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New Permian Corals

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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 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 calyx.

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, Texas.


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 colum 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 corallite.

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.

Explanation of Plate 1. [A higher-resolution PDF version of this figure is available.]
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 sections.
3a-d--Specimen (Univ. Kansas no. 24842). a, Longitudinal section. b-d, Transverse sections.
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.

line drawings of specimens

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 here.

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. proliferum.

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 paleontologiss 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 longitundinal 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.

Explanation of Plate 2. [A higher-resolution PDF version of this figure is available.]
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 sections.
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 sections.
5--Combined transverse and longitudinal sections of specimen (Univ. Kansas no. 74166).

line drawings of specimens


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 from Leonardophyllum 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 frorn 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 of Lophophyllidium, 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 Oklahoma.

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.

Explanation of Plate 3. [A higher-resolution PDF version of this figure is available.]
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 sections.

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 sections.

line drawings of specimens

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 hav- ing 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 Stylidophyllum from 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 of Waagenophyllum 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 to Heritschia include Corwenia densicolumella Dobrolyubova (1936, p. 140), C. cf. lipoensis Huang (Dobrolyubova, 1936, p. 142), Waagenophyllum chitralicum Smith (1935, p. 37), W. muricatum Douglas (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. indicum from 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 U.S.S.R.

Explanation of Plate 4. [A higher-resolution PDF version of this figure is available.]
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).

line drawings of specimens


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 Kansas.

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 diff rs 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.

Explanation of Plate 5. [A higher-resolution PDF version of this figure is available.]
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 sections.

line drawings of specimens


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), 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 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 of Sochkineophyllum, 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.

Explanation of Plate 6. [A higher-resolution PDF version of this figure is available.]
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 sections.
4a-d--Specimen (Univ. Kansas no. 74144). a, Longitudinal section. b-d, Transverse sections.

line drawings of specimens

Explanation of Plate 7. [A higher-resolution PDF version of this figure is available.]
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).

line drawings of specimens

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 septa.

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 of Duplophyllum.

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 of Duplophyllum 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.

Explanation of Plate 8. [A higher-resolution PDF version of this figure is available.]
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).

line drawings of specimens


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 section.

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 manner.

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.

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Kansas Geological Survey, Geology
Placed on web June 18, 2007; originally published June 1941.
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