Two New Plant Genera of 

Pennsylvanian Age from 

Kansas Coal Balls 

By SERGIUS H. MAMA Y 

A SHORTER CONTRIBUTION TO GENERAL GEOLOGY 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 254-D 

Descriptions and illustrations of 

Tyliosperma and Sclerocelyphus 

fructi.ft cations 

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON: 1954 
UNITED STATES DEPARTMENT OF THE INTERIOR 

Douglas McKay, Secretary 

GEOLOGICAL SURVEY 

W. E. Wrather, Director 

• 

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

Washington 25, D. C. - Price 55 cents (paper cover) 
CONTENTS 

Page 

Abstract------------------------------------------------------------------------------------------------------ 81 Introduction__________________________________________________________________________________________________ 81 

Souroo, age of the materiaL ______________________________________________________________________________ -______ 81 

Systeinatiodescriptions_________________________________________________________________________________________ 82 

Sclerocelyphus oviform us Mamay, n. gen., n. sp _______________________________________________ ---- ____ --------- 82 

Tyliosperma orbiculatum Mamay, n. gen., n. sp __________________________________________________ · ___________ - _- 85 

Morphological and systematic considerations _______________________________________________________________ - __ --_- 93 

Literaturecited------------------------------------------------------------------------------------------------ 95 

ILLUSTRATIONS 

Page 

PLATE 21. Photomicrographic illustrations of Sclerocelyphus oviform us Mamay _______________________ - ___ - ____ -- _- Follows 95 

22-23. Photomicrographic illustrations of Tyliosperma orbiculatum Mamay _______________________________ - _--- Follows 95 

FIGURE 2. Camera Iucida drawings of serial peel sections of Tyliosperma orbiculatum Mam&Y---------------------------- 88, 89 

3. Reconstruction showing external aspect of Tyliosperma orbiculatum Mamay ___________________ - ____ - _- __ ---- 92 

4. Partially restored semidiagrammatic median longitudinal section of Tyliosperma orbiculatum Mamay- - --- _----- 92 

III 

A SHORTER CONTRIBUTION TO GENERAL GEOLOGY 

TWO NEW PLANT GENERA OF PENNSYLVANIAN AGE FROM KANSAS COAL BALLS 

BY SERGIUS H. MAMAY 

ABSTRACT 

The strip-mined coal producing area of southeastern Kansas 

has recently been discovered to contain coal balls in great abundance. The fossil plant contents of these coal balls are extremely 

diverse; as a result of the present investigation of this material, 

two new genera: of fructifications may now be added to American 

Pennsylvanian floral assemblages. The generic names Sclerocelyphus and Tyliosperma are proposed for these fructifications. 

Sclerocelyphus is a peculiar sporangium containing numerous 

small spores; in certain features it resembles the megasporangium 

Mittagia seminiformis Lignier. Tyliosperma is a small cupulate 

pteridospermous seed whose structural simplicity suggests a 

primitive position among the seed ferns. 

INTRODUCTION 

This paper deals with two new Pennsylvanian fructifications that were found in coal balls collected at the 

strip pits of the Pittsburgh and Midway Coal Company 

near West Mineral, in southeastern Kansas. Investigations of coal balls from this recently discovered rich 

locality are now being conducted in several American 

paleobotanical laboratories. Baxter (1951) recently 

published a report of this coal ball occurrence; Delevoryas and Morgan (1952) have described a new species 

of Tubicaulis (T. multiscalarijormis) from this locality, 

and a preliminary survey of the flora has recently appeared (Andrews and Mamay, 1952). Although these 

studies are yet in their incipient stages, it has already 

become obvious that the vegetable content of these 

petrifactions represents probably the most diverse 

floral assemblage yet discovered in coal balls. Thus, 

distinct new entities, such as that described by Delevoryas and Morgan and those to be described in this 

paper, may reasonably be expected to be found in some 

abundance as the investigations progress. 

The two fructifications that form the subject of this 

paper entail se~eral fundamental points of morphological interest, whiGh will be discussed at some length in 

the conclusion. Aside from this, further interest lies 

in their appearance as two new generic additions to 

American coal-ball floral assemblages. 

Sclerocelyphus, the first fructification to be discussed, 

is a peculiar sporangium containing numerous small 

spores; thus, it may be either the microsporangium. of a 

heterosporous plant or the fructification of an otherwise unknown homosporous form. Although there 

is some evidence at hand in favor of it being a microsporangium, this cannot be definitely established at 

present. Sclerocelyphus is, in several considerations, a 

most enigmatic fructification. It differs strikingly from 

any fructification hitherto recorded in American fossil 

floras; indeed, the single fossil plant with which it compares favorably as to structural details is the megasporangium described by Lignier (1913) as Mittagia seminiformis, found in coal balls from the Koksfloz in Lower 

Silesia. 

Tyliosperma, the second fructification, is a small, 

simple seed whose organization indicates affinities with 

the pteridosperms. While its gross morphological 

features are reminiscent of such other Paleozoic seeds as 

Conostoma and Lagenostoma, its structural details are on 

the whole simpler and convey the impression of a more 

primitive fructification. Aside from its structural 

simplicity, Tyliosperma is of additional interest in that 

its organic connection with a cupulate organ has been 

observed. Such instances of demonstrable attachment 

between petrified seeds and supporting organs are rare; 

the classical investigation of Lagenostoma lomaxi by 

Oliver and Scott (1904) may be cited here as the outstanding example of this sort of discovery. Unfortunately, however, the single specimen of Tyliosperma 

in which this connection has been demonstrated does 

not shed any light upon the question of the identity 

of the stem that bore these interesting seeds or the 

position they occupied on the parent plant. 

SOURCE, AGE OF THE MATERIAL 

The specimens were found in coal balls collected at 

the strip pits of the Pittsburgh and Midway Coal 

Company, about 4 miles south of West Mineral, Kans. 

(Columbus Quadrangle, Cherokee County). The 

Fleming coal, in which these petrifications occur, lies in 

the upper part of the Cherokee shale, which belongs to 

the Desmoinesian series of the Pennsylvanian as classi81 
82 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY 

fied by the Kansas State Geological Survey (see Abernathy, 1946). The collection was made in March I950 

with Dr. H. N. Andrews, of Washington University, St. 

Louis, Mo., who kindly deposited the specimens with 

me for investigation. 

To the best of my knowledge, both Sclerocelyphus and 

Tyliosperma are unique to coal balls from this locality~ 

SYSTEMATIC DESCRIPTIONS 

Sclerocelyphus oviformus Mamay, n. gen., n. sp. 

Plate 21, figures 1-12 

General description.-A single coal ball (WCB 71I-B) 

provided all the Sclerocelyphus material on hand. A 

preliminary saw cut exposed a group of several intimately associated sporangia; these were investigated by 

means of a closely spaced series of IIO Parlodion peel 

sections. 

During the preparation of the peel series, about 60 

essentially complete sporangia were exposed. These 

all lie fairly close to each other (within an area of not 

more than about 2 em in diameter in any single section), 

and many are appressed to each other with a consequential flattening of contiguous sporangia! wall surfaces. This feature is shown in plate 21, figure I2. 

The orientation of the group of sporangia shown in 

plate 21, figure 10 is of particular interest. These 

sporangia all lie with their axes oriented in nearly the 

same direction; this suggests that they were preserved 

in essentially natural positions without first having 

become detached from the supporting appendages of the 

plant that bore them. Had they been shed prior to 

fossilization, there is little likelihood that they would 

have fallen into as uniformly oriented a group as this. 

Isolated sporangia that were not subjected to 

pressure with others display an essentially spherical 

shape. The largest ones attain diameters of 2 mm. 

Aside from their relatively large proportions, the 

sporangia are characterized by the following combination of features: 

1. A complex sporangia! wall, consisting of two 

distinct tissue layers: an outer layer (a single cell thick) 

of prismatic palisade cells with heavily thickened walls 

(the palisade cells are dark brown, and to the naked eye 

they readily serve to distinguish the sporangia from the 

surrounding matrix); and an inner tissue, consisting of 

several layers of thin-walled parenchymatous cells. 

2. A "cushion" or somewhat cupshaped aggregate of 

sclerified cells in the base of the sporangium. 

3. Slender vascularized pedicels, the xylem of which 

enters into the bases of the sclerotic cushions. 

4. Spore masses consisting of hundreds of small, 

~pherical spores with trilete markings and highly 

listinctive ornamentation. 

The sporangial wall.-The sclerotic outer layer of the 

sporangia! wall shows some variation in thickness (pl. 

21, fig. 12). In places where it is appressed to that of 

an adjacent sporangium, it may not exceed 40,u in 

thickness; otherwise, it reaches a thickness of 100,u. 

As suggested by Hoskins and Cross (1943, p. 127) in 

reference to the sporangia of Bowmanites trisporangiatus, this variation in thickness may well be a growth 

phenomenon, the less thickly developed portions of the 

wall being a reflection of pressure produced during the 

early stages of development of a compact group of 

sporangia. A similar variation in wall thickness is 

present in sporangia of Mittagia seminijormis (see 

Lignier, 1913, p. 54); Lignier also suggested ontogenetic 

appression, writing: 

* * * leur juxtaposition est en realite la cons~quence d'une 

croissance c~te a cate sur un m~me organe c'est-a-dire, en somme, 

qu'ils appartenaient a un m~me sore. 

In one instance the appression of two sporangia was 

apparently great enough to produce a slight interlocking 

of their outer wall cells during ontogeny. This, however, is not sufficiently consistent of occurrence to 

provide grounds for a synangial interpretation of the 

fructification. 

Apart from the slightly thinner surfaces of contact 

between walls of contiguous sporangia, there is no 

systematic thickening of the sporangia! walls that 

would suggest an annulate dehiscence mechanism. 

When viewed in transverse section, cells of the outer 

layer appear irregularly hexagonal in outline, with long 

and short transverse axes reaching 100 and 60,u, respectively (pl. 21, fig. II); the long axes are oriented 

parallel with the polar axis of the sporangium. When 

seen in longitudinal section, these cells present a palisadelike appearance. They are heavily thickened on 

their outer and radial walls, the innermost walls being 

very thin in contrast. The lacunae are small in proportion to the outside dimension of the cells; these are 

shown transversely in plate 21, figure 11 and longitudinally in plate 21, figure 3 (the latter photograph shows a 

group of cells with exceptionally large lacunae). The 

possibility of a functional annulate interpretation of 

these heavily indurated cells meets with two outstanding difficulties: (1) Here the induration is centrifugal with the thinnest walls facing the sporangia! 

cavity (indeed, in many instances the lacunae of the 

wall cells appear to open directly into the sporangia! 

cavity), a condition directly opposite to that found in 

the annuli of modern ferns with their centripetally 

thickened cells; (2) here the indurated cells are not 

restricted to a clearly circumscribed portion of the 

sporangia! wall as in modern fern annuli. 

The sclerified cells of the outer sporangia! layer end 

abruptly at the point of attachment of the sporangia! 
SCHLEROCELYPHUS OVIFORMUS MAMAY, N. GEN., N. SP. 83 

pedicel without any appreciable decrease in size (pl. 21 

fig. 4). This results in a circular aperture approximately 0.5 mm in diameter. 

A lining of very delicate, thin-walled parenchymatous 

cells lies immediately beneath the outer layer of the 

sporangia! wall. This lining reaches a thickness of 

about 100JL and is composed of three or four layers of 

flattened, platelike cells whose greatest dimensions lie 

parallel to the sporangia! surface (pl. 21, fig. 2). In the 

majority of the sporangia the inner tissue is poorly preserved or entirely lacking; its remains are usually 

indicated by a thin membrane surrounding the spore 

mass (pl. 21, fig. 4). The best instances of preservation of this tissue are invariably found in the basal 

portions of the sporangia. There is no evidence of 

vascularization in this tissue. 

The sclerotic cushion.-In a median longitudinal section of the sporangium (pl. 21, figs. 1, 4, 5), the sclerotic 

cushion appears as a conspicuously dark and wedgelike 

or saucer-shaped structure. Its basal portion is normally situated in a position essentially flush with the 

aperture in the sclerotic outer layer of the sporangia] 

wall. It does not, however, completely fill this opening 

nor is it in direct connection with the latter tissue. It 

flares out distally and terminates in a concave distal 

surface that forms a direct continuity with the inner 

surface of the parenchymatous layer of the sporangia! 

wall (pl. 21, fig. 1). It may reach 0. 7 mm in diameter. 

In some instances the sclerotic cushion has been displaced distally beyond the inner limits of the sporangia! 

wall; this is most likely a positional distortion effected 

following the disorganization of the inner parenchymatous lining of the sporangium. 

Individual cells of the sclerotic cushion are fusiform; 

in the center of the cushion, their long axes are directed 

parallel to the polar axis of the sporangium; but proceeding toward the margin of the cushion, they come to 

lie parallel with the surface of the sporangium. The 

cells average 80JL in length and 35JL in width. The 

walls are heavily thickened. Cells of the cushion differ 

markedly from those of the parenchyma that separates 

it from the sclerotic outer wall of the sporangium. 

Vascular elements from the pedicel reach the proximal 

surface of the cushion but apparently do not penetrate 

into it. 

The presence of this unique cushion (in sporangia! 

fructifications) is shared only by Mittagia seminiformis. 

The pedicel.-In only a few instances has the pedicel 

been observed. It is an extremely slender axis not 

exceeding 150JL in diameter (pl. 21, fig. 5). It contains 

a small vascular bundle composed of tiny scalariform 

tracheids that extend as far as the base of the sclerotic 

cushion. The extrastelar parenchyma of the pedicel is 

in organic connection with the soft inner tissue of the 

sporangia! wall; whether or not the epidermis of the 

pedicel forms a direct continuity with the outer layer of 

the sporangia! wall has not been determined. In the 

majority of cases, the sporangia are borne singly upon 

the pedicels; but in one instance, a bifurcated pedicel 

was seen with two very short termini, each bearing-a 

sporangium at its tip. 

In no instance has the pedicel been followed for more 

than 1 or 2 mm from the point of sporangia! attachment. 

Numerous small axes with diarch steles are intimately 

associated with the fructifications. These are somewhat stouter than the pedicels and may have been the 

axes that produced the latter. Such an association is 

not demonstrable, however, nor is it possible to determine the affinities of these axes beyond the assumption 

that they are apparently not of stigmarian origin. 

Dehiscence.-Little evidence is present concerning 

dehiscence of these peculiar sporangia. There is no 

differential development of the sporangia! wall which 

might properly be interpreted as an annulate dehiscence 

mechanism comparable to those of modern ferns. 

In one instance (pl. 21, fig. 10) a sporangia! wall is 

fractured at a point near the base, and the edges are 

curled inward toward the sporangia! cavity. This is 

as might be expected and may represent the normal 

type of dehiscence if a mechanical function may be 

ascribed to the centrifugal thickening of the outer cells 

of the sporangia! wall. If so, the cumulative action of 

all these cells might have forced a fracture and incurling 

of the sporangia! wall, the line of dehiscence depending 

upon loci of weakness in each individual sporangium. 

An alternative speculation is suggested by several 

fragments of sporangia, varying in size, scattered 

through the matrix. The sclerotic outer wall of the 

sporangium probably became extremely brittle upon 

drying and \Yas readily broken into irregular fragments 

upon agitation by the wind and friction with adjacent 

sporangia. 

Spores.-The majority of the sporangia! cavities are 

uniformly filled with essentially spherical spores that 

vary from 45JL to 55JL in diameter. In some instances 

the spores are slightly flattened on their proximal surfaces, the three facets representing surfaces of contact 

with sister spores of the same tetrad; no intact tetrads 

have been observed, however. 

The spore exines are thick (reaching 5JL) and are 

covered with coarse reticulations of .ridges that reach 

3JL in height and 3JL in thickness (pl. 21, figs. 7, 9). 

The reticulations form a pattern of irregular polygons 

with essentially straight sides. They are usually most 

strongly developed on the distal surfaces of the spores, 

but in some instances they extend as far as the triradiate ridges. More usually, however, the reticula-
84 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY 

tions are weakly developed or entirely lacking over the 

greater part of the proximal surface of the spore. 

'rhe triradiate ridges are usually faint, with little 

relief; their rays measure from 20JL to 25JL in length 

(pl. 21, figs. 6, 8). 

Many of the spores contains an inner body, usually 

a dark spherical mass with a smooth or slightly wrinkled surfac·e; this is quite separate from the spore 

exine. These bodies reach about 30JL in diameter; 

they may represent the shrunken remains of an inner 

spore membrane (pl. 21, fig. 6). 

The spores seem to agree generically with Reticulatisporites, as redefined by Schopf, Wilson and Bentall 

(1944, p. 34). 

Discussion.-In the structure of the sporangia! wall 

and size of sporangia, Sclerocelyphus contains certain 

points of similarity to sporangia of the sphenophyllaceous strobilus Bowmanites trisporangiatus Hoskins and 

Cross (1943). The Bowmanites fructification contains 

a prismatic outer layer of heavily thickened cells lined 

by a delicate parenchymatous layer very similar to the 

corresponding tissues in Sclerocelyphus. However, the 

basal sclerotic cushion is a feature not present in 

Bowmanites. Furthermore, there is absolutely no evidence that Sclerocelyphu.s was part of a strobilar unit. 

The compactly arranged groups of sporangia (pl. 21, 

figs. 10, 12) are much too intimately associated to allow 

for any intervening strobilar appendages, had such been 

present. Had the sporangia been shed from a strobilus, 

there is little likelihood that they would have fallen into 

such a compact group; nor would their thinner surfaces 

of sporangia! contact have maintained their original 

juxtaposition with the corresponding surfaces of contiguous sporangia. It is difficult to visualize the group 

of sporangia shown in plate 21, figure 10, becoming 

oriented with their axes all pointed in the same direction unless they grew in such a manner and were preserved in essentially their natural positions.· Detached 

sporangia are found abundantly in coal balls. However, it is seldom that large numbers are found in close 

association without affording some evidence regarding 

the plant organs that bore them. The orientation of 

this particular group of sporangia suggests that they 

were borne much like those of Botryopteris forensis or 

B. globosa, in large clusters terminating the smallest 

ramifications of a highly branched axis, rather than in 

a strobilar fructification. 

The only previously described fructification that compares closely with Sclerocelyphus is Mittagia seminijormis 

Lignier, which was originally described from Lower 

Silesian coal balls equivalent in age, according to Schopf 

(1941, p. 9), to the lowermost portion of the Morrow 

series of the American Pennsylvanian. A second 

occurrence of Mittagia may now be reported. A nodule 

collected at the Kilpatrick hills of Dunbartonshire, 

Scotland, from the Calciferous Sandstone series of the 

Lower Carboniferous by Professor John Walton, of 

Glasgow University, contained a single specimen, apparently referable to Mittagia. The specimen was 

kindly loaned to me by the collector. Unfortunately, 

the preservation is not sufficiently complete to afford 

additonal details to Lignier's original description of 

Mittagia. In Mittagia we find every distinguishing 

feature of Sclerocelyphus sporangia: the sclerotic outer 

wall, the delicate parenchymatous inner wall, and the 

sclerotic basal cushion or "cupule" of Lignier's terminology. Furthermore, the sizes of the two fructifications compare closely, and it would be difficult to distinguish differences of more than mere specific value if 

empty sporangia of Mittagia and Sclerocelyphus were 

placed side by side for comparison. Then the chief 

distinctions would lie in the sporangia! dimensions, 

microscopic details of the sporangia! walls, and the 

presence or absence of a dehiscence mechanism. While 

no clear-cut dehiscence mechanism is apparent in 

Sclerocelyphus, Lignier described a longitudinal line of 

dehiscence in Mittagia. It was marked by an abrupt 

diminution of the sclerotic cells on either side of the 

cleft; presumably the sporangia were two valved as a 

result of this mechanism. 

There are more significant factors, however, which 

preclude any strong likelihood of congenerity between 

Mittagia and Sclerocelyphus: 

1. Spores: Mittagia contains four megaspores, whereas Sclerocelyphus contains numerous small spores; and 

while Mittagia unquestionably represents the female 

fructification of a heterosporous plant, nothing is 

known of its microsporangiate counterpart. To infer 

a congenerity between the two fructifications would 

then necessitate the following assumptions: 

(a) That the male and female organs of the plant 

that bore Mittagia were alike in every detail except 

for the differences between megaspores and microspores. This in itself is not an unreasonable ~ssumption, in considerati.on of what is known about the 

fructifications of certain other Carboniferous heterosporous plants; but in this instance the uniqueness 

of the Mittagia fructification suggests that we may 

be dealing with a very unusual and perhaps entirely 

unknown type of plant. 

(b) That Sclerocelyphus is the male fructification 

of a heterosporous plant whose megasporangia are 

identical with Mittagia rather than an isosporangium 

or the microsporangium of a seed plant. To definitely establish it as any thing other than an isosporangium will necessitate more information than that 

presently at hand. 
TYLIOSPERMA ORBICULATUM MAMAY, N. GEN., N. SP. 85 

2. Comparative ages: Mittagia is geologically much 

older than Sclerocelyphus; the type specimens originated 

from coal beds in the Koksfloz (Namurian series), and 

its geologic range is extended to the basal part of the 

Lower Carboniferous (lowermost Calciferous Sandstone 

series) by the discovery of the Scottish specimen. 

3. Associated fossils: Although there is no need to 

discuss the indecisiveness of associations, it may be of 

some interest to point out that the floras associated with 

Mittagia and Sclerocelyphus, insofar as they are known, 

contain not a single common element. The Silesian 

nodules have been investigated by Kubart (1908) and 

Stur (1885); the flora is almost exclusively a Lyginopteris-Heterangium assemblage with several species of 

each. (See Hirmer, 1927.) Although Lyginopteris has 

been reported in American coal balls, the evidence for 

its presence is rather slender. Certainly it has not yet 

been noted in the Kansas material; Heterangium is not 

rare in American coal ball floras, but it too is yet unknown in the Kansas rna terial. 

The above factors would seem to encourage the 

proposal of Sclerocelyphus as a new genus, at least 

tentatively and pending further evidence pertinent to 

its possible relationships with Mittagia. The striking 

structural similarities between the two fructifications 

are beyond ascription to coincidence; indeed, they 

invite postulation of Sclerocelyphus as the male organ 

of a plant directly descended from that which bore 

Mittagia. Perhaps it is even the male organ of a seed 

bearing plant; for in consideration of the greater age 

of Mittagia, it seems reasonable that the time lapse 

was sufficient for the megasporangium to evolve into 

a bona fide seed. If such should prove to be the case, 

the similarity of Sclerocelyphus to Mittagia would be 

ascribable to "conservatism of male parts," and the 

generic separation of the two fructifications would be 

more completely justified. 

The generic name Sclerocelyphus is proposed with 

reference to the sclerification of the sporangia! wall and 

of the basal cushion (Greek skleros, tough; kelyphos, 

case). The specific name oviformus refers to the shape 

of the sporangia. . 

Generic Diagnosis.-Sporangia large, massive-walled, 

terminally borne on slender pedicels, probably in 

botryoid clusters; spores small and numerous. Sporangia! wall with an outer sclerotic prismatic layer a 

single cell thick and an inner parenchymatous layer 

several cells thick. Base of sporangium containing a 

sclerotic cushion; vascular tissue of the pedicel reaching 

the base of the sclerotic cushion. 

Specific diagnosis. -Sporangia ovoid to globular, with 

average diameters of 2 mm. Outer sclerous layer of 

sporangia! wall composed of irregularly hexagonal 

palisade cells averaging 100~ by 60~ in transverse 

266839-53-2 

dimensions and from 40~ to 100~ in thickness, the 

thinnest cells occurring at points of juxtaposition with 

contiguous sporangia. Inner parenchymatous layer 

of sporangia! wall averaging 100~ in thickness. 

Sclerotic cushion reaching 0. 7 mm in diameter and 

composed of short (averaging 80~ in length) fusiform 

cells. Dehiscence mechanism not known. 

Spores spherical, ranging from 45 to 55~ in diameter. 

Exines thick,. ornamen,ted with coarse reticulations of 

thick ridges; reticulations sometimes complete, sometimes absent from proximal surfaces of spores. Triradiate ridges present, the rays averaging 20~ in length. 

Spores sometimes containing dark inner bodies averaging 30~ in diameter and probably representing shrunken 

inner membranes. 

Stratigraphic occurrence.-Fleming coal of the Cherokee shale (Desmoinesian series of the Pennsylvanian as 

classified by the Kansas State Geological Survey). 

Locality.-Pittsburgh and Midway Coal Company 

strip pits, about 4 miles south of West Mineral, Kans. 

(Columbus quadrangle, Cherokee County). 

Type material.-Slides 2000 to 2014, inclusive, in the 

Washington University paleobotanical collection. The 

slides were made from peels of coal ball WCB 711-B; 

the unmounted peels of this series are entered in the 

Paleozoic plant collection in the U. S. National 

Museum. 

Tyliosperma orbiculatum Mamay, n. gen., n. sp. 

Figures 2-4 and plates 22, 23. 

Material.-Four specimens provide the basis for this 

description. Three were investigated by means of 

longitudinal or oblique longitudinal series, the fourth 

transversely. The first and most complete specimen 

found was encountered in coal ball WCB 711-B during 

the preparation of peels of the Sclerocelyphus specimens. 

It occurred in close association with the Sclerocelyphus, 

lying within a centimeter of the sporangia! cluster. 

This specimen will be referred to as specimen A; it was 

entirely consumed in the preparation of a series of 160 

oblique longitudinal peels (711-B; 100-260). The 

description is based largely on this specimen. 

Specimens B (WCB 717-F, t) and 0 (WCB 717-I, t) 

were longitudinally exposed by the original saw cuts 

of the coal ball. Since approximately one-third of 

each was lost in the saw kerf, the peel series of these 

two specimens are incomplete; they do, however, substantiate the conclusions derived from specimen A. 

Specimen D was exposed transversely, the apical 

portion lying above surface WCB 717-I, b, and the 

basal portion below the contiguous surface WCB 

717-H, t. The saw cut consumed the major portion 

of the nucellar mound and the floor of the micropylar 

chamber; these features are clearly understood, however, 
86 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY 

from the longitudinal series of specimen A. Both 

portions of specimen D were entirely consumed in the 

preparation of peel series 717-I, b 1-22 and 717-H, 

t 1-17. 

General morphology.-The seeds are small, simple, and 

radiospermic. They are nearly spherical, except for 

slight protrusions of the apical lobes and a slight flattening or beveling of the portion of the integument surrounding the chalaza. The seeds reach a maximum of 

3.7 mm in length and 3.5 mm in diameter. 

The free micropylar portion of the integument forms 

a canopy over the micropylar chamber; this canopy is 

divided into seven essentially equal lobes, which extend 

proximally to a level only slightly higher than that of 

the floor of the micropylar chamber. An undetermined 

number of vascular strands lie against the inner surface 

of the integument. 

A broadly conical mound of delicate nucellar tissue 

extends upward into the micropylar chamber. This 

is closely invested by an epidermal sheath that is continuous with the epidermis of the canopy. The epidermal sheath is open at the summit, leaving the very 

tip of the nucellar mound exposed. 

An interesting feature of the seed lies in the presence 

of a sclerified chalaza! cushion that bears a noteworthy 

resemblance to the sclerotic cushions in the bases of the 

Sclerocelyphus sporangia. The basal portion of the 

chalaza! cushion is composed largely of delicate parenchyma, supplied with vascular tissue that originates 

from the cupule; the bulk of the cushion is sclerified. 

The cupule is a deeply lobed structure that closely 

invests at least the basal half of the seed; the distal 

extent of the lobes is not certain. The seed is sessile 

on the cupule; there is a direct continuity between the 

epidermis of the integument and that of the cupule, and 

between the chalaza! cushion and the inner tissues of 

the cupule. 

The integument.-The integument contains an epidermis, a single cell thick; a palisade layer, also a single 

cell in thickness; and an inner layer, several cells thick, 

which makes up the greater bulk of the integument 

(pl. 22, fig. 4). 

The epidermis was extremely delicate; hence, it is 

absent from the outer surfaces of all specimens except 

0 and D. In these it appears as a thin black line that 

is separated from the seed by only a very slight interval, 

perhaps the thickness of a single cell. In plate 23, 

figure 1 (a transverse section through the apical lobes 

of the canopy) the epidermis may be seen extending 

over the inner surfaces of the lobes; it is absent from 

their outer surfaces, however. This condition is seen 

in all sections made through the micropylar chamber 

except specimen 0. The epidermis forms a complete 

lining for the chamber; it also forms the sheath that 

encloses the nucellar mound. The continuity of the 

epidermis from the inner surface of the integumental 

lobes to the sheath is shown in plate 23, figures 8 

and 9. 

Beneath the epidermis there is a palisade layer, in 

most parts only a single cell thick. The palisadelike 

aiTangement of these cells is most distinct in the equatorial portions of the seed but becomes less clearly 

defined as the poles are approached. These cells reach 

maximums of 85J.L in length and 55J.L in diameter. 

They are in sharp contrast with the cells of the inner 

part of the integument by virtue of their cellular contents; the palisade cells contain a brownish substance 

(probably the remains of a mucilaginous product) that 

is absent in the remainder of the integument (pl. 22, 

figs. 4, 6). The palisade layer is not entirely restricted 

to the outer surface of the seed; it extends over the 

inner surfaces and as far as the bases of the integumental lobes. Toward the tips of the lobes, the inner 

and outer palisade layers tend to lose definition as such 

and merge into a rather homogeneous, dark tissue (pl. 

23, figs. 8, 9). This coalescence is accompanied by a 

diminution in cell size. 

The inner tissue of the integument is composed of a 

very homogeneous mass of parenchyma, several cells in 

thickness. Toward the outer surface of the integument, 

they are irregular in shape and arrangement, but those 

toward the inner surface tend to become flattened in 

the plane parallel to the seed surface. Although the 

majority of the sections leave some doubt as to whether 

the inner limits of the integument are preserved, other 

sections (especially in specimens B and 0; see pl. 22, 

fig. 8) show the innermost layer of cells to present a 

perfectly smooth, undisrupted inner surface; probably 

this is the actual inner limit of the integument. The 

inner and outer layers of the integument attain a combined maximum thickness of 320J.L. 

The micropylar portion of the integument is deeply 

divided into seven discrete lobes. Details of this 

feature were afforded by only specimens A and D; there 

are seven lobes in each of these, however. 

Seen in transverse section (pl. 23, fig. 1) and toward 

their tips, the lobes appear as_ a series of rather triangular 

structures; at this point their component cells are 

fairly thick walled and comprise a nearly homogeneous 

tissue system with dark cellular contents. Proceeding 

in the chalaza! direction the ·lobes tend to assume the 

shape of thick crescents with slight concavities on 

their outer surfaces. Here the differentiation between 

inner and outer integumental layers becomes more 

evident. At this level (about halfway toward their 

bases) the lobes are about 600J.L wide and 450J.L thick, 

They reach lengths of nearly 1 mm. 

The chalaza! portion of the integument is perforated 
TYLIOSPERMA ORBICULATUM MAMAY, N. GEN., N. SP. 87 

by a circular aperture approximately 1 to 1.2 mm in 

diameter. The integumental tissue surrounding the 

aperture may gently decrease in thickness toward the 

lip of the aperture or n1ay bear a distinctly beveled 

appearance on its outer surface, as shown in plate 22, 

figure 5. This is probably the result of appression with 

the subtending cupule. 

A median transverse section of the seed shows an 

almost perfectly circular outline (pl. 23, fig. 4). Neither 

ribs nor sutures are evident. However, toward the 

chalaza the outline assumes a roughly polyhedral 

aspect. In (the single specimen sectioned transversely 

(specimen D), this outline appears to have seven or 

eight facets; an exact count is made difficult by the 

partial fragmentation and distortion of the specimen 

at this level. (See pl. 23, figs. 3 and 5.) These facets 

correspond with the beveling of the integument, and 

each probably represents the surface of contact with a 

single cupular lobe, although the cupule is not present 

in specimen D. 

Particular attention was directed toward the vascular 

system, but in neither transverse nor longitudinal 

sections were vascular elements noted within the 

integumental tissues. 

The micropylar chamber and nucellar mound.-By 

virtue of their simplicity, these features, especially the 

mound, stand in marked contrast with corresponding 

structures in other pteridospermous seeds. They are 

best seen in the obliquely longitudinal series of specimen 

A, which is partly represented in the series of diagrams 

presented in figure 2; parts of this series are also ill us~ 

trated in plate 22, figures 1, 2, 3, 5, 7, and 9. 

The micropylar chamber is a broadly conical cavity 

nearly 1 nun high and about 1.5 mm wide at the base. 

From the center of the floor of the chamber, a conical 

mound rises, which is approximately 0.5 mm broad at 

its base and 0.55 mm high; it terminates about 0.4 mm 

below the apical opening between the tips of the integumental lobes. The mound is enclosed in an epidermal sheath a single cell thick. The epidermal layer 

extends over the entire floor of the micropylar chamber 

and forms a direct continuation of the epidermal layer 

of the inner surfaces of the integumental lobes (pl. 23, 

figs. 6-9). The apex of the epidermal sheath is per-. 

forated by an opening about 0.2 mm wide (pl. 23, 

fig. 9). The closed appearance of the apex of the 

sheath as seen in figure 2, diagram 0, is due to the 

obliqueness of the section, which intercepted a slight 

constriction below the apical aperture of the sheath. 

The tissue within the epidermal sheath is an extremely 

delicate parenchyma that represents the apex of the 

nucellus. This fills the sheath nearly as far as its 

sununit, and the tip of the nucellus is exposed only a 

few microns below the aperture. In a median longitudinal section the tissues of the nucellar mound and 

the epidermal sheath are appressed and probably are 

organically coalescent. This in itself presents a novel 

feature among pteridospermous seeds; for here the floor 

of the micropylar chamber, as well as the sheath of 

the nucellar mound, is clearly of integumental origin. 

This represents the highest degree of coalescence 

known to exist between the integument and nucellus in 

Paleozoic seeds. 

It is possible that the specimens on hand were preserved in a slightly immature stage and that with increasing maturity the epidermal sheath became separated from the nucellar mound with the resultant cavity 

functioning as a pollen chamber. This is not demonstrable with the present material, however; and for 

this reason, in addition to the very simple structure of 

the nucellar mound and its enclosing sheath, I hesitate 

to apply the terms "pollen chamber" or "lagenostome" 

to any of these parts. No pollen grains have been 

observed within the micropylar chamber. 

The nucellus.-The basal portion of the nucellus 

forms a conspicuous pad or cushion opposite the chalazal 

aperture of the integument. It has the form of a thick 

saucer (pl. 22, figs. 5, 7, 8); and although it is invariably 

somewhat displaced toward the center of the seed, the 

convexity of its proximal surface corresponds exactly 

with that of the inner surface of the integument, suggesting that it was originally tightly seated in the 

chalazal aperture of the integument. The cushion 

reaches 1.2 mm in width and 0.4 mm in thickness. 

The proximal surface of the cushion consists of 

delicate parenchyma. The basalmost portion is made 

up of parenchyma and very small tracheids that are 

derived from the vascular supply of the cupule. The 

intervening tissue, which comprises the bulk of the 

nucellar cushion, consists of a mass of sclerous cells 

that compare closely in size and appearance with the 

sclerous elements in the basal cushion of Sclerocelyphus 

sporangia. (Compare pl. 21, fig. 4 with pl. 22, fig. 5.) 

Aside from the nucellar mound and the basal cushion, 

the remainder of the nucellus is incompletely preserved; 

the peel series of specimens A and 0, however, afford a 

certain measure of important additional information, 

especially with reference to the vascularization of the 

seed. In these series the margins of the nucellar cushion 

taper off into a thin layer of delicate parenchyma that 

extends upward and toward the base of the micropylar 

chamber (pl. 22, fig. 5). In large, part of this layer 

is badly shrunken, appearing only as a thin line of 

tissue remains; however, in places the organic fusion 

with the integument is quite apparent. Although the 

greater part of the nucellus that remains has shrunk 

away from the integument, there is little question that 

the two tissues were originally confluent as far as the 
88 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY 

FIGURE 2.-Tyliosperma orbiculatum Mamay: A series of semidiagrammatic, camera lucida drawings of several sections from the oblique-longitudinal series of specimen A. 

A-F, the apical structure of the seed and lobation of the cupula. Drawings made from slides as follows: A, 2034; B, 2037; C, 2038; D, 2041 (see also pl. 2, fig. 1, and pl. 3 fig. 8); E, 2044 (see also pl. 2, fig. 3, and pl. 3, fig. 7); F, 2046. 

Tissues shaded as follows: 

Fine stipplings __________________________________________ integument of seed and compact outer layer of cupula Medium stippling _______________________________________ delicate inner tissue of capule Coarse stippling _________________________________________ undifferentiated nucellar tissue 

Solid-black shading ____ --------------------------------- xylem Heavy black line over surface of integument, floor of 

apical chamber, and nucellar mound __________________ integumental epidermis 

Cross hatching ________________ ------ ________ ------------ sclerenchyma 
TYLIOSPERMA ORBICULATUM MAMAY, N. GEN., N. SP. 89 

FIGURE 2.-Continued. 

G-L, the basal structure of the nucellus and vascularization of the seed. Drawings made from slides as follows: G, 2048; H, 2049; I, 2051; J, 2053; K, 2054; L, 20115. 

Parts lettered as follows: ac _________________ apical chamber c __________________ cupula cl _________________ cupula lobe 

iL ________________ integumental lobe nc _________________ nucellar cushion nm ________________ nucellar mound m _________________ megaspore membrane s __________________ sheath ofnucellar mound 

vb _________________ vascular bundle 
90 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY 

tip of the nucellar mound in the micropylar chamber. 

Vascularization.-Since the vascular system was not 

apparent in the sections of the single specimen studied 

in transverse section, investigation of this important 

feature was limited to longitudinal sections; hence, the 

number or disposition of the vascular bundles was not 

determinable, although it is apparent that the seed was 

fairly well supplied with conducting tissue. 

The main vascular system of the cupule, a strand 

approximately 0.2 mm thick, enters through the chalaza 

into the base of the nucellar cushion (fig. 2, diagram J). 

The strand flares out toward the margins of the cushion; 

and narrow strands depart, proceeding distally almost 

as far as the base of the micropylar chamber. A few 

tracheids from the nucellar cushion are shown in plate 

23, figure 2. 

In specimen C, two strands arise on either side of the 

seed. Part of the length of one is organically attached 

along the inner surface of the integument; the other has 

been mechanically displaced from the inner surface of 

the integument. 

Specimen A also shows two strands arising in essentially the same relative positions; the origin of one of 

these from the margin of the nucellar cushion is shown 

in plate 22, figure 7. The strands are limited entirely 

to the inner surface of the integument; close serial 

sections were employed throughout, and no evidence 

of vascular tissue within the integumental layers has 

been noted. Each of the individual slender strands 

have two or three thicknesses of tiny tracheids (pl. 22, 

fig. 4) which do not exceed 15JL in diameter and display 

_closely set annular thickenings. 

If, as in Physostoma and Sphaerostoma, a single strand 

was an accompanying feature of each of the integumental lobes in Tyliosperma, then we may assume that the 

seed contained seven vascular bundles. However, 

pending the discovery of more and better preserved 

specimens, this point will remain conjectural. 

Megaspores.-Specimen B is the only longitudinally 

sectioned specimen that contains a complete megaspore 

membrane (pl. 22, fig. 8). This is seated upon the 

distal surface of the nucellar cushion. It is nearly 

spherical, measuring about 2 mm along its greater 

axis. There is a single deep fold in the apical surface, 

probably brought about by shrinkage. The membrane 

is thin, light brown, and devoid of any organic contents 

or distinguishing markings. 

The median sections of specimen A (pl. 23, fig. 8) 

contain a fragment of a thin membrane, which extends 

across the base in appression with the tissues of the 

nucellar mound. Judging from its similarity in color 

and· texture to the complete megaspore membrane, 

this thin membrane is interpreted as the remains of the 

apical portion of the megaspore. 

The distorted outline of a megaspore may also be 

seen in the transversely sectioned specimen D (pl. 23, 

figs. 4, 5). In these figures the megaspore appears to 

contain a double membrane. The intimate contact 

between the megaspore and the nucellar cushion is 

clearly shown in plate 23, figure 5. 

The cupule.-This structure is known only from 

specimen A. To more clearly illustrate the nature of 

this organ and its relation to the seed, a series of outline 

drawings is presented in figure 2. This series represents 

the more significant stages in the closely spaced series 

of oblique-longitudinal sections made of the specimen. 

The first indication of the presence of a cupule associated with the seed was noted in the level represented by 

diagram A, in which it appears as a pair of irregularly 

shaped but distinct bodies lying near the basal portion 

of the seed. In this plane, however, the specimen is 

incomplete, the greater part of one side of the integument and cupule having been broken off prior to preservation. Not until several sections lower in the series 

does the entire specimen appear. 

At a lower level (diagram E), four lobes are appressed 

to the basal part of the seed, with one extending nearly 

as far as the equatorial plane. At points of contact 

with the cupular lobes, the surface of the integument 

is slightly flattened. This level slightly precedes 

coalescence of the cupular lobes, which present rather 

irregular transverse sectional outlines, excepting their 

flattened surfaces of appression with the seed and with 

each other. They are roughly 0. 7 mm broad and 0. 9 

mm thick. From their free outer surfaces, irregular 

tentaclelike processes proceed at random. These reach 

0.7 mm in length and 0.15 mn1 in thickness; insofar 

as known, they are nonglandular (pl. 22, fig. 6). 

The tissues composing the cupule lobes were apparently very soft and delicate and fall into two distinct 

zones (pl. 22, fig. 6). The outer zone is the better 

preserved; it consists of a rather compact layer two or 

three cells deep. The component cells are thin-walled 

parenchyma. This layer varies in thickness from about 

80JL at points of contact with the seed to 150JL at the 

free surfaces of the lobes. 

The inner parenchyma of the lobes is poorly preserved. At best it can only be said that this appears 

to have been a very loosely aggregated tissue of large, 

thin-walled cells, much less compact than the outer 

tissue layer. 

A single delicate vascular strand not exceeding 100JL 

in diameter lies within each lobe, in a slightly eccentric 

position toward the seed surface (fig. 2, diagram E). 

The preservation of the vascular strands is too poor to 

allow investigation of the more intimate anatomical 

details; however, in several of the sections these are 

seen in longitudinal aspect, and the component tiny 
TYLIOSPERMA ORBICULATUM MAMAY, N. GEN., N. SP. 91 

tracheids exhibit annular or spiral thickenings. The 

tracheids average 15J.L in width. 

At a slightly lower level in the peel series the four 

lobes become laterally united (fig. 2, diagram G); here 

the rim of the nucellar cushion also appears in the 

section. The cupule appears as a single thick pad of 

tissue; since this section was cut through interlobal 

planes on either side of the seed, it does not appear to 

enclose the seed to any great extent. However, in the 

preceding diagrams (fig. 2, diagrams Hand I) additional 

lobes appear on either side of the seed and extend well 

up along the sides. The section shown in diagram I 

is of further interest in that it clearly illustrates the 

tenuous nature of the organic connection between seed 

and cupule. The nucellar cushion is sectioned in a 

nearly median longitudinal plane; extending downward 

from its parenchymatous base are the remains of a 

delicate tissue connecting it with the inner tissues of the 

cupule; the continuity between the epidermal layers of 

the cupule and integument may also be noted. The 

four vascular strands that supplied the lobes shown in 

diagram E now show a tendency to converge toward the 

center of the cupule. This is also illustrated in plate 22, 

figure 7. 

Proceeding further with the series, a fifth vascular 

strand departs to the left, entering a lobe of the cupule 

(diagram I) ; the base of the chalaza} cushion becomes 

deeper, and vascular elements are evident in it. These 

are short, relatively broad tracheids (50JL by 15-20JL) 

with delicate scalariform thickenings (pl. 23, fig. 2). 

They extend as far as the proximal surface of the 

sclerotic portion of the chalazal cushion. 

A further section (fig. 2, diagram J) shows the cupule 

increasing in depth; the tips of additional lobes appear 

above the seed, bringing the total number of lobes to 

at least seven, and perhaps eight. The vascular 

strand at the base of the chalazal cushion now enters 

the inner tissues of the cupule, where it is surrounded 

by the individual strands that supply the lobes. This, 

the central strand, now continues down into the cupule 

and diminishes in width until it is no more than 130J.L 

thick (fig. 2, diagram L). In the last section where 

preservation allows observation (fig. 2, diagram L), 

there is evidence of five minor strands surrounding the 

central one. The outline of the cupule has here increased in depth (to 3 mm) and decreased in width 

(to 2 mm) until it suggests a somewhat obliquely 

sectioned slender axis or pedicel. (See pl. 22, fig. 9.) 

Unfortunately the cupule and its enclosed seed bear 

no distinctive features that might afford a clue as to 

the identity of the plant that bore them. The vascular 

system is too poorly preserved for determination of the 

anatomical structure of the individual vascular strands; 

likewise, details of the pitting of the xylem elements, 

aside from those spiral and annular thickenings mentioned in a previous paragraph, are not known. Neither 

the exact number of cupular lobes nor the extent to 

which they enclosed the seed are known. 

In spite of these gaps in our knowledge of this organ, 

the cupule remains important in that it definitely establishes the cupulate nature of Tyliosperma, a feature that 

has been previously demonstrated in Lagenstoma, Gnetopsis, and Sphaerostoma but only surmised in connection 

with Conostoma and Physostoma. The cupulate nature 

thereby supports the proposal of a lagenostomalian 

affinity for Tyliosperma. Indeed, the cupule itself 

bears a marked resemblance to that of Lagenostoma 

lomaxi, differing only in minor details. The cupule of 

Tyliosperma is more deeply lobed, lacks glandular 

emergences, and apparently does not enclose the seed 

as completely as that of Lagenostoma. 

A reconstruction of Tyliosperma and its cupule, 

based on the series of sections of specimen A, is presented in figure 3. Although the length of the cupular 

lobes is not definitely known, I believe this to be a 

fairly accurate restoration of the general external aspect 

of Tyliosperma. The reconst-r:uction is supplemented 

by figure 4, a partially restored, diagrammatic longitudinal section of the seed and cupule. 

Comparison with other Paleozoic seeds.-The fundamental organizational features of Tyliosperma (its 

radial symmetry, the apparent cohesion between 

nucellus and integument, and the structure of the 

n1icropylar region) as well as its cupulate nature 

suggest affinity with the Lagenostomales as defined by 

Seward (1917, p. 308). It is, however, sufficiently unique 

in several details that its reference to any of the previously described seed genera included in this group 

is clearly unwarranted. 

The deep lobation of the integumental canopy itself 

affords grounds for a generic separation of Tyliosperma 

fron1 Lagenostoma Williamson, Conostoma Williamson, 

Gnetopsis Renault, and the -recently described Coronastoma Neeley (1951); all of the these genera possess unlobed integuments. _The peculiar plilmelike apical 

processes of Gnetopsis as well as the presence of several 

seeds in a single cupule lend additional weight to an 

already clearcut distinction from Tyliosperma; indeed, 

in these features Gnetopsis stands quite apart from any 

lagenostomalian entity. Conostoma, Coronostoma, and 

Lagenostoma, on the other hand, share the distinctive 

feature of a complex pollen receiving mechanism, the 

lagenostome, by virtue of which they cannot be confused with Tyliosperma, in which the lagenostome is 

lacking. 

While the remaining lagenostomalian genera, Physostoma Williamson arid Sphaerostoma Benson, resemble 

Tyli().§perma -in- their lobed integumental canopies,-they 
92 SHORTER CONTRIBUTIONS TO GENERAL GEOLOGY 

FIGURE a.-Reconstruction showing the external aspect of Tyliosperma orbiculatum 

and its cupnle. 

contain important differences from the latter genus. 

The circumscissile dehiscence of the pollen chamber of 

Sphaerostoma, the retention of an epidermal covering 

on the apex of the nucellar central column (the nucellar 

mound of Tyliosperma), and the projection of the megaspore into the nucellar apex of Physostoma are features 

whose absence in Tyliosperma provide grounds for 

generic separation. 

Further distinguishing features among these three 

genera are to be found in their chalaza! regions and 

vascular systems. The conspicuous sclerotic nucellar 

cushion of Tyliosperma has no counterpart in either 

Physpstoma or Sphaerostoma. Furthermore, while these 

two genera contain vascular systems that are well imbedded within their integuments, the vascularization of 

Tyliosperma is clearly of nucellar origin and has not been 

observed to penetrate into the tissues of the integument, 

although it does lie appressed against the inner surface 

of the latter. 

In the light of the comparisons presented, the creation of a new genus for this uniquely simple seed seems 

to be in order. The generic name Tyliosperma (Greek 

tyleion, cushion, bolster; sperma, seed) is proposed, with 

reference to the conspicuous nucellar pad or cushion, 

which constitutes one of its more outstanding distinguishing characters. The specific name orbiculatum 

refers to the shape of the seed. 

Generic diagnosis.-Seeds small, radiospermic, borne 

sessile in lobed cupules. Integument unribbed, apparently non vascularized; free apical portion forming a 

deeply lobed canopy over the micropylar chamber, 

basal portion with a circular chalaza! perforation. 

Micropylar chamber completely lined by an epidermal 

layer continuous with that of the integumental lobes. 

Nucellus fused with integument; all vascular tissue 

apparently of nucellar origin. Basal portion of nucellus 

forming a conspicuous cushion; apical portion projecting into the micropylar chamber as a simple mound, 

encased in a sheath formed by the epidermis of the 

micropylar chamber, except for an apical perforation. 

Specific diagnosis.-Seeds ovoid to spherical, reaching 

3.7 mm in length and 3.5 mm in diameter. Integument 

to 0.35 mm in thickness, composed of a simple epidermis 

one cell thick, a palisade layer one cell thick with darkbrown cellular contents, and a thick inner fleshy layer. 

Lobes of integumental canopy to 0.9 mm in length, 

seven in number. 

Nucellar cushion to 1.2 mm in width, 0.4 mm in 

thickness; cells of innermost portion of the nucellar 

cushion heavily sclerified. Base of cushion receiving 

a supply of small xylem elements from the cupular 

FIGURE 4.-Partially restored semidiagrammatic median longitudinal section of 

Tyliosperma orbfculatum Mamay. Shading as in figure 2. 
MORPHOLOGICAL AND SYSTEMATIC CONSIDERATIONS 93 

vascular system; vascular strands (number unknown) 

departing from margin of the nucellar cushion, proceeding in the apical direction and appressed against the 

inner surface of the integument. Apical mound of 

the nucellus 0.55 mm wide at the base, 0.5 mm high, 

terminating well below the micropylar opening of the 

integument. 

Cupule divided nearly as far as the base of the seed 

into seven, possibly eight thick, fleshy lobes; apical 

extent of the lobes uncertain. Lobes to 0.9 mm thick 

and 0. 7 mm wide; irregular nonglandular processes produced on free outer surfaces· of lobes. Each lobe containing a single, small, vascular strand. Extravascular 

tissues delicate, undifferentiated. Basal portion of 

cupule tapering into a thick, fleshy pedicel. 

Microspores, vegetative parts unknown. 

Stratigraphic occurrence.-Fleming coal of the Cherokee shale (Des Moines group of Pennsylvanian age). 

Locality.-Pittsburgh and Midway Coal Company 

strip pits, about 4 miles south of West Mineral, Kans. 

(Columbus quadrangle, Cherokee County). 

Type material.-The following slides in the Washington University paleobotanical collection made from 

peel sections of coal ball surfaces: WCB 711-B (specimen ,A); WCB 717-F, t (specimen B); WCB 717-I, 

t (specimen C); WCB 717-I, b (specimen D, apical 

portion) and WCB 717-H, t (specimen D, basal portion); 2015-2021 (specimen C); 2022-2031 (specimen 

D); 2032-2060 (specimen A); 2061-2062 (specimen B). 

Unmounted peels of these series are entered in the 

Paleozoic plant collections in the U. S. National 

Museum. 

MORPHOLOGICAL AND SYSTEMATIC 

CONSIDERATIONS 

In consideration of the distinctness with which 

Sclerocelyphus stands apart from all other known 

American Pennsylvanian fructifications, the problem of 

attempting to draw conclusions concerning its affinities 

may be approached only with extreme caution. In 

preceding pages certain evidence bearing against the 

probability of a strobilar origin of these sporangia has 

been presented;. thus, there is serious doubt cast upon 

the possibility that Sclerocelyphus represents the fructification of any of the lycopods, articulates, or cordaitaleans, insofar as these groups are presently 

understood. The solitary terminal manner in which 

the sporangia are borne would also seem to preclude 

their assignment to either the Marattiaceae with their 

superficial synangial fructifications or to the pteridosperms with their complex synangial microsporangiate 

fructifications. 

The maxmer in which the spora:agia are borne in itself 

suggests affinity among the coenopterids, for it is 

chiefly within this vaguely defined and poorly understood group that terminally borne sporangia are to be 

found. The massive exannulate sporangia! walls of 

Sclerocelyphus, however, discount an alliance with the 

botryopterids, whose sporangia possess a strikingly 

leptosporangiate aspect in their delicate annulate 

sporangia! walls (Mamay, 1950, p. 416-417). 

A somewhat closer comparison might be drawn with 

the fructification of Stauropteris oldhamia; however, the 

differences in detail are great, and the presence of 

vegetative remains of Stauropteris in American coal balls 

has not been convincingly illustrated. 

There are, however, coenopterid elements in the 

Kansas flora whose fructifications remain unknown; 

Etapteris, Ankyropteris, and Tubicaulis have been noted 

in these coal balls, and the possibility that one of these 

genera bore Sclerocelyphus cannot be discounted 

entirely. 

If, on the other hand, the notable structural similarities between sporangia of Mittagia and SclerocelyphuB 

may be construed as indicative of natural affinity, then 

Sclerocelyphus must be interpreted as a microsporangium rather than the fructification of a homosporous 

plant. 

As pointed out by Seward (1917, p. 63), histological 

details of the sporangia! wall of Mittagia bear a close 

resemblance to those of the integument of Lagenostoma 

lomaxi. Led by this resemblance, Lignier interpreted 

his first specimen o( Mittagia to be a Lagenostoma; the 

error of this identification was brought to light by the 

subsequent discovery of additional material that proved 

Mittagia to be a megasporangium containing four 

megaspores. 

The sporangia! wall of Sclerocelyphus bears an 

equally striking resemblance to the integument of 

Lagenostoma lomaxi; this may be demonstrated by a 

comparison of plate 21, figure 2 with Oliver and Scott's 

(1904) plate 5, figure 11. 

A second and perhaps more curious point of resemblance between Sclerocelyphus, Mittagia and certain of 

the lagenostomalian seeds lies in their basal structures. 

The sclerotic cushions of Sclerocelyphus and Mittagia 

are disturbingly similar to the nucellar cushions of both 

Lagenostoma and Tyliosperma; this comparison is 

especially notable between Sclerocelyphus and Tyliosperma. (Compare pl. 21, fig. 4.with pl. 22, fig. 5.) 

If these two points of structural similarity between a 

possible microsporangium (Sclerocelyphus), megasporangium (Mittagia), and seed (Tyliosperma) are to be 

interpreted as homologies, we are faced with two 

possibilities in morphological interpretation, the second 

one involving a radical departure from the tenets of 

accepted seed morphology: 

1. That Sclerocelyphus is the male fructification of a 
94 SHORTER CONTRIBUTIONS TO. GENERAL GEOLOGY 

pteridosperm, and Mittagia represents an archaic 

megasporangium that eventually gave rise to a bona 

fide seed. 

2. That the seed in some instances may merely 

represent a highly modified megasporangium whose 

integument has not been derived from an extrasporangia! source but represents the original megasporangia! wall. Such a conjecture has been offered 

by Andrews (1949, p. 493) in connection with the 

morphology of Nucellangium glabrum; he wrote: 

Can it be that we are dealing with a plant in which the protective function of the integument was developed by the sporangium 

wall, that is, the tissue that would normally have evolved into 

a nucellus of the more usual type? 

While it is indeed fascinating to indulge in the consideration of morphological propositions such as those 

presented, they are admittedly of a highly speculative 

nature, and a much more complete fossil record is 

needed for their satisfactory support. These propositions do not surmount the evolutionary difficulties 

entailed in producing the complicated apical structure 

of lagenostomalian seeds by modification of a single 

simple megasporangium. Nor do they compromise 

the organization&] simplicity of Sclerocelyphus with the 

complex synangial organization of the pteridosperm 

male fructifications as we know them. The question 

of natural affinities of Sclerocelyphus cannot, then, be 

satisfactorily answered. 

In Tyliosperma we have a seed entailing two principal 

points of interest: its cupulate nature and its simplicity 

of structure. The cupule adds little to our knowledge 

of Paleozoic seed cupules; it differs from that of Lagenastoma lomaxi only in minor details. It is, however, the 

first reported occurrence of this organ in American coal 

ball literature; the single specimen described here 

reveals the delicate nature of both the cupule itself 

and of the attachment between cupule and seed, thus 

explainjng the rarity of such occurrences in the fossil 

record. 

Several points in the structure of Tyliosperma 

present striking contrasts with other lagenostomalian 

seeds by virtue of their simplicity. Beginning with 

the integument, its apparently nonvascularized and 

relatively undifferentiated nature is unique. Furthermore, the deeply lobed canopy, a feature previously 

unknown among American pteridospermous seeds, 

presents an extremely simple· aspect. In the light of 

Benson's (1904) synangial theory of the evolution of 

seeds, Oliver (1909) has interpreted the canopy of 

Lagenostoma as an advanced state of fusion of originally 

free integumental lobes. If we apply this interpretation to Tyliosperma, it appears to occupy a primitive 

position among the Lagenostomales along with Physostoma and Sphaerostoma. This would necessitate the 

interpretation of the integument of Tyliosperma as 

originating from a partially sterilized synangium, rather 

than from the wall of a single megasporangium. If, on 

the other hand, we choose to interpret the similarities 

between Mittagia, Sclerocelyphus, and Tyliosperma as 

evidence in support of Andrews' conjecture, then the 

free apical lobes of Tyliosperma may be explained as 

apical proliferations of the megasporangium wall. The 

continuity of integumental tissue over the surface of 

the micropylar chamber and nucellar mound would 

seem to be in support of such an interpretation. If the 

evolutionary tendency in this case, as in so many other 

demonstrable instances in the plant kingdom, leads to 

fusion of free parts, then the freedom of the lobes of 

Tyliosperma may be interpreted as indicative of a 

relatively primitive condition, although by no means 

the archaic one. 

The apex of the nucellus in Tyliosperma deserves 

careful consideration, for herein lies an important point 

of contrast with other lagenostomalian entities. In 

the latter seeds, the pollen receiving mechanisms or 

lagenostomes are interpreted as entirely nucellar in 

origin, the coalescence of nucellus and integument 

extending only as far as the base of the micropylar 

chamber. The nucellar apex thus represents a highly 

modified structure. In Tyliosperma, on the other hand, 

the nucellar apex forms a simple, unmodified mound 

of tissue. The epidermal sheath surrounding it is apparently an integumental layer; thus, if this structure 

be interpreted as a simple pollen chamber or lagenostome, it represents one composed of a combination of 

both integumental and nucellar tissues rather than only 

a modified nucellar apex. Accordingly, this entails a 

much greater degree of nucellar-integumental coalescence than that present in the related seeds. These 

features add to the primitive aspect of Tyliosperma; 

along with the apparently nonvascularized, relatively 

undifferentiated integument and the deeply lobed 

canopy, they lend the impression that we are dealing 

with perhaps one of the most primitive pteridospe.cmous 

seeds discovered thus far. 

The affinities of Tyliosperma with the pteridosperms 

pose a perplexing problem. To attribute a lagenostomalian affinity to it does not seem unreasonable. Its 

gross organization and the nature (indeed the mere 

presence) of its cupule support such an alliance. However the simple nucellar apex, nonvascularized integument, exclusively nucellar source of vascular tissue, 

and high degree of coalescence between nucellus and 

integument stand in discord with Seward's concept of 

the Lagenostomales and suggest a necessity for expansion of that concept. 

Little more can be said regarding the affinities of 

Tyliosperma. While its simplicity is of considerable 
MORPHOLOGICAL AND SYSTEMATIC CONSIDERATIONS 95 

morphological interest and while it may possibly lead 

to significant new points in our understanding of the 

early seed plants, Tyliosperma cannot yet be linked 

with any of t)le presently known pteridospermous stem 

or foliage genera. Rather, it adds another organ genus 

to the overall picture of American coal ball floras and 

contributes to the confusion produced in dealing, as the 

paleobotanist must, with detached plant parts. 

LITERATURE CITED 

Abernathy, G. E., 1946, Strip-mined areas in the southeastern 

Kansas coal fields: Kans. State Geol. Surv. Bull. 64 (4), p. 

125-144. 

Andrews, H. N., 1949, Nucellangium, a new genus of fossil seeds 

previously assigned to Lepidocarpon: Annals Mo. Bot. Garden, 

v. 36, p. 479-504. 

Andrews, H. N., and Mamay, S. H., 1952, A brief conspectus of 

American coal ball studies: The Paleobotanist, v. 1 (Sahni 

Memorial Volume), p. 66-72. 

Baxter, R. W., 1951, Coal balls-new discoveries in plant petrifications from Kansas coal beds: Kansas Acad. Sci. Trans., 

v. 54, no. 4, p. 526-535. 

Benson, Margaret J., 1904, Telangium Scotti, a new species of 

Telangium (Calymmatotheca) showing structure: Annals of 

Botany, v. 78, p. 161-177. 

Delevoryas, T., and Morgan, J., 1952, Tubicaulis multiscalarijormis: a new American coenopterid: Am. Jour. Botany, v. 39, 

no. 3, p. 16Q-166. 

Hirmer, M., 1927, tlber Vorkommen und Verbreitung der 

Dolomitknollen und deren Flora: 1st Cong. Carboniferous 

Stratigraphy, Heerlen, p. 289-312. 

Hoskins, J. H., and Cross, A. T., 1943, Monograph of the Paleozoic cone genus Bowmanites (Sphenophyllales): Amer. Midland Naturalist, v. 30, p~ 113-163. 

Kubart, B., 1908, Pflanzenversteinungen enthaltenden Knollen 

aus dem Ostrau-Karwiner Kohlenbecken! K. Akad. Wiss. 

Wien, Math.-naturwiss. Kl., Sitzungber. band 117, p. 573-578. 

Lignier, 0., 1913, Un nouveau sporange seminiforme, Mittagia 

seminiformis, gen. et sp. nov: Soc. Linneenne Normandie 

Mem., v. 24, p. 49-67. 

Mamay, S. H., 1950, Some American Carboniferous fern fructifications: Annals Mo. Bot. Garden, v. 37, p. 409-476. 

Neeley, F. E., 1951, Small petrified seeds from the Pennsylvanian of Illinois: Bot. Gaz., v. 113, p. 165-179. 

Oliver, F. W., 1909, On Physostoma elegans, Williamson, an 

archaic type of seed from the Paleozoic rocks: Annals of 

Botany, v. 23, p. 73-116. 

Oliver, F. W., and Scott, D. H., 1904, On the structure of the 

Paleozoic seed Lagenostoma lomaxi, with a statement of the 

evidence upon which it is referred to Lyginodendron: Royal 

Soc. London, Philos. Trans., Series B, v. 197, p. 193-247. 

Schopf, J. M., 1941, Contributions to Pennsylvanian paleobotany. Mazocarpon oedipternum, sp. nov. and sigillarian 

relationships: Ill. State Geol. Surv. Rep. Inv. 75, p. 3-53. 

Schopf, J. M., Wilson, L. R., and Bentall, R., 1944, An annotated synopsis of Paleozoic fossil spores and the definition of 

generic groups: Ill. Geol. Survey Rep. Inv. 91, p. 5-73. 

Seward, A. C., 1917, Fossil Plants, v. 3, Cambridge. 

Stur, D., 1885, 'tlber die in Flotzen reiner Steinkohle Enthaltenden Stein«Rundmassen und Torf-Spharosiderite: Jahrbuch 

der Kaiserlich-Koniglichen Geol. Reichsanstalt, Wien, v. 35, 

p. 613-648. 

PLATES 21-23 
PLATE 21 

[1, lacuna; p, pedicel,; pw, parenchymatous inner layer of sporangia! wall; ac, sclerotic cushion; aw, sclerotic layer of sporangia! wall.] 

FIGURES 1-12. Sclerocelyphus ovijormus Mamay, n. gen., n. sp. 

1. Longitudinal section of a sporangium, showing the sclerotic cushion and its organic continuity with the soft 

parenchymatous inner tissue of the sporangia! wall. 

Slide 2011. 

2. Section of the sporangia! wall, showing the sclerotic outer layer and the parenchymatous inner layer. 

Slide 2002. 

3. Longitudinally sectioned portion of the sporangia! wall, showing large lacunae in the cells of the sclerotic layer. 

Lacunae open toward the spore cavity. 

Slide 2005. 

4. Median longitudinal section of a sporangium. The sclerotic cushion has been somewhat displaced toward the 

inside of the sporangium. 

Slide 2007. 

5. Longitudinally sectioned sporangium, showing its attachment to a pedicel. 

Slide 2013. 

6. Two spores, one showing the triradiate ridge and a reticulate proximal surface; the other contains a dark inner 

body, probably representing a shrunken endospore. 

Slide 2012. 

7. Spore, showing the characteristic reticulate ornamentation of the exine. The elliptical shape is atypical. 

Slide 2012. 

8. Proximal surface view of a spore. Note the triradiate ridge and the relatively smooth proximal surface. 

Slide 2012. 

9. A more typically spherical spore. 

Slide 2012. 

10. Group of nine longitudinally sectioned sporangia, all oriented in the same direction. Note the fractured and 

inwardly curled wall of the sporangium at the lower left. 

Slide 2013. 

11. Group of transversely sectioned cells of the sclerotic layer of the sporangia! wall. Note the hexagonal cell outlines and the lacunae in cells toward the center of the group. 

Slide 2014. 

12. Group of closely appressed sporangia, shown in transverse section. Note that the sporangia! walls are thinnest 

at surfaces of contact with contiguous sporangia. 

Slide 2001. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 254 PLATE 21 

s pw 

2 X 75 

9 

SCLEROCELYPHUS OVJFORMUS MAMAY, N. GEN., N. SP. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 254 PLATE 22 

1 

4 X 70 

TYLJOSPERMA ORBICULATUM MAMAY, N. GEN., N. SP. 
PLATE 22 

[c, cupule; ch, chalaza! aperture; cl, cupule lobe; i, Integument; m, megaspore; nc, nucellar cushion; Pt, vascular strand.) 

FIGURES 1-!}. Tyliosperma orbiculatum Mamay, n. gen., n. sp. 

1. Oblique-longitudinal section of specimen A, slightly tangential to the nucellar mound. The apical portion of 

this section is shown enlarged in plate 23, figure 8. Successively deeper sections in this series are shown in 

figures 2, 3, 5, 7, and 9. 

Slide 2041. 

2. A slightly deeper section than that shown in figure 1. Two lobes of the cupule are shown at the lower left. The 

apical portion of this section is shown enlarged in plate 23, figure 9. 

Slide 2040. 

3. A slightly deeper section than that shown in figure 2. Four lobes of the cupule are shown; lateral fusion of the 

lobes has begun. The apical portion of this section is shown enlarged in plate 23, figure 7. 

Slide 2044. 

4. Section of the integument. The epidermis is seen as a thin black line at the right. Note the palisade layer, the 

fleshy inner layer, and the narrow vascular strand, which has been partly torn away from its natural position 

against the inner surface of the integument. 

Slide 2019. 

5. A slightly deeper section than that shown in figure 3. Note the sclerified portion of the nucellar cushion. Remnants of the nucellus are shown extending away from the margins of the cushion. The cupule appears here as 

a thick pad of tissue, with fragments of two lobes extending upward at each side of the seed. Four small 

vascular strands are shown in the cupule. 

Slide 2050. 

ti. Single lobe of the cupule, closely appressed against the integument of the seed. Note the distinct palisade layer 

in the integument. The loosely aggregated inner tissue, the more compact outer layer, the small vascular 

strand and irregular nonglandular protuberances on the outer surface of the cupule lobe are shown. 

Slide 2042. 

7. A slightly deeper section than that shown in figure 5. That part of the nucellar cushion below the sclerified zone 

contains vascular elements. Part of a vascular strand is shown against the inner surface of the integument at 

the left. 

Slide 2051. 

8. Tangential-longitudinal section of specimen B, showing the megaspore seated on the nucellar cushion. 

Slide 2061. 

9. Tangential section taken just below the surface of the integument of specimen A. Note the irregular protuberances arising from the surface of the cupule. The vascular bundle indicated by the guide line is the main strand 

of the cupule; it is surrounded by several lateral strands, each of which supplies one lobe of the cupule. 

Slide 2056. 
PLATE 23 

1ac,apical chamber; le,!ntegumental epidermis; il,!ntegumentallobe; m,megaspore; nc,nucellar cushion; nm,nucellar mound; 1,sheath ot nucellar mound.] 

FIGUREs 1..:9. Tyliosperma orbiculatum Mamay, n. gen., n. sp. 

'1. · 

1. Transverse section through the apical chamber of specimen D, showing the seven discrete lobes of the integumental 

canopy, and the nucellar mound in the center. Slide 2029. 

2. Group of small annular tracheids from the nucellar cushion of specimen A. Slide 2051. 

3. Basal transverse section of specimen D, showing the nucellar cushion and the faceted outline of the seed. Slide 

4. 

5. 

6. 

7. 

8. 

9. 

2025. 

Nearly median transverse section of specimen D, showing the megaspore membrane. Slide 2028. 

Transverse section of specimen D, about midway between the sections shown in figures 3 and 4. Slide 2026. 

Tangential-longitudinal section of specimen A, taken near the base of the Clj.nopy. The integumental epidermis 

is clearly shown. Slide 2021. 

Enlargement of the apical portion of the section shown in plate 22, figure 3. Note thP. continuity of the integumental epidermis over the floor of the apical chamber. Slide 2044. 

Enlargement of the apical portion of the section shown in plate 22, figure 1, showing the nucellar mound with its 

epidermal sheath. A fragment of the megaspore is also shown. Slide 2041. 

Enlargement of the apical portion of the section shown in plate 22, figure 2. The aperture at the summit of the 

sheath of the nucellar _mound is shown. Slide 2040. 

0 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 254 PLATIC n 

3 

m 

m 

nc 

5 

6 X 44 

8 X 43 9 X 45 

TYLIOSPERMA ORBICULATUM MAMAY, N. GEN., N. SP.