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New Permian Corals from Kansas, Oklahoma, and Texas

by Raymond C. Moore and Russell M. Jeffords

Originally published in 1941 as Kansas Geological Survey Bulletin 38, Part 3.


Nine new species of Permian rugose corals are referred to eight genera, of which three are new and three have
not previously been reported from America. New species from Lower Permian (Wolfcamp) beds of Kansas and
northern Oklahoma are assigned to Malonophyllum Okulitch and Albritton, known elsewhere from the Leonard
series of western Texas; Lophophyllidium Grabau, seemingly a long-ranging genus of Upper Carboniferous and
Permian age and world-wide distribution; Sochkineophyllum Grabau, previously reported from Middle
Carboniferous (Moscovian) rocks of Russia and Permian strata of China; and two new genera named
Lophamplexus and Heritschia. New species of Middle Permian (Leonard) corals from the Glass Mountains, in
western Texas are referred to Timorphyllum Gerth, described from the Permian of Timor, in the East Indies;
Duplophyllum Koker, previously reported from the Permian of Timor, Australia, and China; and a new genus that
is designated as Leonardophyllum. Most of these forms bear a distinct axial column and all but one species
seem to lack dissepiments.

Permian corals from Asia that heretofore have been referred to Corwenia Smith and Ryder, a genus that is
based on Lower Carboniferous corals, are placed in the new genus Heritschia, and a few other, variously
identified corals are also thought to belong in Heritschia.

The common Late Paleozoic column-bearing corals of North America that have been classed as belonging
to Lophophyllum seem to belong to the genus Lophophyllidium. The stratigraphic usefulness of corals in
correlation of Permian rocks is considered briefly. Numerous taxonomic problems similar to that of so-called
Lophophyllum are encountered in work on the Permian corals.

Corals are widely distributed in marine Permian deposits of the world and studies on these fossils are
comparatively numerous. Some of the published works are elaborate monographs, reflecting both the richness
of the coral faunas and the extensive nature of researches on them. In North America, however, less than a
dozen kinds of Permian corals have been reported, and descriptions are scattered in about half as many papers.

Previous Studies

Important papers on Permian corals outside of North America include the following: from Australia, by Etheridge
(1891) and Hill (1937); from Japan and Korea, by Yabe and Hayasaka (1915, 1916) and Hayasaka (1932); from
China, by Grabau (1922, 1928, 1931, 1936), Chi (1935, 1937, 1938), Yu (1934), and Huang (1932, 1932a); from
Timor, by Beyrich (1865), Rothpletz (1892), Penecke (1908), Gerth (1921, 1922), Koker (1924), and Heritsch
(1937a); from the Salt Range of India, by Waagen and Wentzel (1886), Sen (1931, 1931a, 1931b), and Heritsch
(1937a); from Iran, by Heritsch (1933a), and Douglas (1936); from Turkey, by Abich (1878), and Heritsch
(1937a); from the U.S.S.R., by Toula (1875), Dobrolyubova (1936), Soschkina (1925, 1928, 1932, 1936), and
Yakovlev (1903, 1939); from the Carnic Alps, by Heritsch (1933a, 1936), and Felser (1937, 1937a); from Serbia,
by Heritsch (1933, 1934); from Greece, by Heritsch (1937); and from England, by King (1850).

The earliest published record of Permian corals in North America is a description by B. F. Shumard (1858) of
specimens from the Guadalupe Mountains in western Texas. These were named Campophyllum? texanum and
Polycoelia? sp. No illustrations are given, and because the descriptions are inadequate and the types are lost,
Shumard's species are not recognizable. Girty (1908), in his monograph on the Guadalupian fauna of western
Texas, recognized nine species and one variety of corals, distributed in six genera, as follows: Lindstroemia
permicana Girty, n. sp., L. permicana var. Girty, L. cylindrica Girty, n. sp., L. sp., Zaphrentis? sp., Amplexus sp.,
Campophyllum? texanum Shumard, Cladopora spinulata Girty, n. sp., C.? tubulata Girty, n. sp., and Aulopora sp.
Some of these are not recognizable on the basis of the published descriptions and illustrations but possibly may
be made so by study of his types and supplementary specimens. One of us (R.C. Moore) recently has had
opportunity to examine Girty's types of Guadalupian corals, but lack of sections prevents diagnosis, at least in
the case of the rugose forms. A small horn coral from Permian limestone in northern Mexico has been described
by Haack (1914) under the name of Cyathaxonia girtyi, but because definitive characters are not indicated, this
form cannot now be recognized. Specimens collected by John Skinner from Lower Permian rocks of north-
central Texas and from Upper Permian beds in western Texas were sent by him to Franz Heritsch and on the
basis of this material descriptions of two species (Palaeosmilia schucherti, Waagenophyllum texanum) have
been published by Heritsch (1936a, 1936c). A few silicified corals from the Malone Mountains in western Texas
have been described by Okulitch and Albritton (1937) as the type species, Malonophyllum texanum, of a new
genus. A paper by Heritsch (1937b, p. 315) treating on the zonation of Permian rocks by means of rugose corals
mentions a form called Texanophyllum skinneri, said to occur in the Delaware Mountain formation of west Texas.
We have been unable to learn whether this coral has been described or whether the name is a nomen nudum.
No reference to it is found in any of Heritsch's papers in our file, complete so far as we know, nor is publication of
a description known to J.W. Skinner, for whom the species presumably was named. It has not been possible to
communicate directly with Dr. Heritsch.

No corals from Permian strata of the northern midcontinent region have been described, but Fath (1921, p. 48)
reports a determination by G.H. Girty of specimens found in Lower Permian rocks of Butler County, Kansas, as
Lonsdaleia? n. sp.

In British Columbia, good specimens of corals have been collected from rocks thought to be Upper
Carboniferous or Permian. These fossils were sent to Stanley Smith, British authority on late Paleozoic corals,
who described them under the names Waagenophyllum columbicum, n. sp., and Caninia sp. (Smith 1935).

The lack of extensive published data on North American Permian corals may be ascribed partly to their relative
scarecity in fossil collections. Many localities that have yielded abundant Permian marine fossils of several sorts
lack corals, or they have provided only a few specimens that may be none too well preserved. This observation
is emphasized by an inspection of the large collections from Permian formations of Texas in laboratories of the
U.S. Geological Survey and the University of Texas, or those from the northern midcontinent region in the
University of Kansas and the University of Nebraska.
Stratigraphic Significance of Permian Corals

Only locally and exceptionally in North America are Permian corals abundant. Seemingly, the known Permian
invertebrate faunas of this continent are much less coralline than those of many other regions. Little attention
has been given to search for these fossils, however, and special efforts in field collecting will doubtless result in
important additions of study material. Also, it is likely that all together there are many unstudied specimens in
collections of various surveys, universities, and museums. Careful investigation of all obtainable fossils should
augment considerably the knowledge of the kinds of Permian corals that occur in North America and their
stratigraphic distribution.

Heritsch (1936a, 1936c, 1937a, 1937b) has presented the major outlines of a zonation of rocks belonging to the
Permian system that is based on distribution of rugose coral genera. This seems to be very broadly applicable,
and classification of late Paleozoic marine deposits in all parts of the world eventually may become well
established through verified and amplified information on the nature and occurrence of corals. This group of
fossils may then become entitled to rank with ammonoids, fusulinids, and other especially useful guide fossils in
these rocks. The corals may be expected to supplement the other biologic groups to an important degree,
because noteworthy faunal variations due to ecologic factors are observed. As a whole, corals tend to choose
particular features of habitat, but, in addition, Hill (1938a) has called attention to existence of different types of
coral assemblages that reflect special environmental conditions. Mud deposits that presumably represent off-
shore moderately deep and quiet water are characterized by prevalence of small solitary horn corals. Calcareous
shale, commonly interbedded with thin limestone strata, representing moderately shallow, slightly turbid
conditions, contains robust horn corals of the Caninia type. Warm shallow clear waters, possibly agitated
considerably by waves and currents, characterize a limestone-producing environment, in which colonial reef-
building corals are abundant. The differentiation of these and other sedimentary facies requires somewhat
independent calibration of each type and imposes need for as complete intercorrelation as possible in order to
establish a composite paleontologic zonation of wide applicability.

Scope of the Present Paper

In this paper little attention is given to questions of correlation of Permian formations on the basis of corals or
definition of coral zones in the Permian succession of North America. Attack on such matters is premature at this
time. It is intended merely to describe and illustrate several of the corals that have been studied, and to indicate
some of the taxonomic problems that have been encountered in efforts to classify American species. Several
Permian coral genera that have been defined from study of collections from other continents are here recognized
in North America for the first time.

Our study of Permian corals has been undertaken as a part of the program of work outlined by Ronald K.
DeFord and others, under auspices of the American Association of Petroleum Geologists, in order to obtain
material for a volume to be published on the Permian rocks of the central and southwestern part of the United
States. Since the beginning of the project, the number of fossil corals available for study has increased
considerably, but because the additions came after most of the manuscript for this paper was completed, we
have decided to publish the results now obtained, taking notice of the new material only in so far as it belongs to
species already studied. It is hoped that work on the yet unstudied specimens can be pursued actively and that
our observations can be reported reasonably soon.

Sources of Material Studied

Specimens of Permian corals that have been available for our study have all been obtained from (1) beds of
Wolfcamp age, here classed as Lower Permian, in Kansas and Oklahoma, and (2) from Wolfcamp, Leonard, and
Word deposits in the Glass Mountains, western Texas. The Kansas specimens include material collected by
various geologists of the Kansas Geological Survey during years of mapping and stratigraphic field work, a small
lot of corals from Grand Summit, Kans., in a collection obtained from J.W. Mickle, of Wichita, and a large
collection of corals from the Florence limestone in Butler County, Kansas, that was made after beginning this
study. The west Texas corals were collected by R.C. Moore on trips to the Glass Mountains in 1939 and previous
years, and by R.H. King, J.W. Skinner, C.C. Williams, and us on a trip in March, 1940. Determination of
stratigraphic horizons is precisely made for all the Kansas collections on the basis of very detailed subdivision
and tracing of the Permian rocks in this region. Stratigraphic placement of the western Texas specimens rests on
the field studies and mapping of the Glass Mountains by P.B. and R.E. King (1930), supplemented in part by
personal acquaintance of Skinner with the geologic section of the Glass Mountains.
Methods of Study

External features of the Permian rugose corals serve only in small part as basis for classification, and in some
types they are almost negligible in taxonomic study. Likewise, the mode of growth, whether solitary or colonial, is
believed not to be an important distinction, unless such characters are firmly fixed. Judgments of the taxonomic
value of mode of growth and of various structural characters of corals differ widely. Variability of the corals
themselves, conjoined with the uncertainty of paleontologists as to significant evolutionary trends and
divergences in viewpoints just indicated makes for lack of uniformity in classification and nomenclature of these
fossils. In any case, knowledge of internal structures of the Permian corals is absolutely requisite.

After classification of the corals in our collection on the basis of obvious external features, the specimens were
cut transversely at selected points. Except in a few cases these sections served to establish the position of the
cardinal-counter septal plane and to indicate differentiation of major and minor septa. The sections were
photographed at enlarged scale after polishing or immersion in oil in order to bring out structures clearly. By
marking the septal grooves of major septa the arrangement and order of introduction of the septa were
determined on the exterior of the coral if this was possible. This method usually serves for identification of the
alar septa, which may not be independently identifiable from study of the transverse sections. The segments of
the transversely cut corals were carefully assembled in proper orientation (by means of index marks and
matching septal grooves) and were cemented together. They were then cut longitudinally in a plane
approximately at right angles to that of the cardinal and counter septa. The longitudinal section was oiled and
photographed, the half of the coral containing the counter septum, if identifiable, being used uniformly.

Illustrations of the structures shown in the transverse and longitudinal sections were prepared by drawing on the
photographs with waterproof India ink and then bleaching the photographs so as to leave only the inked lines.
The photographs, though mostly very clear, are less satisfactory for reproduction than the drawings. This is
partly due to the fact that half-tone or full-tone figures, which are required for reproduction of photographs, are
much more expensive than line cuts. Another item for consideration is the lack of clarity of many unretouched
photographs and the presence of adventitious features of several sorts that may obscure interpretation.
Comparative study of the photographs and of the specimens, mounted on the stage of a microscope, serves to
verify identification of significant structures in the sections. Irrelevant but frequently confusing details such as
cleavage lines in calcite and distributed foreign matter are eliminated in making the drawings. Thin sections or
celluloacetate peels, as described by Fenton (1935), are superior in some respects to the polished or oil-
immersed surfaces that were photographed in the course of our study, for the thin sections commonly show
structures more clearly and can be used for direct projection in making photographic enlargements. Preparation
of these sections calls for special techniques and is time-consuming. Also, it is not generally practicable to make
thin sections without destroying too large segments of the corals that are being studied.

Descriptive Terminology

Several attempts have been made to standardize the terminology of coral structures (Grabau, 1922; Hill, 1935;
Sanford, 1939), but none of these has gained general acceptance. These proposed schemes of terminology are
excessively technical and include a great number of precisely defined individual terms. We have adopted a
somewhat simpler, less concise terminology in order to avoid the use of terms not generally understood. It is
therefore necessary to define the meanings of many of the terms used in the following descriptions.

Theca--The outer layer of the corallite has commonly been designated as the epitheca, inasmuch as it was
presumed to be a layer deposited upon the original wall, or theca. In well-preserved specimens of rugose corals
Grabau (1922, p. 4) found, however, that this outer layer extends to the highest part of the calyx,, showing that it
is the first-formed and fundamental unit of the wall. Therefore, we follow Sanford (1939,, p. 305) in designating
this layer as the theca.

Septa--The terminology of the classes of septa is in a very confused state. We use the following designation of
the septa, as given by Hill (1935, pp. 504-505), because these terms have been used in a consistent manner
and have not varied in meaning as is true of the terms primary, secondary, and tertiary. The cardinal, counter,
two alar, and two counter-lateral septa are designated as theprotosepta. A variable number of other septa
added in pairs in the cardinal and counter quadrants and closely resembling the protosepta are designated the
metasepta. The protosepta and the metasepta together are termed the major septa. Short septa alternating
with the major septa, introduced later than the majors, are termed minor septa.
Axial structures--Both Grabau (1922) and Hill (1935) have proposed detailed classifications of the axial
structures of rugose corals. These authors differ widely in terminology, however, and neither has been followed
closely by many other workers. Although Grabau states that a true columella is found only in the Hexacoralla, he
describes the solid rodlike axial structure of the rugose corals as a columella. The term pseudocolumella is used
by him to include all non-solid types of axial structures, most of which are given descriptive names. Hill uses the
term columella for a solid rodlike axial structure and restricts the terms pseudocolumella to the Hexacoralla. The
general term axial complex includes both the columns formed by the twisting of axial ends of the septa (axial
vortex) and those formed by vertical plates in the axial region (axial column).

The terminology used by Grabau and Hill in describing the many types of axial structures is complete and
precise, but we do not think it necessary to use these little-known complex technical terms. Because the names
now used in describing axial structures are in such a confused state, we designate any columnar structure that is
present in the axial region of the corallite as a column and describe the features of each in detail. The column
may or may not be solid.


We are indebted to Ralph H. King and Charles C. Williams, of the University of Kansas, for aid in collecting
specimens, and to Maurice Wallace, of the University of Kansas, for work in sawing and polishing some of the
sections of corals. Ruth Mary Dudley, of the Kansas Geological Survey staff, assisted in supervising work of
N.Y.A. students who were employed for part of the task of making an illustrated card index of Carboniferous and
Permian corals of the world. The literature on Paleozoic corals is extremely large and because it has been
necessary to study early and middle Paleozoic coral genera in order to give foundation for work on Permian
forms, the organization of information from previously published papers has been one of the largest items of
labor up to the present point. We cordially express thanks for assistance received in the bibliographic work,
which would have been somewhat lessened if the extremely valuable Index of Paleozoic Coral Genera, by Lang,
Smith, and Thomas (1940) had been available to us at an earlier date.

Sincere thanks are given to E.H. Sellards and F.B. Plummer, of the Bureau of Economic Geology, University of
Texas, and to Carl O. Dunbar, Peabody Museum, Yale University, for the loan of specimens, which, however, are
mostly unstudied at the present time. Robert E. King has sent us some material from the Wolfcamp beds in the
Glass Mountains, and John W. Skinner has aided us, both by guidance in some of the field collecting in western
Texas and by providing information concerning fossil localities in the Permian rocks near Fife and Dothan in
north-central Texas. Monroe G. Cheney kindly took us to the place, southeast of Santa Anna, Texas, where Mr.
Skinner collected the specimens that were described by Heritsch as Palaeosmilia schucherti, and we were thus
able to procure an excellent collection of topotype specimens. William M. Furnish, of Oklahoma A. and M.
College, has collected and sent to use several lots of Lower Permian corals from northern Oklahoma. Finally, we
wish to acknowledge the kindness of James Steele Williams, of the U.S. Geological Survey, in giving opportunity
to examine the type specimens of Girty's species described from the Guadalupian rocks of western Texas.

The Graduate Research Committee of the University of Kansas is thanked for important aid that is represented
in a grant to defray photographic costs of materials for the illustrated catalog of corals and for the preparation of
drawings given in this paper.
Systematic Descriptions

General Statement Concerning Taxonomy

Classification of Paleozoic corals is now in confused, if not, almost chaotic state. It is generally recognized by the
best-informed students of these fossils that organization of described genera in families has been based very
largely on superficial and even artificial lines and that we are yet far from ready to evaluate with reasonable
accuracy the significant phylogentic evidence that is given in the structures and geologic distribution of the older
corals. Not only do families commonly defined by treatises such as Zittel's (1895) Handbuch der Paläontologie
and later works, include some very slightly related forms, but there can be no doubt as to the polyphyletic nature
of some genera, especially supposedly long-ranging genera of simple structure, like "Amplexus." Schindewolf
(1940) has recently described Lower Carboniferous and Permian species of amplexoid adult nature that are
shown to be derived from wholly different, unrelated stocks, and on this basis he follows Weissermel (1897) in
making new genera out of segments of the stratigraphically almost meaningless assemblage called Amplexus.
Convergence of characters in adult forms is shown to extend to internal features, such as arrangement and form
of the septa, as well as external features, and knowledge of youthful growth stages is necessary in order to
recognize the very unlike origins of seemingly identical end forms. A few authors, such as Grabau, recognize
finely drawn generic distinctions, of which many may be sound and probably some may be unsound. Many
families are proposed. Other authors, such as Hill (1939), are inclined to reduce to synonymy a very large
number of the listed Paleozoic coral genera and to group these in a small number of families. Lang, Smith, and
Thomas (1940) in their recently published Index, record 566 Paleozoic coral genera, exclusive of objective
synonyms, but they make no effort to arrange the genera in families.

The genera and species of Permian rugose corals that are described in this paper are discussed as units,
without indication of their inferred family relationships. They are arranged in approximate order of their increasing
complexity of internal structure, under each of two main groups, (1) genera that have a well-defined column, and
(2) genera that lack a column.

Corals Bearing an Axial Column

Genus MALONOPHYLLUM Okulitch and Albritton, 1937

This genus comprises conical to subcyclindrical solitary horn corals of moderately small size having a well-
defined theca marked by septal grooves. The internal structure corresponds to that of Lophophyllidium, except
that tabulae are absent. Longitudinal sections through the axial column of some specimens indicate a structure
like that of Lophophyllidium, consisting of superposed tabellae-like cones having nearly vertical sides built
around the distal part of the counter septum. The column projects upward strongly into the lower part of the

Genotype--Malonophyllum texanum, Okulitch and Albritton, Leonard series, Middle Permian, northwest end of
Malone Hills, Texas.

Discussion--Description of the genotype of Malonophyllum was not accompanied by sections showing internal
structure. The types of M. texanum are indicated to be silicified specimens, some of which are broken so as to
reveal internal features. Specimens in our collection that lack any evidence of tabulae and dissepiments seem
referable to this genus. Hill (1940, p. 131), indicates that Malonophyllum is to be regarded as a synonym of
Fasciculophyllum Thomson, but offers no discussion of such treatment. The illustrations of Fasciculophyllum
eruca (McCoy), which Hill thinks is most closely related to the genotype of Fasciculophyllum, the types and all
authentic specimens of which are missing, and also to Malonophyllum texanum, show the septa to be somewhat
curved toward the counter septum. The presence of alar fossulae and tabulae, and distinct inequality of length of
the major septa in Fasciculophyllum separate it from Malonophyllum, however.

Occurrence--Wolfcamp series, Grand Summit, Cowley County, Kansas; Leonard series, Malone Mountains,

Plate 1, figures 6-8; plate 7, figure 3

Corallites belonging to this species are steeply conical, short, solitary individuals that are slightly curved in the
plane of the alar septa. The theca is moderately thick, its surface being marked by deep septal grooves and
distinct interseptal ridges that are crossed by small wrinkles and growth lines. The calyx is deep, containing a
prominent spikelike axial column in the lower part. The type specimen is 20.2 mm in length and 11.7 mm in
maximum diameter, at the calyx.

A section of the type just below the calyx shows 24 short major septa. Alternating minor septa are one-third as
long as the major septa. Two other specimens show 20 and 22 major septa, respectively. The cardinal septum is
shortened so as to form an open fossula. The counter septum extends to the axis and thickens to form a solid
column in the young portion of the corallite, but near the calyx the counter septum is not significantly longer than
other major septa. Dissepiments and tabulae are absent. A solid rodlike column in the lower portion of the
corallite is formed by the thickened end of the counter septum. It tapers gradually upward and becomes free in
the lower part of the calyx.

Transverse sections of the early parts of the corallites show the structural elements much thickened by
stereoplasm. The septa are joined at their axial ends to form a large solid column. Successive sections higher in
the corallite reveal a progressive decrease in the amount of steroplasm and a shortening of the major septa. The
counter septum is the last to draw away from the column. Minor septa are not seen in immature portions of the

Discussion--The description and illustrations of the weathered specimens of Malonophyllum texanum Okulitch
and Albritton (1937) indicate that the septa withdraw only slightly from the axial column in the upper part of the
corallite and that they are strongly deflected toward the counter septum.
Malonophyllum kansasense can be distinguished readily from species of Lophophyllidium and Lophamplexus by
the concentration of structural elements and stereoplasm in the lower part of the corallite, and by the absence of
dissepiments and tabulae. The form of the corallite of M. kansasense is distinctly conical, in contrast to the
conical-cylindrical shape of Lophophyllidium dunbari and Lophamplexus eliasi, and it reaches maximum
diameter more rapidly.

Occurrence--Florena shale member, Beattie limestone, Council Grove group, Wolfcamp series, Lower Permian.
Grand Summit, Cowley County, Kansas.

Type--University of Kansas, no. 23291.

Genus LOPHOPHYLLIDIUM Grabau, 1928

Lophophyllum (in part) of authors.

Corals assigned to this genus are solitary individuals of straight or curved, conical to conical-cylindrical form,
having a well-developed longitudinally striated theca that is crossed transversely by growth lines and wrinkles of
varying strength. The calyx is moderately deep, bearing near its center a laterally compressed column that forms
a more or less prominent projection. Septa are straight and mostly thin, the counter septum joined to the axial
column, cardinal septum short but other major septa reaching almost to the center, not joined to the column.
Several thin tabulae extend from the theca to the column, their surfaces being gently convex upward or
somewhat wavy and irregularly curved. Dissepiments are not observed. The axial column is a more or less solid
rodlike structure of laterally compressed form, composed of a thickened portion of the counter septum and of
very steeply upbent extensions of the tabulae, thickened by stereoplasm.

Genotype--Cyathaxonia prolifera McChesney, Upper Carboniferous (Missouri stage), Illinois. Widely reported
from other Upper Carboniferous beds of North America and other continents and, probably erroneously, from
Lower Permian rocks of Russia.

Discussion--The classification and nomenclature of the relatively simple, column-bearing horn corals that are
widely distributed in Carboniferous and Permian deposits have long been in a confused state, and the studies
published by Carruthers (1913), Grabau (1928), Huang (1932), Yii (1933), Stanley Smith (1934), Heritsch
(1936b), Chi (1938), and Hill (1940)--not to mention numerous others--incompletely resolve the difficulties. The
widely accepted reference of American Upper Carboniferous and Permian corals to Lophophyllum seems to be
erroneous. Only brief treatment, touching the most pertinent aspects of these taxonomic problems, is offered

The genus Lophophyllum was proposed by Edwards and Haime (1850) on the basis of a Lower Carboniferous
coral from Tournai, Belgium, that was designated as genotype. This form they described as a new species,
Lophophyllum konincki. Carruthers (1913) showed that Edwards and Haime's genotype of Lophophyllum was a
synonym of Cyathaxonia tortuosa Michelin (1846), described from the same horizon and locality. Thorough study
by Carruthers of topotypes of Lophophyllum tortuosum establishes the significant characters of this genus, as
follows. The corallites are solitary, turbinate, and enclosed by a theca; major septa meet at the center in early
growth stages, one of them, generally the counter septum, is thickened at the inner end so as to form a
moderately prominent axial column that projects upward into the calyx, but the column may be discontinuous;
tabulae that arch upwards in the center to a varying degree occur below the floor of the calyx, and in the mature
part of the corallite dissepiments appear between the tabulate area and the outer wall; unlike Dibunophyllum,
there is no axial zone characterized by more numerous or vesicular tabulae or by a system of vertical lamellae
distinct from the septa.

In 1876 Thomson and Nicholson (p. 297) proposed a new genus, Koninckophyllum, that was based on a new
species called Koninckophyllum magnificum, from Lower Carboniferous rocks of England. This genus was
regarded as including both solitary and compound corals having internal structures essentially the same as
in Lophophyllum except for a greater prominence of the zone of dissepiments and the occurrence of "a
considerable space" between the axial column and the inner margins of the major septa. Thomson and
Nicholson (1876, p. 298) state that "in no case do the tabulae extend to the inner surface of the wall." Among
species assigned to Koninckophyllum are some that are definitely of the Lophophyllum tortuosum type, and
following Carruthers' (1913) restudy of the genotype of Lophophyllum, some workers have concluded
that Koninckophyllum should be suppressed as a synonym of Lophophyllum. On the other hand, Smith (1934),
Vaughn (1915), Heritsch (1936b), Hill (1939), and other students of these corals regard the two genera as
distinct, although closely related. Grabau (1928) and Huang (1932) arbitrarily and incorrectly place corals similar
to the genotype of Lophophyllum L. tortuosum, in Koninckophyllum and use Lophophyllum for corals similar to L.

Edwards and Haime in 1851 (p. 323) described an American column-bearing small horn coral from beds of early
Pennsylvanian age (Pottsville) at Flint Ridge, Ohio. This species they called Cyathaxonia profunda. McChesney
in 1860 (p. 75) described a similar coral from rocks of middle Pennsylvanian age in Illinois, naming
it Cyathaxonia prolifera. Meek (1872, p. 149) transferred McChesney's species to Lophophyllum, and Foerste
(1888, p. 136), working on the Flint Ridge fauna, classed C. profunda as belonging to Lophophyllum. American
paleontologists generally have come to regard L. proliferum as a synonym of L. profundum (Weller, 1898, p.
333), though this may well be an erroneous conclusion. It is very doubtful indeed that L. proliferum occurs in
Permian rocks of other continents, as reported, for example,, by Soschkina (1925, p. 88) in Russia.

Assignment of either "Cyathaxonia" profunda or "Cyathaxonia" prolifera to Lophophyllum is obviously incorrect,

because these species lack dissepiments, so far as known, and, more important, they have a nearly solid axial
column. On the ground that corals of the "Cyathaxonia" prolifera type are generically distinct from Lophophyllum,
Grabau (1928) proposed for them the new name,Lophophyllidium, designating Cyathaxonia
prolifera McChesney as genotype. Critical studies of the internal structure of authentic examples of C.
prolifera are needed in order to determine satisfactorily the characters that should be assigned
to Lophophyllidium. Our studies of Pennsylvanian and Permian corals of this sort, having a solid or
comparatively dense axial column, indicate the existence of several types of internal structures that may have
taxonomic significance. Based on present knowledge, Lophophyllidium is tentatively interpreted to include
solitary horn corals having a compressed column that is connected with the counter septum throughout most of
its length, other major septa (except the short cardinal septum) meeting, or almost meeting the column, but not
joined to it, the interior of the corallite marked by tabulae of sub-horizontal or somewhat convex attitude, and
lacking dissepiments.

In 1928 Grabau (p. 99) differentiated under the name Sinophyllum forms of lophophyllid corals having a thick
axial column joined to the counter septum so as to produce a pendulum-like appearance in transverse sections
of the corallites, ends of the longer septa also being deflected and joined to one another in such manner as to
form an inner wall nearly surrounding the column. Grabau designated Lophophyllum pendulum, described by
him in 1922, (p. 48) as genotype. Many corals from Upper Carboniferous and Lower Permian rocks of North
America also bear a thickened column that is joined to the counter septum and that show presence of a
stereozone formed by inner portions of the major septa, thus being suggestive of Sinophyllum. Huang (1932, p.
23) has described the structure of such a coral (called by him Lophophyllum proliferum) from Pennsylvanian
rocks of Pennsylvania. He observed in it a "wall" of thickened inner septal margins surrounding the thickened
column and, concluding that this structure has no generic significance, he referred both "Lophophyllidium"
proliferum and "Sinophyllum" pendulum to Lophophyllum. Huang included in Lophophyllum only those corals
that he regarded as similar to Lophophyllum proliferum. In the forms discussed by Huang and in others observed
by us, the distal parts of the major septa are not deflected, but are merely dilated by excess deposits of
steroplasm. The thickened septa come laterally into contact and may impinge also on the column but they retain
structural identity. The nature of the axial region corresponds only superficially to that of typical representatives
of Sinophyllum pendulum illustrated by Grabau and by Chi (1938) from China. Heritsch (1936) concludes, from
extensive studies of Carboniferous and Permian corals of the Carnic Alps and other areas,
that Lophophyllidium and Sinophyllum are distinct types of corals. It should be added that Grabau (1928, p. 99)
reports the occurrence of a few dissepiments on the cardinal side of Sinophyllum.

The axial column of the Lower Permian corals that are here referred to Lophophyllidium seems to be formed in
part by thickening of the inner margin of the counter septum, but most importantly it consists of the superposed,
almost vertically deflected parts of successive tabulae. It is the latter that account for the upward elongation of
the column above the floor of the calyx. Deposits of stereoplasm obscure this structure in some specimens, but
polished sections of even relatively dense parts of the column reveal its structural elements when studied under
the microscope. The superposed laminae of the column seem to be no more numerous than the tabulae, and no
basis is seen for designating the steeply inclined plates as tabellae. Transverse sections of some of the
specimens show clearly the dilated inner ends of the major septa, forming a stereozone that surrounds but
generally does not touch the column. No dissepiments have been seen. The observed structures support
identification of the specimens as belonging to Lophophyllidium, but there is a measure of uncertainty in
determination, owing to lack of detailed information as to the structure of authentic examples of the genotype
species, Lophophyllidium proliferum. Also important, but taxonomically less vital, is detailed knowledge of the
internal structure of Lophophyllidium profundum, for if L. proliferum is a synonym of this species, the genotype
of Lophophyllidium should be designated as L. profundum.

Occurrence--Upper Carboniferous and Permian of North America, Europe, and eastern Asia.


Plate 1, figures 1-5; plate 7, figure 4

Corallites belonging to this species have a conical-cylindrical form that is slightly curved in the plane of the alar
septa. The moderately thick theca shows prominent and regular septal grooves and interseptal ridges, the ridges
being slightly broader than the grooves. Numerous fine growth lines and somewhat low annulations run
transverse to the septal grooves. The type specimen in 25.5 mm in length and has a maximum diameter of 12.0
mm at the calyx.

The laterally compressed column projects as a tall spine into the center of the deep calyx. In early stages the
axial column is directly connected to the counter septum, but in the mature portion of the corallite it is free.

Major septa are strong and straight, but excepting the counter septum, they do not reach the column. These
septa vary in number from 19 to 25 in mature forms. Minor septa are inconspicuous, except in the adult portion
of the corallite, where they alternate with the major septa. Only rarely and irregularly are any of the minor septa
more than mere ridges on the interior of the theca.

The partial abortion of the cardinal septum forms a conspicuous open fossula. The tabulae and column are
described in detail in the generic description.

Serial sections of Lophophyllidium dunbari show that tne counter septum reaches the axis at a very early stage.
In succeeding sections the axial end of the counter septum becomes thickened by addition of the upbent edges
of the tabulae. The tabulae become more numerous and prominent in the upper part of the corallite, forming one
or more rows around the axial region. in later sections the major septa become club-shaped or rhopaloid by
thickening at their axial ends. Although the inner ends of these swollen septa are laterally contiguous and
approach close to the column, they retain their structural identity. In the mature stage the rhopaloid nature of the
septa disappears and they become progressively shortened so as to assume an amplexoid form. Tabulae are
entirely absent in the mature stage. In the uppermost sections the counter septum is indistinguishable from the
other septa and the column assumes a cylindrical form.

Discussion--Inasmuch as transverse sections of the mature region of many lophophyllid corals are very similar,
serial transverse sections and a longitudinal section are necessary in order to identify a specimen accurately.
Such sections have not been prepared in previous work on American corals of this type, and many of the forms
referred to Lophophyllum proliferum may belong to Lophophyllidium or related genera.

Lophophyllidium dunbari differs from Lophophyllidium proliferum (McChesney) (1860) in the proportionately
smaller diameter of the column and in the less curved form of the corallite. Information as to the internal features
of the genotype species, originally described by McChesney, is lacking.

Lophophyllidium dunbari differs from Lophophyllidium amygdalophylloidea Huang (1932) in its smaller axial
structure, much shorter minor septa, and fewer tabulae in the longitudinal sections. Lophophyllidium
zaphrentoidea Huang (1932) seems to lack numerous tabulae, and it has longer and more numerous septa
than L. dunbari. Also, L. zaphrentoidea has a more conspicuous cardinal fossula, and it seems to have no
distinct rhopaloid stage. From the Russian form identified by Soschkina (1928) as Lophophyllidium proliferum,
this species differs in the more prominent development of the rhopaloid septa and tabulae during development,
and the more amplexoid septa and smaller column in the mature part of the corallite.

The species is named for Carl O. Dunbar, of Yale University, who was a student of geology at the University of
Kansas. A leader in paleontologic and stratigraphic research, Professor Dunbar is especially noted on account of
contributions to knowledge of Permian fossils and rock formations. He has aided our study of the Permian corals
by loan of many specimens collected in Texas and Mexico.

Occurrence--Florena shale member, Beattie limestone, Council Grove group, Wolfcamp series, Lower Permian.
The type and other specimens were collected by J.W. Mickle and by N.D. Newell near Grand Summit , Cowley
County, Kansas.

Type--University of Kansas, no. 23301.


This genus includes solitary rugose corals of straight or gently curved, conical to cylindrical form, having a well-
developed theca that bears transverse growth lines and wrinkles but no clearly defined septal grooves. Except
the counter septum, which is joined to the axial column, the septa reach only part way to the column. The major
septa are evenly disposed and approximately uniform in length, except the cardinal septum, which is
distinguished by its short length and by its position opposite the counter septum. Minor septa occur between the
major ones.

The axial column is very prominent, being strongly elevated as a sharp point in the center of the calyx. Generally
it shows clearly the component structural elements when studied in longitudinal or transverse section. The most
important elements in the axial column are the sharply upturned central portions of tabulae that form closely
superposed sharp-pointed cones. The walls of the cones are intersected by a main vertical lamina that is
continuous with the counter septum, and by several radial lamellae that (as in Dibunophyllum) converge to meet
the main lamina at different points. Dissepiments are absent.

Genotype--Leonardophyllum distinctum, n. sp., Leonard series, Glass Mountains, western Texas.

Discussion--The described structural features of Leonardophyllum suggest comparison

with Clisiophyllum Dana, Dibunophyllum Thomson and Nicholson, Carcinophyllum Thomson and Nicholson, and
especially Verbeekiella Gerth. All these genera have an axial column that is characterized by arched tabulae or
tabellae intersected by radial lamellae, producing a cobweb pattern in tranverse sections. The genotype species
of Clisiophyllum, Dibunophyllum, and Carcinophyllum, and other species that are assigned without question to
these genera, are distinguished from the Permian corals here considered by a common occurrence of
dissepiments and by other features that need not be reviewed here in detail. Permian species that have been
assigned to the genera mentioned (Clisiophyllum australe Beyrich, C. torquatum Rothpletz, C.
talboti Hosking, Dibunophyllum rothpletzi Gerth, D. tubulosum Gerth, and Carcinophyllum cristatum Gerth) are
all assigned by Hill (1937, p. 54) to Verbeekia (equals Verbeekiella Gerth, the name Verbeekia Penecke being a
homonym). Also, Hill thinks that "Verbeekia" is not closely related to the Lower Carboniferous clisiophyllids, and
probably not derived from them. Verbeekia permica Penecke (1908), which was designated as genotype
of Verbeekia [Verbeekiella], comes from the same locality in Timor as that which yielded the type specimen
of ClisiophylIum australe Beyrich (1865), and studies by Gerth (1921, p. 85), which are accepted by Hill (1937, p.
54), indicate that Verbeekiella permica is a synonym of C. australe. Accordingly, the genotype
of Verbeekiella may be designated as Clisiophyllum australe. The axial column of Verbeekiella seems to
resemble that of Leonardophyllum, but closer inspection shows important distinctions. None of the illustrations
given by Beyrich, Rothpletz, Penecke, or Gerth for C. australe show a connection between the column and the
counter septum, and all indicate occurrence of a distinct wall surrounding the column, so as to separate sharply
the closely spaced arched tabellae and radial lamellae of the column from the differently disposed tabulae and
sepia outside the axial column. In Leonardophyllum there is no definite boundary between the central columnar
structure and the rest of the corallite. The arched or conical laminae of the columnar structure are parts of the
tabulae. They are not distinct elements confined to the columnar region, as in the case of tabellae.

The columnar structure of some lophophyllid types of Permian and Pennsylvanian corals correspond to that
of Leonardophyllum as regards extension and steep upward deflection of tabulae in the central area, but radial
lamellae are lacking in the so-called lophophyllids.

Occurrence--Leonard series, Middle Permian, western Texas.


Plate 2, figures 1-3; plate 8, figure 4

Corallites belonging to this species have a cylindrical form. The thick theca shows transverse growth-lines and
wrinkles but no septal grooves. The incomplete type specimen is 21.2 mm in length and 12.3 mm in diameter. A
cone-shaped column projects into the lower part of the deep calyx.

The major septa are strong and equal in length except for the very slightly shorter cardinal and slightly longer
counter septum. The major septa have a length equal to about one-fifth the diameter of the corallite. The minor
septa are about one-third the length of the major.

The tabulae are irregular and slightly anastomosing. They are nearly horizontal adjacent to the theca but rise
steeply to the column where they become nearly vertical. Dissepiments are absent.

The narrow dibunophyllid column is formed by the intersection of the steeply ascending tabulae, median lamina,
and a few strong radiating lamellae. These produce a cob-web appearance of the column in transverse section.
The tabulae, however, are not limited to the axial portion, but join the end of the counter septum and more rarely
other septa.

Discussion--The dibunophyllid column and steeply up-arched tabulae clearly separate this species from species
of other genera of North American corals. Leonardophyllum distinctum differs fromLeonardophyllum acus, the
only other species referred to this genus, in the less numerous and less closely bundled tabulae and in the more
nearly cylindrical shape of the corallite.

The thick theca and strong septa of the specimen identified by Dobrolyubova (1936, p. 132)
as Verbeekiella cf. rothpletzi Gerth resemble those of L. distinctum. The transverse sections of the Russian
specimen, however, show a different septal development and a column formed by a few very thick radiating
lamellae without a cob-web appearance.

Occurrence--Leonard series, Middle Permian, Hess Ranch, saddle north of Leonard Mountain, Glass
Mountains, Texas.

Type--University of Kansas, no. 74161. About a dozen incomplete specimens in our collection are assigned to
this species.

Plate 2, figures 4, 5, plate 8, figure 3

The description of this species is based on two well-preserved, although incomplete, conical-cylindrical corallites.
The theca is very thick, and shows prominent transverse wrinkles and growthlines, but no septal grooves. The
calyx is not known. The incomplete type specimen is 17.7 mm in length and has a maximum diameter of 11.0
mm at the uppermost part of the fragment.

The strong major septa thicken peripherally. The cardinal septum is only slightly shortened but the counter
septum reaches the column and is directly connected to the median lamina. Other major septa reach the column
and may be slightly twisted in the early stages, but in maturity withdraw from the column and have a length of
about one-fifth the diameter of the corallite. Minor septa are short and thick.

The tabulae are incomplete, closely packed, and anastomosing. They rise steeply at an angle of about 45° to the
column, where they become nearly vertical. Dissepiments are absent.

The axial portion of the corallite contains a broad column formed by the nearly vertical tabulae. In transverse
section the column has a well-defined cob-web appearance caused by the intersection of the tabellae with the
median lamina and the few radiating lamellae.

Discussion--Leonardophyllum acus is distinguished from other coral species in the same manner
as Leonardophyllum distinctum. L. acus can be distinguished from the latter species by the broad column and
closely packed anastomosing tabulae. In transverse section the tabulae are limited to the column, where they
are intersected by the radiating lamellae. The tabulae in L. distinctum broadly surround the column as well as
intersecting the radiating lamellae.

Occurrence--Leonard series, Middle Permian, Hess Ranch, saddle north of Leonard Mountain, Glass
Mountains, Texas.

Type--University of Kansas, no. 74162.

Genus LOPHAMPLEXUS, n. gen.

Solitary conical to subcylindrical, straight, curved, or somewhat irregularly bent corals of moderately small size
are included in this new genus. The theca is well developed, longitudinally grooved, showing position of the
septa, and marked by growth lines and transverse corrugations. The apical part of the corallite, representing
youthful stages of growth, bears internal structures typical ofLophophyllidium, having an axial column that is
composed mainly of vertical extensions of tabulae joined to the counter septum, and surrounding this, thickened
inner parts of the major septa that tend to form a stereozone. The mature part of the corallite is distinguished by
disappearance of the axial column, septa in this region becoming shortened and not distally thickened. Thin
tabulae curve convexly upward, extending to the thecal walls. Dissepiments are not observed.

Genotype--Lophamplexus eliasi, n. sp., Foraker and Beattie limestones, Lower Permian, Kansas and

Discussion--The corals here described are evidently related closely to Lophophyllidium, as indicated by
correspondence in structures of the juvenile parts of the corallite. They are readily distinguished
from Lophophyllidium and similar genera by absence of the column in the upper parts of the corallite and by the
tendency of the counter septum in the mature region to become even shorter than the adjacent major septa. The
cardinal septum is notably shortened in all stages of growth except the earliest. This reduction in length forms a
distinct but not prominent open fossula.

The distinctly amplexoid part of the corallite, as observed in specimens assigiled to Lophamplexus, seems to be
less dominant than in such genera as "Pseudamplexus" Weissermel (1897, p. 878),Pleramplexus Schindewolf
(1940, p. 401), and Pentamplexus Schindewolf (1940, p. 403). The youthful stages of these genera point to
parent stocks that are radically different, one from another, and the amplexoid nature of the mature stages of
each of them is a result of the evolutionary modification that Schindewolf calls convergence. Lophamplexus is
indicated to have been derived from an ancestor belonging to Lophophyllidium or a closely related genus.
Occurrence--Foraker and Beattie limestones, Wolfcamp series, Lower Permian, Kansas and Oklahoma.


Plate 3, figures 2, 3; plate 8, figure I

The principal structural features of this species are described in the generic description. The conical-cylindrical
corallite is slightly curved in the plane of the alar septa. Transverse growth lines and annulations are
conspicuous but septal grooves and interseptal ridges are narrow and somewhat shallow. The type specimen
has a length of 30.4 mm and a maximum diameter of 12.8 mm, measured at a distance of 20 mm from the apex.

The theca is very thin. There are 27 major septa in the uppermost part of the corallite of the type, which shows
no minor septa. Another specimen has very short minor septa alternating with the major septa in a section just
below the top of the column-bearing stage. An open cardinal fossula is visible in all but the most mature parts of
the corallite. Tabulae are incomplete in youthful stages, but above the column some are complete. The complete
tabulae are horizontal in the axial region but they slope downward and outward somewhat steeply near the
periphery. A column is present in the lower 43 percent of the length of the type, and in the lower 69 percent of a
specimen measuring 22.6 mm in length.

Discussion--This species is easily distinguished from described species of lophophyllid corals by the absence of
the column in the mature portion of the corallite. Transverse sections of the juvenile portions resemble those
of Lophophyllidium dunbari, and it is to be noted that sections of the upper parts of the calyx of specimens of L.
dunbari and Malonophyllum kansasense may also show no axial column. Longitudinal sections of these species,
however, do not show the more or less complete tabulae above the termination of the axial column. The coral
that was identified as Amplexus cf.abichi Waagen and Wentzel by Heritsch (1937a, p. 4) and other species of
"Amplexus" lack a prominent column in any part of the corallite.

Lophamplexus eliasi is named for Maxim K. Elias, of the Nebraska Geological Survey. Much work has been
done by Elias on Permian rocks and fossils, especially during the period of about ten years while he was a
member of the Kansas Geological Survey.

Occurrence--Foraker and Beattie limestones, Council Grove group, Lower Permian, Wolfcamp series. The type
was collected by N. D. Newell just east of the county line, near Grand Summit, Cowley County, Kansas, from the
Florena shale member of the Beattie limestone. The other specimen was collected 2 miles west and 1 mile north
of Fairfax, Oklahoma.

Type--University of Kansas, no. 23292.

Genus HERITSCHIA, n. gen.

This genus includes compound corals, individuals being of conical-cylindrical form, growing in sub-parallel
position but not closely adjoining, united only at points of lateral budding. The theca is marked externally by well-
defined longitudinal grooves that indicate position of septa, and at intervals there are moderately well developed
irregular cross-wrinkles. The depth of the calyx is equal to about one-half the diameter of the corallite, the sides
sloping evenly to a central pit that holds a prominent, flattened, and sharply pointed axial column. Septa are
numerous, evenly disposed, and nearly equal in size, the major ones extending only slightly farther toward the
center in the mature regions than the minor ones. A tetrameral arrangement of the septa is not observable in the
calyx, but the position of the plane intersecting the cardinal and counter septa is marked by the long transverse
axis of the column, and these septa may be located also by study of the arrangement of septal grooves on the
exterior of the theca. The septa are thin and strongly flexuous in the peripheral part of the corallite, but thickened
and relatively straight in the inner part. The counter septum is joined to the column but all other septa are slightly
separated from it.

A peripheral zone, equal in width to about one-half of the radius, bears numerous dissepiments, their inwardly
convex surfaces sloping steeply upward and outward. The inner margin of the dissepimental zone, as seen in
transverse sections, is marked by more closely spaced and somewhat thickened dissepiments that, combined
with an abrupt thickening of the septa, form a distinct stereozone.

The intermediate zone that contains the thickened inner margins of the septa is crossed by somewhat widely
spaced, partly anastomosing tabulae that slope steeply upward and outward, except for narrow subhorizontal
portions adjoining the axial structure. The column, joined to the counter septum, is lenticular in crosssection,
being compressed in the plane of the counter and cardinal septa and bearing a median lamina that lies in this
plane. Commonly, the column seems to be composed of a solid deposit of stereoplasm, but sections of several
specimens reveal a dibunophyllid structure consisting of a few radial lamellae that converge to meet the median
lamina at different points. These structures are crossed by steeply arched tabellae that slope nearly vertically
upward and inward. The septa and the median lamina of the column are fluted along curved lines than run
parallel to their distal margins.

Genotype--Heritschia girtyi, n. sp., Florence limestone, Lower Permian, Butler County, Kansas.

Discussion--Structural features of the corals here described under the new generic name of Heritschia are
intermediate between those of Waagenophyllum Hayasaka and Iranophyllum Douglas, on the one hand, and
such forms as Lonsdaleia McCoy and Stylidophyllum Fromentel, on the other. Waagenophyllum, as indicated by
its genotype species, Lonsdaleia indica Waagen and Wentzel, from the middle Productus limestone of the Salt
Range, India, comprises compound corals of closely bundled (phaceloid) habit, containing numerous small
closely spaced long cylindrical corallites. The septa of these corallites are flexuous and fluted as in Heritschia;
they are evenly disposed radially, comprise two orders, and they extend inward from the theca to or almost to the
axial column. Also, as in Heritschia, there is an outer zone of dissepiments sloping outward and upward, an
intermediate zone of tabulae similarly inclined except at their inner margins, and in the axial region closely
spaced tabellae that slope upward and inward. Unlike Heritschia, however, the septa of Waagenophyllum are
strongest in the peripheral zone, next to the theca, and the tabellae and radial lamellae of the column are much
less obscured by stereoplasm than in Heritschia. The axial structure of Waagenophyllum resembles that
of Dibunophyllum. Difference in deposits of stereoplasm in the axial region is not deemed to have generic
significance, and accordingly the chief distinction between Waagenophyllum and Heritschia, based on
comparison of their genotype species, is found in characters of the peripheral zone, where the externally
thickened septa and relatively narrow belt of dissepiments in Waagenophyllum contrast with the thin flexuous
outer parts of the septa and the strongly developed dissepimental zone of Heritschia.

Iranophyllum, which is based on the species Iranophyllum splendens Douglas, was established for solitary
rugose corals having the internal structure of Waagenophyllum and Wentzelella. The septa reach the theca, and
in general the tabulae are inclined proximally towards the axial column in ihe same manner as
in Waagenophyllum. The solitary form of growth, as well as the strong peripheral parts of the septa,
distinguish Iranophyllum from Heritschia.

Corwenia Smith and Ryder, as established on the basis of the Lower Carboniferous genotype
species, Lonsdaleia rugosa McCoy, differs essentially from Heritschia in the attitude of the tabulae and in the
expanded peripheral zone of dissepiments, which is not penetrated by the outer parts of the septa.
In Corwenia the inner parts of the evenly disposed septa are moderately thick, but outside a wall-like belt of
closely spaced dissepimental tissue they are very thin and do not reach the theca. A dibunophyllid column is
separated by a short space from the inner margins of the septa. Some Permian corals have been assigned
to Corwenia, but as interpreted on characters of the genotype and associated Lower Carboniferous
species, Corwenia is clearly unlike the Permian forms. It has a median zone of tabulae that rise towards the
column and are convex upwards, whereas tne tabulae of the median zone in Permian species that have been
referred to Corwenia slope distinctly downward and inward, as in Iranophyllum and Waagenophyllum. The major
septa of the Permian species are not extended to the central column as is the case in the type species, and the
minor septa are in general much more prominent than in Corwenia rugosa. Smith (1935), Douglas (1936), and
Hill (1940) have regarded these Permian species as more closely related to Waagenophyllum than to Corwenia.
We are of the opinion that the Permian corals that have been assigned to Corwenia belong to Heritschia.

According to Heritsch (1936a, p. 147, 1937b, p. 313), the lowermost Permian coral zone of China (Huang, 1932,
p. 10) and some other regions is characterized by the coral Stylidophyllum volzi(Yabe and Hayasaka). The close
packing of the corallites of Stylidophyllum, which imposes a prismatic form and polygonal outline on individuals,
is a main feature in distinguishing Stylidophyllumfrom Heritschia. The colonial growth form of Stylidophyllum is
unlike that of the very loosely packed corallites of Heritschia, but partial correspondence of internal structures
and seeming relationship of Stylidophyllum, Iranophyllum, Heritschia, and Waagenophyllum suggest the
approximate equivalence in age of Stylidophyllum and Heritschia. Undoubted representatives
ofWaagenophyllum seem to occur only in beds of Middle and Late Permian age.
On the basis of published figures and descriptions, we refer to Heritschia the Permian species Corwenia
chiutsingensis Chi (1931, p. 45), C. chiuyaoshanensis Huang (1932, p. 43), C. parachihsianensis Huang (1932,
p. 51), C. lipoensis Huang (1932, p. 52), Waagenophyllum columbicum Smith (1935, p. 38), and W.
persicum Douglas (1936, p. 20). Species doubtfully assigned toHeritschia include Corwenia
densicolumella Dobrolyubova (1936, p. 140), C. cf. lipoensis Huang (Dobrolyubova, 1936, p.
142), Waagenophyllum chitralicum Smith (1935, p. 37), W. muricatumDouglas (1936, p. 22),
and Lonsdaleia (Waagenella [sic]) omiensis Yabe and Hayasaka (1915, p. 104).

Heritsch (1936a, p. 145) differentiated Waagenophyllum columbicum, W. katoi, and W. chitralicum as a group
that is distinct from the typical forms of Waagenophyllum, inasmuch as the peripheral parts of the septa are
notably reduced in thickness, tending in some cases to disappear. The subhorizontal or somewhat upward and
inward sloping attitude of the tabulae of Heritschia columbica is more pronounced than in H. girtyi and the
dibunophyllid column is both broader and less obscured by steroplasm, but essential structural features
correspond closely. Although W. chitralicum has a prominent stereozone, like that of Heritschia, thickening of the
peripheral parts of the septa indicate closer relationship with Waagenophyllum. W. katoi differs from Heritschia in
the peripheral thickening of the septa, the type of column, and the form of growth.

As interpreted by Heritsch (1936a, p. 148; 1937b, p. 315-316) from a survey of Permian corals of the world, the
zone of Waagenophyllum is the upper middle part of the Permian system. W. indicumfrom the Middle Productus
limestone of the Salt Range in India certainly represents a horizon far above the base of the
Permian. Waagenophyllum texanum from the Capitan limeestone of West Texas and other species undoubtedly
belonging to Waagenophyllum are likewise obtained from beds that are much younger than lowermost
Permian. Heritschia girtyi, on the other hand, comes from beds of late Wolfcamp age. The Florence limestone of
Kansas has also yielded Pseudoschwagerina (Moore, 1940, p. 314). On the basis of this and much other
paleontologic and stratigraphic evidence, the placement of H. girtyi in the upper part of the succession of beds in
the Wolfcamp series, generally classed by American geologists as Lower Permian, is well fixed. This is not the
case in referring to H. columbica, inasmuch as the rocks that yielded this fossil are identified as "Permian or
possibly Upper Carboniferous." It is reasonable to infer that they are Lower Permian, and this seems to apply
also to the rocks containing other foreign species that are tentatively assigned to Heritschia. Stratigraphic
distribution, as well as paleontologic grounds, is thought to give basis for separation
of Heritschia from Waagenophyllum.

The genus Heritschia is named in honof of Franz Heritsch, of Wien (Vienna), who has contributed numerous
important studies of Permian corals.

Occurrence--Wolfcamp series, Lower Permian, Kansas; Lower Permian (?), British Columbia, Japan, Iran, and


Plate 4, figures 5-8; plate 7, figures 1, 2; plate 8, figure 5

Lonsdaleia? n. sp., GIRTY, 1919, in Fath, Kansas Geol. Survey, Bulletin 9, p. 48.

This species is based on abundant well-preserved material permitting serial sections of numerous specimens.
The corallites are conical near the apex but rapidly become cylindrical. Reproduction is by lateral budding,
attachment being at point of budding only. Colonies consist of many or few parallel cylindrical corallites. The
larger corallites reach a length of 80 mm and a diameter of nearly 20 mm.

The theca is thin, showing septal grooves and interseptal ridges crossed by prominent wrinkles. The type
specimen has 30 major septa and a like number of minor septa. The major septa are of equal length, except for
the counter and cardinal septa, and the minor septa are nearly as long as the major. An inconspicuous fossula is
formed by the shortening of the cardinal septum. The structure of the septa, column, tabulae, and dissepiments
are described under discussion of the genus.

Discussion--The strongly flexuous septa and complex dissepimental area, which are distinctive features
of Heritschia girtyi, are observed also in Heritschia columbica (Smith). The species from Canada has only a
slight indication of a stereozone, however, and its lacks the compact thickened column. The Chinese
species, Heritschia lipoensis (Huang, 1932), and Heritschia parachihsianensis(Huang, 1932), have a distinctly
open column and relatively straight septa. The peripheral dissepimental zone is less well developed and minor
septa are not more than half as long as the major septa.

The first collected specimens were found in 1918 by A.E. Fath, at a locality near Leon, Kansas, in the course of a
cooperative study of the El Dorado oil field by the U.S. Geological Survey and Kansas Geological Survey. The
specimens were examined by Girty who identified them as Lonsdaleia? n. sp. The corals that were collected by
Fath have not been seen by us. A small number of specimens was collected by M.K. Elias in 1932 while working
for the Kansas Geological Survey in Butler County. In 1940. R.C. Moore obtained a fairly large collection of these
corals near Leon, the lot containing parts of about 20 colonies and 200 or more corallites. Only one other
species of coral, which has not yet been thoroughly studied, is now known to be associated with Heritschia
girtyi in the Florence limestone. Several exceptionally well-preserved specimens were collected by A.I. Levorsen
of Tulsa, Okla., from a locality about a mile east of Leon. These were kindly cliven by him to the University of

This species is named in honor of the late George H. Girty, of the U.S. Geological Survey, who for many years
enriched the paleontologic literature on Carboniferous and Permian fossils of North America.

Occurrence--Upper part of the Florence limestone (about 5 feet above top of main chert zone), along road in NE
SE sec. 31, T. 27 S., R. 6 E., about 2 miles southwest of Leon, Butler County, Kansas, and in sec. 23, T. 27 S.,
6E., about one mile east of Leon, also Florence limestone (?), near Augusta, Kansas.

Type--University of Kansas, no. 34191.

Corals Not Bearing a Distinct Axial Column

Genus TIMORPHYLLUM Gerth, 1921

Small, solitary corals of somewhat irregular elongate cylindrical form are typical of Timorphyllum. The theca is
not marked by septal grooves but is crossed transversely by growth lines and wrinkles. A short cardinal septum
lies in an open fossula; the counter septum is elongate, reaching through the center of the corallite, but its distal
part is not distinctly thickened to form a column. Major septa are short, extending inward about one third the
diameter of the corallite; they are evenly spaced and not thickened distally. Minor septa are extremely short.
Somewhat regularly spaced tabulae arch gently upward and extend to the walls of the corallite. Dissepiments
are absent.

Genotype--Timorphyllum wanneri Gerth, Permian of Timor.

Discussion--The described structural features of this small coral readily distinguish it from other types of
Permian corals. Gerth's interpretation of the axial region as consisting of a much flattened columella that is
attached to the counter septum is supported by observation that a central structure projects upward in the lower
part of the calyx. This feature is not shown in our material, however, and transverse section of the corallite show
a barely perceptible thickening of the distal or axial part of the counter septum.

Occurrence--Lower Middle Permian of Bitauni and Basleo-Weslo, Timor (regarded as equivalent to the Artinsk
or Leonard and Word), and Leonard series, Middle Permian, West Texas.


Plate 5, figure 3; plate 8, figure 6.

The description of this species is based on several nearly complete, curved, cylindrical corallites partly
embedded in matrix. Inasmuch as the upper portions are broken, the calyx is not known. Incomplete length of
the type specimen is 35 mm and its average diameter is 5 mm. There are 18 major septa, which are of equal
length except the shortened cardinal septum and elongated counter septum. No minor septa are observed and
there is no stereozone. Complete tabulae occur at regularly spaced intervals. Their axial portions in a space
equal to about one-half the diameter of the corallite are horizontal, but their peripheral portions slope steeply
downward and outward.

Discussion--This species differs from Timorphyllum wanneri Gerth (1921) in that the theca is less wrinkled, and
the peripheral slope of the tabulae is steeper. T. simulans can be distinguished from T.
wanneri var. variabilis Gerth (1921) by its lack of a definitely separated column and the essentially equal length
of the septa.

Occurrence--Leonard series, Middle Permian, saddle north of Leonard Mountain, Glass Mountains, Texas.

Type--University of Kansas, no. 74163, collected by R. C. Moore.


This genus contains solitary conical corals of medium size having a longitudinally grooved theca. The calyx is
moderately deep, not containing a columnar projection. Internally, however, the counter septum is observed to
reach the center of the corallite and to be slightly thickened distally so as to simulate the lophophyllid column.
Three or more pairs of major septa, though shorter than the counter septum, are similarly thickened in the distal
region; other septa are much shorter, the cardinal septum being very short, so as to produce a fairly well marked
open fossula even at an early stage of growth. The counter quadrants are accelerated over the cardinal
quadrants. Tabulae of somewhat irregular and slightly anastomosing form extend across the interior of the
corallite, being arched gently upward. No dissepiments are observed.

Genotype--Pleurophyllum [sic] artiense Soschkina, Lower Permian (Artinsk), Government of Perm (Molotov),

Discussion--A distinctive feature of this genus is the thickened distal portion of an odd number of major septa,
which comprise the counter septum and three or more additional pairs of septa. Deposits of stereoplasm may
make the apical part of the corallite nearly solid. The thickening of septa towards their inner or axial margins,
which produces a club-shaped cross section, designated rhopaloid by some British paleontologists, is by no
means confined to Sochkineophyllum. It is a prominent peculiarity of Tachylasma, Plerophyllum, and some other
genera that are represented in Lower Carboniferous and probably Upper Carboniferous, as well as in Permian
rocks. In the genera other than Sochkineophyllum, the distribution of the rhopaloid thickening differs from that
ofSochkineophyllum, the unusually elongate counter septum being an especially distinctive feature
of Sochkineophyllum.

Hill (1940, p. 131) questionably includes Sochkineophyllum as a synonym of Fasciculophyllum Thomson, stating
that Sochkineophyllum "closely resembles Fasciculophyllum but some account of its [Sochkineophyllum],
variability is necessary before we can decide whether it is a synonym. The figured syntype of Sochkineophyllum
artiense only differs [from "Fasciculophyllum" eruca, which is not the genotype species] in that different
metasepta become long and rhopaloid." These two genera are similar in having a shortened cardinal septum
and an elongated counter septum; also, in both there is an absence of dissepiments, and several pairs of long
rhopaloid septa are observed. Hill describes Fasciculophyllum, however, as a form possessing alar fossulae,
globose tabellae, and rudimentary septa or minor septa of the type called contratingent (joined to adjacent major
septum outward from the cardinal septum).

The spelling of this generic name calls for brief consideration. The International Rules of Zoological
Nomenclature (Article 8h) provide that generic names may be based on patronymics and recommend that the
exact form of writing the proper name (using Roman letters) be employed. Thus, such genera
as Mülleria, Stålia, Krøyeria, and Ibañezia are recognized as correctly formulated. Special difficulty is
encountered, however, in transliterating some names from Russian, Chinese, and other languages that do not
use the Roman alphabet. Lack of uniformity naturally arises from an absence of an agreed procedure in
transliteration. We find that Yakovlev, and Jakowlew, as published in various non-Russian literature, all refer to
the same person, and that several divergent spellings of the same Chinese name may be given when written in
Roman letters. M.K. Elias has pointed out to us that the paleontologist for whom Sochkineophyllum was named
is a woman and that the surname of Russian women always bears the ending -a, as Soschkina. Published
spellings of this name include Sochkine (Soschkina, 1925), Soschkina (1928, 1936), Sochkina (Soschkina,
1932), and Soshkina (Nickles, Siegrist , and Tatge, 1938; Licharew and others, 1939; Lang, Smith, and Thomas,
1940). Grabau adopted the orthography as given in Soschkina's own rendering of her name in 1925, published
in French. Study of the Rules indicates that the spelling used by Grabau is not emendable (Moore, Weller and
Knight, 1941).

Occurrence--Upper Carboniferous (Moscovian), China; Lower Permian (Wolfcamp), Kansas; Middle Permian
(Artinsk), U.S.S.R.


Plate 3, figure 1; plate 7, figure 5

Description of this species is based on a very well preserved, nearly straight conical corallite, which has a length
of 22 mm and a maximum diameter (at the calyx) of 17 mm. The theca, which is of medium thickness, is marked
by prominent wrinkles and growth lines and by low septal grooves. Calyx 10 mm in depth.

The major septa are of unequal length in the mature part of the corallite. The cardinal septum extends only one-
fourth of the distance from theca to axis, whereas the counter septum, only slightly thickened, reaches to the
axis. Twelve other major septa extend nearly to the axis and are thickened distally. These elongated septa are
arranged in three groups, each consisting of four similar septa, two located on one side of the counter-cardinal
plane and two in corresponding position on the other side of this plane. If a transverse section of the corallite is
oriented with the counter septum at the top, one of these groups comprises the second and fourth septa on each
side of the counter septum, another group contains the seventh and ninth septa left of the counter and the sixth
and seventh to the right of the counter; the third group consists of the thirteenth and fourteenth septa on the left
and the tenth and eleventh on the right of the counter septum. Other major septa are shorter than the counter
and the 12 other long septa, their average length being about one-third that of the counter septum. Minor septa
are about one-third as long as the shorter major septa. In the type specimen there is a total of 29 major septa,
which are arranged in the following order, proceeding clockwise: counter septum, 12 major septa, cardinal
septum, 15 major septa, and back to the counter septum.

Transverse sections of the immature portions show major septa reaching nearly to the axis, being joined by
steroplasm and thickening at their distal ends. A fossula is formed by shortening of the cardinal septum. Thin
incomplete tabulae are seen but there are no dissepiments.

Discussion--No Late Paleozoic species resemblinc, Sochkineophyllum mirabile has been described from rocks
of this continent. The new species resembles Sochkineophyllum artiense (Soschkina, 1925) in the elongation
and thickening of several of the major septa and the extended nature of the counter septum. S. mirabile has a
greater number of long major septa, however, and has loncer minor septa. Sochkineophyllum tenuiseptatum
(Soschkinal, 1925) and Sochkineophyllum kansuense Grabau (1928) both lack the very prominent pairs of
elongated septa that are observed in the Kansas species. The unequal length of the major septa, lack of distinct
axial column, and more conical form of the corallite distinguish this species from representatives of
Malonophyllum and Lophophyllidium.

Occurrence--Florena shale member, Beattie limestone, Council Grove group, Wolfcamp series, Lower Permian.
Collected near highway on county line, Grand Summit, Cowley county, Kansas.

Type--University of Kansas, no. 23302.

Genus DUPLOPHYLLUM Koker, 1924

Elongate conical-cylindrical solitary corals having a well-developed theca that externally bears more or less
strongly developed transverse wrinkles and growth lines but no septal grooves are typical of Duplophyllum.
Septa are comparatively numerous, consisting of two orders, the major ones reaching to the center except near
the calyx; the arrangement of the septa is symmetrical, or nearly so, with respect to the cardinal-counter plane.
The position of the cardinal septum in the immature region is said by Koker to be occupied by a complex of short
septa that are joined together, an open space separating these from the center, but in the middle and upper parts
of the coral this structure disappears and a single cardinal septum is identified extending to the center. A
peculiarity of the genus is the tendency of minor septa to unite at their distal extremities with adjoining major

Genotype--Cyathophyllum? zaphrentoides Etheridge, Jun., Carboniferous?, New South Wales, Australia.

Discussion--Several problems are almost immediately encountered in studying Duplophyllum. In the light of
present information, there is ample room for doubt both as to validity of the genus and, if validity is established,
as to structural characters assigned to it. The original illustrations of Cyathophyllum? zaphrentoides do not
especially resemble figures of the corals from the Permian of Timor that were designated as Duplophyllum
zaphrentoides by Koker. Seemingly the latter specimens provided the basis for all her observations and generic
diagnosis. Etheridge's (1891, pl. 10, figs. 4-6) figures of C.? zaphrentoides indicate a coral that has very
numerous straight, relatively thick septa extending to the center, except in the case of short secondary septa, not
joined along their distal edge to one of the neighboring septa. Transverse sections show numerous, somewhat
regularly spaced curved tabulae and a peripheral zone that is characterized by stereoplasmic thickening of all
elements. There are numerous regularly spaced connections between septa, probably dissepiments. One
cannot rely too much on the published description and figures of C.? zaphrentoides by Etheridge, and because
Koker does not compare critically her Timor specimens with examples of the Australian species, available
evidence points to distinctness rather than identity of structure and classificatory status. Chi (1938, p. 164)
concludes that the specimens described by Koker are not conspecific with the doubtful and indefinite form from
New South Wales. Because Koker seemingly based the generic description on the Timor specimens, Chi thinks
that Koker's genotype should be designated as Duplophyllum, n. sp. Koker (1924, p. 22) describes the Timor
specimens in the text under the heading Duplophyllum cf. zaphrentoides (Etheridge). In the caption for the text
figure and in the explanation of plates, however, the forms are listed as Duplophyllum zaphrentoides(Etheridge).
Thus, it seems that Koker regarded her specimens as conspecific with those from New South Wales. C.?
zaphrentoides Etheridge must therefore be regarded as the genotype ofDuplophyllum.

There is question as to existence in the genotype species of characters ascribed to the genus by Koker, and
consequently there is doubt as to the generic diagnosis.

The "cardinal complex" of the immature region in a Timor specimen (or specimens?) studied by Koker may be
adventitious. It is not a normal structure in corals and is regarded as of doubtful significance. The tendency of
minor septa to unite distally with adjacent majors, shown in at least part of Koker's illustration of a Timor
specimen, associated with other characters, may have generic importance.

Fused septa of the type described are reported by Chi (1938, p. 165, pl. 1, figs. 5a-c) in Permian corals from
Yunnan, China, called Duplophyllum compactum. Similar fused septa are a characteristic structural feature also
of some of our corals from the Leonard series of the Glass Mountains. Transverse sections of specimens
illustrated by Koker show the notably uneven crooked attitude of some of the septa. This is not seen in Chi's
species (D. compactum), but is an especially noteworthy feature of the minor septa and a few of the major ones
in the West Texas specimens. The septa of these Leonard corals show a tendency to twist slightly in the axial
region, suggesting the axial structure of Clisiophyllum. No longitudinal sections are given by Etheridge for the
genotype species ofDuplophyllum nor by Koker for the Timor coral that she identified as belonging to this
species. Longitudinal sections of the West Texas corals are difficult to interpret, owing to partial disruption of the
internal structures. Tabulae are definitely identified, though most of them are broken and somewhat displaced.
Undisturbed portions of the tabulae indicate that they arch upward in the central part of the corallite. There are
no dissepiments.

Occurrence--The genotype species of Duplophyllum occurs in the Carboniferous? of Australia. The species from
Timor that was studied by Koker is reported to have been collected from Wesleo, Timor. Chi's Duplophyllum
compactum was obtained from the Maping limestone. The American corals that are here placed in Duplophyllum
are from the Leonard series. If all these references of corals to Duplophyllum are correctly made, they indicate
that the genus ranges from Carboniferous? to Leonard, and that it occurs in eastern Asia, Australia, the East
Indies, and North America.


Plate 5, figures 1, 2; Plate 6, figures 1-4; plate 8, figure 7

This species is represented by straight or irregularly curved, solitary, nearly cylindrical corallites, having growth
lines and coarse wrinkles but no longitudinal markings. The theca is thick in all stages of growth. The calyx is
unknown. Some of the specimens indicate various stages of rejuvenation, as two definite thecal walls separated
by matrix are shown in transverse section.

The type has 28 strong major septa in the upper part of the corallite, alternating with thin crooked minor septa. In
this section the major septa are equal in length except for the counter septum, which extends to the axis but is
not thickened to form a column. Minor septa are one-third the length of the major, except for one on each side of
the counter septum that are nearly as long as the major septa. Transverse sections lower in the corallite show
crooked major septa joined at the axis. The septa may unite in irregular groups or tend to twist together at their
axial extremities.

Individual major septa thicken near the periphery but vary irregularly in thickness throughout the rest of their
length. Irregularly curved minor septa thicken towards the axis. Many of the minor septa directly join the adjacent
major septum or are closely connected to it by tabulae.

Thin, somewhat anastomosing tabulae spaced at regular intervals rise steeply near the periphery and again
close to the counter septum where this is extended to the axis. There is, however, neither thickening of the
septum nor tabular structure forming an axial column. Lower in the corallites the septa join at the axis but do not
form a column. In these lower sections it is difficult to recognized the cardinal-counter plane.

Nearly all of the specimens show one or more stages of rejuvenation. This rejuvenation seems to follow so
closely the mature stage that only rarely are the nearly straight prominent septa of maturity seen in transverse

Discussion--Most of the specimens studied are difficult to interpret, owing to the broken and displaced internal
structures. This is particularly true of the longitudinal sections. The type, however, has not been affected in this

There is no described species of rugose coral from the Late Paleozoic rocks of this continent with which this
species could be confused. Duplophyllum cf. zaphrentoides (Etheridge) Koker (1924) from Timor is most closely
related to D. septaritgosum, but differs in the less prominent theca, more even thickness of the individual septa,
and less crooked character of both the major and minor septa. Duplophyllum compactum Chi (1938) may be
distinguished from the Leonard species by the more numerous septa, peripheral sterozone, and much less
crooked septa.

Occurrence--Upper 400 feet of Leonard series, Middle Permian, at Clay Slide, 2 miles west of Iron Mountain,
Glass Mountains, Texas,

Type--University of Kansas, no. 74141.


Abich, H., 1878, Eine Bergkalkfauna aus der Araxesenge bei Djoulfa in Armenien: Geol. Forsch. Kaukasischen
Ländern, tome 1, pp. 85-90, pls.

Beyrich, E., 1865, Uber eine Kohlenkalkfauna von Timor: Abh. d. K. Akad. d. Wiss., pp. 61-98, pls. 1-3, Berlin.

Carruthers, R.G., 1913, Lophophyllum and Cyathaxonia: Revision notes on two genera of Carboniferous corals:
Geol. Mag., n. ser., dec. 5, vol. 10, no. 2, pp. 49-56, pl. 3.

Chi, Y.S., 1931, Weiningian (Middle Carboniferous) corals of China: Palaeontologia Sinica, ser. B, vol. 12, fasc.
5, pp. 1-70, pls. 1-5.

Chi, Y.S., 1935, Notes on some Carboniferous and Permian corals of Dr. Erik Norin's collection from Sinkiang
Province: Geol: Soc. China Bull., vol. 14, pp. 23-34, pls. 1, 2.

Chi, Y.S., 1937, On some siple corals from the Permian of Yungsin, Kiangsi: Geol. Soc. China Bull., vol. 17, no.
1, pp. 83-108, pls. 1-4.

Chi, Y.S., 1938, Permian corals from southeastern Yunnan: Geol. Soc. China Bull., vol. 18, no. 2, pp. 155-190,
pls. 1, 2.

Diener, Carl, 1898, The Permocarboniferous fauna of Chitichun: Himalayan fossils, vol. 1, pt. 3., Palaeontologia
Indica, ser. 15, pp. 1-105, pls. 1-13.

Diener, Carl, 1903, Permian fossils of the central Himalayas; Himalayan fossils, vol. 1, pt. 5, Palaeontologia
Indica, ser. 15, pp. 1-204, pls. 1-10.

Diener, Carl, 1911, Anthracolithic fossils of the Shan States: India Geol. Survey Mem., n. ser., vol. 3, mem. 4,
pp. 2-67, pls. 1-7.

Dobrolyubova, T.A., 1936, Rugosa corals of the Middle and Upper Carboniferous and Permian of the North Ural:
Polar Comm. Acad. Sci. U.S.S.R. Trans., fase. 28, pp. 77-146 (Russian) 146-158 (English), text figs. 1-81.

Douglas, J.A., 1936, A Permo-Carboniferous fauna from southwest Persia (Iran): India Geol. Survey Mem.,
Palaeontologia Indica, n. ser., vol. 22, no. 6, pp. 1-59, pls. 1-5.

Edwards, H.M., and Haime, J., 1850, A monograph of the British fossil corals, Part 1. Introduction: Palaeont.
Soc., pp. i-lxxxv, 1-71, pls. 1-11.

Edwards, H.M., and Haime, J., 1851, Monographie des Polypiers Fossiles des Terrains Palaeozoïques,
precedée d'un tableau général de la classification des polypes: Arch. Mus. Hist. nat. Paris, tome 5, pp. 1-502,
pls. 1-20.

Edwards, H.M., and Haime, J., 1852, A monograph of the British fossil corals, part 3--Corals from the Permian
formation and the Mountain limestone: Palaeont. Soc., pp. 147-210, pls. 31-46.

Enderle, Julius, 1900, Uber eine Anthracolithische Fauna von Balia Maaden in Kleinasien: Beitr. z. Pal. u.
Geol.; Mitt. Paläeont. Inst. Univ. Wien, B. 13, Heft 2, pp. 49-109, pls. 4-8.

Etheridge, R. Jun., 1891, A monograph of the Carboniferous and Permo-Carboniferous Invertebrata of New
South Wales. Part 1, Coelenterata: New South Wales Geol. Survey Mem., Palaeont., no. 5, pp. 1-64, pls. 1-11.

Felser, K.O., 1937, Rugose Korallen aus dem Oberkarbon-Perm der Karnischen Alpen zwischen Schulterkofel
und Tressdorfer Hohe: Mitt. Naturwiss. Ver. f. Steiermark, Bd. 74, pp. 1-20, pl. 1.

Felser, K.O., 1937a, Mitteilung über einige stratigraphisch wichtige Korallen aus den obersten
Nassfeldschichten und dem Trogk6fekalk: Akad. Wiss. Wien, Math.-naturwiss, Kl., anx., Jg. 74, pp. 60-63.
Fenton, M.A., 1935, Nitrocellulose sections of fossils and rocks: Am. Midland Naturalist, vol. 16, pp. 410-412.

Foerste, A.F., 1888, Notes on Paleozoic fossils: Denison Univ., Sci. Lab., Bull. 3, pp. 117-136.

Fromentel, E. de, 1861, Introduction à l'étude des polypiers fossiles, pp. 1357.

Gerth, H., 1921, Die Anthozoen der Dyas von Timor: Paläontologie von Timor, Lief. 9, Abt. 16, pp. 67-147, pls.
145-150, Stuttgart.

Gerth, H., 1922, Der Palaeontologische Character der Anthozoenfauna des Perm von Timor: Jaarb. Mijnwezen
Ned.-Indië, 1920, Verh. 3, pp. 1-32, pls. 1, 2, Leiden.

Girty, G.H., 1908, The Guadalupian fauna: U.S. Geol. Survey, Prof. Paper 58, pp. 1-651, pls. 1-31.

Girty, G.H., 1913, A report on Upper Paleozoic fossils collected in China in 1903-1904: Research in China, vol.
3, Carnegie Inst. Washington pub. 54, pp. 297-334.

Grabau, Amadeus, 1922, Palaeozoic corals of China: Part 1, Tetraseptata: Palaeontologia Sinica, ser. B, vol. 2,
fasc. 1, pp. 1-70, pl. 1, text figs. 1-73.

Grabau, Amadeus, 1928, Palaeozoic Corals of China; Part 1, Tetraseptata: Palaeontologia Sinica, ser. B, vol. 2,
fase. 2, pp. 1-151, pls. 1-6, text figs. 1-22.

Grabau, Amadeus, 1931, The Permian of Mongolia: Nat. History Central Asia, vol. 4, pp. 1-665, pls. 1-35.

Grabau, Amadeus, 1936, Early Permian fossils of China, pt. 2: Palaeontologia Sinica, ser. B, vol. 8, fasc. 4, pp.
1-441, pls. 1-31.

Gregorio, Antoine de, 1930, Sul Permiano di Sicilia: Ann. Geol. Pal., 52 Livr., pp. 1-70, pls. 1-21.

Haack, Von, H.W., 1915, Uber eine marine Permfauna aus Nordmexiko nebst Bemerkungen über Devon
dasselbst: Deutschen geol. Gesell. Zeitschr., Bd. 66, pp. 482-504, pls. 38-39, text figs. 1, 2.

Hayasaka, I., 1924, On the fauna of the anthracolithic limestone of Omi-Mura in the western part of Echigo:
Tohoku Imp. Univ. Sci Repts.. Sendai, Japan, 2d ser., (Geol.), vol. 8, no. 1, pp. 1-82, pls. 1-7.

Hayasaka, I., 1932, Two new species of Permian Pachypora from the Kitakami Mountains region, Japan:
Formosa History Soc., Trans., vol. 22, nos. 118, 119, pp. 1-3, pls. 1, 2.

Heritsch, Franz, 1933, Waagenopltyllum indicum aus Belacrkva im westlichen Serbien: Ann. Géol. de la
Péninsule Balkanique, tome 11, fasc. 2, pp. 214-238, pls. 1, 2.

Heritsch, Franz, 1933a, Rugose Korallen aus dem Trogköfelkalk der Karawanken und der Karnischen Alpen:
Prirodoslovne Razprave, Knjiga 2. Ljubljana, 13, IV, pp. 42-55, pls. 5, 6.

Heritsch, Franz, 1934, Korallen aus dem oberen Perm von Likodra im westlichen Serbien: Bull. Service Geol.
Royaume Yougoslavie pour I'année 1933, tome 3, fasc. 2., pp. 27-43, pl. 1, text figs.

Heritsch, Franz, 1936, Korallen der Moskauer-, Gshel- und Schwagerinen-Stufe der Karnischen Alpen:
Palaeontographica, Beit. z. Naturges. d. Vorzeit, Bd. 83, Abt. A, pp. 99-162, pls. 14-18, text figs. 1-6, Stuttgart.

Heritsch, Franz, 1936a, A new species of Waagenophyllum from the Permian of the Glass Mountains, Texas:
Am. Jour. Sci., 5th ser., vol. 31, pp. 141-148, fig. 1.

Heritsch, Franz, 1936b, Lophophyllum, Lophophyllidium, and Sinophyllum: Centralblatt f. Min. etc., Jahrg. 1936,
Abt. B, no. 9, pp. 408-415.

Heritsch, Franz, 1936c, A new rugose coral from the Lower Permian of Texas, with remarks on the stratigraphic
significance of certain Permian coral genera: Am. Jour. Sci., 5th ser., vol. 32, pp. 134-144, pls. 1, 2.
Heritsch, Franz, 1937, Rugose Korallen aus dem Perm von Euböa: Akad. Athenon, Praktika, tome 12, nos. 3-4,
pp. 209-213, text figs. 1-5.

Heritsch, Franz, 1937a, Rugose Korallen aus dem Salt Range, aus Timor und aus Djoulfa mit Bemerkungen
über die Stratigraphie des Perms: Akad. Wiss. Wien, Math.-naturwiss, Kl. Sitzungsber., Abt. 1, 146, B. 1 und 2,
Heft, pp. 1-16, pls. 1, 2.

Heritsch, Franz, 1937b, Die rugosen Korallen und die Stratigraphie der Permformation: F.E. Suess-Festschrift
der Geol. Gesell. Wien, Mitt. Bd. 29, pp. 307-328.

Heritsch, Franz, 1938, Die stratigraphische Stellung des Trogköfelkalkes: Neues jahrb., Beilage-Band 79, Abt.
B, pp. 63-186, pls. 3-8, text figs. 1, 2.

Hill, Dorothy, 1935, British terminology for rugose corals: Geol. Mag., vol. 72, no. 857, pp. 481-519, text figs. 1-

Hill, Dorothy, 1937, The Permian corals of western Australia: Royal Soc. Western Australia Jour., vol. 23, pp.
43-63, pl. 1, text figs. 1-12.

Hill, Dorothy, 1938, Euryphyllum: a new genus of Permian zaphrentoid rugose corals: Royal Soc. Queensland
Proc., vol. 49, no. 2, pp. 23-28, pl. 1, text figs. 1-17.

Hill, Dorothy, 1938a, A monograph on the Carboniferous rugose corals of Scotland, Part 1: Palaeont. Soc., vol.
91, part 3 (for 1937), pp. 1-78, pls. 1, 2, 2 text figs.

Hill, Dorothy, 1939, A monograph on the Carboniferous rugose corals of Scotland, Part 2; Paleont. Soc. (for
1938), pp. 79-114, pls. 3-5.

Hill, Dorothy, 1940, A monograph on the Carboniferous rugose corals of Scotland, Part 3: Palaeont. Soc. (for
1940) vol. 94, pp. 115-204, pls. 6-11.

Hinde, G.J., 1890, Notes on the palaeontology of western Australia, corals and Polyzoa: Geol. Mag., new ser.,
dec. 3, vol. 7, pp. 194-204, pls. 8-8a.

Huang, T.K., 1932, Permian corals of southern China: Palaeontologia Sinica, ser. B, vol. 8, fasc. 2, pp. 1-163,
pls. 1-16.

Huang, T.K., 1932a, Some Uralian corals from northern Kuangsi collected by Dr. V.K. Ting in 1930: Geol. Soc.
China, Bull., vol. 12, no. 1, pp. 113-118, pl. 1.

King, P.B., and KING, R.E., 1930, The geology of the Glass Mountains, Texas: Texas Univ. Bull. 3038, pp. 1-
245, pls. 1-44, text figs. 1-43.

King, Wm., 1850, The Permian fossils of England: Palaeont. Soc., pp. 1-258, pls. 1-28.

Koker, E.M.P., 1924, Anthozoa uit het Perm van het Eiland Timor. I. Zaphrentidae, Plerophyllidae, Cystiphyllidae,
Amphiastraeidae: Jaarb, mijnwezen Ned.-Indië, Verh., Gravenhage, pp. 1-50, pls. 1-11, text figs. 1-26.

Koninck, L., de, 1862, Descriptions of some fossils from India: Quart. Jour. London Geol. Soc. Proc., vol. 19,
pp. 1-19, pls. 1-8.

Lang, W.D., Smith, Stanley, and Thomas, H.D., 1940, Index of Paleozoic coral genera: British Mus. Nat.
History, pp. 1-231, London.

Lange, Erich, 1927, Eine mittelpermische Fauna von Guguk Bulat (Padanger Oberland, Sumatra); Geolog.
Mijbouwk., Genootsch 7,1925, voor Nederland en Kolonien, verh. Geol. ser., Deel 7, pl. 5, text figs. 1-10.

Licharew, B., et al., 1939, The atlas of the leading forms of the fossil fauna U.S.S.R.: Central Geol. and Prosp.
Inst., vol. 6, Permian, pp. 1-228, pls. 1-56; text figs. 1-113.

McChesney, J.H., 1860-1865, Descriptions of new fossils, from the Palaeozoic rocks of the western states: pp.
1-96, pls. 1-9, Chicago. [pp. 1-56 (1859) issued Jan. 3, 1860; pp. 57 -76, issued May 24, 1860; pp. 77-96 issued
Feb., 1861; pls. issued April, 1865. Reissued in revised and rearranged form in Chicago Acad. Sci. Trans., pp. 1-
57, pls. 1-9, 1867.]

Martin, K., 1881-1883, Die Versteinerungs-Fuerenden Sedimente Timors, Beitr. zur Geologie öst-Asiens und
Australiens, Band. 1: Sammluilgen des Geol. Reichs-Museums Leiden, ser. 1, B. 1, pp. 1-64, pls. 1-3, Leiden.

Meek, F.B., 1872, Report on the paleontology of eastern Nebraska: Hayden, Final Report, U.S. Geol. Survey
Nebraska, U.S. 2d Cong., House Ex. Doc. 19, pp. 140-244, pls. 1-11.

Michelin, J.L.H., 1841-1848, Iconographie Zoophytologique, description par localités et terrains des polypiers
fossiles de France et pays environments: pp. 1-348, pls. 1-129, Paris. [pp. 1-40, 1841; pp. 42-72, 1842; pp. 73-
104, 1843; pp. 105-144, 1844; pp. 145-184, 1845; pp. 185-248, 1846; pp. 249-328, 1847; pp. 329-348, 1848.]

Moore, R.C., 1940, Carboniferous-Permian boundary: Am. Assoc. Petroleum Geologists Bull., vol. 24, pp. 282-
336, text figs. 1-5.

Moore, R.C., Weller, J.M., and Knight, J.B., 1941, Erroneous emendation of generic names: Jour.
Paleontology, vol. 15 (in press) [Note: 1942, v. 16; no. 2; p. 250-261].

Netschajew, A., von, 1894, Die Fauna der permischen Ablagerungen des östlichen Theils des europaischen
Russlands: Soc. Sci. Kasan, Trans., vol. 27, art. 4, pp. 1-503, pls. 1-12.

Nickles, J.M., Siegrist, M., and Tatge, E., 1938, Geol. Soc. America, Bibliography and index of geology
exclusive of North America, vol. 5, pp. 1-510.

Okulitch, Vladimir J., and Albritton, Claude C., Jr., 1937, Malonophyllum, a new tetracoral from the Permian
of Texas: Jour. Paleontology, vol. 11, pp. 24-25, pl. 4, figs. 15-17.

Ozawa, Y., 1925, Palaeontological and stratigraphical studies on the Permo-Carboniferous limestone of Nagato,
Part 2, Paleontology: Coll. Sci., Tokyo Jour., Imp. Univ., vol. 45, art. 6, pp. 1-90, pls. 1-14.

Penecke, Karl A., 1908, Ueber eine neue Korallengattung aus der Permformation von Timor: Jaarb. mijnwezen.
Ned. Indië, Zeven en Dertigste Jaargang, Wetensch. Gedeelte, sec. F, p. 1, pp. 657-659, text figs. 1, 2.

Rothpletz, A., 1892, Die Perm-, Trias- und Jura-Formation auf Timor und Rotti im indischen Archipel:
Palaeontographica, vol. 39, pp. 57-89, pls. 9-14.

Salee, A., 1913, Contribution à l'étude des polypiers du calcaire carbonifère de la Belgique, Part 2, Le groupe
des Clisiophyllides: Louvain Univ., Inst. Géol., Mem., tome 1, pp. 177-293, pls. 4-11.

Sanford, W.G., 1939, A review of the families of tetracorals: Am. Jour. Sci., vol. 237, pp. 295-323, 401-423, text
figs. 1-16.

Schenk, E.T. and McMasters, J.H., 1936, Procedure in taxonomy: pp. 1-72, Stanford Univ. Press, Palo Alto,

Schindewolf, Otto H., 1940, "Konvergenzen" bei Korallen und bei Ammoneen: Fortschr. Geologie u.
Palaeontologie, Bd. 12, Heft 41, pp. 389-492, 1 pl., text figs. 1-33.

Sen, A., 1931, On the occurrence of Lonsdaleia canalifera, Mans. in the Productus Limestone beds of the Salt
Range: Geol. Min. Met. Soc. India, Quart. Jour., vol. 3, pp. 35-36, pl. 6.

Sen, A., 1931a, On the identity between Lonsdaleia Indica, Waag. and Wentz., and Lonsdaleia Virgalensis,
Waag. and Wentz: Geol. Min. Met. Soc. India, Quart. Jour., vol. 3, pp. 9-11, pl. 1.
Sen, A., 1931b, On the development of the Genus Waagenophyllum Yabe and Hayasaka, from the Productus
Limestone beds of the Salt Range: Geol. Min. Met. Soc. India, Quart. Jour., vol. 3, pp. 125-134, pls. 9-10.

Shumard, F.B., 1858, Notice on fossils from the Permian strata of New Mexico and Texas: Acad. Sci. St. Louis
Trans., vol. 1, pp. 290-297.

Smith, Stanley, 1915, The genus Lonsdaleia and Dibunophyllum Rugosum (McCoy); Geol. Soc. London Quart.
Jour., vol. 71, pp. 218-272, pls. 17-21.

Smith, Stanley, 1934, An appendix in F.R.C. Reed, Anthracolithic fauna of the southern Shan States: India Geol.
Survey Rec., Vol. 67, pp. 83-134, pls. 1, 2.

Smith, Stanley, 1934a, Descriptions of two Anthracolithic corals, Waagenophyllum columbicum n. sp.
and Caninia sp. from British Columbia, and of some species of Waagenophyllum from the Tethys: Geol. Soc.
America, Proc., 1933, p. 375.

Smith, Stanley, 1935, Two Anthracolithic corals from British Columbia and related species from the Tethys: Jour.
Paleontology, vol. 9, pp. 30-42, pls. 8, 9.

Smith, S., and Ryder, T.A., 1926, The genus Corwenia gen. nov.: Annals and Mag. Nat. History, vol. 17, ser. 9,
pp. 149-159, pls. 5, 6.

Sochkina, E., 1932, The Lower Permian corals of the Oufimskoe Plateau: Soc. naturalistes Moscou Bull., géol.
sec., col. 10, pt. 2, pp. 251-267, pl. 1, text.

Sochkine, E., 1925, Les coraux du Permien inférieur (étage d'Artinsk) du versant Occidental de l'Oural: Soc.
naturalistes Moscou Bull., géol. sec., tome 3, n. ser., tome 33, pp. 76-104, pls. 1-3.

Soschkina, E., 1928, Die unterpermischen Korallen vom westlichen Abhang des nordlichen Uralgeberges: Soc.
naturalistes Moscou Bull., géol. sec., vol. 5, pp. 339-393, pl. 12, text figs.

Soschkina, E., 1936, New species of the Artinskian (Lower Permian) corals from the Aktubinsk region, south
Ural: Petroleum Geol.-Prosp. Inst., tr. e. B., no. 61, pp. 27-40, text figs. 1-13.

Stache, Guido, 1883, Fragmente einer afrikanischen Kohlenkalkfauna aus dem Gebiete der West-Sahara:
Akad. Wiss. Wien, Math.naturwiss. KI., Denkschr., Bd. 46, pp. 369-418, pls. 1-7.

Stuckenberg, A., 1888, Anthozoen und Bryozoen des oberen mittelrussischen Kohlenkalks: Comité Géol. Mem.
vol. 5, no. 4, pp. 1-44 (Russian), pp. 4554 (German), pls. 1-4.

Stuckenberg, A., 1895, Korallen und Bryozoen der Steinkohlenablagerungen des Ural und des Timan: Comité
Géol. Mém., vol. 10, no. 3, pp. 1-175 (Russian) pp. 180-244 (German), pls. 1-24.

Stuckenberg, A., 1904, Anthozoen und Bryozoen des unteren Kohlenkalks von Central-Russland: Comité Géol.
Mém., n. ser., livr. 14, pp. 1-67 (Russian), pp. 68-109 (German), pls. 1-9.

Thomson, J., and Nicholson, H.A., 1875, Contributions to the study of the chief generic types of the
Palaeozoic corals: Annals and Mag. Nat. History, ser. 4, vol. 16, pp. 424-429, pl. 12.

Thomson, J., and Nicholson, H.A., 1876, Contributions to the study of the chief generic types of Palaeozoic
corals: Annals and Mag. Nat. History, ser. 4, vol. 17, pp. 60-70, 123-128, 290-305, 451-462, pls. 6-8, 12, 14-17,

Thomson, J., and Nicholson, H.A., 1876a, Contributions to the study of the chief generic types of the
Palaeozoic corals: Annals and Mag. Nat. History, ser. 4, vol. 18, pp. 68-72, pls. 1-3.

Toula, Franz, 1875, Eine Kohlenkalk-Fauna von den Barents-Inseln (Nowaja-Semlja N.W.): Akad. Wiss. Wien,
Math.-naturwiss. Kl., Delikschr., Bd. 71, Abt. 1, Heft 5, pp. 527-608, pls. 1-6.
Vaughn, Arthur, 1915, Correlation of Dinantian and Avonian: Palaeontological section: Geol. Soc. London
Quart. Jour., vol. 71, pp. 34-52, pls. 3-7.

Waagen, William, and Wentzel, J., 1886, Salt Range fossils, vol. 1, Productus Limestone fossils, pt. 6,
Coelenterata: India Geol. Survey Mem., Paleontologia Indica, ser. 13, pp. 835-924, pls. 97-116, text figs. 26-32.

Weissermel, W., 1897, Die Gattung Columnaria und Beiträge zur Stammesgeschichte der Cyathophylliden und
Zaphrentiden: Deutsche geol. Gesell. Zeitschr., vol. 49, pp. 865-888, text figs. 1-3.

Weller, Stuart, 1898, A bibliographic index of North American Carboniferous invertebrates, U. S. Geol. Survey,
Bull. 153, pp. 1-653.

Yabe, H., and Hayasaka, I., 1915, Palaeozoic corals from Japan, Korea and China: Geol. Soc. Tokyo Jour., vol.
22, no. 261, pt. 1, pp. 55-70, pl. 5; vol. 22, no. 263, pt. 2, pp. 79-91, pl. 6; vol. 22, no. 264, pt. 3, pp. 93-109, pls.
7, 8; vol. 22, no. 265, pt. 3, pp. 127-139, pls. 9-10.

Yabe, H., and Hayasaka, I., 1916 Palaeozoic corals from Japan, Korea, and China: Supp. 3 and 4: Geol. Soc.
Tokyo Jour., vol. 23, no. 271, pp. 57-75, pl. 6.

Yakovlev, N., 1903, Die Fauna der oberen Abtheilung der palaeozoischen Ablagerungen im Donez-Bassin, II,
Die Korallen: Comité Géol. Mém., n. ser., livr. 12, pp. 1-8 (Russian), pp. 9-16 (German), pl. 1, text figs. 1-11.

Yakovlev, N., 1939, Nouveaux genres de coraux Tabulata du permien inferieur de l'Oural et du bassin du
Donetz: U.S.S.R. Bull., vol. 24, no. 6, pp. 622-632, text figs. 1, 2.

Yu, C.C., 1933 (1934), Lower Carboniferous corals of China: Palaeontologia Sinica, ser. B, vol. 12, fase. 3, pp.
1-212, pls. 1-24.

Yu, C.C., 1934, Descriptions of corals collected from the Maping and Huanglung limestones in South China:
Acad. Sinica, Mem. National Research Inst. Geol., 14 (C), pp. 55-72, pls. 9-13.

Zittel, K.A., von, 1876, Handbuch der Palaeontologie: Bd. 1, pp. 203-236.
Explanation of Plate 1
All figures 2.5 times natural size.

Lophophyllidium dunbari, n. sp., Florena shale member, Beattie limestone, Council Grove group,
Wolfcamp series, Lower Permian, Grand Summit, Cowley County, Kansas.
1a-e—Transverse sections of specimen (Univ. Kansas no. 24841).
2a-d—Type specimen (Univ. Kansas no. 23301). a, Longitudinal section. b-d, Transverse
3a-d—Specimen (Univ. Kansas no. 24842). a, Longitudinal section. b-d, Transverse
4a-d—Transverse sections of specimen (Univ. Kansas no. 23303).
5a-c—Transverse sections of specimen (Univ. Kansas no. 23304).

Malonophyllum kansasense, n. sp., Florena shale member, Beattie limestone, Council Grove group,
Wolfcamp series, Lower Permian.
6a-d—Specimen (Univ. Kansas no. 24843) from Grand Summit, Cowley County, Kansas.
a, Longitudinal section. b-d, Transverse sections.
7a-d—Type specimen (Univ. Kansas no. 23291) from along highway east of county line,
near Grand Summit, Cowley County, Kansas. a, Longitudinal section. , b-d, Transverse sections.
8a-d—Specimen (Univ. Kansas no. 21941) from road cut on highway in sec. 15, T. 31 S.,
R. 8 E., Cowley county, Kansas. a, Longitudinal section. b-d, Transverse sections.

New Permian Corals from Kansas, Oklahoma, and Texas, by Raymond C. Moore and Russell M.
Originally published July 1941 as Kansas Geological Survey Bulletin 38, Part 3.
Available online at
June 2007
Explanation of Plate 2
All figures 2.5 times natural size.

Leonardophyllum distinctum, n. gen., n. sp., Leonard series, Middle Permian, saddle north of
Leonard Mountain, Glass Mountains, Texas.
1a-b--Specimen (Univ Kansas no. 74165). a, Combined transverse and longitudinal
sections. b, Transverse section.
2a-c--Type specimen (Univ. Kansas no. 74161). a, Longitudinal section. b, c, Transverse
3--Longitudinal section of specimen (Univ. Kansas no. 74164) showing the calyx.

Leonardophyllum acus, n. gen., n. sp., Leonard series, Middle Permian, saddle north of Leonard
Mountain, Glass Mountains, Texas.
4a-d--Type specimen (Univ. Kansas no. 74162). a, Longitudinal section. b-d, Transverse
5--Combined transverse and longitudinal sections of specimen (Univ. Kansas no. 74166).

New Permian Corals from Kansas, Oklahoma, and Texas, by Raymond C. Moore and Russell M.
Originally published July 1941 as Kansas Geological Survey Bulletin 38, Part 3.
Available online at
June 2007
Explanation of Plate 3.
All figures 2.5 times natural size.

Sochkineophyllum mirabile, n. sp., Florena shale member, Beattie limestone, Council Grove group,
Wolfcamp series, Lower Permian, near Grand Summit, Cowley County, Kansas.
1a-e—Type specimen (Univ. Kansas no. 23302). a, Longitudinal section. b-e, Transverse

Lophamplexus eliasi, n. gen., n. sp.

2a-d—Specimen (Univ. Kansas no. 67581) from the Hughes Creek shale member, Foraker
limestone, Council Grove group, Wolfcamp series,, Lower Permian, 2 miles west and 1 mile north
of Fairfax,, Okla. a, Longitudinal section. b-d, Transverse sections.
3a-f—Type specimen (Univ. Kansas no. 23292) from the Florena shale member, Beattie
limestone, Council Grove group, Wolfcamp series, Lower Permian, along highway east of county
line, near Grand Summit, Cowley County,. Kansas. a, Longitudinal section. b-f, Transverse

New Permian Corals from Kansas, Oklahoma, and Texas, by Raymond C. Moore and Russell M.
Originally published July 1941 as Kansas Geological Survey Bulletin 38, Part 3.
Available online at
June 2007
Explanation of Plate 4.
All figures 2.5 times natural size, except as indicated.

Corwenia rugosa (McCoy), Lower Carboniferous, north Wales and north Welsh border, England.
1a-b—Specimen, after Smith and Ryder (1926, pl. 5) x1.5 a, Transverse section (fig. 2). b,
Longitudinal section (fig. 4).
Waagenophyllum indicum (Waagen and Wentzel), Middle Productus limestone, Upper Permian,
Salt Range, India.
2a-c—Specimen after Waagen and Wentzel (1886, pl. 101) x 16.5. a, Transverse section
(fig. 1c). b, Longitudinal section (fig. 3c). c, Column and peripheral part of section (figs. 3a, b).
Heritschia columbica (Smith), Permian (or Upper Carboniferous), British Columbia, Canada.
4—Transverse section of specimen after Smith (1935, pl. 9, fig. 1), x 2.
Heritschia girtyi, n. gen., n. sp.
5a-b—Type specimen (Univ. Kansas no. 3491) from upper part Florence limestone, Chase
group, Wolfcamp series, Lower Permian, along road in NE SE sec. 31, T. 27 S., R. 6 E., about 2
miles southwest of Leon, Butler County, Kansas. a, Transverse section. b, Longitudinal section.
6a-b—Transverse sections of specimen (Univ. Kansas no. 77821) from ?Florence
limestone, Chase group, Wolfcamp series, Lower Permian, near Augusta, Kansas. a, Transverse
section just below the calyx. b, Transverse section 12.5 mm lower in coralite.
7—Longitudinal section of a specimen (Univ. Kansas no. 34192).
8—Longitudinal section of specimen (Univ. Kansas no. 34193).

New Permian Corals from Kansas, Oklahoma, and Texas, by Raymond C. Moore and Russell M.
Originally published July 1941 as Kansas Geological Survey Bulletin 38, Part 3.
Available online at
June 2007
Explanation of Plate 5.
All figures 3 times natural size.

DuplophyIlum septarugosum, n. sp., upper 400 feet of Leonard series, Middle Permian, at Clay
Slide, 2 miles west of Iron Mountain, Glass Mountains, Texas.
1a-e—Type specimen (Univ. Kansas no. 74145). a, Longitudinal section of uppermost part
of corallite. b, Longitudinal sections of lower portion. c-e, Transverse sections.
2—Transverse sections of specimen (Univ. Kansas no. 74146).
Timorphyllum simulans, n. sp., Leonard series, Middle Permian, saddle north of Leonard Mountain,
Glass Mountains, Texas.
3a-c—Type specimen (Univ. Kansas no. 74163). a, Longitudinal section. b-c, Transverse

New Permian Corals from Kansas, Oklahoma, and Texas, by Raymond C. Moore and Russell M.
Originally published July 1941 as Kansas Geological Survey Bulletin 38, Part 3.
Available online at
June 2007
Explanation of Plate 6.
All figures 3 times natural size.

Duplophyllum septarugosum, n. sp., upper 400 feet of Leonard series, Middle Permian, at Clay
Slide, 2 miles west of Iron Mountains, Texas.
1a-c--Transverse sections of a specimen (Univ. Kansas no. 74141).
2a-c--Transverse sections of a specimen (Univ. Kansas no. 74142,).
3a-e--Specimen (Univ. Kansas no. 74143). a, Longitudinal section. b-e, Transverse
4a-d--Specimen (Univ. Kansas no. 74144). a, Longitudinal section. b-d, Transverse

New Permian Corals from Kansas, Oklahoma, and Texas, by Raymond C. Moore and Russell M.
Originally published July 1941 as Kansas Geological Survey Bulletin 38, Part 3.
Available online at
June 2007
Explanation of Plate 7.
All figures 3 times natural size.

Heritschia girtyi, n. gen., n. sp. Nearly perfect corallites weathered from the upper part of the
Florence limestone in sec. 23, T. 27 S., R. 6 E., Butler County, Kansas; collected by A.I. Levorsen.
1a, b—A small specimen showing the typical elongate cylindrical form of individuals
belonging to colonies of the species. a, Side view. b, Drawing of one half of the calyx, showing the
strongly projecting column and the flat-bottomed vertical-walled nature of the central pit. (Univ.
Kansas no. 77551).
2—A complete but very short individual having about the maximum observed diameter of
the corallites in the species. View from the cardinal side, tipped so as to show part of the deep calyx
and the terminal portion of the laterally compressed column. (Univ. Kansas no. 77552).

Malonophyllum kansasense, n. sp., from the Beattie limestone in sec. 15, T. 31 S., R. 8 E., near
Grand Summit, Kans.
3—Side view of the type specimen. Position of transverse sections shown on plate 1 is
indicated. (Univ. Kansas no. 21941).

Lophophyllidium dunbari, n. sp., from the Beattie limestone in sec. 15, T. 31 S., R. 8 E., near Grand
Summit, Kans.
4—Side view of the type specimen, showing alar septal groove and indicating position of
transverse sections given in plate 1. (Univ. Kansas no. 23301).

Sochkineophyllum mirabile, n. sp., from the Beattie limestone in sec. 15, T. 31 S., R. 8 E., near
Grand Summit, Kans.
5—Side view of the type specimen, showing conical form of the corallite and the shallow
longitudinal grooves that are crossed by fine growth lines. The position of the transverse sections
given on plate 3 is indicated. (Univ. Kansas no. 23301).

New Permian Corals from Kansas, Oklahoma, and Texas, by Raymond C. Moore and Russell M.
Originally published July 1941 as Kansas Geological Survey Bulletin 38, Part 3.
Available online at
June 2007
Explanation of Plate 8.
All figures 3 times natural size.

Lophamplexus eliasi, n. gen., n. sp., from the Council Grove group, Lower Permian, in southern
Kansas and northern Oklahoma.
1—Side view of a specimen from the Hughes Creek shale member of the Foraker
limestone, 1 mile north and 2 miles west of Fairfax, Osage county, Okla. The position of transverse
sections shown on plate 3 is indicated. (Univ. Kansas no. 67581).
2—Side view of the type specimen, the cardinal septum in the middle part of the view. The
top of the specimen is a cut and polished section; other transverse sections shown on plate 3 are
located at positions marked. The geniculation slightly above midheight marks rejuvenation by
calicinial budding. Beattie limestone, in sec. 15, T. 31 S., R. 8 E., near Grand Summit, Kans. (Univ.
Kansas no. 23292).

Leonardophyllum acus, n. gen., n. sp., from the lower part of the Leonard series at saddle north of
Leonard Mountain, Glass Mountains, north of Marathon, Tex.
3—Apical part of corallite, the top being a polished section (plate 2, figure 5). The
transversely wrinkled but otherwise smooth nature of the surface of the corallite is shown. (Univ.
Kansas no. 74166).

Leonardophyllum distinctum, n. gen., n. sp., from the lower part of the Leonard series on the Hess
ranch, Glass Mountains, north of Marathon, Tex.
4—Side view of typical specimen showing cylindrical form and the nearly smooth,
transversely corrugated surface. (Univ. Kansas no. 74165).

Heritschia girtyi, n. gen., n. sp., from the upper Florence limestone in sec. 23, T. 27 S., R. 6 E.,
Butler County, Kansas.
5—Vertical view of the deep calyx of the specimen shown in plate 7, figure 1a, showing the
sharp-walled appearance of the pit and the laterally compressed form of the column. (Univ. Kansas
no. 77551).

Timorphyllum simulans, n. sp., from the lower part of the Leonard series at saddle north of Leonard
Mountain, Glass Mountains north of Marathon, Tex.
6—Side view of a corallite showing the nearly smooth, transversely wrinkled surface.
(Univ. Kansas no. 7416-22a).

Duplophyllum septarugosum, n. sp., from the Leonard series at Clay Slide, 2 miles west of Iron
Mountain, Glass Mountains, north of Marathon, Tex.
7—Side view of specimen showing transverse corrugations and fine growth lines. The
position of transverse sections shown on plate 6 is indicated. (Univ. Kansas no. 74143).

New Permian Corals from Kansas, Oklahoma, and Texas, by Raymond C. Moore and Russell M.
Originally published July 1941 as Kansas Geological Survey Bulletin 38, Part 3.
Available online at
June 2007