New Permian Corals from Kansas, Oklahoma, and Texas

8/14/2019 New Permian Corals from Kansas, Oklahoma, and Texas

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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.

Abstract

Nine new species of Permian rugose corals are referred to eight genera, of which three are new and three havenot previously been reported from America. New species from Lower Permian (Wolfcamp) beds of Kansas andnorthern Oklahoma are assigned to MalonophyllumOkulitch and Albritton, known elsewhere from the Leonardseries of western Texas; LophophyllidiumGrabau, seemingly a long-ranging genus of Upper Carboniferous andPermian age and world-wide distribution; SochkineophyllumGrabau, previously reported from MiddleCarboniferous (Moscovian) rocks of Russia and Permian strata of China; and two new genera namedLophamplexusand Heritschia. New species of Middle Permian (Leonard) corals from the Glass Mountains, inwestern Texas are referred to TimorphyllumGerth, described from the Permian of Timor, in the East Indies;DuplophyllumKoker, previously reported from the Permian of Timor, Australia, and China; and a new genus thatis designated as Leonardophyllum. Most of these forms bear a distinct axial column and all but one speciesseem to lack dissepiments.

Permian corals from Asia that heretofore have been referred to CorweniaSmith and Ryder, a genus that is

based on Lower Carboniferous corals, are placed in the new genus Heritschia, and a few other, variouslyidentified corals are also thought to belong in Heritschia.

The common Late Paleozoic column-bearing corals of North America that have been classed as belongingto Lophophyllumseem to belong to the genus Lophophyllidium. The stratigraphic usefulness of corals incorrelation of Permian rocks is considered briefly. Numerous taxonomic problems similar to that of so-calledLophophyllumare encountered in work on the Permian corals.


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Introduction

Corals are widely distributed in marine Permian deposits of the world and studies on these fossils arecomparatively numerous. Some of the published works are elaborate monographs, reflecting both the richnessof the coral faunas and the extensive nature of researches on them. In North America, however, less than adozen 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); fromChina, by Grabau (1922, 1928, 1931, 1936), Chi (1935, 1937, 1938), Yu (1934), and Huang (1932, 1932a); fromTimor, 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), andYakovlev (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? texanumandPolycoelia? 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 westernTexas, 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? texanumShumard, Cladopora spinulataGirty, n. sp., C.? tubulata Girty, n. sp., andAulopora sp.Some of these are not recognizable on the basis of the published descriptions and illustrations but possibly maybe made so by study of his types and supplementary specimens. One of us (R.C. Moore) recently has hadopportunity to examine Girty's types of Guadalupian corals, but lack of sections prevents diagnosis, at least inthe case of the rugose forms. A small horn coral from Permian limestone in northern Mexico has been describedby Haack (1914) under the name ofCyathaxonia girtyi, but because definitive characters are not indicated, thisform 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) havebeen published by Heritsch (1936a, 1936c). A few silicified corals from the Malone Mountains in western Texashave been described by Okulitch and Albritton (1937) as the type species, Malonophyllum texanum, of a newgenus. A paper by Heritsch (1937b, p. 315) treating on the zonation of Permian rocks by means of rugose coralsmentions 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 ofa description known to J.W. Skinner, for whom the species presumably was named. It has not been possible tocommunicate 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, asLonsdaleia?n. sp.

In British Columbia, good specimens of corals have been collected from rocks thought to be UpperCarboniferous 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 relativescarecity in fossil collections. Many localities that have yielded abundant Permian marine fossils of several sortslack corals, or they have provided only a few specimens that may be none too well preserved. This observationis emphasized by an inspection of the large collections from Permian formations of Texas in laboratories of theU.S. Geological Survey and the University of Texas, or those from the northern midcontinent region in theUniversity of Kansas and the University of Nebraska.


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Stratigraphic Significance of Permian Corals

Only locally and exceptionally in North America are Permian corals abundant. Seemingly, the known Permianinvertebrate faunas of this continent are much less coralline than those of many other regions. Little attentionhas been given to search for these fossils, however, and special efforts in field collecting will doubtless result inimportant additions of study material. Also, it is likely that all together there are many unstudied specimens incollections of various surveys, universities, and museums. Careful investigation of all obtainable fossils shouldaugment 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 thePermian 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 wellestablished through verified and amplified information on the nature and occurrence of corals. This group offossils may then become entitled to rank with ammonoids, fusulinids, and other especially useful guide fossils inthese 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 chooseparticular features of habitat, but, in addition, Hill (1938a) has called attention to existence of different types ofcoral 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. Calcareousshale, 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 agitatedconsiderably 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 somewhatindependent calibration of each type and imposes need for as complete intercorrelation as possible in order toestablish 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 ordefinition of coral zones in the Permian succession of North America. Attack on such matters is premature at thistime. It is intended merely to describe and illustrate several of the corals that have been studied, and to indicatesome of the taxonomic problems that have been encountered in efforts to classify American species. SeveralPermian 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 obtainmaterial for a volume to be published on the Permian rocks of the central and southwestern part of the UnitedStates. Since the beginning of the project, the number of fossil corals available for study has increasedconsiderably, but because the additions came after most of the manuscript for this paper was completed, wehave decided to publish the results now obtained, taking notice of the new material only in so far as it belongs tospecies already studied. It is hoped that work on the yet unstudied specimens can be pursued actively and thatour 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, andWord deposits in the Glass Mountains, western Texas. The Kansas specimens include material collected byvarious geologists of the Kansas Geological Survey during years of mapping and stratigraphic field work, a smalllot of corals from Grand Summit, Kans., in a collection obtained from J.W. Mickle, of Wichita, and a largecollection of corals from the Florence limestone in Butler County, Kansas, that was made after beginning thisstudy. The west Texas corals were collected by R.C. Moore on trips to the Glass Mountains in 1939 and previousyears, and by R.H. King, J.W. Skinner, C.C. Williams, and us on a trip in March, 1940. Determination ofstratigraphic horizons is precisely made for all the Kansas collections on the basis of very detailed subdivisionand tracing of the Permian rocks in this region. Stratigraphic placement of the western Texas specimens rests onthe field studies and mapping of the Glass Mountains by P.B. and R.E. King (1930), supplemented in part bypersonal acquaintance of Skinner with the geologic section of the Glass Mountains.


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Methods of Study

External features of the Permian rugose corals serve only in small part as basis for classification, and in sometypes they are almost negligible in taxonomic study. Likewise, the mode of growth, whether solitary or colonial, isbelieved not to be an important distinction, unless such characters are firmly fixed. Judgments of the taxonomicvalue of mode of growth and of various structural characters of corals differ widely. Variability of the coralsthemselves, conjoined with the uncertainty of paleontologists as to significant evolutionary trends anddivergences 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 werecut transversely at selected points. Except in a few cases these sections served to establish the position of thecardinal-counter septal plane and to indicate differentiation of major and minor septa. The sections werephotographed at enlarged scale after polishing or immersion in oil in order to bring out structures clearly. Bymarking the septal grooves of major septa the arrangement and order of introduction of the septa weredetermined on the exterior of the coral if this was possible. This method usually serves for identification of thealar septa, which may not be independently identifiable from study of the transverse sections. The segments ofthe transversely cut corals were carefully assembled in proper orientation (by means of index marks andmatching septal grooves) and were cemented together. They were then cut longitudinally in a planeapproximately at right angles to that of the cardinal and counter septa. The longitudinal section was oiled andphotographed, 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 thephotographs 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 ispartly due to the fact that half-tone or full-tone figures, which are required for reproduction of photographs, aremuch more expensive than line cuts. Another item for consideration is the lack of clarity of many unretouchedphotographs 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 toverify identification of significant structures in the sections. Irrelevant but frequently confusing details such ascleavage lines in calcite and distributed foreign matter are eliminated in making the drawings. Thin sections orcelluloacetate 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 showstructures 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 makethin 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 areexcessively technical and include a great number of precisely defined individual terms. We have adopted asomewhat simpler, less concise terminology in order to avoid the use of terms not generally understood. It istherefore 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 waspresumed to be a layer deposited upon the original wall, or theca. In well-preserved specimens of rugose coralsGrabau (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 designatingthis layer as the theca.

Septa--The terminology of the classes of septa is in a very confused state. We use the following designation ofthe septa, as given by Hill (1935, pp. 504-505), because these terms have been used in a consistent mannerand 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 septaadded in pairs in the cardinal and counter quadrants and closely resembling the protosepta are designated themetasepta. The protosepta and the metasepta together are termed the major septa. Short septa alternatingwith the major septa, introduced later than the majors, are termed minor septa.


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Axial structures--Both Grabau (1922) and Hill (1935) have proposed detailed classifications of the axialstructures of rugose corals. These authors differ widely in terminology, however, and neither has been followedclosely by many other workers. Although Grabau states that a true columella is found only in the Hexacoralla, hedescribes the solid rodlike axial structure of the rugose corals as a columella. The term pseudocolumella is usedby him to include all non-solid types of axial structures, most of which are given descriptive names. Hill uses theterm columella for a solid rodlike axial structure and restricts the terms pseudocolumella to the Hexacoralla. Thegeneral term axial complex includes both the columns formed by the twisting of axial ends of the septa (axialvortex) 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 andprecise, but we do not think it necessary to use these little-known complex technical terms. Because the namesnow used in describing axial structures are in such a confused state, we designate any columnar structure that ispresent in the axial region of the corallite as a column and describe the features of each in detail. The columnmay or may not be solid.

Acknowledgments

We are indebted to Ralph H. King and Charles C. Williams, of the University of Kansas, for aid in collectingspecimens, and to Maurice Wallace, of the University of Kansas, for work in sawing and polishing some of thesections of corals. Ruth Mary Dudley, of the Kansas Geological Survey staff, assisted in supervising work ofN.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 beennecessary to study early and middle Paleozoic coral genera in order to give foundation for work on Permianforms, the organization of information from previously published papers has been one of the largest items oflabor 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 ofTexas, and to Carl O. Dunbar, Peabody Museum, Yale University, for the loan of specimens, which, however, aremostly unstudied at the present time. Robert E. King has sent us some material from the Wolfcamp beds in theGlass Mountains, and John W. Skinner has aided us, both by guidance in some of the field collecting in westernTexas and by providing information concerning fossil localities in the Permian rocks near Fife and Dothan innorth-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 thusable 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, wewish to acknowledge the kindness of James Steele Williams, of the U.S. Geological Survey, in giving opportunityto 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 representedin a grant to defray photographic costs of materials for the illustrated catalog of corals and for the preparation ofdrawings given in this paper.


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Systematic Descriptions

General Statement Concerning Taxonomy

Classification of Paleozoic corals is now in confused, if not, almost chaotic state. It is generally recognized by thebest-informed students of these fossils that organization of described genera in families has been based verylargely on superficial and even artificial lines and that we are yet far from ready to evaluate with reasonableaccuracy 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 Palontologieand later works, include some very slightly related forms, but there can be no doubt as to the polyphyletic natureof 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 areshown to be derived from wholly different, unrelated stocks, and on this basis he follows Weissermel (1897) inmaking new genera out of segments of the stratigraphically almost meaningless assemblage calledAmplexus.Convergence of characters in adult forms is shown to extend to internal features, such as arrangement and formof the septa, as well as external features, and knowledge of youthful growth stages is necessary in order torecognize the very unlike origins of seemingly identical end forms. A few authors, such as Grabau, recognizefinely drawn generic distinctions, of which many may be sound and probably some may be unsound. Manyfamilies are proposed. Other authors, such as Hill (1939), are inclined to reduce to synonymy a very largenumber of the listed Paleozoic coral genera and to group these in a small number of families. Lang, Smith, andThomas (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 increasingcomplexity 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 ofLophophyllidium, except

that tabulae are absent. Longitudinal sections through the axial column of some specimens indicate a structurelike that ofLophophyllidium, consisting of superposed tabellae-like cones having nearly vertical sides builtaround the distal part of the counter septum. The column projects upward strongly into the lower part of thecalyx.

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

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

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


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MALONOPHYLLUM KANSASENSE., n. sp.

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 theplane of the alar septa. The theca is moderately thick, its surface being marked by deep septal grooves anddistinct interseptal ridges that are crossed by small wrinkles and growth lines. The calyx is deep, containing aprominent 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 aslong as the major septa. Two other specimens show 20 and 22 major septa, respectively. The cardinal septum isshortened so as to form an open fossula. The counter septum extends to the axis and thickens to form a solidcolumn in the young portion of the corallite, but near the calyx the counter septum is not significantly longer thanother major septa. Dissepiments and tabulae are absent. A solid rodlike column in the lower portion of thecorallite is formed by the thickened end of the counter septum. It tapers gradually upward and becomes free inthe lower part of the calyx.

Transverse sections of the early parts of the corallites show the structural elements much thickened bystereoplasm. The septa are joined at their axial ends to form a large solid column. Successive sections higher inthe 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 thecorallite.

Discussion--The description and illustrations of the weathered specimens ofMalonophyllum texanum Okulitchand Albritton (1937) indicate that the septa withdraw only slightly from the axial column in the upper part of thecorallite and that they are strongly deflected toward the counter septum.Malonophyllum kansasense can be distinguished readily from species ofLophophyllidium and Lophamplexus bythe concentration of structural elements and stereoplasm in the lower part of the corallite, and by the absence ofdissepiments and tabulae. The form of the corallite ofM. kansasense is distinctly conical, in contrast to theconical-cylindrical shape ofLophophyllidium dunbariand Lophamplexus eliasi, and it reaches maximumdiameter 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 ofvarying strength. The calyx is moderately deep, bearing near its center a laterally compressed column that formsa more or less prominent projection. Septa are straight and mostly thin, the counter septum joined to the axialcolumn, 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 solidrodlike structure of laterally compressed form, composed of a thickened portion of the counter septum and ofvery steeply upbent extensions of the tabulae, thickened by stereoplasm.

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

Discussion--The classification and nomenclature of the relatively simple, column-bearing horn corals that arewidely distributed in Carboniferous and Permian deposits have long been in a confused state, and the studiespublished by Carruthers (1913), Grabau (1928), Huang (1932), Yii (1933), Stanley Smith (1934), Heritsch


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(1936b), Chi (1938), and Hill (1940)--not to mention numerous others--incompletely resolve the difficulties. Thewidely accepted reference of American Upper Carboniferous and Permian corals to Lophophyllum seems to beerroneous. Only brief treatment, touching the most pertinent aspects of these taxonomic problems, is offeredhere.

The genus Lophophyllum was proposed by Edwards and Haime (1850) on the basis of a Lower Carboniferouscoral 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 ofLophophyllum was a

synonym ofCyathaxonia tortuosa Michelin (1846), described from the same horizon and locality. Thorough studyby Carruthers of topotypes ofLophophyllum tortuosum establishes the significant characters of this genus, asfollows. The corallites are solitary, turbinate, and enclosed by a theca; major septa meet at the center in earlygrowth stages, one of them, generally the counter septum, is thickened at the inner end so as to form amoderately 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 maturepart 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 lamellaedistinct from the septa.

In 1876 Thomson and Nicholson (p. 297) proposed a new genus, Koninckophyllum, that was based on a newspecies called Koninckophyllum magnificum, from Lower Carboniferous rocks of England. This genus wasregarded 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 "aconsiderable space" between the axial column and the inner margins of the major septa. Thomson andNicholson (1876, p. 298) state that "in no case do the tabulae extend to the inner surface of the wall." Amongspecies assigned to Koninckophyllum are some that are definitely of the Lophophyllum tortuosum type, andfollowing Carruthers' (1913) restudy of the genotype ofLophophyllum, some workers have concludedthat Koninckophyllum should be suppressed as a synonym ofLophophyllum. On the other hand, Smith (1934),Vaughn (1915), Heritsch (1936b), Hill (1939), and other students of these corals regard the two genera asdistinct, although closely related. Grabau (1928) and Huang (1932) arbitrarily and incorrectly place corals similarto the genotype ofLophophyllum L. tortuosum, in Koninckophyllum and use Lophophyllum for corals similar to L.

proliferum.

Edwards and Haime in 1851 (p. 323) described an American column-bearing small horn coral from beds of earlyPennsylvanian 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, namingit 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. Americanpaleontologists generally have come to regard L. proliferum as a synonym ofL. profundum (Weller, 1898, p.333), though this may well be an erroneous conclusion. It is very doubtful indeed that L. proliferum occurs inPermian 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 axialcolumn. 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 ofC.prolifera are needed in order to determine satisfactorily the characters that should be assignedto Lophophyllidium. Our studies of Pennsylvanian and Permian corals of this sort, having a solid orcomparatively dense axial column, indicate the existence of several types of internal structures that may havetaxonomic significance. Based on present knowledge, Lophophyllidium is tentatively interpreted to includesolitary horn corals having a compressed column that is connected with the counter septum throughout most ofits 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, andlacking dissepiments.

In 1928 Grabau (p. 99) differentiated under the name Sinophyllum forms of lophophyllid corals having a thickaxial column joined to the counter septum so as to produce a pendulum-like appearance in transverse sectionsof the corallites, ends of the longer septa also being deflected and joined to one another in such manner as toform an inner wall nearly surrounding the column. Grabau designated Lophophyllum pendulum, described by


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approach close to the column, they retain their structural identity. In the mature stage the rhopaloid nature of thesepta disappears and they become progressively shortened so as to assume an amplexoid form. Tabulae areentirely absent in the mature stage. In the uppermost sections the counter septum is indistinguishable from theother 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 dunbaridiffers from Lophophyllidium proliferum (McChesney) (1860) in the proportionatelysmaller diameter of the column and in the less curved form of the corallite. Information as to the internal featuresof the genotype species, originally described by McChesney, is lacking.

Lophophyllidium dunbaridiffers from Lophophyllidium amygdalophylloidea Huang (1932) in its smaller axialstructure, much shorter minor septa, and fewer tabulae in the longitudinal sections. Lophophyllidiumzaphrentoidea Huang (1932) seems to lack numerous tabulae, and it has longer and more numerous septathan L. dunbari. Also, L. zaphrentoidea has a more conspicuous cardinal fossula, and it seems to have nodistinct 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 ofKansas. A leader in paleontologic and stratigraphic research, Professor Dunbar is especially noted on account ofcontributions to knowledge of Permian fossils and rock formations. He has aided our study of the Permian coralsby 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 , CowleyCounty, Kansas.


Genus LEONARDOPHYLLUM, n. gen.

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. Exceptthe counter septum, which is joined to the axial column, the septa reach only part way to the column. The majorsepta are evenly disposed and approximately uniform in length, except the cardinal septum, which isdistinguished by its short length and by its position opposite the counter septum. Minor septa occur between themajor ones.

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

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

Discussion--The described structural features ofLeonardophyllum suggest comparisonwith Clisiophyllum Dana, Dibunophyllum Thomson and Nicholson, Carcinophyllum Thomson and Nicholson, andespecially Verbeekiella Gerth. All these genera have an axial column that is characterized by arched tabulae ortabellae intersected by radial lamellae, producing a cobweb pattern in tranverse sections. The genotype speciesofClisiophyllum, Dibunophyllum, and Carcinophyllum, and other species that are assigned without question tothese genera, are distinguished from the Permian corals here considered by a common occurrence ofdissepiments and by other features that need not be reviewed here in detail. Permian species that have beenassigned to the genera mentioned (Clisiophyllum australe Beyrich, C. torquatum Rothpletz, C.


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talbotiHosking, Dibunophyllum rothpletziGerth, D. tubulosum Gerth, and Carcinophyllum cristatum Gerth) areall assigned by Hill (1937, p. 54) to Verbeekia (equals Verbeekiella Gerth, the name Verbeekia Penecke being ahomonym). Also, Hill thinks that "Verbeekia" is not closely related to the Lower Carboniferous clisiophyllids, andprobably not derived from them. Verbeekia permica Penecke (1908), which was designated as genotypeofVerbeekia [Verbeekiella], comes from the same locality in Timor as that which yielded the type specimenofClisiophylIum 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 ofC. australe. Accordingly, the genotypeofVerbeekiella may be designated as Clisiophyllum australe. The axial column ofVerbeekiella seems to

resemble that ofLeonardophyllum, but closer inspection shows important distinctions. None of the illustrationsgiven by Beyrich, Rothpletz, Penecke, or Gerth forC. australe show a connection between the column and thecounter septum, and all indicate occurrence of a distinct wall surrounding the column, so as to separate sharplythe closely spaced arched tabellae and radial lamellae of the column from the differently disposed tabulae andsepia outside the axial column. In Leonardophyllum there is no definite boundary between the central columnarstructure and the rest of the corallite. The arched or conical laminae of the columnar structure are parts of thetabulae. 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 thatofLeonardophyllum as regards extension and steep upward deflection of tabulae in the central area, but radiallamellae are lacking in the so-called lophophyllids.

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

LEONARDOPHYLLUM DISTINCTUM, n. sp.

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 andwrinkles but no septal grooves. The incomplete type specimen is 21.2 mm in length and 12.3 mm in diameter. Acone-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 longercounter septum. The major septa have a length equal to about one-fifth the diameter of the corallite. The minorsepta 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 risesteeply 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 rarelyother septa.

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

The thick theca and strong septa of the specimen identified by Dobrolyubova (1936, p. 132)

as Verbeekiella cf. rothpletziGerth resemble those ofL. distinctum. The transverse sections of the Russianspecimen, however, show a different septal development and a column formed by a few very thick radiatinglamellae without a cob-web appearance.

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

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


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LEONARDOPHYLLUM ACUS, n. sp.

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. Thecalyx is not known. The incomplete type specimen is 17.7 mm in length and has a maximum diameter of 11.0mm at the uppermost part of the fragment.

The strong major septa thicken peripherally. The cardinal septum is only slightly shortened but the counterseptum reaches the column and is directly connected to the median lamina. Other major septa reach the columnand may be slightly twisted in the early stages, but in maturity withdraw from the column and have a length ofabout 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 thecolumn, 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 transversesection the column has a well-defined cob-web appearance caused by the intersection of the tabellae with themedian lamina and the few radiating lamellae.

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

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


Genus LOPHAMPLEXUS, n. gen.

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

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

Discussion--The corals here described are evidently related closely to Lophophyllidium, as indicated bycorrespondence 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 thetendency of the counter septum in the mature region to become even shorter than the adjacent major septa. Thecardinal septum is notably shortened in all stages of growth except the earliest. This reduction in length forms adistinct but not prominent open fossula.

The distinctly amplexoid part of the corallite, as observed in specimens assigiled to Lophamplexus, seems to beless 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 toparent stocks that are radically different, one from another, and the amplexoid nature of the mature stages ofeach of them is a result of the evolutionary modification that Schindewolf calls convergence. Lophamplexus isindicated to have been derived from an ancestor belonging to Lophophyllidium or a closely related genus.


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Occurrence--Foraker and Beattie limestones, Wolfcamp series, Lower Permian, Kansas and Oklahoma.

LOPHAMPLEXUS ELIASI, n. sp.

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

The principal structural features of this species are described in the generic description. The conical-cylindricalcorallite 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 specimenhas 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 showsno minor septa. Another specimen has very short minor septa alternating with the major septa in a section justbelow the top of the column-bearing stage. An open cardinal fossula is visible in all but the most mature parts ofthe corallite. Tabulae are incomplete in youthful stages, but above the column some are complete. The completetabulae are horizontal in the axial region but they slope downward and outward somewhat steeply near theperiphery. A column is present in the lower 43 percent of the length of the type, and in the lower 69 percent of aspecimen measuring 22.6 mm in length.

Discussion--This species is easily distinguished from described species of lophophyllid corals by the absence ofthe column in the mature portion of the corallite. Transverse sections of the juvenile portions resemble those

ofLophophyllidium dunbari, and it is to be noted that sections of the upper parts of the calyx of specimens ofL.dunbariand 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 coralthat was identified asAmplexus cf.abichiWaagen and Wentzel by Heritsch (1937a, p. 4) and other species of"Amplexus" lack a prominent column in any part of the corallite.

Lophamplexus eliasiis named for Maxim K. Elias, of the Nebraska Geological Survey. Much work has beendone by Elias on Permian rocks and fossils, especially during the period of about ten years while he was amember of the Kansas Geological Survey.

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

of Fairfax, Oklahoma.


Genus HERITSCHIA, n. gen.

This genus includes compound corals, individuals being of conical-cylindrical form, growing in sub-parallelposition 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 developedirregular cross-wrinkles. The depth of the calyx is equal to about one-half the diameter of the corallite, the sidessloping evenly to a central pit that holds a prominent, flattened, and sharply pointed axial column. Septa arenumerous, evenly disposed, and nearly equal in size, the major ones extending only slightly farther toward thecenter in the mature regions than the minor ones. A tetrameral arrangement of the septa is not observable in thecalyx, 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 theexterior of the theca. The septa are thin and strongly flexuous in the peripheral part of the corallite, but thickenedand relatively straight in the inner part. The counter septum is joined to the column but all other septa are slightlyseparated from it.

A peripheral zone, equal in width to about one-half of the radius, bears numerous dissepiments, their inwardlyconvex surfaces sloping steeply upward and outward. The inner margin of the dissepimental zone, as seen intransverse sections, is marked by more closely spaced and somewhat thickened dissepiments that, combinedwith 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


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spaced, partly anastomosing tabulae that slope steeply upward and outward, except for narrow subhorizontalportions 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 thisplane. Commonly, the column seems to be composed of a solid deposit of stereoplasm, but sections of severalspecimens reveal a dibunophyllid structure consisting of a few radial lamellae that converge to meet the medianlamina at different points. These structures are crossed by steeply arched tabellae that slope nearly verticallyupward and inward. The septa and the median lamina of the column are fluted along curved lines than runparallel 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 ofHeritschia areintermediate between those ofWaagenophyllum Hayasaka and Iranophyllum Douglas, on the one hand, andsuch forms as Lonsdaleia McCoy and Stylidophyllum Fromentel, on the other. Waagenophyllum, as indicated byits genotype species, Lonsdaleia indica Waagen and Wentzel, from the middle Productus limestone of the SaltRange, India, comprises compound corals of closely bundled (phaceloid) habit, containing numerous smallclosely 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 theaxial column. Also, as in Heritschia, there is an outer zone of dissepiments sloping outward and upward, anintermediate zone of tabulae similarly inclined except at their inner margins, and in the axial region closelyspaced tabellae that slope upward and inward. Unlike Heritschia, however, the septa ofWaagenophyllum are

strongest in the peripheral zone, next to the theca, and the tabellae and radial lamellae of the column are muchless obscured by stereoplasm than in Heritschia. The axial structure ofWaagenophyllum resembles thatofDibunophyllum. Difference in deposits of stereoplasm in the axial region is not deemed to have genericsignificance, and accordingly the chief distinction between Waagenophyllum and Heritschia, based oncomparison of their genotype species, is found in characters of the peripheral zone, where the externallythickened septa and relatively narrow belt of dissepiments in Waagenophyllum contrast with the thin flexuousouter parts of the septa and the strongly developed dissepimental zone ofHeritschia.

Iranophyllum, which is based on the species Iranophyllum splendens Douglas, was established for solitaryrugose corals having the internal structure ofWaagenophyllum and Wentzelella. The septa reach the theca, andin general the tabulae are inclined proximally towards the axial column in ihe same manner asin 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 genotypespecies, Lonsdaleia rugosa McCoy, differs essentially from Heritschia in the attitude of the tabulae and in theexpanded 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 ofclosely spaced dissepimental tissue they are very thin and do not reach the theca. A dibunophyllid column isseparated by a short space from the inner margins of the septa. Some Permian corals have been assignedto Corwenia, but as interpreted on characters of the genotype and associated Lower Carboniferousspecies, Corwenia is clearly unlike the Permian forms. It has a median zone of tabulae that rise towards thecolumn and are convex upwards, whereas tne tabulae of the median zone in Permian species that have beenreferred to Corwenia slope distinctly downward and inward, as in Iranophyllum and Waagenophyllum. The majorsepta of the Permian species are not extended to the central column as is the case in the type species, and theminor septa are in general much more prominent than in Corwenia rugosa. Smith (1935), Douglas (1936), andHill (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 closepacking of the corallites ofStylidophyllum, which imposes a prismatic form and polygonal outline on individuals,is a main feature in distinguishing Stylidophyllumfrom Heritschia. The colonial growth form ofStylidophyllum isunlike that of the very loosely packed corallites ofHeritschia, but partial correspondence of internal structuresand seeming relationship ofStylidophyllum, Iranophyllum, Heritschia, and Waagenophyllum suggest theapproximate equivalence in age ofStylidophyllum and Heritschia. Undoubted representativesofWaagenophyllum seem to occur only in beds of Middle and Late Permian age.


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On the basis of published figures and descriptions, we refer to Heritschia the Permian species Corweniachiutsingensis 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 Corweniadensicolumella 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 groupthat is distinct from the typical forms ofWaagenophyllum, inasmuch as the peripheral parts of the septa arenotably reduced in thickness, tending in some cases to disappear. The subhorizontal or somewhat upward andinward sloping attitude of the tabulae ofHeritschia columbica is more pronounced than in H. girtyiand thedibunophyllid column is both broader and less obscured by steroplasm, but essential structural featurescorrespond closely. Although W. chitralicum has a prominent stereozone, like that ofHeritschia, thickening of theperipheral parts of the septa indicate closer relationship with Waagenophyllum. W. katoidiffers from Heritschia inthe 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, thezone ofWaagenophyllum is the upper middle part of the Permian system. W. indicumfrom the Middle Productuslimestone of the Salt Range in India certainly represents a horizon far above the base of thePermian. 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 lowermostPermian. Heritschia girtyi, on the other hand, comes from beds of late Wolfcamp age. The Florence limestone ofKansas has also yielded Pseudoschwagerina (Moore, 1940, p. 314). On the basis of this and much otherpaleontologic and stratigraphic evidence, the placement ofH. girtyiin the upper part of the succession of beds inthe Wolfcamp series, generally classed by American geologists as Lower Permian, is well fixed. This is not thecase in referring to H. columbica, inasmuch as the rocks that yielded this fossil are identified as "Permian orpossibly Upper Carboniferous." It is reasonable to infer that they are Lower Permian, and this seems to applyalso to the rocks containing other foreign species that are tentatively assigned to Heritschia. Stratigraphicdistribution, as well as paleontologic grounds, is thought to give basis for separationofHeritschia from Waagenophyllum.

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

Occurrence--Wolfcamp series, Lower Permian, Kansas; Lower Permian (?), British Columbia, Japan, Iran, andU.S.S.R.

HERITSCHIA GIRTYI, n. sp.

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. Thelarger 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 typespecimen has 30 major septa and a like number of minor septa. The major septa are of equal length, except forthe counter and cardinal septa, and the minor septa are nearly as long as the major. An inconspicuous fossula isformed by the shortening of the cardinal septum. The structure of the septa, column, tabulae, and dissepimentsare described under discussion of the genus.

Discussion--The strongly flexuous septa and complex dissepimental area, which are distinctive featuresofHeritschia girtyi, are observed also in Heritschia columbica (Smith). The species from Canada has only aslight indication of a stereozone, however, and its lacks the compact thickened column. The Chinesespecies, Heritschia lipoensis (Huang, 1932), and Heritschia parachihsianensis(Huang, 1932), have a distinctly


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open column and relatively straight septa. The peripheral dissepimental zone is less well developed and minorsepta 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 acooperative study of the El Dorado oil field by the U.S. Geological Survey and Kansas Geological Survey. Thespecimens were examined by Girty who identified them as Lonsdaleia?n. sp. The corals that were collected byFath have not been seen by us. A small number of specimens was collected by M.K. Elias in 1932 while workingfor 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 otherspecies of coral, which has not yet been thoroughly studied, is now known to be associated with Heritschiagirtyiin the Florence limestone. Several exceptionally well-preserved specimens were collected by A.I. Levorsenof Tulsa, Okla., from a locality about a mile east of Leon. These were kindly cliven by him to the University ofKansas.

This species is named in honor of the late George H. Girty, of the U.S. Geological Survey, who for many yearsenriched 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 NESE 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.


Corals Not Bearing a Distinct Axial Column

Genus TIMORPHYLLUM Gerth, 1921

Small, solitary corals of somewhat irregular elongate cylindrical form are typical ofTimorphyllum. The theca isnot marked by septal grooves but is crossed transversely by growth lines and wrinkles. A short cardinal septumlies in an open fossula; the counter septum is elongate, reaching through the center of the corallite, but its distalpart is not distinctly thickened to form a column. Major septa are short, extending inward about one third thediameter 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. Dissepimentsare absent.

Genotype--Timorphyllum wanneriGerth, Permian of Timor.

Discussion--The described structural features of this small coral readily distinguish it from other types ofPermian corals. Gerth's interpretation of the axial region as consisting of a much flattened columella that isattached to the counter septum is supported by observation that a central structure projects upward in the lowerpart of the calyx. This feature is not shown in our material, however, and transverse section of the corallite showa 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 Artinskor Leonard and Word), and Leonard series, Middle Permian, West Texas.

TIMORPHYLLUM SIMULANS, n. sp.

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

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

Discussion--This species differs from Timorphyllum wanneriGerth (1921) in that the theca is less wrinkled, and


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the peripheral slope of the tabulae is steeper. T. simulans can be distinguished from T.wannerivar. variabilis Gerth (1921) by its lack of a definitely separated column and the essentially equal lengthof 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.

Genus SOCHKINEOPHYLLUM Grabau, 1928

This genus contains solitary conical corals of medium size having a longitudinally grooved theca. The calyx ismoderately deep, not containing a columnar projection. Internally, however, the counter septum is observed toreach 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 distalregion; other septa are much shorter, the cardinal septum being very short, so as to produce a fairly well markedopen fossula even at an early stage of growth. The counter quadrants are accelerated over the cardinalquadrants. Tabulae of somewhat irregular and slightly anastomosing form extend across the interior of thecorallite, being arched gently upward. No dissepiments are observed.

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

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 maymake 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 nomeans confined to Sochkineophyllum. It is a prominent peculiarity ofTachylasma, Plerophyllum, and some othergenera that are represented in Lower Carboniferous and probably Upper Carboniferous, as well as in Permianrocks. In the genera other than Sochkineophyllum, the distribution of the rhopaloid thickening differs from thatofSochkineophyllum, the unusually elongate counter septum being an especially distinctive featureofSochkineophyllum.

Hill (1940, p. 131) questionably includes Sochkineophyllum as a synonym ofFasciculophyllum Thomson, statingthat 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 ofSochkineophyllumartiense only differs [from "Fasciculophyllum" eruca, which is not the genotype species] in that differentmetasepta become long and rhopaloid." These two genera are similar in having a shortened cardinal septumand an elongated counter septum; also, in both there is an absence of dissepiments, and several pairs of longrhopaloid 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 majorseptum outward from the cardinal septum).

The spelling of this generic name calls for brief consideration. The International Rules of ZoologicalNomenclature (Article 8h) provide that generic names may be based on patronymics and recommend that theexact form of writing the proper name (using Roman letters) be employed. Thus, such generaas Mlleria, Stlia, Kryeria, and Ibaezia are recognized as correctly formulated. Special difficulty isencountered, however, in transliterating some names from Russian, Chinese, and other languages that do notuse 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 tothe same person, and that several divergent spellings of the same Chinese name may be given when written inRoman letters. M.K. Elias has pointed out to us that the paleontologist for whom Sochkineophyllum was namedis a woman and that the surname of Russian women always bears the ending -a, as Soschkina. Publishedspellings 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, publishedin French. Study of the Rules indicates that the spelling used by Grabau is not emendable (Moore, Weller andKnight, 1941).

Occurrence--Upper Carboniferous (Moscovian), China; Lower Permian (Wolfcamp), Kansas; Middle Permian


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(Artinsk), U.S.S.R.

SOSCHKINEOPHYLLUM MIRABILE, n. sp.

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 lengthof 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 theaxis. Twelve other major septa extend nearly to the axis and are thickened distally. These elongated septa arearranged in three groups, each consisting of four similar septa, two located on one side of the counter-cardinalplane and two in corresponding position on the other side of this plane. If a transverse section of the corallite isoriented with the counter septum at the top, one of these groups comprises the second and fourth septa on eachside of the counter septum, another group contains the seventh and ninth septa left of the counter and the sixthand seventh to the right of the counter; the third group consists of the thirteenth and fourteenth septa on the leftand the tenth and eleventh on the right of the counter septum. Other major septa are shorter than the counterand the 12 other long septa, their average length being about one-third that of the counter septum. Minor septaare 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, cardinalseptum, 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 bysteroplasm and thickening at their distal ends. A fossula is formed by shortening of the cardinal septum. Thinincomplete tabulae are seen but there are no dissepiments.

Discussion--No Late Paleozoic species resemblinc, Sochkineophyllum mirabile has been described from rocksof this continent. The new species resembles Sochkineophyllum artiense (Soschkina, 1925) in the elongationand thickening of several of the major septa and the extended nature of the counter septum. S. mirabile has agreater 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 ofelongated 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 ofMalonophyllum 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.


Genus DUPLOPHYLLUM Koker, 1924

Elongate conical-cylindrical solitary corals having a well-developed theca that externally bears more or lessstrongly developed transverse wrinkles and growth lines but no septal grooves are typical ofDuplophyllum.Septa are comparatively numerous, consisting of two orders, the major ones reaching to the center except nearthe 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 shortsepta that are joined together, an open space separating these from the center, but in the middle and upper partsof the coral this structure disappears and a single cardinal septum is identified extending to the center. Apeculiarity of the genus is the tendency of minor septa to unite at their distal extremities with adjoining majorsepta.

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

Discussion--Several problems are almost immediately encountered in studying Duplophyllum. In the light ofpresent 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 ofCyathophyllum? zaphrentoides do not


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especially resemble figures of the corals from the Permian of Timor that were designated as Duplophyllumzaphrentoides by Koker. Seemingly the latter specimens provided the basis for all her observations and genericdiagnosis. Etheridge's (1891, pl. 10, figs. 4-6) figures ofC.? zaphrentoides indicate a coral that has verynumerous 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, somewhatregularly spaced curved tabulae and a peripheral zone that is characterized by stereoplasmic thickening of allelements. There are numerous regularly spaced connections between septa, probably dissepiments. Onecannot rely too much on the published description and figures ofC.? zaphrentoides by Etheridge, and because

Koker does not compare critically her Timor specimens with examples of the Australian species, availableevidence 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 fromNew South Wales. Because Koker seemingly based the generic description on the Timor specimens, Chi thinksthat Koker's genotype should be designated as Duplophyllum, n. sp. Koker (1924, p. 22) describes the Timorspecimens in the text under the heading Duplophyllum cf. zaphrentoides (Etheridge). In the caption for the textfigure 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, andconsequently 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 beadventitious. It is not a normal structure in corals and is regarded as of doubtful significance. The tendency ofminor septa to unite distally with adjacent majors, shown in at least part of Koker's illustration of a Timorspecimen, 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 fromYunnan, China, called Duplophyllum compactum. Similar fused septa are a characteristic structural feature alsoof some of our corals from the Leonard series of the Glass Mountains. Transverse sections of specimensillustrated by Koker show the notably uneven crooked attitude of some of the septa. This is not seen in Chi'sspecies (D. compactum), but is an especially noteworthy feature of the minor septa and a few of the major onesin the West Texas specimens. The septa of these Leonard corals show a tendency to twist slightly in the axialregion, suggesting the axial structure ofClisiophyllum. No longitudinal sections are given by Etheridge for thegenotype 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 theinternal 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 areno dissepiments.

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

DUPLOPHYLLUM SEPTARUGOSUM, n. sp.

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 growthlines and coarse wrinkles but no longitudinal markings. The theca is thick in all stages of growth. The calyx isunknown. Some of the specimens indicate various stages of rejuvenation, as two definite thecal walls separatedby 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. Inthis section the major septa are equal in length except for the counter septum, which extends to the axis but isnot thickened to form a column. Minor septa are one-third the length of the major, except for one on each side ofthe counter septum that are nearly as long as the major septa. Transverse sections lower in the corallite show


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New Permian Corals from Kansas, Oklahoma, and Texas

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