A LOWER PENNSYLVANIAN FLO-RA FROM
UTAH AND ETS STRATEGRAPHIC SIGNIFECAN-CE

Thesis for the 099199 of Ph. D.
MECHEGAN STATE UNWERSETY
WiEiiam D. Tidwell
1966

 

x‘ r1 6.515

 

W . .1. p
6‘..- 4.“. a—o .. . « is l
I 1‘ ' 1‘
1V111Li-tc - «633

University

 

aw ~—

This is to certify that the

thesis entitled

A Lower Pennsylvanian Flora from Utah
and its Stratigraphic Significance

presented by

William D. Tidwell

has been accepted towards fulfillment
of the requirements for

Ph . D. degree mm

Major professor

Date November 1, 1966

0-169

 

 

ABSTRACT

A LOWER PENNSYLVANIAN FLORA FROM UTAH
AND ITS STRATIGRAPHIC SIGNIFICANCE

by
William D. Tidw ell

The Manning Canyon shale on the eastern slopes of Lake
Mountain in Central Utah contains a unique, lowermost Pennsyl-
vanian flora within its upper shales. This flora contains both

Mississippian and Pennsylvanian plant species. Crossopteris

 

gen. nov. is the most abundant form with Calamites (mesocalamites)

 

also occurring with relatively high frequency.
The flora consists of thirty-six genera and sixty—eight

species. Thirteen new species and two new genera; Cross0pteris

 

and Rigbyocarpus, are described. The new species are Neurop-

 

teris ampelinos, Crosscmteris utahensis, Crossopteris undulatus,

Crossopteris mcKnightii, Alloiopteris cruciatus, SJJhenopteridium

 

Zaitzeffis, Lefldostrobus skemmatos, Cordaicarpus globosus,

 

Cordaicarpus gayshuleri, Cordaicarpus manningcanensis,

 

Cordaicarpus binutus, Cornucarpus discissus, and Rigbyocarpus

 

 

ebracteatus. Five species were recombined.

The occurrence of Sphenopteridium dissecth Sphenopteris

 

gothanica, Diplothmema arnoldi, Gnetopsis anglica, and Ting’a sp.

are reported for the first time from North America. This is also
the first record of Telanqium affine in the United States.

The Manning Canyon shale is‘ composed of shales with
quartzites, sandstones and limestones which appear to have been
deposited in an embayment with transgressive-regressive cycles
alternating between lagoonal and paludal environments. The

flora indicates a fresh or brackish swamp environment.

A LOWER PENNSYLVANIAN FLORA FROM UTAH

AND ITS STRATIGRAPHIC SIGNIFICANCE

by

William D. Tidwell

A THESIS
Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of
DOCTOR OF PHILOSOPHY
Department of Geology

1966

ACKNOWLEDGEMENTS

The writer wishes to express his appreciation to Dr. Aureal
T. Cross, Department of Geology and Botany, Michigan State
University, for his assistance and interest during this study.

He is also indebted to Dr. Jane E. Smith, Dr. James Fisher,
Dr. C. E. Prouty of the Department of Geoloqy; and Dr. R.
Mericle and Dr. G. W. Prescott of the Department of Botany,
Michigan State University for their service as Guidance Committee
members and for reading of the manuscript.

To Dr. Harold B. Stonehouse, he owes a special debt of
gratitude for his critical reading of the manuscript and other
valuable assistance.

The writer is grateful for the assistance, advice and opinions
of other paleobotanists and paleontologists. These include Dr.

S. H. Mamay and his assistant, Arthur Watts, of the U. S. Geo-
logical Survey; Dr. Helen Duncan and Dr. McKenzie Gordon Jr.
also of the Survey; Dr. Chester A. Arnold who gave much of his
valuable time and the use of his specimens; Dr. Herman Becker;
Dr. Joseph Wood of the University of Missouri; Dr. D. C.
McLaren and Dr. D. C. McGregor of the Geological Survey of
Canada; Dr. Francis Hueber at the National Museum, Dr. Kent

ii

McKnight of the Botany Department of Brigham Young University
and Dr. J. Keith Rigby of the same institution.

The author also wishes to thank James B. Zaitzeff and
Gary G. Thompson of the Department of Geology, Michigan State
University for reading of the manuscript and assistance in the
field.

He wishes to acknowledge not only the assistance of his
wife, Ann, in typing of the manuscript and drafting of the plates,
but to express a sincere thanks for her many years of encourage-

ment and companionship.

iii.

TABLE OF CONTENTS

Page
ACKNOWLEDGEMENTS ii
INTRODUCTION 1
LOCATION AND AREA 3
PREVIOUS WORK 5
GEOLOGY
Stratigraphy and Lithology 7
Structure 12
FLORAL COMPOSITION l4
COMPARISON WITH OTHER NORTH AMERICAN FLORAS
OF MISSISSIPPIAN AND PENNSYLVANIAN AGE 20

COMPARISON WITH EUROPEAN CARBONIFEROUS FLORAS 28

 

 

AGE OF THE MANNING CANYON SHALE FLORA 38
PALEOECOLOGY 46
SUMMARY 49
SYSTEMATIC PALEOBOTANY 51
Genus Lepidodendron 51
Genus Lepidostrobus 57
Genus Lepidocarpon 60

 

Genus Lepido stroboPhyllum 6 3

 

iv

TABLE OF CONTENTS Cont.

Genus Lepidcmhyllum
Genus Stigmaria

Genus Archeocalamites
Genus Calamites
Genus Asterophyllites
Genus Palaeostachya
Genus Calamostachys
Genus Ti_n_g_i__

Genus Sphenopteris
Genus Corynepteris
Genus Alloiopteris
Genus R_I_1_o_<_i_e_a_

Genus Zeilleria

Genus Sphenopteridium
Genus Adiantites
Genus Diplothmema
Genus Mariopteris

Genus Neuropteris

 

Genus Crossopte ris
Genus Odo ntopte ris

Genus Co rdaites

64
65
66
68
’78
82
83
84
86
91
94
95
100
101
106

108

119
122
131

l32

TABLE OF CONTENTS Cont.

 

 

 

 

 

 

 

 

 

 

Page
Genus Cordaianthus 134
Genus Cordaicarpus . 135
Genus Trigonocaxjgus 143
Genus Holcospermum 146
Genus Cornucarpus 147
Genus Rigbyocamus 148
Genus Lagenospermum 149
Genus Gnetopsis 151
Genus Telanqium 153
Genus Aulacotheca 155
REFERENCES 158
PLATES » 175

Table

II.

III.

IV.

TAB LES

Floral List and Relative Abundance

General Stratigraphic Nomenclature of the
Carboniferous in Europe and North America

Stratigraphic Ranges of Plant Species in the
Manning Canyon Shale

Seed Chart

Seed Chart

vii

Page

16A

31

41
143A

149A

FIGURES

Figure

1. Index map

2. Paleotectonic map

3. Stratigraphic section

4. Structural map and section

5. Bolsters of Lepidodendron aculeatum, I_._. obovatum

 

and I: Volkmannianium.

 

6. Dichotomies of Mariopteris and Diplothmema fronds;

 

Page

4A
10A
11 A

13A

55

schematic drawings of Archeocalamites, Mesocalamites

 

and Calamites.

 

69

'7. Illustrations of Cordaianthus, Rigbyocarpus, Zeilleria, '76

Calamites (Calamitina) Lepidostrobus and Lepidocarpon

8. Illustrations of Corynepteris, Sphenopteridium and
Sphenggteris
9. Illustrations of Neuropteris and Crossopteris

10. Illustration of Crossopteris utahensis

viii

96

118

128

PLATES

EXPLANATION OF PLATE I
PLATE I

EXPLANATION OF PLATE II
PLATE II

EXPLANATION OF PLATE III
PLATE III

EXPLANATION OF PLATE IV
PLATE IV

EXPLANATION OF PLATE V
PLATE V

EXPLANATION OF PLATE VI
PLATE VI

EXPLANATION OF PLATE VII
PLATE VII

EXPLANATION OF PLATE VIII
PLATE VIII

EXPLANATION OF PLATE IX
PLATE IX

EXPLANATION OF PLATE X

PLATE X

ix

Page
17 5
l7 6 ‘
177
178
17 9
180
181
182
183
184
185
186
187
188
189
190
191
192
193

194

PLATES Cont.

EXPLANATION OF PLATE XI
PLATE XI

EXPLANATION OF PLATE XII
PLATE XII

EXPLANATION OF PLATE XIII

PLATE XIII

Page
195
196
197
198
199

200

INTRODUCTION

The Manning Canyon Shale is a time-transgressive formation
in central Utah. Previous geologic investigations and studies of the
fossil fauna of this formation indicate a late Chesterian and early
Springerian age (Moyle, 1958).

The presence of plants in the Manning Canyon Shale has been
previously mentioned by Gilluly (1932), Moyle (1958), Hyatt (1956),
and Calderwood (1951). A reconnaissance of the flora and its
related stratigraphic position was attempted previously by the
author (Tidwell, 1962). The present study is a re-evaluation and
enlargement of that study with special emphasis on the systematics
of these fossil plants, and correlation of the assemblage with other
known fossil floras of this age.

The flora from the Manning Canyon Shale as described in this
study has two aspects of paramount importance.

(1) Other floras of this approximate age are rare in western
United States. Only two other Lower Pennsylvanian floras and one
Upper Mississippian flora is known from this vast region. The state
of preservation of these floras is generally too poor to allow detailed
differentiation. Consequently, the number of species from each is
small. In comparison, the flora from the Manning Canyon Shale is
well preserved and consists of a relatively diverse assemblage of

plants.

(2) The flora occurs in rocks situated stratigraphically between
strata containing fossil invertebrate fauna of Lower Pennsylvanian
age near the top of the formation and a Mississippian invertebrate
assemblage near the middle. The flora is transitional and the time
boundary between these two periods is probably located near the
plant horizon.

Type specimens of fossil plant material from the Manning Canyon
Shale used in this study will be on deposit at the U. S. National Museum,
Washington D. C. , and a representative suite of specimens are also
placed in the repository at Michigan State University, East Lansing,

Michigan.

LOCATION AND AREA

The Manning Canyon Shale is exposed in central and northern
Utah within the eastern Great Basin and the western Central Rocky
Mountain physiographic provinces. Most of the plant collections
were made from this formation in exposures on Lake Mountain and
the Traverse Mountains which are located directly west of the
Wasatch range (fig. 1). The Wasatch Mountains mark the western
boundary of the Central Rocky Mountain province. They are
separated by the undrained basin of Cedar Valley from the southern
Oquirrh Mountains to the west. The Traverse Mountains are
located to the north and northeast of Lake Mountain. They are a
small, subdued, east-west branch of the Oquirrh Mountains, and
separate Utah Valleu from Salt Lake Valley.

Lake Bonnevifle occupied the valleys adjoining Lake Mountain
during the Pleistocene, and many well developed shoreline features
of the lake remain, particularly along the eastern face of Lake
Mountain.

Manning Canyon Shales may be overlain by Bonneville sands
and gravels, or by quartzite, sandstone and limestone talus which
is derived from overlying strata. The only reasonably unweathered
exposures of the shale are in clay pits where they are quarried.

The clay pits on Lake Mountain (pl. 1, fig. 1; p1. II, fig. 2) are

approximately thirteen miles southwest of Lehi, Utah, near state
highway 68. The clay pits on Traverse Mountains (pl. 11, fig. 2)
are about nine miles west of Lehi and about two miles north of

state highway 73 (fig. 1).

 

 

II

 

 

f
A»
‘a

/

 

 

 

 

CLAY PIT A
4, SOLDIER CANYON
5 MANNING CANYON

2. CLAY PIT C
3. CLAY PIT 0

 

TEXT Fm.
INDEX MAP

 

 

 

 

 

 

 

 

MTS.
4579‘“

 
  
   

25W

 

TRAVERSE

Atty?

\ 3 sdtvmnow

PREVIOUS WORK

The Manning Canyon Shale is the name given to a certain sequence
of rocks exposed in the Manning Canyon on the western slopes of the
Oquirrh Mountains by Gilluly (1932). Various sections of the formation,
some of them essentially complete, have been measured and reported
by Nolan (1935), Bissell and Hansen (1935), Baker (1947), Bullock (1951),
Calderwood (1951, ms) and many others. Studies of the Manning Canyon
Shale include a selenium abundance by Robertson (1950, ms) and one by
Brimhall (1963); a study of the physical and chemical sedimentary
characteristics by Herbertson (1950, ms), a study of the Mississippian-
Pennsylvanian time boundary by Sadlick (1955, ms), studies of the
clays by Orneales (1953, ms) and Hyatt (1956), a paleoecological and
sedimentation study by Moyle (1958), a preliminary study of the flora
and its Stratigraphic position by Tidwell (1962), and a study of cyclothems
in the Manning Canyon Shale by Prince (1963).

Very little is known about the megascopic fossil plants from
rocks of the Mississippian and Pennsylvanian systems of the western
continental United States. Five upper paleozoic leaf floras besides
the preliminary survey of the Manning Canyon Shale flora (Tidwell,
1962) have been described with some detail from the region west of

the Great Plains. These studies include the Permian Hermit Shale

flora of northern Arizona (White, 1929); a Permian (?) flora from
Colorado (Arnold, 1941); an Early Pennsylvanian flora from the
Weber (‘2) Formation of Central Colorado (Read, 1934); a Penn-
sylvanian flora from the Spotted Ridge Formation of Central Oregon
(Read and Merriam, 1940; Arnold, 1953; Mamay and Read, 1956)

and a Mississippian flora from the Uinta Mountains of Utah reported

by Arnold (1962).

GEOLOGY

Stratigraphy and Lithology: The exposures. of the Manning Canyon
Shale at the type locality designated by Gilluly (1982) are poor (fig. 1).
Therefore, Moyle (1958) designated Soldier Canyon, located a few
miles north in the Oquirrh Range, as the "type section" (pl. II, fig. 1.).

The formation is a time-transgressive unit, containing the
Mississippian-Pennsylvanian boundary within its limits. It is predomi-
nately shales with interbedded limestones, orthoquartzite and some
siltstone (fig. 3). The shales are black, gray, brown and red,
weathering to black, purple, brown and yellow. They are fissile,
calcareous and platy in many localities. The limestones are thin-to
thick-bedded, light blue-gray to dark blue-gray, usually weathering
light gray and brown to yellow, and contain abundant fossils. The
light gray to light tan orthoquartzites show variable characters of
grain size, cementation and porosity as well as crossbedding and
ripple marks.

The lower part of the formation on Lake Mountain is essentially
shale with some limestone and quartzite, whereas the upper half is
predominately quartzitic with a few shale and limestone beds. The

following is a modified stratigraphic section after Calderwood (1951).

Oquirrh Formation.

Conformable contact.

Manning Canyon Shale.

 

Bed
No. Description Feet
9. Limestone, shaly, sen dy to silty, also calcareous sand-

stones. Limestones dark blue-gray on fresh surfaces,
weather gray-brown. Sandstones shaly and weather

reddish brown to flesh colored 95
Base of a ledge of mudstones, argillaceous calcareous

shales, sandy and silty limestones, siltstones, and
interbedded orthoquartzites. Numerous leaf impressions,
mud cracks, and other indications of shallow water-
environment. Above are limestones, dark blue-gray to
brown, sandy, rough weathering, shaly fragments and
limestone, dense black, weather pink and lavender. . . . . 173
Shale at base, to shaly limestone. Shales tan, yellow-

brown, buff and tan, some dark gray. Limestones, dark
gray, weather pale blue-gray to gray-blue. Banded ortho-
quartzites, flesh colored to reddish brown (on weathered

SlJ-rface)OOOOOOOOOOOOOOOOO.0.OOOOOOOOOOOOOOOOOOOOCOOO 139

Medial limestone of the Manning Canyon: limestone,

dark gray to medium dark gray, dense, hard, weathers

pale somber gray to gray-blue and light to light medium

gray; nonfossiliferous lower part, few brachiopods upper

part, well bedded in beds, average 18" - 24". . . . . . . . . . . . . 93
Series of arkosic (?) orthoquartzite, subgraywackes,
orthoquartzites and chloritic to micaceous shales. . . . . . . . 128
Shale, brown, red brown, purplish brown, containing.

lingulid brachiopods; soft, argillaceous clay shales. Beds
contain seams of calcite and gypsum. Some yellow, ochre,
and limonite-stained shales present; most beds in this unit

are clay shales 92
Base of ledge is brown-weathering, gray to pink ortho-
quartzites; seamed with white quartz, well bedded in

6-inch to 2-foot layers. Arkosic in part, locally cross-
bedded 94
TOp of black shales, base of dense, hard, very fine-

grained orthoquartzite, pink to tannish gray with many

white quartz seams; interbedded black shale similar to

underlyj—ngShales..000......OOOOOOOOOOOOOOOOOOOOO... 140

1. Base: shale, brown and black, fissile, and interbedded
l- to 3-foot beds of dense black siliceous limestone; shale
weathers black and very dark brown. Some siliceous
ironstone concretions present..... .. .. __l'_7_6_

Total 1, 180

Conformable contact.

Great Blue Limestone
A lithologic summary of the Lake Mountain section is as

follows: (after Moyle, 1958)

 

 

 

Rock Type Thickness in feet 26).
Shales 1231 64. 5
Quartzites, arkose and graywackes 600 31. 4
Limestones 79 4. l
1910 100. O

The formation is underlain by the ridge-forming Chesterian Great
Blue Limestone and overlain by the limestones of the Morrowian
portion of the Oquirrh Formation. The Great Blue Limestone and
Manning Canyon Shale contact on Lake Mountain is taken at the top of
a dark gray, light gray-weathering cherty limestone. The contact with
the Oquirrh formation is not readily discernible, but a point beneath
a dominant, medium gray brown-weathering quartzite was used.

Baker (1947) measured 1, 645 feet of Manning Canyon Shale in

the Wasatch Mountains. This section includes coarser sediments

 

 

\/

NORTHEAST . .
NEVADA HIGHLAN

 

 

     

 

 

 

FREMONT
ACCESSWAY

 

 

 

(After Bissell. I962)

Fig 2' Paleotectonic map of Cordilleran area during Pennsylvanian and Early Permian
time

than those characterizing the rocks at Lake Mountain which is one
indication of an eastward source of much of the mate rial. Stokes
and Cohenour (1956)report that possibly as much as 800 feet of the
section is Manning Canyon Shale in wells located at the Mounds in
southern Carbon County, Utah. They state that the resemblance of
the section at the north end of the San Rafael Swell (Fig. l) to that
of the Wasatch Mountains indicates a deeper trough which entered
from the southwest and received more sediments than the area on
the Swell to the south. Zabrisky (personal communication) stated
that only fifteen feet of the Manning Canyon shale occurs in wells
drilled north and west of Price, Utah, (fig. 1). This variation
appears to be due to a thick accumulation of coarser materials in
the Oquirrh Basin along the flanks of the emerging Emery Uplift
(fig. 2) with a concurrent thinning of the formation over the

uplift and then a thickening again in the direction of the Paradox
Basin.

The Manning Canyon Shale was probably deposited in an
embayment which was alternating lagoonal and paludal, dominately
marine to the west and nonmarine to the east and south. The
embayment was bordered by three other late Paleozoic positive

areas in addition to the Emery Uplift. These uplifts were also

 

GENERALIZED
SECTION

CLAY PIT A

Legend

QUART ZIT E

MANNING

    

Fig 3. Stratigraphic section of Manning Canyon Shale, Lake Mt.
with relcaonships of clay we: A and clay pit C.

possible sources for the sediments comprising the Manning Canyon
Shale. Sediments in the Oquirrh and lake Mountain areas may have
been derived in part from the Western Utah Highland (Bissell, 1962)
which was located along the southwestern border of the embayment
(fig. 2).

The northern part of the embayment may have received sediments
from the northeastern Nevada Highland (Bissell, 1962). The highland,
located in northeastern Nevada and northwestern Utah (fig. 2), may
have been active during the Antler Orogeny, a late Mississippian-
early Pennsylvanian disturbance. Dott (1955) reported that coarse
conglomerates and quartzites in the Tonka Formation of eastern
Nevada grade laterally eastward into shales.

The Uncompahgre uplift to the southeast may also have contributed
some sediments to the embayment.

Structure: In the area of Lake Mountain, the strata are deformed
into a major syncline and anticline with many minor folds; thrust, tear
and normal and reverse faults (fig. 4a). According to Bullock (1951),
Lake Mountain is a broad syncline with a large anticline to the north.

The Lake Mountain syncline can be traced approximately eleven
miles. It is a northwest-southeast trending, slightly assymmetrical

syncline with maximum dips on the steepest limb of nearly 550 (fig. 4b)

and a northwest plunge of about 140 (Bullock, 1951). The north end
of the syncline is in structural continuity with the Bingham Syncline
in the Oquirrh Mountains, and the two appear to be parts of the same
fold.

A broad northwest trending anticline is exposed at the northern
end of Lake Mountain and in the southern portion of the Traverse
Mountains (Bullock, 1951). Minor folds and faults complicate the

main folding.

BLUE L8.

MANNING curves In
[El cent

         

  

Is.

a

TEXT FIG. 4

40 Structural Map of

.3... t. A / .
............-..... fi/uh.ww%wfl/.////x//// //

Mountain

Lake

  

m
mm»
.m
as.
us...
18....
O
Y
an
O-
U
m.
s n.
b .Ar
4 I

1...... / x
w .1........ ,,//. 2% /////W.: . i

 

 

 

 

 

FLORAL COMPOSITION

The plant fossils of the Manning Canyon Shale consist of
impressions of stems, portions of fronds, and detached pinnules,
isolated seeds and other dissociated plant remains. Nearly all of
these fossils are compressions. Very little of the original
carbonaceous material is preserved. The remains have been
secondarily replaced with limonite and hematite, resulting in
reddish or brownish coloration of the impressions. This replacement
shows little cellular structure and cuticle of the original plants.

Minute quartzite crack-fillings, arranged in box-like structures
which are filled with limonite and hematite occur on the stem and
leaf remains. The quartz appears to have filled minute joints or
cracks which developed in the replacing materials.

The Manning Canyon Shale flora is represented by sixty-eight
species and thirty-six genera. Twenty-four species are of fern or
fern-like foliage. Ten species of lycopods, eleven species with
calamitian affinities, fifteen species of seed types, five cordaitian
species and three species related to microsporangiate structures
are included.

The synopsis of Table I reveals the most common groups to be

represented are crossopterids, calamitean forms and seed types.

These comprise the majority of the compressions. Crossopteris

utahensis. the most abundant fossil form present, accounts for at

 

least seventy percent of the total number of collected specimens.
The lycopods, although represented by many form genera of stems,
leaves and fructifications, account for less than 5% of all the
material collected. Of these, the majority of compressions are
Lepidocarpon and Lepidophyllum. The sphenopterids are frequent
and diversified. Their associated genera, Rhodea and Adiantites,
are also present in small quantities. Other species are relatively
rare and any collections from the Lake Mountain or Traverse
Mountain clay pits would result in an accumulation of ten or twelve
species of the more common types, particularly _C_3. utahensis.

The relative abundance of species does not necessarily infer
similar relationships of sparseness or profuseness to the actual
numbers of the plants. A megaflora generally gives a good record
as to the abundance of the lowland flora, but an inadequate record
of the "upland" plant types. The usual plant remains encountered
as megafossils are of plants which grew along waterways or in
low, moist areas and were preserved in situ or near the locality
in which they grew. Very few "upland" forms are represented and

these were probably transported by wind or water.

The microfossil flora should, by comparison, give a more
accurate indication as to the numbers and plant types present.
Unfortunately, this is not always true. Spores and pollen may be
destroyed by oxidation, abrasion and metamorphism. Samples
were collected from the various lithologies in the type section
of the Manning Canyon Shale and from different clay pits. These
were processed for spore-pollen analysis, but were found to be

barren.

TABLE I Cont.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

AB F
Palaeostachya (‘2) sp.
Calamostachys (‘2) sp.
MVP) sp.
Mariopteris muricata Schloth
Diplothmema obtusiloba (Brgnt. ) White X
Dipmmema trifoliata (Artis) White
Diplothmema arnoldi Stockmans 85 Williere
fighenopteris (?Renaultia schatzlarensis (Stur) Zeiller
Sphenopteris gothanica (7) Dolianiti) comb. nov. X'
ngnngterIS angustissima (Stbg) Nemejc
Alloiomeris cruciatus sp. nov.
thenopteridium dissectum Goppert X
Smhenopteridium Zaitzeffium sp. nov. X
Rhodea vesperfina Read
PM sp.
Adiargites sp. X

 

Adiantites Mardia) tenufolius var. defoliolatus
(White) comb. nov.

Zeilleria (‘2) sp.

Neuropteris g1 gantea Sternberg

Neuropteris heterophjlla Brongniart

Neuromeris cf. pocahontas White

NMMNEU

>4

NNNNN

TABLE I Cont.

 

 

AB
Neuropteris ampelinos sp. nov.
Crossopteris utahensis sp. nov.
Crossomeris undulatus sp. nov. X

 

Crossopteris mclmiqhtii sp. nov.
Cyclppteris dilatata Lindley 85 Hutton
Odontomeris sp.

Cordaites princi is (Germar) Geinitz
Cordaites sp. A

Cordaites sp. B

Cordaites sp. C

Cordian’ghus gseudofluitans Kidston
Cordaicarpus llobosus sp. nov.
Cordaicarpus elongatus (Newberry)
Cordaicarpus jayshuleri

Cordaicarpus manninqcanensis sp. nov.
Cordaicarpus cordatus (Jackson) comb. nov.
Maicarpus binutus sp. nov.
Trigonocarpus noegqerathi (Stnbg. ) Brongniart
_‘_I‘__1;iqonocarpus s p. A

lriqonocarpus sp. B

Trigonocarpus sp. C

Holcospermum s p.

I><II><II><Ii><I

TABLE I Cont.

 

 

 

AB
Cornucarpus discissus sp. nov.
Riqbyocarpus ebracteatus gen. & sp. nov.
Lagenospermum bifurcatus (Stopes) comb. nov.
Gnetopsis anglica Kidston X

 

Telanqium affine (L.& H) Benson
Aulacotheca campbelli (White) Halle

Aulacotheca hemingwayi (7) Halle

I><II><II><I*1:J

COMPARISON WITH OTHER NORTH AMERICAN FLORAS
OF MISSISSIPPIAN AND PENN 'SYLVANIAN
AGE

A comparison of the Manning Canyon Shale flora with other
Mississippian and Pennsylvanian floras is very difficult because of
the inadequate knowledge of these floras in the United States. Not
only are floras associated with many of the important coal seams
virtually unknown, but the existing information is often of little
value for comparative purposes. Lesquereux's Coal Flora is
essentially the only reference available for much of the Appalachian
region and many of his figures and descriptions are incomplete.
Thus it becomes necessary to rely upon the European literature
for modern sources of information on Carboniferous floras. The
inherent disadvantages of this were pointed out by Arnold (1949).
He states (p. 152), "It not only reflects the serious lag in the
development of American paleobotany, but it renders the task of
correctly interpreting the smaller and more isolated American
floras unduly difficult. "

The comparison of the Manning Canyon Shale flora with the
Chesterian plant materials from the Uinta Mountains of Utah
(Arnold, 1962); with the Spotted Ridge flora of Central Oregon

(Arnold, 1953); Mamay and Read, 1956). and with the Mosquito

Range flora from Colorado (Read, 1934) is of particular interest
because of their geographic distribution. Fossil plant localities
in the Manning Canyon Shale are geographically midway between
the Spotted Ridge flora (500 miles northwest) and the Mosquito
Range flora (600 miles east). The Mississippian Uinta flora
lies about 150 miles northeast of Lake Mountain.

Archeocalamites radiatus (=Asterocalamites scrobiculatus)
and Rhgd_e_a sp. are the only species common to both previously
described Utah Carboniferous floras. The genera Cardiopteris
(=Frygpsis) and Caulopsis, present in the Uinta flora, are not
foundin the Manning Canyon Shale collections and with the exception
of Archeocalainites, Rhodea andLepidodendrom none of the 36
genera represented in the Manning Canyon Shale flora have been
reported from the Uinta Mountain locality.

If Lacey and Eggert (1964) are correct in referring the
Lgdodendron from the Uinta flora to I_._. vOIMannianum, then
the floras would have three species in common. However, after
examining Arnold's specimens, the author believes their preserv-
ation is too poor for specific identification.

A flora possibly equivalent to the Uinta Mountain flora is

located near the base of the Manning Canyon Shale in the Soldier

Canyon section. This very poorly preserved flora has a
stratigraphic position similar to that of the Uinta flora; but it has
not been extensively collected or studied.

Species common to the Manning Canyon Shale and the Spotted
Ridge flora are Calamites (lVlesocalamites) hesperius and A_s_tg__ro-
phyllites equisetiformis. However, there are striking generic
differences between these two floras. The genera Dicrancmhyllum,
Pecopteris and thllotheca in the Oregon flora have not been
recognized in the Manning Canyon Shale, and conversely , in the
Oregon flora, Neuropteris. Alliopteris, clearly identifiable
lycopsids, Sphenopteris, Mariopteris and several other forms
present in the comparatively well-preserved Manning Canyon Shale
flora have not been found. Whether these generic differences are
due to differences in preservation, the author is unable to say, but
from information available, the floras appear to be distinct.

Mamay and Read (1956) considered the Spotted Ridge flora to
be Pennsylvanian; but were uncertain as to whether the flora
represented Upper, Middle, or Lower Pennsylvanian. Arnold (1953)
thinks its affinities are more closely related to floras of early
Pennsylvanian age.

The species common to the Mosquito Range flora and the Manning

Canyon Shale flora are Asterophyllites longifolius, Neuropteris

heterophylla (7), Neuropteris gigantea, and Stigmaria ficoides
(=§_. verrucosa). There are thirteen otherspecies described
from the Mosquito Range which Read places in the Lower
Pennsylvanian.

These floras are floristically distinctive as well as
geographically separated. The distinct difference is probably
due to a difference in their time of growth, for the Manning
Canyon Shale flora appears to be older. The Oregon and Colorado
Carboniferous floras are very poorly preserved. Many of the
plants which grew in these areas, and which may have been
represented by plant fragments deposited during the accumulation
of these sediments have not been preserved. Therefore, these
fossil floras, as they are known today, may give a biased record
of the Pennsylvanian vegetation of those areas.

The two Mississippian floras from Arkansas, - the Stanley-
Jacks fork and the Wedington, - have no species correlative with
those found in the Manning Canyon Shale. The Stanley-Jack fork
flora, thought to be Pennsylvanian by White (1937), is now con-
sidered as Mississippian by Mamay (in Miser and Hendricks, 1960).

Similarity between the Manning Canyon Shale flora and the
Chesterian flora from Illinois (Iacey and Eggert, 1964) is indicated

by the common occurence of the species Archeocalamites radiatus,

 

 

Lepidodendron volkmannianum and Stigmaria ficoides. L. 17913;
mannianum from the Manning Canyon Shale is an isolated specimen
in quartzitic material which is stratigraphically lower than the
horizon from which the remainder of the collection was made.

Rhodea vespertina is the only species common to the
Mississippian Pocono and Price floras of eastern United States
(Arnold, 1949; Read, 1955) and the Manning Canyon Shale. These
Appalachian floras contain the characteristic Mississippian genera
Tripwerisd Cardioptergs and Lepidodendropsis which do not
occur in the Manning Canyon Shale.

The Michigan Coal Basin flora (Arnold, 1949) has ten forms
from a total of forty-five species in common with the Manning
Canyon Shale. These are Lepidodendron agulreatum, Lepidocarpon
linearfolium, Stigmaria ficoides. Asterophyllites equisetiformis,
N. or. heterophylla, _l_\T. gigantea (?) Sphenopteris obtusiloba,
Acuiacotheca campbelli; Trigonocarpus noquerathi. and
qu'daites principalis. Arnold (1949) considers the Michigan; Coal
Basin flora to be late Westphalian A to Westphalian B in age.

J ongmans (1937), in his subdivision of the Pottsville, interprets
the Pocahontas seams to be equivalent to the European Namurian age
and subdivides its coal seams into Namurian A, B or C (table II).
The coal seams underlying seam #1 which he attributes to Namurian

A has one species in common with the Manning Canyon Shale. Seams

1-3 (Namurian B) and seams 4-6 (Namurian C) have five species and
one species comparable, respectively. The New River Series
(Westphalian A) has two species in common, whereas Kanawha
(Westphalian B) has five and Alleghany (W estphalian C) has one.
The number in common may not be important as the particular
species involved. Coal seams 1-3 contain Alloiopteris sternbergi
and Sphenopteris obtusiloba which are characteristic of this and
higher stratigraphic horizons in Europe. Neuropteris gigantea
and Sphenopteris schatzlarensis are found in the Kanawha (Westphalian
B) and Alleghany (Westphalian C). They are indicative of Upper
Namurian and Westphalian age in Europe.
J ohngmans (1937) states that the Pocahontas seam #3 on the
basis of its principal plant components, Neuropteris pocahontas,
N4 sthehani and A_lloiopteris sternberqi, has a Namurian or older
Westphalian A character. He places the Pocahontas Coal in an
age equivalent to the middle and upper portions of the Namurian
or perhaps as late as the transition into the lowest Westphalian A.
Lepidodendron aculeatum, Lepidophyllmn sp. , Lepidostrobus
variabilis, Diplothmema obtusiloba, Neuropteris heterophylla, and
Asterophyllites wisetiformis are species common to both the
Stanley Cemetery flora of Indiana (Wood, 1963) and the Manning
Canyon Shale flora. Wood (1963) considers the Stanley Cemetery

flora to be Westphalian B in age.

Bell (1938, 1944) divided the Upper Carboniferous strata of
Nova Scotia into three groups based on‘ their distinctive floral
components. These are the Canso Group, the Riversdale Group
and the Cumberland Group.

The Canso Group has three species, Telanqium affine, Archeo-
galamites radiatus (=Asterocalamites scrobiculatus) and Calamites
(Mesocalamites) cistiiformis in common with the Manning Canyon
Shale flora. According to Bell (1944) the poorly preserved calamitean
stems are fairly abundant in the Canso Group, but other fossil plant
remains are scarce. Bell (1944) states that the flora of the Canso
group is possibly equivalent to Dix's (1933) floral zone A of southern
Wales which is comparable to Lower Namurian (Table II).

The Riversdale Group contains five species common with the
Manning Canyon Shale flora. They are Sphenopteris obtusiloba,

_S_. schatzlarensis, A. _eguisetiformis, Lepidostrobus variabilis‘1

and Cordaites principalis. Bell (1944) considers the Riversdale

 

Group to be no older than late Namurian. or younger than Westphalian
A. He favors Westphalian A, and bases this on Neuropteris
schlehani and Mariopteris acuta which are the most characteristic
forms present. He also cites the first appearance of Cordaites

principalis as evidence for this age assignment.

The Cumberland Group which he studied has the same seven
floral components in common with the Manning Canyon as those in
the Riversdale Group and in addition, _Lepidodendron aculeaturn
and Allpiopteris @orynepteris) sternbemi are found in both the
Cumberland and Manning Canyon Shale floras. Bell (1944) cites
this flora as indicative of late Westphalian A or Early Westphalian
B, because - "a number of species reach their acme of development
in late Westphalian B or early Westphalian C time. " He gives
more importance to the presence of Asterophyllites equisetiformis,
Neuropteris tenuifolia and Lepidodendron dichotomum than to the
"rare survival" of Neuropteris schlehani which is not present in
the Westphalian B of Europe. Lonchopteris is the index form for
this time interval in Europe, but it is rare in North America.

Bell (1944) attributes this "to the non-deposition of sediment
throughout most of Westphalian B time. "

The Pictou Group of New Brunswick, which Bell (1962) states
is Westphalian C in age, has Asterophyllites longifolius, Stigmaria

ficoides and Neuropteris heterophylla in common with the Manning

 

Canyon Shale flora.

COMPARISON WITH EUROPEAN CARBONIFEROUS FLORAS

Crookall (1934) outlined the three following factors to be used in
comparing fossil floras: l) the occurrence of certain assemblages
of species; 2) the frequency of individual forms composing the
assembalges; and 3) the absence of other assemblages. The problem
encountered in using his second criterion is the lack of uniform
quantitative data for the various described floras. Occasionally an
author will enumerate relative abundance, but generally only a floral
list is given.

Great Britain: The following species are found in both the floras
of the Upper Carboniferous of Great Britain and the flora of the

Manning Canyon Shale.
R* S* Y* L*

 

 

 

 

 

 

 

Asterophyllites equisetiformis 8 5 8 6
A. Lonqifolius 1 2 l
Cordaites mincigalis ' 3 9 6
Corynepteris sternberqi 1 4
Gnetopsis anglica 3
Lepidostrobus variabilis 2 2 2
Mario pteris muricata l 3 l
Neuropteris gigantea 3 10 3

 

Neuropteris hete rophylla l 2 10 4

29

 

 

 

 

 

 

 

Sphenopteris obtusiloba (‘2)
Sphenopteris schatzlatensis 6 l
Sphenofieris trifoliata _ 7
Stigmaria ficoides 6 6 10 7
Trigonocarpus noegqerathi 4 2 1
Lepidodendron aculeatum 4 4 9 5
Lepido dendron obovatum 2 9 4
Lepidostrobophyllum majus 5 2 1

 

(*Time intervals: R=Radstockian; S=Staffordian; Y=Yorkian; and
L=Lanarkian. Quantitative data: 1-2 = very rare; 3-4 fairly rare;
5-7 = fairly common; 8-10 = very common. )

The six species in common with the Radstockian are from a total of
132 species reported from Great Britain. Twelve from 168 species in
Staffordian, sixteen species in the Yorkian out of 392 , and nine species
of 91 for the Lanarkian are also in common.

Pecopterids as a group are characteristic of the Radstockian Series;
an abundance of Linopteris species marks the Staffordian; and Lonchopteris,
though rare, is typical of the Yorkian. The Lanarkian has several typical

forms of Sphenopteris (Crookall, 1931-32). None of these forms occur in

 

the Manning Canyon Shale.
Species from the Manning Canyon Shale which correspond to the Lower

Namurian of Great Britain are:

Eumorphoceras gqne (Namurian A)
Cordaites principalis
Lepidostrobus variabilis
Mesocalamites cistiiformis

(From a total of 51;
after Dix, 1933)

Reticuloceras Zone (Namurian B)
Calamites cf. cistiiformis
Neuropteris qiqantea
_N. cf. heterophllla
Co rdaites principalis
Stigmaria ficoides

(From a total of 26+
after Dix, 1933;
Lacey, 1951)

Of the species listed for the Lower Carboniferous of Great
Britain (Crookall, 1934), the species (2. cistiiformis , Astero-
calamites scrobiculatus ercheocalamites radiatus), Sphenopteris
affine (=Telangium affine), Sphenopteridium dissectum, and
Lepidodendron volkmannianum also occur in the Manning Canyon
Shale flora.

The largest number of common species are located in Dix's

floral zones C, E, and G (Table I1). These species, as well as

those already given, are composed largely of long ranging species

Lower
Carb.

Upper Carboniferous

.HLPWE H.

 

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meadow . UR WWQMowwMMH , , womflg _ Oommflnmfimm Zo<m moofim Some Ssflam d. m
U _.m__ -Wmmmfloofimb .w EoQUoHQ
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v.
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31

(table III) found throughout either the Lower or Upper Carboniferous.

Bflgium; The Namurian of Belgium as studied by Stockmans and
Williere (1952-53) has a few species in common with the Manning
Canyon Shale. The Assise de Chokier (Namurian A) has only three
species, Stigmaria ficoides, Lepidodendron obovatum and Archeo-
calamites cf. radiatus in common from a total of 77 species. The
Assise d'Andenne (Namurian B) has ten in common from 176 species.
These are Lepidodendron obovath _L. variabilis, Trigonocarpus
noegqerathi. Gnetopsis anglica, Lepidostrobophyllum majus,
Sphenopteris (=Diplothmema) obtusiloba, Corynepteris angustissima,
N europteris qiqantea, Cantheliophorus aff. linearifolius (= Lepi__d_-
carpon linearifolium). and Aulacotheca hemingwayi.

Ancion and Van Leckwij ck (1947 ) described a flora from Zone
de Gilly (uppermost Namurian B) which has five species, Astero-
phygites equisetiformis, A. anqustissima, Neuropteris qiqantea,
Sphenopteris obtusilobaJ and Aulacotheca heminqwayi, corresponding

to species collected from the Manning Canyon Shale.

France: Two species are common to the Namurian A of France
as reported by Carpentier (1928) from the Basse-loire. They are
Asterocalamites scrobiculatus (=Archeocalamites radiatus), and
Calamites cistiiformis. Carpentier recorded 34 species in this

flora.

Carpentier (1907) listed about 50 species for coals of
Valenciennes. The following are species foundin both the
Valenciennes beds and the Utah flora. These are divided into
Carpentier's Zones.

Zone A2 - Base of B1

Mariopteris muricata
Sphenopteris trifoliata
_S_. sternberg’

_S_. obtusiloba
Neurgptergheterophylla

N. g1 gantea
Asterophyllites equgetfiormm

Trignocarpus noeggerathi
Lepidodendron aculeatum
L4 obovatum

L volkmannianum

 

 

Zone A2, which is stratigraphically younger, is characterized

by Zeiller (1888). It contains:
Neuropteris heterophylla
Mario; teris muricata

Sphenopteris trifoliata

Sphenopteris trifoliata is also reported from Zone B1 to B2.

Again, the species common to this flora and the Manning
Canyon Shale flora are only long ranging species with the exception
of L. volkmannianum which appears out of place in the Valenciennes
flora. The number of species corresponding to those from the
Manning Canyon Shale, is fairly small (12 of about 50) in the
lower coal beds, becoming less numerous upward, indicating
very little relationship between these two floras.

Germany: Gothan (1931) listed six species from the Namurian
A in the Westphalian-Rhine basin. None of these are found in the
Manning Canyon Shale.

The Namurian B and lower Namurian C of this same region
has Lepidodendron aculeatum, Asterophyg’gs longifolius,
Myteris sternberqi (- Corynepteris anqustissima) Cordaites
Malia, Neuropteris qiqantea. and Sphenopteris schatzlar-
_e_r_1__s_i_s_ in common from a total of 38 species.

Leggewie (1933) lists Asterophyllites charaeformis, Cordaites
principalis and Lepidodendron obovatum among 24+ others from
the upper Namurian. These three are also present in the Utah
material.

Zimmermann (1958) reports 62 species from the Culm of

Waldenburger. Of this number, Sphenopteridium dissectum,

 

Asterocalamites scrobiculatus (=Archeocalamites radiatus),
and Stigmaria ficoides are found in the Manning Canyon Shale
flora. In a second list for his Namurian flora, he lists 52 species
of which Asterocalamites scrobiculatus (=A_rcheocalamites
radiatus). Lepidodendron v_p_ll_a_nannianum4 I; aculeatum and
Stigmaria ficoides are also present in the Utah flora.

He reports Smhenopteridium dissectum as only from the
Culm, and Lepidodendron volkmannianum and L_._. aculeatum
as only from the Namurian.

Netherlands: J ongmans (1955) synthesized the information
on the coal floras from the Anatolie region. Sixteen species
of these floras are common with the Manni ng Canyon Shale

flora. These are given in chart form with the age where they

 

 

were found in the Netherlands. Namurian Westphalian
DiantAB C A B C D

Asterocalamites scrobiculatus X ?

I-Archeocalamites radiatus)

Mesocalamites cistiiformis X

 

Asterophyllites charaeformis
A: eduisetiformis

A; longifolius

Lepidodendron aculeatum

_L. obovatum
Lepidostrobophyllum majus

Lepidostrobus variabilis

Alloiopteris sternbergi (-_C_._ angustissirna)
Neuropteris gigantea X
_N. heterophylla

 

 

 

 

 

 

 

 

NMNN NNNMN

NNNNNNNMNM

I><Ii><II><Ii><H><I MNNN
NM

 

 

N amurian Westphalian

 

 

Diant A B C A B C D
Renaultia schatzlarensis X X X X
Sphenopteris obtusiloba X X X X X
Spheno pteris trifoliolata X
Cordaites principalis ~ X X 'X 7
Jongmans total species 25 19 30 ‘132 “ 147 124 123

 

Bulgaria: Hartung (1935) reported on the Carboniferous floras

from Bulgaria which share the following species with the Utah flora.

 

 

Namurian Westphalian

 

C
Lepidodendron obovatum Rare
Lepidostrobufivariabilis
Stigmaria ficoides X

 

Astero phyllites longifolius
Spheno pteris schatzlarensis
Neuropteris gigantea

Co rdaites principalis

NNNNNNN tb
NNNNNMN
MN NNNN

Russia: The mutual species between the floras of the Russian

"Coal Measures" and the Manning Canyon Shale are listed as to the

levels used by the Russian stratigraphers for the Namurian.

Level C

Level C

Level C

2
l

l

4

Lepido dendron aculeatum

1:5 obovatum

Lepidostrobqghyllum maius

Stigmaria ficoides

Asterocalamites scrobiculatus (=Asterocalamites
Radiatus)

Stigmaria ficoides

Asterocalamites scrobiculatus

Stigmaria ficoides

Asterocalamites scrobiculatus
Asterophyllites charaeformis
Sphenopteris Erienaultia) schatzlarensis
Lepidostrobophyllum majus '

5

Level C1 Stigmaria fico ides

Asterophyllites charaeformis
A. equisetiformis
Neuropteris gigantea

Stockman and Williere (1953) place of to of of D. Zalessky as
comprising all of Namurian A and a part of Namurian B. J ongmans
(1939) gave his equivalents as C? equalling Namurian B and into

2

Namurian C, C} as part of Namurian C and C 2 as entering into

the Westphalian.

AGE OF THE MANNING CANYON SHALE FLORA

It might be possible to determine the Mississippian-Penn-
sylvanian boundary, which has not been accurately determined in
the type section of the Manning Canyon Shale, by using fossil
invertebrates or fossil plants.

Fossil fish in the Manning Canyon Shale are not well
enough known to be of any value for stratigraphic differentiation
of this formation.

Sadlick (1955) identified as Springerian an assemblage of
fragments of the fossil invertebrates Schizophoria cf. _S. texans
and Spirifer cf. _S_. occidentalis in the uppermost unit from the
Soldier Canyon section. He also reports a Chesterian assemblage
702 feet lower stratigraphically than this Springerian unit. Helen
Duncan (personal communication) writes:

"According to Gordon (MacKenzie Gordon Jr.) the lower

400 feet of the Manning Canyon in the type area contains

an upper Mississippian fauna as pointed out by Girty.

Fossil invertebrates are generally poorly preserved and

relatively uncommon in the upper part of the Manning

Canyon, but a diagnostic Pennsylvanian fauna has been

found near the top of the formation. A considerable

thickness of the Manning Canyon in the type area has not

been precisely dated on evidence from invertebrate fossils. "

The Mississippian-Pennsylvanian boundary has been placed by

some workers at the top of the dark gray-blue limestone, commonly

termed the "medial limestone", which lies about 450 feet above the

 

base of the formation. Others place the boundary 200 to 300 feet
higher in the formation at the arkosic and graywacke zone.

The fossil plant horizon is located within the zone between the
Springerian fauna and the Chesterian assemblage. However, the
plants occur near the top of the section, stratigraphically closer
to the position of the Springerian fauna.

In order to establish the age of this flora, the stratigraphic
ranges of its various species were studied and are plotted on
Table III. In compiling the stratigraphic ranges, floral lists were
studied as well as ranges recorded by Gothan and Remy (1957) and
Dix (1932). The chart shows the probable ranges for all but new
species, although a number of gaps may be noted for species not
frequently reported.

Eighteen of the thirty-five species listed on table III have
stratigraphic ranges throughout all or most of the Carboniferous.
Therefore, species like Asterophyllites longifolius, _A._’ eguiseti-
formis, Cordaites principalis, and Neuropteris gigantea would
only indicate a probable Upper Carboniferous age.

First appearances of so many. species, however, is important.
The two vertical columns on table III, Namurian A and Namurian B,

contai n the majority of first appearances.

Index species are also important. Sphenopteris (7Renaultia)

 

schatzlarensis and Corynepteris anqusjissima are indicative of
uppermost Carboniferous of Europe. Bell (1944) reported both
from Canada. He reports Sphenopteris schatzlarensis from the
Riversdale group of Late Namurian or younger Westphalian A,
and Alloiopteris sternbera_i (=9. anqustissima) from the
Cumberland group which he considers to be late Westphalian A
or early Westphalian B. J ongmans (1937 ) reports _A. sternbergi
from the Pocahontas coal seams 1-3 which ”are equivalent to
Namurian B.

Neuropteris pocahontas and Marioptegiimuricata are also
important index forms for the Lower Pennsylvanian anngpper
Carboniferous. N. mcahontas characterizes the lower Pottsville,
whereas _l§_/I_2 muricata is important in the lower Upper Carbon-
iferous of Europe.

The Manning Canyon Shale flora appears to be transitional
between Mississippian and Pennsylvanian ages.

Excluding the ubiquitous Stigmaria ficoides, there are six
species in this flora which are known to characterize the Miss-
issippian. Three of these, _S_Lhenopteridium dissectum,

Sphenopteris gothanica and Telangium affine-are fairly common,

TABLE III
STRATIGRAPHIC RANGES OF 34 PLANT SPECIES OF THE MANNING
CANYON SHALE

Mis sis sippian Pennsylvanian

Namurian Westphalian
Diant A B C A B C D

Archeocalamites

 

 

radiatus
Asterophyllites _ — _-

 

 

charaefo rmis
Aste rophyllites

 

 

 

equis etifo rmis
As te roph yllite s

 

 

 

lo ngifolius
Calamites (Mesocalamites) 7 ?

 

 

 

 

he spe rius
Calamites (Mesocalamites)
cistiiformis
Lepidodendron

 

 

 

 

 

 

obovatum
Lepido dendron

 

 

 

aculeatum
Lepido dendron

 

 

volkmannianum
Lepidostrobus _ _ _____
variabilis 1
Lepidostrobophyllum
maj us
Stigmaria ficoides
Mariopteris _____ _ _. _
muricata
Sphenopteris (Renaultia) __ _
schatzlarensis
Sphenopteris
gothanica (7)
Corynepteris _ __ _ _ _ _ _—
angustissirna
Diplo thmema ..... -
obtusiloba
Diplo thmema — - - -
trifo liata
Diplo thmema

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

"’ arnowi

 

41

TABLE III Cont.

 

 

 

 

 

Mississippian Pennsylvanian
N amurian Westphalian
Dinant A B C A B

Sphenopteridium

dissectum
Rho dea

vespertina _.
Adiantites (Wardia)
Tenufolius var. r; 7

 

defoliolatus
Neuropteris

gi gantea
Neuropteris

heterophylla (7)
N europteris c_:_f_._

pocahontas
Co rdaites

principalis
Co rdaianthus

ps eydofluitans
Triqqno carpus

no eqqerathi
LacLenospe rmum

bifurcatus
Gentopsis

anglica
Telanqium

affine
Aulacotheca

campbelli
Aulacotheca

heminqwayi(7)

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

h-——

 

whereas three; Rhodea vespertina, Lepidodendron volkmannianum,
and Archeo calamites radiatus are rare. Twenty-two Pennsylvanian
species are present.

In this transitional flora, other typical Mississippian genera
such as Cardiopteris (Fryopsis), Tr_iphyllopteris, Rhaccmteris, and
Lepidodendropsis are lacking; but also conspicuously missing are
Pecopteris; true Alethopteris, Sphenophyllum, Lepidophlo ios.
Bothrodendron, CaulppterisL Megaphyton and Lonchopteris which
are genera found in the Pennsylvanian or younger strata.

Sph enopteris and Diplothmema forms are highly diversified,
although only two species, one Mississippian and one Pennsylvanian
are common. Rhgplea is limited, as is Adiantites. Sphenopteridium
is represented in the flora by two relatiVely abundant species.
Mesocalamites, the dominant calamitean form, is, in itself;
tansitional between Archeocalamites and true Calamites. Calamites,
abundant in the Middle and Upper Pennsylvanian, is rare in the
Mississippian, whereas Archeocalamites is common. Odontopteris,
fairly common in the Manning Canyon flora, is reported by Crookall
(1931‘.- 32) as generally characterizing the Radstockian (Table II);

and although 1... aculeatum and L, obovatum* two of the

 

Lepidodendraceae in the flora, range throughout most of the

Pennsylvanian; they are more frequent in the lower series.

Crossopteris utahensis however, suggests an older flora.
White (1900) consistly thought Neuropteris pocahontas to be a
relatively primitive form of the neuropterids. He based this 1
upon the partial basal attachment of the uppermost pinnules on
the pinnae. He considered this form to be one of the intermed-
iate forms between Neuropteris and an ancestral form common
to the alethopterids and neuropterids. White considered it to be
significant that Neuropteris pocahontas was common only in the
lowermost Pottsville. Crossopteris utahensis is similar to N4
pocahontas in the manner of its small pinnule attachment, but
appears to be more primitive in its more persistent half-basal
attachment and its lobing which is mo re alethopteroid-pecopteroid
than neuropteroid. If White's hypothesis is correct, then Crossopteris
utahensis should be from a horizon older than Lower Pottsville;
However, intermediate forms would be expected to occur in a
transitional flora. Also, since this is a form not previously re-
ported, its range has not been established.

The Mississippian form Archeocalamites radiatus is known to
extend into Westphalian A. Telanqium affine has been previously
reported only from the Lower Carboniferous of Europe and from
the Lower Namurian of Canada (Bell, 1944). Rhodea vespertina ,

reported by Read (1955), from the basal Mississippian Pocono

Formation, is like Sphenopteridium dissectum and the other
Mississippian forms in that they are perhaps "hold'overs"
from an older, diversified flora, some species of which were
still extant at the time the Manning Canyon Shale flora was
being deposited. I

In summary, the plants indicate a lowermost Pennsylvanian
age, and the horizon from which they were collected is close to the
Mississippian-Pennsylvanian boundary. The flora occupies a
zone between Read and Mamay's (1964) floral zone 3 which is
characterized by Fryppsis (=Cardiop_teris) and Sphenopteridium
and their floral zone 4 with Neuropteris pocahontas and Mariopteris
eremopteroides as the zonal species.

The presence of Sphenopteridium, Telangiurn and Archeo-
calamites from the Mississippian in association with components
of a Pennsylvanian flora indicate a very early Namurian age for
this flora, probably Namurian A or possibly as young as Namurian

B.

46

PALEOECOLOGY

The study of the structure, development and affinities of the
Manning Canyon Shale plant types, especially lycopods and
calamitens, found in association with the Pennsylvanian coals of
eastern North America, indicates a paludal environment. Potonie'
(1910) has pointed out that the enlarged bases of many of the older
Sigillarians or other lycopods are analogous to the basal enlarge-
ments of the black gum, bald cypress and other extant forms
growing in tropical and warm temperate swamps. The Lepido-
dendron leaf scars were, in effect, lenticels, or gaseous exchange
surfaces augmenting plant aeration at least when water covered the
root systems (stigmarian rhizomes and rootlets). TheSe shallow
root systems extended horizontally for some distance, thus
functioning effectively as absorptive organs above the toxic
air-exhausted deeper layers (White, 1913). The hollow interior
of stigmarian appendages and the absence of root hairs also
indicate an aqueous habitat.

White (1913) cites the air chambers in calamitean roots, the air
chambers for seed flotation, the production of two kinds of spores
in lycopods and Calamiteans, requiring water to insure fertili-

zation as evidences for humid, swamp environment.

The vegetation at the time of deposition of the Manning
Canyon Shale consisted of pteridosperms, ferns, lycopods and
calamitens growing in or near the water, depending upon the
needs of each individual plant. Forms such as Cordaites
probably grew in areas at the edges of the swamps, possibly
even in upland areas. This vegetation was established in a
region with a uniform climate characterized by generally mild
temperatures. The temperatures were possibly subtropical.

The presence of marine invertebrates and the cyclic nature
of the sediments indicates the swamps were close to the sea.

The repeated vertical change in the sediments from silty, sandy
I and clayey shales to calcareous shales and limestones suggests
two overall shifts in the sedimentary environments from shallow
. to deeper marine.

The plant remains which are not in situ have not been
transported far from their place of growth. This is inferred by
(1) their random orientation and; (2) the occurrence of many
complete plant organs, particularly leaves. Specimens of
Stigmaria ficoides with rootlets attached have been uncovered.
The radially extended positions of the interlacing rootlets,
passing outward from the parent root in all directions and oblique

to the bedding planes indicates the roots are in situ. Several

intact Lepidostrobules, calamitean cones, calamitean and fern-
like foliage have been collected. If these plant organs had been
transported far, they would have been degraded or broken, or
perhaps destroyed.

Most of the fossil plant remains are found in the laminated
silty and clayey shales which were probably deposited in a
lagoonal environment from silt-laden waters. These plant
materials were incorporated into the mud and silt layers. At
some later time, iron-rich ground waters percolating through
the sediments and seeping along joints, fault planes and bedding

surfaces replaced the original plant remains with iron oxides.

SUMMARY

The fossil flora from the Manning Canyon Shale of central
Utah is important because of its floral content, stratigraphic
position and geographical location. The flora contains 68 species
of Mississippian and Pennsylvanian age, indicating that these
strata are transitional between these two periods. Six species
have a Mississippian range, twenty two are known Pennsylvanian
species, twenty one have no definite range, five are longer
ranging species. This flora contains two new genera, thirteen
new species and one species (Tingla' s_p, ) which has been previously
reported only from the Lower Permian of China.

Stratigraphically, the flora lies near the Mississippian and
Pennsylvanian boundary, with a Springerian fauna above and a
Chesterian fauna below. The plants indicate a lone rmost Penn-
sylvanian age and probably represent Namurian A or possibly
as late as Namurian B in European terminology. The flora is
compared here with other Carboniferous floras described from
Europe and North America, but none of these floras appear to be
similar to the Manning Canyon Shale flora. There is some
similarity in long-ranging species present in those floras but

none has the overall aspect of the Utah flora. The flora of Canso

group in Maritime Canada may prove to be the most closely
related but further study of both floras will be. required to
demonstrate this.

The flora is located geographically between the Spotted
Ridge flora of Oregon and the Mosquito Range flora of Colorado.
These three represent essentially the only known Pennsylvanian
floras from Western United States. One small Mississippian
flora is reported by Arnold (1962) from the Uinta Mountains of
Utah.

The flora appears to have been depOSited under swampy
conditions with a rather humid environment in an embayment
which was progressively silting in.

The flora fio m the Manning Canyon Shale exhibits a number
of significant paleobotanical features. Further study of this
flora and other potential Pennsylvanian floras in Western United
States is necessary if a more comprehensive understanding of

the Pennsylvanian paleobotany of this vast region is to be attained.

SYSTEMATIC PALEOBOTANY
Order LEPIDODENDRALES
Genus LEPIDODENDRON Sternberg 1820
Lepidodendron obovatum Sternberg

pl. III fig. 5; text fig. 5B
pl. XI fig. 4

1820 Lepidodendron obovatum Sternberg, -Flora der Vorwelt,

v. 1, fasc. l, p. 20, 23, pl. VI, fig. 1, pl. VII, fig. 1A.

1875-77 Lepidodendron rhodeanum Stnbg. -Stur, Culm-flora der
Ostrauer und Waldenburger Schichten, p. 283, pl. EQIIV, figs.

1, 2, 3.

1951 Lepidodendron obovatum Stockmans and Willie' re. - Quelques
Végétaux Namuriens et Westphaliens du Charbonnage d'Aiseau-
Presle, pl. 2, fig. 11.

1962 Lepidodendrgn sp. A. Tidwell. -Brigham Young University
Geol. Studies. Vol. 9(2), p. 99, pl. 5, fig. 2.

Description: Scars transversely diamond-shaped, bolsters
prominent, 10 mm. long, 5 mm. wide. Elliptical leaf scar located
in the upper one-half of bolster; wider than high; lateral angles
acutely pointed and lower angles obtusely rounded. Lingule scar
at upper edge of leaf scar. Vascular strand and parichnos occur
in lower portion of leaf scar. Large aerencyhma areas on each
side of upper portion of keel. Keel is raised near lower edge of

leaf scar and near base of bolster, but is obscure between.

Transverse markings on the keel are absent on some bolsters and
not well preserved on others.

Discussion: In discussing Lepidodendron mosaic¥umg J ongmans
(1938) states, "These figures show that they can be compared with
Lepidodendron obovatum Sternberglwith which L_._._ rhodeanum
Sternberg is very nearly related, if not identical. "

This form occurs frequently in the Manning Canyon Shale.
Fragments of this species often occur as isolated bolsters or
specimens with a small number of bolsters present.

One specimen (Pl. XI, fig. 4) is a branched portion of a
stem with what appears to be a strobilus or a cluster of sterile
leaves attached. No spores have been obtained from this specimen.

Lepidodendron aculeatum Sternberg

pl. III fig. 7; text fig. 5A
1820 Lepidodendron aculeatum Sternberg 1820. - Flora der Vorwelt,
pl. 1, p. 20, 23;, pl. VI, fig. 2, pl. VIII, figs. lBa, b.
1962 Lepidodendron sp. B. Tidwell. - Brigham Young Geol. Studies,
v. 9 (2), p. 99, pl. 5, fig. 4.

Description: Bolsters prominent, contiguous, rhomboidal with
rounded sides and pointed at top and bottom; 14 mm. wide and 30 mm.
long, acute ends asymmetrical; median line straight or slightly
curved. Leaf scar cuneate with lateral angles and basal angles

acute; top obtusely rounded. Leaf scar situated toward right side of

bolster. Right lateral angle of scar nearly touches right margin of
bolster and occupies one-half of the distance to the left margin.
Vascular scar appears to be in lower portion of leaf scar; parichnos
areas are not preserved; aerenchyma tissue present; keels obscure,
but present. One form has faint transverse irregular lines.

Discussion: The bolsters of Lepidodendron aculeatum are
large with the elongated ends passing beyond the ones directly
above and below, causing the top and bottom ends of the same
bolster to curve in opposite directions, thus giving the volsters a
rather sinuous appearance.

The preservation of Lepidodendron aculeatum (pl. III fig. 7)
is relatively poor. The leaf scars, ligule scars and the keels
are present. A composite figure (Text fig. 5A) made from
several spe cimens illustrates a more can plete bolster.

L. aculeatum is separated from L. obovatum by its more
elongated bolster, although they often intergrade and may at times
be very difficult to distinguish from one another.

Lepidodendron volkmannianum Sternberg
pl. 111 fig. 1; text fig. 5C
1957 Lepidodendron \_7_o_l‘l_<:mannianum Stnbg. -G0than & Remy,

Steinkohlenpflanzen, p. 36; pl. 25.

1964 Lepidodendron volkmannianum Stnbg. -Lacey & Eggert,
Am. Jour. Bot. 51, p. 979, figs. 5-13.

Description: Bolsters somewhat inverted pear-shape; upper
portion ovate, lower somewhat blunt, inserted contiguously above
the bolster below. Leaf scar elliptical to rhomboidal situated
in upper portion of bolster. Bolsters are aligned horizontally
and vertically.

Discussion: The specimens of this species were collected
from a quartzite bed about 200 feet below the plant horizon where
the remainder of the specimens were obtained.

Lacey and Eggert (1964) reported specimens of L. volkmannianum
showing gradations of leaf cushion shape and size on a single stem,
making it possible to assign species to it which were originally
considered to be distinct. Using this criterion, they were able to
assign most of White's (1937) new species from the Wedington Shale
to .L_..‘_ volkmannianum.

Lepidodendron volkmannianum is characteristic of the Lower
Carboniferous and rarely extends into Namurian A. Namurian A

would be roughly equivalent to Chesterian age.

EXPLANATION OF TEXT FIG. 5

Fig. 5A Lepidodendron aculeatum Sternberg (2X) --------- 52
Fig. 5B Lepidodendron obovatum Sternberg (5X) ---------- . 51

 

Fig. 5C Lepidodendron volkmannianum Sternberg (5X) ----- 53

ueuu: scnn
. ‘ LEAF son

A“ vnecuun BOAR

I

if}: ' nsnencnmn new:
A: EEL

e‘\ '

' I

1" i

I)

 

TEXT FIG.

5

Genus LEPIDOSTROBUS Brongniart 1828

Although the majority of the species of Lepidostrgbus are
heterosporous, a few have been described as homosporous.
Abbott (1963) has proposed the generic name Lepidostrobopsis
for such "unisexual" strobili. This genus would contain "unisexual"
strobili, either microsporangiate or megasporangiate, whose
overall characteristics are otherwise similar to Lepidostrobus.
Although this taxon appears to be useful, the problem arises into
what genus should a specimen be placed that (1) has no spores, or
(2) is incomplete with only the apex, containing microspores, or
a strobilus base with only megaspores present. These should
perhaps be placed with Lepidostrobus until more refined criteria
are delineated.

Lepidostrobus variabilis Lindley & Hutton
pl. IlIfig. 3

1831 _Lfipidostgrrobus variabilis, Lindley 85 Hutton. -The Fossil Flora
of Great Britain, v. 1, pls. 10 & 11.
1963 Iepidosfitrgbus variabilis, L. &H.-Wood, Ind. Geol. Surv. Bull.
29, p. 37, pl. 2, figs. 4, 5, and 6.

Description: Strobili linear-oblong, at least 100 mm. long and
18 mm. wide in specimens studied. Margins are essentially parallel;

base and apexrounded or truncated. Axis 4 mm. wide. Sporophylls

wide, fusiform in cross-section, 3 mm. wide, 2 mm. high.
Sporophst crushed together, spirally arranged, inserted
obliquely to the axis near base, becoming more oblique near

apex. Bracts about 6 mm. long, 1 mm. wide; acuminate-
lanceolate to linear, overlapping distal laminae of 3 to 4 spor-
ophylls above. Pedicel 3-7 mm. long with sporangium on upper
surface; crestline of sporophyll slightly curved upward above keel.

Discussion: No spores are present. Specimen preserved as
limonite replacement with minute, quartz, box-like structures
transcending the sporophylls obliquely across the surface of the
cone.

Lepidostrobus variabilis L. &. H. is an extremely variable
form. Into this group are placed Lepidostrobus cones which have
sporophylls resembling sterile leaves and are often difficult to
separate.

In 1880, Lesquereaux wrote, "from the figures of the species
given by European authors, it seems evident that different kinds
have been described under the common name. " Zeiller (1888)
also found this species to vary as to size and disposition of the
sporophylls.

Specimens from Greene County, Indiand, housed at the

University of Missouri (Wood, 1963) and the Geological Survey

of Canada compare favorably with the Utah material. Bell (1944)
thought that his specimens of _LL variabilis from the Cumberland
Group, the Riverside Group and the Stellarton Group may have
included more than one species. But, examination of the types
does not indicate great variations.

Gothan and Weyland (1959) discuss}; variabm “s and mention
that its axis corresponds to the structure of Lepidodendron
oldhamius Williamson. Its axis also shows a central stele with
pith similar to Lepidodendron vasculare Binn. Hirmer (1927) also
notes L. variabilis may be the strobilus of L. oldhamius.

Stockmans and Williere (1953) report that the strobili of the
arborescent lycopodiales are relatively common in the Namurian
of Belgium. In the quarry at Rieudotte in particular, they found

perfect types of I_.._ variabilis associated with Lepidodendron

 

obovatum and Lepidophloios laricinus.
Lepidostrobus skemmatos sp. nov.
pl. Illfig. 2; text fig. 7E
Description: Atrobilus oblong with parallel margins tapering
to form a pointed apex. Strobilus incomplete, basal portion is
missing. Axis 2 mm. wide. Sporophylls crowded and crushed;

spirally arranged, normal to the axis. Bracts linear-elongate

to linear-lanceolate, at least 11 mm. long, 2 mm. wide at base,
and overlapping 3-6 sporophylls distally. Pedicels 6-7 mm. long
with sporangia on upper surface, crestline about hcrizontal with
slight rise toward bract.

Discussion: No spores are present. This type varies from
Epipmtmbm variabilis in shape of the bract, more compact
structure, sporophylls more nearly perpendicular to the axis,
pointed apex and larger size.

Lepidosytrquus skemmatos is similar to Lepidostrobus
Luger; Lx. , but L. Mis smaller, of more oblong-oval
shape, and apex is not as pointed.

Genus LEPIDOCARPON Scott 1900

Lepidocamn is a genus of seed-bearing lycopods founded by
Scott (1900, 1901) for petrified cones of both Upper and Lower
Carboniferous age. However, Schopf (1941) emended Scott's
original description to include all such compressions within the
genus. Schopf (1940) transferred to Lepidocarpon all twelve
compression species previously assigned to Cantheliophorus
by Bassler (1919).

Bassler (1919) based his genus upon (1) two sac-like sporangia

borne on short sporangiophoric stalks per sporophyll; and (2) a -

plate of sterile sporophyll tissue ascending from the ventral
midline of the pedicel and to which the sporangiophores were
attached laterally and distally. Schopf (1941) states Bassler has
not properly shown that the two sac-like bodies exist and the
"median" plate to be other than the compressed form of the
Lepidocarpon integument.

Lepidocarpon is used for fructifications with strobilar habit
which shed their entire sporophylls by disintegration of the
strobilus at maturity. An integument completely envelopes the
megasporangium which provides the large "seed-like" megaspore
with a distinct integumentary organ. A slit between the two
membranes at the crest corresponds morphologically. to a
micropyle. Only one megaspore matures per sporophyll. The
remainder of the tetrad aborts (Schopf, 1941).

Lepidocarpon linearifolium (Lesq) Schopf

pl. V fig. 3 text fig. 7F
1880 Lepidophyllum linearifolium Lesquereux, Coal Atlas, 2nd
Geol. Surv. Penn. , Rept. of Prog. , pp. 452-454, pl. 69, fig. 39.
1919 Cantheliophorus linearifolius (Lesq.) Bassler - Botanical
Gaz. Vol. 67, p. 97, pl. IX, figs. , 1, 8-10.

1940 Lepidocarpon linearifolium (Lesq. ) Schopf - Schopf, J. , in

J anssen, 1940, Illinois State Museum Sci. Pub. Vol. I, p. 43.
1949 Lepidocarpon linearifolium (Lesq. ) Schopf. -Arnold, Univ.
Michigan Contr. Mus. Paleont. , v. VII, p. 174, pl. IX, figs. 2-3.

Description: No strobilus. Single megasporophyll with
sporangium showing ligule. The pedicel is perpendicular to the
axis and extends 11 mm. to the upturned distal lamina. Distal
lamina is 30-35 mm. long, extending from base of heel to apex
of sporophyll (distal lamina). Vascular bundle enters directly
from the axis, and can be seen nearly to heel where it disappears.
Lateral lamina extends above and around the sporangium, and forms
an investing structure. Ligule obscure. Megasporangium small,
10 mm. long, 4 mm. wide, sporangial cavity about 2. 5 mm. high.
Sporangial wall extends vertically above the megaspore for l. 5 mm.
to formthe megasporangial ridge. Sporangium appears to be
attached for nearly its entire length along the upper surface of the
pedicel.

Discussion: Isolated sporophylls of Lepidocarpon lineari-
folium are very abundant in the Manning Canyon Shale. These
sporophylls are nearly exact duplicates of L. linearifolium
specimens described by Arnold from the Lower Pennsylvanian of

Grand Ledge, Michigan.

Arnold (1949) states the most conspicuous feature of L_.§p_i_._d_:
carpon @earfioHM' is the long slender terminal bract lamina
which generally extends horizontally, but in many cases is at a
45° angle to the pedicel. The latter form is the general type
encountered in the Manning Canyon Shale.

The specimen described by Lesquereux consists of the distal
lamina and part of the pedicel. If a megasporangium was present,
he did not illustrate it.

The stratigraphi‘ range of L._ linearifolium is at present
undefined.

Genus LEPIDOSTROBOPHYLLUM Hirmer 1927)

Lepidostrobophyllum rpaju_s_ (Brognt.) Hirmer
Pl. III, fig. 6

1928 Lepidophyllum m Brongnt Prodrome
1880 Lepidophyllum majus Brognt. -Lesquereux, Coal Flora,
2nd Geol. Surv. Pennsylvania, Rpt. of Prog., p. 499.
1927 Lepidostrobophyllum majus(Brognt.) Hirmer - Handbuch
der Palaobotanik, Bd. 1, p. 193, fig. 213.

Description: Lamina of sporophyll large, rather oblong,
parallel margins tapering near tip to a lanceolate shape with an

acute apex. Single midrib; sporangiopho res obovate, obtusely

rounded at the base.

Discussion: Lesquereux (1880) states that the blades are
generally 7 -9 cm.long and 13-16 cm. wide. He reports them as
rare in the American Coal Measures. They are also rare in the
Manning Canyon Shale.

Crookall (1925) in comparing the specimens of L. _r_n_ajps_, he
had collected, with L. missouriensis, as figured by White (1899)
states that the latter species should be combined with L. MES.-
Abbott (1963) in discussing her new genus Lepidostrobopsis, in which

she placed L, missouriensis, reports the difference between the

 

two is that the sporophyll of L, mambears hair or cila along
its margins, whereas the margins of L. missouriensis are smooth.
Crookall (1925) reports that L. floccus throughout the
Coal Measures of Great Britain.
Genus LEPIDOPHYLLUM Brongniart 1828
Lepidophyllum longifolium Brongniart
pl. IV, fig. 5
1939 Lepidophyllum longifolium Brongniart; J anssen, Leaves and
Stems from Fossil Forests, p. 62, fig. 44.
A large number of the sterile leaves of the ancient lycopods
are found isolated and in a f ‘ragmental condition. Some of these

from the Manning Canyon Shale are referable to this species.

Although incomplete, some measure 4 mm. wide and over 20 mm.

65

long; complete length indeterminable. A single midrib traverses
the length of this linear, grass-like leaf.

Lepidophyllum sp.

pl. III fig. 4

These sterile leaves are smaller than Lepidophyllpm Logan;-
_f_9_l_i__u_r_r_1_, 5-6 cm. long, 4 mm. wide. The leaves are elongate
triangular in shape with a flat, truncated base and an acute apex.
A single vein is continuous from base to apex. Stomatal grooves
parallel the vein.

Leaves of this type have been found attached to L. obovatum
where they may constitute the base of a strobilus with sporangia
in the axils or they may be sterile foliage. The preservation is
not usfficient to determine if spores are present.

Genus STIGMARIA Brongniart 1822
Stigmaria ficoides (Sternberg) Brongniart
, pl. IX, fig. 5
1820 Variolaria ficoides Sternberg. -Flora der Vorwelt, vol. 1,
fasc. 1, pp. 23-26, pl. 12, figs. 1-3.
1822 Stigmaria ficoides (Stnbg.) Brongniart. -C1ass. distr. veget.
foss. Mem. Mus. Hist. Nat. V. 8, pl. 1, p. 219, fig. 7.

1877 Stigmaria verrucosa Miller. - Mer. Paleozoic Fossils, p. 40

 

1899 Stigmaria verrucosa White.-U. S. Geol. Surv. Prof. Paper

 

185-D, p. 82.

1962 Stigmaria verrucosa Tidwell. -Brigham Young University Geol.
Studies, v. 9 (2), p. 99, pl. 5, fig. 1.

Description: A few, rather large-sized specimens were
collected. Rhizome with spirally disposed circular scars of
rootlets, each with a raised upper rim and a central cicatrix.
Appendages (rootlets) still attached.

Discussion: Our knowledge of Stigmaria, the rhizome of
certain Lepidodendreae, is still very imperfect, although fossils
of Stigmaria are among the most common Carboniferous specimens.
The idfficulty lies in the inability to place specimens of Stigmaria
into natural species or to relate them to the genus of lycopods to
which they were attached. They have been found in connection
with the stems of both Sigillaria and Lepidodendron.

The species name Siipmaria verrucosa is invalid as the
species term was derived from Phytolithus verrucosus Martin
(1809), a term used prior to 1820; which is the priority date for
paleobotanical nomenclature.

Order EQUISETALES

Genus ARCHEOCALAMITES Stur. 1875
Archeo calamites radiatus (Brgnt.) Stur.

pl. V, fig. 7, text fig. 6C

1828 Calamites radiatus Brongr_u’ar_t, -Prod. Hist. Veg. Foss. I,
p. 122, pl. 26, figs. 1,2.

1875 Archeocalamites radiatus (Brongnt.) Stur. -Culmflora Abh.
‘K. Geol. Reinchsanst. v. 8, pl. 1, figs. 3-8; pl. 2, 3, 4, pl. 5,
figs 1 & 2.

1964 Archeocalamites radiatus (Brongnt) Stur. -Boureau,
Sphenophyta, Noeggerathiophyta. Traite de Peleobotanique III,
p. 209, figs. 186, 187, 188.

Description: Compression. Ribs straight, narrow, passing
directly through the nodes. The foliage consists of long, narrow,
linear leaves which bifurcate several times. The leaves arise
from nodes. .

Discussion: Archeocalamites radiatus is the only species of
Archeocalamites from which the leaf-bearing shoots are known.
The dividing of the leaves by repeated bifurcations is contrary to
the general leaf types found in the Equisetales. Both the unbranched
part of the archeocalamitean leaf and its segments are extremely
narrow; more or less filiform. Halle (1925) points out that the
leaves are given off in great numbers in radially symmetrical nodes.
They are ascending or slightly spreading, attaining lengths of 10 to

12 cm. or more. They regularly bifurcate (pl. V, fig. 7) into

two similar halves with the entire leaf having a uniform appearance.
Halle (1925) states that the branching of this species is sparse, and
its lateral shoots are finely divided.

Archeocalamites has a particularly primitive appearance with
its bifurcated leaves and opposite ribs (text fig. 6C). This
primitive group was possibly derived from the Hyeniales of the
Middle Devonian. Its foliage has changed during this transition
from the leaves of Hyeniales which were thickened to Archeo-
calamites which are disposed in a plane (Boureau, 1965).

lestikow (1959) demonstrated that Calamites scrobiculatus
Schloth. and Asterocalamites scrobiculatus (Schloth.) Zeiller
should be assigned to Archeocalamites radiatus.

Lacey and Eggert (1964) placed all of White's (1937b) astero-
calamitean species from Arkansas, Bell's Canadian asterocala-
mitean forms and Arnold's (1962) specimens from the Uinta flora,
Utah, with Archeocalamites radiatus.

Archeocalamites radiatus has been reported from the lower
Carboniferous up into Namurian A which approximates the Miss
issippian and the very lowest Pennsylvanian.

Genus CALAMITES Suckow, 1784

(Hist. Comm. Acad. Elect. Sci. , Litt. Theod. -Palat. V: 355, Mannheim)

Fig.
Fig.

Fig.

Fig.

Fig.

6A
6B

6C

6D

6E

EXPLANATION OF TEXT FIG. 6

Illustrates the dichotomies of a Mariopteris frond.
Illustrates the dichotomy of a Diplothmema frond.
Schematic diagram of the ribs of Archeocalamites
passing directly through the nodes.

Schematic diagram of Mesocalamites showing the
alternation of some ribs passing directly through
the node and other ribs alternating.

Schematic diagram of the alternating ribs of true

Calamites.

69

Calamites(Mesocalamites) hesperius Arnold
pl. V, figs. 2,5

1953 Calamites hesperius Arnold. -Pa1aeontographica, V 93B,
pp. 62-63, pl. 24, figs. 1, 6-8.
1956 Mesocalamites hesperius (Arnold) Mamay and Read. - U. S.
Geol. Surv. Prof. Paper 274-1, p. 215, pl. 34, figs. 4, 4a.
1964 Calamites (Mesocalamites) hesperius Arnold. -Boureau
I Traite de Paleobotanique III, Sphenophyta, Noeggerathiophyta.
p. 247, fig. 217.

Description: Pith cast compression. Ribs nearly straight

 

with rounded or flattened surfaces; ribs 1. 5 mm. wide, some
terminating in acute or rounded points and alternating; others
truncated and pass directly across the node. Tubercles oval-
elongate near rib apex; smaller oval tubercles near rib base.
Internodes 2. 4 - 3. 0 cm. in length. No branch-scars are present.

Discussion: Arnold (l953)informally discussed the affinities

 

of this species with the Mesocalamites group of Hirmer. Mamay
and Read (1956) recombined it, fiving this taxon official status.
Boureau (1964), however, reduced the genus Mesocalamites to a
_subgeneric position.

The mesocalamitian group, because of its series of alternating

ribs and ribs crossing directly through the node, occupies a

morphological and stratigraphic position between Archeocalamites
and the other subgenera of Calamites.

Archeocalamites with its ribs extending straight through the
nodes, is considered to be a primitive calamitean form. The other
subgenera of CalamitesL such as Stylocalamites Weiss, Diplo;-
calamites Gothan, Crucicalamites Gothan and Calamophyllites
Grand'Eury have most of their ribs alternating.

Mesocalamites resembles a mixture of these two types, with
the opposing ribs more or less predominant over the alternating
ribs (text fig. 6D). The vertical range of Archeocalamites is
essentially lower Carboniferous through Namurian A, whereas
Calamites ranges through the Carboniferous and Permian. Misp-
calamites is positioned more or less transitionally from Namurian
A. into Westphalian A.

Calamites (Mesocalamitesl cistiiformis Stur
p1. IV fig. 3, pl. V fig. 1
1877 Calamites cistiiformis Stur. -Culm Flora II, K. Geol.
Reichsanst. Sbh. Wien, VIII, p. 200, pl. 4, figs. 5,6.
1917 Calamites cistiiformis Stur. - Kidston and J ongmans, Monograph
of Calamites of Western Europe, Rijpsop van Delfst. Meded. , v. 1

pp. 192-195

1964 Calamites(Mesocalamites) cistiiformis Stur. -Boureau,
Sphenophyta, Noeggerathiophyta. Traite de Paleobotanique III,
p. 245.

Description: Internodes longer than broad, straight, narrow,
alternating or passing straight over the node. The alternating
ribs end in acute apices. Tubercles appear to be oval; furrows
straight, ribs 1 mm. wide.

Discussion: The width of the specimen illustrated in pl. IV
fig. 3 is 24 mm. and its internodes are 42 mm. long.

There is a strong similarity between 9. cistiiformis and g.
_<_:_i_§gi_ Brongnt. J ongmans and Kukuk (1913) stated that these two
species might eventually be united.

Kidston and J ongmans (1917), however, state that although
many authors would unite g. cistiiformis and _C. _<_:_i_§_t_i_, Q. plat;-
formis differs from Q. gist; in the constant occurrence of some
of the ribs passing through the nodal line and some alternating.
The ends of the alternating ribs are sharp pointed in g. Cistiiformis
whereas they are rounded in _Q. p18];

_C, cistiiformis is a typical Namurian form, especially for
Namurian A. However, Gothan, Leggewie and Schonefeld (1959)

found it frequently in Namurian B and C. They state that Q. cistiformis

is not found in the lower Westphalian. Leggewie and Schonefeld
(1961) report its occurrence in the Homoceras zone (Hl & H2)
(Table II).
Calamites (Stylocalamites) sp.
pl. IV ,, fig. 7

Description: Compression of a pith cast. Ribs narrow,

 

relatively straight, alternating; constricted at the nodes to form
two small branch scars.

Discussion: None of the available literature describes or
illustrates any species comparable to this form. Not enough
material is available to make a new type. The internodes are
considerably longer (74 mm.) than broad (15 mm.) and the ribs
are approximately the same size as the intervening furrows. Of
the number of specimens attributable to thisspecies, very few show
evidence of branch scars.

This species is somewhat similar to the form illustrated by
Lesquereux (1880) as the stem of Asterophyllites graciles.

Calamites LCalamatina7) sp.
pl. V fig. 4, text fig. 7D
Description: Incomplete specimen of pith cast showing large

branch scars and leaf scars. Internodes between nodes with branch

scars are 30 mm in length, whereas the internodes between nodes
bearing leaf scars are about two-thirds as long. Ribs poorly
preserved. Branch scars roughly rhombohedral in outline with the
umbilicus small, oval, approximately 6 mm. wide. Two nodes
bearing branch scars occurring together are followed by at least
two nodes bearing leaf scars. The specimen is incomplete and it

is not known whether the cycle repeats itself. Branch scars appear
to alternate around the stem with those on the node below not being
aligned with the scars above.

Discussion: The specimen is too incomplete to make a com-

 

parison with specimens previously figured. The manner of the cycle
of branch scars and leaf scars appears to be unique. No similar
illustration or description was found in the literature. This
specirnan may, however, constitute an abnormality of some species
already known.

The branch scars are actually too widely spaced to be placed
with the subgenus Calamitina and too close, large and numerous to
coincide with Diplocalamites.

The shape of the branch scar of this species is similar to those
described and figured by J ongmans (1956) for Calamites pyriformis

although somewhat reverse to it in outline. The shape of the branch

Fig

Fig.

Fig.

Fig.

Fig.

Fig.

.'7A

7B

7C

7D

7E

7F

EXPLANATION OF TEXT FIG. '7

Co rdaianthus pseudofluitans (‘2) Kidston 134

A strobulis in aXil of a bract. Note bifurcated

 

sporophylls.
Rigbyocarpus ebracteatus sp. nov. 148
Zeilleria (?) sp. (2. 5X) lOO

Dotted lines indicate fructifications. Note

the thickened terminus of the veins

Calamites (Calamitina) sp. (nearly l/ZX) '74
Two nodes with branch scars with two lower

nodes showing leaf scars.

Lepidostrobus skemmatos sp. nov. 59
Megasporangium and sporophyll with a distal

lamina figured apartto illustrate its shape and size

Lepidocarpon linearifolium (1X) Schopf 61

76

 

 

 

 

 

. .
I \u n «I
. ._I .n . u
.\ ... M. . .x. F
a \ i
w .s lo all \s T _ u D
no a, o. v - s
N I |\ ~ v ~
.I a“... v .n
a
H

/ A”
i/ / \tM/V A
.. N\x\\\ E

V \ a

TEXT FIG. 7

scars is also similar to those of Calamites discifer Weiss, but

the specimen differs in other ways. He separated Q. pyriformis
from g. semigchularis and Q. wedekindi on the form of the branch
scars.

This species differs from _Q. wedekindi on the shape of the leaf
scars which are elongate-oval in that species and transversely
oval. on the Manning Canyon specimen. These two have a similar
distance between scars.

Genus ASTEROPHYLLITES Brongniart 1822
(Class. , Mem. Mus. Hist. Nat. V. 8, p. 210)

Differentiating between the genera Asterophyllites and Annularia
is often very difficult. They are similar in that both have linear-
lanceolate, single-veined leaves which are more or less united at
their bases with a sheath.

Abbott (1958) in her comprehensive work on these genera,
separates them on width-length ratios and the position of the leaves
in the whorl in relation to their axis. In Asterophyllites the leaves,
which are of equal length, are usually cupped around the axis. In

Annulariag the leaves, of equal or unequal length, radiate from the

 

node. She states that these orientations are not always constant,
but a contrary orientation would be an exception and probably due

to preservation.

She defines the margins of the leaves from Asterophyllites as
essentially parallel or straight, gradually tapering to an acute apex,
whereas the leaves of Annularia are lanceolate to obovate or
spatulate.

Asterophyllites equisetiformis (Schloth) Brongniart

pl. V fig“ 8, pl. IV fig. 1

1828 ero h ' es eguisetiformis (Schloth.) Brongniart. —
Prodrome d' hist. veg. foss., p. 159, 176.
1958 Asterophyllites eguisetiformis (Schloth.) Brongniart. -
Abbott, Bull. Amer. Paleo. No. 174, p. 299, pl. 35, fig. 4:; pl. 36
figs 12, l5, 19, 30; pl. 39, figs. 46, 47, 49-51; pl. 43, fig. 63; pl.
47, fig. 78.
1962 Asterophyllite 8 _sp, it; Tidwell. - Brigham Young University
Geol. Studies, v. 2(2), p. 98, pl. 4., figs. 6,8.

Description: Leaves linear-lanceolate with nearly parallel
margins. They are 5-6 mm. in length with a length-width ratio of
20 to l. The single vein occupies nearly 1/2 the width of each leaf.
They occur in whorls with the average 12 per whorl. The nodes from
which the whorls originate appear to shorten toward the end of the
branch causing the leaves from each preceeding whorl to overlap
the succeeding whorl, giving the tip a "paint brush" effect which is

characteristic of this species.

I‘V

Discussion: Asterophyllites equisetifprmis is the most
abundant calamitean foliage found in the Manning Canyon Shale.
However, it appears to be a somewhat smaller form than normal
A, equisetiformis in both size and number per whorl.

Abbott (1958) lists the stratigraphic range for this species
as Pottsville through Allegheny, although Stockmans and Williere
(1952) report it from the Namurian of Belgium.

Asterophyllites charaeformis (Sternberg) Goeppert

pl. IV, fig. 2
1825 Bechera charaeformis Sternberg. -Flora der Vorwelt I, v. 4,
p. 30, pl. 55, figs. 8,5.
1844 Astgpphyiitgsglgmeformis (Sternberg) Goeppert, in
Wimmer, Flora von Schlesian Preuss, und Oster, Anth. , Breslau,
p. 198.

Description: Leaves 3 mm. long with 4-10 leaves per whorl.
Their width-length ratio is 1:4. Leaves are falcate, wide at base.
Their basal attachment is nearly perpendicular to the stem but
cups somewhat distally about the axis. The vein is nearly one-half
the width of the leaf.

Discussion: This form is rare in the Manning Canyon Shale.

 

Its distribution as given by Abbott (1958) is Pottsville to lower

Allegheny. According to Leggewie and Schonefeld (1961),

Asterophyllites charaeformis should not occur in Namurian C,
but one would encounter Asterophyllites unguis Jongmans and
Stockmans, With which Asterophyllites charaeformis is often
confused. Because the difference between these two is not
clear, the Utah specimens were placed with A, charaeformis.
Asterophyllites longifolius (Stnbg.) Brongniart
pl. IV, fig. 4
1825 Bruckmannia longifolia Sternberg. -Flora der Vorwelt,
V. 1, no. 4, p. 45, pl. 58, fig. 1.
1828 Asterophyllites longifolius (Sternberg) Brongniart. -
Prodrome d'Hist. Veget. Foss. , p. l59-l‘76.
1880 Asterophyllites longifolius (Stnbg.) Brongnt. -Lesquereux,
Coal Flora, Sec. Geol. Surv. Pennsylvania, Rept. P, p. 36, v. l.
1958 Asterophyllites long1_f' olius (Stnbg.) Brgnt. -Abbott, Bull
Am. Paleont., 38 (174), p. 303, pl. 40, fig. 53, pl. 42, fig. 60,
chart L
1962 Asterophyllites longifolius (Stnbg.) Brongniart. -Tidwell,
Brigham Young University Geol. Studies, v. 9 (2), p. 98, pl. 4
fig. 7.
Description : Leaves approximately 33 mm. long and covering

one internode and most of another (20 mm. between internodes).

Leaves curve upward on some and parallel the rachis on others.
Some may be distorted due to preservation and others are lost;
venation obscure.

Discussion: Asterophyllites longifolius is very rare in the
flora from the Manning Canyon Shale. The specimen illustrated
on P1. IV fig. 4 is very f ragmental with incomplete whorls,
although the length of the leaves is clear.

Lesquereux (1880) states that Asterophyllites longifolius bears
some resemblance to Asterophyllites rigidus Geinitz, and gives the
broader stems with shorter internodes and more rigid leaves of
A, rig'dus as the major distinctions. Miklausen (1949) illustrates
how these factors are not constant, and concludes by using
Asterophyllites longLf' olius forma rigida Weiss for at least one of
his specimens.

The age range of Asterophjflites longifolius is Pottsville to
Upper Allegheny (Abbott, 1958).

Genus PALAEOSTACHYA Weiss 1879
(Steinkohl. Calamar. I. Abh. 2. geol. Spezial K. v. 2 (l):lOB-lO5)
Palaeostachya (‘2) sp.
pl. IV fig. 6
Description: The main stem from which the branches arise

appears to be a mesocflamitean form with some ribs alternating

and some extending through the nodes. Ribs taper into the node
forming round cicatrices from which the branches are attached.
The branches are also ribbed and give rise to a whorl of strobili
at each node. Each strobilus consists of alternating sterile bracts
and sporangiophores in its lower portion, but is comprised of
sterile bracts only in its upper portion. Sporangiophores are
peltate and located in the axils of the sterile bracts. Axis of the
strobilus is ribbed, but not too well preserved. The sterile
bracts are generally oblique to the axis, occasionally saucer-
shaped and rarely at right angles, turning upward abruptly near
the tip of the bract. No spores were preserved.

Discussion: Although the species is of the Palaeostachya type,
it is placed with this genus provisionally. The lack of sporangio-
phores with the sterile bracts near the cone apex is not clearly
understood. This form is similar to Palaeostachya elongata (Presl)
Weiss, but the latter species does not have'the feature of only
sterile bracts in the upper portion of its strobili.

Genus CALAMOSTACHYS Schimper 1869
(Traite, V. l, p. 328)
Calamostachys (?) sp.
pl. V, fig. 6
Description: Small, incomplete cone, 15 mm. long, with

alternating sterile and fertile sporophst which are associated

with foliage of the Asterophyllites equisetiformis type. Fertile
whorls are situated midway between the sterile whorls. The
sporangiophores are of a cruciate shape and are borne at right
angles to the cone axis. No spores were obtained from this
specimen.

Discussion: No illustrations or descriptions were found in

 

the literature of a calamitean cone which was similar to the form
described here. However, the form is too incomplete to give it
a new species name or to assign it to any other species.
Genus .THVGIA Halle 1925
(Geol. Surv. China, Bull. '7, pp. 4-6)
wasp.
_pl. VI, fig. 3

Description: Frondlike. Leaves obcuneate to linear-oblong

 

with rather truncated apices. These apices are dissected through
narrow incisions to form 2 to 3 linear lobes with obtuse apices
which may again be divided near their apices. Leaves alternate,
oblique, attached with a broad base; veins fine, bifurcating mostly
in the lower part, supplying one to two veinlets to each lobe.
Discussion: Tingia is characterized by frond-like anisophyllous
shoots with a thick axis. The leaves are arranged in four rows, two

on the upper and two on the lower side of the axis. The leaves on

the upper side are broadly obcuneate-obovate to oblong or linear,
deeply lobed. Those on the lower side similar, but narrower and
often more deeply dissected. The venation of this genus is nearly
parallel to the margins. I

These characters fit Ill-9919.51)- except for the anisophyllous
structure and the broad axis. The anisophyllous structure,
although not present, may eventually be discovered with further
collecting. However, this structure is rarely uncovered, even in
known species (Stockmans - personal communication).

Msp. is extremely close to Tingia carbonice Halle, the
type species for the genus, and differs only on the size and the

s lender axis of Msp. In fact, they are so close that 11491.31.

sp. looks like a small replica of the specimen figured by Halle
(1927) on plate 62, fig. 6. rI‘i_n_giit_sp. appears to be from near the
apex of a frond which may account for its smaller size.

The genus T_i_n_giQ_is presently known only from the Lower
Permian of China. Darrah (1938) has reported Tingia from the

Permian of Texas, but Mamay (1964) believes Darrah's specimens

belong to another genus unrelated to Tingia.

86

Orders FILICALES AND CYCADOFILICALES

Genus Sphenopteris Brongniart 1822

 

The genus Sphenopteris constitutes a heterogenous group and
assignment here does not imply systemic relationship between the
species which are included in it. Brongniart (1822) defined the
genus as "pinnules cuneiformes, arrondies ou lobees a' l'extremite'
et a' nervures palmees ou rayonnantes de la base de la pinnule".

Sphenopteris includes the sterile organs of species with general
similarity of shape, their pinnule attachment and their venation.

It is simply a provisional group which probably includes both ferns
and pteridosperms.

The pinnules are mostly small, wedge-shaped, often rounded
generally more or less contracted at the base. They are rarely
simple and are generally divided into angular or rounded lobes with
obtuse or acute apices. A single midvein in each pinnule gives
rise to simple or dichotomizing secondary veins which extend into
each segment or lobe.

Sphenopteris originally included species whose fructification

 

are now known. These species have been placed with genera based
upon the structure and arrangement of their fructifications.

Sphenopteris is distributed throughout the Mississippian and

 

Pennsylvanian.

Sphenopteris (Renaultia) schatzlarensis (Stur) Zeiller

 

pl. VII, fig. 4a, text-fig. 8C
1885 Hapalopteris schatzlarensis Stur. Pars. - Die Farne: Carb. -
Flora d. Schatzlarer Schichten I Farne: K- K. Geol. Reichsanst,
Wien, XI, p. 58, text-fig. 2, pl. XXXEX, figs 7, 7a; pl. XL, figs.
2-6.
1899 Sphenopteris (Renaultia) Schatzlarensis(Stur) Zeiller. -Flore
Foss. bassin houil. d'Heraclee, p. 15, pl. 1, fig. 2.
1923 Sphenopteris (?Renaultia) schatzlarensis Stur. -Kidston,
Fossil plants of the Carboniferous rocks of Great Britain: Mem.
of Geol. Surv. Breat Brit., Paleont. v. 11, pt. 2, p. 123, pl. XXIX,
fig. 4; pl. XXXfigs. 1, la.

Descriptions: Antepenultimate pinnae incomplete, rachis some-

 

what flexuous. Penultimate pinnae: alternate, normal to the primary
rachis, lanceolate, rachis flexuous. Ultimate pinnae: alternate,
oblique, lanceolate with rounded apices. An ultimate pinna arises

at each flexuous curve of the secondary rachis. Pinnules: alternate
oblique, deltoid, 2-3 lobes which spread in more mature forms;
complete basal attachment, decurrent, contracted slightly above;
apices rounded, obtuse, sometimes acute. Venation: midrib arising

acutely from rachis, dividing to supply each lobe with one veinlet.

Discussion: Sphenopteris schatzlarensis belongs to the
narrow, delicate-leaved sphenopterids. Its fertile remains

belong to the Renaultia-type. The structure of its sporangia is

 

still not certain. Zeiller (1899) reported a fertile specimen which
Kidston (1923) notes as being imperfectly preserved and therefore
having insufficient characters to warrant placing S. schatzlarensis
in Renaultia.

Boweria schatzlarensis has larger pinnules with segments
wider in prOportion to their length and generally entire as compared

with _S_. schatzlarensis. According to Bell (1944), the frond of

 

_S_. schatzlarensis is more delicately out than that of Boweria

schatzlarensis, and the two species should no longer be confused

 

with one another.

Kidston (1923) restricts this species to the Westphalian;
whereas Daber (1955) places it from Namurian B to Westphalian D.
Bell (1944) records occurrences of Sphenopteris schatzlarensis in
Cumberland and Riversdale groups in Canada.

Sphenopteris gothanicaj?) Dolianiti) comb. nov.
pl. VI, fig. 6; text fig. 8D

1954 Adiantites gothanica Dolianiti.-B01. Div. Geol. Mineral.

 

148, p. 41, pl. 1, II.

Description: Frond at least bipinnate. Penultimate pinnae not
complete; rachis felxuous. Ultimate pinnae; somehwat (oblong with
constricted base, alternate, slightly decurrent to secondary rachis.
Terminal pinnule missing. Pinnule: triangular, deltoid or rhomboidal,
dividing into 2-4 spreading cuneate segments which are in turn
split into 2-3 cuneate, rounded lobes; margins on outer segments
tapering toward base, attached with a short, broad footstalk,
constricted base, decurrent, acute, rarely obtuse apex on each
segment; alternate, oblique, rarely normal to the rachis. Venation:
distinct, decurrent, arises from rachis acutely to enter the pinnules
at a sweeping angle; bifurcates to supply each segment with two
veinlets.

Discussion: Gothan (1928) stated that Berry's (1922) Palmatopteris

 

furcata from Peru was incorrectly identified and that it resembles

a narrow-leaf Adiantites and renamed it Sphenopteris paracasica

 

Gothan. Colianiti (1954) reported Gothan believes his S. paracasica
is really Adiantites and requested that he use Adiantites paracasica
rather than create a new species. However, Dolianiti separated
his species from Gothan's on the basis of the small "nervils" on
his specimens.

Gothan's description is incomplete and his illustration is too

poor to determine venation or anything else about his species except

size and shape. It appears to be smaller than the Manning Canyon
specimen and smaller and more flabellate than, Dolianiti's species.
Gothan's specimen has truncated apices, whereas Adiantites
gothanica Dolianiti has more-slender pinnules and lobes with acute
apices. Gothan's form does resemble an Adiantites, but Dolianiti's
appears to be a Sphenopteris.

A similar form is Sphenopteris pseudo-furcata Kidston which
also has two veinlets per lobe. The pinnules of S. pseudo-furcata
are not as deeply incised or as spread out as those of Sphenopteris

gothanica. This species is fairly abundant in the Manning Canyon

 

Shale flora and is placed provisionally with Sphenopteris gothanica
although it lacks the "nervils" which are so important in Dolianiti‘s
description. This may be due to lack of preServation or they may
have never been present. These "nervils", as defined by Dolianiti
in his species, may also be due to some vagary of preservation.
They do not appear to be consistent along the main veins and seem
to cross one another rather than anastomosing into networks which
Dolianiti describes. Dolianiti gives no idea as to how consistent
these "nervils" are.

Dolianiti (1954) gives the age for this species as Lower Carbon-

iferous.

Genus CORYNEPTERIS Baily 1860

Corynepteris was founded for unique fertile foliage placed with
the leptosporangiate ferns. Their sporangia are united in sori,
with 6-8 situated along the axis. The axis is represented by the
f form genus Zygopteris, Zeiller (1883) created the genus M
Mfor fructifications similar to those included in Corynepteris,
but Grand Eurya also included fructifications belonging to As_te_r;<_>_-_-
mpg; Dober (1955) provided a new interpretation of Saccopteris
which would have included Corynepteris, but Danze (1956) comments
that Stur considered Saccopteris as not only a fructification, but a
complete genus in the botanical sense, and therefore, Saccopteris
should be placed in synonymy with Corynepteris.

Potonie' (1899) proposed Alloiopteris for the sterile foliage; but

 

according to Danze (1956) these two genera. can now be united.
However, Kidston (192 3) states that "most probably the fossils
placed in Alloiopteris are only sterile examples of members of the
genus Corynepteris, but until their fructification is known, they
cannot be placed in the latter genus. " Therefore, new species of
this type without fructifications should be placed with. Alloiopteris
until their fructifications become known. Those with fructification

known may be properly assigned to Corynepteris.

Ii

Corynepteris angustissima (Sternberg) Nemejc
p1. VII, fig. 5, text-fig. 8a

1823 Aspidium angustissimum Sternberg. -Versuch einer geognotischen
botanischen Darstellung d. Flora d. Vorwelt, v. 1, fasc. 2, p. 29,
pl. 23, fig. la & b.
1825 Pecopteris angustissima sternberg. Versuch. v. I, fasc. 4,
p1. XVIII.
1854 Aspenites sternbergi Ettingshausen. -Die Steinkohlenflora von
Radnitz, K. -k. Geol. Reichsanst. , Abh. Wien. V. 2, p. 42, pl. XX
figs. 2,3.
1899 Alloiopteris sternbergi Potonie, N. -Lehrbuch der Pflanzen
palaontologie, p. 139.
1899 Sphenopteris (Corynepteris) sternbergi Zeiller. - Mem. Soc.
Geol. France No. 21, p. 24, pl. IIfigs. 8,9.
1923 Corynepteriszygopteris sternbergi Kidston.-Mem. Geol. Surv.
Great Britain, Paleont. III, p. 301, pl. LXXIV, figs. 1,5; p1. IJQCVII,
figs. 1,2.
1938 Corynepteris anqustissima Nemejc. - Rev. Karvon. a. Perm.
Pal. Bohemiae, part 2: no. 16, pp. 15 (Czech) 41, (Engl), pl. Ifigs

3, 4; p1. III figs 8-10, text-fig. 4, 5, 9b.

1955 Saccopteris sternbergi Daber. — Pflanzengeograph. Besond
Karbonflora Zwickau-Lagauer. Geologie Berlin, Beih, No. 13
(1955), p. 22, pl. IV, fig. 2.
1956 Corynepteris anqustissima Danze. -Contribution a L'etudes
des Sphenopteris, fougeres sphenopteridiennes; Etudes Geologiques,
Houill. du. Bassin du Nord et du Pas-de-calais, p. 325-331, p1.
LII, figs. 4-6; pl. LIV, fig. 1,2; pl. LV, fig. 3, 3a

Description: Small, assymmetrical, pecopteroid leaflets
with complete basal attachment, united, sloping forward, 4-6
teeth, alternate perpendicular to the rachis. Venation: midvein
decurrent, arising acutely, branching into two equal dichotomies,
which subdivide again to supply a veinlet to each tooth.

Discussion: The French authors use Corynepteris angustissima

 

which Sternberg figured as Pecopteri; but, as Danze (1956) points
out, which has pinnules similar to those for which Ettinghausen

created Asplenites sternbergi. The Germans use Alloiopteris

 

sternbergi designated by Potonie (1899) for the form. White (1899)
created Alloiopteris winslovii for similar forms. Thus we have
three names with the specific differences being whether you speak
French, German or English. However, Alloiopteris winslovii may

be a valid species, based on fairly consistent larger size.

Corynepteris antuqtissima differs from Cormepteris erosa on
smaller pinnules which have fewer and more rounded lobes or teeth.
Corynepteris angustissima has a wide distribution and is a
readily recognized species. It has been reported from Asia Minor,

Canada, Central and Western Europe.

This species ranges from Namurian B through Westphalian,
although it is rare in the Namurian, becoming fairly abundant in
the younger Carboniferous strata.

Genus ALLOIOPTERIS Potonie 1899
Alloiopteris cruciatus sp. nov.
pl. VII, figs. 3, 7

Description: Pinnules: small (3 mm long), essentially the

 

same size, assymmetrical, slightly inclined forward, sphenop-
teroid with (complete basal attachment, decurrent, alternate,
united, normal to the rachis. Venation: assymmetrical; midvein
decurrent along rachis, essentially paralleling the lower pinule
margin, dividing four to five times. Two branches of midvein to
lower side of pinnule, three branches to the upper side. These
secondary veins may again divide supplying each lobe or tooth with
a veinlet.

Discussion: The uniform, assymmetrical pinnule attributed

 

to this species compose long, linear pinnae which are characteristic

of the genus Alloiopteris. They are larger than pinnules of most
species assigned to this genus. They are larger, more erect with

more branching of the veins than Corynepteris anqustissima Stnbg.

 

They are more erect and not so broad as g, coralloides Gutb.

 

and are not as deeply dissected as Corynflteris essinqlii Andra.

 

The venation also differs from these three forms. The midvein

branches more than in C_._ anqustissima and _C_. coralloides. The

 

venation pattern in _A_. cruciatus differs from _(_3_. essinglii in being
more distant between branches of the midrib and broader, more
dichotomising secondary branching.

_A_. cruciatus is very close to A, quercifolia (Goepp) Pot.
in pinnule shape, but A; quercifolia has more elongated pinnules with
a number of secondary veins arising from the midrib.

Alloiopteris cruciatus cannot be assigned to Corynepteris because
its fructifications are presently unknown.

Genus RHODEA Presl 1838
The genus R_hqc_l_e_a_1 was proposed by Presl (1838) for sterile,

fernlike forms which resemble Sphenopteris, but are separated

 

from it by having more deeply incised, linear, decurrent, delicate
pinnules containing a single vein which is not always visible. The
segment is greatly reduced, giving the appearance of vascular

strands surrounded by thin laminae. Kidston (1923) arbitrarily

Fig.
Fig.

Fig.

Fig.

EXPLANATION OF TEXT FIG. 8

 

 

 

 

Page
8A Corynepteris angustissima (Stnbg.) Nemejc (10X) 92
8B fighenopteridium dissectum Goppert (2. 5X) 102
8C Sphenopteris (Renaultia) schatzlarensis (Stur)
Seiiler 87

8D Sphenopteris gothanica (?)(Dolianiti') comb. nov. (2X) 88

96

 

separated Rhodea from Sphenopteris on the basis of the width

 

of the lamina that borders the single central vein of the segment.
Presl (1838) placed Rhodea together with Hymenophvllites in
Hemenophyllaceites, and subsequently, had placed it in synonymy
w ith Hmenophyllites. Stur (1879) re-established it as a separate
genus.
Kidston (1923) assigned the fertile forms to Zeilleria,
Urnatopteris and some species of Telangium.
A. Carpentier (1929) reported a fructification for _l3_h_p_d_ea
gutheri and considered it a pteridosperm. Gothan and Weyland (1954)
placed R_h_o_de_a_ among the Sphenopteridaceae and among the
pteridosperms, although they included the ferngeilleria frenzli.
Danze (1956) preferred limiting 31.9293 to pteridosperms and
reserved Zeilleria for the ferns. This would complicate the genus
by mixing fertile and sterile forms. Rhodea, like Sphenopteris,
is a heterogeneous genus and should, perhaps be handled the same
way by separating the species as their fructifications are discovered.
_thdea was originally considered to be restricted to the
Lower Carboniferous, but, although it is more common in the
Mississippian and Lower Carboniferous rocks, species belonging

to this genus also occur stratigraphically higher.

Rhodea vespertina Read
p1. VII, fig. 8
1955 Rhodea vespertina Read. - F loras of the Pocono Formation
and Price Sandstone: U. S. Geol. Surv. Prof. Paper 263, p. 22;
pl. 3, figs. 3,4; pl. 4, figs. 1-4, pl. 16, fig. 6.

Description: Frond incomplete. Tertiary pinnae: alternate
to subopposite, oblique, linear-lanceolate, slender axis. Pinnules:
slender, delicate, finely divided, lobed, linear, rounded apex;
alternate, oblique, single vein.

Discussion: Read (1955) reports this species as the most
widely distributed and mo st abundant plant species in Lower
Mississippian of the Appalachian trough. Arnold (1962) reports
a Rhodea sp. which he suggests may be conspecific. His figured
specimen is s’milar to those from the Manning Canyon Shale.

_thdea sp.

Description: Tertiary pinnae: linear-lanceolate, incomplete.
Pinnules: small, erect, alternate, oblique, open, elongate, distant,
forking at 450 two or four times to form elongated segments or lobes
with but a slight change in width of the lamina. Apices of these
segments are rounded. Lamina rather thick, of nearly equal width
in all parts of the pinnule. The lower-most pinnule shows a zig-zag

(flexuous) pattern of branching. This appears to be present in

varying degrees in the other pinnules, but not as conspicuous.
Venation arises from rachis, decurrent, forks to fill each lobe
with a vein.

Discussion: This form is similar to White's Bho_d_e_a
qoepperti (Ettingshausen) Stur reported from the Stanley Shale
of Arkansas. White (1937 ) has originally thought his form was
Sphenopteris qersdorfii based on the "zig-zag" pattern of its
branching and on the general shape and size of its pinnules; but
he has decided in favor of Ii. goepperti. Rhodea sp. differs
from both on its more slender, delicate pinnules. This m
form approaches closely the illustrations Kidston (1923) gives
for the sterile foliage of Telangium bifida. Assignment of this
species, however, must await more complete specimens.

This species is also similar to Rhodea lemayi Broussier and
Bertrand. However, 13.. Wis larger with more variable
pinnules as compared to _Rhggga sp.

Genus ZEILLERIA Kidston 1884
Zeilleria (7) sp.
p1. VI, figs. 1,4; text-fig. 7C
Description: Frond rather delicate; rachis slender, rigid.

Ultimate pinnae alternate, oblique, linear-lanceolate, close,

overlapping. Pinnules alternate, oblique, finely divided, deeply
incised into 2 to 4 lobes or segments. Lobesslender, forming only
a slight wing of lamina on each side of the single vein in each

lobe; apices of the lobes are rounded. Fruiting structures of some
nature terminate the vein in each lobe, some hanging pendant or
nearly pendant.

Discussion: Rhodea-like forms, but fih_o_d_ea is for sterile
foliage. Therefore, this species is provisionally placed with
Zeilleria because that genus was proposed for fertile fern foliage
with the sporangia terminating the pinnule or each veinlet.

Genus SPHENOPTERIDIUMSchimper 1874

Sphenopteridium is a form genus which includes fern-like
foliage with laterally dichotomizing fronds which are simply
pinnate. Ultimate pinnae of these fronds are sessile or slightly
pinnate. Ultimate pinnae of these fronds are sessile orslightly
petioled. They are largely oval near frond base, becoming more
elongate and lanceolate near frond center. Pinnules shorter and
subtrilobed to trilobed. Pinnules are wedge-shaped with truncate
or rounded apex. Veins parallel the margins of the pinnule and

dichotomize.

The foliage attributed to this genus was initially described
by Goeppert (1852) under Cyclopteris and attached to the
neuropterids. Schimper (1874) redescribed them under
Wand placed them with sphenopterids. Pontine
(1899) and Gothan and Weyland (1954) have placed this genus with
Archeopteridales. Gothan and Weyland (1954) placed Sphenopter-
idiu_m_ with pteridosperms and assign Calathiops -1ike fructifications
to it.
Sphenopteridium differs from Sphenopteris by its veins
radiating from the pinnule base and by its lacking a distinct midrib.
Sphenopteridium dissectum (Gopp) Schimper
pl. VII, fig. 2, text fig. 8b
1852 Cyclopteris dissecta Goeppert. -Foss. Fl. des Uberganesgeb.
Nova Acta Acad. Caes. Lep. Car. Nat. Cur. Suppl., p. 161, taf.
XIV, fig. 3.
1874 Sphneopteridium dissectum (Goepp.) Schimper. - Traite Pal.
Veg. v. 3, p. 488, pl. CVII, fig. 12.
1875 Archeopteris dissecta (Goepp.) Stur. -Cu1m Flora, Pt. 1 K.
Geol. Reichsanst. Abh. v. 8, p. 61.
1899 Sphenopteridium dissectum (Goepp.) Schimper. -Potonie,

Lehrbuch der Pflanzenpalaeontologie, p. 130, text-fig. 119.

1962 Diplothmema subdecipiens, White. - Tidwell, Brigham Young

 

University Geol. Studies, v. 9 (2), p. 97 , pl. 3, fig. 6.

Description: Frond at least tripinnate. Penultimate pinnae
lanceolate, normal to rachis, alternate, closely spaced. Ultimate
pinnae triangular or broadly lanceolate, alternate at nearly right
angles to the rachis. Pinnules cuneate, bisublobate to trilobate
with lobes having truncated, obtuse or rounded apices. Small
pinnules are decurrent and broadly attached, constricted above.
Venation: no central midvein, several veins arise from rachis and
curve upward and outward, divinding and furnishing many veinlets
per lobe.

Discussion; Sphenopteridium dissectum differs from Sphen;
opteridium paclyrrachis (Goepp. ) Schimper by having narrower
ultimate pinnae and from Sphenopteridium collombianum (Schimper)
Potonie' by having more triangular pinnules.

Sphenopteridium dissectum is similar to the polymorphous
Diplothmema subdecipiens which has elongated, lobed, truncated
to rounded pinnules; but differs from it on the midrib which _D;

subdecipiens has and S. dissectum lacks.

 

 

Sphenopteridium dissectum is considered to be Lower Carboniferous
in age. Danze-Corsin (1960) gives its age as Dinantian (Mississ-
ippian). Daber (1959) states its age as Middle Visean or Upper

Mississippian.

Sphenopteridium zaitzeffium sp. nov.
p1. VII, fig. 12, text fig. 9B
1962 Sphenopteris (Diplothmema) spinosa Goeppert. -Tidwell,
Brigham Young University Geol. Stud., v. 9(2), p. 97 , pl. 3,
fig. 5.

Description; Tripinnate (?). Pinnae flabelliform, deltoid
to lanceolate, alternate, attached by broad, although rather
constricted stalked-base. Pinnules constricted above, decurrent
below on the pinna rachis, attached with a somewhat constricted
base, alternate, deltoid, flabellate, divided into two spreading
lobes which in turn divide into three to four cuneate segments with
obtuse apices, most are entire. Venation: veins often obscure in
the lamina. No midvein is present. Veins enter pinnules as a
cluster from the rachis, dichotomously divided to fill each lobe

and segments with finer veins.

Discussion: This form from the Manning Canyon Shale has

 

some similarities with Diplothmema furcatum (=Palmatopteris
furcata), but differs from it by having more rounded and spread
out pinnules, and also in the venation. _I_). furcatum has a single
vein per lobe.

Sphenopteridium Zaitzeffium varies from the other Spheri-

opteridium species by its mo re flabellate shape of both pinnules

 

and pinnae, and its somewhat broader footstalk. This species
was named for the author's colleague, James. B. Zaitzeff.
Genus ADIANTITES Boeppert 1836
This genus has had a rather confused history. According to
Danze (1956), Goeppert (1836) had originally placed his new genus
with the neuropterids and had used it again in 1852 for a division

of Cyclopteris in opposition to the division of Neuropteroides.

 

 

White (1904) stated that Goeppert had originally proposed
Adiantites to include species of Ginkgo. Geinitz (1854) and
Zeiller (1883) both used Adiantites in synonymy with Neuropteris

and Cyclopteris.

 

Schimper (1869) prOposed the new genus Adiantides for

 

foliage similar to Goeppert's Adiantites. Adiantites was sub-

 

 

sequently placed in synonymy with Adiantites by Kidston (1887)
who had created Macrosphenopteris, a closely related genus.
Adiantites, as presently used, appears to be synonymous

with Aneimites, Ettingshausen (1865) raised the subgenus Aneimites,

 

described by Dawson (1860) to generic level. The usage of

Aneimites as a genus persists in North America. White (1904)

 

rejected Aneimites as being "untenable in its restricted application

 

as employed by Schimper, Stur and others and as now generally

recognized. " He stated that "the emended genus is indistinguish-
able from the American plant to which Dawson gave the name

Aneimites. " and concluded to use Aneimites rather than Adiantites.

 

Danze (1956) comments that the European paleobotanists have
dropped Aneimites into synonymy with Adiantites. To illustrate

how confusing this has become, Read (1964) mentions "Aneimites-

 

like Adiantites. "

 

Gothan and ‘Weyland (1954) describe Adiantites as a valid
genus, and retained Potonie's (1899) concept of placing it in the
Archaeopteridales.

It would seem that Adiantites should be conserved over
Aneimites. .

’ Read (1955) defines Adiantites as rather lax, angular fronds
which are several times pinnate, owing to a climbing habit.
"Pinnae alternate. Pinnules: alternate, usually cuneate, apically
truncate, attached at the base by a footstalk or sessile, the base
constricted. Venation derived from a single strand or plexus of
strands at the base and radiating to supply the lamina by repeated
dichotomy. Texture usually coriaceous. "

Adiantites sp.
pl. VI, fig. 5

Description: Generally detatched pinnules cut into two lobes.

 

Oblong-cuneate to deltoid with rounded apex, stalked, venation
consists of several veins of equal size radiating from the base
and bifurcating as they extend to the margins.

Discussion: Adiantites sp. is close to Adiantites ungeri
Read and A. antiguus (Ettingshausen) Stur, but is much smaller.
This species may be new but the specimens are not complete
enough to establish specific differentiation.

Adiantites (Wardia) tenufolius Goeppert var. difoliolatus

(White) comb. nov.

p1. VII, fig. '11

1943 Aneimites Mardia) tenuifolius (Goeppert) var. difoliolatus
White.- U. S. Geol. Surv. Prof. Paper l97-C, p. 101, pl. 36,
figs. 8, ll.
_. 1962 Aneimites (Wardia) tenuifolius (Goeppert) var difoliolatus
White. -Tidwell, Brigham Young University Geol. Studies. v. 9 (2),
p. 98, pl. 2, fig. 4.

Description: Specimen small. Pinnules oblong in larger and

 

cuneate in 'smaller forms, 5 to 10 mm. long, 2 to 5 mm. wide.
Contracted to a narrow base; divided into two or three unequal
cuneate lobes which are obliquely truncated or rounded. Obscurely
striated with nervation indistinct.

Discussion: Specimen very small and delicate, having both

 

difoliate and trifoliate lobation. They are lower Pennsylvanian

in age.

Genus DIPLOTHMEMA Stur 187 7
Stur (1877-7 9) proposed Diplothmema to include many
unrelated species which he regarded as an absolutely natural
genus. White (1943) in the discussion of this genus enumerates
the complications and restrictions imposed upon it by various
authors which have resulted, in White's Opinion, in a genus

without any of its original components. He concludes that

 

Diplothmema should include non only the round-pinnuled species
placed in it by Zeiller (1888), but other round-pinnuled species
with bipartitely divided, but not distinctly quadripartite fronds,
which are not definitely referable to Mariopteris as originally
defined.

Corsin (1932) pointed out that he could see no fundamental
differences between the fron structure of D_ip10thmema and

Mariopteris and that they were intergradiational.

 

Danze-Corsin (1953) reserved the term D_iplothmema
"(diplo=two, thmema: section)" for those plants whose frond is
definitely composed of two symmetrical sections bearing the
primary rachis and lacks a quadripartite aspect (text-fig. 6B).
She considered the primary rachis to be in general naked, but
occasionally partially covered with pinnae. The pinnae of this

genus have sphenopteroid pinnules.

White (1943) in his synopsis states that Diplothmema,
Mariopteris and Palmatopteris were similargin their major
structural features and habit of grovth. They were apparently
climbers or lianas with relatively slender, flexuous, aerial
trunks or axes.

Diplothmema trifoliolata (Artis) Stur

 

p1. VII, fig. 1
1825 Filicites trifoliolatus Artis. -Antediluvian Phytology, p. 11,
p1. XI.
1828 Sphenopteris trifoliolata (Artis) Brongniart. -Prodrome d‘une
histOire des Végéteaux Fossiles, p. 50.
1880 Pseudopecppteris trifoliolata (Artis) Lesquereux. -Coal Flora,
Penn. 2nd Geol. Survey, Report P, Vol. 1, p. 217.
1885 Diplothmema trifoliolata (Artis) Stur. -K. Geol. Reichsanstalt
Abh. Vol. 11, p. 346, pl. 19, figs. 1-4.
1943 Diplothmema trifoliata (Artis) Stur. -White, U. S. Geol. Surv.
Prof. Paper 197-C, p. 98, pl. 32.
1962 Diplothmema trifoliolata (Artis) Stur. -Tidwell, Brigham
Young University Geol. Studies, v. 9(2), p. 96, pl. 3, fig. 7.

Description: Ultimate pinnae alternate; open atnearly right

 

angles, ovate lanceolate, narrow flexuous rachis, Pinnules alternate,

small, oblique, uppermost ovate-cuneate, remainder ovate,
constricted near base. Largest are lobed, forming three to
five ovate lobes, becoming pediculate with the terminal lobe
larger and unequally deltoid. Venation indistinct, although
primary nerve is strongly decurrent.

Discussion: The pinnules of I_)_. trifoliolata are smaller

 

and more distant than those of _D. obtusiloba Brgnt. and the plant
is more lax. The trifoliate segmentation of the pinnules frequently
occurs on'specimens of 12. obtusiloba; but these are probably
dependent upon their position within the frond. This has probably
lead to some erroneous conclusions, (Kidston, 1903). Kidston
(1923-25) doubted that Sphenopteris trifoliolata existed on
continental Europe but that the various authors using this taxon
were actually identifying S. obtusiloba.

Kidston (1923-25 reports S. obtusiloba (=_I__). obtusiloba) as
occurring in Westphalian.

Diplothmema obtusiloba (Brongniart) Stur
pl. VII, fig. 10

1829 Sphenopteris obtusiloba, Histoire des vegetaux fossiles,
p. 204, pl. LIII, fig. 2.
1877 Diplothmema obtusilobum (Brongniart) Stur. -Die Culmflora

K. k. Geol. Reichsanstalt Abh., B. 8, Heft, 2, p. 230.

1943 Diplothmema obtusiloba (Brgnt. ) Stur. -White, U. S. Geol.
Surv. Prof. Paper l97-C, p. 97, pl. 30, fig. 4; pl. 35, fig. 7,9.
1962 Diplothmema spectabilis White. -Tidwell, Brigham Young
University Geol. Studies, v. 9(2) p. 96, pl. 3, fig. 8.

1962 Diplothmema obtusiloba (Brgnt.) Stur. -Tidwell, Brigham,

 

Young University Geol. Studies, v. 9 (2), p. 97, pl. 3, fig. 9.
Description: Quadripinnate. Penultimate pinnae ovate

to oblong-lanceolate, alternate, at nearly right angles to primary

rachis. Ultimate pinnae oblique to nearly right angles to the rachis,

alternate oblong to broadly lanceolate. Pinnules stalked, alternate

sometimes contiguous, decurrent; pinnules near center of pinna

are oblong to oval with less prominent lobes, and those near

pinna apex are more or less rounded and entire. Venation indistinct.

Midrib originates low on rachis and branches outward towards

margins. In the more or less orbicular pinnules the veinlets

radiate from the base without any clearly defined midvein.

Discussion: The similarity between this form and D. trifol—

 

iolata (Artis) Stur was discussed under the later species.
Diplothmema arnoldi Stockmans 31 d Williere

1956 Diplothmema arnoldi, Sto ckmans and Williere. -Vegetaux de

 

1a Zone d'Oupeye d"Argenteau-Sarolay. Publ. Assoc. Etude.

Paleont. no. 25, pl. A.

Description: Penultimate pinnae about 2 cm. wide. Rachis
appears to be slightly winged. Ultimate pinnae; parallel borders,
10+ mm. long, 4-6 mm. wide, alternate, normal to the rachis.
Pinnules: triangular, some more or less circular, divided into
lobes which are subdivided into digitate segments. The number of
lobes per pinnule diminish to two in direction of pinnae-apex.
Venation: mid-vein distinct, divides twice supplying each digitate
segment and lobe with one lateral vein.

Discussion: This is the first reported occurrence of this

 

species in North America, and perhaps the first outside of
d'Argenteau-Sarolay where it was originally described. Stockmans
& Williere (1956) report this form from the "Zone d'Oupeye" which
is equivalent to lowermost Westphalian A.

Genus MARIOPTERIS Zeiller

Mariopteris was created by Zeiller (1878) as a genus to include

 

certain ferns which, among other characters, are distinguished
by a peculiar dichotomizing of their pinnae.

The frond of Mariopteris is composed of quadripartite pinnae;

 

that is, they are divided into four parts. The primary pinnae are
attached to the rachis by a naked stalk which bifurcates, dividing
the frond into two equal segments. These two short, primary

petioles are also naked; and again divide, forming the secondary

pinnae are borne the tertiary pinnae, which in turn support the
pinnules.

The pinnules of Mariopteris are pecopteroid to sphenopteroid,
generally sub-triangular, with complete basal attachment or
constricted at the base. They are always decurrent and sometimes
confluent. The pinnules are entire, lobed, or with margins undulated,
or more or less toothed. The basal pinnule on the posterior side
is usually distinctively larger than the others and divided into two
prominent lobes which may be entire or dentate.

The nervation is sphenopteroid with oblique ascending veins.

It is generally immersed and often difficult to trace. The mid-vein
extends to near the apex of the pinnule. The secondary or lateral
veins arise acutely and divide once or twice before reaching the
margins.

The fructification of Mariopteris is at present essentially
unknown. Gothan (1935) reported Calathiops Bernhardti as having
affinity with Mariopteris acuta. Danze-Corsin (1953) accepts this
as confirming the hypothesis Mariopteris belongs to the pterido-
sperms. Daber (1955) states the clarification of this relationship
has not been demonstrated, and that the type of fructification
belonging to Mariopteris is still unknown; although Mariopteris is

undoubtedly a pteridosperm.

White (1943) discusses the difficulty of differentiating
Mariopteris species due to the variation of size and form of
the pinnules in different positions within the quadripartite
frond and in younger as compared to older fronds of the same
plant. The species also are very intergradational. White
thought one could distinguish the various species by arranging
them according to their stratigraphic position. He considered
the forms to be confined to and even to be characteristic of these
stages. To do this, however, would require very fine and subtle
differentiation in order to divide the long-ranging species. To
use these forms stratigraphically would require an expert to
differentiate these subtle differences between the newly created
species. These differences may also not be of sufficient magnitude
on which to base a specific distinction.

Mariopteris muricata (Schlotheim) Zeiller
p1. VI, figs. 2,8; p1. VII, fig. 6

1804 Schlotheim, "Flora d. vorwelt" pp. 54,55, p1. XII, figs. 21,
23.
1836 Pecopteris muricata Brongniart "Hist. des Veget. Foss. "

Vol. I, p. 352, pl. xcv, figs. 3,4.

1880 Pseudopecopteris muricata Brngn. -Lesquereux, Coal

 

Flora, Vol. I, p. 203, p1. XXXVII, fig. 2. .

1886-88 Mariopteris muricata (Schl.) Zeiller. -Flore fossile du
basin houillier de Valenciennes, Etude Gites Mineraux de
France, pl. XX, fig. 2,3; pl. XXI, fig. 1.

1953 Mariopteris muricata (Schl. ) Zeiller. -Danze-Corsin;

 

Les Mariopteris du Nord de La France; EfJudes Geol. Houill.

 

du Bassin du Nord et du Pas-de-calais; pp. 122-135, pl. XXII
to XXXVIII.

Description: Fragmental. Ultimate pinnae: alternate,

 

oblique, lanceolate or triangular. Terminal pinnule: broadly
triangular, acute apex, sublobate base. Pinnules: ovate-
lanceolate to sub-triangular, acute apex, lower margin of

young pinnae strongly decurrent, upper margin constricted,
complete basal attachment with an assymmetrical base, margins
entire, crenulate becoming toothed, alternate, close, oblique,
larger pinnules near base becoming lobed with five lobes with
acute apices forming, spreading out somewhat fan-shaped.

11 mm. long, 4 mm. wide. Venation: obscure, one strong midvein
arising obliquely from the rachis near lower margin of the pinnule,
decurrent, swinging through center and terminating before

reaching apex. Lateral veins arise from midvein alternately,

forking 3-4 times, some arching strongly, others cruising to
the margins rather obliquely.

Discussion: The pinnules of Mariopteris muricata are

 

 

toothed, but this dentation presents a peculiar aspect in that it
is irregular. The typical l\_/I_. muricata pinnule will have two to
six teeth which are slightly developed except for the basal
pinnule which is relatively well-developed. The pinnules of
_l_\_/i_. muricata, as can be seen in pl. VI, fig. 2, are clearly
assymmetrical due to the uneven development of the teeth with
those below being better developed than the teeth above.
‘Mariopteris muricata is very similar to Mariopteris acuta

and Marippteris nervosa. Lutz (1938) described the fundamental

 

difference between _1\/_i_. muricata and_l\11_. _agpta is the stratigraphic
level in which one finds these plants. White (1899) proposed much
the same difference for M, muricata andA/i. nervosa. He confined
M. muricata to the Pottsville series, while M. nervosa characterized
the Allegheny series, rarely being found within and near the top of
the Pottsville series.

Kidston (1923-25) distinguishedM, nervosa from_l\i. muricata
by the mo re or less triangular or sub-triangular form of its
decurrent pinnules; by . the pinnules of M. nervosa being invariably

united and with the exception of the posterior basal pinnule and

occasionally the corresponding pinnule on the anterior side, they
rarely have lateral lobes.

The definition of Mariopteris nervosa as given by Danze-
Corsin (1953) illustrates the difference between it and _M_. muricata
as basically M, nervosa having entire margins and the margins of
1\_/1_. muricata being toothed.

Danze-Corsin (1953) separates l\_/1_. muricata from M. acuta

 

by the pinnules of l_\_/I_; muricata (1) being closer on the rachis;
(2) being broader and slightly contracted at the base; (3) having
slightly fewer teeth; (4) having more apparent venation, illustr-
ating clearly the finer secondary veins; (5) having its tertiary
pinnae slightly distant. The fourth difference however, is
dependent largely upon preservation. If the preservation Of both
is excellent, then the clearness of the venation may be a specific
difference. Kidston (1923-25) mentions the venation of_lyl_. muricata
as being immersed.
In general, l\_/I_. muricata is wider and thicker than _M_. w
The specimen illustrated on plate VII, fig. 6, was loaned to the
author by Dr. C. A. Arnold of the University of Michigan. This
specimen exhibits pinnules of the upper portion of a secondary
pinnae. The pinnules are more elongated with rounder apices than
those of the ultimate pinnae figured on plate VI, fig. 2 and 8. The

venation is the same for both specimens.

Fig.

Fig.

Fig.

Fig.

9A

9B

9C

9D

EXPLANATION FOR TEXT FIG. 9

 

 

 

Page
Crossopteris utahensis sp. nov. (5X) ----------- 125
Terminus of a penultimate pinnae.
Sphenppteridium zaitzeffium sp. nov. (5X) ----- 104
This species has no midvein. All veins arise
from the rachis and bifurcate several times be-
fore reaching the margin.
Odontopteris sp. (8X) ------------------------ 131
Neuropteris ampelinos sp. nov. -------------- 121

 

(4. 5X)

118

 

TEXT FIG. 9

Mario pteris muricata is essentially a Westphalian form, rarely
extending down into the Namurian.

Carpentier (1907) reports it from his 1B and 2A zones which
are Upper Numerian A. Crookall (1933) reports one specimen from
the Lanarkian of Great Britain.

Genus NEUROPTERIS Brongniart 1822

Neuropteris qiqantea Sternberg

pl. 1X, fig. 4
1821 Osmunda qiqantea Sternberg.-Versuch, v. i, fasc. ii, p. 33,
pl. xxxii
l8 Neuropteris qiqantea Sternberg.-ibid, v. i, fasc. iv, p. xvi.
1962 Neuropteris qiqantea Sternberg. - Tidwell, Brigham Young
Univ. Geol. Studies, v. 9 (2), p. 93, pl. 1, fig. 7.

Description: Specimen at hand fragmentary. Pinnules flat,

 

falcate, alternate, nearly right angles to the rachis, contiguous,
sessile, slightly cordate at base, rounded apex; 25 mm. long, 8 mm
wide. Strong mid-vein continuous to nearly two thirds pinnule length.
Lateral veins arise from midrib at a sharp angle and subdivide
several times to form very fine, very close veinlets.

Discussion: Neuropteris lunata White and_N_, gigantea are very
similar. _N. Ma, however, has more narrow pinnules than _l\_l,
qiqantea which are proportionately less acute. The curvature, when

present in _l\_l_. lunata, is more uniformly distributed, slightly

crescentic, through the whole length rather than being expressed
as an upward turn near the apex (Crookall, 1959).

The material from the Manning Canyon Shale applicable to the
species appear to have broader pinnules and more falcate shape
than pinnules of __N_. Ma.

Neuropteris qiqantea has been recorded from the Westphalian
A through D (Crookall, 1959). Gothan and Remy (1957) shows this
form to be distributed from Namurian C. to Westphalian D
definitely, and possibly into Namurian B.

Neuropteris heterophylla Brongniart

p1. VIII, fig. 2; p1. IX, fig. 3
1709 Lithosmunda minor Scheuchzer. — Herbarium Diluvianum,
p. 15, p1. iv. fig. 3.
1833 Neuropteris heterophylla Sternberg. -Versuch, v. i, fasc.
v-vi, p. 73.
1962 Neuropteris heterophylla. - Tidwell, Brigham Young Univ.
Geol. Studies, v. 9 (2), p. 93, pl. 1, figs. 3,8.

Description: Bi- to tripinnate. Pinnules alternate, oblique,
ovate to oblong, attached by single point, those on upper pinnae
odontopteroid, 15 mm. long, 7 mm. wide. Pinnules have a distinct
midrib with lateral vein distinct, close (35 to 40 per cm. ), arising

at an acute angle, oblique, strongly arched on some, divides two or

three times, contacts margins obliquely. Unipinnate near apex with
the terminal pinnule lanceolate or rhomboidal, 30 mm. long, 6 mm.
wide, sharply to bluntly pointed, slightly lobed on margins.
Discussion: _l_\l_. heterophylla and _N_. tenuifolia are similar.
_N__._ heterophylla differs from N. tenufolia in having stronger and
fewer lateral veins and in having pinnules shorter in relation to
width.
_N_, heterophvlla's stratigraphic range is from Westphalian
A through Westphalian D.
Neuropteris cf. pocahontas White
pl. VIII, figs. 4,6
1900 Neuropteris mcahontas, White. - U. S. Geol. Surv. 20th Ann.
Rept., p. 888, pl. CLXXXIX, figs. 4,49; pl. CXCI, figs. 5,5a.
Specimens attributable to this species are small and often in-
complete. They have the characteristic attachment and pinnule
shape of _l\_l. pocahoan and the Neuropteris manner of lobing. N.
pocahontas characterizes the lower Pottsville of the Appalachian
region.
Neuropteris ampelinos sp. nov.
pl. IX, fig. 2, text-fig. 9D
Description: Frond: bi-or tri (f?) pinnate. Pinnae: alternate,

right angles to rachis, but curving sharply upward, rachis flexuous,

no terminal pinnules present. Pinules: lanceolate, apex acute
to pointed, rarely obtuse, asymmetrical base with single point
attachment on the larger (mature) pinnules, pinnules nearer the
pinnae apex have 1/2 basal attachment. Pinnules 9-15 mm. long,
3 mm. wide, closely spaced, although gaps occur due to preservation,
sometimes contiguous or overlapping, oblique, alternate, no small
(young) pinnules present. Venation: midrib strong, raised, con-
tinuous from 3/4 to near apex, decurrent. Laterals: fine, close
(54 per cm. ), arise acutely, strike margins generally at an
oblique angle, sometimes right angles, after arching strongly and
forking 2-3 times. ’

Discussion: Neuropteris ampelinos differs from Crossopteris
utahensis in its finer venation, single point attachment and viney,

lax habit. _N. ampelinos and N, pocahontas are separated from one

 

another on the finer venation, one-half basal attachment of some of
its pinnules, and the more lax habit of _N_. ampelinos.
Genus CROSSOPTERIS Gen. nov.

Description: Tri- or quadripinnate forms with large triangular
to broadly lanceolate pinnae. Pinnules: irregular, polymorphous.
Small pinnules near pinnae apex decurrent with complete basal
attachment, upper margins may be constricted forming an asym-
metrical base. Upper margins in larger pinnules become constricted,

creating a one-half basal attachment which may or may not be

slightly decurrent and in some forms having single point attachment,
pinnatilobate pinnules have single point attachment. Lobing is
pec0pteroid-alethopteroid. Venation distinct with midrib in larger
pinnules continuous but not extending to apex of pinnule. Midrib
lacking or very indistinct in smaller pinnules. Secondary veins
arise acutely, arch strongly and strike margins obliquely, after
forking 2-3 times, rarely four. Subsidiary veins arise directly
from rachis in pinnule with partial or entire basal attachment.

Discussion: This genus is similar to Neuropteris in venation,
general pinnule shape, size and in nearly reaching single point
attachment; but differs from it on several points:

(1) The manner of lobing. In Crossopteris, the first larger
(mature) pinnules have undulating margins (text-fig. 10B). These
marginal indentations become deeper in each succeeding pinnule in
the direction of the pinnae-base, until small (young) pinnules are
distinct from the preceding lobes. This pecopteroid manner of
lobing is similar 'to Pecopteridium armosi and many Alethopteris
species such as, _A_. helenae LX and A, _gib_§p_r_1_i LX.

In Neuropteris, the first lobe forms at the base of the larger
(mature) pinnule, and a lobe or small pinnule is added to each
succeeding large pinnule in sequence towards the base of the

pinnae.

(2) The attachment of the small (young) pinnules. In Crossop-
t_e_r;_is they are strongly decurrent with contractions of their upper
margins near the pinnae apex (text. fig. 10A, 10D) whereas in
Neuropteris, particularly N. pocahontas, N. gpliqua and N.
schlehani these features are generally not as pronounced. N,
condrusiana Stockmans and Williere has small pinnules near
pinnae apex which are very similar to Crossopteris type; however,
its larger pinnules are likeN, m.

Crossopieris varies from Alethopteris, Pecopteris, and

 

Neuralethopteris on venation and manner of attachment. The mid-

 

vein is not continuous to the pinnule apex as in Alethopteris and
Neuralethopteris; and the secondary veins arch more strongly and

are more oblique than in Alethopteris (text fig. 10C). Although the

 

small pinnules have complete basal attachment (Alethopteroid) in
Crossopteris, the larger pinnules have either a single or nearly
a single point attachment rather than a persistent complete basal
attachment as in Alethopteris and Pecopteris.

The lanceolate or oblong, irregular, polymorphous pinnules of

Crossopteris vary from the relatively uniform Pecopteris pinnules

 

which have parallel or weakly convergent, lateral margins with
rounded, rarely pointed apices. The venation of Pecopteris is also

simple or grouped by twos, threes, rarely fours.

Odontopteris is also close, because the majority of the small
pinnules in Crossopteris have odontopteroid venation, but Crossop-
teris varies from Odontgiteris by its pecopteroid-alethopteroid
lobing.

Basically, therefore, Crossopteris is characterized by (1) its
pecopteroid-alethopteroid manner of lobing; (2) its complete
(alethopteroid) 1/2 basal or single point (neuropteroid) attachment
(text fig. 10C); and (3) its neuropteroid-odontopteroid venation.

This new genus was named in honor of Dr. Aureal T. Cross
of Michigan State University for all of the advice and assistance he
so willingly gave.

Crossopteris utahensis sp. nov.
pl. IX, figs. 1,6; pl. X: text figs. 9a,10.

Description: Frond: tri-perhaps quadripinnate. Antepenultimate
pinnae, large, triangular to broadly lanceolate, terminal portion
missing. Penultimate pinnae: alternate, linear-lanceolate to tri-
angular; mostly oblique, although on some specimens they appear
normal to the rachis; margins taper rapidly to form an acute apex,
close, contiguous, overlapping, some slightly distant. Ultimate
pinnae: lanceolate, tapering somewhat to base, pinnitilobate (undu-
lating margins), attached with single point, base assymmetrical,
although some are obtuse, alternate, both oblique and normal to the

rachis. Pinnules: generally small (5-30 mm. long, 3-8 mm. wide),

irregular polymorphous, alternate, rarely subalternate; oblique,
few at right angles; distant (2mm. ) with some close, contiguous or
overlapping, laterally unequal. Pinnules nearer apex of pinnae
elliptic, rarely ovate, decurrent with complete basal attachment,
upper margins constricted forming an asymmetrical base. The
upper margin becomes more constricted in each succeeding pinnule
in direction of pinnae-base until margin is constricted to near mid-
rib. Larger pinnules both oblong and lanceolate with obtuse to
acute apices, attachment one-half base, eventually acquiring single
point attachment in pinnatilobate pinnules.

Pecopteroid-alethopteroid manner of lobing. Venation: dis-
tinct, thin, regular. Midvein distinct in larger (mature) pinnules
but does not reach apex of pinnule, lacking or veryindistinct in
smaller (young) forms. Laterals arise acutely, arch and reach
margins obliquely (38 per cm.) after forking 2-3 times, rarely
four. Laterals strike the margins of the pinnatilobate forms at
nearly right angles. Subsidiary veins arise directly from the rachis.
Terminal pinnule: linear-lanceolate, laterally unequal, sublobate,
apex generally acute, rarely rounded.

Discussion: Crossopteris utahensis is the most abundant form
in the Manning Canyon Shale. This species is comparable in

abundance to Neuropteris mcohantas which characterize the Lower

127

Pottsville of the Appalachians and Neuropteris schlehani which
characterizes the lower Westphalian of Europe.
Neuropteris, Alettgpterisi Callipteris, Callipteridium,

Odontofieris, and perhaps Pecopteris are considered to have had

 

 

a common ancestor. This concept came from the observance of
the intermediate forms between these intergradational groups.

Neuropteris schlehani is an example of these intermediate forms.

 

It intergrades with Alethgiteris, and at times is distinguished

 

from Alethopteris lonchitica or A; decurrens only with difficulty.

 

A more primitive form 'of Crossopteris, perhaps the ancestor
to _C_. utahensis, may have been the ancestor for Neuropteris which
may'in turn, have given rise to) Callipteridium and Mixoneura
through the neurocallipterid: Neuropieris pocahontas. This
primitive form may have also given rise to Alethopteris through
theneuralethopterid form Neuropteris biformis LX or through a form
similar to Neuropteris schlehani.

By grouping the veins slightly, evolving more uniform pinnules
and achieving complete basal attachment, the primitive form of

Crossopteris may have become Pecopteris.

 

Jackson (1917) described. a form he related to Alethopteris sp.
from the Lower Pennsylvanian of Indiana. This form has an attach-
ment of its pinnules similar to the attachment of Crossopteris. Its

variation from _C_. utahensis is by its wider pinnules. It is repre-

Fig. 10A

Fig. 10B

Fig. 10C

Fig. 10D

EXPLANATION OF TEXT FIG. 10
Page

Crossopteris utahensis sp. nov. (7X) ---------- 125
This is a small pinnule from the upper portion

of a penultimate pinnae. Note complete basal
attachment with lower margin decurrent and the

odontopteroid venation.

Crossopteris u_t,ahensis sp. nov. (1X) ---------- 125
Penultimate pinnae illustrating the 1/2 basal
attachment becoming single point attachment in

the pennatilobate pinnules.

Crossopteris utahensis sp. nov. (5X)-----‘- ----- 125
Lower portion of pinnule showing the 1/2 basal

attachment and the neuropteroid venation.

Crossopteris utahensis sp. nov. (7. 5X) -------- 125
Small pinnule of an upper portion of a penultimate
pinnae or a small ultimate pinnae. This pinnule
would be situated below the small pinnule in text

fig. 10A.

128

 

 

 

 

D

TEXT Fm.

sented by only the upper portion of a pinnae. Its venation is
neuropteroid.

In the preliminary study of the Manning Canyon Shale flora,
several fra gmental specimens with decurrent pinnules were placed

with Alethopteris (Tidwell, 1962). These specimens (Alethopteris

 

sp. '8 B, C, and E) are now placed in the polymorphic form:

Crossopteris utahensis.

 

Crossopteris mcKniqhtii sp. nov.

pl. VIII, fig. 5
1962 Neuropteris sp. A cf N, _f_a_l_c__a_t_t_a, Tidwell, Brigham Young
Univ. Geol. Studies. v. 9 (2), pl. 1, figs 1,9.

Description: Frond: bipinnate, perhaps tripinnate. Penultimate
pinnae: large, linear-lanceolate. Ultimate pinnae; incomplete.
Pinnules: alternate, linear to linear lanceolate (ratio 6:1), rather
erect, distant, sessile, and decurrent small (young) pinnules, base
asymmetrical, some pinnules normal, others oblique to the rachis.
Larger pinnules with 1/2 basal attachment, becoming attached with
single point in the pinnatilobate forms. Apex pointed, although
roundly pointed on some. Smaller pinnules odontopteroid, tri-
angular in outline. Venation: distinct, midvein continuous to 5/6th
or to near apex and decurrent on the rachis. Laterals: thick,
close (30 per cm. ), arise acutely, arch rather strongly and meet

the margins obliquely.

Discussion: Crossopteris mcKnightii varies from Crossop-
teris utahensis by the shape and size ratio of its pinnules, the
more pointed shape of its apex, and its more strongly arched
venation. The author (Tidwell, 1962) had originally thought this
variety had affinities with Neuropteris foilcata. but Crossopteris
mcKnightii lacks the single point attachment except in pinnatilobe
pinnules.

The species was named for my good friend and colleague, Dr.
Kent McKnight of the Botany Dept. at Brigham Young University
who has contributed significantly to this study.

Crossopteris undulatus sp. nov.
p1. VIII, fig. 3

Description: Frond: at least bipinnate. Ultimate pinnae not
complete. Pinnules: alternate, oblique to normal to the rachis,
distant, oblong, acute to rounded apices, some pinnules curved,
upper and lower margins parallel, both constricted at the base to
form an asymmetrical base with single point attachment in the
larger pinnules, 18-25 mm. long, 5-8 mm. wide. Smaller pinnules
odontopteroid with complete basal attachment. Alethopteroid-
pecopteroid lobing. Venation: midrib broad, distinct, often in
relief, continuing to nearly 5/6th the pinnule length. Lateral veins

very thin, fine, close, numerous (56-64 per cm. ), arising at an

131

acute angle, arching strongly, forming 2-3 times, and striking
margins at nearly right angles. Subsidiary veins direct from rachis
in smaller pinnules.

Discussion: This form is closely related to S, utahensis in its
pinnule shape, size, small pinnule attachment and lobing but is
distinctive in its finer, more numerous venation and single point
attachment of the larger pinnules.

Genus ODONTOPIERIS Brongniart 1828

Odontopteris sp.

 

pl. VII, fig. 1; text fig. 9C

Description: Bi - or tri- (’2) pinnate. Ultimate pinnae linear,
tapering to a rounded apex, alternate, normal to the secondary
rachis, decurrent, rachis striated. Pinnules: broadly elliptic,
thickened in middle, with acute to obtuse apices, laterally unequal,
complete basal attachment, decurrent, upper margin slightly con-
tracted, distant, oblique, alternate, 6 mm. long, 3 mm. wide.
Venation: veins arising direct from rachis generally, midrib, if
present, obscure, veins arch and subdivide one to two times before
striking margins obliquely.

Discussion: This species is frequently encountered in collections
of the Manning Canyon Shale. Although other odontopteroid forms

are also encountered, they are not considered here because of their

132

similarity to odontopteroid pinnules attached to Crossopteris

 

utahensis.

This species is similar to Odontopteris peyerimhoff Bertrand,

 

but not sufficient material is available to be conclusive.
Order: CORDAITALES
Genus CORDAITES Unger 1850

Paleobatanists have long considered the name Cordaites for
leaves. Grand "Eury subdivided this genus on the basis of leaf
characters. Eu-cordaites contained leaves which were broad, with
rounded apex and with strong veins alternating with weaker. _Pga;
co rdaites had long, linear, grass-like leaves; and Dory-cordaites
included broad-lanceolate leaves with acute apices and rather fine,
equal veins.

Stockmans and Williere (1953) states that this system is not
useful in practice and that these divisions should therefore be re-
united with Cordaites, a concept with which the author agrees.
These divisions have been used as subgenera by some authors, but
in general have been largely ignored.

Cordaites principalis (Germar) Geinitz
p1. XI, fig. 5
1855 Cordaites prinpipalis (Germar) Geinitz. Vers. d. Steinhf. in

Sachsen p. 41, pl. XXI, figs. 1,2.

133

1949 Cordaites principalis (Germar) Geinitz. -Arnold, Univ. Mich.
Contr. Mus. Paleont. v. 7, p. 223, p1. XXX, figs. 1-3; pl. XXXI,
fig. 1. i

1962 Cordaites communis? 1x. -Tidwell, Brigham Young Univ.

 

Geol. Studies, v. 9 (2) p. 97, pl. 4, fig. 1,2.

Description: Leaves spatulate in outline, tapering to a thickened
apex, long, attaining a length of 22 mm. , and sometimes 20 mm.
wide, narrowed towards base. Apex rounded or obtuse (‘2).
Venation: irregular, nine veins per centimeter. Strong vein alter-
nating with from 3 to 5 weaker veins.

Discussion: The chief distinguishing feature of this species
consists of a number of strongly marked parallel veins running down
the leaf, between each pair of strong veins one to fiVe (usually three
to five) finer, parallel veins are found.

Cordaites principalis is a wide-ranging species from Namurien
B into Lower Permian.

Cordaites sp. A
pl. XI, figs. 1, 6

Incomplete specimens, long, linear leaves without either apex
or base. The venation is too indistinct for definite placement of
these forms into species, or for determining their relationships to

one another.

Cordaites sp. B
pl. XI, fig. 3

Incomplete leaves of broad spatulate shape without a definite
base or apex. Venation indistinct. Separated from the other
Cordaites species on shape which is similar to the shape for
Cordaites lingulatus Grand'Eury. The venation too incomplete to
make a positive identification.

Cordaites (‘2) sp. C
pl. XI, fig. 2

Description: Long, tapering to thin base, apex rounded, veins
of about the same strength, dichotomizing as they proceed toward
the apex.

Discussion: Cordaites (?) sp. C. rare in the Utah flora. This
species is placed provisionally with Cordaites, but may belong to
the genus Ginkophyton.

Genus CORDAIANTHUS Grand ‘Eury 1877
Cordaianthus pseudofluitans (?) Kidston
pl. XIII, fig. 7, text fig. 7A
1950 Cordaianthus pseudofluitans Kidston. -Florin, Acta Horti
Berg. , Bd. 15 (6) p. 112.
1951 Cordaianthus pseudofluitans Kidston. Florin, Acta Horti

Berg., Bd. 15 (11), pp. 307-9, figs. 15, 16.

135

Description: Cordaitean inflorescences with maximum width of
35 mm. Bracts alternate or subopposite, 45 to 90 degrees to axis,
20 mm. long, although not complete. Sterile scale leaves spirally
arranged, oppressed, imbricated, linear-lanceolate to lanceolate,
10 mm. long, 1-1. 5 mm. wide. Megasporophylls three to four per
strobilus, stalk-like, unbranched, 16 mm. long, bifurcated at the
distal end. _

Discussion: Cordaianthus pseudofluitans is considered by Florin
(1950) to be geologically older and more primitive than the older
forms of Cordaianthus. They have widely projecting elongated,
lateral fertile appendages which are repeatedly forked by "cruciate
dichotomy" and carry more than one ovule (or seed). Cordaianthus
zeilleri, Florin (1951) considers as the more modern cordaitean
fructification. It is characterized by very short, unbranched and
uniovulate megasporophylls, concealed among the sterile scales of
the flower axis. _C_, pitcairniae, _C_._ lindleyi and _C_._ longibracteatus
of Westphalian age are intermediate, resembling S, pseudofluitons
in their megasporophyll appearance which are only occasionally
bifurcated at their apices and are always uni-ovulate.

Florin (1951) gives a Westphalian age for Q, pseudofluitans.

SEEDS, CUPULES AND SPORANGIA

Genus CORDAICARPUS Geinitz 1862

13 6

The identification and classification of flattened, bilaterally
symmetrical seeds using as criteria the external shape of the nucellus
and the integuments surrounding it are subject to a high degree of
uncertainty. The integuments may be partially destroyed or flattened
or separated from each other or the nucellus during the process of
deposition and preservation. The surface of the seed or its integu-
ments may also acquire different textures, and the resulting seed
may assume an appearance quite different from other members of
its species or genus, making identification difficult.

Another difficulty arises from the lack of a uniform classification
system. In many of the older works, seeds are vaguely described
and inadequately figured, causing frequent misapplication of names.
Generic descriptions were often interpreted differently, and the pre-
carious application of names originally used to designate structuraly
preserved types for seed impressions has added to the confusion.

Brongniart (182 8) made the first attempt at classification by
giving the generic name Cordaicarpon to " Fruits comprimes,
lenticulaires, cordiformes ou reniformes termines par une pointe
peu aigue". This name was later changed to Cordaicarpus by
Geinitz (1862).

Goeppert (1864) used the term Cardiocarpon to include seeds of

Cardiocarpus and Samaropsis types. White (1899) used Cardiocarpon

and Samaropsis as synonyms or at least placed Samaropsis as a
subgenus of Cardiocarpon. He described seeds under Cardiocarpon
and Cordaicarpon using Brongniart’s Cardiocarpon in the synonymies
of both.

Kidston (1909) included Cardiocarpon and Cordaispermum under
Cordaic_a,rpus, and distinguished it from Samaropsis by the absence
of lateral wings and from Cardiocarpus by its more cordate outline.

Lesquereux (1880, 1884) Arber (1914) and Seward (1917) made
attempts at correcting this situation. »

Lesquereux (1880) used four generic divisions for seeds. These
were Cardiocarpus, Rhabdocarpus, 'i‘iigonocgpus and Carpolithes.
In 1884, he defined (_Jagdiocagus as " seeds of various shapes,
composed of a compressed, gen erally cordiform or oval acute or
acuminate nucelus surrounded by a flattened, fibrous border or a
membraneous wing and often narrowed at the base into a short pedicel
or acute appendage. " To Cordaicarpus he referred seeds of various
shapes and sizes which were "hypothecially considered until now
by their association to Cordaites as the fructifications of these plants. "

Lesquereux (1880) recognized a difference between Samaropsis

whose nucelus was superposed to or enclosed into a kind of samara

or thin membraneous scale; and Cardiocarpus with a " generally
narrower margin. " He noted transitional forms between these
t wo groups but included both under Cardiocarpus.

Arber (1914) defined Cardiocarpus as not-winged, and
referred all platysperms with a "triangular or heart-shaped
nucellus, surrounded by a more or less circular wing" to
Samaropsis. She restricted Samai'opsis to seeds which were

"not longer than broad or broader than long, " and created a new
genus Samarospermum for those forms having a very long and
narrow wing. I

Although Arber's classification simplifies generic identi-
fication, the relationship between the broad, membraneous
wing and a small fiberous coating should be more distinct.
Seward (1917) attempted to classify seeds by (l) restricting
Cardiocamug to petrified seeds exhibiting characters described
by Brongniart (188D. and Bertrand (1908) (2) including those seeds
with broader and more clearly defined border under Samaropsg,‘ ‘
(he also restricted the designation of Samaropsis to Paleozoic
seeds) and (3) placing platyspermic seeds, preserved as casts
or impressions with a comparatively narrow border enclosing an

ovate or co rdate—ovate nucellus and having a rounded or cordate

base, with Cordaicarpus. In his classification, the choice between
Cordaicarpus and Samaropsis is much the same as Lesquereux
(1880) proposed. Seward also mentions that seeds preserved with
a narrow border could be assigned only to Cordaicarpus, although
many are undoubtedly incomplete or immature. He also comments
that CordaicarpusL thought more suggestive of a cordaitean alliance,
may in some cases be a pteridosperm.

The problems encountered with Sewards classification are
(l) identification of the transitional forms between Samaropsis and
Cordaicarpus, and (2) the lack of constant characters by which to
distinguish Cardiocarpus from Cordaicarpus - i. e. , distinguish
a petrification from a cast or impression.

Seward's system has the advantages of (1) being more universal,
(2) separating petrifaction genera from impression genera, (3)
eliminating unnecessary generic names and (4) having more
definite descriptions.

The generic names used in the study follow Seward's classif-
ication system.

Co rdaicarpus elongatus (Newberry) comb. nov.
pl. XII figs 6,11, Table IV

1873 Cardiocarpon elorigatum Newberry, Geol. Surv. Ohio Paleont-

ology, v. 1, p. 373, pl. 43, fig. 5.

Description: Seed: 15 mm. long, 10 mm. wide, cuneate to
cordate with a slightly truncated base; widest portion of seed
across center, tapering to an acute apex. Wings thin around
base, thickening toward apex. Wings emarginate near micro-
pylar end, 3 mm. wide near micropyle. Micropyle faint.
Nucellus cordate, rounded base, acute apex.

Discussion: This species is similar to Q. cordatus but
differs from it in size and shape. 9, elongatus is widest near
the middle of the seed, whereas _C_, obtusum is widest near the
base.

Cordaicarpus sp. globosus sp. nov.
pl. XIIfig. 2, table IV

Description: Seed about 8 mm long, 7 mm. wide; cordate
in outline. Nucellus cordate with a rounded base and an acuminate
apex. Wings somewhat uniformly surrounding base and thickening
toward apex, emarginate near micropyle.

Discussion: This species differs from Cardiocarpon acuminatum

 

Jackson by having a more cordate outline and emarginate wings, and
from Cardiocarpus late-alatus Lx by having a smaller wing, a

broader nucellus, and a slightly smaller size.

Cordaicappus cordatus (Jackson) nov. comb.
pl. XII figs. 8, 9, 12, 13, 14; Table IV
1916 Cardiocarpon cordatum Jackson, Proc. Indiana Acad. Sci.
p1. X, fig. 10.
1916 Cardiocarpon obtusum Jackson, Proceedings of the Indiana
Acad. Sci., pl. X, figs. 13-14, p. 426.

Description: Small, cordate seeds with rounded base and
acute apex. 9-10 mm. long and 5-7 mm. wide. Nucellus cordate,
rounded base and acute apex. Micropyle faint. Wings thin around
base and expanding in direction of apex; emarginate near apex,

0. 1 mm. wide around base, becoming 2 mm. wide near apex.

 

Discussion: C. cordatus is one of the most abundant seeds
in the Manning Canyon Shale flora. This form varies from 5;
gr_'_a_c_:_i_l_e Jackson, which it closely resembles, by its smaller
size, more acute nucellus and more truncate base.

In examining the Jackson types, the author found no difference
between his S. cordatus and _C_. obtusium. The specimen in
plate XII, fig. 9, 13, appears to be the nucellus of C_3. cordatus
with the wings missing.

Cordaicarpus iayshuleri sp. nov.

pl. XII figs. 3, 4, 5 Table V

Description: Seed nearly round, although some specimens tend
toward a cordate shape; broad, rounded base, slightly acute apex.
10-14 mm. long, 7-12 mm. wide. No wings are present.

Discussion: This species is similar to Cardiocarpon
circulare Lx and Cardiocarpon subcirculare Jackson in shape
and size, but varies from both by not having a wing.

This species is named for my friend and colleague, James
B. Shuler of Greenville, South Carolina.

Cordaicarpus manningcanensis sp. nov.
pl. XIII fig. 2, Table V

1962 Cardiocarpon sp. A. , Tidwell. Brigham Young University
Geol. Studies, v. 9(2), p. 99, pl. 3, fig. 3.

Description: Seed oval outline, truncate to cordate at base,
8 mm. long, 7 mm. wide with greatest width near center.
Nucellus cordate, cordate at base, apex acute. Inner and outer
integuments appear to be present. Nucellus divided by a broad,
flattened ridge, extending from microphyle to base. Wings less
than 1 mm. at base, and about 2 mm. wide at apex, sharply
emarginate apex.

Discussion: This species is similar to Q. moreiranum White

 

in shape, but is much larger with narrower wings. This species

also lacks the acuminate apex which characterizes 9. minus Newberry.

Cordaicarpusbinutus sp. nov.

pl. XII fig. 3; table IV
1962 Cardiocarpon sp. B. , Tidwell. Brigham Young University
Geol. Studies, v. 9, (2) p. 99, pl. 3, fig. 2.

Description: Seed ovate to cordate outline. Size varies
slightly from 15 mm. long, 11 mm. wide to 10 mm. long, 10 mm.
wide for smaller specimens. Greatest width occurs slightly
below center on some specimens and nearer the base on others.
Base is cordate, although one specimen has an obtuse-rounded
shape. Two reniform depressions occur near the basal walls of
the nucellus. Wings are 0. 5 to 1 mm. wide near base and
expanding to l to 2 mm. at the apex. Nucellus cordate in outline,
pointed down.

Discussion: These specimens are separated from S. cordatum

 

by the larger size and the two depressions near the base.
Genus TRIGONOCARPUS Brongniart 1828
The genus Trigonocarpus was established by Brongniart (1828)
to include radially symmetrical seeds characterized by three
longitudinal ribs. Hoskins and Cross (1946) restricted this genus to
seeds known from external characteristics. This revision was

necessary to avoid the conflict between seed compressions and

TAB LE IV

 

 

 

 

 

 

 

 

 

 

 

 

 

 

cordatus qlobosus domains binutus
Name (Jackson) sp. nov. (Newberry) sp. nov.
.comb. nov. comb. nov.
. (2X) (1x1 (1. 5X) (2X)
Shape Ovate to Cordate Cuneate- Ovate to
cordate cordate cordate
Base Rounded Rounded Semi- Cordate
truncated
Apex Acute Acute Acute
Size:
Length 8 mm 8 mm 15 mm 10-15 mm.
Width 6 mm 7 mm 10 mm 10-11 mm
Nucellar Co rdate Cordate Ovate— Co rdate
Shape Cordate
Base Rounded Rounded Truncated
,Apex Acuminate Acuminate Acute Pointed
Wings Expanded Expanded Expanded Expanded
toward apex toward apex toward apex toward apex
Shape Emarginate at Emarginate Emarginate
apex. Thin at apex
around base
Size Base:0. 5 mm Base: 0. 5 mm Base: 0 Base:0. 5-1 mm
Apex:l mm 1 Apex: 1 mm Apex: 2 mm Apex: 2 mm
No line as ' two reniform
Jackson depressions
Misc. Described NC NC near base
Small depress
ion near base
Stalked NC NC NC NC
Micropule Faint Present Present NC

 

 

 

 

 

143A

 

144

impressions and those seeds which are preserved as petrifactions.

Stockmans and Williere (1953) mention the difficulty in making
specific determinations in Triqonocarpus-type seeds. They considered
the arbitrary system of measurements as the best way of placing
these seeds into species. It would seem, however, that these
seed forms would be too intergradational as to size for this system
to be of value.

Triqinocarpus noeggerathi (Stnbg.) Brongniart
p1. XIII fig. 14

1880 Trigonocarpus noeggerathi Brgnt. -Lesquereux, Coal Flora,
2nd Geol. Surv. Penn. Rept. P, p. 584; pl. LXXXV fig. 1.
1949 "Trigonocarpus" noeggerathi (Stnbg.) Brgnt. -Arnold, Univ.
Mich. Contr. Mus. Vol. VII, p. 214, pl. XXIX, fig. 3.

Description: Seed elliptical with one longitudinal ridge

 

visible. Micropylar portion of the specimen is missing. The
specimen measures 1. 4 cm. wide, and at least 2. 5 cm. long. The
remainder of its exposed surface is smooth.

Discussion: This specimen matches exactly a specimen from

 

the Michigan Coal Basin identified to this species by Dr. C. A.
Arnold.
Arber (1914) placed this species in a new genus Schizospermum.

This is based upcn the appearance of three valves at one end of a

cast and a foramen or opening at the other. Neither of these
features are observable in the specimen under consideration.

Stockmans and Williere (1955) found Trigonocappus specimens
resembling ’_I‘_,_ noeggerathi T. parkinsoni Brgnt; and '_I‘__:_ schultzianus
Goepp, in the Zone de Bioul in the lowermost Namurian A of
Belgium. They state it would be dangerous for them to attribute
them to these species, because this would imply they were known.
The reason they hesitate to place them with these species is
that the plants which the seeds have been associated with are not
present in the beds from which the seeds were collected.

Trigonocarpus sp. A.
pl. XIIIfig. 11

Only one specimen of this type has been found. The seed is
compressed and the surface shows three longitudinal ridges,
although its probable number is six. These ridges are 1 mm wide
and divide the visible surface into four equal parts and converge

t oward the micropylar end. The surface between the ridges is
smooth. The specimen is 1. 5 cm. wide and 3 cm. long. It appears
to be surrounded by a marginal band. The micropylar type is missing.
Trigonocarpus sp. B

p1. x111 fig. 10

Smooth scleratesta and a portion of the micropyle of
Trigonocarpus forma B described by Arber (1914). The sarcotesta
can be observed as a thin film partially imbedded in the matrix
surrounding the sclerotesta. The specimen here described is
about twice the size of _T‘i‘iqonocarpus parkinsoni as illustrated
by Arber (pl. VI , FIG. 2). The micropyle of Trionocarpus
parkinsoni is much langer and thinner than for the Manning Canyon
Shale specimen.

This form is also similar in size and shape to Triponocarpus
ampuilaeforma LX.

Trigonocai'pus sp. C
pl. XIII figs. 9, 13

Seed, 3.4 cm long, 1. 5 - 2. 5 cm. wide, flask shape, micropyle
short, apex truncated. One specimen with 4 longitudinal ridges
visible. Preservation is poor as to internal features. This species
is similar to Trigonocarpus giavatus (Sternberg), but appears to be
wider and generally larger than the latter species.

Genus HOLCOSPERMUM Nathorst
Holcospermum sp.
p1. XIII, fig. 8
1962 (‘2) Cardiocarpus sp. , Tidwell - Brigham Young University

Geol. Studies, V. 9 (2), p. 102, pl. 1, fig. 4.

Deggi‘iption: Seed: elongately elliptical to oblong, 10-20 mm.
long, 5-7 mm. wide, broadest part occurs about middle, base and
apex rounded. Seed bears two prominent longitudinal ribs which
gradually approach each other at base and apex. Fine striations
or grooves appear between the ridges, traversing the length of
the seed.

Genus CORNUCARPUS Arber 1914

Arber (1914) originally included in this genus small seeds,
not winged, triangular shaped, with a short stalk at one end and
two well-marked, projecting horns from the other. Halle (1927) redefined
the genus for "platyspermous seeds, with or without wings, provided
at the apex with two acute projecting horns, which are at least
as long as they are broad and more often longer. " I

The definition should perhaps be emended slightly to include
all seeds with prominent horn-like or whip-like projections from
their micropylar end.

Cornucarpus discissus sp. nov.
pl. XIII, fig. 1; table V

1962 Cornucarpus sp. A. Tidwell. -Brigham Young University Geol.

 

Studies v. 9 (2), p. 100, pl. 3, fig. 4.
Description: Broad oval outline, base slightly rounded and

pointed. 10 mm. long, 8 mm. wide with greatest width near center

Nucellus oblanceolate, obtuse base tapering to a cut apex. Micropyle
extends into upper end of nucellus. Wings are 1 mm. wide near
base, increased to 1. 5 mm. near apex and. extending beyond into long
narrow "whip-like" projections. These projections are about 11 mm.
long and bifurcate near their tip into two dichotomies which curl

back on themselves. Stalked base.

Discussion: This species unlike any species attributed to this

 

genus. They differ from others by the long "whip-like" projections
which often recurve on themselves. The purpose these projections
may have served is uncertain.

Genus RIGBYOCARPUS Gen. nov.

Rigbyocagpus ebracteatus sp. nov.

pl. XIII, figs. 5,6; text fig. 7B tableV

Description: seed; small, 8 mm. long, 2 mm. wide at the base

 

somewhat "jug-shaped", elongated, ribbed, apex cut. Bracts beginning
near base of seed, reaching up around seed. Bracts are 10 mm.

long. , tapering to a pointed apex. Small stalk at base which may

have been point of attachment.

Discussion: This seed is only found isolated. The bracts

 

(pl. XIII fig. 6) have been uncovered with the seed portion missing.
The bracts are generally complete and attached to one another

which would indicate the seed was not lost due to excessive moveme nt.

The mature seed may have been shed enclosed within the bracts,
but later broke away.

This genus was named for Dr. J. Keith Rigby of Brigham
Young University, Provo, Utah; in recognition of his friendship,
interest and aid in this study. '

Genus LAGENOSPERMUM Nathorst 1914

Nathorst (1914) introduced Lagenospermum for small, fusiform
or elongated seeds, characterized by being completely enclosed
within a cupule, and having six (‘2) conspicuous, longitudinal ribs
or ridges. Two species, L_agenospermum sinclairi Arber and
_L., _a_r_'_be_r_'i_ Nathorst, are typified by a long stalk terminated with
a cupule.

Lagenospermum bifurcatus (Stopes) comb. nov.
1914 Pterispermostrobus bifurcatus Stopes, -Geological Survey
of Canada Memoir 41, p.p. 74-77, p1. XVII, fig. 45; pl. XXV,
fig. 69 and text-fig. 15.
1962 Pterispermostrobus sp. A-Tidwell-Brigham Young University
Geol. Studies v. 9 (2), p. 100, pl. 4, fig. 9.

Description: Seeds ribbed with oblong or lanceolate outline

 

rounded base and acute apex, 6 mm long, 3. 5 mm. wide. The

cupule which contained the seed is often preserved. The seed

TABLE V

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Cordaicarpus Cordaicarpus Cornucarpus Egbyocarpus
Name Manninpcane- jayshuleri discissus ebracteatus
§i_s_ (A) sp. nov. sp. nov. gen & sp.
nov.
. <
m (o) <
_ (2X)“ (2X) (3x)
Shape Oval Cordate Broad oval Elongate
bottle shape
Base Truncate- rounded slightly
co rdate rounded
Apex Acute Acute pointed Acute
Size:
Length 8 mm 12 mm 10 mm 8 mm
Width 7 mm 10 mm 8 mm 2 mm
' Nucellus
Shape Cordate Oblanceolate
Base Cordate NC Obtuse NC
Apex Acute Cut
Wings Widening to- Expands into
ward apex narrow whip-
slightly NC like horns NC
Size Emarginate Base: 1 mm
Base: 1 mm Apex: l. 5. mm
Apex: 2 mm
Divided by Horns: 11 mm Seed: ribbed and
Misc. broad, flattened long. surrounded by
ridge NC Bifurcate and long bracts.
recurved Bracts 10 mm
' long.
Stplked NC NC Present Present
Micro pyle Pres ent Faint Present NC

 

 

 

 

149A

forms a depression or mold in the cupule 4 mm. long and 2 mm
wide. The cupule has 5-6 "teeth" or lobes projecting toward
apex. These often converge to form an acute apex. Each cupule
terminates an equally dichotomized branch.

Discussion: This species from the Manning Canyon Shale
resembles closely Pterismrmosprpbus bifurcatus from the Fern
Ledges of New Brunswick. The specimen described by Stopes
(1914) was only a carbonized film in which structural details
are indiscernible. It could not be determined whether the film was a
fructification or a seed or a spo rangium. Therefore, she created
the new genus Pterispermostrobus for fructifications of this nature.

The specimens here described appear to be closely comparable
to Stope's specimen. The Manning Canyon Shale forms contain
"ribbed" seeds and therefore should be placed with the genus
Lagenospermum.

The Utah specimens are smaller than Laqenospermum
imparirameum, described by Arnold (1939) for fossils from the
Pocono Formation, and their bracts or lobes are not as spread
out. They also differ in the manner in which the stalks bifurcate.
The stalks of I._,_ bifurcatus generally divide equally, whereas in

_I___.._ imparirameum and _I_.,, sinclairi Aber, they are unequally

 

divided.

Nathorst established the genus Lagenospermum for fructifications
resembling Laqenostoma, but lacking internal structure. Arnold
(1939) suggests that the affinities of this genus are apparently
with the lyginopterid pteridosperms. Carpentier (1925) found
forms similar to L. kidstoni with Sphenopteris striata. Lobes on
ii, kidstoni are rounder. Its tests is smooth with only slight
longitudinal ridges.
Genus GNETOPSlS Renault and Zeiller 1884

Gentppsis was originally proposed for small petrified seeds
and impressions which Renault and Zeiller believed belonged to
some Cretaceous plant. The early paleobotanists used Gnetopsis
to indicate an affinity between these Paleozoic seeds and the
Gnetales. Depape and Carpentier (1913) state that if this relationship
actually existed, the cup-shaped organ containing several seeds
would have been equivalent to two capillary leaves. They suggest
the existence of several important characters which Gnetopsis
eliptica Oliver and Salisbury has in common with Lagenostomales.
No affinities between Gnetopsis and the gentaleans have actually
been established.

Seward (1917) prOposed that Gnetopsis is a transition form
between the Radiospermae and the Platyspermae. He does not

explain the reason for this proposed relationship. Gnetopsis is

a pteridosperm, but nothing is known of its vegetative parts.
Gnetopsis anglica Kidston
pl. XIII, fig. 4

1917 Gnetopsis anglica Kidston, Seward, Fossil Plants, p. 318,

 

 

figs. 494 E, F, G,H.

Description: Seed oval, elongate, with terminal'appendages"
attached to the micropylar end. These terminal appendages
appear to be partially united on some species or divided into four
components in others. No nucellus or other internal structure
is discernible. The length of these small seeds is 4 mm. Their
maximum width through the median area is 2 mm. Their
"appendages: are 25 mm. or more in length.

Discussion: The Utah specimens correspond closely to

 

Seward's (1917) description and illustrations. He cites the size
of Gnetopsis apglica as being 4 mm. in length with appendages
extending to 3.2 cm. The appendages do not show the hair-like
structure which are prominent on the French specimens.

The specimens figured and described by Stockmans and
Williere (1953) are similar to the Manning Canyon Shale specimens
in size and shape, but vary from them by having the hair-er
attachment to the appendages and a longitudinal ridge traversing

the seed body. They remark on the form described as g. hexagona

153

by Depape & Carpentier (1913) which had six distinctive ridges as
having been transferred into g. anglica by Seward (1917). They)
stated that g. anglica must vary according to the preservation of
its envelope.

Depape and Carpentier (1913) reported their form (9, hexagona)
from the Stephanian of Commentry. Stockman and Willere (1953)
report their specimens of g. apgNga from Namurian B.

Genus TELANGIUM Benson 1904

Stur (1875-77), Kidston (1887) and others described under the
name Calvmmathotheca, impressions which they considered to
be sporangia. Because Calymmathotheca had been originally applied
to fossil seeds, Benson (1904) concluded that sporangia should not
be included in it. Therefore, she proposed the name Telangium
for certain petrified synangia from the lower Coal Measures. She
considered Telangium to be applicable to microsporangia Of a
pteridoseprm, possibly quincmteris.

Seward (1917) noted the similarity of Telangium and Crossotheca
, and recalls Scott's (1908) interpretation that Telangium cannot
be generically separated from Crossotheca. Kidston (1906) regarded

Telangium scotti Benson as a pteridosperm microsporangiate

 

structure, but not related to Lyginopteris. He differentiated

Telangium from Crossotheca on the basis of the presence of a

 

 

single locule in Telangium as compared to the double locule of
Crossotheca. Later he also noted (1923) that the microsporangia

of Telangium stand upright on the rachis Whereas in Crossotheca

 

 

they are attached below.

Walton (1931) deplored the practice of including impressions as
well as petrified structureless compressions in the genus Telangium
as well and the petrifactions showing internal structure only, for
which the genus had been originally proposed. He expressed the
opinion that a new genus should be created for impressions and
structureless compressions, especially for the specimen of a
fructification recorded by Carpentier for Sphenopteris striata.

Telangium affine (I... & H.) Benson

pl. XII, figs. 1, 10, 16
1832 Sphenopteris affinis Lindley and Hutton, " Fossil Flora"
Vol. 1; p1. XLV. .
1887 (Ely-Inmatpiheca affinis (L. &H. ) Kidston, Trans. Roy. Soc.
Edinburgh, Vol. EQCXII: p. 145, pl. 1X, figs. 18-22.
1904 Telangium affine (L. &H.) Benson, Ann. of Bot. Vol. XVIII:
p. 164; pl. XI, fig. 12, text-fig. 33.

Description: Rachis dichotomizes several times, terminating
in two short dichotomies and the resulting branches are terminated

with small upright synangium. The synangia are 3 mm. long and

155

about 4 mm. wide. The synangia are united to one another at their
bases and form a cup-shaped synangium. The lower portion of this
synangium appears to be solid and merges into the supporting rachis.
Six sporangia seem to form a synangium. Each sporangia is 3 mm.
long and 0. 5 mm. wide.

Discussion: Telangium affine is known from the lower Carbon-
iferous of Europe. Bell (1938) described some sterile foliage
from Canso Group which he attributed to this Species.

Genus AULACOTHECA Halle 1933

This name was proposed for elongated, cylindrical or narrowing
club-shaped, seed-like, spore-bearing bodies with marked longitudinal
furrows and ridges. The spores from Aulacotheca are of the
Whittleseya type

Hemingway (1941) reports that these spores are large, smooth,
oval bodies with a longitudinal germinal slit which vary in size from
100 to 300 U, depending upon the species represented. These spores
are larger than those previously reported for pteridosperms which
rarely exceed 50 U in diameter. Hemingway therefore proposed
that perhaps two distinctive families of pteridosperms are represented.

The type of pteridosperm to which Aulacotheca belongs has
not been decided. Hemingway (1941) reported that it was invariably

found associated with the fronds of Alethopteris, usually of the

 

A; lonchitica group. Dix (1932) recorded a form of this type,
later named Aulacotheca dixiana by Hemingway, as being attached
to Neuropteris schlehani. Hemingway (1941) thinks that it is not
attached, but rather the foliage and carpons in closely falling
together, have simulated organic connection. However, this

association supports the concept that Aulacotheca is the micro-

 

sporangiate organ of certain members of the Medullosaceae.
Aulacotheca has not been found closely associated with any
particular foliage type in the Manning Canyon Shale. Alethofieris

lonchitica and definite Neuropteris schlehani have not been

 

uncovered as yet. Therefore, Aulacotheca may have also occurred
on other pteridosperms.
On the stratigraphic range of this genus, Arnold states that

"Since Aulacotheca extends throughout the entire vertical extent

 

of the Pottsville group in the Appalachian region, its use as a
horizon marker within the Pottsville is limited, although White‘s
comment that the sporocarps in the higher beds usually are larger
than those found at lower levels may be based upon genuine
dflierence."
Aulacotheca campbelli (White) Halle
pl. XIII fig. 12

1900 Whittlesefi Campbelli White 20th Ann. Report U. S. G. S. ,

 

p. 905, p1. cxc, figs. 9-11

1933 Aulacotheca Campbelli (White) Halle K. Svenska Vet. Akad.
Hand. p. 20, 39.

1949 Aulacotheca Campbelli (White) Halle, Arnold. Cont. Mus.
Paleo, Univ. of Mich. p. 207, pl. XXIV, figs. 4-6.

Description: Compression, elongated, companulate male-

 

spore bearing structure, terminal position broad with parallel
margins, eventually tapering to a pointed attachment. "Tubes"
long and narrow and overlap because of compression. Six lobes
or "apices at the terminal end give the appearance of six locules.
No spores observed.

Discussion: Halle (1933) mentioned that he was certain
Whittlesefl campbelli White from the Pottsville Formation of the
Anthracite Coal Basin was attributable to this genus, although he
had not seen any specimens. Arnold (1949) makes definite comp-
arisons for material from Michigan and also discusses the variation
in size of this species. He states that the smaller sporocarp
generally occur in the lower horizons of the Pottsville, whereas the
larger forms are found nearer the top.

J ongmans (1937) listed Aulacotheca from two localities in the
Pocahontas coal series and one from the New River Series.

Aulacotheca Hemingwayi Halle

p1. Xn, fig. 7

1933 Aulacotheca heminqwayi Halle, K. Svenka Vet. Akad. Handl.
Vol. XII(6), p. 36, pl. 8, figs. 1-16.

1941 Aulacotheca hemingwayi Halle, Hemingway, Ann. of Bot.

N. S. , vol. V, p. 198, pl. V, figs. 1-9.

Description: Small (18-25 mm. long, 4-4.5 mm. wide near
apex), microsporangiate structure with 9(?) locules. Surface
longitudinally striated. General shape oblong, broad at the apex,
tapering toward a narrow base. Apex dentate. No spores observed.

Discussion: The Aulacotheca specimens from Utah referred

 

provisionally to A. hemingwayi were placed with this species

because of their size and shape. The size range of the Utah specimens
is close to the range outlined for A, hemingwapi by Arnold (1941).

He gives its size as 4. 5-5. 5 mm X 28—30 mm.

A, hemingwayi is separated from A, elongata (Kidston) Halle
on size and its more prominent ridges. The Utah specimens have a
more truncated apex.

J ongmans (1937) reports A. heminpwayi from Pocahontas coal
seams 1-3 which he places in Namurian B. He states that from White's
figures, he supposed A, campbellis should be placed with A, hemingwayi
with White's specific name having priority. Darrah in a sub-note
to J ongman's paper thinks they are distinct because A, hemingwaj'g

has " rather broader aspect and a somewhat different ornamentation. "

REFERENCES

Abbott, Maxine L. , 1958. The American species of Asterophyllitcfi,

 

Annularia, and Sphenomyllum: Bull. American Paleont.

 

v. 38, no. 174, pp. 289-388.

, 1963. Lycopod fructifications from the Upper Freeport
(No. 7) coal in southeastern Ohio: Palaeontographica,

v. 112 B, pp. 93-118, pls. 25-32.

Ancion, Ch. and vanLeckwij ck, W. , 1947. Contribution a l'etude
de la stratigraphie du bassin d'Andenne; niveaux Greseux
et horizons marins du namurien: Soc. Geol. Belgique Ann.
t. 70, pp. B 266-B 306, lpl.

Arber, E. A. N. , 1914. A revision of the seed impressions of the
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pp. 592-608.

Fig. 1

Fig. 2

EXPLANATION OF PLATE I

North View of Clay pit A on Lake Mountain. Plant
horizon occurs in the shales along the eastern face

of the pit. (right side).

Clay pit C looking northwest. Plants were collected

from southwestern face of the pit. (left side).

175

 

PLATE I

 

 

Fig. 1

Fig. 2

EXPLANATION OF PLATE II

View looking east at the type section of the Manning
Canyon Shale in Soldier Canyon. The median lime-
stone appears to the right in the figure beneath the

lone tree on the lower skyline.

A View of the northern end of Clay Pit A. The plant
horizon is situated in the ledges to the right in the

illustration.

177

 

PLATE II

EXPLANATION OF PLATE III

 

 

 

Page
Lepidodendron volkmannianum Sternberg 53
fig. 1 Specimen showing the characteristic horizontal
and vertical alignment of the leaf bolsters (1X)
Lepidostrobus skemmatos sp. nov. 59
fig. 2 Strobilus showing its compacted sporophst and
its general shape (pointed apex) incomplete (IX)
Lepidostrobus variabilis L. & H. 57
fig. 3 Strobilus demonstrating its oblong shape, its
axis and crowded sporophylls. Note the quartz
box-work near its apex. (1X)
Lepidophyllum sp. 65

 

fig. 4 Sterile leaf; midvein and stomatal grooves are visible.
The base of the leaf is at the base of the illustration,
tapering to a pointed apex (2X)
Lepidodendron obovatum Sternberg 51
fig. 5 Leaf bolsters on a stem. Note broad-as-long size
of bolsters with elliptical leaf scar in upper one-
half of bolster. A bolster near base shows the
bascular and parichnos scars (1X)
Lepidostrobophyllum majus 63
Fig. 6 Broad, lanceolate lamina of sporophyll with
fertile portion missing (1X)
Lepidodendron aculteatum Sternberg 52

fig. 7 Bolsters with leaf scar Visible. Ligule scar is

I_J__°-_ LI I1 I flu! « n

. .. .
31¢. 11.x:

 

in! O
\Itfi‘yunlv

§
v .
tn:-

1‘.

 

PLATE III

EXPLANATION OF PLATE IV

 

 

 

 

Page
Asterophyllites equisetiformis (Schloth.) Brongniart 79
fig. 1 Branches with whorls of leaves attached.
Asterophyllites charaeformis (Stnbg.) Goeppert 80
fig. 2 Branch with whorled leaves (1X)
Calamites (Mesocalamites) cistiiformis Stur 72
fig. 3 Stem with two nodes and part of another showing
straightness of ribs and how some ribs alternate
at the node while others pass through the nodes,
particularly at the middle node (1X)
Asterophyllites longifolius (Stnbg.) Brongniart 81
fig. 4 Small stem with leaves attached at nodes. Note
how leaves overlap the node above (1X)
Lepidophyllum longifolium Brongniart 64

 

fig. 5 Basal portion of leav illustrating the central
midvein and the two stomatal grooves on each side (1X)
Palaeostachya sp. 82
fig. 6 Portion of stem, branching at each node. The nodes
of the branches support the calamitean cones. The
dark areas near the cone bases are the fertile part,
whereas the lighter areas near the apex are the
sterile portions (1X)

Calamites (Stylocalamites) sp. 74

 

fig. 8 Stem with two nodes (1X)

tr .1,
153

 

. \

.w.

.7
huh

hum
,R

I,
..

"an 6,1.

 

‘3
r
at

21

 

PLATE IV

EXPLANATION OF PLATE V

 

Page

Calamites(Mesocalamitefi) cistiiformis Stur 72
fig. 1 (1X)

Calamites( Mesocalamites) hesperius Arnold 71

 

fig. 5 Two nodes of a stem showing the broad, straight ribs,
some of which alternate at nodes while others go
directly across (1X)
fig. 2 An enlargement of fig. 5 showing the alternation
between some ribs continuing straight through the
nodes and others which alternate at the nodes (1. 5X)
Lepidocarpon linearifolium (Lx) Schopf 61
fig. 3 Specimen of the megasporangium with the sporophyll
still attached (1X) '
Calamites(Calamitina?) sp. 74
fig. 4 Specimen with the branch scars distinct at two nodes.
The leaf scars which occur on the nodes below the
branch scars are not readily observable.
Calamostachps (‘2) sp. 83
fig. 6 The strobilus is associated with Asterophyllites
equisetiformis. (1X)
Archeocalamites radiatus (Brgnt.) Stur 66
fig. 7 Stem with the bifurcating leaves arising from each node.
The straight ribs are Visible at upper portion of the
stem, but the non-alternation of these ribs at the nodes

be seen here. (1X)

18

 

 

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.. 0.55.0.1. Iain“ 1

 

PLATE V

EXPLANATION OF PLATE V Cont.
Asterophyllites equisetiformis (Schloth.) Brongniart 79
fig. 8 Stem with branches arising from each node. The
leaves are whorled at each node with internodal
area becoming shorter distally eventually
achieving the "bottle brush" affect near the tips

of the branches. (1X)

183A

EXPLANATION OF PLATE VI
Page

Zeilleria sp. 100
fig. 4 Upper penultimate pinnae showing fructification (1X)
fig. 1 Enlargement of fig. 4 (2X)

Mario pte ris muricata Schloth. 114

 

fig. 2 Fragment of ultimate pinnae (1X)
fig. 8 Enlargement of fig. 2 (s.5X)
Tingiasp. 84
fig. 3 Upper pinnules from an antepenultirnate or a
penultimate segment. The deeply dissected lobing
can be observed in pinnules on right side of rachis (1X)

Adiantites sp. 106

 

fig. 5 Isolated pinnule illustrating typical Adiantites
venation (1X)
Sphenopteris ppthanica (‘2) (Dolianitii) comb. nov. 88
fig. 6 Antepenultimate segment. Note that two veinlets
per lobe are distinctly Visible.
Diplothmema arnoldi Stockmans & Williere 111

fig. 7 Pinnules and ultirrate pinnae (1X)

, 135

 

PLATE VI

EXPLANATION OF PLATE VII

 

Page
Diplothmema trifoliata (Artis) White 109
fig. 1 Specimen showing several ultimate pinnae (1X)
Sphenopteridium dissectum Goeppert 102

 

fig. 2 Specimen illustrating general pinnule shape and
venation. Note veins arise direct from rachis. No
central midvein is present. (1X)

Alloiopte ris cruciatus s p. nov. 94

 

fig. 3 General pinnule outline and venation are observable
on this specimen. Note the uniform size of the
pinnules and their assymmetrical shape (1X)

fig. 7 Specimen with only the venation visible. The outline
of the pinnules on specimen are generally lacking (2X)

Sphenopteris (?Renau1tia) schatzlarensis (Stur) Zeiller 87

 

fig. 4 This species is illustrated in the lower right portion
of this figure. Asterophyllites equisetiformis
occupies the remainder of the figure (1X)
Corynepteris angustissima (Stbg) Nemejc 92
fig. 5 Specimen showing the toothed aspect of the species (2X)

Mario pteris muricata Scloth 114

 

fig. 6 Pinnules from near apex of penultimate pinnae (1X)

187

 

PLATE VII

EXPLANATION OF PLATE VII Cont.

 

Rhodea vespertina Read . 99
fig. 8 (1X)
Rhodea sp. 99

fig. 9 Elongated segments and pinnules distinctly
showing venation (3X)
Diplothmema obtusiloba (Brgnt. ) White 110
Fig. 10 Portion of an antepenultimate pinnae (1X)
Adiantites Wardia tenuifolius var. Difoliolatus (White comb. nov.
fig. 11. Penultimate segment of a frond. (1X) 107
Sphenopteridium Zaitzeffium sp. nov. 104

fig. 12 Antepenultimate segment. Notice the flabelliform

lax pinnules. (1X)

187A

EXPLANATION OF PLATE VIII

Odo ntopte ris sp.

 

fig. 1 Ultimate pirmae (2X)
Neuropteris heterophylla Brongniart
fig. 2 Terminal pinnule (1X)

Crossopteris undulatus sp. nov.

 

fig. 3 Penultimate segment (1X)

Neuropteris cf. pocahontas White

 

 

fig. 6 Penultimate pinnae (1X)
fig. 4 Enlargement of fig. 6 showing the single point
attachment (1. 5X)

Crossopteris mckniqhtii gen. & sp. nov.

 

fig. 5 Antepenultimate of penultimate pinnae
One-half basal attachment, alethopteroid-
pecopteroid lobing and neuropteroid venation

can be seen. (1X)

//-Q '3 //189

Page

131

120

130

121

129

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PLATE VIII

EXPLANATION OF PLATE 1X
Crossopteris utahensis gen. & sp. nov. 125
fig. 1, 6 Penultimate segments illustrating one-half
basal attachment, alethopteroid-pecopteroid
lobing and odontopteroid-neuropteroid venation
(1X)
Neuropteris ameplinos sp. nov. 121
fig. 2 Penultimate segment. (1X)
Ngmropteris hetergphylla Brongniart 120
fig. 3 Fragment of penultimate or ultimate pinnae (1X)
Neuropteris qiptantea Sternberg 119
fig. 4 Fragment showing four falcate pinnules (1X)
Stipmaria ficoides (Stnbg.) Brongniart 65

fig. 5 Portion of rhizome with rootlets attached (1X)

191

 

PLATE IX

EXPLANATION OF PLATE X

Crossopteris utahensis gen. sp. nov. 125
An antepenultimate frond illustrating the various characters of
the genus. The polymorphous aspect of this species can be seen

by comparing the pinnules from various pinnae. The terminous

of this frond is missing. (X 1/2)

193

X HIV'Id

 

EXPLANATION OF PLATE XI

Co rdaites sp. A 133

 

figs. 1, 6 Portion of leaves illustrating the poor
preservation. (1X)
Cordaites (2) sp. C 134
fig. 2 Long, linear leaf with rounded apex (1X)

Co rdaites sp. B 134

 

fig. 3 Leaf, illustrating the broad, spatulate shape (1X)
Lepidodendron obovatum Sternberg 51
fig. 4 Branched stem, showing leaf scars on left
branch and cluster of leaves on tip of right
branch. These leaves are possibly the lower
portion of a cone, or sterile leaves still attached.
(1X)
Cordaites gincipalis (germar) Geinitz 132

fig. 5 Spatulate leaf (X 1/2)

195

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bur-In}? - mi...

 

EXPLANATION OF PIA TE XII
Telanpum affine (L. & H.) Benson 154
figs. 1, 10 Specimens showing branching (IX)
of fertile frond and sporangia
fig. 16 Enlargement of fig. 10 (2X)

Cordaicappus globosus sp. nov. 140

 

fig. 2 Figure illustrating shape and relationship of
nucellus to the wings. (2X)

Cordaicarpus iayshuleri sp. nov. 141

 

figs. 3, 4, 5 Illustration showing striated surfaces of
seeds and lack of wings. (2X)

Cordaicarpus elongatus (Newberry) comb. nov. 139

 

fig. 6, ll, 14 These specimens- illustrate general shape
and size of wings. (2X)
Aulacotheca hemingwafl (2) Halle 157
fig. 7 Specimen of the sporocarp.
Cordaicarpus cordatus J ackson comb. nov. 141
fig. 12 Seed illustrating shape with wings attached (1X)
fig. 8 Enlargement of fig. 12
fig. 13 Nucellus with wings missing
fig. 9 Enlargement of fig. 13
Lagenospermum bifurcatus (Stopes) comb. nov. 149
fig. 15 Seed cupules, some with seed still present (1. 5X)

fig. 17 Enlargement of fig. 15. (3X)

/"i7/193

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PLATE XII

EXPLANATION OF PLATE XIII

 

Page
Cornucarpus discissus sp. nov. 147
fig. 1 Seed with bifurcating appendages. Note recurving
of the bifurcations. (1X)
Cordaicarpus manningcanensis sp. nov. 142

 

fig. 2 Seed showing rounded shape, distinct micropule. (1X)

Cordaicarpus binutus sp. nov. 143

 

fig. 3 Seed with a depression showing where the
reniform bodies were harbored. (XI)

Gnetopsis anglica Kidston 152

 

fig. 4 Seed, withiappendages. Note longitudinal folding
of seed coat along right edge of seed. (4X)
Rigbyocarpus ebracteatus sp. nov. 148
fig. 5 Seed surrounded by bracts (2X)
fig. 6 Bracts with the seed missing (2X)
Cordaianthus pgeudofluitans (I?) Kidston 134

fig. 7 Cordaitean strobilis without seeds attached (1X)

 

Holcospermum sp. 146
fig. 8 (1X)
Trignocarpus sp. C 146

 

figs. 9, 13 Specimen illustrating size and general shape
(fig. 9 shows ribs) (1X)
Trignocarpus sp. B 145

fig. 10 Sarcotesta with a portion of the micropyle present.

 

PLATE XIII

EXPLANATION OF PLATE XIII Cont.

 

Trignocarpus sp. A 145
fig. 11(1X)
Aulacotheca campbelli (White) Halle 156

 

fig. 12 Three micro-sporangiate structures on the

specimen illustrating the ribbing (2X0

b

Trignocarpus noeggerathi (Stnbg.) Brongniart 144

 

fig. 14 Specimen showing a single longitudinal ridge

(1X)

200