Paleozoic Origin of the Cycads 

GEOLOGlCAL SURVEY PROFESSIONAL PAPER 934 

Paleozoic Origin of the Cycads 

By SERGIUS H. MAMAY 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 934 

M egasporophylls and associated plant parts from the 

Upper Pennsylvanian and Lower Permian Tacks of the 

southwestern United States are interpreted as a 

lineage of cycads) derived from, the pteridosper1ns. 

Cycadean foliage is thought to be derived 

from Paleozoic taeniopterids 

UI~.~ITED STATES GOVERNMENT PRINTING OFFICE.. WASHINGTON 1976 
UNITED STATES DEPARTMENT OF THE INTERIOR 

THOMAS S. KLEPPE, Secretary 

GEOLOGICAL SURVEY 

V. E. McKelvey, Director 

Library of Congress Cataloging in Publication Data 

Mamay, Sergius H. 1920-

Paleozoic origin of the cycads. 

(Geological Survey professional paper; 934) 

Bibliography: p. 

Includes index. 

1. Cycadales, Fossil. 2. Paleobotany--Paleozoic. 2. Paleobotany--Southwest, New. I. Title. II. Series: United 

States. Geological Survey. Professional paper; 934. 

QE976.2.M35 561 '.591 76-14384 

For sale by the Superintendent of Documents, U.S. Government Printing Office 

Washington, D.C. 20402 

Stock Number 024-001-02831-0 
CONTENTS 

Page 

~bstract ----------------------------------------- 1 

Introduction -------------------------------------- 1 

~cknowledgments _ ___ _ ___ _ ___ _ __ __ __ _ ___ _ __ _ ___ _ __ 3 

Ter.nUnology ------------------------------------- 4 

Sources of material ------------------------------- 4 

Previous work ------------------------------------ 5 

Systematic descriptions --------------------------- 6 

Spermopteris sp _:_____________________________ 7 

Archaeocycas Mamay -------------------------- 8 

Archaeocycas whitei Mamay ------------------- 8 

Phasmatocycas Mamay ------------------------ '12 

Phasmatocycas kansana Mamay ---------------- 12 

?Phasmatocycas sp ---------------------------- 21 

?Phasmatocycas spectabilis Mamay, n. sp -------- 22 

Systematic descriptions-Continued 

?Cycadean male cones ------------------------- Cycadospadix yochelsoni Mamay, n. sp ----------

Associated plants ---------------------------------

D:istribution of the fossils -------------------------

Morphologic and evolutionary considerations --------

Interrelationships of the ~merican fossils -------

~n alternative interpretation of Archaeocycas ---

Relationships of the ~merican fossils to other 

fossil cycads -------------------------------

Evolution of the pinnate habit in cycadalean 

foliage ------------------------------------

Conclusions --------------------------------------

References cited ----------------------------------

Index --------------------------------------------

ILLUSTRATIONS 

[Plates follow index] 

PLATE 1. Archaeocycas whitei Mamay. 

Page 

23 

24 

26 

28 

29 

29 

33 

36 

39 

43 

44 

47 

2. Archaeocycas whitei Mamay, ?Phasmatocycas sp., Spermopteris sp., Cycadospadix yochelsoni Mamay, n. sp., 

3. 

4. 

5. 

FIGURE 1. 

2. 

3. 

4. 

5. 

6. 

7. 

8. 

9. 

10. 

11. 

and ?cycadean male cone. 

Phasmatocycas kansana Mamay. 

,Seed cuticles of Phasmatocycas kansana Mamay. 

?Phasmatocycas spectabilis Mamay, n. sp., and Taeniopteris sp. 

Page 

Reconstruction of part of seed-bearing leaf of Spermopteris coriacea -------------------------------- 2. 

Diagrams showing hypothetical reduction of the megasporophyll and evolution of the ovulate cone implied by living cycads ----------------~------------------------------------------------------ 3 

Semidiagrammatic reconstruction of megasporophyll of Archaeocycas ------------------------------ 11 

Semidiagrammatic reconstruction of megasporophyll of Phasmatocycas ------------------------------ 18 

Stratigraphic correlations showing vertical distribution of late Paleozoic cycads and related plants ____ 2.9 

Diagrams showing hypothetical evolutionary development of Cycas-like megasporophyll from spermop- terid pteridosperm ___ _ ___ _ _ ___ _ __ _ _ ___ _ _ _ _ __ _ _ ___ _ ___ _ _ __ _ ____ ___ _ _ __ __ __ __ __ __ _ _ __ ___ _ ___ __ 31 

Semidiagrammatic representation of hypothetical evolution of Cycas-like megasporophyll from Archaeocycas -------------------------------------------------------------------------------------- 33 

Semidiagrammatic representation of hypothetical evolution of carpellike organ from Archaeocycas ____ 35 

Reconstruction·s of Dioonitocarpidium ------------------------------------------------------------- 37 

Reconstructions of Palaeocycas-Bjuvia ----------------------------------------------------------- 39 

Diagrams depicting stUggested evolution of cycadalean leaves from taeniopterid ancestral forms -------- 43 

III 

PALEOZOIC ORIGIN OF THE CYCADS 

By SERGIUS H. MAMAY 

ABSTRACT 

Upper Paleozoic rocks in the southwestern United States 

contain several taxa of fossil plants that appear to represent a lineage from the Pennsylvanian pteridosperms. to the 

modern cycads. The oldest genus, Spermopteris Cridland and 

Morris, is a megasporophyll that has surficial seeds borne in 

two rows along the sides of presumably abaxial fac,es of 

taeniopterid foliage; it was first found in the Virgilian Lawrence Formation of Kansas, and a poorly preserved specimen 

is now known from the Leonardian Wellington Formation of 

Oklahoma. 

The putatively derivative cycadean genera Phasmatocycas 

Mam:a.y and A rchaeocycas Mamay occur in lower Leonardian 

rocks of Kansas and Texas. P hasmatocycas (type-species: P. 

kansana Mamay) is a fertile axis that has two lateral rows 

of naked ovules. In P. kansana, the ovules are represented by 

well-preserved double cuticles similar to those of the Jurassic 

cycad Beania; small resinoid spherules, interpreted as remains 

of glands, alternate with the ovules. Taeniopterid foliage associated with P. kansana bears abundant spherules identical 

with those on the fertile axis; on the basis of the spherules 

and the known seed habit of Taeniopteris, Phasmatocycas is 

reconstructed as a cycadlike. megasporophyll with an elaminar, 

fertile base and a t31eniopterid distal lamina. The glands may 

be primitive nectaries, poss.ibly involved in an early stage in 

the development of entomophily. Phasmatocycas is regarded 

as hav.ing been derived from a spermopterid ancestor through 

reduction of numbers of seeds and loss of the basal foliar 

lamina. Althoug'h the type specimen of Phasmatocycas is from' 

the Wellington Formation of Kansas, a poorly preserved 

specimen is known from the younger Vale Formation of Texas, 

where it is associated with structures strongly reminiscent of 

male cycadean cones. 

Archaeocyca.s Mamay, typified by A. whitei Mamay, is approximately contemporary with Phasmatocycas kansana, occurring at two localities in the Belle Plains Formation of 

Texas. It resembles Phasmatocycas in having a basal s·eedbearing area and a sterile distal lamina. In Archaeocycas, 

however, the seeds are attached to the basal lamina, which 

is inrolled and at least partly encloses. the seed. A rchaeocycas 

probably represents a form intermediate between Spermopteris 

and Phasmatocycas. Alternativ·ely, Archaeocycas might be a 

fertile appendage evolving in the direction of a carpellike organ through proliferation and fusion of the laminar margins; 

this alternative, however, is not preferred. 

Associated with the cycadean megasporophylls are a single 

specimen of Cycadospadix (C. yochelsoni Mamay, n. sp.) and 

various other plants. C. yochelsoni cannot be determined as 

cycadalean or bennettitalean, although it is possible that it is 

either and that the two orders may have been distinct as 

long ago as the Early Permian. Among other plant associates, only Taeniopteris is common to all the cycad-bearing 

assemblages; this strengthens the hypothesis that Taeniopteris was the leaf of primitive cycads, as indicated by Spermopteris. 

Stratigraphic distribution of the plants supports the idea 

of derivation of the cycads from the Pennsylvanian spermopterids and of a lineage to the modern cycads through the 

Triassic genera Palaeocycas-Bjuvia and Dioonitocarpidium. 

Horizontal and vertical distribution of the Paleozoic taxa emphasizet the importance of the lower Leonardian of the southwestern United States as a temporal and geographic locale 

for rapid and dramatic plant evolution. 

Modern cycadean leaves are thought to be derived from 

taeniopterid stock in the Paleozoic and Mesozoic through 

progressive incision of the laminar margin. In one line of 

foliar evolution, shallow marginal dentation, in which each 

tooth receives only one· vein, becomes progressively deeper, 

resulting in the cycadaceous leaf type in which each leaflet has 

a :single vein; a parallel .series of known forms involved several veins to a segment and could have produced the zamiaceous leaf type·. The Stangeria leaf type could have arisen 

from a compound taeniopterid with little modification, whereas 

marginal incision of a compound taeniopterid might have 

produced the Bowenia type of leaf. 

Cumulative evidence indicates that the cycads extend back 

into the Late Pennsylvanian instead of only the Late. Triassic; 

thus they rival the conifers as the oldest extant seed plants. 

INTRODUCTION 

The tenor of this paper is appropriately established by reference to some of T. M. Harris' perceptive comments on the fossil history of the cycads. 

Harris (1961, p. 322) :wrote as follows: 

The fossil Cycads consist of a good many genera of isolated 

organs classified on more or less good evidence as Cycadales, 

a very few genera with two known organs and one genus 

with more than two organs. If we are right in our synthesis 

of these, plants with two or more· organs, and also right in our 

pre:sent classification of, say, half of the isolated organs (and 

I think it unreasonable to hope for more), then we can say 

that by Jurassic times the family had probably completed its 

evolution. * * * The known genera are no more primitive than 

the living ones ·and they do not help in linking the Cycadales 

with any other family, and for this link one looks first at the 

1 
2 PALEOZOIC ORIGIN OF THE CYCADS 

Pteriodosperms. If we are to find progressive evolution it must 

be before the Jurassic. 

Harris' remarks succinctly outline a particularly 

enigmatic deficiency in the paleobotanical record, in 

regard to which considerable speculatory discussion 

has been published. A brief supplement to Harris' 

statements is in order here as background to the 

present article, which may advance the status of our 

knowledge of the evolution of the cycads. 

The order Cycadales is a small group of gymnosperms; most authors regard it as consisting of only 

nine living genera, although Johnson (1959) recognized a tenth. The geologic history of the Cycadales 

is little known and perplexing, despite the abundant 

Mesozoic occurrences of fossil leaves generally attributed to the cycads. Intermixed and often closely 

associated with these fossils are leaves of the extinct 

Bennettitales or cycadeoids-plants that reached 

their zenith during the Jurassic and disappeared 

before the end of the Mesozoic. The distinction between the foliage of the two groups is based largely 

on features of the epidermis, and particularly on the 

stomatal characters. Otherwise, leaves of the cycadeoids and cycads are so similar that they are difficult 

to differentiate on the basis of gross external 

morphology. 

Reproductive structures of the cycadeoids were 

preserved in some abundance and have been investigated in considerable detail (Delevoryas, 1968a, b). 

However, little is known of either the polleniferous 

or ovulate mechanisms of the fossil cycads; accordingly, their phylogeny is speculative. The often-cited 

Late Triassic genera Dioonitocarpidium von Lilienstern and Palaeocycas Florin ("Bjuvia" Florin) and 

the Jurassic Cycadospadix Schimper are the oldest 

fossils regarded to represent true cycadean ovulate 

appendages. Of much more significance is the Middle 

Jurassic genus Beania Carruthers, a lax conelike 

structure with biovulate, spirally disposed peltate 

appendages. Beania is consistently compared with 

female cones of the modern zamioid cycads (Arnold, 

1953, p. 62). The Jurassic existence of cones with 

advanced morphologic features such as those found 

in Beania indicates a considerable pre-Jurassic evolutionary history for the cycads; clearly, Harris had 

this circumstance in mind while writing the comments quoted above. The Jurassic cycads, along with 

the sparse but significant evidence of Triassic relatives, nurture a reasonable speculation that true 

cycads existed during Paleozoic time. 

Harris' designation of the pteridosperms as the 

probable source of a "link" between the Cycadales 

and other taxa expresses a long-established line of 

thought, but one that has been developed on tenuous 

evidence. I feel that concrete evidence of a late 

Paleozoic cycadalean stock may now be contributed 

to the paleobotanical record. The cycadalean stock 

indeed has its beginnings in the pteridosperms, as 

speculated, and more specifically, in the taeniopterid 

pteridosperms. 

In 1960, Cridland and Morris described the genus 

Spermopteris (fig. 1), a taxonomic novelty based on 

fertile pteridospermous foliage from the Upper 

Pennsylvanian of Kansas. Spermopteris was shown 

to be the ovulate phase of a member of the formgenus Taeniopteris, representatives of which are 

common in upper Paleozoic rocks. The seed-bearing 

specimens were identified with Taeniopteris coriacea 

Goeppert, which became the basionym for the type 

species, Spermopteris coriacea ( Goeppert) Cridland 

and Morris. Spermopteris is characterized by having 

"a row of seeds borne on the abaxial surface of each 

side of the mid vein" ( Cridland and Morris, 1960, p. 

855) on what otherwise were normal-appearing appendages. The authors limited their discussion of the 

significance of their material to a few statements 

regarding the infrageneric relationships of certain 

taeniopterid forms and refrained from speculating 

on the possibly broader evolutionary implications of 

Sperm,opteris. It is my opinion, however, that Spermopteris is one of the earliest known elements of evidence bearing on the paleontologic history of the 

cycads from the standpoint of ovulate reproductive 

structures. This opinion is bolstered considerably by 

the characteristics of the Permian material disFIGURE !.-Reconstruction of part of fertile l~af of Spermopteris coriacea, showing oblique view of upper (ventral) 

surface, and at lower right, mode of seed attachment to 

lamina (from Cridland and Morris, 1960, fig. 10). Approximately X 3. 
INTRODUCTION 3 

cussed here, and considered with those of Spermopteris they form the basic theme of this paper. 

The Paleozoic plant collections of the U.S. Geological Survey contain several Permian specimens 

of ovuliferous appendages that resemble Spermopteris in the general disposition of their seeds but 

differ conspicuously in details of the foliar lamination. Also present in the collections are other forms 

that may be related and are therefore included in 

this paper. Because of the stratigraphic and geographic relationships of the two suites of specimens, 

it seems reasonable that some of the morphological 

resemblances reflect phylogenetic alliances rather 

than fortuities or parallelisms. Thus this paper is 

intended as a sequel to the Cridland and Morris work 

on Spermopteris. All these specimens have been considered critically with the purpose of stressing certain morphological features that they have in common with each other, with other fossils regarded to 

represent true cycads, and with the modern Cycadales. A certain amount of speculation is entailed in 

this presentation, but the ensuing pages will also 

describe tangible evidence for the presence, in the 

late Paleozoic record, of plants directly related to 

the living recent cycads. 

In the literature dealing with the modern cycads, 

ev0lutionary advancement of the various genera is 

usually interpreted in terms of comparative morphology of the female cones. The megasporophylls of 

Cycas are held to be the most primitive ovulate organs of the order, because they retain leaflike features indicative of foliar derivation and are produced like leaves in loose terminal crowns instead of 

in compact cones (fig. 2; Chamberlin, 1965, p. 145-

150). Because they are produced like leaves they are 

also shed like leaves, as individual units rather than 

as more or less complete cones. Thus one would 

logically look for individual detached fossil megasporophylls in searching for primitive fossil cycads 

or their progenitors. This assumption is borne out 

here, inasmuch as the material on which this study 

is based consists. entirely of single detached appendages that provide no evidence regarding modes of 

attachment to the parent plants. Fortunately the 

individual cycadalean megasporophyll, though simply constructed, is sufficiently diagnostic that the incomplete nature of the fossils treated here does not 

detract substantially from their phylogenetic 

significance. 

ACKNOWLEDGMENTS 

The late David White is in no small measure responsible for this paper, because he collected much 

~ 

~ D 

FIGURE 2.-Hypothetical reduction of the megasporophyll and 

eVIOlution of the cone· as implied by living cycads. A, Leaf? 

like megasporophyll of Cycas revoluta; F, loose terminal 

crown of megasporophylls (C. revoluta), not compacted into 

a cone. B, Megasporophyll of Cycas media, the leaflets reduced to serrations; G, terminal aggregate of megasporphyUs of same sp·ecies, in a more conelike arrangement. C, 

Megasporophyll of Dioon edule, in which the lamina is entire-margined; H, loosely compacted cone of D. edule. D, 

Megasporophyll of Macrozamia miquelii, the lamina reduced to a spine; I, tightly compact cone of same species. 

E, Highly reduced megasporophyll of Zamia floridana, having no resemblance to a leaf; J, extremely compact cone of 

Z. floridana. (All from Chamberlain, 1930, p. 290, with 

slight modifications of figure' explanations.) 

of the critical material. Although he published nothing known to refer directly to this material, his 

curatorial markings on specimens and certain other 

kinds of evidence show that he was aware of the importance of some of the material described here and 

almost certainly suspected its cycadlike affinities. 

Thus White's considerable influence on the Paleozoic 

paleobotany of North America continues, even now, 

to exert itself. 

Thanks are extended to Dr. J. A. Wilson, University of Texas, for information regarding the Patterson locality at Lawn, Tex., and to C. 0. Patterson 

for permission to collect on his property. Ralph 
4 PALEOZOIC ORIGIN OF THE CYCADS 

Howe and Mart Russell of Seymour, Tex., kindly permitted me to collect on the "Emily Irish" land during 

several field seasons. T. R. Tinsley of Stamford, Tex., 

gave me some of the specimens described and showed 

me the fossil plant deposit in Haskell County. A. D. 

Watt, U.S. Geological Survey, assisted me in the 

field in 1961, 1963, 1967 and 197 4. During my first 

collecting trip to the Lawn locality, I was accompanied by C. B. Read and E. L. Yochelson, both of 

the U.S. Geological Survey. Read also gave me valuaable instruction in general procedures in locating 

fossiliferous deposits in the southwestern Permian 

red beds. 

TERMINOLOGY 

The literature dealing with cycadophytes is somewhat confusing with regard to the terminology applicable to the foliar organs. Some authors prefer 

the terms "leaf" and "leaflet," whereas, others use 

"front" and "pinna," or "frond-like leaf" ; occasionally the alternative terms are even used interchangeably in the same publication. Thus, this area of terminology seems basically to involve a matter of 

personal choice, especially in routinely descriptive 

approaches. I prefer "leaf" to "frond," particularly 

because this study is fundamentally concerned with 

entire-margined taeniopterid foliar organs of unknown origins or homologies. 

My use of the term "megasporophyll" for the fertile structures discussed here is commonly accepted 

usage, yet it invites a statement of qualification because of an article by Meeuse (1963), in which the 

traditional interpretation of the ovuliferous appendage of Cycas was challenged. 

Sporne (1965, p. 111-112) presented a consensus 

on the evolutionary status of Cycas, reiterating the 

substance of Chamberlain's hypothesis as presented 

in figure 2 of this paper. Sporne wrote "It is widely 

accepted by morphologists that the Cycas sporophyll is the primitive type among cycads, and that, 

during the evolution of the group, there has been a 

reduction in the number of ovules to two * * * ." 

Meeuse (1963, p. 122), however, presented the interesting but improbable hypothesis that the ovulate 

appendage of Cycas is "an· organ of dual natp.re combining axial and foliar characteristics" and the "socalled 'megasporophyll' represents an abnormally 

developed whole' cycadalean strobilus." In attempting to demonstrate that "the conventional identification of the female reproduction organ of Cycas with 

a 'megasporophyll' (defined as a leaf homologue 

bearing marginal ovules) is fallacious," Meeuse invoked unconvincing evidence, such as structural 

aberrancy or resemblance of the ovuliferous organ 

of Cycas to a teratological case. He furthermore rejected Palaeocyca.~ Florin as a cycad, primarily on 

the basis of its entire-margined taeniopterid leaves. 

Because of the taeniopterid form of the lamina of 

SpeTmopteTis, its clearly megasporophyllar nature, 

and the evolutionarily simple processes of reduction 

of parts and incision of foliage necessary for derivation of the Cycas type of fertile appendage, Meeuse's 

hypothesis is herewith rejected in favor of traditional concepts. I regard the term "megasporophyll" as 

correctly applicable to Cycas and use it here 

accordingly. 

The ovulelike bodies on the basal part of the lamina in A1·chaeocycas are described as attached to the 

lower (abaxial) surface of the megasporophyll, but 

the choice of this word-in preference to the contrasting term "upper" adaxial-is an arbitrary one, 

based on homology with the abaxial positons of the 

sporangia of most ferns. The few specimens available are detached, and thus it is impossible to determine their orientation in relation to the apex of the 

parent plant. Even though proof is lacking, however, the specimen shown in lateral view in plate 2, 

figure 7, lends the strong impression of being oriented in its natural attitude, that is, with its adaxial 

surface toward the top of the illustration. 

SOURCES OF MATERIAL 

The specimens described here came from eight 

localities in Lower Permian outcrop areas, one in 

Oklahoma, three in Kansas, and four in Texas. The 

localities are described below, the numbers referring 

to U.S. Geological Surv~y paleobotany localities; 

quotation marks indicate parts of locality descriptions that were taken from the U.S. Geological Survey (USGS) locality register: 

6233: "St. L. & S. F. Ry. in cut % mile west of station at 

Perry, Oklahoma. Coli. David White, Sept. 6, 7, 8, 

1911. Permian." According to the geologic map of 

Oklahoma (Miser, 1954) this locality is within an 

undivided Permian rock interval indicated as "Pwa" 

(including the Admire Group through the Wellington Formation). As far as I am aware, no subsequent collections have been recorded from this locality. 

8298: "Elmo limestone. Wellington formation,. Permian. Insect Hill, 3% mi. SE of Elmo, Kansas, Barn lot on 

S. side of road, and adjacent localities. Coli. C. B. 

Read, 1931." The Elmo Limestone Member of Dunbar (1924) of the Wellington Formation is of Early' 

Permian (Leonardian) age. I have not visited this 

area. 
SOURCES OF MATERIAL 5 

8868: "Carleton, Kansas. Wade Sterling Ranch, 4 miles south 

of Carleton, Dickinson Co., Kansas (Wellington Fm.). 

D. White, 1909." 

8869: "3% miles south of Banner, Kansas (Elmo Is.). Coli. 

by David White, 1909." This collection was probably 

made very close to locality 8298, and almost certainly is from the same rock unit. 

8877: "Wichita formation, 4';4, miles south-ea:st of Fulda, 

Texas. Collected by D. White." This locality is 

within the Lower Permian Belle Plains Formation 

(Romer, 1958,. map facing p. 178), and although the 

locality register and specimen labels lack the date of 

the collection, a brief discussion of the collecting 

activities by White in his paper on Gigantopteris 

americana ( 1912, p. 495) indicates that the collection was made in 1910. White wrote' "the exact locality being the bank of the stream at the crossing of 

the old road, one·-fourth miles south of the ford of the 

Little Wichita River, 4 miles southeast of Fulda, a 

station in Baylor County." In 1955 and 1957, I attempted to relocate this important fossil deposit, but 

the vague nature of the locality description and the 

lack of distinctive physiographic features in this 

badlands terrain made it impossible to be certain 

of one's position. Furthermore, the plant-bearing 

shale is easily eroded and the original deposit probably has been totally destroyed or obscured. A few 

nondescript fossil plant fragments were· found in the 

supposed vicinity of White's locality, but none of 

these equaled White's material in quality of preservation. 

8959: "Baylor County, Texas.. East-central part of Emily 

Irish grant, south side of Salt Fork of Brazos River, 

16 to 16% miles scaled due S.E. of Seymour. Shale 

in upper part of Belle Plains Formation. Coli. Read 

and Ervin. October, 1940." This locality is on prop-:-

erty now owned by Roland Howe, Fort Worth, Tex., 

and the plant deposit is by far the richest and most 

diversified one known in the Permian of the southwestern United States. I have collected from it on 

five different occasions, in 1955, 1957, 1961, 1963, 

1967, and 1974. Actually, the plant deposit is at 

about the middle, rather than the upper part, of the 

Belle Plains Formation and is regarded as the approximate equivalent of the Valera Shale. A detailed 

locality description and a measured section of the 

"Emily Irish" exposure have been published ( Mamay, 1968, p. I2-I3). 

10057: C. 0. Patterson property, Taylor County, Tex. The 

site is reached by starting at the theater in Lawn, 

Tex., going 1.5 miles (2.4 km) west, 2.9 miles ( 4.8 

km) ·south, 1.1 (1.8 km) miles west, and 0.5 (0.9 

km) mile south to the remains of the old home of 

C. 0. Patterson. Fossils were found in a quarry and 

its vicinity, in a pasture against the east slope of 

a low hill as viewed west-southwest from the west 

side of the house. Fossil plants were especially 

abundant in a small lens of light gray shale about 

100 yards (915 m) north of the quarry, and considerable material was collected in April 1955 by 

C. B. Read, E. L. Yochelson, and myself. According 

to Meyers and Morley ( 1929b), the Lawn deposits 

are in the lower part of the Vale Formation, in the 

Lower Permian Clear Fork Group. (See Wilson, 

1953, p. 456-459 for further discussion of this locality.) In 1957, A. D. Watt and I revisted this locality, hoping to obtain additional material. We found 

that the Patterson home had recently been destroyed 

by fire and was then in the process of being rebuilt 

on another site, and our collecting plans were completely thwarted because the pasture in which the 

fossil deposits occurred had been deep-plowed as 

part of a mesquite-eradication program. Small 

chips of the plant-bearing matrix and the underlying caliche bed were scattered over a fairly extensive area, but the plant beds may now be regarded as having been destroyed and lost to science. 

10064: Haskell County, Tex. This locality is approximately 

3 miles (4.8 km) north of Stamford, Tex., and just 

north of the Jones-Haskell County line. To reach 

the locality, go north and northwest on U.S. Highway 380, starting at the city square in Stamford. 

After 2.25 miles (3.7 km) turn north on an unpaved crossroad; approximately 1 mile (1.6 km) 

north, plant fossils occur in red and gray, mediumto fine-grained sandstone which is exposed in the 

road shoulders and nearby pastures along both 

'sides of the road. According to Meyers and Morley 

1929a), exposures in this area are within the 

boundaries of the Vale Formation (Lower Permian 

Clear Fork Group) , and their geographic proximity 

to other Vale exposures would suggest a high position in the Vale F·ormation. The presence of plant 

fossils in these beds was told to me by T. R. Tinsley 

of Stamford, and in his company, A. D. Watt and 

I collected plants there in 1961, 1963, and 1967. 

PREVIOUS WORK 

This study is limited to Paleozoic cycadean fructifications and their possible affinities with the early 

Mesozoic cycads, and because only a few examples 

are known, there is no need for an extensive review 

of past investigations of all fossil cycad parts. Several examples of petrified stems and other vegetative 

remains thought to have cycadean affinities are 

known from the Paleozoic. These are largely from 

the Old World, and their occurrences were summarized adequately by Arnold (1953, p. 61), who 

speculated that "There were probably some cycads 

in existence during the middle or late Carboniferous and by the end of the Paleozoic era they were 

fairly numerous,." 

The existence of true cycads in North America 

during Late Triassic time was demonstrated by 

Delevoryas and Hope (1971), who described the new 

genus Leptocycas from the Pekin Formation of 

North Carolina. The material consists of leaves and 

stem parts, both with typical cycadean cuticular 

characteristics; also found was a structure resembling a male co~e, attached to a stem. Unfortunately, nothing is known of the ovulate parts. Delevoryas 
6 PALEOZOIC ORIGIN OF THE CYCADS 

and Hope's reconstruction of Leptocycas gracilis 

shows a sparsely foliate plant with pinnate leaves 

and a slender trunk with long internodes. Stem 

features of Leptocycas differ from the modern cycad 

trunks which have dense armors of closely spaced 

persistent leaf bases. 

Gould's (1971) anatomical .study of petrified stem 

material from Upper Triassic beds of Arizona and 

New Mexico provided additional important evidence 

of cycads in the American Triassic. This investigation centered upon the taxon Lyssoxylon grigsbye 

Daughtery from the Chinle Formation. Lyssoxylon 

was originally regarded as a member of the Bennettitales (Daughtery, 1941), but various anatomical 

characters led Gould to reinterpret it as a true cycad. 

The first recorded hint of Paleozoic cycads in 

North America appeared in a list of Permian plants 

published by White (1912, p. 508). White listed the 

taxon ucycadospadix? sp." from a locality supposedly near Carlton, Kans. This listing was not substantiated by an illustration or description, but I 

found in White's collections a specimen that I believe to have been the basis for that identification; 

it is probably the most important specimen involved 

in this study. 

In 1916, Bassler described a specimen of Plagiozamites planchardi (Renault) Zeiller from a coal seam 

in the Upper Pennsylvanian Conemaugh Group in 

West Virginia; this species is known from several 

Lower Permian localities in the Old World but had 

not been reported previously in North America. 

Originally Plag·iozamites was regarded as a true 

cycad leaf, although conclusive proof has never been 

found. Bassler's material neither substantiates nor 

refutes a cycadean affinity for the genus, but the 

Pennsylvanian occurrence is noteworthy. 

The present paper culminates several years of 

study and six collecting trips. My interest was 

initially aroused by the presence of cycadlike specimens in White's Permian collections from the southwestern United States and was further stimulated 

by Cridland and Morris' ( 1960) description of 

Spermopteris. My ideas were announced in four 

brief publications (U.S. Geological Survey, 1968; 

Mamay, 1969a, 1971, 1973), in which the concept of 

late Paleozoic origin of cycads from Pennsylvanian 

spermopterids was set forth. 

One of my articles (Mamay, 1969a) was immediately preceded in the same journal by an article by 

Taylor (1969a), in which a new type of polleniferous cone was described from Upper Pennsylvanian 

coal ball petrifications from Illinois. Taylor (p. 294) 

identified his material as Pennsylvanian cycads with 

the statement "The fossil record of true cycads is 

extended from the Upper Triassic to the Upper 

Pennsylvanian" ; he ( 1969b) later reiterated this 

interpretation. In a subsequent publication, Taylor 

(1970) named the cones Lasiostrobus polysacci, at 

the same time modifying his interpretations so as to 

permit consideration of the possibility that the cones 

are coniferalean rather than cycadalean. Taylor's 

reason (1970, p~ 682) for moderation of his originally unequivocal viewpoint was the discovery of additional specimens, some showing features of the apex. 

Taylor's altered stance emphasizes the questionable taxonomic position of Lasi.ostrobus and the necessity for its reevaluation. I regard it as coniferophytic in the bulk of its characteristics and reject it 

as a cycad. 

In my preliminary announcements (Mamay 

1969a, b), I said that two new genera were recognized in this investigation. Formal names were not 

proposed ; the genera were merely designated "new 

genus A" and "new genus B." Sporne (1971) discussed the new plants briefly, drawing attention .to 

their as yet incomplete nomenclatural status; he 

also introduced an incorrect finality into some of my 

then-tentative ideas on angiosperm origins and accredited me with the discovery of "The origin of the 

carpel" (Sporne, 1971, p. 14). So that these possibly controversial fossils might be discussed with 

legitimate names, I proposed and validated the generic names Archaeocycas and Phasmatocycas (Mamay, 1973). 

SYSTEMATIC DESCRIPTIONS 

Four genera are recognized and discussed in this 

paper. Two of these-Spermopteris Cridland and 

Morris, 1960, and Cycadospadix Schimper, 1870-

are represented by one specimen each. The genus 

Archaeocycas Mamay, 1973, is represented by eight 

specimens ; both counterparts of three of these are 

present. Phasmatocycas Mamay, 1973, is based on 

one specimen, both counterparts of which were 

found; a number of other specimens are provisionally assigned to this genus. Inasmuch as the formalization of the names Archaeocycas and Phasmatocycas in 1973 was an expedient designed to make 

names available while full descriptions and discussions were in preparation for this paper, the diagnoses are repeated here in the interest of 

completeness. 

With the possible exception of Cycadospadix, the 

specimens above are clearly of megasporophyllar 

morphological nature. They are presented in the con-
SYSTEMATIC DESCRIPTIONS 7 

text of an evolutionarily interrelated complex of 

plants, which collectively appears to fill an important hiatus in the early history of the cycads. Two 

other plant structures, one foliar and one possibly a 

microsporangiate cone, are discussed because of 

their possible pertinence to the theme of this 

investigation. 

Division PTERIDOSPERMOPHYT A 

Order PTERIDOSPERMALES 

Genus SPERMOPTERIS Cridland and Morris, 1960 

Spermopteris sp. 

Plate 2, figure 2 

DESCRIPTION 

This identification is based on one small specimen, 

poorly preserved on a chip of greenish fine-grained 

shale from the Perry, Okla., locality. The specimen 

represents a taeniopterid leaf fragment 3.0 em long 

and 11.0 mm wide. It is preserved as an impression 

with only a few minor traces of adherent carbonaceous matter, which is restricted to the midrib. 

The midrib is conspicuously marked with parallel 

longitudinal striations and is very broad in proportion to the breadth of the lamina, measuring about 

2.5 mm in width. The leaf fragment does not appreciably diminish in width from one end to the other. 

The venation is poorly preserved, but the delicate 

ultimate veins may be seen by careful observation of 

a few small p~rts of the lamina. The veins are closely spaced, occasionally forked, and gently decurrent. 

Insofar as observable, the venation of this specimen 

is characteristically taeniopterid. 

The chief significance of this spe.cimen is in the 

presence of two uninterrupted rows of closely contiguous swellings that are roughly oblong or ovoid in 

shape and of very minor relief. One row lies along 

each side of the midrib, but the swellings are best 

seen along the right side of the specimen (pl. 2, fig. 

2). The swellings average about 2.5 mm long and 1.5 

mm wide. The basal ends of the swellings are not 

clearly separable from each other lengthwise and 

substantially coalesce to form a fairly conspicuous, 

uninterrupted but slightly undulating trough parallel to the midrib and about halfway out in the lamina 

on each side of the midrib. The sides of the swellings 

form shallow parallel depressions and generally follow the decurrence of the lateral veins. The swellings 

show no evidence of extending beyond the edges of 

the lamina; rather, they appear to be truncated at 

the leaf margins. 

DISCUSSION 

I am satisfied that this fossil represents a specimen of Spermopteris, probably preserved before the 

seeds had matured. Several features of the Oklahoma specimen suggest a congeneric relationship 

with material of Spe\-mopteris that was regarded by 

Cridland and Morris to represent immature ontogenetic stages. In one of their specimens ( Cridland 

and Morris, 1960, fig. 2), the positions of supposedly 

immature seeds are clearly shown, but the general 

proportions of the specimen are somewhat smaller 

than those of the Oklahoma specimen. Cridland and 

Morris illustrated a second specimen ( Cridland and 

Morris, 1960, fig. 9, p. 859), which they cited as a 

fertile leaf seen from the adaxial side, with "swellings which indicate the positions of immature seeds." 

This specimen and the Oklahoma specimen compare 

favorably in such characters as width of the leaf; 

proportional widths of lamina and midrib; density 

and attitude of the secondary veins; size, shape, and 

distribution of the seeds; and termination of the 

seeds apices exactly at the margins of their respective parent laminae. To judge from the indistinct 

seed outlines in the Oklahoma specimen, it may be 

assumed that the surface of this specimen represents 

the upper or adaxial surface of the leaf and, as in 

the case of Cridland's and Morris' specimen, the 

thickness of the foliar tissue obscured the details of 

the seeds in the fossils. 

The Oklahoma specimen conforms with the generic diagnosis of Spermopteris in having a row of 

seeds or seedlike bodies on each side of the midrib 

of a taeniopterid leaf, but specific alliance cannot be 

attempted because of the limitations of material. 

However, a few remarks about specific identifications are added here because there is some evidence 

that this particular specimen was given more than 

passing attention by David White. 

In discussing the relationships of Spermopteris, 

Cridland and Morris (1960, p. 859) commented on 

Von Gutbier's ( 1835, p. 73) original description of 

Taeniopteris abnormis, wherein he mentioned but 

did not illustrate specimens with shallow crosswrinkles (."flache Querfalten") toward the tips of 

the leaves. Apparently Cridland and Morris regarded this condition as a possible point of comparison with Spermopteris but did not pursue the idea 

further because of the unavailability of additional 

information. White (1912, p. 507) published a provisional list of plants from the Perry, Okla. locality. 

This list includes Taeniopteris multinervis Weiss and 

T. abnormis Gutbier, but no indication was given of 

the number of specimens of each that were present. 

The original collection contains just nine specimens 

of Taeniopteris, mostly very badly preserved. As the 

Oklahoma Spermopteris specimen is the only taeni-
8 PALEOZOIC ORIGIN OF THE CYCADS 

opterid in that suite that can be distinguished from 

the others on morphological grounds, it seems likely 

that White had studied this very specimen, equated 

its seedlike swellings with Gutbier's "flache Querfalten," and identified T. abnormis in the Oklahoma 

collection. In the same paper, White ( 1912, p. 506) 

also listed T. abnormis among the plants collected at 

the Fulda, Tex. locality (USGS locality 8877), but I 

have not been able to ascertain the basis for that 

identification. 

This occurrence extends the known geographic 

range of Spermopteris from eastern Kansas to 

north-central Oklahoma. At the same time, the 

stratigraphic range of the genus is extended upward 

from the Upper Pennsylvanian (Virgilian) to the 

Lower Permian, where it is associated with Gigantopteris americana in flora zone 14 (Read and Mamay, 1964). 

Occurrence.-USGS paleobotany locality 6233. 

Figured specimen.-USGS 6233-1 (pl. 2, fig. 2). 

Division CYCADOPHYTA 

Order CYCADALES 

Genus ARCHAEOCYCAS Mamay 

New genus B. Mamay, 1969b. 

Archaeocycas Mamay, 1973. 

Type spec1~es.-Archaeocycas whitei Mamay. 

Generic diagnosis.-Bilaterally symmetrical fertile appendage (megasporophyll) with broad midrib; several pairs of apparently sessile, closely appressed ovules borne in two lateral rows on surface 

of reduced basal part of lamina; ovules closely invested by lamina, each with a small circular scar of 

attachment ta lamina near midrib. Distal part of appendage expanded as flattened sterile foliar blade. 

Archaeocycas whitei Mamay 

Figure 3; plate 1; plate 2, figures 5-7 

Archaeocycas whitei Mamay, 1973, p. 687, fig. 1, a-c. 

Specific diagnosis.-Megasporophylls cuneiform, 

ovate, or lanceolate, 1.7-2.3 em long, with greatest 

widths of 1.0 em. Midribs straight, flat, faintly 

marked with closely set parallel striations; midrib 

widths uniform or increasing distally from 1.0-1.3 

mm at the bases to 3.0 mm at distal termini of fertile areas ; midribs flaring out conspicuously or becoming obscure in distal, laminar parts of megasporophylls; sides of midribs more or less concavely 

scalloped through appression with ovule bases. 

Proximal fertile area 1.2-1.6 em long, 3.0-5.0 mm 

wide at base, nearly uniform in width or increasing 

distally to width of 9.0 mm. Ovules produced in opposite pairs, four to six ovules in each row. Ovules 

oblong, oblanceolate, rhombic or rhomboidal, tightly 

appressed with each other, obliquely directed forward at broad angles to the rachis. Basalmost ovules 

proportionately shorter, broader than distal ones; 

ovules 1.5-3 mm long (measured along lines of appression between contiguous ovules), 1.5-3.0 mm 

wide (measured along outer free margins and along 

lines of appress,ion with the rachis), reaching 5 mm 

in greatest (diagonal) dimension. Ovules each with 

a small circular shallow depression or attachment 

scar, 0.7-1.0 mm in diameter, uniformly placed on 

the upper surfaces of ovules near the center of area 

of ovular appression with the midrib. Upper (adaxial) surfaces of ovules closely invested by foliar 

lamina ; extent of lamina over lower (abaxial) ovular surfaces unknown; laminar investment of ovules 

showing fine closely spaced parallel striations perpendicula~ to midrib. 

Megasporophylls abruptly modified into sterile 

laminar structures immediately distal to distal pair 

of ovules, and contiguous with ovules; extent, venation, and margination of distal laminae obscure. 

Holotype.-USGS 8877-1 (pl. 1, fig. 5). 

Paratypes.-USGS 8877-2, 3, 4 (pl. 1, fig. 3, 4; 

pl. 2, fig. 7) ; USGS 8959-1, 2 (pl. 1, fig. 1, 2; pl. 2, 

fig. 5, 6). 

Occurrences.-USGS paleobotany localities 8877, 

8959. 

DESCRIPTION 

This description is based on eight specimens, six 

of which were collected by White at the Fulda locality (USGS paleobotany locality 8877) in 1910. Although I could not find White's locality, I was fortunate in discovering two additional specimens of 

Archaeocycas at the "Emily Irish" locality (USGS 

paleobotany locality 8959) in 1963; the two associated floras are substantially the same and presumably occupy the same stratigraphic position within 

the Belle Plains Formation. Although White never 

published an account of his specimens, it is obvious 

that he recognized them as unique or important, because I found them carefully trimmed, encircled in 

crayon marks, and segregated from the remainder 

of his Fulda collection in a separate tray. 

The specimens are preserved as compressions with 

a moderate amount of relief that aids somewhat in 

their interpretation. Only small amounts of carbonaceous residue remain, but enough of this is 

present to provide some critical information. Three 

of the Fulda specimens are casts (pl. 1, figs. 4, 5 ; pl. 

2, fig. 7), two are molds (pl. 1, figs. 1, 3), and the 

remaining one is available as both counterparts. 
SYSTEMATIC DESCRIPTIONS 9 

Each of the two "Emily Irish" specimens consists of 

both counterparts; these are shown in plate 1, figures 1 and 2, and plate 2, figures 5 and 6. Unfortunately no cuticularized material is preserved; matrix 

samples from both localities were processed for palynological analyses, with negative results. 

The specimens are each characterized by a fertile 

basal area, which is neatly and symmetrica.lly "compartmented" (pl. 1, figs. 4, 5; pl. 2, fig. 7), and a 

distal, usually poorly preserved laminar extension. 

A superficial examination of this material might 

prompt the conclusion that the paired inflated objects borne along the basal extent of the midrib are 

vegetative pinnules, but close inspection necessitates 

another interpretation. If these were ordinary pinnules rather than casts and molds of expanded threedimensional bodies, the abrupt transition from a 

basal area of distinct pinnulation to the rather 

vague, poorly defined distal lamination would contrast markedly with the conventional type of pinnate foliar appendage. Again, if these were ordinary 

pinnules, one might expect some overlapping or 

other irregularities of attitude such as is found ordinarily in pinnate plant fossils. Instead, these bodies 

show no sign of overlapping but present the appearance of firmly attached solid objects of considerable 

depth; they abutted laterally against each other, 

with the result that contiguous sides were noticeably 

flattened (pl. 1, figs. 4, 5; pl. 2, fig. 7). In the absence of compelling alternative evidence, these objects are regarded as the fillings of ovules, probably 

buried in an immature stage of development; consequently, the term "ovule" is applied herewith. 

The casts are the most informative of the available specimens. Although most of the casts are lacking in carbonaceous residues, most of which adhered 

to the molds when the specimens were split, all the 

casts show remnants of a substantial thickness of 

coalified organic material separating contiguous 

ovules. These films, which average about 0.15 mm in 

thickness, are clearly seen on plate 1, figures 4 and 5, 

and plate 2, figures 6 and 7. The holotype (pl. 1, fig. 

5) is particularly interesting in this respect. When 

tilted and viewed edgewise, the ovules are seen to 

reach thicknesses of 1.0 mm or slightly more, and the 

presence of coaly films beneath the edges of the 

ovules indicates that the ovules are actually the sedimentary casts or mud fillings of hollow bodies that 

were of sufficient substance to resist compaction. 

Most of the specimens were deposited with their flat 

surfaces normal to the surface of the enclosing sediment; the specimen shown in plate 2, figure 7, however, was deposited edgewise on the bedding surface 

and therefore gives some idea of the thickness of the 

ovules, which are seen nearly end-on in a side view 

of the sporophyll. 

The coaly films separating the ovules from each 

other are interpreted as the organic residue of part 

of the investing lamina, which originally was deeply 

creased so as to intrude downward between the 

ovules. When the sporophylls were compressed, the 

foliar tissue was "pinched" between the flattening 

and laterally expanding ovules and is now preserved 

as a thin coalified film. The remainder of the foliar 

residue adhered to the counterparts, or molds; there 

the grooves between ovules lack any coaly residue. 

This distribution of organic residue between counterparts of a given specimen is well demonstrated on 

plate 2, figures 5 and 6. 

The scalloped outlines of the midribs are also delimited by a thin coaly film that separates the ovules 

from the midribs (pl. 1, figs. 4, 5; pl. 2, fig. 6). Tilting and edgewise observation of these specimens 

show that the coaly film continues downward and 

toward the median axis of the midrib. Thus the midrib seems to have been compressed into a flangelike 

form, partly covering the upper surface of the foliar 

lamina in the fertile region of the sporophyll. Compression of the midrib probably caused an entrapment of foliar tissue between the rachis and ovules, 

similar to that between the ovules; the result is a 

gently curved longitudinal continuity of the coaly 

films separating contiguous ovules (pl. 2, figs. 5, 6) . 

These features favor the interpretation of these 

basal objects as originally distended tightly appressed bodies rather than flattened vegetative pinnules. They are emphasized in order to justify the 

diagnosis of the basal objects as ovules. Unfortunately, none of the specimens are exposed in bottom 

(abaxial) vi~w, so that details of the opposite sides 

of the sporophylls and ovules are not known. 

The ovules are arranged in opposite pairs. The 

specimen shown on plate 2, figures 5 and 6, has only 

four pairs, the smallest number of ovules observed. 

Some specimens have five pairs each (pL 1, figs. 3-

5), and others have six, the largest number observed 

(pl. 1, figs. 1, 2; pl. 2, fig. 7). The shape of the ovules 

varies somewhat, those at the basal end of the sporophyll being proportionately the longest in the direction parallel to the axis of the sporophyll ; these variations in size and shape of ovules are shown on plate 

1, figure 5. The overall length of the fertile areas of 

the sporophylls ranges from about 1.2 em in the 

smallest specimen (pl. 2, fig. 5) to 1.6 em in the 

largest (pl. 2, fig. 7). The aggregate of ovules on a 

given sporophyll niay be of an almost uniform width 
10 PALEOZOIC ORIGIN OF THE CYCADS 

(pl. 1, fig. 3) or may show a wedgelike shape, the 

width increasing in the distal direction (pl. 1, figs. 4, 

5). 

Many of the ovules have a single small circular 

depression or scar; these range from 0.7 to 1.0 mm 

in diameter. Each of the eight ovules shown on plate 

2, figures 5 and 6, has one of these marks; these are 

most clearly shown on the mold (pl. 2, fig. 5). The 

uppermost pair of ovules on plate 1, figure 4, shows 

these marks as distinct, fairly deep pits. Two others 

are seen on plate 2, figure 7, where they are partly 

filled with a carbonized residue. The scars are uniformly placed, next to the midrib at approximately 

the middle of the ovule. The scars on the specimen 

shown on plate 2, figure 5, are of additional morphological interest because each one has a centrally located circular scar, approximately one-third of its 

own diameter. 

Because of their occurrence, one to an ovule, these 

marks are interpreted as remnants of points of attachment to the lamina. There is no evidence of a 

stalk, so the ovules are regarded to have been sessile, 

with a small point of attachment; the subsidiary 

marks on the scars shown on plate 2, figure 5, may 

represent points of entry of vascular strands into 

the bases of the ovules. The position of these' marks, 

relative to the lamina and rachis, is essentially identical with the position of seed attachment in 

Spermopteris, as reconstructed by Cridland and Morris (1960, fig. 10). 

Evidence of a laminar element in the ovuliferous 

area of the sporophyll is variously shown in the 

suite of specimens but is best seen on the specimens 

illustrated on plate 1, figures 1 and 2. These are 

counterparts of the same sporophyll; figure 1 shows 

the mold, figure 2, the cast. 

The mold has retained much of the coalified residue of the specimen, and many of its details are 

relatively obscure. However, the right half of the 

photograph shows not only the grooves separating 

the ovular impressions, but also a series of closely 

spaced parallel ridges and grooves, oriented at a 

broad angle decurrently to the midrib. These are indicative of a system of fine foliar venation. 

The cast shows an important additional detail, 

indicated by the arrow in plate 1, figure 2. In this 

specimen, the grooves between contiguous ovules are 

clearly shown, but the most distal groove is incomplete; that is, it does not -extend entirely to the margin of the lamina, as do all the others. This is evidently a function of the fact that this particular 

groove is beyond the most distal ovule, and the absence of the pressure of a contiguous ovule permitted 

the marginal part of the involute lamina to flatten 

out and merge with the distal sterile lamina. This 

demonstrates a continuity of lamina between fertile 

and sterile parts of the sporophyll. 

On the basis of the foregoing, it is well established 

that the ovules of Archaeocycas were borne on a 

laminar blade, which seems to have at least partly 

enclosed the ovules. The evidence bearing on extent 

of the enclosure is incomplete, however. 

All but one of the specimens show remnants of a 

distal foliar lamina immediately above the uppermost pair of ovules. The longest lamina is shown on 

plate 2, figure 7, where it extends about 6 mm beyond 

the last ovule, and constitutes nearly one-third of the 

total length of the specimen. The complete extent of 

the lamina or features of its margination are not 

known. As shown on plate 1, figures 3-5, and plate 2, 

figures 5-7, it forms as a distal continuation of the 

midrib, which normally flares out rather abruptly; 

the striations of the midrib also diverge to present a 

vague suggestion of a fanlike system of fine veins. 

The specimen shown on plate 2, figures 5 and 6, 

differs from the others in that the strong midrib 

essentially retains its identity as far as its broken 

distal end, and the lamina evidently decreases in 

width distally, instead of expanding as all the others 

do. The lamina shows faint striations, decurrently 

directed away from both the midrib and the ovules, 

so that the orientation of the photograph seems to 

be correct; that is, the morphologically proximal end 

of the specimen is directed downward on the plate. 

This conclusion is corroborated by comparisons of 

the shapes and decurrent attitudes of the ovules of 

this specimen with all the other specimens. The 

acute distal tapering of the lamina seems anomalous 

when compared with the other specimens; this, however is not an accident incurred when the matrix 

was split, because the laminar margin along the upper left of the photograph is entire and intact. Thus, 

in this particular specimen the distal laminar extension was significantly smaller than those seen in the 

other specimens; and perhaps this one is nearly complete except for its very tip. 

.Another variation in the form of the distal lamina 

is seen in the holotype (pl. 1, fig. 5). In this specimen, the distal lamina is very broad in proportion to 

the widest part of the fertile area. This is indicated 

by a conspicuous folding of the lamina, which is seen 

in the photograph to begin in the middle of the midrib at a level approximately even with the uppermost pair of ovules. The resultant crease p·roceeds 

obliquely toward the upper right of the photograph, 

meanwhile deepening and continuing as far as the 
SYSTEMATIC DESCRIPTIONS 11 

\ 

\ 

1 

I 

J 

broken distal edge of the lamina. This folding or 

distortion had no noticeable effect on the symmetrical 

arrangement of the two rows of ovules below; the 

intactness of the symmetry may be interpreted to 

mean that the ovuliferous part of the appendage was 

quite rigid and capable of resisting distortion. The 

absence of the lowermost ovule on the right side of 

this specimen is due to breakage during splitting of 

the matrix, for when the specimen is tilted at a high 

angle and viewed from the side, remnants of the 

broken base of the missing ovule are evident. 

There is no extensive information regarding the 

manner of attachment of Archaeocycas. A minor bit 

of evidence, however, is seen at the basal end of the 

specimen shown on plate 2, figure 7. At the left end 

of the specimen, the broken end of the midrib increases abruptly in thickness, and remnants extend 

toward both the top and bottom of the figure. This 

indicates that the base of the sporophyll occurs immediately next to the proximal pair of ovules, and 

the sporophyll therefore had no extended stalk. It 

also suggests connection to a supporting axis whose 

orientation would parallel the left edge of the plate. 

In consideration of the fact that this particular 

specimen was laterally compressed in preservation, 

the basal enlargement of the rachis might be taken 

to indicate that the parent axis was perpendicular 

not only to the direction of the midrib but to the 

natural plane of the lamina as well. Thus the 

Archaeocycas specimen represents a lateral appendage equivalent to a leaf, rather than a subdivision of 

a compound leaf. 

A partly diagrammatic restoration of a sporophyll 

of A1·chaeocycas is presented in figure 3. 

DISCUSSION 

Cycadean affinity of Archaeocycas.-Morphological interpretations and taxonomic assignment of 

Archaeocycas revolve about consideration of the following unique combination of structural features, 

which, insofar as I know, occurs in no other Paleozoic plant and validates the status of Archaeocycas 

as a new genus: 

1. It is a bilaterally symmetrical flattened leaflike 

appendage. 

FIGURE B.-Reconstruction of megasporophyll of Archaeocycas 

whitei, showing upper (?adaxial) surface. Fertile bas_al 

part of megasporophyll rendered transparent to show ovules 

(broken lines) ; darkened circular areas represent points 

of attachment of ovules to lamina. Shape and extent of 

sterile distal lamina hypothetical. For view of lower surface of megasporophyll see figure 7A. 
12 PALEOZOIC ORIGIN OF THE CYCADS 

2. It is clearly differentiated into two areas, the distal one which is sterile, and the basal one which 

is fertile. The basal or fertile part of the lamina is somewhat reduced, and on its lower surface it bears two lateral rows of large paired 

conspicuous bodies that are interpreted as 

ovules; the fertile lamina seems to have invested the ovules tightly and was involute so as 

to at least partly enclose the lower surfaces of 

the ovules. A simple small circular scar occupies the same relative position on each ovule. 

The small circular scars on the fertile laminae are 

difficult to interpret in terms other than scars of attachment; likewise, the large size of the bodies 

attached thereto strongly suggests ovules. Additional evidence in support of their ovular nature is found 

in their resemblance to the seeds of Spermopteris 

and the manner in which the bodies were borne. The 

crowded closely appressed arrangement and shapes 

of the ovules of Archaeocycas are probably a function of immaturity at the time of preservation, and 

a function of the way in which the ovules were enclosed by the foliar lamina. When the plant matured, 

the fertile base of the sporophyll probably elongated 

rapidly to accommodate the increasing size of the 

ovules. The fertile lamina may have ceased to grow 

as the rachis and ovules enlarged, with the result 

that only inconspicuous distorted remnants of the 

fertile lamina remained to maintain the attachment 

of ovules to sporophyll. Thus, mature sporophylls, if 

found fossilized, could be difficult to recognize as 

later ontogenetic stages of the material upon which 

this description of Archaeocycas is based. 

Phyllospermy:--the condition in which seeds were 

borne directly upon more or less unmodified leaves 

(Sahni, 1920, p. 300)-is a phenomenon that became 

well established during late Paleozoic time. Several 

examples appear repeatedly in the literature as the 

result of important work by Halle ( 1929). Spermopteris is a clear example of phyllospermy; the Texa~:~ 

Permian has recently produced two distinctive but 

probably unrelated forms (Padgettia Mamay, 1960; 

Tinsleya Mamay, 1966) , and now Archaeocycas may 

be added to the growing list. Archaeocycas stands 

apart from all other fossils of this general nature in 

having its ovules restricted to the basal part of the 

sporophyll. From the standpoint of symmetry of 

ovular insertion, striking comparisons may be drawn 

between Archaeocycas and Spermopteris, but the 

latter genus shows no evidence of sharp segregation 

of the reproductive function to the basal part of the 

sporophyll. The trend toward reduction of the basal 

part of the lamina, as demonstrated in Archaeocycas, 

is another important point of distinction between 

Archaeocycas and Spermopteris. A continuation of 

that trend obviously could result in a megasporophyll 

much like that of Cycas, and for this reason I regard 

Archaeocycas as an extremely significant stage in 

the evolution of the cycads. In recognition of these 

Permian fossils as representatives of a precursive 

cycadean stock, I have therefore applied the generic 

name Archaeocycas. The species A . whitei is named 

for David White, who found most of the specimens 

on which the description is based. 

Genus PHASMATOCYCAS Mamay 

New genus R. Mamay, 1969a. 

Phasmatocycas Mamay, 1973. 

Type species.-Phasmatocycas kansana Mamay. 

Generic diagnosis.-Fertile axis with two lateral 

rows of sessile, broadly attached gymnospermous 

ovules; ovules simple, with blunt, funnel-shaped 

micropyles, two distinct cuticular layers, and a thick 

megaspore membrane; laminar tissue unknown, interseminal appendages lacking. 

Pbasmatocycas kansana Mamay 

Figure 4; plates 3, 4 

Phasmatocycas kansana Mamay, 1973, p. 689, fig. 1, d-g. 

Specific diagnosis.-Axial fragment 2.5 em long, 

relatively stout, 2.5-4.0 mm wide, rather abruptly 

broader at (?)basal end of specimen; surface of axis 

with obscure narrow longitudinal striations, lacking 

other characteristic markings. 

Bilateral arrangement of ovules symmetrical ; no 

evidence of ovules between lateral rows. Ovules inserted alternately to suboppositely, extending essentially perpendicularly to axis, closely arranged as to 

juxtapose against but not overlap each other. Ovules 

2.8-4.0 mm long, 2.0-2.6 mm wide, ovoid to oblong, 

broadly attached with point of attachment to axis 

nearly as long as greatest width of ovule; ovular 

apices blunt, some specimens with a shallow apical 

cleft or notch. 

Cuticular envelopes slightly smaller than impressions of ovules, lacking ornamentation; cuticles apparently complete except for micropylar openings; 

apices of cuticles gradually decreasing in diameter, 

without marked constriction, terminating in micropylar openings 0.2-0.5 mm wide; specialized chalaza! 

features absent. Outer cuticle very thin, diaphanous, 

with elongate cells reaching 20 by 80 JLm (micrometres) in dimensions, parallel to long axis of ovule. 

Inner cuticle apparently thicker and separate from 

outer; cells thick walled, walls abruptly increasing 
SYSTEMATIC DESCRIPTIONS 13 

in thickness toward the micropylar area; cells in 

micropylar area essentially isodiametric, averaging 

10 ~-tm in diameter. Megaspore membrane essentially 

filling cavity within cuticles; membrane dense, lacking ornamentation. 

Small (0.25-0.33 mm in diameter) spherical sessile bodies with resinlike luster regularly alternating 

with ovules in each row; one such body partly imbedded in axial tissue at point between and approximately level with bases of two adjacent ovules. 

Holotype.-USGS 8869-1 (pls. 3, 4). 

Occurrence.-USGS paleobotany locality 8869. 

DESCRIPTION 

Phasmatocycas kansana is based on one specimen 

from the Banner, Kans., locality. Fortunately, both 

counterparts of the specimen are available, and although only a short fragment of the axis was found, 

it is sufficiently well preserved that some important 

details are observable. The specimen is preserved 

in a matrix of very hard dark-gray limestone of poor 

fissibility and apparently not very rich in plant 

material. 

Both counterparts of the specimen are shown on 

plate 3 (figs. 1, 2). As the axis is broken at both 

ends, it is impossible to ascertain what the total 

length of the fertile structure was, but it may have 

been considerably more than the 2.5 em preserved 

here. 

The axis itself lacks any conspicuous morphological features, and only the fact that one end of the 

fragment is somewhat broader than the other gives 

one any basis for differentiating between the distal 

and proximal ends; I assume that the wider end 

(4.0 mm) is proximal and the narrow end (2.5 mm) 

is distal. Parts of each of the axial counterparts 

carry flecks of a fairly thick carbonaceous residue, 

but no traces of a cuticle are preserved. 

Faint, closely spaced longitudinal striations appear in a few small areas on the axial surface; these 

are distributed over nearly the entire width of the 

axis and probably are best interpreted as striations 

caused by wrinkling of the original surface or as 

impressions of the vascular system of the axis. There 

are no traces of a foliar lamina between or projecting 

beyond the apices of the attached seeds, and, in consideration of the generally good preservation of the 

specimens, it may be concluded that this fragment is 

indeed lacking in any sort of foliar lamina. Indirect 

evidence regarding foliation o.f Phasmatocycas is 

presented on subsequent pages. 

The most impressive feature of Phasmatocycas is 

its two lateral rows of seeds, the near-perfect alinement of which attests to the bilateral morphology 

of the appendage and precludes any interpretation 

of the seed insertion as a spiral system. Furthermore, the axial surface between the two rows of 

seeds shows no evidence whatsoever of scars of seed 

attachment that might provide a basis for challenging a bilateral interpretation for the seed arrangement of Phasmatocycas. 

The seeds are mostly preserved as flattened molds, 

each of which contains a double seed cuticle and 

megaspore membrane on one or the other of its 

counterparts. The outlines of the molds are usually 

slightly wider and longer than the contained cuticles. 

This preservational feature is well shown in the detached seed at the left of figure 1, plate 3; here the 

mold is about 2.5 mm wide and the enclosed cuticle 

is slightly less than 2.0 mm wide. The consistent 

presence of these flattened, slightly hollow seed cavities in the matrix surrounding the completely flattened cuticles suggests that the outermost layers of 

the seeds were fleshy integumental layers that were 

not adapted to fossilization. These integuments did 

resist compaction to a certain extent, however~ and 

resulted in production of the slight cavities in the 

limestone, left there as molds when the integumental 

tissues disintegrated. Flakes of fairly thick coalified 

residue on the surfaces of some of the cuticles attest 

to the original presence of such an integumental 

layer. 

In general proportions, most of the seed molds are 

less than twice as long as they are broad ; their broad 

bases and blunt apices thus lend them a nearly oblong shape in most instances. Most of the seed bases 

are only slightly narrower than the widest parts of 

the seeds, and the actual point of attachment may 

even slightly exceed the width of the seed. The 

broadly sessile nature of the ovules is best seen on 

plate 3, figure 4. The same figure shows another interesting, although perhaps not especially significant 

feature of the seed molds-a shallowly bifid or 

notched apex that appears in several of the specimens. These notches range form 0.2 to 0.3 mm in 

depth and tend to give one the initial impression that 

these are the molds of platyspermic seeds, because 

several previously described genera of supposedly 

platyspermic seeds have apical notches. Cridland and 

Morris ( 1960) described and illustrated similar 

apical notches in the seeds of Spermopteris. They 

speculated (Cridland and Morris, 1960, p. 857) that 

the apical cleft "may have been caused by fracture 

of the micropyle during compression or may be a 

natural feature of the seeds." In the case of Phasmatocycas, I prefer to interpret the apical clefts as a 
14 PALEOZOIC ORIGIN OF THE CYCADS 

mechanical result of compression rather than a 

natural feature of the seeds. This seems to be a preferable interpretation because no consistent feature 

of the cuticles indicates a cleft micropyle in 

P hasmatocycas. 

Associated with the axis are 21 seeds; one is detached but the others are preserved in their original 

positions of attachment. It is not possible to ascertain whether or not they were arranged in morphologically opposite pairs, because some are subopposite or slightly alternate to each other, perhaps as 

a result of slight preservational distortion of position. Nearly all the seeds, however, extend outward 

from the axis at right angles (pl. 3, figs. 1, 2) and 

give the impression that they were firmly attached. 

They are so close to each other that their sides are in 

physical contact with each other, but there is little 

overlapping. The seeds are essentially the same size 

and in the same condition of preservation from one 

end of the specimen to the other. Thus it would seem 

that all the ovules matured simultaneously. 

All the seed cuticles are, in part, only loosely adherent to the limestone matrix. Consequently it was 

possible to remove three nearly complete cuticles and 

parts of two others. These were cleared of coalified 

remnants of integumental tissues and other humic 

components by the standard palynological clearing 

technique, using a mild Schultze's solution followed 

by washing and treatment with potassium hydroxide. This method cleared the specimens sufficiently 

for micruscopic examination. The cuticles were then 

mounted on glass slides in a synthetic resin mounting medium. Some of the specimens are shown on 

plate 4. The evidence for an ovulate nature of the 

inflated bodies in Archaeocycas is partly inferential, 

but Phasmatocycas has genuine gymnospermous 

seeds. The description of the cuticles on the following pages leaves no room for doubt. 

The best cuticular specimen, shown on plate 4, 

figure 3, is nearly complete except for the absence 

of a large patch of material within the almost-intact 

margin of the lower half of the seed. This specimen 

is 3.2 mm long and 1.8 mm wide. The broken speci- · 

men shown on plate 4, figure 4, is 3.3 mm long, and 

in its entirety it must have been at least 2.2 mm 

wide. Another nearly complete specimen, not illustrated, is 2.0 mm wide, but its length is not known. 

This one is the broadest of the cuticular specimens 

removed from the matrix ; other larger specimens 

are still attached to the axis. 

As shown on plate 4, figure 3, the megaspore membrane is a heavily cutinized thick envelope. It is variously folded and wrinkled, as indicated by the dark 

stripes where two or more thicknesses of the membrane are involved. Thus, the light-colored patch 

below the middle of the seed shown on plate 4, figure 

3, represents a single layer of megaspore membrane; 

the darker area above it consists of a double layer, 

and the still darker stripes represent folded areas of 

four layers. Although the chalaza! outlines of the 

megaspore membranes are quite plain, their apical 

contours are difficult to differentiate because of optical interference by the thickened surrounding cuticles in the micropylar areas of the seeds. On plate 4, 

figures 4 and 5, however, the dim outline of the apex 

of the megaspore is seen passing over the upper part 

of the seed approximately 0.5 mm below the seed 

apex, or the area that presumably constituted the 

bottom of the pollen chamber. Although the megaspores are heavily cutinized membranes, they do 

not have any conspicuous or characteristic 

ornamentation. 

It is clear from the apical areas of the seeds that 

two distinct cuticles are present. This is shown in 

all of the photographs on plate 4. The cuticles are 

difficult to differentiate through the entire length of 

the cleared specimens, however, and they may not 

have been sufficiently developed in the basal parts of 

the seeds to be preserved. In the specimen shown on 

plate 4, figure 3, the faint outlines of cuticular cells 

are observable down to about the midsection of the 

seed; in that area, only one layer is visible, however, 

and the differentiation into two layers is not apparent below approximately the apical 0.75 mm of 

the seed. The specimen shown on plate 4, figure 4, 

seems to be the best prese·rved from this standpoint, 

for cellular outlines of the isodiametric type that 

characterize the inner cuticular layer are visible 

proximally about as far as the median part of the 

seed. 

At about the level of the "shoulders" of the seeds, 

or 0.7 to 0.8 mm below the s.eed apices, the inner 

cuticle thickens abruptly, whereas the outer cuticle 

remains very thin and barely visible in many places. 

Apparently the outer cuticle is separate from the 

inner; the distinction between the two layers is 

shown on plate 4, figures 1 and 5. Most cells of the 

outer cuticle are elongate, reaching lengths of 50-80 

fLm, and have widths of one-fourth or less of the 

length; these cells are oriented with their long axes 

parallel to the length of the seed. The isodiametric 

cells of the inner cuticle average about 10-15 !Lm in 

any direction; their walls are very thick; particularly 

in the area immediately above the distal surface of 

the megaspore membrane. As shown on plate 4, figures 1 and 5, this wall thickening decreases abruptly 

in the terminal 0.2-0.3 mm of the micropyle. 
SYSTEMATIC DESCRIPTIONS 15 

The micropylar narrowing of the cuticular envelope is shown on plate 4. All the illustrated specimens 

show a gentle tapering of the apical "funnel." The 

actual openings of the micropylar canals range from 

0.2 to 0.5 mm in width. The tips of the micropyles 

shown on plate 4, figures 1, 2, and 3, are slightly 

damaged, but the one shown on figures 4 and 5 is 

essentially intact and shows no evidence of a bifid 

tip. If any specialized pollen-receiving structures 

were within the seed apices, no evidence of such is 

preserved in the fossils. The present material permits only the assumption that the seeds of Phasmatocycas had a simple pollen chamber, accessible 

through a short, evidently unspecialized micropylar 

canal, which was delimited by the thick inner cuticle. The cleared specimens show no evidence of internal vasculature or pollen grains within the micropylar areas. 

Interpretation of the morphological nature of the 

two cuticles of P hasmatocycas is aided by reference 

again to two of the preservational features of these 

seeds. First, the presence of flat cavities or seed 

molds in the matrix, somewhat larger than the cuticular outlines, indicates that a thick integument 

originally surrounded the seeds. This deduction is 

supported by the second pertinent feature, the presence of thick flecks of coalified organic residue on 

the outer surfaces of the cuticles. Thus, the absence 

of any trace of an outer integumental cuticle, which 

should have directly lined the seed molds, indicates 

that the thin outer cuticles of the cleared specimens 

represent the inner cuticular lining of the integument. Accordingly, the thickened inner cuticles are 

interpreted as the surface of the nucellus. 

These interpretations are supported to some extent by Harris' ( 1941) study of the seeds of the 

Jurassic cycad Beania. Attention is directed to a 

comparison of the cleared specimens of Phasmatocycas (pl. 4) with Harris' (1941, pl. 5, fig. 14) 

preparation of the inner cuticles of a seed of Beania 

gracilis. In both instances, very similar sets of thickened membranes are preserved. The Beania contains 

a thick megaspore membrane surrounded by a relatively thick cuticle (nucellar), which in turn is enclosed by a thin inner integumental cuticle. The 

"apical spike" of Harris' Beania is not present in 

Phasmatocycas, nor does the nucellar cuticle of the 

Bean~a flare upward into the conelike projection 

present in Phasmatocycas. These differences in details of apical morphology of the two genera of 

seeds reduce the exactness of comparison, but the 

similarities point up the fact that the Phasmatocycas 

seeds are similar to those of the Jurassic cycads. 

The excellent preservation of seed cuticles suggested the possibility that a palynological examination of the matrix might corroborate a cycadalean 

interpretation of Phasmatocycas. Chips of the limestone were submitted to R. A. Scott who reported 

(written commun., 1968) as follows: 

The search for palynological evidence for the presence of 

cycadophytes was unsuccessful. Only a very few apparently 

monosulcate grains were seen; these were considered as unreliable because they 1) could be folded fern spores or 2) 

resembled the pollen of gingkophytes more closely than that 

of cycadophytes. 

Because of poor preservation, these results do not in any 

way preclude the presence of cycadophyte pollen; they simply fail to establish it. 

One other feature of Phasmatocycas invites close 

attention. Both counterparts of the holotype bear 

either the impressions or the actual substance of a 

number of tiny, almost perfectly spherical bodies, 

which measure 0.25-0.33 mm in diameter. Where 

the actual spherule is present on one counterpart, the 

impression is usually visible on the opposite counterpart. These bodies do not seem to have been compressed or distorted from a spherical shape during 

the process of preservation. The spherules are dark 

brown to black, and although most have lustrous surtrous surfaces, faint rugosity is evident on some 

specimens. Some are translucent and reddish brown 

suggestive of a resinoid composition. 

One might first be tempted to interpret these 

spherules as a fortuitous concentration of foreign 

objects not related to Phasmatocycas. However, their 

distribution indicates that they are not all randomly 

placed, which necessitates the conclusion that the 

spherules are a natural part of the fertile axis. 

A few of the spherules are present on the axis in 

random positions, whereas a few others are in the 

matrix a short distance away from the specimen. 

The most significant aspect of their distribution, 

however, is the fact that many of the spherules occur 

on the axis at the bases of the seeds, and most of 

these occupy positions exactly between two adjacent 

seeds ; none occur directly on the seeds. This regular 

arrangement is shown on plate 3, figures 3, 4, and 

5. Plate 3, figure 3, shows this feature particularly 

well. Such a positional relationship between the 

seeds and spherules can hardly be accidental. Rather, 

it indicates that the spherules are a natural feature 

of Phasmatocycas and, inasmuch as they are intercalated between adjacent seeds, they should be approximately equal in number to the 21 seeds found 

on the type specimen. By actual count, the specimen 

contains the remains or impressions of 24 spherules. 
16 PALEOZOIC ORIGIN OF THE CYCADS 

The slight disparity between numbers of spherules 

and seeds may reflect an originally weak attachment 

of spherules to the axis, some having been displaced 

from their natural positions and shifted to other 

locations during burial of the specimen. 

If one accepts the distributional pattern of the 

spherules as evidence that they were originally attached to the axis of P hasmatocycas in two rows like 

the rows of seeds, one spherule placed between each 

two successive seeds, one then faces the problem of 

interpretation of these objects. 

Because of their uniformity in size, shape, and 

distribution, the spherules are best interpreted as 

the remnants of regularly placed glands; the spherules represent the coagulated acellular contents. 

These hardened before burial and retained their 

naturally spherical shapes during induration of the 

matrix. Regardless of their functions, the glands 

constitute a point of comparison with associated leaf 

remains; this comparison warrants further elaboration because of its inferential bearing on the 

morphological identity and systematic placement of 

P hasmatocycas. 

DISCUSSION 

Evidence for a megasporophyl!ar interpretation 

of Phasmatocycas.-Phasmatocycas kansana shows 

no direct evidence of attachment to a foliar lamina. 

Because of the strikingly cycadlike nature of this 

fertile organ, however, it is well to examine closely 

whatever evidence is available in the direction of establishing or refuting a megasporophyllar character 

for Phasmatoc'ycas. Certain compelling evidence is 

indeed present in the form of associated taeniopterid 

leaves bearing resinoid spherules that seem identical 

with those on the ovuliferous axis of Phasmatocycas. 

The small assemblage of plants associated with the 

holotype of P hasmatocycas kansana contains seven · 

fragments of narrow taeniopterid leaves, most of 

which are poorly preserved; the largest specimen is 

about 10 em long. All these specimens bear either the 

impressions or the actual substance of small spherules, which compare closely in size with those of the 

fruiting axis. Their preservation is mostly inferior 

to preservation of the glandular remains 011 Phasmatocycas, but in two of the leaves, the spherules are 

identical with those of the fertile specimen in color 

and surface features. Distribution of the spherules 

or glands varies; in some of the leaves, the glands 

are more or less evenly scattered over the surfaces, 

but in one specimen they are concentrated near the 

margins of the lamina. Only a few of the glands appear directly on the midribs. 

Another small collection from a nearby site also 

has taeniopterid material containing resinoid spherules. This collection (USGS paleobotany locality 

8298) was made "31j2 mi. SE of Elmo"; the sources 

of this material and of the holotype of Phasmatocycas (USGS paleobotany locality 8869; "31/2 mi. S. of 

Banner") are probably the same outcrop, because 

Dunbar (1924, p. 174) pointed out that Elmo was "at 

one time called Banner City." Lithologic comparisons suggest that the two collections were made from 

the same rock unit. 

This collection contains nine fragments of taeniopterid foliage, which is not distinguishable from the 

leaves from locality 8869. Most of these leaves are 

poorly preserved, but some show evidence of at least 

a few of the spherical bodies. One of these specimens, 

by contrast, is quite well preserved and is remarkable because of the large number of resinoid glandular remains or hemispherical impressions borne on 

its surface. This specimen is shown on plate 5 (figs. 

5-7). 

Plate 5, figure 5, shows the entire specimen, which 

is 6.5 em long and 1.4 em wide at its broadest point. 

The basal part of the specimen is missing, but the 

fragment clearly shows a stout midrib and simple 

typically taeniopterid venation. 

Plate 5, figure 6, a higher magnification, shows 

the many small spherical bodies, more or less evenly 

distributed over the entire lamina. A small part of 

the lamina is shown at 20 magnifications on plate 5, 

figure 7. The glands average about 0.25 mm in diameter and are so abundant that most of them are within 0.5 mm of another. They are restricted to the narrow laminar areas between the closely set veins, and 

the midrib itself is devoid of glands. The spherules 

are easily removed from the matrix, and many of 

them were detached, presumably when the specimen 

was collected. The positions of missing sphe,rules, 

however, are clearly indicated by hemispherical depressions in the intercoastal areas. Most of the 

glandular bodies are virtually uncompressed, and in 

their general physical characteristics they are remarkably similar to those attached to Phasmatocycas kansana. 

Excellent examples of the glandular remains also 

appear on taeniopterid foliage contained in USGS 

collection 8868, from a locality near Carleton, Kans. 

Attention was originaly directed to spherical bodies on taeniopterid foliage by Sellards (1901, 1908). 

In 1901, Sellards described some Permian taeniopterid material from the Banner City locality; this 

included specimens identified as Taeniopteris coriacea Goeppert, T. coriacea var. linearis, Sellards 
SYSTEMATIC DESCRIPTIONS 17 

var., T. newberriana Fontaine and I. G. White, and 

T. sp. Much of that paper was devoted to description 

of "interneural bodies" and "scars on or near the 

rachis" on the foliage. The scars were compared with 

insect damage and fungal remains, but no definite 

interpretations were adopted. 

The "interneural bodies" were described (Sellards, 

1901, p. 5) as being "small, oval, resistant bodies, 

situated between the veins, half immersed in the epidermis of the frond, nearly globular in shape, some 

smooth on top, but more often showing a slit across 

the top." To judge from Sellards' illustrations, those 

bodies are identical with the spherules described 

here, with the exception of the distal slits; I have not 

observed such slits in the material at my disposal. 

Sellards briefly compared these bodies with dots on 

other genera from other localities. In attempting to 

analyze them in terms of morphology and function, 

he appeared to favor interpretation of the bodies as 

sporangia or a new type of fructification; in his subsequent paper (Sellards, 1908, p. 447), however, he 

regarded the question of function of the objects as 

an equivocal matter. Cridland and Morris (1960, p. 

859) commented that the bodies "have never been 

shown to be anything but globules of iron oxide." 

Much of Sellards' paper of 1901 was incorporated 

without alteration in his subsequent summary article 

on the late Paleozoic plants of Kansas (Sellards, 

1908). Although no new or supplemental information regarding the taeniopterid complex or the "interneural bodies" was presented, that paper provided 

important information regarding the biological significance of the spheres. Sellards described 26 species 

(among nine genera) of plants represented by foliar 

organs, from the Banner City locality. "Interneural 

bodies" were discussed only in connection with the 

taeniopterids in the Banner City flora; comparable 

objects were not noted on any of the associated taxa. 

This consistency of association is borne out by my 

observations. The genera Callipteris, Glenopteris, 

and Odontopteris are present in the material I have 

examined from the vicinity of Banner City, but these 

show no evidence of spherical bodies. 

From the foregoing discussion, one can scarcely 

avoid the deduction that the spherical objects are a 

natural biological attribute of the taeniopterid leaves 

from the Elmo Limestone Member of Dunbar (1924) 

at or near the Banner City plant locality. Their consistency of size, shape, and placement on the lamina 

between veins militates against their origin as accidents of mineralization or organic degradation. They 

are most likely glandular in origin, and their re--

stricted occurrences on just one foliar form genus in 

a rather diverse generic assemblage demand scrutiny 

of possible relationships between these leaves and 

any associated plant remains on which similar 

glandular bodies occur. 

The resemblances between the glands of Phasmatocycas kansana and those of the associated taeniopterid leaves lead to examination of the obvious 

inference, that is, that the fruiting axis and the 

leaves are parts of the same species of plant, if not 

the same morphological unit. Several circumstances 

add support to this inference; these and previously 

noted evidence are summarized here: 

1. The glands are not found on any of several associated genera, although many of the associated taeniopterid leaf specimens show evidence 

of their presence. 

2. Neither the occurrence nor distribution of the 

glands on Phasmatocycas and the associated 

taeniopterid leaves is reasonably ascribable to 

accident; these objects are doubtless a biologically controlled phenomenon. 

3. The cycadlike fundamental morphology of Phasmatocycas requires only a terminal foliar part 

to complete the qualitative morphological requirements of a cycadean megasporophyll. 

4. A seed-bearing taeniopterid leaf (Spermopteris 

Cridland and Morris) is already known; its 

geographic and stratigraphic occurrences and 

morphologic details make it a logical predecessor for a Phasmatocycas-like appendage with 

terminal taeniopterid foliation. 

5. The putative Triassic cycads Dioonitocarpidium 

Lilienstern and Bjuvia Florin both have been 

supposed to have had taeniopterid foliage. 

Viewed as an aggregate, these points of circumstantial evidence provide the basis for a tentativ~ 

reconstruction, in which Phasmatocycas is presented 

as a cycadalean megasporophyll with a taeniopterid 

foliar lamina distal to the basal seeds. This is shown 

in figure 4. 

Cycadean affinity of Phasmatocycas.-Phasmatocycas and Archaeocycas resemble each other fundamentally in their bilateral symmetry, each bearing 

two lateral rows of paired seedlike objects. In the 

case of Archaeocycas, substantial indirect evidence 

indicates that these objects are the remains of seeds, 

partly enclosed by a reduced lamina and attached to 

it a short distance from the rachis. In Phasmatocycas, on the other hand, the bilaterally paired objects 

are unequivocably the remains of gymnospermous 

seeds, as established by their well-preserved megaspore membranes and double cuticles. 
18 PALEOZOIC ORIGIN OF THE CYCADS 

P hasmatocycas shows no development of a lamina 

in the ovuliferous region, so that if it is considered 

in the light of a reduction series, Spermopteris being 

the hypothetical ancestor, it would represent the ultimate stage in elimination of a foliar element in the 

fertile area of the appendage; the result would entail direct attachment of the seeds to a naked midrib. 

Only one specimen of Phasmatocycas is available, 

and absolute proof of attachment of a terminal foliar 

lamina is lacking. However, if my appraisal of indirect evidence and my resultant reconstruction of 

Phasmatocycas are correct, then Phasmatocycas may 

be regarded as the perfect fulfillment of a theoretical 

reduction series, beginning with the completely laminar Spermopteris and possibly involving Archaeocycas as an intermediate form. (See fig. 6). 

Detailed comparisons of Phasmatocycas with other fossil gymnosperms in order to derive taxonomic 

conclusions would be pointless because no close similarities exist among the fossils. The seed-bearing 

groups known to exist in the Permian include the 

pteridosperms, the early conifers ( cordaiteans and 

walchians), and the ginkgophytes, none of which 

constitutes a logical taxonomic receptacle for Phasmatocycas. This fossil shows no evidence of the cupular seed investment, complex pollen-receiving mechanism, or other critical features of the pteridosperms. A pteridospermous alliance for P hasmatocycas may be dispensed with, except in the sense that 

it was probably derived from the pteridospermous 

Spermopteris. 

It is equally obvious that Phasmatocycas is no 

cordaitean or conifer. It lacks any evidence of the 

complex investiture of spirally arranged bracteate 

appendages that appear in cordaitean cones. It has 

no resemblance to the short shoots involved in the 

evolution of coniferous cone scales or of Ginkgo-like 

fructifications. There likewise appears to be no reason to postulate a Bennettitalean alliance for Phasmatocycas; Harris' (1954, p. 290) observations on 

the absence of a cutinized megaspore membrane in 

the cycadeoids, considered against the well-developed 

membrane in Phasmatocycas, weigh heavily against 

an already remote point of possible speculation. 

The remaining approach lies in comparisons of 

qualitative features of Phasmatocycas with other 

gymnosperms, irrespective of relative ages. Such an 

FIGURE 4.-Reconstruction of megasporophyll of Phasmatocycas. showing bilaterally arranged basal ovules, alternating glands, and taeniopterid distal lamina. Relative length 

of distal lamina reduced; venation and glands on lamina 

shown less dense than in actual specimens. From Mamay, 

1969a, figure lA. 
SYSTEMATIC DESCRIPTIONS 19 

examination reveals that, in points of fundamental 

morphology, Phasmatocycas is more like a megasporophyll of the modern genus Cycas than any other gymnospermous fructification, living or fossil. In 

all the living cycads, the ovules (usually only two) 

are bilaterally arranged, but Cycas usually has two 

or more pairs of ovules, indicating a multiovulate 

ancestral megasporophyll. Phasmatocycas at least 

hypothetically performs that role. 

The seeds of Phasmatocycas were apparently of 

simple construction and are more like those of the 

cycads than of the pteridosperms or cordaites. The 

cuticular details of P hasmatocycas are sufficiently 

similar to those of the Jurassic cycad B eania to 

warrant increased emphasis of the other cycadlike 

features of Phasmatocycas. 

In its overall qualitative aspects, then, the one extant specimen of Phasmatocycas is remarkably like 

a megasporophyll of Cycas. Only the terminal leaflike proliferation of the Cycas megasporophyll is 

missing from Phasmatocycas, but evidence for its 

presence, already discussed, is substantial. Conversely, such a terminal proliferation is indeed present in 

Archaeocycas, which I believe to represent an evolutionary form just one step below Phasmatocycas 

in this primitive cycadalean lineage. 

The name Phasmatocycas derives from the modern 

generic name Cycas and the Greek "phasma," meaning apparition or specter. 

In connection with a cycadean interpretation of 

Phasmatocycas, it is of interest that White's (1912 

p. 508) list of provisional plant identifications from 

the Permian of Kansas includes reference to ucycadospadix? sp., C," the "C" referring to Carlton, 

Kans. (according to the USGS paleobotany locality 

register, this is locality 8868, "4 miles south of Carlton"). The Carlton collection contains nothing that 

might suggest a cycadean structure; conversely, 

P hasmatocycas is the only cycadlike specimen in the 

Banner collection, but White's list indicated nothing 

cycadlike from Banner. In attempting to determine 

the exact location of the Banner plant beds, certain 

facts have been found that may explain the above 

inconsistency and that consequently may identify 

White's "Cycadospadix? sp." as the specimen described here as the holotype of Phasmatocycas. 

The town of Banner no longer exists under that 

name; its site is now occupied by the town of Elmo, 

which formerly was less than a mile west of Banner 

on the Missouri Pacific Railroad. Carlton still exists 

and is about 3 miles west of the former town of 

Banner. Collection 8868 was made "4 miles south of 

Carlton," and collection 8869 was made "31/2 miles 

south of Banner." As Banner was just 3 miles east 

of Carlton, the two fossil localities were less than 4 

miles apart. The collections could have been, and 

probably were, made the same day (collecting dates 

for the two localities are given only as "1909" in the 

USGS locality register). Parts of the two collections 

are lithologically indistinguishable, and it thus is 

possible that the two collections were mixed and mislabeled, and that White consequently identified a 

cycadlike fossil from Carlton instead of Banner. The 

two collections are very similar, and I am reasonably 

certain that White had in hand the holotype of Phasmatocycas when he made the identification of "Cycadospadix? sp.". If this analysis is not correct, then 

an important part of the Carlton collection was lost 

and White failed to notice this important specimen 

in the Banner· collection. Either alternative seems 

unlikely, and the combination of circumstances is 

even more difficult to envisage. 

Function of the glands.-The glandular bodies are 

so regularly placed between the seeds of Phasmatocycas and so abundant on the associated taeniopterid 

leaves-putatively the distal laminae of megasporophylls-that speculative teleological comments are 

in order. The function of the glands will likely remain conjectural at best, but the important biologic 

process of entomophily is indeed suggested by certain indirect evidence. 

The presence of glandlike bodies on taeniopterid 

leaves is not a ne·w idea, for Halle's (1927) memoir 

on Paleozoic plants from Shansi, China, included 

descriptions of two species of Taeniopteris that had 

"black dots" between the veins. Halle ( 1927, pl. 37, 

fig. 1) also showed similar bodies on the type specimen of T. multinervis from the Permian of Saxony. 

Halle regarded the dots as probably glandular but 

made no comments on their possible functions. In a 

later paper, however, Halle (1929) presented a 

lengthy and pertinent discussion of that possibility 

of entomophily in the Paleozoic; his argument was 

based largely on the presence of glandular bodies on 

other kinds of foliage, but it may be applied to Taeniopteris as well. 

Halle's discussion (1929, p. 18-23) aimed primarily at trends in seed position in the pteridosperms, but 

modes of pollination were also considered. He wrote 

as follows: 

There remains the possibility of insect pollination. Oliver 

and Scott, in their famous memoir 'On the structure of the 

Palaeozoic seed Lagenostoma Lomaxi etc.' draw attention in 

this connection to the capitate glands found on the cupule of 

Lagenostoma Lomaxi and on various parts of Lyginopteris oldhamia. Following a suggestion of the late Sir Joseph Hooker, 
20 PALEOZOIC ORIGIN OF THE CYCADS 

the authors briefly discuss the possibility of insect pollination in the Lagenostoma s·eeds * * * Insect pollination is not 

peculiar to the Angiosperms, since it has been found to occur in several genera of living Cycads ( cf. Seward, Fossil 

Plants, vol. 3, 1917, p. 28). Several groups of insects being 

known to have existed in Carboniferous times, the possibility 

of entomophily in Paleozoic plants cannot be denied. The frequent occurrence of structures which ~seem to repres·ent external glands is, indeed, a striking feature in the seed-bearing leaves of Pteridosperms known at present. . . . But the 

relative frequency of these gland-like structures among plants 

which were certainly or probably Pteridosperms, coupled with 

the app.earance at about the same time of a ll'ich insect-life is 

suggestive. Additional knowledge of their distribution in different groups of Paleozoic plants is therefore desirable. 

Thus there is evidence of a marked trend toward 

production of glandiferous foliage and accessory 

parts among late Paleozoic seed plants, and P hasmatocycas provides an excellent additional example. 

The variety of insect life in the Paleozoic was 

pointed out by Carpenter ( 1952, p. 16) who listed 

nine orders of insects known from Permian strata 

and two orders from older rocks. Carpenter commented "Altogether, the lower Permian insect fauna 

was very diverse-more so, in fact, than any other 

insect fauna known." 

Not only was the Paleozoic insect fauna taxonomically diverse, but according to Carpenter 

(1971), it showed significant adaptive modification 

of mouth parts. Carpenter commented as follows 

(1971,p. 1238, 1240,1241): 

A very interesting :feature of the Palaeodictyoptera, Megasecoptera and Diaphanopterodea that has been discovered in 

recent year.s is the pres~mce of piercing-sucking mouthparts. 

There can be no question that beaks in the Paleodictyoptera 

and the other two orders mentioned were used for piercing and 

for sucking liquid nutriment. What the types of food were, we 

have no way of knowing; presumably they were of plant 

origin. The most likely source's were probably some of the 

succulent lycopods that were extensive and abundant during' 

the Carboniferous and Permian.1 

Carpenter ( 1971, p. 1241) continued thusly: 

The significance of this large array of hemipterous insects 

in the Paleozoic is obvious: there must have been abundant 

sources of liquid plant food available. With the Palaeodictyoptera, Megasecoptera and Diaphanopterodea also present, it is 

perhaps no exaggeration to state that nearly half the sp.ecies 

of Paleozoic insects had piercing-sucking mouthparts. 

1 It is doubtful that the lycopods, although abundant in the Paleo- zoic constituted an important source of food for herbivorous insects be- cau~e of certain anatomical features of the lycopods. Their; structural strength derived mainly from a thick outer periderm layer · which was nonconductive and therefore not very succulent. Likewise, the thick hypodermal layers of the leaves probably discouraged foraging insects. 

Possibly some soft-skinned amnhibians were plagued by these sharp- beaked insects, just as man suffers mosquito bites to~ay: If one adheres to the idea of an herbivorous habit for these Paleozoic msects, however, 

alternative sources of food other than the lycopods deserve considerati?n. There seems to have bet>n a compatibility of size between the suckmg beaks of some of the Paleozoic insPcts and the micropylar canals of some of the pteridospermous seeds. Perhars the conte:nts of the polle:n chambers were attractive to the insects, who used their beaks to obtam these substances, but it is very doubtfnl that enough seeds were pro- duced to sustain a sizeable insect population. Thus, externai glands on the plants seem to constitute a logical source of insect nutriment. 

Considering the sizable number of Paleozoic insects that had specialized-feeding mouthparts and 

their coexistence with many kinds of seed plants, it 

seems reasonable to assume that the feeding habits 

of at least some of these insects were involved in the 

pollination of seed ferns and precursive cycads. The 

contents of the external glands were possibly attractive to insects and thus served the function of nectaries. It is intriguing to imagine pollen-dusted 

insects, having pre·viously been attracted to the polleniferous organs by odor or other stimuli, visiting 

the putative taeniopterid distal lamina of Phasmatocycas for the food contained within the abundant 

glands. Thence, they would descend along the see~ 

bearing axis, feeding on the interseminal glands and 

meanwhile depositing pollen on the micropyles or 

pollen drops of the ovules. The regular placement of 

the glands of Phasmatocycas would render each 

ovule equally attractive to the pollen-bearing insects, 

thus assuring pollination of all. 

A mechanism for initial visual attraction of insects to a female infructescence of Phasmatocycas 

could conceivably have been one similar to that postulated by Leppik ( 1971, p. 172) for the Triassic 

Palaeocycas integer. This plant supposedly had a 

central crown of megasporophylls surrounded by a 

closely set group of leaves. According to Leppik, 

Such patterns must have been clearly distinctive to the foodsearching phytophagous insects * * * the adaptive form and 

arrangement of these clusters of sporophylls must have made 

them characteristic attractants to food searching insects, allowing them to distinguish thes·e from the less palatable ferns 

and horse tails. 

The question of insect pollination among the cycads is variously argued. Seward (1917, p. 28) 

wrote: 

Recent observations point to the probability that insects 

play a part in the pollination of cycadean ovules. K1·aus drew 

attention to the strong smell emitted by the microstrobili of 

Dioon edule and noticed that small bees were attracted to the 

ripe strobili of Mac1·ozatmia, while odourless cones of a neighboring Ceratozamia received no attention. Pearson and Rattray 

have obtained evidence that beetles and weevils act as pollinators to species of Encephalartos. 

The reports of insect pollination in Encephalartos 

are interesting, but Coulter and Chamberlain (1910, 

p. 141) observed that "It is generally accepted that 

the cycads are wind-pollinated." Both Chamberlain 

(1935, p. 127-128) and Gawssen (1944, p. 56) 

pointed out that the supposedly pollinating beetle 

actually bores into the female gametophyte of Encephalartos, and the gametophyte is destroyed either 

in the process or by the hatching larvae. In this inefficient procedure, the ovules, although possibly 
SYSTEMATIC DESCRIPTIONS 21 

pollinated, can mature only if the boring ovipositing 

process is not extensively damaging or if the beetle 

eggs fail to hatch. Crepet ( 1972, p. 1055) further 

discussed this interaction in presenting conjectures 

on the possibility of animal pollination of the Mesozoic cycadeoids. 

Even though there is significant skepticism toward 

entomophily among the cycads, an affinity of sorts 

exists between certain modern cycads and certain 

insects. Perhaps this is the distant reflection of a 

dubiously successful experiment that began with 

Phasmatocycas, other Paleozoic plants, and insect 

pollinators unknown. 

?Phasmatocycas sp. 

Plate 2, figure 1 

DESCRIPTION 

This account is based on a single specimen from 

the Haskell County, Tex., locality; it was found by 

Mr. T. R. Tinsley, who presented it to me for study. 

The specimen consists only of an impression in red 

siltstone; it lacks any organic matter but shows sufficient character to warrant discussion relative to 

Phasmatocycas and perhaps Spermopteris. 

The specimen consists of a fragment of a stout 

straight axis, which bears the impressions of two 

lateral rows of oval objects that probably represent 

seeds. The axial fragment is 5 em long, about 5.0 mm 

wide at its broadest point, and shows little tapering 

throughout its length. 

Along either side of the axis is a row of shallow 

oval impressions closely resembling the symmetrical 

system of placement of the seeds in Phasmatocycas 

kansana. Toward what I presume to be the top of 

the axis, the impressions are evenly paired, hut the 

positional relationships between the units in the two 

rows become somewhat obscure toward the opposite 

end of the axis where preservation is less complete. 

There are 24 impressions in all, 14 on the right and 

10 on the left side, part of which is missing. It is 

evident that one row contained approximately the 

same number of impressions as the other. 

The impressions are those of small round to oval 

bodies, 3.0-4.0 mm long and 2.0-3.0 mm wide. They 

show no tapering or beaklike apices, but evidently 

reflect broad attachments to the central axis by bases 

nearly as wide as their greatest widths, like seeds of 

Phasmatocycas kansana. The impressions are closely arranged but neither abut upon nor overlap each 

other; there are intervals of 1.0-1.5 mm between 

adjacent impressions. No evidence of accessory 

structures or scars of attachment of additional seedlike bodies is present on the surface of the axis between the lateral rows. 

One additional feature of this specimen remains 

for comment. A slightly elevated ridge almost completely surrounds the specimen except for the basal 

area where preservation is extremely obscure·. This 

ridge more or less outlines the axis at a distance of 

approximately 3 mm beyond the apices of the seedlike impressions. It is best seen at the presumed apex 

of the specimen, where it passes over the tip of the 

axis and lends a tonguelike outline to the organ (pl. 

2, fig. 1). The rounded apical outline resembles the 

tip of a leaf or pinna, but if the ridge does in fact 

represent the margin of a laminar appendage, there 

are no impressions. of veins to confirm this. 

Figured specimen.-USGS 10064-1 (pl. 2, fig. 1). 

Occurrence.-USGS paleobotany locality 10064. 

DISCUSSION 

This specimen is only provisionally assigned to 

Phasmatocycas because of its preservational deficiencies. Nonetheless it merits attention because its 

symmetrical bilateral construction conforms with 

the cycadlike pattern of the other fossils under consideration. Its central axis is more robust than that 

of P hasmatocycas kansana in proportion to the size 

of the seedlike impressions; although the seedlike 

impressions are not as closely arranged as the seeds 

of P. kansana, this specimen is, in most respects, almost exactly what one might expect to see if the 

ho!otype of P hasmatocycas kansana had been preserved in the same manner as the specimen from the 

Vale Formation. I thus have no reservations in regarding the Vale specimen as a seed-bearing axis, 

and further, as a representative of some taxon in 

the Permian complex of ancestral cycads. 

If we could establish beyond doubt that the Vale 

specimen is correctly assignable to Phasmatocycas, 

then the question of specific status would pose difficulties because only one specimen is available from 

each of the two localities concerned. The size and 

other differences between Phasmatocycas kansana 

sp. may fall within a reasonable range of specific 

variation, but until more material becomes available 

this problem must remain unanswered. 

Of more moment is the matter of the generic assignment of the Vale specimen. Were it not for the 

vague suggestion of the outline of a foliar lamina 

surrounding the axis and its complement of presumed seeds, there would be a rather stronger case 

for assigning the specimen to an unnamed species of 

P hasmatocycas without the provisional question 

mark. This outline must be considered, however. If 
22 PALEOZOIC ORIGIN OF THE CYCADS 

it actually represents the margin of a leaf blade 

rather than an anomalous sedimentary feature of 

the rock, then the Vale specimen cannot be regarded 

as a Phasmatocycas but as something more akin to 

Spermopteris. The seedlike bodies of the Vale specimen appear to have been attached directly to the 

axis; in this respect, the specimen differs from 

Spermopteris, whose seeds were attached to the 

lamina. Conversely, the mode of seed attachment relates the specimen to Phasmatocycas. If we assume 

that a lamina was present and the seeds were attached directly to the axis, then we may speculatively regard the Vale specimen as a derivative of 

Spermopteris, in which the foliar lamina retained its 

identity but in which the seeds had undergone a 

"phyletic slide" in the direction of the axis, to the 

extent of actually being attached to the latter. This 

possibility is interesting in the light of relative ages 

of material considered here. The Vale specimen is 

the youngest of the entire complex, but if it indeed 

retains the Spermopteris-like foliar lamina, it may 

represent a conservative line of cycad evolution, as 

opposed to the older Phasmatocycas, which has lost 

the foliar blade in the fertile area. 

1Phasmatocycas specbbilis Mamay, new species 

Plate 5, figures 1-4 

Specific diagnosis.-Fertile axes large, robust, as 

long as 20 em or more; widths as much as 1.0 em or 

slightly more, distally tapering very gradually. Bilaterally inserted rows of seeds conspicuous, the 

seeds as long as 7.0 mm, as wide as 5.0 mm; seeds 

oval or ovoid with blunt or slightly pointed apices, 

broadly attached by bases 2.0-3.0 mm wide, oppositely paired or slightly alternating in position, perpendicular to the axis or slightly ascending in attitude, crowded so as to abut against one another or 

more laxly arranged with intervening spaces 1.0-

2.0 mm wide. 

Holotype.-USGS 8959-3 (pi: 5, fig. 1). 

Paratypes.-USGS 8877-5, 6, 7 (pl. 5, fig. 2, 3, 4). 

Occurrences.-USGS paleobotany localities 8877, 

8959. 

DESCRIPTION 

As was the case with Archaeocycas, my attention 

was originally directed toward the existence of this 

large fertile structure by a suite of specimens collected by David White at the locality south of Fulda, 

Tex. (USGS paleobotany locality 8877), where he 

also found the first specimens of Archaeocycas. Like 

Archaeocycas, these 10 specimens were found segregated in a separate tray, and some had been marked 

with crayon; thus it is obvious that White had noted 

these as fossils of special interest and probably had 

intended ultimately to describe them. In retrospect 

of these events, it is rather strange that he made no 

reference to them in his list of plants from the Fulda 

beds, published in his paper on Gigantopteris americana ( 1912). 

In 1963, I found one small specimen of ?P. spectab·ilis, at the "Emily Irish" locality (USGS paleobotany locality 8959), and in 1967, I found several 

more specimens at the same place. The specimens 

from White's locality are better preserved and have 

more of a carbonaceous residue in a somewhat more 

durable matrix. There is no doubt, however, that the 

two suites of material represent the same taxon. 

None of the axial surfaces between the lateral 

rows of seeds show any features that would aid in 

characterizing this species. Some of the specimens, 

particularly those from the Fulda locality, retain 

thin films of coalified plant residue and show vaguely 

defined longitudinal striations (pl. 5, fig. 4), but 

these may be shrinkage phenomena and probably are 

meaningless. The longitudinal striations are also 

seen in a few small areas on the holotype (pl. 5, fig. 

1). Attention is directed to the "Emily Irish" specimen shown on plate 5, figure 4, because it has a 

rather thick film of coalified material adherent to 

the interseminal surface of the axis. This film is 

fragmented into many small, nearly cubical pieces 

but is particularly significant in that it shows no conspicuous points of interruption or discontinuity that 

would indicate places of attachment of seeds or accessory structures. This and all other observable features of this suite of specimens show that in ?P. 

spectabilis we are again dealing with a bilaterally 

symmetrical organ. Unfortunately the seeds are 

poorly preserved and contain no cutinized parts; pollen analyses were attempted with samples from both 

localities, but results were negative. Nonetheless, the 

arrangement of the seeds conforms closely with the 

patterns seen in the other materials discussed in this 

paper. 

The outstanding morphological feature of ?Phasmatocycas spectabilis is its large size. Some indicat{on of this is provided by the holotype, shown on 

plate 5, figure 1. This incomplete specimen is broken 

at both ends, measures 20 em in length, and has the 

attached remains of about 60 seeds.. As there is 

scarcely any diminution in its width from one end to 

the other, one has little basis for estimating the complete length of this specimen; it could have been considerably longer than 20 em, however. Approximately 20 smaller fragments were found closely associ-
SYSTEMATIC DESCRIPTIONS 23 

ated with the holotype, but these cannot be established as parts of the same specimen. None of the 

shorter fragments exceeds the width of the axis of 

the holotype (approximately 1.0 em) ; probably this 

was the maximum width attained by this species. 

White's specimens from Fulda, one of which is 

shown on plate 5, figure 2, substantially confirm a 

maximum axial width of approximately 1.0 em. 

The best Fulda specimen, shown on plate 5, figure 

2, is 10.8 em long and has a maximum width of 2.3 

em; the latter measurement embraces the two lateral rows of seed. The longest specimen in this suite 

(pl. 5, fig. 4) is 11.5 em long, and the re.maining 10 

are considerably shorter. Among the "Emily Irish" 

specimens, none approaches the holotype in total 

length. None of the other "Emily Irish" specimens 

show any unusual dimensions. The specimen shown 

on plate 5, figure 3, however, shows a definite decrease (from 3.5 to 2.0 mm) in width of the central 

axis and may represent the terminal part of the 

fruiting region. 

The seeds are oval or ovoid, and those of the Fulda 

specimens generally appear to be somewhat more 

pointed than those from the "Emily Irish" locality. 

Exact dimensions are difficult to ascertain because 

the specimens are generally poorly preserved, but 

their size ranges from 5.0 to 9.0 mm in length and 

4.0 to 7.0 mm in width. The largest seeds present in 

the collection are_ attached to the specimen shown on 

plate 5, figure 4; the fifth seed from the bottom in 

the right-hand row, for example, is 9.0 mm long and 

7.5 mm wide, and others on this specimen are of comparable sizes. Some of the seeds on the holotype are 

considerably smaller and seem disproportionately 

small in relationship to the size of the axis. 

Some of the seeds show vague suggestions of bifid 

apices, but such structures are not a consistent feature and may merely be the result of splitting of the 

integuments at the apices during compression. The 

presence of a thick integument is indicated by the 

"Emily Irish" specimens shown on plate 5, figures 2 

and 3. A few of the seeds are represented only by 

flattened coalified films. Others, however, are more 

or less completely delimited by deep cavities in the 

matrix; these are evidently cavities left behind by 

the decay of the seed integuments, and the extent of 

the cavities is indicative of some considerable bulk 

to the original seed coats. 

The seeds are uniformly sessile, being attached by 

very broad bases. In other features there is a certain 

degree of variation, all of which could be attributable to different relative positions on the parent axes. 

In most specimens the seeds are so closely inserted as 

to abut against each other (pl. 5, fig. 2) , but in others 

they are considerably more widely spaced (pl. 5, fig. 

1) . Most of the seeds extend away from the axis perpendicularly, but in a few specimens they are directed forward at broad angles; the latter condition 

is most prevalent in specimens where the seeds are 

loosely arranged, as in the holotype (pl. 5, fig. 1). In 

some specimens, the seeds seem to be arranged in 

distinctly opposite pairs, but most commonly they 

appear to alternate with each other. 

In none of the seeds is there any indication of 

cupular or bracteate investment; some seeds show 

irregular outlines and have erratic projections from 

their overall oval shapes, but I attribute these to 

preservational distortions. Furthermore, this species shows no remnants of the interseminal resinoid 

spheroids that appear on Phasmatocycas kansana. 

A final important feature of ?P. spectabilis is the 

absence of evidence of a foliar lamina. Because a 

complete specimen is not available, the presence or 

absence of a distal foliar lamina cannot be 

established. 

DISCUSSION 

?Phasmatocycas spectabilis is of considerable 

morphological interest because of its large size, the 

limits of which are yet unknown. More importantly, 

it establishes the presence in the Permian of still another representative of the cycadalean complex with 

which this paper is concerned. ?P. spectabilis repeats 

with remarkable clarity the bilateral symmetry and 

other qualitative features of the fruiting organs 

under study and thereby emphasizes the probability 

of phylogenetic derivation from Spermopteris. or a 

similar ancestral form. 

The absence of cuticularization is unfortunate; because of this absence, I refer this species to Phasmatocycas with the provisory question mark. Were 

cuticles present, I am reasonably confident that an 

unqualified generic assignment to Phasmatocycas 

could be justified. 

The specific name spectabilis refers to the overall 

showiness of these large fructifications. 

?Cycadean male cones 

Plate 2., figure 4 

DESCRIPTION 

An interesting element in the Haskell County 

flora is represented by 20 ... odd impressions or molds 

of axial fragments which were collected and given 

to me by Mr. Tinsley. The specimens are stout and 

unbranched, with no evidence of the apical or basal 
24 PALEOZOIC ORIGIN OF THE CYCADS 

organization. The specimen shown on plate 2, figure 

4, is the most informative of the lot. 

The axes have surface patterns of closely imbricated, spirally arranged rhomboidal impressions 

with considerable depth; in most specimens the impressions are elongate transversly to the axis. The 

impressions range from 7.0 to 10.0 mm in width and 

average about 5.0 mm in height. Extending away 

from most of the axial specimens and ascending upward in gently curved attitudes are molds of spinelike appendages, 2.0-3.0 em long; these extend across 

bedding planes in a manner suggesting that the 

appendages were stiff and rigid at the time of burial 

and resisted compaction. Several are seen clearly in 

the illustrated s1pe.cimen. 

Figured specimen.-USGS 10064-2 (pl. 2, fig. 4). 

Occurrence.-VSGS Paleobotany locality 10064. 

DISCUSSION 

These robust axes resemble to a certain extent the 

impressions of the characteristic bark patterns of 

the arborescent lycopods. However, the depth of the 

prominent surface impressions is much greater than 

that of fossil lycopod leaf cushions, which ordinarily 

show only a moderate amount of relief; these are 

more like the impressions that would be made by 

the distal faces of cycadean microsporophylls. If the 

Texas specimens are to be regarded as lycopod remains, this isolated appearance is anomalous with 

the known history of arborescent lycopods in North 

America. I am aware of no representatives of that 

complex in American rocks younger than those, of 

the Belle Plains Formation, where lycopod remains 

occur rarely in an otherwise rich plant assemblage. 

It is therefore highly probable that the giant lycopods became extinct on this continent before the end 

of Belle Plains time, and identification of the Texas 

specimens with the lycopods seems unwarranted. 

The preferred alternative of tentatively treating 

these specimens as cycadean male cones is encouraged by their geographic and stratigraphic proximity to Phasmatocycas and its putative foliage, Taeniopteris. That indirect evidence finds support in the 

resemblance of the Texas material to the specimen 

of the .Jurassic Androstrobus zamioides figured by 

Schimper (1870-1872, -pl. 72, fig. 1), especially in 

size, general proportions of the specimens, and the 

overall appearance of their surface markings. The 

cycadlike aspects of the Texas specimens is further 

accentuat~d by the molds of spinelike, apparently 

rigid extensions of the individual members. These 

may well represent the attenuated termini of microsporophylls, like those produced in the male cones of 

Cycas circinalis (Schuster, 1932, fig. lOA) or of 

Macrozamia (Johnson, 1959). All factors taken into 

consideration then, I prefer at least provisionally to 

regard the Texas specimens as the essentially modern-appearing male cones of an early cycad rather 

than as representatives of a lingering remnant of 

th~ giant lycopod lineage. Only the absence of evidence of microsporangia on the individual units of 

these strobiloid structures restrains me from applying a formal binomial indicative of cycadean affinity. 

Several kinds of presumably male strobiloid fructifications are present in other Permian collections 

under study; some are apparently new, but none can 

be identified as cycadean. Among these, those most 

like fossil cycadean male cones are the strobiloi¢1. 

structures from Fulda, Tex., described by White 

(1912, p. 500-501) as the "supposed polleniferous 

strobili" of Gigantoptwris americana. These were 

composed of reniform bracts inserted in distichous 

rows, with "oval, pendant sacs" on the lower surfaces of the bracts. A few reniform bractlike organs 

are indeed present in the Fulda collection, but my 

examination of the original specimens does not substantiate either White's interpretation of these 

bracts as the actual appendicular components. of the 

strobili, or his observation that the cone scales are 

distichously arranged. Furthermore, the material is 

too poorly preserved to allow observation of any 

details of sporangia! structure or disposition. Nonetheless, the Fulda specimens present the overall appearance of rather loosely constructed cones, probably with spirally arranged appendages. The specimens illustrated by White (1912, pl. 48, figs. 3, 4) 

are remarkably similar to some of the Jurassic male 

cycadean cones, Androstrobus wonnacotti Harris, 

illustrated by Thomas and Harris (1960, pl. 2, figs. 

8-13) ; the similarity is most marked if one turns 

White's illustrations upside down and compares his 

figure 4 with Thomas and Harris' figure 10 or 12. 

The absence of epidermal details or in situ pollen in 

the Fulda specimens, however, renders further comment on this similarity pointless. 

Order uncertain (CYCADALES or BENNETTITALES) 

Genus CYCADOSPADIX Scbimper, 1870 

Cycadospadix yocbelsoni Mamay, new species 

Plate 2, figure· 3 

Specific diagnosis.-Appendage with short, broad 

peduncle and lanceolate lamina. Margin of lamina 

entire from broadest point to base, deeply pinnatifid 

distally; lamina with heavy, unbranched ribs, several to the undissected part, one each to the distal 

lobes. 
SYSTEMATIC DESCRIPTIONS 25 

Appendage 21 mm long, 12 mm broad at widest 

part of lamina. Peduncle 5 mm wide, 4 mm or more 

long. Incised part of lamina with eight teeth along 

each margin; teeth as much as 3 mm long, the longest about halfway towards the sharply pointed tip. 

Teeth directed obliquely forward at broad angles. 

Holotype.--USGS 10057-1. 

Occurrence.-USGS paleobotany locality 10057. 

DESCRIPTION 

Only one specimen of this species has been found ; 

the better of its counterparts is shown on plate 2, 

figure 3. The incompletely preserved peduncle is a 

very broad and flat structure, the distal end of which 

extends as a broad wedge up into the base of the 

lamina and appears to have formed a slight ridge or 

suture at the line of juncture with the lamina. There 

is no indication of the nature of the vascular system 

of the peduncle other than a series of fine, closely 

spaced parallel surface striations that continue distally as far as the tip of the lamina. 

The lamina flares out abruptly and at a broad 

angle from its point of confluence with the peduncle, 

soon reaching its maximum width of 12 mm. Between the base of the lamina and its widest point the 

margin remains entire. The broad, parallel, distally 

directed ribs probably represent the original venation of this appendage. Each side of the undissected 

part of the lamina contains six such ribs, which depart from the distal end of the peduncle in a pseudopalmate fashion. Those that enter the margin lobes 

of the remaining two-thirds of the lamina, however, 

are arranged in a more typically pinnate manner. A 

distinct midrib is not evident in the lamina; the only 

indication of structural differentiation in the median 

part of the appendage is the weak parallel striations 

that extend into the lamina from the peduncle and 

continue to the tip. 

The lobes of the dissected part of the lamina vary 

in angles of departure, the more proximal ones being 

directed forward at about 45° and the distal ones 

extending out at right angles from the median line 

of the appendage. The lobes number eight or nine to 

each margin. The longest ones are about 3 mm long 

and less than 1 mm wide; they decrease in length so 

that the last one or two directly beneath the spicate 

tip of the lamina are slightly less than 1 mm long. 

DISCUSSION 

Cycadospadix Schimper is an artificial genus of 

isolated bractlike or megasporophyll-like organs, 

known from only a few occurrences, mostly of Mesozoic age. As stated by Harris ( 1932b, p. 96), it is 

characterized by "a stalk expanded distally and 

bearing several finger like branches; the fossils resemble in miniature the megasporophylls of Cycas." 

The resemblance to Cycas led Schimper (1870-1872, 

p. 207) to express the opinion that there could be no 

doubt as to a cycadean affinity for his material. 

Schimper referred two species to Cycadospadix: C. 

Hennocquei (Pomel) Schimper and C. Moraeanus 

(Pomel) Schimper. In his diagnosis of the genus, 

Schimper drew attention to scars on the sides of the 

petiole which he interpreted as indications of points 

of seed attachment; this was purely speculative, and 

Schimper's illustration of C. H ennocquei shows no 

seeds attached. Emberger (1944, fig. 346) published 

a drawing of a specimen of that species that had a 

seed attached to the petiole, but Harris (1961, p. 

316) stated that "no seed is known attached." 

Harris ( 1932b, p. 97-98) described Cycadospadix 

dactylota on the basis of three specimens from the 

Rhaetic of Greenland. The cuticles were preserved 

in his material, and on the strength of stomata of the 

bennettitalean type, Harris emended the generic 

diagnosis of the genus, thus inferentially assigning 

all species of Cycadospadix to the Bennettitales; 

Harris interpreted his specimens as the basal scales 

of a bennettitalean "flower" rather than megasporophylls. Florin ( 1933, p. 34) rejected Harris' emendation of Cycadospadix and proposed a new genus, 

Bennettitolepis, for the Greenland material, typified 

by the recombination B. dactylota (Harris) Florin. 

Harris evidently prefers his original designation, 

because in 1961 (p. 316-317) he referred to the 

Greenland material only as Cycadospadix and made 

no mention of Bennettitolepis. 

In his extensive article on the Mesozoic cycads, 

Florin ( 1933) restudied N athorst's original material 

of Cycadospadix integer from the Rhaetic of Sweden. 

Florin was able to determine the cuticular details, 

and on the basis of epidermal structure, he concluded 

that the organs were in no way related to the Bennettitales but instead showed close affinity with the 

modern Cycadales. He proposed a new genus, Palaeocycas, typified by the species of P. integer (Nathorst) Florin, and reconstructed an entire plant. 

Florin's (1933, p. 91) often-reproduced reconstruction shows a plant with a dense aggregate of terminal megasporophylls of the Palaeocycas integer type 

and taeniopteroid leaves called Bj-uvia simplex Florin. A single megasporophyll is shown as having a 

broad entire-margined lamina and two pairs of seeds 

attached to the sides of a thick stalk. Harris ( 1961, 

p. 315) remarked that none of the figured specimens 

has seeds attached, although bulges on the stalks of 
26 PALEOZOIC ORIGIN OF THE CYCADS 

Palaeocycas are suggestive of points of seed attach 

ment. 

The Texas specimen conforms well with the general concept of Cycadospadix but, as is the case with all 

other taxa heretofore assigned to Schimper's genus, 

there is no evidence of seed attachment. The broad 

stalk of C. yochelsoni suggests a stout seed-bearing 

axis, but no evidence of scars of attachment is visible; less than 5 mm of the stalk is preserved, however, and the possibility that more complete specimens would demonstrate evidence of seed attachment cannot be discounted. The distal laminar part 

of the appendage is remarkably similar to the top 

of a Cycas megasporophyll. This resemblance is such, 

in fact, that if C. yochelsoni had been found attached 

to any of the seed-bearing organs described here, 

there would scarcely be room for doubt as to a cycadalean affinity. 

The cuticle of C. yochelsoni is not preserved; thus, 

no evidence of stomatal or other epidermal features 

is available to aid in interpreting relationships of the 

specimen. Furthermore, laminar form is of little 

assistance. With its distally laciniate and basally entire lamina, C. yochelsoni is intermediate between 

Bennettitolepis, a presumed cycadeoid, anp. Palaeocycas, a presumed cycad. A short while before Florin 

reassigned Cycadospadix integer to the new genus 

Palaeocycas because of epidermal characteristics, 

Harris ( 1932b, p. 96) suggested that the undissected 

margin of the lamina of C. integer probably was 

sufficient basis for its reference to another genus. 

Similarly, additional material from the Vale Formation may necessitate a reassessment of the generic 

status of C. yochelsoni. 

Regardless of the taxonomic histories of organs 

originally assigned to Cycadospadix but later reassigned as the result of discovery of anatomical characteristics more definitive than gross external form, 

the original name Cycadospadix retains a considerable degree of utility as an ordinally noncommittal 

receptacle for structures that could be either cycadeoidean bracts or cycadalean megasporophylls. Consequently, the Texas specimen is appropriately retained in Cycadospadix. The question of its true systematic position-bennettitalean or cycadaleanmay, for the time being, be examined principally in 

the light of indirect evidence, some of it negative but 

all of it significant. The _stratigraphic and geographic relationships between Cycadospadix yochelsoni 

and the other materials considered here are overwhelmingly indicative of a cycadalean alliance for C. 

yochelsoni. Conversely, the absence of credible evidence of bennettitalean plants in the Paleozoic of 

North America militates against a bennettitalean interpretation of the Texas specimen. If C. yochelsoni 

is indeed a bennettitalean appendage, it stands 

strangely isolated-biotically, chronologically, and 

geographically. If its resemblance to the distal lamina of a Cycas megasporophyll is taken at face value, 

however, it assumes a harmonious niche in an assemblage of fossils whose collective impact is to 

extend the known history of the cycads downward 

through the greater part of two geologic systems. 

Cycadospadix yochelsoni is named after E. L. 

Yochelson, who found the specimen. 

ASSOCIATED PLANTS 

The following summary enumerates the plants associated with the fossils described here. It is presented to emphasize that the cycadlike fossils share 

little in the way of floral associates that could be 

considered pertinent to questions of taxonomic affinity. The overall effect of this resume is to strengthen 

the circumstantial evidence that the sporophyll of 

P hasmatocycas kansana had a taeniopterid lamina. 

The Perry, Okla., flora (USGS paleobotany locality 6233), of which the specimen of Spermopteris is 

part, was discussed briefly by White ( 1912, p. 506-

507) in connection with occurrences of Gigantopteris americana. He listed the following seven genera: Pecopteris, Gigantopteris, Taeniopteris, Annularia, Sphenophyllum, Walchia, and Araucarites. I 

have confirmed the presence of all of those genera 

except the two arthrophytes ; if present, they are too 

poorly preserved for positive identification. The 

identification of ATaucarites is doubtful, but the collection contains a few small seeds that may have 

been the basis for White's identification; two are 

partly cutinized and resemble somewhat the seeds of 

Phasmatocycas kansana. Two specimens of the 

asymmetrically winged seeds attributed by White to 

Gigantopteris are present, and I have identified 

Odontopteris and Callipteris. This is a typical lower 

Permian assemblage, referable to Zone 14 of Read 

and Mamay (1964). 

·The Banner, Kans., flora, supposedly the source 

of the holotype of Phasmatocycas kansana (USGS 

paleobotany locality 8869) was indicated by White 

(1912, p. 507-509) to contain the following six genera: Callipteris, Glenopteris, Taeniopteris, Sphenophyllum, N oeggerathia ?, and Carpolithes. The collection also contains specimens of cordaitea,n leaves 

and several small undetermined sporangia, but I am 

hesitant to verify a noeggerathialean element in the 

collection. This is a sparse assemblage, but by virtue 
ASSOCIATED PLANTS 27 

of the presence of the genus Glenopteris it is also 

assignable to Zone 14. The small collection from 

Insect Hill (USGS paleobotany locality 8298) contains no elements that are not present in the Banner 

assemblage. Sellards (1908), however, described the 

following additional genera from the vicinity of 

"Banner City" in his review of the Permian flora of 

Kansas: Pecopteris, Aphlebia, N europteris, Odontopteris, Signlaria, Cordaites, Cordaianthus ?, Rhabdocarpos, Cardiocarpon, and Aspidiopsis. 

White ( 1912, p. 505-506) recognized 14 genera 

associated with Gigantopteris americana at the Fulda, Tex., plant beds (USGS paleobotany locality 

8877). These are Diplothmema, Pecopteris, Odontopteris, N europteris, Taeniopteris, Annularia ?, Sphenophyllum, Sigillaria, Cordaites, Poacordaites, Walchia, Gomphostrobus ?, Aspidiopsis, and Araucarites. 

Names of plants from a nearby locality, 21;2 miles 

south of Fulda ("Castle Hollow," locality unnumbered) were incorporated in the same list because of 

the presumable identical stratigraphic positions of 

the two deposits. The "Castle Hollow" beds apparently contained most of the genera present in the Fulda 

beds, as well as the two additional genera Aphlebia 

and Sigillariostrobus. Thus the total number of genera from the two localities, as noted by White, is 17. 

My examination of White's Fulda collection substantiates his identifications with the exceptions of 

Sigillaria, Poacordaites, and Diplothmema. Conversely, this collection contains Stigmaria, Aphlebia, 

Annularia, Discinites (Mamay, 1954), and a curious apparently new peltate structure that in some 

features resembles the genus Daubreeia Zeiller. I 

have not located White's original "Castle Hollow" 

collection and must assume that it has been lost. Its 

absence, however, only reduces the total list of genera known from that area by one, Sigillariostrobus. 

The overall aspect of this flora is unusual only from 

the standpoint of the presence of the Daubreeia-like 

structure, Discinites, Archaeocycas, and ?Phasmatocycas. The flora is recognized as an assemblage typical of Zone 14. 

The most diverse flora involved in this study is 

that of the "Emily Irish" deposit (USGS paleobotany locality 8959). These beds are regarded as the 

approximate stratigraphic equivalent of the Fulda 

beds; they contain all the genera recognized in 

White's collection with the exception of the DaUbreeia-like organ. The "Emily Irish" flora is richer 

than the Fulda flora, however, in numbers of species 

as well as genera. It contains an unusually diverse 

spectrum of callipterids in addition to Labatannularia, Sphenopteris, and the curious genus Russellites (Mamay, 1968). Also present are several types 

of undescribed microsporangiate fructifications. Gigantopteris americana is a conspicuous ele.ment in 

this flora, which is referable to Zone 14. 

The Taylor County, Tex., flora (USGS paleobotany locality 10057) in which the specimen of Cycadospadix yochelsoni was found, is somewhat younger 

than the Fulda and "Emily Irish" assemblages and 

has been assigned to the youngest of the recognizable upper Paleozoic floral zones, Zone 15. It contains essentially the same generic association as the 

older flora but is distinguishable from it on the basis 

of several notable specific differences; the most striking of these differences is the appearance in the 

Lawn flora of a new, as-yet-undescribed species of 

Gigantopteris (G. n. sp. B; Read and Mamay, 1964, 

pl. 19, fig. 2). In addition to Gigantopteris and Cycadospadix, the flora contains Annularia, Aphlebia, 

Taen1:opteris, Sphenophyllum, Pecopteris, Gomphostrobus, Walchia, Callipteris, Odontopteris, and several taxa of unidentified seeds. Some of the odontopterids are of an extremely large-pinnulled type reminiscent of certain elements in the Siberian Angara 

flora, also of Permian age. The flora also contains 

some curious hooked appendages of unknown alliance, two specimens that resemble liverworts, and 

what appears to be a new genus of conifers. A description of this flora and related assemblages from 

other parts of the Vale Formation is being prepared. 

The Haskell County, Tex., exposures (USGS paleobotany locality 10064) are higher in the Vale Formation than the Lawn deposit, and, accordingly, 

theJy contain the youngest flora involved in this 

study. This flora is also the simplest in terms of generic composition, although its taxonomically limited 

aspect is not regarded as a function of limited material. The known plant association consists only of 

the single specimen of ?Phasmatocycas sp., Taenioptwris, walchian conifers, an undescribed new species 

of Gigantoptwris, and the axial specimens described 

as ?cycadean male cones. This sparse list of plant 

types is the result of not only my personal examination of literally hundreds of specimens scattered over 

an area of several acres, but it also reflects the careful collecting activities, over a period of many years, 

of Mr. T. R. Tinsley, who gave me the specimen of 

?Phas,matocycas sp. as well as my choice of any other specimens in his collection. Mr. Tinsley had 

searched the site and immediate environs frequently, 

in the hope of finding "new" fos~ils. On my first visit 

there, I was astonished by not only the great numbers of specimens lying loose on the surface, but also 

by the predominance of specimens of Gigantopteris 
28 PALEOZOIC ORIGIN OF THE CYCADS 

n. sp. This taxon constituted possibly all but 1 percent of the total number of specimens examined; 

walchian conifers were next in frequency. Only a 

few specimens of Taeniopteris have been found at 

this site. By virtue of the numbers of specimens examined, I believe that a reasonable representation of 

the original flora is preserved at the fossil site. 

After elimination of the obviously pteridophytic 

or coniferous elements in the preceding assemblages, 

only seven pteridophyllous genera (Callipteris, Gigantopteris, Glenopteris, N europteris, Odontopteris, 

Russellites, and Taen1:opteris) remain for consideration as possible foliar components of the plants that 

bore the cycadlike structures under investigation. 

From the standpoint of its pinnately compound morphology and parallel-veined foliar segments, Russellites is the most cycadlike of these genera. The possibility of cycadean affinity for Russellites has been 

considered briefly (Mamay, 1968, p. I10), but as it 

is known from only one site (USGS paleobotany 

locality 8959), it is ineligible for consideration as a 

common ancestral form for both P hasmatocycas and 

A rchaeocycas. 

Odontopteris, Callipteris, and N europteris are all 

regarded as pteridosperms; although they are stratigraphically and geographically widespread, none appears in the Haskell County flora. Although Gigantopteris is abundant at the Haskell County site 

and in other parts of Texas and Oklahoma, the genus 

is not known in Kansas; likewise Glenopteris, abundant in a small area of Kansas, is not known outside 

of that State. The distributions of Glenopteris and 

Gigantopteris, viewed in terms of paleophysiography 

and other physical factors, led Read and Mamay 

(1964) to present some conclusions regarding limited geographic provinciality of certain Permian 

plants in North America. 

The conclusions from the foregoing remarks are 

that Taeniopteris is the least common botanical de-

,nominator; it is the only foliar element present at 

all the localities where the cycadlike remains have 

been found. Although inconclusive in itself, the evidence of consistent association must always be given 

careful consideration in drawing general conclusions; in this instance it is particularly germane 

because of the convincing evidence of the known 

seed-bearing habit of Spermopteris and the accessory evidence provided by the resinoid spherules of 

Phasmatocycas and associated taeniopterids. 

DISTRIBUTION OF THE FOSSILS 

Stratigraphic positions of the fossils are shown in 

figure 5. The limited vertical distribution of the 

plants adds substantially to the distinctiveness of the 

guide assemblages that characterize floral zones 11 

to 15 of Read and Mamay ( 1964). The scarcity of 

taxa in the Wolfcampian interval is clearly a function of limited collections. Wolfcampian beds have 

not yet yielded large amounts of good fossil plant 

mat.erial, and the few collections on hand have 

cast no light on the cycadean complex under 

investigation. 

Geographically, the plants are distributed over a 

small part of the southwestern United States, extending only from north-central Texas to east-central 

Kansas. Although a very small area is involved, in 

terms of general geographic distribution of Paleozoic floras, the distribution of these plants may be 

significant from the standpoints of ecologic tolerances and evolutionary plasticity. 

Read and Mamay (1964, p. K16-K18) recognized 

three Permian floral provinces in Leonardian rocks 

of the southwestern United States; these provinces 

were based on lateral distributions of the genera Gigantopteris, Glenopteris, and Supaia, as related to 

paleophysiography, paleoedaphics, floristic characteristics, and other factors. The Glenopteris flora is 

restricted to a small province in eastern Kansas, and 

because of its consistently close association with 

saline rocks, the flora was presumed to have evolved 

in a physiologically xeric environment. Immediately 

south and southwest, the Glenopteris flora is displaced, without evidence of overlap, by the Gigantopteris flora, which occupied a rather less limited geographic province having presumably less rigorous 

environmental conditions. A third province, that of 

the Supaia flora, existed west of the ancestral Rocky 

Mountains; this province seems to have been characterized by harsh ecologic conditions which produced an impoverished flora. 

The plants described in this paper were not restricted to any one of the three Permian floral provinces, but collectively they occupied all three. Although Spe1·mopteris probably evolved before the 

beginning of the differentiation of upper Paleozoic 

floral provinces, it first appeared on the eastern margin of the area later to become the province of Glenopteris, then migrated southwest, and eventually 

was found in north-central Oklahoma in the Gigantopteris province. Phasmatocycas appe·ars in the 

Glenopteris province and, assuming that my identifications are correct, in the Gigantopteris province as 

well. Only Archaeocycas and Cycadospadix are presently limited to one province--that of Gigantopteris. 

Taeniopteris, however, the one floral element in common association with all the others, is found in all 
DISTRIBUTION OF THE FOSSILS 29 

c 

(/) 

Q) 

·.::: 

Q) 

(f) 

c 

ro 

"2 

ro 

c 

0 

Q) 

....J 

-(/) 

~Q) 

oc 

-o i.J...N 

North Texas 

Choza Formation~ ?Cycadean 

r-. male cones 

Vale Formati..:>n l ?Phasmatocycas sp. 

Arroyo/ I Cycadospad1x 

Formation yoche/som 

Leuders 1! 

Formation 

I Archaeocycas 

V) white/ 

Oklahoma 

I 

I 

I 

I 

Kansas 

[ 

I 

I 

I -~ 

E 

a; 

a.. 

.... 

Q) 

""" ;:::: Clyde Formation 1 .. 

,.... Ul ~,..... !PhasmafOCI/CaS Q) "0 g..~ 1-----------l :/ 

c- ~ o Belle Plains'*- spectabt!/s 

Wellington 

Formation 

I 

rspermoptens sp. 

Wellington 

Formation 

~- Phasmatocycas 

1 kansana 

~ 

0 

....J 

c 

-~ c ._ ro Q) > a.>, 

Q.(/) 

~ c 

c 

Q) 

a.. 

c 

ro 

·a. 

E 

ro 

~ 0 

~ 

c 

-~ 

~e~~ c3 Formation 

ci. 

a. (Y) (/) 

..... V) 

Q) ;:::: 

§ ~ 

N::2-

U 

I 

Lawrence 

Formation 

I 

I 

Spermo-

~pterls cor/acea 

j 

FIGURE 5.-Vertical distribution of late Paleozoic cycads and related plants. Modified from Dunbar and others (1960), Moore 

and others (1944), and R-ead and Mamay (1964). 

three provinces; it is a conspicuous element in the 

Supaia flora, three species having been described 

from the Hermit Shale of the Grand Canyon, Ariz. 

(White, 1929). Thus Taeniopteris was chronologically persistent, geographically adventuresome, and 

ecologically resilient. If those qualities reflect a genetically robust constitution for the taeniopterids, the 

phylogenetic acquisition of such traits would equip 

any Paleozoic derivatives of the taeniopterids advantageously for worldwide dispersal during the 

Mesozoic. 

MORPHOLOGIC AND EVOLUTIONARY 

CONSIDERATIONS 

. INTERRELATIONSHIPS OF THE AMERICAN FOSSILS 

Appraisal of the evolutionary significance of these 

fossils is best begun by an attempt to weigh their 

relationships with each other. If their stratigraphic 

positions were such that a vertically linear sequence 

of forms could be used as a framework for phylogenetic analyses, those interpretations would be a 

relatively simple matter. As shown in figure 5, however, a linear phylogeny is not apparent among these 

plants because two of the principal forms, Phasmatocycas and Archaeocycas, are clearly representative of different evolutionary stages; in their earliest 

appearances, however, they occur in essentially synchronous deposits. Thus they probably represent separate evolutionary tangents, derived from a common 

spermopterid ancestor. 

The simplest phyletic facet of this association is 

in the relationship between Spermopteris and Phasm.atocycas. The role of Spermopteris as the morphologic equivalent of, if not the actual archetypic plant 

from whieh Phasmatocycas kansana arose, becomes 

apparent upon examination of all the pertinent facts . 

The stratigraphic relations fit the demands for such 

a phylogenetic affinity, for Spermopteris, the ancestor, is considerably olde,r than Phasmatocycas, the 

descendant. The geographic proximity of known occurrences of the two genera adds substance to a 

hypothetical genetic relationship. Seeds. of the two, 
30 PALEOZOIC ORIGIN OF THE CYCADS 

as far as they are known, are similar. The intimate 

association of the genera with sterile taeniopterid 

foliage emphasizes the probability of physical identity. This probability is virtually converted into reality 

by the presence of the regularly distributed resinoid 

spheroids on the axis of Phasmatocycas kansana and 

of similar bodies on only taeniopterid foliage; only 

the actual organic continuity between Taeniopteris 

and Phasmatocycas remains to be found. The total 

effect of the foregoing lines of evidence is to promote 

acceptance of P hasmatocycas as a cycadean mega .. 

sporophylJ with basal ovules and a taeniopterid distal lamina, as reconstructed in figure 4. 

Phyletic derivation of Phasmatocycas from Spermopteris would involve the morphogenetic processes 

of migration of seed positions, partial sterilization 

of the megasporophyll, and reduction of the lamina. 

According to Cridland and Morris ( 1960, p. S55), 

the seeds were confined to the terminal regions of the 

megasporophyll of Spermopteris but in all the cycads and in the reconstruction of Phasm,atocycas the 

seeds were basal. As seed position varies considerably among the pteridosperms that have simple, noncupular phyllospermy, there is no compelling reason 

to believe that te1rminal seed positions need have 

been phyletically immutable among the spermopterids. If some spermopterid megasporophylls originally were only basally fertile, or became basally fertile 

through distal sterilization of completely fertile 

megasporophylls or through basal migration of originally terminal seeds, one has found a phyletically 

immediate antecedent for Phasmatocycas. Eventual 

organogenetic elimination of the basal fertile part of 

the lamina would then effect a migration of the 

points of seed attachment from laminally superficial 

positions to positions along the sides of the foliar 

midrib, which would thus become the fertile peduncle 

of the megasporophyll. This process would preserve 

the original bilateral disposition of the laminally 

borne seeds of Spermopteris and would be reflected 

in the bilateral ovuliferous habit of all cycadean 

megasporophylls, living and fossil. 

This hypothetical series of modifications is shown 

diagrammatically in figure 6. The ultimate derivative 

form (fig. E), representing Phasmatocycas kansana, is essentially a modern Cycas-type of megasporophyll, except for certain minor morphologic 

divergences such as its relatively large number of 

ovules and its entire-margined lamina. Inasmuch as 

it epitomizes a cycadean megasporophyllar form that 

had substantially completed the important qualitative phases of its evolution by the onset of Leonardian time, closely related taxa might reasonably be 

expected to be found ultimately in any post-Paleozoic rock unit. Even though a certain amount of 

doubt attends the identity of the Haskell County 

specimen from the Vale Formation, the presence 

there of P hasmatocycas may well be expected. Considered in relation to other pertinent material, the 

Vale specimen would more logically be a true Phasmatocycas than an intermediate form with Spermopteris-like foliation; nonetheless, the possibility of 

affinity with Spermopteris cannot be discounted 

without more satisfactorily preserved material. 

With the exception of Archaeocycas, which probably represents an evolutionary stage similar to that 

shown in figure 6D, the age relationships of the 

known elements in this evolutionary series suggest 

that forms intermediate between Spermopteris and 

P hasmatocycas must be sought in Wolfcampian 

rocks, where evidence of cycadean fossils is yet 

absent (fig. 5). 

The specimens of ?Phasmatocycas spectabilis are 

impressive because of the large size of the sporophylls and the great numbers of seeds. These features make it difficult to assess possible affinities between ?P. spectabilis and its contemporary, P. 

kansana, which is known from only one small specimen. Although the bilateral distribution of the seeds 

of the Texas specimens strongly suggests their assignment to Phasmatocycas, the absence of information regarding ovular details precludes the positive 

generic identification. Organic evidence for correlation of the fertile axes with a given foliar type, such 

as presented by the resinoid spherules in the Kansas 

material, is not available in the Texas specimens. 

However, some of the associated Texas taeniopterid 

leaves are so large (laminae as much as 10.0 em 

wide and midribs as broad as 15.0 mm) that their 

establishment as distal laminae of the large fructifications of ?P. spectabilis would not seem disproportionate. 

Accepting the probability that ?P. spectabilis and 

P. kansana are congenerically related, the impressive 

quantitative differences and essentially contemporaneous stratigraphic relationships of the two taxa 

would seem to discourage the concept of a linear 

phylogenetic alliance. More likely they are coordinate descendents of a common spermopterid ancestor, ?P. spectabilis having undergone reduction of a 

considerable length of foliar lamina while retaining 

a large complement of seeds. In the latter respect, P. 

spectabilis is to be regarded as the most prolific 

megasporophyllar type yet discovered among the 

cycads. It is also to be regarded as primitive, if the 

concept of reduction of numbers of parts is to be 
MORPHOLOGIC AND EVOLUTIONARY CONSIDERATIONS 31 

A B c D 

FIGURE 6.-Hypothetical evolutionary development of primitive Cycas-like megasporophyll from Spermopteris-like pteridosperm. A, Dorsal view of ancestral spermopteroid megasporophyll Wiith taeniopterid lamina and superficial seeds 

(small solid-black areas at seed bases indicate points of attachment to lamina). B, Derivative form with reduced num~ 

bers of seeds. C, Further reduction, with seeds restricted to basal area of lamina. D, Form with seed production essentially constant, but basal part of lamina ,reduced, with basalmost seeds attached directly to stalk of megasporophyll. 

E, Ultimate Cycas-like derivative (=Phasmatocycas), with basal part of lamina completely reduced and all seeds attached to megasporophyll stalk. 

retained as indicative of a trend in the direction of 

evolutionary advancement. 

Because of its morphologic intermediacy, Archaeocycas whitei stands out as the most intriguing and 

phylogenetically important taxon found in this study. 

It is a curious mixture of both primitive and advanced features; thus the nature of its morphologic 

origins and the direction of its phyletic destiny seem 

apparent. The critical characteristics of this plant 

are consistently observable in such a large suite of 

specimens that they cannot be coincidental ; thus, an 

obvious set of interpretations may be set forth with 

confidence. 

The spacing of the small circular scars of seed 

attachment is remarkably regular in Archaeocycas, 

showing that the seeds were attached to the lamina 
32 PALEOZOIC ORIGIN OF THE CYCADS 

only a short distance away from the midrib. This 

method of seed attachment is very much like that in 

Sper·mopteris, as diagrammatically shown at the 

lower right of Cridland and Morris' ( 1960) figure 

10 (fig. 1 of this paper). Thus Archaeocycas retains 

the primitive feature of laminally borne seeds; by 

virtue of this characteristic, A rchaeocycas may be 

regarded as a primitive megasporophyll, derived 

from Spermopteris or a Spermopteris-like ancestor. 

Judged on its laminar morphology, Archaeocycas 

is distinctly intermediate between Spermopteris and 

the Cycas-type of megasporophyll. Although its lamina was still essentially complete, the basal part had 

become sharply modified and partly reduced, forming a closely fitting indusiumlike covering for the 

ovules. The sterile distal part of the lamina is known 

to have been abruptly expanded immediate,ly above 

the fertile area. Its distal details are incompletely 

known, but there is no reason to doubt its foliar nature; the specimens shown on plate 1, figures 1, 2, 

and plate 2, figures 5, 6, show hints of a taeniopterid 

type of venation. The extent to which the ovules were 

enclosed by the laminar "indusium," is unknown, 

but, in keeping with the cycadean theme of this paper, I would venture to guess that the lower surfaces 

of the ovules were largely exposed. 

Judged next on the basis of positions and numbers 

of seeds, Archaeocycas contrarily demonstrates a 

distinctly advanced morphological level, particularly 

in comparison with Spermopteris. Although the 

seeds of Spermopteris are said to be restricted to 

terminal parts of the sporophyll, those of Archaeocycas are definitely basal and therefore of substantially modern Cycas-like appearance. Furthermore, 

the number of ovules of Archaeocycas is not known 

to range beyond the extremes of 8 and 12 on a given 

sporophyll, with an average of 10. This modest level 

of ovule production is again an essentially modern 

feature; inasmuch as it is consistent among a handful of specimens from two different localities, this 

seems to have been a stable genetic feature of at 

least the type species, Archaeocycas whitei. Comparison of this characteristic would seem to indicate 

a higher level of advancement for Archaeocycas than 

either Spermopteris or Phasmatocycas. In Spermopteris, the number of seeds on a given sporophyll is 

not evident from Cridland and Morris' (1960) account, but evidently the seeds were produced in 

large numbers; the type specimen of Phasmatocycas kansana had 21 or more seeds, ?P. sp. had 24 or 

more, and ?P. spectabilis produced 60 or more. 

All morphologic features considered, then, Archaeocycas must be the derivative of some plant with 

the salient attributes of Spermopteris. It needed only 

to undergo morphogenetic reduction of the basal 

indusiumlike covering of the ovules and migration 

of the points of seed attachment to the midrib to 

attain the morphologic equivalence of a modern 

Cycas megasporophyll. Archaeocycas is, in fact, a 

classic example of evolutionary intermediacy; it is 

the type of fossil that would eventually have been 

predicted on a paper diagram, had it not been discovered in actual substance. In the vernacular, it is 

a critical "missing link." 

In terms of the hypothetical evolutionary sequence 

diagrammed in figure 6, Archaeocycas would most 

closely approximate the stage depicted in 6C or 6D; 

the chief qualitative difference is that the laminar 

"indusium" extending around the ovules is not 

shown in the diagram. A more complete representation of my visual concept of Archaeocycas and its 

evolutionary development into a Cycas-like megasporophyll is given in figure 7. The progressively 

deeper marginal incision of the distal lamina, as 

shown in forms C and D, represents a necessary 

step in the derivation of a pinnate appendage from 

an entire-margined one; certain fossil evidence for 

such a process is discussed below. 

The kinds or degrees of natural affinity that existed between A rchaeocycas and P hasmatocycas are 

difficult to weigh, primarily because both genera 

appeared at approximately the same time. Were it 

chronologically intermediate, Archaeocycas might 

be regarded as phylogenetically intermediate between Spermopteris and Phasmatocycas, in much 

the same manner that it seems intermediate between 

Spermopteris and Cycas; it might even be construed 

as a common ancestor of both Phasmatocycas and 

Cycas. The essentially comparable ages of the two 

Permian fossils, however, introduces the need for 

caution in reading their relationships in terms of a 

.simple linear phylogeny. This picture is further beclouded by the fact that ?Phasmatocycas spectabilis 

occurs at the same two localities and in the same 

beds in which Archaeocycas has. been found. A clearer understanding of the affinities between P. kansana 

and ?P. spectabilis would be helpful because, as previously stated, their phylogeny probably is not a 

linear one. If one is not derived from the other, it is 

possible that a common ancestor, intermediate between Spermopteris and those two Permian species, 

exists in older Permian rocks, and that the same 

taxon was ancestral to A rchaeocycas as well. In the 

light of age relationships of the fosils, then, it seems 

preferable to regard Archaeocycas as the result of a 
MORPHOLOGIC AND EVOLUTIONARY CONSIDERATIONS 33 

D 

FIGURE 7.-Hypothetical evolution of Cycas-like megasporophyll from ATchaeocycas (modified from Mamay, 1969a, fig. 1). 

A, Presumed dorsal view of megasporophyll, showing partial enclosure of paired basal seeds by sporophyllar lamina 

(extent of enclosure conjectural). B, C, Hypothetical derivatives, showing progressive reduction of basal lamina and 

resultant attachment of basalmost seeds to midTib, or stalk, of sporophyll; margin of distal lamina of form depicted 

in C has become slightly· incised. D, Cycas-like ultimate form, with basal lamina completely reduced and all seeds attached directly to stalk of sporophyll; distal lamina deeply pinnatifid, substantially as. in Cycas Tevoluta. 

line of descent independent from that which produced Phasmatocycas. 

Cycadospadix yochelsoni is difficult to interpret 

from the standpoint of phyletic position. Almost any 

of the necessarily indirect bases for assaying its 

natural relationships to the other fossils discussed 

here are obscured by the possibility that this species 

is actually a cycadeoidean bract rather than part of 

a cycadean megasporophyll. Nonetheless, be it cycadean or cycadeoidean, its presence in Lower Permian 

rocks is chronologically precocious, like that of its 

cycadean associates. Cycadospadix yochelsoni is further intriguing in that it would not appear biologically incongruous if it were established as the distal 

extension of one of the terminally incomplete fructifications described in preceding pages; rather, it 

would seem harmonious in such a position and would 

complete the portrayal of a Cycas-like megasporophyll in any of those instances. 

AN ALTERNATIVE INTERPRETATION OF 

ARCHAEOCYCAS 

The sketches presented in figure 7 were made on 

the premise that the ovules of Archaeocycas were 

incompletely enclosed by the fertile lamina. Placement of the margins of the lamina was rendered 

speculative because the few specimens available for 

this study were preserved only in lateral or ventral 

view, and delicacy of matrix and preservation limited mechanical preparation to a point just short of 

destruction of the specimens. Thus the lower or dorsal aspects of the megasporophyll of Archaeocycas 
34 PALEOZOIC ORIGIN OF THE CYCADS 

are not known. The lack of evidence of constriction 

of foliar tissue immediately distal to the most distal 

pairs of ovules, however, suggests that at least that 

part of the lamina was incompletely inrolled and 

that the plant was therefore gymnospermous. 

With the morphology of Spermopteris and other 

cycadalean entities in mind, then, I applied the most 

conservative interpretation of Archaeocycas and depicted it as a fertile leaf with superficial ovules and 

incompletely involute margins. As such it represents 

as uncomplicated as possible a theoretical derivative 

of a Spermopteris-like ancestor, yet with significant 

attributes of a primitive Cycas-like megasporophyl 

Only a modicum of morphogenetic reduction of the 

lamina was necessary to produce a Phasmatocycas 

or Cycas type of megasporophyll; thus Archaeocycas, if interpreted correctly, stands as an important 

Permian intermediate between the Pennsylvanian 

Spwrmopteris and the modern Cycas. 

The uncertain extent of involution of the megasporophyll of Archaeocycas introduces the necessity 

for considering an alternate interpretation. It seems 

at least remotely possible that, even if not coalescent 

in the known specimens of Archaeocycas, the opposing margins of the fertile base of the lamina were 

in an initial stage of proliferation rather than reduction, so that they subsequently met and began phylogenetic fusion. Continuation of the process of fusion 

would eventually have resulted in an organ in which 

the ovules were completely enclosed. This structure 

would thus show basic characteristics of an angiospermous carpel; as illustrated in my preliminary 

article (Mamay, 1969a, fig. 1H), this hypothetical 

organ was said by Uhl and Moore (1971, p. 990) to 

resemble the carpel of the palm Nypa. 

This theoretical process is illustrated in figure 8, 

adapted from the preliminary article (Mamay, 

1969a, fig. 1). Its significance lies in the inference 

that plants having at least one important angiospermlike reproductive feature could have been derived on a more or less direct line from the late 

Paleozoic pteridosperms, and in the corollary evidence from the fossil record, that the carpel is indeed a modified seed-bearing leaf. Thus the suggestion emerges that possibly the angiosperms and cycads are closely related through a common ancestry 

-the spermopterid pteridosperms-or even that the 

angiosperms arose directly from the cycads during 

the late Paleozoic. The latter idea could be particularly attractive to those who espouse the concept of 

Paleozoic angiospermy. (See Eames, 1959.) At the 

same time it would shed doubt on my arbitrarily 

delineated distinction between dorsal and ventral 

sides of A'rchaeocycas, because it is accepted that 

angiosperm ovules are produced on the adaxial (ventral) side of the megasporophyll (Eames, 1961, p. 

186) ; thus, either the ovules of Archaeocycas were 

attached to the wrong side of the lamina or my terminology has been incorrectly applied. 

The possibility that the specimen of Cycadospadix 

in the Texas flora is a Bennettitalean appendage 

resuscitates the question of relationships between 

the cycads and cycadeoids. Chamberlain ( 1935, p. 

158, figs. 171, 172) proposed that both groups arose 

from a common ancestor among the Paleozoic pteridosperms and presented a series of drawings depicting two reduction series leading to the cycads and 

cycadeoids; the hypothetical ancestral megasporophyll had a terminal ovule and several lateral ones 

intercalated between foliar segments. In one series 

the terminal ovules aborted, and continued reduction of parts resulted in several of the modern types 

of cycadean megasporophylls; in the other series, 

some of the terminal ovules were retained, all others 

were aborted, and with the reduction of laminar 

tissue, the ovule-bearing stalks and sterile "interseminal scales" of Bennettites were produced. Neither the ancestral nor the intermediate forms have 

been discovered, so that Chamberlain's hypothesis 

remains unsubstantiated. Delevoryas (1968b) 

agreed that the pteridosperms are the most likely 

source of the cycadeoids, but he also pointed out that 

the cycads and cycadeoids have always been distinct 

orders. If a common ancestry is involved among the 

pteridosperms, one probably must look for it in prePennsylvanian sediments. Certainly the present 

study sheds no light on the question, and if the Texas 

specimen of Cycadospadix is indeed bennettitalean, 

it is more than ever apparent that the two orde:r.s 

were distinct entities for as long as either can be 

recognized. 

It would be impractical to attempt to review here 

all the thoughts that have been published in regard 

to the ancestry of the angiosperms. Suffice it to say 

that all the various groups of gymnosperms-particularly the pteridosperms-have at one time or 

other been regarded as probable progenitors of the 

flowering plants. The cycads were especially favored 

as angiosperm ancestors by Chadefaud (1936) who 

interpreted the fertile megasporangiate organs as 

homologous and wrote ( Chadefaud, 1936, p. 590-

591): 

Or, la feuille carpellaire des Angiospermes est manifestement un feuille fertile a ovules lateraux * * * 

On en devra evidenment conclure que les Angiospermes sont 

plus etroitement apparentees aux Cycadales qu'a aucun autre 
MORPHOLOGIC AND EVOLUTIONARY CONSIDERATIONS 35 

A - B ---? C ---? D 

FIGURE 8.-Hypothetical evolution of carpellike organ from Archaeocycas (modified from Mamay, 1969a, fig. 1). A, Presumed dorsal view of megasporophyll, showing partial enclosure of ovules by sporophyllar lamina. B- D, Hypothetical 

derivatives, showing progressive proliferation and fusion of margins of lamina, resulting in complete enclosure of 

ovules (D). 

groupe de Gymnospermes. En particulier, c'est aux Cycadales 

et non aux Bennettitales qu'il faut les rattacher. 

Chadefaud's ideas have not been widely accepted 

by subsequent writers. Eames (1961, p. 464-467) reviewed the possible origins of angiospermy and 

omitted the cycads from consideration. Cronquist 

( 1968, p. 40- 46) again considered the possibilities 

and eliminated all the gymnosperms except the pteridosperms ( Lyginopteridales) as likely angiosperm 

forerunners, meanwhile urging caution toward acceptance of the lyginopterids in that capacity. Cronquist conceded, however, that the cycads probably 

arose from the pteridosperms; this view implicitly 

indicates a distant but common ancestry for the 

cycads and angiosperms. 

Carlquist (1969, p. 353), in commenting on fossil 

evidence on floral theories, discounted any evidence 

that "Cordaitales, Ginkoales, Coniferales, Cycadeoidales or even Cycadales ever had leaf-like megasporophylls." Carlquist also commented that "The 

chance of finding a fossil that shows ancestral carpel form is vanishingly small-and would we recognize it if we saw it, or would we misidentify it because of preconceptions?" 

Carlquist's comment on the "vanishingly small" 

chance seems incongruous with the steadily increasing ranks of paleobotanists, the attendant increase 

in collecting activities, and the fluid overall state of 

fossil botany. And perhaps an early "ancestral carpel form" may be represented by an Archaeocycaslike plant, and only better material is needed to establish its identity as such. 

The foregoing discussion is not intended as my 

endorsement of the idea that Archaeocycas is a car-
36 PALEOZOIC ORIGIN OF THE CYCADS 

pel or even an early form in the evolution of the 

carpel; I regard the cycadalean interpretation as 

more consistent with the attendant facts and circumstances, and particularly with the morphology of 

Spermopteris. The discussion is rather intended to 

point up the increasingly evident fact that Permian 

time produced many botanical innovations (see Mamay, 1962, 1966, 1968; White, 1912), some of which 

may have involved carpellike seed enclosure and the 

beginnings of angiospermy. 

RELATIONSHIPS OF THE AMERICAN FOSSILS 

TO OTHER FOSSIL CYCADS 

Appraisal of the natural affinities between the 

American taxa and other fossil cycad fructifications 

ish not substantially more complicated than that of 

the interrelationships of the American fossils themselves. The problem is that so few points of comparison exist; only a few post-Paleozoic ovulife·rous 

cycadean fructifications are recorded, and recently 

Harris ( 1961) has reviewed reasons for doubting 

the validity of some of those. 

The most thoroughly understood of the Mesozoic 

ovuliferous cycad fructifications is the genus Beania 

Carruthers from the Jurassic of Yorkshire, England; our knowledge of Beania is the result principally of the many contributions of T. M. Harris 

and H. H. Thomas. Beania is commonly regarded as 

the most advanced of the known Mesozoic cycads, 

for its seeds are of essentially modern aspect and the 

female infructescence, although long, lax and not 

very conelike, is composed of simple biovulate sporophylls of zamioid construction. Thus Beania does not 

bear any si·gnificant morphological resemblance to 

any of the American fossils and is probably not as 

closely related to Phasmatocycas or Archaeocycas 

as it is to the modern genera; if related within a 

simple infrafamilial phylesis, Beania is certainly 

closer to Phasmatocycas than to Archaeocycas, because of the more modern aspects of Phasmatocycas. 

If Beania arose from a member of the spermopterid 

stock of which the American fossils are clearly the 

progeny, the evidence for such derivation has not 

begun to be recovered; when found, it may be difficult to recognize as such. Beania, then, is of little aid 

in interrelating the American fossils with fossils of 

the Old World or of the post-Paleozoic. Furthermore, the Jurassic Period was sufficiently younger 

than the Permian, and cycad evolution apparently 

so rapid, that even if a more complete spectrum of 

female fructifications were known from the Jurassic, 

probably none of those fossils would be very enlightening on the phylogenetic implications of the 

American material. We must therefore consider the 

Triassic cycads. 

Only two Triassic taxa need to be considered here: 

Dioonitocarpidium Lilienstern 1928 and Palaeocycas 

Florin 1933. Both genera have been widely cited as 

credible examples of Triassic cycads, but in each 

instance challenges have been directed at cycadean 

interpretations. 

Ruhle von Lilienstern described Dioonitocarpidium on the basis of Upper Triassic (Keuper) material from Bavaria. It is a composite taxon, 

consisting of taeniopterid sterile foliage (Danaeopsis angustifolia Schenk) and pinnate sporophylls 

(Dioonites pennaeformis Schenk), each sporophyll 

supposedly bearing a basal pair of ovules. The type 

species is D. pennaeforme (Schenk) Lilienstern. Lilienstern's original reconstruction of the apical part 

of a plant is presented here in figure 9A. Although 

this reconstruction has been duplicated by various 

authors (Gothan and Weyland, 1954; Krause}, 1950; 

Magdefrau, 1956), it has not been universally accepted. It was criticized by Florin (1933, p. 122), 

who expressed doubt that a plant with simple taeniopterioid leaves would produce pinnate sporophylls. 

Florin furthermore pointed out that the foliage 

(Danaeopsis angustifolia) may not even be gymnospermous and that the name Dioonitocarpidium possibly embraces two unrelated plants, one a fern, the 

other a cycadophyte. 

An element of doubt as to morphological detail attends Lilienstern's reconstruction of the fertile base 

of a megasporophyll (fig. 9B). The drawing depicts 

a broad stalk with slender hairy pinnae and a pair 

of basal seeds. Within bounds of artistic license the 

reconstruction is reasonably acceptable, except for 

what appear to be anomalous subsidiary flanges of 

tissue extending from the sporophyll stalk below the 

attached ovules and continuing distally to the upper 

limits of the sketch. This improbable feature was not 

discussed in Lilienstern's text and is possibly attributable to a drafting error. The probability of an 

error gains support from the fact that the detailed 

reconstruction of the sporophyll base (see fig. 9C) 

was modified in two textbooks by Gothan and Weyland (1954, p. 288; 1964, p. 314), so as to eliminate 

the superfluous flange, although Lilienstern's overall 

reconstruction of the plant is faithfully reproduced 

in both the cited instances. 

Inasmuch as the Gothan and Weyland references 

included no explanation of the discrepancies between 

their reconstruction and the original, I inquired of 

Professor Weyland as to a possible reason. Weyland 

(written comm., 1968) advised me that "The illus-
MORPHOLOGIC AND EVOLUTIONARY CONSIDERATIONS 37 

c 

D 

FIGURE 9.-Reconstructions of Dioonitocarpidium Lilienstern. A, Apical part of plant of D. pennaeforme, showing taeniopteroid sterile foliage and pinnate sporophylls (from Lilienstern, 1928, fig. 1). B, Lilienstern's reconstruction of basal 

part of megasporophyll, showing pair of seeds and peculiar subsidiary flangelike structure beneath the pinnae (from 

Lilienstern, 1928, pl. 6, fig. 4). C, Modified reconstruction of basal part of megasporophyll of D. pennaeforme (from 

Gothan and Weyland, 1964, fig. 208 a). D, Reconstruction of megasporophyll of D. keuperianum (Krasser) Krause!, 

showing two pairs of basal seeds and pinnate lamina (from Krause!, 1949, fig. 4). 

tration of the sporophyll originated with Gothan 

and, so far as I know, it is not published elsewhere. 

Lilienstern's reconstruction probably seemed incorrect to him. However, I do not know where the specimen forming the basis of Gothan's illustration is 

located today." Whatever the exact basis of Gothan's 

illustration was, it is apparent that he disagreed 

with Lilienstern's depiction of the fertile base of a 

sporophyll, meanwhile accepting the overall reconstruction of Dioonitocarpidium as a plant with pinnate sporophylls and taeniopterid sterile leaves. The 

modifications introduced by Gothan into the reconstruction of the individual megasporophyll have resulted in a morphologically more reasonable form 

than the original of Lilienstern. 

Two additional species of Dioonitocarpidium-D. 

keuperianum (Krasser) Krausel, and D. liliensterni 

Krausel-were subsequently described; these also 

are from the Upper Triassic Keuper beds of Bavaria. 

D. keuperianum (Krausel, 1949) was reported as a 

megasporophyll with a pinnate distal part and a 

slightly pinnatifid basal flange of laminar tissue to 

the surface of which "several" seeds were allegedly 

attached; Krause} published a reconstruction of the 

sporophyll, which appears here as fig. 9D. In a later 

publication, Krausel ( 1953, p. 106) discussed additional material of D. keuperianum, which was said to 

"partly confirm, partly broaden and correct" his 

earlier description of that species. Krausel (1953, p. 

106) stated that seeds were plainly visible on one of 
38 PALEOZOIC ORIGIN OF THE CYCADS 

the specimens, and that at least five were attached to 

one side of the sporophyll. Unfortunately, the photographic illustrations in both publications are of 

poor quality and do not permit verification of the 

critical details of Krausel's reconstruction or of his 

diagnosis (Krausel, 1949, p. 48) of D. lceuperianum. 

A similar circumstance concerns the third species, 

D. liliensterni. In 1953, Krausel described D. liliensterni on the basis of a single sporophyll. This species differed from Krausel's concept of D. lceuperianum in that its basal, supposedly fertile part was 

much longer and broader than that of D. lceuperianum 

and was entire-margined. Krausel (1953, p. 107) 

stated that three seeds were attached to one side of 

this organ. Here again the photographic illustrations 

fall short of convincing demonstration of attached 

seeds. It may well be that the broad base of the leaf 

in question truly represents a reproductive specialization, but the requisite seeds remain in question. 

Thus, in consideration of the deficiencies of illustration, a certain amount of reservation appears necessary in drawing comparisons of D. lceuperianum and 

D. liliensterni. 

If one assumes that the descriptions and reconstructions of the three species of Dioonitocarpidium 

are accurate, then one must weigh the question of 

their taxonomic relationships, because their modes 

of seed attachment show as much difference, fundamentally, as the difference between the seed attachment of Archaeocycas and Phasmatocycas. In D. 

pennaeforme the seeds are sessile and supposedly 

are attached directly to a naked sporophyll stalk, as 

is the case with Phasmatocycas; conversely the seeds 

of D. liliensterni and D. keuperianum are supposedly 

attached to a basal laminar extension of the sporophyll, similar to the manner of seed attachment in 

Archaeocycas. Thus the Dioonitocarpidium concept 

may in fact consist of two genera, differentiable on 

the basis of characteristics of the fertile bases of the 

sporophylls. From this speculative standpoint, one 

might postulate two Permian-Triassic evolutionary 

lines of cycads, one leading from Phasmatocycas to 

Dioonitocarpidium pennaeforme and an attendant 

reduction in number of seeds, the other leading from 

Archaeocycas to D. liliensterni and D. lceuperianum. 

Admittedly this is a tenuous approach to phyletic 

considerations because of the many deficiencies in 

the physical evidence on hand. The American and 

European taxa may very well not be related by way 

of a linear phylesis. However, they show approximately comparable morphological qualities and seem 

to express similar evolutionary trends. Both groups 

contain members with laminally produced seeds, a 

fact which, through conservative interpretation, suggests that both groups were derived from a spermopterid stock. Thus it is not entirely unrealistic to 

suppose that the two groups had a common ancestry, 

if not a lineal relationship. The evidence of common 

association with taeniopterid foliage seems to bear 

out the foregoing lines of thought. 

The one Triassic taxon remaining for consideration is Palaeocycas-Bjuvia, a cycadean plant from 

the Rhaetian (Upper Triassic) of Sweden, which 

was described and reconstructed by Florin in 1933; 

his reconstructions appear here as figure 10. Like 

Dioonitocarpidium, Palaeocycas (for the sake of 

brevity of reference, Florin arbitrarily selected this 

as the name for the entire plant) is a composite 

genus. It consists of large taeniopterid leaves (Taeniopteris gigantea N athorst), to which Florin applied the binomial Bjuvia simplex, and presumed 

megasporophylls with ovate entire-margined distal 

laminae; Florin named the latter Palaeocycas integer. No seeds were found attached to the megasporophylls, but the stalks of these organs each bore two 

pairs of prominent lateral protrusions. Florin interpreted the latter as points of seed attachment and 

reconstructed the sporophyll as a leaflike appendage 

with four basal seeds attached to the naked stalk. 

Because of their association in the same rock strata 

and similar epidermal anatomy in the leaves and 

sporophylls, Florin deduced that the two organs belonged to the same plants. The reconstruction of this 

plant shows a stout armored trunk and loose terminal crown of leaflike megasporophylls, which strikingly resemble those of the modern Cycas. The outstanding discrepancy between Palaeocycas and Cycas is in the architecture of their respective fronds; 

those of Palaeocycas are simple, whereas those of 

Cycas are pinnate. 

The circumstantial nature of the evidence for 

Florin's synthesis of Palaeocycas and Bjuvia into 

one plant was discussed fully by Harris (1961). Harris' comments were mildly skeptical; his emphasis 

was on the absence of seeds as an important negative 

factor. Notwithstanding this type of criticism, however, Florin's reconstruction continues to appear in 

recent textbooks (Sporne, 1965, p. 106; Emberger, 

1968, p. 457) as an acceptable Mesozoic cycad. Indeed, this is possibly the most widely reproduced example of paleobotanical synthesis; Florin's reconstruction seems to have enjoyed classic status from 

the time of its original publication. 

The relevance of Palaeocycas to the present study 

is in the important resemblances that are seen between this Triassic plant and my concept of Phas-
MORPHOLOGIC AND EVOLUTIONARY CONSIDERATIONS 39 

B 

FIGURE 10.-Reconstructions of Palaeocycas-Bjuvia. A, Hypothetical reconstruction of entire plant. B, Single megasporophyll, reconstructed with two pairs of basal seeds. 

From Gothan and Weyland, 1964, fig. 209. 

matocycas; a much more convincing set of arguments can be presented in support of direct phyletic 

relationships between Phasmatocycas and Palaeocycas than is possible in comparing Dioonitocarpidium with Archaeocycas and Phasmatocycas. Although Palaeocycas and Phasmatocycas are both incompletely known, the gaps in our knowledge, which 

appear in different areas of their diagnostic images, 

are considerably minimized by taxonomically significant similarities. Indeed, their reconstructions 

are mutually corroborative on the grounds of complementarily indirect evidences. 

In both genera, sterile taeniopterid leaves and 

megasporophylls are regarded as attributable to a 

common parent on similar bases: ( 1) consistent association in the same strata, and (2) morphological 

features shared by both pairs of organs-epidermal 

characteristics in the case of Palaeocycas and glands 

in the case of Phasmatocycas. The seeds are absent 

from the sporophyll of Palaeocycas but its distal 

lamina is known, whereas the reverse is true for 

Phasmatocycas; its seeds are known but the distal 

lamina is not. Nonetheless, the evidence for correlation of parts in each instance points toward two 

plants with very similar ovulate sporophylls and 

both with taeniopterid foliage. Although my reconstruction of the megasporophyll of Phasmatocycas 

entails a longer, more leaflike distal lamina and more 

seeds than in P:alaeocycas, I believe that the ove·rall 

aspects of the two plants were probably very similar 

and that, with only minor modifications, Florin's 

habit reconstruction of Palaeocycas might represent 

Phasmatocycas as well. The sporophyllar differences 

between the two genera are of minor quantitative 

nature and may be ascribed to reductive evolution in 

the direction of Palaeocycas, the younger of the two 

genera. From the standpoint of all morphologic evidence as well as stratigraphic relationships, a Palaeocycas-like plant would be a credible derivative of 

Phasmatocycas. Therein lies the most apparent relationship between American and other fossil cycads. 

Archaeocycas must be related to Palaeocycas in 

much the same way as it is related to Phasmatocycas. 

EVOLUTION OF THE PINNATE HABIT IN 

CYCADALEAN FOLIAGE 

Considerable indirect evidence favors the assumption that all the known pre-Jurassic cycads, with the 

exception of the Upper Triassic Leptocycas, bore 

entire-margined taeniopterid foliage. One thus is 

prompted to seek fossil evidence of an evolutionary 

process leading from simple undivided laminae to 

pinnate leaves of the type that characterizes most of 

the modern cycads. Gradual incision or laciniation 

of an originally Taeniopteris-like foliar lamina seems 

the most logical theme. As hypothetically proposed 

on preceding pages, the taeniopterid distal lamina 

of the sporophyll of Archaeocycas or Phasmatocycas 

became incised to produce a Cycas-like megasporophyll, and there is no good reason to discount a coordinate modification of the sterile leaves as well. 

The essence of such an evolutionary sequence among 

the cycads was briefly touched upon by Gaussen 

( 1944, p. 20) , who wrote: "Pour passer de la feuille 

de Palaeocycas a celle des Cycadites ou du type Cycas, il faut que le limbe se dechere en pinnules et que 

dans chacune la nervure fine se transforme en une 

epaisse." 

Taeniopteris or closely allied forms are abundant 

in the late Paleozoic fossil record, and among these 

are some whose margination suggests, and would 

easily fit into, an evolutionary series that produced a 

Cycas-type of leaf in which each laminar division 

contains a single vein. Taeniopteris serrulata, from 

the Lower Shihhotse Series (Lower Permian) of 
40 PALEOZOIC ORIGIN OF THE CYCADS 

Shansi, China (Halle, 1927, p. 160-161, pl. 42, figs. 

13-18, pl. 64, fig. 13), is a species that well might 

represent an early evolutionary stage in such a 

series. T. serrulata has a simple leaf and is quite 

ordinary in most morphological aspects; however, 

it has a finely serrate margin. The marginal teeth 

are very small, but each one is occupied by a single 

vein ending ( cf. fig. 11B) . 

A further step in this evolutionary process is exemplified by the lamina of Taeniopteris? koreanensis 

Cheong and Lee, 1970, from the upper part of the 

Sadong Formation in Korea. The associated plants 

indicate an Early Permian age for T.? koreanensis 

and place it in a harmonious stratigraphic light 

within this hypothetical evolutionary series. Previously given the new generic name Serratopteris 

(Cheong, 1969), this material consists of a "probably unipinnate" leaf, in which all but the basal part 

of the lamina is deeply divided into a series of essentially equal incipient "pinnules" that extend 

about halfway from the margin of the lamina to the 

midrib (cf. fig. 11C). As in T. serrulata, each division received a single vein. The illustrations suggest 

that T.? koreanensis is·a compound leaf and that the 

serrations represent incipient pinnules rather than 

pinnae, but this point is not entirely clear. It is also 

not known whether the deep serrations extend to the 

tips of the laminae, because the specimens are incomplete. Notwithstanding, this species is a near-perfect 

intermediate between a normal, entire-margined 

Taeniopteris and a Cycas leaf with its single-veined 

leaflets; T. serrulata would be the expectable starting point. Interpretation of these plants as an infallibly credi'ble series is beclouded by the uncertainty whether T.? koreanensis is a simple or compound 

leaf, but certainly the two examples demonstrate the 

possibility of a regular increasing process of laminar 

laciniation in the Paleozoic, each resultant ultimate 

foliar division supplied by a single vein, as in the 

modern monotypic family Cycadaceae (classification 

of Johnson, 1959). Culmination of the process producing T. serrulata might be envisioned in the Jurassic genus Paracycas Harris. A natural segregate 

from the form-genus Cycadites, Paracycas has 

Cycas-like pinnation and venation and is grouped 

among the cycads by Harris ( 1964, p. 65) . 

With the exception of Stangeria, all other genera 

of modern cycads are characterized by pinnae or 

pinnules that lack midribs and have several to many 

longitudinal dichotomous veins; those genera were 

placed by Johnson ( 1959) in the family Zamiaceae. 

Stangeria, monotypifying the family Stangeriaceae, 

differs markedly in that its pinnae each have a pronounced midvein with a dense system of transverse, 

parallel, dichotomously divided secondary veins. 

Just as putatively early stages in the development 

of the Cycas type of leaf are present among the 

Paleozoic taeniopterids., there are knc wn fossil forms 

that may be regarded as possibly early representatives of the zamioid type of foliage. Halle's Taeniopteris? serrata (Halle, 1927, p. 161, pl. 42, figs. 9-12), 

a pinnate form associated with T. serrulata in the 

Permian Shansi flora, has a lamina with an irregularly serrate margin. In this species, however, the 

marginal areas between adjacent teeth receive three 

or four veins, rather than one as in T. serr11Jata ( cf. 

fig. 11E). 

Taeniopteris pseudobrevis is a species described 

by Barnard ( 1967, p. 725, pl. 61, figs. 1-4; 1970, p. 

37-38, pl. 5, figs. 1, 4, text-figs. 3B, D) from Upper 

Triassic rocks of Afghanistan. In this plant, the 

foliar lamina is regularly corrugated at intervals of 

2-3 mm and is incised to as much as one-sixth the 

total width of the blade ( cf. fig. 11F). Each partial 

segment contains the ends of several simple veins. 

This leaf thus gives the impression of a simple taeniopterid leaf well on the way toward becoming a pinnate one; the multiple-veined segments would each 

correspond to a single leaflet of a Zamia-like leaf. 

Zamioid foliage appears more fully blown in the 

Upper Triasic of North Carolina, in the genus Leptocycas Delevoryas and Hope. The leaves of Leptocycas were compared with the common Jurassic 

genus Pseudoctenis, which, according to Harris 

(1964, p. 71), "would include the leaves of more 

than one genus of Recent Cycads." The lamina in 

this case is completely divided into essentially equal 

linear segments, each containing several parallel 

veins. This leaf is interpreted here as a remarkably 

precocious end product of laminar dissection, and 

consonantly Leptocycas, if truly a zamioid cycad, 

represents the earliest known appearance of the 

zamioid lineage. 

Jurassic time, during which explosive development of the cycadophytes took place, produced a 

broad variety of foliar types. Some forms are regularly pinnate and quite cycadlike in most aspects; 

others, however, show significant intermediate steps 

in the laciniation, or pinnation, of originally entiremargined taeniopterid laminae. The specimen illustrated by Seward (1917, fig. 615) as Pterophyllum 

(Anomozamites) Nilssoni from the Middle Jurassic 

beds of Yorkshire, England, is a particularly interesting form. It has a very irregular but basically 

taeniopterid lamina, some parts of which are divided as far as the midrib into unequal truncated 
MORPHOLOGIC AND EVOLUTIONARY CONSIDERATIONS 41 

segments; the apical one-third of the lamina is 

nearly entire, however. This species was later placed 

in Anomozamites by Harris (1969, p. 80) and is regarded as bennettitalean. The other species of Anomozamites, A. thomasi Harris, has nearly uniform 

segmentation of the lamina. 

The genus Nilssonia is especially pertinent to the 

rationale of this theory of cycadean foliar evolution, 

because Nilssonia species show extremes of variable 

laminar dissection that occur in no other contemporary plant. Harris' (1932b, p. 40-52) descriptions 

of the Lower Jurassic species of Nilssonia from East 

Greenland revealed interesting examples. In some, 

the lamina may be consistently entire (N. undulata 

Harris), but in others (N. polymorpha Schenk; cf. 

fig. 11G) the lamina is entire or deeply incised into 

irregular segments. Usually the segments on one side 

of the leaf have no regular relationship to those on 

the other. InN. incisoserrata Harris, (1932a; p. 49-

52) the leaf is mostly "incisoserrate," uniform segmentation reaching only part of the distance to the 

midrib ( cf. fig. 11F) ; this form resembles Taeniopteris pseudobrevis Barnard in the partial segmentation of the lamnia. N. obtusa (N athorst) Harris 

usually has entire leaves, but some have deep incisions at long intervals. Asama ( 1968) pointed out 

a trend observed in some Japanese forms of Nilssonia in which the entire-margined Early Jurassic 

N. orientalis supposedly gave rise to the smaller Late 

Jurassic N. orientalis var. minor. Asama assumed 

the series to culminate in the still smaller, Early 

Cretaceous N. schaumburgensis, in which the lamina 

is dissected into irregular segments as in N. tenuine'i·vis. Asama's article, however, was intended to 

demonstrate a decrease in laminar size with the passage of time, and did not express any ideas regarding cycadean leaf derivation from an entiremargined ancestral form. 

Other examples of irregular segmentation in Nilssonia are widely documented, but only one more, N. 

tenuinervis Seward, need be discussed here because 

of its established cycadean affinity. Harris (1941) 

presented compelling evidence that Androst'i·obus, 

a cycadean male cone; B eania, a female cone; and the 

leaf Nilssonia are parts of the same plant and that 

the plant is certainly a cycad. Thomas and Harris 

(1960) defined specific relationships with the conclusion that Beania mamayi, Androstrobus wonnacotti and Nilssonia tenuinervis, all of Middle Jurassic age, belonged to the same plant. They (p. 147) 

stated that 

the Nilssonia plant is to be classified as a member of the 

Cycadales, but though nearest to the Zamioideae in its female 

cones, it should not be included in the Zamioideae because the 

female cones and probably the male cones differ from the 

cones of all Zamioideae in their lax and open construction. 

In the same article, Thomas and Harris ( 1960, p. 

150) concluded that Androstrobus prisma Thomas 

and Harris and Pseudoctenis lanei Thomas emend. 

Harris, were male cone and leaf of the same plant. 

Unlike Nilssonia tenuinervis, Pseudoctenis lanei is 

in most aspects remarkably similar to leaves of 

Zamia or Encephalartos, and Thomas and Harris 

( 1960, p. 153) remarked on the zamioid aspects of 

this combination of organs: 

Taking the two fossils together the agreement with the 

Zamioideae is much enhanced, and without speculating about 

the unknown female cone, we can say that in Pseudoctenis 

lanei with Androstrobus prisma we have a plant which is so 

like them that it may prove to belong to one of these living 

genera. 

Harris' conclusions regarding the relationships 

between Nilssonia tenuinervis, Androstrobus wonna.cotti and B eania mamayi led him to publish a 

restoration of a cycad bearing those three organs 

(Harris, 1961, text-fig. 2). It shows the irregular 

construction of the taeniopterid leaves, with long 

laminar segments separated by clefts extending as 

far as the midribs. 

The morphologic variability discussed here clearly 

expresses the manifestation of a trend involving 

evolutionary modification of leaves constructed on 

the simple taeniopterid pattern. The oldest taeniopterids are of late Paleozoic age and are mostly entiremargined; in the Permian, however, the early beginnings of marginal incision are seen. The trend progresses to the extent that, by Late Triassic time, 

laminar dissection was complete in at least one example; by Middle Jurassic time it was complete in 

several taxa, although resultant foliar segments 

were not always of uniform size. To be sure, certain 

anomalies appear in this trend: the apparent precocity in Leptocycas and the conservatism in Nilssonia. These express the vagaries of morphogenetic 

rates within a group of rapidly evolving organisms. 

Nonetheless, the degree of laminar dissection seems 

generally consistent with decreasing geologic age of 

materials. Furthermore, the evolutionary process 

involved stages leading to two of the basic types of 

cycadean foliage-the Cycas type having singleveined leaflets, and the Zamia type with multipleveined divisions. It is even feasible to postulate derivation of the Stangeria type of leaf from Paleozoic stock, for pinnately compound taeniopterids are 

known, and each leaflet of a Stangeria leaf is basically equivalent to a simple Taeniopteris leaf in overall 
42 PALEOZOIC ORIGIN OF THE CYCADS 

PALEOZOIC MESOZOIC CENOZOIC 

A 

I 

H 
MORPHOLOGIC AND EVOLUTIONARY CONSIDERATIONS 43 

shape, venation, and margination. Thus, a pinnately 

compound taeniopterid leaf could evolve into a Stangeria leaf with only relatively minor modifications. 

Further, the bipinnate Bowenia type of leaf might 

have evolved from a pinnate taeniopterid ancestor 

through incision of the leaflets into multiveined 

segments. 

Intimate association of leaves and fructifications 

has been of utmost importance in tracing the course 

of early cycadean evolution; cuticular comparisons, 

although not always possible, have also constituted 

critical evidence. In all instances where fossil leaves 

and reproductive parts have been suspected or demonstrated to represent the same plant and that plant 

has been interpreted as a cycad, the leaves have been 

taeniopterid or almost certainly of taeniopterid derivation. This circumstance strengthens the proposition that the cycads arose from taeniopterid pteridosperms and that modern cycad leaves have been 

derived from either simple or pinnate taeniopterid 

leaves through relatively little evolutionary modification, or through an evolutionary process no more 

complicated than laminar dissection. 

Substantially the same statement may be made in 

regard to cycadeoid foliage. Taeniopteris has long 

been accepted as an artificial (form) genus representing Cycadales, Bennettitales, and Pentoxylales; 

nomenclatural and taxonomic adjustments have been 

made to accommodate the situation (Harris, 1969, p. 

68; Sahni, 1948, p. 50). Although conclusions cannot 

yet be drawn relative to a possibly common ancestry 

between the Cycadales and Bennettitales or the age 

of the ancestral complex, it seems reasonable, on the 

basis of present knowledge of Bennettitalean leaves, 

to suppose that they too arose from a simple taeniopFmuRE H.-Suggested evolution of cycadalean leaves beginning with (A), an entire-margined ances:bral taeniopterid 

leaf and, through progressively deeper incision of margins 

resulting in two basic types of cycadalean leaves (D, cycad~ aceous type; I, zamiaceous type). All diagrams are based 

?n ~nown fossil or modern forms. The cycadaceous lineage, 

mdiCated by heavy arrows, consists of forms, in which 

marginal teeth and ultimate foliar segments involve: a single vein each (B, cf. Taeniopteris serrulata; C, cf. T.? 

k?reanensis; D, cf. Cycas sp.). The zamiaceous lineage, indiCated by light arrows, consists of forms in which marginal 

incisions and ultimate foliar segments involve several veins 

each (E, cf. Taeniopteris? serrata; F, cf. T. pseudobrevis 

or Nilssonia incisoserrata; G, cf. Nilssonia tenuinervis or 

N. polymorpha; H, cf. N. tenuicaulis; I, cf. Zamia sp.); 

forms such as E and G might have been derived independently from A, whereas H may have arisen from either F 

or G. Both cycadaceous and zamiaceous lineages evolved 

substantially concurrently. Ages of forms: A, Pennsylvanian-Permian; B, E, Permian; C, Triassic; F, G, TriassicJurassic; H, Jurassic; D, I, Mesozoic-Cenozoic. 

terid leaf form through a process parallel to that 

proposed here for the cycads. The sum of presently 

available evidence, then, suggests that Taeniopteris 

constitutes a logical source of evolutional raw material for derivation of the open-veined cycadophytic 

leaf, be it bennettitalean or cycadalean. 

CONCLUSIONS 

The morphology, age, and geographic occurrences 

of the fossils discussed here combine to present a 

significant set of conclusions, bearing on various aspects of the history of some of the gymnosperms. 

Archaeocycas and Phasmatocycas are sufficiently 

cycadlike to be classified among the cycads ; P hasmatocycas is particularly convincing because of its 

twin-ranked, well-preserved seed cuticles. At the 

same time, these genera are so logically derivable 

from Spermopteris that the cycadean lineage is 

clearly extended to the Pennsylvanian pteridosperms; this conclusion corroborates the opinions of 

Harris (1961) and others and injects additional 

credence into the few known occurrences in the Paleozoic of petrified wood with cycadlike anatomical 

features (Cycadoxylon; Andrews, 1940). These fossils eminently represent the pre-Jurassic "progressive evolution" postulated by Harris (1961, p. 322). 

Thus, the stratigraphic continuity of the cycadalean 

lineage is extended from the Upper Triassic to the 

Upper Pennsylvanian, and the cycads join the conifers as one of the two oldest groups of extant seed 

plants. 

On the basis of present information, we cannot 

know when the spermopterid ancestors of the cycads 

ceased to be pteridosperms and became cycads, even 

though the relationship between the two taxa seems 

clear. Concordantly, the possible impact of the Paleozoic fossils on the broad classification of the cycads 

cannot yet be assayed, and a revised system will not 

be attempted here; such a system will depend ultimately on the acquisition of knowledge of other 

parts of the Permian and Triassic cycads, such as 

the male organs and vascular anatomy. Nonetheless, 

Spermopteris, Phasmatocycas and Archaeocycas appear to uphold the concept that Cycas, with its leaflike megasporophylls, is the primitive genus among 

extant cycadales; this appraisal has been the popularly accepted one, although Thomas (1947, p. 250; 

1953, p. 582) held the contrasting view that Zamia, 

rather than Cycas, is the primitive member of the 

Cycadales. 

The evidence for taeniopterid foliation among the 

early cycads and the indications that gradual lami-
44 PALEOZOIC ORIGIN OF THE CYCADS 

nar dissection of entire leaves produced the pinnate 

types of modern cycad foliage are an important 

facet of this study, but they also pose a perplexing 

problem-the origin of taeniopterid foliage-whose 

solution will necessitate extensive searches of prePennsylvanian rocks. Taeniopteris appears "suddenly" in Pennsylvanian rocks and gives no clues as to 

its ancestry; it is a conspicuous element in the Permian and Triassic floras. In keeping with his 

theories of leaf evolution, Asama (1960, p. 258) believes that the simple leaf of Taeniopteris is an example of an "enlarged leaf," formed from an originally pinnate form. Kon'no, Asama, and Rajah 

( 1970, p. 559), however, stated that no substantial 

evidence yet exists in the Cathaysian flora for this 

opinion, and I know of none in the Angara. European, or American floras. It is conceivable, of course, 

that Asama's notions are correct and that intermediate forms ancestral to Taeniopteris simply have not 

been found. 

Geologic age and geologic occurrence of the fossils 

discussed here are significant considerations, for 

they contribute to the ever-growing number of indications that the Lower Permian deposits of the 

southwestern United States contain proof that this 

area was an important center of botanical evolution 

and radiation. Late Pennsylvanian plants such as 

Spwrmoptwris emphasize the importance of that time 

in respect to the beginnings of major innovations in 

plant form. Wolfcampian time produced the Permian guide fossil, Callipteris, but this genus has no 

more morphological significance than that of a 

younger type of seed-fern with a "marker" type of 

foliage. 

Leonardian time, however, saw the introduction 

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different forms, some of which leave no grounds for 

fnterpretation as to antecedents or progeny; Gigantopteris americana White and Russellites taeniata (Darrah) Mamay are prime examples. Reproductive methods of these two ·genera are yet speculative. They are probably seed plants, but this speculation is based only on their age and the otherwise 

demonstrated progress of plant evolution as shown 

by fossils. 

Thus far, the advent of cycads is the most important botanical event attributable to the Leonardian or even to the Permian System. It emphasizes 

the importance of Leonardian rocks as the physical 

representation of a time interval worthy of close 

examination in the search for progressive evolution 

among higher plants. 

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46 PALEOZOIC ORIGIN OF THE CYCADS 

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Gaz., v. 110, no. 1, p. 47-80. 

Schimper, W. P., 1870-1872, Traite de paleont6logie vegetale 

ou la flore du monde primitif: Paris, J. B. Bailliere et 

Fils, 3 v., and atlas containing 110 pls. 

Schuster, J., 1932, Cycadac.eae, 1'n Engl·er, Adolph, Das 

Pflanzenreich: Leipzig and Berlin, W. Engelmann, v. 4, 

pt. 1, p. 1-168. 

Sellards, E. H., 1901, Permian plants-Taeniopteris of the 

Permian of Kansas: Kansas Univ. Quart., v. 10, no. 1, 

p. 1-12, pl. 1-4. 

--- 1908, Fossil plants of the Upper Paleozoic of Kansas: 

Kansas Univ. Geol. Survey, v. 9, p. 386-480, pls. 44-69. 

Seward, A. C., 1917, Fossil plants: London, Cambridge Univ. 

Press, v. 3, 656 p. 

Sporne, K. R., 1965, The morphology of gymnosperms: London, Hutchinson Univ. Library, 216 p. 

--- 1971, The mysterious origin of flowering plants: London, Oxford Univ. Press, 16 p. 

Taylor, T. N., 1969a, Cycads--evidence from the Upper Pennsylvanian: Science, v. 164, no. 3877, p. 294-295. 

--- 1969b, Evidence for the pres.ence of cycads in the 

Upper Pennsylvanian: Internat. Bot. Cong., 11th, Seattle, 

1969, Abstracts, p. 216. 

--- 1970, Lasiostrobus gen. n., a staminate strobilus of 

gymnospermous affinity from the Pennsylvanian of North 

America: Am. Jour. Botany, v. 57, no. 6, pt. 1, p. 670-

690. 

Thomas, H. H., 1947, The history of plant form: British 

Assoc. Adv. Sci. Reports, v. 4, no. 15, p. 243-254. 

--- 1953, The relationships between the Bennettitales and 

Cycadales: Internat. Bot. Cong., 7th, Stockholm, 1950, 

Proc., p. 581-582. 

Thomas, H. H., and Harris, T. M., 1960, Cycadean cones of 

the Yorkshire Jurassic: Senckenbergiana Lethaea, v. 41, 

no. 1-6, p. 139-161, pl. 1-4. 

Uhl, N. W., and Moore, H. E., 1971, The palm gynoecium: 

Am. Jour. Botany, v. 58, no. 10, p. 945-992. 

U.S. Geological Survey, 1968, Geological Survey Research, 

1968: U.S. Geol. Survey Prof. Paper 600-A, 371 p. 

Von Gutbier, C. A., 835, Abdri.icke und Versteinerungen des 

Zwickauer Schwarzkohlengebirges und seiner Umgebungen: Zwickau, G. Richter, 80 p., 11 pl. 

White, C. D., 1912, The characters of the fossil plant Gigantopteris Schenk and its occurrence in North America: 

U.S. Natl. Mus. Proc., v. 41, no. 1873, p. 493-516. 

--- 1929, Flora of the Hermit ·shale, Grand Canyon, Arizona: Carnegie Inst. Washington Pub. 405, 221 p., 51 pl. 

Wilson, J. A., 1953, P.ermian vertebrates from Taylor County, 

Texas: Jour. Paleontology, v. 27, no. 3, p. 456-4 70. 
INDEX 

[Italic page numbers indicate descriptions and major references] 

Page 

A 

abnormis, Taeniopteris ---------------- 7, 8 

americana, Gigantopteris ___ 5, 8, 24, 26, 27, 44 

Androstrobus ------------------------- 41 

prisma --------------------------- 41 

wonnacotti ------------------------ 24,41 

zamioides _____ --------- __ ________ 24 

angustifolia, Danaeopsis -------------- 36 

Annularia ---------------------------- 26,27 

Anomozamites ------------------------ 41 

thomasi -------------------------- 41 

Nilssoni, Pterophyllum ------------ 40 

(Anomozamites) Nilssoni, Pterophyllum 40 

Aphlebia ----------------------------- 27 

Araucarites -------------------------- 26, 27 

Archaeocycas __ 4, 6, 8, 10, 11, 12, 14, 17, 18, 19, 

22, 27, 28, 29, 30, 31, 32, 33, 34, 35, 36, 38, 39, 43 

whitei _______ 8, 11, 12, 31, 32,; text-fig. 3; 

Aspidiopsis 

pls. 1, 2 

27 

B 

Beania ----------------------- 2, 15, 19, 36, 41 

gracilis --------------------------- 15 

mamayi -------------------------- 41 

Bennettites --------------------------- 34 

Bennettitolepis ------------------------ 25, 26 

dactylota ------------------------- 25 

Bjuvia -------------------------- 2, 17, 38,39 simplex __________________________ 25, 38 

Bowenia ------------------------------ 43 

c 

Callipteris 17, 26, 27, 28,44 

<Jardiocarpon -------------------------

Carpolithes _ --------------------------

27 

26 

Ceratozamia -------------------------- 20 

circinalis, Cycas ___ ------ _______ ------ 24 

Cordaianthus ------------------------- 27 

Cordaites ----------------------------- 27 

coriacea, Spermopteris ---------------- 2 

Taeniopteris ---------------------- 2, 16 

linearis, Taeniopteris ------------ 16 

Cycadites ----------------------------- 40 

Cycadospadix ----------- 2, 6, 25, 26, 27, 28, 34 

dactylota ------------------------- 25 

Hennocquei ----------------------- 25 

integer --------------------------- 25, 26 

Moraeanus ------------------------ 25 

yochelsoni ------------------- 26, 33; pl. 2 

sp ------------------------------- 6, 19 

Cycadoxylon -------------------------- 43 

Cycas ----------- 3, 4, 12, 19, 25, 26, 30, 31, 32, 

33, 34, 38, 40, 43 

circinalis ------------------------- 24 

media ______ --- __ -- _ -------------- revoluta _________________________ _ 

sp -------------------------------

D 

3, 33 

43 

dactylota, Bennettitolepis ------------- 25 

Cycadospadix --------------------- 25 

Danaeopsis angustijolia ---------------

Page 

36 Daubreeia _ __ _ __ _ __ _ __ _ __ __ __ _ __ __ _ __ _ 27 

Dioon edule --------------------------- 3, 20 

Dioonites pennacformis --------------- 36 

Dioonitocarpidium -------- 2, 17, 36, 37, 38, 39 

keuperianum --------------------- 37, 38 

liliensterni ----------------------- 37,38 

pennaeforme ------------------ 36, 37, 38 Diplothmema _ __ _ __ _ __ _ __ _ ____ ___ __ _ __ 27 

Discinites ----------------------------- 27 

E 

edule, Dioon -------------------------- 3, 20 

Enccphalartos ______ ------------------ 20, 41 

F 

jloridana, Zamia 3 

G 

gigantea, Taeniopteris ---------------- 38 

Gigantopteris ---------------------- 26, 27,28 

americana ------------- 5, 8, 24, 26, 27, 44 

n. sp ---------------------------- 27 

Ginkgo ------------------------------- 18 

Glenopteris -------------------- 17, 26, 27,28 

Gomphostrobus ----------------------- 27 

gracilis, Beania ----------------------- 15 

Leptocycas ___ ------··-------------

grigsbye, Lyssoxylon -----------------

H 

H ennocquei, Cycadospadix 25 

incisoserrata, Nilssonia --------------- 41,43 

integer, Cycadospadix ----------------- 25,26 

Palaeocycas ------------------- 20, 25, 38 

K 

kansana, Phasmatocycas _ 12, 21, 23, 26, 29, 30, 

32; text-fig. 4; pls. 3, 4 

keuperianum, Dioonitocarpidium ______ 37, 38 

koreanensis, Taeniopteris ------------- 40, 43 

L 

Lagenostoma 20 

Lomaxi --------------------------- 19 

lanei, Pseudoctenis -------------------- 41 

Lasiostrobus -------------------------- 6 

polysacci ------------------------- 6 

Leptocycas ------------------- 5, 6, 39, 40,41 

gracilis -----------~--------------- 6 

liliensterni, Dioonitocarpidium -------- 37, 38 

linearis, Taeniopteris coriacea --------- 16 

Lobatannularia ----------------------- 27 

Lomaxi, Lagenostoma ----------------- 19 

Lyginopteris oldhamia ---------------- 19 

Lyssoxylon --------------------------- 6 

grigsbye -------------------------- 6 

M 

Macrozamia 

miquelii _________ ------------------

mamayi, Beania ----------------------

media, Cycas -------------------------

minor, Nilssonia orientalis ------------

miquelii, Macrozamia -----------------

Moreanus, Cycadospadix --------------

multinervis, Taeniopteris -------------

N 

Neuropteris 

newberriana, Taeniopteris -------------

Nilssonia ___ -------------------------- incisoserrata _________ ------------

obtusa ______ - ________ ------ _- ----

orientalis ______ -------------------

Page 

20,24 

3 

41 

3 

41 

3 

25 

7, 19 

27,28 

17 

41 

41,43 

41 

41 

minor ------------------------ 41 

polymorph a ________________ ------- 41, 43 

schaumbttrgensis ------------------ 41 

tenuicaulis ------------------------ 43 

tenuinervis ----------------------- 41,43 

undulata ------------------------- 41 

N oeggerathia _____________ -- ___ ------- 26 

Nypa -------------------------------- 34 

0 

obtusa, Nilssonia --------------------- 41 

Odontopteris ------------------- 17, 26, 27,28 

oldhamia, Lyginopteris --------------- 19 

orientalis, Nilssonia ----------------- 41 

minor, Nilssonia ----------------- 41 

p 

Padgettia ---------------------------- 12 

Palaeocycas ------------- 2, 4, 25, 26, 36, 38, 39 

integer ----------------------- 20, 25, 38 

Paracycas ---------------------------- 40 

Pecopteris ---------------------------- 26,27 

pennaeforme, Dioonitocarpidium ---- 36, 37,38 

pennaeformis, Dioonites --------------- 36 

Phasmatocycas __ 6, 13, 14, 15, 16, 17, 18, 19, 20, 

21, 22, 23, 24, 27, 28, 29, 30, 

31,32,3~34,36,3~39,43 

lcansana --------- 12, 21, 23, 26, 29, 30, 32; 

text-fig. 4; pls. 3', 4 

spectabilis ________________ 22, 30, 32; pl. 5 

sp ----------------------------- 27; pl. 2 

Plagiozamites ------------------------- 6 

planchardi --------------------- 6 

planchardi, Plagiozamites ------------- 6 

Poacordaites -------------------------- 27 

polymorpha, Nilssonia ---------------- 41, 43 

polysacci, Lasiostrobus ---------------- 6 

prisma, Androstrobus ----------------- 41 

pseudobrevis, Taeniopteris --------- 40, 41, 43 

Pseudoctenis -------------------------- 40 

lanei ----------------------------- 41 Pterophyllum ___ ------ _____ -- _ -- 40 

47 
48 

Page 

R 

revoluta, Cycas ----------------------- 3, 33 

Rhabdocarpos ------------------------ 27 

RusseUites ---------------------------- 27,28 

taeniata -------------------------- 44 

s 

schaumburgensis, Nilssonia 41 

serrata, Taeniopteris ----------------- 40,43 

Serratopteris ------------------------- 40 

serrulata, Taeniopteris ------------- 39, 40,43 

Sigillaria ----------------------------- 27 

Sigillariostrobus --------------------~-- 27 

simplex, Bjuvia ----------------------- 25,38 

spectabilis, Phasmatocycas ___ 22, 30, 32; pl. 5 

Spermopteris ------ 2, 3, 4, 6, 7, 8. 10, 12, 13, 17, 

18, 21, 22, 23, 26, 28, 29, 

30, 31, 32, 34, 36, 43, 44 

coriacea -------------------------- 2 

sp ------------------------------ 7; pl. 2 

INDEX 

Page 

Sphenophyllum ------------------------ 26 

Sphenopteris ------------------------- 27 

Stangeria ------------------------- 40, 41,43 

Stigmaria ---------------------------- 27 

Supaia 28,29 

T 

taeniata, Russellites ------------------ 44 

Taeniopteris _____ 2, 7, 19, 24, 26, 28, 29, 30, 39, 

41, 43, 44 

abnormis ------------------------- 7, 8 

coriacea -------------------------- 2, 16 

linearis ----------------------- 16 

gigantea 38 

koreanensis ---------------------- 40, 43 

multinervis ------------------------ 7,19 

newberriana ---------------------- 17 

pseudobrevis ------------------- 40, 41,43 

serrata --------------------------- 40.43 

serrulata --------------------- 39, 40,43 

sp -------------------------------- 17 

Page 

tenuicaulis, Nilssonia ----------------- 43 

tenuinervis, Nilssonia ----------------- 41, 43 

thomasi, Anomozamites --------------- 41 

Tinsleya ------------------------------ 12 

u 

undulata, Nilssonia 41 

w 

Walchia 26,27 

whitei, Archaeocycas -------- 8, 11, 12, 31, 32; 

text-fig. 3; pls. 1, 2 

wonnacotti, Androstrobus ------------- 24, 41 

y 

yochelsoni, Cycadospadix ________ 26, 33; pl. 2 

z 

Zamia 40, 41,43 

floridana ___ ______ ___ __ ____ _______ ____ _ 3 

sp ------------------------------- 43 

zamioides, Androstrobus -------------- 24 

*U.S.GOVERNMENT PRINTING OFFICE: 1976 0-211-317/144 
PLATES 1-5 

Contact photographs of the plates in this report are available, at 

cost, from U.S. Geological Survey Library, Federal Center, 

Denver, Colorado 80225 
FIGURES 

PLATE 1 

1-5. Archaeocycas wlvitei Mamay (p. 8). 

1. Megasporophyll exposed in oblique-lateral position, showing carbonaceous remnants of original tissues, indusiumlike laminar covering of ovules and faint striations suggestive of fine, parallel venation. Paratype, USGS 8959-1b. X 8.5. 

2. Counterpart of specimen illustrated in figure 1, showing adaxial view of six closely appressed ovules. 

Note incomplete fold in lamina at upper (right) edge of most distal ovule (arrow), indicating flattening out of inrolled lamina covering the ovules. Paratype, USGS 8959-la. X 8.5. 

3. Mold of adaxial side of megasporophyll, showing stout midrib, five pairs of oppositely attached ovules, 

and distal expansion of sterile lamina. Paratype, USGS 8877-3. X 6. 

4. Cast of adaxial side of megasporophyll, showing stout midrib, five pairs of opposite ovules, and coalified residue of laminar tissue that originally intruded between contiguous ovules. Several ovules show 

circular scars of attachment; these are best shown in the distal (uppermost) pairs of ovules. Paratype, USGS 8877-2. X 6. 

5. Cast of adaxial side of megasporophyll, with nine ovules (lower right ovule broken off). Note fairly 

strong development of sterile distal lamina, with fold extending distally and diagonally from midrib. 

Holotype, USGS 8877-1. X 6. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 934 PLATE 1 

ARCHAEOCYCAS 
FIGURE 

PLATE 2 

1. ?Phasmatocycas sp. (p. 21). 

Fragmentary impression of stout axis with evidence of approximately 12 pairs of opposite seedlike bodies 

and suggestion of outline of surrounding foliar lamina. USGS 10064-1. X 2. 

2. Spermopteris sp. (p. 7). 

Fragmentary impression of taeniopterid leaf, bearing two marginal rows of ovoid swellings suggestive of 

immature ovules. USGS 6233-1. X 2. 

3. Cycadospadix yochelsoni n. sp. (p. 24). 

Small appendage with broad peduncle and deeply pinnatifid lamina. Holotype, USGS 10057- 1. X 3. 

4. ?Cycadean male cone (p. 23). 

Fragmentary mold of conelike organ, with deep rhomboidal impressions, probably representing distal faces 

of microsporophylls. Note distally curved spinelike molds, apparently impressions of rigid attenuated 

termini of microsporophylls. USGS 10064-2. X 1. 

5-7. Archaeocycas whitei Mamay (p. 8). 

5. Mold of adaxial side of megasporophyll with four pairs of ovules. Each ovule shows a circular scar of 

attachment near the rachis; three lower left ovules in turn have tiny central markings, probably representing vascular strands from lamina to base of ovules. USGS 8959-2a. X 6. 

6. Counterpart of specimen shown in figure 5. USGS 8959-2b. X 6. 

7. Cast of megasporophyll, exposed .in lateral view. Most proximal ovule partly broken away; contiguous 

ovule showing attachment ·scar near midrib. Note expanded base of midrib, indicative of perpendicular attachment to an axis; also note fragmentary but apparently well-developed sterile lamina at 

right. Paratype, USGS 8877- 4. X 6. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 934 PLATE 2 

1 

4 

?PHASMATOCYCAS, SPERMOPTERIS, CYCADOSPADIX, AND ARCHAEOCYCAS 
FIGURES 

PLATE 3 

1-5. Phasmatocycas kansana Mamay (p. 12). 

1, 2. Counterparts of holotype, showing stout axis with two lateral rows of seeds, one displaced from original position. Figure 1, USGS 8869-1a; figure 2, USGS 8869- 1b. X 4. 

3. Seed on USGS 8869-1b, oriented with apex upward. Spherical glandlike bodies at base of seed (indicated by arrows) one at either side. X 21. 

4. Part of USGS 8869-1b, enlarged. Second complete seed from top at left side appears in figure 3. 

X 9. 

5. Line tracing of figure 4, showing distribution of glandlike bodies relative to seeds. Solid black circles represent actual glands; unshaded circles represent impressions of glands, X 9. 
GEOLOGICAL SUHVEY 

4 

PHASMATOCYCAS 

PROFESSIONAL PAPER 934 PLATE 3 

3 
FIGURES 

PLATE 4 

1-5. Phasmatocycas kansana Mamay (p. 12). 

Chemically cleared cuticular systems of three seeds removed from holotype (USGS 8869-1b). 

1, 2. Apical region of seed fragment, showing short, relatively broad micropyle and two cuticles. Outer 

cuticle (inner integument) is thin and delicate; inner layer (nucellar integument) is thicker, with 

isodiametric, heavy-walled cells. Note absence of any pollen-chamber mechanisms. Fig 1, X 100; 

fig. 2, X 30. 

3. Nearly complete specimen, showing double nature of cuticle to approximate "shoulders," and heavy 

megaspore membrane. Light area below center represents damage in which part of megaspore 

membrane ·is missing; darkest streaks represent folds in megaspore membrane, in which at least 

three thicknesses of membrane are present. X 30. 

4, 5. Two views of third specimen, fig. 5 showing details of micropylar area, double cuticle, and top of 

megaspore membrane seen as a dark arc across bottom of figure. Fig. 4, X 30; fig. 5, X 100. 
GEOLOGICA L SURVEY 

2 

5 

PHASMATOCYCAS 

; ~ 

1(.. t 

·;'I 

PROFESSIONAL PAPER 934 !'LATE 4 

3 
FIGURES 

PLATE 5 

1-4. ?Phasmatocycas spectabilis Mamay, n. sp. (p. 22). 

1. Unusually long, stout axis with about 60 seeds, borne in two lateral rows. Holotype, USGS 8959-3. 

XL 

2-4. Paratype specimens, clearly demonstrating lack of evidence of seed attachments between lateral rows. 

Fig. 2, USGS 8877-5, X 1; fig. 3, USGS 8877-6, X 2; fig. 4, USGS 8877-7, X 1. 

5-7. Specimen of Taeniopteris sp., associated with holotype of Phasmatocycas kansana, and showing abundant 

spherical, intercostal, resinoid glandlike bodies. Compare those in fig. 7 with similar bodies shown in pl. 

3, fig. 3. Figure 5, X 2; fig. 6, X 5; figure 7, X 20. USGS 8869-2. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 934 PLATE 5 

6 

5 

?PHASMATOCYCAS AND TAENIOPTERIS