Chapter XXXIX: Cycadophytan Fronds (1)

Occasional reference is made to Cycadean fronds in the account of flowers and stems but it is seldom that genera or species founded on leaves can be definitely correlated with particular types of reproductive organs or stems. As in the case of Ferns and Pteridosperms so also with detached leaves believed to be Cycadean, a large number of generic names have been employed for impressions which afford no information with regard to anatomical characters except, in some of the more favourably preserved specimens, a few facts as to the epidermal cells. Though association often suggests original connexion it is inadvisable except in well established cases to extend to fronds generic terms based on reproductive shoots. The designation _Zamites_ has long been used for fronds that are clearly not closely related to recent species of _Zamia_, and were it not an old established genus the significance of which is not likely to be misunderstood, it would be wiser to substitute for it some name implying no affinity with any existing type. On the other hand the employment by some authors of such generic names as _Encephalartos_ and _Ceratozamia_ is not warranted by the evidence furnished by the imperfect material. Prof. Newberry[1424] described as _Encephalartos_? _denticulatus_ a piece of a frond from Rhaetic beds in Honduras characterised by lanceolate pinnae (30 × 6 mm.) gradually narrowed towards the acute apex and abruptly contracted at the base: the method of attachment of the pinnae appears to agree with that in the genus _Zamites_. In this case there is no valid reason for assuming a relationship with _Encephalartos_ or with any recent type. An impression from Lower Cretaceous, Dakota, beds in Kansas described as _Encephalartos cretaceus_ Knowlt. ex Lesq. ms.[1425] consists of a piece of lamina, 9 × 4 cm., obovate-oblong and with a cuneate base, a serrate margin and thick diverging veins: the specimen is too incomplete to serve as a record of any Cycadean genus. The generic name _Encephalartopsis_ was applied by Fontaine[1426] to some imperfect pinnae from the Potomac beds characterised by a linear-elliptical lamina with a spinous margin and slightly diverging and occasionally anastomosing veins. The figured examples of the type-species, _E. nervosa_, suggest pinnae like those of _Ctenis_; but in the absence of a rachis the method of attachment of the segments cannot be ascertained. Saporta[1427] named a specimen from the Miocene flora of Koumi, Greece, _Encephalartos Gorceixianus_ because of its resemblance in habit to some species of the recent genus; but the designation _Zamites_ would be more appropriate. Ettingshausen recorded an imperfect impression of a pinna from Tertiary rocks in Styria as _Ceratozamia Hofmanni_[1428] although it is by no means certain that the fragment is even Cycadean. The genus _Taeniopteris_ was described in the second volume of this book as probably a Pteridophyte, though of uncertain systematic position: it has, however, been shown by Mr Thomas[1429] that the Jurassic species _T. vittata_ was almost certainly borne on a stem with reproductive organs constructed on the Bennettitalean plan. Further research may enable us to fix the position of other species but as yet _T. vittata_ is the only representative of the genus which there is good reason for assigning to the Bennettitales.

It is undoubtedly true that Cycadean plants, using the term in a wide sense to include the Bennettitales as well as the Cycadales, bulked largely in Upper Triassic, Rhaetic, Jurassic-Wealden floras; the Bennettitales probably reached their maximum development as regards wealth of form and geographical range in the latter part of the Jurassic period and in the earliest phase of the Cretaceous epoch. In Triassic floras Cycadean plants are represented almost solely by fronds but the very close resemblance between Keuper species and forms that in Jurassic rocks are found in association with fertile shoots leaves little doubt as to the affinity of Keuper and Rhaetic species.

The evidence obtained from Permo-Carboniferous strata is much more meagre, at least as regards Cycadean leaves: the occurrence of certain morphological Cycadean features is revealed by petrified vegetative organs of Palaeozoic plants, and the Cycadean plan of organisation is conspicuous in many Carboniferous and Permian seeds. The discovery of frond-impressions identical in external characters with Mesozoic genera may be accepted as a substantial indication that genera already existed possessing foliage of the Cycadean type, though we have no certain information with regard to the nature of the other organs of the parent-plants. A few examples of Palaeozoic species are included among those selected in illustration of the different genera, namely _Plagiozamites Planchardi_, _Sphenozamites Rochei_, _Pterophyllum Fayoli_, _P. Cambryi_: among other recorded instances of Palaeozoic species are _Pterophyllum Cottaeanum_ Gutb.[1430], a Permian type similar in habit to _Ctenis_ but without anastomosing veins, a feature in which it resembles _Pseudoctenis_; the Carboniferous species _Pterophyllum inflexum_ Eich.[1431] from the Altai mountains, transferred by Zeiller to _Dioonites_; _P. blechnoides_ Sand.[1432] from the Stephanian of Oppenau; _Pterophyllum Grand’Euryanum_[1433] from Upper Carboniferous beds in France, and _P. gonorrachis_ Goepp.[1434] from Silesia. An examination of the type-specimens of the Carboniferous species _Cycadites gyrosus_ Goepp. and _C. taxodinus_ Goepp.[1435] in the Breslau Museum led me to regard the material as too imperfect to determine.

A conclusion that is forced upon us by a consideration of the geological range of Cycadean fronds is that at the close of the Wealden period, a period very closely linked in the character of the vegetation with the preceding Jurassic floras, there appears to have been a relatively sudden decrease in the number of members of the Cycadophyta: the decline in the fortunes of Cycadean plants is coincident with the rise and remarkably rapid extension of the Angiosperms. From Middle and Upper Cretaceous and from Tertiary beds very few Cycadean remains have been obtained and many of them are represented by fragmentary fossils that afford no definite evidence of affinity to recent genera. The antiquity of the Cycadales, that is the section represented by existing Cycads, cannot be determined; but it would seem probable that if the Cycads apart from the Bennettitales existed in Jurassic and Lower Cretaceous floras they occupied a very subordinate position in comparison with the extinct Bennettitales. There are no data pointing to any widespread occurrence of the Cycadales in the Northern Hemisphere in Tertiary times at all comparable with the geographical range of Tertiary ancestors of the solitary survivor of the Ginkgoales.

The following records of Tertiary Cycadean fronds illustrate the paucity of the records. Reference has already been made to _Encephalartos Gorceixianus_ Sap. of Miocene age, a species that has no claim to be regarded as an example of the recent South African genus. The specimen described by Saporta and Marion as ? _Zamites palaeocenicus_[1436] from the Eocene of Gelinden is too imperfect to serve as a trustworthy record. A more satisfactory species, similar in habit to _Zamites gigas_, is that on which Saporta founded the species _Zamites epibius_[1437] from Lower Miocene beds at Bonnieux (Vaucluse), France. Another Tertiary species is mentioned by Krasser[1438] from Pliocene strata in Brazil as _Zamia praecedens_ Krass. ex Ett. +MS.+ Ettingshausen has described a Tertiary species from New South Wales, either Lower Miocene or Upper Eocene in age, as _Anomozamites Muelleri_[1439], characterised by truncate segments with simple veins and set obliquely to the rachis.

While certain form-genera of fronds can only be referred to the Cycadophyta, in other cases it is possible to assign fronds to a section of this comprehensive group characterised by a particular type of fertile shoot and by certain well defined epidermal features.

The investigation of the cuticular structure of various Cycadean fronds by Nathorst[1440] and especially by Mr Thomas[1441] has supplied a basis of classification which affords the best criterion of affinity so far available. The majority of fronds are placed in the Bennettitales while the three genera _Ctenis_, _Nilssonia_, and _Ctenopteris_ (or _Ptilozamites_[1442]) are placed in the Nilssoniales.

I. =Bennettitales.= Epidermal cells characterised by sinuous walls and generally rectangular; the cuticle is thin; the stomata, confined to the lower surface of the pinnae, tend to be arranged at right-angles to the veins and are on a level with the epidermis or very slightly depressed; two large laterally placed subsidiary cells more or less surround the guard-cells and these are provided with thickenings of a definite shape (figs. 594, 609).

Genera: _Ptilophyllum_, _Zamites_, _Otozamites_, _Dictyozamites_, _Pterophyllum_ and _Anomozamites_, _Taeniopteris_, _Pseudocycas_.

II. =Nilssoniales.= Epidermal cells with straight walls, not sinuous, irregular in form, rounded, hexagonal, or rectangular; the stomata are below the level of the epidermis; the cuticle may be thin or thick. There is no regular arrangement of the stomata; the guard-cells are surrounded by 6–8 subsidiary cells (fig. 625) which often form an overarching canopy; thickening lamellae like those on the guard-cells of the Bennettitales are seldom present.

Genera: _Nilssonia_, _Ctenis_, _Ctenopteris_.

It is a noteworthy fact that the representatives of the smaller group, the Nilssoniales, in their cuticular features, are more akin than the Bennettitales to modern Cycads. Until definite evidence is obtained as to the nature of the reproductive organs of _Nilssonia_, _Ctenis_, and _Ctenopteris_ it is impossible to say how closely these genera agree in essential characters with existing members of the Cycadales. If, as has been suggested, the fertile shoots known as _Beania_[1444] belong to _Nilssonia_ their resemblance in plan of construction to the cones of recent genera, much greater than in the case of the flowers of the Bennettitales, is in accordance with the evidence of the epidermal characters.

Reference was made in the second Volume of this book to several genera founded on fronds which through lack of evidence as to the nature of the reproductive organs cannot be assigned with certainty either to Ferns or Cycads: it was stated that the genera _Ptilozamites_ and _Ctenopteris_ are probably Cycadean, and the structure of the epidermal cells in the latter genus lends support to this view[1445]. Among other genera of doubtful position not included in the following descriptions of fronds is _Zamiopsis_ of Fontaine[1446] founded on large compound fronds from the Potomac formation: the venation and form of the pinnae are more Fern-like than in _Ctenopteris_, but in general habit the two genera are not very dissimilar. The generic name Zamiopsis is misleading as the species bear no resemblance to _Zamia_ or _Zamites_.

+I. Bennettitales.+

=PTILOPHYLLUM.= Morris.

The generic name _Ptilophyllum_ was instituted in 1840 for some specimens of pinnate fronds from Cutch: in this genus Morris[1447] included with the Indian leaves the English Jurassic species _Pterophyllum pecten_ Lind. and Hutt.[1448] (figs. 587, etc.) and some other forms. He defined _Ptilophyllum_ as follows: ‘Fronds pinnate; pinnae closely approximated, linear, lanceolate, more or less elongate, imbricate at the base, attached obliquely; base semicircular or rounded; veins equal, slender, parallel.’ Morris adds that he instituted a new genus in preference to _Zamites_ because of the ‘oblique insertion of the pinnae and their overlapping each other at the base.’ In a later paper Morris[1449] states that the pinnae of _Ptilophyllum_ fronds are ‘sometimes auriculed in the upper and sometimes in the lower part’ of the base of the lamina. In his catalogue[1450] he adopted Endlicher’s genus _Palaeozamia_ instead of _Ptilophyllum_. Without discussing the generic nomenclature adopted by various authors for the Indian types and similar fronds it is important to refer briefly to the treatment of Morris’s species by Oldham and Morris and by Feistmantel. In the first of the series of Memoirs on Gondwana floras[1451] _Ptilophyllum_ is retained for a section of _Palaeozamia_ together with _Otozamites_ and _Sphenozamites_ as other sectional subdivisions: in the subgenus _Ptilophyllum_ are included _Palaeozamia acutifolia_ and _P. cutchensis_ (fig. 588, A, C), also _P. affinis_, _P. rigida_, and _P. bengalensis_. The last species is in all probability an _Otozamites_: _P. affinis_ and _P. rigida_ are almost certainly indistinguishable from _P. cutchensis_. Feistmantel[1452] dealt in detail with the genus _Ptilophyllum_: he wrote, ‘with Schimper and Schenk I therefore look upon this genus as an Indian type especially characterised by its ... more or less slender leaves, angustate towards the apex and base, and petiolate, with regularly adfixed leaflets.... The leaflets are equal to each other, for the most part elongate linear, and _auriculate and free at the upper angle at the base_, but adfixed at the lower angle, and each is decurrent behind the upper angle of the leaflet next below it, thus the leaflets are almost imbricate. The veins are rather numerous, simple, and forked, and more or less divergent[1453].’ Feistmantel distinguishes certain varieties of _P. cutchense_ (fig. 588, A, C), none of which appear to be well defined. A specimen from the Rajmahal Hills with unusually long pinnae, the frond having a breadth of 8 cm., is described as _Ptilophyllum acutifolium_ var. _maximum_[1454], but it differs in no important feature from the smaller and commoner form. The next point to be considered is the variability of certain species referred by Feistmantel to _Otozamites_. He figures specimens from the Jabalpur group as _O. Hislopi_ Feist. ex Old. +MS.+, _O. gracilis_ (Kurr), _O. angustatus_ Feist. (fig. 588, B) and _O. distans_[1455]. An examination of the figured specimens leads me to regard _O. Hislopi_ and _O. gracilis_ as identical with the _Ptilophyllum_ fronds: the pinnae exhibit no distinguishing features and there is no reason for a specific, still less a generic, separation. _Otozamites angustatus_ is indistinguishable from _Otozamites_ sp. as figured from the Madras coast and from _Ptilophyllum cutchense_, _P. cutchense_ var. _curvifolium_ and var. _minimum_. The drawings reproduced in fig. 6_a_, Pl. +x.+ of the Madras flora[1456] and in fig. 8_a_, Pl. +vi.+ of the Jabalpur flora[1457] showing auriculate bases are inaccurate: in all the fronds named the pinnae are straight with rounded edges precisely as in _Ptilophyllum_. The conclusion forced upon me by a comparison of the actual specimens is that the Indian fronds are not separable into well-defined species and should all be included in _Ptilophyllum cutchense_. Moreover in this comprehensive species should be included the specimens described by Feistmantel as _Otozamites Hislopi_ (fig. 589), _O. angustatus_, and _O. gracilis_. It may well be that a more detailed investigation of the numerous forms comprised in this protean species, particularly if specimens are obtained from which cuticular preparations can be made, may lead to the recognition of additional species or well-defined varieties. The resemblance between the various forms of _P. pecten_ from the Jurassic strata of Yorkshire and those of _P. cutchense_ from India is very striking, and at least in many cases no specific separation is possible so far at least as the form of the fronds and pinnae is concerned. The occasional close association of _Ptilophyllum_ fronds and _Williamsonia_ flowers is an important agreement between the English and Indian fronds (fig. 590).

In the first part of the Catalogue of Jurassic plants from Yorkshire the opinion was expressed[1458], based on an examination of Morris’s type-specimen of _Ptilophyllum cutchense_ and of a large number of English and other fronds identical with or closely allied to _Pterophyllum pecten_ Lind. and Hutt., that the Indian and European fronds belong to the same genus. In a later paper[1459] it was maintained that _P. cutchense_ and _P. acutifolium_ are probably identical with the English type, and a drawing was published—reproduced in fig. 591—of Morris’s type-specimen. Dr Halle[1460] has discussed the genus _Ptilophyllum_ and his investigations lead him to a different conclusion; he, like Zeiller and some other authors, employs _Ptilophyllum_ in Feistmantel’s sense. The pinna-base is said to agree in its asymmetrical form with that in _Otozamites_, while it differs from the symmetrical base of _Zamites_ pinnae. In _Ptilophyllum_ both edges of the pinna-base are said to bend down in joining the rachis; at the upper corner the base thus becomes rounded and for some distance free from the rachis; it is also sometimes a little auriculate; at the lower edge the pinna is decurrent on the upper surface of the rachis. In a subsequent paper Halle[1461] repeats the view that the decurrence of the pinnae by their lower edges is an important distinguishing feature of _Ptilophyllum_, thus agreeing with Feistmantel whose illustrations appear to be confirmatory. The drawings in Feistmantel’s memoirs are, however, misleading and in some cases incorrect. An examination of a photograph of Morris’s type-specimen of _P. cutchense_ led Halle to conclude that the two edges of the pinna-base bend downwards on joining the rachis as described by Feistmantel. This feature is not shown in the drawing reproduced in fig. 591: it is clear that either the drawing is incorrect or that there has been some mistake in the interpretation of the photograph. Through the courtesy of Dr Halle I have been able to examine the actual print: when viewed in its correct position the two edges of the pinnae appear to bend down as described by Halle, but if it is examined in the reverse position the lower angle of the pinnae is seen to be slightly rounded as in fig. 591, the apparent decurrence being due to a confusion between the appressed lower edge of one pinna, which is faintly shown, and the stronger downward trend of the upper edge of the pinna next below. The upper edges of the pinnae are more prominent because they are less appressed to the rachis while the lower half of the base is closer to the rachis and is frequently, though not in Morris’s specimen, overlapped by the upper edge of the next lower pinna. A re-examination of the type-specimen in the British Museum confirms this interpretation. The pinnae of _Ptilophyllum_ are characterised by their attachment to the upper face of the rachis which they almost completely cover; the upper angle is rounded and in a few cases auriculate (fig. 592); the lower angle of the base is slightly rounded and not infrequently hidden by the imbrication of the adjacent pinna; it is occasionally auriculate (fig. 593). The pinnae are attached by nearly the whole base, but the upper angle is free. The veins are parallel, sub-parallel or, especially in the proximal portion of the lamina, oblique. The pinnae are linear, varying considerably in relation of length to breadth and in the form of the apex; they are straight or more or less falcate. The epidermal cells of such _Ptilophyllum_ fronds as have been examined are characterised by strongly looped or sinuous walls; the stomata, confined to the lower surface, are roughly circular and the guard-cells are at right-angles to the veins and not appreciably sunk. Fig. 594 represents the appearance of a stoma in surface-view: ‘on either side of the central slit-like pore are two elliptical or hemispherical structures; they are somewhat flattened when they abut on the pore, and have rounded ends.... Between these and the subsidiary cells lie two other thickened patches, more or less hemispherical in shape, and apparently overlying the central structures.’ On the analogy of similar appearances in recent Cycads Mr Thomas[1462] interprets the two pairs of thickened patches as belonging to the upper and lower sides of the highly inclined guard-cells. This author calls special attention to the abundance on some of the fronds included in the aggregate species _P. pecten_ of regular rows of circular hair-scars preserved as small annulate projections, ·03–·04 mm. in diameter. A comparison of the cuticles of different forms of _Ptilophyllum pecten_ enabled Thomas to recognise more than one type: for one of these the name _Ptilophyllum hirsutum_ is proposed. It is by such work as this that we may hope to discover differentiating characters.

]

The different forms of _Williamsonia_ flowers found in association with fronds of the _Ptilophyllum_ habit also point to the inclusion of more than a single species under the group-species _P. pecten_. As additional evidence is obtained further analysis will be possible, but in dealing with impressions which include specimens transitional from one form of frond to another, the most convenient and to my mind the most logical course is to treat a species as an aggregate- or group-species. Some authors believe that the two fronds described by Phillips as _Cycadites pecten_ and _C. pectinoides_ are distinct species[1463], but there would seem to be no adequate reason for this view. The fronds described by Heer[1464] from the Lower Cretaceous of Greenland as species of _Zamites_ and similar leaves from Graham Land included in that genus by Halle[1465] are in my opinion inseparable from _Ptilophyllum_, and this applies equally to _Otozamites Hislopi_ (Old.) (fig. 589) and _O. abbreviatus_ as figured by Feistmantel and by Halle. The nomenclature of Cycadean fronds having the habit of _Ptilophyllum pecten_ is a great difficulty: after carefully reconsidering the whole question and comparing Feistmantel’s figured specimens with the large series of English fronds the conclusion reached is that the characters exhibited by ordinary impressions do not admit of any satisfactory grouping under well-defined specific types. In the first place, as already indicated, the use by some authors of the generic names _Ptilophyllum_, _Otozamites_, and _Zamites_ creates a false impression of the degree of difference between the numerous forms of frond agreeing more or less closely with the specimens on which were founded the species _Ptilophyllum acutifolium_, _P. cutchense_, _P. pecten_, and _P. pectinoides_ (figs. 587, 591, 595, etc.). In his important memoir on the Jurassic flora of Graham Land Halle[1466] discusses the limitation of _Ptilophyllum_, _Zamites_, and _Otozamites_. He employs _Zamites_ for fronds with linear pinnae attached to the upper face of the rachis by a base which is more or less, but often very little, rounded and always asymmetrical, with or without a basal callosity: fronds of the type _Z. gigas_ he includes in the section _Euzamites_, while _Z. borealis_ and similar forms (fig. 597) are referred to a second section, _Subzamites_. It is in the sense of Halle’s section _Euzamites_ that the generic name _Zamites_ is employed in this chapter. On the other hand the fronds grouped by Halle as _Subzamites_ have pinnae with the basal angles of the lamina very slightly rounded precisely as in _Ptilophyllum_ as seen in fig. 596 (_cf._ fig. 598 which represents fronds referred by Halle to _Zamites_); and they are not distinguished by any feature of generic importance from _Ptilophyllum_ as defined on page 519. The species _Zamites pusillus_, _Z. Anderssoni_, and _Z. antarcticus_ (fig. 598) described by Halle from Graham Land as well as Heer’s Arctic forms[1467] _Z. borealis_ (fig. 597), _Z. speciosus_, _Z. brevipennis_, and others are transferred to _Ptilophyllum_ as types agreeing very closely with _P. pecten_ and in some cases not clearly distinguishable from it even specifically.

_Otozamites_ is the name employed by Halle for fronds with pinnae having contracted, asymmetrical and auriculate, bases, the anterior lobe being more developed than the posterior. The asymmetry of the pinna-base is considered an essential feature. As Halle states it is very difficult in some instances to draw a distinction between _Otozamites_ and _Ptilophyllum_. As used in this chapter _Otozamites_ signifies fronds with pinnae characterised by an auriculate base, a lamina usually broader than in _Ptilophyllum_, and by more spreading veins (fig. 604). The not infrequent occurrence of auriculate pinnae on fronds (fig. 603, A) which cannot be separated from typical examples of _P. pecten_ illustrates the narrow dividing line as regards the form of the pinna-base between _Ptilophyllum_ and _Otozamites_. The Indian species _O. Hislopi_ (fig. 589) and _O. abbreviatus_, to which Halle refers some Antarctic fronds, cannot be distinguished from the English _P. pecten_ or Feistmantel’s and Morris’s Indian fronds described as _P. cutchense_ and _P. acutifolium_: these forms are therefore included in _Ptilophyllum_.

_Ptilophyllum_ (_Williamsonia_) _pecten_ (Phillips).

This designation is employed in a wide sense for a group of fronds exhibiting a considerable range in size, in the relative breadth and length of the linear pinnae, and in other features. Under _P. pecten_ are included (i) the English fronds from Yorkshire first described by Phillips[1468] as _Cycadites pecten_ and _C. pectinoides_, the former from the Middle shale, the latter from the Lower shale of the Yorkshire coast, together with the Stonesfield slate specimens named by Sternberg[1469] _Polypodiolites pectiniformis_ (fig. 595) and by Brongniart[1470] and Lindley and Hutton[1471] _Zamia pectinata_; (ii) the Indian specimens already considered and (iii) numerous examples recorded under different names from Jurassic strata in many countries. Fronds from the Yorkshire coast named by Brongniart[1472] _Zamia Goldiei_, though regarded by some authors as examples of _Otozamites_, are probably referable to _P. pecten_. Andrae’s _Pterophyllum rigidum_[1473] (fig. 596) from Steierdorf is almost certainly a form of _Ptilophyllum pecten_. The specimen shown in fig. 587 is one of the few examples of fronds apparently preserved in their original position attached in a cluster to a _Williamsonia_ (_Bucklandia_) type of stem. The range in size and form of the pinnae is illustrated in figs. 588, 592, 593.

Fronds linear, tapering gradually towards the base and apex, often characterised by a marked uniformity in breadth. The pinnae, short or comparatively broad or long and narrow, are straight or more or less falcate; the apex is obtuse or acute or the upper margin of the lamina may be almost straight and the lower edge curved abruptly upwards at the apex; the pinnae are usually attached obliquely to the rachis but may be almost at right-angles; with the exception of the upper angle, the whole of the base is attached to the frond-axis; the base of the lamina may be symmetrical, both angles being slightly rounded, or asymmetrical, the upper or less frequently the lower corner being auriculate (figs. 592, 593). There is no basal callosity on the lamina nor is there a median sinus. The veins are more or less spreading at the base but for the most part parallel. The features of the epidermal cells and stomata are mentioned in the general account of the genus. Fertile shoots of plants with this type of frond are described under _Williamsonia_. Very little is known of the stems which bore _Ptilophyllum_ fronds, but as stated on page 488 Indian specimens show leaves of _Ptilophyllum cutchense_ attached to a piece of stem (fig. 579) having the characters of _Bucklandia_ and characterised by a xylem-cylinder denser than in recent Cycads; the structure of the wood at least in the Indian stem is rather pycnoxylic than manoxylic.

The fronds grouped under _Ptilophyllum pecten_ are very widely distributed in Jurassic floras; they are recorded from many localities in Europe, from Turkestan[1474], India, Graham Land[1475], Patagonia[1476], and elsewhere.

As thus defined this ‘species,’ or more correctly this group of forms, undoubtedly includes more than one species in the strict sense, but without additional data it is maintained that the recognition of clearly defined specific types or varieties is beyond our power. It may be urged that in view of the wide geographical range of the _Ptilophyllum pecten_ type of frond and the admitted probability that several species in the narrower sense are represented, distinctive specific names should be retained even though under such designations are included forms that, so far as can be seen from impressions, exhibit no constant distinguishing features. My purpose is to emphasise the futility of attempting to found well-marked species on the available material. The student must decide for himself what course to pursue, whether to retain such a specific name as _cutchense_ for the Indian fronds or to employ that and other specific names as designations of geographical types differing in no clearly defined or constant characters from examples of the English _Ptilophyllum pecten_. A comparative examination of the cuticular membranes in the comparatively few cases where that is possible would probably furnish a basis for a satisfactory subdivision of the group-species.

The names _Ptilophyllum Anderssoni_, _P. boreale_, and _P. antarcticum_ are retained for certain forms selected from a number of closely allied types partly on the ground that these fronds exhibit some more or less well-marked distinctive characters and in part as a recognition of the existence of geographical forms.

_Ptilophyllum boreale_ (Heer).

Heer[1477] described numerous well-preserved impressions of pinnate fronds from the Lower Cretaceous plant-beds of Kome, Greenland, which he referred to several species of _Zamites_ though the differences between them are hardly of specific rank. An inspection of several of the figured specimens in the Stockholm Museum showed that Heer’s drawings are in the main accurate. _Zamites borealis_ (fig. 597) is characterised by small linear pinnae attached to the upper face of the rachis, the stoutness of which is a noteworthy feature; the veins are parallel and simple. In the shape of the pinnae, including the base-characters, this species agrees closely with _P. pecten_. The slightly falcate or straight pinnae are 1·7–2 mm. broad and reach a length of 15 mm.; in _Z. speciosum_ Heer, probably specifically identical with _P. boreale_, the pinnae are relatively longer, and in _Z. brevipennis_ Heer they resemble the shorter pinnae of some of the narrow fronds of _P. pecten_.

A preparation of the cuticle of a pinna of _P. boreale_ made by Mr Thomas from a specimen in the Stockholm Museum shows a marked difference between the upper and lower epidermis: the cells on the upper surface of the lamina have very thick and sinuous walls precisely as in _P. pecten_, but the cells are often broader than long and not elongated parallel to the long axis of the pinna; those of the lower epidermis are thinner and less clearly preserved: the stomata, which appear to be like those of _P. pecten_, are scattered and not in rows.

_Ptilophyllum antarcticum_ (Halle).

The fronds from the Jurassic rocks of Graham Land, described by Halle as _Zamites antarcticus_[1478], are very similar to some forms of _P. pecten_ and to _P. boreale_ and other Greenland forms; the linear subacute pinnae are attached to the upper face of the rachis at a wide angle and the base of the lamina is truncate and may be very slightly constricted. The veins are dense, as many as 5 in 1 mm., and they are occasionally forked near the base (fig. 598, A).

_Ptilophyllum Anderssoni_ (Halle).

This Graham Land species (fig. 598, B), referred by Halle to _Zamites_[1479], is of the same general type as _P. pecten_, but is characterised by a coarser venation and by the wider angle of attachment of the pinnae.

_Ptilophyllum Dunkerianum_ (Goeppert).

Goeppert[1480] proposed the name _Pterophyllum Dunkerianum_ for some specimens from the Wealden of North Germany which were afterwards figured by Dunker[1481]. Miquel[1482] included the species in his genus _Dioonites_ and this name has been adopted by other authors. Attention has been drawn to the inconsistent use of the title _Dioonites_[1483], and I have previously employed the name in a sense similar to that in which it has been adopted by Nathorst, that is for fronds with long and narrow pinnae without any basal constriction and not auriculate, attached more or less at right-angles to the upper face of the rachis; but so defined _Dioonites_ differs in no essential particular from forms of _Ptilophyllum_ or from fronds referred by authors to Schimper’s genus _Ctenophyllum_. The name _Ctenophyllum_[1484] was instituted for certain fronds differing in some points from _Otozamites_ and _Dioonites_. One such type is _Ctenophyllum_ (_Ptilophyllum_) _pecten_ (Lind. and Hutt.): this is quoted in Zittel’s _Handbuch_ as a typical representative of the genus. Schimper followed Feistmantel in his definition of _Ptilophyllum_, a definition which is not in accordance with the characters of the fronds on which it was founded by Morris. Fontaine, on the other hand, has applied _Ctenophyllum_ to fronds of a different type which are now included in the genus _Pseudoctenis_. There would seem to be no adequate ground for the retention of _Ctenophyllum_ as a generic designation.

The Wealden species _P. Dunkerianum_ is characterised by the following features: rachis fairly stout, pinnae approximate, linear 2–3 mm. broad and reaching a length of 11 cm. or more, gradually narrowed towards the apex, attached in two almost contiguous rows to the upper face of the frond-axis: the lower margin of the lamina may be slightly decurrent, _e.g._ in the apical part of the frond or very slightly broadened and bluntly rounded. The pinnae are attached at right-angles or, near the apex, obliquely; veins 5–6, parallel. The epidermal cells as figured by Schenk[1485] have very sinuous walls and are identical with those of the English and Indian forms of the group-species _P. pecten_; the stomata are confined to the lower surface. If Schenk’s drawings of the stomata are correct they differ from those of other species of the genus in their simpler structure; there are no subsidiary cells and the guard-cells show no cuticularised bands. In habit this species resembles _Encephalartos Ghellinckii_ Lehm. (fig. 382). The superficial resemblance of the narrow linear pinnae to those of _Cycadites_ led Dunker and Schenk to refer to that genus some specimens which have since been described as identical with _P. Dunkerianum_.

=ZAMITES.= Brongniart.

In the first instance the generic name _Zamites_ was used in addition to _Zamia_ for certain pinnate fronds including species[1486], such as _Z. Bechei_, which are now regarded as typical examples of _Otozamites_. Subsequently Brongniart gave up _Zamia_ for fossil fronds and applied _Zamites_ to fronds with entire pinnae, not truncate at the apex and not decurrent but slightly constricted at the base. Braun’s two genera _Podozamites_ and _Pterozamites_ were relegated to the position of subgenera. The name _Podozamites_ has been employed by Schenk[1487] for specimens now included in _Zamites_, and _Zamites_ is used by him[1488] for some fossils which are examples of _Podozamites_ as generally understood. Goeppert’s definition[1489] of _Zamites_ includes fronds with pinnae of the _Otozamites_ type, and this author pertinently compares _Zamites_ with recent _Encephalartos_ leaves. Pomel[1490] proposed the name _Crossozamia_ for certain fronds of the _Zamites_ type, but this genus with several others instituted by the same author has not been adopted. Bornemann[1491] described _Zamites_ as comprising species with a greater or less resemblance to the fronds of recent Zamias.

As defined below, _Zamites_ fronds may be compared with those of some species of _Encephalartos_, _Ceratozamia_, and _Macrozamia_. There has been considerable difference of opinion with regard to the range of form in the pinnae that it is advisable to include in _Zamites_. The name _Zamiophyllum_ was proposed by Nathorst[1492] for a Wealden species, described by Ettingshausen as _Pterophyllum Buchianum_ (fig. 601, A–C), characterised by a decrease in the breadth of the linear pinnae towards the point of attachment and, according to Nathorst’s description, by the lateral attachment of the pinnae. An examination of specimens of this type from English rocks[1493] enabled me to show that the pinnae are attached to the upper face of the rachis. Zeiller[1494] has also included _Zamiophyllum_ in _Zamites_, but Halle’s recent definition[1495] of the latter genus excludes fronds of the _Zamiophyllum_ type. Schimper instituted the name _Glossozamites_[1496] for fronds bearing pinnae with rounded and slightly contracted bases and borne on grooves on the upper surface of the rachis (fig. 601, F). In venation the pinnae agree with those of _Otozamites_ but the base is not auriculate. Kurr’s Liassic species _Pterophyllum oblongifolium_[1497] included by Schimper in _Glossozamites_ is probably an _Otozamites_. Another species referred to this genus is Schenk’s _Podozamites Zitteli_[1498] from the Urgonian of Austria (fig. 601, F): in this species the slightly falcate pinnae with obtuse apices have rounded and not auriculate bases and there is some evidence of a basal callosity. This species agrees so closely with such a typical _Zamites_ as _Z. gigas_ that it is difficult to see on what grounds the retention of _Glossozamites_ is desirable. A Portuguese specimen referred by Saporta[1499] to _Glossozamites_, _G. brevis_, is founded on a single specimen very like a leaflet of _Otozamites Klipsteinii_ (Dunk.); and _G. parvifolia_ Yok.[1500] from China has no claim to be included among Cycadean fronds. Feistmantel’s _Glossozamites Stoliczkanus_[1501] is almost certainly a leaf of _Cordaites_. For certain fronds originally described as _Zamites_ Zeiller[1502] has instituted the genus _Plagiozamites_. An important question as to the type of frond which may conveniently be included in the genus _Zamites_ was raised by the application of this name by Heer to some Lower Cretaceous fronds from Greenland, _Z. borealis_ (= _Ptilophyllum boreale_, fig. 597), and others which differ in the form of the pinnae from species usually regarded as typical of the genus. Halle[1503] accepts the Greenland species as examples of _Zamites_ though he distinguishes them by a sectional name _Sub-Zamites_; he defines _Zamites_ as including fronds with pinnae attached to the upper face of the rachis with a contracted and always symmetrical base though in some cases the basal contraction is exceedingly small; there is a more or less distinct basal callosity. He recognises two types, (i) _Eu-Zamites_, _e.g._ _Zamites gigas_ (fig. 599), in which the pinnae are strongly contracted basally and have a callosity, and (ii) _Sub-Zamites_, _e.g._ _Z. borealis_[1504], etc., in which the pinnae are not so broadly rounded at the base and retain the same breadth to the point of insertion where they are ‘very rapidly and very little contracted,’ with or without a basal callosity. The basal callosity in these forms is not shown in many of the pinnae and is at most but a slight rounding of the angles of the truncate base. The specimens referred by Halle to _Sub-Zamites_ do not appear to differ in any feature worthy of generic rank from _Ptilophyllum_.

In the following definition _Zamites_ is employed in a sense more or less in accordance with Brongniart’s usage but with the inclusion of such forms as _Z. Buchianus_, the type of Nathorst’s _Zamiophyllum_, and the exclusion of Heer’s Arctic and Halle’s Antarctic fronds referred by them to _Zamites_.

Fronds broadly lanceolate reaching a length of over 60 cm.; pinnae more or less oblique or at right-angles to the rachis (fig. 599), attached to the upper surface but not completely covering it, linear or linear-lanceolate, acuminate or obtuse, usually abruptly contracted at the rounded base and more rarely _(e.g._ _Z. Buchianus_, fig. 601) gradually tapering to the proximal end, with or without a callosity at the symmetrical base; veins divergent at the base, simple or dichotomously branched, for the most part parallel to the edges of the lamina and slightly divergent in the apical region. The presence of a basal callosity, such as is best seen in the pinnae of recent species of _Macrozamia_, is not a feature of great importance and cannot be easily recognised in many impressions. In the process of fossilisation the pinnae are often flattened against the surface of the rachis and this may produce transverse wrinklings of the lamina suggestive of a basal thickening. In some cases stems of the _Bucklandia_ type occur in connexion with _Zamites_ fronds (fig. 541). An account of these stems, and of inflorescences of _Williamsonia_ which were borne by some at least of the plants with _Zamites_ leaves is given in Chapter +xxviii.+ In such examples of _Zamites_ as have been examined the epidermal cells have sinuous walls and the stomata[1505], confined to the lower face of the lamina, occur in rows with their long axis at right-angles to that of the pinnae.

_Zamites_ ranges from Rhaetic to Lower Cretaceous strata.

_Zamites truncatus_ Zeiller.

A Rhaetic species from Tonkin[1506] characterised by oval linear pinnae given off at a wide angle, alternate and not contiguous; apex obtusely truncate, gradually narrowed to a cuneate base which is rounded or elliptical and has a more or less definite callosity. The pinnae vary from 3 to 5 cm. in length and from 5 to 13 mm. in breadth.

_Zamites gigas_ (Lindley and Hutton).

The specific name _gigas_ is retained in preference to that of _Mantelli_ employed by Brongniart[1507] on the ground that he afterwards discarded it in favour of the designation proposed by Lindley and Hutton[1508], and because their name has been generally adopted.

Fronds large, exceeding 60 cm.; broadly linear lanceolate; the comparatively slender rachis bears alternate linear lanceolate pinnae with a rounded and usually slightly swollen base and an acuminate apex. At the apex of the frond the pinnae are narrow and linear (fig. 599) and almost parallel to the rachis; in the lower part they are shorter and relatively broader and attached approximately at right-angles. The numerous veins diverge from the centre of the base but for the most part are parallel to the edge of the lamina. The form of the epidermal cells and the structure of the stomata have recently been described by Mr Thomas. The external features of the stem (fig. 541) are described under the genus _Williamsonia_. In the case of fronds of this type from English Jurassic rocks it would be legitimate to speak of them as _Williamsonia gigas_, but in view of the fact that such fronds usually occur as detached specimens and without any associated flowers it is advisable, as Nathorst maintains, to retain the non-committal genus _Zamites_.

This type of frond is widely distributed in Jurassic strata. The occurrence of many forms agreeing generally with the type-specimen but differing from it in features that are not constant or of real morphological importance renders accurate specific delimitation very difficult. Species that appear to be indistinguishable from _Zamites gigas_ by any well-marked characters are, _Zamites Feneonis_[1509] Brongn., _Z. Moreaui_[1510] Brongn., _Z. Renevieri_[1511] Heer, and _Z. claravallensis_[1512] Sap.; similarly _Z. Schmiedelii_ And.[1513] is probably identical with _Z. gigas_. The Lower Cretaceous species _Zamites bohemicus_[1514] Vel. and _Z. iburgensis_[1515] Hos. and von d. Marck represent very similar forms.

_Zamites recta_ (Tate).

This species was originally described by Tate[1516] as _Palaeozamia_ (_Otozamites_) _recta_ from Wealden strata in South Africa and subsequently transferred to _Zamites_[1517]; it bears a very close resemblance to _Z. gigas_. The fronds bear alternate linear pinnae attached to the upper face of the rachis by a slightly contracted and swollen base. The lamina has an acuminate asymmetrical apex and the upper edge is slightly falcate; the larger pinnae are over 6 cm. long and nearly 1 cm. broad; the veins are frequently forked as they converge towards the base of the lamina. No clear evidence of association of these fronds in the Uitenhage series of South Africa with _Williamsonia_ flowers has been discovered, but a specimen[1518] in the Tate collection in the British Museum may be a badly preserved cluster of bracts belonging to a _Williamsonia_. The rachis of this species shows some peculiar features in the form of two rows of alternate cushions in some partially decayed specimens. One of these is shown in fig. 600; the cushions are raised oval projections with a flat top, and pieces of the rachis without pinnae might easily be mistaken for a Coniferous stem.

_Zamites Carruthersi_ Seward.

A species founded on specimens from the Wealden of Sussex[1519] and recorded from Kimmeridge beds in Scotland[1520], characterised by linear or linear-elliptical pinnae attached obliquely to the outer part of the upper surface of the rachis, somewhat abruptly narrowed at the proximal end but slightly broadened at the actual base (fig. 601, D, E). The pinnae appear to have been caducous and, as in fig. 601, D, the position of an absciss-layer is occasionally visible. In habit the fronds bear a close resemblance to those of _Encephalartos longifolius_ Lehm. The veins diverge from the base and are for the most part parallel, though divergent at the bluntly rounded apex. If, as may be the case, a specimen figured by Hugh Miller[1521] from Scotland as _Zamites_ and subsequently named by Richards[1522] _Podozamites Milleri_ is identical with _Z. Carruthersi_, the specific name _Milleri_ has priority.

_Zamites Buchianus_ (Ettingshausen).

A Wealden and Lower Cretaceous species[1523] (figs. 601, A–C; 602) represented in several European localities, also in North America and Japan, reaching a length of over 70 cm.; in habit very similar to _Ceratozamia mexicana, Macrozamia Macleayi_ and some other recent Cycads. The rachis has a fairly broad median groove on the upper surface; pinnae alternate, opposite or sub-opposite, from 3 to 20 cm. long and from 1·5 to 2 cm. broad, linear, generally narrowed towards the base, but in the more slender segments the reduction in breadth is less obvious; attached obliquely to the rachis, slightly thickened and broadened at the base (fig. 601, C), separated from the rachis by a distinct absciss-layer and leaving an elliptical scar; usually inclined at about 45° but the angle varies considerably in different parts of a frond (fig. 602); apices generally tapering to a point, or more or less obtusely rounded; veins numerous, parallel, and not as a rule prominent. It is by no means unlikely that specimens figured by Goeppert[1524] and some other authors as _Pterophyllum saxonicum_ or _Dioonites saxonicus_ are examples of this species. Fontaine[1525] speaks of _Dioonites Buchianus_ as one of the most widely distributed and characteristic members of the Potomac flora and it is described from Japan by Yokoyama[1526] and Nathorst[1527]. This type appears to be especially characteristic of Wealden strata.

=OTOZAMITES.= Braun.

Braun[1528] proposed the name _Otozamites_ for certain Mesozoic fronds formerly included in _Zamites_, one of his types being _Otozamites obtusus_ (Lind. and Hutt.) (fig. 603, B) originally regarded by Brongniart as a Fern and named _Filicites Bucklandi_[1529]. The auriculate form of the base of the pinnae and the spreading veins were emphasised in the definition of the genus. As in the case of many other Cycadean fronds the limits of the genus are not always easy to define, but as described below the genus is on the whole fairly distinctive. It is a very widely spread Jurassic type and extends from Triassic to Lower Cretaceous rocks. The supposed Cretaceous species from Greenland, _O. groenlandica_ Heer[1530], is probably not a plant-impression but a polished groove in the rock.

Fronds pinnate, reaching a length of 50 cm. or more in some species; pinnae alternate, separate or contiguous and imbricate, attached by a portion of the base to the upper surface of the rachis, long and narrow (fig. 603, A), broadly oval or almost orbicular, apex acute or obtuse, base auriculate and asymmetrical[1531], the anterior lobe being more prominent than the posterior edge of the lamina which is usually rounded. The veins radiate from the base and pass obliquely with occasional branching to the edge of the pinna; in the more linear pinnae the veins may be parallel or nearly so. Zigno[1532] figured a piece of an _Otozamites_ frond from Jurassic Italian strata in connexion with a _Williamsonia_ and the actual specimen in the Padua Museum amply justifies the impression produced by the published drawing. Wieland’s investigations[1533] in Mexico have brought to light many cases of association of _Otozamites_ fronds and _Williamsonia_ flowers.

The structure of the epidermis is partially described by Schenk[1534] and more fully by Thomas[1535]: the epidermal cells have sinuous walls and the stomata, apparently confined to the lower surface, have guard-cells with hemispherical or spindle-shaped thickened patches like those of some _Zamites_ fronds. In one species, _O. Feistmanteli_ Zig., Thomas found about 100 stomata in 1 sq. mm. of lamina forming almost contiguous lines between the veins. In the account of the genus _Ptilophyllum_ reference is made to the occurrence of pinnae with asymmetrical and auriculate bases, and it is only by the comparison of a large number of specimens that a distinction can be drawn between fronds that should be assigned to _Otozamites_ and forms of _Ptilophyllum_ which exhibit a well-marked tendency towards the _Otozamites_ type of pinna (_cf._ figs. 592, 593). The variation in the form of the apices of pinnae and the relative position of the pinnae in different parts of the same frond are features worthy of notice in the determination of species[1536]. The different appearance presented by an _Otozamites_ frond as viewed from the upper and lower face is illustrated in fig. 604. There are no recent Cycads in which the segments have auriculate bases, but in this feature as in the sinuous epidermal walls _Otozamites_ agrees with some species of the Fern _Aneimia_, _e.g._ _A. rotundifolia_ Schrad. (fig. 223, Vol. +ii.+ p. 288).

_Otozamites Bechei_ Brongniart.

There is considerable confusion in the nomenclature of this species described by Brongniart[1537] from Jurassic strata as _Filicites_ _Bechei_: in it Brongniart included a specimen from the Lias of Axminster (Dorsetshire) previously figured by De la Beche as a fossil Fern. It was on the Axminster specimen that Lindley and Hutton founded the species _Otopteris obtusa_[1538], and as there is no doubt as to the specific identity of their type-specimen (fig. 603, B) and De la Beche’s fossil, Brongniart’s designation has prior claim[1539]. _Otozamites Bechei_ is, perhaps, best regarded as a comprehensive type or a group-species in which numerous _Otozamites_ fronds described by authors, on inadequate grounds, as distinct species may well be included. Fronds agreeing generally with _O. Bechei_ were very widely spread in Rhaetic and Jurassic floras.

Specimens from the Middle Jurassic rocks of Yorkshire have been described as _Otozamites obtusus_ var. _ooliticus_[1540] to denote a slight difference in the form of the pinnae from the Liassic fronds from Dorsetshire; but the distinctive features of the variety _ooliticus_ are unimportant and hardly worthy of consideration in a general account of the species interpreted in a wide sense.

Fronds pinnate; pinnae usually more or less falcate, occasionally straight and with parallel sides, attached obliquely to the upper side of the rachis; imbricate or separate, the upper edge of the base of the lamina strongly auriculate, the lower edge rounded; apex obtuse; veins strongly divergent especially in the lobed base and extending obliquely to the upper and lower edge of the lamina.

The specimen, from the Lias of Lyme Regis, drawn in outline in fig. 605, is an almost perfect frond: the pinnae are obtusely pointed, slightly falcate, and there is a prominent lobe on the upper edge of the base of the laminae.

The Jurassic species _Otozamites graphicus_[1541] (Leck. ex Bean +MS.+), _O. vicetinus_ Zig.[1542], _O. Hennocquei_[1543] (Pom.), _O. recurrens_ Sap., _O. Terquemi_[1544] Sap., _O. linearis_[1545] Halle, are some of many examples of fronds agreeing closely with _O. Bechei_, or in the case of _O. Terquemi_ with _O. graphicus_. The fronds described by Halle from Graham Land as _O. linearis_ afford a good illustration of the range of variation in the pinnae: a characteristic feature is the considerable length, exceeding 20 cm., of the narrow linear fronds. The impressions from the Kome (Cretaceous) beds of Greenland described by Heer as _Glossozamites Schenkii_[1546] are probably closely allied to _O. Bechei_. The Rhaetic species _O. Bucklandi_ Schenk[1547], _O. indosinensis_ Zeill.[1548], and _O. Polakii_ Krass[1549], illustrate older examples of closely allied types. Zeiller records _Otozamites_ pinnae similar to those of _O. indosinensis_ from Rhaetic beds in Persia[1550].

_Otozamites Beani_ (Lindley and Hutton).

Founded on a specimen in the Scarborough Museum from the Middle Jurassic of Yorkshire[1551] which is indistinguishable from Leckenby’s type-specimen of _Otopteris mediana_[1552] in the Sedgwick Museum, Cambridge. Long and narrow fronds exceeding 20 cm. in length, characterised by the broadly oval, deltoid or sub-orbicular pinnae, separate or partially imbricate and attached by a broad auriculate base; apex bluntly rounded; veins numerous and spreading from the base of the lamina. The pinnae, which may be 3·5 cm. long and nearly 2 cm. broad, are narrower and longer in the distal part of the frond (fig. 606).

The Italian Jurassic species _Otozamites molianus_ Zig.[1553], recorded also from Bornholm, is a very similar type. The generic name _Cyclozamia_ suggested by Pomel for this form of frond has not been adopted though it is applied by Schimper to _O. Bunburyanus_ Zig.

_Otozamites Bunburyanus_ Zigno.

A Jurassic species[1554] similar in the long and narrow form of the frond to _O. Beani_ but distinguished by the much smaller segments and by their more orbicular lamina (fig. 606, B). In habit the fronds agree closely with the Fern _Nephrolepis Duffi_. Leckenby’s species, _Otopteris tenuata_[1555], is probably the same as Zigno’s _O. Bunburyanus_, the type-specimen of which in the Padua Museum consists of a long and narrow frond with leaflets not exceeding 8 cm. in length; the rachis is hidden by the imbricate auriculate bases of the leaflets. Feistmantel[1556] refers to this species some pieces of very narrow fronds with overlapping pinnae from Upper Gondwana beds in India. A still smaller form is described by Möller[1557] from Bornholm as _O. tenuissimus_, and _O. Bunburyanus_ is recorded from the same flora. _O. Feistmanteli_ Zig.[1558] agrees in the form of the frond and in its short and broad pinnae with _O. Bunburyanus_.

_Otozamites bengalensis_ (Oldham and Morris).

This Indian species from the Rajmahal Hills[1559] and specimens of the same type from the Cutch flora described as _O. contiguus_ Feist. afford another illustration of long and narrow fronds with short and relatively broad pinnae. The drawings published by Oldham and Morris accurately represent the specimens: the longest frond is 21 cm. long and neither end is complete (fig. 607); it is 9 mm. broad at the narrower end and 1·5 cm. at the broader end. The rachis is represented by a deep and broad groove; the actual bases of the leaflets are not preserved, but their position shows that they were attached to the upper face: this is clearly seen in the specimen described by Feistmantel as _O. contiguus_ which shows also that the lamina is auriculate at the upper edge of the base. The pinnae vary in shape; the smaller ones are characterised by a strongly curved lower margin and the upper edge is slightly curved or straight, while the larger leaflets have more parallel edges and blunter rounded apices: the latter form is well shown in the specimens unnecessarily distinguished by Oldham and Morris as var. _obtusa_.

_Otozamites Klipsteinii_ (Dunker).