.. 

Revision of Lithostrotionella 

(Coelenterata, Rugosa) from 

the Carboniferous and Permian 

G E 0 L 0 G I C A L S U R V ~ Y P R 0 F E S S I 0 N A L P A p . E R. l 2 4 7 
Revision of Lithostrotionella 

(Coelenterata, Rugosa) from 

the Carboniferous and Permian 

By WILLIAM J. SANDO 

GEOLOGICAL SURVEY PROFESSIOI'JAL PAPER 1247 

A revision of colonial rugose corals based on 

restudy of Hayasaka~ North American species 

and the literature on other species 

referred to Lithostrotionella 

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON 1983 · 
UNITED STATES DEPARTMENT OF THE INTERIOR 

JAMES G. WATT, Secretary 

GEOLOGICAL SURVEY 

Dallas L. Peck, Director 

Library of Congress Cataloging in Publication Data 

Sando, William Ja.Sper. 

Revision of Lithostrotionella (Coelenterata, Rugosa) from the Carboniferous and Permian. 

(Geological survey professional paper ; 124 7) 

"A revision of colonial rugose corals based on restudy of Hayasaka's North American species and the literature on other spedes 

referred to Lithostrotionella." 

Includes bibliographical references and index. 

Supt. of Docs. no.: I 19.16: 

1. Lithostrotionidae. 2. Paleontology-Carboniferous. 3. Paleontology- Permian. I. Title. II. Series. 

QE778.S24 563'.6 81-607828 

AACR2 

For sale by the Distribution Branch, U.S. Geological Survey, 

604 South Pickett Street, Alexandria, VA 22304 
CONTENTS 

Page Page 

Abstract -------------------------------------------- 1 Systematic paleontology-Continued 

Introduetion ----------------------------------------- 1 Family Lonsdaleiidae Chapman, 1893 ----------------- 36 

Acknowledgments --------------------------------- 2 

The status of Lithostrotionella -------------------------- 2 

Classification ----------------------------------------- 4 

Phylogeny ------------------------------------------- 6 

Genus Thysanophyllum Nicholson and Thomson, 1876 _ 36 

Genus Lonsdaleia McCoy, 1849 ------------------- 37 

Subgenus Actinocyathus d'Orbigny, 1849 ________ 37 

Family Durhaminidae Minato and Kato, 1965----------- 37 

Genus Kleopatrina McCutcheon and Wilson, 1963 ____ 37 

Systematic paleontology-------------------------------- 8 Subgenus Kleopatrina McCutcheon and Wilson, 

Family Lithostrotionidae d'Orbigny, 1851 -------------- 9 1963 --------------------------------- 38 

Genus Stelechophyllum Tolmachev, 1933 ------------ 9 

Genus Aulostylus Sando, 1976 -------------------- 15 

Family Acrocyathidae new family --------------------- 15 

Genus Acrocyathus d'Orbigny, 1849 ---------------- 15 

Family Petalaxidae Fomichev, 1953 ------------------- 23 

Undetermined lithostrotionelloid corals---------------- 38 

Additional taxa ------------------------------------ 39 

Register of USGS localities for Hayasaka (1936) type specimens 

of Lithostrotionella species ------------------------ 40 

Locality data for USNM specimens not ~ayasaka types ______ 41 

References cited -------------------------------------- 42 

Genus Petalaxis Milne-Edwards and Haime, 1852 ____ 23 Index ------·----------------------------------------- 49 

ILLUSTRATIONS 

[Plates follow index] 

PLATE 1,2. Stelechophyllum microstylum (White). 

3. Stelechophyllum banffense (Warren)?. 

4. Stelechophyllum banffense (Warren)? and Stelechophyllum sp. indet. 

5. Acrocyathusjloriformisjloriformis d'Orbigny. 

6. Acrocyathusjloriformisjloriformis d'Orbigny?. 

7. Acrocyathus floriformis floriformis d'Orbigny? and Acrocyathus jloriformis jloriformis d'Orbigny. 

8-11. Acrocyathus jloriformis jloriformis d'Orbigny. 

12. Acrocyathus jloriformis hemisphaericus (Hayasaka). 

13. Acrocyathus jloriformis hemisphaericus (Hayasaka) and Acrocyathus jloriformis hemisphaericus (Hayasaka)?. 

14. Acrocyathus jloriformis hemisphaericus (Hayasaka). 

15. Acrocyathus proliferus (Hall). 

16. Acrocyathus proliferus (Hall) and Acrocyathusjloriformisjloriformis d'Orbigny. 

17. Acrocyathus pilatus n. sp. and Acrocyathus girtyi (Hayasaka). 

18. Petalaxis simplex (Hayasaka) and Petalaxis wyomingensis n. sp. 

19. Petalaxis exiguus n. sp. and Petalaxis tabulatus (Hayasaka). 

20. Petalaxis occidentalis (Merriam) and Lonsdaleia (Actinocyathus) berthiaumi (Merriam). 

Page 

FIGURE 1. Morphology of the type species of the principallithostrotionelloid genera-------------------------------------------- 7 

2. Postulated phylogeny of the principal genera of lithostrotionelloid corals-------------------------------------------- 8 

3. Variation in maximum number of major septa and maximum corallite diameter in 56 coralla of Acrocyathusjloriformis ______ 18 

TABLE 

Page 

TABLE 1. Hayasaka (1936) type specimens of Lithostrotionella ---------------·--------------------------------------------- 3 

III 

REVISION OF LITHOSTROTIONELLA (COELENTERATA, RUGOSA) 

FROM THE CARBONIFEROlTS i\ND PERMIAN 

By WILLIAM J. SANDO 

ABSTRACT 

Species of predominantly massive colonial rugose corals from the 

Carboniferous and Permian that were referred previously to 

Lithost1·otionella Y abe and Hayasaka are reassigned to the following 

genera: Acrocyathus d'Orbigny (including probable junior synonym 

Lithost1·otionella Y abe and Hayasaka), Stelechophyllum Tolmachev 

(including junior synonym Eolithostrotionella Zhizhina), Petalaxis 

Milne-Edwards and Haime (including junior synonyms Hillia de Groot 

and Eastonoides Wilson and Langenheim), Aulostylus Sando, 

Kleopatrina McCutcheon and Wilson, Lonsdaleia McCoy (subgenus 

Actinocyathus d'Orbigny), and Thysanophyllum Nicholson and Thomson. One of the species referred to Cystolonsdaleia Fomichev is 

reassigned to Petalaxis. 

Stelechophyllum and A ulostylus are referred to the Family 

Lithostrotionidae d'Orbigny. A new family, the Acrocyathidae, is 

created for the genus Acrocyathus. Petalaxis is placed in the Family 

Petalaxidae Fomichev, Thysanophyllum and Lonsdaleia are referred 

to the Family Lonsdaleiidae Chapman, and Kleopatrina is assigned to 

the Family Durhaminidae Minato and Kato. 

The principal lithostrotionelloid genera are Stelechophyllum, 

Aulostylus, Acrocyathus, and Petalaxis. Stelechophyllum ranges in 

age from Late Devonian(?) into the late Visean and is represented in 

the Carboniferous by 15 nominal species allocated to five species 

groups; the genus occurs in the U.S.S.R., U.S.A., Canada, and Mexico. 

A ulostylus is represented by two middle Tournaisian species in the 

U.S.A. and Canada and a possible species from the Visean of China. 

Acrocyathus is represented by 14 nominal species (one new) from the 

lower to upper Visean of the U.S.S.R., U.S.A., Canada, and China. 

Petalaxis is represented by 41 nominal species (six new) allocated to 

five species groups that range from the upper Visean into the Permian; 

the genus occurs in the U.S.S.R., U.S.A., Canada, Spain, Japan, and 

possibly China and Spitzbergen. 

Stelechophyllum may have been derived from Endophyllum in the 

Devonian. A ulostylus, Acrocyathus, and Petalaxis are regarded as offshoots from the Stelechophyllum stock in the Carboniferous. 

Lonsda/,eia, Thysanophyllum, and Kleopatrina do not seem to be 

closely related to Stelechophyllum and its derivatives. 

Hayasaka's type specimens of Lithostrotionella species from North 

America are reassigned to Acrocyathus, Stelechophyllum, Petalaxis, 

andAulostylus and are revised specifically. Three new species and two 

new subspecies are based on specimens studied by Hayasaka. 

INTRODUCTION 

Inadequate description of taxa is one of the principal 

deterrents to progress in paleontology. The study of 

fossil corals has been particularly plagued by this problem because many taxa were proposeq before thinsection techniques· became widely used and thus were 

founded largely or entirely on external skeletal features. 

Until these extant taxa are redescribed and adequately 

understood, new taxa cannot be proposed without risk 

of duplication. 

In North America, massive colonial rugose corals 

originally referred to Lithostrotion Fleming and later to 

Lithostrotionella Y abe and Hayasaka are noteworthy 

examples of inadequate description. After the genus 

Lithostrotionella was founded by Y abe and Hayasaka 

(1915, 1920) on material from the Carboniferous of 

China, Hayasaka studied North American corals in the 

large U.S. Geological Survey collection of fossils (now 

housed in the U.S. National Museum of Natural History 

in Washington, D.C.). Although Hayasaka's visit to the 

United States was in 1927, his monograph on North 

American Lithostrotionella was not published until 

1936. Hayasaka prepared thin sections from all the 

holotypes of his North American species, but the thin 

sections were small and only a few paratypes were sectioned. Consequently, Hayasaka's species concepts were 

based largely on the holotypes, and many paratypes 

were incorrectly identified. Moreover, the geologic ages 

of these specimens were poorly known at the time the 

study was made. Subsequently, the name Lithostrotionella came into wide use for Carboniferous and Permian colonial Rugosa throughout. the world. 

I beeame acquainted with the problem of Hayasaka's 

type specimens some years ago during the course of investigations of North American Mississippian coral 

faunas. Easton's (1973) study of Acrocyathus and his 

conclusion that Lithostrotiorwlla is a junior synonym 

lent impetus to a restudy of the Hayasaka material. All 

Hayasaka's type specimens were sectioned and restudied in detail except for four para types that could not 

be found; most of Hayasaka's specimens are illustrated 

in this report. Other described and undescribed 

specimens of Carboniferous and Permian age in the collections of the U.S. National Museum of Natural History 

were also investigated in order to broaden the base for 

taxonomic conclusions. All locality data were rechecked 
2 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

and were revised as necessary on the basis of the latest 

available information (see locality register on p. 40). 

As work proceeded on the Hayasaka material, it 

became apparent that the classification of this material 

had broader implications concerning the validity of 

Lithostrotionella as a taxonomic concept and concerning the scope and phylogeny of other genera. Accordingly, the present study was expanded to include an evaluation of all corals that have been described under the 

name of Lithostrotionella; this evaluation was based on 

a study of the systematic literature on these corals as of 

the end of 1978. I found that most of the specimens 

originally referred to Lithostrotionella by Hayasaka 

(1936) should be reassigned to Stelechophyllum, 

A ulostylus, Acrocyathus, and Petalaxis (table 1). Corals 

assigned by other authors to Lithostrotionella also 

belong in these genera, but a few are assigned to 

Thysanophyllum, Kleopatrina, and Lonsdaleia. 

ACKNOWLEDGMENTS 

I am indebted to P. M. Kier and R. E. Grant of the 

U.S. National Museum of Natural History for making 

Hayasaka's types and other specimens available to me 

for study. I am also grateful to P. Semenoff-TianChansky of the Museum National d'Histoire Naturelle in 

Paris and to N. V. Kabakovich and T. A. Sayutina of the 

Paleontologic Institute of the U.S.S.R. Academy of 

Sciences in Moscow for courtesies extended to me during visits to study material in their care. Many of the 

ideas expressed in this paper were discussed with colleagues at the Eighth International Congress of Carboniferous Stratigraphy and Geology in Moscow and the 

Third International Symposium on Fossil Corals in 

Paris, 1975; N. P. Vasilyuk, E. W. Bamber, M. Minato, 

M. Kato, P. K. Sutherland, and J. Fedorowski provided 

useful suggestions and criticisms. M. Minato and M. 

Kato provided me with photographs of the holotype of 

the type species of Lithostrotionella and translations of 

some of Hayasaka's short papers in Japanese. E. C. 

Wilson provided information on some Pacific Coast 

localities and specimens. B. L. Mamet identified 

foraminifers in some collections. I am also indebted to 

the following for searching collections in their care for 

type material: J. A. Stitt, University of Missouri; L. S. 

Kent, Illinois Geological Survey; H. R. Cramer, Emory 

University; A. Horowitz, Indiana University; D. B. 

Macurda, University of Michigan; and R. R. Shrock, 

Massachusetts Institute of Technology. K. R. Moore and 

H. I. Saunders searched the collections of the U.S. 

Geological Survey and U.S. National Museum of Natural 

History for specimens used in this study. K. R. Moore 

also made thin sections and photographs of the 

specimens. 

THE STATUS OF LITI/OSTROTIONELLA 

Yabe and Hayasaka (1915, p. 94-96, 133-184) originally proposed Lithostrotionella as a subgenus of 

Lithostrotion for a cerioid coral having a lamellar columella and differing from true Lithostrotion by having a 

lonsdaleoid dissepimentarium. They described, but did 

not illustrate, the type species (by monotypy) L·ithostrotion (Lithostrotionella) unicum from the 

Carboniferous(?) of Yun-nan Province, China. They suggested that Petalaxis Milne-Edwards and Hairne might 

be a senior synonym but concluded that Petnlaxis is 

identical with true Lithostrotion. In 1920, the same 

authors (Yabe and Hayasaka, p. 11, pl. 9, figs. 12a, b) illustrated two thin sections of the type species, but both 

thin sections were cut obliquely with respect to the axes 

of the corallites, making it difficult to interpret internal 

structures. 

Chi (1931, p. 28-29) raised the name to generic rank 

for a species from the Middle Carboniferous W einingian 

System of Kuechow Province, China, and Yu (1933, p. 

101-1 02) described two species of Lithostrotionella from 

the Lower Carboniferous of Kuechow Province. 

Dobrolyubova (1935b, p. 10-20) described species from 

the Middle Carboniferous of the Moscow Basin and, 

later (1936a, p. 9, 28, 66), from the Upper Carboniferous 

of the Urals. This early work set the stage for 

Hayasaka's (1936) monograph of Lithostrotionella in 

North America. 

Meanwhile, Stelechophyllum Tolmachev was :ignored, 

even by Soviet students, until Hill (1956, p. F286) 

treated it as a distinct genus and· Dobrolyubova and 

Kabakovich (1962) used the name for corals from the 

Kuznetsk Basin. The interpretation of Petalaxis MilneEdwards and Haime at that time was based on an erroneous designation of Petalaxis portlocki MilneEdwards and Haime as type species by Hill (1940, p. 

165, 166), who had overlooked Roemer's (1883:, p. 387, 

388) designation of P. maccoyanus Milne-Edwards ~nd 

· Haime as type species. P. portlocki is a Lithostrotion, 

whereas P. maccoyanus has a lonsdaleoid dissepimentarium and cannot be referred to Lithostrotion. 

Recent work has not unequivocally resolved the morphology of the type species or the most practical limits 

of the generic concept. Easton (1973) revived 

Acrocyathus d'Orbigny, redescribed the type specimen 

of its type species, and concluded that Lithostrotionella 

is a junior synonym. By this action, Easton simply 

transferred the highly variable group of corals previously referred to Lithostrotionella to Acrocyathus. 

Minato and Kato (1974, p. 67-75) restudied the 

holotype of Lithostrotionella unicum. All that remains 

of the original type material is one of the oblique thin 

sections originally illustrated by Yabe and Hayasaka 

(1920, pl. 19, fig. 12a). Minato and Kato concluded that 
THE STATUS OF LITHOSTROTIONELLA 3 

TABLE 1. -Hayasaka {1936} type specimens ofLithostrotionella 

Hayasnka Revised stratigraphic level, age, and 

species Kind of Revised USNM USGS and location (see Register of USGS Localities Plate and 

identification type specimen identification No. Loc. No. (p. 40) for detailed data) page 

americana ___ Holotype Acrocyathus jloriformis floriformis d'Orbigny 120240 2333-PC St. Louis Limestone, Late Mississippian Pl. 5; p. 17 

(Vis~an), Kentucky. 

Para type A crocyathus jloriformis jloriformis d'Orbigny? 120241 12UB-PC St. Louis Limestone, Late Mississippian Pl. 6; p. 17 (Vis~an), Missouri. ____ do ________ A crocyathus jloriformis jloriformis d'Orbigny? 162001 498-PC St. Louis Limestone, Late Mississippian Pl. 6; p. 17 (Vis~an), Illinois. 

do Indet. lithostrotionelloid coral 174371 3858-PC (green McCloud Limestone, Early Permian, ---------- label) California. 

____ do-------- Stelechophyllum banffense (Warren)? 174372 71308-PC Alapah Limestone(?), Late Mississippian Pl. 3; p. 13 (Vis~an), Alaska. ____ do ________ lndet. lithostrotionelloid coral 174373 3890-PC (green McCloud Limestone, Early Permian, ---------- label) California. ____ do ________ Stelechophyllum banffense (Warren)? 174374 970-PC Alapah Limestone(?), Late Mississippian Pl. 3; p. 13 (Vis~an), Alaska. ____ do ________ Stelechophyllum banffense (Warren)? 174375 3024-PC Little Flat Formation, Late Mississippian Pl. 4; p. 13 (Vis~an), Idaho. ____ do ________ Acrocyathus jloriformis jloriformis d'Orbigny 174376 499-PC St. Louis Limestone, Late Mississippian Pl. 5; p. 17 (Vis~an), Illinois. 

castelnaui ____ Holotype Acrocyathus jloriformis jloriformis d'Orbigny 120235 499-PC St. Louis Limestone, Late Mississippian Pl. 7; p. 17 (Vis~an), Illinois. 

Para type Acrocyathus jloriformis hemisphaericus 120236 3282-PC Hillsdale Member of Greenbrier Limestone, Pl. 14; p. 00 

(Hayasaka) Late Mississippian (Visean), West Virginia. 

____ do-------- Acrocyathus jloriformis jloriformis d'Orbigny 161989 2226-PC St. Louis Limestone, Late Mississippian Pl. 9; p. 17 (Vis~an), Illinois. ____ do ________ Acrocyathus floriformis jloriformis d'Orbigny 161990 346C-PC Residual chert from Tuscumbia Limestone, Pl. 8; p. 17 Late Mississippian (Vis~an), Alabama. ____ do ________ Acrocyathus jloriformis jloriformis d'Orbigny 161991 643-PC St. Louis Limestone(?), Late Mississippian Pl. 8; p. 17 (Vis~an), Missouri. 

____ do-------- Acrocyathus jloriformis hemisphaericus 161992 2020A-PC Greenbrier Limestone, Late Mississippian Pl. 14; p. 00 (Hayasaka) (Vis~an), Virginia. ____ do ________ Acrocyathus floriformis floriformis d'Orbigny 161993 3159-PC Newman Limestone, Late Mississippian Pl. 10; p. 17 (Visean), Virginia. ____ do ________ Acrocyathus floriformis jloriformis d'Orbigny 161994 2013B-PC Greenbrier Limestone, Late Mississippian Pl. 9; p. 17 (Vis~an), Virginia. ____ do ________ Acrocyathus jloriformis floriformis d'Orbigny? 174377 3946-PC (gTeen Unknown, probably St. Louis Limestone, Pl.7;p.17 

label) Mississippi Valley region. 

floriformis ___ Holotype ______ Stelechophyllum banffense (Warren) 120242 3760-PC Peratrovich Formation, Late Mississippian See Armstrong (Vis~an), Alaska. (1970a) 

girtyi _______ ____ do ________ Acrocyathus girtyi (Hayasaka) 120243 4801H-PC (green Little Flat Formation, Late Pl. 17, p. 21 label) Mississippian (Vis~an), Utah. 

Paratype ______ Petalaxis exiguus n. sp. 162002A, B 3856-PC (g:reen McCloud Limestone, Early Pemian, Pl. 19; p. 28 label) California. 

____ do-------- Stelechophyllum microstylum (White)? 162003 3864-PC Lodgepole Limestone, Early Mississippian Pl. 2; p. 11 (Tournaisian), Montana. 

----do Acrocyathus pilatus n. sp. 162004 499-PC St. Louis Limestone, Late Mississipian Pl. 17; p. 19 

(Visean), Illinois. 

----do Specimen lost 2111-PC Meridian Range, Montana. ---------- ----do ----do --------------------------------- 5077 A-PC (green Weber Canyon, Utah. ---------- label) 

____ do-------- ____ do--------------------------------- Granitville, Utah. ---------- hemisphaerica Holotype Acrocyathus jloriformis hemisphaericus 120237 1148-PC St. Louis Limestone, Late Mississippian Pl. 12; p. 19 

(Hayasaka) (Visean), Illinois. 

Paratype ______ Acrocyathus jloriformis jloriformis d'Orbigny 120238 3283-PC Hillsdale Member of Greenbrier Limestone, Pl. 11; p. 17 Late Mississippian (Vis~an), West Virginia. ____ do ________ Stelechophyllum sp. indet. 120239 5892-PC Madison Limestone, Early Mississippian Pl. 4; p. 15 (Tournaisian), Utah. 

____ do-------- Acrocyathus floriformis jloriformis d'Orbigny 161995 83-PC Newman Limestone, Late Mississippian Pl. 11; p. 17 (Vis~an), Kentucky. 

____ do-------- Stelechophyllum microstylum (White) 161996 1439-PC Lodgepole Limestone, Early Mississippian Pl. 2; p. 11 (Tournaisian), Idaho. 

____ do-------- Acrocyathus jloriformis floriformis d'Orbigny 161997 2020-PC Greenbrier Limestone, Late Mississipian Pl. 10; p. 17 (Visean), Virginia. 

----do Acrocyathus jloriformis hemisphaericus 161998 932A-PC St. Louis Limestone, Late Mississippian Pl. 12; p. 19 

(Hayasaka) (Visean), Missouri. 

----do Acrocyathus jloriformis hemisphaericus 161999 2222A-PC St. Louis Limestone, Late Mississippian Pl. 13; p. 19 

(Hayasaka) (Vis~an), Illinois. 

do Acrocyathus jloriformis hemisphaericus 162000 2222C-PC St. Louis Limestone, Late Mississippian Pl. 13; p. 19 

(Hayasaka)? (Visean), Illinois. multiradiata Holotype Stelechophyllum microstylum (White) 120244 490-PC Lodgepole Limestone, Early Mississipian Pl. 1; p. 11 (Tournaisian, Utah. 

Para type Stelechophyllum microstylum (White) 162005 104-PC Lodgepole Limestone, Early Mississippian Pl. 1; p. 11 (Tournaisian), Idaho. 

simplex -·----- Holotype Petalaxis simplex (Hayasaka) 120249 5893-PC Little Flat Formation, Late Pl. 18; p. 26 Mississippian (Visean), Utah. 

Para type Petalaxis wyomingensis n. sp. 120675 745Z-PC (brreen "Wells" Formation, Late Mississippian or Pl. 18; p. 26 label) Early Pennsylvanian (Namurian), 

Wyoming. tabulata _____ Holotype Petalaxis tabulatus (Hayasaka) 120246 1476-PC Aspen Range Formation, Late Mississippian Pl. 19; p. 26 (Vis~an), Idaho. 

tubifera _____ _ ___ do ________ A ulostylus tubiferus tubiferus (Hayasaka) 120247 5894-PC Mission Canyon Limestone, Early See Sando 

Para type 

Mississippian (Tournaisian), Montana (1976) A ulostylus tubiferus eotubiferus Sando 120248 3290-PC Lodgepole Limestone, Early Mississippian See Sando 

(Tournaisian), Montana. (1976) vesicularis Holotype Stelechophyllum banffense (Warren) 120245 3747C-PC Peratrovich Formation, Late Mississippian See Armstrong 

(Visean), Alaska. (1970a) Para type Specimen lost 970-PC(?) Porcupine-Arctic section, Alaska{?) 
4 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

the geologic age of the type specimen is probably Carboniferous. According to these writers, Lithostrotionella and Acrocyathus are distinct and separate 

genera, Stelechophyllum is a junior synonym of 

Lithostrotionella, Eolithostrotionella is a junior 

synonym of Acrocyathus, and Petalaxis is a junior 

synonym of Lithostrotion. 

The single surviving thin section of the holotype 

(Minato and Kato, 1974, pl. 15, fig. 1) does not permit an 

unequivocal evaluation of all the critical internal 

characters of the type species of Lithostrotionella. 

However, the corallite in the upper left-hand corner of 

the slide has what appear to be conical tabulae like those 

of Acrocyathus and shows no peripheral tabellae. The 

tabularium is certainly not that of Stelechophyllum or 

Petalaxis. Although none of the corallites show clearly 

defined septal lamellae, septal lamellae are not present 

in all corallites of some species of Acrocyathus. 

I conclude that the type species of Lithostrotionella is 

most likely a species of Acrocyathus; therefore, I regard 

Lithostrotionella as a probable junior synonym of 

Acrocyathus. This action, although debatable, is the 

most practical solution to the nomenclatural problem 

because both Acrocyathus and Stelechophyllum are 

founded on well-described and well-illustrated 

specimens whose ages are well documented. Although 

the holotype of the type species of Petalaxis is lost and a 

neotype remains to be selected, the species is well 

established on topotypes (Sutherland, 1977). 

CLASSIFICATION 

Species that have been referred previously to 

Lithostrotionella Y abe and Hayasaka are reassigned to 

the following genera in this report: Acrocyathus d'Orbigny (including probable junior synonym Lithostrotionella Yabe and Hayasaka), Stelechophyllum 

Tolmachev (including junior synonym Eolithostrotionella Zhizhina), Petalaxis Milne-Edwards and Haime 

(including junior synonyms Hillia de Groot and 

Eastonoides Wilson and Langenheim), A ulostylus 

Sando, Kleopatrina McCutcheon and Wilson, 

Lonsdaleia McCoy (subgenus Actinocyathus d'Orbigny), 

and Thysanophyllum Nicholson and Thomson. One of 

the species referred to Cystolonsdaleia Fomichev is 

reassigned to Petalaxis. 

Recent classification of the generic taxa recognized 

herein is in a state of flux, and there has been little 

agreement on their familial relations. Acrocyathus was 

referred to the Family Lonsdaleiidae Chapman by Hill 

(1956) (as a questionable senior synonym of Lithostrotionella), Easton (1973), and by Ivanovskiy (1975) (as a 

junior synonym of Lonsdaleia). Dobrolyubova and 

Kabakovich (1962) regarded Acrocyathus as a questionable junior synonym of Lithostrotion and referred it 

to the Family Lithostrotionidae d'Orbigny. 

Lithostrotionella was referred to the Family 

Lonsdaleiidae by Hill (1956), Easton (1973) (as a junior 

synonym of Acrocyathus), de Groot (1964), Armstrong 

(1970a, b; 1972a, b), and Ivanovskiy (1975). It was placed 

in the Family Lithostrotionidae by Easton and 

Gutschick (1953) (as a junior synonym of Lithoi~trotion) 

and by Dobrolyubova and Kabakovich (1962). Easton 

(1958) placed Lithostrotionella in the Family Li.thostrotionidae Grabau. The genus was placed in Family 

Petalaxidae Fomichev by Pyzh'anov (1964) (as a junior 

synonym of Petalaxis) and by Onoprienko (1970, 1976). 

Stelechophyllum was referred to the Family Lithostrotionidae by Hill (1956) and to the Family Lonsdaleiidae 

by Ivanovskiy (1975) (as a junior synonym of L'l~thostrotionella). Dobrolyubova and Kabakovich (196~~, 1966) 

placed this genus in the Family Endophyllidae Torley 

because of its inferred derivation from Endophyllum, a 

Devonian genus that they recognized as ranging into the 

Lower Carboniferous. 

Eolithostrotionella was referred to the Family 

Lithostrotionidae by Zhizhina (1956) and in Bul'vanker 

and others (1960), by Vasilyuk (1960), and by 

Dobrolyubova and Kabakovich (1962) (as a junior 

synonym of Lithostrotionella). It was placed in the 

Family Lonsdaleiidae by Easton (1973) (as a questionable junior synonym of Acrocyathus), by Degtyarev 

(1973), and by Ivanovskiy (1975) (as a junior synonym of 

Lithostrotionella). Dobrolyubova and Kabakoviclh (1966) 

regarded it as a junior synonym of Stelechophyllum and 

placed it in the Family Endophyllidae. Pyzh'anov (1964), 

Onoprienko (1970), and Kozyreva (1974) placed this 

genus in the Family Petalaxidae. Fedorowski and 

Gorianov (1973) placed it in the Family Lonsdaleiidae. 

Petalaxis was referred by Hill (1956) (as a. junior 

synonym of Lithostrotion) and by Ivanovskiy (19'75) (as a 

junior ·

4 synonym of Lithostrotion) to the Family 

Lithostrotionidae, on the basis of an erroneous typespecies designation. Fomichev (1953), Dobrolyubova and 

Kabakovich (1962), Pyzh'anov (1964), Onoprienko 

(1970), and Kozyreva (1974) placed this genus in the 

Family Petalaxidae. Fedorowski and Gorianov (1973) 

placed it in the Family Lonsdaleiidae. 

Hillia was referred to the Family Lonsdaleiida.e by de 

Groot (1964) (as a subgenus of Lithostrotionella) and by 

Easton (1973) (questionably). Ivanovskiy (19'75) regarded it as a questionable junior synonym of Lithostrotion and placed it in the Family Lithostrotionidae. 

Eastonoides was referred to the Family Lonsdaleiidae 

by Wilson and Langenheim (1962). 

A ulostylus was placed in the Family Lithostrotionidae 

by Sando (1976). 
CLASSIFICATION 5 

Kleopatrina was referred to the Family Durhaminidae 

Minato and Kato by Minato and Kato (1965) and by 

Ivanovskiy (1975). 

Lonsdaleia was placed in the Family Lonsdaleiidae by 

Hill (1956), Vasilyuk (1960), Dobrolyubova and 

Kabakovich (1962), de Groot (1964), and I vanovskiy 

(1975). 

Thysanophyllum was referred to the Family 

Lonsdaleiidae by Hill (1956), Armstrong (1970a, b), 

Degtyarev (1973), and Ivanovskiy (1975). Dobrolyubova 

and Kabakovich (1962) placed it in the Family Lithostrotionidae. Pyzh'anov (1964) and Onoprienko (1970, 1976) 

referred it to the Family Petalaxidae. 

In this report, the following classification is used: 

Order Rugosa Milne-Edwards and Haime, 1850 

Suborder Columnariina Rominger, 1876 

Family Lithostrotionidae d'Orbigny, 1851 

Genus Stelechophyllum Tolmachev, 1933 (junior synonym 

Eolithostrotionella Zhizhina in Fomichev, 1955) 

S. ascendens species group 

S. ascendens (Tolmachev, 1924) 

S. ascendens simplex Dobrolyubova in 

Dobrolyubova and Kabakovich, 1966 

S. ascendens ascendens (Tolmachev, 1924) 

S. megalum (Tolmachev, 1924) 

S. grande (Tolmachev, 1924) 

S. venukoffi (Tolmachev, 1924) 

S. venukojji venukoffi (Tolmachev, 1924) 

S. venukoffi altaicum (Tolmachev, 1924) 

S. microstylum species group 

S. microstylum (White, 1880a) 

S. circinatum (Easton and Gutschick, 1953) 

S. longiseptatum (Lisitsyn, 1925) 

S. banffense (Warren, 1927) 

S. micrum species group 

S. micrum (Kelly, 1942) 

S. lochmanae (Armstrong, 1962) 

S. ergunjaicum (Onoprienko, 1976) 

S.? mclareni species group 

S.? mclareni (Sutherland, 1958) 

S.? birdi (Armstrong, 1970a) 

S.? niakensis (Armstrong, 1972a) 

S.? macouni species group 

S.? macouni (Lambe, 1899) 

S. sp. indet. 

S.? sp. indet. 

Genus Aulostylus Sando, 1976 

A. tubiferus (Hayasaka, 1936) 

A. tubiferus tubiferus (Hayasaka, 1936) 

A. tubiferus eotubiferus Sando, 1976 

A.? sp. 

Family Acrocyathidae new family 

Genus Acrocyathus d'Orbigny, 1849a (?junior synonym 

Lithostrotionella Yabe and Hayasaka, 1915) 

A. floriformis d'Orbigny, 1849a 

A. floriformis floriformis d'Orbigny, 1849a 

A. floriformis hemisphaericus (Hayasaka, 1936) 

A. pilatus n. sp. 

A. proliferus (Hall in Hall and Whitney, 1858) 

A. girtyi (Hayasaka, 1936) 

A. pennsylvanicus (Shimer, 1926) 

Order Rugosa- Continued 

Suborder Columnariina-Continued 

Family Acrocyathidae- Continued 

Genus Acrocyathus-Continued 

A. utkae (Degtyarev, 1973) 

A. rotai (Zhizhina in Bul'vanker and others, 1960) 

A. cystosus (Zhizhina in Bul'vanker and others, 

1960) 

A. lissitzini (Zhizhina in Bul'vanker and others, 1960) 

A. hsujiulingi (Yoh, 1961) 

A.? unicus (Yabe and Hayasaka, 1915) 

A.? shimeri (Crickmay, 1955) 

A.? grechovkae (Degtyarev, 1973) 

A.? zhizhinae (Vasilyuk, 1960) 

A. spp. indet. 

Family Petalaxidae Fomichev, 1953 

Genus Petalaxis Milne-Edwards and Haime, 1852 (junior 

synonyms Hillia de Groot, 1963, and Eastonoides 

Wilson and Langenheim, 1962) 

P. simplex species group 

P. simplex (Hayasaka, 1936) 

P. wyomingensis n. sp. 

P. tabulatus (Hayasaka, 1936) 

P. bailliei (Nelson, 1960) 

P. flexuosus species group 

P. flexuosus (Trautschold, 1879) 

P. donbassicus (Fomichev, 1939) 

P. mokomokensis (Easton, 1960) 

P. exiguus n. sp. 

P. brokawi (Wilson and Langenheim, 1962) 

P. monocyclicus (de Groot, 1963) 

P. sexangulus (de Groot, 1963) 

P. taishakuensis (Yokoyama, 1957) 

P. immanis Kozyreva, 1974 

P. belinskiensis Fomichev, 1953 

P. major (de Groot, 1963) 

P. fomichevi n. sp. 

P. grootae n. sp. 

P. wagneri species group 

P. wagneri (de Groot, 1963) 

P. perapertuensis (de Groot, 1963) 

P. radians (de Groot, 1963) 

P. santaemariae (de Groot, 1963) 

P. cantabricus (de Groot, 1963) 

P. orboensis (de Groot, 1963) 

P. occidentalis (Merriam, 1942) 

P. vesiculosus species group 

P. vesiculosus (Dobrolyubova, 

1935a) 

P. lisitschanskensis (Fomichev, 

1953) 

P. exilis Kozyreva, 1974 

P. confertus Kozyreva, 197 4 

P. persubtilis Kozyreva, 1974 

P. korkhovae Kozyreva, 1974 

P. mirus Kozyreva, 1974 

P. evidens Kozyreva, 197 4 

P. maccoyanus species group 

P. maccoyanus Milne-Edwards and 

Haime, 1851 

P. stylaxis (Trautschold, 1879) 

P. mohikanus (Fomichev, 1939) 

P. celadensis (de Groot, 1963) 
6 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Order Rugosa-Continued 

Suborder Columnariina-Continued 

Family Petalaxidae-Continued 

Genus Petalaxis-Continued 

P. maccoyanus species group-Continued 

P. elyensis (Wilson and Langenheim, 1962) 

P. dobrolyubovae n. sp. 

P. donetsensis n. sp. 

P. ivanovi (Dobrolyubova, 1935a) 

P.? spp., indet. 

Family Lonsdaleiidae Chapman, 1893 

Genus Thysanophyllum Nicholson and 

Thomson, 1876 

T. astraeiforme (Warren, 1927) 

Genus Lonsdaleia McCoy, 1849 

Subgenus Actinocyathus d'Orbigny, 

1849a 

L. (A.) berthiaumi (Merriam, 1942) 

L. (A.) peratrovichensis (Armstrong, 1970a) 

Family Durhaminidae Minato and Kato, 1965 

Genus Kleopatrina McCutcheon and Wilson, 

1963 

Subgenus Kleopatrina McCutcheon and 

Wilson, 1963 

K. (K.)? dilatata (Easton, 1960) 

K. (K.)? uralica (Dobrolyubova, 

1936a) 

K. (K.)? wahooensis (Armstrong, 

1972b) 

Undetermined lithostrotionelloid corals 

The foregoing classification of lithostrotionelloid corals follows Hill (1956) at the ordinal and subordinallevel. 

Placement of Stelechophyllum in the Family Lithostrotionidae also follows Hill (1956). Aulostylus is placed in 

the Lithostrotionidae because of its close relationship to 

Stelechophyllum. The recognition of Family Petalaxidae 

including only Petalaxis and Cystolonsdaleia conforms 

to Fomichev's (1953) original definition and the subsequent acceptance by Dobrolyubova and Kabakovich 

(1962). A new family, the Acrocyathidae, is proposed to· 

include only the genus Acrocyathus, which appears to 

have been derived from Stelechophyllum. Only a few 

species are allocated to Thysanophyllum, Lonsdaleia 

(Actinocyathus), and Kleopatrina (Kleopatrina), which 

are not closely related to the main stock of lithostrotionelloid corals. Thysanophyllum and Lonsdaleia are 

placed in the Family Lonsdaleiidae following Hill (1956), 

and Kleopatrina is placed in the Family Durhaminidae 

following Minato and Kato (1965). 

PHYLOGENY 

Hill's (1938, p. 35) statement that "little is known of 

the phylogeny of the Rugosa" is still valid today. In the 

absence of general agreement on the composition of 

families, it is very difficult to sketch out even the main 

phylogenetic lines. Hill (1938, p. 35-36) listed five 

methods of approach to the phylogeny of genera and 

species in Rugosa: (1) morphologic comparison without 

strict attention to detailed stratigraphic chronology; (2) 

the adult "Formenreihe" method used by Vaughan in his 

studies of the Rugosa, in which adult charaeters are 

traced through a stratigraphic sequence of species; (3) 

the "Formenreihe" method used by Carruthers in his 

classic studies of Zaphrentis, in which Haeckel's law of 

recapitulation is used on a stratigraphic succession of 

species; ( 4) deduction of phylogeny of individua1 species 

from their ontogeny and checking this against stratigraphic evidence; and (5) deduction of phylogeny of individual species from ontogeny without detailed 

stratigraphic evidence. Unfortunately, few studies have 

been made of colonial Rugosa in the Carboniferous by 

any of these methods. Nevertheless, it is useful · to 

speculate on how the lithostrotionelloid corals are 

related phylogenetically. 

The lithostrotionelloid corals may be divided into two 

groups of genera, those that seem to be closely related 

phylogenetically, and those that seem to be independent 

unrelated forms. In the first group are species allocated 

to Stelechophyllum, Aulostylus, Acrocyath,us, and 

Petalaxis (fig. 1), which are characterized by a 

lonsdaleoid dissepimentarium, tabulae that range from 

tent-shaped to conical to horizontal, and an axial structure that ranges from a simple axial plate ordinarily connected to the counter septum to a complex spider-web 

structure made up of axial plate, vertical axial tabellae, 

and septal lamellae. 

The origin of the first group of lithostrotionelloid corals is uncertain. The earliest member of this group is 

Stelechophyllum, which has a lonsdaleoid dissepimentarium, a simple columella, and tent-shaped tabulae. According to Soshkina, Dobrolyubova, and Kabakovich 

(1962, p. 342), Stelechophyllum was derived from Endophyllum Milne-Edwards and Haime, a Middle Devonian coral that they considered to range into the Tournaisian of N ovaya Zemlya. The occurrence of Endophyllum 

in the Carboniferous apparently was based on Gorskiy's 

(1935, p. 49-56; 1938, p. 21-24) description of species 

from the Etroeungtian beds of Novaya Zemlya, which 

Gorskiy regarded as lower Tournaisian (most Carboniferous and Devonian stratigraphers now regard the 

E troeungtian as latest Devonian). True Middle Devonian Endophyllum has a lonsdaleoid dissepimentarium, 

major septa that approach the axis of the corallites but 

do not meet or form a columella, and flat tabulae that 

have down-turned margins (see Jones, 1929). Gorskiy's 

(1935, 1938) colonial species are more like 

Stelechophyllum than Endophyllum in the nature of 

their tabulae but do not have a columella. Some species 

of Stelechophyllum have a poorly developed columella 

(for example, S. ascendens, S. megalum, and S. grande). 

It seems more reasonable to refer Gorskiy's species 
PHYLOGENY 7 

AULOSTYLUS 

PETALAXIS 

ACROCYATHUS STELECHOPHYLLUM 

FIGURE 1.-Morphology of the type species of the principal lithostrotionelloid genera. Acrocyathus: longitudinal and transverse thin 

sections ( x 4) of A. florijormis floriformis d'Orbigny (USNM 120235). Stelechophyllum: longitudinal and transverse thin section ( x 4) of 

S. venukoffi altaicum (Tolmachev) (from Dobrolyubova and Kabakovich, 1966). Petalaxis: longitudinal and transverse thin sections (x 4) 

ofP. maccoyanus Milne-Edwards and Haime (from Fedorowski and Gorianov, 1973). Aulostylus: longitudinal and transverse thin sections 

(x 6) of A. tubiferus tubiferus (Hayasaka) (from Sando, 1976). 
8 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

questionably to Stelechophyllum and to regard them as 

possible derivatives of the Middle Devonian Endophyllum. 

If the Famennian forms in N ovaya Zemlya are referred questionably to Stelechophyllum, then there is a gap 

until the middle Tournaisian, when true Stelechophyllum 

appears and ranges into the upper Visean (fig. 2). In the. 

middle and upper Tournaisian, the axial structure was 

modified to an aulos in Aulostylus, which is a closely 

related offshoot of Stelechophyllum. Increased complexity of the axial structure and the formation of conical 

tabulae took place in Acrocyathus, which ranges through 

most of the Visean. Acrocyathus seems to be an offshoot 

of Stelechophyllum rather than a member of the 

Lonsdaleiidae, although it may have been the ancestor 

of Lonsdaleia (Actinocyathus). The earliest species of 

Petalaxis, which occur in the upper Visean, are similar 

to Stelechophyllum in the morphology of their septa and 

columella, but they have horizontal tabulae. Petalaxis . 

continues upward to the top of the Middle Carboniferous 

(Moscovian), but there is a gap in its record of occurrence in the Upper Carboniferous before it is seen again· 

in the Permian. The Permian species are very similar to 

Middle Carboniferous species. The significance of the 

gap is not now understood; the Permian forms may be 

merely homeomorphs in an unrelated phylogenetic 

stock. 

The second group of lithostrotionelloid corals includes 

two phylogenetic stocks that do not seem to be related 

to Stelechophyllum. Some species previously assigned to 

Lithostrotionella are herein placed in Lonsdaleia and 

Thysanophyllum, which are members of the Family 

Lonsdaleiidae. The origin of Lonsdaleia, which first appears at the top of the Tournaisian (Hill, 1940, p. 151), 

has been variously linked to Clisiophyllum (Vaughan, 

1905, p. 184), Carcinophyllum (Vaughan, 1906, p. 148), 

and Thysanophyllum (Carruthers, in Garwood, 1912, p. 

563; Smith, 1916, p. 235). However, E. W. Bamber 

(written commun., 1980) has pointed out that 

Acrocyathus pennsylvanicus and A. shimeri have axial 

structures and tabulation similar to those of the North 

American species of Lonsdaleia (Actinocyathus), that is, 

L. (A.) stelcki and L. (A.) peratrovichensis. Although 

these similarities suggest derivation of Lonsdaleia (Actinocyathus) from Acrocyathus in the Visean of North 

America, they are inconsistent with the earliest occurrence of Lonsdaleia in the upper Tournaisian of Great 

Britain. Thysanophyllum is characterized by a discontinuous axial structure and may be polyphyletic (Hill, 

··1940, p. 161). For the present, the origins of Lonsdaleia 

and Thysanophyllum and their possible relationships to 

the Stelechophyllum stock remain obscure. 

· Other -Carboniferous corals previously assigned to 

Lithostrotionella are placed herein in Kleopatrina, a 

predominantly Permian genus. According to Minato and 

Kato (1965, test-fig. 4), Kleopatrina was derived from 

Durhamina, and all the members of the Durhaminidae 

were ultimately derived from the Lithostrotionidae in 

the Lower Carboniferous. Hence, Kleopatrina does not 

seem to be closely related to the main stock of lithostrotionelloid corals. 

SYSTEMATIC PALEONTOLOGY 

Morphologic terminology is generally that of Hill 

(1956). The term "tent-shaped" refers to tabulae that 

slope peripherally from their intersection with the columella at a slight angle to a distinct rounded shoulder, 

where they are vertical (see Stelechophyllum in fig. 1). 

These tabulae ordinarily terminate peripherally at a 

zone of horizontal tabellae without reaching the inner 

margin of the dissepimentarium and rest one upon the 

other. The term "conical" refers to tabulae that slope 

peripherally from their intersection with the columella 

at a moderate to steep angle without distinct shoulders 

and reach the inner margin of the dissepimentarium (see 

LOWER CARBONIFEROUS /y-"'/ MIDDLE CARBONIFEROUS UPPER CARBONIFEROUS 

DEVONIAN PEFIMIAN Serpu- Tournaisian Visean khovian Bashkirian Moscovian Kasimovian Gzhelian 

ACROCYATHUS (ACROCYATHIDAE) 

? /sTELECHOPH'(LLUM (LITHOSTROTIONIDAE) 

~-- \ \ PETALAX/S. (PETALAXIDAE) 

7 

----..;_ _____ ~· 

AULOSTYLUS 

(LITH,STROTIONIDAE) 

No record ofl these corals 

FIGURE 2.- Postulated phylogeny of the principal genera of lithostrotionelloid corals. 
SYSTEMATIC PALEONTOLOGY 9 

Acrocyathus in fig. 1). Convex or bell-shaped tabulae are 

like tent-shaped tabulae but are horizontal at the top 

where they intersect the columella. All types of tabulae 

are commonly seen in the same specimen, but ordinarily 

one type dominates. Sections that are not in the plane of 

the axis show tabulae that appear to be progressively 

flatter as one approaches the periphery of the 

tabularium. 

Because skeletal microstructure has not been described in most of the species diagnosed herein, no attempt is made to include it in the diagnoses of genera. 

Septal microstructure is described according to the 

classification of Kato (1963) only in descriptions of type 

material of new species and subspecies. 

The repositories for material studied by the author are 

the U.S. National Museum of Natural History, Washington, D.C. (USNM), and the Museum National d'Histoire 

Naturelle, Paris, France (MNHN, FL). Other material 

referred to is in the Rijksmuseum van Geologie en 

Mineralogie at Leiden, The Netherlands (RGM). 

Occurrence data are limited to information taken from 

descriptive studies given in the synonymies of the taxa 

concerned, unless otherwise indicated. No attempt was 

made to search out all occurrences in faunal lists or 

other similar citations of the taxa. 

Order RUGOSA Milne-Edwards and Haime, 1850 

Suborder COLUMNARIINA Rominger, 1876 

Family UTHOSTROTIONIDAE d'Orbigny, 1851 

Genus STELECHOPHYLLUM Tolmachev, 1933 

Stylophyllum Tolmachev, 1924, p. 316; 1931, p. 606; Fomichev, 1931, 

p. 43, 72 (not Reuss, 1854). 

Stelechophyllum Tolmachev, 1933, p. 287 (replacement name for 

Stylophyllum Tolmachev); Soshkina, Dobrolyubova, and 

Kabakovich, 1962, p. 342; Dobrolyubova and Kabakovich, 1962, p. 

122; 1966, p. 130; Minato and Kato, 1974, p. 69. 

Eolitho.strotionella Zhizhina in Fomichev, 1955, p. 303, 304; 

Zhizhina, 1956, p. 40. 

Lonsdaleia McCoy. Lisitsyn, 1925, p. 68. 

Lithostrotion Fleming. White, 1880a (1883), p. 159 [part]; Keyes, 

1894, p. 124; Lambe, 1899, p. 220; 1901, p. 176; Tolmachev, 1924, 

p. 314 [part]; Warren, 1927, p. 46 [part]; Easton, 1944, p. 53; 

McLaren and Sutherland, 1949, p. 631; Bassler 1950, p. 213; 

Crickmay, 1955 (1961), p. 12 [part]; Nelson, 1960, p. 122 [part]. 

Lithostrotion [Lithostrotionella]. Bassler, 1950, p. 221; Easton and 

Gutschick, 1953, p. 19. 

Lithost'rotionella Yabe and Hayasaka. Hayasaka, 1936, p. 61, 62, 64, 

65, 67, 68 [part]; Kelly, 1942, p. 354, 356, 357 [part]; Easton, 1958, 

p. 31; Nelson, 1960, p. 112, 113, 119, [part]; 1962, p. 170; Bamber, 

1961, p. 107 [part]; 1966, p. 9, 14, 17 [part]; Armstrong, 1962, p. 

38, 39 [part]; 1970a, p. 29, 32, 35 [part]; 1970b, p. 16, 19, 20, 21 25, 

26; Easton, 1963, p. 297; Sando, 1969, p. 309; Onoprienko, 1976, p. 

29 [part]. 

Lithostrotion [Lithostrotionella] [Thysanophyllum] Sutherland, 1958, 

p. 95. 

Type species. -Stylophyllum venukofi Tolmachev, 

1924, p. 318, pl. 19, figs. 9, 10 and 1931, p. 607, pl. 23, 

fig. 2 (by designation of Tolmachev, 1933, p. 287). Lower 

Carboniferous (Tournaisian), Kuznetsk Basin, U.S.S.R .. 

Diagnos·is.- Cerioid corals with tabular to 

hemispherical growth form. Septa of two orders. Major 

septa thin, ordinarily extending across tabularium to 

columella but discontinuous or absent in dissepimentarium. Minor septa absent to variably developed. Columella ordinarily a simple, smooth or serrated, strong, 

continuous axial rod or plate that may have been derived 

from one or more septa, but thysanophylloid variants 

are known. Tabularium ordinarily consisting of an axial 

series of bell-shaped to tent-shaped complete tabulae 

resting one upon the other and a weak peripheral series 

of horizontal or inclined, concave-upward tabellae ("split 

tabulae" of Soviet authors). Dissepimentarium 

lonsdaleoid, commonly showing traces or crests of 

discontinuous septa in transverse section. Increase axial, peripheral, and intermural. 

Discussion. -Tolmachev's (1924, 1931) original 

diagnosis of this genus described a coral similar to 

cerioid Lonsdaleia but without an axial plate and having 

a false columella formed by arched tabulae. Some of 

Tolmachev's figures of transverse sections suggest that 

an axial rod is present, but the point is ambiguous. Hill 

(1940, p. 164; -1956, p. 286) and Cotton (1973, p. 195) 

followed the original diagnosis. However, Fomichev 

(1931, p. 43, 72) restudied Tolmachev's types and concluded that Tolmachev's longitudinal sections are cuts 

that do not intersect the axes of the corallites. Fomichev 

found a columella like that of Lithostrotion in 

Tolmachev's transverse sections. These observations 

were confirmed by Fomichev's study of topotypes. 

Subsequent study by Dobrolyubova and Kabakovich 

(1962, 1966) on specimens from the Kuznetsk Basin confirmed Fomichev's findings and also showed that the 

Kuznetsk Basin species form a genetic series from 

weakly to strongly columellate forms. 

Most authors (Hill, 1956, p. F286; Soshkina, 

Dobrolyubova, and Kabakovich, 1962, p. 342; 

Dobrolyubova and Kabakovich, 1962, p. 122-124; 1966, 

p. 130-157; Cotton, 1973, p. 195) regarded 

Stelechophyllum as a distinct genus, but Wang (1950, p. 

212), Ivanovskiy (1967, p. 34), and Minato and Kato 

(1974, p. 69-71) regarded it as a junior synonym of 

Lithostrotionella Y abe and Hayasaka. In my opinion, 

Stelechophyllum is a distinct genus and Lithostrotionella 

is a probable junior synonym of Acrocyathus d'Orbigny. 

Stelechophyllum is differentiated from Acrocyathus 

and Petala-Xis by the nature of its axial structure and 

tabula.rium (fig. 1). In Stelechophyllum, the axial structure consists of a simple columella in the form of a rod or 

plate unmodified by septal lamellae or axial tabellae. 

The major septa may or may not join the columella, and 

the columella may be slightly serrated where the septa 

fall short of it. The tabularium in Stelechophyllum con-
10 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

sists of an axial series of convex to tent-shaped tabulae 

with well-defined shoulders that rest one upon the other 

and a weak peripheral zone of more or less horizontal, 

ordinarily concave-upward tabellae. Acrocyathus has a 

complex axial structure composed of an axial plate, axial 

lamellae, and steeply inclined tabulae or tabellae. The 

major septa seldom extend to the axial complex. The 

tabulae are essentially conical and ordinarily extend 

without shoulders to the periphery of the tabularium; 

peripheral tabellae are rare. Petalaxis has an essentially 

simple columella like that of Stelechophyllum, although 

some species have impersistently developed axial 

tabellae. The tabulae in Petalaxis are essentially 

horizontal. 

Eolithostrotionella Zhizhina was proposed for corals 

that differ from Lithostrotion by having lonsdaleoid 

dissepiments and from Lithostrotionella by having 

arched, rather than horizontal, tabulae. Although the 

morphology of the type species of Lithostrotionella is in 

doubt, it seems to have conical tabulae like those of 

Acrocyathus; at any rate, the tabulae are not horizontal. 

The type species of Eolithostrotionella has an axial 

structure identical with the type species of 

Stelechophyllum and is here placed in the synonymy of 

Stelechophyllum. 

Species of Stelechophyllum are distinguished on differences in mature corallite diameter, number of major 

septa at maturity, presence or absence and length of 

minor septa (weak or strong), development of the columella (absent, weak, or strong), major septal extensions into the tabularium (weak or strong), major septal 

extensions in the dissepimentarium (weak or strong), 

shape of the axial tabulae (flat, tent-shaped, convex), 

spacing of the tabulae, development of peripheral 

tabellae (absent, weak, or strong), number of rows of 

dissepiments, size and shape of dissepiments (large, 

small, inflated, flattened), and ratio of tabularium width 

to corallite diameter. 

The species of Stelechophyllum fall into five major 

groups: 

1. S. ascendens group, including ascendens, megalum, 

grande, and venukoffi. This group is characterized 

by its large corallites, moderate number of major 

septa, and by having a variably developed columella. The group is exclusively Tournaisian. 

2. S. microstylum group, including microstylum, circinatum, longiseptatum, and banffense. This group 

is characterized by its large corallites, large 

number of major septa, and well developed columella. The group is Tournaisian and Visean. 

3. S. micrum group, including micrum, lochmanae, 

and ergunjaicum. This group is characterized by 

its small corallites and small number of major septa. The group is Tournaisian and Visean. 

4. S.? mclareni group, including mclareni, b'irdi, and 

niakense. This group is characterized by its strongly polymorphic corallites. The group is exclusively 

Visean. 

5. S.? macouni group, including only S.? macouni from 

the Visean. This group is characterized by its 

weakly lonsdaleoid dissepimentarium. 

Occurrence.- Upper Devonian (Famennian)(?) and 

Lower Carboniferous, middle Tournaisian to upper 

Visean. U.S.S.R., Canada, U.S.A., Mexico. 

Stelechophyllum ascendens species group 

Stelechophyllum ascendens (Tolmachev) 

Stylophyllum ascendens Tolmachev, 1924, p. 319, pl. 19, figs. 14, 15; 

1931, p. 608, pl. 23, fig. 4. 

Stelechophyllum ascendens (Tolmachev). Dobrolyubova and 

Kabakovich, 1962, p. 123, pl. C-5, fig. 11, pl. C-6, figs. la, b; 1966, 

p. 136. 

Diagnosis. - Stelechophyllum with corallite diameter 7 

to 12 mm and 13 to 17 major septa that seldom extend 

into the dissepimentarium. Major septa join the columella or end a short distance from it or from the axis 

when a columella is absent. Minor septa weakly 

developed or absent. Columella weakly developed or absent from some corallites. Dissepimentarium composed 

of a single row of large inflated dissepiments. Axial 

tabulae flat to strongly convex, spaced 0.5 to 1 mm 

apart. Peripheral tabellae absent or weakly developed. 

Ratio of tabularium width to corallite diameter 0.5 to 

0.6. Increase is peripheral. 

Discussion. -This species is distinguished by its weak 

development of minor septa and columella. 

Occurrence. -Lower Carboniferous, upper Tournaisian. Nizhnetersin Horizon, Kuznetsk Basin, U.S.S.R. 

Stelechophyllum ascendens simplex Dobrolyubova, 

in Dobrolyubova and Kabakovich 

Stelechophyllum ascendens simplex Dobrolyubova, in Dobrolyubova 

and Kabakovich, 1966, p. 137, pl. 23, figs. 1, 2, text-fig. 7. 

Diagnosis. -Stelechophyllum ascendens in which most 

of the corallites are without a columella and have 

tabulae that range from flat (without columella) to tentshaped (with columella). 

Description of holotype.- See Dobrolyubova and 

Kabakovich (1966, p. 137). 

Discussion. -This form is the most primitive in the 

Russian Stelechophyllum sequence (Dobrolyubova and 

Kabakovich; 1966, p. 134). 

Occurrence. -Lower Carboniferous, Tournaisian. 

Nizhnetersin Horizon, Kuznetsk Basin, U.S.S.R. 

Stelechophyllum ascendens ascendens (Tolmachev) 

Stelechophyllum ascendens ascendens (Tolmachev). Dobrolyubova and 

Kabakovich, 1966, p. 139, pl. 24, figs. 1, 2; pl. 25, figs. 1, 2; pl. 28, 

fig. 3; text-figs. 8; 9. 
SYSTEMATIC PALEONTOLOGY 11 

Diagnosis. -Stelechophyllum ascendens in which most 

of the corallites have a columella and axial tabulae and 

peripheral tabellae are present. 

Description of lectotype.- See Tolmachev (1924, p. 

319). 

Occu.rrence. -Lower Carboniferous, upper Tournaisian. Nizhnetersin Horizon, Kuznetsk Basin, U.S.S.R. 

Stelechophyllum meqalum (Tolmachev) 

Stylophyllum megalum Tolmachev, 1924, p. 319, pl. 19, figs. 11, 12; 

1931, p. 608, pl. 23, fig. 3. 

Stelechophyllum megalum (Tolmachev). Dobrolyubova and Kabakovich, 

1962, p. 123, pl. C-5, fig. 1; 1966, p. 143, pl. 26, figs. la-c, 2, textfig. 10. 

Diagnosis. -Stelechophyllum with corallite diameter 7 

to 14 mm and 13 to l9 major septa that most commonly 

extend from the columella across the tabularium and into the dissepimentarium. Minor septa generally well 

developed. Columella variable in form and thickness, 

commonly thick and lenticular, in some corallites replaced by ends of one or more ·major septa, absent from 

many corallites. Dissepimentarium consists of 1 to 3 

poorly defined rows of large and small inflated dissepiments. Axial tabulae tent-shaped, spaced about 5 in 

1 mm. Peripheral tabellae well developed. Ratio of 

tabula.rium width to corallite diameter about 0.4. Increase is peripheral. 

Description of lectotype.- See Tolmachev (1924, p. 

319). 

Occurrence. -Lower Carboniferous, upper Tournaisian. Nizhnetersin Horizon, Kuznetsk Basin, U.S.S.R. 

Stelechophyllum grande (Tolmachev) 

Lithostrotion grande Tolmachev, 1924, p. 315, pl. 19, fig. 13; 1931, pl. 

22, fig. 6. 

Stelechophyllum grande (Tolmachev) Dobrolyubova and Kabakovich, 

1966, p. 146, pl. 27, figs. 1, 2; text-figs. 11-14. 

Diagnosis. - Stelechophyllum with corallite diameter 9 

to 18 mm and 13 to 20 major septa that most commonly 

extend from the columella across the tabularium but 

seldom extend into the dissepimentarium. Minor septa 

absent to weakly developed. Columella variable in form 

and thickness, commonly thick and lenticular, in some 

corallites replaced by ends of one or more major septa, 

absent from some corallites. Dissepimentarium consists 

of 1 to 4 rows of small to large dissepiments. Tabulae 

spaced 0.5 to 1 mm apart, variable in form. Peripheral 

tabellae poorly· developed. Ratio of tabularium width to 

corallite diameter about 0.4 . Increase is peripheral. 

Description of lectotype.- See Tolmachev (1924, p. 

315). 

Occurrence. -Lower Carboniferous, upper Tournaisian. Nizhnetersin Horizon, Kuznetsk Basin, U.S.S.R. 

Stelechophyllum venukoffi (Tolmachev) 

See subspecies below for synonymy. . 

Diagnosis. -Stelechophyllum with corallite diameter 

5.5 to 15 mm and 12 to 20 major septa that ordinarily extend from the columella across the tabularium but 

seldom extend into the dissepimentarium. Minor septa 

absent to well developed. Columella absent to very 

thick. Dissepimentarium composed of 1 to 4 rows of 

small to large dissepiments. Axial tabulae convex to 

tent-shaped, spaced about 0.5 mm apart. Peripheral 

tabellae poorly to well developed. Ratio of tabularium 

width to corallite diameter about 0.3 to 0.4. Increase is 

peripheral, axial, or intermural. 

St,elechophyll~ venukoffi venukoffi (Tolmachev) 

Stylophyllum venukoffi Tolmachev, 1924, p. 318, pl. 19, figs. 9, 10; 

1931, pl. 23, fig. 2. 

Stelechophyllum venukoffi (Tolmachev). Dobrolyubova and 

Kabakovich, 1962, p. 124. 

Stelechophyllum venukoffi venukoffi (Tolmachev). Dobrolyubova and 

Kabakovich, 1966, p. 156, pl. 28, figs. 1, 2, text-fig. 15. 

Diagnos1~s. -Stelechophyllum venuko.ffi with minor 

septa absent or weakly developed and predominantly 

large dissepiments. 

Description of lectotype. -See Tolmachev (1924, p. 

318). 

Occurre'nce. -Lower Carboniferous, upper Tournaisian. Nizhnetersin Horizon, Kuznetsk Basin, U.S.S.R. 

Stelechophyllum venukoffi altaicum (Tolmachev) 

Lithostrotion altaicum Tolmachev, 1924, p. 314, pl. 19, figs. 7, 8; 1931, 

pl. 23, fig. 1. 

Stelechophyllum venukoffi altaicum (Tolmachev). Dobrolyubova and 

Kabakovich, 1966, p. 157, pl. 29, figs. 1-3, pl. 30, fig. 1. 

Diagnosis. -Stelechophyllum venukoffi with minor 

septa well developed and variable, predominantly small 

dissepiments. 

Description of lectotype.- See Tolmachev (1924, p. 

314). 

Discussion. -This subspecies is similar to 

Stelechophyllum microstylum (White) but has fewer major septa, fewer extensions of the septa in the 

dissepimentarium, and a lower ratio of tabularium width 

to corallite diameter. 

Occurrence. -Lower Carboniferous, upper Tournaisian. Nizhenetersin Horizon, Kuznetsk Basin, U.S.S.R. 

Stelechophyllum microstylum species group 

Stelechophyllum microstylum (White} 

Plates 1 and 2 

Lithostrotion microstylum White, 1880a (1883), p. 159, pl. 40, fig. 7a; 

Keyes, 1894, p. 124; Easton, 1944, p. 53, pl. 13, figs. 1-3, pl. 17, 

fig. 1; Bassler, 1950, p. 213, Nelson, 1962, p. 170. 

Lithostrotionella microstyla (White). Bamber, 1961, p. 110, pl. 8, figs. 

4a, b, pl. 9, figs. 1a-h; Bowsher, 1961, pl. 110, figs. 4, 5a-c; 

Carlson, 1964, p. 663, pl. 110, figs. 4, 6. 
12 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

?Lithostrotionella cf. microstyla (White). Bamber, 1961, p. 121, pl. 9, 

figs. 2a-e. 

Lithostrotionella microstylum (White). Easton, 1963, p. 297; Bamber, 

1966, p. 9, pl. 1, figs. 5a-g, pl. 2, figs. 1-3, pl. 3, figs. 1-3. 

Lithostrotionella jasperensis Kelly, 1942, p. 356, pl. 51, figs. 3, 6; 

Nelson, 1960, p. 112, pl. 21, figs. 1-4; 1961, pl. 1, figs. 1-3; 1962, p. 

170; Stensaas and Langenheim, 1960, p. 184, text-figs. 9a, b, lOa, 

b; Easton, 1963, p. 297. 

Lithostrotion [Lithostrotionella] jasperensis (Kelly). Bassler, 1950, p. 

221. 

Lithostrotionella conjluens Easton, 1958, p. 31, pl. 1, fig.12, pl. 2, figs. 

8, 9; Armstrong, 1962, p. 39, text-fig. 18, pl. 4, figs. 1-5, Nelson, 

1962, p. 170; Easton, 1963, p. 297. 

?Lithostrotionella girtyi Hayasaka, 1936, p. 65 [part]. 

Lithostrotionella hemisphaerica Hayasaka, 1936, p. 61 [part]. 

Lithostrotionella multiradiata Hayasaka, 1936, p. 67. 

Diagnosis. -Stelechophyllum with corallite diameter 4 

to 17 mm and 15 to 22 major septa that most commonly 

extend across the tabularium from the columella and into the dissepimentarium. Minor septa well developed. 

Columella ordinarily very well developed. Dissepimentarium composed of 1 to 10 rows (ordinarily 5 rows) of 

small to large, commonly flattened dissepiments. Axial 

tabulae convex to tent-shaped, spaced 2 to 4 in 1 mm. 

Peripheral tabellae well developed. Ratio of tabularium 

width to corallite diameter 0.4 to 0.5. Increase is intermural(?). 

Description of neotype.- According to Easton (1944, 

p. 54), the holotype of S. microstylum was probably lost 

in a fire. USNM specimen 66838, described and illustrated by Bamber (1961, p. 110, pl. 8, figs. 4a, b, pl. 9, 

figs. la-h; 1966, p. 10-11, pl. 1, figs. 5a-g, pl. 2, figs. 3a, 

b), is here designated neotype for this species. 

Discussion. -This is the most widespread species of 

Stelechophyllum in North America, ranging from the 

Mississippi Valley area to western Canada, where its occurrence is exclusively middle and upper Tournaisian 

(upper Kinderhookian and lower Osagean). S. 

microstylum is distinguished from the Russian species 

S. megalum by its greater maximum corallite diameter, 

more numerous major septa, ordinarily stronger columella, and more numerous, generally less inflated 

dissepiments. It is separated from S. banffense by its 

greater maximum corallite diameter, stronger minor 

septa, more numerQUS septal extensions into the 

tabularium, less flattened axial tabulae, and less inflated 

dissepiments. 

Bamber (1966, p. 12-14) has discussed the synonymy 

of Lithostrotionella jasperensis Kelly and Lithostrotionella confluens Easton with S. microstylum,. on the 

basis of study of the type material of these taxa. Three 

of Hayasaka's (1936) specimens are here referred 

definitely to S. microstylum. These are the holotype and 

paratype of Lithostrotionella multiradiata Hayasaka, 

USNM 120244 and 162005, respectively, and a paratype· 

of L. hemisphaerica Hayasaka, USNM 161996. A fourth 

specimen, a paratype of L. girtyi Hayasaka, USNM 

162003, is referred questionably to S. microstylum. 

These specimens are all illustrated herein. 

Occurrence. -Lower Carboniferous, middle and upper 

Tournaisian. Chouteau Limestone and N orthview Shale, 

Missouri, U.S.A.; Gilmore City Limestone, Iowa, 

U.S.A.; Lodgepole Limestone, Montana(?), Idaho, and 

Utah, U.S.A.; Joana Limestone, Nevada, U.S.A.; 

Keating Formation and Lake Valley Limestone, New 

Mexico, U.S.A.; Represo and Venada Formations, 

Sonora, Mexico; Banff and Pekisko Formations, Alberta, Canada. 

Stelechophyllum circinatum (Easton and Gutschick) 

Lithostrotion (Lithostrotionella) circinatus Easton and Gutschick, 

1953, p. 19, pl. 3, figs. 5, 6 [part]. 

Lithostrotionella circinatus (Easton and Gutschick). Nelson, 1962, p. 

170; Easton, 1963, p. 297; Sando, 1969, p. 309, pl. 38, figs. 1-7. 

Diagnosis. -Stelechophyllum with corallite diameter 9 

to 13 mm and 21 to 30 major septa that most commonly 

extend from the columella across the tabularium but 

seldom extend into the dissepimentarium. Minor septa 

absent to poorly developed. Columella absent to very 

well developed. Dissepimentarium composed of 1 to 4 

rows of small to large, commonly flattened dissepiments. Axial tabulae flat to tent-shaped, spaced 4 in 

1 mm. Peripheral tabellae well developed. Ratio of 

tabularium width to corallite diameter 0.4 to 0.5. Increase is peripheral. 

Description of type specimens. -See Easton and 

Gutschick (1953, p. 19). 

Discussion. -This species is similar to S. micr"ostylum 

but differs in having more major septa, a columella that 

is variably developed, and fewer extensions of the major 

septa into the dissepimentarium. S. circinaturn occurs 

at a slightly higher stratigraphic level than S. 

microstylum. 

Occurrence. -Lower Carboniferous, upper Tournaisian. Redwall Limestone, Arizona, U.S.A. 

Stelechophyllum longiseptatum (Lisitsyn) 

Lonsdalia longiseptata Lisitsyn, 1925, p. 68, p. 1, fig. 4. 

Eolithostrotionella longiseptata (Lisitsyn). Zhizhina, 1956, p. 40, pl. 9, 

figs. la, b. 

Diagnosis. -Stelechophyllum with corallite diameter 

15 to 20 mm and 24 to 32 major septa that most commonly extend from the columella across the tabularium 

and into the dissepimentarium. Minor septa well 

developed. Columella well developed. Dissepimentarium 

composed of 2 to 5 rows of small to large inflated 

dissepiments. Axial tabulae convex to tent-shaped, 

spaced 0.2 to 1 mm apart. Peripheral tabellae well 

developed(?). Ratio of tabularium width to eorallite 

diameter 0.4 to 0.5. Mode of increase unknown. 
SYSTEMATIC PALEONTOLOGY 13 

Description of holotype.- See Zhizhina (1956). 

Discussion. -This species is similar to S. circinatum 

but differs in having larger corallites, more major septa, 

a stronger columella, and in having the major septa commonly extending into the dissepimentarium. 

· Occurrence. -Lower Carboniferous, middle Visean. 

Zone C1vd Donetz Basin, U.S.S.R. 

Stelechophyllum banffense (W arran) 

Plate 3; plate 4, figures 1, 2 

Lithostrotion banffensis Warren, 1927, p. 46, pl. 3, figs. 5, 6, pl. 5; 

Criekmay, 1955, 1961, p. 12, pl. 1, figs. 13, 14. 

Lithostrotionella banffensis (Warren). Kelly, 1942, p. 354; Bamber, 

1966, p. 17, pl. 3, fig. 5; Armstrong, 1970a, p. 29, pl. 10, figs. 1-8, pl. 

13, figs. 1-7; 1970b, p. 16, pl. 3, figs. 5-8, pl. 4, figs. 1-4, pl. 10, fig. 

1. 

Lithostrotionella banffense (Warren). Nelson, 1960, p. 119, pl. 23, figs. 

4, 5; 1961, pl. 17, figs. 1, 2. 

Lithostrotionella cf. banffensis (Warren). Bamber, 1961, p. 133, pl. 10, 

figs. 3a-d. 

Lithostrotionella jloriformis Hayasaka, 1936, p. 64, pl. 17, figs. 1a, b. 

Lithostrotionella vesicularis Hayasaka, 1936, p. 68, pl. 14, figs. 3a, b. 

?Lithostrotionella americana Hayasaka, 1936, p. 62 [part]. 

?Lithostrotionella aff. L. banffensis (Warren). Armstrong, 1970b, p. 

19, pl. 4, figs. 5, 6. 

Diagnosis. -Stelechophyllum with corallite diameter 6 

to 13 mm and 19 to 26 major septa of variable length 

that seldom join the columella and seldom extend into 

the dissepimentarium. Minor septa absent to poorly 

developed. Columella well developed. Dissepimentarium 

composed of 1 to 6 rows of small to large inflated 

dissepiments. Axial tabulae ordinarily broadly tentshaped, spaced 2 to 4 in 1 mm. Peripheral tabellae poorly to well developed. Ratio of tabularium width to corallite diameter 0.5 to 0.6. Increase is peripheral. 

Description of lectotype.- See Nelson (1960) and 

Bamber (1966). 

Discussion. -This highly variable species is common in 

rocks of early and middle Visean age in western Canada 

and Alaska. It appears to have been derived from S. 

microstylum, from which it differs in having smaller corallites, weaker major and minor septa, somewhat flattened tabulae that are variable in form, ordinarily fewer 

rows of dissepiments, and a larger ratio of tabularium 

width to corallite diameter. 

Arp1strong (1970a, p. 29-31) has discussed the 

synonymy of Lithostrotionellafloriformis Hayasaka and 

Lithostrotionella vesicularis Hayasaka with S. banffense 

on the basis of study of the type material. Hayasaka's 

type specimens are illustrated in Armstrong's paper. 

Three para types of Lithostrotionella americana (USNM 

174372, 174374, and 174375) are here referred to the 

species with query and are illustrated herein. 

Oceurrence.- Lower Carboniferous, lower and middle 

Visean. Mount Head Formation, Alberta, Canada; 

Prophet Formation, British Columbia, Canada; 

Peratrovich, Nasorak, Kogruk, and Alapah(?) Formations, Alaska, U.S.A.; Little Flat Formation, Idaho, 

U.S.A.(?). 

Stelechophyllum micrum species group 

Stelechophyllum micrum (Kelly) 

Lithostrotionella micra Kelly, 1942, p. 357, pl. 50, fig. 7; Nelson, 1960, 

p. 113, pl. 21, figs. 5, 6; 1961, pl. 6, figs. 1-3; Bamber, 1966, p. 14, 

pl. 3, figs. 4a-e. 

Lithostrotion [Lithostrotionella] micra (Kelly). Bassler, 1950, p. 221. 

Lithostrotion micra (Kelly). Brindle, 1960, pl. 10, fig. 2. 

Diagnos1:s.-Stelechophyllum with corallite diameter 2 

to 6 mm and 10 to 15 major septa that most commonly 

extend. from the columella across the tabularium but 

seldom extend into the dissepimentarium. Minor septa 

poorly developed. Columella poorly to well developed. 

Dissepimentarium composed of 1 to 2 rows of large inflated dissepiments. Axial tabulae convex, spaced 2 in 1 

mm. Peripheral tabellae poorly developed. Ratio of 

tabularium width to corallite diameter about 0.6. Increase is peripheral(?). 

Descript-ion ofholotype.- See Kelly (1942) and Bamber 

(1961). 

Discussion.-This species is similar to Stelechophyllum 

lochmctnae and S. ergunjaicum. S. lochmanae has larger 

corallites and stronger major septa. S. ergunjaicum has 

slightly more major septa and only one row of 

dissepiments. 

Occurrence.- Lower Carboniferous, upper Tournaisian. Pekisko and Shunda Formations, Alberta, Canada. 

Stelechophyllum lochmanae (Armstrong) 

LithostTotionella lochmanae Armstrong, 19()2, p. 38, pl. 4, figs. 6-8, 

text fig. 17. 

Diagnos'1:s.-Stelechophyllum with corallite diameter 5 

to 8 mm and 13 to 15 major septa that most commonly 

extend from the columella across the tabularium and into the dissepimentarium. Minor septa well developed. 

Columella well developed. Dissepimentarium composed 

of 1 to 2 rows of large inflated dissepiments. Axial 

tabulae convex, spaced 3 in 1 mm. Peripheral tabellae 

poorly developed. Ratio of tabularium width to corallite 

diameter about 0.6. Mode of increase unknown. 

Description of type material.- See Armstrong (1962). 

Discussion.-This species is similar to S. micrum and 

S. ergunja,icum. S. micrum has smaller corallites and 

weaker minor septa, and S. ergunJ·aicum has weaker 

minor septa and fewer extensions of the major septa into the dissepimentarium. 

Occurrence.- Lower Carboniferous, middle Tournaisian. Keating Formation and Lake Valley Limestone, 

New M:exico and Arizona, U.S.A. 

Stelechophyllum ergunjaicum (Onoprienko) 

Lithostrotionella ergunjaicum Onoprienko, 1976, p. 29, pl. 11, figs. 3, 4. 
14 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Diagnosis. -Stelechophyllum with corallite diameter 4 

to 6 mm and 15 to 16 major septa that most commonly 

extend from the columella across the tabularium but 

seldom extend into the dissepimentarium. Minor septa 

poorly developed. Columella well developed. Dissepimentarium composed of a single row of large inflated dissepiments. Axial tabulae convex to tentshaped, spaced 0.1-1.2 mm apart. Peripheral tabellae 

well developed. Ratio of tabularium width to corallite 

diameter 0.5 to 0.6. Mode of increase unknown. 

Description of type material.- See Onoprienko (1976). 

Discussion. -This species is similar to the North 

American species S. lochmanae and S. micrum. S. 

ergunjaicum differs from S. micrum by having convex 

to tent-shaped tabulae, better developed peripheral 

tabellae, and only one row of dissepiments. The Russian 

species differs from S. lochmanae by having smaller corallites, weaker minor septa, fewer extensions of the major septa into the dissepimentarium, and only one row of 

dissepiments. 

Occurrence.- Lower Carboniferous, upper Visean. 

Utaveem Suite, East Chukotka, Chegitun' River basin, 

U.S.S.R. 

Stelechophyllum? mclareni species group 

Stelechophyllum? mclareni (Sutherland) 

Lithostrotion sp. McLaren and Sutherland, 1949, p. 631, pl. 103, figs. 

1-9. 

Lithostrotion [Lithostrotionella] [Thysanophyllum] mclareni 

Sutherland, 1958, p. 95, pl. 33, figs. la-g. 

Lithostrotionella mclareni (Sutherland). Armstrong, 1970b, p. 21, pl. 

5, figs. 4, 7, 8-10, pl. 6, figs. 1, 2, 7-9. 

?Lithostrotionella aff. L. mclareni Sutherland. Armstrong, 1970a, p. 

35, pl. 8, figs. 8, 9. 

Diagnosis.-Stelechophyllum? with corallite diameter 

3.5 to 4.4 mm and 12 to 15 major septa of variable length 

that may or may not join the columella and may or may 

not extend into the dissepimentarium. Minor septa absent to well developed. Columella weakly developed (or 

discontinuous) to strongly developed. Dissepimentarium 

composed of 1 to 3 rows of small to large inflated 

dissepiments. Axial tabulae flat to tent-shaped, depending on strength of columella, spaced 2 in 1 mm 

Peripheral tabellae absent to poorly developed. Ratio of 

tabularium width to corallite diameter about 0.5. Increase is peripheral. 

Description of holotype.- See McLaren and Sutherland (1949). 

Discussion.- McLaren and Sutherland (1949), 

Sutherland (1958), and Armstrong (1970b) have discussed in detail the extreme morphologic variability in 

this species. According to E. W. Bamber (written commun., 1980), who has studied the species, serial sections 

of the corallites show vertical discontinuity of the septa 

in the· dissepimentarium and vertical discontinunity of 

the colummella so that some sections have the appearance of Sciophyllum in corallites that are mainly 

characterized by the Stelechophyllum structure. The 

amount of variation is notable in this species. 

S.? mclareni is similar to S.? birdi and S.? niakense, 

from which it differs in corallite diameter, in number of 

major septa, and in the degree to which the major septa 

extend into the dissepimentarium Armstrong's (1970a) 

Lithostrotionella aff. L. mclareni has characters that 

place it close to S.? birdi and is here referred to S.? 

mclareni with query. 

Occurrence.- Lower Carboniferous, Visean. Prophet 

Formation, British Columbia, Canada; Kogruk Formation, Alaska, U.S.A. 

Stelechophyllum? birdi (Armstrong) 

Lithostrotionella birdi Armstrong, 1970a, p. 32, pl. 7, figs. 1-4, pl. 8, 

figs. 1-7; 1970b, p. 20, pl. 4, fig. 1, pl. 5, figs 1-3, 5, 6. 

Diagnosis. -Stelechophyllum? with corallite diameter 

3.5 to 7.5 mm and 15 to 20 major septa of variable length 

that seldom join the columella and may or may not extend into the dissepimentarium. Minor septa absent to 

poorly developed. Columella absent to weakly developed. Dissepimentarium composed of 1 to 2 rows of 

mostly large inflated dissepiments. Axial tabulae flat to 

flattened tent-shaped, spaced 2 in 1 mm. Peripheral 

tabellae absent to poorly developed. Ratio of tabularium 

width to corallite diameter about 0.6. Increase is 

peripheral. 

Description of holotype.- See Armstrong (1970a). 

Discussion.- This species shows intracolony variation 

similar to that of S.? mclareni (see discussion of S.? 

mclareni). It is distinguished from S.? mclareni by its 

larger corallites and larger number of major septa. Its 

extreme morphologic variation makes generie placement questionable. 

Occurrence.- Lower Carboniferous, Visean. Peratrovich-and Kogruk Formations, Alaska, U.S.A. 

Stelechophyllum? niakense (Armstrong) 

Lithostrotionella sp. A. Armstrong, 1970b, p. 25, pl. 7, figs. 1-6. 

Lithostrotionella sp. B. Armstrong, 1970b, p. 26, pl. 7, figs. 7-9. 

Lithostrotionella.niakensis Armstrong, 1972a, p. A23, pl. 4, figs. 2, 6, 

pl. 9, figs. 1-3, pl. 9, figs. 1-6. 

Diagnosis. -Stelechophyllum? with corallite diameter 

2.5 to 4.7 mm and 9 to 14 major septa of variable length 

that seldom extend from the columella across the 

tabularium but commonly extend into the dissepimentarium. Minor septa poorly to well developed. Columella 

absent to poorly developed. Dissepimentarium composed of 1 to 2 rows of mostly large inflated dissepiments. Axial tabulae flat to tent-shaped, spaced 2 in 

1 mm. Peripheral tabellae absent to poorly developed. 

. Ratio of tabularium width to corallite diameter about 

0.6. Increase is prob~bly peripheral. 
SYSTEMATIC PALEONTOLOGY 15 

Description of holotype.- See Armstrong (1972a). 

Discussion. -S.? niakense is similar to S.? mclareni 

but differs in having fewer major septa, more extensions 

of the major septa into the dissepimentarium, and less 

polymorphic variation between corallites. S.? niakense 

differs from S.? macouni in having larger corallites, 

more major septa, and fewer extensions of the major 

septa to the columella. 

Occurrence. -Lower Carboniferous, Visean. Kogruk 

Formation, Alaska, U.S.A. 

Stelechophyllwn macouni species group 

Stelechophyllwn? macouni (Lambe) 

Lithostrotion macouni Lambe, 1899, p. 220; 1901, p. 176, pl. 14, figs. 

11, 11a, 11b. · 

L.ithostrotion? macouni Lambe. Nelson, 1960, p. 122, pl. 23, figs. 1, 2 

[part]. 

Lithostrotionella macouni (Lambe). Armstrong, 1970b, p. 23, pl. 12, 

figs. 1-6. 

Diagnosis. -Stelechophyllum? with corallite diameter 

1.9 to ~~.8 mm and 9 to 11 major septa that commonly extend from the columella across the tabularium and into 

the dissepimentarium. Minor septa absent to poorly 

developed. Columella ordinarily well developed. 

Dissepimentarium composed of a single row of small inflated dissepiments. Axial tabulae nearly flat to ~onical, 

spaced 3 in 1 mm. Peripheral tabellae poorly developed. 

Ratio of tabularium width to corallite diameter 0.7. 

Mode of increase unknown. 

Description of lectotype.- GSC 4327 is here designated 

lectotype for this species. See Nelson (1960) and Armstrong (1970b) for description of this specimen. 

Discussion. -This species is distinguished from all 

other species of Stelechophyllum by its smaller corallite 

diameter, fewer major septa, and very weakly developed 

lonsdaleoid dissepimentarium. Its morphology approaches that of Lithostrotion (Lithostrotion), which is 

unknown in North America 

Occurrence. -Lower Carboniferous, Visean. Prophet 

Formation, British Columbia, Canada. 

Stelechophyllwn sp. indet. 

Plate 4, figures 3, 4 

Lithost·rotionella hemisphaerica Hayasaka, 1936, p. 61 [part]. 

Discussion.- An indeterminate species of 

Stelechophyllum is represented by USNM 120239, which 

is a paratype of Lithostrotionella hemisphaerica 

Hayasaka. This specimen has corallites 9 to 12 mm in 

diameter, 18 to 20 major septa, and an impersistent columella, but the corallites are too badly crushed to permit 

evaluation of all specific characters. 

Occurrence. -Lower Carboniferous, Tournaisian. 

Madison Limestone, Utah, U.S.A. 

Stelechophyllwn? sp. indet. 

Lithostrotionella sp. Bamber, 1961, p. 129, pl. 10, figs. 2a-d. 

Discussion.- Bamber's (1961) specimen has corallites 

4 to 8 mm in diameter, 19 to 21 major septa, a simple columella, and tabulae that appear to be of the 

Stelechophyllum type. Formal recognition of this species 

is deferred pending discovery of more specimens. 

Occurrrence. -Lower Carboniferous, Visean. Prophet 

Formation, British Columbia, Canada. 

Genus AULOSTYLUS Sando, 1976 

Aulostylus Sando, ·1976, p. 427. 

Type species. - Lithostrotionella tubifera Hayasaka, 

1936, p. 69, pl. 16, figs. 1a, b, 2. Lower Carboniferous 

(Tournaisian), Montana. 

Diagnosis.- See Sando (1976). 

Discu,ssion. -This genus was established for lithostrotionelloid corals that have an aulos and a weak columella. Two middle Tournaisian species from the U.S.A. 

and Canada and a possible representative from the 

Visean of China are the only described species. The 

species formerly referred to Lithostrotionella are listed 

below; pertinent information on them has been given by 

Sando (1976). 

Aulostylus tubiferus (Hayasaka) 

Lithostrotionella tubifera Hayasaka, 1936, p. 69, pl. 16, figs. 1a, b, 2; 

Smith and Yii, 1943, p. 42; S~ndo, 1963, p. 1076. 

Lithostrotion [Lithostrotionella] tubifera (Hayasaka). Bassler, 1950, 

p. 220. 

A ulina tubifera (Hayasaka). Hill, 1940, p. 190; Bamber, 1961, p. 161, 

pl. 12, figs. 3a-d. 

Aulostyl·us tubiferus (Hayasaka). Sando, 1976, p. 428, figs. 3, 4. 

Description of holotype. -See Sando (1976). 

Diagnosis.- See Sando (1976). 

Discussion. -Sando (1976) recognized two subspecies, 

A. tub1jeru.s tubiferus and A. tubiferus eotubiferus. The 

holotype of A. tubiferus tubiferus is probably from the 

lower part of the Mission Canyon Limestone in Montana 

rather than from the W oodhurst Member of the 

Lodgepole Limestone as stated by Sando (1976). 

Aulostylus? sp. 

Lithostrotionella sp. A Lo and Chao, 1962, p. 184, pl. 19, fig. 1. 

AulostyLus? sp. Sando, 1976, p. 431. 

Family ACROCYATHIDAE new family 

Diagnos1:s. - Cerioid and fasciculate colonial Rugosa 

that have two orders of septa, a columella that ranges 

from a simple axial plate joined to the counter septum to 

a complex spiderweb structure made up of axial plate, 

septal lamellae, and axial tabellae; ordinarily complete 

conical tabulae; and a lonsdaleoid dissepimentarium. 

Type genus. -The type and only included genus is 

Acrocyathus d'Orbigny, 1849. 

Genus ACROCYATHUS d'Orbigny, 1849 

Astraea Castelnau, 1843, p. 45 (not Fischer von Waldheim, 1830, pl. 

31, fig. 3). 
16 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Acrocyathus d'Orbigny, 1849a, p. 12; 1850, p. 160; 1852, p. 184; 

Thevenin in Boule and others, 1906, expl. pl. 6; 1907, p. 90; 1923, 

p. 90; Easton, 1973, p. 130, 132 [part]; Minato and Kato, 1974, p. 

70 [part]. 

Lithostrotion Fleming. Milne-Edwards and Haime, 1851, p. 433, 483 

[part]; Owen, 1852, expl. pl. 4; Hall in Hall and Whitney, 1858, p. 

667; Milne-Edwards, 1860, p. 423 [part]; Owen, 1862, p. 364; Rominger, 1876, p. 111; White, 1880a (1883), p. 159 [part]; 1880b, p. 

506; 1882, p. 401; Weller, 1898, p. 329 [part]; Ulrich, 1905, p. 33; 

Butts, 1917, p. 46; 1926, p. 176; 1941, p. 239; Shimer, 1926, p. 27 

[part]; Morse, 1930, p. 104; Weller, 1931, p. 274; Kelly, 1942, p. 

361 (part]; Allen and Lester, 1954, p. 101. 

?Lithostrotion (Lithostrotionella) Yabe and Hayasaka, 1915, p. 133, 

1920, p. 11; Bassler, 1950, p. 217, 221. 

Lithostrotionella Yabe and Hayasaka. Hayasaka, 1936, p. 58, 61, 62, 

65 (part]; Kelly, 1942, p. 352 [part]; Parks, 1951, p. 180; Weller and 

others, 1952, p. 84; Allen and Lester, 1954, p. 101; Nelson, 1960, p. 

117, 118 (part]; 1961, pl. 17 (part]; Yoh, 1961, p. 8; Bamber, 1961, 

p. 107 [part]; 1966, p. 19 [part]; Armstrong, 1962, p. 39 [part]; 

1970a, p. 31 [part]; Wu in Yu and others, 1963, p. 86; Nations, 

1963, p. 1257; Minato and Kato, 1974, p. 72 (part]. 

Lonsdaleia McCoy. Crickmay, 1955, 1961, p. 13. 

Eolithostrotionella Zhizhina. Vasilyuk, 1960, p. 95; Zhizhina in 

Bul'vanker and others, 1960, p. 250, 251, 252; Degtyarev, 1973, p. 

192, 193. 

Type species. -Acrocyathus floriformis d'Orbigny, 

1849, p. 12 (by monotypy). Lower Carboniferous 

(Visean), Indiana, U.S.A. 

Diagnosis. -Ordinarily cerioid colonial corals with 

. tabular to hemispherical growth form; one fasciculate 

species is known and cerioid-fasciculate coralla are rare. 

Septa of two orders. Major septa thin, seldom extending 

across tabularium to columella and ordinarily discontinuous or absent in dissepimentarium. Cardinal septum 

short and situated in a fossula formed by downwarped 

tabulae in some species. Counter septum commonly long 

and extending to columella. Minor septa ordinarily absent or poorly developed. Columella ranging from a simple axial plate joined to counter septum to a complex 

spiderweb structure made up of an axial plate, septal 

lamellae, and axial tabellae or upturned edges of 

tabulae. Tabulae ordinarily complete, conical, without 

well-defined shoulders, but may be nearly horizontal in 

some species. Peripheral tabellae rare. Dissepimentarium lonsdaleoid, commonly showing traces or crests 

of discontinuous septa in transverse section. Increase 

peripheral. 

Discussion. - d'Orbigny's (1849a) original proposal and 

subsequent citations (1849b, 1852) of Acrocyathus provided only brief diagnoses of the genus, and Thevenin's 

(in Boule, 1906, 1907, 1923) reinvestigation of the type 

specimen added only illustrations of the exterior of the 

specimen. Most earlier authors (Milne-Edwards and 

Haime, 1851, p. 432, 433; 1852, p. 192; Lindstrom, 1883, 

p. 5, 11; de Koninck, 1872, p. 26; Thevenin, in Boule, 

1907, 1923, p. 90; Sanford, 1939, p. 405; Soshkina and 

others, 1962, p. 336) regarded Acrocyathus as a junior 

synonym of Lithostrotion Fleming, although de 

Fromentel (1861, 304) thought it was a Diphyphyllum 

Lonsdale, and ,Wang (1950, p. 212) placed it in 

synonymy with Lonsdaleia McCoy. More recently, some 

authors (Hill, 1956, p. F307; lvanovskiy,-1967, p. 34, and 

Cotton, 1973, p. 12) linked it questionably with Lithostrotionella Y abe and Hayasaka. 

Easton (1973) redescribed in detail the type specimen 

and illustrated thin sections of it, thus providing the first 

adequate basis for a determination of the morphology 

and affinities of the type species. Easton regarded 

Acrocyathus as a senior synonym of Lithostrotionella 

and espoused a broad generic concept that included 

species placed herein in Stelechophyllum Tolmaehev and 

Petalaxis Milne-Edwards and Haime. Minato and Kato 

(1974, p. 70-71) regarded Acrocyathus as a distinct 

genus distinguished from Lithostrotionella by having a 

complex axial structure rather than a simple ax:ia[ plate. 

In my opinion, Acrocyathus is a distinct genus 

separated from Stelechophyllum and Petalax1:s by its 

complex axial structure and complete conical tabulae. It 

is distinguished from Lithostrotion by its lonsdaleoid 

dissepimentarium and complex axial structure and from 

Lonsdaleia by its lack of a distinct separation of axial 

and peripheral series of tabellae. Although the morphology of the type species of Lithostrotionella is in 

doubt, Lithostrotionella is regarded as a probable junior 

synonym (see p. 4). 

Eolithostrotionella is included in the synonymy of 

Acrocyathus because some of the Russian species were 

referred to that genus. The type species of Eol·ithostrotionella is a Stelechophyllum. 

Species of Acrocyathus are distinguished on differences in mature corallite diameter, number of major 

septa at maturity, development of major septa, development of minor septa, complexity of the columella, shape 

and spacing of the tabulae, size and number of 

dissepiments, number of rows of dissepiments, and ratio 

of tabularium width to corallite diameter. Most of the 

species of Acrocyathus have a cerioid corallum, but one 

fasciculate species, A. proliferus, is includE~d here 

because of its internal morphologic similarity to cerioid 

species and the presence of transitional forms. Cerioid 

species included here are the abundantly represented A. 

floriformis from the Eastern United States; A. pilatus 

from the Mississippi Valley; A. girtyi from the VI/ estern 

United States; A. pennsylvanicus from western Canada; 

A. utkae, A. rotai, A. cystosus, and A. lissitzini from the 

U.S.S.R.; and A. hsuJ'iulingi from China. Questionable 

forms include A.? unicus and A.? zhizhinae, whose morphology is in doubt, and A.? shimeri and A.? grechovkae, 

which have essentially horizontal tabulae like Petalaxis 

but have a complex columella characteristic of 

Acrocyathus. 
SYSTEMATIC PALEONTOLOGY 17 

Occurrence. -Lower Carboniferous, lower to upper 

Visean. U.S.S.R., U.S.A., Canada, China. 

Acrocyathus floriformis d'Orbigny 

Plates 5-14, 16 

Not Astraea mamillaris Fischer von Waldheim, 1830, pl. 31, fig. 3 

Astraea mamillaris Castelnau, 1843, p. 45, pl. 24, fig. 5. 

Acrocyathu,sjloriformis d'Orbigny, 1849a, p. 12; 1850, p. 160; 1852, p. 

184; Thevenin in Boule and others, 1906, expl. pl. 6, pl. 6, figs. 1, 2; 

1907, p. 90; 1923, p. 90, expl. pl. 6, pl. 6, figs. 1, 2; Easton, 1973, p. 

130, pl. 1, figs. 1a-f. 

Acrocyathu,s sp. Easton, 1973, p. 132, pl. 1, figs. 2a-b. 

Not Ax1:nura canadensis Castelnau, 1843, p. 49, pl. 24, fig. 4. 

Lithost?·otion mamillare Milne-Edwards and Haime, 1851, p. 433, pl. 

13, fig. 1 [part]; Hall in Hall and Whitney, 1858, p. 667, pl. 24, figs. 

5a, b; Rominger, 1876, p. 111, pl. 55, upper-right figure [part]. 

LithostTotion mamillare (Castelnau). White, 1880a (1883), p. 159, pl. 

40, figs. 6a, b; 1880b, p. 506, pl. 6, figs. 1, 2; 1882, p. 401, pl. 52, 

fig. 3. 

Not LithostTotion mamillare(?) Meek, 1864, p. 5, pl. 1, figs. 4-4b. 

Lithost1·otion canadense Milne-Edwards and Haime, 1851, p. 483, pl. 

13, fig. 1 [part]; Milne-Edwards, 1860, p. 423; Owen, 1862, p. 364, 

fig. 6; Butts, 1926, p. 176, pl. 58, figs. 12, 13; Morse, 1930, p. 104, 

pl. 9; Weller, 1931, p. 274, pl. 36, figs. 1a, b. 

Lithost1·otion canadense (Castelnau). Weller, 1898, p. 329 [part]. 

Lithost1·otion? canadense Ulrich, 1905, p. 33, pl. 3, figs. 1, 2. 

Lithost1·otion "canadensis" (Castelnau). Butts, 1941, p. 239, pl. 129, 

fig. 3. 

Lithostrotion basaltiforme Owen, 1852, expl. pl. 4, pl. 4, figs. 5, 6; 

Butts, 1917, p. 46, pl. 11, figs. 1, 2. · 

Lithost?·otionella castelnaui Hayasaka, 1936, p. 58, pl. 11, figs. 1, 2 

[part]; Weller and others, 1952, p. 84, pl. 1, figs. 6, 7; Allen and 

Lester, 1954, p. 101, pl. 26, fig. 1. 

Lithostrotian [LithostTotionella] castelnaui (Hayasaka). Bassler, 1950, 

p. 217 [part]. 

Lithost1•otionella a?nericana Hayasaka, 1936, p. 62, pl. 14, figs. 1, 2 

[part]. 

Lithostrotion [Lithostrotionella] a?nericanum Bassler, 1950, p. 217 

[part). 

Lithostrotionella hemisphaerica Hayasaka, 1936, p. 61, pl. 12, fig. 1, 

pl. 13, figs. 1, 2 [part). 

Lithostrotion [Lithostrotionella} hemisphericum (Hayasaka). Bassler, 

1950, p. 217 [part). 

Not Lithostrotionellajloriformis Hayasaka, 1936, p. 64, pl. 17, fig. 1. 

Material studied. -MNHN 1140 (holotype), FL 411; 

USNl\1 756, 3779, 8211, 13669, 15526, 17071, 17848, 

37466, 39654, 42695, 42766, 71646, 98102, 120235, 

120236, 120237, 120238, 120240, 120241, 135092, 

135094, 135096, 135097, 135172, 135173, 135174, 

135177, 135179, 135300, 135301, 135302, 136704, 

161989, 161990, 161991, 161992, 161993, 161994, 

161995, 161997, 161998, 161999, 162000, 162001, 

166604, 166605, 174376, 174377, 216198, 216199, 

216200, 216202, 216203, 216204, 216205, 216206, 

216207, 216208, 216210, 216211, 216212, 216214. 

Diagnosis. -Predominantly cerioid Acrocyathus with 

corallite diameter 10 to 30 mm and 20 to 40 major septa 

that are ordinarily short and do not extend across the 

tabula.rium to the columella and seldom extend into the 

dissepimentarium. Cardinal fossula ordinarily well 

developed. Counter septum commonly joined to columella. Minor septa weakly developed as crests on 

dissepiments. Columella highly variable, ranging from a. 

simple axial plate joined to the counter septum to a complex spiderweb structure made up of an axial plate, septal lamellae, and axial tabellae or upturned edges of 

tabulae; thysanophylloid variants are known. Dissepimentarium composed of 1 to 5 rows of large and 

small, ordinarily inflated dissepiments. Tabulae complete, conical, sharply deflected upward at columella and 

without shoulders or with poorly defined shoulders and 

spaced about 1 mm apart. Peripheral tabellae rare. Axial tabellae common. Ratio of tabularium width to corallite diameter 0.4 to 0.6. Increase is peripheral. 

Descript1:on of holotype. -See Easton (1973). 

Disc'ussion. -This species is characterized by the 

highly variable structure of the columella and the great 

range in corallite diameter and number of major septa 

(fig. 3). The variation in the latter two characters appears to be continuous and does not permit recognition 

of separate species or subspecies. The species has been 

divided into two subspecies on the basis of the complexity of the columella (see below). Similarities between 

Acrocyathus floriformis and the fasciculate species A. 

proliferus are discussed under A. proliferus. 

Occurrence. -A. floriformis is abundant in and 

characteristic of the St. Louis Limestone and correlative 

formations in the Southeastern Province of Sando and 

others (1975, 1977). Data on its occurrence are noted 

under th~ two subspecies of the species. 

Acrocyathus floriformis floriformis d'Orbigny 

Plates 5-11; plate 16, figure 2 

Not AstTaea ·mamillaris Fischer von Waldheim, 1830, pl. 31, fig. 3. 

AstTaea marnillaris Castelnau, 1843, p. 45, pl. 24, fig. 5. 

Acrocyathu,sfloriformis d'Orbigny, 1849a, p. 12; 1850, p. 160; 1852, p. 

184; Thevenin in Boule, 1906, expl. pl. 6, pl. 6, figs. 1, 2; 1907, p. 90; 

1923, p. 90, expl. pl. 6, pl. 6, figs. 1, 2; Easton, 1973, p. 130, pl. 1, 

figs. 1a-f. 

?Lithostrotion mamillare Milne-Edwards and Haime, 1851, p. 433, pl. · 

13, fig. 1 [part]; Rominger, 1876, p. 111, pl. 55, upper right figure 

[part]. 

Not Lithostrotion mamillare (Castelnau). White, 1880a (1883), p. 159, 

pl. 40, figs. 6a, b; 1880b, p. 506, pl. 6, figs. 1, 2; 1882, p. 401, pl. 52, 

fig. 3. 

Not Lithostrotion mamillare(?) Meek, 1864, p. 5, pl. 1, figs. 4-4b. 

?Lithosf:rotion canadense Milne-Edwards and Haime, 1851, p. 483, pl. 

13, fig. 1 [part]. 

Lithostrotion canadense Butts, 1926, p. 176, pl. 58, figs. 12, 13. 

Lithostrotion? canadense Ulrich, 1905, p. 33, pl. 3, figs. 1, 2. 

?Lithostrotion basaltiforme Owen, 1852, expl. pl. 4, pl. 4, figs. 5, 6. 

Lithostrotion basaltiforme Owen. Butts, 1917, p. 46, pl. 11, figs. 1, 2. 

Lithostrotionella castelnaui Hayasaka, 1936, p. 58, pl. 11, figs. 1, 2. 

Lithost'rotion [Lithostrotionella] castelnaui (Hayasaka). Bassler, 1950, 

p. 217. 

Lithostrotion.ella a?nericana Hayasaka, 1936, p. 62, pl. 1.4, figs. 1, 2 

[part]. 
18 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

40~------------~------------~------------~--------------------------~------------~ 

35 

c::x: 

Ia.. 

w 

C/) 

a: 

0 

-, 

c::x: 

~ 

L.L 

0 

a: 

w 

Ill 

~ 

:::) 

z 

25 

~ 

:::) 

~ 

X 

c::x: 

~ 

J5L--L~--~~--L--L--~~~--L-~~--~~--~~--~~~--~~--~~--~~~--~-~--~~ 

10 15 20 25 30 35 40 

MAXIMUM CORALLITE DIAMETER (MM) 

FIGURE 3.-Variation in maximum number of major septa and maximum corallite diameter in 56 coralla of Acrocyathus floriformis. Polygon 

defines field of ¥ariation of coralla from one locality (Livingston, Tenn.). Numbers beside dots indicate more than one corallum. 

Lithostrotion [Lithostrotionella] americanum Bassler, 1950, p. 217. 

Lithostrotionella hemisphaerica Hayasaka, 1936, p. 61, pl. 12, fig. 1, 

pl. 13, figs. 1, 2 [part]. 

Lithostrotion [Lithostrotionella] hemisphericum (Hayasaka). Bassler, 

1950, p. 217 [part]. 

Not Lithostrotionellafloriformis Hayasaka, 1936, p. 64, pl. 17, fig. 1. 

Description of holotype.- See Easton (1973). 

Material studied. -MNHN 1140 (holotype), FL 411; 

USNM 13669, 71646, 120235, 120238, 120240, 

120241(?), 161989, 161990, 161991, 161993, 161994, 

_161995, 161997, 162001(?), 174376, 174377(?), 216205(?), 

216206. 

Diagnosis. -Acrocyathus floriformis having an axial 

structure ranging from a simple axial plate to a complex 

structure composed of an axial plate, a few axial 

lamellae, and a few concentric intercepts of axial 

tabellae or upturned edges of axial tabulae in transverse 

section. 

Discussion. -This predominant subspecies of the 

widespread species A. florijorniis is distinguished from 

A. floriformis hemisphaericus by its less complex columella. The two subspecies occur in the same formations 

in the same geographic areas. 
SYSTEMATIC PALEONTOLOGY 19 

Occurrence. -Lower Carboniferous, middle or upper ' 

Visean. St. Louis Limestone, Indiana, Illinois, Iowa, 

Kentucky, Missouri, Tennessee, U.S.A.; Greenbrier 

Limestone and Newman Limestone, Virginia, U.S.A.; 

Hillsdale Member of Greenbrier Limestone, West 

Virginia, U.S.A.; Newman Limestone, Kentucky, 

U.S.A.; Tuscumbia Limestone, Alabama, U.S.A. 

Acrocyathus floriformis hemisphaericus (Hayasaka) 

Plates 12-14 

Lithostrotion mamillare (Castelnau). White, 1880a, p. 159, pl. 40, figs. 

1a, b. 

Lithostrotion "canadensis" (Castelnau). Butts, 1941, p. 239, pl. 129, 

fig. 3. 

Lithostrotion castelnaui Hayasaka, 1936, p. 58 [part]. 

Lithostrotion [Lithostrotionella] castelnaui (Hayasaka). Bassler, 1950, 

p. 217 [part]. 

Lithostrotionella hemisphaerica Hayasaka, 1936, p. 61, pl. 12, 13, figs. 

2a, b. [part]. 

Lithostrotion [Lithostrotionella] hemisphericum (Hayasaka). Bassler, 

1950, p. 217. 

Material studied.- USNM 8211, 98102, 120236, 

120237 (holotype), 161992, 161998, 161999, 162000(?). 

Diagnosis.-Acrocyathus. floriforrnis having an axial 

structure ranging from a simple axial plate to a very 

complex structure composed of an axial plate, many axial lamellae, and many concentric intercepts of axial 

tabellae or upturned edges of axial tabulae in transverse 

section. 

Description of holotype. -The specimen is a fragment 

of a large, silicified, cerioid, hemispherical colony. It is 

about 15 em high and was more than 18 em in diameter. 

Internal structures are poorly preserved. 

Corallites are polygonal, have straight double-layered 

walls as much as 0.25 mm thick, and range from 10 to 15 

mm in diameter at maturity. Increase is peripheral. The 

major septa seldom extend to the columella but commonly extend into the dissepimentarium. The cardinal 

septum is shorter than the other major septa, and the 

counter septum commonly joins the columella. There are 

26 to 32 major septa at maturity. Minor septa are 

relatively well developed at the corallite walls and as 

crests on the dissepiments. The columella ranges from a 

simple axial .plate joined by a few septal lamellae in 

young corallites to a very complex structure composed 

of an axial plate, many septal lamellae, and as many as 4 

concentric traces of axial tabellae; the columella is as 

much as 2.5 mm in diameter. Tabulae are ordinarily 

complete, conical, curved upward to the columella, and 

spaced 0.25 to 1 mm apart. The dissepimentarium is 

weakly lonsdaleoid and consists of 2 to 3 rows of inflated 

to flattened dissepiments of varying sizes. There are ordinarily 2 or 3 dissepiments in 1 mm. The ratio of the 

tabularium width to the corallite diameter is about 0.6 to 

0. 7. Septal microstructure has been destroyed by 

silicification. 

Discussion. -This subspecies is distinguished by its 

compact, very complex columella. The holotype (pl. 12, 

figs. 1, 2) is a poor example of the species because of the 

poor preservation of internal structures. A paratype, 

USNM 161H98, is a much better example. This specimen 

pl. 12, figs. 3, 4) has as many as 5 tabellar rings around 

the medial plate. The other paratypes have fewer 

tabellar rings than USNM 161998, but all have a more 

complex columella than is found in A. floriforrnis 

floriforrn is. 

The transverse section of this subspecies mimics that 

ofLonsdaleia(Actinocyathus) because of its complex columella and lonsdaleoid dissepimentarium. It is readily 

distinguished from Lonsdaleia (Actinocyathus) in 

longitudinal section by lacking a distinct continuous 

separation of axial and peripheral tabellae. The 

longitudinal section (pl. 12, fig. 3) also shows that the 

complexity of the columella ranges from a simple structure like that pf A. floriformis floriforrnis to a complex 

structure in the same corallite. 

Occu'rrence. -Lower Carboniferous, middle or upper 

Visean. St. Louis Limestone, Illinois, Indiana, and 

Missouri, U.S.A.; Newman Limestone and Greenbrier 

Limestone, Virginia, U.S.A.; Hillsdale Mamber of 

Greenbrier Limestone, West Virginia, U.S.A. 

Acrocyathus pilatus n. sp. 

Plate 17, figures 1, 2 

Lithostrotionella girtyi Hayasaka, 1936, p. 65 [part]. 

Material studied.- USNM 162004 (holotype). 

Descript1:on of holotype. -The specimen is a fragment 

of a small, cerioid, hemispherical corallum. It was more 

than 4.5 em high and more than 8 em in diameter. The 

outer part of the specimen is silicified. 

Corallites are polygonal, have straight, doublelayered, beaded walls 0.25-0.5 mm thick, and range from 

8 to 11 mm in diameter at maturity. Increase is 

peripheral. The major septa most commonly extend to 

the columella and may or may not extend into the 

dissepimentarium. The cardinal septum is elongated in 

some corallites. There are 17 to 19 major septa at 

maturity. Minor septa are poorly developed as spines on 

the corallite walls. The columella is complex, composed 

of a medial plate, septal lamellae, and conical axial 

tabellae and is as much as 2 mm in diameter; it is ordinarily thickened by stereoplasm. Tabulae are ordinarily complete, conical, uncurved, and spaced about 0. 7 to 

0.8 mm apart. The dissepimentarium is weakly 

lonsdaleoid and consists of 1 or 2 rows of inflated 

dissepiments of variable size. There are ordinarily 2 or 3 

dissepiments in 1 mm. The ratio of the tabularium width 

to the corallite diameter is about 0.6 to 0.7. Septal 

microstrueture is obscured by recrystallization, seemingly diffusotrabecular. 
20 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Discussion. -This species is most similar to 

Acrocyathus floriformis hemisphaericus by virtue of its 

complex columella but is distinguished by having fewer 

tabellar rings and by the stereoplasmic thickening of the 

columella. 

Occurrence. -Lower Carboniferous, middle Visean. 

St. Louis Limestone, Illinois, U.S.A. 

Acrocyathus proliferus (Hall in Hall and Whitney) 

Plate 15; plate 16, figures 1, 3, 4 

Lithostrotion mamillare Milne-Edwards and Haime, 1851, p. 433, pl. 

13, figs. la, b. [part]; Rominger, 1876, p. 111, pl. 55 [part]. 

Lithostrotion canadense Milne-Edwards and Haime, 1851, p. 483, pl. 

13, figs. la, b. [part]. 

Lithostrotion canadense Castelnau. Weller, 1898, p. 329 [part]. 

Lithostrotion prolijerum Hall in Hall and Whitney, 1858, p. 668, pl.24, 

figs. 6a-c; Rominger, 1876, p. 111, pl. 55; Butts, 1917, p. 45, pl. 10, 

figs. 15-17; 1926, p. 176, pl. 58, fig. 14; Morse, 1930, p. 102, pl. 8; 

Weller, 1931, p. 276, pl. 37, figs. la, b; Kelly, 1942, p. 361, pl. 51, 

figs. 1, 4; Allen and Lester, 1954, p. 101, pl. 26, fig. 2; not Davis, 

1956, p. 31, pl. 2, figs. 3a, b. 

Lithostrotion? prolijerum Hall. Ulrich, 1905, p. 32, pl. 3, figs. 3-7. 

Lithostrotionella prolijera (Hall). Butts, 1941, p. 239, pl. 129, figs. 1, 

2; Weller and others, 1952, p. 84, pl. 1, fig. 5. 

Lithostrotion [Lithostrotionella] prolijerum (Hall). Bassler, 1950, p. 

217. 

Material studied.- USNM 841 (neotype), 4587, 37469, 

37470, 37471, 39655, 42667, 42705, 42714, 42845, 

49941, 52681, 60306, 71647, 135085, 135087, 135091, 

135095, 135098, 135168, 135176, 216201, 216207, 

216209, 216213, 239233. 

Diagnosis. -Fasciculate Acrocyathus with corallite 

diameter 10 to 25 mm and 26 to 49 major septa. Internal 

morphology like that of Acrocyathus floriformis, but 

dissepimentarium is consistently weakly lonsdaleoid. Increase is peripheral. 

Description of neotype. -I have been unable to locate 

the specimens originally described and illustrated by 

Hall (in Hall and Whitney, 1858). I have therefore 

selected USNM 841, from the St. Louis Limestone, 

Hardin County, Ill., as neotype for the species. 

The neotype is a fragment of a large phaceloid corallum. The specimen is about 12 mm high and 17 x 9 mm 

in diameter. 

Corallites are cylindrical, have single-layered walls 

about 0.2 mm thick, and range from 12 to 15 mm in 

diameter at maturity. Increase is peripheral. The major 

septa do not extend to the columella but do extend into 

the dissepimentarium, most commonly reaching the corallite wall. There are 26 to 29 major septa at maturity. 

The cardinal septum is slightly shorter than the other 

major septa and is in a fossula formed by downwarped 

tabulae. Minor septa are poorly developed as spines on 

the corallite walls. The columella ranges from a simple 

axial plate to a structure made up of axial plate, a few 

axial lamellae, and a few concentric traces of axial 

tabellae in transverse section. Tabulae are ordinarily 

complete, conical, curved upward at the columella, and 

spaced about 1 mm apart. The dissepimentarium is 

weakly lonsdaleoid and consists of 1 to 3 rows of inflated 

dissepiments of variable size. There is ordinarily 1 

dissepiment in 2 mm. The ratio of the tabularium width 

to the corallite diameter is about 0.6 to 0.8. Septal 

microstructure is diffusotrabecular. 

Discussion. -Colonies of the fasciculate A. JJroliferus 

show the same range in variation of internal structures, 

such as the columella and tabulae, as the cerioid A. 

florijormis. Both forms have a shortened cardinal septum and commonly a long counter septum. The only consistent internal difference is in the dissepimentarium, 

where the fasciculate forms have fewer rows of 

dissepiments, and the dissepimentarium is only weakly 

lonsdaleoid. Increase in both the fasciculate and cerioid 

forms is peripheral. In the fasciculate forms, the offsets 

tend to arise in clusters of several new individuals at 

levels of rejuvenation of the parent corallite. 

Form variation was studied in 263 USNM specimens 

from the St. Louis Limestone and equivalent beds in 

Alabama, Indiana, Illinois, Iowa, Kentucky, :Missouri, 

Tennessee, Virginia, West Virginia, Georgia, and 

Michigan. Of these specimens, 154 are cerioid A. 

floriformis and 95 are fasciculate A. proliferus. Both 

forms occur together at two localities. Ordinarily cerioid 

and fasciculate forms are discrete colonies:, but 14 

specimens show both growth forms in the same colony 

(pl. 16). In most of these, cerioid corallites gave rise to 

fasciculate corallites, but in one colony, a fasciculate 

form became cerioid at a later stage. 

The form variation noted above presents problems in 

taxonomic treatment of these corals. Some authors (for 

example, Hill, 1940, p. 151, 166) place cerioid and 

fasciculate forms of Lithostrotion and Lonsdaleia in the 

same genus. Others (for example, Kato, 1966, p. 

100-101; Sando, 1975, p. C20) regard the two growth 

forms as subgenera. The evidence with respect to form 

variation in "Lithostrotionella" floriforrnis and 

"Lithostrotion" proliferum suggests a strong genetic 

relationship between the two forms. Indeed, the basic 

similarities in internal structure and the presence of 

transitional forms might lead to the conclusion that the 

two forms are merely ecologic variants of the same 

biologic species. However, the rarity of transitional 

forms, the lack of detailed information on geographic 

and stratigraphic distribution of the morphotypes, and 

·the obvious practicality of recognizing the two growth 

forms as separate entities lead me to maintain them as 

separate species. The treatment is different from my 

treatment of Lithostrotion and Lonsdaleia, because in 

North America, Lithostrotion is represented almost entirely by the fasciculate- form (Siphonodendron), and 

fasciculate and cerioid Lonsdaleia are separated from 
SYSTEMATIC PALEONTOLOGY 21 

each other in different faunal provinces. Furthermore, 

no transitional forms are known between the subgenera 

of either Lithostrotion or Lonsdaleia in North America. 

Occurrence. -Lower Carboniferous, middle or upper 

Visean. St. Louis Limestone and equivalent strata, Illinois, Georgia, Kentucky, Tennessee, Alabama, and 

Michigan, U.S.A. 

Acrocyathus girtyi (Hayasaka) 

Plate 17, figures 3, 4 

Lithostrotionella girtyi Hayasaka, 1936, p. 65, pl. 13, figs. 3a, b. 

?Lithost'rotionella ~P· Parks, 1951, p. 180, pl. 29, figs. 2a, b. 

Material studied.- USNM 120243 (holotype). 

Description of holotype. -The specimen is a fragment 

of a large hemispherical corallum. It was more than 15 

em in diameter and attained a height of more than 8 em. 

The outer part of the specimen is silicified. 

Corallites are polygonal, have straight double-layered 

walls as much as 0.3 mm thick, and range from 8 to 11 

mm in diameter at maturity. Increase is peripheral. Except for the counter septum, the major septa are short 

and do not extend to the columella and seldom extend into the dissepimentarium. There are 18 to 21 major septa 

at maturity. Minor septa are absent or poorly developed. 

The columella is a simple axial plate connected to the 

counter septum in young corallites but is composed of an 

axial plate, septal lamellae, and upturned edges of 

tabulae in mature corallites. The columella is ordinarily 

thickened by stereoplasm and attains a maximum 

?iameter of 2 mm. Tabulae are ordinarily complete, conIcal, turned up at the columella, and spaced 0.5 to 1 mm 

apart. The dissepimentarium is strongly lonsdaleoid and 

consists of a single row of large inflated dissepiments. 

There are ordinarily 1 or 2 dissepiments in 2 mm. The 

ratio of the tabularium width to the corallite diameter is 

about 0.6 to 0.7. Septal microstructure is fibronormal. 

Discussion. -This species has a columella and corallite 

diameter like that of A. pilatus but is distinguished from 

the latter by its strongly lonsdaleoid dissepimentarium, 

single row of dissepiments, and septa that seldom extend to the columella. The specimen illustrated by Parks 

(1951) from the "Brazer Limestone" in Utah may belong 

here, but the lack of information on morphological 

details prevents certain identification. 

Occurrence. -Lower Carboniferous, middle or upper 

Visean. Little Flat Formation, Utah, U.S.A. 

Acrocyathus pennsylvanicus (Shimer) 

Lithostrotion pennsylvanicum Shimer, 1926, p. 27, pl. 5, figs. 3-5 

[part]. 

Lithostrotionella pennsylvanica (Sh.imer). Kelly, 1942, p. 352, pl. 50, 

figs. 1, 2, 5, 6, 8; Bamber, 1961, p. 145, pl. 11, figs. lb, c, 3a-f; 

1966, p. 19, pl. 4, figs. la-c, 2a-c. 

Lithostrotion [Lithostrotionella] pennsylvanicum (Shimer). Bassler, 

1950, p. 221. 

Lonsdaleia pennsylvanica (Shimer). Crickmay, 1955, 1961, p. 13, pl. 1, 

figs. 11, 12. 

Lithostrotionella pennsylvanicum (Shimer). Nelson, 1960, p. 117, pl. 

22, figs. 4-6; 1961, pl. 17, figs. 3, 4. 

Diagnosis. - Cerioid Acrocyathus with c_orallite 

diameter 10 to 17 mm and 18 to 23 major septa that ordinarily approach the columella but do not reach it and 

seldom extend into the dissepimentarium. Minor septa 

weakly developed. Columella complex, composed of an 

axial plate, septal lamellae, and axial tabellae or upturned edges of tabulae. Dissepimentarium composed of 

1 to 3 rows of large and small inflated dissepiments. 

Tabulae complete, conical, sharply deflected upward at 

columella, without shoulders or with poorly defined 

shoulders, and spaced about 0.5 mm apart. Ratio of 

tabularium width to corallite diameter 0.5 to 0.8. Mode 

of increase unknown. 

Description of lectotype.- See Nelson (1960) and 

Bamber (1966). 

Discu,ssion. -This species falls within the range of 

Acrocyathus floriformis on corallite diameter and 

number of major septa. A. pennsylvanicus is distinguished from A. floriformis by lacking differentiation of 

the cardinal and counter septa, by having fewer rows of 

dissepiments, and by its larger ratio of tabularium width 

to corallite diameter. 

.o~cu1-rence.- Lower Carboniferous, middle or upper 

VIsean. Mount Head Formation, Alberta, Canada; 

Prophet Formation, British Columbia, Canada. 

Acrocyathus utkae (Degtyarev) 

Eolithostrotionella utkae Degtyarev, 1973, p. 192, pl. 1, figs. la, b. 

Diagnosis.- Cerioid Acrocyathus with corallite 

diameter 1 to 15 mm and 22 to 25 major septa that approach the columella but ordinarily do not reach it and 

seldom extend into the dissepimentarium. Minor septa 

absent. Columella composed of a thickened axial plate 

commonly joined by counter and cardinal septa, septal 

lamellae, and concentric traces of upturned tabulae in 

transverse section. Dissepimentarium composed of 1 to 

3 rows of large inflated dissepiments. Tabulae complete 

conical to nearly horizontal, and spaced about 0.5 m~ apart. Ratio of tabularium width to corallite diameter 

about 0.6. Mode of increase unknown. 

D~scrip~ion of h?lotype.- See Degtyarev 1973). 

D~~c~':'s~~n. - T?Is species is similar to A.? grechovkae 

but differs m haVIng larger corallites, more major septa, 

a more complex columella, and conical tabulae. 

.o~cu~rrence. -Lower Carboniferous, lower or middle 

VIsean. Zapadnouralsk Horizon, Ural Mountains, 

U.S.S.R. 

Acrocyathus rotai (Zhizhina in Bul'vanker and others) 

Eolithostrotionella rotai Zhizhina in Bul'vanker and others 1960 

251, pl. 61, figs. 2a, b. ' ' p. 
22 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Diagnosis.- Cerioid Acrocyathus with corallite 

diameter 15 to 20 mm and 22 to 26 major septa that 

commonly reach the columella but seldom extend into 

the dissepimentarium. Minor septa absent or poorly 

developed. Columella seemingly simple to complex. 

Dissepimentarium composed of 1 to 3 rows of mostly 

large inflated dissepiments. Tabulae complete, conical, 

and spaced about 0.5 mm apart. Ratio of tabularium 

width to corallite diameter about 0.6. Mode of increase 

unknown. 

Description of holotype. -See Zhizhina (in Bul'vanker 

and others, 1960). 

Discussion. -At first glance, the transverse section of 

this species suggests Stelechophyllum, but the tabulae 

are clearly of the Acrocyathus type, and the axial structures in some corallites appear to be complex. 

Occurrence. -Lower Carboniferous, middle or upper 

Visean. Zone C1vf, Donetz Basin, U.S.S.R. 

Acrocyathus cystosus (Zhizhina in Bul'vanker and others) 

Eolithostrotionella cystosa Zhizhina in Bul'vanker and others, 1960, p. 

250, pl. 61, figs. la, b. 

Diagnosis.- Cerioid Acrocyathus with corallite 

diameter 20 to 30 mm and 28 to 38 major septa that may 

or may not extend to the columella and seldom extend 

into the dissepimentarium. Minor septa poorly 

developed. Columella seemingly simple to complex. 

Dissepimentarium composed of as many as 5 rows of 

dissepiments of varying sizes. Tabulae complete, conical, and spaced about 0.5 mm apart. Ratio oftabularium 

width to corallite diameter about 0.4 to 0.5. Mode of increase unknown. 

Description of holotype. - See Zhizhina (in Bul'vanker 

and others, 1960). 

Discussion. -This species is distinguished by its broad 

dissepimentarium. 

Occurrence.- Lower Carboniferous, lower to upper. 

Visean. Zones C1vb to C1vf, Donetz Basin, U.S.S.R. 

Acrocyathus lissitzini (Zhizhina in Bul'vanker and 

others) 

Eolithostrotionella lissitzini Zhizhina in Bul'vanker and others, 1960, 

p. 252, pl. 61, figs. 3a, b. 

Diagnosis.- Cerioid Acrocyathus with corallite 

diameter 15 to 20 mm and 18 to 25 major septa that approach the columella but seldom reach it and seldom extend into the dissepimentarium. Minor septa absent. 

Columella seemingly simple to complex. Dissepimentarium composed of 1 to 3 rows of dissepiments of varying sizes. Tabulae complete, conical, and spaced about 1 

mm apart. Ratio of tabularium width to corallite 

diameter about 0.4 to 0.5. Mode of increase unknown. 

Description of:holotype. -See Zhizhina (in Bul'vanker 

and others, 1960),. 

Occurrence. -Lower Carboniferous, middle or upper 

Visean. Zone C1vf, Donetz Basin, U.S.S.R. 

Acrocyathus hsujiulingi (Yoh) 

Lithostrotionella hsujiulingi Yoh, 1961, p. 8, 16, pl. 13, figs. la-c. 

Diagnosis.- Cerioid Acrocyathus with corallite 

diameter averaging 6 mm and about 16 major septa that 

approach the columella but seldom reach it and do not 

extend into the dissepimentarium. Minor septa well 

developed. Columella composed of a thickened axial 

plate joined to the counter septum, septal lamellae, and 

concentric traces of tabulae in transverse section. 

Dissepimentarium composed of a single row of dissepiments of varying sizes. Tabulae complete, conical, 

and spaced about 0.5 mm apart. Ratio of tabularium 

width to corallite diameter about 0.5. Mode of increase 

seemingly peripheral. 

Description of holotype.- See Yoh (1961). 

Discussion. -This species differs from the other 

Chinese species, A.? unicus, in having fewer major septa, fewer rows of dissepiments, and a somewhat more 

complex columella. 

Occurrence. -Lower Carboniferous(?), Kwangsi Province, China. 

Acrocyathus? unicus (Y abe and Hayasaka) 

Lithostrotion (Lithostrotionella) unicum Yabe and Hayasaka, 1915, p. 

133; 1920, p. 11, pl. 9, figs. 12a, b. 

Lithostrotionella unica Yabe and Hayasaka. Wu in Yii and others, 

1963, p. 86, pl. 24, figs. 7a, b. 

Lithostrotionella unicum Yabe and Hayasaka. Minato and Kato, 197 4, 

p. 72, pl. 15, fig. 1. 

Diagnosis. - Cerioid Acrocyathus? with corallite 

diameter averaging 7 mm and about 22 to 26 major septa that approach the columella but seldom reaeh it and 

seldom extend into the dissepimentarium. Minor septa 

well developed. Columella seemingly a simple axial plate 

augmented by a few axial tabellae and septal ]ameBae. 

Dissepimentarium composed of 1 to 3 rows of large inflated dissepiments. Tabulae complete, conical, and 

spaced 6 to 7 in 2 mm. Ratio of tabularium width to corallite diameter about 0.5. Mode of increase unknown. 

Description of holotype. -See Y abe and Hayasaka 

(1915, 1920) and Minato and Kato (1974). 

Discussion. -This species, the type of Lithostrotionella, is referred to Acrocyathus with query because 

of uncertainty concerning the morphology of the type 

specimen (see previous discussion on page 4). 

Occurrence.- Carboniferous(?), Yun-nan Province, 

China. 

Acrocyathus? shimeri (Crickmay) 

Lithostrotion pennsylvanicum Shimer, 1926, p. 27 [part]. 

Lonsdaleia shimeri Crickmay, 1955, 1961, p. 13, pl. 1, figs. 9, 10. 
SYSTEMATIC PALEONTOLOGY 23 

Lithostrotionella shimeri (Crickmay). Nelson, 1960, p. 114, pl. 21, figs . 

. 9-15, pl. 22, figs. 1-3; 1961, pl. 14, figs. 3-5, pl. 16, figs. 8, 9, pl. 17, 

fig. 5, pl. 18, figs. 4, 5; Bamber, 1961, p. 137, pl. 10, figs. 4a-d, pl. 

11, figs. 1a, d-j, 2a-d; Armstrong, 1962, p. 39, pl. 3, figs. 13-15, 

text-fig. 19. 

Lithostrotionella pennsylvanica (Shimer). Armstrong, 1970a, p. 31, pl. 

9, figs. 1-3. 

Diagnosis. - Cerioid Acrocyathus? with corallite 

diameter 7 to 13 mm and 17 to 30 major septa that commonly extend across the tabularium to the columella and 

seldom extend into the tabularium. Minor septa ordinarily well developed. Columella complex, composed 

of an axial plate, septal lamellae, and the upturned 

edges of tabulae. Dissepimentarium composed of a 

single row of inflated dissepiments. Tabulae complete, 

essentially horizontal, but deflected upward at columella, and spaced about 0. 75 mm apart. Ratio of 

tabularium width to corallite diameter about 0.4 to 0.5. 

Increase is peripheral. 

Description of holotype.- See Nelson (1960). 

Discussion.-This species is distinguished by its nearly 

horizontal tabulae, which approach the morphology of 

Petalaxis. It differs from the species of Petalaxis by its 

complex columella. 

Occurrence. -Lower Carboniferous, middle and upper 

Visean. Mount Head Formation, Alberta and British 

Columbia, Canada; Peratrovich Formation, Alaska, 

U.S.A.; Black Prince Limestone, Arizona, U.S.A. 

Acrocyathus? grechovkae (Degtyarev) 

Eolithostrotionella grechovkae Degtyarev, 1973, p. 193, pl. 1, fig. 2, pl. 

2, figs. 1a, b. 

Diagnosis.-Cerioid Acrocyathus? with corallite 

diameter 10 to 12 mm and 18 to 24 major septa that approach the columella but seldom reach it and seldom extend into the dissepimentarium. Minor septa absent. 

Columella ordinarily a simple thickened axial plate surrounded by intersections of upturned tabulae in 

transverse section and occasionally abutted by a few 

septal lamellae. Dissepimentarium composed of 2 to 3 

rows of large inflated dissepiments. Tabulae complete, 

nearly horizontal at the periphery, but deflected upward 

at the columella, and spaced about 0.5 mm apart. Ratio 

of tabularium width to corallite diameter about 0.6. 

Mode of increase unknown. 

Desc,ription of holotype.- See Degtyarev (1973). 

DisC'ussion. -This species is similar to A.? shimeri in 

having nearly horizontal tabulae but differs in having 

fewer major septa, no mino~ septa, more rows of 

dissepiments, and a less compl~x columella. 

Occurrence.- Lower Carboniferous, lower or middle 

Visean. Zapadnouralsk Horizon, Ural Mountains, 

U.S.S.R. 

Acrocyathus? zhizhinae (Vasilylik) 

Eolithostrotionella zhizhinae Vasilyuk, 1960, p. 95, pl. 25, figs. 1, 1a. 

Diagnosis.- Cerioid-fasciculate Acrocyathus? with 

corallite diameter 13 to 20 mm and 25 to 27 major septa 

that extend to the columella but do not extend into the 

dissepimentarium. Minor septa well developed, contratingent. Columella seemingly simple to complex, absent in some corallites. Dissipimentarium composed of 1 

to 4 rows of dissepiments of varying sizes. Tabulae complete, flat to conical, and spaced 0. 75 to 1 mm apart. 

Ratio of tabularium width to corallite diameter about 

0.5. Mode of increase unknown. 

Description of type material.- See Vasilyuk (1960). 

Discussion.-The original description and illustrations 

of this species leave some doubt concerning the structure of the columella and the tabulae. The species_ is 

therefore referred to Acrocyathus with a query. The 

long contratingent minor septa are noteworthy. 

Occurrence. -Lower Carboniferous, lower to middle 

Visean. Zones C1vc and C/d, Donetz Basin, U.S.S.R. 

Acrocyathus spp. indet. 

The following taxa are not adequately described and illustrated for specific identification: 

Lithostrotionella americana Hayasaka(?). Nelson, 1960, 

p. 118, pl. 23, fig. 3. Lower Carboniferous, Visean. 

Mount Head Formation, Alberta, Canada. 

Lithostrotionella shimeri (Crickmay). Nations, 1963, p. 

1257, pl. 176, figs. 1, 2. Base of Black Prince 

Limestone in pebble derived from Escabrosa 

Limestone, Arizona, U.S.A. 

.Family PETALAXIDAE Fomichev, 1953. 

Genus PETALAXIS Milne-Edwards and Haime, 1852 

Stylaxis Milne-Edwards and Haime, 1851, p. 452 [part] (not McCoy, 

1849, p. 119). 

Petalaxis Mil:ne-Edwards and Haime, 1852, p. 204 [part]; MilneEdwards, 1860, p. 440 [part]; not Barrois, 1882, p. 305; Roemer, 

1883, p. 387, 388 [part]; Lindstrom, 1883, p. 12; Stuckenberg, 

1888,. p. 20 [part]; 1895, p. 74 [part]; Yanishevskiy, 1900, p. 89[?]; 

Etheridge, 1900, p. 17; Grosch, 1909, p. 5; Yabe and Hayasaka, 

1915, p. 9•1 (32); Bolkhovitinova, 1915, p. 63[?]; Gabounia, 1919, p. 

39[?]; not Chi, 1931, p. 28; Dobrolyubova, 1935b, p. 10; Heritsch, 

1939, p. 18; not Hill, 1940, p. 165; not Lang, Smith, and Thomas, 

1940, p. 97; Kolosvary, 1951, p. 39[?]; Fomichev, 1953, p. 449; not 

Hill, 1956, p. F 282; Soshkina, Dobrolyubova, and Kabakovich, 

1962, p. 339; de Groot, 1963, p. 81; Onoprienko, 1970, p. 3; 

Fedorowski and Gorianov, 1973, p. 58; not Minato and Kato, 197 4, p. 

68; Kozyreva, 1974, p. 24 [part]; Sutherland, 1977, p. 185. 

Lithostrotion Fleming. Eichwald, 1861, p. 149; Trautschold, 1879, p. 

36 [part]; Bassler, .1950, p. 222 [part). 

Lithostrotion (Petalaxis) Fomichev, 1931, p. 43. 

Lithostrotion (Lithostrotionella) Merriam, 1942, p. 377 [part]; Bassler, 

1950, p. 220, 222, 235, 252; Easton, 1960, p. 578. 

Lithostrotionella Yabe and Hayasaka. Chi, 1931, p. 28; YO., 1933 

(1934], p. 102[?]; Dobrolyubova, 1935a, p. 10; 1935b, p. 10, 12; 

1936a, p. 28; Hayasaka, 1936, p. 65, 70 (part]; Heritsch, 1937, p. 

164; Fomichev, 1939, p. 60; Minato, 1955, p. 99[?]; Yokoyama, 

1957, p. 78; Nelson, 1960, p. 114 [part]; Yamagiwa, 1961, p. 102[?]; 

Bamber, 1961, p. 107 (part]; de Groot, 1963, p. 80 (part]; Wu in Yii 
24 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

and others, 1963, p. 86[?]; Minato and Kato, 1974, p. 68; 

Onoprienko, 1976, p. 29 [part]. 

Lithostrotionella (Hillia) de Groot, 1963, p. 86. 

Lonsdaleia McCoy. Dobrolyubova, 1935a, p. 12; 1935b, p. 29. 

Cystolonsdaleia Fomichev, 1953, p. 464 [part]. 

Eastonoides Wilson and Langenheim, 1962, p. 511. 

Type species. -Stylaxis M'coyana Milne-Edwards and 

Haime, 1851, p. 453, pl. 12, figs. 5, 5a, equals Petalaxis 

M'coyana Milne-Edwards and Haime, 1852, p. 205, 

equals Petalaxis maccoyanus Milne-Edwards and Haime 

(by subsequent designation of Roemer, 1883, p. 387, 

388). Middle Carboniferous (Moscovian), Moscow Basin, 

U.S.S.R. 

Diagnosis.-Cerioid colonial corals with tabular to 

hemispherical growth form. Septa of two orders·. Major 

septa thin, ordinarily extending into the tabularium but 

seldom reaching the columella and ordinarily discontinuous or absent in the dissepimentarium, but species 

with weakly lonsdaleoid dissepimentarium are known. 

Minor septa absent to well developed. Columella ordinarily a simple axial plate joined to counter and (or) 

cardinal septum but composed of axial plate, impersistent vertical axial tabellae, and a few axial lamellae in 

some species. Tabulae complete and incomplete, essentially horizontal, flat, concave, or convex, turned up at 

columella in some species. Peripheral clinotabellae may 

be present. Dissepimentarium ordinarily lonsdaleoid but 

may be mostly regular in some species. Increase 

peripheral and intermural. 

Discussion. -Milne-Edwards and Haime (1852, p. 204) 

proposed Petalaxis for corals previously described and 

named Stylaxis M'coyana and Stylaxis portlocki (Milne 

Edwards and Haime, 1851, p. 452) because Stylaxis had 

become a junior synonym of Lithostrotion, and these 

two species were distinct. P. M'coyana, described and illustrated in 1851, was distinguished by having septa 

that were interrupted by the dissepiments and did not 

reach the corallite wall. The type specimen was collected 

by V erneuil at Kolomna on the Oka River in the Moscow 

Basin, U.S.S.R., where it is now known that beds of 

Middle Carboniferous (Moscovian) age are exposed. 

Subsequently, Hill (1940, p. 167) determined that P. 

portlocki, from -the Lower Carboniferous of England, is 

aLithostrotion. Milne-Edwards (1860, p. 440) diagnosed 

the genus Petalaxis for the first time, basing the 

diagnosis on the previous description of P. M'coyana. 

Lang, Smith, and Thomas (1940, p. 97) concluded incorrectly that Petalaxis is a replacement name for 

Nematophyllum and that, therefore, Petalaxis is a junior 

synonym of Lithostrotion. Hill (1940, p. 167) also attempted to p~ace Petalaxis in the synonymy of Lithostrotion by selecting P. portlocki as type species. However, 

the foregoing nomenclatural actions were invalidated by 

Roemer's (1883, p. 387, 388) earlier designation of P. 

M'coyana as the type species of Petalaxis. 

Although the name Petalaxis was used by several 

earlier Russian authors (Stuckenberg, 1888, 1895; 

Yanishevskiy, 1900, Bolkhovitinova, 1915; Gabounia, 

1919, Fomichev, 1931), there was confusion over the 

meaning of the name for many years. In the meanwhile, 

Yabe and Hayasaka (1915) had proposed Lithostrotionella, and some authors (for example, Dobrolyubova, 

1935a, b, 1936a) used this name in preference to 

Petalaxis for corals with a simple columella and 

lonsdaleoid dissepiments, regardless of the nature of the 

tabulae. Fomichev (1953, p. 449-463) sought to clarify 

the usage of Petalaxis by regarding P. maccoyanus as 

the type species and describing some other Middle Carboniferous corals under the name of Petalaxis; he noted 

that forms with horizontal tabulae are characteristic of 

the Middle Carboniferous in contrast to the tent-shaped 

tabulae of Early Carboniferous corals. 

Kozyreva (197 4, p. 24-26) summarized the history of 

Petalaxis and stressed the importance of horizontal 

tabulae in the definition of the genus. She included in the 

genus some North American species that actually 

belong to Stelechophyllum, Acrocyathus, and 

Thysanophyllum. 

Sutherland (1977) reviewed the Petalaxis problem and 

pointed out that the type specimen has been lost. He 

based his discussion of the genus on a study of topotypes 

at the Paleontological Institute in Moscow. Sutherland 

called attention to the fact that the topotypes have a 

variable axial structure that is complex in some corallites. He also stressed the horizontal tabulae as a 

distinguishing feature of Petalaxis, which occurs 

predominantly in the Middle Carboniferous. 

Petalaxis is distinguished from Acrocyathus and 

Stelechophyllum chiefly by its horizontal tabulae and by 

its columella. In Acrocyathus, the tabulae are conical, 

without well-defined shoulders, and in Stelecho~ohyllum, 

the tabulae are tent-shaped, with well-defined shoulders 

and a peripheral zone of more or less horizontal tabellae. 

In Petalaxis, the columella is most commonly a simple, 

thickened or unthickened axial plate connected to the 

counter and (or) cardinal septum, but some speeies have 

impersistent vertical axial tabellae and rare septal 

lamellae. In Acrocyathus, the columella ranges from a 

simple axial plate to a complex spiderweb structure composed of axial plate, septal lamellae, and axial tabellae or 

upturned tabulae. In Stelechophyllum, the columella is 

always a simple axial plate or rod unmodified by axial 

tabellae or septal lamellae. 

Fomichev (1953, p. 464) proposed Cystolonsda.leia as a 

subgenus of Petalaxis for corals from the Middle and 

Upper Carboniferous of the Donetz Basin that have a 

columella composed of a medial plate, a few septal 

lamellae, and axial tabellae similar to the Early Carboniferous Lonsdaleia. The type speeies of 
SYSTEMATIC PALEONTOLOGY 25 

Cystolonsdaleia is C. lutugini Fomichev from the Moscovian of the Donetz Basin. Fomichev also included 

Lonsdaleia portlocki {Stuckenberg) of Dobrolyubova 

(1935a, b) and Lonsdaleia ivanovi Dobrolyubova (1935a, 

b) from the Moscovian of the Moscow Basin and indicated that species attributed to Stylidophyllum from 

the Pe:rmian of China by Yoh and Huang (1932) and 

Huang (1932) might belong to the new subgenus. The 

Permian species were later reassigned to genera of the 

Family W aagenophyllidae by Minato and Kato (1965). 

Petalaxis maccoyanus, the type species of Petalaxis, 

has an impersistently developed complex axial structure 

with the same morphologic elements as the axial structure of most of the Moscovian species assigned to 

Cystolonsdaleia by Fomichev. It seems impractical to 

separate these species generically from P. maccoyanus. 

The type species of Cystolonsdaleia, C. lutugini, differs 

from the other Moscovian species by having a complex 

spiderweb columella with inner zone of tabellae distinct 

from an outer zone of regular concave tabulae. Accordingly, Cystolonsdaleia is retained as a separate genus for 

this complex species, whereas the other species are placed in Petalaxis. 

De Groot (1964, p. 86) proposed Hillia as a subgenus 

of Lithostrotionella for corals from the Bashkirian of 

Spain that differ from Lithostrotion by having major 

septa that fall short of the columella, horizontal tabulae, 

and a columella connected to the cardinal septum; they 

differ from true Lithostrotionella by having a poorly 

developed lonsdaleoid dissepimentarium. 

In other corals that have a columella arising from a 

single septum, that septum is the counter septum 

wherever the septa can be identified. De Groot gave no 

evidence for her conclusion that the septum in Hillia is 

the cardinal septum, and her illustrations of species 

assigned to Hillia show no basis for that conclusion. The 

columella appears to be the same as that seen in many 

species of Petalaxis. Moreover, the name Hillia is preoccupied by Hillia Grote (1883) and Hillia Mallock (1929), 

according to Cotton (1974, p. 13), and a replacement 

name has not been proposed. In my opinion, the reasons 

for separating Hillia de Groot from Petalaxis are insufficient, and I am reassigning the species of Hillia to 

Petalaxis, recognizing them only as a species group 

within that genus. 

Eastonoides was proposed by Wilson and Langenheim 

(1962, p. 511) for a Permian coral that has a columella 

composed of an axial plate connected to the counter septum augmented by rare axial tabellae, slightly sloping 

tabulae, and a vertically discontinuous lonsdaleoid 

dissepimentarium. Lonsdaleia ivanovi Dobrolyubova 

was included as the only other known species referred to 

Eastonoides. The essential features of this genus are so 

similar to the type species of Petalaxis that I regard 

Eastonoides as a junior synonym of Petalaxis, which includes four other Permian species. 

Species of Petalaxis are distinguished on differences 

in mature corallite diameter, number of major septa at 

maturity, development of minor septa, complexity and 

development of the columella, major septal extensions 

into the tabularium, major septal extensions into the 

dissepimentarium, shape ·and completeness of the 

tabulae, spacing of the tabulae, number of rows of 

dissepiments, size and shape of the dissepiments, and 

the ratio of tabularium width to corallite diameter. 

The species of Petalaxis may be arranged in five 

species groups: 

1. P. simplex group, including simplex, wyomingensis, 

tabulatus, bailliei, and tschucoticus. This group 

comprises simple forms having mostly complete 

tabulae turned up at the columella and a simple 

columella; these forms occur in the Lower Carboniferous (Visean) and lowest Middle Carboniferous (Namurian). 

2. P. jlexuosus group, including flexuosus, donbassicus, 

mokmnokensis, exiguus, brokawi, monocyclicus, 

sexan~7ulus, taishakuensis, immanis, belinskiensis, 

major, fomichevi, and grootae. This group comprises forms having a simple columella and a narrow dissepimentarium (large ratio of tabularium 

width to corallite diameter); these forms occur in 

the Middle Carboniferous (Bashkirian and Moscovian) and Permian.· 

3. P. wagneri group, including wagneri, perapertuensis, radians, intermedius, santaemariae, cantabricus, orboensis, and occidentalis. This group 

comprises forms having a simple columella and a 

weakly lonsdaleoid dissepimentarium; these forms 

occur in the Middle Carboniferous (Bashkirian and 

Moscovian) and Permian. 

4. P. vesiculosus group, including vesiculosus, 

lisitschanskensis, exilis, confertus, persubtilis, 

korkhovae, mirus, and evidens. This group comprises forms having a simple columella and a narrow dissepimentarium (small ratio of tabularium 

width to corallite diameter); these forms occur in 

the Middle Carboniferous (Bashkirian and Moscovian). 

5. P. maccoyanus group, including maccoyanus, 

styla:xis, mohikanus, celadensis, elyensis, 

dobrolyubovae, donetsensis, and ivanovi. This 

group comprises forms having a complex columella; these forms occur in the Middle Carboniferous (Moscovian) and Permian. 

Occurrence.- Lower Carboniferous (Visean), Middle 

Carboniferous (Namurian, Bashkirian, and Moscovian), 

and Permian. U.S.S.R., U.S.A., Canada, Spain, Japan, 

China(?), Spitzbergen(?). 
26 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Petala.xis simplex species group 

Petala.xis simplex {Hayasaka) 

Plate 18, figures 1-3 

Lithostrotionella simplex Hayasaka, 1936, p. 70, pl. 14, figs. 4a, b; 

Bamber, 1961, p. 155, pl. 12, figs. 2a-c. 

Lithostrotion [Lithostrotionella] simplex (Hayasaka). Bassler, 1950, 

p. 220. 

Material studied.- USNM 120249 (holotype). 

Description of holotype. -The specimen is a fragment 

of a corallum of indeterminate shape in limestone 

matrix. It was more than 9 em in diameter and more 

than 4 em in height. 

Corallites are polygonal, have straight, slightly 

beaded, double-layered walls as much as 0.3 mm thick, 

and range from 6 to 9 mm in diameter at maturity. Except for the counter septum, the major septa are short, 

do not extend to the columella, and seldom extend into 

the dissepimentarium. There are 16 to 18 major septa at 

maturity. Minor septa are absent or poorly developed. 

The columella is a simple thin axial plate that may or 

may not be connected to the counter septum. Tabulae 

are ordinarily complete and essentially horizontal; some 

are turned up at the columella to produce concentric 

traces in the transverse sections of some corallites. 

Tabulae are irregularly spaced 0.5 to 1 mm apart. The 

dissepimentarium is strongly lonsdaleoid and consists of 

a single row of large inflated dissepiments. There are ordinarily about 2 dissepiments in 2 mm. The ratio of the 

tabularium width to the corallite diameter is about 0.6 to 

0. 7. The mode of increase is indeterminate. "Septal 

microstructure is fibronormal. 

Discussion. -Comparison with P. wyomingensis is 

made under the discussion of that species. 

Occurrence.- Lower Carboniferous, upper Visean. 

Little Flat Formation, Utah, U.S.A. 

Petalaxis wyomingensis, n. sp. 

Plate 18, figures 4, 5 

Lithostrotionella simplex Hayasaka, 1936, p. 70 [part]. 

Material studied.- USNM 120675 (holotype). 

Description of holotype. -The specimen is a fragment 

of a corallum of indeterminate shape in limestone 

matrix. It was more than 7 em in diameter and more 

than 4 em in height. 

Corallites are polygonal, have sinuous, beaded, 

double-layered walls as much as 0.6 mm thick, and range 

from 7 to 9 mm in diameter at maturity. The major septa 

ordinarily extend to the columella but seldom extend into the dissepimentarium. There are 17 to 18 major septa 

at maturity. Minor septa are absent. The columella is a 

simple thickened axial plate ordinarily connected to the 

counter septum. It is as much as 0.7 mm in short 

diameter and 1 mm in long diameter. Tabulae are complete or incomplete, essentially horizontal, turned up at 

the columella, and irregularly spaced 0.25 to 0.5 mm 

apart. The dissepimentarium is strongly lonsdaleoid and 

consists of 1 to 3 rows of inflated dissepiments of varying sizes. Ordinarily, there are 1 to 3 dissepiments in 2 

mm. The ratio of tabularium width to the corallite 

diameter is about 0.5 to 0.6. The mode of increase is indeterminate. Septal microstructure is fibronormal. 

Discussion.-The holotype of this species was a 

paratype of Lithostrotionella simplex Hayasaka. P. 

wyomingensis is similar to P. simplex (Hayasaka) but 

differs in having longer major septa, a thickened columella, thicker corallite walls, more rows of 

dissepiments, a smaller ratio of tabularium width to corallite diameter, and more tabulae. 

Occurrence.- Middle Carboniferous, Namurian. 

"Wells" Formation, Wyoming, U.S.A. 

Petalaxis tabulatus (Hayasaka) 

Plate 19, figures 5-7 

Lithostrotionella tabulata Hayasaka, 1936, p. 69, pl. 17, fig. 2. 

Lithostrotion [Lithostrotionella] tabulatum (Hayasaka). Bassler, 1950, 

p. 220. 

Material studied.- USNM 120246 (holotype), USNM 

216197 (paratypes). 

Description of holotype.-The specimen is a fragment 

of a corallum of indeterminate shape. It was more than 

12 em in diameter and 14 em in height. 

Corallites are polygonal, have straight, beaded, and 

denticulate double-layered walls as much as 0.4 ·mm 

thick, and range from 8 to 10 mm in diameter at maturity. The major septa ordinarily extend to the columella 

and into the dissepimentarium. There are 16 to 18 major 

septa at maturity. Minor septa are ordinarily well 

developed. The columella is a simple, serrated, thickened axial plate connected to one or more major septa. 

Tabulae are mostly complete, essentially horizontal, 

commonly turned up at the columella, and irregularly 

spaced 0.25 to 0.5 mm apart. The dissepimentarium is 

composed of 1 to 2 rows of inflated dissepiments of varying sizes. There are 2 to 4 dissepiments in 2 mm. The 

ratio of tabularium width to corallite diameter is about 

0.6 to 0. 7. The mode of increase is peripheraL Septal 

microstructure is obscure, seemingly amorphous. 

Discussion. -The transverse section of this species 

looks like that of a Stelechophyllum. It is placed in 

Petalaxis because of its horizontal tabulae. The species 

differs from P. wyomingensis in having a serrated, less 

thickened columella, to which more major septa are attached, thinner corallite walls, a larger ratio of 

tabularium width to corallite diameter, more extensions 

of the major septa into the tabularium, and welldeveloped minor septa. 

Occurrence.- Lower Carboniferous, upper Visean. 

Aspen Range Formation, Idaho, U.S.A. 
SYSTEMATIC PALEONTOLOGY 27 

Petalaxis bailliei (Nelson) 

Lithostrotionella bailliei Nelson, 1960, p. 114, pl. 21, figs. 7, 8; 1962, 

pl. 14, figs. 1, 2. 

Lithostrotionella cf. bailliei Nelson. Bamber, 1961, p. 126, pl. 10, figs. 

la-g. 

Diagnosis. -Petalaxis with corallites 5 to 6 mm in 

diameter and 17 to 18 major septa that ordinarily extend 

to the columella but seldom extend into the dissepimentarium. Minor septa ordinarily well developed. Columella a simple thickened axial rod or plate formed by 

extension of the counter septum. Dissepimentarium 

composed of a single row of large inflated dissepiments. 

Tabulae ordinarily complete, essentially horizontal, and 

irregularly spaced 3 in 1 mm. Ratio of tabularium width 

to corallite diameter about 0.5. Mode of increase 

unknown. 

Description of holotype.- See Nelson (1960). 

Discussion. -This species is distinguished from P. 

wyomingensis and P. tabulatus by its smaller corallite 

diameter. 

Occurrence.- Lower Carboniferous, Visean. Mount 

Head Formation, Alberta, Canada; and Prophet Formation, British Columbia, Canada (E. W. Bamber, written 

commun., 1980). 

Petalaxis tschucoticus (Onoprienko) 

Lithostrotionella tschucotica Onoprienko, 1976, p. 30, pl. 10, fig. 6, pl. 

11, figs. 1, 2. 

Diagnosis.-Petalaxis with corallite diameter 5 to 7 

mm and 15 to 17 major septa that ordinarily extend to 

the columella but seldom extend into the dissepimentarium. Minor septa poorly developed. Columella a simple axial plate connected to cardinal and counter septa. 

Dissepimentarium composed of a single row of large inflated dissepiments. Tabulae ordinarily complete, 

horizontal, commonly turned up at columella, regularly 

spaced 0.3 to 0.5 mm apart. Ratio of tabularium width to 

corallite diameter about 0.5 to 0.6. Mode of increase 

unknown. 

Description of type material.~ See Onoprienko (1976). 

Discussion.-This species differs from P. bailliei by 

having fewer major septa, a less robust columella, and 

weaker minor septa. 

Occurrence.- Lower Carboniferous, Namurian. 

Yunon Suite, West Chukotka, U.S.S.R. 

Petalaxis flexuosus species group 

Petalaxis flexuosus (Trautschold) 

LithostTotionjlexuosum Trautschold, 1879, p. 37, pl. 5, figs. 7a, b. 

Lithostrotionella jlexuosa (Trautschold). Dobrolyubova, 1935a, p. 11, 

pl. 3, figs. 1, 2; 1935b, p. 18, pl. 3, figs. 1, 2. 

?Lithostrotionellaflexuosa (Trautschold). Heritsch, 1939, p. 30, pl. 13, 

fig. 5, pl. 19, fig. 5. 

?Lithostrotion [Lithostrotionella] jlexuosum Trautschold. Bassler, 

1950, p. 235. 

Diagnosis.-Petalaxis with corallite diameter 5 to 7 

mm and 14 to 17 major septa that approach t~e columella but seldom reach it and commonly extend into 

the dissepimentarium. Minor septa well developed. Columella a simple thickened axial plate ordinarily connected to the counter septum and to the cardinal sep~m 

in some corallites. Dissepimentarium composed of 1 to 2 

rows of generally small inflated dissepiments of varying 

sizes. Tabulae complete and incomplete, horizontal to inclined, nat, concave or convex, irregularly spaced 0.25 

to 1 mm apart. Ratio of tabularium width to corallite 

diameter 0.6 to 0.7. Mode of increase unknown. 

Description of type material.- See Trautschold (1879). 

The diagnosis is taken largely from Dobrolyubova's 

(1935b) descriptions and illustrations of topotypes. 

Discussion. - Petalaxis flexuosus differs from P. 

stylaxis by lacking a complex columella and by having 

better development of the minor septa, more extensions 

of the major septa into the dissepimentarium, and 

generally smaller dissepiments. 

Fomiehev (1953, p. 453) and Kozyreva (197 4, p. 25) 

plac·ed P. flexuosus in the synonymy of P. mac_co¥an_us 

Milne-Edwards and Haime because of general similarity 

in morphology. I regard these , two species as distinct 

because P. flexuosus does not seem to have axial tabellae 

like P. maccoyanus. However, Sutherland (1977, p. 187) 

pointed out that the complexity of the axial structure is 

a variable feature in topotypes of P. maccoyanus; hence, 

Dobrolyubova's (1935b) description may be based on 

simple corallites in a variable colony. 

Heritsch's (1939) specimen is referred questionably to 

the species because no longitudinal section is available. 

Occurrence.- Middle Carboniferous, Moscovian. 

Myachkovo Horizon (C2 

4), Moscow Basin, U.S.S.R. 

Kings Bay, Spitzbergen(?). 

Petalaxis donbassicus (Fomichev) 

Lithostrotionella donbassica Fomichev, 1939, p. 60, pl. 9, figs. 4a, b. 

Lithostrotion [Lithostrotionella] donbassica (Fomichev). Bassler, 

1950, p. 222. 

Diagnosis. -Petalaxis with corallite diameter 5 to 7 

mm and 14 to 17 major septa that approach the columella but seldom reach it and commonly extend into 

the dissepimelitarium. Minor septa well developed. Columella a strongly thickened axial plate ordinarily connected to the counter septum. Dissepimentarium com-

. posed of 1 to 2 rows of inflated dissepiments of varying 

sizes. Tabulae ordinarily complete, horizontal, irregularly spaced 0.25 to 0.5 mm apart. Ratio of tabularium 

width to corallite diameter 0.5 to 0.6. Mode of increase 

unknown. 

Description of type material.- See Fomichev (1939). 

Discussion.-This species is very similar to Petalaxis 

flexuos1.tS, from which it differs in having better develop~ 
28 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

ment of the minor septa, a thicker columella, a wider 

dissepimentarium (smaller ratio of tabularium width to · 

corallite diameter}, and more complete tabulae. 

Occurrence.- Middle Carboniferous, Moscovian. Svita 

C2

6 (L5}, Donetz Basin, U.S.S.R. 

Petalaxis mokomokensis (Easton) 

Lithostrotion [Lithostrotionella] rrwkorrwkensis Easton, 1960, p. 578, 

text-figs. 9, lO. 

Diagnosis.-Petalaxis with corallite diameter about 7 

mm and 15 major septa that ordinarily are short and do 

not extend to the columella or into the dissepimentarium. Minor septa poorly developed. Columella a thin 

axial plate connected to the counter septum. 

Dissepimentarium composed of 1 to 2 rows of large inflated dissepiments. Tabulae complete and incomplete, 

essentially horizontal, mostly concave, irregularly 

spaced 0.25 to 0. 75 mm apart. Ratio of tabularium width 

to corallite diameter about 0.7. Mode of increase 

unknown. 

Description of type material.- See' Easton (1960). 

Discussion. -This species is distinguished from Middle 

Carboniferous species from the Soviet Union by its thinner columella and more generally concave tabulae. 

Occurrence.~ Permian. Arcturus Formation, Nevada, 

U.S.A. 

Petalaxis exiguus n. sp. 

Plate 19, figures 1-4 

Lithostrotionella girtyi Hayasaka, 1936, p. 65 [part]. 

Lithostrotion [Lithostrotionella] girtyi (Hayasaka). Bassler, 1950, p. 

220 [part]. 

Material studied.- USNM 162002B (holotype) and 

USNM 162002A (paratype). 

Description of holotype. -The specimen is a small complete silicified corallum 3.5 em in diameter and 2 em in 

height. 

Corallites are polygonal, have straight to rounded, 

beaded, sinuous, double-layered walls as much as 0.4 

mm thick, and range from 10 to 12 mm in diameter at 

maturity. Except for the counter and cardinal septa, the 

major septa extend about halfway to the columella and 

are present in the dissepimentarium only as crests. 

There are 14 to 15 major septa at maturity. Minor septa 

are absent. The columella is a simple thickened axial 

plate, commonly connected to both counter and cardinal 

septa. Tabulae are complete or incomplete,. essentially 

horizontal, concave to convex, and irregularly spaced 

0.5 to 1 mm apart. The dissepimentarium is strongly 

lonsdaleoid and consists of 1 to 2 rows of mostly large inflated dissepiments. The ratio of tabularium width to 

corallite diameter is about 0.6. Increase is peripheral. 

Septal microstructure is fibronormal. 

Discussion.-The holotype and para type of this new 

species were paratypes of Lithostrotionella girtyi 

Hayasaka. The two type specimens are so small that 

they may actually represent immature coralla, although 

no larger coralla were found at the type locality. The 

description of the holotype serves as a diagnosis for the 

species, because the paratype is essentially identical 

with the holotype. 

Petalaxis exiguus is distinguished from P. mokomokensis, another Permian form, by its diminutive corallum, 

thickened columella, larger corallite diameter, and 

smaller ratio of tabularium width to corallite diameter. 

Occurrence.- Permian. McCloud Limestone, California, U.S.A. 

Petalaxis brokawi (Wilson and Langenheim) 

Lithostrotionella sp. Langenheim and others, 1960, p. 151. 

Lithostrotionella brokawi Wilson and Langenheim, 1962, p. 512, pl. 

88, figs. 7, 8. 

Diagnosis. - Petalaxis with corallite diameter 4 to 5 

mm and 10 to 15 major septa that ordinarily are short 

and do not extend to the columella and seldom extend into the dissepimentarium. Minor septa ordinarily well 

developed. Columella a simple thickened axial plate ordinarily connected to the counter septum. Dissepimentarium composed of 1 to 2 rows of mostly large inflated 

dissepiments. Tabulae mostly complete, inclined slightly 

toward columella, and irregularly spaced 0.25 to 0.5 mm 

apart. Ratio of tabularium width to corallite diameter 

about 0;5. Mode of increase unknown. 

Description of type specimens.- See Wilson and 

Langenheim (1962). 

Discussion. - Petalaxis brokawi is distinguished from 

P. mokomokensis by its smaller corallites, thickened columella, ·well-developed minor septa, and smaller ratio of 

tabularium width to corallite diameter. 

Occurrence.- Lower Permian. Ely Limestone, 

Nevada, U.S.A. 

Petalaxis monocyclicus (de Groot) 

Lithostrotionella rrwnocyclica de Groot, 1963, p. 85, pl. 17, figs. la-c. 

Diagnosis. -Petalaxis with corallite diameter 5 to 8 

mm and 20 to 24 major septa that ordinarily approach 

the columella but do not reach it and seldom extend into 

the dissepimentarium. Minor septa absent. Columella a 

simple thin axial plate connected to the counter septum 

and to the cardinal septum in some corallites. Dissepimentarium composed of 1 to 3 rows of elongate 

dissepiments of varying sizes. Tabulae complete, essentially horizontal, concave, turned up at the columella, 

and irregularly spaced 0.25 to 1 mm apart. Ratio of 

tabularium width to corallite diameter about OA to 0.5. 

Mode of increase peripheral. 

Description of holotype.- See de Groot (19641[1963]). 

Discussion. -This species differs from the Middle Carboniferous and Permian species diagnosed on previous 

pages in this report by having tabulae that are regularly 
SYSTEMATIC PALEONTOLOGY 29 

concave and turned up at the columella. It also has more 

major septa. 

Occurrence. -Middle Carboniferous, Bashkirian. Santa Maria Limestone, Palencia, Spain. 

Petalaxis sexangulus (de Groot) 

Lithostrotionella sexangulus de Groot, 1963, p. 84, pl. 16, figs. 3a-c, 

4a, b. 

Diagnosis. -Petalaxis with corallite diameter 3 to 4 

mm and 13 to 15 major septa that ordinarily extend 

about halfway to the columella and seldom extend into 

the dissepimentarium. Minor septa poorly developed. 

Columella a simple slightly thickened axial plate connected to the counter septum and to the cardinal septum 

in some corallites. Dissepimentarium composed of a 

single :row of small dissepiments. Tabulae mostly complete, essentially horizontal, but commonly nearly tentshaped, turned up at the columella, and irregularly 

spaced 0.25 to 1 mm apart. Ratio of tabularium width to 

corallite diameter about 0.8. Mode of increase unknown. 

Description of type material.- See de Groot (1964 

[1963]). 

Discussion. -This species is distinguished from other 

members of the P. flexuosus species group by its very 

large ratio of tabularium width to corallite diameter and 

its tabulae that are commonly nearly tent-shaped and 

turned up at the columella. 

Occurrence. -Middle Carboniferous, Moscovian. 

Palencia, Spain. 

Petalaxis taishakuensis (Y okoyarila) 

Lithostrotionella taishalcuensis Yokoyama, 1957, p. 78, pl. 10, figs. 

1-4. 

Diagnosis.-Petalaxis with corallite diameter 3 to 4 

mm and 13 to 15 major septa that ordinarily are short 

and do not extend to the columella and seldom extend into the dissepimentarium. Minor septa well developed. 

Columella· a simple axial plate connected to the counter 

septum and joined by 2 or 3 other major septa. 

Dissepi.mentarium composed of 1 to 2 rows of inflated 

dissepiments of varying sizes. Tabulae complete and incomplete, essentially horizontal, and irregularly spaced 

0.25 to 1 mm apart. Ratio of tabularium width to corallite diameter about 0.6 to 0. 7. Mode of increase 

unknown. 

Description of holotype.- See Yokoyama (1957). 

Discussion. -This species is distinguished from P. sexangulus by having its columella joined by several major 

septa, by having stronger major septa, by its tabulae not 

being turned up at the columella, by its smaller ratio of 

tabularium width to corallite diameter, and by having 

more rows of dissepiments. 

Occurrence. -Middle Carboniferous. Dangyokei For· 

mation, Hiroshima,· Japan. 

Petalaxis immanis Kozyreva 

Petalaxis imm.anis Kozyreva, 1974, p. 30, pl. 2, figs. 4a-c, 5a, b. 

Diagnosis. -Petalaxis with corallite diameter 8 to 18 

mm and 17 to 18 major septa that ordinarily approach 

the columella but do not reach it and seldom extend into 

the dissepimentarium. Minor septa absent or poorly 

developed. Columella a simple, strongly thickened axial 

plate connected to counter septum. Dissepimentarium 

composed of 2 rows of small inflated dissepiments. 

Tabulae complete, essentially horizontal, turned up at 

columella, and irregularly spaced 0.25 to 0.5 mm apart. 

Ratio of tabularium width to corallite diameter about 0.5 

to 0.6. lVIode of increase unknown. 

Description of type material.- See Kozyreva (197 4). 

Discussion. -This species is distinguished from all 

other members of the P. flexuosus species group by its 

very large corallites and its strongly thickened columella. 

Occurrence.- Middle Carboniferous, Bashkirian. 

Horizon b1-b2 , Voronezh anteclise, U.S.S.R. 

Petalaxis belinslciensis Fomichev 

Petalaxis mccoyana var. belinskiensis Fomichev, 1953, p. 457, pl. 31, 

figs. 3a, b. 

Diagnosis.-Petalaxis with corallite diameter 7 to 12 

mm and 1H to 24 major septa that approach the columella but seldom reach it and commonly extend into 

the dissepimentarium. Minor septa well developed. Columella a simple thickened axial plate ordinarily connected to the counter septum. Dissepimentarium composed of 1 to 3 rows of inflated dissepiments of varying 

sizes. Tabulae mostly complete, essentially horizontal, 

and spaced 0.25·to 0.5 mm apart. Ratio of tabularium 

width to corallite diameter about 0.4 to 0.5. Mode of increase unknown. 

Descr·iption of type material.- See Fomichev (1953). 

DisCY.~sion.- This species is distinguished from P. 

donbassicus by having larger corallites, more major septa, and a broader dissepimentarium. It lacks the complex columella that characterizes P. mccoyana. 

Occurrence.- Middle Carboniferous, Moscovian. L5 

Limestone, Donetz Basin, U.S.S.R. 

Petalaxis major (de Groot) 

Lithostrotionella maccoyana forma major de Groot, 1963, p. 83, pl. 16, 

figs. 2a, b. 

Diagnosis.-Petalaxis with corallite diameter 5 to 9 

mm and 17 to 22 major septa that closely approach the 

columella but seldom reach it and that commonly extend 

into the dissepimentarium. Minor septa well developed. 

Columella a simple axial plate, seldom dilated, connected to the counter septum. A few septal lamellae 

present in some corallites but no axial tabellae. 

Dissepimentarium composed of 1 to 3 rows of inflated 
30 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

dissepiments of varying sizes. Tabulae mostly incomplete, horizontal, concave, and irregularly spaced 

0.25 to 0.5 mm apart. Ratio of tabularium width to corallite diameter about 0.6. Increase peripheral. 

Description of holotype.- See de Groot (1964 [1963]). 

The holotype is RGM 112726. 

Discussion.-This species differs from P. mccoyanus 

by lacking a complex columella and by its larger corallite 

diameter and more numerous major septa. It differs 

from P. vesiculosus by having longer major septa, more 

major septa, by the presence of septal lamellae, and by 

having more rows of dissepiments. 

Occurrence.- Middle Carboniferous, Moscovian. 

Vanes Formation, Palencia, Spain. 

Petalaxis fomichevi n. sp. 

Petalaxis maccoyana Milne-Edwards and Haime. Fomichev, 1953, p. 

453, pl. 31, figs. la-d, 2a-d. 

Diagnosis.-Petalaxis with corallite diameter 5 to 8 

mm and 13 to 22 major septa that extend about half way 

to the columella and commonly extend into the 

dissepimentarium. Minor septa well developed. Columella a simple thickened axial plate ordinarily connected to the counter septum and to the cardinal septum 

in some corallites. Dissepimentarium composed of 1 to 3 

rows of inflated dissepiments of varying sizes. Tabulae 

complete and incomplete, flat to convex, horizontal, and 

spaced 0.25 to 0.5 mm apart. Ratio of tabularium width 

to corallite diameter about 0.5 to 0.7. Mode of increase 

unknown. 

Description of type material.- See Fomichev (1953). 

The holotype is here designated specimen number 57 of 

Fomichev (1953, p. 456). 

Discussion.-This species differs from P. maccoyanus 

by lacking a complex columella and other morphological 

details. It is distinguished from P. flexuosus by having 

more major septa, more regular tabulae, and more rows 

of dissepiments. 

Occurrence.- Middle Carboniferous, Moscovian. L5 

Limestone, Donetz Basin, U.S.S.R. 

Petalaxis grootae, n. sp. 

Lithostrotionella maccoyana (Edwards and Haime). De Groot, 1963, p. 

82, pl. 16, fig. 1a, b. 

Diagnosis.-Petalaxis with corallite diameter 4 to 6 

mm and 14 to 16 major septa that approach the columella but seldom reach it and seldom extend into the 

dissepimentarium. Minor septa well developed. Columella a simple axial plate, curved in some corallites, 

and ordinarily connected to the counter septum. 

Dissepimentarium composed of 1 to 3 rows of inflated 

dissepiments of varying sizes. Tabulae complete and incomplete, essentially horizontal, flat, concave or convex, 

and spaced 0.25 to 0. 75 rrim apart. Ratio of tabularium 

width to corallite diameter about 0.6 to 0.7. Mode of increase unknown. 

Description of type material.- See de Groot (1964 

[1963], p. 83). The holotype is here designated· RGM 

112721. 

Discussion.-This species differs from P. maccoyanus 

by lacking a complex columella. It is distinguished from 

P. flexuosus by having longer major septa, a thinner, 

commonly curved columella, more rows of disse:piments, 

and more regularly spaced tabulae. 

Occurrence.- Middle Carboniferous, Moscovian. 

Vanes Formation and Cotarraso Limestone, Palencia, 

Spain. 

Petalaxis wagneri species group 

Petalaxis wagneri (de Groot) 

Lithostrotionella (Hillia) wagneri de Groot, 1963, p. 88, pl. 18, figs. 

1-3. 

Diagnosis. -Petalaxis with corallite diameter 3 to 4.5 

mm and 14 to 16 major septa that closely approach the 

columella but seldom · reach it and extend into the 

dissepimentarium, ordinarily being attached to the corallite wall. Minor septa ordinarily well developed. Columella a simple axial plate connected to counter or cardinal septum. Dissepimentarium weakly lonsdaleoid, 

composed of a single row of small inflated dissepiments. 

Tabulae complete or incomplete, essentially horizontal, 

mostly concave, and spaced 0.5 to 1 mm apart. Ratio of 

tabularium width to corallite diameter about 0. 7 to 0.8. 

Increase peripheral. 

Description of type material.- See de Groot (1964 

[1963]). 

Discussion.-This species is similar to P. maccoyanus 

but has smaller corallites, fewer major septa, a weakly 

lonsdaleoid dissepimentarium, and a simple columella. 

Occurrence.- Middle Carboniferous, Bashkirian. 

Perapertu Formation, Palencia, Spain. 

Petalaxis perapertuensis (de Groot) 

Lithostrotionella (Hillia) perapertuensis de Groot, 1963, p. 89, pl. 19, 

figs. la-c, 2a-d. 

Diagnosis. -Petalaxis with corallite diameter 3.5 to 

4.5 mm and 14 to 19 major septa that closely approach 

the columella but seldom reach it and ordinarily extend 

across the dissepimentarium to the corallite wall. Minor 

septa ordinarily well developed. Corallite walls strongly 

dilated. Columella a simple thickened axial plate connected to counter or cardinal septum. Dissepimentarium 

weakly lonsdaleoid, composed of 1 to 2 rows of small inflated dissepiments. Tabulae incomplete, essentially 

horizontal, mostly concave, and spaced 0.25 to 0.5 mm 

apart. Ratio of tabularium width to corallite diameter 

0.6 to 0.8. Increase peripheral. 

Description of type material.- See de Groot (1964 

[1963]). 
SYSTEMATIC PALE ONTOLOGY 31 

Discussion.-This species differs from P. wagneri by 

having more major septa, more tabulae, and strongly 

dilated corallite walls. 

Occurrence.- Middle Carboniferous, Bashkirian. 

PerapertU Formation, Palencia, Spain. 

Petalaxis radians (de Groot) 

Lithostrotionella (Hillia) radians de Groot, 1963, p. 89, pl. 20, figs. 

la-c. 

Diagnosis.-Petalaxis with corallite diameter 7 to 9 

mm and 15 to 18 major septa that closely approach the 

columella but seldom reach it and extend into the 

dissepimentarium, ordinarily being attached to the corallite wall. Minor septa ordinarily well developed. Columella a simple axial plate connected to the counter or 

cardinal septum. Dissepimentarium weakly lonsdaleoid, 

composed of 1 to 3 rows of inflated dissepiments of varying sizes. Tabulae incomplete, horizontal, mostly convex, and irregularly spaced 0.25 to 1 rrim apart. Ratio of 

tabularium width to corallite diameter about 0.5 to 0.6. 

Mode of increase unknown. 

Description of type material.- See de Groot (1964 

[1963]).. 

Discussion. -This species differs from P. wagneri by 

having larger corallites, more rows of dissepiments, and 

less regular tabulae. 

Occurrence.- Middle Carboniferous, Bashkirian. 

Perapertu Formation, Palencia, Spain. 

Petalaxis intermedius (de Groot) 

Lithostrotionella (Hillia) intermedia de Groot, 1963, p. 90, pl. 20, figs. 

2a, b, 3. 

Diagnosis. -Petalaxis with corallite diameter 5 to 7 

mm and 16 to 19 major septa that closely approach the 

columella but seldom reach it and extend into the 

dissepimentarium, commonly being attached to the corallite wall. Minor septa well developed. Columella a simple axial plate connected to counter or cardinal septum. 

Dissepimentarium weakly lonsdaleoid, CQmposed of 1 to 

4 rows of mostly small inflated dissepiments. Tabulae 

mostly incomplete, flat, concave or convex, horizontal, 

and irregularly spaced 0.25 to 0. 75 mm apart. Ratio of 

tabularium width to corallite diameter about 0. 7 to 0.8. 

Increase peripheral. 

Description of type material.- See de Groot (1964 

[1963]). 

Discussion. -This species differs from P. wagneri by 

having larger corallites, more major septa, and more 

tabulae. 

Occurrence.- Middle Carboniferous,_ Bashkirian. 

Perapertu Formation, Palencia, Spain. 

Petalaxis santaemariae (de Groot) 

Lithostrotionella (Hillia) santaemariae de Groot, 1963, p. 91, pl. 21, 

figs. la-e. 

Diagnosis.-Petalaxis with corallite diameter 3.5 to 8 

mm and 16 to 19 major septa that closely approach the 

columella but seldom reach it and extend into the 

dissepimentarium, ordinarily attached to the corallite 

wall. Minor septa poorly developed. Corallite walls 

strongly dilated. Columella a simple thickened axial 

plate connected to counter or cardinal septum. 

Dissepimentarium weakly lonsdaleoid, composed of a 

single row of small inflated dissepiments. Tabulae mostly complete, horizontal, mostly concave, and irregularly 

spaced 0.25 to 1 mm apart. A peripheral zone of 

clinotabellae may be present. Ratio of tabularium width 

to corallite diameter about 0.8. Increase peripheral. 

Description of holotype.- See de Groot (1964 [1963]). 

Discu,ssion.-This species differs from P. perapertuensis by having larger corallites, poorly developed minor 

septa, and fewer tabulae. 

OccU'rrence.- Middle Carboniferous, Bashkirian. Santa Maria Limestone, Palencia, Spain. 

.Petalaxis cantabricus (de Groot) 

Lithostrotionella (Hillia) cantabrica de Groot, 1963, p. 92, pl. 22, figs. 

1-4. 

Diagnosis. -Petalaxis with corallite diameter 4 to 8 

mm and 19 to 28 major septa that closely approach the 

columella but seldom reach it and extend into the 

dissepimentarium, ordinarily being attached to the corallite wall. Minor septa well developed. Cora!lite wall 

strongly dilated and denticulate. Columella a simple 

commonly thickened axial plate connected to counter or 

cardinal septum. Dissepimentarium weakly lonsdaleoid, 

composed of 1 to 3 rows of small inflated dissepiments. 

Tabulae mostly complete, horizontal, concave, and 

spaced 0.2!5 to 1 mm apart. A peripheral zone of 

clinotabellae may be present. Ratio of tabularium width 

to corallite diameter about 0.6 to 0.7. Increase 

peripheral. 

Description of type material.- See de Groot (1964 

[1963]). 

Discussion. -This species is distinguished from all 

others in the P. wagneri species group by having more 

major septa. 

Occurrence.- Middle Carboniferous, Bashkirian to 

Moscovian(!'). Santa Maria Limestone, Perapertu Formation, and Celada Limestone, Palencia, Spain. 

Petalaxis orboensis (de Groot) 

Lithostrotionella orboensis de Groot, 1963, p. 85, pl. 17, figs. 2a-d. 

Diagnosis. -Petalaxis with corallite diameter 4.5 to 8 

'mm and 16 to 21 major septa that closely approach the 

columella but seldom reach it and extend into the 

dissepimentarium, commonly being attached to the corallite wall. Minor septa well developed. Columella a simple axial plate connected to counter or cardinal septum; 
32 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

columella absent or discontinuous in some corallites. 

Dissepimentarium weakly lonsdaleoid, composed of 1 to 

3 rows ofinflated dissepiments of varying sizes. Tabulae 

complete and incomplete, generally horizontal but 

turned up at columella, and spaced 0.25 to 0.5 mm apart. 

Ratio of tabularium width to corallite diameter about 

0.7. Mode of increase unknown. 

Description of type material.- See de Groot (1964 

[1963]). 

Discussion. -This species is distinguished from others 

in the P. wagneri group by having tabulae that are 

turned up at the columella and by having the columella 

absent or discontinuous in some corallites. -

Occurrence. -Middle Carboniferous, Moscovian. Orbo 

Limestone, Palencia, Spain. 

Petalaxis occidentalis (Merriam) 

Plate 20, figures 1, 2 

Lithostrotion (Lithostrotione.lla) occidentalis Merriam, 1942, p. 377, 

pl. 56, figs. 2, 4, 7, 8, 11. 

Lithostrotion [Lithostrotionella] occidentale Merriam. Bassler, 1950, 

p. 252. 

Diagnosis.-Petalaxis with corallite diameter 4.5 to 

6.5 mm and 16 to 19 major septa that closely approach 

the columella but seldom reach it and extend into the 

dissepimentarium, commonly being attached to the corallite wall. Minor septa well developed. Corallite wall 

dilated and denticulate. Columella a simple thickened 

axial plate connected to the counter or cardinal septum. 

Dissepimentarium weakly lonsdaleoid, composed of 1 to 

2 rows of small inflated dissepiments Tabularium consists of a peripheral zone of clinotabellae and a periaxial 

zone of mostly complete, horizontal, flat or concave 

tabulae spaced 0.25 to 0. 75 mm apart. Ratio of 

tabularium width to corallite diameter about 0.6. Mode 

of increase unknown. 

Description of holotype.- See Merriam (1942). The 

holotype is USNM 143440; new thin sections of this 

specimen are illustrated herein for clarification of morphologic details. 

Discussion.-This species differs from P. perapertuensis by having larger corallites, a thicker columella, and 

well-developed clinotabellae. 

Occurrence.- Permian. Coyote Butte Formation, 

Oregon, U.S.A. 

Petalaxis vesiculosus species group 

Petalaxis vesiculosus (Dobrolyubova) 

Lithostrotionella vesiculosa Dobrolyubova, 1935a, p. 11, pl. 2, figs. 3, 

4; 1935b, p. 1.9, pl. 2, figs. 3, 4. 

Lithostrotion [Lithostrotionella] vesiculosa Dobrolyubova. Bassler, 

1950, p. 222. 

Diagnosis.-Petalaxis with corallite diameter 6 to 12 

mm and 10 to 14 major septa that may or may not extend to the columella and seldom extend into the 

dissepimentarium. Minor septa ordinarily well 

developed. Columella a simple thickened axial plate ordinarily connected to the counter and cardinal septa. 

Dissepimentarium composed of 2 to 6 rows of inflated 

dissepiments of varying sizes. Tabulae complete or incomplete, horizontal, concave or convex, and irregularly 

· spaced 0.25 to 1 mm apart. Ratio of tabularium. width to 

corallite diameter about 0.3 to 0.4. Mode of increase 

peripheral. 

Description of holotype.- See Dobrolyubova (1935b). 

Discussion. -This species is distinguished from all 

species of the P. flexuosus and P. wagneri groups by its 

broader dissepimentarium. 

Occurrence.- Middle Carboniferous, Moscovian. 

Myachkovo Horizon (C2

4), Moscow Basin, U.S.S.R. 

Petalaxis llsitschanskensis Fomichev 

Petalaxis vesiculosa (Dobrolyubova) var. lisitschanskensis Fomichev, 

1953, p. 462, pl. 32, figs. 3a, b. 

Diagnosis.-Petalaxis with corallite diameter 12 to 15 

mm and 16 to 21 major septa that approach the columella and may or may not reach it and do not extend into the dissepimentarium. Minor septa ordinarily well 

developed. Columella a simple thickened axial plate ordinarily connected to the counter septum. Dissepimentarium composed of 1 to 5 rows of inflated dissepiments 

of variable size. Tabulae complete or ineomplete, 

horizontal, concave or convex, and irregularly spaced 

0.25 to 0. 75 mm apart. Ratio of tabularium width to corallite diameter about 0.4. Mode of increase unknown. 

Description of holotype.- See Fomichev (1953). 

Discussion.-This species is distinguished from P. 

vesiculosus by having larger corallites, more major septa, and generally fewer rows of dissepiments. 

Occurrence.- Middle Carboniferous, Moscovian. L4 

and L6 Limestones, Donetz Basin, U.S.S.R. 

Petalaxis exilis Kozyreva 

Petalaxis exilis Kozyreva, 1974, p. 26, pl. 1, figs. la-d. 

Diagnosis. -Petalaxis with corallite diameter 5 to 11 

mm and 10 to 14 major septa that ordinarily are short 

and do not extend to the columella or into the dissepimentarium. Minor septa absent. Columella a simple 

thickened axial plate ordinarily connected to the counter 

septum and to the cardinal septum in some eorallites. 

Dissepimentarium composed of 1 to 3 rows of mostly 

large inflated dissepiments. Tabulae complete:, horizontal or slightly elevated toward columella, and irregularly 

spaced 0.5 to 1.25 mm apart. Ratio of tabularium width 

to corallite diameter about 0.4. Increase probably 

. peripheral. 

Description of holotype.- See Kozyreva (197·i). 

Discussion.-This species is distinguished from P. 

vesiculosus by having fewer septa that extend to the col-
SYSTEMATIC PALEONTOLOGY 33 

umella, by lacking minor septa, by having fewer rows of 

dissepiments, and by having fewer tabulae. 

Occurrence.- Middle Carboniferous, Bashkirian. 

Horizon b4, Voronezh anteclise, U.S.S.R. 

Petalaxis confertus Kozyreva 

Petalaxis confertus Kozyreva, 1974, p. 27, pl. 1, figs. 2a, b, 3a; b. 

Diagnosis.-Petalaxis with corallite diameter 8 to 12 

mm and 19 to 20 major septa that commonly approach 

the columella but do not extend into the dissepimentarium·. Minor septa absent. Columella a simple 

thickened axial plate connected to the counter septum. 

Dissepimentarium composed of 1 to 3 rows of inflated 

dissepiments of varying sizes. Tabulae mostly incomplete, varying in attitude, and irregularly spaced 

0.25 to 1 mm apart. Ratio of tabularium width to corallite diameter ·about 0.4 to 0.5. Mode of increase 

unknown. 

Desc·ription of type material.- See Kozyreva (197 4). 

Discussion.-This species is distinguished from P. exilis by having more major septa, longer major septa, and 

by its incomplete tabulae. 

Occurrence.- Middle Carboniferous, Bashkirian. 

Voronezh anteclise, U.S.S.R. 

Petalaxis persubtilis Kozyreva 

Petalaxis persubtilis Kozyreva, 1974, p. 27, pl. 1, figs. 4a-c, 5, 6. 

Diagnosis.-Petalaxis with corallite diameter 9 to 15 

mm and 15 to 17 major septa that commonly approach 

the columella and may or may not extend into the 

dissepimentarium. Minor septa absent or variably 

developed. Columella absent or weakly developed as a 

simple axial plate connected to the counter septum. 

Dissepimentarium composed of 2 to 4 rows of large inflated dissepiments. Tabulae complete, horizontal, commonly weakly turned up at columella, and irregularly 

spaced 0.25 to 1 mm apart. Ratio of tabularium width to 

corallite diameter 0.3 to 0.4. Increase peripheral. 

Description of type material.- See Kozyreva (197 4). 

Discussion.-This species is distinguished from P. confertus by having larger corallites, more major septa, 

more rows of dissepiments, and a smaller ratio of 

tabularium width to corallite diameter. 

Occurrence.- Middle Carboniferous, Bashkirian. 

Horizon b1, Voronezh anteclise, U.S.S.R. 

Petalaxis korkhovae Kozyreva 

Petalaxis korkhovae Kozyreva, 1974, p. 28, pl. 1, figs. 7a-c. 

Diagnosis. -Petalaxis with corallite diameter 9 to 15 

mm and 16 to 18 major septa that ordinarily extend 

about halfway to the columella but seldom extend into 

the d:issepimentarium. Minor septa absent or poorly 

developed. Columella a simple thickened axial plate connected to the counter septum. Dissepimentarium composed of 3 to 4 rows of inflated dissepiments of varying 

sizes. Tabulae complete, horizontal, mostly flat, and irregularly spaced 0.25 to 1 mm apart. Ratio of 

tabularium width to corallite diameter 0.3 to 0.4. Increase peripheral. 

Description of type material. -See Kozyreva (197 4). 

Discu.ssion.-This species is distinguished from P. persub til is by having a stronger columella, shorter major 

septa, and :flatter tabulae. 

Occu'rrence.- Middle Carboniferous, Bashkirian. 

Horizon b11 Voronezh anteclise, U.S.S.R. 

Petalaxis mirus Kozyreva 

Petalaxis mirus 1\,ozyreva, 1974, p. 29, pl. 2, figs. la-c. 

Diagnosis. - Petalaxis with corallite diameter 7 to 15. 

mm and H> to 18 major septa that ordinarily extend 

about halfway to the columella but seldom extend into 

the dissepimentarium. Minor septa absent or very poorly developed. Columella a simple thickened axial plate 

connected to the counter septum in most corallites but 

absent in some. Dissepimentarium consists of 1 to 3 

rows of large elongate dissepiments. Tabulae complete, 

horizontal, and spaced about 1 mm apart. Ratio of 

tabularium width to corallite diameter about 0.3 to 0.4 

Increase peripheral and intermural. 

Description of type material.-. See Kozyreva (197 4). 

Discussion. -This species is distinguished by its 

polymorphic coralla, which include the morphologies of 

Petalaxis, Sciophyllum, and Thysanophyllum. 

Occurrence. -Middle Carboniferous, Bashkirian. 

Horizon b4 , Voronezh anteclise, U.S.S.R. 

Petalaxis evidens Kozyreva 

Petalaxis evidens Kozyreva, 1974, p. 30, pl. 2, figs. 2a, b, 3. 

Diagnosis. - Petalaxis with corallite diameter 8 to 16 

mm and 18 to 22 .major septa that extend about twothirds of the way to the columella but seldom extend into 

the dissepimentarium. Minor septa absent or very poorly developed. Columella a simple thickened axial plate 

connected to the counter septum. Dissepimentarium 

consists of 1 to 3 rows of large inflated dissepiments. 

Tabulae complete and incomplete, horizontal, concave, 

turned up at the columella, and irregularly spaced 0.25 

to 1 mm apart. Ratio of tabularium width to corallite 

diameter about 0.3 to 0.4. Mode of increase unknown. 

Desc·ript'ion of type material.- See Kozyreva (197 4). 

Discussion.-This species is distinguished from P. 

korkhovae by having more major septa, longer major 

septa, and by having its tabulae turned up at the columella.. 

Occurrence. Middle Carboniferous, Bashkirian. 

Horizon b4, Voronezh anteclise, U.S.S.R. 
34 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Petalaxis maccoyanus species group 

Petalaxis maccoyanus Milne-Edwards and Haim.e 

Stylaxis M'coyana Milne-Edwards and Haime, 1851, p. 453, pl. 12, 

figs. 5, 5a. 

Petalaxis M'coyana Milne-Edwards and Haime, 1852, p. 205; MilneEdwards, 1860, p. 440; Roemer, 1883, p. 387. 

Lithostrotion Portlocki Milne-Edwards and Haime. Eichwald, 1861, p. 

149, 150. 

Lithostrotion Mac-Coyanum Milne-Edwards and Haime. Eichwald, 

1861, p. 150. 

Lithostrotion maccoyanum (Milne-Edwards and Haime). Bassler, 

1950, p. 222. 

Petalaxis mccoyana Milne-Edwards and Haime. Fedorowski and 

Gorianov, 1973, p. 58, pl. 12, figs. 4a, b, text-figs. 20a, b; 

Sutherland, 1977, p. 185, figs. 1-7. 

NotPetalaxis maccoyana Milne-Edwards and Haime. Fomichev, 1953, 

p. 453, pl. 31, figs. la-d, 2a-d. 

Not Lithostrotionella maccoyana (Edwards and Haime). de Groot, 

1963, p. 82, pl. 16, figs. la, b. 

Diagnosis. -Petalaxis with maximum corallite 

diameter 5.4 to 7.8 mm and 13 to 18 major septa that approach the columella but seldom reach it and seldom extend into the dissepimentarium. Minor septa poorly 

developed. Columella a thickened axial plate connected 

to the counter septum augmented in some corallites by 

vertical axial tabellae and rare, poorly developed septal 

lamellae. Dissepimentarium composed of 1 to 2 rows of 

small inflated dissepiments Tabulae mostly incomplete, 

essentially horizontal, flat, concave, or convex, and irregularly spaced 0.5 to 1 mm apart. Ratio of tabularium 

width to corallite diameter about 0.6. Increase 

peripheral. 

Description of type material.- See Milne-Edwards 

and Haime (1851). According to Sutherland (1977, p. 

185), the holotype and paratype are lost and a neotype 

may have to be designated from a topotype lot in the collections of the Paleontological Institute of the Academy 

of Sciences of the USSR in Moscow. Pending further 

work by Sutherland, the diagnosis is based largely on 

descriptions and illustrations of topotypes published by 

Fedorowski and Gorianov (1973) and Sutherland (1977). 

Discu,ssian.-This species, the type of Petalaxis, has a 

narrow dissepimentarium like species of the P. flexuosus 

group but is distinguished by its variable columella, 

which ranges from a simple thickened axial plate to a 

complex structure made up of axial plate, vertical 

tabellae, and rare septal lamellae. Specimens of similar 

general morphology but lacking a complex axial structure were referred to the species by Fomichev (1953) 

and de Groot (1964) but are here excluded from the 

species. The only described material included in the 

species is from the Moscow Basin. 

Occurrence.- Middle Carboniferous, Moscovian. 

Myachkovo Horizon, Moscow Basin, U.S.S.R. 

Petalaxis stylaxis (Trautschold) 

Lithostrotion stylaxis Trautschold, 1879, p. 36, pl. 5, figs. 6a-c. 

Petalaxis stylaiis Stuckenberg, 1888, p. 21, pl. 3, figs. 17-21. 

?Petalaxis stylaxis Trautschold. Bolkhovitinova, 1915, p. 63, pl. 5, 

fig. 1. 

Lithostrotionella stylaxis (Trautschold). Dobrolyubova, 1935a, p. _10, 

pl. 1, figs. 1-5, pl. 2, figs. 1, 2, 5; pl. 13, figs. 1-3; 1935b, p. 14, pl. 

1, figs. 1-5, pl. 2, figs. 1, 2, 5, pl. 13, figs. 1-3; Fomichev, 1939, p. 

60, pl. 9, figs. 6a, b; not Wu and Zhao, 1974, p. 271, pl. 137, 

figs. 5, 6. 

?Lithostrotionella stylaxis (Trautschold). Heritsch, 1937, p. 164, figs. 

1-4; 1939, p. 29, pl. 18, fig. 7, pl. 21, figs. 17, 18. 

?Lithostrotion [Lithostrotionella] stylaxis (Trautschold). Bassler, 

1950, p. 235. 

Diagnosis. -Petalaxis with corallite diameter 3.5 to 9 

mm and 11 to 18 major septa that approach the columella but seldom reach it and seldom extend into the 

dissepimentarium. Minor septa absent to well 

developed. Columella ranging from a simple axial plate 

ordinarily connected to the counter septum to a complex 

structure in which the axial plate is augmented by impersistent vertical axial tabellae and rare septal 

lamellae. Dissepimentarium composed of 1 to 3 rows of 

inflated dissepiments of varying sizes. Tabulae complete 

and incomplete, horizontal, flat, concave or convex, and 

irregularly spaced 0.25 to 2 mm apart. Ratio of 

tabularium width to corallite diameter 0.4 to 0. 7. Mode 

of increase unknown. 

Description of type material.- See Trautschold (1879). 

The diagnosis is taken largely from Dobrolyubova's 

(1935b) descriptions and illustrations of topotypes. 

Discussion. -Dobrolyubova (1935a, b) recognized 

three morphological variants in her study of topotypes: 

Lithostrotionella stylaxis, L. stylaxis var. 1, and L. 

stylaxis var. 2. These were treated like separate species 

in a morphological series. The three morphotypes were 

collected from the same horizon at different geographic 

locations, all in the Moscow Basin. Pending further 

study, I have included all three morphotypes in the concept of Petalaxis stylaxis. 

Bolkhovitinova's (1915) specimen is included here with 

a question because of inadequate description and illustration. Heritsch's (1937, 1939) specimens are referred questionably to the species because no longitudinal 

sections are available. 

Petalaxis stylaxis differs from P. maccoyanus by having slightly larger corallites, an unthickened columella, 

and some well-developed minor septa. 

Occurrence.- Middle Carboniferous, Moscovian. 

Myachkovo Horizon (C2

4), Moscow Basin, U.S.S.R. Corakalk, Kings Bay, Spitzbergen(?). Arabian Desert(?). 

Petalaxis mohikanus (Fomichev) 

Lithostrotionella mohikana Fomichev, 1939, p. 60, pl. 9, figs. 5a, b. 

Petalaxis mohikana Fomichev, 1953, p. 459, pl. 32, figs. la, b, 2a, b, pl. 

33, fig. 1. 
SYSTEMATIC PALEONTOLOGY 35 

Diagnosis.-Petalaxis with corallite diameter 6 to 10 

mm and 13 to 18 major septa that closely approach the 

columella and commonly extend into the dissepimentarium. Minor septa well developed. Columella ordinarily a simple axial plate but augmented by rare short septal lamellae and a few vertical axial tabellae in some corallites. Dissepimentarium composed of 1 to 3 rows of 

mostly large inflated dissepiments. Tabulae complete 

and incomplete, essentially horizontal, commonly turned 

up at columella, and spaced 0.25 to 0.5 mm apart. Ratio 

of tabularium width to corallite diameter about 0.4 to 

0.5. Mode of increase unknown. 

Desc'ription of type material.- See Fomichev · (1953). 

Discussion.-This species differs from P. maccoyanus 

by having larger corallites, more extensions of the major 

septa into the dissepimentarium, well-developed minor 

septa, and more tabulae. 

Occurrence. -Middle Carboniferous, Moscovian. M3 

Limestone, Donetz Basin, U.S.S.R. 

Petalaxis celadensis (de Groot) 

Lithostrotionella celadensis de Groot, 1963, p. 82, pl. 15, figs. 2a-d. 

Diagnosis. -Petalaxis with corallite diameter 3.5 to 6 

mm and 12 to 16 major septa that extend about halfway 

to the columella but seldom extend into the dissepimentarium. Minor septa absent or poorly developed. Columella ranging from a simple axial plate connected to 

the counter septum to a complex structure composed of 

an axial plate, vertical axial tabellae, and a few septal 

lamellae. Dissepimentarium composed of 1 to 2 rows of 

large inflated dissepiments. Tabulae mostly incomplete, 

flat to concave, horizontal, and irregularly spaced 0.25 

to 0. 75 mm apart. Ratio of tabularium width to corallite 

diameter about 0.6. Mode of increase unknown. 

Desc'ription of holotype.- See de Gro9t (1964 [1963]). 

Discussion.-This species differs from P. maccoyanus 

by having smaller corallites, fewer and shorter major 

septa, weaker minor septa, and more tabulae. 

Occurrence.- Middle Carboniferous, Moscovian. 

Celada Limestone, Palencia, Spain. 

Petalaxis elyensis (Wilson and Langenheim) 

Eastorwides elyensis Wilson and Langenheim, 1962, p. 512, pl. 88, 

figs. 4-6. 

Diagnosis. -Petalaxis with corallite diameter 4 to 5 

mm and 10 to 15 major septa that extend about halfway 

to the columella. Major septa are attached to the corallite wall except where impersistent lonsdaleoid 

dissepiments are present. Minor septa poorly developed, 

contratingent. Columella a thickened axial plate connected to the counter septum and augmented by rare impersistent vertical axial tabellae and very rare septal 

lamellae. Dissepimentarium consists of a single row of 

impersistent inflated dissepiments. Tabulae mostly complete, generally flat, locally sagging or domed, and 

spaced 0.5 to 1 mm apart. Tabularium width equals corallite diameter, except where dissepiments are 

developed. Mode of increase unknown. 

Description of type material. -See Wilson and 

Langenheim (1962). 

Discussion. -This species is distinguished from most 

other species of Petalaxis by having an impersistent 

lonsdaleoid dissepimentarium and impersistent axial 

tabellae. P. elyensis is the type species of Eastonoides 

Wilson and Langenheim, which was proposed to include 

the type species and Lonsdaleia ivanovi Dobrolyubova, 

which Fomichev (1953, p. 464) included in 

Cystolonsdaleia. These species have the general morphology of P. maccoyanus, the type species of Petalaxis, 

which also has impersistent axial tabellae. Thus, 

Eastonoides is regarded as a junior synonym of 

Petalaxis, and P. elyensis is included in the P. maccoyanu.s species group. 

Occurrence. -Lower Permian. Ely Limestone, 

Nevada, U.S.A. 

Petalaxis dobrolyuhovae n. sp. 

Not Styla:r:is portlocki Milne-Edwards and Haime, 1851, p. 453. 

Not Petolazis portlocki Milne-Edwards and Haime, 1852, p. 204, pl. 

38, figs. 44, 4a; Haime, 1860, p. 441. 

Petolazis portlocki Edwards and Haime. Stuckenberg, 1888, p. 22, pl. 

2, figs. 44-49. 

Lonsdakia portlocki (Stuckenberg). Dobrolyubova, 1935a, p. 12, pl. 9, 

figs 1-4, not pl. 10, figs. 1, 2; 1935b, p. 29, pl. 9, figs. 1-4, not pl. 

10, figs. 1, 2. 

Not Cystolonsdaleia portlocki (Dobrolyubova). Fomichev, 1953, p. 467, 

pl. 32, figs. 4a-v. 

Not Lonsdale'ia portlocki (Stuckenberg) densiconus de Groot, 1963, p. 

79, pl. 15, fig. 1. 

DiagnosiH. -Petalaxis with corallite diameter 5 to 11 

mm and 11 to 17 major septa that extend halfway to or 

closely approach the columella but seldom reach it and 

commonly extend into the dissepimentarium. Minor septa absent to well developed. Columella ranging from a 

simple axial plate connected to the counter septum to a 

complex structure composed of axial plate, vertical axial 

tabellae, and rare septal lamellae augmented by 

stereoplasm in some corallites to produce a netlike 

structure in transverse section. Dissepimentarium composed of 1 to 2 rows of inflated dissepiments of varying 

sizes. Tabulae mostly incomplete, flat, concave, or convex, horizontal, and irregularly spaced 0.25 to 1 mm 

apart. Ratio of tabularium width to corallite diameter 

about 0.6 to 0.7. Mode of increase unknown. 

Description of type material. -See Dobrolyubova 

(1935b). The holotype is the specimen illustrated by 

Dobrolyubova (1936b) as her plate 9, figures 1 and 2. 

Disc1tssion.-Dobrolyubova's illustrations suggest 

that more than one species may be present in the type 

lot. I exclude the specimen figured as her plate 10, 

figures I and 2, because it seems much more complex 
36 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

than the other two specimens illustrated. The remaining 

figured specimens are notably different from one 

another but are retained in the species concept pending 

further work on the type material. 

Cystolonsdaleia portlocki (Dobrolyubova) Fomichev 

does not belong here and is described below as a new 

species. Lonsdaleia portlocki (Stuckenberg) densiconus 

de Groot is a separate species assigned to 

Cystolonsdaleia Fomichev. 

P. dobrolyubovae differs from P. maccoyanus by having larger corallites, more abundant complex columellae, and more extensions of the major septa into the 

dissepimentarium. 

Occurrence.- Middle Carboniferous, Moscovian. 

Myachkovo Horizon, Moscow Basin, U.S.S.R. 

Petalaxis donetsensis n. sp. 

Cystolonsdaleia portlocki (Dobrolyubova). Fomichev, 1953, p. 467, pl. 

32, figs. 4a-v. 

Diagnosis.-Petalaxis with corallite diameter 8 to 10 

mm and 13 to 16 major septa that approach the columella but do not ordinarily reach it and seldom extend 

into the dissepimentarium. Minor septa absent or poorly 

developed. Columella ranging from a simple axial plate 

connected to the counter septum to a more complex 

structure composed of axial plate and impersistent vertical axial tabellae. Dissepimentarium composed of 2 to 

4 rows of inflated dissepiments of varying sizes. Tabulae 

mostly incomplete, mostly concave, horizontal, and irregularly spaced 0.25 to 1 mm apart. Ratio of 

tabularium width to corallite diameter about 0.3 to 0.6. 

Mode of increase unknown. 

Description of holotype.- See Fomichev (1953). The 

holotype is specimen number 36 of Fomichev. 

Discussion.- This species differs from P. 

dobrolyubovae by having more rows of dissepiments, 

weaker minor septa, a smaller ratio of tabularium width 

to corallite diameter, and by its less abundant complex 

columellae. 

Occurrence.- Middle Carboniferous, Moscovian. N 1 

Limestone, Donetz Basin, U.S.S.R. 

Petalaxis ivanovi (Dobrolyubova) 

Lonsdaleia ivanovi Dobrolyubova, 1935a, p. 12, pl. 11, figs. 1, 2; 

1935b, p. 31, pl. 11, figs. 1, 2. 

Diagnosis. -Petalaxis with corallite diameter 6 to 6.5 

mm and 13 to 14 major septa that closely approach the 

columella but seldom reach it. Major septa are attached 

to the corallite wall, except where impersistent 

lonsdaleoid dissepiments are present. Minor septa well 

developed. Columella ranging from a simple axial plate 

to a complex structure consisting of axial plate, vertical 

axial tabellae, and a few septal lamellae commonly reinforced by stereoplasm to form a thick rod or netlike 

structure in transverse section. Dissepimentarium impersistent, composed of a single row of dissepiments of 

varying sizes where present. Tabulae mostly incomplete, horizontal, flat, concave, or convex, and irregularly spaced 0.25 to 1 mm apart. Tabularium width equals 

corallite diameter except where dissepiments are present. Mode of increase unknown. 

Description of holotype. -See Dobrolyubova (1935b ). 

Discussion.-This species differs from P. elyensis by 

having larger corallites, stronger minor septa, and a better developed complex columella. 

Occurrence. -Middle Carboniferous, Moscovian. 

Myachkovo Horizon, Moscow Basin, U.S.S.R. 

Petalaxis? spp. indet. 

The following taxa are referred questionably to 

Petalaxis and are not diagnosed because of insufficient 

illustration or description of morphological details: 

Lithostrotionella tingi Chi, 1931, p. 28, pl. 4, figs. 6a, b; 

Wu in Yu and others, 1963, p. 86, pl. 24, figs. 6a, b. 

Middle Carboniferous, Moscovian. Laokunchai 

Limestone, Kueichou, China. 

Lithostrotionella aff. stylaxis Trautschold. 

Dobrolyubova, 1936b, p. 127, figs. 55, 56. Middle 

Carboniferous, Moscovian. Ural Mountains, 

U.S.S.R. 

Lithostrotionella sp. indet. Yamagiwa, 1961, p. 102, pl. 

5, figs. 1-3. Lower Permian. Atetsu Limestone, 

Okayana, Japan. 

Lithostrotionella kitakamiensis Minato, 1955, p. 88, pl. 

4, figs. 2, 7, 8, 10, pl. 34, figs. 2, 3. Middle Carboniferous, Bashkirian. Nagaiwa Series, Iwate, 

Japan. 

Lithostrotionella spiniformis Yu, 1933 [1934], p. 102, pl. 

21, figs. 2a, b; Wu in Yu and others, 1963, p. 87, pl. 

24, figs. 5a, b. Lower Carboniferous, Visean. 

Shangssu Limestone, Kueichou, China. 

Petalaxis timanicus Stuckenberg. Heritsch, 1939, p. 18, 

pl. 2, fig. 8, pl. 3, fig. 1, pl. 15, figs. 2, 3, pl. 21, figs. 

8-16. Carboniferous or Permian. Cora Limestone, 

Spitzbergen. 

Petalaxis timanicus Stuckenberg. Kolosvary, 1951, p. 

39, pl. 9, figs. 5-7. Permian, Hungary. 

Petalaxis maccoyana var. multiseptata Fomichev, 1953, 

p. 458, pl. 31, fig. 4. Middle Carboniferous, Moscovian. L5 Limestone, Donetz Basin, U.S.S.R. 

Petalaxis maccoyana forma orlovkensis Fomichev, 1953; 

p. 456. Middle Carboniferous, Moscovian. L5 and L6 

Limestone, Donetz Basin, U.S.S.R. 

Family LONSDALEIIDAE Chapman, 1893 

Genus THYSANOPHYLLUM Nicholson and Thomson, 1876 

Thysanophyllum Nicholson and Thomson, 1876, p. 150; Hill, 1940, p. 

160. 
SYSTEMATIC PALEONTOLOGY 37 

Type species.-Thysanophyllum orientale Nicholson and 

Thomson, 1876, p. 150. Lower Carboniferous, Scotland. 

Discussion.- Thysanophyllum is distinguished by its 

cerioid corallum having corallites with a lonsdaleoid 

dissepimentarium and lacking an axial structure or having a weak impersistent axial plate. The genus is probably polyphyletic like other colonial rugose corals 

without a persistent axial structure, corals such as 

Diphyphyllum and Pseudodorlodotia. 

Thysanophyllum astraeiforme (W arran) 

Diphyphyllum aJJtraeiformis Warren, 1927, p. 44, pl. 3; figs. 2, 3. 

Lithostrotionella aJJtraeiformis (Warren). Kelly, 1942, p. 352; Bamber, 

1961, p. 152, pl. 12, figs. 1a, b; Nelson, 1961, pl. 18, figs. 1-3. 

Lithostrotionella (Thysarwphyllum) aJJtraeijormis (Warren). Nelson, 

1960, p. 115, pl. 22, figs. 7-10. 

Thysarwphyllum aJJtraeiforme (Warren). Bamber, 1966, p. 23, pl. 4, 

figs. 3a-b, 4a-c; Armstrong, 1970a, p. 37, pl. 11, figs. 5-8; 1970b, 

p. 28, pl. 9, figs. 1-6, text-fig. 33. 

Desc.-'ription of lectotype.- See Bamber (1966). 

Discussion.-This species is rather common in 

western Canada and Alaska, where it ranges from 

Mamet Zone 13 into 16i; it is most common in Mamet 

Zones 14 and 15. These occurrences are of middle and 

late Visean age. The structure of the columella, where 

present, suggests relationship to Acrocyathus. 

Genus LONSDALEIA McCoy, 1849 

Lonsdaleia McCoy, 1849, p. 11; Smith, 1916, p. 218; Hill, 1940, p. 151; 

Sando, 1975, p. C20. 

Type species. -Erismatolithus Madreporites 

(duplicatus) Martin, 1809, equals Lonsdaleia duplicata 

(Martin). Lower Carboniferous, England. 

Diagnosis.- See Cotton (1973, p. 117). 

Discussion.-The genus is subdivided into two 

subgenera on the basis of growth form: Lonsdaleia 

(Lonsdaleia), phaceloid, and Lonsdaleia (Actinocyathus), 

cerioid. 

Subgenus ACTINOCYATHUS d'Orbigny, 1849 

Actinocyathus d'Orbigny, 1849a, p. 12. 

Stylidnphyllum de Fromentel, 1861, p. 316. 

?Protolomdalia Lisitsyn, 1925, p. 68. 

?Sublonsdalia Lisitsyn, 1925, p. 68. 

?Protolomdaleia Lang, Smith, and Thomas, 1940, p. 106. 

?Sublonsdaleia Lang, Smith, and Thomas, 1940, p. 128 

Type. species. - Cyathophyllum crenulare Phillips, 

1836, equals Erismatolithus Madreporites (jloriformis) 

Martin, 1809, equals Lonsdaleia floriformis (Martin). 

Lower Carboniferous, England. 

Diagnosis.-Cerioid Lonsdaleia. 

Discussion.- See Sando (1975, p. C20) for discussion 

of the type species and synonymic placement of 

Stylidophyllum. Protolonsdalia (nomen vanum Protolonsdaleia) and Sublonsdalia (nomen vanum 

Sublonsdaleia) are both inadequately founded on type 

material that has been lost (N. P. Vasilyuk, oral commun., 1975); they may be junior synonyms of Actinocyathu.s, but neotypes need to be described before a 

final decision can be made. 

Lc•nsdaleia (Actinocyathus) berthiaumi (Merriam) 

Plate 20, figures 3, 4 

Lithostrotion(Lithostrotionella) berthiaumi Merriam, 1942, p. 378, pl. 

56, figs. 9, 10; Bassler, 1950, p. 252. 

Descript'ion of holotype.- See Merriam (1942). 

Discussion. -Lonsdaleia (Actinocyathus) was 

previously represented· in North America by a single 

species, L. (A.) stelcki Nelson (1960, p. 119, pl. 23, figs. 

6-10), as determined by Sando (1975, p. C21). Nelson's 

species is restricted to the upper Visean and lower 

Nammian (Mamet Zones 16s, 17, and 18). Examination 

of the holotype (USNM 132988) and a topotype (USNM 

132989) of Merriam's species reveals that it is very 

similar to Nelson's species but differs in having fewer 

tabulae. New thin sections of Merriam's holotype 

(USNl\1 132988) are figured herein to provide a better 

basis for interpreting this specimen. 

Merriam (1942, p. 379) stated that the type material of 

L. berthiaumi is from the Permian Coyote Butte Formation of Oregon, a stratigraphic level that is inconsistent 

with other North American occurrences of the genus to 

which it is now assigned. According to E. C. Wilson 

(written commun., 1979), exposures in the type area are 

poor, geologic structure is complex, and beds in the area 

range from Devonian to Permian. I conclude that the 

specimens are probably from beds of Carboniferous (late 

Visean or early Namurian) age. 

lonscl.aleia (Actinocyathus) peratrovichensis (Armstrong) 

Lithostrotionella peratrovichensis Armstrong, 1970a, p. 35, pl. 12, 

figs. 8-11. 

Description of holotype.- See Armstrong (1970a). 

Discussion.- Examination of the holotype (USNM 

160493) of L. peratrovichensis reveals a morphology 

consistent with Lonsdaleia (Actinocyathus). The species 

differs from the widely distributed L. (A.) stelcki 

(Nelson) by its smaller corallite diameter, thicker corallite walls, fewer dissepiments, and tabulae of varying 

form, some of which are inclined upward toward the columella. 

The presence of L. (A.) peratrovichensis in beds of late 

Visean age: and the nature of its tabulae suggest that it 

is ancestral to L. (A.) stelcki. 

Family DURHAMINIDAE Minato and Kato, 1965 

Genus KLEOPATRINA McCutcheon and Wilson, 1963 

Ptolema'i..a McCutcheon and Wilson, 1961, p. 1020 (not Osborn, 1908, 

p. 267). 

Kleopatrina McCutcheon and Wilson, 1963, p. 299; Minato and Kato, 

1965, p. 67 (replacement name for Ptolemaia McCutcheon and 

Wilson). 
38 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Type species.-Ptolemaia ftatateeta McCutcheon and 

Wilson, 1961, p. 1025, pl. 123, figs. 1-6. Lower Permian, 

·Nevada. 

Diagnosis.- See Minato and Kato (1965). 

Discussion. -This genus is found predominantly in 

Permian strata, although Bamber and Macqueen (1979, 

p. 11) reported Kleopatrina (Porfierevella) from beds of 

Moscovian Age in British Columbia. The two species 

questionably referred here (see below) are the first 

possible representatives from the Upper Carboniferous. 

Subgenus KLEOPATRINA McCutcheon and Wilson, 1963 

Type species.-As for genus Kleopatrina. 

Diagnosis.- See Minato and Kato (1965). 

Kleopatrina (Kleopatrina)? dilatata (Easton) 

Lithostrotion [Lithostrotionella] dilatata Easton, 1960, p. 578, textfigs. 7, 8. 

Description of holotype.- See Easton (1960). 

Discussion. -Judging from Easton's description and 

illustrations, the type material of this species has most 

of the essential features of Kleopatrina (Kleopatrina). 

However, the types do not seem to have a complex axial 

structure, which casts doubt on the taxonomic placement. The type material needs to be reinvestigated. 

Occurrence.- Permian. Arcturus and Rib Hill Formations, Nevada, U.S.A. 

Kleopatrina (Kleopatrina)? uralica (Dobrolyubova) 

Lithostrotionella uralica Dobrolyubova, 1936a, p. 28, pl. 13, figs. 

33-35. 

Description of type material. -See Dobrolyubova 

(1936a). 

Discussion.- Dobrolyubova's description and illustrations of this species suggest that it may be an early 

representative of Kleopatrina (Kleopatrina). This 

species occurs in beds of Late Carboniferous age, 

whereas all other known representatives of the 

subgenus are from the Permian. The type material 

needs to be reinvestigated. 

Kleopatrina (Kleopatrina)? wahooenis (Armstrong) 

Lithostrotionella wahooenis Armstrong, 1972b, p. 14, pl. 5, fig. 1, pl. 

6, figs. 1-5, pl. 7, figs. 1-3. 

Description of holotype and paratypes.- See Armstrong (1972b). · 

Discussion.-Armstrong's holotype and para types 

have the essential morphological characters of 

Kleopatrina (Kleopatrina). Differences between these 

specimens and the holotype of K. (K.) ftatateeta seem to 

be of specific ·rank only. However, Armstrong's species 

is from beds that are much older than those in which 

other known species of the subgenus are found. The occurrence is Middle Pennsylvanian (Atokan) Mamet Zone 

21, which equates with high Bashkirian or low Moscovian of the U.S.S.R. Because of possible homeomorphism with the Permian forms, the placement of this 

species in Kleopatrina (Kleopatrina) is made with question. 

UNDETERMINED LITHOSTROTIONELLOID CORALS 

The following taxa are not identified generically 

because of inadequate specimens or inadequate illustrations and descriptions: 

Lithostrotionella americana Hayasaka, 1936, p. 62 

(Para types, USNM 17 4371 and 17 4373). Permian. 

McCloud Limestone, California, U.S.A. 

Lithostrotionella flexuosa (Trautschold). Heritsch, 1940, 

p. 72, pl. 2, figs. 1, 2. Upper Carboniferous, 

Yugoslavia. 

Petalaxis grandis Heritsch, 1939, p. 27, pl. 2, figs. 4, 5, 

pl. 19, fig. 11. Carboniferous or Permian, Spitzbergen. 

Petalaxis? inconfertus Lonsdale. Yanishevskiy, 1900, p. 

89. Carboniferous, Ural Mountains, U.S.S.R. 

Petalaxis indigae Stuckenberg, 1895, p. 76, pl. 19, fig. 7. 

Upper Carboniferous, Timan, U.S.S.R. 

Lithostrotionella kuechouensis Yii, [1933] 1934, p. 101, 

pl. 21, figs. 1a, b; Wu in Yii and others, 1963, p. 87, 

pl. 25, figs. 1a, b. Lower Carboniferous, Visean. 

Shangssu Limestone, Kueichou, China. 

Petalaxis kunthi Stuckenberg, 1888, p. 23; 1895, p. 78, 

pl. 12, fig. 7; Heritsch, 1939, p. 25, pl. 11, fig. 1, pl. 

13, figs. 3, 7, pl. 19, fig. 4. Upper Carboniferous, 

Moscow Basin and Ural Mountains, U.S.S.R. Carboniferous or Permian, Spitzbergen. 

Petalaxis sibiricus Gabounia, 1919, p. 39, pl. 2, fig. 2, pl. 

3, fig. 1; Fomichev, 1931, p. 44, 72, pl. 1, figs. 2a-e; 

1955, p. 303, pl. 80, figs. 5, 6. Lower Carboniferous, 

Kuznetsk Basin, U.S.S.R. 

Lithostrotionella stylaxis (Trautschold). Rukhin, 1938, 

p. 40, pl. 5, figs. 7-9. Lower Carboniferous, Siranka 

River basin, U.S.S.R. 

Lithostrotionella stylaxis (Trautschold). Wu and Zhao, 

1974, p. 271, pl. 137, figs. 5, 6. Middle Carboniferous, Wei-Ling Group, Kueichou, China. 

Petalaxis timanicus Stuckenberg, 1895, p. 76, pl. 12, fig. 

5. Upper Carboniferous, Ural Mountains and Timan, 

U.S.S.R. 

Lithostrotionella tingi Chi. Yii, . Lin and Fan, 1962, p. 

24, pl. 4, figs. 3a, b. Middle Carboniferous, Sinkiang, 

China. 

Styllophyllum tolmachevi Rukhin, 1938, p. 41, pl. 5, figs. 

10, 11. Lower Carboniferous, Omulevka River, 

U.S.S.R . 

. Petalaxis uchtensis Stuckenberg, 1895, p. 77, pl. 12, fig. 

2, pl. 16,fig. 4. Upper Carboniferous, Timan, 

U.S.S.R. 
SYSTEMATIC PALE ONTOLOGY 39 

Lithostrotionella cf. spiniformis Yii. Wu, 1964, p. 35, 

79, pl. 4, figs. 7, 8. Lower Carboniferous, Visean. 

Hunan, China. 

Lithostrotionella sp. B. Lo and Chao, 1962, p. 184, pl. 

22, fig. 6. Lower Carboniferous, Visean. Ching-hai, 

China. 

Lithostrotionella sp. Kolosvary, 1951, p. 38, pl. 10, figs. 

4-9,pl. 18, figs. 1-3. Carboniferous or Permian, 

Hungary. 

Lithostrotionella sp. Dobrolyubova, 1936a, p. 29, pl. 13, 

figs. 36, 37. Upper Carboniferous, Ural Mountains, 

U.S.S.R. 

Lithostrotionella sp. undet. Minato, 1955, p. 87, pl. 1, 

fig. 3, pl. 34, fig. 11, pl. 37, fig. 9. Lower Carboniferous, I wate, Japan. 

Lithostrotionella? sp. Ross and Ross, 1963, p. 418, pl. 

49, figs. 1, 2, 4, 10. Upper Carboniferous, Virgilian. 

Gaptank Formation, Texas, U.S.A. 

ADDITIONAL TAXA 

While the manuscript was being processed for publication, several additional studies of lithostrotionelloid corals described under the names of Hillia, Lithostrotionella, Petalaxis, Stelechophyllum, and 

Thysanophyllum came to my attention. Translations of 

the descriptions of these taxa that are in Chinese and 

Russian papers were not available at the time, preventing a complete evaluation of them. Moreover, taxa in the 

Chinese papers are poorly illustrated. These additional 

taxa are listed below, by publication. 

Fan, Y. N., 1978, in Southwest China Geological Institute, [Paleontological atlas of southwestern 

China, Sichuan Province]: v. 2, 684 p., 191 pls. (In 

Chinese.) Seep. 182-184. 

Lithostrotionella maccoyana (Edwards and Haime) 

L. sexangula de Groot 

L. mui (Lo) 

L. fugimotoi (Igo) 

L. orboensis de Groot 

L. orboensis regularis Fan, n.subsp. 

L. pinguis Fan, n. sp. 

L. awenggouensis Fan, n. sp. 

L. vesiculosa vesiculosa Fan, n. subsp. 

?L. sp. 

Stelechophyllum ascendens ascendens (Tolmachev) 

S. ascendens simplex Drobolyubova 

Gorskiy, I. 1., 1978, Korally Srednego Karbona zapadnogo sklona U rala [The Middle Carboniferous corals 

of the western slope of the Urals]: Akademiya Nauk 

SSSR, Otdelenie Geologii, Geofiziki, i Geokhimi, Izdatelstvo "Nauka," 223 p., 23 pls., 43 figs. See p. 

150-152. 

Lithostrotionella stylaxis (Trautschold) subsp. 

'Uralica Gorskiy, n. subsp. 

L. flexuosa (Trautschold) 

Guo, Z. C., 1976, in Northeastern Geological Institute 

and Geological Bureau of Inner Mongolia, [Paleontological atlas of northern China, part of Inner 

Mongolia]: v. 1, 502 p., 232 pls. (In Chinese.) Seep. 

86. 

Lithostrotionella ivanovi Dobrolyubova 

L. banffense (Warren) 

Wang, H. D., 1978, in Stratigraphic-Paleontological 

Working Team of Guizhou Province, [Paleontological atlas of southwestern China, Guizhou Province]: v. 2, 638 p., 165 pls. (In Chinese.) See p. 

133-140. 

Lithostrotionella kueichouensis Y ii 

L. kuechouensis magna Wang, n. subsp. 

L. baijinensis Wang, n. sp. 

L. multivesiculata Wang, n. sp. 

L. spiniformis Yii 

L. changshunensis Wang, n. sp. 

L. stylaxis (Trautschold) 

L. tingi Chi 

L. dushanensis Wang, n. sp. 

L. elegantula (Wu and Zhao) 

L. jiaruoensis Wang, n. sp. 

H'illia perapertuensis de Groot 

H. mi~rwr (Wu and Zhao) 

Wilson, E. C., 1982, Wolfcampian rugose and tabulate 

corals (Coelenterata: Anthozoa) from the Lower 

Permian McCloud Limestone of northern California: 

Los Angeles County Natural History Museum Contributions in Science no. 337, 90 p., 48 figs. See p. 

65-73. 

Petala.xis allisonae Wilson, n. sp. 

P. besti Wilson, n. sp. 

P. kennedyi Wilso;n, n. sp. 

P. pecki Wilson, n. sp. 

P. sutherlandi Wilson, n. sp. 

Xu, 1977, in Central Southern Geological Institute and 

Geological Bureaus of Henan, Hubei, Hunan, 

Guangdong, and Guangxi Provinces, [Paleontological atlas of central southern China]: v. 2, 856 

p., 253 pls. (In Chinese.) Seep. 202-203. 

Lithostrotionella hsujiulingi Y oh 

L. micra Kelly 

L. maccoyana (Edwards and Haime) 

Thysanophyllum ascendens ascendens (Tolmachev) 

T. ascendens simplex (Dobrolyubova) 

Yii, C. C., Lin, I. T., Huang, C. H., and Tsai, T. S., 1978, 

[Early Carboniferous stratigraphy and corals of 

eastern Xinjiang]: Chinese Academy of· Geological 

Sciences, Editorial Committee of Professional 

Papers of Stratigraphy and Paleontology, Profes-
40 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

sional Papers of Stratigraphy and Paleontology No. 

5, p. 1-70, 10 figs., 16 pis. (In Chinese with English 

summary.) See p. 29, 38-40. Peking, Geological 

Publishing House) 

Lithostrotionella shimeri (Crickmay) 

L. pennsylvanica (Shimer) 

L. crassus n. sp. 

REGISTER OF USGS LOCALITIES FOR 

RAYASAKA (1936) TYPE SPECIMENS OF 

LITHOSTROTIONELLA SPECIES 

83-PC: Newman Limestone, StLouis Limestone Member. West side 

of Roundstone Creek, 300ft (100 m) above creek, about 0.5 mi (0.8 

km) south of site of town of Langford near intersection of Roundstone Creek and Renfro Creek, southeast quarter of Wildie 

quadrangle, Rockcastle County, Ky. Collected by F. B. Weeks, 

June 29, 1896. See Gualtieri (1968) for geologic map. 

104-PC: Lodgepole Limestone (Zone C1 of Sando and others, 

1969).Sec. 12, T. 11 S., R. 44 E., Montpelier quadrangle, Bear 

Lake County, Idaho. Collected by G. R. Mansfield, June 12, 1911. 

This collection is composed of two lots, one that has a fauna from 

the upper part of the Lodgepole Limestone and another that contains Faberophyllum and is from the Aspen Range Formation. 

346C-PC: Boulders of residual chert derived from Tuscumbia 

Limestone (St. Louis Limestone equivalent) at unconformity at top 

of Tuscumbia Limestone. Along pike about 0.5 mi (0.8 km) 

southwest of Cherokee, probably in SE1/4 sec. 34 or SW1/4 sec. 35, 

T. 3 S., R. 14 W., Cherokee quadrangle, Colbert County, Ala. Collected by P. V. Roundy, 1911. See Butts (1926) for geologic map of 

the area. See McCalley (1896, p. 152-153, 158) and Butts (1926, p. 

175) for description of the geology. 

490-PC: Float from Lodgepole Limestone (Zone C1 of Sando and 

others, 1969). Old Laketown Canyon, near center of sec. 32, T. 13 

N., R. 6 E., Randolph quadrangle, Rich County, Utah. Collected by 

G. B. Richardson, 1912. See Richardson (1941) for geologic map 

and Sando and others (1959) for discussion of geology. 

498-PC: St. Louis Limestone, at base. Outskirts of Maeystown, probably in bed of Maeystown Creek about 1 mi (1.6 km) southeast of 

Maeystown in SW1/4 sec. 1, T. 4 S., R. 11 W., Selma quadrangle, 

Monroe County, Ill. Collected by G. H. Girty, September 8, 1912. 

499-PC: St. Louis Limestone, basal beds just above cephalopod bed. 

Ravine in river bluff about 1 mi (1.6 km) southeast of Maeystown, 

probably in SE1/4 sec. 1 or NE1/4 sec. 12, T. 4 S., R. 11 W., Selma 

quadrangle, Monroe County, Ill. Collected by Weller and Girty, 

September 8, 1912. 

643-PC: St. Louis Limestone(?). On Tebo Creek about 1.5 mi (2.4 km) 

east of Leesville, Leesville quadrangle, Henry County, Mo.(?). Collected by W. P. Jenney, 1891. 

Original locality data indicate that the stratigraphic unit was 

probably the Cherokee limestone or Seneca chert of Jenney (1894), 

which Jenney thought to be of Warsaw or St. Louis age. According 

to Wilmarth (1938, p. 415), subsequent authors regarded the 

Cherokee limestone as Boone Formation and the Seneca chert as 

the Grand Falls Chert Member of the Boone. According to Wilson 

(1922, p. 180), most of Henry County is underlain by the Cherokee 

Shale of Pennsylvanian age, but the Burlington and Keokuk 

Limestones are exposed beneath the Cherokee in the valley of 

Tebo Creek. 

Hayasaka's specimen attributed to this locality is surely a St. 

Louis Limestone form. Thus, the locality data may be erroneous or 

the specimen may be a residual fragment derived from the St. 

Louis Limestone during post-St. Louis, pre-Cherokee erosion. 

There are no other specimens in the collection. 

932A-PC: Float from St. Louis Limestone. In cut of St. Louis and San 

Francisco Railroad in NE1/4 SE1/4 or SE1/4 NE1/4 sec. 10, T. 4 N., R. 

5 E., Kirkwood quadrangle, St. Louis County, Mo. Collected by G. 

H. Girty, August 8, 1904. See Fenneman (1911, pl. 1) for geologic 

map of the area. 

970-PC: Alapah Limestone(?). 141st meridian, easterly spur from 

ridge between headwaters forks of Incog Creek, about 2 mi (3.2 

km) northwest from station 966, Table Mountain quadrangle, 

northeastern Alaska. Collected by A. G. ,Maddren, July 1, 1912. 

1148-PC: Float from St. Louis Limestone. 3 mi (4.8 km) west of 

Tennessee, Ill., in NW1/4 SW1/4 sec. 19, T. 5 N., R. 4 W., Colchester 

quadrangle, McDonough County, Ill. Collected by Henry Hinds, 

November 22, 1912. See Hinds (1919) for geologic map. 

1211B-PC: Oolite in lower part of St. Louis Limestone. Mosher quarry 

(Ste. Genevieve Lime and Quarry Company) at Macy (Mosher Station) about 2 mi (3.2 km) southwest of Ste. Genevieve on Illinois 

Southern Railroad in T. 38 N., R. 9 E., Weingarten quadrangle, 

Ste. Genevieve County, Mo. Collected by E. 0. Ulrich, November 

2, 1903. See Weller and St. Clair (1928) for geologic map of the 

area. 

1439-PC: Lodgepole Limestone, Woodhurst Member (Zone C1 of 

Sando and others, 1969). In low saddle on west slope of hill 

(altitude 8,172 ft (2,490 m)) west of Georgetown Canyon, NW1/4 

sec. 34, T. 10 S., R. 44 E., Slug Creek quadrangle, Bear Lake 

County, Idaho. Collected by G. H. Girty, November 11, 1914. This 

locality was mapped as Madison Limestone by Mansfield (1927, pl. 

6). Megafossils associated with the coral specimen are all compatible with a Zone C1 determination. 

1476-PC: Aspen Formation (Mamet foraminifer Zone 14). SW1/4 sec. 

8, T. 5 S., R. 43 E., at altitude 6,650 ft (2,015 m) in point projecting 

north from ridge south of Grays Lake, Lanes Creek quadrangle, 

Caribou County, Idaho. Collected by E. L. Jones and G. H. Girty, 

November 5, 1914. This locality was mapped as Brazer Limestone 

by Mansfield (1927, pl. 4). The file card by Girty indicates "near 

top" of the Brazer, but this locality is in a fault block, so the position is doubtful. Microfossils in the collection indicate Mamet Zone 

14, and megafossils indicate Zone E of Sando and others (1969). 

2013B-PC: Greenbrier Limestone, Hillsdale Member, near base. In 

cut of Norfolk arid Western Railroad along Indian Creek 1.5 mi 

(2.4 km) northeast of C~ Bluff, Tazewell quadrangle 

(30-minute) and Pounding Mill quadrangle (15-minute), Tazewell 

County, Va. Collected by G. H. Girty, June 17, 1920. See Campbell 

(1897) for geologic map of the area. 

2020-PC: Greenbrier Limestone, Hillsdale Member, at base. Forks of 

road 0.5 mi (2.4 km) southeast of Shrader, Tazewell quadrangle 

(30-minute), Tazewell County, Va. Collected by Harnsberger, 

September 1916. See Campbell (1897) for geologic map of the area. 

2020A-PC: Greenbrier Limestone, Hillsdale Member, about 25ft (7.5 

m) above base. Same geographic location as 2020-PC. See Girty 

(1923, p. 470) for faunal list and description of the locality. 

2222A-PC: St. Louis Limestone, 0.75 mi (1.2 km) northeast of Hicks, 

probably in NE1/4 sec. 19, T. 11 S., R. 8 E., Equality quadrangle, 

Hardin County, Ill. Collected by Charles Butts, 1916. See Weller 

(1920, pl. 1) for geologic map of the area. 

2222C-PC: Same as 2222A-PC. 

2226-PC: St. Louis Limestone. 0.75 mi (1.2 km) north of Big Creek 

School, probably in SE1/4 sec. 21, T. 11 S., R. 8 E., Equality 

quadrangle, Hardin County, Ill. Collected by Charles Butts, 1916. 

See Weller (1920, pl. 1) for geologic map of the area. 

2333-PC: St. Louis Limestone. Near top of divide near Allen Springs, 

8 to 10 mi (12.8 to 16 km) northwest of Scottsville, Allen Springs 
LOCALITY DATA FOR USNM SPECIMENS NOT HAYASAKA TYPES 41 

quadrangle, Allen County, Ky. Collected by E. W. Shaw, 1917. See 

Moore (1963) for geologic map of the area. 

3024-PC: Little Flat Formation (Mamet foraminifer Zone 14). NW1/4 

sec. 13, T. 7 S., R. 40 E., altitude 6,650 ft (2,015 m), Henry 

quadrangle, Caribou County, Idaho. Collected by P. V. Roundy, 

June 29, 1916. This locality is mapped as Brazer Limestone by 

Mansfield (1927, pl. 3). Megafossils in the collection indicate Zone 

E of Sando and others (1969), and microfossils are Mamet Zone 14. 

3159-PC: Newman Limestone, at base. 1.5 (2.4 km) mi northeast of 

Cleveland, Carbo quadrangle (7 1/2-minute), Russell County, Va. 

Collected by Charles Butts, March 1912. See Campbell (1899) for 

geologic map of the area. 

3282-PC: Hillsdale Member of Greenbrier Limestone, lower 50.:::.75 ft 

(15-23 m). In low hills and flats of the "little levels" within 1 mi (1.6 

km) west and southwest of Mill Point, Marlintown quadrangle 

(15-minute), Pocahontas County, W.Va. Collected by W. A. Price 

and G. H. Girty, September 23, 1920. See Price (1929) for geologic 

map of the area and Girty (1923, p. 456) for faunal list and description of the locality. The collection contains corals, bryozoans, and 

brachiopods listed by Girty. 

3283-PC: Hillsdale Member of Greenbrier Limestone, lower 70ft (21.2 

m). Limestone ledges at Mill Point in small hill (altitude 

2,2.60-2,330 ft, 685-706 m) just north of residence of Mr. Wallace, 

east of Stamping Creek and north of highway from Mill Point to 

Buckeye, Marlinton quadrangle (15-minute), Pocahontas County, 

W.Va. Collected by W. A. Price and G. H. Girty, September 22, 

1920. See 3282-PC for further information on this locality. 

3290-PC: Lodgepole Limestone, Woodhurst Member (Zone C1 of Sando and others, 1969). Near Lime Spur, probably' in sec. 17, 18, 19, 

or 20, T. 1 N., R. 2 W., Jefferson Island quadrangle, Jefferson or 

Madison County, Mont. Collected by D. C. Bard. See Chelini (1965, 

fig. 5) for geologic map of the area. Collection contains megafossils 

that indicate Zone C1 of Sando and others (1969) and algae that indicate Mamet Zones 7-8. 

3747C-PC: Peratrovich Formation, limestone and chert member (upper Meramecian). South shore of Shelikof Island at entrance to 

Soda Bay, about on parallel55°15', southeastern Alaska. Collected 

by G. H. Girty, June 9, 1918. See Armstrong (1970a, p. 29-31) for 

discussion. 

3760-PC: Peratrovich Formation, limestone and chert member (upper 

Meramecian). West end of south shore of Madre de Dios Island, 

southeastern Alaska. Collected by G. H. Girty, June 18, 1918. See 

Armstrong (1970a) for discussion. 

3856-PC (green label): McCloud Limestone (Lower Permian). Crest of 

Limestone ridge 0.5 mi (0.8 km) north of James ranch in NE 1

/4 sec. 

22, T. 33 N., R. 4 W., Redding quadrangle, Shasta County, Calif. 

Collected by J. S. Diller, July 5, 1902. The occurrence of 

Heterocaninia? gahbi (Meek) and Omphalotrochus whitneyi (Meek) 

in the collection confirm the McCloud assignment. 

3858-PC (green label): McCloud Limestone (Lower Permian). 1 mi (1.6 

km) northwest of Lillienthal, probably in NE 1

/4 sec. 27, T. 33 N., R. 

4 W., Redding quadrangle, Shasta County, Calif. Collected by 

Stanton and Richardson, July 8, 1902. The locality was mapped as 

McCloud Limestone by Diller (1906). The only other fossil in the 

collection is Heritschioides? sp. The locality number was erroneously given as 3859 by Hayasaka (1936, p. 64). 

3864-PC: Float from Lodgepole Limestone, Woodhurst Member (Zone 

C1 of Sando and others, 1969). Unknown canyon on south side of 

Little Rocky Mountains, Phillips County(?), Mont. See Knechtel 

(1959, pl. 52) for geologic map of the area. 

The stratigraphic level assigned to this locality is the only one in 

which corals of this kind occur in the Little Rocky Mountains. The 

canyon in which the specimen was found was said to be the one 

containing the powerline to Zortman, but on recent maps of the 

area, I can find no powerline shown in any of the canyons on the 

south side of the Little Rocky Mountains. 

3890-PC (green label): McCloud Limestone (Lower Permian). Summit 

of Gray Rock Ridge northwest of road above Black Diamond mine, 

probably in NE1

/4 sec. 5, T. 33 N., R. 4 W., Redding quadrangle, 

Shasta County, Calif. Collected by Storrs and Washburne. See 

Diller (1906) for geologic map of the locality. Brachiopods in the 

collection indicate a Permian age. The locality number was erroneously given as 3896 by Hayasaka (1936, p. 64). 

3946-PC (green label): Probably from St. Louis Limestone of the 

Mississippi Valley region. The specimen was said to come from the 

Santa Anna Mountains, Orange County, Calif., which is an area of 

post-Paleozoic rocks. It is a museum specimen attributed to Dr. 

Steven Bowen of Los Angeles, Calif. The specimen has all the 

characteristics of corals that are abundant in the St. Louis 

Limestone of the Mississippi Valley region. The locality number 

was erroneously given as 3446 by Hayasaka (1936, p. 61). 

4801H-PC (green label): Little Flat Formation (Member B of 

Brazer Limestone of Mullens and Izett, 1964). Probably in sec. 30, 

T. 10 N., R. 2 E., Paradise quadrangle, Cache County, Utah. Collected by :E. M. Kindle, August 12, 1907. 

This specimen was erroneously attributed by Hayasaka (1936, p. 

66) to USGS Mesozoic loc. 4801H, which is from the Monte de Oro 

Formation (Upper Jurassic), "western portion of the Oroville plant 

beds, north side of Feather River, 3 to 4 mi above Oroville, California." Foraminifera in the specimen are Mamet Zones 13-15 (probably 14) and are Western United States species. 

5892-PC: Madison Limestone (Zone C1 of Sando and others, 1969). 

Ridge west of Holiday Park, probably in sec. 26, 34, or 35, T. 1 N., 

R. 8 E., Coalville quadrangle, Summit County, Utah. The collection contains megafossils that indicate Zone C1 of Sando and 

others (1H69). 

5893-PC: Little Flat Formation (Member B of the Brazer 

Limestone of Mullens and Izett, 1964). Probably in sec. 5 or 6, T. 9 

N., R. 2 E., Paradise Canyon, Paradise quadrangle, Cache County, 

Utah. Collected by E. Finch. The collection contains 

Faherophyllum sp. (Zone F of Sando and others, 1969) and 

foraminifers of Mamet Zone 14. 

5894-PC: Float probably from lower part of the Mission Canyon 

Limestone (Zone C2 of Sando and others, 1969). Crest of ridge east 

of Old Baldy at altitude of about 9,000 ft (2, 727 m), probably in sec. 

26 or 27, T. 7 S., R. 3 W., Varney quadrangle, Madison County, 

Mont. Collected by R. W. Richards. See Hadley (1969) for geologic 

map of the area. 

7130B-PC: Cobble in river wash derived probably from the Alapah 

Limestone. Along Hulahula River, Mount Michelson quadrangle, 

northeastern Alaska. Collected by E. de K. Leffingwell, February 

1908. 

7452-PC (green label): "Wells" Formation (this may actually be a 

Mississippian unit related to the Humbug Formation). South end of 

Cokeville Butte, jt1st west of phosphate mine 1.5 mi (2.4 km) northeast of Cokeville, probably in NW1

/4 sec. 4, T. 24 N., R. 119 W., 

Cokeville quadrangle, Lincoln County, Wyo. Collected by G. H. 

Girty, July 22, 1909. Foraminifers in the specimen are Mamet 

Zones 18-20. 

LOCAUTY DATA FOR USNM SPECIMENS NOT 

HAYASAKA TYPES 

USNM No. Stratigraphic unit Locality 

Acrocyathus floriformis floriformis 

13669 St. Louis Limestone_____ Mt. Pleasant Iowa 

71646 St. Louis Limestone_____ Rainbow Mountain, Ala. 

[equivalent]. 

216206 St. Louis Limestone_____ Livingston, Tenn. 

Acrocyathus floriformis hemisphaericus 

8211 St. Louis Limestone_____ Monroe County, Ind. 

98102 St. Louis Limestone_____ 1.25 mi (2 km) north of 

~quivalent]. Cleveland, Va. 
42 

USNM No. 

756 

3779 

15526 

17071 

17848 

37466 

39654 

42695 

42766 

135092 

135094 

135096 

135097 

135172 

135173 

135174 

135177 

135179 

135300 

135301 

135302 

136704 

166604 

166605 

216198 

216199 

216200 

216202 

216203 

216204 

216205 

216207 

216208 

216210 

216211 

216212 

216214 

841 

4587 

37469 

37470 

37471 

39655 

42667 

42705 

42714 

REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

LOCALITY DATA FOR USNM SPECIMENS NOT 

HA YASAKA TYPES -Continued 

Stratigraphic unit Locality 

Acrocyathus floriformis subspecies undet. 

St. Louis Limestone _____ St. Francisville, Mo. 

Lower Carboniferous ____ Sugar Creek, Iowa ____ do________________ Near Prairie du Rocher, Ill. ___ do________________ Clear Springs, Harrison 

St. Louis Limestone ____ _ ____ do _______________ _ 

----do _______________ _ ____ do _______________ _ 

____ do _______________ _ ____ do _______________ _ ____ do _______________ _ 

____ do _______________ _ ____ do _______________ _ ____ do _______________ _ 

____ do _______________ _ 

County, Ind. 

Kentucky 

St. Louis, Mo. 

Elizabethtown, Ky. 

Eddyville, Ky. 

Clarksville, Tenn. 

Monroe County, Ind. 

Weaver's Branch of Des 

Moines River, near St. Francisville, Clark 

County, Mo. 

Livingston, Tenn. 

Do. 

Do. 

Hillside north of South 

Fork of Holston River, 

5 mi (8 km) north 

of Kingsport, Tenn. 

Mississippian __________ Millpoint, Pocahontas 

County, W.Va. 

St. Louis Limestone _____ Iowa ____ do________________ Horse and Muddy Creeks, 

Dade County, Mo. 

Mississippian __________ Elgin, Ill 

____ do________________ Caney Fork River, Dekalb 

St. Louis Limestone ____ _ ____ do _______________ _ 

Mississippian _________ _ ____ do _______________ _ 

St. Louis Limestone ____ _ 

County, Tenn. 

Livingston, Tenn. 

Do. 

Franklin, Ky. 

Do. 

1 mi (1.6 km) west of Red 

Hill Post Office, 

Marshall, County, Ala. ____ do________________ River bluff 1 mi (1.6 km) 

south of Maeystown, Ill. ____ do________________ Do. 

____ do________________ Rock Creek, Hancock 

County, Ill. 

Mississippian __________ Bloomington, Ind. ____ do________________ Slick Rock Creek, Barren 

County, Ky. 

St. Louis Limestone _____ 2 mi (3.2 km) west of 

Ste. Genevieve, Mo. ____ do________________ Livingston, Tenn. ____ do________________ Do. 

Mississippian __________ Near Forbuss, Fentress 

County, Tenn. ____ do________________ About 1 mi (1.6 km) east of 

Little Crab, Standingstone quadrangle, 

Tennessee 

Greenbrier Limestone ___ 0.5 mi (0.8 km) southeast of 

Shrader, Pounding Mill 

quadrangle, Virginia ____ do________________ Millpoint, Pocahontas 

County, W.Va. 

Acrocyathus proliferus 

St. Louis Limestone_____ Hardin County, Ill. 

Mississippian ---------- Gallatin County, Ill. 

St. Louis Limestone _____ Marion, Ky. 

Mississippian __________ Saginaw Bay, Mich. 

____ do---------------- Belleview. Mich. 

St. Louis Limestone_____ Elizabethtown, Ky. ____ do________________ Eddyville, Ky. ____ do________________ Do. 

____ do________________ Clarksville, Tenn. 

LOCALITY DATA FOR USNM SPECIMENS NOT 

HA YASAKA TYPES-Continued 

USNM No. Stratigraphic unit Locality 

Acrocyathus proliferus-Continued 

42845 ____ do ________________ Scott County, Ill. 

49941 ____ do ________________ Clarksville, Tenn. 

49942 ____ do ________________ Eddyville, Ky. 

52681 Mississippian __________ Hardin County, Ky. 

60306 St. Louis Limestone _____ Eddyville, Ky. 

71647 St. Louis Limestone _____ Blount Springs, Bount 

[equivalent]. County, Ala. 

135085 Hillsdale Member of the Opposite Bishop High 

Greenbrier Limestone. School on Virginia 

Highway 16, Tazewell 

County, Va. 

135087 St. Louis Limestone _____ 1 mi (1.6 km) east of 

Rockwell, Ky. 

135091 St. Louis Limestone _____ 2.5 mi (4 km) southeast of 

[equivalent]. Bandy, Pounding Mill 

quadrangle, Virginia 

135095 St. Louis Limestone _____ Eddyville, Ky. 

135098 ____ do ________________ · St. Francisville, Mo. 

135168 ____ do ________________ 0.75 mi (1.2 km) east of 

Columbia, Ill. 

135176 ____ do ________________ Eddyville, Ky. 

216201 ____ do ________________ Maeystown, Ill. 

216207 ____ do ________________ Livingston, Tenn. 

216209 ___ do ________________ do. 

216213 Greenbrier Limestone ___ Right river cut, 1 mi. 

(1.6 km) northeast of · Cedar Bluff, Va. 

239233 St. Louis Limestone _____ 2 mi (3.2 km) northeast of 

Mills Springs, Ky. 

REFERENCES CITED 

Allen, A. T., Jr., and Lester, J. G., 1954, Contributions to the paleontology of northwest Georgia: Georgia Geological Survey Bulletin 

62, 166 p., 42 pls. 

Armstrong, A. K., 1962, Stratigraphy and paleontology of the 

Mississippian System in southwestern New Mexico and adjacent 

southeastern Arizona: New Mexico Bureau of Mines and Mineral 

Resources Memoir 8, 99 p., 12 pls., 41 figs. 

--1970a, Mississippian rugose corals, Peratrovich Formation, 

west coast, Prince of Wales Island, southeastern Alaska: U.S. 

Geological Survey Professional Paper 534, 44 p., 13 pls., 30 figs. 

--1970b, Carbonate facies and the lithostrotionoid corals of the 

Mississippian Kogruk Formation, DeLong Mountains, northwestern Alaska: U.S. Geological Survey Professional Paper 664, 

38 p., 37 figs., 14 pls. 

--1972a, Biostratigraphy of Mississippian lithostrotionoid corals, 

Lisburne Group, arctic Alaska: U.S. Geological Survey Professional Paper 743-A, 28 p., 9 pls., 25 figs. 

--1972b, Pennsylvanian carbonates, paleoecology, and rugose colonial corals, north flank, eastern Brooks Range, arctic Alaska: 

U.S. Geological Survey Professional Paper 747, 21 p., 8 pls., 16 

figs. 

Bamber, E. W., 1961, Mississippian corals from northeastern British 

Columbia, Canada: Princeton, N.J., Princeton University, Ph.D. 

thesis, 199 p., 14 pls., 1 fig. 

--1966, Type lithostrotionid corals from the Mississippian of 

western Canada: Canada Geological Survey Bulletin 135, 28 p., 4 

pls. 

Bamber, E. W., and Macqueen, R. W., 1979, Upper Carboniferous and 

Permian stratigraphy of the Monkman Pass and southern Pine 

Pass areas, northeastern British Columbia: Canada Geological 

Survey Bulletin 301, 27 p., 40 figs. 
REFERENCES CITED 43 

Barrois, Charles, 1882, Recherches sur les terrains anciens des 

Asturies et de la Galice: Societe Geologique du Nord Memoire, v. 

2, no. 1, 630 p., 20 pis. 

Bassler, R. S., 1950, Faunal lists and descriptions of Paleozoic corals: 

Geological Society of America Memoir 44, 315 p., 20 pis. 

Bolkhovitinova, M. A., 1915, 0 Kamennougol'nykh korallakh i 

mshannakh Moskovskoi gubernii [On· Carboniferous corals and 

Bryozoa of Moscow province]: Obshchestvo Lyubitelei Estestvoznaniya Antropologii i Etnografii, Moscow, Geologicheskoe 

Otdelenie Zapiski, v. 3, p. 61-81, pis. 5, 6. 

Boule, Marcellin, and others, 1906, 1907, Types du Prodrome de 

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44 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

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Page 

A 

Abstract ------------------------------------

Acknowledgements --------------------------

Acrocyathidae --------------------------- 1, 5, 6, 15 

Acrocyathus --------------- 1, 2, 4, 5, 6, 7, 9, 15, 20, 37 

cystosus ------------------------------ 5, 16, 22 

floriformis ------------------- 5, 16, 17, 19, 20, 21 

florijormis _____ 3, 5, 7, 16, 17, 41; pl. 5, 6, 7, 8, 9, 

10, 11, 16 

hemisphaericus ____ 3, 5, 18, 19, 41; pl. 12, 13, 14 

girtyi ------------------------- 3, 5, 16, 21; pl. 17 

grechoykae ---------------------------- 5, 21, 29 

hsujiulingi ----------------------------- 5, 22 

lissitz·ini ----------------------------- 5, 16, 22 

pennsylvanicus ---------------------- 5, 8, 16, 21 pilatus _____________________ 3, 5, 16, 19, 21; pl. 17 

prolijm·us ----------------- 5, 16, 20, 42; pl. 15, 16 

rotai -·-------------------------------- 5, 16,21 

shimeri -------------------------- 5, 8, 16, 22, 23 

unicwr ------------------------------- 5, 16, 22 

utkae -------------------------------- 5, 15, 21 

zhizhinae ----------------------------- 5, 16, 29 

spp ----------------------------------- 5, 17, 

spp. indet ------------------------------- 29 

Additional taxa ----------------------------- 39 

Actinocyathus --------------------------- 1, 4, 6, 97 

(Actinocyathus), Lonsdaleia ----------- 1, 4, 6, 8, 19, 97 

berthiaumi, Lonsdaleia ----------------- 6, 18, 97 

perat.,·ovichensis, Lonsdaleia -------------- 6, 8, 97 

stelcki, Lonsdaleia ----------------------- 8, 37 

Alabama ·------------------------------------ 3 

Alapah Limestone ---------------------------- 3 

Alaska -·------------------------------------ 3 

allisonae, Petalaxis -------------------------- 99 

altaicum, Lithostrotion ----------------------- 11 

Stelechophyllum venukoffi ---------------- 5, 7, 11 

americana, Lithostrotionella ---------- 3, 13, 17, 23, 38; 

pl. 3, 4, 5, 6 

americanum, Lithostrotion [Lithostrotionella] ____ 17, 18 

ascendens, Stelechophyllum ------------------- 5, 6, 10 

Stelechophyllum ascendens ---------------5,10, 39 

simplex--------------------------- 5, 10, 99 

Stylophyllum ---------------------------- · 10 

Thysannphyllum ascendens ---------------- 99 

simplex ----------------------------- 99 

ascendens, Stelechophyllum -------------- 5, 10, 99 

Thysanophyllum ---------------------- 99 

simplex, Stelechophyllum ---------------- 5, 10, 39 

Thysanophyllum ---------------------- 99 

Aspen Range Formation -------------------- 3, 26, 40 

Astraea -·---------~------------------------- 15 

mamillaris ------------------------------ 17 

astraeijorme, Thysanophyllum ---------------- 6, 97 

astraeijormis, Lithostrotionella ---------------- 97 

Lithostrotionella (Thysanophyllum) _____________ 97 

A ulina tubifera ----------------------------- 15 

AulostylWJ ------------------------- 1, 2, 4, 5, 6, 7, 15 

tubiferus ------------------------------- 5, 15 

eotubiferus ------------------------- 3, 5, 15 

tubiferus ------------------------- 3, 5, 7, 15 

sp -----------------------------------5, 15 

awenggouensis, Lithostrotionella --------------- 99 

Axinura canadensis-------------------------- 17 

B 

baijinensis, Lithostrotionella ---------.--------- 99 

INDEX 

[Italic page numbers indicate major references] 

Page 

bailliei, Lithostrotionella --------------------- 27 

Petalaxis ----------------------------- ~i, 25, 27 

basaltiforme, Lithostrotion -------------------·- 17 

banffense, Lithostrotionella ------------------- 19, 39 

Stelechophyllum --------------- 3, 5, 10, 19; pl. 3, 4 

banfjensis, Lithostrotion ---------------------·- 19 

Lithostrotionella ------------------------·- 19 

belinskiensis, Petalaxis ------~-------------- !), 25, 29 

Petalaxis mccoyana ----------------------·- 29 

berthiaumi, Lithostrotion {LithostrotioneUa] ----·- 97 

Lonsdaleia (Actinocyathus) --------- 6, 18, 97; pl. 20 

besti, Petalaxis ------------------------------ 99 

birdi, Lithostrotionella ----------------------- 1.1; 

StelechophyUum ----------------------- 5, 10, 14 brokawi, Lithostrotionella __________________________ 28 

Petalaxis _______________________________ >), 25, 28 

c 

California ---------------------------------·-

canadense, Lithostrotion --------------·------·- 17, 20 

canadensis, Axinura -------------------------- 17 

canadensis, Lithostrotion -------------------·- 17, 19 

cantabrica, LithostrotioneUa (Hillia) ------------- 91 

cantabricus, Petalaxis ---------------------- 5, 25, 91 

CarcinophyUum ---------------------------·-- 8 

castelnaui, Lithostrotion --------------------- 17, 19 

Lithostrotion[Lithostrotionella] ------------ 17, 19 

Lithostrotionella -------------- 3; pl. 7, 8, 9, 10, 14 

celadensis, LithostrotioneUa ------------------- 95 

Pelalaxis ----------------------------- 5, 25, 95 

changshunensis, Lithostrotionella -------·-----·-- 99 

Cherokee Shale ----------------------------·- 40 

China------------------------------·------ 1, 2, 16 

circinatum, Stelechophyllum ---------------- 5, 10, 12 

circinatus, Lithostrotion (LithostrotioneUa) ----·-- 12 

Lithostrotionella ------------------------·-- 12 

Classification -------------------------------- 4 

Clisiophyllum ------------------------------ 8 

Columnariina -------------------------------

confertus, Petalaxis ------------------------ 5, 25, 99 

confluens, Lithostrotionella ------------------- 12 

crassus, Lithostrotionella -------------------·-- 40 

crenulare, Cyathophyllum -------------------- 37 

Cyathophyllum crenulare --------------------- 37 

Cystolonsdaleia ----------------------- 1, 4, 6, 24, 35 

lutugini -------------------------------- 25 

portlocki --------------------------------- 95 

cystosus, Acrocyathus ---------------------- 5, 16, 22 

Eolithostrotionella ---------------·-------- 22 

D 

densiconus, Lonsdaleia portlocki --------·-----·-- 36 

dilatata, Kleopatrina ------------------~----- 6 

Kleopatrina (Kleopatrina) ----------------- 98 

Lithostrotion [Lithostrotionella] ------------ 98 

Diphyphyllum ----------------------------- 16, 37 

dobrolyubovae, Petalaxis --------------·--- 6, 25, 95, 36 

donbassica, Lithostrotion [LithostrotioneUaj _____ 27 

Lithostrotionella -----------------·-------- 27 

donbassicus, Petalaxis ----------------·--- 5, 25, 27, 29 

donetsensis, Petalaxis ----------------·------- 25, 96 

duplicata, Lonsdaleia ----------------·-------- 37 

duplicatus, Erismatolithus Madreporites ·-------- 97 

Durhaminidae ------------------------- 1, 5, 6, 8, 97 

Durhamina ------------------------·-------- 8 

dushanensis, Lithostrotionella ----------------- 99 

Page 

E 

Eastonoides ----------------------------- 1, 4, 5, 24 

elyensis -------------------------------- 95 

elegantula, Lithostrotionella ------------------- 99 

elyensis, Eastonoides ------------------------- 95 

Petalaxis -------------------------- 6, 25, 95, 36 

Endophyllidae ------------------------------ 4 

Endophyllum ------------------------------ 1, 4, 6 Eolithostrotionella ______________________ 1, 4, 5, 9, 16 

cystosus -------------------------------- 22 

grechovkae ------------------------------ 29 

lissitzini -------------------------------- 22 

longisirptata ----------------------------- 12 

rotai ----------------------------------- 21 

utkae ---------------------------------- 21 

zhizhinae ------------------------------- 29 

eotubiferus, Aulostylos tubiferus -------------- 3, 5, 15 

ergunjaicum, Lithostrotionella ----------------- 19 Stelechophyllum ________________________ 5, 10, 19 

Erismatolithus M adreporites ( duplicatus) -------- 97 

M adreporites (floriformis) ----------------- 97 

evidens, Petalaxis ------------------------- 5, 25, 99 

exilis, Petalaxis ------------------------ 5, 25, 92, 33 

exiguus, Petalaxis ------------------3, 5, 25, 28; pl. 19 

F 

Faberophyllum ----------------------------- 40 

flexuosa, Lithostrotionella -------------------- 27, 39 

flexuosum, Lithostrotion ---------------------- 27 

Lithostrotion [Lithostrotionella] ------------ 27 

fiexuosus, Petalaxis ------------5, 25, 27, 29, 30, 32, 34 

floriformius, Acrocyathus ----------5, 16, 17, 19, 20, 21 

Acrocyathusfloriformis ___ 3, 5, 7, 16, 17, 41; pl. 5, 6, 

7, 8, 9, 10, 11, 16 

Lithostrotionella ----------------- 3, 13 17, 18, 20 

Lonsdaleia ------------------------------ 97 floriformis, AcroC?Yathus _____ 3, 5, 7, 16, 17, 41; pl. 5, 

6, 7, 8, 9, 10, 11, 16 

hemisphaericus, Acrocyathus __ 3, 5, 18, 19, 41; pl. 12, 

13, 14 

(florijormis), Erismatolithus M adreporites _______ 97 

jomichevi, Petalaxis ------------------------- 5, 90 

"Formenreihe" method ----------------------- 6 

jomichevi, Petalaxis -------------------------- 25 

jtatateeta, Kleopatrina (Kleopatrina) ------------ 98 

Ptole7naia ------------------------------ 38 

fugimotoi, Lithostrotionella ------------------- 99 

G 

girtyi, Acrocyathus -----------------3, 5, 16, 21; pl. 17 

Lithostrotion [LithostrotioneUa] ------------ 28 

Lithostrotionella _________ 3, 12, 19, 28; pl. 2, 17, 19 

grande, Lithostrotion ------------------------ 11 

Stelechophyllum --------------------- 5, 6, 10, 11 

grandis, Petalaxis --------------------------- 98 

grechovkae, Acrocyathus --------------------5, 21, 29 

Eolithostrotionella ----------------------- 29 

Greenbrier Limestone ------------------------ 3 

Hillsdale Member ------------------------ 3 

Granitville, Utah ---------------------------- 3 

grootae, Petalaxis ------------------------- 5, 25, 90 

H 

hemisphaerica, Lithostrotionella __ 3, 12, 15, 18, 19; pl. 2, 

4, 10, 11 

49 
50 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Page 

hemisphericum, Lithostrotion [Lithostrotionella] ____ 17, 

18, 19 

hemisphaericus, Acrocyathusfloriformis ____ 3, 5, 18, 19, 

41; pl. 12, 13, 14 

Hillia ------------------------------- 1, 4, 5, 25, 39 

minor ---------------------------------- 99 

perapertuensis --------------------------- 99 

(Hillia), Lithostrotionella --------------------- 24 

cantabrica, Lithostrotionella --------------- 91 

intermedia, Lithostrotionella --------------- 91 

perapertuensis, Lithostrotionella ----------- 90 

radians, Lithostrotionella ----------------- 91 

santaemariae, Lithostrotionella ------------ 91 

wagneri, Lithostrotionella ----------------- 90 

Hillsdale Member, Greenbrier Limestone -------- 9 

hsujiulingi, Acrocyathus --------------------- 5, 22 

Lithostrotionella ------------------------ 22, 39 

Humbug Formation-------------------------- 41 

Idaho ------------------------------------- 3, 40 

Illinois ---------------------------------- 3, 20, 40 

immanis, PetaUu;is ------------------------ 5, 25, 29 

inconfertus, PetaUu;is ------------------------ 98 

indigae, PetaUu;is --------------------------- 98 

intermedia, Lithostrotionella (Hillia) ----------- 91 

intermedius, PetaUu;is ----------------------- 25, 91 

Introduction -------------------------------- 1 

ivanavi, Lithostrotionella --------------------- 99 

Lonsdaleia ---------------------------25,25,26 

PetaUu;is ----------------------------- 6, 25, 96 

J 

jasperensis, Lithostrotion {Lithostrotionella] _____ 12 

Lithostrotionella ------------------------- 12 

jiaruoensis, Lithostrotionella ------------------ 99 

K 

kennedyi, PetaUu;is -------------------------- 99 

Kentucky ---------------------------------- 3, 40 

kitakamiensis, Lithostrotionella --------------- 96 

Kleopatrina ----------------------- 1, 2, 4, 5, 6, 8, 97 

dilatata -------------------------------- 6 

uralica --------------------------------- 6 

wahooensis ------------------------------ 6 

(Kleopatrina) ----------------------------- 98 

dilatata ----------------------------- 98 

ftatateeta ---------------------------- 98 

uralica ------------------------------ 98 

wahooensis -------------------------- 98 

(Porfierevella) --------------------------- 38 

(Kleopatrina), Kleopatrina -------------------- 99 

dilatata, Kleopatrina --------------------- 98 

ftatateeta, Kleopatrina -------------------- 98 

uralica, Kleopatrina ---------------------- 98 

wahooensis, Kleopatrina ------------------- 98 

korkhovae, PetaUuJis ----------------------- 5, 25, 99 

kuechouensis, Lithostrotionella ---------------- 98, 39 

magna, Lithostrotionella ------------------ 99 

Kuechow Province, China --------------------- 2 

kunthi, PetaUu;is ---------------------------- 98 

Kuzn~tsk Basin ----------------------------- 2 

L 

lisitschanskensis, PetaUuJis ----------------- 5, 25, 92 

PetaUuJis vesiculosa ---------------------- 92 

lissitzini, Acrocyathus --------------------- 5, 16, 22 

Eolithostrotionella ----------------------- 22 

Lithostrotion ----------------- 1, 2, 4, 9, 16, 20, 23, 24 

altaicum -------------------------------- 11 

banffensis ------------------------------- 19 

basaltijorme ------------------------------ 17 

canadense ------'------------------------- 17, 20 

canadensis ----------------------------- 17, 19 

castelnaui ------------------------------ 17, 19 

jlexuosum ------------------------------- 27 

Page 

grande --------------------------------- 11 

M ac-Coyanum --------------------------- 94 

maccoyanum ---------------------------- 94 

macouni -------------------------------- 15 

mamillare --------------------------- 17, 19, 20 

microstylum ---------------------------- 11 

pennsylvanicum ------------------------- 21, 22 

Portlocki ------------------------------- 94 

proliferum ------------------------------ 20 

styUu;is --------------------------------- 94 

sp ------------------------------------- 14 

(Lithostrotion) --------------------------- 15 

(Lithostrotionella) ----------------------- 16, 23 

circinatus --------------------------- 12 

occidentalis -------------------------- 92 

unicum ----------------------------- 2, 22 

[Lithostrotionella] ------------------------ 9 

americanum ------------------------ 17, 18 

berthiaumi -------------------------- 97 

castelnaui --------------------------- 17, 19 

dilatata ----------------------------- 98 

donbassica --------------------------- 27 

flexuosum --------------------------- 27 

girtyi ------------------------------- 28 hemisphericum ____________________ 17, 18, 19 

jasperensis -------------------------- 12 

micra------------------------------- 19 

mokomokokensis ---------------------- 28 

occidentale --------------------------- 92 

pennsylvanicum ---------------------- 21 

proliferum --------------------------- 20 

simplex ----------------------------- 26 

styUu;is ----------------------------- 94 

tabulatum --------------------------- 26 

tubijera ----------------------------- 15 

vesiculosa --------------------------- 92 

[Thysanqphyllum] --------------------- 9 

mclareni ------------------------- 14 

(PetaUu;is) ------------------------------ 29 

(Lithostrotion), Lithostrotion ------------------ 15 

Lithostrotionella -------- 1, 2, 4, 5, 8, 9, 15, 16, 22, 29, 39 

americana------------ 3, 13, 17, 23, 38; pl. 3, 4, 5, 6 

astraeiformis ---------------------------- 97 

awenggouensis --------------------------- 99 

baijinensis ------------------------------ 99 

bailliei --------------------------------- 27 

banffense ------------------------------- 19 

banffensis ------------------------------- 19 

birdi ----------------------------------- 14 

brokawi -------------------------------- 28 

castelnaui -------------------- 3; pl. 7, 8, 9, 10, 14 

celadensis ------------------------------- 95 

changshuensis --------------------------- 99 

circinatus ------------------------------ 12 

confluens ------------------------------- 12 

crassus --------------------------------- 40 

donbassica ------------------------------ 27 

dushanensis ----------------------------- 99 

elegantula ------------------------------ 99 

ergunjaicum ---------------------------- 19 

flexuosa ----------------------------- 27, 38, 39 

flo'riformis ----------------------3, 19, 17, 18, 20 

fugimotoi ------------------------------- 99 

girtyi -------------------3, 12, 19, 28; pl. 2, 17, 19 

hemisphaerica ______ 3, 12, 15, 18, 19; pl. 2, 4, 10, 11 

hsujiulingi ----------------------------- 22, 39 

ivanovi --------------------------------- 99 

jasperensis ------------------------------ 12 

jiaruoensis ------------------------------ 99 

kitakamiensis --------------------------- 96 

kuechouensis ---------------------------- 98, 39 

magna -----------·------------------- 99 

lochmanae ------------------------------ 19 

maccoyana --------------------------- 90, 34, 39 

major ------------------------.------- 29 

macouni -------------------------------- 15 

mclareni -------------------------------- 14 

micra --------------------------------- 19, 39 

microstyla ------------------------------ 11 

microstylum ---------------------------- 12 

Page 

mohikana ------------------------------- 94 

monocyclica ----------------------------- 28 

mui ------------------------------------ 99 

multiradiata ------------------------ 3, 12; pl. 1 

multivesiculata -------------------------- 99 

niakensis ------------------------------- 14 

orboensis ------------------------------- 91, 39 

regularis ---------------------------- 99 

pennsylvanica ------------------------ 21, 23, 40 

pennsylvanicum ------------------------- 21 

peratrovichensis ------------------------- 97 

pinguis --------------------------------- 99 

prolijera -------------------------------- 20 

sexangula ------------------------------- 99 

sexangulus ------------------------------ 29 

shimeri -------------------------------- 29, 40 

simplex---------------------------- 3, 26; pl. 18 

spiniformis ----------------------------- 96, 39 

styUu;is --------------------------- 94, 36, 38, 39 

uralica ------------------------------ 99 

tabulata -------------------------------- 3, 26 

taishakuensis ---------------------------- 29 

tingi -------------------------------- 96, 38, 39 

tschucotica ------------------------------ 27 

tubijera -------------------------------- 3 

unica ---------------------------------- 22 

unicum -------------------------------- 2, 22 

uralica --------------------------------- 98 

vesicularis ------------------------------ 3, 19 

vesiculosa ------------------------------- 92 

vesiculosa --------------------------- 99 

wahooensis ------------------------------ 98 

sp ------------------------------- 15, 21, 28, 39 

sp. A ----------------------------------- 14 

sp. B ----------------------------------- 14 

sp. indet -------------------------------- 96 

sp. undet ------------------------------- 99 

(Hillia) --------------------------------- 24 

cantabrica--------------------------- 91 

intermedia -------------------------- 91 

perapertuensis ----------------------- 90 

radians ----------------------------- 91 

santaemariae ------------------------ 91 

wagneri ----------------------------- 90 

(Thysanophyllum) astraeijormis ------------ 97 

[Lithostrotionella], Lithostrotion ---------------

americum, Lithostrotion ------------------ 17, 18 

berthiaumi, Lithostrotion ---~-------------- 97 

castelnaui, Lithostrotion ------------------ 17, 19 

dilatata, Lithostrotion -------------------- 98 

donbassica, Lithostrotion ------------------ 27 

flexuosum, Lithostrotion ------------------- 27 

girtyi, Lithostrotion ---------------------- 28 

hemisphericum, Lithostrotion ___________ 17, 18, 19 

jasperensis, Lithostrotion ------------------ 12 

micra, Lithostrotion ---------------------- 19 

mokomokensis, Lithostrotion --------------- 28 

occidentale, Lithostrotion ------------------ 92 

pennsylvanicum, Lithostrotion ------------- 21 

proliferum, Lithostrotion ------------------ 20 

simplex, Lithostrotion -------------------- 26 

styUu;is, Lithostrotion -------------------- 94 

tabulatum, Lithostrotion ------------------ 26 

tubifera, Lithostrotion ---------------'------ 15 

vesiculosa, Lithostrotion ------------------- 92 

[Thysanophyllum], Lithostrotion ------------ 9 

mclareni, Lithostrotion ---------------- 14 

(Lithostrotionella), Lithostrotion -------------- 16, 23 

circinatus, Lithostrotion ------------------ 12 

occidentalis, Lithostrotion ----------------- 92 

unicum, 'Lithostrotion -------------------- 2, 22 Lithostrotionidae __________________________ 1, 4, 5, 6 

Little Flat Formation ---------------- 3, 13, 21, 26, 41 

Locality data, USNM specimens not Hayasaka 

types---------------------------- 41 

lochmanae, Lithostrotionella ------------------ 19 

Stelechophyllum ----------------------- 5, 10, 19 

Lodgepole Limestone ------------------------ 3, 40 

longiseptata, Eolithostrotionella --------------- 12 

Lonsdalia ------------------------------- 12 
Page 

longiseptatum, Stelechophyllum -------------- 5, 10, 12 

Lonsdaleia ---------------- 1, 2, 4, 5, 6, 8, 16, 20, 24, 97 

duplicata ------------------------------- 37 

jloriformis ------------------------------ 97 

ivanavi ------------------------------25, 35, 96 

pennsylvanica --------------------------- 21 

portlocki -------------------------~----- 25, 95 

densiconus -------------------------- 36 

shimeri ---------~----------------------- 22 

(Actinocyathus) ------------------------8, 19, 97 

berthiaumi --------------------- 18, 97; pl. 20 

peratrovichensis --------------------- 6, 8, 97 

stelcki ------------------------------ 8, 37 

(Lonsctaleia) ----------------------------- 97 

(Lonsdale·ia), Lonsdaleia ---------------------- 97 

Lonsdaleiidae ---------------------------- 4, 5, 6, 96 

Lonsdalia. ----------------------------------

longiseptata ----------------------------- 12 

lutugini, Cystolonsdaleia --------------------- 25 

M 

maccoyana, Lithostrotionella --------------- 90, 34, 39 

Petalaxis ------------------------------- 90, 34 

major, Lithostrotionella ------------------- 29 

multi:Jeptata, Petalaxis -------------------- 96 

orlovkensis, Petalaxis --------------------- 96 

Mac-Coyanum, Lithostrotion ------------------ 94 

maccoyanum, Lithostrotion ------------------- 94 

maccoyanus, Petalaxis ___ 2, 5, 7, 24, 25, 27, 30, 94, 35, 36 

macouni, Lithostrotion ----------------------- 15 

Litho3trotionella ------------------------- 15 

Stelechophyllum ----------------------- 5, 10, 15 

Madison Limestone -------------------------- 3 Madreparites (duplicatus), Erismatolithus _______ 97 

(floriformis), Erismatolithus --------------- 97 

magna, Lithostrotionella kuechouensis ---------- 99 

major, Lithostrotionella maccoyana ------------ 29 

Petalaxis ----------------------------- 5, 25, 29 

mamillare, Lithostrotion ------------------ 17, 19, 20 

mamillaris, Astraea ------------------------- 17 McCloud Limestone 3 mccoyana, Petalaxis ·_·_·_·_·_·_·_·_·_·_·_·_·_·_·_·_·_·_·_·_·_·~~-·-· 29, 94 

belinskiensis, Petalaxis -------------------- 29 

mccoyanus, Petalaxis ------------------------ 90 

mclareni, Lithostrotion [Lithostrotionella] 

[Thysanophyllum] ----------------- 14 

Lithostrotionella ------------------------- 14 

Stelechophyllum ----------------------- 5, 10, 14 

M'coyana, Petalaxis ------------------------- 24, 94 

Stylaxis -------------------------------- 24, 94 

megalum, Stelechophyllum ---------------- 5, 6, 10, 11 

Stylophyllum ---------------------------- 11 

Meridian Range, Mont ----------------------- 3 micra, Lithostrotion [Lithostrotionellaj _________ 19 

Lithostrotionella ------------------------ 19, 19 

microstyla, Lithostrotionella ------------------ 11 

microsty/.um, Lithostrotion -------------------- 11 

Lithostrotionella ------------------------- 12 

Stelechophyllum --------------- 3, 5, 10, 11; pl. 1, 2 

micrum, Stelechophyllum ------------------- 5, 10, 19 

minor, Hillia ------------------------------- 99 

mirus, Petalaxis -------------------------- 5, 25, 99 

Mission Canyon Limestone -------------------- 3 

Missouri ----------------------------------- 3, 40 

mohikarm, Lithostrotionella ------------------- 94 

Petalaxis ------------------------------- 94 

mohikanus, Petalaxis ---------------------- 5, 25, 94 

mokomokensis, Lithostrotion[Lithostrotionella] ___ 28 

Petalaxis ----------------------------- 5, 25, 28 

monocyclica, Lithostrotionella ----------------- 28 

monocyc/.icus, Petalaxis --------------------- 5, 25, 28 

Moscow Basin ------------------------------ 2 

mui, Lithostrotionella ------------------------ 99 

multiradiata, Lithostrotionella -------------3, 12; pl. 1 

multiseptata, Petalaxis maccoyana ------------- 96 

multivesiculata, Lithostrotionella -------------- 99 

N 

Namurian --------------------------------

INDEX 

Page 

Nematophyllum ----------------------------- 24 

Newman Limestone------------------------- 3, 40 

niakense, Stelechophyllum -------------·------- 10, 14 

naikensis, Lithostrotionella -------------------- 14 

Stelechophyllum ------------------·-----·-- 5 

0 

occidentale, Lithostrotion [Lithostrotionella] _____ 92 

occidentalis, Lithostrotion (Lithostrotionella) _____ 92 

Petalaxis ------------------------ 5, 25, .12; pl. 20 

orboensis, Lithostrotionella ------------------- 91, 39 

Petalaxis ----------------------------- 5, 25, 91 

regularis, Lithostrotionella ---------------- 99 

arientale, Thysanophyllum -------------------- 37 

orlovkensis, Petalaxis maccoyana --------------- 96 

p 

pecki, Petalaxis ----------------------·-------- 99 

pennsylvanica, Lithostrotionella ------------ 21, 23, 40 

Lonsdaleia ------------------------------- 21 

pennsylvanicum, Lithostrotion ---------------- 21, 22 

Lithostrotion [Lithostrotionella] ------------ 21 

Lithostrotionella ------------------------- 21 

pennsylvanicus, Acrocyathus -------------- 5, 8, 16, 21 

perapertuensis, Hillia ------------------------ 99 

Lithostrotionella (Hillia) ------------------ 90 

Petalaxis ----------------------- 5, 25, 90, 31, 32 

Peratrovich Formation ------------------------ 3 

peratrovichensis, Lithostrotionella ------------- 97 

Lonsdaleia (Actinothyathus) -------------- 6, 8, 97 

persubtilis, Petalaxis ------------------------ 5, 25, 99 

Petalaxidae ----------------------------- 1, 4, 5, 29 

Petalaxis _____ 1, 2, 4, 5, 6, 7, 9, 16, 22, 28, 24, 26, 27, 28, 

29, 30, 31, 32, 33, 39 

allisonae -------------------------------- 99 

bailliei ------------------------------- 5, 25, 27 

belinskiensis -------------------------- 5, 25, 29 

besti ----------------------------------- 99 

brokawi ------------------------------ 5, 25, 28 

cantabricus --------------------------- 5, 25, 91 

celadensis ----------------------------- 5, 25, 95 

confertus ----------------------------- 5, 25, 99 

dobrolyubovae --------------------·-- 6, 25, 95, 36 

donbassicus ----------------------·-- 5, 25, 27, 29 

donetsensis ----------------------------·- 25, 96 

elyensis --------------------------- 6, 2:5, 95, 36 

evidens -------------------------------- 5, 25, 99 

exiguus -----------------------3, 5, 25, 28; pl. 19 

exilis ------------------------------ 5, :~5, 92, 33 

jlexuosus ------------------5, 25, !!7, 29, ao, 32, 34 

fomichevi -----------------------·-------- 5, 90 

jomichevi -----------------------------·-- 25 

grandis ---------------------------------- 98 

grootae ------------------------------- 5, 25, 90 

immanis ------------------------------ 5, 25, 29 

inconfertus ------------------------------ 98 

indigae --------------------------------- 98 

intermedius ---------------------------- 25, 91 

ivanovi ------------------------------- 6, 25, 96 

kennedyi ------------------------·--------- 99 

korkhovae ------------------------------ 5, 25, 99 

kunthi -------------------------------------98 

lisitschanskensis -----------------------· 5, 25, 92 

maccoyana ----------------------------- 90, 34 

multiseptata ----------------------·--- 96 

orlovkensis --------------------------- 96 

maccoyanus --------2, 5, 7, 24, 25, 27, 30, 94, 35, 36 

major -------------------------------- 5, 25, 29 

mccoyana ------------------------------ 29, 94 

belinskiensis ------------------------- 29 

mccoyanus ------------------------------- 90 

M'coyana ------------------------------- 24, 94 

mirus -------------------------------- 5, 25, 99 

mohikana -------------------------------- 94 

mohikanus ----------------------·------- 5, 25, 94 

mokomokensis -------------------·------- 5, 25, 28 

monocyclicus --------------------------·- 5, 25, 28 

occidentalis ----------------------5, 25, 92; pl. 20 

51 

Page 

orboensis ----------------------------- 5, 25, 91 

pecki ----------------------------------- 99 . 

perapertuensis -------------------5, 25, 90, 31, 32 

persubtilis ---------------------------- 5, 25, 99 

portlocki ------------------------------- 2, 95 

radians ------------------------------ 5, 25, 91 

santaemariae --------------------------5,25,91 

sexangulus ---------------------------- 5, 25, 29 

sibiricus -------------------------------- 98 

simplex ----------------------- 3, 5, 25, 26; pl. 18 

stylaxis ----------------------------5, 25, 27,94 

sutherlandi ----------------------------- 99 

tabulatus -------------------3, 5, 25, 26, 27; pl. 19 

taishakuensis -------------------------- 5, 25, 29 

timanicus ------------------------------ 96, 98 

tschucoticus ---------------------------- 25, 27 

uchtensis ------------------------------- 98 

vesiculosa lisitschakensis ------------------ 92 

vesiculosus -------------------------5, 25, 30, 92 

wagneri ------------------------ 5, 25, 90, 31, 32 

wyomingensis --------------- 3, 5, 25, 26, 27; pl. 18 

spp ------------------------------------ 6 

spp, indet ------------------------------- 96 

(Petalaxis), Lithostrotion --------------------- 29 

Phylogeny --------------------------------- 6 

pilatus, Acrocyathus ------------- 3, 5, 16, 19, 21; pl. 17 

pinguis, Lithostrotionella --------------------- 99 

Porcupine-Arctic section ---------------------- 3 

(Porfierevella), Kleopatrina ------------------- 98 

portlocki, Cystolonsdaleia --------------------- 95 

Lonsdaleia ----------------------------- 25, 95 

Petalaxis ------------------------------- 2, 95 

Stylaxis -------------------------------- 24, 95 

densicionus, Lonsdaleia ------------------- 36 

Portlocki, Lithostrotion ----------------------- 94 

prolifera, Lithostrotionella -------------------- 20 

proliferum, Lithostrotion --------------------- 20 

Lithostrotion [Lithostrotionella] ____________ 20 

proliferus, Acrocyathus --------- 5, 16, 20, 42; pl. 15, 16 

Protolonsdaleia ----------------------------- 97 

Protolonsdalia ------------------------------

Pseudodorlodotia ----------------------------

Ptolemaia ----------------------------------

ftatateeta -------------------------------

R 

97 

37 

97 

38 

radians, Lithostrotionella (Hillia) -------------- 91 

Petalaxis ----------------------------- 5, 25, 91 

References cited ---------------------------- 42 

Register of USGS localities, Hayasaka (1936) 

Lithostrotionella type specimens _____ 40 

regularis, Lithostrotionella orboensis ----------- 99 

rotai, Acrocyathus -------------------------5, 16,21 

Eolithostrotionella ----------------------- 21 

Rugosa----------------------------------- 1, 5, 6 

s 

St. Louis Limestone ----------------------- 3, 20, 40 

santaemariae, Lithostrotionella (Hillia) --------- 91 

Petalaxis ----------------------------- 5, 25, 91 

Sciophyllum ------------------------------- 14, 33 

sexangula, Lithostrotionella ------------------- 99 

sexanguls, Lithostrotionella ------------------- 29 

Petalaxis ----------------------------- 5, 25, 29 

shimeri, Acrocyathus ------------------5, 8, 16, 22,23 

Lithostrotionella ------------------------ 29, 40 

Lonsdaleia ------------------------------ 22 

sibiricus, Petalaxis -------------------------- 98 

simplex, Lithostrotion [Lithostrotionella] -------- 26 

Lithostrotionella -------------------- 3, 26; pl. 18 

Petalaxis ----------------------3, 5, 25, 26; pl. 18 

Stelechophyllum ascendens --------------- 5, 10, 39 

Siphonodendron ----------------------------- 20 

spiniformis, Lithostrotionella ----------------- 96, 39 

Status of Lithostrotionella -------------------- 2 

stelcki, Lonsdaleia (Actinocyathus) ------------- 8, 37 
52 REVISION OF LITHOSTROTIONELLA FROM THE CARBONIFEROUS AND PERMIAN 

Page 

Stelech!Yphyllum ____ 1, 2, 3, 4, 5, 6, 7, 9, 16, 22, 24, 26, 39 

ascendens ------------------------------ 5, 6, 10 

ascendens ------------------------- 5, 10, 39 

simplex--------------------------- 5, 10, 39 

banffense --------------------- 3, 5, 10, 19; pl. 3, 4 

· birdi --------------------------------- 5, 10, 14 

circinatum ---------------------------- 5, 10, 12 

ergunjaicum -------------------------- 5, 10, 19 

grande----------------------------- 5, 6, 10, 11 

lochmanae ---------------------------- 5, 10, 19 

longiseptatum ------------------------- 5, 10, 12 

macouni ------------------------------ 5, 1,0, 15 

mclareni ----------------------------- 5, 10, 14 

megalum --------------------------- 5, 6, 10, 11 

microstylum ------------------ 3, 5, 10, 11; pl. 1, 2 

micrum ------------------------------ 5, 10, 19 

niakense ------------------------------- 10, 14 

niakensis ------------------------------- 5 

venukoffi ----------------------------- 5, 10, 11 

altaicum --------------------------- 5, 7, 11 

venukoffi ---------------------------- 5, 11 

sp indet ---------------------------- 5, 15; pl. 4 

Stylaxis ------------------------------------ 29 

M'coyana ------------------------------ 24, 94 

portlocki ------------------------------- 24, 95 

Lithostrotion ---------------------------- 94 

Lithostrotion [Lithostrotionella] ------------ 94 

Lithostrotionella ------------------- 94, 36, 38, 39 

Petalaxis -------------------------- 5, 25, 27, 94 

uralica, Lithostrotionella ------------------ 99 

Stylid!Yphyllum ----------------------------- 25, 97 

StyliY[Jhyllum ------------------------------- 9 

ascendens ------------------------------- 10 

megalum ------------------------------- 11 

tolmachevi ------------------------------ 98 

venukoffi -------------------------------- 9, 11 

Sublonsdaleia ------------------------------- 97 

Sublonsdalia ------------------------------- 97 

sutherlandi, Petalaxis ------------------------ 99 

Systematic paleontology ---------------------- 8 

Page. 

T 

tabulata, Lithostrotionella -------------------- 3, 26 

tabulatum, Lithostrotion [Lithostrotionella] ______ 26 

tabulatus, Petalaxis ------------- 3, 5, 25, 26, 27; pl. 19 

taishakuensis, Lithostrotionella ---------------- 29 

Petalaxis ----------------------------- 5, 25, 29 

ThysaniY{Jhyllum ----------- 1, 2, 4, 5, 6, 8, 24, 33, 96, 39 

ascendens ascendens ---------------------- 99 

simplex ----------------------------- 99 

astraeiforme ---------------------------- 6, 97 

orientale ------------------------------- 37 (Thysan!Yphyllum) astraeiformis, Lithostrotionella 97 

[Thysanophyllum], Lithostrotion [Lithostrotionella 9 

mclareni, Lithostrotion[Lithostrotionella] ____ 14 

timanicus, Petalaxis ------------------------- 96, 38 

tingi, Lithostrotionella -------------------- 96, 38, 39 

tolmachevi, Styl!Yphyllum --------------------- 98 

tschucotica, Lithostrotionella ------------------ 27 

tschucoticus, Petalaxis ----------------------- 25, 27 

tubifera, Aulina ----------------------------- 15 

Lithostrotion [Lithostrotionella] ------------ 15 

Lithostrotionella ------------------------- 3 

tubiferus, Aulostylus ------------------------- 5, 15 

Aulostylus tubiferus -------------------3, 5, 7, 15 

eotubiferus, Aulostylus -------------------3, 5, 15 

tubiferus, Aulostylus -------------------3, 4, 7, 15 

Tuscumbia Limestone ------------------------ 3, 40 

u 

uchtensis, Petalaxis --------~----------------- 38 

Undetermined lithostrotionelloid corals --------- 38 

unica, Lithostrotionella ---------------------- 22 

unicum, Lithostrotion (Lithostrotionella) -------- 2, 22 

Lithostrotionella ------------------------- 2, 22 

unicus, Acrocyathus ----------------------- 5, 16, 22 

U.S.S.R. ----------------------------------- 16 

uralica, Kleopatrina ------------------------- 6 

Kleopatrina (Kleopatrina) ----------------- 98 

Page 

Lithostrotionella ------------------------- 98 

stylaxis ----------------------------- 99 

Utah-------------------------------------- 3, 40 

utkae, Acrocyathus ------------------------ 5, 16, 21 

Eolithostrotionella ----------------------- 21 

v 

venukoffi, StelechiY[Jhyllum ------------------ 5, 10, 11 

Stelech!Yphyllum venukoffi ----------------- 5, 11 

Stylophyllum ---------------------------- 9, 11 

altaicum, Stelech!Yphyllum ---------------- 5, 7, 11 

venukoffi, Stelech!Yphyllum ----------------- 5, 11 

vesicularis, Lithostrotionella ------------------ 3, 19 

vesiculas a Lithostrotion [Lithostrotionella] ------- 92 

Lithostrotionella ------------------------- 92 

vesiculosa --------------------------- 99 

lisitchanskensis, Petalaxis ----------------- 92 

vesiculosa, Lithostrotionella ---------------- 99 

vesiculosus, Petalaxis ------------------- 5, 10, 30, 92 

Virginia 3 

w 

wagneri, Lithostrotionella (Hillia) -------------- 90 

Petalaxis ----------------------- 5, 25, 90, 31, 32 

wahooensis, Kleopatrina ---------------------- 6 

Kleopatrina (Kleopatrina) ----------------- 98 

Lithostrotionella ------------------------- 98 

Weber Canyon, Utah------------------------- 3 

"Wells" Formation------------------------- 3, 26,41 

West Virginia------------------------------- 3 

Wyoming ---------------------------------- 3 wyomingensis, Petalaxis __________ 3, 5, 25, 26, 27; pl. 18 

z 

Zaphrentis --------------------------------- 6 

zhizhinae, Acrocyathus --------------------- 5, 16, 29 

Eolithostrotionella ----------------------- 23 
PLATES 1-20 

Contact photographs of the plates in this report are available, at cost, from U.S. Geological Survey Library, 

Federal Center, Denver, Colorado 80225 
PLATE 1 

[All figures x 4] 

FIGURES 1, 2. Stelechophyllum microstylum (White) (p. 11). 

USNM 120244 (holotype of Lithostrotionella multiradiata Hayasaka). USGS loc. 490-PC, Lodgepole Limestone, Utah. 

1. Longitudinal thin section, USNM 120244b. 

2. Transverse thin section, USNM 120244a. 

3, 4. Stelechophyllum microstylum (White) (p. 11). 

USNM 162005 (paratype of Lithostrotionella multiradiata Hayasaka). USGS loc. 104-PC, Lodgepole Limestone, Idaho. 

3. Transverse thin section, USNM 162005a. 

4. Longitudinal thin section, USNM 162005b. 
PROF'ESSIOt-;AL PAPER 1247 PLATE 1 

2 

3 4 

STELECHOPHYLLUM MICROSTYLUM (WHITE) 
PLATE 2 

[All figures x 4] 

FIGURES 1, 2. Stelechophyllum microstylum (White) p. 11). 

USNM 161996 (paratype of Lithostrotionella hemisphaerica Hayasaka). USGS loc. 1439-PC, Lodgepole Limestone, Idaho. 

1. Transverse thin section, USNM 161996a. 

2. Longitudinal thin section, USNM 161996b. 

3. 4. Stelechophyllum microstylum (White)? (p. 11). 

USNM 162003 (paratype of Lithostrotionella girtyi Hayasaka). USGS loc. 3864-PC, Lodgepole Limestone, Montana. 

3. Transverse thin section, USNM 162003a. 

4. Longitudinal thin section, USNM 162003c. 
GEOLO<:ICAL Sl. lt\.I<:Y PROFESSIONAL PAPER 1247 PLATE 2 

2 

1 

3 4 

STELECHOPHYLLUM MICROSTYLUM (WHITE) 
PLATE 3 

[All figures x 4] 

FIGURES 1, 2. Stelechophyllum banffense (Warren)? (p. 13). 

USNM 174372 (paratype of Lithostrotionella americana Hayasaka). USGS Loc. 7130B-PC, Alapah Limestone(?), Alaska. 

1. Transverse thin section, USNM 17 4372a. 

2. Longitudinal thin section, USNM 17 4372b. 

3, 4. Stelechophyllum banffense (Warren)? (p. 13). 

USNM 17 437 4 (para type of Lithostrotionella americana Hayasaka). USGS Loc. 970-PC, Alapah Limestone(?), Alaska. 

3. Transverse thin section, USNM 17 437 4a. 

4. Longitudinal thin section, USNM 17 437 4b. 
CEOL()(;[('i\L Sl"R\"EY PROFESSIO:-.IAL PAPER 1247 PLATE 

2 

3 4 

STELECHOPHYLLUM BANFFENSE (WARREN)? 
PLATE 4 

[All figures x 4) 

FIGURES 1, 2. Stelechophyllum banffense (Warren)? (p. 13). 

USNM 174375 (paratype Lithostrotionella americana Hayasaka). USGS Loc. 3024-PC, Little Flat Formation, Idaho. 

1. Transverse thin section, USNM 174375a. 

2. Longitudinal thin section, USNM 17 4375c. 

3, 4. Stelechophyllum sp. indet. (p. 15). 

USNM 120239 (paratype of Lithostrotionella hemisphaerica Hayasaka). USGS Loc. 5892-PC, Madison Limestone, Utah. 

3. Longitudinal thin section, USNM 120239c. 

4. Transverse thin section, USNM 120239a. 
PROFESSIONAL PAPER 1247 PLATE .J 

2 

1 

3 4 

STELECHOPHYLLUM BANFFENSE (WARREN)? AND STELECHOPHYLLUM SP. INDET. 
PLATE 5 

[All figures x 4] 

FIGURES 1, 2. Acrocyathus floriformis floriformis d'Orbigny (p. 17). 

USNM 120240 (holotype of Lithostrotionella americana Hayasaka). USGS Loc. 2333-PC, St. Louis Limestone, Kentucky. 

1. Transverse thin section, USNM 120240a. 

2. Longitudinal thin section, USNM 120240b. 

3, 4. Acrocyathus floriformis jloriformis d'Orbigny (p. 17). 

USNM 174376 (paratype of Lithostrotionella americana Hayasaka). USGS Loc. 499-PC, St. Louis Limestone, Illinois. 

3. Longitudinal thin section (cut slightly off axis of corallite), USNM 174376b.· 

4. Transverse thin section, USNM 17 4376a. 
GEOLOC ICAL SURVEY 

3 

PROFESS IONAL PAPER 1247 PLATE 5 

4 

ACROCYATH US FLORIFORMIS FLORIFORMIS D'ORBIGNY 

2 
PLATE 6 

[All figures x 4) 

FIGURES 1, 2. Acrocyathusflorijormisflorijormis d'Orbigny? (p. 17). 

USNM 162001 (paratype of Lithostrotionella americana Hayasaka). USGS Loc. 498-PC, St. Louis Limestone, Illinois. 

1. Transverse thin section, USNM 162001a. 

2. Longitudinal thin section, USNM 162001b. 

3,4. Acrocyathus floriformis floriformis d'Orbigny? (p. 17). 

USNM 120241 (paratype of Lithostrotionella americana Hayasaka). USGS Loc. 1211B-PC, St. Louis Limestone, Missouri. 

Thysanophylloid variants in a polymorphic corallum. 

3. Longitudinal thin section, USNM 120241b. 

4. Transverse thin section, USNM 120241a. 
GEOLOGICAL SURVEY 

3 

PROFESSlO::\AL PAPER 1247 PLATE 6 

4 

ACROCYATHUS FLORIFORMIS FLORIFORMIS D'ORBIGNY? 

2 
PLATE 7 

[All figures x 4) 

FIGURES 1, 2. Acrocyathu.s floriformis jloriformis d'Orbigny? (p. 17). 

USNM 174377 (paratype of Lithostrotionella castelnaui Hayasaka). USGS 3946-PC (green), unknown locality, probably St. 

Louis Limestone, Mississippi Valley region. 

1. Transverse thin section, USNM 174377a. 

2. Longitudinal thin section (cut slightly off axis of corallite), USNM 174377b. 

3, 4. Acrocyathu.s floriformis floriformis d'Orbigny (p. 17). 

USNM 120235 (holotype of Lithostrotionella castelnaui Hayasaka). USGS Loc. 499-PC, St. Louis Limestone, Illinois. 

3. Longitudinal thin section, USNM 120235c. 

4. Transverse thin section, USNM 120235a. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1247 PLATE 

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ACROCYATHUS FLORIFORMIS FLORIFORMIS D'ORBIGNY? AND 

ACROCYATHUS FLORIFORMJS FLORIFORMIS D'ORBIGNY 

2 
PLATE 8 

[All figures x 4) 

FIGURES 1, 2. Acrocyathus floriformis floriformis d'Orbigny (p. 17). 

USNM 161991 (paratype of Lithostrotionella castelnaui Hayasaka). USGS Loc. 643-PC, St. Louis Limestone(?), Missouri. 

1. Transverse thin section, USNM 161991a. 

2. Longitudinal thin section, USNM 161991b. 

3, 4. Acrocyathus floriformis floriformis d'Orbigny (p. 17). 

USNM 161990 (paratype of Lithostrotionella castelnaui Hayasaka). USGS Loc. 346C-PC, residual chert from Tuscumbia 

Limestone, Alabama. 

3. Longitudinal thin section, USNM 161990b. 

4. Transverse thin section, USNM 161990a. 
GEOLOGICAL S URVEY l'IWFESS!Ol'\A L PAPE R 1247 PLATE 8 

3 4 

ACROCY ATHUS FLORIFORMIS FLORIFORMIS D'ORBIGNY 
PLATE 9 

[All figures x 4) 

FIGURES 1, 2. Acrocyathusfloriformisjloriformis d'Orbigny (p. 17). 

USNM 161994 (paratype of Lithostrotionella castelnaui Hayasaka). USGS 2013B-PC, Greenbrier Limestone, Virginia. 

1. Transverse thin section, USNM 161994a. 

2. Longitudinal thin section, USNM 161994b. 

3, 4. Acrocyathus jloriformis jlorijormis d'Orbigny (p. 17). 

USNM 161989 (paratype of Lithostrotionella castelnaui Hayasaka). USGS Loc. 2226-PC, St. Louis Limestone, Illinois. 

3. Transverse thin section, USNM 161989a. 

4. Longitudinal thin section, USNM 161989c. 
GEOLOG ICAL SURVEY PROFESS!Ol'\AL PAPER 1247 PLATE 9 

2 

3 4 

ACROCYATHUS FLORIFORMIS FLORIFORMIS D'ORBIGNY 
PLATE 10 

[All figures x 4] 

FIGURES 1, 2. Acrocyathusfloriformisfloriformis d'Orbigny (p. 17). 

USNM 161997 (paratype of Lithostrotionella hemisphaerica Hayasaka). USGS Loc. 2020-PC, Greenbrier Limestone, 

Virginia. 

1. Transverse thin section, USNM 161997a. 

2. Longitudinal thin section, USNM 161997b. 

3, 4. Acrocyathus floriformis floriformis d'Orbigny (p. 17). 

USNM 161993 (paratype of Lithostrotionella castelnaui Hayasaka). USGS Loc. 3159-PC, Newman Limestone, Virginia. 

3. Longitudinal thin section, USNM 161993b. 

4. Transverse thin section, USNM 161993a. 
GEOLOG ICA L SURVEY PIWFESSJ<lJ 'AI. l'i\PEH 1~·!7 !'LATE 10 

3 4 

ACROCYATHUS FLORIFORMIS FLORIFORMIS D'ORBIGNY 
PLATE 11 

[All figures x 4] 

FIGURES 1, 2. Acrocyathusjloriformisjloriformus d'Orbigny (p. 17). 

USNM 161995 (paratype of Lithostrotionella hemisphaerica Hayasaka). USGS Loc. 83-PC, Newman Limestone, Kentucky. 

1. Transverse thin section, USNM 161995a. 

2. Longitudinal thin section, USNM 161995b. 

3, 4. Acrocyathus jloriformis jloriformis d'Orbigny (p. 17). 

USNM 120238 (paratype of Lithostrotionella hemisphaerica Hayasaka). USGS Loc. 3283-PC, Hillsdale Member of Greenbrier Limestone, West Virginia. 

3. Longitudinal thin section, USNM 120238d. 

4. Transverse thin section, USNM 120238b. 
GEOLOGICA l " SURVEY . 

/ 

1 

ACROCYATHUS FLORIFORMIS FLORI FORM IS D'ORBIGNY 
PLATE 12 

[All figures x 4] 

FIGURES 1, 2. Acrocyathus jlorijormis hemisphaericus (Hayasaka) (p. 19). 

Holotype, USNM 120237. USGS Loc. 1148-PC, St. Louis Limestone, Illinois. 

1. Transverse thin section, USNM 120237a. 

2. Longitudinal thin section, USNM 120237c. 

3, 4. Acrocyathus jloriformis hemisphaericus (Hayasaka) (p. 19). 

Paratype, USNM 161998. USGS Loc. 932A-PC, St. Louis Limestone, Missouri. 

3. Longitudinal thin section, USNM 161998b. 

4. Transverse thin section, USNM 161998a. 
GEOLOGICAL SURVEY PROFESSIO:-.IAL PAPER 1247 PLATE 12 

3 4 

ACROCYATHUS FLORIFORMIS HEMISP HAERICUS (HA YASAKA) 
PLATE 13 

[All figures x 4] 

FIGURES 1, 2. Acrocyathus jloriformis hemisphaericus (Hayasaka) (p. 19). 

Paratype, USNM 161999. USGS Loc. 2222A-PC, St. Louis Limestone, Illinois. 

1. Transverse thin section, USNM 161999a. 

2, 3. Longitudinal thin section, USNM 161999b. 

4-6. Acrocyathusjloriformis hemisphaericus (Hayasaka)? (p. 19). 

Paratype, USNM 162000. USGS Loc. 2222C-PC, St. Louis Limestone, Illinois. 

4, 5. Longitudinal thin section, USNM 162000b. 

6. Transverse thin section, USNM 162000a. 
GEOLOGICAL SURVEY 

4 

5 

PROFESSIONAL PAPER 1247 PLATE 1:~ 

1 

6 

ACROCYATHUS FLORIFORMIS HEMISPHAEHICUS (HAYASAKA) AND 

ACROCYATHUS FLORIFORMIS HEMISPHAERICUS (HAYASAKA)? 

2 

3 
PLATE 14 

[All figures x 4] 

FIGURES 1, 2. Acrocyathusfloriformis hemisphaericus (Hayasaka) (p. 19). 

USNM 161992 (paratype of Lithostrotionella castelnaui Hayasaka). USGS Loc. 2020A-PC, Greenbrier Limestone, Virginia. 

1. Transverse thin section, USNM 161992a. 

2. Longitudinal thin section, USNM 161992b. 

3, 4. Acrocyathus floriformis hemisphaericus (Hayasaka) (p. 19). 

USNM 120236 (paratype of Lithostrotionella castelnaui Hayasaka). USGS Loc. 3282-PC, Hillsdale Member of Greenbrier 

Limestone, West Virginia. 

3. Longitudinal thin section, USNM 120236b. 

4. Transverse thin section, USNM 120236a. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1247 PLATE 14 

1 

3 4 

ACROCYATHUS FLORIFORMIS HEMISPHAERICUS (HAYASAKA) 
PLATE 15 

FIGURES 1-3. Acrocyathus proliferus (Hall) (p. 20). 

Neotype, USNM 841. St. Louis Limestone, Illinois. 

1. Transverse thin section, x 4, USNM 841a. 

2. Longitudinal thin section, x 4, USNM 841b. 

3. Side view of corallum, x 1. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1247 PLATE 15 

ACROCYATHUS PROLIFER US (HALL) 
PLATE 16 

FIGURES 1. Acrocyathus proliferus (Hall) (p. 20). 

USNM 216207. St. Louis Limestone, Tennessee. 

Transverse thin section, x 2, USNM 216207a, showing crowding of corallites by profuse budding. 

2. Acrocyathus floriformis floriformis d'Orbigny (p. 17). 

USNM 161994. Greenbrier Limestone, Virginia. 

Transverse thin section, x 2, USNM 161994a, showing cylindrical tendency of some corallites in a predominantly cerioid 

corallum. 

3, 4. Acrocyathus proliferus (Hall) (p. 20). 

USNM 216209. St. Louis Limestone, Tennessee. _ 

3. Longitudinal thin section, x 1, USNM 216209b showing cylindrical corallites in a predominantly fasciculate corallum. 

4. Transverse thin section, x 2, USNM 216209a showing prismatic tendency in crowded corallites in a predominantly fasciculate corallum. 
GEOLOGICAL SURVEY PROFESS!Oi\'AL PAPER 12-17 PLATE 16 

3 

2 4 

ACROCYATHUS PROLIFERUS (HALL) AND ACROCYATHUS FLORIFORMIS FLORIFORMIS D'ORBIGNY 
PLATE 17 

[All figures x 4] 

FIGURES 1, 2. Acrocyathus pilatus n. sp. (p. 19). 

Holotype, USNM 162004 (paratype of Lithostrotionella girtyi Hayasaka). USGS Loc. 499-PC, St. Louis Limestone, Illinois. 

1. Transverse thin section, USNM 162004a. 

2. Longitudinal thin section, USNM 162004b. 

3, 4. Acrocyathus girtyi (Hayasaka) (p. 21). 

Holotype, USNM 120243. USGS Loc. 4801H-PC (green), Little Flat Formation, Utah. 

3. Longitudinal thin section, USNM 120243b. 

4. Transverse thin section, USNM 120243a. 
GEOLOGICAL SURVEY PROFESSlOI'\AL PAPER 1247 PLATE 17 

2 

3 4 

ACROCYATHUS PILATUS N. SP. AND ACROCYATHUS GIRTYI (HAYASAKA) 
• 

PLATE 18 

[All figures x 4] 

FIGURES 1-3. Petalaxis simplex (Hayasaka) (p. 26). 

Holotype, USNM 120249. USGS 5893-PC, Little Flat Formation, Utah. 

1. Transverse thin section, USNM 120249a. 

2, 3. Longitudinal thin section, USNM 120249b. 

4, 5. Petalaxis wyomingensis n. sp. (p. 26). 

Holotype, USNM 120675 (paratype of Lithostrotionella simplex Hayasaka). USGS 7452-PC (green), "Wells" Formation), 

Wyoming. 

4. Longitudinal thin section, USNM 120675b. 

5. Transverse thin section, USNM 120675c. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1247 PLATE 18 

2 

3 

1 

4 5 

PETALAXIS SIMPLEX (HAYASAKA) AND PETALAXIS WYOMINGENSIS N. SP. 
PLATE 19 

FIGURES 1-4. Petalaxis exiguus n. sp. (p. 28). 

Holotype, USNM 162002B (paratype of Lithostrotionella girtyi Hayasaka). USGS Loc. 3856-PC (green), McCloud Limestone, California. 

1. Transverse thin section, x 4, USNM 162002Ba. 

2, 3. Longitudinal thin section, x 4, USNM 162002Bb. 

4. Oblique view of corallum, x 1. 

5-7. Petalaxis tabulatus (Hayasaka) (p. 26). 

Holotype, USNM 120246. USGS Loc. 1476-PC, Aspen Range Formation, Idaho. 

5. Longitudinal thin section, x 4, USNM 120246b. 

6. Longitudinal thin section, x 4, USNM 120246c. 

7. Transverse thin section, x 4, USNM 120246a. 
GEOLOG ICAL SURVEY PROFESSIO!\"A L PAPER 1247 PLATE 19 

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3 

4 

5 

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PETALAXIS EXIGUUS N. SP. AND PETALAXIS TABULATUS (HAYASAKA) 
PLATE 20 

[All figures x 4] 

FIGURES 1, 2. Petalaxis occidentalis (Merriam) (p. 32). 

Holotype, USNM 143440. Coyote Butte Formation, Oregon. 

1. Transverse thin section, USNM 143440c. · 

2. Longitudinal thin section, USNM 143440e. 

3, 4. Lonsdaleia (Actinocyathus) berthiaumi (Merriam) (p. 37). 

Holotype, USNM 132988. Unknown formation, Oregon. 

3. Longitudinal thin section, USNM 132988d. 

4. Transverse thin section, USNM 132988c. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1247 PLATE 20 

3 4 

PETALAXIS OCCIDENTALIS (MERRIAM) AND LONSDALEIA 

( ACTINOCYATHUS) BERTHIAUMI (MERRIAM) 

2 

'