that it was permeated with vascular 
bundles. The woody cylinder surrounding the pith, in the 
specimen figured, consists of two rings (figs. 5 and 6); it is 
everywhere pierced by medullary rays, which are often so large 
as to break the continuity of the wood (fig. 6). The sides of the 
wood-cells parallel to the medullary rays are covered with disks 


1 Schimper, Trait. pal. vég. vol. iii. p. 544, 


YATESIA. 167 


in two or three rows (pl. lx. fig. 13). The inner surface of the 
woody cylinder exhibits numerous narrow grooves, being the 
meshes for the passage outwards of the vascular bundles to 
the leaves. These meshes are larger and more regular on the 
outer surface of the wood. A very thin layer of cortical tissue 
separates the wood from the base of the petioles. The bases 
of the petioles spring from this layer, at mght angles to the stem. 
Externally they present a rhomboid form, the horizontal diameter 
of which is but little more than the perpendicular.” 

Although the Potton and Leighton Buzzard sands, in which 
the examples of this species have been found, are of Lower 
Greensand age, the plant may be reasonably included in the 
present list as possibly a member of the Wealden flora. 


47029. Portion of a stem preserved in oxide of iron. The 
inner face of the wood is shown, with the elongated medullary 
ray cavities; an impression of this face would present a similar 
appearance to the medullary cast of Bucklandia. Pith large, 
surrounded by a zone of wood 1:4cm. in width, and consisting 
of two concentric rings. The transverse section does not show 
any well-marked medullary rays traversing the wood. The 
petiole bases fairly distinct; some of them terminate apically 
in a manner suggestive of a clean surface from which the frond 
has been detached. Leighton Buzzard. Morris Coil. 


V. 221. Two smaller specimens. A distinct variation in the 
size of the leaf bases: ¢f. V. 2610. Leighton Buzzard. 
Morris Coll. 


Cf. Yatesia Morrisii, Carr. 
[Fig. 9.] 


The following specimens in the Beckles Collection, from the 
Wealden rocks of Sussex, while agreeing fairly closely with 
Yatesia Morrisii, possess certain points of resemblance to the 
genus Bucklandia. The external characters correspond to some 
extent with those in Saporta’s genus Cylindropodium,' but the 


1 Saporta, Pal. Frane. vol. ii. p. 265, pl. xlix. 


168 YATESIA. 


specimens from the French beds are in a better state of 
preservation, and admit of a more complete diagnosis, than the 
English examples, 


Fig. 9.—Cf. Yatesia Morrisii (nat. size). (V. 26100.) 


V. 26103. Fig. 9. 


Leaf bases very prominent and considerably waterworn 
(Fig. 9, a and 6). At the upper end of the specimen, as seen 
in Fig. 9a, the outer surface of the wood is exposed; in a view 
of the transverse section (Fig. 9c) there is seen to be a single 
ring of wood, with the bundles separated by broad primary 
medullary rays, the spaces in the structureless cast representing 
the xylem bundles. The small development of wood points to 
a young stem, in which no second cambium had been formed. 
Of, Carruthers’ figures of Y. Morrisii (pl. lv. figs. 8, 5, and 6)." 
Sussex. Beckles Coll. 


1 Carruthers (1). 


TRUNCI. 169 


V. 2607. Two specimens. The leaf bases present for the most 
part a different appearance to those in the other specimens; they 
are closer together and not separated by the deep grooves shown 
in Fig. 9 (V. 26100). Some of the bases, however, project exactly 
as in the preceding example, and there can be little doubt as to 
their specific identity. There is a distinct disparity in the size 
of the petiole bases. Sussex. Beckles Coll. 


V. 2610. Waterworn stem with leaf bases as in V. 26100. 
There is only one ring of wood, but this is no doubt merely 
a matter of age, and shows that the stem was younger than that 


of 47029 ( Yatesia Morrisic). 
V. 2610a. Smaller example. 


V. 2610c. Two impressions of the worn surface of a stem, or 
possibly a large cone. Cf. V. 26100 (Fig. 9). Sussex. 
Beckles Coll. 


V. 2612. An impression of the outer surface of a stem, or less 
probably of a cone. Cf. V. 2607, also V. 2749a (Bucklandia 
anomala, Stokes and Webb).’ Sussex. Beckles Coll. 


Trunci (Cycadaceze). 
Cf. ‘ Dracena Benstedtii,” Konig. 
[Pl. XII. Figs. 4 and 5.] 


In a paper on Mesozoic Angiosperms, contributed to the 
Geological Magazine in 1886 by Starkie Gardner, we find the 
following statement: ‘‘The stems of Hndogenites erosa, so common 
in the Wealden and Neocomian, are now known to be cycadeous, 
and it is probable that the Dracena-like stems from Tilgate Forest 
and elsewhere, so often referred to by Mantell, are referable to 
the same group.”? Endogenites erosa is now recognized as a fern 
(Vol. I. p. 148); but the Dracena-like stems are in all probability, 
as Gardner suggests, cycadean. Unfortunately no reasons are 
given for this opinion. In the Second Report of the Committee 
on British Tertiary and Secondary Beds, Gardner writes:* ‘“‘ We 


0, U8): 
2 Gardner (A. 1), p. 201. 
3 Gardner (A. 2), p. 243. 


170 TRUNCI. 


are not able to speak with certainty regarding the supposed 
liliaceous or Dracena-like stems from the Wealden, so frequently 
mentioned by Mantell, since it is not easy now to identify the 
particular specimens referred to by him.’’ Mantell? refers to the 
fossil stems discovered by Bensted at Maidstone, as nearly related 
to Yucca or Dracena. On examining the large specimens of stems 
in the British Museum, from the Iguanodon quarry” at Maidstone, 
I was struck by their resemblance in external characters to the 
stem of such recent cycads as Zamia Loddigesit, Miq., Z. Skinners, 
Warsz., and Z. pumila, L. 

In 1868 Carruthers referred to the Maidstone fossils under the 
name Dracena Benstedtii of Konig,*® and expressed his opinion that 
they exhibit a closer resemblance to the stem of a Pandanus than 
to that of a Dracena; but he refers to certain specimens in the 
British Museum which appear to show the remains of internal 
woody tissue, and thinks it possible that a closer examination 
might not lend support to the comparison with either mono- 
cotyledonous genus. In one or two of the Maidstone Kentish 
Rag stems, there are portions of what closely resembles woody 
tissue showing well-marked rings of growth, but a section cut 
from this wood-like material proves it to be simply a deposit of 
carbonate of lime formed in such a way as to closely simulate the 
structure of wood. One of these so-called Dracena stems was 
figured and briefly described in the Geologist for 1862,‘ but the 
drawing does not give a very accurate idea of the specimen. 
In a footnote to a paper by Bensted, Mackie® points out 
the absence of any figure or description of Dracena Benstedti 
by Konig. Morris’ gives this name as Konig’s, but adds after 
the author’s name ‘ British Museum,” and gives no reference to 
any published account. In attempting to trace the geological 
history of Monocotyledons, we are confronted on every hand with 


1 Mantell (1), vol. i. p. 186. 
2 Bensted. 
3 Carruthers (7), p. 154 (footnote). 
4 Mackie, Geologist, vol. v, p. 401, pl. xxi. 
5 Bensted, Geologist, Joc. cit. p. 336. 
6 Kénig was sometime Keeper in the Mineralogical Department of the British 
Museum. 
7 Morris (A.), Brit. Foss. p. 8. 


TRUNCI. 7s 


exceedingly doubtful fossils which cannot be relied upon as 
satisfactory records; many of the supposed oldest monocotyledonous 
plants have been shown to be either inorganic fossils, or to belong 
to some other class of plants. These Maidstone Lower Greensand 
stems, and the smaller Wealden examples of what appear to be 
the same form of plant, do not afford any trustworthy evidence 
of the existence of angiospermous plants at this horizon. The 
resemblance to Dracena or Pandanus does not bear the test of 
any careful comparison with the recent genera; in the fossils we 
have none of the regular transversely elongated leaf-scars so 
characteristic of these living monocotyledons. The method of 
branching and the general surface characters are much more in 
harmony with certain species of the genus Zamia. It may perhaps 
be advisable to institute a new generic name for this form of 
fossil stem, but for the present we are chiefly concerned with the 
small Wealden examples, and need not introduce any new term. 


V.-d162. -Pl Xi. Big. 5. 

This specimen is probably the impression of a stem at a point 
where branching is taking place; the surface is deeply and 
irregularly wrinkled, and studded with round or oval pro- 
minences showing no regularity of disposition. On the surface 
of the specimen there is a small amount of carbonaceous matter, 
which probably represents altered cortical tissue. A comparison 
of this specimen with Dracena, and with Zamia Loddigesi or 
Z. Skinneri, shows a much more striking resemblance to the latter 
genus. In these forms of Zamia, as in the fossil stems, there 
is not the characteristic armour of petiole bases, but a surface 
marked by transverse and irregular wrinklings, with here and 
there small knob-like protuberances. There is some slight 
resemblance to Saporta’s Changarniera inqurenda’; but this is 
described as a leaf, and not a stem structure. Very similar to 
some of the Maidstone specimens. Ecclesbourne. Rufford Coll. 


V. 2350. Pl. XII. Fig. 4. 

A portion of the specimen shown in the figure. The surface 
is slightly convex, suggesting a small segment of a large stem. 
The surface markings resemble those of V. 3162, but in this 


1 Saporta, Pal. Frang. vol. iv. p. 246, pl. cclxx. fig. 2, etc. 


ee TRUNCI. 


case they assume the somewhat more definite form of transversely 
elongated elliptical areas. Were it not for the convexity of the 
specimen, one might, perhaps, be inclined to regard it as a pith 
cast showing impressions of medullary rays. It closely resembles 
the stems referred to as Dracena Benstedtii. A very thin layer of 
a mineral substance occurs on the surface of the stem. LEccles- 
bourne. Rufford Coll. 


V. 2822. Part of a smaller stem than V. 3162; at one end 
it shows the same form of branching as in the figured specimen 
(Pl. XII. Fig. 5). Just below the place of origin of the large 
branch there is a smaller branch or lateral appendage. Inter- 
rupted transversely running ridges and numerous circular and 


elliptical scars constitute the surface features. Ecclesbourne. 
Rufford Coll. 


V. 2170. An impression of what seems to be the surface of 
a stem like V. 2322, ete. Ecclesbourne. Rufford Coll. 


Trunci, etc. (incertz sedis). 


V. 2807. Small specimen of some cylindrical structure ; surface 
marked with irregular longitudinal ridges; apparently crushed. 
Mr. Rufford suggests that this may be the axis on which a 
williamsonian fructification was borne. Ecclesbourne. 


Rufford Coll. 


V. 2259. Similar specimen, but more like some pith cast. 
Cf. Fittonia insignis, Sap. (Pal. Franc. vol. 1. pl. lvi.). Kecles- 
bourne. Rufford Coll. 


V. 2133. This specimen shows several more or less rhomboidal 
scaly structures, which may be the petiole bases of a cycadean 
stem. Ecclesbourne. Rufford Coll. 


V. 3237. Possibly the impression of a petiole base, showing 
what appear to be the impressions of vascular bundles, and other 
smaller black spots, which may be gum canals. Lcclesbourne. 


Rufford Coll. 
V. 21382a. V. 3187. Ecclesbourne. Rufford Coll. 


WITHAMIA. 173 


Specimens of Doubtful Position. 


Genus WITHAMIA, gen. nov. 


In the second volume of the Plantes Jurassiques, Saporta figures 
two specimens under the generic name Cycadorachis,' C. armata,’ 
Sap., from the Lower Kimmeridgian, and C. abscissa,* Sap., from 
the same horizon. The former species is represented by a fairly 
stout axis bearing four spinous recurved appendages, having the 
appearance of rose thorns. This is not unnaturally compared 
with the rachis of a cycadean petiole, in which, as in recent 
species of Cycas and Dioon, the lower pinne are reduced to 
spiny processes. 

The genus is thus defined :— 

‘¢ Rachides frondium foliolis destitute: vel etiam frondium partes 
infer, petioli dicte, sive nude sint, sive aculeis armate, aut 
ad basin insertionis causa paullo dilatatam squamatis e tomento 
piloso constantibus ad utrumque latus predite videantur.” 

Such a provisional genus like that of Rachiopteris among ferns, 
is a useful institution, and the species Cycadorachis abscissa may 
well be included in it; but the discovery by Mr. Rufford of 
several specimens very similar to Saporta’s C. armata, negatives 
the suggested relationship to a cycadean frond. 

In the Ecclesbourne (Hastings) specimens there are large leaf- 
like structures attached to the axis in the axils of the spines, 
and, without attempting to speak definitely as to the precise 
nature of these two kinds of appendages, it would seem unwise 
to retain a generic designation indicating a cycadean alliance. 
Although it is held by some a wrong course to adopt, I propose to 
substitute, in the case of Cycadorachis armata, Sap., and the almost 
identical fossils from the English Wealden, a new generic name in 
place of that instituted by Saporta. To retain Saporta’s genus, with 


1 Pal. Franc. vol. ii. p. 193. 
2 Ibid. p. 196, pl. exvii. fig. 1. 
3 Ibid. p. 198, pl. exiv. fig. 3. 


174 WITHAMIA. 


the recently discovered specimens before us, would be practically 
equivalent to assigning the plant to a position which appears to 
be entirely at variance with the facts. I propose, therefore, to 
institute the new genus Withamia for these spiny axes with 
leaf-like appendages, and in doing so to place on record some 
slight recognition of the immensely important service which 
Witham of Lartington rendered to paleobotanical science. The 
Internal Structure of Fossil Vegetables‘ is widely known as a 
classic work marking the beginning of a new method of in- 
vestigation; but so far as I am able to discover, the name of 
the author of this epoch-making book has not been made use 
of as a genus of plants. We may define Withamia as follows :— 

A woody axis bearing two rows of spiny appendages, in the 
axils of which are borne flat leaf-like appendages. 


Withamia Saporte, gen. et spec. nov. 
[Pl OTs Bigs, doand)2); Pl. oVii Fie. 13] 


Type. Specimens in the British Museum. 

There is a very strong likeness between Saporta’s species, 
C. armata, and the English specimens as regards the axis and 
recurved spines, but the absence of any leaf-like appendages in 
the former, and the difference in geological age, render it advisable 
to adopt a new specific name for the present examples of the 
genus. I have chosen as a specific designation the name of the 
author who first described this form of fossil plant. The species 
may be defined as follows :— 

Axis having a breadth of about lem., striated longitudinally, 
bearing stout recurved spines arranged laterally in two rows, 
and at slightly irregular intervals. In the axils of the spinous 
processes there are attached more or less orbicular or obcuneate 
leaf-like structures, having a distinct flabellate (Cyclopteris type) 
venation. 


1 Edinburgh, 1833. 

2 For Saportaia on Pl. II, substitute Withamia; the former name being too 
nearly identical with Saportea (Fontaine and White, The Permian or Upper 
Carboniferous Flora. Harrisburg, 1880). 


WITHAMIA. lige 


A purely provisional genus like Withamia seems decidedly 
preferable for the present species, to one which in any way 
implies a definite botanical position. It is by no means clear 
how such a plant can well be included in the Cycadacee; and 
we have no evidence of sufficient value to enable us to assign 
the species to any other particular group. In the English Wealden 
rocks there has not so far been found any trustworthy record 
of an angiospermous plant. The Conifers and Ferns may be 
mentioned as possible groups in which to include this species, 
but as yet we have not sufficient evidence to warrant the selection 
of a generic name, which would imply a connection with one or 
other of these sub-classes. 

In a letter written about a fortnight before his death, the 
Marquis of Saporta wrote to me at some length in answer to 
an expression of doubt on my part as to the cycadean nature 
of his Cycadorachis armata, and our more perfect English species. 
The following sentences are taken from his letter, written on 
January 10th, 1895'; his words may be quoted in full; they 
are valuable, not merely as giving the opinion of one so well 
qualified to speak on such a question, but as some of the last 
from the ready pen of this indefatigable and accomplished student. 

“Je suis en effet ravi l’apprendre que vous avez rencontré 
dans votre Wealdien une portion de fronde, encore munie de 
pinnules en place de mon Cycadorachis armata (Pal. frang. 
Cycadées, p. 195, pl. exvii. fig. 1). Point de doute relativement 
a lV’étroite conformité de votre échantillon avec le mien les épines 
sont égales des deux parts et distribuées de la méme facgon sur 
le rachis. Du Kimmeridgien au Wealdien la distance verticale 
n’est pas telle que la méme espéce de Cycadées n’ait pu se 
maintenir et réparaitre sans changement appréciable. Le type 
est assurément curieux, et mérite d’obtenir une dénomination 
générique. A votre place je donnerais 4 ce type de Cycadées le 
nom d’ Acanthozamites. Remarquez d’abord, cher Monsieur, qu’il 
n’y a dans la présence de ces épines acérées disposées le long du 
rachis de la fronde rien d’insolite et pour en étre persuadé vous 
n’avez qu’ad consulter la figure 1, pl. xi. du volume des Cycadées 
de mes plantes Jurassiques. Cette figure represente la base d’une 


1 The Marquis of Saporta died at Aix-en-Provence on January 26th, 1895. 


176 WITHAMIA. 


fronde de Cycas avec les épines, qui sauf la dimension plus petite 
sont pareilles a celles du type fossile Wealdien et Kimmeridgien. 
Seulement au lieu d@’un Cycas ou @un Dioon, comme je le 
présumais, nous avons ici un type eteint dont les frondes portaient. 
En méme temps des épines et a leur aisselle vers milieu du rachis 
des folioles, sans doute caduques dont le ressemblance avec celles 
du Sphenozamites latifolius, Brong. (Pal. frang. Cycadées, pl. exvil. et 
pls. exii. and exiii. pour les folioles de Sphenozamites), doit étre prise 
en considération. Il l’agit seulement de décider si la présence de 
ces épines constitue un caractére générique ou seulement spécifique, 
puisque nous savons par |’échantillon de ma planche cxi., que le 
rachis du Sphenozamites latifolius n’était pas armé d’appendices 
épineux. Je crois que dans V’incertitude on est fondé a reconnaitre 
au moins dans cette particulité Vindice d’un sous-genre ou section 
a part quw’il est naturel de désigner par un dénomination a part, 
comme serait celle de Acanthozamites que je proposais plus haut ou 
toute autre a votre convenance. 

Mon sentiment est done ici que le rapprochement avec les 
Phyllocladus waurait aucune vraisemblance tandis que celui avec 
les Cycadées et les Sphenozamites en particulier dont étre adopté 
comme le plus naturel.” 

My reason for not adopting the genus Acanthozamites, as 
suggested by Saporta, is that it suggests a cycadean affinity 
which is hardly supported by the nature of the specimens. As 
regards the spines, these in themselves are by no means opposed 
to a cycadean rachis, but the structures in their axils seem to 
me quite inconsistent with the morphological character of any 
recent cycadean frond. Mr. Carruthers suggested to me that the 
specimens show some resemblance to certain ferns, and expressed 
an opinion in favour of the /ilicine as the most likely plants 
with which to compare the fossil species. We have various recent 
fern fronds which are more or less spinous'; but the general habit 
of the fossil form, the nature of the spines, their disposition on 
the axis, and their definite relation to the leaf-lke structures 
constitute important points of divergence from any living / wlicine. 


1 An interesting form of fossil fern, Gleichenia Hantonensis, is figured by 
Starkie Gardner from the Eocene plant-beds of Bournemouth, in which strongly 
recurved climbing organs are preserved. [Gardner (2), p. 60.] 


WITHAMIA. 7/7 


In answer to my question as to the probability of such a fern- 
affinity, Mr. Baker, of the Kew Herbarium, wrote as follows: 
‘‘T should not think this very curious fossil is likely to be a 
fern. Phyllocladus seems far more likely. But, of course, without 
flower and fruit there can be no certainty. The climbing stem 
and hooked prickles recall Calamus.” I had suggested the 
possibility of the Wealden plant being compared with the New 
Zealand conifer, Phyllocladus,' in the recent species of which we 
have small scaly leaves subtending flattened cuneiform branches 
(phylloclades). If the leaves were modified into climbing-hooks, 
we should have a fairly close approximation to Withamia, but 
the evidence at hand does not allow of any great weight being 
attached to such a comparison. In a palm such as Calamus or 
Desmoncus we find somewhat analogous spines, but in these 
monocotyledonous plants, there is not the same relation as regards 
position on the axis between hooks and leaf segments as in 
Withamia. For the present, then, I propose to leave the position 
of Withamia an open question, in view of the difficulty of deciding 
the morphological value of the stout recurved spines and leaf-like 
appendages, and the insufficient evidence afforded by incomplete 
vegetative structures. 


Wa2ziod, Plt, Pig; 2. 

Length of axis 12°2cm., about lem. broad, striated longi- 
tudinally. In the axil of each stout recurved hook there occurs 
a portion of a leaf-like appendage; these leaves or phylloclades 
are imperfectly preserved, but enough is seen to demonstrate the 
flabellate venation, and to suggest a form similar to that of 
the detached ‘‘leaf” represented in Pl. IL. Fig. 1 (V. 2195). 
The markings shown on the surface of the uppermost left-hand 
spine are merely cracks, and not the remains of any original 
structure; the two highest spines are attached to the axis in 
a manner indicative of an alternate arrangement, the middle pair 


are opposite, and the lowest subopposite. Ecclesbourne. 
Rufford Coll. 


We29t5e Pie Bie. 1. 
This well-preserved Cyclopteris-like leaf appears to have been 


1 Species occur in New Zealand, Tasmania, and Borneo. 


178 WITHAMIA. 


sessile; the veins are numerous and clearly shown. The irre- 
gularity of the margin is probably an original character; in some 
other examples of these ‘‘leaves”’ the margin is much more indented, 
and the form of the ‘‘leaf” longer and narrower: cf. V. 2134e, 
V. 2798, etc. There is a distinct similarity between this speci- 
men and an unusually entire Ginkgo leaf. Saporta’s figure of a 
Sphenozamites latifolius' leaf agrees very closely with the Wealden 
specimen; if this form of leaf were attached to Withamia ( Cycado- 
rachis) armata it would make the resemblance between the French 
and English species still more striking. Ecclesbourne. 


Rufford Coll. 


V. 2134c. Pl. V. Fig. 1. 

Axis 18cm. in length, and 9mm. broad. The longitudinal 
striations very clearly shown, also the stout nature of the hooks. 
Very incomplete fragments of the flat appendages in the spine 
axils. The spines are less regularly placed than in V. 2184 
(Pl. II. Fig. 2) and farther apart. Cf. Cycadorachis armata, Sap. 
Ecclesbourne. Rufford Coll. 


V. 2805. A short piece of an axis with two well-preserved 
recurved spines. No leaves shown. 


V. 28050. 20cm. long, showing seven spines. Traces of 


flattened appendages in the axils of some of the spines. Eccles- 
bourne. Rufford Coll. 


V. 21345. Portion of a large leaf, apparently about 7-8 cm. 
in length. A good specimen of Sphenopteris Kontainet, Sew., 
on the same piece of rock. 


V. 2184c, V. 2184d, V. 2134¢, and V. 2134f. Portions of 


‘‘leaves,’”? showing venation. Ecclesbourne. Rufford Coll. 
? to} “ 


V. 2182, V. 2732, and V. 2798. Specimens of ‘‘leaves” Some 
have a more cuneiform shape than the example figured (V. 2915, 
Pl. IL. Fig. 1). Ecclesbourne. Rufford Coll. 


1 Loe. cit. pl. exiii. 


BECKLESIA. 179 


V. 2923. Axis 43 cm. long, 1-1:3cm. broad. Portions of four 
hooks seen on one side, smaller than those of 2134a (Pl. V. Fig. 1). 
The appearance of this specimen is suggestive of a hollow axis, 
but this is probably due to the preservation of the cortex apart 
from the internal woody tissue. Ecclesbourne. Rufford Coll. 


Genus BECKLESIA, gen. nov. 


The specimens included under this genus are difficult to describe 
with any completeness, on account of the fragmentary and 
imperfect nature of the material. So far as I have been able 
to discover, it is impossible to include these fossils in any known 
genus; the above name is therefore proposed as a convenient 
generic term, and one which does not imply any exact botanical 
position. The National Museum owes some of its valuable 
examples of Mesozoic plants to the enthusiasm of the late 
Mr. Beckles; I have therefore made use of his name as a generic 
designation. As a specific name for the few examples referred to 
the genus Becklesia, the term anomala may be adopted. 


Becklesia anomala, gen. et spec. nov. 
[Pl. XIV. Figs. 2 and 3.] 


Type. Fragments, British Museum; from Ecclesbourne, near 
Hastings. 

The type species of the genus may be defined as follows :— 

Axis comparatively broad, giving off (on one side?) stout and 
stiff branches, attached to the axis of higher order in different 
positions, either laterally or on the surface, and at irregular 
intervals. On one surface the branches show a number of 
parallel longitudinal striations, and on the other surface a broad 
median rib with a small groove on either side. 

The specimens are, however, too imperfect to admit of any 
satisfactory generic or specific diagnosis. 

In his monograph on fossil cycadean stems, Carruthers makes 
a brief reference to a specimen found at Maidstone, which is 
spoken of as possibly a bennettitean frond. This fossil bears 


180 BECKLESIA. 


in some respects a distinct resemblance to the present species. 
Carruthers thus describes the Maidstone specimen: ‘‘It is a very 
large leaf, with numerous long linear segments, attached very 
obliquely to the rachis. The segments are simple on the upper 
part of the frond, but the lower ones give off, at regular distances, 
several long and slender ultimate segments.’’?! Mr. Carruthers 
afforded me an opportunity of examining these Lower Greensand 
specimens, and suggested that the Wealden examples figured in 
Pl. XIV. Figs. 2 and 3, represent the lateral segments of the 
Maidstone ‘‘frond”’ with their lateral long and slender segments. 
Although there undoubtedly exists a distinct resemblance, yet 
one cannot speak at all positively as to the identity of the two 
sets of fossils. In the Wernsdorf flora of the northern Carpathians, 
an abundant and characteristic species is that described by 
Ettingshausen as Thuctes Hoheneggeri, Ett.,2 and afterwards by 
Schenk as Frenelopsis Hoheneggert (Htt.).2 Of the specimens so 
named, some of those figured by the latter author present a more 
or less close resemblance to Becklesia anomala; this is especially 
the case with those represented in Schenk’s pl. v. figs. 1 and 2. 
Frenelopsis was proposed by Schenk as a generic name for plants 
having a similar habit to the recent genus Frenela, and possessing 
among other characters cylindrical articulated branches bearing 
small scaly leaves. The majority of the Wernsdorf examples of this 
genus show these characters very clearly, but those in pl. v. figs. 1 
and 2 are apparently without them, and in some degree conform 
to the present species. Without examining Schenk’s material, 
it is impossible to speak definitely as to the exact nature of these 
particular examples; it may be that a difference in age or manner 
of preservation, is sufficient to account for the apparent absence 
of the articulations and small leaves. Schenk‘ speaks of these two 
specimens (pl. v. figs. 1 and 2) as older examples, in which the 
leaves are only partially preserved. The chief point of contact 
between the Wealden fossils and those from the Wernsdorf beds, 
described as older portions of Frenelopsis Hoheneggeri, lies in the 


1 Carruthers (1), p. 697 (footnote). 

2 Ettingshausen (A. 4), Abh. k.-k. geol. Reichs. vol. i. Abth. iii. No. 2, 
p. 26, pl. i. figs. 6 and 7. 

3 Schenk (A. 3), Paleeontographica, vol. xix. p. 18, pls. iv.—vil. 

4 Loe. cit. p. 14. 


BECKLESIA. 181 


long and stiff lateral branches. In the English specimens there 
are no signs of any articulations or of leaf structures; so that 
they cannot well be included in the genus Frenelopsis. The 
typical form of the genus is well illustrated by Ettingshausen’s 
figures, as also by those of Heer! and Saporta.” Jn Fontaine’s 
Potomac Flora, several specimens are referred to Schenk’s genus, 
but these have recently been transferred by Nathorst to a new 
genus, Pseudofrenelopsis,t on the ground that the American forms 
have been incorrectly interpreted by Fontaine. As regards habit, 
there is some slight resemblance between Becklesia and Camptopteris 
spiralis, Nath.,° from Bjuf. On the whole, however, Carruthers’ 
specimens offer the greatest similarity to the following fragments, 
the nature of which must be left entirely unsettled. 


V. 2361. Pl. XIV. Fig. 2. 


In the portion of the specimen represented in the figure the 
characters of the lateral segments are fairly clearly shown. A small 
piece of the branch at the right-hand upper corner of the drawing, 
shows the parallel striation and apparently woody nature of the 
segments; a little below, this branch is crossed by another in 
which the broad median ridge may be seen. Most of the lateral 
segments are flattened, and do not present such distinct surface 
features. 


Vv 286lese BPochye Pig. '3: 

The flattened main axis fairly distinct, with the irregularly 
placed lateral branches. The third branch from the top does not 
appear to arise laterally, but rather from the exposed face of the 
broader axis. 


V. 23614. Smaller fragment. Ecclesbourne. Rufford Coll. 


1 Heer (A. 6), Secc. Trab. Geol. Portugal, p. 21, pl. xii. figs. 3-7. (The 
specimens figured by Heer in his Flor. foss. Arct., and referred to Frenelopsis, 
are probably not true examples of this genus.) 

2 Saporta (1), pp. 113 and 139, pl. xxi. figs. 9-11, and pl. xxvi. fig. 16. 

3 Fontaine (A. 2), p. 213, pls. xcv.-xcix., exi., exii., and clxviil. 

4 Nathorst (5). 

5 Ibid. (A. 1), p. 33, pl. iii. 


182 BECKLESIA. 


Cf. Becklesia anomala, sp. nov. 
[Pl. XIV. Fig. 1.] 


V. 2608. Pl. XIV. Wig. 2. (4 ‘mat. size.) 


The nature of this specimen is very doubtful, and its imperfect 
preservation does not allow of any accurate description. Length 
46cm., breadth about lem. From the central flattened axis a 
number of comparatively straight lateral branches are given off 
at irregular intervals; many of these appendages are separated 
from one another by about 1°5cm., and have a breadth of 3cm.; 
they are linear in form and of a uniform breadth; one branch, in 
which the tip is not shown, measures 13cm. There appear to be 
a small number of parallel veins in each segment. In some cases 
the branches appear to bifurcate close to the point of attachment 
to the central axis. Possibly we have here a larger specimen of 
Becklesia, but the occurrence of branches on both sides of the axis 
makes it difficult to be at all certain as to specific or even generic 
identity with the previous specimens. Among recent plants there 
is a form of MMacrozamia heteromera, M. heteromera var. glauca, 
Moore,’ in which the bifurcate pinne bear a certain resemblance 
to the Wealden fossil, but in the former the more regular dis- 
position of the segments affords an important point of divergence. 
Cf. Schenk’s Frenelopsis Hoheneggeri (Ett.),? as shown in pl. v. 
figs. 1 and 2; also Camptopteris spiralis, Nath.® Beckles Coll. 


V. 2359. Two detached forked segments, probably the same as 
V. 2608. 


1 Moore, p. 4. 
2 Schenk, Joe. cit. 
3 Nathorst, doc. cit. 


DICHOPTERIS. 183 


Genus DICHOPTERIS, Zigno. 
[Mem. Instit. Veneto, vol. xii. p. 217, 1864.] 


This genus is defined by Zigno as follows :— 

‘“‘Frons bipartita, bipinnata, rachide primaria, crassa, striata. 
Pinne liber, pinnatifide, alterne, vel subopposite. Pinnule 
coriaceee integerrime, seepe basi angustate, in rachides alatas 
decurrentes. Nervi «quales pauci, simplices, interdum furcati, 
e rachide seriatim orti, ad apicem marginemve pinnularum 
flabellatim excurrentes. Sori rotundi, prominuli, sparsi. Capsule 
(Sporangia) ovato-globose, sessiles, vel subsessiles, annulo completo 
cinctee. Filices elastic, rachide crassa bipartita, facie Gleichenia- 
cearum.” 

Zigno’s genus is classed by Solms-Laubach? with WVilssonia, 
Thinnfeldia, and others, which ‘‘have been shifted backwards 
and forwards by different authors from cycads to ferns, and from 
ferns to cycads.” In looking over the references to Dichopteris 
by various writers, we find a considerable difference of opinion, 
both as regards the necessity for such a generic designation, in 
distinction to the much older genus Pachypteris of Brongniart, 
and as to the affinity of the plants described under this name. 
Schimper, in the first volume of the Trait. pal. vég.,’ includes 
Dichopteris under Pachypteris, and remarks that it is ‘impossible 
to doubt the identity of the two genera”; in the third volume ® 
of the same work, he includes the former among the ferns as an 
independent genus. Saporta includes some species of Dichopteris 
in his genus Seleropteris‘; e.g. the two plants originally described 
by Phillips from the Lower Sandstone and Shale of the Yorkshire 
Coast as Sphenopteris lanceolata,’ Phill., and Neuropteris levigata,® 
Phill., but the nature of these species has been a matter of 


1 Fossil Botany, p. 87. 

2 p. 492. 

3 p. 490. 

4 Saporta, Pal. Frang. vol. i. p. 364. 
5 Phillips (A. 2), p. 200, pl. x. fig. 6. 
6 Ibid. p. 201, pl. x. fig. 9. 


184 DICHOPTERIS. 


much discussion and need not be considered here. Feistmantel ' 
describes certain plant remains from the Gondwana flora of 
India, which closely resemble Zigno’s species; he prefers to go 
back to Brongniart’s Pachypteris, and extends the original definition 
so as to make it embrace, not only plants with the ultimate 
segments “‘ enerviis vel uninerviis”’? but those in which the veins 
are more numerous. In Schenk’s monograph on Die fossile Flora 
der Grenzschichten ... . a specimen is figured as Dichopteris 
incisa, Schenk,® but, as Feistmantel suggests,‘ the characters do 
not seem to agree with Zigno’s genus. The larger and more 
perfect specimens of Dichopteris figured by Zigno’ would seem to 
favour the inclusion of such plants among the Filicine; but, as 
Schenk points out,® the fructification is too indistinct to be of 
any taxonomic value. It is safer, therefore, while expressing 
a bias towards the pteridophytic nature of the genus, to speak 
of it as occupying a somewhat doubtful position. 


Dichopteris, sp. Cf. D. levigata (Phill.).’ 
[Pl. XII. Fig. 6.] 


V. 3145. Part of a single pinna, showing the coriaceous 
ultimate segments without any distinct venation. 

Cf. Dichopteris Visianica, Zig., D. levigata (Phill.), and 
Scleropteris Pomelii, Sap.6 Ecclesbourne. Rufford Coll. 


1 Foss. Fl. Gond. vol. i. p. 29. 

2 Brongniart (A. 8), Hist. vég. foss. p. 166. 

3 Schenk (A. 1), p. 121, pl. xxviii. figs. 5-8. 

4 Loe. cit. p. 30. 

5 (1), pls. xii. and xiii. 

6 (A. 8), Schenk’s Handbuch, p. 41. 

7 =Neuropteris levigata, Phill., Pachypteris levigata (Phill.), Scleropteris 
levigata (Phill.). 

& Saporta, Joe. cit. pl. xlvii. 


CONIFER. 185 


Order CONIFER Z. 


Stem much branched, leaves usually small and simple. Flowers 
unisexual and without a perianth, plants moneecious or dicecious. 

The past history of the Conzfere is but imperfectly known, and, 
owing to peculiar difficulties connected with the determination of 
fossil forms, the evidence of paleobotany as to the development 
and geological distribution of these plants, must be accepted with 
the greatest caution. It would take us far beyond the limits of 
the present work to discuss at length the distribution in time of 
coniferous types. In the Paleeozoic rocks there are various repre- 
sentatives of this Class, and we have an example, in such an 
extinct genus as Cordaites, of a synthetic type in which coniferous 
characteristics are combined with certain structural features met 
with in other Orders of gymnosperms. As a general rule, fossil 
conifers are perhaps the most unsatisfactory plants with which the 
paleobotanist has to deal: structureless and imperfectly preserved 
fragments of broken twigs, isolated cones, leaves or seeds, have 
usually to be determined separately, and it is only in comparatively 
rare instances that we are in a position to connect cones and vege- 
tative branches. Coniferous wood, with its mineralized tissues more 
or less well defined, is met with in rocks of nearly every age, 
but here, again, the stems or thick branches must be determined as 
far as possible from histological structure alone, and without any 
leafy twigs or reproductive organs. Goppert,’ Kraus,’ Kleeberg,! 
Felix,’ Schenk,” Knowlton,*® and others have attempted to devise 
convenient methods of classifying and identifying fossil Conifere by 
means of the peculiarities of structure presented by the secondary 
wood and the distribution of resin ducts. For the most part, 
however, fossil conifers are represented by structureless casts or 
impressions of leafy branches, occasionally bearing characteristic 
cones or other forms of reproductive organs. 

In treating of the Cycadacee, some general account was attempted 
of the difficulties and possible sources of error which ought to be 


1 For references see Solms-Laubach’s Fossil Botany. 

2 Zittel (A.), Handbuch, p. 848. 

5 Knowlton (A. 2), Bull. U.S. Geol. Surv. No. 56, 1889. (See also Géppert 
and Menge, Die Flora des Bernsteins, vol. i., and Conwentz.) 


186 CONIFERZ. 


kept in mind in the identification of fossil specimens. It may be 
useful to draw attention to similar difficulties in the case of 
Conifere, which have not always been observed by palobotanical 
writers. 

If we examine the external characters of older branches of recent 
conifers from which the leaves have been detached, it will be found 
impossible to institute on such a basis any useful classification. It 
happens, not infrequently, that the leaves and cortical tissues 
become readily detached from the surface of the wood, leaving 
a smooth axis in place of the corticated branch or stem. A good 
example of this is afforded by such specimens as those represented 
in Pl. XVII. Figs. 4-6. Occasionally we have to deal with pith 
casts having the surface covered with lozenge-shaped prominences, 
simulating elongated leaf bases. A good example of such a 
medullary cast is afforded by Weiss’ genus Zylodendron, of which 
the true nature was pointed out by Potonié! in 1887. Again, in 
some specimens of the Triassic Voltz’a* we have smaller pith casts 
of similar form. In his Zntroduction to the Study of Paleontological 
otany, Balfour* calls attention to the unnecessary multiplication of 
fossil species, and illustrates the need for careful observation of the 
characters of recent stems, by reference to the striking differences 
presented by a branch of Araucaria imbricata, Pay., when the bark 
is viewed intact, and after it has been more or less completely 
stripped off the surface of the wood. In Araucaria Cunninghami, 
Ait., we find equally striking contrasts between the younger 
branches, with their stiff faleate leaves, the slightly older stems, on 
which only the rhomboidal leaf bases are left, the smooth surface of 
the wood, from which overlying tissues are readily detached, and 
finally, the surface features presented by a pith cast, 

In the long needles of Pinus and the broad flabellately veined 
leaves of Ginkgo, we have sufficiently well-marked characters to 
enable us in most cases to arrive at a generic determination. In 
many instances, however, it is a hopeless task to attempt to found 
any accurate determination on leaves alone. Among recent genera 
we have a deciduous habit in such plants as Lariz, Ginkgo, Taxodium 


1 Potonié (2). 
2 Seward (4). 
3 Balfour, p. 4 


Ss ee 


CONIFERS. 187 


distichum, Rich. ; also, to a certain extent, in Sequoia sempervirens, 
Endl., Thuja occidentalis, L., Libocedrus decurrens, Torr., etc.!; but 
in most species the leaves remain on the tree for more than one 
year. Occasionally, the manner of occurrence of detached leaves or 
leafy shoots in a fossil state may afford evidence of the existence of 
deciduous species. A careful examination of branches of recent 
conifers bearing vegetative leaves, enables us to realize the im- 
possibility of relying for accurate determination or comparison on 
such uncertain characters as leaf form or arrangement. The uni- 
veined leaves of Podocarpus in some forms of the genus, may be 
confused with the foliage of araucarian species, in which the veins 
are imperfectly preserved ; in such a plant as Podocarpus andina, 
Popp., the long narrow leaves agree closely with those of some 
forms of Cephalotaxus, and the detached leaves of either bear a strong 
resemblance to single pinne of Cycas. A specimen of an unnamed 
species of Cephalotaxus in the British Museum possesses leaves 
measuring 11cm. in length and 4mm. broad, a close approach to 
the pinnee of Cycas species. The genus Agathis, e.g. A. Australis, 
Salisb., cannot be readily distinguished from some forms of Arau- 
caria, if we have only the leaves to guide us. The large leaves of 
Agathis Dammara, Rich., and the broad pinne of Podozamites 
cannot always be separated with certainty, at least in such 
specimens as do not show distinct venation. If we have not the 
general habit of the tree, or characteristic differences of colour to 
help us, it is practically impossible to discriminate with accuracy 
between the leafy twigs of many recent genera. Without entering 
into any detailed comparison of living forms from this point of 
view, we may note the close agreement between Zaxodium distichum, 
Rich., Sequova sempervirens, Endl., and Taxus baccata, L.; between 
Cryptomeria, sp., and Araucaria, sp.; between different genera of 
the Cupressine, etc. Similar examples might be readily multiplied ; 
but an examination of the recent species will at once demonstrate 
the futility of attempting generic distinction on such data, and will 
emphasize the unfortunate habit of some writers of applying to 
fossil fragments the unaltered names of recent genera. Another 
pitfall as regards leaf form, is the fairly common occurrence of 
heterophylly among coniferous plants. Writing in 1803, Lambert 


1 Stark. 


188 CONIFERZ. 


says:' “‘T must here observe a remarkable peculiarity belonging to 
the Conifere of the Southern Hemisphere, which is, that while the 
trees are young their leaves are long and divaricating, but when 
they become old enough to bear fruit, those leaves fall off, and are 
succeeded by short scales, closely imbricated on the branches, so 
that, seeing them in their different states, one could hardly suppose 
it possible that they could belong to the same species.” This 
young form of leaf may be retained for some few years before the 
adult foliage is developed, and thus present a possible source of 
error in the determination of fossil branches. In such a species as 
Pinus pinea, L., we have the young leaves retained for some con- 
siderable time previous to the development of the needles and short 
shoots. An interesting case of this difference between the leaves 
of young and adult plants was pointed out to me at the Royal 
Gardens, Kew: the young plants of the new species of Widdring- 
tonia, W. Whyter, Rend.,? from Nyassa-Land, bear comparatively 
long needles, in marked contrast to the small scale leaves of the 
older tree. A striking instance of a similar kind is afforded by 
some specimens of Araucaria excelsa, R. Br., in the Herbarium of 
the British Museum : there is a seedling with its long and spreading 
leaves, an older specimen with narrow and spreading leaves, and 
another with the stiff leaves of the adult plant. In Daecrydium 
Kirkii, F. Muell.,* from New Zealand, we find a marked difference 
between the small and closely adpressed leayes, and the much 
larger and more spreading leaves of other branches; also, in 
D. elatum, Wall., D. Westlandicum, Kirk.,* and other forms, there 
is a striking disparity in the leaf form. In Podocarpus cupressina,® 
R. Br., there is a decided difference between the young and old 
forms of leaves. In Athrotaxis selaginoides, Don.,° we have various 
forms of leaf, from the longer and more openly disposed to the 
smaller and closer leaves. Among other species exhibiting similar 
differences in the shape and size of the leaves, we may note the 
well-known Juniperus Chinensis, L., Biota orientalis, Endl., Juni- 
perus Bermudiana, L., Glyptostrobus heterophyllus, Endl., ete. 


1 Lambert, p. 89. 

2 Rendle, p. 60. 

3 Hooker (1), pl. mecxix. 

4 Tbid. pl. Meexviil. 

5 Brown and Bennett, pl. x. 
§ Don, pl. xiv. 


CONIFERS. 189 


Examples of heterophylly have been recorded among fossil forms— 
e.g., Voltzia heterophylla, Brong.,' from the Bunter beds; and the 
specimens of Sequoia Tournalit (Brong.), figured by Gardner® from 
the Bagshot beds of Bournemouth, show a considerable difference 
in the form of the leaves. The microscopical examination of the 
epidermal cell-outlines of fossil coniferous leaves has been success- 
fully adopted in some instances, e.g. by Zeiller* in the case of 
Frenelopsis Hoheneggert (Ett.), and by Schenk ‘ in several instances. 

In addition to the similarity of leaf form in different species and 
genera, and the heterophylly in the same species, it is important 
to note the common occurrence of more than one method of leaf 
arrangement in the same tree. Masters,’ in his useful paper in 
the Journal of the Linnean Society for 1891, has drawn attention to 
this variation in leaf arrangement among recent species of conifers. 

In describing eycadean flowers, it was pointed out how difficult 
it is in some cases to distinguish between the cones of cycads and 
those of certain genera of conifers, when we have only external 
form to guide us. The seeds of Cephalotaxus, Ginkgo, Torreya, and 
other genera may be easily mistaken for those of Cycas and other 
cycads. There is in many instances, the same difficulty in identifying 
the detached cones of recent conifers as in determining detached 
leafy twigs. Schimper and Mougeot, recognizing the difficulty of 
discriminating between fossil cones, suggested the general generic 
term Strobilites,’ which they used in a somewhat similar sense to 
that in which I have used the more comprehensive genus Conites. 

Hitherto the number of Conifere recorded from English Wealden 
strata has been extremely small. In addition to isolated cones 
described by Carruthers, Gardner, and others, we have only one 
species represented by a leaf-bearing branch—Sphenolepidium Kur- 
rianum (Dunk.). The Rufford Collection has enabled us to recognize 
as British plants several of the species previously described from 
Germany and elsewhere, and to make several additions to the list 
of Wealden Conifer. 


 Brongniart (5), p. 451. See also Schimper and Mougeot, p. 22, pls. vi.-xiv. 

2 Gardner (2), pl. v. 

3 Zeiller (3), p. 231, pl. xi. 

* Schenk (A. 1), Fl. foss. Grenz. Keup. Lias, and (A. 2) Paleontographica, 
vol. xix. 

> Masters, p. 244. 

6 Schimper and Mougeot, p. 31. 


190 ARAUCARITES. 


PryoiwEx — ABIETINEZ—ARAU- . Araucarites cf. Conites elegans (Carr.), 
CARINE. etc. 
Araucarites, sp. 
Prnoipe#—ABIETINEZ—ABIETINE. Pinites Dunkeri, Carr. 
Pinites Carruthersi, Gard. 
Pinites Solmsi, sp. nov. 
Pinites Ruffordi, sp. nov. 
Pinites, sp. 
PrnoiwExZ—ABIETINEZ—TAxopDINz Sphenolepidium Kurrianum (Dunk.). 
Sphenolepidium Sternbergianum (Dunk.). 
Sphenolepidium ct. S. subulatum (Heer). 
PrnoiwEx#—CUPRESSINEZ . . . . Thuites, sp. 
TaxoDIEZ—PopocaRPE# . . . . Nageiopsis, sp., cf. NV. heterophylla, Font. 
GENERA ET SPECIES INCERTH SEDIS . Pagiophyllwm crassifolium, Schenk. 
Pagiophyllum, sp. 
Brachyphyllum obesum, Heer. 
Brachyphyllum spinosum, sp. nov. 

In the above list of Conifere described in this volume, I have 
suggested the probable position of various species in the classifica- 
tion adopted in Engler and Prantl’s recent work. It must, 
however, be admitted that at present we cannot feel great con- 
fidence in the attempts to determine, even the approximate affinities 
of such provisional genera as Sphenolepidium, Pagiophyllum, and 
Brachyphyllum. 


Genus ARAUCARITES, Presl. 
[Flor. Vorwelt, Fasc. vii. p. 203, 1838.] 


In the present instance this genus is used as a convenient 
designation for certain female cones which resemble, in their 
form and structure, those of the recent genus Araucaria. 


Araucarites (Conites), sp. Cf Conites elegans (Carr.) 
and Kaidocarpon minor, Carr. 


RES Xie owics- 5] sande2=]] 


It has already been pointed out that the specimen figured by 
Carruthers as probably a male cone of Bucklandia has little or no 
claim to be regarded as cycadean; it very closely resembles the 
specimens figured in Pl. XII. Figs. 1 and 2, and like them should 


ARAUCARITES. 191 


probably be referred to the genus Araucarites. Another fossil 
which is probably identical with these Wealden cones is that 
described by Carruthers as Aaidocarpon minor from the Potton 
beds of Bedfordshire; the type specimen’ of this species in the 
Woodwardian Museum shows the general characters of a somewhat 
waterworn female araucarian cone. The same form of cone, but 
one belonging to another species, is illustrated by a beautiful 
specimen in the York Museum, which has been described and 
figured by Carruthers from the Coralline Oolite of Malton, in York- 
shire, under the name of Araucarites Hudlestoni.2, Mr. Carruthers 
tells me he is disposed to regard some of the fossils described 
by him as monocotyledonous inflorescences, as more probably 
araucarian cones. It is proposed to discuss elsewhere, at greater 
length, the value of several of the published records of supposed 
monocotyledonous plants from Jurassic and Wealden strata. 


W. 2180: - Pl XII, Fig. 2: 

In this specimen we have a view of the proximal ends of the 
scales, their broad and flattened form is clearly seen, also the 
lozenge-shaped cavity in which the seeds were originally situated. 
The form of the scales and the more or less globose form of the 
cone, present a strong resemblance to the female strobili of species 
of Araucaria; e.g., ef. the figure given by Martius in pl. ex. of 
his Flora Brasiliensis, with the specimens in our Pl. XII. Figs. 
1 and 2. 


ViocL80q) (Rls hl, Fic. 1. 

The stout central axis is clearly shown, with the spirally 
disposed points of insertion of the broad scales. Breadth of axis 
about 4cem.; the scales probably wider towards the distal end, 
showing prominent lateral angles or wings with a slightly convex 
and wrinkled upper and lower surface. In one part of the 
specimen the impressions of the scale apices show a prominent. 
distal end with a central dot, as in V. 2148. Some of the scales 
show a clearly defined cavity, originally occupied by the small 
seeds which were narrowed towards the cone axis. There are 
three small seeds which seem to have fallen out from this cone. 
Ecclesbourne. Rufford Coll. 


1 The specimen has not been figured. 
? Carruthers (8). 


192 ARAUCARITES. 


V. 2331. Two specimens, in the form of oblique sections, of 
a cone, probably belonging to the same species as V. 2180. Cf. 
Carruthers,’ pl. vi. figs. 1 and 9. Ecclesbourne. Rufford Coll. 


V. 21804. Two specimens, showing an impression of the cone 
surface. In size, and as regards the form of the scales, very 
similar to Carruthers’ supposed bucklandian male cone, pl. liv. 
fig. 6. Cf. V. 2148. Possibly this specimen should be referred 
to another species; it is smaller than V. 2180 (Pl. XII. Fig. 2). 
Ecclesbourne. Rufford Coll. 


V. 2148. The impressions of the distal ends of the scales show 
a small central depression, corresponding to an umbo on the 
scale apex. This and other examples should be compared with 
Araucarites Pippingfordensis (Ung.), the original of which is in 
the Museum Collection. Ecclesbourne. Rufford Coll. 


V. 22652. Scales clearly preserved, showing in side view 
a longitudinal depression, suggesting a shrinking of the seed 
cavity. V. 2265. <A well-preserved, but smaller specimen. 
Ecclesbourne. Rufford Coll. 


V. 2277. Cf. V. 2148. V.3185. The thick central axis clearly 
shown, also impressions of scales in side view. LEcclesbourne. 


Rufford Coll. 


V. 3173. Cone in cross section; scales with seeds clearly pre- 
served. Other specimens referred to this form of Araucarites : 


V. 2245, V. 2268, V. 2263, V. 2264, V. 2279. 


Araucarites, sp. 


V. 2266 and V. 2280. Two specimens of small subspherical 
cones imperfectly preserved ; of the same form as the preceding 
examples, but considerably smaller. Ecclesbourne.  Rufford Coll. 


1 Carruthers (8). 
2 Ibid. (1). 


PINITES. 193 


Genus PINITES, Endlicher. 


[Synopsis Coniferarum, 1847, p. 283.] 


In dealing with detached and imperfect cones, in which the 
scales have a flattened form like those of -Adzes and certain 
species of Pinus, it is difficult, or indeed impossible, to arrive at 
a very accurate generic determination. The use of Endlicher’s 
genus in a wide sense is, therefore, a matter of convenience, and 
in most cases preferable to the application of the generic name of 
Pinus to detached cones which cannot be referred with absolute 
confidence to a narrowly defined recent genus. 

Endlicher defines this genus as follows :— 

‘‘Folia, amenta staminigera et strobili, diversis Pinuum speciebus 
similes.”” The term is a convenient one to adopt, if we do not 
confine its use within the limits of the genus Pinus as defined in 
modern systematic works. 

Several detached cones have been described by Carruthers, 
Gardner, and others from the Wealden rocks of England, under 
the generic name Pinites. Their general character justifies the 
choice of this genus, but an examination of several of the type 
specimens lends no support to the existence of so many distinct 
species as have been described. In the 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, 
Starkie Gardner figures and describes the following new species: 
Pinites valdensis, P. Carrutherst, P. eylindroides, and P. Pottoni- 
ensts, from the Wealden rocks of the Isle of Wight and the 
Lower Greensand of Potton. The Potton specimens are pro- 
bably of Wealden age. In the case of Pinites cylindroides, from 
Potton, Gardner describes the solitary specimen as being ‘in 
excellent condition, certainly not derived from any older bed.” 
An inspection of the type specimen in the Woodwardian 
Museum, Cambridge, leads me to unhesitatingly describe it as 
distinctly worn and rolled, and imperfectly preserved. The figure 
does not convey a very accurate idea of the actual fossil; the scales 
are very imperfect, and their half-moon form spoken of by the 
author of the species, is almost certainly due to wearing, and 
cannot, I believe, be accepted as an original character. 

0 


194 PINITES. 


Pinites cylindroides may possibly be identical with P. Dunkeri, 
Carr., and even with P. Carrutherst, P. Pottoniensis, and P. valdensis. 
The type specimen on which the species P. Pottoniensis is founded 
is too small to admit of any specific diagnosis, and does not appear 
to me to have any claim to be regarded as specifically distinct from 
P. cylindroides. It must be admitted, however, that there is 
considerable risk in attempting to discriminate between these 
imperfect and detached cones; but it is surely a mistake to 
multiply the number of species without stronger evidence for the 
existence of any real specific differences. 

A careful revision of the cones of Mesozoic Abietinee is very 
desirable; the number of species would no doubt be considerably 
reduced. 

In the Rufford Collection there are some specimens of fairly 
well-preserved cones attached to their branches, and in addition 
to these, several isolated specimens in other Wealden collections, 
which differ in their greater length from those preserved in the 
position of growth. The larger detached cones I have referred 
to Carruthers’ species Pinites Dunkeri; the others, with their 
branches and leaves, are placed in a new species, P. Solmsi. 


Pinites Dunkeri, Carruthers, 


1853. <Abietites Dunkeri, Mantell, Geol. I. Wight, p. 452. 

1866. Pinites Dunkeri, Carruthers, Geol. Mag. vol. iii. p. 542, pl. xxi. figs. 1 
and 2. 

1867. Pinites Dunkeri, Carruthers, Journ. Bot. vol. vy. p. 14, pl. lix. figs. 1 
and 2. 

1870. <Abietites Dunkeri, Schimper, Trait. pal. vég. vol. ii. p. 307. 

1878. Pinites Dunkeri, Dixon’s Geol. Sussex, p. 279. 

1886. Pinites Dunkeri, Gardner, Rep. Brit. Assoc. p. 5. 

1889. Pinites Dunkeri, Bristow, Geol. I. Wight, p. 268. 


Type. Isolated cones. British Museum. 

The following definition is given by Carruthers for this species :— 

‘Cone elongated cylindrical; scales broad, with a rounded and 
thin apex; axis slender; seeds oval compressed.” 

The largest cone referred to this species has a length of over 
33cm. and is 83cm. broad. Cones of a similar form have been 


PINITES. 195 


described by Velenovsky' under the name of Pinus longissima, 
having a length of 3lem. and 38cm. broad. It is difficult to 
decide in many instances between P. Dunkeri and P. Carruthersi 
as the most suitable species to which to refer the specimens. 


46654. Portion of a long cone, with the scales partially ex- 
panded, showing some clearly preserved cavities from which the 
small oval seeds have fallen. Very similar to Pinites Carruthersi, 
Gard. Several specimens with this registered number containing 
iron pyrites, and very friable. Brook, I. Wight. Bowerbank Coll. 


Pinites Carruthersi, Gardner. 
[Pl. XX. Fig. 5.] 
1886. Pinites Carruthersi, Gardner, Brit. Assoc. Rep. 1886, p. 4, fig. 6. 


Type. Imperfect cone. Woodwardian Museum, Cambridge. 

Gardner speaks of the type specimen as a long cylindrical cone 
with numerous persistent, leathery, imbricated scales, tapering 
towards the base, with scales thicker than those of Pinites valdensts, 
Gard., but thin at the edges, smooth, without a keel, and with 
entire rounded margins, 

This form of cone is very similar to that represented by the 
more perfect specimens which I have included under a new species, 
Pinites Solmst. In P. Andrei, Coem.,? P. Coemansi, Heer,’ and 
other Mesozoic forms, the same type occurs; as a rule, however, 
it is impossible to determine the precise affinities of these detached 
examples. 


V. 2611, “Pie XX. Fig. 5. V. 2852. 

Cones in a crumbling condition, partly preserved in iron 
pyrites. Larger than those of V. 2146 (Pl. XIX. Fig. 1), but 
it is possible that they both belong to the same species. Eccles- 
bourne. Rufford Coll. 


V. 2266. Two smaller specimens, probably belonging to this 
species. Ecclesbourne. Rufford Coll. 


1 Velenovsky (A. 1), p. 26, pl. i. figs. 14-17. 
* Coemans (A.). 
° Saporta, Pal. France. vol. iii. p. 474, pl. exci. figs. 6 and 7. 


196 PINITES. 


Pinites Solmsi, sp. nov. 
[Pl. XVIII. Figs. 2and3. Pl. XIX.] 


Type. Cones attached to leaf-bearing branches. British Museum. 

Some of the specimens referred to this species, closely resemble 
Pinites Carruthersi, Gard., but in view of the much more perfect 
nature of the Rufford material, and the doubtful identity of 
Gardner’s type, I have ventured to found a new species. The 
specific name Solms? has been adopted as a slight record of 
Graf zu Solms-Laubach’s services to Mesozoic paleeobotany. The 
species may be thus defined :— 

Short lateral branches covered with well-marked elongated bases 
of the scale leaves, in the axil of which are borne the short shoots 
with long needle-like leaves. Cones oblong in form, with broad 
scales similar to those of the Strobus section of the recent genus 
Pinus, or those of Picea and Abies. 

In Pinites Carruthersi the scales have a similar form, but 
slightly larger, and with a more flattened thin upper border. In 
a few specimens of this species the needles are in place, but do 
not show the manner of attachment of the leaves with sufficient 
clearness, to enable us to determine how many needles are borne 
on each short shoot. It is possible, indeed, that the leaves arise 
direct from the large branches, as in Abies and Picea, but the 
form and size of the needles are much more in accordance with 
the characters of the genus Pinus. 


V. 21465) PI ie 

Portions of four unripe cones, apparently in place; possibly the 
three uppermost cones are in their natural position, and the lower 
one displaced. The clearly marked impressions of the bracts show 
their rounded outline very distinctly: cf. P. Carruthersi, Gard., 
and P. Andrei, Coem., as figured by Gardner.’ Similar to Advetites 
ellipticus, Font.,? but smaller. The surface of the cones is 
marked by a number of fine longitudinally running striations. 
The branches are covered with well-preserved decurrent leaf 
cushions. There are no leaves in their position of growth, but 
several fragments of needles occur on the rock surface. Eecles- 
bourne. Rufford Coll. 


1 Gardner, Joc. cit. fig. 1. 
2 Potomac Flora, pl. exxxiil. figs. 2-4. 


PINITES. 197 


V. 2169. Pl. XVIII. Fig. 2. 


Here again the leaf cushions are clearly preserved; at the ends 
of the short lateral branches there are borne clusters of long 
needles, but it is impossible to make out with certainty the actual 
leaf arrangement, or manner of attachment to the leaf-bearing axes. 
A few of the leaves show an acuminate apex. The general habit 
of the specimen is similar to that of Cedrus or Larix, but the 
greater length of the branches and the form of the leaves offer 
a still stronger resemblance to Pinus. If we compare young 
branches of some species of Pinus with this and other specimens 
we find a very close agreement. The portion of a cone below the 
main branch probably belongs to this species. There is some 
resemblance to Leptostrobus longifolius, Font.’ Ecclesbourne. 

Rufford Coll. 


V. 21470. Pl. XIX. Fig. 4. 


With some of the scales in this specimen there appear to be 
associated narrow and pointed structures, similar to the semin- 
iferous scales, and shorter and broader bract scales of such a 
form as Zsuga Douglasii, Sab. This appearance is, however, 
probably deceptive, and is the result of our seeing some of 
the bracts edgewise. There can be very little doubt as to the 
identity of this cone with those in V. 2146 (Pl. XIX. Fig. 1). 
Eeclesbourne. Rufford Coll. 


V. 2146a. Pl. XIX. Fig. 3. 


In this specimen we have two female cones which appear to be 
in place, and a branch continued above them, on the upper portion 
of which there appear to be the remains of imperfectly preserved 
structures, which may possibly be male cones. There is not, 
however, sufficient evidence on which to found any very definite 
statement. Ecclesbourne. Rufford Coll. 


V. 2255.72 XVITL. Fig: 3. 


Part of a somewhat smaller cone, and a cluster of needles borne 
on a short lateral branch. Probably the same species as the larger 
specimens. Kcclesbourne. Rufford Colt. 


1 Loe. cit. pl. cii. figs. 1 and 2. 


198 PINITES. 


Weoiay. Pl. XTX Mie. 2, 


Part of a cone like that of V. 2147a (Pl. XIX. Fig. 4), 
attached to a branch bearing the characteristic leaf cushions. 
The difference between this specimen and V. 2146 (Pl. XIX. 
Fig. 1) is due to the scales being open in the present example. 


V. 21475. Open cone, and portions of branches with well- 
preserved leaf cushions. Cf. Pl. XIX. Fig. 1. Eccleshourne. 
Rufford Coll. 


V. 1069. Branches with leaf cushions fairly distinct ; cones 
imperfectly preserved. Somewhat similar to the specimens de- 
scribed in Volume I. as rhizomes of Onychiopsis Mantelli (Brong.).! 
Ecclesbourne. Rufford Coll. 


V. 3167. In this specimen the leaf cushions are clearly shown, 
and the limits of annual growth are suggested by the closer 
arrangement of the cushions in certain parts of the branch. 
Keclesbourne. Rufford Coll. 


V. 3165. Very small needles, like those of V. 2255 (Pl. XVIII. 
Fig. 8). A fairly long branch with short leaf-bearing lateral 
branches. 


V. 2270. A cone in longitudinal section. 


V. 2291. Possibly a different plant, but too imperfect to deter- 
mine with any certainty. Ecclesbourne. Rufford Coll. 


Other specimens of cones and branches referred to this species: 
V. 1069a, V. 10692, V. 1069c, V. 1069¢, V. 2291, V. 3165, 
V. 3168. 


Pinites, sp. 


V. 2922. <A single winged seed. Numerous fragments of 
Onychiopsis Mantelli (Brong.) on the same piece of rock. LEccles- 
bourne. Rufford Coll. 


D Voli Lepao2:. 


SPHENOLEPIDIUM. 199 


Pinites Ruffordi, sp. nov. 


V. 2804. <A specimen of coniferous wood with the minute 
structure clearly preserved, and showing the characters of the 
genus Pinites. It is proposed to publish elsewhere a detailed 
description of the anatomy of this specimen of Pinites, under 
the name of Pinites Ruffordi. The annual rings are very clearly 
marked; resin ducts fairly numerous; the tracheids in radial 
section show either a single row of bordered pits, or a double 
row having the arrangement characteristic of the genus Pinites. 
Eeclesbourne. Rufford Coll. 


Genus SPHENOLEPIDIUM, Heer. 
[Sece. Trab. Geol. Portugal, 1881, p. 19.] 


The generic name Sphenolepis, proposed by Schenk in 1871,' was 
changed by Heer to Sphenolepidium, on account of the previous use 
of the former name by Agassiz as a genus of fishes. Heer’s new 
term is adopted by Schenk in his account of fossil Conifere con- 
tributed to Zittel’s Handbuch.” The species of this genus have been 
included by Schenk and others in the family Zazodiee, but Solms- 
Laubach® considers that the botanical nature of these fossils is too 
imperfectly known to admit of any precise localization among 
existing subdivisions of the Group Conifere. Previous writers have 
drawn attention to the resemblance of the Wealden species, Spheno- 
lepidium Kurrianum, to Athrotaxis, and Sequoia has also been 
referred to as the nearest living genus. There is nothing in the 
nature of the fossil cones of this genus, so far as I am able to judge 
from the published figures, and an examination of fairly well-pre- 
served English specimens, to stand in the way of a comparison with 
these two living genera. As regards the leaf form and arrangement, 
and the general habit of the fossil species, there is a very close 


1 Paleontographica, vol. xix. p. 243. 
2 Zittel (A.), p. 304. 
> Fossil Botany, p. 71. 


200 SPHENOLEPIDIUM. 


resemblance between Sphenolepidium Kurrianum (Dunk.) and 
species of Athrotaxis, e.g. A. laxifolia, Hook, and A. cupressoides, 
Don.1 Both Athrotaxis and Sequoia are placed by Eichler? in the 
section Pinoidee-A bietinee-Taxodiine, and it would seem highly 
probable that the Wealden form bears a close relationship to these 
recent genera, especially to Don’s genus Athrotaxis. 

We may define Sphenolepidium as follows: Branches alternate, 
with spirally disposed and decurrent leaves, cones small, oblong 
and spherical, borne on short lateral branches. 


Sphenolepidium Kurrianum (Dunk.). 
[Pl. XVII. Figs. 7 and 8; Pl. XVIII. Fig. 1.] 


1839. ? Muscites imbricatus, Romer, Verstein. Ool. Geb. p. 9, pl. xvii. fig. le. 
1846. TZhuites (Cupressites?) Kurrianus, Dunker, Wealdenbildung, p. 20, 
pl. vii. fig. 8. 
? Lycopodites, Dunker, loc. cit. p. 20, pl. viii. fig. 8. 
Thuites Germari, Dunker, loc. cit. p. 19, pl. ix. fig. 10. 
1847. Widdringtonites Kurrianus, Endlicher, Synopsis, p. 272. 
1848. Thuites Kurrianus, Bronn, Index nomencl. p. 1271. 
1849. Brachyphyllum Kurrianum, Brongniart, Tableau, p. 107. 
1849. Brachyphyllum Germari, Brongniart, loc. cit. p. 107. 
1850. Widdringtonites Kurrianus, Goppert, Foss. Conif. p. 176. 
1850. Widdringtonites Kurrianus, Unger, Gen. spec. plant. foss. p. 342. 
Thuites Germari, Unger, loc. cit. p. 348. 
1851. Widdringtonites Kurrianus, Ettingshausen, Abh. k.-k. geol. Reichs. 
vol. i. Abth. iii. No. 2, p. 25. 
W iddringtonites Haidingeri, Ettingshausen, loc. cit. p. 26. 
Araucarites Dunkeri, Ettingshausen, Joc. cit. pl. ii. fig. 10; pl. ii. fig. 1. 
1854. Thuites Kurrianus, Morris, Brit. foss. p. 24. 
1861. Widdringtonites Kurrianus, Hildebrand, Verbreit. Conif. p. 296. 
W iddringtonites Haidingeri, Hildebrand, loc. cit. p. 296. 
1870. Widdringtonites Kurrianus, Schimper, Trait. pal. vég. vol. ii. p. 329. 
1870. ? Araucarites hamatus, Trautschold, Nouv. Mém. Soc. Nat. Moscou, 
vol. Aunt poi, °pla xx. era. 
1871. Sphenolepis Kurriana, Schenk, Paleontographica, vol. xix. p. 243, 
pl. xxxvil. figs. 5-8; pl. xxxviii. fig. 1. 


1 Don. 
* Engler and Prantl (Conifere), p. 84. 


SPHENOLEPIDIUM. 201 


1875. Thuites (Cupressites) Kurrianus, Topley, Weald, p. 409. 
1881. Sphenolepidium Kurrianwn, Heer, Secc. Trab. Geol. Portugal, p. 19, 
pl. xii. fig. 14; pl. xiii. figs. 15 and 80; pl. xviil. figs. 1-8. 
1881. ? Thuites Choffati, Heer, loc. cit. p. 11, pl. x. fig. 8. 
1884. Sphenolepidium Kurrianum, Schenk in Zittel’s Handbuch, p. 304, fig. 210. 
1885. ? Sphenolepis Kurriana, Hosius and Von der Marck, Paleontographica, 
vol. xxvi. p. 216, pl. xliv. fig. 209. 
1889. Sphenolepidium Kurrianum, Fontaine, Potomac Flora, p. 260, ? pl. exxvi. 
figs. 1-6 ; pl. exxviii. figs. 1 and 7; pl. exxix. figs. 1, 2, 4, 6, and 8; 
2 pl. cxxx. fig. 11; pl. exxxi. fig. 4; pl. elxvii. fig. 2. 
? Sphenolepidium Virginicum, Fontaine, loc. cit. p. 259, pl. exxv. fig. 4, 
and pl. elxvi. fig. 6. 
? Athrotaxopsis expansa, Fontaine, loc. cit. p. 241, pl. exxxv. figs. 15, 
18, and 22. 
1894. Sphenolepidium Kurrianum, Saporta, Flor. foss. Port. p. 114, pl. xxii. 
figs. 3-5. 
1895. ?Sphenolepidium Kurrianum, Kerner, Jahrb. k.-k. geol. Reichs. 
vol. xlv. Hefti. p. 51, pl. iv. fig. 2. 


Type. Vegetative branch. ? Berlin Museum. 

Dunker thus defines the species :— 

‘‘Thuites ramulis erectis irregulariter pinnatis, compressiusculis 
utrimque subcarinatis, foliolis crassiusculis imbricatis irregulariter 
dispositis elongatis subflexuosis apice acutis dorso carinatis sub- 
distantibus.”’ 

The small fragment figured by Romer as Jfuscites imbricatus, 
Roém., is probably identical with Sphenolepidium Kurrianum 
(Dunk.); Schenk calls attention to this resemblance, but, not having 
seen the type specimen, hesitates to express any decided opinion. 
Although there is a strong probability of Romer’s specimen being 
a leafy twig of the present species, it would hardly be wise to 
enforce the rule of priority as regards the specific designation 
without more trustworthy data. The other fragment figured by 
Romer! as Muscites falcifolius, Rom., and compared by Dunker 
with Sphenolepidium Kurrianum, is too small to identify with ~ 
certainty, and does not bear such a strong resemblance to Dunker’s 
species as does M imbricatus. 

Ettinghausen’s species Widdringtonites Haidingert is no doubt 
correctly included by Schenk in the present species. The specimens 
figured by Ettingshausen as Araucarites curvifolius agree so closely 


1 Romer, F. A. (A.); Verstein. Ool. Geb. pl. xvii. fig. le. 


202 SPHENOLEPIDIUM, 


with S. Xurrianum that there cannot be much doubt as to their 
specific identity. It has already been pointed out,! that one of the 
specimens referred by Schenk to this species is no doubt a fertile 
frond of Onychiopsis Mantelli (Brong.). Some of the numerous 
fragments figured by Fontaine from the Potomac beds, as examples 
of S. Kurrianum, suggest a plant with a habit somewhat different 
to that of Dunker’s species. Without attempting to discuss the 
exact nature of all Fontaine’s fragments, it is probably safe to 
assert that the present species is represented in the Potomac 
Flora. Fontaine’s specimens are all without cones, but the 
small cones figured by him as Athrotaxopsis expansa, Font., may 
in all probability be referred to S. Aurrianum. The specimen 
figured by this author as Sequota gracilis, Heer,” bears a decided 
resemblance to the present species. The fragments of cone-bearing 
twigs figured by Fontaine as a new species, S. Virginicum, are 
compared by him with 8. Aurrianum. I have included these 
specimens in the synonomy as probably identical with the present 
species. Saporta’s Portuguese examples of the species are for 
the most part small fragments of twigs; but there can be little 
or no doubt as to their specific identity with the English plant. 
The numerous specimens figured by Heer from the Lower 
Cretaceous rocks of Greenland as Cyparassidium gracile, Heer,* 
agree so closely with S. Kwrrianum, that one feels tempted to 
regard the two as identical. Heer notes the resemblance as 
regards leaf form. and disposition between Cyparissidium, Wid- 
dringtonites, Glyptostrobus, Athrotaxis, and Sequoia; but adds that 
the form of the cones in Cyparissidium is quite distinct from 
that in the other genera, and more allied to Cunninghamia. 
Although there are some slight differences between the cones 
figured by Heer and those of Sphenolepidium, the points of 
difference do not appear to be very wide. Ettingshausen figures 
some specimens from Niederschoena under the name of Frenelites 
Reiehii, Ett.;* these bear a strong resemblance to the present 
species, and it is difficult to determine on what grounds the 


B Wolk, dl jos 625 

2 Fontaine, Potomac Flora, pl. exxvi. 

3 Heer (A. 3), Fl. foss. Arct. vol. vi. pl. i., and Fl. foss. Arct. vol. iii. (2) 
pl. xix. etc. 

4 Ettingshausen (A. 8), p. 246, pl. i. fig. 10. 


SPHENOLEPIDIUM. 203 


comparison with Frenela is made. Possibly Yokoyama’s Japanese 
species, Cyparissidium (?) Japonicum,' may be closely allied to 
S. urrianum, but the preservation of the specimens is too 
imperfect to allow of any satisfactory comparison. We may 
define the species as follows :— 

Branching alternate; leaves ovate, acuminate, or triangular ; 
keeled dorsally; cones small, borne on clusters of short slender 
branches, globose or oblong; scales broad and short, thick, with 
an elongated lozenge-shaped depression at the apex. 


V. 2318. Pl. XVII. Figs. 8 and 8a. 

This specimen appears to be practically identical with that 
figured by Schenk in his pl. xxxviii. figs. 10 and 11.* The 
oblong cone and the broad scales with the elliptical transversely 
elongated scars are well shown. The leaves of the fertile 
branches have an elongated oval form, with acute tips, and are 
closely adpressed to the stem. ‘he cone is similar to that figured 
by Heer in pl. xiv.,? but in his specimen the leaves are less 
adpressed to the branch. Compare also Fontaine’s S. Virginicum ; 
this species may, however, be identical with S. Sternbergianum. 
Ecclesbourne. Rufford Coll. 


V. Zated: (Pl; XVIIT. Fig. 1. 

This specimen shows very clearly the connection between the 
thicker and more slender branches. Compare the thicker portion 
with V. 23164. Ecclesbourne. Rufford Coll. 


V. 23162) PPG Ric. 7, 

The clustered fertile branches and small cones represent a 
characteristic feature of the species, and may be compared with 
those of such a recent plant as <Athrotaxis laxifolia. Cf. 
V. 231382, etc. 


V. 23134, V. 23160, V. 23164. Cones; some with fairly well- 


preserved scales. 


V. 2316c¢. Part of a thicker unbranched stem with clearly 
preserved leaves. This specimen, if examined without reference 


1 Yokoyama, p. 229, pl. xx. figs. 3, 6, and 13, and pl. xxiv. fig. 4. 
2 Paleeontographica, vol. xix. 
3 Sece. Trab. Geol. Port. 1881. 


204 SPHENOLEPIDIUM. 


to the more slender branches of S. Auwrrianum, would probably 
be referred to Brachyphyllum, and it serves to illustrate the great 
difficulty in attempting to discriminate between the various 
provisional genera of fossil conifers. 


V. 2316¢. Fragment of a thick axis. Cf. Brachyphyllum obesum, 
Heer. Ecclesbourne. Rufford Coll. 


V. 2303. The adpressed leaves very clearly defined. The 
smaller branches appear to be identical with the fertile branch of 
V. 2313 (Pl. XVII. Fig. 8). The long unbranched axis of this 
and other specimens suggests an open habit of branching. 


V. 2308a. Probably the same species, but the leaves are rather 
less closely adpressed to the stem. Of. Schenk, pl. xxxvii. fig. 5.’ 


V. 23030. A much branched specimen. Towards the lower 
part of the thickest branch the leaves are seen to be shorter 
and more crowded. Cf. also V. 23808, and Wiaddringtonites 
Haidingeri, Ett., which Schenk has referred to S. Kurrianum. 


V. 2303c. Long branches with the leaves less clearly preserved. 
Ecclesbourne. Rufford Coll. 


V. 2253. Portions of branches. The leaves showing longi- 
tudinal striations, as in V. 2750 (Pl. XVII. Fig. 5). Eccles- 
bourne. Rufford Coll. 


V. 2285. Probably S. Kurrianum. Ecclesbourne. Rufford Coll. 


V. 2286. Cones with open scales, probably belonging to this 
species. V. 718. Fragments of branches. Ecclesbourne. 


Rufford Coll. 


V. 2303. Slender branches with leaves rather less adpressed 
to the stem than in some examples of the species. This form 
suggests a passage to the more open leaves of S. Sternbergianum. 


Cf. V. 21894 (Pl. XVI. Fig. 5). Ecclesbourne. Rufford Coll. 


? Sphenolepidium Kurrianum. 
V. 3343. Two large specimens. Slender branches given off 
from an axis 3cm. in diameter. 


1 Schenk, Joc. cit. 


SPHENOLEPIDIUM. 205 


Sphenolepidium Sternbergianum (Dunk.). 


[Pl]. XVI. Figs. 4-6.] 


1846. DMuscites Sternbergianus, Dunker, Wealdenbildung, p. 20, pl. vii. fig. 10. 
1848. Muscites Sternbergianus, Bronn, Index nomencl. p. 759. 
1849. Jwniperites Sternbergianus, Brongniart, Tableau, p. 108. 
1850. Muscites Sternbergianus, Unger, Gen. spec. plant. foss. p. 42. 
1851. Araucarites Dunkeri, Ettingshausen, Abh. k.-k. geol. Reichs. vol. i. 
Abth. iti. No. 2, p. 27, pl. ii. figs. 2, 3, 7, and 8. 
Araucarites curvifolius, Ettingshausen, Joc. eit. p. 28, pl. ii. figs. 11, 
13.14, 172211, 
1861. Araucarites Dunkeri, Hildebrand, Verbreit. Conif. p. 276. 
Araucarites curvifolius, Hildebrand, ibid. p. 276. 
1870. Widdringtonites Dunkeri, Schimper, Trait. pal. vég. vol. ii. p. 329. 
Widdringtonites curvifolius, Schimper, ibid. p. 329. 
1871. Sphenolepis Sternbergiana, Schenk, Paleontographica, vol. xix. p. 243, 
pl. xxxvii. figs. 3 and 4; pl. xxxvii. figs. 3-13. 
1881. Sphenolepidium Sternbergianum, Heer, Secc. Trab. Geol. Portugal, p. 19, 
pl. xiii. fig. lw; pl. xiv. figs. 2-8. 
1884. Sphenolepidium Sternbergianum, Schenk in Zittel’s Handbuch, p. 304, 
fig. 210a, 0, and e. 
1885. Sphenolepis Sternbergiana, Hosius and Von der Marck, Paleeontographica, 
vol. xxvi. p. 218, pl. xliv. figs. 206—208.1 
1889. Sphenolepidiun Sternbergianum, Fontaine, Potomac Flora, p. 261, 
? pl. exxi. figs. 8, 10, and 11; pl. cxxx. fig. 9. 
1894. Sphenolepidium Sternbergianum, Saporta, Flor. foss. Port. p. 114, 
pl. xxii. figs. 1 and 2; p. 139, pl. xxvii. fig. 14; p. 193, pl. xxxiil. 
fig. 18. 


Type. Fragments of small twigs. 

Dunker gives the following definition of the species :— 

‘‘Muscites caule virgato subflexuoso, foliis bifariis imbricatis 
patentibus ovato-lanceolatis subfalcatis.”’ 

Brongniart substituted /wniperites as a more appropriate generic 
name than Jfuseites; and Ettingshausen renamed Dunker’s plant 
Araucarites Dunkert. The specimen figured in Ettingshausen’s 


1 These fragments, from the Neocomian of Toénsberg, are very small and 
imperfect ; fig. 206 is probably a fragment of S. Sternbergianum, but it would 
be unwise to make any definite statements on such slender evidence. 


206 SPHENOLEPIDIUM. 


pl. ii. fig. x. seems to be identical with what he called Widdring- 
tonites Haidingert, and which we have included, following Schenk’s 
example, as a synonym of S. Sternbergianum. Most of the speci- 
mens figured by Ettingshausen as Araucarites curvifolius (Dunk.), 
must no doubt be included in the present species, and are not 
identical with Dunker’s Lycopodites curvifolius. Some of the 
species, on the other hand, are the same as Dunker’s type, 
and do not appear to belong to Sphenolepidium Sternbergianum. 
Fontaine’s small specimens agree with this species, but it is 
impossible to discriminate with any degree of certainty between 
the numerous and very similar twigs, which he figures from the 
Potomac beds under different specific names. 

If we compare the figures of Sequoia ambigua, Heer,’ with 
Sphenolepidium Sternbergianum (Dunk.), we find a striking re- 
semblance; e¢.g., Heer, pl. xxi. fig. 3, and Schenk, pl. xvi. 
figs. 8 and 4. It is exceedingly difficult to come. to any satis- 
factory conclusion as to the specific identity of these various 
coniferous twigs; and the striking resemblance between the 
branches of certain recent forms, should sufficiently demonstrate 
the futility of attempting to carry our comparisons too far in 
the case of fossil fragments. It would seem, however, that if 
Heer’s Greenland plant is not identical with the present species, 
it is closely allied to it. 8S. Sternbergianum differs from the 
preceding species chiefly in the more open and linear leaves. 
As regards the cones of the two species, it is not quite clear how 
far Schenk’s description of a specific difference holds good; the 
material from the English beds is hardly sufficiently well preserved 
to enable us to give any satisfactory detailed diagnosis. 


V; 2311. Pl. XVI. Fig. 4. 

An imperfectly preserved specimen showing two cones attached 
to short branches. The difference in the leaf form from that of 
S. Kurrianum is clearly shown. Ecclesbourne. Rufford Coll. 


V. 2144. Pl. XVI. Fig. 6. 
Well-preserved branches with spreading and falcate leaves. 
Ecclesbourne. Ruffurd Coll. 


1 Heer, Fl. foss. Arct. vol. iii. (2) pl. xxi. etc. 


SPHENOLEPIDIUM. 207 


V.: 2139¢.) PIZXVEI. Fig: 5. 

Smaller specimen, with more crowded leaves, leaf form dis- 
tinctly preserved. Compare with the upper part of V. 2139, in 
which the leaves are less closely arranged. Very similar to the 
specimens figured by Schenk! in pl. xxxvii. figs. 3 and 4. 


V. 2139. 20cm. in length. No doubt the same form as 
Muscites Sternbergianus, Dunk. The thicker branch well pre- 
served, showing clearly defined leaves, with the distal end free, 
and the broader basal portion adpressed to the axis. Several 
delicate branches shown at the upper end. Cf. Schenk,’ pl. 


9 


xxxviii. fig. 8. Ecclesbourne. Rufford Coll. 


V. 2141. Probably the same species. Cf. V. 2144, also 
Ettingshausen’s figures of Araucarites curvifolius,? some of which 


Schenk refers to S. Sternbergianum, LEcclesbourne. Rufford Coll. 


V. 2289, V. 22894. Larger examples with imperfect fragments 
of cones. 


V. 2289c. Imperfectly preserved, probably specifically identical 
with the above specimens. Compare the finer branches with 
Dunker’s Lycopodites, pl. vii. fig. 8.3 


V. 2289¢d. Imperfect fragment showing the leaves in side and 
surface view. Ecclesbourne. Rufford Coll. 


V. 2290, V. 2315. Small curved and closely crowded leaves, 
probably identical with the form represented in Ettingshausen’s 
figures of Araucarites Dunkert. Cf. also V. 2189a, ete. Possibly 
this form should be regarded as a distinct species, but the evidence 
is hardly sufficient to justify a separation from S. Sternbergianum. 
Ecclesbourne. Rufford Coll. 


V. 2811a. Cone; fairly well preserved. 


V. 2919. The leaves show very clearly the single median vein. 
Ecclesbourne. Rufford Coll. 


E Loc cit. 
2 Ettingshausen (A. 4). 
3 Dunker, Wealdenbildung. 


208 SPHENOLEPIDIUM. 


Cf. Sphenolepidium (Sequoia) subulatum (Heer). 
[Pl. XVI.. Fig. 3.] 


There are a few specimens in the Rufford Collection which 
differ from the typical S. Sternbergianum (Dunk.), in having more 
closely arranged, longer and narrower leaves; they agree very 
well with Heer’s Greenland species Sequoia subulata,! and are 
probably identical with this form, and with the specimens described 
by Saporta from Portugal as Sequoia subulata var. Lusitanica.’ 
Saporta’s variety appears to be hardly justified by the very small 
differences which he notes between the Portuguese and Arctic 
forms. There would seem to be no satisfactory evidence to 
warrant the use of the generic name of Sequoia. 

Ettingshausen’s fig. 18, pl. ii.2 [Araucarites curvifolius (Dunk.) | 
bears a strong likeness to the specimen represented in our Pl. XVI. 
Fig. 3 (V. 2140). It is not improbable that some of the specimens 
figured by Fontaine as Sequoia Reichenbachii, Heer,* are identical 
with the present species. Possibly the following specimens might 
reasonably be referred to as constituting a variety of Spheno- 
lepidium Sternbergianum. 


VW. 2140. ° Pl XVI. Fig. 3. 

Branches fairly closely set. Leaves narrow, linear and keeled, 
slightly curved or almost straight. At first sight the branches 
appear to have a bilateral form as regards the leaf arrangement, 
but this may be simply due to manner of preservation. ccles- 
bourne. Rufford Coll. 

V. 20938. Broken fragments. Leaves fairly distinct. Cf. S. 
Sternbergianum and Ettingshausen’s pl. i. fig. 18. Ecclesbourne. 

Rufford Coll. 


V. 1069. Smaller specimen. LEcclesbourne. Rufford Coll. 


Sphenolepidium, sp. 
V. 2281, V. 2283. Specimens of cones which cannot be referred 


with any certainty to a particular species of the genus. 


1 Heer, FI. foss. Arct. vol. iii. (2) p. 102, pls. xxvii.-xxix. 
2 Saporta (1), p. 177, pl. xxxiii. figs. 7-12. 
% Ettingshausen, Joc. cit. 

4 Fontaine, Potomac Flora, pl. exviil. ete. 


THUITES. 209 


Genus THUITES, Brongniart. 


[Tableau, 1849, p. 71.] 


Thuites valdensis, sp. nov. 
[Pl. XX. Fig. 6.] 


Type. Single specimen of a leafy twig. British Museum. 

Although it is impossible to give a complete or entirely satis- 
factory definition of this species from the single specimen in the 
Rufford Collection, it may be convenient to adopt a new specific 
term, as there seems to be little doubt that we have to do 
with fragments of a conifer different from any known species. 
The characters may be summarized as follows :— 

Branching opposite, leaves in whorls, adpressed to the stem, 
two in each whorl, keeled dorsally and with comparatively 
blunt apices. 

Some of the specimens referred by Ettingshausen to Frenelopsis 
Hoheneggert (Ett.), and, in a less degree, a few of those figured 
by Schenk under Ettingshausen’s specific name, show a distinct 
resemblance to the present specimen. 


V. 2188. Pl. XX. Fig. 6. 

Preservation good. The leaves and opposite branching are 
clearly seen. Cf. Mrenelopsis ramosissima, Font., Potomac Flora, 
pl. xcv.,! etc., also F. Hoheneggert (Ett.).2 In describing the 
specimens subsequently referred by Schenk to the genus Frene- 
lopsis, Ettingshausen adopts the generic name TZhuites in the 
wide sense, as including various forms of Cupressinee ; it would 
probably be wiser to retain Zhuites for such examples as those 
figured by Ettingshausen. In instituting the species 7. Hohen- 
eggert, Ettingshausen speaks of the branching as alternate, and 
the leaves as ‘‘quadrifariam imbricatis’”; but the specimen 
shown in his pl. i. fig. 7 has clearly only two leaves in some 
of the verticils; indeed, there is a striking similarity between 
this German specimen and the single example from the English 


1 Fontaine (A. 2), Potomac Flora. 
2 Ettingshausen (A. 4), Abh. k.-k. geol. Reichs. vol. i. Abth. iii. 1852, pl. i. 
figs. 6 and 7. 


P 


210 NAGEIOPSIS. 


beds. Turning to Schenk’s description, we find that he notes 
the presence of rows of small tubercles on the branches as an 
additional character, and describes the leaves as decussate and 
opposite, and not four in each whorl as stated by Ettingshausen. 
It may be that under Frenelopsis Hoheneggert (Ett.) we have 
more than one species; the younger fragments, such as those 
figured by Ettingshausen, and a few of those described by Schenk, 
agree very closely with the English specimen of TZhuites, but 
the larger branches of Schenk may perhaps belong to another 
plant. Ecclesbourne. Rufford Coll. 


Genus NAGEIOPSIS, Fontaine. 


[Potomac Flora, 1889, p. 194.] 


Fontaine proposes this generic title for one of the most largely 
developed and characteristic Potomac plants. He compares the 
leaves with those of Podozamites, but is enabled by the large and 
numerous specimens at his disposal to recognize distinct coniferous 
features. He defines the genus as follows :— 

‘‘Trees or shrubs with leaves and branches spreading in one 
plane; leaves varying much in size and shape, those towards 
the base of the twigs sometimes smaller than those higher up, 
distichous mostly, or rarely subdistichous, opposite and persistent, 
attached by a short, slightly twisted foot-stalk, usually to the 
side of the twig, more rarely slightly within the margin on 
the upper or under surface of the stem, either attenuated towards 
the base or abruptly rounded off there, at their ends acute or 
sub-acute; nerves several, coalescing at base to form a foot-stalk, 
forking immediately at the base or a short distance above, then 
approximately parailel to near the tips of the leaves, where 
they are somewhat crowded together, but do not converge to a 
union, ending in or near the extremity.” 

The genus Podocarpus is divided by Eichler’ into four sections, 
of which section i. is MWageva, formerly regarded as a distinct 
genus.2 In this form of Podocarpus the leaves have numerous 
veins, and not a single midrib as in other species of the same 
genus. 


Engler and Prantl, p. 104. 


i Tahies 
2 —. G. Gordon, The Pinetum, p. 135 (1858). 


PAGIOPHYLLUM. 211 


Nageiopsis ¢f. N. heterophylla, Font.’ 
[Pl. XII. Fig. 3.] 


The few small fragments from the English beds bear a strong 
resemblance to the specimen figured by Fontaine as Magevopses 
heterophylla, and most probably belong to this species. 


V. 3190. Pl. XII. Fig. 3. 


Compare Fontaine, pl. lxxxvi. figs. 6 and 7, etc. There are 
several equal veins in each leaf which converge somewhat towards 
the apex; the leaves are gradually tapered distally, and towards 
the point of attachment. The actual manner of attachment to 
the branch is not clearly shown. Near Hastings. Rufford Coll. 


V. 2123, V. 2362. 
Probably the same species, but much more imperfect. Kccles- 
bourne. Rufford Coll. 


Conifers incertz sedis. 


Genus PAGIOPHYLLUM, Heer. 
[Sece. Trab. Geol. Portugal, 1881, p. 11.] 


Saporta? includes in the tribe Araucarinee two genera, Pachy- 
phyllum and Araucaria; the former representing the extinct types, 
the latter the living species. Pachyphyllum was first instituted 
by Pomel? as a section of his genus Dforeania, including I. brevi- 
folia, Pom., as the typical species. . 

Saporta figures some examples of the genus in which portions 
of cones are preserved, and is thus able to give a fairly detailed 
diagnosis. He places Pachyphyllum close to Araucaria (Eutacta), 
Agathis, and Cunninghamia. The general appearance of the 


1 Potomac Flora, p. 201, pls. Ixxxiv.-lxxxvi. 
2 Pal. Franc. vol. iii. p. 372. 
5: Pomel; p:.21. 


D2 PAGIOPHYLLUM. 


branches referred to this genus suggests an araucarian habit, 
and there is a decided probability that we may consider the fossil 
forms as closely allied to the recent genus. As regards the 
English specimens, in the absence of fossil cones we have no 
very satisfactory evidence as to the relationship to modern forms. 
Heer substituted Pagtophyllum for Pachyphyllum, on the ground 
that the latter name had already been assigned to a genus of 
orchids. The following concise definition is given in Zittel’s 
Handbuch: : ‘‘Leaves spirally arranged, leathery, thick, triangular, 
lanceolate ; spreading or closely imbricate; decurrent at the base.” 
Solms-Laubach? justly considers Pagiophyllum a purely artificial 
and provisional genus. 


Pagiophyllum crassifolium (Schenk). 
[Pl. XVI. Figs. 1 and 2.] 


1871. Pachyphyllum crassifolium, Schenk, Paleontographica, vol. xix. p. 240, 
pl. xl. fig. 8. 

1874. Pachyphyllum erassifolium, Schimper, Trait. pal. vég. vol. il. p. 570. 

1834. ? Pachyphyllum crassifolium, Saporta, Pal. Frang. vol. ii. p. 609, 
pl. cexxvi. fig. 1. 

1884. Pagiophyllum crassifolium, Schenk in Zittel’s Handbuch, p. 276. 


Type. Small and imperfect fragment of a branch. Gottingen 
Museum. 

Schenk defines the species as follows :— 

‘‘Folia in ramulo spiraliter disposita, trigona crassa conica 
falcata basi sessilia decurrentia.”’ 

The specimens referred by Saporta to Schenk’s species were 
obtained from some limestone rocks of Upper Jurassic age, in the 
neighbourhood of Grenoble; it is by no means certain that they 
are specifically identical with the Wealden type. 

We may slightly extend Schenk’s diagnosis :— 

Leaves spirally arranged; sessile, with broad base, triangular, 
somewhat falcate, keeled on the dorsal surface; the leaf lamina 
marked by fine parallel lines. Branching alternate. 


1p. 275. 
* Fossil Botany, p. 77. 


PAGIOPHYLLUM. 213 


Vi2808. “BIO kvl Bie. 1. 

Well-preserved specimen; the thick, falcate, and keeled leaves 
are closely arranged on the branches, reminding one of Cryptomeria 
Japonica. This form is very similar to the smaller specimens, 
compared with Lycopodites curvifolius, Dunk., and referred to 
Sphenolepidium Sternbergianum. Ecclesbourne. Rufford Coll. 


V. 21420. Pl. XVI. Fig. 2. 


The left-hand fragment agrees closely with Sphenolepidium 
Sternbergianum ; the leaves seen edgewise appear to be narrow, 
and show traces of a median keel. Cf. V. 2289. 

Towards the upper end of the large branch the tips of the 
falcate leaves are distinctly preserved; in the lower part they are 
seen in side view, and present a broader triangular appearance. 


V. 2142 and V. 21424. Here the leaves appear to be narrower, 
but this is largely due to the fact that they are seen edgewise, 
and not so directly as in V. 2747. KEcclesbourne. Rufford Coll. 


V. 2747. Cf. V. 21420 (Pl. XVI. Fig. 2). The flattened leaves 
seen in this view do not show the falcate form so distinctly in 
other specimens. LEcclesbourne. Rufford Coll. 


Pagiophyllum, sp. 
PPE xB’ 3!) 
V. 2288. Pl. XX. Fig. 3. 


In this specimen the leaves are fairly well preserved; they 
appear to be broader than those of the specimens referred to 
P. crassifolium. The fragment is, however, too small and im- 
perfect to admit of more exact determination. Ecclesbourne. 


Rufford Coll. 


V. 2148, V. 2317, V. 2931. 

These fragments may possibly be portions of branches of the 
preceding species, but they are too fragmentary to determine with 
any accuracy. 


214 BRACHYPHYLLUM. 


Genus BRACHYPHYLLUM, Brongniart. 
[Tableau, 1849, p. 69.] 


Brongniart proposed this name for conifers with alternate leaves 
disposed in a spiral, short, fleshy, and inserted by a broad and 
rhomboidal base. Schimper! extends this definition, and speaks 
of the genus as differing in its characters from all living forms ; 
he points out the striking resemblance between old branches of 
Brachyphyllum, with the leaves in the form of hexagonal or 
pentagonal cushions, and certain strobili of cycads and conifers. 
The surface features of a branch from which the leaves have 
fallen resemble those of Lepédodendron. Saporta® further extends 
Brongniart’s definition; and remarks that probably no genus of 
conifers has given rise to more confusion and uncertainty than 
the present genus. It has been compared with several recent 
genera, but we cannot regard Brachyphyllum, with its numerous 
species from various geological horizons, as more than a purely 
provisional genus, the actual botanical position of which is very 
uncertain; probably more than one family of Conifere being 
represented by the forms referred to under this generic name. 
Schenk* has drawn attention to the too comprehensive nature of 
the genus as used by Saporta, Heer, and others, but does not 
suggest any more precise definition of the generic characters. 

As Fontaine remarks, there is a striking resemblance between 
some forms of Evhinostrobus, Brachyphyllum, and Paleocyparis. 
Saporta, in describing the characteristics of Echinostrobus, points 
out that it differs from Brachyphyllum in having the leaves less 
thick, more pointed, and less completely adnate to the stem. We 
may compare both of these fossil genera with certain species of 
the recent genus Athrotavis. The thick fleshy leaves, with their 
broad rhomboidal bases and spiral arrangement, constitute the 
leading features of this artificial genus of fossil conifers. 


1 Trait. pal. vég. vol. ii. p. 334, 
2 Pal. Franc. vol. iii. p. 310. 
3 Zittel’s Handbuch, p. 301. 


| 
; 


Or 


BRACHYPHYLLUM. De) 


Brachyphyllum spinosum, sp. nov. 
[Pl. XVII. Figs. 1-6.] 


Type. Large specimens from LEeclesbourne, near Hastings. 
British Museum. 

This specific term is proposed as a convenient designation for 
what is probably a new species of Brachyphyllum. It is possible 
that some of the Wealden specimens previously recorded from 
other localities, and referred to this genus, may be fragments of 
the present species; but of this there is no proof. Seeing that 
no fossils have been, so far, described in which the characters of 
the large branches in the Rufford Collection are represented, we 
must institute a new term. The species may be defined as 
follows :— 

Leaves fleshy, with a median keel on the convex surface; the 
surface may be finely striated; leaf-scars rhomboidal, contiguous, 
and spirally arranged. Some of the short lateral branches have 
a pointed thorn-like form, and are clothed with the characteristic 
fleshy leaves with pointed apices. Two or three of the spiny 
branches arise approximately at the same level from the parent 
axis. 

The short, stiff, leaf-bearing thorns or pointed branches constitute 
the most prominent feature of the species. One may compare the 
thicker branches, with their large rhomboidal leaf bases, with small 
cycadean stems or twigs of Lepidodendron. In the absence of any 
reproductive organs, it is impossible to assign the species to any 
definite position among the Conifere. It is no easy task to 
determine the connection, if any, between such large specimens 
as that represented in Pl. XVII. Fig. 1, and the numerous 
smaller portions of branches. The specimen shown in Pl. XVII. 
Fig. 6 (V. 2135)! is regarded as specifically identical with PI. 
XVII. Fig. 1 (V. 2746), the striking difference being probably 
due to the destruction of the cortical tissues in the former case. 
The small piece of branch represented in Fig. 5 (V. 2750), shows 
very clearly the woody axis detached in the lower part of the 
specimen from the cortical tissues and leaves. 


1 Represented in the figure one-third natural size. 


216 BRACHYPHYLLUM. 


This form of Brachyphyllum does not appear to have been 
hitherto recorded from rocks of Upper Jurassic or Lower Cretaceous 
age, unless we may refer to this species some of the small twigs 
described by Saporta, Heer, Fontaine, and others from Wealden 
strata. The spinous branches and general habit of B. spinosum 
distinguish it from previously figured examples with similar leaves 
and leaf cushions. Velenoysky! figures some specimens from 
the Perucer beds of Bohemia under the name of Echinostrobus 
squamosus, Vel.: these have similar leaves to those of B. spinosum ; 
but coniferous branches with this form of leaf are too abundant in 
Jurassic and Cretaceous rocks to admit of any strict comparison. 
Brachyphyllum crassicaule, Font.,? B. obesum, Heer,*? and others 
may be referred to as similar in leaf form to the present species. 
Among the numerous conifers figured by Saporta from Jurassic 
strata, we have such species as B. nepos, Sap.,* and B. Desnoyersii§ 
(Brong.), and other forms which resemble the Wealden species in 
a greater or less degree. We may compare also Pagiophyllum 
cirtnicum,® Sap. Possibly the latter genus might be a suitable 
designation for the English specimens, but the distinction between 
Brachyphyllum and Pagiophyllum is in many cases by no means 
well marked, and neither term is more than a convenient generic 
name which does not imply any precise botanical affinity. 


VO 27/46," PUOXVI. Fie. 1, 

The main axis of this large specimen has a breadth of 2cm., 
and is covered with polygonal areas representing the impressions 
of large scale leaves, very similar to those in thicker branches of 
species of Athrotaxis, Thujopsis, etc., among recent genera. These 
are the remains of short and broad leaves preserved in the form 
of carbonaceous impressions on the matrix, immediately in contact 
with the thick axis. The two lowest branches are 4em. apart; 
their surface markings are identical with those on the main branch; 
from each of the lateral branches there are given off short tapered 
stiff branches covered with similar leaf impressions. Each of 


1 (A. 1) Gym. bohm. Kreid. p. 16, pl. vi. figs. 3, 6, 7, 8. 

* Potomac Flora, p. 221, pl. 100, fig. 4, pl. cix. etc. 

° Heer (A. 6), Secc. Trab. Geol. Portugal, 1881, p. 20, pl. xvii. figs. 1-4. 
* Pal. Frang. vol. iii. p. 356, pl. clxviii. ete. 

> Ibid. p. 331, pl. elsiii. ete. 

§ Ibid. p. 402, pl. clxxx. ete. 


BRACHYPHYLLUM. alot 


these small branches ends in a distinct spinous apex, and in the 
axils of some of them are seen the indistinct impressions of smaller 
leafy twigs. A still larger specimen, very similar to the above, 
is in the possession of Mr. Rufford; it shows two branches, the 
longest of which has a length of 42cm., and gives off numerous 
stiff, pointed branches like those shown in V. 2746. These 
spinous branches appear to be given off approximately at the 
same level, and probably there were three in each pseudo-whorl. 
Some of the large leaves in these specimens show fine longitudinal 
striations. Ecclesbourne. Rufford Coll. 


Waziso.’ Pl XVII. Fig. 6. . (4 nat. size.) 

This long and narrow branched axis I have referred to Brachy- 
phyllum spinosum, on the grounds that it represents a decorticated 
specimen of the same plant which is more perfectly preserved in 
V. 2746 (Pl. XVII. Fig. 1). The breadth is fairly uniform, about 
5-6 mm.; the numerous short spinous branches are clearly marked, 
and appear to have been given off in groups of twos or threes ; one 
sees in some cases two branches lying lengthwise in the sandstone, 
and the base of a third in the form of a round scar in the sub- 
stance of the main branch. There are no signs of any leaf bases 
or leaves in this example, the exposed surface of which probably 
represents the face of the woody axis. In specimen V. 2750 
(Pl. XVII. Fig. 5) we have a good example of the marked 
difference in breadth and surface characters between the leafy 
branch and the decorticated woody axis. The thorn-like processes 
in this specimen are regarded as the decorticated spinous branches 


of V.,2746 (Pl. XVII. Fig. 1). Ecclesbourne. Rufford Coll. 


V. 2240. Pl. XVII. Fig. 4. (A portion of the specimen 
shown in the figure.) 

At the upper end of the specimen we have what appears to be. 
a pith cast, surrounded by a woody cylinder, in the form of a 
dusty substance, representing the remains of wood tissue. The 
figured portion shows two spinous branches and the base of a 
third; probably there may have been four such branches in each 
pseudo-whorl. It may be noted that the spinous appendages in 
this specimen and in V. 2135 (Pl. XVII. Fig. 6), are more nearly 
at right angles to the larger axis than in V. 2746 (Pl. XVII. 
Fig. 1). Ecclesbourne. Rufford Coll. 


218 BRACHYPHYLLUM. 


V. 3180. Pl. XVII. Fig. 3. 

This thicker branch resembles a small cycadean axis with well- 
marked bases of petioles. Probably it is a portion of B. spinosum. 
At the edges of the cast there are here and there the impressions 
of the narrow distal ends of scale leaves; also at one place there 
is shown the point of attachment of a lateral branch. Length 
15cm.; diameter 1°5cm. LEcclesbourne. Rufford Coll. 


V. 2750. Pl. XVII. Figs. 5 and 5a. 

A small piece of a branch, showing elearly preserved leaves 
with longitudinal striations (5a). At each end there projects the 
impression of the woody axis, or possibly of a large pith-cavity, 
such as one finds in the genus Araucaria. It is difficult, not to 
say impossible, to distinguish between fragments of this species 
and those of B. obesum. 


V. 27460. Axis 13cm. long, showing the clearly defined out- 
lines of short and broad leaves. Immediately above and below 
the point of attachment of a spinous branch, there are impressions 
of much more slender leafy twigs. Ecclesbourne.  Luffurd Coll. 


? Brachyphyllum spinosum. 


Vie2206) Ob xy igw2: 

The broad leaves of this specimen show very clearly the fine 
striations similar to those already noted, and identical with the 
surface characters represented by Saporta in species of Pagio- 
phylium and Brachyphyllum. 


V. 2750a, V. 2751. Fragments. Ecclesbourne. Rufford Coll. 


Brachyphyllum obesum, Heer. 


1881. Brachyphyllum obesum, Heer, Secc. Trab. Geol. Portugal, p. 20, pl. xvii. 
figs. 1-4. 
1889. ? Brachyphyllum ecrassicaule, Fontaine, Potomac Flora, p. 221, pl. 100, 
fig. 4; pl. cix. figs. 1-7. 
1894. Brachyphyllum obesum, Saporta, Flor. foss. Port. pp. 112, 138, pl. xxi. 
figs. 1-7; pl. xxvii. figs. 7 and 8. 
Brachyphyllum obesiforme, Saporta, loc. cit. p. 176, pl. xxxi. figs. 12 
and 13. 
Brachyphyllum obesiforme var. elongatum, Saporta, loc. cit. p. 176, 
pl. xxxi. fig. 14. 


BRACHYPHYLLUM. 219 


Type. Portions of vegetative branches. ; 

Heer instituted this species from some fragments of coniferous 
branches from the Lower Cretaceous rocks of Almargem, Portugal ; 
and defined it as follows :— 

‘‘Br. ramis alternis, ramulis numerosis, aggregatis, crassis, 
brevibus, apice obtusis, foliis rhombeis, dense imbricatis, dorso 
leviter striatis.” 

In his recent monograph on the Portuguese flora, Saporta 
proposes a new specific name, obesiforme, for some examples of 
Brachyphyllum, which he considers may be distinguished from 
Heer’s species by their more slender branches, which are less thick- 
set, more elongated, and subdivided. He speaks of the difference 
between the two forms as slight, and admits that one may be 
merely a variety of the other. If we compare Saporta’s figures with 
those of B. obesum given by Heer, it must be admitted that the 
grounds for a specific distinction are extremely slight, and we may 
not unreasonably regard the two sets of specimens as specifically 
identical. Another specimen figured by Saporta is spoken of as 
B. obesiforme var. elongatum; this, again, appears to be too closely 
allied to such examples of B. obesiforme as are figured on pl. xxxi. 
figs. 12 and 18 to be entitled to a separate designation. The new 
specific term instituted by Fontaine for some Potomac specimens is 
perhaps a somewhat unnecessary addition to specific nomenclature ; 
the author of the species does not, apparently, draw attention to 
the very close resemblance between the fossils he describes as 
B. crassicaule and those already figured by Heer and Saporta as 
B. obesum. The figures in Fontaine’s pl. cix. may include more 
than one specific form, but the evidence is too meagre to admit 
of exact determination; figs. 4 and 5 resemble the form named 
by Saporta B. confusum,' but the difference between this species 
and B. obesum is very small. The agreement between Fontaine’s 
pl. cix. fig. 1 (B. crassicaule) and Heer’s pl. xvii. fig. 2, is very 
striking: compare also Fontaine, pl. cix. fig. 2, Heer, pl. xvii. 
fig. 3, and Saporta, pl. xxxi. fig. 12. Fontaine’s species, B. par- 
ceramosum,” comes very near to the specimen figured by Saporta as 
B. obesiforme var. elongatum. 


1 Flor. foss. Portugal, p. 112, pl. xxi. fig. 8. 
2 Potomac Flora, pl. ex. fig. 4. 


220 BRACHYPHYLLUM. 


I believe we cannot sufficiently discriminate between the several 
specimens figured by the above authors, and the similar fragments 
from the English Wealden strata, to arrive at any trustworthy 
specific distinctions or diagnoses of well-marked specific types. 
To group together all these forms may be unwise, and it has 
already been pointed out that we find certain points of difference 
between some of the specimens, which suggest either specific 
distinctions or varieties of the same type. I venture, therefore, 
to make use of Heer’s specific name B. obesum in a somewhat 
more comprehensive sense than has been adopted by Saporta. 

Among recent conifers, e.g. Cupressus Lawsoniana, Parl., etc., 
we find a considerable difference in the appearance of the branches 
depending on the development of numerous or few lateral branches 
in the axils of the leaves; we have the closely set lateral branches 
as in some forms of B. crassicaule, as figured by Fontaine, and 
the more elongated branches without the closely set lateral shoots, 
as in B. obesiforme var. elongatum, Sap. To unite such forms under 
one name, especially in the absence of cone-bearing branches, can 
hardly be regarded as an unwarranted extension of the limits of 
a fossil species of which our accurate knowledge is extremely 
small. The Jurassic species Brachyphyllum gracile, Brong.,! appears 
almost identical with some forms of . obeswm; such comparisons 
might, however, be considerably increased, but without leading 
to any satisfactory conclusions. 

Many of the specimens referred to B. obesum agree very closely 
with B. spinosum, sp. nov., and it is, I believe, almost impossible 
to feel much confidence in our attempts to distinguish between 
small specimens of plants of this particular form. Possibly it 
would have been better to make use of Fontaine’s specific term 
erassicaule for some of the following examples, and to have 
included others under B. obesum; but if we examine such a series 
as is represented by the following specimens, the difficulty of 
accurate determination becomes apparent: V. 8348 (Pl. XVII. 
Fig. 9), V. 2137 (Pl. XX. Fig. 1), V. 2137a (Pl. XX. Fig. 2), 
V. 2337 (Pl. XX. Fig. 4). 


V. 2137. Pl. XX, Bigs 1. 
Cf. Brachyphyllum obesiforme var. elongatum, Saporta, pl. xxxi. 


1 Saporta, Pal. Frang. vol. iv. p. 365, pl. clxviii., clxx., and clxxi. 


BRACHYPHYLLUM. 221 


fig. 14; also Fontaine’s Athrotaropsis expansa, pl. exiii. figs. 5 and 6, 
and A. grandis, Font., pl. exvi. figs. 1-4. This specimen does not 
show the leaf form very clearly, but the manner of branching is 
well illustrated; similar to the smaller example (V. 3348) figured 
in Pl. XVII. Fig. 9. Ecclesbourne. Rufford Coll. 


V213ia. PIO XX. Fig. 2. 

Leaves distinct ; the habit agrees with that of the smaller speci- 
men (V. 3848) shown in Pl]. XVII. Fig. 9. Cf. Saporta, pl. xxxi. 
fig. 14, and Fontaine, pl. cexi. fig. 7. 


V. 3348. Pl. XVII. Fig. 9. 

Leaves very indistinct. Cf. Brachyphyllum obesum, Heer, 
pl. xvii. fig. 2, and Saporta, pl. xxxiv. fig. 8; also B. crassicaule, 
Font., pl. exii. fig. 6. Ecclesbourne. Rufford Coll. 


WV: 2837. Pl. XX. Fig. 4. 
Leaves well shown, with faint indications of longitudinal 
striations. Similar to V. 2816a, etc. Ecclesbourne. Rufford Coll. 


V. 23162. Leaves agree closely with those of Sphenolepidium 
Kurrianum (Dunk.), but the branching is apparently different. 
Cf. V. 2316¢ (S. Kurrianum). Ecclesbourne. Rufford Coll. 


V. 21387a. Cf. Saporta, pl. xxxi. fig. 14, and Fontaine’s B. 
crassicaule, pl. exi. fig. 7. Ecclesbourne. Rufford Coll. 


V. 22920. Cf. Saporta, pl. xxxiv. fig. 8. Branches in this 
specimen closer together and thicker. Similar habit to V. 2292. 
Ecclesbourne. Rufford Coll. 


V. 2307. Similar to V. 23160, etc. Branch 18cm. long; 
imperfectly preserved. 


V. 3312. Broad axis with two small branches; similar habit to 
that of V. 2292a. Cf. Fontaine, pl. cxi. fig. 7. Ecclesbourne. 
Rufford Coll. 


Other specimens of branches or leaves referred to this 
species: V. 2257, V. 2303, V. 23032, V. 2310, V. 2883, 
V. 3188, V. 3192. Ecclesbourne. Rufford Coll. 


222 CONITES. 


Genus CONITES, Sternberg. 


On page 113 of the present volume I have suggested the revival 
of Sternberg’s genus Conites for cones of doubtful position, and 
have included under this generic name two species previously 
described by Carruthers as examples of cycadean strobili. From 
an examination of additional specimens in the Rufford Collection, 
it would seem that the species Conites elegans should be 
referred to the genus Araucarites, as being, in all probability, 
a female araucarian cone. Owing to the different manner of 
preservation of the Rufford specimens, it would, perhaps, be 
somewhat rash to speak of them as specifically identical with 
Carruthers’ cones from the Isle of Wight, but there is a strong 
likelihood of their close relationship, if not specific identity. 
Possibly the institution of a new species of Araucarites would 
be the most convenient course to adopt in dealing with these 
new Wealden specimens; but the various strobili hitherto described 
as species of Cycadeostrobus, Araucarites, and Kaidocarpon require 
careful revision, and I prefer, therefore, to refrain from adding 
a new specific name until the several forms have been more 
thoroughly examined. 


Conites armatus, sp. nov. 
[Pl. IX. Fig. 7.] 


Type. Imperfectly preserved impression of flattened cone. 
British Museum. 

The single specimen to which I have assigned the above specific 
name is too imperfect to admit of any complete diagnosis, but 
the very distinctly marked and characteristic spinous processes 
render it convenient to have some descriptive designation, even 
in the absence of those more important characters on which a 
satisfactory specific definition could be founded. 


V. 23388. Details very obscure; the long recurved spines are 
in all probability the apical prolongation of the cone scales, 
similar to those in the cones of such recent species as Araucaria 
Bidwilli, Hook., A. Cooki, Brown, Pinus Coulteri, Don, P. 
Sabiniana, Dougl., ete. Ecclesbourne. Rufford Coll. 


CONIFEROUS WOOD. 223 


CONIFEROUS WOOD. 


The occurrence of coniferous wood of Wealden age has long 
been known in the case of the so-called Pine-raft of Brook Point, 
in the Isle of Wight. This seems to have been first observed by 
Webster in 1811,! and was afterwards described by Mantell in 
1846; the latter writer compares the numerous coniferous trunks 
and associated fossils, with the rafts of drifted trees carried 
down by the waters of the Mississippi. In addition to the 
fossil wood, with its tissues more or less perfectly preserved in 
carbonate of lime, there are numerous deposits of lignite at various 
horizons in the Wealden strata, in which the lgnitic material 
obviously consists of the wood of coniferous trees. In the Medals 
of Creation, Mantell writes:* ‘‘In the Wealden deposits of Sussex, 
Kent, and Surrey, I have not observed a single fragment of 
coniferous wood.”’ More recently, in Dixon’s Geology of Sussex, 
we find that the occurrence of wood similar to that of the recent 
genus Pinus is recorded, both in the form of brittle jet and as 
mineralized fossil wood.‘ 

In the British Museum Collection, there are several good 
specimens of lignite in which the characters of coniferous wood 
are clearly seen, and numerous examples of wood with the tissues 
for the most part imperfectly preserved. 

By far the most perfectly preserved specimen of coniferous wood 
is that previously mentioned as Pinites Ruffordi, sp. nov., and 
which I hope to describe in detail elsewhere. In addition to 
this, the following specimens may be mentioned :— 


V. 701. Specimens of lignite, or perhaps more accurately de- 
scribed as jet; the annual rings clearly seen at one end of the 


large block. Hastings. Dawson Coll. 


V. 704. Lignite. Hastings. Dawson Coll. 


1 Bristow (A), Geol. I. Wight, pp. 6 and 262. 
2 Mantell (2), p. 92. 

3 Tbid. (1), vol. i. p. 168. 

4 Dixon (A:), p. 279. 


224 CONIFEROUS WOOD. 


V. 706. A fairly large piece of wood with patches of resinous 
material or amber traversing the mass longitudinally; some of 
these suggest the presence in the wood of large groups of 
parenchymatous tissue, such as Conwentz has described as 
‘‘abnormes Holz-parenchym”’ in the case of Pinus succinifera 
(Gopp.)! from the North German amber beds. The microscopic 
structure is very imperfectly preserved. Kcclesbourne. 

Rufford Coll. 

V. 707. Wood partially converted into lignite. Ecclesbourne. 

Dawson Coll. 

V. 713 (Dawson Coll.), V. 2283, V. 2237. 

Specimens of wood showing little or no internal structure. 


Ecclesbourne. Rufford Coll. 


V. 2247 and V. 2247a. In the latter specimen the structure 
is fairly well preserved; annual rings are distinct, but less so than 


in V. 2804 (Pinites Ruffordi). 


V. 2326. A specimen of wood which has undergone com- 
paratively little alteration. Ecclesbourne. Rufford Coll. 


38374. Small piece of lignite. Fairlight. Manteli Coll. 


(Spirangium Jugleri (Ett.). In the Rufford Collection 
there are several exceedingly well-preserved specimens of this 
fossil which merit careful examination; but if we accept the view 
that they are the eggs of fishes and not plant structures, this is 
not the place for any descriptive account of them. For informa- 
tion as to the nature of Spirangium reference may be made to 
the following sources :—Ettingshausen, Ueber Pal@obromelia, em 
neues Fossiles Pflanzengeschlecht, Abh. k.-k. geol Reichs., vol. 1. 
Abth. 1. p. 1; Schenk, Paleontographica, vol. xix. 1871, p. 247; 
Schenk, Die fossilen Pflanzenreste (Schenk’s Handbuch, vol. iv. 
1888), p. 186; Nathorst, Ofvers. hongl. Vetensk.-Akad. Forhand. 
1879, No. 3; Renault and Zeiller, Compt. Rend. vol. cvii. 1888, 
p. 1022; Saporta, Pal. Frang. vol. iv. 1891, p. 38; Seward, 
A new British Carboniferous fossil, MVaturalist, 1894, p. 238; 
A. Hollick, Remarks on a paper by Dean in the Zrans. New 
York Acad. Sct. vol. xiii. 1893, p. 115, ete. ] 


1 Conwentz, p. 61. 


. 
. 
. . 
ae =e De, 


ADDENDA TO VOL. I. 229 


AVDENDA TO VOLE EL 


Nathorstia valdensis (=Leckenbya valdensis), gen. et sp. nov. 
Volo ie py ane VALS Bis. 5, and Pl. LX. Figs. 2 and Qa. 

After the publication of the first volume of the Wealden 
Catalogue, it was pointed out to me by Prof. Nathorst that his 
name had been previously made use of by Heer for certain 
marattiaceous ferns from the Cretaceous strata of Pattorfik, in 
Greenland. This oversight on my part was corrected in a short 
note published in the Geological Magazine for 1894,? and the 
generic name Leckenbya suggested as a substitute for WVathorstia. 


Phyllopteris acutifolia (= Sagenopteris acutifolia). Vol. I. p. 148, 
Pi, EX. Bies.6: 

The generic name Phyllopteris was chosen for certain small 
isolated leaflets, on account of the apparent absence of any 
anastomosis of the lateral veins. The examination of more 
perfect specimens has enabled me to detect true reticulate venation, 
and to confirm Mr. Rufford’s opinion that the leaflets should be 
included in the genus Sagenopteris. 


Weichselia Mantelli (Brong.). Vol. I. p. 114. 

In the synonymy of this species, I included a plant named by 
Nathorst Weichselia erratica, as probably identical with the English 
Wealden form. Prof. Nathorst* has expressed some doubt as to 
the correctness of this suggestion, his species being found in 
a deposit which is probably of Upper Cretaceous age; he adds 
that there are no Wealden strata in Sweden. 


Sagenopteris Mantel (Dunk.). Vol. I. p. 132. 

In speaking of this species I referred to a plant described by 
Velenovsky under the name of Thinnfeldia variabilis as probably 
identical with S. Muntelli, and added, with regard to his use of the 


1 Fl. foss. Arct. vol. vi. 1882. 
2 Geol. Mag. 1894, p. 384. 
3 Letter, June 25, 1894. 


226 ADDENDA TO VOL. I. 


genus Thinnfeldia, that he made no reference to the resemblance 
of the leaves to the genus Sagenopteris. Velenovsky’s error 
was pointed out to him by Nathorst,’ with the result that he 
afterwards acknowledged his mistake,? a correction which I un- 
fortunately overlooked. 


CANADA. 


Sir William Dawson has pointed out to me, that I appear to 
have done injustice to Canadian geologists in the brief notice 
of the Kootanie plant beds in Vol. I. p. xxxi’ It was not 
my intention to give a complete historical sketch of these 
Cretaceous deposits, but I am glad to take the opportunity of 
calling attention to a paper by Dawson On the Correlation of 
early Cretaceous Floras in Canada and the United States, and on 
some new plants of this period. In this contribution we have a 
summary of the work of Richardson, G. M. Dawson, and others, 
and a description of some new plants from the Kootanie forma- 
tion of the Rocky Mountains. After giving a list of Kootanie 
species, Dawson discusses the age of the flora, and points out 
that while some of the plants must be regarded as Jurassic, the 
majority have a Lower Cretaceous facies. On the whole, he 
concludes that ‘‘ the Kootanie flora belongs to the lowest portion 
of the Cretaceous, and may be a little older than that of the 
main part of the Potomac formation.” * 

Among the plants described by Dawson we find the following 
Wealden forms: yuisetites Lyelli, Mant., Pecopteris Browniana, 
Dunk., Sphenolepidium, sp. The pinna figured as Pecopteris 
Browniana 1s very imperfectly preserved, and does not afford 
very satisfactory proof of the occurrence of Dunker’s Wealden 
species in the Kootanie flora. In another form, Cladophlebis 
falcata, Font., we have one of those ferns which it is very 
difficult to separate from the widely distributed C. Whitbyensis 
(Brong.) and C. <Albertsii (Dunk.). The fragment figured as 


1 Letter, June 25, 1894. 

* Velenovsky (A. 2), Abh. k. bohm. Ges. Wiss. vol. ii. 1888, p. 5. 
3 Letter, Oct. 17, 1894. 

4. Dawson (2), p. 938. 


ADDENDA TO VOL. I. DIRT 


Sphenopteris latiloba? Font., I am disposed to regard as identical 
with |S." Aiztontesew.: ef. Vol. I. p.111, Fig. V1; and Pl. VI. 
Fig. 2. The specimens figured by Dawson are for the most 
part very small and imperfectly preserved, and the task of deter- 
mination has necessarily been extremely difficult. 


AMERICA. 


Prof. Lester Ward writes to me as follows in reference to the 
geological age of the Potomac formation ! :— 

‘‘As you may know, I have been engaged for several years 
on our Potomac formation, and I have established the fact that 
it is by no means so simple a group as might be supposed from 
what has been thus far said about it. I have been able to 
subdivide it into no less than six somewhat distinct horizons, 
each of which is fairly well marked off stratigraphically, and 
has its own peculiar flora. The uppermost of these subdivisions 
embraces the well-known Amboy clays of New Jersey, which 
have yielded a very rich flora, a monograph of which had been 
nearly completed by Dr. Newberry at the time of his death, 
and has been edited by Dr. Arthur Hollick, and has now gone 
to press to be published by the U.S. Geological Survey. The 
difference between the lowest and highest of these beds is very 
great, and the flora is correspondingly different. I suppose that 
the series as a whole includes nearly the entire Lower Cretaceous. 

“Tt is only with the older Potomac that your Wealden of 
England can properly be compared. It is in that that we have 
the remarkable cycad trunks, so much like those of the Isle 
of Purbeck, which I suppose are really Jurassic; although, as 
you show, they are most intimately connected with the overlying 
Wealden.” 


PORTUGAL. 


Reference was made in Volume I. (p. xxiii.), to Saporta’s 
important work on the fossil flora of Portugal, and a few notes 
were added with regard to some of the more interesting species. 


1 Letter, July 3, 1894. 


228 ADDENDA TO VOL. I. 


Since this was written a large monograph’ has been published, in 
which the Portuguese floras receive a full and careful treatment ; 
this work is of considerable scientific importance, and its value 
is enhanced by the fact of its being the last important contribution 
to paleobotany by that indefatigable worker whose loss is so deeply 
deplored. 

Beginning with strata of infra-Liassic age, Saporta describes a 
number of plants from beds referred to the following geological 
horizons: Niveau de Sinémurien (Couches a Gryphea obliqua), 
Niveau du lusitanien, Niveaux du néo-jurassique (Couches a Lima 
alternicostata, ptérocérien et portlandien), Niveaux infra-crétaciques 
(du valanginien 4 l’aptien et de l'urgonien a V’albien). In summing 
up the characteristics and affinities of the neo-Jurassic floras, 
Saporta draws attention to the occurrence of species characteristic 
of the Corallian and Kimmeridgian, and associated with these he 
recognizes various Wealden types such as occur in Northern 
Germany, the Carpathians, and in North America. To discuss at 
length the numerous points suggested by this extremely valuable 
memoir, would take us far beyond the limits of the present work, 
but some portions of the monograph more immediately germane 
to our present subject must be briefly dealt with. In looking 
through the excellent plates, we find a large number of very small 
specimens referred to a great variety of species, and on examining 
the evidence on which many of the determinations are based, it 
would seem that the number of specific names might well be 
considerably reduced. 


Neo-Jurasstc SpEcrEs. 
Sphenopteris dissectifolia, Sap., p. 19, pl. iii. fig. 9; pl. vill. fig. 2; 
and pl. x. fig. 9. 

Saporta notes a resemblance to the Wealden species Ruffordia 
Gépperti (Dunk.); we may also draw attention to a resemblance 
with Onychiopsis elongata (Geyl.). 

S. marginata, Sap., p. 20, pl. vii. fig. 6. While recognizing 
the striking agreement of this fern with Onychiopsis Mantelli 
(Brong.), Saporta prefers to consider it a distinct type. It is 
impossible to be quite certain as to the precise nature of the 


1 Saporta (1). 


—— 


ADDENDA TO VOL. I. 229 


specimen, but it is difficult to understand on what grounds it can 
be separated from such a form as O. Mantellv. 

S. Mantelli neojurassica, p. 21, pl. vii. ete. 

S. (Davallia) Mantelli, Brong., p. 72, pls. xv. and xviii. 

After referring to the wide range and distribution of this fern, 
Saporta describes specimens of fertile fronds, which he compares 
with those of the recent species Davallia gibberosa, Sw., and 
D. coneinna, Schrad. The agreement of the fossil and recent 
specimens, leads the author to refer the Wealden fern to the genus 
Davallia, or at least to a sub-genus of the Davralliee. The figured 
examples of fertile pinne are less perfect than those which I 
have described in Volume I. (p. 50, Pl. III. Figs. 2-4); and the 
comparison made by Yokoyama in the case of Onychiopsis elongata, 
and by myself as regards O. Mantelli, with the recent genus 
Onychium, is, I believe, a much nearer approach to the truth than 
if we adopt the conclusions of Saporta. 

The following are a few of the numerous instances in which mere 
fragments are referred to specific types, or on which new species 
are founded :—Sphenopteris trifida, Sap., p. 26, pl. x. fig. 20; 
S. pedicellata, Sap., p. 26, pl. x. fig. 21; S. minima, Sap., p. 26, 
pl. xiv. fig. 16; S. trapezotdea, Sap., p. 27, pl. xi. fig. la; 
S. acutidens, Sap., p. 27, pl. x. fig. 14; Cladophlebis minor, 
Sap., p. 80, pl. iv. fig. 14; C. obtustloba, Sap., p. 30, pl. xiv. 
fig. 15. To carry the niceties of determination so far, and to 
institute new species on almost microscopic fragments of pinne 
or pinnules, is, I venture to think, a retrograde rather than a 
progressive method in paleobotany. A large number of other 
cases might be cited illustrating this method of determination, 
but the above may serve as examples of this dangerous practice. 
The specimens named S. microclada, Sap., p. 23, pl. vi., may 
be compared with S. Fontaine’, Sew., and those described as 
Scleropteris sinuata, Sap. (p. 45, pls. vii. and viii.), with Spheno- 
pteris Iittoni, Sew. 

Any traces of angiospermous plants in strata of Jurassic age 
are of special interest. Saporta, in speaking of fossil angiosperms, 
writes: ‘‘En dépit du nombre restreint des espéces et de l’extréme 
rareté des échantillons, la présence de végétaux anglospermiques 
dans la flore portugaise néo-jurassique est cependant certaine.”’? 


1 Saporta, Joc. cit. p. 56. 


230 ADDENDA TO YOL. T. 


The fragment referred to as Rhizocaulon vetus, Sap. (p. 57, pl. x. 
fig. 22), is very small, and in itself hardly satisfactory as an 
example of a monocotyledonous species. Other fragments of 
parallel-veined leaves are referred to five species of Poacites; it 
may be that we have in them portions of monocotyledonous leaves, 
but the specimens are so small and fragmentary, that one cannot 
feel much confidence in them as trustworthy evidence in so 
important a matter. 


Iyrra-Cretaceous (Valanginian to Aptian). 
Adiantum aneimiefolium, Sap., p. 82, pl. xv. fig. 21. 

There is apparently no evidence that this fragment belongs to 
the genus Adiantum: cf. Ruffordia Gépperti var. latifolia (Vol. 1. 
Pl. VI. Figs. 1 and 1a). Similarly there does not appear to be 
sufficient reason for naming the fragment represented in pl. xvi. 
fig. 14 Marattia minor, Sap. Oleandridium tenerum, Sap., p. 85, 
pl. xv. fig. 3; pl. xvi. fig. 18, as Saporta points out, closely 
resembles Zeniopteris Beyrichit (Schenk). The leaf figured as 
Glossozamites brevior, Sap., pl. xvi. fig. 82, may be a pinna of 
Otozamites K lipsteinit (Dunk.). 

Saporta has drawn my attention to the resemblance of Cyclopitys 
Delgadot, Sap. (p. 91), to the specimen figured by me in Vol. I. 
(p. 19, Pl. I. Fig. 7) as probably an equisetaceous sheath. 

From these infra-Cretaceous strata we have several specimens 
referred to Ehizocaulon and Poacites, which may be portions of 
monocotyledonous leaves, but the leaf fragment spoken of as 
Yuccites fractifolius, Sap., p. 110, pl. xix. fig. 20a, cannot be 
accepted as a certain monocotyledon. A few specimens are 
described as probably dicotyledons, but the existence of such 
plants at this horizon is considered as still problematical. 


Iyrra-Creracrous (Urgonian to Albian). 

Sphenopteris circalensis, Sap., p. 126, pls. xxiv. and xxv.: 
cf. Onychiopsis elongata (Geyl.). SS. cuneifida, Sap., p. 127, 
pl. xxii. fig. 5. This closely resembles the finely divided form 
of Ruffordia Gopperti (Vol. I. Pl. IV.). 

An interesting series of specimens is described and figured 
under the name of Jsoetes Choffati, Sap., p. 134, pls. xxiv., xxv., 
and xxvil.; these fossils show a striking resemblance to this 
recent genus of vascular cryptogams. Among the specimens 


EE ——— 


9 


ADDENDA TO VOL. I. 251 


figured as Protorrhipis Choffati, Sap., p. 144, pls. xxii., xxvi., 
and xxvil., there are some which suggest leaves very similar 
to those described by Bartholin from Bornholm under the name 
of Hausmannia Forchhammeri, Barth. 

Sphenopteris tenurfissa, Sap., p. 161, pl. xxviii. fig. 4. In all 
probability this is identical with Ruffordia Gépperti (Dunk.). 
Saporta himself noted the close agreement; but he does not 
accept my determination of the broad-leaved fronds figured in 
Vol. I. Pl. VI. as a variety of Dunker’s species. 

Adiantum eximium, Sap., p. 164, pl. xxviii. fig. 18; pl. xxxi. 
fig. 6. Probably identical with R. Géppertd var. latifolia. 

In speaking of Cycadites Saporta, sp. nov. (p. 30), I have 
drawn attention to the very close similarity of the Portuguese 
specimens named by Saporta Cycadites tenuisectus. The numerous 
dicotyledonous leaves described by Saporta are of considerable 
interest, but need not be dealt with here, as we have no trace 
of angiospermous fossils in our Wealden strata, which occupy 
a lower horizon than the beds from which Saporta’s specimens 
were obtained. 

In taking a review of the whole flora described in his monograph, 
Saporta calls attention to the remarkable series of types which are 
represented in the plant-bearing beds ranging from the Corallian 
to the Cenomanian; the series is practically continuous, and 
without any distinct break or hiatus in the succession of genera 
and species. The flora is compared with that of the Potomac beds 
of America; a close comparison of the two sets of plants! has 
also been recently instituted by Prof. Lester Ward. 


JAPAN. 


Yokoyama followed up his account of the Jurassic plants from 
Kaga, Hida, and Echizen, by a memoir on the Mesozoic plants 
from Kotzuke, Kii, Awa, and Tosa.? Nathorst*® had previously 
described several species from some of these localities, and con- 
cluded that the plant-bearing strata should be classed as transition 


1 Science, March 29, 1895. 
* Yokoyama. 
3 Nathorst (A. 3), Denkschr. k. Ak. Wiss. vol. lvii. 1890, p. 43. 


232 ADDENDA TO VOL. I. 


beds between the Jurassic and Cretaceous systems. Yokoyama 
supports Nathorst’s view, but prefers to regard the Japanese beds 
as corresponding to the whole Neocomian series, and to the Potomac 
of America. The following species are mentioned in Yokoyama’s 


paper :— 


Thyrsopteris, sp. Podozamites lanceolatus, L. and H., 
Dicksonia Tosana, Yok. var. minor, Heer. 

Dicksoniopteris Nawmanni, Nath. P. lanceolatus var. latifolia, Nath. 
Onychiopsis elongata (Geyl.). P. pusillus, Velen. 

O. elegans, Yok. Zamiophyllum Buchianum (Ett.). 
Adiantites Yuasensis, Yok. Z. Buchianum var. angustifolia, Font. 
Pteris (?), sp. Z. Naumanni, Nath. 

Sphenopteris tenuicula, Yok. Glossozamites parvifolius, Yok. 
Pecopteris Browniana, Dunk. Nilssonia Johnstrupi, Heer. 

P. Geyleriana, Nath. N. Schaumburgensis, Dunk. 

P. cf. virginiensis, Font. NV. pterophylloides, Yok. 

Cladophlebis Nathorsti, Yok. Ptilophyllum ct. Cutchense, Morr. 
Macroteniopteris (?) marginata, Nath. | Cyparissidium (?) Japonicum, Yok. 
LIycopodites, sp. Torreya venusta, Yok. 


Podozamites, sp. 


Some of the specimens described as Podozamites show a strong 
resemblance to Fontaine’s WVagevopsis; but it is no easy matte” 
to decide between these two genera when we have to deal with 
small specimens. 


AUSTRIA. 


Fritz Von Kerner! has recently described some fossil plants 
from the Island of Lesina, off the Austrian coast, in the Adriatic 
Sea; the flora is regarded as Lower Cretaceous in age. Among 
the species recorded from these beds we have Dvoonites cf. 
Saxonicus (Reich.), p. 49, pl. iv. fig. 6: this species I have 
included as a synonym of Zamites Buchianus (Ett.); but it is 
difficult to decide from the indistinct photograph of Kerner’s 
fragment how far his specimen may be regarded as identical with 
the Wealden species. A coniferous twig figured by Kerner as 
Pagiophyllum rigidum, Sap., resembles fairly closely some of the 
English specimens of P. crassifolium, Schenk: ef. Kerner’s figure, 
pl. iv. fig. 8, and my Pl. XVI. Fig. 2. 


1 Kerner. 


CONCLUSIONS. 2338 


A very small and imperfect specimen is figured as a piece of 
Sphenolepidium Kurrianum (Dunk.), (pl. iv. fig. 2); this does not 
seem to be a typical example of the species, but it is impossible 
to accurately determine so small a fragment. 


SPITZBERGEN. 


In 1883 Nathorst' published a short account of some plant- 
bearing beds in Spitzbergen, which had formerly been classed as 
Cretaceous, but which he preferred to consider as uppermost Jurassic 
in age. Having lately had an opportunity, through the kindness of 
Professor Nathorst, of examining the Spitzbergen fossils, I am 
disposed to think that we shall find in his forthcoming monograph 
on this flora a certain amount of evidence in favour of its being 
regarded as Wealden. j 


CONCLUSIONS. 


Prior to the acquisition of the Rufford Collection by the British 
Museum, the following plants had been recorded from the Wealden 
strata of England. Such alteration as I have suggested with 
regard to any of the species are noted in brackets. 


Chara (=C. Knowiltoni, sp. nov.). Pinites Mantelli, Carr. 

Lquisetites Lyelli, 8. and W. P. patens, Carr. 

LE. Burchardti, Dunk. Cycadeostrobus elegans, \ [ = Conites 

Onychiopsis Mantelli (Brong.). Carr. | elegans 

Sphenopteris Fittoni, sp. nov. (= 8. | C. ovatus, Carr. (Carr.) ]. 
Mantelli, Brong.). C. tumidus, Carr. [ = Conites tumidus, 

Tempskya Schimperi, Corda. (Carr.)]. 

Teniopteris Beyrichii (Schenk). Bucklandia anomala (S. and W.). 

Weichselia Mantelli (Brong.). Fittonia squamata, Carr. 

Sphenolepidium Kurrianum (Dunk.). Yatesia Morrisii, Carr. 

Araucarites Pippingfordensis (Ung.). Bennettites Saxbyanus (Brown). 

Pinites Dunkeri, Mant. Dioonites Brongniarti (Mant.) (= Cyca- 

P. Carruthersi, Gard. dites Brongniarti, Mant.). 

P. valdensis, Gard. Nilssonia Schaumburgensis (Dunk.). 


1 Nathorst (6). 


234 


The following list includes the species described in Volume I., 
with the addition of those dealt with in Volume II. 


} SPECIES CONFINED TO ENGLAND. 


THALLOPHYTA. 


t+ Algites valdensis, Sew. 


CHAROPHYTA. 


t Chara Knowltoni, Sew. 


BRYOPHYTA. 
+ Marchantites Zeilleri, Sew. 


PTERIDOPHYTA. 


Equisetites Lyelli, Mant. 

E. Burchardti, Dunk. 

E. Yokoyame, Sew. 
Onychiopsis Mantelli (Brong.). 
O. elongata (Geyl.). 

t Acrostichopteris Ruffordi, Sew. 
Matonidium Gopperti (Ett.). 
Protopteris Witteana, Schenk. 
huffordia Gopperti (Dunk.). 
R. Gopperti var. latifolia, Sew. 

+ Cladophlebis longipennis, Sew. 
C. Albertsii (Dunk.). 

C. Browniana (Dunk.). 


GYMNOSPERM 2. 


Cycadites Romeri, Schenk. 
+ C. Saporte, Sew. 
Dioonites Dunkerianus (Gopp.). 
D. Brongniarti (Mant.). 
Nilssonia Schaumburgensis (Dunk.). 
Otozamites Klipsteinit (Dunk.). 
+ O. Klipsteinii var. superbus, Sew. 
t O. Klipsteinit var. longifolius, Sew. 
Otozamites, sp. Ci. O. Reibeiroanus, 
Heer. 
O. Géppertianus (Dunk.). 
Zamites Buchianus (Ett.). 
t+ Z. Carruthersi, Sew. 
Z. Carruthersi var. latifolius, Sew. 


CONCLUSIONS. 


t A. catenelloides, Sew. 


Cladophlebis Dunkeri (Schimp.). 

Sphenopteris Fontainei, Sew. 

S. Fittoni, Sew. 

Weichselia Mantelli (Brong.). 

Teniopteris Beyrichii (Schenk). 
t+ T. Beyrichii var. superba, Sew. 
t T. Dawsoni, Sew. 

Sagenopteris Mantelli (Dunk.). 
T S. acutifolia, Sew. 

Mierodictyon Dunkeri (Schenk). 

Dictyophyllum Rémeri, Schenk. 
{ Leckenbya valdensis, Sew. 

Tempskya Schimperi, Cord. 


Anomozamites Lyellianus (Dunk.). 
Cycadolepis (Dory-Cycadolepis and 
Eury-Cycadolepis) . 
Carpolithes, sp. 
t Androstrobus Nathorsti, Sew. 
Bucklandia anomata (S. and W.). 
t Fittonia Ruffordia, Sew. 
t+ Bennettites Saxbyanus (Brown). 
t B. Gibsonianus, Carr. 
Bennettites, sp. 
t+ B. (Williamsonia) Carruthersi, Sew. 
+ B. (Williamsonia) Carruthersi var. 
latifolius, Sew. 
t+ Yatesia Morrisii, Carr. 


CONCLUSIONS. 935 


Sphenolepidium Kurrianum (Dunk.). | + Pinites Dunkeri, Carr. 
S. Sternbergianum (Dunk.). Tt P. Carruthersi, Gard. 
Cf. S. (Sequoia) subulatum, Heer. t P. Solmsi, Sew. 
Sphenolepidium, sp. t P. Ruffordi, Sew. 
Pagiophyllum crassifolium (Schenk). Cf. Nageiopsis heterophylla, Font. 
Pagiophyllum, sp. Tt Thuites valdensis, Sew. 
t+ Brachyphyllum spinosum, Sew. Conites (Araucarites), sp. 
B. obesum, Heer. tT C. armatus, Sew. 


PLANTA INCERTA SEDIS. 


Specimen A. (Vol. I. p. xxxv. Pl. I. Fig. 7). 
Specimen B. (Vol. f. p. xxxv. Pl. I. Figs. 8 and 9). 
+ Withamia Saporte, Sew. 
+ Becklesia anomala, Sew. 
Cf. Dichopteris levigata (Phill.). 


In the accompanying table an attempt is made to show the 
geographical range of such species as are not confined to the 
Wealden rocks of England. The occurrence of the same species 
in different regions is not regarded as necessarily proving homo- 
taxial strata. A more detailed consideration of the geological 
correlation of the plant-bearing strata, in which Wealden types 
occur, will be undertaken elsewhere, as there are still a few 
species to be described from the English beds. Such plants as 
Onychiopsis Mantelli (Brong.), Datonidium Gépperti, Schenk, 
Ruffordia Goépperti (Dunk.), are by no means confined to one 
geological horizon; and it is possible that we have other species, 
the range of which is not strictly limited to true Wealden strata. 


In the following list are added the chief districts or localities 
in each country from which the plants have been obtained. The_ 
names of the principal authors of the several Wealden floras (or 
floras containing Wealden species), are given below, with numbers 
referring to the bibliographies at the end of Part I. and Part II. ; 
those in the former being printed with A. after the author’s name, 
those in the latter with B. 


et I all i —_—ae ae 


(qq2.M PUL sox04g) v/yUoUn vepunpyon 

srererere (CoyTIN(T) snunypahT sazvumzomoupy 
eee eee eeteeeseeseeee (495) snumryang sayy 
rereeressess IOaTT ‘snuvouiagay sazumMz0j(—) 2 
tereseereres (ony) snupydsaddoy) sapzuUmz0jQ 
creanahcessrGex (rin) nuasdy y sazvunz0ji9 
tereeseee(oyTN(T) Ssusuabumquennyagy DULOSS]UAT 


ia ia) Sot) ate x oon sia8 afaie ree ees vee 500 x neta Sabnacoodnanoce (que) AQADUDUOMT sapruoouy 
ae ae p06 os 900 S00 cid 900 B00 a6 mee bob cio seseeerereees Adon ‘snumuayunyy say uooug 
rae ae Aa sae BEE ela faiatat eee eee eee eee cece eee a * “00 ‘ds ‘py sodny sap~pnahg 
oes asa Age eee eee sina arin eee An eee x eee wale ves saasvansseresslenree- Tg “waUlony sajipwalig 
cue eee Gad eee oe aa a diets eee eee soe male fark oe CELIO Chi fo) 3] “np hydo.agay sisdovbon 2 


4 


sees vrata “wuniyofissnia unjhydobog 
seeeseeerecseee ToaTT ‘wunsago wnpphydhyavug 
“+ (yuncq) wnunrbusquiagg unyndajouaydy 
seeees (CTT) wnunrwny Uunrprdajousydy 
PEO URC FICCIE II 91116) “wad nya! vhysduay, 
sree sagt ‘ds yo “was ‘siswapjva vhquayaiT 
srreereesess yrrgyag ‘euauoy wnpphydohjarq 
sreseeeesees (srtayog) wayung wohporpo.wryy 
Rristalentelstsleniets (-yunqq) Yaqun yr srvajdouabhoy 
seeeereecece (yuoyog) nya Mag ELON fi 
ROP DOSUICHOBOCOD (-Suoig) YJIQUD JY VV]ASYIV AY 
testeeerrereees sao “dg “UOMUT siuagdouaydgy 
apOnoedoDene (-duryog) 1ayuUn(y siqauydopng 
sesreresenes (omy) vupiunoug siga,ydopnyg 
AA GocenneTIorit) (-yunq) Us74ag) siqayydopnig 
strseeeceeeroeens (ormed) auaddoy vripLofinyy 


ae 
Oo. 

KK 
KK 
Le he 
KW 
Oo 

KKK 
KK 


nw 
tal 
i 
ww 


K 
24 
Oa. 


KKK KK KKK 
K 


= eae a oe det ae ey HR Ane ante eae ini sen EC UREREC LS Seale “pupary A 82.0090040.4. 
an eee eee oe see tee eee og x x x ee x eh dee e er eer ereeee (495) yuaddoy unrpruozD yy 
500 out ono x ses tee tee te ooo Sch aco 206 500 X seetevearsereseese (Thay) wynhuoja srsdovyahuge 


weececeecoccesr (-Suorg) Yjaqunyyr sasdorvyahug 
toreeeeeeees cag ds ‘wuphoyox sagyasinby 
Hireeeesecseees stingy “UgpmyaIng savas nbey 
see recveecerese see “VUB IL “any sagijas nba 


a 

tal 

KA 
ial 
tal 
wn 
tal 


al 


hw 
a. 
HMMM MRR MMM HK KKK KKK RRR 


*satoadg Jo 4stT 


“BISSUY 


*a0uRay 
“purlsuq 


*eolIOULy 
“wot u0g 
pur uapang 

*elysnVy 

“Auvutiey 

"mUnIS [aq 

*[esn 10g 


*puepusety 


CONCLUSIONS. E37 


Enetanp. Isle of Wight, Kent, Surrey, Sussex, and Bedfordshire. [Stokes 
and Webb (A.), Mantell (A. 1-7), Carruthers (A. 3 and 4), 
(U8, dl, By G, emel ts). Topley (A. 1.), Gardner (A. 1 and 2), 
Bristow (A.), Peyton (A.), ete. ] 

Portrucan. Cercal, Bellas, Torres-Vedras, etc. [Heer (A. 6), Saporta (B. 1).] 

FRANCE. Beauvais. [Brongniart (A. 4). ] 

Breicrum. Hainaut. [Coemans (A.), Bommer (B. 1 and 2).] 

Germany. Deister, Quedlinburg, Tentoburgerwald, and other localities in 
N.W. Germany, [Dunker (A. 2), Schenk (A. 2 and 4), Hosius 
and Von der Marck (A. 1 and 2).] 

AUSTRIA. North Carpathians. [Ettingshausen (A. 4), Schenk (A. 3).]— 
Lesina I. [Kerner (B.).] 

Russia. Klin. [Trautschold (A. 3).] 

Bornuoim. [Bartholin (A.) and (B.).] 

SWEDEN. Hor (Scania). [Nathorst (A. 4).] 

America. Virginia, Maryland, Montana. [Fontaine (A. 2 and 3), New- 
berry (A. 1), Knowlton (A. 2).] 

CANADA. Rocky Mountains (Kootanie R.). [Dawson (B. 2).] 

GREENLAND. Kome (Nugsuaks Peninsula). [Heer (B.).] 


JAPAN. Kaga, Hida, Echizen, Kii, ete. [Yokoyama (A. 2) and (B.), 
Nathorst (A. 3). ] 
AFRICA. Geelhoutboon (South Africa). [Tate (A.).] 


AvstrauiA. [Tenison- Woods (A.). ] 
New Zeauanp. Prov. Auckland. [Unger (A. 3).] 


The quéstions of geological age suggested by the above table, 
will be discussed in a subsequent communication on the Wealden 
floras, which it is intended to lay before the Geological Society 
at an early date. With a view to discover if the manner of 
occurrence of the fossil plants, or the association of different 
genera and species, would afford any evidence as to the relative 
positions of growth of the various floral types, I wrote to Mr. 
Rufford asking him to give me such information as his accurate 
knowledge of the Hastings district! enabled him to contribute with 
regard to this question. I cannot do better than reproduce the 
main facts which he communicated to me. He writes that he is’ 
unable to discover any reliable data as to the relative altitudes at 
which the plants grew, but adds the following useful information 


as to the occurrence of some of the characteristic species in the 
different beds :— 


1 For a geological section of this district see Vol. I. p. xvii. 


238 CONCLUSIONS. 


Farriuient Crays. 


(a) Fine clay ironstone; ferns well preserved, no cycads or 
conifers. This bed occupies about the same geological position as (d), 
although separated from it horizontally; it contains Marchantites, 
Liquisetites Burchardti, Onychiopsis Mantelli, Ruffordia Gopperti, 
Sphenopteris Fittoni, and other ferns. 

(6) Fine blue clay. The ‘‘cycad bed,” containing Zamites, 
Anomozamites, Otozamites, Fittonia, etc.; ferns rare. A few yards 
farther on, and about the same horizon, there have been found 
coniferous twigs, etc. 

(c) Porous sandstone, with Prnites; no cycads or ferns. This 
bed occupies a lower horizon than (a) and (8). 

(d) Blue clays, sometimes sandy. ‘‘Fern bed’; no cycads; 
occasionally leaves of Pinites, also Onychiopsis Mantelli, Ruffordia, 
Sphenopteris Fittoni, Cladophlebis, ete. 

(e) For the most part reddish ironstone, with some grey sand- 
stone merging into sandy clay. About the same horizon as (d); 
plants very abundant— Onychiopsis, Ruffordia, Sphenopteris, Clado- 
phlebis, Teniopteris, Protopteris; also Dioonites Brongniarti, Zamites, 
Otozamites, Benneitites (Williamsonia), Pagiophyllum, and other 
conifers, etc. This bed is of somewhat coarser material than the 
others, and contains a greater mixture of plants; Pagiophyllum 
crassifolium and other conifers are very abundant. 


In reading the above notes by Mr. Rufford, we find that with 
the exception of bed (e), in which the various classes of plants are 
well represented, the ferns, cycads, and conifers are not usually 
intermixed. The partial or complete separation of ferns, cycads, 
and conifers, may be due either to the nature of the plant material, 
which might be sorted by the water by reason of some differences 
in weight, or to the relative adaptability to longer or shorter 
transport by water; or the result of the plants growing in 
different districts and at different elevations. The more delicate 
fern fronds would probably be carried to a greater distance 
than the heavier and larger pieces of cycadean or coniferous 
plants. On the other hand, the gymnosperms may well have 
been more abundant on higher ground and in drier situations 
than the Filicine. Assuming the bed (e) to occupy the lowest 
horizon in the series, it would appear that the material com- 
posing the sandstones and ironstones was laid down in somewhat 


CONCLUSIONS. 239 


shallow water, and the fossils embedded in the strata were 
derived from a wide area, embracing localities rich in both 
ferns and gymnosperms. The petrological nature of (d) and the 
absence of cycads suggest deeper water; while the fine blue clay 
of (6) may have been derived from rocks in an area characterized 
by the predominance of cycads. It would, however, be difficult to 
reconstruct the conditions of growth of the several plants without 
a very careful examination of the rocks and their fossil contents, 
and at best our conclusions would probably not possess any great 
scientific value. 

In an old work by Unger,’ we read that in the Wealden period 
there were ‘‘ small wet islands, covered with forests, inhabited by 
the largest and most terrible monsters of the primitive world. 
The atmosphere was filled with moist vapour, and carbon dioxide 
exhalations favourable to the prodigious propagation of the 
amphibian race, and to the development of ferns, cycads, 
conifers, and some monocotyledons.’”? He goes on to say that 
‘La triste sauvagerie de cet intérieur de forét est encore redoublée 
par celle de ses habitants, parmi lesquels le gigantesque /ywanodon 
a crété osseuse et la monstreux Hy/gosaurus tiennent la premi€re 
place.” 

In a more recent monograph on the Wealden period, Schenk ? con- 
fidently speaks of the climate as undoubtedly tropical, and refers to 
the occurrence of tree ferns, the abundance of cycads, and other facts 
in support of this conclusion. It would be extremely difficult, 
or indeed impossible, to give any approximate estimate of the 
temperature in Northern and Central Europe during the Wealden 
period. The general characters of the vegetation would certainly 
seem to point to a tropical climate, and there can be little doubt 
that the temperature was considerably higher than the Wealden 
districts enjoy at the present day. 

In discussing the climate of a past age, in which no living species 
of plants existed, and in attempting to make use of fossil plants 
as indices of climatal conditions, we have to bear in mind the 
great danger of drawing conclusions from a comparison of extinct 
and living forms. It is superfluous to point out the lesson so 
clearly taught by recent plants, that closely allied species frequently 


1 Unger, p. 29. 
3 Paleontographica, vol. xix. p. 256. 


210 CONCLUSIONS. 


occur under very different conditions of temperature. In the 
present instance, the numerous species of cycads naturally suggest 
conditions similar to those most favourable or essential to the 
living representatives of the Cycadacee; but we must remember 
that the so-called cycadean fronds from Mesozoic rocks are nearly 
always found apart from the stems and reproductive structures, 
and we are still to a large extent in the dark as to the exact 
nature and structure of these extinct cycadean plants. 

Looking at the Wealden plants collectively, we notice a very 
striking agreement with the flora of the underlying Jurassic 
strata, and it would be difficult to point to any well-marked or 
essential difference between the plant-life of the two periods. 
The evidence of palsobotany certainly favours the inclusion of 
the Wealden rocks in the Jurassic series. 

One of the most attractive and difficult problems which is 
suggested to a botanist by such a flora as that of the Wealden 
period, is the evolution of angiospermous plants. A reviewer has 
happily expressed this in the following words’: ‘In the folds of 
the Wealden we imagine the secret of the evolution of angiosperms 
must be locked. It is as if we stood at the mouth of a great 
river flowing from an unexplored interior, whose flotsam we 
anxiously interrogated for clues as to the nature of the unknown 
Hinterland; yet nothing reaches us from beyond the coast-belt, 
which we have already explored.’”’” Among the English species 
there are none which can be regarded as the earliest angiosperms, 
and we search in vain among the abundant samples of the Wealden 
vegetation for any fragments of monocotyledonous or dicotyledonous 
plants. In the Potomac beds of America, which include strata of 
Jurassic and Lower Cretaceous age, we have several undoubted 
angiospermous species; and again, in the closely parallel series of 
Portuguese rocks, dicotyledons and monocotyledons are fairly 
abundant. The true Wealden vegetation would seem to have 
been without any examples of the highest class of plants, and 
may be looked upon as the last of the Mesozoic floras in which 
the gymnosperms represented the limit of plant development. 
One genus, however, carries us a few steps towards the next 
stage in botanical evolution ; the inflorescence of Bennettites marks 


1 Nature, July 26, 1894, p. 294. 


CONCLUSIONS. 241 


a distinct advance in the differentiation of reproductive structures 
beyond the characteristic cycadean type. Were we in a position 
to speak of the anatomical structure, or to describe more fully 
the reproductive organs, of Wealden plants, it might be that we 
should be able to recognize a distinct foreshadowing of angio- 
spermous characters; unfortunately, however, the extreme scarcity 
of mineralized plant tissues precludes any such treatment of plant 
types. In spite of the somewhat disappointing nature of the 
flora from the point of view of angiosperm development, we 
may reasonably hope that a more detailed comparison of floras 
possessing a Wealden facies will enable us to add something of 
value to the history of plant evolution, and to the facts of plant 
distribution. 


LIST OF WORKS QUOTED. 


[With two or three exceptions the works quoted in Part I. are not repeated in the 
following list. The letter (A.) after authors’ names in the footnotes of the 
present volume signifies that the work referred to is in the bibliography of 
Part I. Under Feistmantel, Heer, and Zittel, I have given the dates of the 
several parts of vols. i. and ii. of the Memoirs of the Geological Survey of 
India (second series), of the Flora fossilis Arctica, and of Schimper and 
Schenk’s volume on fossil plants in Zittel’s Handbuch. ] 


Balfour, J. H. Introduction to the Study of Paleontological Botany. 
Edinburgh, 1872. 

Bartholin, C. T. Nogle i den bornholmske Juraformation forekom- 
mende Planteforsteninger. [Bot. Tvdssk., vol. xix. 1894, p. 87.] 

Bennett, J. J. See Brown, R. 

Bensted, W. H. (1) On the Kentish Rag as exhibited in the Iguanodon 
Quarry at Maidstone. [Proc. Geol. Assoc. vol. i. 1860, p. 57.] 

Bensted, W. H. (2) Notes on the Geology of Maidstone. [Geologist, 
vol. v. 1862, p. 294.] 

Berger, H. A. C. Die Versteinerungen der Fische und Pflanzen im 
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244 LIST OF WORKS QUOTED. 


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(a) Jurassic (Oolitic) Flora of Kach. Pt. i. 1876. 
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246 LIST OF WORKS QUOTED. 


Géppert, H. R. (1) Ueber die fossilen Cycadeen iiberhaupt, mit Riick- 
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Géppert, H. R. (2) Beitrige zur Kenntniss fossiler Cycadeen. [ Vewes 
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Goppert, H. Rk. (3) Ueber die gegenwirtigen Verhiltnisse der Palion- 
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Goppert, H. R. (5) Zur Flora des Quadersandsteins in Schlesien. 
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Goppert, H. R. (6) Monographie der fossilen Coniferen. [Vat. Verhand. 
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Goppert, H. R. and Stenzel, G. Die Medullosex. [Palcontographica, 
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Gordon, EL. G. The Pinetum. London, 1858. 

Grand’ Eury, M. C. Géologie et paléontologie du bassin houiller du 
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(ii.) Mit Einem Anhang iiber versteinerte Hélzer der 
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Vol. II. Winterthur, 1871.—(i.) Fossile Flora der Biren-Insel. 
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(ii.) Flora fossilis Alaskana ; Fossile Flora von Alaska. 
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(iii.) Die Miocene Flora und Fauna Spitzbergens. [Jdid. 
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LIST OF WORKS QUOTED. 247 


being a description of the plants collected by Mr. Edward 
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Vol. III. Ziirich, 1875.—(i.) Beitriige zur Steinkohlen-Flora. der 


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(iii.) Nachtriige zur Miocenen Flora Grénlands.  [Lbid. 
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(iv.) Uebersicht der Miocenen Flora der arctischen Zone. 
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Vol. IV. Ziirich, 1877.—(i.) Beitriige zur fossilen Flora Spitz- 


Vol. 


Vol. 


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(ii.) Beitrige zur Jura-Flora Ost Sibiriens und des Amur- 
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(iii.) Ueber die Pflanzen-Versteinerungen von Andé in 
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V. Ziirich, 1878.—(i.) Die Miocene Flora des Grinnell- 
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(iii.) Primitize floree fossilis Sachalinensis. Miocene Flora 
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(iv.) Ueber fossile Pflanzen von Novaja Semlya. [A’. Svensk. 
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(v.) Beitriige zur Miocenen Flora von Sachalin. [Jbid. 
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VI.—Flora fossilis Grénlandica. Die fossile Flora Gronlands. 

Th. i. Ziirich, 1882. 

(i.) (a) Flora der Kome Schichten, pp. 1-19 } 
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(iii.) Nachtriige zur fossilen Flora Grénlands. [X. Svensk. 
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(iv.) Beitriige zur Miocenen Flora von Nord - Canada. 
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(v.) Untersuchung iiber fossile Hélzer aus der arctischen 
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pis. i.—xlvii. 


248 LISE OF WORKS QUOTED. 


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250 LIST OF WORKS QUOTED. 


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eee eee ——E———— 


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Solms-Laubach, Graf zu. See Capellini. 

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Williamson, W.C. (1) On the Distribution of Fossil Remains on the 
Yorkshire Coast, from the Lower Lias to the Bath Oolite 
inclusive. [Zrans. Geol. Soc. (2) vol. v..1840, p. 223.] 


252 LISY OF WORKS QUOTED. 


Williamson, W. C. (2) On the Scaly Vegetable Heads or Collars 
from Runswick Bay, supposed to belong to Zamia gigas. 
[Proc. Yorks. Phil. Soc. vol. i. 1847, p. 45.] 

Williamson, W. C. (3) Contributions towards the History of Zamia 
gigas, L. and H, [Zinn. Trans. vol. xxvi. 1870, p. 663.] 

“ates, J. Notice of Zamia gigas. [Proc. Yorks, Phil. Soc, vol. i. 1855, 
p. 37.] 

Yokoyama, M. Mesozoic Plants from Kozuke, Kii, Awa, and Tosa. 
[Journ. Coll. Sct. Japan, vol. vii. pt. iii. 1894, p. 201.] 

Young, G. and Bird, J. A Geological Survey of the Yorkshire Coast. 
Whitby, 1822. 

Zeller, R. See Renault and Zeiller. 

Zeiller, R. (1) Bassin houiller et Permien d’Autun et d’Epinac. Paris, 
1890. 

Zeiller, R. (2) Notes sur la flore des couches permiennes de Trienbach 
(Alsace). [Bull. Soc. Géol. France, vol. xxii. (3) 1894, p. 163.] 

Zeiller, R. (3) Observations sur quelques cuticles fossiles. [Ann. Set. © 
Nat. 1882, p. 217.] 

Zigno, A. (1) Flora fossilis formationis Oolithice. Vol. ii. Padova, 
1873-85. 

Zigno, A. (2) Monografia del genere Dichopteris nuovo genere di felce 
fossile. [J/ém. instit. Veneto, vol. xii. 1864. ] 

Zittel, K. A. Handbuch der Paleontologie. Abtheilung ii.: Paleeo- 
phytologie. Schimper, W. P. and Schenk, A. Munich and 
Leipzig, 1890. 


Lieferung i. - - pp. 1-152 - 1879. 
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Roh it CORNY MAINT G5 O58 er enagu: 


INDEX OF GENERA, SPECIES, ETC. 


[Synonyms are printed in italies. ] 


Abies, 193. 

Abietites ellipticus, 196. | 

Acanthozamites, 175, 176. 

Acrostichopteris Ruffordi, 234. 

Adiantites Klipsteinii, 60. 
Yuasensis, 232. 

Adiantum aneimizfolium, 230. 
eximium, 231. 

Agathis, 187, 211. 
Australis, 187. 
Dammara, 187. 

Alethopteris, 5. 

Algites, 114. 
catenelloides, 234. 
valdensis, 234. 

Androstrobus, 108-112. 
Balduini, 109, 111. 
borealis, 109. 
Guerangeri, 109, 112. 
Nathorsti, 110-112, 234. 
Sibiricus, 109. 
zamioides, 109. 

Aneimidium Klipsteini, 60. 


Anomozamites, 36, 50, 51, 53, 90-94. 


gracilis, 53. 

Lindleyanus, 150. 

Lyellianus, 91-94, 234, 236. 

minor, 53, 150. 

Muelleri, 14. 

Nilssoni, 92, 94. 

Schaumburgensis, 53. 
Aralia, 24. 


Araucaria, 114, 187, 190, 191, 211, 218. 


Bidwilli, 222. 

Cooki, 222. 

Cunninghami, 186. 

excelsa, 188. 

imbricata, 186. 
Araucarites, 190-192, 235. 

curvifolius, 201, 205, 207, 208. 

Dunkeri, 200, 205. 

hamatus, 200. 

Hudlestoni, 191. 

Phillips, 112. 

Pippingfordensis, 192, 233. 
Asterophyllites, 155. 


Athrotaxis, 199, 200, 202, 216. 
grandis, 221. 
lexifolia, 200, 203. 
selaginoides, 188. 

Athrotaxopsis expansa, 201, 202, 221. 


Baiera, 5. 
Balanophore, 150. 
Becklesia, 179-182. 
anomala 179-182, 235. 
Bennettitez, 6. 
Bennettites, 6, 7, 9, 96, 97, 116, 117, 
120, 123, 134-164. 
Carruthersi, 141, 145, 157-164, 
234, 
etrusca, 153. 
Gibsonianus, 120, 136, 138-140, 
142-144, 158, 160. 
Moriérei, 137, 145, 157. 
Saxbyanus, 139-144, 233, 234. 
Biota orientalis, 188. 
Blastolepis, 150. 
falcata, 150, 155. 
otozamitis, 150, 154. 
Bolbopodium, 120. 
Bowenia, 3, 4, 20. 
Brachyphyllum, 204, 214-221. 
confusum, 219. 
crassicaule, 216, 218-221. 
Desnoyersii, 216, 
Germari, 200. 
gracile, 220. 
Kurrianun, 200. 
nepos, 216. 
obesiforme, 218-220. 
obesum, 190, 204, 216, 218-221, 
230, 236. 
parceramosum, 219. 
spinosum, 190, 215-218, 234. 
Bucklandia, 99, 100, 121-132, 165, 
167, 190. 
anomala, 123, 169, 233, 234, 236. 
Mantelli, 124, 125, 127. 
Milleriana, 165. 


254 


Calamus, 177. 

Calpoxylon, 13. 
Camptopteris spiralis, 182. 
Cardiopteris, 63. 

Carpolithes, 101-107, 115. 
Carpolithus Mantelli, 106. 
Cedrus, 197. 

Cephalotaxus, 103, 187, 189. 

Fortunei, 26. 

Ceratozamia, 3, 17, 78, 87, 95. 

Hoffmanni, 15. 

Mexicana, 21, 82. 
Changarniera inquirenda, 171. 
Chara Knowltoni, 233, 234. 
Cladophlebis Albertsii, 226, 234, 236. 

Browniana, 234, 236. 

Dunkeri, 234, 236. 

falcata, 236. 

longipennis, 234. 

minor, 229. 

Nathorsti, 232. 

obtusiloba, 229. 

Whitbyensis, 226. 
Clathraria, 121, 122, 124, 127, 166. 

anomala, 122, 123. 

Galtiana, 126. 

Lyelli, 119, 122-126, 128, 130, 

131, 142-144. 

Mantelli, 124. 

strigata, 10. 
Clathropodium, 120. 

Conifer, 6, 138, 108, 118, 115, 136, 
138, 185-224. 
Conites, 113-116, 189, 222. 

armatus, 222, 230. 

elegans, 116, 116, 188, 190. 

tumidus, 233. 

Cordaitez, 9. 
Cordaites, 9, 18, 63, 99. 

lancifolius, 10. 
Crossozamia, 76. 

Cryptomeria, 187. 

Japonica, 213. 
Ctenis, 20. 

falcata, 10, 89. 

Leckenbyi, 52. 
Ctenophyllum, 37, 40, 41, 59, 90. 

latitolium, 90. 

pectinata, 73. 
Cunninghamia, 202, 211. 
Cupressinee, 187. 

Cupressus Lawsoniana, 220. 
Cycadacee, 2-172. 
Cycadeoidea, 120, 139, 140. 

etrusca, 158. 

Gibsoni, 142. 

Morrisii, 166. 

Saxhyana, 139. 

Yatesvi, 166. 


ALPHABETICAL INDEX. 


Cycadeoidez, 119, 120. 
Cycadeomyelon, 120, 126, 130, 131. 
Cycadeostrobus, 114, 115. 

Brunonis, 114. 

elegans, 116, 116, 233. 

ovatus, 116, 233. 

truncatus, 116. 

Cycadinocarpus, 102, 103. 

Cycadites, 11, 19, 20, 23-35, 43, 48, 
106, 108, 120. 

acinaciformis, 33. 
alatus, 25. 
Brongniarti, 30, 31, 48, 44, 47, 
48, 50, 233. 
constrictus, 27. 
Dicksoni, 25, 27. 
Escheri, 14, 27. 
gramineus, 26. 
gyrosus, 10, 11, 26. 
Heerii, 31, 32, 44. 
lanceolatus, 146. 
linearis, 28. 
Morrisianus, 20, 29, 31, 34, 36, 
42, 44. 
Nilssoni, 23. 
palmatus, 23. 
planicosta, 27. 
Rajmahalensis, 33. 
rectangularis, 32. 
Rémeri, 27-30, 32, 234, 236. 
Saportee, 28, 29-35, 44-46, 66, 
231, 234, 236. 
Saxbyanus, 134, 1389. 
Sibiricus, 26. 
Steenstrupi, 105. 
taxodinus, 10, 11, 25. 
tenuisectus, 30, 231. 
Unjuga, 27. 
zamioides, 23, 26. 
Cycadolepis, 94-101, 234. 
hirta, 96-98. 
villosa, 96, 97, 160. 
Cycadorachis, 173. 
abscissa, 173. 
armata, 173, 175, 178. 
Cycadospadix integer angustior, 97. 
Cycadospermum, 102. 
obovatum, 106. 

Cycadoxylon, 8. 

Cycas, 3, 4, 20, 24, 29, 78, 95-97, 
103, 106, 108, 118, 122-124, 136, 
173, 176, 187, 189. 

Beddomei, 20. 

Cairnsiana, 20. 

circinalis, 16, 25, 124. 
media, 19. 

revoluta, 19, 28, 124, 130. 
Siamensis, 28. 


Cyclopitys Delgadoi, 230. 


ALPHABETICAL INDEX. 


Cyclopteris frondosa, 62. 
Kilipsteinti, 58, 60. 
Mantelli, 60. 

Cyclozamites, 58. 

Cylindropodium, 120, 166, 167. 

Cynara integrifolia, 146. 

Cyparissidium gracile, 202. 
Japonicum, 203, 282. 


Dacrydium elatum, 188. 

Kirkii, 188. 

Westlandicum, 188. 

Davallia, 229. 
concinna, 229. 
gibberosa, 229. 

Desmoncus, 177. 

Dichopteris, 183, 184. 
levigata, 184. 

Visianica, 184. 

Dicksonia Tosana, 232. 

Dicksoniopteris Naumanni, 232. 

Dictyophyllum Rémeri, 234, 236. 

Dictyozamites, 4. 

Dioon 3s) LiselSsn9o0e0 97, 1095 e111), 

118, 173. 

edule, 22. 

Dioonites, 18, 31, 35-49, 71, 75, 80. 
abietinus, 32, 42, 48, 83, 85. 
Brongniarti, 31, 39, 47-49, 233, 

234, 
Buchianus, 79, 80, 82, 85. 
Dunkerianus, 31, 34, 40-47, 234, 
236. 

Goppertianus, 70, 71. 

Kotoei, 47-49. 

Kurrii, 39. 

Lyellianus, 91. 

Saxonicus, 79, 232. 
Dory-Cycadolepis, 96-98, 234. 
Dracena, 170, 171. 

Benstedti, 118, 169, 170, 172. 


Echinostrobus squamosus, 216. 
Encephalartopsis, 21, 87. 
Encephalartos, 3, 17, 21, 22, 44, 45, 
66, 76, 78, 87, 111, 118, 131. 

Altensteinii, 111. 

Catfer, 18, 22, 87. 

cretaceus, 22. 

cycadifolius, 22, 45, 87. 


Ghellinckii, 26, 22, 29, 44, 45, 87. 


Gorceixianus, 14, 22. 
horridus, 22, 87. 
Lehmanni, 22, 69, 82. 
longifolius, 21, 88. 
pungens, 87, 111. 
Endogenites erosa, 169. 


ho 
Or 
On 


Eolirion, 150. 

Equisetites, 102, 106. 
Burchardti, 102, 233, 234, 236,238. 
Lyelli, 226, 233, 234, 236. 
Yokoyame, 102, 234, 236. 

Kuphorbiacee, 121, 124. 

Eury-Cycadolepis, 96, 98-101, 234. 


Filicine, 4, 8, 11, 51, 57, 176, 184. 
Filieites Bucklandi, 56. 
Fittonia, 98, 100, 120, 128, 128, 129, 
131-134. 
insignis, 133, 172. 
Rigauxi, 99. 
Ruffordi, 132, 133, 234. 
squamata, 99, 132, 239. 
Flores, 108-112, 146-164. 
Frenela, 180. 
Frenelites Reichii, 202. 
Frenelopsis, 180, 181, 209. 
Hoheneggeri, 180, 182, 189, 209, 
210. 
ramosissima, 209. 
Fricia, 110. 


Ginkgo, 13, 108, 178, 186, 189. 

Gleichenia Hantonensis, 176. 

Glossozamites brevior, 230. 
oblongifolius, 65. 
parvifolius, 232. 
Stoliczkanus, 62. 

Glyptostrobus, 202. 
heterophyllus, 185. 

Goniolina, 151. 

Gymnosperme, 2. 


Hausmannia Forchhammeri, 231. 
Hisingera Manteili, 47. 
Hyleosaurus, 239. 


Iguanodon, 239. 
Isoetes Choffati, 230. 


Juniperites Sternbergianus, 205. 
Juniperus Bermudiana, 188. 
Chinensis, 206. 


Kaidocarpon minor, 190, 191. 
Kaloxylon, 8. 


Larix, 103, 186, 197. 
Leckenbya, 226. 


256 


Leckenbya valdensis, 225, 234, 236. 
Lepidodendron, 215. 
Lepidofloyos, 117. 
laricinus, 117. 
Leptostrobus longifolius, 197. 
Libocedrus decurrens, 187. 
Liliacez, 121. 
Lycopodites, 200, 207, 232. 
curvifolius, 206, 213. 
Lyginodendron, 8. 


Macroteniopteris marginata, 232. 

Macrozamia, 3, 78, 95, 101, 123. 
Denisoni, 18. 

Douglasii, 99, 105. 
Dyeri, 96. 
heteromera, 5, 182. 
Macleayi, 21, 82. 
spiralis, 43. 

Mantellia, 119, 120, 135. 

Marattia minor, 230. 

Marattiacez, 2, 7. 

Matonidium Gopperti, 234-236. 

Medullosa, 119. 

Leuckarti, 8. 

Megalozamia, 98. 

Microcyeas, 3. 

Microdictyon Dunkeri, 234, 236. 

Moreania, 211. 
brevifolius, 211. 

Muscites faleifolius, 201. 
imbricatus, 200, 201. 
Sternbergianus, 205, 207. 

Myelopteris, 7. 

Myeloxylon, 7, 8. 


Nageia, 210. 
Nageiopsis, 210, 211, 282. 
heterophylla, 190, 211, 235, 236. 
Nathorstia, 225. 
valdensis, 225. 
Neuropteris, 5, 16. 
levigata, 183, 184. 
Nilssonia, 4, 16, 23, 25, 37, 48, 50-56, 
90, 183. 
eequalis, 50. 
Bergeri, 26. 
brevis, 25, 50. 
Brongniarti, 47. 
elongata, 50. 
Johnstrupi, 232. 
pecten, 42. 
pterophylloides, 232. 
pygmea, 14. 
Schaumburgensis, 53-56, 232-234, 
236. 
Serotina, 14. 


ALPHABETICAL INDEX. 


Noeggerathia, 11-13, 16. 
Noeggerathiace, 13. 


Oleandridium tenerum, 230. 
Onychiopsis elongata, 228-232, 234, 
Mantelli, 198, 202, 228, 229, 232- 
236, 238. 
Oolithes, 107. 
spheericus, 107. 
Ophioglossacez, 7. 
Otopteris, 56-58, 62. 
lanceolata, 73. 
Otozamites, 40, 56-75, 77, 94, 98, 150. 
angustifolius, 72, 73. 
Beanii, 58, 61, 62, 64-67, 69. 
Bucklandi, 57, 58. 
Bunburyanus, 48. 
decorus, 63. 
Goéppertianus, 19, 21, 48, 70-74, 
1338, 234. 
gramineus, 57, 49. 
Klipsteinii, 18, 60-70, 89, 230, 
234, 236. 
lagotis, 63. 
latifolius, 64. 
latior, 72, 73. 
marginatus, 69. 
Molinianus, 62. 
obtusa, 57. 
Reglei, 58. 
Reibeiroanus, 70, 234, 236. 


Pachyphyllum, 211, 212. 
crassifolium, 212. 
levigata, 184. 
Pachypteris, 183, 184. 
Pagiophyllum, 211-218, 216, 218. 
crassifolium, 190, 212, 213, 232, 
235, 236. 
rigidum, 232. 
Paleobromelia, 224. 
Paleeozamia pecten, 147. 
recta, 88. 
Palissya, 26. 
Pandanus, 147, 171. 
Pecopteris Browniana, 226, 232. 
decipiens, 33. 
Geyleriana, 232. 
linearis, 71. 
virginiensis, 232. 
Phyllocladus, 176, 177. 
Phyllopteris acutifolia, 225. 
Pinites, 193-199. 
Andrei, 195, 196. 
Carruthersi, 190, 1938, 195, 196, 
233, 235. 
Coemansi, 195. 


ALPHABETICAL INDEX. 51 


Pinites cylindroides, 193, 194. 


Dunkeri, 190, 194, 195, 233, 235. 


macrocephala, 114. 

Mantelli, 233. 

patens, 233. 

Pottoniensis, 193, 194. 

Ruffordi, 190, 199, 228, 224, 235. 

Solmsi, 190, 194-198, 235. 

valdensis, 193-195, 233, 235. 
Pinus, 103, 186, 193, 196, 223. 

Coulteri, 222. 

longissima, 195. 

pinea, 188. 

Sabiniana, 222. 

succinifera, 224. 
Plagiozamites, 12. 

Platylepis, 120. 
Poacites, 230. 
Podocarpus, 187, 210. 

andina, 187. 

cupressina, 188. 

Podocarya, 147, 150, 151, 152. 
Podozamites, 76, 77, 187, 210, 232. 

distans, 94. 

lanceolatus, 94, 282. 

pusillus, 232. 

Reinii, 64. 

Poroxylon, 8. 
Protopteris, 117. 

punctata, 118. 

Witteana, 234, 236. 
Protorrhipis Choffati, 231. 
Pseudofrenelopsis, 181. 
Pteridophyta, 2. 

Pteris, 232. 

Pterophyllum, 10-12, 18, 19, 25, 31, 
35-37, 43, 48-51, 71, 75, 80, 91, 
92 


abietinum, 42, 48. 
equale, 48. 
angustifolius, 48. 
blechniforme, 63. 
blechnoides, 10. 
Braunianum, 40. 
Braunii, 36, 38, 40. 
Brongniarti, 39, 47. 
Buchianum, 39, 75, 79. 
Carnallianum, 81. 
Combrayi, 13. 
Cottzanum, 10. 
Dunkerianun, 32, 87, 38, 42, 44. 
Fayoli, 12. 
gonorrachis, 10. 
Goppertianum, 44, 70. 
Grand’ Euryanum, 11. 
Humboldti, 80. 
inconstans, 86, 38. 
inflexus, 10. 

Jaegeri, 13, 35, 37, 38. 


Pterophyllum Lyellianum, 91. 


majus, 50. 

medianum, 36. 
Nilssoni, 92. 
oblongifolium, 65, 68. 
pecten, 40. 
Richthofeni, 49. 
Saxonicum, 79, 80. 
Schaumburgense, 53. 


Pterozamites, 76. 
Ptilophyllum, 40, 41, 51, 56, 59. 


acutifolium, 18. 
Cutchense, 41, 232. 
oligoneurum, 49. 


Ptilozamites, 52, 53, 64. 


Heeri, 53. 


Rachiopteris, 173. 


aspera, 8. 


Rhizanthee, 153. 
Rhizocaulon, 230. 


vetus, 230. 


Ruffordia Gépperti, 228, 230, 231, 234, 


235, 238. 


Sagenopteris, 226. 


acutifolius, 225, 234. 
Mantelli, 225, 234, 236. 


Saportea, 174. 
Saportaia, 174, 
Schizoneura, 155. 
Scleropteris, 183. 


levigata, 184. 
Pomelii, 184. 


Sequoia, 199, 200, 202. 


ambigua, 206. 
gracilis, 202. 
Reichenbachii, 208. 
sempervirens, 187. 
subulata, 208. 
Tournalii, 189. 


Sigillaria, 117. 


Brardii, 133. 


Spermaphyta, 2. 
Sphenolepidium, 199-208. 


Kurrianum, 189, 190, 199-204, 
206, 221, 233, 235, 236. 

Sternbergianum, 190, 205-208, 
213, 235, 236. 

subulatum, 190, 208, 234. 

Virginieum, 201-203. 


Sphenolepis, 199. 


Kurriana, 200, 201. 


Sphenopteris acutidens, 229. 


Brongniarti, 63. 
circalensis, 230. 
cuneifida, 230. 


n 


Sphenopteris dissectifolia, 228. 
Fittoni, 227, 229, 233, 234, 236, 
238. 
Fontainei, 101, 178, 229, 234. 
lanceolata, 183. 
latiloba, 227. 
Mantelli, 60, 229. 
Mantelli neojurassica, 229. 
marginata, 228. 
microclada, 229. 
minima, 229. 
pedicellata, 229. 
sinuata, 229. 
tenuicaula, 232. 
tenuifissa, 231. 
trapezoidea, 229. 
trifida, 229. 
Sphenozamites, 57, 176. 
latifolius, 176, 178. 
Rochei, 11. 
Spirangium Jugleri, 224. 
Stangeria, 3, 4, 20, 97. 
paradoxa, 16. 
Stangerites, 16. 
Stenzelia, 7. 
Strobilites, 115, 189. 


Teniopteris, 4, 16, 55. 

Beyrichil, 55, 56, 230, 233, 234, 

236. 

Dawsoni, 234. 
Taxodium distichum, 186. 
Taxus, 138. 

baceata, 187. 

Tempskya Schimperi, 2838, 234, 236. 
Thinnfeldia, 183, 226. 

variabilis, 225. 

Thuites, 190, 209, 210. 

Choffati, 201. 

Germari, 200. 

Hoheneggeri, 180. 

Kurrianus, 200, 201. 

valdensis, 209, 210, 235. 
Thuja occidentalis, 187. 
Thujopsis, 216. 
Thyrsopteris, 232. 
Titanophyllum, 13. 

Torreya, 189. 

venusta, 232. 
Trunci, 116, 169-172. 
Tsuga Douglasii, 197. 
Tylodendron, 150, 186. 


Vlospermum, 102 


Voltzia, 130, 186. 
heterophylla, 189. 


ALPHABETICAL INDEX. 


Walchia, 26. 

Weichselia erratica, 225. 

Mantelli, 67, 225, 233, 234, 236. 
Weltrichia, 150, 151. 
Widdringtonia Whytei, 188. 
Widdringtonites curvifolius, 206. 

Haidingeri, 200, 201, 204, 206. 

Kurrianus, 200. 

Williamsonia 9, 96, 97, 120, 146-164. 
acuminata, 155, 156, 166. 
angustifolia, 150, 165. 
Blanfordi, 155. 

Bucklandi, 154. 

cretacea, 155, 156. 

elocata, 155. 

Forchhammeri, 155. 

Gagnierei, 154, 160. 

gigas, 145,147, 150,154, 158-160, 

163. 

Italica, 156, 160. 

Leckenbyi, 147, 150, 154, 155. 

microps, 15d. 

minima, 164, 155. 

Moriérei, 137, 154. 

pictaviensis, 154. 

Pougneti, 154, 155. 

recentior, 165, 156. 

Riesii, 155, 156. 

virginiensis, 97, 155. 

Zeilleri, 154, 155. 

Withamia, 173-179. 
armata, 178. 

Saporte, 174-179, 235. 


Yatesia, 120, 165-169. 
Joassiana, 165, 166. 
Morrisii, 166-170, 238, 234. 

Yucca, 170. 

Yuccites, 150, 151. 
fractifolius, 230. 


Zamia, 3, 4, 20, 76, 78, 87, 112, 118, 
Wale 


angustifolia, 20. 
cycadifolia, 82. 
furfuracea, 17. 

gigas, 94, 146-148, 153. 
elobuliferus, 81. 
Lindeni, 22. 

Loddigesii, 118, 170, 171. 
macrocephala, 113. 
Mantelli, 146. 

media, 81. 

muricata, 5. 

picta, 4, 64. 

pumila, 170. 

pygmea, 64. 


to 
Go 
Ne) 


ALPHABETICAL INDEX. 


Zamia Skinneri, 5, 64, 118, 170, 171. | Zamites carbonarius, 11, 12. 


Wallisii, 5. Carruthersi, 86-89, 234. 
Zamiee, 3, 6. Dunkerianus, 42. 
Zamiophyllum, 75, 77. epibius, 14. 

Buchianum, 40, 79, 232. falcatus, 56. 

Naumanni, 79, 80, 86, 90, 232. familiaris, 110. 
Zamiostrobus, 113, 114. gigas, 149, 150. 

elegans, 115. Gopperti, 80. 

mirabilis, 118. ; Goppertianus, 70. 

ovatus, 115. gramineus, 57, 72. 

Saportanus, 14, lanceolatus, 147. 
Zamites, 12, 32, 36, 75-90. Lyellianus, 91. 

acutipennis, 36. Mandelslohi, 62. 

eequalis, 92. Milleri, 79. 

affinis, 88. Montana, 94. 

arcticus, 15. Montanensis, 94. 

Bechii, 76. Planchardi, 12. 

borealis, 36. proximus, 37. 

brevifolius, 18, 56. regularis, 12. 

Brongniarti, 47. Schenkii, 81. 

Buchianus, 19, 21, 79-86, 88, 90, tenuinervis, 63, 65, 68, 69, 88. 

232, 234, 236. tertiarius, 14. 


Bucklandi, 56. 


PRINTED BY STEPHEN AUSTIN AND £0NS, HERTFORD. 


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d ’ 


EXPLANATION OF PLATES. 


Aut the figured specimens are preserved in the British Museum 
(Natural History), their registered numbers being quoted in 
square brackets. The figures are drawn natural size, except in 
a few cases where the enlargement is stated. With the excep- 
tion of those represented on Plates XIII. and XIV., the figured 


specimens are from the Rufford Collection. 


PLATE I. 


Fic. 1. Otozamites Goppertianus (Dunk.). Portion of a frond. Page 70. 
[V. 2123.] 


Fig. 2. Otozamites. Géppertianus (Dunk.). The lower and middle 
portion of a frond. P. 70. [V. 2360. ] 


Fia. 3. Otozamites Klipsteinii (Dunk.). Terminal portion of a frond. 
P64 [V. 2336.] 


Fia. 4. Otozamites Klipsteinii (Dunk.). P. 65. [V. 2745a. | 


Plate J 


BM.WEHALDEN PLANTS. 


West, Newman inp 


GM Woodward del et lith . 


Otozamites 


- 


PLATE II. 


* Fia. 1. Withamia armata, gen. et sp. nov. Single detached leaf, — 
showing well-marked flabellate venation. P. 177. [V. 2915.] 


* Fic. 2. Withamia armata, gen. et sp. nov. Axis bearing recurved 
spines and imperfect leaves in their axils, P.177. [V. 2134] _ 


Fic. 3. Dioonites Dunkerianus (Gopp.). Terminal portion of a frond. — 
P. 46. [V. 2823. ] 


Fic. 4. Otozamites, sp. Cf. O. Klipsteinit (Dunk.). Basal portion of — 
a young frond. P. 69. [V. 2734] 


* The name Saportaia was unfortunately printed on the plate before the — 
previous use of Saporte@a was discovered (p. 174). 


B.MWEALDEN PLANTS. 


West,Newman imp. 


G.M.Woodward del.etlith. 


ics Se 


tes. 4. Otozar 


100N1 


ia, 3.D 


LA ~Saporta 


Ine}, Il, 


IMiie, 


Ine Bt 


EWG 4 


Inve, 459, 


Fia. 6. 


KEL, Fla 


PEATE ALT: 


Zamites Buchianus (Ett.). Page 85. [V. 2262. ] 


Zamites Buchianus (Ett.). Apex of a single pinna.  P. 83. 
[V. 2363. | 


Zamites Buchianus (Ett.). Apex of a single pimna, P. 84. — 
[V. 2123¢.] 


Zamites Buchianus (Ett.). Portion of a large frond, showing 
manner of attachment of a pinna. P. 83. [V. 2227.] 


Zamites Buchianus (Ett.). Apical portions of two pinne. 
P. 82. [V. 2120.] 


Dioonites Dunkerianus (Gépp.). Portion of a frond, showing 
the form and manner of attachment of the pinne. P. 46. 
[V. 3218. ] 


Cycadites Saporte, sp. nov. Portion of a frond, showing the 
venation and arrangement of the pinne. P. 34. [V. 2124a.] 


Plate Il. 


B.MWEALDEN PLANTS. 


panto 
iene 
oe 


West Newman imp. 


Figs.1-5, Zamites. Fig. 6. Dioonites. Fis.7.Cycadites. 


G M Woodward del et lith. 


, 
4 oy = i 
Th Me 
is ‘ 
4 
¥ 1 
i ’ 
. ' 
i d 
¥ 
coat 
‘ } 
. 
‘ — 
j ‘ 
t 
t 
. 
\ 
\ 
t\ 
5 o 
~ 
F yen 
) 
1 nA 
. t 
a 
a 
x ry 


hee er i 
ad 


PLATE IV. 
Fig. 1. Zamites Buchianus (Ett.). Young frond. Page 84. [V. 2125d.] 


Fic. 2. Portion of a large frond, showing the position of the pinn in 
the rachis. P. 84. [V. 2125c. ] 


Fic. 3. Terminal portion of a very small frond. P. 84. [V. 2925. ] 


Fies. 4 and 5. Portions of a large frond, showing manner of attach- 
ment of the pinne. P. 84. [V. 21232. ] 


Plate IV. 


B.M.WEALDEN PLANTS. 


West, Newman imp 


GM. Woodward del.ethth 


a 


PLATE V. 


Fic. 1. Withamia armata, gen. et sp. nov. Axis with strongly recurved 
spines, with the basal portions of leaves in their axils. P. 178. 
[V. 2134a. ] 


Fia. 2. Cycadolepis. Single scale. P. 99. [V. 2929.] 


Plate V. 


BMWEALDEN PLANTS. 


ey 

5 

re] 

a 

= ® 
O, 
oO 
ro 
(eo) 
re} 
a0) 
O 
ea 
OO) 
3 
ol 
0) 
a 

cms) 

4 

q 9 

re VD 

E 

E 

3 

= 

al 

1) 


a 


PLATE VI. 
Fic. 1. Zamites, sp. P. 89. [V. 2743. ] 


Fies. 2 and 3. Zamites Carruthersi, sp. nov. The apical and basal 
portions of a pinna. P. 88. [V. 2123c.] 


Fic. 4. Zamites Carruthersi, sp. nov. Portion of a frond showing the 
manner of attachment and venation of the pinne. P. 88. 
[V. 2123d.] 


Fies. 5 and 5a. Cycadites Saporte, sp. nov. Portion of a large frond. 
In 5a part of a single pinna is slightly enlarged, showing 
the apex and single vein. P. 34. [V. 2797.] 


Fics. 6 and 6a. Cycadolepis, A detached scale ; in 6a the incurved 
distal margin is represented. P. 99. [V. 2699. } 


B.M WEALDEN PLANTS. Plate VI. 


GM Woodward del.et hth. West, Newman imp. 
Figs.1-4. Zamites. Figs. 5&5a.Cycadites Mg 6.Cycadolepis. 


Fie. 


Fic. 


Fic. 


ire: 


Fic. 


laurel, 


Fie. 


Fia. 


Fic. 


5. 


PLATE VII. 


of the longer and narrower type. 


O. Klipsteinii (Dunk.), var. superbus. 


. Otozamites Klipsteinti (Dunk.), var. longifolius. Single pinna 
P. 68. (See also Fig. 6.) 


[V. 2122.] 


Single pinna of the 


shorter and broader type, with lobed margin. P. 66. 


Single broad and short pinna. 


iP. 67. 


Part of a frond with smaller pinne. 


[V. 2122a.] 
[2912a. ] 


Cf, Pl. I. Fig. 4. P. 66. 
[V. 2126a.] 


Part of a frond showing smaller pinne. Cf. Pl. I. Fig. 4, 


and Pl. I. Fig. 3. P. 67. 


O. Klipsteinii var. longifolius. 
pinna. 7, Bigs I) P.1638: 


[V. 2745.] 


Single longer and narrower 


[V. 2122.] 


O. Klipsteinii var. superbus. Shorter pinna showing distinctly 


auriculate base. P. 67. 


[V. 2740.] 


Two pinne from a large frond. P. 66. [2126c. ] 


Large frond, probably not fully expanded; venation and 


form of the pinna distinct. 


B66: 


[V. 2170.] 


Plate Vili 


i 


> 


B.M.WEALDE 


p 


West, Newman im 


| 
H 


LC 


Ww ard pet 


M Woox 


G 


SNS Saaret 


. 
ie (cAine Oe Ba ee 


rT. i 
‘ oy 
ie 
a > , 
) of .: 
> ’ : ’ 
‘ € 
= , i 5 
Fi a 
. . : 
> t fy 
ft f 
1 " - 
¥ 
; ' Sr 
Py 
4 . 
r 20 . 
we 
ant / 
on ’ \ 
‘ x, ‘ a 
i 
i 5 
: ' 
* M : 
ra = 
ig 7 
z a 4 1 
j ’ 
y } 
\ 7 
i Ns 


’ me 
é = 

+ 

a, 

b, 4 

i ae 
% } 

a 4% uy os 
y 4 

* 

py { 
4 i 
4 * 
ey Fi j 
» { 
x 
iy 
i i ‘P 
e F ‘ 
i 
: ’ 
7, FE wala 
i ; 
. ww — 
i i i f 
. 

i ‘ 


PLATE VIII. 


Fic. 1. Zamites Buchianus (Ett.). Large frond, from a photograph 
by Mr. Gepp. One-sixth nat. size. P. 82. [V. 2120.] 


Fie. 2. Cycadites Saporte, sp. nov. From a photograph by Mr. Gepp. 
One-fourth nat. size. P. 33. [V. 2777.] 


Plate wine 


B. M. WEALDEN PLANTS. 


A. Gepp, phot. ad nat. 


Bice 23 CycAninEes: 


Fic. 1. ZAMITES. 


PLATE IX. 


Fic. 1. Androstrobus Nathorsti, sp. nov. Showing central axis and 
flattened sporophylls. P. 111. [V. 2810. ] 


Fig, 2. Androstrobus Nathorsti, sp. nov. End view of sporophylls. 
Pe. [V. 2811.] 


Fics. 3 and 4. Two sporophylls, slightly enlarged, showing pollen-sac 
impressions. P. 110, [V. 2701. ] 


Fig. 5. Seed-like body. Cf. Oolithes, Carruthers. P.107. [V. 2796a.] 


Fie. 6. Fittonia Ruffordi, sp. nov. Portion of a stem with well- 
preserved petiole bases. P. 133. [V. 2238. ] 


Fic. 7. Conites armatus, sp. nov. Badly preserved cone, showing 
the recurved spinous terminations of the scales. P. 222. 


[V. 2338.] 


a 
P. 


aml 
L 


BALDEN PLAN’ 


N 


BMV 


West, Newman imp 


el.etlith. 


M Woodward d: 


Fig.6.Fitt 


trobus. 


dros 


awal 


A 


ifs. 1-4/ 


ie 


Fia. 


PLATE X. 


Bennettites (Williamsonia) Carruthersi, sp. nov. 


1. Unexpanded fructification showing external bracts, and in 
Fig. la the reticulate lamellar projections from the inner 
face of a bract. Fig. 1b represents the conical basal cavity 
of the fructification. P. 157. [V. 3177.] 


Fic. 2. Basal view of a slightly larger fructification. P.159. [V. 3202.] 


Fig. 


Fie 


Fic. 


3. A small fructification in longitudinal section, showing a 
spherical boss at the base, and a few involucral bracts with 
the thread-like interstitial organs internal to the bracts. 
iP S2'60: [V. 21290.] 


. 4, The basal portion of a longer and expanded fructification, in 
the centre the base of the central boss, surrounded by the 
reticulately marked peripheral tissue. P. 159. [V. 3201.] 


5. Expanded bracts near the base of a fructification. P. 160. 
[V. 2129c.] 


Plate X. 


BMWEHALDEN PLANTS. 


West Newman i 


Qa 
oO 


Bennettite 


oe 


7 


iar ee 


eer 


Fic 


Fia. 


FIG. 


Gs 


PLATE XI. 


Benuettites (Williamsonia) Currutherst, sp. nov. 
’ 


. 1. Large expanded bracts below the base of a fructification, the 
base of the central boss, traces of interstitial organs, and 
the reticulate peripheral tissue. P. 161. 


the central boss of a fructification. 


P16). 


[V. 2793.] 


2. Bracts surrounding the conical cavity originally occupied by 


[V. 2129d.] 


3. Bennettites (Williamsonia) Carruthersi var. latifolius. Short 


and broad bracts seen from the under-side; at a lower level 
portions of the reticulations are shown. P. 163. [V. 21297.] 


4, The base of the central boss, surrounded by expanded 


interstitial organs, and below these, in one place, some 


of the reticulations are visible; and at a still lower level 


portions of short and broad bracts. 


P. 163. 


[V. 21296. ] 


BMWHALDEN PLANTS. 


GM Woodward del.et hth. 


Bennettites. 


West 


Newman imp 


analy 


PLATE XII. 


Fie. 1. Araucarites (Contes), sp. Central axis of cone and imperfect 


scales. P. 191. [V. 2180z. ] 
Fic. 2. Araucarites (Conites), sp. nov. Proximal ends of bracts. 
P19). [V. 2180.] 
Fic. 3. Nageiopsis, sp. A branched specimen with well-preserved 
leaves. P. 211. [V. 3190. ] 
Fig. 4. Cycadean trunk. P. 171. [V. 2350. ] 
Fia. 5. Cycadean trunk showing branching. P. 171. [V. 3162. ] 


Fic. 6. Dichopteris, sp. P. 184. [V. 3145. ] 


BM.WEHALDEN PLANTS. Plate XI] 


GM Wocdward del-etlith 


West, Newman imp 
Pigsdee, Conites: F'1g.3, Nageiopsis. 
Fig.6,Dichopteris. Figs 4-5,Trunci. 


Fig. 


Fia. 


Fia. 


Fia. 


Fic. 


Fia. 


PEACE, Sherr. 


. Macrozamia heteromera, Moore. Single branched pinna. P. 5. 


(Royal Gardens, Kew.) 


. Macrozamia heteromera. Single pinna. P.5. (Royal Gardens, 


Kew.) 


. Encephalartos Ghellinckii, Lem. Pp. 20, 22, 29, etc. (British 


Museum Herbarium.) 


. Encephalartos Ghellinckii, Lem. Single pinna from the under- 


side. P. 29. 


. Encephalartos Ghellinckii, Lem. Cross section of a pinna, 


showing the revolute edges. P. 29. 


. Encephalartos cycadifolius, Lehm. Portion of frond. P. 22. 


(Kew.) 


— 


BM.WHALDEN PLANTS. Plate XIIL. 


ate 


Se eo 
are aoe 
ee 
GM Weedward del et hth. West, Newman imp. 


Migs.a 6,linecephalartos. Figs.1-2,Macrozamia. 


nt 


PLATE XIV. 


Fia. 1. Cf. Becklesia anomala, gen. et sp. nov. P. 182. [V. 2608. ] 
. (Beckles Coll.) 


Fia. 2. Becklesia anomala, gen. et sp. nov. P. 179. [V. 2361a.] 


Fic. 3. Becklesia anomala, gen. et sp. nov. P. 179. [V. 23610. ] 


BM.WHALDEN PLANTS. Plate XIV. 


GM .Woodward delet lith. 


West Newman rmp. 


Becklesia. 


- 


PLATE XV. 
Bennettites, sp. 
P. 144. [V. 3177.] 
Fig. 1. Surface view of stem, showing position of the inflorescences. 
Fic. 2. Bracts of a single inflorescence. 


Fie. 3. Involucral bracts, and the central cavity with reticulate 
markings. 


Fic. 4. Wax cast of Fig. 3. 


Fies. 5-7. Enlarged portions of the surface shown in Fig. 3. P. 145. 


BMWEALDEN PLANTS. Plate XV. 


GM Woodward &J.Cameron del et hth. West, Newman intp 


Bennettites. 


Fig. 


Fic. 


Fie. 


Fic. 


Fic. 


janes, 


6. 


PLATE: XVI. 


. Pagiophyllum crassifolium (Schenk). Branch with leaves well 


preserved. P. 213. [V. 2803. ] 


. Pagiophyllum crassifolium (Schenk). Preservation less perfect, 


and leaves more indistinct than in Fig. 1. P. 213. 
[V. 2142q. ] 


. Cf. Sphenolepidium subulatum (Heer). P. 208. [V. 2140.] 


. Sphenolepidium Sternbergianum (Dunk.). Imperfect female 


cones, with expanded scales. P. 206. [V. 2311.] 


. Sphenolepidium Sternbergianum (Dunk.). Branches showing 


spreading leaves. P. 207. [V. 2139a. ] 


Sphenolepidium Sternbergianum (Dunk.). Leaves and leaf 
bases clearly shown. P. 206. [V. 2144. ] 


BM WEALDEN PLANTS. Plate XVI. 


GM Woodward &J.Cameron delet lith. West Newman imp. 


Figs.1—2,Pagiophyllum. Figs 3-6,Sphenolepidium. ! 


none =P hing az . Taal. ey re 


are ae er 
kA fol of ‘ : re A : 
WOT A Fy 
a CY la 
pie ‘ 
oa +e 
aay on i” 
Cy erst he. seine on 
oe eh a" y eek 
wy WN a > 
nt ee Aa an th, oe i . 
: ‘ LA . I 7 
Ate : fs - 
; any oy ar - 
ae 2 ? - ro r 
an a ‘ A bl g ’ 4 
i e : 
* * . >i 
Ae - 
. r . . ." ry 
a , > 
+ ~ 4 
¢ Peo at * te 
+i ‘oh “4 
‘ 
~% 
Sy 
ie | 
i} 
_ 


as 
? 2 { ‘To ee 
- ' 
ms j { , 
a 7 ea or ~ 
4 ee 
G Wc 6. yee 
ue: 
. ~ A MAG ? e 


a ‘ Lay in’ pened : 7 , Pal 
Bitar ys Si ee ei. 4 ; 14 Dit STOO 4 Le] os \ 


a ’ bonnes veh Ties Vaan 40 ee 
— af ie iL “he ag i ' 7 
_ ; Fy > 4 
pasite eget Te Ge Gh Renner at's Pay hohe ies = 
“a ty : 
ay: + of 
a ; % - ia ~ ; 
, 
« a 
te 
LOR 
-_— - 
; you 
e: ak Y 
us > 


PRATE: XVil. 


Fic. 1. Brachyphyllum spinosum, sp. nov. Large branched specimen 
with leaves, leaf-scars, and thorn-like branches. P. 216. 
[V. 2746.] 


Fic. 2. ? Brachyphyllum spinosum. Leaves slightly enlarged, showing 
the form and striate structure. P. 218. [V. 2296. ] 


Fic. 3. Brachyphyllum spinosum. Portion of a thick branch with 
leaf bases. P. 218. [V. 3180.] 


Fic. 4. Brachyphyllum spinosum. Portion of a decorticated axis with 
three branches. P. 217. [V. 2240. ] 


Fies. 5 and 5a. Brachyphyllum spinosum. Branch showing decorti- 
cated axis. P. 218. [V. 2750.] 


Fie. 6. Brachyphyllum spinosum. One-third nat. size. Decorticated 
specimen showing the spinous branches. P. 217. [V. 21365.] 


Fia. 7. Sphenolepidium Kurrianum (Schenk). Cluster of female cones. 
P. 203. [V. 2316. ] 


Fias. 8 and 8a. Sphenolepidium Kurrianum (Schenk). Single cone 
more perfectly preserved. P. 203. [V. 2213.] 


Fic. 9. Brachyphyllum obesum, Heer. Small twig. P. 221. [V.3348.] 


Plate XVIL. 


~ GM Woodward 


West, Newman imp 


IOhiag, 


PLATE XVIII. 


Fig. 1. Sphenolepidium Kurrianum (Schenk). Large specimen with 
well-preserved leaves. P. 203. [V. 2316d. ] 


Fic. 2. Pinites Solmsi, sp. nov. Branches with leaf bases and long 
needles. P. 1977. [V. 2169. ] 


Fic. 3. Pinites Solmsi, sp. nov. Female cone with short branch and 
narrower needles. P. 197. [V. 2255. ] 


B.M.WEHALDEN PLANTS. Plate XVIII. 


GM Woodward del et hth. West,Newmanimp. 


Fig.l. Sphenolepidium. Figs.2&3.Pinites 


PLATE XIX. 
Pinites Solmst, sp. nov. 


Fig. 1. Branch with well-preserved leaf bases, and cones with un- 
expanded scales. P. 196. [V. 2146. |] 


Fic. 2. Branch and cone with partially expanded scales. P. 198. 
 [V. 2147.] 


Fie. 3, Branch bearing two female cones, and in the upper portion 
small indistinct structures. P. 197. [V. 2146a. ] 


Fia, 4. Cone with partially expanded scales. P. 197. [V. 2147a.] 


Plate ZIX 


B.M.WEALDEN PLANTS. 


ewman Imp. 


con West N 
Picante s: 


GM Woodward del.et hth 


Jeng, il, 


Fie. 2. 


PLATE XX. 


Brachyphyllum obesum, Heer. Twig showing manner of 
branching. P. 220. [V. 2137.] 


Bruchyphyllum obesum. Stouter branch with well-preserved 
leaves, .P. 221. [V. 2137a.] 


Pagiophyllum, sp. Twig with broad leaves. P. 213. [V. 2288.] 


Brachyphyllum obesum. Comparatively thick branch. P. 221. 
[V. 2337. ] 


Pinites Carruthersi, Gard. Detached female cone. P. 195. 
[V. 2611. ] 


Thuites valdensis, sp. nov. Twig with distinctly preserved 
branches and leaves. P. 209. [V. 2138. ] 


B.M.WEALDEN PLANTS. Plate XX. 


GM.Woodward &J.Cameron del et ith. West, Newman imp. 


Pigs.1.2&4. Brachyphyllum. Fig.3.Pagiophyllum. 
. 5 Pimaites- 6. Thuites. 


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Birds, in the Collection of the British Museum. Cichlo- 
morphe: Part V., containing the families Paride and 
Laniide (Titmice and Shrikes) ; and Certhiomorphe 
(Creepers and Nuthatches). By Hans Gadow, M.A., Ph.D. 
Pp. xiii., 386. Woodeuts and 9 coloured Plates. [With 
Systematic and Alphabetical Indexes.] 1883, 8vo. 17s. 


BRITISH MUSEUM (NATURAL HISTORY). 3 


Catalogue of the Birds in the British Museum —continued. 


Vol. TX. Catalogue of the Passeriformes, or Perching Birds, 
in the Collection of the British Museum. Cinnyrimorphe, 
containing the families Nectariniide and Meliphagide 
(Sun Birds and Honey-eaters). By Hans Gadow, M.A., 
Ph.D. Pp. xii., 310. Woodcuts and 7 coloured Plates. 
[With Systematic and Alphabetical Indexes.] 1884, 
8vo. 14s. 

Vol. X. Catalogue of the Passeriformes, or Perching Birds, 
in the Collection of the British Museum. /ringilliformes : 
Part L, containing the families Dicwide, Hirundinide, 
Ampelide, Mniotiltid, and Motacillide. By R. Bowdler 
Sharpe. Pp. xiii.,682.. Woodcuts and 12 coloured Plates. 
[With Systematic and Alphabetical Indexes.] 1885, 
8vo. 12. 2s. 

Vol. XI. Catalogue of the Passeriformes, or Perching Birds, 
in the Collection of the British Museum. /ringilliformes : 
Part IL., containing the families Corebide, Tanagride, 
und Icteride. By Philip Lutley Sclater, M.A., F.R.S. 
Pp. xvii, 431. [With Systematic and Alphabetical 
Indexes.| Woodcuts and 18 coloured Plates. 1886, 
8vo. 1d. 

Vol. XII. Catalogue of the Passeriformes, or Perching 
Birds, in the Collection ofthe British Museum. Fringilii- 
formes; Part III., containing the family Fringillide. By 
R. Bowdler Sharpe. Pp. xv., 871. Woodcuts and 16 
coloured Plates. [With Systematic and Alphabetical 
Indexes.] 1888, 8vo. 1é. 8s. 

Vol. XIII. Catalogue of the Passeriformes, or Perching 
Birds, in the Collection of the British Museum. Sturni- 
formes, containing the families Artamide, Sturnide, 
Ploceide, and Alaudide. Also the families Atrichiidse 
and Menuride. By R. Bowdler Sharpe. Pp. xvi., 701. 
Woodecuts and 15 coloured Plates. [With Systematic 
and Alphabeticai Indexes.] 1890, 8vo., 1d. 8s. 


Vol. XIV. Catalogue of the Passeriformes, or Perching 
Birds, in the Collection of the British Museum. Oligo- 
myode, or the families Tyrannide, Oxyrhamphide, Pipride, 
Cotingide, Phytotomide, Philepittide, Pittide, Xenicide, 
and Eurylemide. By Philip Lutley Sclater, M.A., 
F.R.S. Pp. xix., 494. Woodcuts and 26 coloured Plates. 
[With Systematic and Alphabetical Indexes.] 1888, 
Svo. 1. 4s. 

Vol. XV, Catalogue of the Passeriformes, or Perching 
Birds, in the Collection of the British Museum. T'racheo- 
phone, or the families Dendrocolaptide, Formicariide, 
Conopophagidz, and Pteroptochide. By Philip Lutley 
Sclater, M.A., F.R.S. Pp. xvii., 371. Woodcuts and 20 
coloured Plates. [With Systematic and Alphabetical 
Indexes.| 1890, 8vo. 1/. 

A 2 


4 LIST OF PUBLICATIONS OF ‘THE 


Catalogue of tne Birds in the British Museum—continued. 

Vol. XVI. Catalogue of the Picariz in the Collection of the 
British Museum. OUpupe and Trochili, by Osbert Salvin. 
Coracia, of the families Cypselide, Caprimulgide, Podar- 
gide, and Steatornithide, by Ernst Hartert. Pp. xvi. 
703. Wocdcuts and 14 coloured Plates, [With Systematic 
and Aiphabetical Indexes.] 1892, 8vo. 12, 16s. 

Vol. XVII. Catalogue of the Picaria# in the Collection of 
the British Museum. Coracie (contin.) and Halcyones, 
with the families Leptosomatide, Coraciide, Meropide, 
Alcedinide, Momotidex, Totida, and Coliida, by R. Bowdler 
Sharpe. Bucerotes and Trogones, by W. R. Ogilvie 
Grant. Pp. xi., 522. Woodeuts and 17 coloured Plates. 
[With Systematic and Alphabetical Indexes.] 1592, 
Svo. 1é. 10s. 

Vol. XVIII. Catalogue of the Picariz in the Collection of 
the British Museum. Scansores, containing the family 
Picide. By Edward Hargitt. Pp. xv., 597. Woodcuts 
and 15 coloured Plates. [With Systematic and Alpha- 
betical Indexes.| 1890, 8vo. 12. 6s. 

Vol. XIX. Catalogue of the Picarie in the Collection of 
the British Museum. Scansores and Coccyges : contain- 
ing the families Rhamphastide, Galbulide, and Bucconide, 
by P. L. Sclater ; and the families Indicatoride, Capitonide, 
Suculide, and Musophagide, by G. E. Shelley. Pp. xii, 
484: 13 coloured Plates. [With Systematic and Alpha- 
betical Indexes.| 1891, 8vo. 1/. 5s. 


Vol. XX. Catalogue of the Psittaci, or Parrots, in the 
Collection of the British Museum. By 'T. Salvadori. 
Pp. xvii., 658: woodcuts and 18 coloured Plates. [With 
Systematic and Alphabetical Indexes.] 1891, 8vo. 1/. 10s. 


Vol. XXI. Catalogue of the Columbe, or Pigeons, in the 
Collection of the British Museum. By T. Salvadori. 
Pp. xvii., 676: 15 coloured Plates. [With Systematic and 
Alphabetical Indexes.] 1893, 8vo. 1d. 10s. 


Vol. XXII. Catalogue of the Game Birds (Pterocletes, 
Galline, Opisthocomi, Hemipodii) in the Collection of 
the British Museum. By W. R. Ogilvie Grant. Pp. xvi., 
585: 8 coloured Plates. [With Systematic and Alpha- 
betical Indexes.| 1893, 8vo. 1/. 6s. 


Vol. XXIII. Catalogue of the Fulicaria (Rallide and 
Heliornithidz) and Alectorides (Aramid, Eurypygide, 
Mesitide, Rhinochetide, Gruide, Psophiide, and Otididz) 
in the Collection of the British Museum. By R-. Bowdler 
Sharpe. Pp. xiii, 353: 9 coloured Plates. [With Syste- 
matic and Alphabetical Indexes.] 1894, 8vo. 20s. 

List of the Specimens of Birds in the Collection of the British 
Museum. By George Robert Gray :— 

Part IJII., Section I. Ramphastide. Pp. 16. [With 

Index.} 1855, 12mo. 6d. 


BRITISH MUSEUM (NATURAL HISTORY). 5 


List of Specimens of Birds in the British Museum—continued. 
Part ILI., Section IT. Psittacide. Pp. 110. [With Index. ] 
1859, 12mo. 2s. 


Part III., Sections III. and IV. Capitonide and Picide. 
Pp. 137. [With Index.] 1868, 12mo. ls. 6d. 


PartIV. Columbe. Pp.73. [With Index.] 1856, 12mo. 
Ls. 9d. 


Part V. Galline.- Pp. iv., 120. [With an Alphabetical 
Index.] 1867, 12mo. 1s. 6d. 


Catalogue of the Birds of the Tropical Islands of the Pacific 
Ocean in the Coilection of the British Museum. By George - 
Robert Gray, F.L.S., &e. Pp. 72. [With an Alphabetical 
Index.] 1859, 8vo. 1s. 6d. 


REPTILES. 


Catalogue of the Tortoises, Crocodiles, and Amphisbenians in the 
Collection of the British Museum. By Dr. J. E. Gray, 
F.R.S., &ce. Pp. viii., 80. [With an Alphabetical Index. ] 
1844, 12mo. Is. 


Catalogue of Shield Reptiles in the Collection of the British 
Museum. By John Edward Gray, F.R.S., &c. :— 
Appendix. Pp. 28. 1872, 4to. 2s. 6d. 


Part II. Emydosaurians, Rhynchocephalia, and Amphis- 
benians. Pp. vi., 41. 25 Woodcuts. 1872, 4to. 3s. 6d. 


Hand-List of the Specimens of Shield Reptiles in the British 
Museum: By Dr. J. EH. Gray, F.R.S., F.L.S., &c. Pp. iv., 
124. [With an Alphabetical Index.] 1873, 8vo. 4s. 

Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles 
in the British Museum (Natural History). New Edition. By 
George Albert Boulenger. Pp. x., 311. 73 Woodcuts and 6 
Plates. [With Systematic and Alphabetical Indexes.] 1889, 
8vo. lis. 

Catalogue of the Specimens of Lizards in the Collection of the 
British Museum. By Dr. J. HE. Gray, F.R.S., &e. Pp. xxviii., 
2389. [With Geographic, Systematic, and Alphabetical Indexes. | 
1845, 12mo. 3s. 6d. 


Catalogue of the Lizards in the British Museum (Natural His- 
tory). Second Edition. By George Albert Boulenger :— 

Vol. I. Geckonide, Kublepnaride, Uroplatide, Pygopodide, 
Agamide. Pp. xii., 436. 32 Plates. [With Systematic 
and Alphabetical Indexes.] 1885, 8vo. 20s. 

Vol. Il. Iguanide, Xenosauride, Zonuride, Anguide, 
Anniellide, Helodermatide, Varanidx, Xantusiide, Teiide, 
Amphisbenide. Pp. xiii, 497. 24 Plates. [With 
Systematic and Alphabetical Indexes.] 1885, 8vo, 26s, 


6 LIST OF PUBLICATIONS OF THE 


Catalogue of the Lizards in the British Museum—continued. 

Vol. III. Lacertide, Gerrhosauride, Scincide, Anelytro- 
pide, Dibamide, Chameleontide. Pp. xi, 575. 40 
Plates. [With a Systematic Index and an Alphabetical 
Index to.the three volumes.| 1887, 8vo. 1. 6s. 

Catalogue of the Snakes in the British Museum (Natural History). 
By George Albert Boulenger, F.R.S. :— 


Vol..I., containing the families Typhlopide, Glauconiide, 
Boide, Ilysiide, Uropeltide, Xenopeltide, and Colubride 
aglypha, part. Pp. xiii, 445: 26 Woodcuts and 28 
Plates. [With Systematic and Alphabetical Indexes. | 
1893, 8vo. IZ. Ls. 


Vol. IL., containing the conclusion of the Colubride aglyphe. 
Pp. xi. 382: 25 Woodcuts and 20 Plates. [With 
Systematic and Alphabetical Indexes.| 1894, 8vo. 17s. 6d. 


Catalogue of Colubrine Snakes in the Collection of the British 
Museum. By Dr. Albert Giinther. Pp. xvi, 281. [With 
Geographic, Systematic, and Alphabetical Indexes.] — 1858, 
12mo. 4s. : 


BATRACHIANS. 


Catalogue of the Batrachia Salientia in the Collection of the 
British Museum. By Dr. Albert Giinther. Pp. xvi., 160. 12 
Plates. [With Systematic, Geographic, and Alphabetical 
Indexes.] 1858, 8vo. 6s. 


Catalogue of the Batrachia Gradientia, s. Caudata, and Batrachia 
Apoda in the Collection of the British Museum. Second 
Edition. By George Albert Boulenger. Pp. viii, 127. 9 
Plates. {With Systematic and Alphabetical Indexes.] 1882, 
Svo. 9s. 


FISHES. 


Catalogue of the Fishes in the Collection of the British Museum. 
By Pr. Albert Giinther, F.R.S., &e. :— 

Vol. VII. Physostomi (Heterophygii, Cyprinide, Gono- 
rnynchide, Hyodontide, Osteoglosside, Clupeide, Chiro- 
centride, Alepocephalide, Notopteride, Halosauridz). 
Pp. xx., 512. Woodcuts. [ With Systematic and Alpha- 
betical Indexes.] 1868, 8vo. 8s. 


Vol. VIII. Physostomi (Gymnotide, Symbranchide, Mure- 
nide, Pegaside), Lophobranchii, Plectognathi, Dipnoi, 
Ganoidei, Chondropterygii, Cyclostomata, Leptocardii. 
Pp. xxv., 549. [With Systematic and Alphabetical 
Tndexes.| 1870, 8yo, 8s. 6d, 


BRITISH MUSEUM (NATURAL HISTORY). 7 


Catalogue of the Fishes in the British Museum. Second edition. 
Vol. I. Catalogue of the Perciform Fishes in the British 
Museum. Vol. I. containing the Centrarchide, Percide, and 
Serranide (part), By George Albert Boulenger, F.R.S. Pp. 
xix., 394. Woodeuts and 15 plates. [With Systematic and 
Alphabetical Indexes.] 1895, 8vo. 15s. 

List of the Specimens of Fish in the Collection of the British 
Museum. Part J. Chondropterygii. By J.E. Gray. Pp.x., 
160. 2 Plates. [With Systematic and Alphabetical Indexes. ] 
1851, 12mo. 3s. 


Catalogue of Fish collected and described by Laurence Theodore 
Gronow, now in the British Museum. Pp. vi., 196. [With a 
Systematic Index.] 1854, 12mo. 3s. 6d. 


Catalogue of Lophobranchiate Fish in the Collection of the British 
Museum. By J. J. Kaup, Ph.D., &c. Pp. iv., 80... 4 Plates, 
| With an Alphabetical Index.] 1856, 12mo, 2s. 


MOLLUSCA. 


Guide to the Systematic Distribution of Mollusea in the British 
Museum. Part I. By John Edward Gray, Ph.D., F.R.S., 
&e. Pp. xii., 230. 121 Woodcuts. 1857, 8vo. 5s. 


List of the Shells of the Canaries in the Collection of the British 
Museum, collected by MM. Webb and Berthelot. Described 
and figured by Prof. Alcide D’Orbigny in the “ Histoire 
Naturelle des Iles Canaries.” Pp. 32. 1854, 12mo, Is. 


List of the Shells of Cuba in the Collection of the British Museum, 
collected by M. Ramon de la Sagra. Described by Prof. Alcide 
@ Orbigny in the * Histoire de Pile de Cuba.” Pp. 48. 1854, 
12mo. ls. 


List of the Shells of South America in the Collection of the British 
Museum. Collected and described by M. Alcide D’Orbigny in 
the “ Voyage dans PAmériqne Méridionale.” Pp. 89. 1854, 
t2mo. 2s. 


Catalogue of the Collection of Mazatlan Shells in the British 
Museum, collected by Frederick Reigen. Described by Philip 
P. Carpenter. Pp. xvi., 532. 1857, 12mo. 8s. 


List of Mollusca and Shells in the Collection of the British 
Museum, collected and described by MM. Eydoux and Souleyet 
in the “ Voyage autour du Monde, exécuté pendant les années 
“ 1836 et 1837, sur la Corvette ‘ La Bonite,’” and in the 
* Flistoire naturelle des Mollusques Ptéropodes.” Par MM. 
P. C. A. L. Rang et Souleyet. Pp. iv., 27... 1855, 12mo. 8d. 

Catalogue of-the Phaneropneumona, or Terrestrial Operculated 
Mollusca, i in the Collection of the British Museum. By Dr. L. 
Pfeiffer. Pp. 324. [With an Alphabetical Index. ] 1852, 


J2mo. 5s, 


8 LIST OF PUBLICATIONS OF THE 


Nomenclature of Molluscous Animals and Shells in the Collection 
of the British Museum. PartI. Cyclophoride. Pp. 69. [With 
an Index.} 1850, 12mo. Is. 6d. 


Catalogue of Pulmonata, or Air Breathing Mollusca, in the Col- 
lection of the British Museum. PartI. By Dr. Louis Pfeiffer. 
Pp. iv., 192. Woodeuts. 1855, 12mo. 2s. 6d. 


he ae of' the Auriculide, Proserpinidw, and Truncatellide in 
the Collection of the British Museum. By Dr. Louis Pfeiffer. 
Pp. iv., 150. Woodecuts. 1857, 12mo. 1s. 9d. 


List of i Mollusca in the Collection of the British Museum. By 
John Edward Gray, Ph.D., F.R.S., &c. 
Part I. Volutide. Pp. 28. 1855, 12mo. 6d. 
Part II. Olivide. Pp. 41. 1865, 12mo. Is. 


Catalogue of the Conchifera, or Bivalve Shells, in the Collection 
of the British Museum. By M. Deshayes :— 
: Part I. Veneride, Cyprinide, Glauconomide, and Petri- 
colade. Pp. iv., 216. 1853, J2mo. 3s. 
Part IJ.  Petricolade (concluded); Corbiculade. Pp. 
217-292. [With an Alphabetical Index to the two 
parts.| 1854, 12mo. 6d. 


BRACHIOPODA. 


Catalogue of Brachiopoda Ancylopoda or Lamp Shells in the 
Collection of the British Museum. [J/ssued as ‘* Catalogue of 
the Mollusca, Part TV.”] Pp. iv., 128 25 Woodcuts. | With 
an Alphabetical Index.| 1858, 12mo. 3s. 


POLYZOA. 


Catalogue of Marine Polyzoa in the Collection of the British 
Museum. Part III. Cyclostomata. By George Busk, F.R.S. 
Pp. vili., 39. 38 Plates. [With a Systematic Index.] 1875, 
8vo. 5s. 


CRUSTACEA. 


Catalogue of Crustacea in the Collection of the British Museum. 
Part I. Leucosiade. By Thomas Bell, V.P.R.S., Pres. L.S., 
&e. Pp. iv., 24. 1855, 8vo. 6d. 


Catalogue of the Specimens of Amphipodous Crustacea in the 
Collection of the British Museum. By C. Spence Bate, F.R.S., 


&e. Pon, iv., 399. 58 Plates, [With an Alpbabetical Index. | 
1862, 8vc, I. Ss. 


BRITISH MUSEUM (NATURAL HISTORY). 9 


ARACHNIDA. 


Descriptive Catalogue of the Spiders of Burma, based upon the 
Collection made by Eugene W. Oates and preserved in the 
British Museum. By T. Thorell. Pp. xxxvi., 406. [With 
Systematic List and Alphabetical Index.] 1895, 8vo. 10s. 6d. 


MYRIOPODA. 


Catalogue of the Myriapoda in the Collection of the British 
Musum. By George Newport, F.R.S., P.E.S., &e. Part I. 
Chilopoda. Pp. iv., 96. [With an Alphabetical Index. ] 
1856, 12mo. 1s. 9d. 


INSECTS. 
Coleopterous Insects. 


Nomenclature of Coleopterous Insects in the Collection of the 
British Museum :-— 

Part IV. Cleride. By Adam White. Pp. 68. [With 
Index.] 1849, 12mo. ls, 8d. 

Part V. Cucujide, &c. By Frederick Smith. [Also issued 
as “ List of the Coleopterous Insects. Part ].”] Pp. 25, 
1851, 12mo. 6d. 

Part VI. Passalide. By Frederick Smith. Pp. iv., 23. 
1 Plate [With Index.] 1852, 12mo. 8d. 

Part VII. Longicornia, I. By Adam White. Pp. iv., 174. 
4 Plates. 1853, 12mo. 2s. 6d. 

Part VILI.-Longicornia, Il. By Adam White. Pp. 237. 
6 Plates. 1855, 12mo. 3s. 6d. 

Part IX. Cassidide. By Charles H. Boheman, Professor of 


Natural History, Stockholm. Pp. 225. [With Index. ] 
1856, 12mo. 3s. 


Illustrations of Typical Specimens of Coleoptera in the Collection 
of the British Museum. Part I. Lycide. By Charles Owen 
Waterhouse. Pp. x., 83. 18 coloured Plates. [With Syste- 
matic and Alphabetical Indexes.| 1879, 8vo. 16s. 


Catalogue of the Coleopterous Insects of Madeira in the Collection 
of the British Museum. By 'T. Vernon Wollaston, M.A., F.L.S. 
Pp. xvi., 254: 1 Plate. [With a Topographical Catalogue and 
an Alphabetical Index.] 1857, 8vo. 3s. 


Catalogue of the Coleopterous Insects of the Canaries in the Collec- 
tion of the British Museum. By T. Vernon Wollaston, M.A., 
F.L.S. Pp. xiti., 648. [With ‘Topographical and Alphabetical 
Indexes.| 1864, 8vo. 10s. 6d. 

Catalogue of Halticide in the Collection of the British Museum. 
By the Rev. Hamlet Clark, M.A., F.L.S. Physapodes and 
(Edipodes. Part I. Pp. xii., 301. Frontispiece and 9 Plates, 
1860, 8vo. 7s. 


10 LIST OF PUBLICATIONS OF THE 


Catalogue of Hispide in the Collection of the British Museum. 
By Sloseph 'S. Baby. MB.Ss gcc.’ Part: 54 Pps xy lg) we 
Plates. [With an Alphabetical Index.] 1858, 8vo. 6s. 


Hymenopterous Insects. 


List of the Specimens of Hymenopterous Insects in the Collection 
of the British Museum. By Francis Walker, F.L.S.:— 

Part If. Chalcidites. Additional Species. Appendix. 
Pp. iv., 99-237. 1848, 12mo. 2s. 

Catalogue of Hymenopterous Insects in the Collection of the 
British Museum. By Frederick Smith. 12mo. :— 

Part I. Andrenide and Apide. Pp. 197. 6 Plates. 1853, 
2s. 6d. 

Part II. Apide. Pp. 199-465. 6 Plates. [With an 
Alphabetical Index.| 1854, 6s. 

Part III. Mutillide and Pompilide. Pp. 206. 6 Plates. 
1855, 6s. 

Part IV. Sphegide, Larride, and Crabronide. Pp. 207- 
497. 6 Plates. [With an Alphabetical Index.] 1856, 
6s. 

Part V. Vespide. Pp. 147. 6 Plates. [With an Alpha- 
betical Index.] 1857, 6s. 

Part VI. Formicide.’ Pp. 216.14 Plates. [With an 
Alphabetical Index.] 1858, 6s. 

Part VII. Dorylide and Thynnide. Pp. 76. 3 Plates. 
[With an Alphabetical Index.] 1859, 2s. 

Descriptions of New Species of Hymenoptera in the Collection 
of the British Muserm. By Frederick Smith. Pp. xxi., 240. 
[ With Systematic and Alphabetical Indexes.] 1879, 8vo. 10s. 


List of Hymenoptera, with descriptions and figures of the Typical 
Specimens in the British Museum. Vol. I., Tenthredinide and 
Siricide. By W. F. Kirby. Pp. xxviii., 450. 16 Coloured 
Plates. [With Systematic and Alphabetical Indexes.] 1882, 
8vo. 1/. 18s. 


Dipterous Insects. 


List. of the Specimens of Dipterous Insects in the Collection of 
the British Museum. By Francis Walker, F.L.S. 12mo. :— 
Part IV. Pp. 689-1172. [With an Index to the four parts, 
and an Index of Donors.] 1849. 6s. 
Part VII. Supplement III. Asilide.. Pp. ii.. 507-775. 
1855. 3s. 6d. 


Lepidopterous Insects. 


Tllustrations of Typical Specimens of Lepidoptera Heterocera in 
the Collection of the British Museum :— 
Part Il. By Arthur Gardiner Butler. Pp. xviii, 82. 
41-60 Coloured Plates, [With a Systematic index. ] 
1879, 4to. 22, 10s, 


BRITISH MUSEUM (NATURAL HISTORY). 11 


Illustrations of Rypical Specimens of Lepidoptera Heterocera 
—continued. 
Part V. By. Arthur Gardiner Butler. Pp. xii, 74. 
78-100 Coloured Plates. [With a Systematic Index. ] 
1881, 4to. 27. 10s. 


Part VI. By Arthur Gardiner Butler. Pp. xv., 89. 
101-120 Coloured Plates. [With a Systematic Index. | 
1886, 4to. 24. 4s. 


Part VII. By. Arthur Gardiner Butler. Pp. iv., 124. 
121-138 Coloured Plates. [With a Systematic List. ] 
1889, 4to. 2/. 


Part VIII. The Lepidoptera Heterocera of the Nilgiri 
District. By George Francis Hampson. Pp. iv., 144. 
139-156 Coloured Plates. | With a Systematic List. | 
1891, 4to. 22. 


Part IX. The Macrolepidoptera Heterocera of Ceylon. By 
George Francis Hampson. Pp. v., 182. 157-176. 
Coloured Plates. [With a General Systematic List of 
Species collected in, or recorded from, Ceylon.j| 18938, 
Ato. 21. 2s. 

Catalogue of Diurnal Lepidoptera of the family Satyride in the 
Collection of the British Museum. By Arthur Gardiner Butler, 
F.L.S., &. Pp. vi. 211. 5 Plates. [With an Alphabetical 
Index.] 1868, 8vo. 5s. 6d. 


Catalogue of Diurnal Lepidoptera described by Fabricius in the 
Collection of the British Museum. By Arthur Gardiner Butler, 
F.L.S., &c.. Pp. iv., 303. 3 Plates. 1869, 8vo. 7s. 6d. 


Specimen of a Catalogue of Lyczenid in the British Museum. By 
W.C. Hewitson. Pp. 15. 8 Coloured Plates, 1862, 4to. 1/. 1s. 


List of Lepidopterous Insects in the Collection of the British 
Museum. Part J. Papilionide. By G. R. Gray, F.L.S. 
Pp. 106. [With an Alphabetical Index.] 1856, 12mo. 2s. 


List of the Specimens of Lepidopterous Insects in the Collection 
of the British Museum. By Francis Walker. 12mo. :— 
Part VI. Lepidoptera Heterocera. Pp. 1258-1507. 1855, 


3s. 6d. 
Part X. Noctuide. Pp. 253-491. 1856, 3s. 6d. 
Part XIJT. ———— Pp. 765-982. 1857, 3s. 6d. 
Part XIII. —————_ Pp. 983-1236. 1857, 3s. 6d. 
Part XIV. ———— Pp. 1237-1519. 1858, 4s. 6d. 
Part XV. Pp. 1520-1888. [With an Alpha- 


betical Index to Parts IX.-XV.] 1858, 4s. 6d. 
Part XVI. Deltoides. Pp. 253. 1858, 3s. 6d. 
Part XTX. Pyralides. Pp. 799-1036. [With an Alpha- 
betical Index to Parts XVI.-XIX.] 1859, 3s. 6d. 
Part XXI. Geometrites. Pp. 277-498. 1860, 3s. 
Part XXII. Pp. 499-755. 1861, 3s. 6d. 
Part XXIII. ———-———_ Pp. 756-1020, 1861, 3s. 6d. 


12 LIST OF PUBLICATIONS OF THE 


List of Specimens of Lepidopterous Insects—continued. 

Part XXIV. —— Pp. 1021-1280. 1862, &s. 6d. 

Part XXV. —— Pp. 1281-1477. 1862, 3s. 

Part XXVI. Pp. 1478-1796. {With an 
Alphabetical Index to Parts XX.-XXVI.] 1862, 4s. 6d. 

Part XXVIUI. Crambites and Tortricites. Pp. 1-286. 
1863, 4s. 

Part XXVIII. Tortricites and Tineites. Pp. 287-561. 
1863, 4s. 

Part XXIX. Tineites. Pp. 562-835. 1864, -ts. 

Part XXX. — Pp. 836-1096. [With an Alpha- 
betical Index to Parts XXVIL-XXX.] 1864, 4s. 

Part XXXII. Supplement. Pp. 1-321. 1864, ds. 


Part XXXII. Part 2. Pp. 322-706. 1865, 
as. 

Part XXXIIT. —— Part 3. Pp. 707-1120. 1865, 
6s. 

Part XXXIV. —— Part 4. Pp. 1121-1533. 1865, 
5s. 6d. 


Part XXXV. Part 5. Pp. 1534-2040. [ With 
an Alphabetical Index to Parts XXXI-XXXYV.] 1866, 
7s. 


Neuropterous Insects. 


Catalogue of the Specimens of Neuropterous Insects in the Collec- 
tion of the British Museum. By Francis Walker. 12mo. :— 
Part I. Phryganides—Perlides. Pp. iv., 192. 1852, 2s. 6d. 
Part 1. Sialide—Nemopterides. Pp. 1., 193-476. 1853, 
3s. 6d. 
Part ilf. Termitide—Ephemeride. Pp. ii., 477-585. 1853, 
ls. 6d. 
Catalogue of the Specimens of Neuropterous Insects in the Col- 
lection of the British Museum. By Dr. H. Hagen. Part I. 
Termitina. Pp. 384. 1858, 12mo. 6d. 


Orthopterous Insects. 


Catalogue of Orthopterous Insects in the Collection of the British 
Museum. Part 1. Phasmide. By Jobn Obadiah Westwood, 
F.L.S., &. Pp. 195. 48 Plates. [With an Alphabetical 
Index.] 1859, 4to. 3Z. 

Catalogue of the Specimens of Blattariz in the Collection of the 
British Museum. By Francis Walker, F.L.S., &. Pp. 259. 
[With an Alphabetical Index.] 1868, 8vo. 5s. 6d. 

Catalogue of the Specimens of Dermaptera Saltatoria [Part I.] 
and Supplement to the Blattariz in the Collection of the British 
Museum. Gryllide. Blattarie. Locustide. By Francis 
Walker, F.L.S., &c. Pp. 224, [With an Alphabetical Index. | 
1869, vo, 5s. 


BRITISH MUSEUM (NATURAL HISTORY). 13 


Catalogue of the Specimens of Dermaptera Saltatoria in the 
Collection of the British Museum. By Francis Walker, 
F.L.S., &e.— 


Part II. Locustide (continued). Pp. 225-423. 
Alphabetical Index.] 1869, 8vo. 4s. 6d. 


Part III. Locustide (continued).—Acrididz. Pp. 425-604. 
[ With an Alphabetical Index.] 1870, 8vo. 4s. 


Part IV. Acridide (continued). Pp. 605-809. 
Alphabetical Index.] 1870, 8vo. 6s. 


Part V. ‘Tettigide.—Supplement to the Catalogue of Blat- 
tarie.—Supplement to the Catalogue of Dermaptera 
Saltatoria (with remarks on the Geographical Distribution 
of Dermaptera). Pp. 811-850; 43; 116. [With Alpha- 
betical Indexes.| 1870, 8vo. 6s. 


[With an 


[ With an 


Hemipterous Insects. 


List of the Specimens of Hemipterous Insects in the Collection of 
the British Museum. By W. S. Dallas, F.L.S. Part II. Pp. 
369-590. Plates 12-15. 1852, 12mo. 4s. 

Catalogue of the Specimens of Heteropterous Hemiptera in the 
Collection of the British Museum. By Francis Walker, F.L.S., 


&e. 8vo.:— 
Part I. Scutata. Pp. 240. 1867. 5s. 
Part II. Scutata (continued). Pp. 241-417. 1867. 4s. 


Part III. Pp. 418-599. [With an Alphabetical Index to 
Parts I., II., IlI., and a Summary of Geographical 


Distribution of the Species mentioned.|] 1868. 4s. 6d. 
Part IV. Pp. 211. [Alphabetical Index.] 1871. 6s. 
Part V.. Pp. 202. Stead 1872. 5s. 
Part VI. Pp. 210. —- 1873. 5s. 
Part VII. Pp. 2138. —— 1873. 6s. 
Part VIII. Pp. 220. ——— 1878. 6s. 6d. 


Homopterous Insects. 


List of the Specimens of Homopterous Iusects in the Collection of 
the British Museum. By Francis Walker. Supplement. Pp. 
ii., 369. [With an Alphabetical Index.] 1858, 12mo. 4s. 6d. 


VERMES. 


Catalogue of the Species of Entozoa, or Intestinal Worms, con- 
tained in the Collection of the British Museum. By Dr. Baird, 
Pp. iv., 182. 2 Plates. [With an Index of the Animals in 
whieh the Entozoa mentioned in the Catalogue are found; and 
an Index of Genera and Species.] 1853, 12mo. 2s. 


ANTHOZOA. 


Catalogue of Sea-pens or Pennatulariide in the Collection of the 
British Museum. By J. E. Gray, F.R.S., &. Pp. iv., 40. 
2 Woodeuts. 1870, 8vo. 1s. 6d. 


14 LIS! OF PUBLICATIONS OF THE 


Catalogue of Lithophytes or Stony Corals in the Collection of the 
British Museum. By J. E. Gray, F.R.S., &e. Pp. iv., 51. 
14 Woodcuts. 1870, 8vo. ds. 

Catalogue of the Madreporarian Corals in the British Museum 
(Natural History). Vol. I. The Genus Madrepora. By 
George Brook. Pp. xi., 212. 35 Collotype Plates. [With 
Systematic and Alphabetical Indexes, Explanation of Plates, 
and a Preface by Dr. Ginther.] 1898, 4to. 1/. 4s. 


BRITISH ANIMALS. 


Catalogue of British Birds in the Collection of the British 
Museum. By George Robert Gray, F.L.S., F.Z.S., &e. Pp. 
xii., 248. [With a List of Species.] 1863, 8vo. 3s. 6d. 

Catalogue of British Hymenoptera in the Collection of the British 
Museum. Second edition. Part I, Andrenide and Apide. 
By Frederick Smith, M.E.S. New Issue. Pp. xi, 206. 11 
Plates. [With Systematic and Alphabetical Indexes.| 1891, 
Svo. 6s. 

Catalogue of British Fossorial Hymenoptera, Formicide, and 
Vespide in the Collection of the British Museum. By Frederick 
Smith, V.P.E.S. Pp. 236. 6 Plates. [With an Alphabetical 
Index.] 1858, 12mo. 6s. 

A Catalogue of the British Non-parasitical Worms in the Collec- 
tion of the British Museum. By George Johnston, M.D., Edin., 
F.R.C.L. Ed., Li.D. Marischal Coll. Aberdeen, &c. Pp. 360. 
Woodecuts and 24 Plates. [With an Alphabetical Index.] 
1865, 8vo. 7s. 

Catalogue of the British Echinoderms in the British Museum 
(Natural History). By F. Jeffrey Bell, M.A. Pp. xvii., 202. 
Woodcuts and 16 Plates (2 coloured). [With Table of Con- 
tents, Tables of Distribution, Alphabetical Index, Description 
of the Plates, &c.} 1892, 8vo. 12s. 6d. 

List of the Specimens of British Animals in the Collection of the 
British Museum; with Synonyma and References to figures. 
12mo. :— 

Part I. Centronie or Radiated Animals. By Dr. J. E. 
Gray. Pp. xiii, 178. 1848, 4s. 

Part IV. Crustacea. By A. White. Pp. iv., 141. (With 
an Index.) 1850, 2s. 6d. 

Part V. Lepidoptera. By J. F. Stephens. 2nd Edition. 
By H. T. Stainton and EK. Shepherd. Pp. iv., 224. 1856, 
Ls, Od. 

Part VI. Hymenoptera. By F. Smith. Pp. 134. 1851, 
2s. 

Part VII. Mollusca, Acephala, and Brachiopoda, By Dr. 
J, Gray. Pp. iv. 16” WSais as. oe 


BRITISH MUSEUM (NATURAL HISTORY). 15 


List of the Specimens of British Animals—continued.  _ 
Part VIII. Fish. By Adam White. Pp. xxii., 164. 
(With Index and List of Donors.) 1851, 3s. 6d. 
Part IX. Eggs of British Birds. By George Robert Gray. 
Pp. 143. 1852, 2s. 6d. 
Part XI. Anoplura or Parasitic Insects. By H. Denny. 
Pps iy, ol 1852, 1s. 


Part XII. Lepidoptera (continued.) By James F. Stephens. 
Pp. iv., 54. 1852, 9d. 


Part XIII. Nomenclature of Hymenoptera. By Frederick 
Smith. Pp. iv., 74. 1853, 1s. 4d. 


Part X1V. Nomenclature of Neuroptera. By Adam White. 
Pp. iv., 16. 1853, 6d. 

Part XV. Nomenclature of Diptera, I. By Adam White. 
Pp. iv., 42. 18538, 1s. 

Part XVI. Lepidoptera (completed). By H. T. Stainton. 
Pp. 199. [With an Index.] 1854, 3s. 


Part XVII. Nomenclature of Anoplura, Euplexoptera, and 
Orthoptera. By Adam White. Pp. iv.,17. 1855, 6d. 


PLANTS. 

A Monograph of Lichens found in Britain: being a Descriptive 
Catalogue of the Species in the Herbarium of the British 
Museum. By the Rev. James M. Crombie, M.A., F.LS., 
F.G.S., &¢. Part I. Pp. viii., 519: 74 Woodcuts. [With 
Glossary, Synopsis, Tabuiar Conspectus, and Index.] 1894, 8vo. 
16s. 

A Monograph of the Mycetozoa: being a Descriptive Catalogue 
of the Species in the Herbarium of the British Museum. By 
Arthur Lister, F.L.S. Pp. 224. 78 Plates and 51 Woodcuts. 
[With Synopsis of Genera aud List of Species, and Index. | 
1894, 8vo. 15s. 


List of British Diatomacez in the Collection of the British Museum. 
By the Rev. W. Smith, F.L.S., &c. Pp.iv., 55. 1859, 12mo. 1s. 


FOSSILS. 
Catalogue of the Fossil Mammalia in the British Museum (Natural 
History). By Richard Lydekker, B.A., F.G.S. :-— 

Part I. Containing the Orders Primates, Chiroptera, Insec- 
tivora, Carnivora, and Rodentia. Pp. xxx., 268. 33 
Woodeuts. [With Systematic and Alphabetical Indexes. | 
1885, 8vo. 5s. 

Part I]. Containing the Order Ungulata, Suborder Artio- 
dactyla. Pp. xxii., 324. 89 Woodeuts. [With Systematic 
and Alphabetical Indexes.| 1885, 8vo. 6s. 

Part III. Containing the Order Ungulata, Suborders Peris- 
sodactyla, ‘Toxodontia, Condylarthra, and Amblypoda. Pp. 


16 LIST OF PUBLICATIONS OF THE 


Catalogue of the Fossil Mammalia—continued. 

xvi., 186. 30 Woodcuts. [With Systematic Index, and 
Alphabetical Index of Genera and Species, including 
Synonyms.| 1886, 8vo. 4s. 

Part IV. Containing the Order Ungulata, Suborder Probos- 
cidea. Pp. xxiv., 235. 32 Woodcuts. [With Systematic 
Index, and Alphabetical Index of Genera ‘and Species, 
including Synonyms.]| 1886, 8vo. 5s. 

Part V. Containing the Group Tillodontia, the Orders Si- 
renia, Cetacea, Edentata, Marsupialia, Monotremata, and 
Supplement. Pp. xxxv., 345. “55 Woodeuts. [With 
Systematic Index, and Aiphabetical Index of Genera and 
Species, including Synonyms.] 1887, 8vo. 6s. 

Catalogue of the Fossil Birds in the British Museum (Natural 
History). By Richard Lydekker, B.A. Pp. xxvii., 368. 75 
Woodcuts. [ With Systematic Index, and Alphabetical Index of 
Genera and Species, including Synonyms.| 1891, 8vo. 10s. 6d. 


Catalogue of the Fossil Reptilia and Amphibia in the British 
Museum (Natural History). By Richard Lydekker, B.A., 
F.G.S. :— 


Part I. Containing the Orders Ornithosauria, Crocodilia, 
Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. 
Pp. xxviil., 309. 69 Woodcuts. [With Systematic Index, 
and Alphabetical Index of Genera and Species, including 
Synonyms.| 1888, 8vo. 7s. 6d. 

Part II. Containing the Orders Ichthyopterygia and Sau- 
ropterygia. Pp. xxi., 307. 85 Woodcuts. [With Syste- 
matic Index, and Alphabetical Index of Genera and 
Species, including Synonyms.] 1889, 8vo. 7s. 6d. 

Part III. Containing the Order Chelonia. Pp. xviii. 239. 
53 Woodcuts. [With Systematic Index, and Alphabetical 
Index of Genera and Species, including Synonyms. ] 1889, 
8vo. 7s. 6d. 

Part IV. Containing the Orders Anomodontia, Ecaudata, 
Caudata, and Labyrinthodontia; and Supplement. Pp. 
xxiii, 295. 66 Wocdeuts. [With Systematic Index, 
Alphabetical Index of Genera and Species, including 
Synonyms, and Alphabetical Index of Genera and Species 
to the entire work.] 1&90, 8vo. 7s. 6d. 

Catalogue of the Fossil Fishes in the British Museum (Natural 
Hisicry). By Arthur Smith Woodward, F.G.S., F.Z.S.:—- 

Part I. Containing the Elasmobranchii. Pp. xlvil., 474. 13 
Woodeuts and 17 Plates. {With Alphabetical Index, and 
Systematic Index of Genera and Species.] 1889, 8vo. 21s. 

Part II. Containing the Elasmobranchii (Acanthodii), Holo- 
cephali, [chthyodorulites, Ostracodermi, Dipnoi, and 'Teleo- 
stomi (Crossopterygii and Chondrostean Actinopterygii). 
Pp. xliv., 567. 58 Woodcuts and 16 Plates. [ With 
Alphabetical Index, and Systematic Index of Genera and 
Species.| 1891, 8vo. 21s. 


BRITISH MUSEUM (NATURAL HISroRy)}. 17 


Catalogue of the Fossil Fishes—continued. 

Part III. Containing the Actinopterygian Teleostemi of 
the Orders Chondrostei (concluded), Protospondyli, 
Aetheospondyli, and Isospondyli (in part). Pp. xlii., 544. 
45 Woodeuts and 18 Plates. [With Alphabetical Index, 
and Systematic Index of Genera and Species.] 1895, 8vo 
21s. 


Systematic List of the Edwards Collection of British Oligocene and 
Eocene Mullusca in the British Museum (Natural History), 
with references to the type-specimens from similar horizons 
contained in other collections belonging to the Geological 
Department of the Museum. By Richard Bullen Newton, 
F.G.S. Pp. xxviii., 365. [With table of Families and Genera, 
Bibliography, Correlation-table, Appendix, and Alphabetical 
Index.] 1891, 8vo. 6s. 


Catalogue of the Fossil Cephalopoda in the British Museum 
(Natural History). By Arthur H. Foord, F.G.S. :— 


Part I. Containing part of the Suborder Nautiloidea, con- 
sisting of the families Orthoceratide, Endoceratide, Actino- 
ceratide, Gomphoceratide, Ascoceratide, Poterioceratide, 
Cyrtoceratide, and Supplement. Pp. xxxi., 344. 651 
Woodcuts. [With Systematic Index, and Alphabetical 
Index of Genera and Species, including Synonyms. ] 
1888, 8vo. 10s. 6d. 


Part II. Containing the remainder of the Suborder Nauti- 
loidea, consisting of the families Lituitide, Trochoceratide, 
Nautilidz, and Supplement. Pp. xxviii., 407. 86 Wood- 
cuts. [With Systematic Index, and Alphabetical Index 
of Genera and Species, including Synonyms. | 1891, 8vo. lis. 


A Catalogue of British Fossil Crustacea, with their Synonyms and 
the Range in Time of each Genus and Order. By Henry 
Woodward, F.R.S. Pp. xii., 155. [With an Alphabetical 
Index.] 1877, 8vo. ds. 


Catalogue of the Blastoidea in the Geological Department of the 
British Museum (Natural History), with an account of the 
morphology and systematic position of the group, and a revision 
of the genera and species. By Robert Etheridge, jun., of the 
Departmert of Geology, British Museum (Natural History), 
and P. Herbert Carpenter, D.Se,, F.R.S., F.U.S. (of Eton 
College). [With Preface by Dr. H. Woodward, Tabie of 
Contents, General Index, Explanations of the Plates, &c.] Pp. 
xv., 022. 20 Plates. 1886, 4 to. 25s. 


Catalogue of the Fossil Sponges in the Geological Department of 
the British Museum (Natural History). With descriptions of 
new and little known species. By George Jennings Hinde, 
Ph.D., F.G.S. Pp. viii., 248. 38 Plates. [With a Tabular 
List of Species, arranged in Zoological and Stratigraphical 
sequence, and an Alphabetical Index.] 1883, 4to. 1/. 10s. 

o 89533. B 


18 LIST OF PUBLICATIONS OF THE 


Catalogue of the Fossil Foraminifera in the British Museum 
(Natural History). By Professor T. Rupert Jones, F.R.S., 
&e. Pp. xxiv., 100. [With Geographical and Alphabetical 
Indexes.] 1882, 8vo. és. 

Catalogue of the Palsozoic Plants in the Department of Geology 
and Paleontology, British Museum (Natural History). By 
Robert Kidston, F.G.S. Pp. viii., 288. [With a list of works 
quoted, and an Index.] 1886, 8vo. 5s. 

Catalogue of the Mesozoic Plants in the Department of Geology, 
British Museum (Natural History). he Wealden Flora. 
By A. C. Seward, M.A., F.G.S., University Lecturer in 
Botany, Cambridge. , 

Part I. Thallophyta—Pteridophyta. Pp. xxxviii.. 179: 17 
Woodcuts and 11 Plates. [With Preface byDr. Woodward, 
Alphabetical Index of Genera, Species, &c., Explanations 
of the Plates, &c.] 1894, 8vo, 10s. 

Part II. Gymnosperme. Pp. viii. 259. 9 Woodeuts and 
20 Plates. [With Alphabetical Index, Explanations of the 
Plates, &c.} 1895, 8vo. 15s. 


GUIDE-BOOKS.. 
(To be obtained only at the Museum.) 


‘A General Guide to the British Museum (Natural History), 
Cromweli Road, London, 8.W. [By W.H. Flower.] With 2 
Plans, 2 views of the building, and an illustrated cover. Pp. 80. 
1895, 8vo. 3d. 


Guide to the Galleries of Mammalia (Mammalian, Osteological, 
Cetacean) in the Department of Zoology of the British Museum 
(Natural History). [By A.Giinther.] 5th Edition. Pp. 126. 
57 Woodcuts and 2 Plans. Index. 1894, 8vo. 6d. 


Guide to the Galleries of Reptiles and Fishes in the Department of 
Zoology of the British Museum (Natural History). [By A. 
Giinther.] 3rd Edition. Pp. iv.,119. 101 Woodcuts and 1 
Plan. Index. 1893, 8vo. 6d. 


Guide to the Shell and Starfish Galleries (Mollusea, Echinoder- 
mata, Vermes), in the Department of Zoology of the British 
Museum (Natural History). [By A. Giinther.] 2nd Edition. 
Pp. iv., 74. 51 Woodeuts and 1 Plan. 1888, 8vo. 4d. 


A Guide to the Exhibition Galleries of the Department of Geology 
and Paleontology in the British Museum (Natural History), 
Cromwell Road, London, $.W. [New Edition. By Henry 
Woodward. |— 

Part I. Fossil Mammals and Birds. Pp. xii, 103. 119 
Woodeuts and 1 Plan. 1890, 8vo. 6d. 

Part II. Fossil Reptiles, Fishes, and Invertebrates. Pp. 
xii., 109. 94 Woodeuts and 1 Plan. 1890, 8vo. 6d. 


- 


BRITISH MUSEUM (NATURAL HISTORY). 19 


“Guide to the Collection of Fossil Fishes in the Department of 
Geology and Paleontology, British Museum (Natural History), 
Cromwell Road, South Kensington. [By Henry Woodward. | 
2nd Edition. Pp. 51. 81 Woodcuts. Index. 1888 8vo. 4d. 

-Guide to Sowerby’s Models of British Fungi in the Department of 
Botany, British Museum (Natural History). By Worthington 
G. Smith, F.L.S. Pp. 82. 93 Woodcuts. With Table of 
Diagnostic Characters and Index. 1893, 8vo. 4d. 

‘Guide to the British Mycetozoa exhibited in the Department of 
Botany, British Museum (Natural History). By Arthur Lister, 
F.L.S. Pp. 42. 44 Woodcuts. Index. 1895, 8vo. 3d. 

_A Guide to the Mineral Gallery of the British Museum (Natural 
History). [By L. Fletcher.] Pp 32. Plan. 1895, 8vo. 1d. 

-An Introduction to the Study of Minerals, with a Guide to the 
Mineral Gallery of the British Museum (Natural History), 
Cromwell Road, S.W. By L. Fletcher. Pp. 120. With 
numerous Diagrams, a Plan of the Mineral Gallery, and an 
Index. 1895, 8vo. 6d. 

‘The Student’s Index tothe Collection of Minerals, British Museum 
(Natural History). [New Edition.] Pp. 33. Witha Plan of 
the Mineral Gallery. 1895, 8vo, 2d. 

An Intreduction to the Study of Meteorites, with a List of the 
Meteorites represented in the Collection. [By L. Fletcher. ] 
Pp. 94. [Witha Plan of the Mineral Gallery, and an Index to 
the Meteorites represented in the Collection.| 1894, 8vo. 6d. 

An Introduction to the Study of Rocks. [By L. Fletcher.] Pp. 
118. [With plan of the Mineral Gallery, table of Contents, 
and Index.] 1895, 8vo. 6d. 


W. H: FLOWER, 
Director. 
_British Museum 
(Natural History), 
Cromwell Road, 
London, S.W, 


December Ist, 1895, 


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