Tinskya^ a New Genus of 

Seed-bearing Callipterid

Plants From the Permian of

\ North-Central Texas

GEOLOGICAL SURVEY PROFESSIONAL PAPER 523-E
Tinsleya, a New Genus of 

Seed-bearing Callipterid 

Plants From the Permian of 

North-Central Texas

By SERGIUS H. MAMAY

CONTRIBUTIONS TO PALEONTOLOGY

GEOLOGICAL SURVEY PROFESSIONAL PAPER 523-E

Tinsleya texana, the first known example of 

seed-bearing foliage in the family Callipterideae^ 

is described; biotic relationships and evolutionary 

implications are discussed

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1966
UNITED STATES DEPARTMENT OF THE INTERIOR 

STEWART L. UDALL, Secretary

GEOLOGICAL SURVEY 

William T. Pecora, Director

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

Washington, D.C. 20402 - Price 30 cents (paper cover)
CONTENTS

Abstract

Introduction _ _

Source of material

Age of material

Previous studies _

Description.

General

Foliar laminae

Page 

______________ El

1

2

2

2

______________ 3

_____________ 3

___________ .. 3

___________ 3

Description Continued

Biotic relationships of Tinsleya

Evolutionary significance _

References cited

Index

Page 

__________ E4

6

8

__________ 10

____---.__ 10

___-____-. 11

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________ 14

__________ 15

ILLUSTRATIONS

[Plates follow index]

PLATES 1-3. Compressions of Tinsleya texana.

FIGURE 1. Venation of Tinsleya texana. 

2. Seeds of Tinsleya texana., __

Page 

E5

7
CONTRIBUTIONS TO PALEONTOLOGY

TINSLEYA, A NEW GENUS OF SEED-BEARING CALLIPTERID PLANTS FROM THE

PERMIAN OF NORTH-CENTRAL TEXAS

By SERGIUS H. MAMAY

ABSTRACT

Tinsleya texana, n. gen. and sp., is described on the basis of 

a large suite of plant fossils from the upper part of the Lueders 

Limestone of the Wichita Group (Lower Permian) near Lake 

Kemp in Baylor County, Tex.

In general aspect, Tinsleya is similar to the genus Callipteris 

but differs in details of venation and polymorphism of the frond. 

Tinsleya is further unique in having seeds attached either apically or marginally on a few pinnules. This occurrence is the 

first known of seeds of any member of the Oallipterideae and 

establishes the group as truly pteridospermous. This type of 

fruiting habit may eventually lend important fossil support to 

the classical theory of derivation of the angiospermous carpel.

INTRODUCTION

Permian plant assemblages have been recorded from 

many geographic localities in both the Northern and 

Southern Hemispheres but principally from the southern areas constituting Gondwana land. The southern 

floras are generally very limited in numbers of species 

and have commonly been regarded as depauperate as 

the result of having existed under postglacial conditions. They are largely dominated by the ubiquitous 

genus G-lossopteris, whose problematical fructifications 

have attracted considerable interest during the past 15 

years. (See Plumstead, 1952.)

In contrast with the southern floras, the northern contemporary assemblages show considerably more taxonomic diversity and are generally in greater need of intensive investigation. Important among the northern 

Permian floras are the callipterids (Callipterideae of 

Doubinger, 1956, p. 98), a highly variable and presumably artificial complex of pteridophyllous plants that 

are stratigraphically restricted to rocks of late Paleozoic age but which are broadly distributed geographically. The species Callipteris conferta (Sternberg) 

Brongniart has attracted more interest than any other 

single entity in this group, inasmuch as it makes its first

appearance in rocks of earliest Permian age and, consequently, has been accepted as an important index fossil.

Callipteris and related taxa are generally regarded 

as members of the pteridosperm complex, but prior 

to the present paper no factual evidence has been presented to substantiate an ovuliferous reproductive 

habit. Indeed, the evidence bearing on the reproduction of Callipteris has been limited until now to Remy's 

(1953) description of Thuringia callipteroides from the 

Permian (Lower Rotliegende) coal-bearing strata 

near Crock, Thuringia, in Eastern Germany, and to Roselt's (1962) more recent discovery, also from the Lower 

Rotliegende of Thuringia, of organically connected 

male fructifications to which he applied the name Callipterianthus amhardti. Thuringia was described as a 

synangial fructification at least 28 mm long and about 

3.5 mm broad. It consists of several sporangia embedded in a solid groundmass of sterile tissue and contains winged spores about 50/i in diameter. Preservation of the material permitted investigation of epidermal features, and, on the basis of epidermal similarities 

and association, Remy suggested that Thuringia is in 

fact the male fructification of Callipteris conferta. Although Remy's interpretation seems reasonable, the organic connection between fructification and foliage remains to be demonstrated.

Roselt's material, Callipterianthus, consists of a single frond fragment bearing both sterile and fertile 

penultimate pinnules, so there is no doubt as to their 

relationship. The sterile parts are of the deeply dissected sphenopterid type, similar to Callipteris scheibei 

Gothan, C. owydata (Gb'ppert) Zeiller, or C. naumanni 

(Gutbier) Sterzel. As no epidermal comparisons were 

possible because of incomplete preservation, positive 

identification was precluded, but Roselt speculated that 

Callipteriantlius was ascribable to Callipteris nauEl
E2 CONTRIBUTIONS TO PALEONTOLOGY

manni. The penultimate fertile pinnae are somewhat 

longer than the sterile ones and have rather flexuose axes 

bearing undivided to twice-divided fertile "trusses."

o

The fructifications consist usually of five, or in some six 

or seven, terminal groups of microsporangia, 2-2.5 mm 

long, which apparently dehisced by slits along the inner 

surfaces of the sporangial walls. The microspores are 

round to oval, having dimensions of 20ju,-24ju, by 16ju.-18ju. 

The present paper is based on a large assemblage of 

fragmentary foliar compressions and impressions of a 

CalUpt&ris-like plant having extremely variable foliar 

characteristics. Most of the fragments are sterile, but 

a few specimens demonstrate the attachment of large 

fertile-appearing structures that are interpreted here 

as evidence of the female reproductive organs. These 

plants occur with a few other taxa, principally walchian 

conifers. A brief announcement of the discovery of 

this material was published in 1963 (U.S. Geol. Survey, 

1963), while the study of Tinsleya was in a very preliminary stage. The most significant specimens were 

found subsequently, during the fall field season of 1963. 

In view of its apparent dominance in this florule and its 

significant bearing on the matter of reproductive habits 

among the Callipterideae, however, this plant Tinsleya, new genus is regarded as of sufficient importance 

to warrant separate description at this time.

ACKNOWLEDGMENTS

I was assisted in the field during the 1957, 1961, and 

1963 seasons by Arthur D. Watt, U.S. Geological Survey. Our thanks are due to Mr. Fred Parkey, general 

manager of Wichita County Water Improvement District No. 2, who granted permission to collect on Lake 

Kemp grounds, where the material discussed in this 

paper was found.

SOURCE OF MATERIAL

The Tinsleya material was accumulated during four 

collecting trips in 1955, 1957, 1961, and 1963. It has 

been found at only one locality thus far and represents 

at least 75 percent of the specimens obtained from this 

particular site, where it occurs in association with a 

sparse assemblage of walchian conifers, a gigantopterid 

and a few other pteridophylls. The productive outcrop 

is approximately 6 miles north of Mabelle, in Bay lor 

County, Tex. It is reached by following U.S. Highway 183-283 north from Mabelle to a point 0.4 mile 

south of the highway bridge under which the Wichita 

River flows eastward from the spillway of Lake Kemp. 

At this point one sees a conspicuous exposure of predominantly tan sedimentary rocks in a low east-facing 

bluff about 125 yards west of the highway; this is the 

plant-bearing site. As exposed at this outcrop, the

sequence consists of approximately 20 feet of siltstone 

and sandstone, mostly thin bedded with variable fissility, ranging in color from wThite to gray, bluish gray, 

or tan and having considerable limonitic streaking. 

The section is capped by a limestone bed that is light 

gray in color, ranges from 2 to 3^2 feet in thickness, 

and contains minor amounts of sand. This bed is the 

only conspicuous limestone in the vicinity of the plant 

beds and doubtless is the unit mapped by Garrett, Lloyd, 

and Laskey (1930) as the Mabelle Limestone of Romer 

(1928). This fossiliferous site will be referred to here 

as the Lake Kemp locality (USGS paleobotany loc. 

10,000).

The exposure was originally created by a readout, and 

the old roadbed itself is still evident at the bottom of 

the bluff. Although finely comminuted plant debris is 

scattered sparsely through the entire sequence below 

the limestone unit, the best preserved plant material was 

found in a 1-foot unit of tan and bluish-gray siltstone, 

the base of which is about 5 feet above the old roadbed. 

The beds in approximately the lower half of the section 

show minor angular unconformities and discordant 

bedding planes that suggest a channel-fill type of 

deposition. Associated with and above the main plantbearing strata are a number of thin bone beds that 

contain fragmentary remains of a fresh-water fauna of 

small amphibians and paleoniscoid fishes. These beds 

and the presence of land-plant fossils indicate that the 

rock sequence at this locality represents a deltaic depositional environment in keeping with the generally 

accepted interpretations of the Permian rocks of this 

area.

AGE OF MATERIAL

The geology of Baylor County has not been mapped 

recently, or in detail. The only county geologic map 

available (Garrett, Lloyd, and Laskey, 1930) is based 

exclusively on surface exposures of a "few limestone 

units, most of which are not named. The stratigraphic 

position of the Lake Kemp strata relative to Romer's 

Mabelle Limestone, however, places the plant beds at a 

level slightly above the middle of the Lueders Formation (usage of Garrett, Lloyd, and Laskey, 1930). 

Romer's geologic map of north-central Texas (1958, 

facing p. 178) does not differentiate between the Lueders 

and the underlying Clyde Formation, but nonetheless 

it substantiates a high position in the Lueders for the 

Lake Kemp plant beds. Thus the plants occur near the 

middle of the lower Permian which, according to Kummel (1961, p. 570-571) and Dutro (1961), is equivalent 

to the Artinskian interval of the Ural section. In terms 

of the mid-European section, the Lueders Limestone 

(of Leonard? age) would fall within the limits of the 

Tholey beds (Kummel, 1961, p. 572-573).
TIN 8 LEY A, A NEW GENUS OF CALLIPTERID PLANTS E3

PREVIOUS STUDIES

A short list of fossil-plant species from a lower Lueders locality approximately 3 miles south of the Lake 

Kemp locality was published by Read (1943, p. 484), 

but inasmuch as Read's paper was primarily an analysis 

of the depositional environments of the outcrops 

involved in his study, the paleobotanical aspects were 

touched upon only cursorily. Otherwise, the plant content of the Lueders Limestone has not been treated in 

publication.

Fossil plants from other nearby exposures in Baylor 

County have, however, been more completely described. 

The Belle Plains Formation, slightly older than the 

Clyde Formation, provided the material on which 

White's (1912) important paper on Gigantopteris was 

based. The same strata yielded the first known Permian specimens of Discinites (Mamay,.1954). Farther to 

the southeast in Young County, the Moran Formation 

of Wolf camp (Sakmarian equivalent) age produced a 

novel form of ovuliferous neuropteroid foliage described by Mamay (1960) as Padgettia readi.

DESCRIPTION 

GENEBAL

Most of the Tinsleya specimens consist of detached 

incomplete penultimate pinnae. The collection, however, contains a few larger more complete specimens 

which demonstrate that Tinsleya produced large compound fronds of at least bipinnate architecture; it is 

likely that the fronds were, in fact, tripinnate. The 

largest fertile specimen (pi. 1, fig. 4) obviously represents the terminal part of a frond and consists of a 

terminal pinna and four laterals; although the two 

basal laterals at the lower left cannot be shown to be 

connected to the main rachis, their positions relative to 

the latter are strongly suggestive of an original connection. This specimen measures 19.5 cm from the tip of 

the terminal pinnules to the broken base of the rachis. 

A larger but sterile specimen is shown on plate 3; this 

specimen is nearly 24 cm long and consists of a penultimate rachis with parts of 14 pinnae attached.

Penultimate rachises are relatively stout, reaching an 

observed maximum of 1 cm in width and averaging 

about 0.5 cm in most specimens. Ultimate rachises 

depart decurrently at angles ranging from about 30° 

(pi. 1, fig. 4) to 60° (pi. 2, fig. 3); none, however, 

approach a right angle of departure from the penultimate rachis. The rachises lack any distinguishing 

features such as glands or ornamentation.

FOLIAB LAMINAE

The pinnules vary considerably in size, shape, and 

margination, as shown on plates 1-3. Observation of 

many specimens showing transitional pinnule form,

however, shows that these pinnules all represent the 

same plant and that a fairly high degree of foliar polymorphism is present in Tinsleya. Some pinnules, for 

example, show a length-width ratio of only 2:1, 

whereas others, such as those shown on plate 1, figure 

4, are much narrower and have length-width ratios of 

4:1 or 5:1. The broadest pinnules in the collection are 

12 mm broad and 24 mm long; others reach lengths of as 

much as 40 mm, and the narrowest specimens are only 

2.5 mm wide. Notwithstanding this variability, all the 

pinnules observed thus far demonstrate the relatively 

broad basal attachment to the rachis, the gracefully decurrent departure (generally at an angle of about 45°), 

and the proximally directed curvature of the lamina 

that contribute toward morphological characterization 

of the jutieri-typs of the Callipteris complex. The 

specimens shown on plate 2, figure 3, are particularly 

reminiscent of C. jutieri, as illustrated by Doubinger 

(1956,pi. 6,figs. 1,la).

Most of the pinnules are entire margined and obovate 

in outline (pi. 2, fig. 3), but marginal lobation and 

incision occur in varying degrees. All the pinnules of 

the specimens shown on plate 2, figure 3, are entire 

margined with rounded or nearly rounded tips. On 

plate 2, figure 2, the basal pinnule at the left is entire 

margined whereas the succeeding pinnule has a definite 

sinus at the median part of the tip, as though marginal 

growth had been arrested abruptly during ontogeny. 

(See also fig. IB.)

A slightly more pronounced degree of marginal lobation is shown on plate 2, figure 4, where pinnules at the 

left side of the illustrated pinna bear shallow apical 

sinuses that divide the tips of the lamina into three or 

four bluntly rounded lobes. Much deeper incisions, resulting in long, relatively narrow linear lobes that arise 

usually from the basal-proximal areas of the pinnules 

are shown on plate 1, figure 4, a part of which is shown 

enlarged on plate 1, figure 2. This trend toward 

pinnatifid margination of Tinsleya pinnules seems to 

be expressed most acutely by the specimen shown on 

plate 1, figure 5. Here several pinnules are generally 

irregularly pinnatind along the whole of their margins; 

thus some pinnules have as many as seven or eight lobes. 

This advanced degree of lobation approaches the condition characteristic of such species as G. Mbractensis 

Zeiller and G. lyratifolia Goeppert (compare pi. 1, fig. 

5 with Doubinger, 1956, figs. 9 and 10). These species, 

known principally from European deposits, have deeply 

and generally regularly pinnatifid pinnules that seem to 

separate them as a group from forms such as C. conferta, 

where the margins are entire. Doubinger (1956, p. 

100) has presented a phylogenetic scheme in which depth 

of foliar incision or lobation apparently reflects proportionate degrees of evolutionary advancement. If the
E4 CONTRIBUTIONS TO PALEONTOLOGY

lobate type is to be considered as having been derived 

from the entire-margined type, then one would expect 

to find forms in which transitional marginations are 

combined; Tinsleya might logically be construed as 

representing such a form.

A foliar feature of primary importance in distinguishing the Callipterideae from other pteridophyllous 

forms is found in the presence of subsidiary or rachial 

pinnules that arise directly from the penultimate 

rachises in the spaces intermediate between the successive pinna rachises. Although much of the Tinsleya 

material available for this study consists only of detached pinnae, the collection contains a fairly large 

number of specimens consisting of penultimate rachises 

with pinnae attached. Several of these rachises clearly 

show the attachment of rachial pinnules, and the 

callipteroid nature of Tinsleya is thus established.

Eachial pinnules are illustrated on plate 1, figure 1, 

and plate 2, figures 1 and 3. The specimen shown on 

plate 1, figure 1, consists of a fragment of a very stout 

penultimate rachis, to which two pinnae are attached, 

one at either side; the basal part of a third pinna departs to the left side of the specimen at about the middle 

of the upper margin of the photograph. Directly below 

this pinna fragment is shown the greater part of a 

rachial pinnule, the upper (distal) edge of which is 

partly confluent with the base of the pinna rachis. 

Three additional rachial pinnules are attached to this 

specimen.

The fertile specimen shown on plate 2, figure 1, has 

several rachial pinnules attached, which are somewhat 

larger than those shown on plate 1, figure 1; these, however, are rather fragmentary and do not show spatial 

relations satisfactorily. On plate 2, figure 3, however, 

the larger of the two frond fragments (trending toward 

the upper right of the photograph) bears several rachial 

pinnules; the one most clearly shown appears near the 

basal end of the penultimate rachis and extends toward 

the top of the photograph.

Rachial pinnules are opposite to alternate, depending 

on the relative positions of insertion of the ultimate 

pinnules between which they arise. Most commonly 

they arise singly and are approximately the same size 

and shape as the pinnules of the ultimate pinnae above 

and below. Examples are present where two discrete 

rachials originate from the space between two successive 

ultimate pinna rachises, but in such places proximity 

to the apical end of a penultimate rachis is usually 

accompanied by increasing confluence of multiple 

rachials. This confluence may be so marked that the 

penultimate rachis is virtually completely winged in 

its apical region, and laminar tissue extends without 

interruption from one pinna to the next. This feature 

is well illustrated on plate 3.

The strong tendency toward formation of subsidiary 

laminar outgrowths in Tinsleya apparently extends to 

the ultimate possible level of development; this point 

is illustrated by the pinna shown on plate 1, figure 5. 

Here are seen several pinnulets, or secondary pinnules, 

arising from the ultimate pinna rachis between successive pinnules. These, of course, are to be regarded as 

the morphological homologs of the rachial pinnules discussed in the preceding paragraphs; furthermore, their 

presence implies a need for caution in interpreting the 

rank of foliar appendages among the callipterids.

VENATION

Callipterid foliar venation is typically pinnate, with 

a usually pronounced midrib giving off secondary veins 

that may or may not divide dichotomously before reaching the pinnule margin. A few accessory veins that 

arise directly from the ultimate rachis may also enter 

the base of the pinnule. These features are particularly 

characteristic of callipterid forms having large relatively undissected "alethopteroid" pinnules, such as occur in Callipteris conferta. (See Doubinger, 1956, 

fig. 6.)

Although Tinsleya resembles a typical "alethopteroid" form of Callipteris in most of its foliar features, it 

shows a marked tendency toward the Odontopteris type 

of open venation. The vascular system of Odontopteris 

pinnules characteristically lacks a pronounced midvein; 

instead, in most species it has several independent veins 

entering the lamina directly from the ultimate rachis. 

These veins usually divide dichotomously from one to 

three times before reaching the pinnule margin. In 

Odontopteris minor Brongniart there is a tendency toward formation of a midrib, but species referable to 

Odontopteris predominantly lack this character.

The ultimate venation in Tinsleya is generally not 

well preserved, but its features are shown remarkably 

well in several specimens (fig. 1), and thus there is no 

reason for hesitancy in presenting its description on 

the basis of the material on hand.

From four to eight thin veins arise from the ultimate rachis and enter the base of the pinnule. These 

veins depart from the rachis with a gentle decurrence 

equal to that of the lamina and usually are evenly 

spaced and approximately equal in strength. The 

veins extend entirely to the margins of the pinnules; 

most terminate in the rounded pinnule apices, but some 

end in either the adrachial or abrachial margins of the 

lamina.1 The veins are either simple or forked; some

1 The terms "adrachial" and "abrachial" are introduced here in a 

sense analogous to the commonly used terms "adaxial" and "abaxial" 

because of the absence here of a midrib that would separate the pinnules 

into two well-defined virtually equal linear divisions. "Adrachial," 

then, refers to the approximate linear half of the pinnule that creates 

an acute angle with the rachis, whereas "abrachial" refers to the 

opposite half.
TINSLEYA, A NEW GENUS OF CALLIPTERID PLANTS E5

C

1 cm

FIGURE 1. Variation in venation of Tinsleya pinnules, showing tendency toward unequal development of the 

veins. Drawings adapted from USNM specimens as follows: A, 42565; B, 42559b; C, 42564; D, 42563. 

Black dots on B and D represent probable remains of glandular structures.
E6 CONTRIBUTIONS TO PALEONTOLOGY

individuals divide three or four times during their extent. Thus a pinnule that receives only four or five 

veins from the rachis may produce as many as 18 or 20 

vein termini at its tip.

All the veins of a given pinnule do not necessarily 

divide at equal intervals or rates. Rather, it is more 

usual for the basalmost one or two veins not to divide 

at all; dichotomies occur mostly among the veins that 

enter the lamina toward the adrachial margin. Thus, 

even though there is a virtually equal spacing of veins 

at the pinnule bases, the concentration of veins becomes 

greater adrachially than abrachially, as the tip of the 

pinnule is approached. This feature of the venation 

is shown in figure 1.

The drawings in figure 1 also illustrate a striking 

degree of variability in the character of the ultimate 

venation. In some pinnules one or more of the veins 

are slightly more robust than the others as they enter 

the base of the lamina (fig. 1A ; upper pinnule in fig. 

ID). Other pinnules show a more pronounced differential in relative thickness of the veins (figs. IB, C], 

and in a few specimens one of the veins is approximately 

equivalent in thickness to the vascular strand of the 

ultimate rachis from which it originates (lower pinnule 

in fig. ID). The thicker veins persist as such for only 

a relatively short distance into the lamina, beyond which 

they divide into several less robust strands, which in 

turn dichotomize one to three times to provide the 

greater part of the vascularization of the pinnule. Because the thicker veins invariably enter the pinnule 

well toward the adrachial edge at its base., pinnules with 

uneven development of basal venation always have the 

greatest concentration of veins in their adrachial areas.

Unequal development of the veins has not been observed in the rachial pinnules.

REPRODUCTIVE STRUCTURES

Three specimens of Tinsleya (fig. 2; pi. 1, figs. 2,4, 6; 

pi. 2, fig. 1) are of considerable morphologic interest 

in that they bear on their pinnules large specialized 

structures of apparently reproductive nature; a fourth 

specimen (pi. 2, fig. 4) has certain characteristics that 

are suggestive of immature forms of the same 

structures.

Near the apex of one of the distal pinnae of the holotype (pi. 1, fig. 4), one of the pinnules at the right is 

seen to bear a single urn-shaped structure, which is imbedded basally in the lamina of the tip of the pinnule 

but extends out beyond it; this structure is shown enlarged on plate 1, figure 2. As a matter of convenience 

and with no morphological implications intended at 

this point, this and other similar structures to be discussed here will be referred to as the "seeds."

This particular seed is 15 mm long and 8 mm wide at 

its broadest point, about one-third of the distance away 

from its rounded base. Apically the specimen gently 

tapers, diminishing to a width of only 3 mm. Its apical 

end terminates in three lobate projections, each about 

1.0 mm long and slightly less than 0.5 mm wide at the 

tip. The overall attitude of the seed coincides with the 

graceful decurrence of the parent pinnule, which on its 

lower margin bears two long, narrow foliar lobes; one 

of the latter originates near the base of and seems to 

"subtend" the seed.

Although the preservation of the pinnule and its 

seed is insufficiently good for observation of minute details, there can be no doubt as to the original organic 

continuity between the two, or that the base of the seed 

was indeed imbedded in the tip of the lamina. Further, 

even though some of the f usinized film representing the 

original organic constituency of the plant was removed 

from the surface of the specimen when the matrix was 

split, ample evidence still remains, in the form of a 

much thicker coaly film, that the seed itself was originally thicker and bulkier than the lamina of the parent 

pinnule. There is also a distinct suggestion in the specimen that the seed received a substantially concentrated 

vascular supply from the pinna rachis. Careful examination of the pinnule, as shown on plate 1, figure 2, 

shows a dark albeit not sharply delineated fairly 

broad line of coalification extending from the pinna 

rachis, traversing the length of the pinnule, and entering the base of the seed; there it flares out to attain a 

breadth of approximately half that of the seed. Individual veins are difficult to discern in this pinnule, but 

the presence of this median area of apparently increased 

coalification is indicative of an increased concentration 

of vascular strands, as would be necessary to supply the 

additional physiological demands of a terminal seed or 

other type of propagule.

A second fertile specimen is illustrated on plate 2, 

figure 1. This specimen is noteworthy in that it shows 

evidence of attachment of at least seven seeds. This 

number of seeds borne on a relatively small frond fragment suggests a foliar dimorphism in Tinsleya, the 

fertile function being ordinarily restricted to a limited 

proportion of the foliage.

So far as can be seen, all the seeds present on this 

specimen are imbedded in the adrachial areas of the 

pinnules and protrude in the direction of the apex of 

the frond. Apical extensions are quite prominent in 

some of the seeds, particularly those at the low^er left 

and upper left of the specimen. These further demonstrate a roughly urnlike shape and a constricted neck, 

whereas the seeds at the lower left and upper right of 

the photograph show lobate apices similar to that seen 

in plate 1, figure 2. (See also fig. 2.) The largest seeds
TINS LEY A, A NEW GENUS OF CALLIPTERID PLANTS E7

FIGURE 2. Seeds of Tinsleya, showing relationship to pinnules and apical lobation of investitures, Unshaded ovoid 

areas apparently represent the originally bulky central parts of the seeds comparable to the specimen shown on 

plate 1, figure 6. Urnlike shape of the investiture is best shown in drawing in center. Specimens depicted in 

bottom and top drawings apparently bore two seeds each, but details are poorly preserved. All drawings adapted 

from parts of USNM 42558. (See pi. 2, fig. 1.)
E8 CONTRIBUTIONS TO PALEONTOLOGY

on this specimen are nearly 2.0 cm long and approximately 0.8 cm wide.

A third fertile specimen is shown twice actual size on 

plate 1, figure 6. It consists of only one detached pinnule bearing a single apical seed which, like that shown 

on plate 1, figure 2, is "subtended" by a narrow extension 

of the lamina. The ovoid seed is 8 mm long and 5 mm 

wide at its broadest point. Its apex is rather sharply 

truncated. This shape may represent the original form 

of the seed or it may be due to faulty preservation, inasmuch as careful excavation of both counterparts revealed no apical lobations such as are present in other 

seed specimens. Even though it may be an incomplete 

specimen, this seed is important in that it demonstrates 

a considerable thickness relative to that of the lamina. 

The coalified residue of the seed is nearly 1.0 mm thick 

in places, whereas the thickness of that of the supporting lamina is negligible. These relative amounts of 

coalification of parts (lamina versus seed) strengthen 

the interpretation suggested by the holotype (pi. 1, fig. 

2), that is, that the structures discussed here were originally of much more substance than the supporting foliar 

laminae and in all likelihood were of a propagative 

nature.

One other specimen is available that may be interpreted as an ovuliferous frond part; it is shown on plate 

2, figure 4. Even though the physical evidence of organic differentiation in this specimen is vaguely preserved and limited to small surface depressions and 

small patches of slightly heavier carbonaceous films, the 

positions of these features are such that they lend a 

certain amount of support to an interpretation of Tinsleya as a plant with ovulif erous fronds.

The fourth specimen consists of an incomplete ultimate pinna, 7 cm long, having parts of seven pinnules 

on the right side and an equal number on the left. Of 

the pinnules on the right, the three basal ones are incomplete and show no notable characteristics that might be 

interpreted as evidence of fruiting structures, but the 

two next higher pinnules each have in the medial areas 

of their apices a small ovoid depression, partly covered 

with a fusinized film of carbon, suggesting as in other 

specimens, the original presence of a bulky ovoid body 

embedded in the apex of the pinnules. Counting next 

from the bottom of the left side of the specimen, pinnules 2,3,4, and 6 each have in approximately the medial 

part of the pinnule apex a small dark area, elongate 

parallel to the length of the pinnule and created by a 

slightly thickened film of fusinized carbon. The consistent placement of these coalified areas and ovoid 

depressions both in relation to the pinnule apices and 

to the positions of the larger structures described on the 

three previously discussed specimens suggests that in

this specimen we may be dealing with an early ontogenetic phase of development of the seeds of Tinsleya.

The matrix from the Lake Kemp fossil locality contains no cuticularized material such as leaf epidermis 

or spore coats. Spore analyses showed samples of this 

matrix to be barren (R. A. Scott, written commun., 

1963) as are most plant-bearing sedimentary rocks 

from the Permian of north Texas. Had megaspore 

membranes been preserved and observable in the structures described as seeds in the preceding paragraphs, 

there would be little doubt as to their true morphologic 

nature. However, the absence of megaspore membranes 

in this material need not necessarily invalidate the interpretation of these large bulky bodies on the foliage 

of Tinsleya as ovuliferous structures. Certain aspects 

of these fossils that weigh heavily in favor of such an 

interpretation in spite of the incomplete preservation 

are discussed in succeeding pages.

MORPHOLOGICAL INTERPRETATIONS

An interpretation of the obviously specialized foliageborne reproductive structures of Tinsleya as evidence 

of bona fide seeds may not meet with universal acceptance among botanists particularly those who adhere 

strictly to morphological definitions because it is not 

possible to demonstrate the presence in these structures 

of either megaspore membranes or embryos. The evidence discussed here is partly circumstantial, but in 

presenting my views it should be stated at the outset 

that the entire concept of the existence of true seeds in 

the Paleozoic phanerogamous assemblages is likewise 

based partly on circumstantial evidence.

A seed is defined by plant morphologists as a "matured ovule," and consequently the presence of an embryo is a technical requisite to the classification of an 

organ as a seed. It is a well-known, broadly discussed, 

and perennially enigmatic fact, however, that until now 

not one indisputable embryo has been recognized in a 

Paleozoic "seed." This circumstance has given rise to 

the often debated concept of the "prephanerogams" 

(Emberger, 1944, p. 269), in which the existence of 

seeds in the Paleozoic is denied. Paleozoic seeds commonly contain megaspore membranes, so they at least 

fulfill the basic requirements for classification as ovules. 

Furthermore, many of the Paleozoic seeds are extremely 

well preserved as petrifactions, such as those found in 

the abundant coal balls from Carboniferous coal seams 

in Europe, Great Britain, the United States, and the 

Soviet Union. These seeds have been investigated in 

detail, and some (Physostoma, Conostoma, Lagenostoma, and PacJiytesta, as examples) are known to have 

had extremely complex and specialized anatomical devices, such as micropyles, pollen chambers, and plinths, 

in the apical regions. Such characteristics are doubt-
TINSLEYA, A NEW GENUS OF CALLIPTERID PLANTS E9

less structural adaptations for the phanerogamous process of pollination, and, but for the absence of demonstrable embryos, they eminently distinguish the organs 

as true seeds.

The literature dealing with Paleozoic seeds is extensive, and the notion that seed-bearing plants (the 

cordaiteans and pteridosperms) were conspicuous elements in the Paleozoic flora is generally accepted; it is 

reflected in all modem classifications of fossil woody 

plants except that of Emberger. Arnold (1948) thoroughly discussed the subject of Paleozoic seeds and the 

absence of visible embryos, giving particular attention 

to the times of fertilization and shedding of the seeds. 

He arrived at the well-taken conclusion (p. 463) that 

"It therefore seems unduly dogmatic and arbitrary to 

exclude from the seed category these ripened ovules that 

occur in such copious quantities in the rocks."

A brief review, based on acceptance of Arnold's philosophy regarding the phanerogamous nature of Paleozoic seeds, of the various positions and modes in which 

they were borne on the parent plant seems appropriate 

at this point, apropos of interpretation of Tinsleya and 

its fructifications.

The seed-bearing organs of the Paleozoic cordaiteans 

and early conifers were all generally conelike structures 

that need not be considered here. The pteridosperms, 

however, showed a variety of seed-bearing habits that 

appears to entail more than one evolutionary trend and 

suggests a complex set of evolutionary relationships.

Halle (1929, p. 18-22) reviewed the seed-bearing 

habits of certain Paleozoic plants in which organic connection of seeds to foliage was known. He demonstrated a striking series of modification of seed position 

in which lower Upper Carboniferous seeds were terminal 011 reduced foliar segments (Lyginopteris oldhamia 

Binney; Aneimites fertilis White), whereas some seeds 

were terminal on ordinary pinnules in the middle Upper 

Carboniferous (Neuropteris heterophylla Brongniart; 

N. obliqua Brongniart). The trend toward decreased 

heterophylly and increased proximity of seed to lamina 

was continued by the upper Upper Carboniferous 

Pecopterix pluckeneti Schlotheim, in which the seeds 

were borne marginally on nearly normal pinnules. The 

migration of seed position was culminated during the 

Permian in such plants as Alethopteris norinii Halle, 

Sphenopteris tennis Schenk, and Emplectopteris triangularis Halle, in which the seeds were attached directly 

to the surfaces of unreduced foliar segments or, as in 

Alethopteris norinii^ to the rachis.

Subsequent to Halle's important publication, other 

examples of seed-bearing foliage have been described 

from upper Paleozoic rocks. On the basis of one large 

specimen from the upper Stephanian near Decazevilfe, 

France, Doubinger and Vetter (1950) described a seedbearing odontopterid plant; they named it Odontopteris 

Bourranensis. The plant bore small seeds, 5-7 mm long 

and 2-3 mm wide, which were attached by short pedicels 

to the bases of the pinnules in "axillary" positions.

Kon'no (1950) erected the genus Shirakiopteris on 

the basis of seed-bearing foliage from the middle Permian beds of the Pen-hsi-hu coal fields of Manchuria. 

The fronds bore small cupulate seeds on the lower surfaces of slightly reduced pinnules. The cupules are 

shallow structures with pointed marginal lobes.

The upper Paleozoic rocks of the United States have 

yielded two additional examples of seed-bearing pteridospermous foliage Spermopteris Cridland and Morris (1960) and Padgettia Mamay (1960). The Spermopteris material [type species S. coriacea (Goeppert) 

Cridland and Morris] was collected from the Ireland 

Sandstone Member of the Upper Pennsylvania (Virgil) Lawrence Formation in Douglas County, Kans. 

It consists of fertile leaves that would be referable to 

the form genus Taeniopteris in the sterile condition. 

The leaves bear a row of apparently platyspermic seeds 

on each side of the midvein, the seeds being oriented 

perpendicularly to the midvein. The seeds are apically 

notched and seem to resemble isolated seeds known as 

Diceratosperma carpenteriana Andrews.

Padgettia readi Mamay was found in the material in 

the Lower Permian (Wolfcamp) Moran Formation of 

Young County, Tex. It consists of many neuropterid 

pinnules, some of which contain one to several seeds 

arranged parallel to the midribs and apparently embedded within the tissues of the pinnules; the apices of the 

seeds were truncated exactly at the pinnule margins.

In spite of the unconventional morphological aspects 

of the various seed-bearing habits briefly discussed 

above, significant evidence continues to accumulate in 

support of Halle's ideas on the evolution of the pteridospermous seed position in the late Paleozoic. The 

fact that seeds were produced directly on the foliar 

lamina has been established by recorded fossil occurrences and has lost much of its original bizarreness in 

the eyes of the paleobotanist. In view of this and 

other evidence, then, I have no hesitation in regarding 

the foliar inclusions of Tinsleya as seeds. In further 

support of this interpretation the following points are 

set forth:

1. The previously discovered evidence of fructifications 

among plants of the Callipteris alliance is distinctly indicative of pteridospennous microsporangiate structures rather than fern sporangia.

2. The structures on Tinsleya are too large for fern 

sporangia and totally unlike any organs known to 

be pteridospennous microsporangiate fructifications. Rather, they are of a size and shape comparable to any number of Paleozoic seeds.

208-079 O - 66 - 2
E10 CONTRIBUTIONS TO PALEONTOLOGY

3. No evidence has been found in the Paleozoic of 

vegetative propagules among the ferns with which 

the Tinsleya seeds might be compared.

4. One specimen of T'msleya (pi. 1, fig. 2) clearly shows 

apical lobations that resemble apical structures in 

other Paleozoic seeds. Leisman (1964) has demonstrated a fine example of apical tentaculation in 

Physostoma calcaratum, a seed from the Middle 

Pennsylvanian of Kansas. In compressed form, 

such a seed apex could bear a significant resemblance to the apex of this Tinsleya seed. It cannot 

be stated, of course, whether the urnlike, apically 

lobate investiture of the seed of Tinsleya was 

actually homologous with the integument of 

Physostoma or other seeds. One must admit the 

possibility that it is a cupulelike or indusiumlike 

covering within which the true seed was contained. 

Whatever the true morphological nature of this 

covering may have been, however, it is apparent 

that the apical aperture was sufficiently broad to 

permit an easy avenue of access for the pollen.

TAXONOMY

The generic segregation of Tinsleya is amply justified 

by its fructifications and by some of its vegetative features as well. Attention has already been directed to 

some vegetative similarities between Timleya and 

Callipteris jutieri Zeiller. However, the absence of a 

midrib in pinnules of Tinsleya, its strong tendency 

toward odontopterid venation and unequal development 

of the veins, and the polymorphism of its foliage are 

features that combine to distinguish Tinsleya from 

Callipteris jutieri. The differences between Tinsleya 

and other species of Callipteris, particularly the 

sphenopterid types such as C. strigosa Zeiller, are even 

more obvious and need no discussion here. It appears 

that the primary vegetative feature of importance 

shared by Tinsleya and members of the callipterid complex is the rachial or subsidiary pinnule, but this feature 

in itself must be regarded as subordinate to the reproductive mechanisms demonstrable on Tinsleya.

Tinsleya is therefore proposed here as a new genus 

because retention of the material within the genus 

Callipteris would necessitate expansion of the generic 

diagnosis of Callipteris to include the reproductive 

habit described here; this taxonomic retention would 

imply possession of seeds by all presently known species of Callipteris and would thus involve a completely 

unwarranted set of assumptions regarding the fruiting 

habits of a large number of species. Although meager, 

the evidence bearing on the male fructifications of the 

callipterids suggests that the callipterid complex consists of possibly several natural genera, and it is likely 

that additional discoveries of the female parts will

ultimately corroborate polygeneric intrafamilial relationships.

The genus Tinsleya is named for Mr. T. R. Tinsley 

of Stamford, Tex., who conducted me to important fossiliferous exposures. Because the genus is yet monotypic, a combined generic-specific diagnosis is given.

Class GYMNOSPERMAE

Order PTERIDOSPERMAE

Family CALLIPTERIDEAE

Genus TINSLEYA Mamay, n. gen.

Type species: Tinsleya texana Mamay, n. sp.

Figures 1, 2; plates 1-3 

COMBINED DIAGNOSIS

Fronds of generally callipteroid aspect, with laminally borne seeds.

Fronds large, bipinnate, possibly tripinnate. Penultimate rachises stout, as much as 1.0 cm wide; ultimate 

rachises opposite to alternate, decurrent at angles of 

30°-60°. Pinnae as much as 15 cm or more in length, 

with 18 or more pinnules, usually closely spaced. 

Pinnules polymorphic but mostly obovate with broad 

basal attachment, gently decurrent, opposite to alternate, mostly entire margined with rounded apices; some 

pinnules lobed, shallowly or deeply, apically or marginally. Pinnules as much as 4.0 cm long and 12 mm 

wide, with length-width ratios of 2:1-5:1. One or 

two rachial pinnules arising from penultimate rachises 

between successive pinnae; rachial pinnules relatively 

long and narrow, approximately equal in size to nearest 

normal pinnules, or short and broad, roughly triangular 

in shape; some small rachial pinnulets produced from 

ultimate rachises between pinnules; rachial pinnules 

confluent toward frond apices, producing winged 

rachises.

Venation open, mostly odontopteroid. Four to eight 

decurrent evenly spaced veins entering pinnule bases 

directly from ultimate rachises, mostly terminating at 

apical margins of pinnules; veins simple or forking two 

to four times; dichotomies occurring mostly in veins of 

adrachial parts of the pinnules. Some veins unequally 

developed, one or more being dominant; stronger veins 

entering adrachial part of pinnule and dividing several 

times; venation usually most concentrated in adrachial 

half of pinnule. Venation of rachial pinnules always 

equally developed.

Seeds urn shaped, as much as 2.0 cm long and 0.8 cm 

wide, consisting each of an oval thickened inner body 

invested by a cupuliform sheath, possibly integumentary; sheath terminating in a usually narrowed neck 

with several digitate apical projections. Seeds borne 

one or two to a fertile pinnule, deeply or only slightly
TINS LEY A, A NEW GENUS OF CALLIPTERID PLANTS Ell

imbedded in pinnules, either apically or marginally on 

adrachial side of lamina. 

Male parts unknown.

Holotype: USNM 42554.

Paratypes: USNM 42553, 42555-42562.

Stratigraphic occurrence: Upper part of Lueders Limestone, 

Wichita Group, Lower Permian (Leonard?) Series.

Geographic occurrence: Lake Kemp roadcut, Baylor County, 

Tex.; USGS paleobotany loc. 10,000.

BIOTIC RELATIONSHIPS OF TINSLEYA

A surprisingly diverse representation of several 

groups of both plants and animals is recognizable in association with Tinsleya in the Lake Kemp deposit, in 

spite of the very limited extent of the fossiliferous exposure and the generally poor quality of preservation 

of the fossils. Most of these organisms still must be 

identified below the generic level; in some, however, this 

incomplete taxonomy is due not to poor quality of 

preservation but to the presence of distinctive morphologic features that will eventually lead to establishment 

of new taxa. A provisional list of the plants associated 

with Tinsleya follows:

?Cordaitean leaf, apparently a new taxon.

Gigantopteris n. sp. A. (See Read and Mamay, 1964, pi. 19,

fig.l.)

Gom.phostrobus Mfidus Geinitz. 

G. sp.

Neuropteris spp. 

"iOdontopteris sp. 

Samaropsis spp.

Other similar seeds, generically undetermined. 

'iSaportaea sp. 

Taeniopteris sp.

Walchia piniformis (Schlotheim) Sternberg. 

W. n. spp. (possibly two). 

Walchiostrobus sp.

The composition of this florule is not unusual; all the 

previously established genera are well known in Permian assemblages from widely scattered localities in the 

Northern Hemisphere.

The fossil-plant assemblage seems to reflect an association of the walchian complex, with an undergrowth 

of pteridophylls. The walchians are conspicuously 

represented by handsome foliar remains, including two 

large slabs approximately 12 by 14 inches in dimension, 

completely covered by leafy branches derived from a 

single plant; some specimens are distinctly different 

from the piniformis type of walchians and warrant 

further study. Also representing the walchian conifers 

are several large specimens of male cones reaching 8 

inches in length and more than an inch in diameter.

The presumptive cordaitean leaf indicated in the 

preceding floral list may not really represent the cordaites inasmuch as the venation pattern is distinctly 

asymmetrical, the veins predominantly bending toward

one margin of the leaf in a manner somewhat reminiscent of an insect wing. This element is rare in the flora.

The neuropterids and generalized seed types found in 

the flora are commonplace in late Paleozoic floras. The 

gigantopterid (Gigantopteris n. sp. A), however, is of 

more than usual interest, inasmuch as it contains a complicated vascular system featuring four orders of venation. One complete frond and several smaller fragments have been found thus far, only at the Lake Kemp 

locality. The complete frond demonstrates a bifurcative mode of laminar division which, along with the 

unique venation pattern, sets this entity apart from any 

gigantopterid described thus far from America or the 

Eastern. Hemisphere.

Animal remains recognized thus far include representatives of the following groups: polychaete 

worms, pelecypods, myriapods, estherids, fishes, and 

amphibians.

The polychaetes are represented by abundant specimens of the coiled tubes of Spirorbis, mostly attached 

to foliage of Tinsleya. Pelecypods are found as rare 

molds and casts that are neither identifiable taxonomically nor significant in terms of ecologic implications 

(John Pojeta, Jr., oral commun., 1965). The faint 

impression of one myriapod specimen, probably referable to the genus Xylobius, has been noted, and poorly 

preserved estherid valves are fairly common.

Teeth and small disarticulated skeletal fragments of 

both fish and amphibian origin are found throughout 

the exposure and are particularly concentrated in two 

or three conspicuous and fairly continuous bone beds 

that reach thicknesses of as much as 6 inches. The 

vertebrate genera have not yet been identified, but according to Walter Dalquest (written commun., 1964) 

the fishes are paleoniscoid and, together with the amphibians, definitely constitute a fresh-water fauna. The 

concentrations of bone in this exposure indicate a rich 

vertebrate population, in terms of numbers of individuals, if not of species, fairly close to the site of ultimate 

deposition.

To the evidence of fresh-water vertebrates of parts of 

the fauna may be added the fact that the estherids indicate fresh to brackish aquatic environments, and the 

myriapod specimen certainly suggests proximity to 

land. Thus the overall aspect of the f aunal assemblage 

provides grounds for the assumption that deposition of 

this f ossiliferous unit occurred in a deltaic environment, 

in probably brackish but certainly not highly saline 

water. The physical evidence of channeling in the sediments adds support to this interpretation.

The spirorbids inject an extremely interesting ecological note into the plant-animal association observed 

at this locality. As previously mentioned, the coiled 

tubes of Spirorbis are found in some abundance in the
E12 CONTRIBUTIONS TO PALEONTOLOGY

Lake Kemp biota; however, only a few isolated specimens have been seen lying free in the matrix. The 

great majority of Spirorbis specimens are attached to 

Tinsleya foliage; none has been seen attached to any of 

the associated plants. A few specimens of Tinsleya 

carry such dense concentrations of Spirorbis tubes that 

the entire leaf specimens present a generally rugose appearance; this condition is well demonstrated by the 

leaf fragment shown on plate 1, figure 1. The specimen shown on plate 3, figure 1, also carries a large number of /Spirorbis tubes, some of which are particularly 

well shown on the pinnules at the right in the vicinity 

of the locality label. The substrate selectivity oil the 

part of these spirorbids is so striking that coincidence 

may safely be discounted and an explanation sought in 

terms of physiological relations between Spirorbis and 

Tinsleya.

The attachment of spirorbids to Paleozoic plants is 

observed commonly, and Barrois (1904) cited numerous 

examples. Trueman (1942) indicated the frequent occurrence of Spirorbis tubes attached to plants, marine 

shells, and nonmarine lamellibranchs in the British 

fresh-water and marine Coal Measures. Malaquin 

(1904) presented an excellent discussion of the growth, 

habitat requirements, and adaptability of Spirorbis. 

He also pointed out that fossil plants were often observed as a substrate for these worms. None of the 

preceding authors, however, indicated examples of the 

degree of selectivity shown by the Lake Kemp /Spirorbis 

in attaching itself only to leaves of Tinsleya and avoiding all associated plants.

Seward (1910, p. 102-104) described and figured a 

fascinating example of the Spirorbis-fossil plant association that may contain an explanation of the 

Spirorbis-Tinsleya relationship. He discussed a Carboniferous specimen of Lepidodendron in which Spirorbis tubes are attached to the parichnos scars of the leaf 

cushions. Seward's more pertinent remarks (p. 103) 

follow:

The fact 'that wiith one exception all the Spirorbis shells 

on the specimen of Lepidodendron, of which two leaf cushions are shown in fig. 146, E, occur on the large parichnos 

scars on the cheeks of the cushions, suggests the possibility 

that the escape of gases from the parichnos tissue may 

have rendered the position attractive to the Spiror'bis. It 

can hardly be ^accidental that the shells occur on the parichnos strands. This fact recalls the view held by Binney 

and accepted with favour by Darwin that Lepidodendron 

and other coal-forest trees may have lived with the lower 

parts of the stems in sea water.

The final statement in Seward's remarks clearly reflects the opinion that the Carboniferous Spirorbis was 

necessarily a marine organism and that the Lepidodendron substrate was a living plant at the time that the 

Spirorbis became attached to the parichnos scars.

According to Trueman (1942, p. 317), however, most of 

the Coal Measure spirorbids "lived under non-marine 

(probably fresh water) conditions * * *"; Haack 

(1923, p. 589-590) pointed out that S. pusillus Martin 

occurs in the Carboniferous of Belgium, northern 

France, England, and Germany in not only marine but 

also in brackish and fresh-water deposits and that one 

Triassic species, S. zimmermanni, may have been euryhaline. Malaquin (1904) presented evidence that the 

Carboniferous spirorbids were adapted to fresh as well 

as salt-water habitats. Weller (1957, p. 333), in his 

treatment of the Pennsylvania!! of Illinois, discussed 

the pelecypod Aviculopecten rectilaterarius Cox:

* * * It also occurs more sparingly in gray calcareous 

shale, associated with a few common marine species, and 

in ironstone concretions with Spirorbis which has never been 

observed as a member of a normal marine Pennisylvanian 

fauna. This worm appears to have been a fresh-water species with considerable tolerance for brackish conditions.

Ill consideration of the foregoing series of comments 

on the salinity tolerances of the Carboniferous spirorbids, the conclusion is inescapable that the Lepidodendron-Spirorbis association described by Seward 

need not have been produced in marine water, but that 

a fresh-water or brackish environment was probably a 

more logical setting.

Seward's inference that the Lepidodendron was alive 

at the time of attachment of the spirorbids is also open 

to question. The Lepidodendron may in fact have been 

alive and partly submerged in fresh or slightly brackish 

water and gases produced by life functions and escaping through the parichnos openings may have attracted 

the spirorbid larvae, but it strikes me as equally possible 

that the Lepidodendron was dead and that the escaping 

gases that attracted the Spirorbis larvae to the parichnos were actually the products of decay. Whichever 

alternative is correct, the strikingly preferential location of the Spirorbis tubes on the parichnos scars can 

hardly be explained in terms other than attractiveness 

to the young animals of some sort of gas. In my 

opinion this thought contains the preeminently logical 

explanation of the preferential attachment of Spirorbis 

to Tinsleya in the Lake Kemp collection.

Considering the great abundance of Tinsleya in this 

plant assemblage, it is not difficult to visualize a virtually pure, dense stand of this plant, surrounding and 

extending up to the edge of a pool or sluggish watercourse, with the less common associated plants growing 

farther away from the water on slightly higher ground. 

Some of the fronds of Tinsleya may have dangled into 

the water while still alive and photosyntliesizing; perhaps the oxygen emitted through the stomata attracted 

the settling larvae of Spirorbis, which attached themselves to the submerged leaves. A necessary adjunct to
TINSLEYA, A NEW GENUS OF CALLIPTERID PLANTS E13

this set of conditions would have been protracted submersion of the substrate foliage for a period of time 

sufficiently long for the spirorbids to build their tubes. 

Whether or not a continuing supply of the originally 

attractive gas was necessary to the continued growth of 

the worms is a moot point, but it remains possible that, 

once attached to the Tinsleya pinnules, the worms continued to grow even though the leaves became detached 

and dropped into the water.

On the other hand, this selectivity on the part of 

Spirorbis may be hypothetically explained in terms of 

quality of gaseous decay products. Inasmuch as the 

Spirorbis-l&den foliage of Tinsleya occurs not only in 

the same beds but on the same bedding surfaces with 

parts of other, unrelated plants that show no evidence 

of attached Spirorbis tubes, it may logically be assumed 

that detached parts of the whole floral assemblage were 

washed into the site of sedimentation at approximately 

the same time; let us assume further that none of the 

plant debris carried Spirorbis tubes until this time. If 

we then recognize that decay of the plant debris began 

soon after detachment from the parent plants, the 

preferential attachment of Spirorbis may be explained 

as the result of an element of attraction to the animals 

in the gaseous decay products of Tinsleya that was 

absent from the decay products of the other plants. 

Belative amounts of cutin, starch, chlorophyll, and 

other plant substances may have produced a complex 

system of subtly differing decay products, only those 

of Tinsleya offering the peculiarly attractive ones that 

might explain the substrate selectivity of the Spirorbis 

larvae. Acceptance of this alternative explanation 

must, of course, be predicated on the assumption that 

disintegration of the Tinsleya foliage was forestalled 

for a period of time sufficient for completion of the 

worm tubes; otherwise the association of plant and 

animal could not have been preserved as fossils.

Admittedly the alternative explanations offered here 

must be oversimplifications of a complicated ecosystem 

involving conditions and rates of growth of the spirorbids, as well as conditions for and rates of decay of the 

plants. The valid explanation may never be known, 

but it is hoped that description of this peculiar example 

of selective epiphytism will draw attention to similar 

examples that will eventually shed light on the overall 

problem.

EVOLUTIONARY SIGNIFICANCE

The recorded fossil occurrences are far from sufficiently complete to permit conclusions as to the systematic positions and possible derivative forms of such 

plants with seed-bearing foliage, such as Tinsleya, 

Padgettia, and others. These plants nonetheless, are of 

sufficient interest to invite a certain amount of speculation as to their places in the succession of spermatophytes. The diverse types of Paleozoic seed-bearing 

habits strongly indicate that the interrelationships of 

these plants, if ever fully understood, will not prove to 

have been simple. It is not the intent to treat here any 

major part of the phylogenetic possibilities offered by 

that complex. Kather, I wish to discuss briefly the 

possible implications of the seed-bearing habit of 

Tinsleya, because it entails an intriguing element of 

morphological nature that eventually may contribute 

toward an understanding of the ultimate sources and 

evolutionary development of the angiospermous carpel. 

Any discussion of the morphological nature of the 

angiosperm carpel must ultimately hark back to the 

classical concept based on Goethe's (1790) interpretation of the carpel as a green leaf with marginal ovules. 

According to the classical view, the carpel formed by 

inrolling of the margins of the fertile leaf and fusion 

of the opposite margins; this process produced the hollow carpel, the ovules being contained within. Several 

other theories have been advanced and much has been 

written in attempts to explain the basic nature of the 

carpel. Eames, however, has thoroughly reviewed the 

various theories and expressed his summary opinion 

(1961, p. 218) :

* * * When evidence from all fields is considered, none of 

of the twentieth century concepts of the nature of the carpel 

can displace the classical view that the carpel is a fertile 

lateral appendage of a determinate stem tip. In details of 

position and origin on the stem, and of ontogeny and 

anatomy, it is like the leaf and is clearly of leaf rank.

Eames' conclusions are apparently derived primarily 

from evidence based 011 modern plants, for even though 

he discusses the theory of caytonialean origin of the 

carpel, he points out what he considers to be eminent 

weaknesses in that hypothesis the palmate venation 

of the modern carpel as opposed to the pinnate venation 

of Caytonia fruits and the double integument of angiosperms as opposed to the single integument of Caytonia.

Andrews (1961) also cites the possibility of caytonialean derivation of the carpel through lateral fusion 

of two "carpels" of the Pilophorospertna or Caytonia 

types, but he admits that this concept is speculative because the later stages are unknown. He also states (p. 

179) that "* * * the concept of the carpel as an enfolded leaflike structure, although long favored by many 

botanists, finds no supporting evidence in the form of 

gymnospermous plants, living or fossil, which may be 

regarded as ancestral.'1 ' It is true that fossil evidence 

of conduplication or involution of megasporophylls 

(enfolding) is still lacking, but the beginnings of such 

a line of evidence become more evident, particularly in 

the Permian rocks, as the phenomenon of the seed-bearing leaf becomes more of a reality.
E14 CONTRIBUTIONS TO PALEONTOLOGY

Determined attempts to uncover direct fossil evidence 

to support the classical theory have obviously been 

negligible, if one excludes the works of Zimmerman, 

recently summarized (1959) in a revised edition of his 

"Die Phylogenie der Pflanzen." As such evidence for 

the foliar origin of the carpel, Zimmerman (p. 510-511) 

cited the Pennsylvanian sporophyll Aorangiophyllum 

pendulatuvn Mamay, which has large monosporangia at 

the tips of the lobes of the pinnules. This plant is a 

remarkable factual facsimile of a hypothetical form proposed by Zimmerman, and even though Acrangiophyllum is not a megasporophyll but is either a microporophyll or isoporophyll, Zimmerman regards it as an 

ancestral form in the evolution of the foliar carpel. 

Substitution of the sporangia of AcrangiophyUum by 

seeds would result in a fertile foliar organ, a considerable step closer to approximation of the hypothetical 

carpel of the classical theory. (See Melville, 1964, fig. 

la). Although it retains fernlike venation and pinnulation, Tinsleya otherwise approximates closely the 

hypothetical form of primitive carpel in that it bears 

seeds on the margins of the lamina and thus, along with 

other fertile forms discussed here, suggests that continued investigations of late Paleozoic floras with emphasis on those of the Permian may ultimately lend 

undeniable support to the classical theory of carpellary 

evolution.

REFERENCES CITED 

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INDEX

[Italic numbers indicate descriptions]

Page

Acrangiophyllum- -__-_-_ ___________________ E14

pendulatum... ______________________ 14

Alethopteris norinii... _________________________ 9

AneimitesfertUis -- __________________ 9

arnhardti, Callipterianthus.--- __________ 1

Aviculopecten rectilaterarius..- __________ 12

bibractensis, Callipteris..... 

bifidus, Gomphostrobus..... 

Bourranensis, Odontopteris.

calcaratum, Physostoma...- 

Callipterianthus... -______

arnhardti........ ______

Callipterideae..............

. -- .- 3

_ __. 11

_____________ 9

_____________ 10

_____________ 1

1 

_____________ 10

Callipteris...- - ----. .... - 1,2,3,4,9,10

bibractensis.-...-- ______________ 3

conferta_______________________ 1,3,4

jutieri...........................___ _ 3,10

lyratifolia ......... _______________ 3

naumanni.--- ____._____________________ 1

oxydata.---.............. _______________ 1

scheibei. __________________________ 1

callipteraides, Thuringia.. _______________ 1

carpenteriana, Diceratosperma _______________ 9

Caytonia _______________________ 13

conferta, Callipteris..... __________________ 1,3,4

Conostoma......................... ___________ 8

coriacea, Spermopteris...---------------------- 9

Diceratosperma carpenterianaDiscinites_________

Emplectopteris triangularis _ 

fertilis, Aneimites..- _____

Page

Gigantopteris.................. __________ E3

sp. A._____ ___ 11

Glossopteris... ____._.__________.._ 1

Gomphostrobus bifidus... ______ __________ 11

Sp 11

Gymnospermae____-___________ 10

heterophylla, Neuropteris..- _____________ 9

jutieri, Callipteris.-- _______________ 3,10

Lagenostoma...... __

Lep idodendron _ _ _ _

Lyginopteris oldhamia. 

lyratifolia, Callipteris 

naumanni, Callipteris...

Neuropteris heterophylla.

spp _ __ _

norinii, Alethopteris-__

Odontopteris..- _______.

Bourranensis _ 

minor..______

sp~. 

oldhamia, Lyginopterisoxydata, Callipteris --.

12

Pachytesta.. ________________________ 8

Padgettia--.- _____________________________ 9,13

readi .-- ____________________________ 3,9

Pecopteris pluckeneti_______________ 9

pendulatum, Acrangiophyllum--.. _______ 14

Physostoma. ________________________________ 8,10

calcaratum_____-___________ 10

Pilophorosperma.------------ ________---.___ 13

Page

piniformis, Walchia. __________________________ Ell

pluckeneti, Pecopteris ________________ ___ 9

Pteridospermae____________________ 10

pusttlus, Spirorbis----------. ___ ___________ 12

readi, Padgettia______________________ 3,9

rectilaterarius, Aviculopecten... __.--__________- 12

Samaropsis spp-_------____ ___ - 11

Saportaeasp.-. - ...____ _ 11

scheibei, Callipteris- ~. _- _ __ 1

Shirakiopteris.......-.-.... ______ -- 9

Spermopteris--------- ____ __- 9

coriacea____ _____---___ ___ 9

Sphenopteristenuis.------- _________ ____ 9

Spirorbis-..... . .. - 11,12, 13

pusttlus---------------------------------- 12

zimmermanni________.___-.-____ 12

Taeniopteris.---------- _____- _ - 9

Sp__ 11

tennis, Sphenopteris_.___________-__--_-.____ 9

texana, Tinsleya.--. __________10; pis. 1-3

Thuringia-..- __________________- 1

callipteroides- ______-______ 1

Tinsleya.----__----- 2,3,4,6, 8,9, 10, 11,12,13,14

texana.--- ______________-__ 10; pis. 1-3

triangularis, Emplectopteris...------.-. 9

Walchia piniformis.. - ______ 11

Walchia spp.------------- _____ 11

Walchiostrobus sp____________ 11

Xylobius ----. ___ 

zimmermanni, Spirorbis.

E15
PLATES 1-3
PLATE 1

[All figures natural size unless otherwise indicated]

FIGURES 1-5. Tinsleya texana Mamay, n. gen., n. sp. (p. E10).

1. Specimen showing rachial pinnules and rugose appearance of lamina, 

caused by numerous epiphyllous spirorbids. Paratype, USNM 42553.

2. Enlargement of fertile pinnule of holotype, showing urn-shaped seed 

with apical lobations and long, narrow lobations of pinnule. Holotype, USNM 42554. X 2.

3. Frond fragment showing venation. Paratype, USNM 42555.

4. Large frond fragment, with fertile pinnule near top center (enlarged in 

fig. 2). Holotype, USNM 42554.

5. Part of pinna, showing much-dissected pinnules and rachial pinnulets. 

Paratype, USNM 42556.

6. Fertile pinnule, bearing ovoid seed at apex, subtended by narrow lobe 

of pinnule. Paratype, USNM 42557.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 523-E PLATE 1

^s&'iJl ^S^L^Jafci

X2

COMPRESSIONS OF TINSLEYA TEXANA
PLATE 2

[All figures natural size unless otherwise indicated]

FIGURES 1-4. Tinsleya texana Mamay, n. gen., n. sp. (p. E10).

1. Fertile specimen with several seeds attached. Paratype, USNM 42558.

2. Sterile pinna showing slight degree of lobation and shallow sinuses in 

pinnule apices. Paratype, USNM 42559a. X 2.

3. Part of large frond consisting entirely of entire-marginal pinnules and 

showing several rachial pinnules. Burrow across basal part of frond 

excavated by unknown Recent invertebrate. Paratype, USNM 42560.

X >72.

4. Part of pinna with shallow apical lobations and indications of immature 

seeds shown by thickened apical carbonaceous areas in pinnules at 

left and apical depressions in pinnules at right. Paratype, USNM 

42561. X 2.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 523-E PLATE 2

X2

COMPRESSIONS OF TINSLEYA TEXANA
PLATE 3

[Specimen figured natural size]

FIGURE 1. Tinsleya texana (p. E10).

Large sterile specimen showing irregular margination of lamina, confluence of 

rachial pinnules toward apex of frond, and attachment of numerous spirorbid tubes, particularly on right-hand parts of frond. Paratype, USNM 

42562.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 623-E PLATE 3

COMPRESSIONS OF TINSLEYA TEXANA