Coelenterata of the Amsden Formation 

(Mississippian and Pennsylvanian) 

of Wyoming 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 848-C 

Coelenterata of the Amsden Formation 

(Mississippian and Pennsylvanian) 

of Wyoming 

By WILLIAM J. SANDO 

THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) 

OF WYOMING 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 848-C 

Descriptions and illustrations of 11 taxa 

of corals and 3 of hydrozoans 

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON 1975 
UNITED STATES DEPARTMENT OF THE INTERIOR 

ROGERS C. B. MORTON, Secretary 

GEOLOGICAL SURVEY 

V. E. McKelvey, Director 

Library of Congress Cataloging in Publication Data 

Sando, William Jasper. 

Coelenterata of the Amsden Formation (Mississippian and Pennsylvanian) of Wyoming. 

(The Amsden Formation (Mississippian and Pennsylvanian) of Wyoming) (Geological Survey professional 

paper ; 848~C) 

Bibliography: p. 

Includes index. 

Supt. of Docs. no.: I 19.16:848-C 

1. Coelenterata, Fossil. 2. Paleontology-Carboniferous. 3. Paleontology-Wyoming. I. Title. II. Series. III. 

Series: United States. Geological Survey. Professional paper ; 848-C. 

QE777.S35 563'.5'09787 74-32116 

For sale by the Superintendent of Documents, U.S. Government Printing Office 

Washington, D.C. 20402 

Stock Number 024-001-02633-3 
CONTENTS 

Page Page 

Abstract ---------------------------------------- G1 

1 

4 

4 

Systematic paleontology-Continued 

Introduction _______________________ -------_ ------ Class Anthozoa-Continued 

Systematic paleontology -------------------------- Genus Caninia Michelin in Gervais, 1840 ---

Genus Turbinatocaninia Dobrolyubova, 1970_ 

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

Genus Pleurosiphonella Chudinova, 1970 ----

Genus Multithecopora Yoh, 1927 -----------

Genus Duncanopora n. gen ----------------

G18 

19 

20 

23 

25 

25 

28 

31 

Class llydrozoa ------------------------------ Genus Chaetetes Fischer von W aldheim in 

Eichwald, 1829 ------------------------

Class Anthozoa ------------------------------ Genus Amplexus Sowerby, 1814 -----------

5 

7 

7 

7 

8 

FIGURE 

Genus Zaphrentites lludson, 1941 ----------

Genus Barytichisma Moore and Jeffords, 1945 

Genus Neokoninckophyllum Fomichev, 1939 _ 10 

Ref,erences cited - _ -------------------------------

Index -------------------------------------------

1. 

2. 

3. 

4. 

5. 

6. 

ILLUSTRATIONS 

[Plates follow index] 

PLATE 1. Chaetetes. 

2. Chaetetiform bryozoans. 

3. Barytichisma amsdenense (Branson and Greger) and "Amplexus" sp. indet. 

4, 5. Neokoninckophyllum hamatilis n. sp. 

6. N eokoninckophyllum inconstans n. sp. 

7. Caninia cf. C. nevadensis (Meek) and Turbinatocaninia? sp. 

8. Lonsdaleia (Actinocyathus) stelcki (Nelson). 

9. Pleurosiphonella drummondi (Nelson) and Multithecopora? amsdenensis 

n. sp. 

10. Duncanopora duncanae n. sp. 

Scatter diagram showing relation between alar diameter and number of major septa in Barytichisma 

Page 

amsdenense (Branson and Greger) ----------------------------------------------------------- C11 

Scatter diagram showing relation between alar diameter and number of major septa in Neokoninckophyllum hamatilis n. sp ---------------------------------------------------------------------- 12 

Relation between alar diameter and distance above tip in the holotype of N eokoninckophyllum hamatilis 

n. sp --------------------------------------------------------------------------------------- 13 

Scatter diagram showing relation between alar diameter and number of major septa in Neokoninckophyllum inconstans n. sp --------------------------------------------------------------------- 16 

Variation in average corallite diameter per colony and average number of major septa per corallite in 

colonies of Lonsdaleia (Actinocyathus) stelcki (Nelson) from beds of Chesterian age in western 

North America ------------------------------------------------------------------------------ 22 

Corallite diameter and frequency in syringoporoid corals from the Amsden Formation --------------- 26 

TABLES 

Page 

TABLE 1. Distribution of described coelenterates in the Amsden Formation of Wyoming ---------------------- C2 

2. Checklist of Amsden coelenterate collections ------------------------------------------------------- 3 

3. Measurements of Chaetetes wyomingensis n. sp. in transverse section -------------------------------- 6 

4. Ontogeny in the lectotype (UM 2663) of Barytichisma amsdenense (Branson and Greger) ------------ 9 

5. Ontogeny in a topotype (USNM 165091) of Barytichisma amsdenense (Branson and Greger) -------- 10 

6. Ontogeny in the holotype ("USNM 165095) of Neokoninckophyllum hamatilis n. sp ------------------- 14 

7. Ontogeny in the holotype (USNM 165109) of Neokoninckophyllum inconstans n. sp ------------------ 17 

8. Ontogeny in a paratype (USNM 165114) of Neokoninckophyllum inconstans n. sp ------------------- 18 

9. Ontogeny in a paratype (USNM 165115) of Neokoninckophyllum inconstans n. sp ------------------- 19 

10. Variation in colonies of Lonsdaleia (Actinocyathus) stelcki (Nelson) from the United States and Canada 23 

III 

THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

COELENTERATA OF THE AMSDEN FORMATION 

(MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

By WILLIAM ] . SANDO 

ABSTRACT 

The coelenterate faunas of the Amsd.en Formation of 

Wyoming consist of 14 species that represent 11 genera distributed among the chaetetid hydrozoans and the rugose and 

tabulate corals. Five new species and one new genus are 

described in this paper. Coelenterates are rare or absent in 

the Darwin Sandstone, Horseshoe Shale, and Ranchester 

Limestone Members of the Amsden, but they form a significant part of the inv·ertebrate fauna of the Moffat Trail 

Limestone Member. Although both Missis·sippian and Pennsylvanian forms ar·e present in the Amsden, the Moffat Trail 

fauna, which is of Late Mississippian (Chesterian) age, is 

the only coelenterate assemblage that is significant for correlation with other strata of similar age. 

INTRODUCTION 

This report is part of a larger work that deals 

with the biostratigraphy of the Amsden Formation 

of Wyoming. The project was undertaken in the 

belief that the long and complex controversy over 

the age and correlation of the Amsden could be resolved by means of a comprehensive restudy of its 

fauna. This report deals only with the coelenterates 

collected from the Amsden Formation. Details on the 

stratigraphy of the Amsden and the occurrence of 

these fossils are given by Sando, Gordon, and Dutro 

( 197 4) . A summary of the occurrence of the coelenterate fossils of the Amsden is given in table 1 in 

terms of stratigraphic units and localities described 

by Sando, Gordon> and Dutro; locality data are 

recorded in table 2. 

Coelenterates form a relatively minor part of 

the Amsden faunas; the entire assemblage of coelenterate fossils described from the Amsden consists of only 14 species allocated to 11 genera. No 

coelenterates have been found in the Darwin Sandstone Member, the lowest unit of the Amsden. The 

Horseshoe Shale Member, a transgressive red siltstone and shale unit that overlies the Darwin, has 

yielded only an indeterminate Chaetetes and five 

species of corals representing four genera from rare, 

thin marine limestone beds at widely scattered localitie-s. Significant elements of this meager fauna 

are Barytichisma amsdenense (Branson and Greger) from beds of Chesterian age (Anthracospirifer 

welleri-shawi Zone, Composita poposiensis Subzone) 

in the Wind River Range and two new species of 

N eokoninckophyllum from the Rawlins hills, where 

the Horse-shoe Member contains beds of Early Pennsylvanian (Morrowan) age in the upper part (Neokoninckophyllum hamatilis Zone). 

The greatest concentration of coelenterates occurs 

in the richly fossiliferous marine limestone of the 

Moffat Trail Limestone Member ( Caninia Zone) in 

western Wyoming which has yielded eight species 

repres·enting eight genera. The Moffat Trail fauna 

is of Late Mississippian (Chesterian) age. The 

chaetetid hydrozoans in this fauna are represented 

by the first true Chaetetes to be described from the 

Mississippian of North America. The rugose corals 

are represented by a zaphrentoid species, two 

cyathopsid species, and a species of Lonsdaleia 

(Actinocyathus). The tabulate corals are represented by three syringoporoid species assigned to 

three genera, one of which is new. 

The Ranchester Limestone Member, a unit mostly 

of Pennsylvanian age that overlies the Horseshoe 

and Moffat Trail Members, has yielded only a single 

species of Chaetetes. 

Cl 
TABLE !.-Distribution of described coelenterates in the Amsden Formation of Wyoming 

[Localities listed by collection numbers used by Sando, Gordon, and Dutro (1974)] 

Horseshoe Shale Member Moffat Trail Limestone Member 

I 

Ql 

.§'"' ...:IGI 

21 ~--------------~------------------------------------4-----------------------------------------------------------------------------~ID~ 

-5QI 

~=:~=: 

Species 2 3 4 5 7 28b 27 36a 36c 36e 38 39 41 48 I 72 73 74 75 76 77 78 79 80 88 90 91 92 93 103 104 105 109 121 122 124 

Class Hydrozoa: 

Chaetetes wyomingensis n. sp -----------------• ______ --- cf. C. ea;imius Moore 

and Jeffords ------

sp. indet ------------ Class Anthozoa: 

"Amplea;us" sp. indet --- Zaphrentites? sp. indet -- Barytichisma amsdenense 

(Branson and Greger)_ Neokoninckophyllum 1 

hamatilis n. sp --------, X X X X X inconstans n. sp ----- --- --- --- --- --- X 

Caninia cf. C. nevadensis 

(Meek) -------------- __ _ Turbinatocaninia? sp ---- Lonsdaleia (Actinocyathus) 

stelcki (Nelson) ------ Pleurosiphonella drummondi (Nelson) ------- Multithecopora? amsdenensis n. sp -------------- Duncanopora duncanae 

n. sp -----------------• --- ---

X --- --- --- --- X X X X 

-- --- --- --- --- --- --- --- ___ ,___ --- --- --- --- --- --- --- X --- --- X ---

-~- -X I ::: --- --- --- X ---

T X X --- X X X --- --- --- --- --- --- --- --- ---

--- --- --- --- --- --- --- --- --- --- --- --- --- X X --- X X X X --- --- --- --- X --- X X X X X ---

X 

---· --- --- --- X --- X --- --- X 

--- ---I --- --- --- --- X X --- --- X 

X --- X X X ---

X --- X X X X 

X --- X --- ---

X --- X ? T ---

X 

X 

X 

X 

X X --- --- X 

X X --- --- X 

X 

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X --- --- X 

T 

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COELENTERATA OF THE AMSDEN FORMATION OF WYOMING 

TABLE 2.-Checklist of Amsden coelenterate collection 

Name of locality Mountain range County Location 

C3 

Member 

L __ USGS 5441-PC ___ Cherokee Spring_ Rawlins hills ------- Carbon __ Probably SW~ sec. 11, T. Horseshoe 

21 N., R. 88 W. Shale. 

2 ___ USGS 8071-PC --- ___ do ---------- ___ do -------------- ___ do ---- ___ do ------------------- Do. 

3 ___ USGS 3139-PC --- ___ do ---------- ___ do -------------- ___ do---- NW~SW~ sec. 11, T. 21 Do. 

N., R. 88 W. 

4 ___ USGS 3140-PC Cherokee Peak _____ do -------------- ___ do---- NE~ sec. 10, T. 21 N., R. 

88 W. 

5 ___ USGS 3085-PC ---do ---------- ___ do -------------- ___ do---- SE~NE~ sec. 10, T. 21 

N., R. 88 W. 

7 ___ USGS 21727-PC __ Meadow Ranch-- ___ do -------------- ___ do---- SW~NE~ sec. 25, T. 22 

N., R. 88 W. 

17--- USGS 19236-PC __ Long Creek 

Canyon. 

Granite Mountains -- Fremont - SW~ sec. 23, T. 32 N., R. 

26b __ USNM 487 ------- Amsden Hill ____ Wind River Range _____ do ___ _ 94 W. 

Probably SW~ sec. 32, T. 

31 N., R. 99 W. 

27--- USGS 19283-PC __ Cherry Creek --- ___ do -------------- ___ do ____ Probably sec. 30, T. 31 N., 

R. 99 W. 

36a __ UM 2663 --------- ___ do ---------- ___ do -------------- ___ do---- Probably sec. 19, T. 31 N., 

R. 99 W. 

36c __ UN 2664a -------- ___ do ---------- ___ do -------------- ___ do ____ ---do ------------------- 36e __ USNM 487A ------ ___ do ---------- ___ do -------------- ___ do ____ ---do ------------------- 38 ___ USGS 16397-PC __ Little Popo Agie ___ do -------------- ___ do ____ NW~NW ,4 sec. 19, T. 31 

N., R. 99 W. -------------- ___ do---- SE~SE~ sec. 13, T. 31 

Canyon. 39 ___ USGS 14025-PC _____ do ---------- ___ do 

N., R. 99 W. 

4L __ USGS 19156-PC __ Soda Creek ----- Washakie Range ---- T·eton ____ Sec. 5, T. 45 N., R. 110 W _ 

48 ___ YPM 20107 ------- Wiggins Fork --- ___ do -------------- Fremont _ SW~ sec. 7, T. 42 N., R. 

105 w. 

72 ___ USGS 6957-PC ___ Bear Creek ----- Salt River Range ___ Lincoln __ NE~SW ,4SW~ sec. 27, 

T. 33 N., R. 117 W. 

73 ___ USGS 6960-PC --- Moffat Trail ---- ___ do -------------- ___ do ____ NW~NE1,4 sec. 3, T. 33 

74 ___ USGS 22985-PC _____ do ---------- ___ do 

75 ___ USGS 22986-PC _____ do ---------- ___ do 

76 ___ USGS 22987-PC __ _ __ do ---------- ___ do 

77--- USGS 6960A-PC _____ do ---------- ___ do 

78 ___ USGS 6960B-PC _____ do ---------- ___ do 

79 ___ USGS 22981-PC _____ do ---------- ___ do 

80 ___ USGS 2.2982-PC _____ do ---------- ___ do 

88 ___ USGS 22992-PC __ Peak 9837 ------ ---do 

N., R. 117 W. 

-------------- ___ do _______ do ------------------- -------------- ___ do _______ do ------------------- -------------- ___ do _______ do ------------------- -------------- ___ do _______ do ------------------- -------------- ___ do ---- ___ do ------------------- -------------- ---do _______ do ------------------- __ _. ______________ do ---- ___ do -------------------

-------------- ---do ---- NE~ sec. 34, T. 34 N., R. 

117 w. 

90 ___ USGS 22995-PC __ Covey Cutoff ___ do -------------- ---do---- NW1,4NE1,4 sec. 27, T. 34 

N., R. 117 W. 

-----------------do _______ do -------------------

Trail. 

9L __ USGS 22996-PC _____ do ---------- ---do 

92 ___ USGS 6965-PC ______ do ---------- ___ do -----------------do _______ do ------------------- 93 ___ USGS 6951-PC ______ do ---------- ___ do -------------- ___ do ---- ___ do ------------------- 103 ___ USGS 17905-PC Hay·stack Peak _____ do -------------- ---do---- Center sec. 19, T. 34 N., R. 

117 W. 

104 ___ USGS 17906-PC ___ do ---------- ___ do -------------- ___ d.o ---- ___ do ------------------- 105 ___ USGS 17907-PC _____ do ---------- ___ do -------------- ___ do ---- ___ do ------------------- 109 ___ USGS 23002-PC __ South Indian Snake River Range -- ---do---- NE~ sec. 14, T. 38 N., R. 

Creek. 118 W. ' 

12L __ USGS 16208-PC __ Hoback Canyon - Hoback range ------- Teton ---- Sec. 2, T. 38 N., R. 115 W _ 

122 ___ USGS 16209-PC _____ do ---------- ___ do -------------- ---do ---- ___ do ------------------- 124 ___ USGS 18799-PC _____ do ---------- ___ do -------------- ---do---- Sec. 3, T. 38 N., R. 115 W _ 

126 ___ USGS 22998-PC _____ do ---------- ___ do -------------- ___ do---- Sec. 2, T. 38 N., R. 115 W _ 

Do. 

Do. 

Do. 

Ranchester 

Limestone. 

Horseshoe 

Shale. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Moffat Trail 

Limestone. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

The coelenterate faunas of the Horseshoe and 

Ranchester Members are too meager and too poorly 

known to be of much value in correlation of these 

parts of the Amsden Formation. Barytichisma amsdenense (Branson and Greger), from the Mississippian part of the Horseshoe, is unknown outside of 

the Amsden and is the first Mississippian species 

to be assigned to the genus BaryUchism.a, which was 

previously known only from beds of Early Pennsylvanian age. N eokoninckophyllum hamatilis n. sp. 

and N. inconstans n. sp., from the Pennsylvanian 

part of the Horseshoe Member, are also known 

only from the Amsden Formation; other described 

species of N eokoninckophyllurn in North America 

range through most of the Pennsylvanian. The only 

coelenterate found in the Ranchester Member is a 

poorly preserved species of Chaetetes most similar 

to Chaetetes eximius Moore and Jeffords from the 

Early Pennsylvanian of Texas and Oklahoma. 
C4 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

The coelenterate fauna of the Moffat Trail Limestone Member offers a more fruitful basis for correlation. Most of the described species in this fauna 

have been found elsewhere in the western United 

States in the Caninia Zone, also known as Zone K 

in the biostratigraphic scheme of Sando, Mamet, 

and Dutro ( 1969) . This zone has been correlated by 

means of Foraminifera with the middle and upper 

parts of the Chesterian Series of the midcontinent 

region and with the latest Visean (V3cs) and early 

Namurian (E1 and E2) of Europe. Characteristic 

of Zone K are the large cyathopsid corals allocated 

to Caninia nevadensis (Meek) and the related species C. excentrica (Meek), C. enormis Easton, and 

C. montanensis Easton which are known elsewhere 

from the Surrett Canyon Formation and Monroe 

Canyon Limestone of Idaho, the Big Snowy Group 

of southwest and central Montana, the upper parts 

of the Great Blue Limestone and Green Ravine Formation of central Utah, and the Ochre Mountain 

Limestone and Chainman Shale of western Utah. 

Zone K is also characterized by a proliferation of 

distinctive syringoporoid corals. Among th~ Moffat 

Trail representatives of this group is Duncanopora 

duncanae n. sp., a species known from the Surrett 

Canyon Form~ation and Monroe Canyon Limestone 

of Idaho and the Green Ravine Formation of Utah. 

Pleurosiphonella drummondi (Nelson), another Moffat Trail syringoporoid, occurs in the upper part of 

the Etherington Formation of western Canada. 

Another characteristic Zone K coral is Lonsdaleia 

(Actinocyathus) stelcki (Nelson), which occurs in 

the Surrett Canyon Formation and Monroe Canyon 

Limestone of Idaho, the Green Ravine Formation of 

Utah, and the Etherington Formation of western 

Canada. Zone K coelenterates not certainly known 

outside of the Moffat Trail Limestone Member of 

the Am,sden are Chaetetes wyomingensis n. sp., 

Multithecopora? amsdenensis n. sp., and Turbinatocaninia? sp. 

The classification of the chaetetid hydrozoans followed in this paper is taken from Sokolov ( 1950, 

1955). The Rugosa are classified according to Hill 

(1956). Tabulates are regarded as an order, following Hill and Stumm ( 1956) , but the families are 

taken from Sokolov (1962). 

Morphologic terminology follows that of Hill 

(1956) and Hill and Stumm (1956) with the exception of the terms "alar diameter" and "calicular 

angle," which were defined by Sando (1961). The 

terminology of microstructural elements is that of 

Kato (1963). In descriptions of soHtary corals 

where the terms "left" and "right" are used, these 

terms are in reference to the cardinal-counter plane 

when the corallum is viewed from below with the 

cardinal septum oriented toward the observer. 

Photographs and drawings of serial transverse thin 

sections of solitary corals ~are oriented so as to be 

looking up the corallum axis toward the calice and 

the cardinal septum is toward the bottom of the 

plate. 

Specimens studied are in the collections of the 

U.S. National Museum, Washington, D.C. (USNM), 

the Department of Geology, University of Missouri, 

Columbia, Mo., (UM), and the Peabody Museum 

of Natural History, Yale University, New Haven, 

Conn., (YPM). Detailed locality data are given in 

the Register of Amsden Fossil Collections in Sando, 

Gordon, and Dutro, 197 4. 

I am indebted to the late Helen Duncan for technical review of the manuscript and particularly for 

her advice concerning the taxonomy of the Chaetetidae. Thanks are also extended to W. A. Oliver, Jr., 

for his review of the manuscript. Thin sections were 

prepared by K. R. Moore, and photographs were 

made by D. H. Massie (thin sections) and R. H. 

McKinney (exteriors) . 

SYSTEMATIC PALEONTOLOGY 

Phyllum COELENTERATA 

Class HYDROZOA 

Group CHAETETIDA 

Family CHAETETIDAE 

Discussion.-Among the most interesting of the 

Amsden fossils are the colonial organisms classified 

as chaetetids. Although the taxonomic position of 

this poorly known group was doubtful for many 

years, Sokolov ( 1950, 1955) has presented convincing arguments for interpreting it as an ancient 

member of the Class Hydrozoa and has pioneered 

in the discrimination and description of genera. Accordingly, the classification and morphological terminology used by Sokolov are followed in this paper. 

Because of their similarity to the Bryozoa, an 

appreciable number of bryozoan species were formerly allocated to the chaetetids. Aside from these, 

the valid North American record of the Chaetetidae 

is almost exclusively Pennsylvanian, in fact so overwhelmingly so that the group, and particularly the 

genus Chaetetes, has been regarded generally as an 

index to beds of Pennsylvanian age in North America. However, rare Mississippian occurrences were 

recorded by Stouder (1938, p. 284) and H. M. Duncan (in U.S. Geological Survey, 1965, p. A122; 1966, 

p. A112) in the Chester Series of Kentucky. Moreover, the worldwide range of the group is from 

Ordovician to Miocene (Sokolov, 1955, p. 95), and 
COELENTERATA OF THE. AMSDEN FORMATION OF WYOMING C5 

the Chaetetidae are common throughout the Carboniferous in Europe (Sokolov, 1939, p. 411). It 

is not surprising, therefore, to find this group represented in both the Mississippian and Pennsylvanian 

parts of the Amsden Formation. 

The identifiable Amsden forms are all allocated 

to the genus Chaetetes Fis.cher. Two species, both 

similar to Chaetetes milleporaceus Milne-Edwards 

and Haime, are recognized in the Amsden material, 

one in the Mississippian part and the other in the 

Pennsylvanian part of the formation. 

Chaetetiform trepostomatous bryozoans occur 

sparingly in the Moffat Trail Member of the Amsden along with Chaetetes wyomingensis n. sp. These 

bryozoans formed spherical or hemispherical colonies of size comparable to that of the Chaetetes, are 

comrposed of polygonal to rounded-polygonal tubes 

of comparable diameters, and have horizontal partitions similar to the tabulae of Chaetetes. Although 

the bryozoans are not formally described in this 

paper, it seems desirable to point out features by 

which they may be distinguished from the chaetetids 

(compare plates 1 and 2): 

1. The wall structure of the bryozoans is laminar, 

as compared to the trabecular wall structure 

of the chaetitids. 

2. The walls of the bryozoans are thinner (0.01 to 

0.03 mm, average about 0.02 mm) than those 

of the Amsden Chaetetes. 

3. The bryozoans are ·Composed of tubes of two 

distinct sizes (smaller ones are acanthopores 

or mesopores) , whereas the Amsden Chaetetes 

have tubes of relatively uniform diameter. 

4. The bryozoans lack pseudosepta. Reproduction is 

by fission in which vertical partitions arise 

from the floors of horizontal partitions so that 

the vertical partitions are always complete in 

transverse section. In Chaetetes, vertical partions arise from one or more sides of the walls 

of the tubes so that many are incomplete in 

transverse section. 

Genus CHAETETES Fischer von W aldheim in Eichwald, 1829 

Chaetetes wyomingensis n. sp. 

Plate 1, figures 1-3 

Description.-Colony seemingly regularly or irregularly spherical or hemispherical, composed of 

radially diverging cells. The largest colonies (including holotype) were at least 6 em high and 13 em in 

diameter; most of the specimens are fragments of 

smaller size than this, and the actual size of the 

colonies was probably considerably larger than the 

largest measured specimen. The specimens are all 

composed of partly silicified, moderately to strongly 

recry,stallized calcite. 

In transverse section (pl. 1, fig. 1), the cross 

sections of cells are polygonal, mostly hexagonal, 

but rarely quadragonal or pentagonal. They are 

equidimensional to elongate, but not meandroid, and 

fairly uniform in diameter. Visceral spaces are 

angular to rounded. Cell walls are straight and unbeaded. Wall microstructure is faintly trabecular 

(pl. 1, fig. 3). Maximum diameter of the visceral 

space ranges from 0.12 to 0.45 mm; the average 

per colony ranges from 0.28 to 0.35 mm (table 3) . 

Minimum diameter of the visceral space ranges 

from 0.11 to 0.42 mm; the average per colony ranges 

from 0.23 to 0.29 mm (table 3). Wall thickness 

ranges from 0.02 to 0.05 mm; the average per colony 

ranges from 0.30 to 0.36 mm (table 3). Pseudosepta 

are common, present in about 10 to 20 percent of 

the cells. Fission is mostly bipartite, rarely tri- or 

quadripartite; all stages of fission are commonly 

visible. 

In longitudinal section (pl. 1, fig. 2), the cells 

appear as straight to slightly sinuous tubes that 

have generally parallel walls except where fission 

has taken place. There is some evidence for fission 

being more common at some levels of the colony than 

at others. Evidence of complete fission is rare. 

Tabulae are mostly complete, that is, their traces 

ordinarily extend uninterruptedly from one side of 

the wall to the other; incomplete traces apparently 

are the result of poor preservation. Tabulae are 

horizontal and flat to very slightly concave upward. 

Tabular thickness is approximately 0.01 mm but is 

difficult to measure owing to difficulty in differentiating true tabular structure from crystalline overgrowths. Tabulae in adjacent cells may or may not 

be coincident in position; the majority are noncoincident. Tabular spacing is generally relatively 

constant from one cell to another in a given colony. 

Although some colonies exhibit nontabulate zones 

of 1 or 2 mm thickness, these do not persist throughout the colony and appear to be a function of preservation. The average tabular frequency is about four 

tabulae per mm and is relatively constant from one 

colony to another. Individual tabulae are ordinarily 

spaced 0.10 to 0.77 mm apart. 

Discussion.-Chaetetes wyomingensis is the first 

true Chaetetes to be described from the Mississipian 

rocks of North America. Its occurrence in the Amsden Formation is confined to the Caninia Zone of 

the Moffat Trail Member in western Wyoming. 

Foraminifera in three of the five collections are indicative of Foraminiferal Zones 17 or 18, an interval 

of early Namurian (El-E2) age. 
C6 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

TABLE 3.-Measurements (in mm) of Chaetetes wyomingensis n. sp. in transverse section 

[Each parameter is represented by 20 measurements per specimen] 

Maximum diameter Minimum diameter 

USNM 

specimen 

numoer 

of visceral space of visceral space Wall thickness 

Range Average Range Average Range Average 

165082 -- 0.22-0.44 

16.5081 .17-.45 

165083 .22-.45 

165084 .21-.38 

165080 .12-.34 

0.33 0.21-0.42 

.30 .15-.30 

.35 .15-.38 

.30 .15-.30 

.28 .11-.29 

0.29 0.02-0.05 0.036 

.23 .02-.05 .034 

.27 .03-.04 .035 

.23 .02-.04 .030 

.23 .02-.05 .032 

Comparison with other North American species 

is based on descriptions and illustrations of a topotype of Chaetetes milleporaceus Milne-Edwards and 

Haime (Middle Pennsylvanian of Indiana) published 

by Lane and Martin (1966) and of three species 

from the Lower Pennsylvanian of Texas and Oklahoma by Moore and Jeffords (1945). Chaetetes 

wyomingensis is similar to C. milleporaceus in the 

average diameter of its visceral spaces and in its 

average tabular frequency but is readily distinguished by its thinner cell walls (about 0.03 mm vs. 

0.065 mm in C. milleporaceus) and its relatively 

abundant pseudosepta. Moreover, C. milleporaceus 

formed lenticular colonies whereas C. wyomingensis 

appears to have formed large spherical or hemispherical colonies. 

The ri~w Amsden species is also readily differentiated from C. eximius, C. favosus, and C. subtilis 

(all described by Moore and Jeffords, 1945, p. 191-

195) on the basis of its thinner cell walls. C. eximius 

and C. favosus have about the same average visceral 

space diameter as C. wyomingensis, but the Pennsylvanian species have beaded walls. C. subtilis has 

straight walls but has a smaller visceral space diameter. Pseudosepta are rare in C. favosus and C. 

subtilis. Tabulae are also rare and commonly incomplete in these two species, but preservation may 

be a factor in this feature. Tabular frequency in 

C. eximius is greater than that of C. wyomingensis. 

Occurrence ·t:Lnd abundance.-Moffat Trail Limestone Member: collections 72 (2), 77 (1), 78 (1), 

79 (2+), 80 (1), Lincoln County, Wyo. 

Specimens.-Holotyp.e, USNM 165080; paratypes, 

USMN 165081-84. 

Chaetetes cf. C. eximius Moore and Jeffords 

Plate 1, figures 4-7 

Description.-Colony seemingly large, massive, 

exact form unknown, composed of radially diverging cells. The species is known from one completely 

silicified specim·en that was at least 5 em high and 

10 em in diameter. 

In transverse section (pl. 1, fig. 6), the cross sections of cells are polygonal, mostly hexagonal, but 

rarely quadragonal or pentagonal. They are equidimensional to elongate, but not meandroid, and fairly 

uniform in diameter. Visceral spaces are angular 

to rounded. Cell walls are straight and unbeaded. 

Wall microstructure is obscure but seemingly trabecular (pl. 1, fig. 4). Maximum diameter of the 

visceral space ranges from 0.20 to 0.44 mm; the 

average per colony is 0.33. Minimum diameter of 

the visceral space ranges from 0.16 to 0.33 mm; 

the average per colony is 0.26. Wall thickness 

ranges from 0.04 to 0. 7 mm; the average per colony 

is 0.055. Pseudosepta are very rare in some sections 

and common (visible in 10 to 20 percent of the 

cells) in others. Where present, fission is mostly 

bipartite but may be tri-or quadripartite; all stages 

of fission may be seen. 

In longitudinal section (pl. 1, figs. 5, 7), the cells 

appear as straight to slightly sinuous tubes that 

have generally parallel walls. Evidence of complete 

fission is rare. Tabulae are extremely rare in most 

sections, probably owing to poor preservation. They 

are complete or incomplete, mostly horizontal, and 

flat to very slightly concave upward. In well-preserved parts of the colony, the tabulae are spaced 

0.10 to 0.30 mm apart, and the average tabular frequency is about six or seven tabulae per mm. 

Discussion.-This species is based on a single 

completely silicified specimen from the Ranchester 

Member and is of Pennsylvanian age. Because the 

poor preservation of the material leaves some doubt 

as to the frequency of the tabulae, it does not seem 

advisable to give this species a new name. 

Chaetetes cf. C. eximius is similar to C. wyomingensis in visceral space diameter, cell shape, and 

apparently in the form of the colony, but it is readily 

distinguished by its thicker cell walls. Moreover, 

where tabulae are preserved, the average tabular 

frequency is greater in the Pennsylvanian form. 

The Amsden species seems most ·similar to Chaetetes eximius Moore and Jeffords (1945, p. 191, figs. 

207, 208) but has straight instead of beaded walls 

and has slightly thicker walls. Its cells are larger 

than those of C. subtilis Moore and Jeffords (1945, 

p. 194, figs. 211a-d), and it has more numerous 

pseudosepta than that species. It is distinguished 

from C. favosus Moore and Jeffords (1945, p. 193, 

figs. 209, 210) by its straight walls, more numerous 

pseudosepta, and slightly thicker walls. It differs 

from C. milleporaceus Milne-Edwards Haime (see 

Lane and Martin, 1966) in having slightly thicker 

walls, more numerous pseudosepta, and a greater 

tabular frequency. 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C7 

Occurrence and abundance.-Ranchester Limestone Member: collection 17 (1), Fremont County, 

Wyo. 

Specimen.-Figured, USNM 165085. 

Chaetetes sp. indet. 

1918. Chaetetes? Branson and Greger, p. 313, pl. 19, figs. 

21, 22. 

1937. Chaetetes? sp. Branson, p. 652. 

Description.-Two fragmentary, completely silicified specimens approximately 4 mm high by 4 mm 

in diameter cannot be assigned to any known species 

owing to poor preservation of internal structures. 

The general morphology of these specimens is essentially the same as that of the two species described 

previously in this report. As in C. cf. C. eximius, 

evidence of fission is rare and tabulae are rare, probably as the result of silicification. Visceral space 

diameters are within or close to the ranges of both 

species previously described; maximum diameter 

ranges from 0.20 to 0.39 mm and averages 0.28 mm, 

whereas minimum diameter ranges from 0.14 to 

0.31 mm and averages 0.22 mm, as measured in one 

specimen (UM 2664). Wall thickness is close to that 

of C. cf. C. eximius, ranging from 0.04 to 0.07 mm 

and averaging 0.056 mm (UM 2664). 

Morphologically, the species seems to be closer to 

C. cf. C. eximius from the Ranchester Member in 

central Wyoming rather than C. wyomingensis from 

the Moffat Trail Member of western Wyoming. However, crude preservation of internal structures by 

silicification leaves doubt concerning wall thickness 

and number of tabulae. 

Occurrence and abundance.-Horseshoe Shale 

Member: collections 26b ( 1) , 36c ( 1) , Fremont 

County, Wyo. 

Specimens.-Unfigured, USNM 487 and UM 2664. 

Class ANTHOZOA 

Subclass ZOANTHARIA 

Order RUGOSA 

Suborder STREPTELASMA TINA 

Superfamily CY A THAXONIICAE 

Family AMPLEXIDAE 

Genus AMPLEXUS Sowerby, 1814 

"Amplexus" sp. indet. 

Plate 3, figures 26-29 

Description.-This category includes five specimens, four of which appear to be immature forms, 

from a single locality. The largest specimen in the 

lot is the one illustrated on pl[-..te 3. This specimen 

is about 28 mm long and attains a maximum diameter of 14 mm, at which there are 26 short major 

septa. Minor septa are absent. The corallum is 

cylindrical and slightly curved, and the cardinal 

septum is on the concave side of the corallum. Serial 

transverse sections (pl. 3, figs. 26-29) suggest that 

the early part of the corallum (up to a diameter of 

about 9 mm) is characterized by a zaphrentoid septa 

plan like that of the genus Zaph1·entites Hudson. 

Later sections are characterized by amplexoid retreat of the major septa. Tabulae appear to be more 

or less flat and complete, downturned at the margins 

and depressed in the cardinal fossula. Dissepiments 

are absent. 

Discussion.-Easton (1962, p. 31) pointed out 

that the genus Amplexus has been so broadly applied as to include corals of uncertain phylogenetic 

relationship. Hill (1940, p. 147-148) also suggested 

that the group is probably polyphyletic. Moreover, 

the early stages of the type species are unknown, 

and until that is determined, species cannot be assigned to Amplexus with any degree of certainty. 

Easton (1962, p. 31) erected the genus Fasciculiamplexus, based on a species whose early stages are 

known, as a partial solution to this nomenclatural 

problem. According to Easton, Fasciculiamplexus 

may have developed from Fasciculophyllum Thomson by continuation of an amplexoid trend. Although 

the Amsden coral here provisionally called "Amplexus" has many features in common with Fasciculiamplexus, its cardinal fossula is on the concave 

side of the corallum, rather than on the convex side 

as in Easton's genus. I am therefore reluctant to 

refer it to Fasciculiamplexus. Because the Amsden 

material is meager and generally poorly preserved, 

it is impractical to base a new genus on it; hence 

the old designation "Amplexus" is retained. 

Occurrence and abundance.-Horseshoe Shale 

Member: collection 41 (5), Teton County, Wyo. 

Specimens.-USNM 165088. 

Family HAPSIPHYLLIDAE 

Genus ZAPHRENTITES Hudson, 1941 

Zapbrentites 1 sp. indet. 

This category includes four small, imperfectly 

preserved horn corals that have a simple zaphrentoid septal plan and cardinal fossula on the concave 

side of the corallum. The largest specimen is about 

30 mm long and attains a maximum diameter of 

about 15 mm, at which there are 27 major septa. 

In the absence of better material, it is impossible 

to determine whether these specimens represent the 

mature stages of Zaphrentites or are merely immature stages of amplexoid corals such as "A rr1-plexus" 

or Barytichisma. 

Occurrence and abundance.-Moffat Trail Limestone Member: Collections 76 ( 1), Lincoln County; 

121 (2), Teton County. Horseshoe Shale Member: 

Collection 48 (1), Fremont County, Wyo. 

Specimens.-USNM 165089, 165090; YPM 2010. 
C8 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

Genus BARYTICHISMA Moore and Jeffords, 1945 

Baryticbisma amsdenense (Branson and Greger) 

Plate 3, figures 1-25 

1918. Zaphrentis amsdenensis Branson and Greger, p. 313, 

pl. 19, figs. 19, 20. 

1937. Zaphrentis amsdenensis Branson and Greger. Branson, p. 652. 

1950. Zaphrentis amsdenensis Branson and Greger. Bassler, 

p. 220. 

Diagnosis.-Solitary, moderately curved, trochoid 

to ceratoid corals that have a cardinal septum on the 

concave side of the corallum. Corallum as much as 

50 mm in length and 23 mm in maximum diameter 

and having as many as 38 major septa at maturity. 

Calice moderately deep and having a slightly depressed floor. Exterior marked by prominent septal 

grooves and interseptal ridges and transverse wrinkles and fine growth lines. Theca thick. In young 

stages, major septa axially confluent and grouped 

in quadrants that are marked by a well-defined cardinal fossula, alar pseudofossulae, and less readily 

identified counter septum. Ephebic stage characterized by amplexoid retreat of major septa on upper 

surfaces of tabulae. Minor septa absent. Tabulae 

mostly complete, more or less horizontal, but bent 

downward marginally, and deeply depressed in cardinal fossula. Dissepiments absent. Septal microstructure of the feather type of Kato (1963). 

Description of lectotype.-The type lot (UM 

2663) consists of six free coralla of yellowish calcite preservation evidently collected from float. The 

largest specimen in the lot is chosen as the lectotype~; 

this specimen appears to be the one illustrated by 

Branson and Greger, although their illustrations are 

not clear enough to be certain. 

The lectotype (pl. 3, figs. 1-18) is a nearly perfect specimen, lacking only the walls of the calice, 

which were broken off down to the floor of the calice. 

The corallum is 50 mm long and attains a maximum 

diam.eter of 23 mm. The corallum is trochoid, moderately curved, and has an apicai angle of about 

44° and a calicular angle of about 16°. The cardinal 

septum is on the concave side of the corallum and 

is located slightly to the left of the plane of curvature. The transverse section of the corallum is approximately circular throughout the growth. The 

broken calice (pl. 3, fig. 15) reveals 38 major septa 

that exend from the periphery one-half to two-thirds 

of the radius on the axially-depressed tabular floor 

of the calice. The cardinal septum is reduced to a 

low ridge at the end of a parallel-sided fossula that 

has a depressed floor. No minor septa are present. 

Longitudinal ornamentation consists of broad, 

slightly elevated interseptal ridges 1 to 1.5 mm wide 

in the mature parts of the corallum. Transverse ornamentation consists of growth wrinkles 2 to 4 mm 

apart and finer growth lines. The exterior has been 

modified by solution weathering so that all ornamentation is subdued. 

Ontogeny in the lectotype was studied by means 

of 14 serial transverse sections (table 4; pl. 3, figs. 

1-14). Seven thin sections were cut at intervals of 

approximately 5 mm beginning at a level just below 

the floor of the calice down to a level 6.3 mm above 

the tip. The remaining, earliest part of the specimen was studied by the serial peel technique described by Sando (1967). 

The obvious lack of a significant part of the 

counter side of the corallum, probably due to solution weathering, seriously limited the completeness 

of observations made in the earliest 4.3 mm of the 

specimen. The earliest section studied (table 4, section n; pl. 3, fig. 1) revealed only the cardinal 

septum, two alar septa, and a cardinal lateral septum, but it seems probable that the counter and one 

or more counter lateral septa were also present at 

this stage. Moreover, the very earliest part of the 

corallum (brephic stage) does not seem to have been 

preserved. Section i (table 4; pl. 3, fig. 6) is the 

first stage at which septal complements in the counter quadrants appear to be trustworthy. Acceleration in the counter quadrants, which marks the later 

stages, was first noted in this section. 

In the absence of a section in which only the 

protosepta are present, no brephic stage is recognized in the lectotype. The neanic stage (table 4, 

sections f to n; pl. 3, figs. 1-9) is characterized by 

a well-defined cardinal fossula that contains a cardinal septum of variable length and thickness and 

by axially confluent major septa. The neanic stage 

is recognized up to an alar diameter of 12 mm and 

a complement of 28 major septa. Section e (table 

4; pl. 3, fig. 10) records the earliest noted appearance of the mature characters of the corallum 

(ephebic stage). The alar diameter is 14.5 mm, and 

the corallum contains 34 major septa at this level. 

The ephebic stage is characteriz·ed by amplexoid 

retreat of the major septa, a very short cardinal 

septum, and a peripheral ster:eozone 1.5 to 2.5 mm 

thick. Minor septa are absent in all growth stages. 

The septal microstructure is the feather type of Kato 

( 1963) . Tabular traces are visible in all late neanic 

and ephebic transverse sections beginning with section h (table 4; pl. 3, fig. 7), but the nature of the 

tabulae was not studied in longitudinal section. 

Description of other specimens.-Superficial examination of paralectotypes, topotypes, and other 

specimens revealed a morphology similar to that of 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C9 

TABLE 4.-0ntogeny in the lectotype (UM 2663} of Barytichisma amsdenense (Branson and Greger). Brephic stage 

seemingly not preserved 

Section 

a -----

b -----

c -----

d -----

e -----

f -----

g -----

h -----

k -----

1 -----

m -----

n -----

Distance 

above 

tip 

(mm) 

40 

35 

30 

25 

20 

13 

6.3 

5.3 

4.3 

3.3 

2.3 

1.3 

.3 

0-.2 

Alar 

diameter 

(mm) 

23 

22 

19.5 

17.5 

14.5 

12 

7.9 

5.6 

4.4 

3.1 

2.5 

.9 

.6 

.4 

Number 

of major 

septa 

38 

37 

34 

34 

34 

28 

24 

19 

17 

10? 

9? 

6? 

6? 

4? 

Septal 

formula 

K 

10 110 

A-1-A 

717 

c 

K 

10 19 

A-l-A 

717 

c 

K 

919 

A-l-A 

616 

c 

K 

919 

A-l-A 

616 

c 

K 

919 

A-1-A 

616 

c 

K 

717 

A-1-A 

515 

c 

K 

616 

A-1-A 

414 

c 

K 

314 

A-l-A 

313 

c 

K 

314 

A-1-A 

313 

c 

K 

1? 11? 

A-l-A 

212 c 

K 

0? 11? 

A-l-A 

212 

c 

K 

o? I o? 

A-l-A 

111 

c 

K 

o? I o? 

A-l-A 

1!1 

c 

? 

A-1-A 

110 

c 

Ontogenetic 

stage 

Ephebic 

Neanic 

the lectotype. Although a few specimens have nearly 

straight conical coralla, the dominant corallum form 

is a slightly curved cone. The specimens range from 

8 to 45 mm in length and 11 to 21 mm in maximum 

diameter ; all but two of the specimens are incomplete. The cardinal septum is invariably on the concave side of the corallum, mostly in the plane of curvature or slightly to one side, but rarely close to the 

alar plane. Calicular angles range from 15° to 40° 

(average 21.9°) and apical angles range from 25° 

to 48:J (average 37°). Except where abraded or corroded, the exteriors are marked by rather strong 

longitudinal ribbing and transverse wrinkles and 

fine growth lines. Septal complements at various 

growth stages are shown in figure 1. 

One of the topotypes (USNM 165091), a neanic 

specimen about 20 mm long and 12 mm in maximum 

diameter, was serially sectioned (table 5; pl. 3, figs. 

20-25) in order to study early growth stages that 

are unreliably represented in the lectotype. The 

earliest part of the tip was not preserved on this 

specimen, but a four-septum stage (pl. 3, fig. 20) 

represents the brephic part of the corallum. 

A thin section of one of the paralectotypes (UM 

2663 ; pl. 3, fig. 19) illustrates features of the species seen in longitudinal section. The tabulae are 

mostly complete, flat to sagging, and slightly inclined cardinally. Their margins are reflexed downward. Dissepiments are absent. 

Discussion.-Species previously assigned to the 

genus Barytichisma are exclusively Early Pennsylvanian. These include B. crassum, B. repletum, and 

B. callosum, described by Moore and Jeffords (1945, 

p. 131-137) from formations of Morrowan age in 

Texas and Arkansas, and B. zaphrentiforme 

(White), redescribed by Sando (1965, p. E7-E11) 

from the Morgan Formation of Atokan age in 

Colorado. Although Barytichisma atnsdenense seems 

most closely allied to B. callosurn and B. zaphrentiforme by virtue of its cardinal septum on the concave side of the corallum, it differs significantly from 

both of these Pennsylvanian species. B. callosum has 

a cylindrical rather than curved conical corallum, 

a larger number of major ·septa at comparable corallum diameters, and more numerous tabulae than B. 

amsdenense. B. zaphrentiforrr~e is considerably 

larger than B. amsdenense, has a much thicker 

theca, and does not have the coarse external ribbing 

of the Amsden species. B. arnsdenense and B. 

zaphrentiforme have a similar ontogeny and septal 

complement at comparable corallum diameters, but 

the latter has a greater septal number at maturity. 

Ba1-·ytichisma amsdenense is known only from a 

relatively small area in the southern Wind River 
C10 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

Range that includes the Cherry Creek, Amsden Hill, 

and Little Popo Agie River localities. Its exact stratigraphic position is in doubt because all the .specimens were collected from float. However, it occurs 

with fos,sils characteristic of the late Chesterian 

Anthracospirifer welleri-shawi Zone at all localities. 

In three of the collections (26b, g6a, g9) it occurs 

with fossils representative of both the Composita 

poposiensis Subzone and the Carlinia amsdeniana 

Subzone. In two collections (g6e, g8) it occurs with 

fossils representativ~e of the Composita poposiensis 

Subzone. It occurs exclusively with Carlinia amsdeniana Subzone fossils in only one collection (27). 

The known di,stribution suggests that B. amsdenense 

is restricted to the Composita poposiensis Subzone 

and that its occurrence with fossils of the Carlinia 

amsdeniana Subzone at some localities is accidental. 

Occurrence and abundance.-Horseshoe Shale 

Member: collections 26b(2?), 27(16), g6a(6), 

g6e(2), g8(11), g9(2), Fremont County, Wyo. 

Specimens.-Lectotype and paralectotypes 

UM266g; topotypes USNM 165091, 165092; other 

specimens USNM 487, 487A, 16509g, 165094. 

Superfamily ZAPHRENTICAE 

Family AULOPHYLLIDAE 

Subfamily AULOPHYLLINAE 

Genus NEOKONINCKOPHYLLUM Fomicbev, 1939 

Neokoninckopbyllum bamatilis n. sp. 

Plates 4 and 5 

Diagnosis.-Solitary corals having a conical 

neanic stage and a cylindrical ephebic stage. Corallum as much as 75 mm or more in length and 40 

mm in maximum diameter and having as many as 

65 major septa at maturity. Exterior marked by 

subdued interseptal ridges and narrow septal 

grooves and subdued rugae and fine growth lines. 

Corallum supported by talons in early stages. Rejuvenation common. Theca thin. Septal plan pinnate 

in brephic and early neanic stages but radial in 

later stages. Cardinal septum long, on convex side 

of corallum in conical stage, and in a poorly defined 

fossula. Minor septa very short. N eanic axial structure consisting of a medial plate oriented in the 

cardinal-counter plane and continuous with cardinal 

and counter septa, periaxial tabellae and tabulae, 

and a few septal lammellae; medial plate free, very 

thin, sinuous, and discontinuous and septal lamellae 

poorly developed in ephebic stage. Tabularium 

marked by a marginal septal stereozone and horizontal inflated tabulae. Dissepimentarium characterized by as many as 1g rows of steeply inclined 

regular and herringbone dissepiments and rare 

lonsdaleoid dissepiments. Septal microstructure of 

the diffusotrabecular type of Kato ( 196g) . 

TABLE 5.-0ntogeny in a topotype (USNM 165091) of Barytichisma amsdenense (Branson and Greger) 

Distance 

above Alar 

Section tip diameter 

(mm) (mm) 

f ----- 14 11.5 

e ----- 9 8.0 

d ----- 5 6.0 

e ----- 3 3.6 

b ----- 2 2.2 

a ----- 1 0-1 1.3 

1 Actual tip broken off. 

Number 

of major 

septa 

27 

25 

22 

14 

9 

4 

Septal Ontoformula genetic 

K 

717 

A-l-A 

415 

c 

K 

716 

A-l-A 

414 

c 

stage 

K Neanic 

515 

A-1-A 

414 

c 

K 

313 

A-1-A 

212 

c 

K 

112 

A-1-A 

111 

c 

K 

A -1- A Brephic 

c 

Description of holotype.-The holotype (USNM 

165095; pl. 4) is a perfect specimen except for some 

decortication in the lower part of the corallum. The 

corallum is a gently curved cone about 55 mm long 

and g5 mm in maximum diameter. The calicular 

angle is between 25° and goo. The lower 10 m~m of 

the corallum is strongly curved and has the cardinal 

septum on the convex side, approximately in the 

plane of curvature. Above the lower 10 mm, the 

plane of curvature rotates through an angle of about 

goo from its earlier position so that the cardinal 

position, although still located on the convex side 

of the corallum, is about goo to the left of the plane 

of curvature when the corallum is viewed from 

below. Approximately 10 mm above the tip, the 

entire left side of the corallum is extended outward 

to form a supporting structure or talon; the lateral 

surface of the talon is extensively and deeply crenulated. Although the calice is covered with matrix, 

it appears to be only about 10 mm deep. The exterior 

of the corallum is marked longitudinally by broad, 

subdued interseptal costae and narrow septal 

grooves. Transverse ornamentation consists of subdued rugae and fine growth lines. One episode of 

rejuvenation is recorded 20 mm above the tip. 

The internal morphology of this specimen was 

studied by means of seven transverse thin sections 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C11 

cut from the upper part of the corallum (pl. 4, figs. 

9-15) and eight transverse peels made from the 

lower 4.5 mm (pl. 4, figs. 1-8). No longitudinal 

sections were made. Distance above tip, alar diameter, number of major septa, septal formula, and 

ontogenedc stage for each of these sections are recorded in table 6. The relation between number of 

major septa and alar diameter in various sections 

is compared with similar data from paratypes in 

figure 2. 

The brephic stage (table 6, sections a and b ; pl. 

4, figs. 1, 2), characterized by the development of 

the six protosepta (cardinal, counter, two alars, and 

two counter laterals), occupies the lower 0.4 mm 

of the corallum. The earliest section studied, cut 

within 0.1 mm of the preserved tip, revealed a twochambered cup partitioned by a single septum (pl. 

4, fig. 1). Serial studies indicate that this septum 

later differentiated into the cardinal and counter 

septa. In the interval 0.1 to 0.2 mm above the tip, 

the chamber to the right of the initial septum was 

24 

22 

20 

en 

18 

0::: 

UJ 

1-

UJ 16 

~ 

::::; 

...J 14 

~ 

z 

ci 12 

UJ 

1-

UJ 10 ~ 

ct 

0 

8 0::: 

ct 

...J 

ct 6 

4 

2 

0 

0 2 

EXPLANATION 

x Lectotype 

• Paralectotype 

• Topotype 

4 6 8 10 12 14 16 

closed by skeletal deposits, whereas the left-hand 

chamber remained open. Differentiation of the nextformed septa were observed within the skeletal deposits forming the thecal wall in the interval 0.2 

to 0.4 mm above the tip. The alar and two counter 

lateral septa began to differentiate approximately 

simultaneously. At first the protosepta were buried 

in the wall of the primitive cup (pl. 4, fig. 2) ; 

higher in the corallum, but still less than 0.4 mm 

above the tip, open spaces developed between the 

septa, and the large open ~space became the loculus 

between the left alar septum and the left counter 

lateral septa (pl. 4, fig. 3). 

The neanic stage (table 6, sections c-k; pl. 4, figs. 

3-11) is defined a,s the period of growth that began 

with insertion of the metasepta about 0.4 to 0.5 

mm above the tip and ended about 20 mm above the 

tip, where the rate of diameter increase appears to 

have diminished significantly (fig. 3). The total 

major septal complement at the end of the neanic 

stage was about 40 to 4 7. The septal plan was 

• 

18 20 22 24 26 28 30 32 34 36 38 

NUMBER OF MAJOR SEPTA 

FIGURE 1.-Scatter diagram showing relation between alar diameter and number of major septa by means of 30 measurements on 16 specimens of Barytichisma amsdenense (Branson and Greger). Straight lines connect measurements 

made from serial sections of lectotype (UM 2663) and a topotype (USNM 165091). 
C12 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

moderately pinnate and had relatively well defined 

insertion posi,tions up to about 3.5 mm above the 

tip (pl. 4, figs. 3-7), after which the plan tended 

to be radial, and had poorly defined insertion positions (particularly the alar positions). As a radial 

plan developed, the major septa became narrowly 

wedge shaped and withdrew from the axial region 

(pl. 4, figs. 9-11). The septa were uniformly dilated 

in the early neanic sections (pl. 4, figs. 3-8) , but 

dilation moved inward and was confined to the 

tabularium after development of the dissepimentarium (pl. 4, figs. 9-11). Insertion of major septa in 

the cardinal quadrants did not begin until the 

counter quadrants had a total of seven septa, and 

counter acceleration continued throughout the neanic 

stage. The cardinal septum is long and in a poorly 

defined loculus form.ed by slight bowing of adjacent 

cardinal lateral septa. Minor septa were first noted 

about 8 mm above the tip (pl. 4, fig. 9), where they 

are difficult to distinguish from dissepimental 

traces; they attained a maximum length of 0.5 mm 

40 

36 

32 

C/) 

0:: 

LLJ 28 1-

LLJ 

~ 

::::i 

...J 24 

~ 

z 

ci 20 

LLJ 

1-

LLJ 

~ 16 

<( 

Q 

0:: 

<( 12 ...J 

<( 

8 

4 

0 

0 4 

EXPLANATION 

X Holotype 

e Paratypes 

8 12 16 20 24 28 

(about one-ninth the length of the major septa). The 

first tabular traces are evident 0.5 mm above the 

tip (pl. 4, fig. 3), but they did not become abundant 

until8 mm above the tip (pl. 4, fig. 9). Dissepiments 

were first noted 3.5 mm above the tip (pl. 4, fig. 7), 

where they appear in transverse section as a single 

discontinuous ring of traces at the periphery of the 

corallum. By the end of the neanic stage (pt 4, fig. 

11), there were two to three rings of herringbone dissepi·ments. In the early neanic stage (pl. 4, 

figs. 3-5), thP. axial structure was manifested as a 

simple axial swelling of the union of pinnate major 

septa. At a level 2.5 mm ·above the tip (pl. 4, fig. 

6), an axial plate oriented with its long dimension 

approximately in the cardinal-counter plane began 

to differentiate, and a few axial tabulae appeared. 

Gradual withdrawal of most of the major septa produced a more or less discrete axial structure made 

up of a slightly swollen axial plate augmented by 

tabellae and short, irregular septal lamellae by the 

end of the neanic stage (pl. 4, fig. 11). The axial 

32 36 40 44 48 52 56 60 64 

NUMBER OF MAJOR SEPTA 

FIGURE 2.-Scatter diagram showing relation between alar diameter and number of major septa by means of 29 measurements on 12 specimens of N eokoninckophyllum hamatilis n. sp. Straight lines connect measurements made from serial sections of holotype (USNM (165095) and three paratypes (USNM 165097, 165099, 165104). 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C18 

plate was connected to the cardinal and counter 

septa throughout the neanic stage. 

The ephebic stage (table 6, sections 1-o ; pl. 4, 

figs. 12-15) comprises the upper 85 mm of the 

corallum, where increase in corallum length was 

greater than increase in corallum diameter (fig. 8). 

A total of 55 radially arranged major septa, whose 

length is about two-thi~ds the radius of the corallum, were developed by the end of this stage. Minor 

septa are ·Consistently short (maximum length 1 

mm). Aside from increase in size and number of 

component elements, no major changes took place 

in the skeleton from the late neanic stage. The characteristic ephebic transverse section consists of a 

central axial structure, whose radius occupies about 

a third of the radius of the corallum, a periaxial 

tabularium, whose width occupies the middle third 

of the radius of the corallum, and a peripheral dissepimentarium, whose width occupies the outer third 

of the radius of the corallum. The axial structure 

consists of a thin axial plate oriented with its long 

• 

en 24 0:: 

LLJ Talon tLLJ 

~ ". 

:J 20 ...J 

~ 

z 

a::· 16 

LLJ 

tLLJ 

~ 

c( 12 c 

0::: 

5 

c( 8 

4 

axis in the cardinal-counter plane, loosely-arranged 

tabellae, and as many as six irregular septal lamellae. The axial plate was connected to both the cardinal and counter septa in the early ephebic stage 

(pl. 4, figs. 12, 13), but became free in the late 

ephebic stage (pl. 4, figs. 14, 15). The major septa 

are dilated in the tabularium. The dissepimentarium 

consists mostly of regular and herringbone dessepiments, but one lonsdaleoid dissepiment was noted 

in the highest section studied (pl. 4, fig. 15). There 

is a maximum of about nine rings of dissepimental 

traces. 

The theca is very thin throughout growth; it 

never became thicker than about 0.3 mm. Septal 

microstructure is the diffusotrabecular type of Kato 

(1963). 

Description of paratypes.-The paratypes are all 

incomplete specimens that represent various parts 

of the conical or cylindrical stages of the species. 

Measurements of these specimens and the holotype 

suggests that the corallum was conical up to a 

DISTANCE ABOVE TIP, IN MILLIMETERS 

FIGURE 3.-Relation between alar diameter and distance above tip in the holotype of N eokoninckophyllum hamatilis n. 

sp. (USNM 165095). Straight lines fitted visually to points representing brephic, neanic, and ephebic stages in order 

to demonstrate changes in rate of diameter increase. 
C14 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

TABLE 6.-0ntogeny in the holotype (USNM 165095) of Neo- TABLE 6.-0ntogeny in the holotype (USNM 165095) of Neokoninckophyllum hamatilis n. sp koninckophyllum hamatillis n. sp-Gontinued 

Section 

Distance 

above 

tip 

(rom) 

0 ----- 51 

n ----- 40 

m ----- 29 

1 ----- 20 

k ----- 13 

j 10 

i_ 8 

h ----- 4.5 

g ----- 3.5 

f ----- 2.5 

e ----- 1.5 

d ----- 1.0 

c ----- .5 

1 Includes talon. 

Alar 

diameter 

(rom) 

32 

28 

25.5 

23 

11 

7.5 

5.2 

4.5 

4.0 

2.6 

2.1 

1.9 

Number 

of major 

septa 

55 

53 

52 

48 

39 

35 

30 

21 

21 

18 

13 

10 

8 

Septal 

formula 

K 

14115 

A-~-A 11 11 

c 

K 

14115 

A-,-A 

10.10 

c 

K 

13 115 

A-l-A 

10 110 

c 

K 

13!14 

A-1-A 

918 

c 

K 

10 Ill 

A-l-A 

717 

c 

K 

9 110 

A-1-A 

6j6 

c 

K 

718 

A-l-A 

516 

c 

K 

616 

A-i-A 

312 

c 

K 

616 A--A 

312 

c 

K 

6!5 

A-1-A 

211 

c 

K 

413 

A-l-A 

111 

c 

K 

412 

A-l-A 

OIO 

c 

K 

212 

A-l-A 

010 

c 

Ontogenetic 

stage 

Ephebic 

Neanic 

Section 

b -----

a -----

Distance 

above 

tip 

(rom) 

.2 

0-.1 

Alar 

diameter 

(rom) 

1.3 

0.9 

Number 

of major 

septa 

6 

2 

Septal 

formula 

K 111 A--A 

OIO 

Ontogenetic 

stage 

C Brephic 

K 

I 

c 

diameter of about 20 to 30 mm, after which it became 

nearly cylindrical. Several fragmentary specimens 

(pl. 5, fig. 7) suggest that the cylindrical stage was 

sinuous and of considerable length. Maximum length 

attained was greater than 75 mm. The maximum 

diameter observed in the paratypes is about 40 mm, 

at which 65 major septa are present (fig. 2). All 

specimens in which the early part of the corallum 

is preserved exhibit evidence of supporting talons 

like that of the holotype (an example is shown on 

pl. 5, figs. 8, 9) . Calices are rarely preserved except 

in some neanic specimens (pl. 5, fig. 10), where a 

moderately convex tabular floor is surmounted by 

low ridges formed by the axial plate and a few 

septal lamellae. The floor of the cardinal fossula is 

deeply depressed in these specimens. 

Supplementary information on the internal 

morphology of the skeleton was obtained from nine 

transverse thin sections and six longitudinal thin 

sections cut from six paratypes. A typical late 

neanic stage is illustrated on plate 5, figures 1-3. 

The transverse sections (pl. 5, figs. 2, 3) of this 

specimen are similar in most respects to comparable 

sections of the holotype (pl. 4, figs. 10, 12). The 

longitudinal section (pl. 5, fig. 1), cut perpendicular to the cardinal-counter plane, exhibits the same 

three-fold zonation of the corallum seen in transverse section. The axial zone consists of a strong, 

straight, continuous axial plate surrounded by short 

tabellae that slope away from the plate at angles of 

about 45° to 80° from the horizontal. The periaxial 

tabellae rest on larger, more gently sloping, slightly 

convex or concave tabulae that blend into the tabulae of the ta;bularium. The tabularium consists of 

broad, nearly horizontal, inflated tabulae. The dissepimentarium is composed of 2 to 10 rows of inflated, steeply inclined, relatively small dissepiments. 

Fragmentary cylindrical specimens (pl. 5, figs. 

4-6, 11, 12) exhibit a somewhat different axial 

structure in late ephebic longitudinal sections. In 

these specimens, the axial plate is very thin, sinuous, 
COELENTERATA OF THE. AMSDEN FORMATION OF WYOMING C15 

and occasionally discontinuous. Steeply inclined periaxial tabellae are rarely developed, and the tabulae 

of the axial zone are broad, slightly convex, gently 

sloping plates that may or may not have downturned 

peripheral edges. In transverse section, the axial 

plate is difficult to identify, septal lamellae are rare, 

and tabular traces are haphazardly arranged. Tabulae of the tabularium are similar to those of the 

neanic stage. The dissepimentari urn consists of as 

many as 13 rows of steeply inclined, mostly herringbone and regular dissepiments. Lonsdaleoid dissepiments are rare. 

Discussion.-Although N eokoninckophyllum hamatilis shows some similarities to the species described by Moore and Jeffords (1945) from the 

Early Pennsylvanian of Texas and Oklahoma, it is 

clearly distinguishable by its larger corallum and 

details of its internal morphology. The mature stage 

of N eokoninckophyllum simplex Moore and Jeffords 

(1945, p. 159, text figs. 149, 150, 156) is smaller, 

has fewer s·epta, and a narrower dissepimentarium 

made up of fewer rows of dissepiments than the 

Amsden species. Moreover, N. simplex lacks a septal 

stereozone and has a less well defined cardinal fossula. 

N. arcuatum Moore and Jeffords (1945, p. 161, 

text figs. 157, 162-164) is also distinguished from 

N. hamatilis by its smaller size and fewer septa at 

maturity. N. arcuatum has longer minor septa, a 

less we11 defined cardinal fossula, a more regular 

axial structure, a more complex dissepimentarium, 

and its tabularium is not so well differentiated from 

the axial structure as inN. hamatiUs. 

N. gracile Moore and Jeffords (1945, p. 162, text 

figs. 158-161) is smaller, has fewer septa, and has 

a thinner dissepimentarium composed of fewer dissepiments than N. hamatili.~. It is also distinguished 

from the Amsden species by its simpler axial structure. 

N. sp. A. Moore and Jeffords (1945, p. 163, text 

figs. 165a-d) differs from N. hamatilis by its smaller 

corallum having fewer septa and its narrower dissepimentarium having fewer dissepiments. Moreover, its axial structure seems to lack an axial plate 

in the ephebic stage. 

Occurrence and abundance.-Horseshoe Shale 

Member: collections 1(2+), 2(3), 3(26+), 4(20), 

5(4), Carbon County, Wyo. 

Specimens.-Holotype, USNM 165095; paratypes, 

USNM 165096-108. 

Neokoninckophyllum inconstans n. sp. 

Plate 6 

Diagnosis.-Solitary corals having a conical 

neanic stage and a cylindrical ephebic stage. Corallum as much as 60 mm or more in length and 27 

mm in maximum diameter and having as many as 

45 major septa at maturity. Exterior marked by 

subdued transverse and longitudinal ornament. 

Corallum supported by talons in early stages, Rejuvenation common. Theca thin. Septal plan pinnate 

in brephic and early neanic stages but radial in 

later stages. Cardinal septum long, located on convex side of corallum in conical stage, and in a very 

poorly defined fossula. Minor septa very short. 

N eanic axial structure consisting of a medial plate 

oriented in the cardinal-counter plane and continuous with cardinal and counter septa, periaxial 

tabellae and tabulae, and a few septal lamellae. 

Ephebic stage without axial structure or with discontinuous medial plate. Tahularium marked by a 

very weak septal stereozone and horizontal inflated tabulae. Dissepimentarium characterized by 

as many as eight rows of regular, herringbone, 

and lonsdaleoid dissepiments. Septal microstructure 

of the diffusotrabecular type of Kato ( 1963). 

Description of holotype.-The holotype (USNM 

165109; pl. 6, figs. 1-7) is an imperfect specimen 

in which neither the top nor the tip of the corralum 

are preserved. The corallum is a gently curved cone 

that expands from an alar diameter of about 5 mm 

to an alar diameter of about 18 mm in a preserved 

length of about 45 mm. The lower 25 mm of the 

corallum is conical and has an apical angle (apex 

not preserved) of about 40°; the upper part of the 

corallum is approximately cylindrical. The cardinalcounter plane coincides approximately with the 

plane of curvature, and the cardinal septum is on 

the convex side of the corallum. The calice, although not completely preserved, was at least 10 

mm deep. External details are lacking on most of 

the specimen owing to decortication, but the exterior seems to have been characterized by subdued longitudinal and transverse ornamentation. 

The internal morphology of this specimen was 

studied by means of six transverse thin sections (pl. 

6, figs. 1-6) cut at intervals of 5 to 8 mm beginning about 3 mm above the base of the specimen. 

Alar diameter, number of major septa, septa 

formula, and ontogenetic stage for each of the 

sections are recorded in table 7. The relation between number of major septa and alar diameter in 

various sections is compared with data from paratypes in figure 4. 

The earliest section studied in this specimen represents the late neanic stage and is characterized 

by 26 major septa at an alar diameter of 7.3 mm 
C16 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

(table 7, section a; pl. 6, fig. 1). The septal plan 

is radial, and the insertion positions are v·ery poorly marked. Cardinal and counter septa are long and 

join the axial plate. The axial structure consists of 

an axial plate, a few tabellae, and the axial ends of 

three major s·epta. Most of the major septa are 

withdrawn from the axial region. The major septa 

are slightly dilated and wedge shaped within the 

tabularium, which is bounded by a weak stereozone. 

The dissepimentarium is thin (maximum width 1 

mm) and consists of two or three rows of herringbone or regular dissepiments. Short minor septa 

(maximum length 0.5 mm) are present in the 

counter quadrants. 

The neanic stage persists to an alar diameter of 

17 mm, where 34 major septa are developed (table 

7, section d; pl. 6, fig. 4). The latest neanic sections are characterized by a radial septal plan, poorly defined insertion positions, and weak dilation in 

the tabularium (table 7, sections c and d; pl. 6, figs. 

3, 4). A sinuous axial plate is still present and is 

connected to the cardinal and counter septa, but l 

EXPLANATION 

X Holotype 

en e Paratype a::: 20 UJ 

1-

UJ 

~ 

::::i 

...1 

~ 16 

z 

ci 

UJ 

1-

UJ 12 

~ 

~ 

c 

a::: 

<( 

...1 8 

<( 

4 

the axial tabellae are reduced in number and less 

regular, and only one or two short septal lamellae 

represent connections to the other major septa. The 

dissepimentarium is as much as 3 mm wide and 

consists of three or four rows of herringbone and 

regular dissepiments of unequal size. The tabularium 

is broad and exhibits few tabular traces. Minor 

septa remain short. 

The ephebic stage (table 7, sections e and f; pl. 

6, figs. 5, 6) is characterized by the nearly cylindrical form of the corallum. At this stage, increase in 

corallum length was greater than increase in corallum diameter. In the holotype, the ephebic stage 

began when the corallum was approximately 18 mm 

in diameter and contained 36 major septa; the 

maximum number of major septa is 39 at the same 

diameter. The ephebic sections are distinguished 

from late neanic sectons by changes in the axial 

structure and dissepimentarium. The axial structure 

is reduced to a short, sinuous medial plate unconnected to the cardinal or counter septa and augmented by few or no accessory tabellae or septal 

• 

NUMBER OF MAJOR SEPTA 

FIGURE 4.-Scatter diagram showing relation between alar diameter and number of major septa by means of 32 measurements on 11 specimens of Neokoninckopkyllum inconstans n. sp. Straight lines connect measurements made from 

serial sections of holotype (USNM 165109) and three paratyp·es (USNM 165111, 165114, 165115). 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C17 

lamellae. In the dissepimentarium, lonsdaleoid dissepiments appear for the first time in addition to 

herringbone and regular dissepiments. The dissepimentarium is 3 or 4 mm wide, and the radius 

of the tabularium, including the weak axial structure, is 5 or 6 mm. The theca is very thin (0.1 mm) 

and the minor septa are very short as in the neanic 

stage. Septal microstructure is of the diffusotrabecular type of Kato (1963). 

Description of paratypes.-The paratypes are all 

incomplete specimens that represent various parts 

of the conical or cylindrical stage of the species. 

Most of the specimens are partly or completely decorticated. The conical stage is mostly slightly 

curved, rarely straight, and no more than 20 to 25 

mm in length. Apical angles range from about 40° 

to 70° ; there are few specimens with a widely 

flaring corallum (pl. 6, fig. 29). Attachment scars 

or talons were observed on most specimens that 

have undecorticated tips preserved (pl. 6, fig. 28). 

The corallum is subcylindrical beyond a length of 

20 to 25 mm and attained a maximum length of at 

least 60 mm. The maximum diameter observed in 

the paratypes is about 27 mm, at which diameter 

42 major septa are present (fig. 4). Rejuvenation 

is common. The cardinal septum is on the convex 

side of the corallum in specimens with regular 

curvature in the conical stage. N eanic calices exhibit a moderately convex tabular floor surmounted 

by a median boss formed by the axial plate and 

septal lamellae (pl. 6, fig. 28). 

Supplementary information on the internal 

morphology of the skeleton was obtained from 24 

transverse thin sections, four transverse peels, and 

two longitudinal thin sections cut from 11 paratypes. 

Serial transverse sections of a paratype (USNM 

165114, table 8; pl. 6, figs. 8-16) illustrate the 

brephic and neanic stages of the species. The earliest section studied (pl. 6, fig. 8), cut about 0.5 

mm above the tip, revealed a single cardinal-counter 

septum. In the next two sections (pl. 6, figs. 9, 10), 

cut about 1.4 and 1.5 mm above the tip, respectively, the alar septa were observed, followed by addition of metasepta in the right counter quadrant. 

Strangely, two counter lateral septa appeared in 

the right counter quadrant before the appearance 

of any metasepta in the left counter quadrant. The 

earliest neanic section studied (pl. 6, fig. 11), cut 

about 2.5 mm above the tip, showed a complement 

of 16 major septa and a continuation of acceleration in the counter quadrants. In these brephic and 

early neanic sections, the septal plan was dominated 

by the coalesced cardinal and counter septa, but no 

TABLE 7.-0ntogeny in the holotype (USNM 165109) of Neokoninckophyllum inconstans n. sp. Tip of specimen not 

preserved 

Section 

f -------

e -------

d -------

c -------

b -------

a -------

Alar 

diameter 

(mm) 

18 

18 

17 

13 

9.6 

7.3 

Number 

of major 

septa 

39 

36 

34 

32 

28 

26 

Septal 

formula 

K 

1019 A--A 

818 

c 

K 

919 

A-~-A 7 7 

c 

K 

818 

A-~-A 7 7 

c 

K 

818 

A-l-A 

616 

c 

K 

717 

A-l-A 

515 

c 

K 

616 

A-l-A 

515 

c 

Ontogenetic 

stage 

Ephebic 

Neanic 

axial structure was evident. In the next neanic section (pl. 6, fig. 12), the septal plan was decidedly 

radial, and a few tabellae were concentric about the 

axially joined cardinal-counter plate to form a primitive axial structure. Succeeding neanic sections 

(pl. 6, figs. 13-16) were characterized by increased 

independence of a weak axial structure composed of 

an axial plate, axial tabellae, and septal lamellae 

and a radial septal plan composed of withdrawn major septa. A few short minor septa appeared in the 

last three sections of the series (pl. 6, figs. 14-16). 

Tabulae were noted in the earliest sections studied, 

but they did not become abundant until later in the 

series (pl. 6, fig. 12). Slight dilation of the major 

septa occurred in the tabularium. The dissepimentarium consisted of a few rows of herringbone and 

regular dissepiments. 

Serial transverse sections of another paratype 

(USNM 165115; table 9; pl. 6, figs. 18-24) show 

features of the neanic stage of the species similar 

to those described above. 

Typical ephebic transverse sections (pl. 6, figs. 25, 

27, 31, 32) show the absence of a discrete axial 

structure at this stage in the development of the 
C18 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

TABLE B.-Ontogeny in a paratype (USNM 165114) of Neokoninckophyllum inconstans n. sp. 

Section 

h -------

g -------

f -------

e -------

d -------

c -------

b -------

a -------

Alar 

diameter 

(mm) 

8 

6.6 

5.6 

3.9 

2.1 

1.5 

1.2 

.9 

Number 

of major 

septa 

28 

26 

23 

19 

16 

6 

5 

2 

Septal 

formula 

K 

717 

A-,-A 

5 5 

c 

K 

617 

A-1-A 

514 

c 

K 

516 A--A 

414 

c 

K 

414 

A-1-A 

413 

c 

K 

314 

A-1-A 

213 

c 

K 

012 

A-1-A 

OIO 

c 

K 

A--A 011 

0 0 

c 

K 

I 

I 

c 

Ontogenetic 

stage 

Neanic 

Brephic 

skeleton. Ordinarily all the major septa are withdrawn from the axial region, but in some sections 

several septa may extend haphazardly to a position 

near the axis (pl. 6, fig. 25). The septal plan is 

radial and insertion positions are so poorly marked 

that they cannot be identified in most specimens. 

The appearance of numerous lonsdaleoid dissepiments is a noteworthy feature of the ephebic stage. 

Dilation of septa in the tabularium is absent or very 

slight. 

Longitudinal sections of the neanic and ephebic 

stages (pl. 6, figs. 26, 33) illustrate the weak and 

variable development of the axial structure in the 

ephebic stage. These sections also show a weak 

differentiation of the tabulae into axial and periaxial zones. The axial tabulae are broadly convex 

upward and tented only where an axial plate is 

present. The periaxial tabulae are convex, flat, or 

concave. The dissepimentarium consists of as many 

as eight rows of gently to steeply inclined inflated 

dissepiments of greatly unequal size. 

Discussion.-N eokoninckophyllum inconstans is 

distinguished from N. hamatilis, the other Amsden 

species, by its smaller corallum and smaller number of major septa at maturity. Internal features 

that serve to differentiate it from N. hamatilis are 

the abundance of lonsdaleoid dissepiments and the 

absence of a continuous axial structure at maturity. 

N. inconstans is also characterized by less dilation 

of the major septa in the tabularium of the neanic 

and ephebic stages than N. hamatilis. 

N. inconstans is distinguished from the species 

described by Moore and Jeffords (1945) from the 

Early Pennsylvanian of Texas and Oklahoma by its 

discontinuous and poorly developed axial structure 

in the ephebic stage and the presence of numerous 

lonsdaleoid dissepiments. It has a generally larger 

corallum than N. simplex Moore and Jeffords and N. 

gracile Moore and Jeffords. It also has a broader 

dissepimentarium and less well defined tabularium 

than N. gracile Moore and Jeffords. 

Occurrence and abundance.-Horseshoe Shale 

Member: collection 7 (256), Carbon County, Wyo. 

Specimens.-Holotype, USNM 165109; paratypes, 

USNM 165110-122. 

Family CY A THOPSIDAE 

Genus C'ANINIA Michelin in Gervais, 1840 

Caninia cf. C. nevadensis (Meek) 

Plate 7, figures 1-8 

Description.-Corals included here are characterized by their large, solitary corallum that is trochoid 

in the early stages and expands rapidly to a cylindrical form at maturity. Fragmentary, commonly 

decorticated specimens suggest that this species 

commonly attained a length greater than 12 em; 

its diameter in the cylindrical stag·e was ordinarily 

between 5 and 6 em, although one specimen has a 

maxim urn diameter of 8 em. 

Major septa ordinarily number between 50 and 

60 at maturity, but there are 68 major septa in the 

largest specimen. The septa extend about two-thirds 

to three-fourths of the distance from the periphery 

of the corallum to the corallum axis at maturity. 

The cardinal septum is shorter than other major 

septa. Dilation in the cardinal quadrants is present 

in the immature stages of the corallum and may 

also be present in mature stages. In most transverse sections of the cylindrical stage, the major 

septa are not continuous through the dissepimentari urn. Minor septa are very short and confined 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C19 

TABLE 9.-0ntogeny in a paratype (USNM 165115) of Neokoninckophyllum inconstans n. sp. Tip of specimen not 

preserved 

Alar Number 

Section diameter of major (mm) septa 

g ------- 9+ 30 

f ------- 8 28 

e ------- 7 26 

d ------- 5 23 

c ------- 4 19 

b ------- 3.1 15 

a ------- 2.3 11 

Septal Ontoformula genetic 

K 

818 

A-l-A 

515 

c 

K 

717 

A-l-A 

515 

c 

K 

617 

A-1-A 

415 

stage 

C Neanic 

K 

616 

A-1-A 

314 

c 

K 

515 

A-1-A 

213 

c 

K 

414 

A-1-A 

1?12 

c 

K 

113 

A-1-A 

112 

c 

to a zone 1 to 4 mm wide at the periphery of the 

corallum. 

The tabularium occupies about one-third of the 

diameter of the cor all urn in rna ture stages. Tabulae 

are variable in size and shape but are generally flat 

with downturned margins. They are spaced 0.5 to 

2 mm apart; one specimen has about 15 tabulae in 

1 em. No axial structure was observed in mature 

stages, but in some trochoid coralla an impersistent 

axial plate is formed by elongation of the counter 

septum (pl. 7, fig. 5). 

The mature dissepimentarium characteristically 

consists of an outer zone of regular and herringbone dissepiments, a medial zone of lonsdaleoid dissepiments, and an inner zone of regular and herringbone dissepiments. The lonsdaleoid zone seems 

to be vertically and horizontally impersistent in 

some specimens. There may be as many as 10 to 

12 rows of dissepiments at maturity. Dissepiments 

are variable in size and shape, globose to elongate, 

and generally steeply inclined. 

Discussion.-There are no complete specimens in 

the material assigned to this species. The fragmentary nature of the specimens makes characterization of the species and appraisal of individual variation difficult. Sando (1965, p. E26-E27) has pointed 

out the difficulties in using the name Caninia nevadensis and the similarities between this species and 

Caninia excentrica (Meek) and Caninia enormis 

Easton. Present data on coral faunas of Chesterian 

age in the Western United States indicate that 

these faunas are characterized by large Caninias 

having curved conical or cylindrical coralla and a 

dissepimentarium that consists of a lonsdaleoid zone 

between regular and herringbone zones. Collections 

studied thus far do not indicate whether the conical 

forms represent a species distinct from the cylindrical forms. Until such time as better material is 

found, it seems desirable to recognize the two 

growth forms as separate species. 

A fourth species that is allied to this group of 

Chesterian Caninias is Caninia montanensis Easton 

(1962, p. 33, pl. 3, figs. 18-20) from the Heath 

Formation of central Montana. This is a diminutive 

conical form that seems to have been derived from 

the Caninia excentrica-Caninia nevadensis stock 

when the Chesterian sea swept into the Big Snowy 

basin from the miogeosynclinal trough in Idaho. 

Occurrence and abundance.-Moffat Trail Limestone Member: collections 73 (1), 74 (18), 76 (1), 

77(6), 78(3), 79(4), 88(2), 91(5), 92(2), 93(2), 

104 (2), 105 (12), 109 (9), Lincoln County; 124 (2), 

126 (6), Teton County, Wyo. 

Specimens.-USNM 165123-141. 

Genus TURBIN A TOCANINIA Dobrolyubova, 1970 

T urbinatocaninia 1 sp. 

Plate 7, figures 9-19 

Description.-This category includes fragmentary 

specimens of small, subcylindrical to vermiform 

corals. Although most of the specimens appear to 

have a solitary habit, one specimen (pl. 7, fig. 19) 

showed evidence of budding. The coralla range from 

about 4 mm to about 20 mm in diameter. The longest 

specimens observed are about 50 mm in length. 

Major septa range in number from 20 in a corallum 6 mm in diameter to 37 in a corallum 18 mm in 

diameter. The major septa ordinarily extend about 

half the distance from the epitheca to the axis of 

the corallum; they are commonly dilated in the 

cardinal quadrants. The cardinal septum is ordinarily shorter or thinner than the other major 

septa, but the cardinal fossula is poorly defined. 

Minor septa are commonly absent in the younger 

stages of the corallum and never attain a length 
C20 THE AMSDEN FORMA•riON (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

greater than about a third of the length of the 

major septa. 

The axial zones of some coralla are characterized 

by an impersistent, sinuous axial plate formed by 

elongation of the counter septum (pl. 7, figs. 11-13, 

15, 16) ; one to three accessory lamellae that may 

or may not join other major septa are commonly 

present. About half the transverse sections examined show a weak and variable axial structure, 

which seems to be most characteristic of coralla 

having fewer than 30 major septa. 

The tabularium comprises all but 1 to 4 mm of. 

the diameter of the corallum and ordinarily consists of flat tabulae with downturned edges (pl. 7, 

figs. 9, 10, 19). Where a columella is present, the 

tabulae may be slightly tented upward (pl. 7, fig. 

15). Tabulae are spaced 0.5 to 1.5 mm. The dissepimentarium consists ordinarily of one to four rows of 

regular or herringbone dissepiments. However, dissepiments are absent in parts of some young coralla, 

and a few lonsdaleoid dissepiments are present in 

some specimens. 

Discussion.-The material at hand poses difficult 

taxonomic problems owing to its fragmentary nature 

and its morphologic variability. Some of the smaller 

cylindrical fragments were originally interpreted as 

disaggregated corallites of the lithostrotionid genus 

N emistium. This interpretation was abandoned when 

coralla of compar8!ble diameter and morphology 

were found to be parts of larger, distinctly solitary 

specimens having a weak cardinal septum and dilated major septa in the cardinal quadrants. 

The general morphology of this species is suggestive of the genus Caninia as this term is generally 

used in a broad sense. Characteristic features are 

the broad tabularium, peripheral dissepimentarium, 

major septa that ordinarily do not reach the axis of 

the corallum, weak cardinal septum, dilation of 

major septa in the cardinal quadrants, and principally solitary habit. On the other hand, the presence 

of an impersistent columella in some parts of the 

corallum is inconsistent with the commonly accepted 

definition of Caninia. Dobrolyubova (1970) described a similar inconsistency of morphology in 

some corals from· the Visean of the U.S.S.R., for 

which she proposed the genus Turbinatocaninia. 

According to Dobrolyubova (1948), these corals 

form a part of a phylogenetic series from Dibunophyllum through Koninckophyllum to Caninia. 

Occurrence and abundance.-Moffat Trail Limestone Member: collections 73 (7), 75 (1)), 76 (2), 

77(2), 78(3), 79(10), 88(10), 91(4), 92(1), 93(1), 

103(7), 105(63), 109(7}, Lincoln County; 124(4), 

Teton County, Wyo. 

Specimens.-USNM 165144-166. 

Suborder COLUMNARIINA 

Family LONSDALEIIDAE 

Subfamily LONSDALEIINAE 

Genus LONSDALEIA McCoy, 1849 

Subgenus ACTINOCYATHUS d'Orbigny, 1849 

1849. Actinocyathus d'Orbigny, p. 12. 

1861. Stylidophyllum de Fromentel, p. 316. 

Type species.-Cyathophyllum crenulare Phillips, 

1836 equals Erismatolithus M adreporites (floriformis) Martin, 1809 equals Lonsdaleia floriformis 

(Martin). 

Diagnosis.-Cerioid Lonsdaleia. 

Discussion.-Most students of Paleozoic corals 

have included both fasciculate and massive species in 

the genus Lonsdaleia. However, as early as 1861, de 

Fromentel proposed the genus Stylidophyllum to distinguish massive corals with the internal structure 

of Lonsdaleia, whose type species is a fasciculate 

form. In recent years, beginning with Chi (1931), 

many authors have used de Fromentel's concept in 

describing corals of Carboniferous and Permian age 

in China, Japan, Iran, the Carnic Alps, and the 

U.S.S.R. The concept has been applied mostly commonly to Permian corals of the Tethyan region ; most 

of these species have been reallocated most recently 

to Yokoyamaella Minato and Ka.to, lpciphyllum Hudson, and Wentzelophyllum Hudson by Minato and 

Kato (1965). 

Smith (1916, p. 257) studied a specimen that he 

considered to be the holotype of Cyathophyllum crenulare Phillips, the type species of Actinocyathus 

d'Orbigny, and concluded that it "is a characteristic 

specimen of Lonsdaleia floriformis ," the most widely 

known cerioid species of Lonsdaleia. Kato (1966, p. 

102) studied the same specimen, which he designated as lectotype of Phillips' species, and also concluded that it is conspecific with Lonsdaleia floriformis (Martin). Inasmuch as Stylidophyllum de 

Fromentel, 1861, is based on the same species (by 

subsequent designation of Chi, 1931, p. 44), Stylidophyllum is a junior synonym of Actinocyathus. 

Kato (1966, p. 100-101) discussed the question of 

the validity of growth form as a generic character in 

the group of corals ordinarily referred to Lonsdaleia and concluded that the name Actinocyathus 

should be applied to the cerioid forms while retaining Lonsdaleia for the fasciculate forms. Although 

this question is controversial, I agree with Kato that 

growth form is a useful taxonomic distinction in 

this group of corals. However, until there is further 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C21 

clarification of the phylogenetic significance of 

growth form, I propose to give this character only 

subgeneric rank. Accordingly, I divide the genus 

Lonsdaleia McCoy into subgenus Lonsdaleia McCoy 

(fasciculate forms) and subgenus Actinocyathus 

d'Orbigny ( cerioid forms). 

Although both subgenera of Lonsdaleia have been 

reported in beds ranging from Early Carboniferous 

to Permian in age, recent taxonomic revisions of 

Middle Carboniferous and Permian corals have resulted in reallocation of most of the post-Early Carboniferous species formerly assigned to this genus. 

It seems probable that further revisions will result 

in restriction of Lonsdaleia to the Lower Carboniferous. 

Lonsdaleia (Actinocyathus} stelcki (Nelson} 

Plate 8 

1960. Lithostrotionella stelcki Nelson, p. 119-120, pl. 23, 

figs. 6-10. 

1963. Lithostrotionella cf. L. stelcki Nelson. Sando in Dutro 

and Sando, p. 1982. 

1969. Lithostrotionella cf. L. stelcki Nelson. Sando in Sando, 

Mamet, and Dutro, p. Ell. 

Diagnosis.-Actinocyathus having corallites that 

average between 5 and 13 mm in diameter and contain an average of between 14 and 26 major septa. 

Minor septa absent to well developed. Axial structure weakly arachnoid, composed of an axial plate 

that originates as an extension of one or two major 

septa, a variable number of septal lamellae, and 

axial tabellae; entire complex commonly strongly 

reinforced by stereoplasm, forming a solid axial rod 

that is oval to circular in transverse section and 1 

to 2 mm in diameter. Tabulae flat to concave upward. Axial tabellae nearly vertical to vertical. Dissepiments variable in size, lonsdaleoid, in one to 

three poorly defined row.s. Corallite walls strongly 

corrugated. 

Description.-The Amsden specimens are all fragments of colonies; the largest corallum observed in 

the field was about 8 inches high and 2 feet in diameter. Budding was peripheral. Corallite walls are 

strongly corrugated. 

Corallites range from 3.2 to 10.5 mm in diameter; 

average corallite diameter per corallum ranges from 

5.6 to 7.5 mm. Major septa range in number from 

10 to 24; average septal number per corallite per 

corallum ranges from 14.5 to 19.8. 

Major septa are ordinarily confined to the tabularium, except for one or two (presumably counter 

and cardinal) that commonly join the axial plate. 

Extensions into the dissepimentarium as septal 

crests are rare. Minor septa are generally rare or 

absent; where present they are one-fourth to onehalf the length of the major septa. 

The axial structure is composed of an axial plate 

that is continuous with 1 or 2 major septa, 1 to 10 

septal lamellae, and nearly vertical axial tabellae. 

Stereoplasmic thickening of the axial complex is 

variable within the colony; most coralla have at 

least a few corallites with solid axial rods, but many 

corallites show little or no thickening. The axial 

structure is oval to circular in transverse section 

and 1 to 1.5 mm in diameter. 

The tabularium ranges from 1.8 to 5.4 mm in 

diameter and has an average diameter per corallum 

of 3.3 to 4.1 mm. Tabulae are mostly concave upward 

but some are horizontal and a few show slight upward convexity. The tabulae terminate sharply at 

their junctions with the dissepimentarium and the 

axial complex. There are about three to five tabulae 

in 1 mm vertically. 

The dissepimentarium consists of one to three 

variable and poorly defined rows of lonsdaleoid dissepiments. The dissepiments are variable in size, 

moderately to strongly inflated, and inclined from 

the corallite walls at an angle of about 30° to 45° 

from the horizontal. 

Discussion.-Cerioid rugose corals are relatively 

rare in rocks of Chesterian age in western North 

America; the only corals of this description have 

been referred to Lithostrotionella stelcki Nelson. 

Previous allocation of this species to Lithostrotionella Yabe and Hayasaka was due to the weakly 

arachnoid nature of the columella and the pronounced stereoplasmic thickening of the columella 

in many corallites. Careful study of many transverse 

thin sections shows that all elements of the complex 

columella characteristic of Lonsdaleia are present in 

contrast to the simpler columella of Lithostrotionella, 

which lacks persistent axial tabellae and discrete 

septal lamellae. 

In order to determine the degree and nature of 

morphological variation in this species, I studied 

10 colonies of the species from the Western United 

States and 4 colonies from western Canada (through 

the courtesy of E. W. Bamber of the Geological Survey of Canada). These were compared with Nelson's 

holotype and paratype, obtained through the courtesy of C. R. Stelck of the University of Alberta. 

Data from this study are shown in table 10 and 

figure 5. 

Conclusions drawn from the study are as follows: 

1. All the specimens studied have the distinctive 

tabulation and complex axial structure that 
C22 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

characterize the type specimens of Lonsdaleia 

(Actinocyathus) stelcki. The specimens are 

also characterized by their corrugated corallite 

walls and one to three rows of variable lonsdaleoid dissepiments. 

2. Average corallite diameter per colony, average 

number of major septa per corallite per corallum, development of minor septa, and stereoplasmic thickening of the axial structure are 

variable features in the material studied. 

3. A graph of average corallite diameter per colony 

plotted against average number of major septa 

per corallite per corallum (fig. 5) indicates 

that most of the colonies cannot be separated 

into more than one discrete taxonomic entity 

on these characters. Only one colony ( GSC 

49·354) is sufficiently distinct to warrant consideration as a separate taxon. No geographic 

variants are di,scernible on the graph. 

en 

0::: 

IJ.J 

rIJ.J 

~ 

::J 

...J 

~ 

z 

ci 

IJ.J 

rIJ.J 

~ 

<( 

0 

IJ.J 

r-

::J 

...J 

<( 

0::: 

0 

() 

IJ.J 

" <( 

0::: 

IJ.J 

> <( 

I EXPLANATION I • Amsden Formation, Wyoming 

13 

.& Monroe Canyon Limestone, Idaho 

1- I • Surrett Canyon Formation, Idaho 

Q Green Ravine Formation, Utah 

0 Etherington Formation, Alberta 

and ritish Columbia 11 

~ 

9 

1-

USGS 18715 GSC 57916t:"' 

• 7 

• USGS 20249 USNM 165167 El 

USGS 23635 • eusNM 165168 

0 USGS 23636 USGS[:J 

1-

5 

14 

• GSC 53435 16329 

us~s 19897 •

1 

USNM 165169. 

I 

16 18 

I 

USNM 165171 e 

0 GSC 49388 

UA 330 0 

(p'aratype) 

I 

4. Variations in development of minor septa and 

thickening of the columella are similarly unsystematic and seemingly unrelated to corallite 

diameter and septal number (table 10). 

5. These data support inclusion of all the material 

studied in one species with the possible exception of GSC 49354, which is distinguished by 

its much larger corallites and greater septal 

number per corallite. Inasmuch as GSC 49354 

exhibits other characters that unite it with 

the majority of the specimens, it might be 

practical to recognize it as a distinct subspecies. However, formal recognition is deferred pending discovery of more and better 

specimens of this variant. 

Occurrence and abundance.-Moffat Trail Limestone Member: collections 90(6), 92(2), 104(4), 

105 (1), Lincoln County, Wyo. 

Specimens.-VSNM 165167-173. 

I I I 

GSC 49354 0 

-

-

-

UA 329 

t:"' ( holotype) 

-

1 I I 

20 22 24 26 

AVERAGE NUMBER OF MAJOR SEPTA PER CORALLITE 

FIGURE 5.-Variation in average corallite diameter per colony and average number of •major septa per corallite in colonies of Lonsdaleia (Actinocyathus) stelcki (Nelson) from beds of Chesterian age in western North America. Numbers refer to U.S. National Museum collections (USNM), U.S. Geological Survey collections (USGS), Canada 

Geological Survey collections (GSC), and University of Alberta collections (DA). 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C23 

TABLE 10.-Variation ·in colonies of Lonsdaleia (Actinocyathus) stelcki (Nelson) from tke United States and Canada 

"" 

Locality or ~ s specimen Formation Area Ill-. 

number :as 

~s ~._.. 

=~ Ill be ""= O..:s 

o ... 

USNM 165171 _____ Amsden ___ Wyoming -------- 3.5-10.5 

USNM 165168 ----- ___ do ________ do ------------ 3.2-10.0 

USNM 165167 ----- ___ do ________ do ------------ 4.5-9.0 

USNM 165169 ----- ___ do ----- ~--do ------------ 4.5-6.0 

USGS 18715-PC --- Monroe Idaho ------------ 2.8-8.5 

Canyon. 

USGS 19897-PC ___ Surrett ___ do ·------------ 3.5-8.0 

Canyon. 

USGS 23636-PC --- ___ do ----- ___ do ------------ 5.0-8.0 

USGS 2·3635-PC --- ___ do ----- ___ do ------------ 4.5-7.5 

USGS 16329-PG --- Green Utah ------------ 3.5-7.5 

Ravine. 

USGS 20249~PC --- ___ do ________ do ------------ 5.5-9.0 

Holotype, U A 329 __ Etherington British Columbia _ 5.0-9.0 

Paratype, UA 330 ____ do _____ Alberta ---------- 4.0-8.0 

GSC 57916 -------- ___ do ________ do ------------ 5.0-8.0 

GSC 49354 -------- ___ do _____ British Columbia _ 11.0-14.0 

GSC 49388 -------- ___ do _____ Alberta ---------- 5.0-9.0 

GSC 53435 -------- ___ do _____ British Columbia _ 3.0-8.0 

Order TABULATA 

Family SYRINGOPORIDAE 

Genus PLEUROSIPHONELLA Cbudinova, 1970 

Type species.-Pleurosiphonella crustosa Chudinova (1970, p. 105, pl. 38, figs. 1, 2). 

Diagnosis.-Corallum fasciculate, phaceloid or 

dendroid, composed of relatively thick walled, cylindrical corallites that are joined at irregular intervals 

and connected by mural pores or very short connecting tubules. Corallites commonly joined to form 

short chains or clusters at some levels in the corallum. Increase lateral, commonly multiple. Tabulae 

"" ... 

~ ~ 

.~ 1 .~ "" 

s-- '0 :as Ill 

~~~s s-. s~ :as =s ss 'l-lbl) 

~~ ..... s .::!'-' os= 

""~ ""Ill it-' .!!~ .!': -~~~ 111~-o -~ ]be ,.6~-o S!! ""~ ~a ~~ z~ 0> O..:s 

7.5 2.2-5.4 3.9 13-24 

6.4 1.8-4.3 3.3 12-18 

6.8 2.5-5.0 3.6 10-17 

5.6 3.5-4.5 4.1 16-21 

6.9 2.5-4.5 3.6 12-18 

5.8 2.5-4.5 3.6 14-18 

6.3 2.0-4.0 3.1 13-18 

6.1 3.0-5.0 3.9 12-21 

6.2 2.5-4.0 3.4 15-20 

6.6 3.0-4.0 3.5 16-20 

7.0 3.5-5.0 4.4 18-23 

6.4 3.0-4.5 3.8 16-21 

7.0 3.5-4.5 4.1 16-20 

12.6 5.0-6.0 5.6 24-28 

6.9 3.0-4.5 3.8 16-21 

6.1 2.0-4.0 3.2 12-20 

... 

.~ 

~~~~ Sbe 

~~ ~-o> 

~~~~ 

~i =+' 

z~ 

19.8 

15.8 

14.5 

17.6 

15.0 

15.9 

15.5 

14.3 

17.8 

17.4 

20.8 

19.3 

17.6 

25.7 

18.2 

15.3 

Minor septa Axial 

structure 

Variable, Variable, 

generally unthickwell de- ened to 

veloped. solid. 

Generally Variable, 

absent or rarely 

short. solid. ___ do _____ Do. 

Absent or Unthickshort. ened. ___ do ----- Variable, 

generally 

Well dev-eloped. 

Variable, 

generally 

well developed. 

Absent or short. 

Variable, 

generally 

short or absent. 

Variable, 

g·enerally 

present 

but short. 

solid. 

Do. 

Variable, 

rarely 

solid. 

Variable, 

unthickened to 

solid. 

Variable, 

generally 

solid. 

Do. 

Well de- Generally 

veloped. solid. ___ do Do. 

___ do ----- Variable, 

generally 

solid. ___ do ----- Generally 

solid. ___ do _____ Variable, 

generally 

solid. 

Absent or Variable, 

solid. unthickened to 

solid. 

mostly vesicular, ordinarily vertically disposed along 

one side of the corallite wall in such a way as to 

leave a visceral canal along the other side or tabulae 

may be infundibular, having the axis of the funnels 

closer to one side of the corallite than the other. 

Septal spines rare or absent in some species, present 

in others. Wall structure fibrous. 

Discussion.-Nelson (1962, p. 444) distinguished 

a group of three syringoporid species in the Mississippian of western Canada that are characterized 

by eccentric visceral canals, relatively thick walls, 

and corallites joined by anastomosis. He included 
C24 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

Sy1~ingopora virginica Butts, S. magnussoni Nelson, 

and S. drummondi Nelson in this group but did not 

propose a new generic name for it. Chudinova 

( 1970) established the name Pleurosiphonella for 

similar forms from the Tournaisian of Armenia. 

Having noted such corals in the Amsden Formation 

of Wyoming, I examined the type specimens of the 

type species through the courtesy of Dr. Chudinova 

when I was in Moscow in 1971 and concluded that 

the American and Armenian forms are congeneric. 

According to Chudinova (oral commun., 1971), 

the genus is known from the Tournaisian of Armenia and the Tournaisian and Visean of the Ural 

Mountains. Lonsdale ( 1845, p. 592-593) described a 

specimen from the Carboniferous Limestone at 

Ilinsk, U.S.S.R. under the name of Syringopora distans (Fischer von Waldheim) in which he noted eccentric visceral canals. Although this may belong in 

Pleurosiphonella, I have not been able to verify the 

identification. On the other hand, Pickett (1966, 

p. 36, pl. 20, figs. 3, 4) described and illustrated 

Syringopora pachytheca Pickett from the Halls 

Creek Limestone (middle Visean, cu 11,8-y) in the 

New England District, New South Wales, which is 

characterized by an eccentric visceral tube and is 

here included in the genus. Pickett's species is distinguished from other species of Pleurosiphonella 

by its very thick deposits of stereoplasm between 

the visceral canal and the corallite wall. 

According to Nelson (1962, p. 444), the Canadian 

species here referred to Pleurosiphonella are most 

common in the Meramecian and rare in the late 

Osagean. However, a revision of the position of the 

Osagean Meramecian boundary by Macqueen, Bamber, and Mamet (1972) indicates that the Canadian 

occurrences are exclusively post-Osagean, which is 

more in accord with the writer's knowledge of the 

range of the genus in the United States. 

Pleurosiphonella virginica is the most common 

North American species. Nelson (1962, fig. 7) recorded it in the Livingstone (upper part), Mount 

Head, and Etherington (lower part) Formations of 

western Canada in beds of Meramecian and Chesterian age. The species was originally described 

from strata equivalent to the St. Louis Limestone 

in Virginia. In the Western United States, it occurs 

in the Monroe Canyon Limestone of southeast Idaho 

and the Scott Peak Formation of south-central 

Idaho. Its known range in the Western United 

States is in Zones E and F of Sando, Mamet, and 

Dutro (1969), an interval of middle and late Meramecian age (equivalent to V2b through V3b of the 

Visean of Europe). 

Pleurosiphonella magnussoni ranges from the 

upper part of the Livingston Formation through 

the Mount Head Formation and into the lower part 

of the Etherington Formation (Nelson, 1962, fig. 7). 

The age range is middle Meramecian into the early 

Chesterian (equivalent to parts of the middle and 

upper Visean of Europe). This species has not been 

identified in the United States. 

Pleurosiphonella drummondi was recorded by N elson (1962, fig. 7) exclusively in the upper part of 

the Etherington Formation of Chesterian age in 

western Canada. It is described below from the 

Amsden Formation of western Wyoming, where it 

occurs in Foraminiferal Zones 17 and 18, an interval of late Chesterian age (equivalent to E1 and E2 

of the Namurian of Europe). 

Pleurosiphonella drummondi (Nelson} 

Plate 9, figures 1-4 

1962. Syringopora drummondi Nelson, p. 458, pl. 72, figs. 

12-14; pl. 75, figs. 8-11; text fig. 4i. 

Description.-Specimens from the Amsden are 

fragments of colonies; the largest corallum observed 

was at least 6 em high and 16 em in diameter. Corallites are subparallel and connected at irregular intervals of about 1 em by short connecting tubules 

or by mural pores without connecting tubules. 

In transverse sections, corallites have circular to 

oval cross sections and appear to be connected in 

short chains at some levels in the corallum. Corallite 

diameters range from 1.8 to 2.8 mm; average corallite diameters in two measured coralla are 2.4 mm 

and 2.3 mm, respectively (fig. 6). Corallites are 

spaced as much as 4 mm apart; the average corallite 

frequency in two measured coralla is 7.8 and 8. 7 

corallites per square centimeter (fig. 6). Wall thickness ranges from 0.1 to 0.5 mm. The microstructure 

of the wall appears to be fibrous, with fibres radially 

arranged. Tabular traces are numerous and concentric, having the center of concentricity located 

closer to one side of the corallite. Septal spines are 

abundant in one specimen and seemingly absent in 

another, although poor preservation may be a factor 

in the latter. Where present, the spines are as much 

as 0.3 mm long and located on both the corallite wall 

and on tabulae. 

Occurrence and abundance.-Moffat Trail Limestone Member: collections 78 (1), 109 (2), Lincoln 

County, Wyo. 

Specimens.-USNM 16517 4-176. 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C25 

Family MULTITHECOPORIDAE 

Genus MUL TITHECOPORA Yob, 1927 

Multitbecopora 1 amsdenensis n. sp. 

Plate 9, figures 5-9 

Description.-The Amsden specimens are all 

fragments of coralla ; the largest corallum was at 

least 5 em high and 11 em in diameter. The coralla 

are all phaceloid to subdendroid; most of the corallites are subparallel. Corallites are subcylindrical; 

their cross sections range from circular to rounded 

polygonal. The epitheca of individual corallites is 

marked by rugae, fine growth lines, and commonly 

shows abundant talons. Corallites are connected by 

short connecting tubules or by mural pores where 

the corallite walls are in close contact with one 

another. There are commonly one or two connecting 

tubules or mural pores at a given level in a corallite, 

although there may be as many as four. These connections between corallites occur commonly at intervals of about 4 mm. Increase is commonly multiple; 

as many as three offsets may arise at one level in 

a corallite. 

Corallites range from 1.5 to 2.7 mm in diameter; 

average corallite diameter measured in thin sections 

of five coralla ranges from 1.9 to 2.2 m·m (fig. 6). 

Corallites are spaced as much as 5 mm apart, and 

average corallite frequency measured in five coralla 

ranges from 7.1 to 13.3 corallites per square centimeter (fig. 6) . Wall thickness is extremely variable, 

ranging from 0.1 to 1.0 mm in five coralla. Concentric layers of stereoplasm are common in thickened 

walls of some corallites, but the visceral cavity ordinarily occupies at least one half to two-thirds the 

diameter of the corallite. 

Tabulae are relatively sparse and erratically distributed in groups of 2 to 15 spaced at vertical intervals of 2 to 10 mm. They are ordinarily complete 

and horizontal and flat to upwardly concave. In 

transverse sections the tabular intercepts are generally concentric, and the centers of concentricity 

range from axial to excentric in position. 

Septal spines are rare, no more than 0.1 mm in 

length, and arise from the walls of the corallites. 

None were observed within the stereoplasmic thickening of the corallite walls. 

Discussion.-The exact taxonomic position of the 

new species is in doubt. The general form of the 

corallites, the thick corallite walls, the nature of the 

tabulae, and the wrinkled epitheca are features that 

suggest placement in the Family Multithecoporidae 

as interpreted by Sokolov (1950, p. 28; 1955, p. 199; 

1962, p. 237). Although the species is referred provisionally to Multithecopora Yoh, it lacks the extreme thickening of the corallite walls and consequent narrowing of the visceral cavity to one-half 

to one-third the diameter of the corallite that characterize other species referred to this genus. The 

Amsden species is also distinguished by having septal spines and moderately abundant connections 

between corallites in the form of short connecting 

tubules and mural pores. 

The Amsden species is also similar to species assigned to Neomultithecopora Lin (see interpretation 

by Kachanov, 1967) but differs in lacking extreme 

thickening of the corallite walls and in the absence 

of spines within the zone of stereoplasmic thickening. 

Relationship to the Devonian genus Syringoporella 

Kettner is obscure because of the lack of detailed 

information on this genus. The Devonian forms 

appear to be characterized by much fewer tabulae, 

thicker connecting tubules, and they apparently 

lack the lamellar wall structure that characterizes 

the Carboniferous forms. 

Multithecopora? amsdenensis is associated with 

foraminifers of Foraminiferal Zones 17 and 18, an 

interval of Chesterian age equivalent to E1 and E2 

of the Namurian of Europe. In addition to its occurrence in the Moffat Trail Member of the Amsden 

Formation, it has also been found in the Big Snowy 

Group at Baldy Mountain in southwest Montana. 

Occurrence and abundance.-Moffat Trail Limestone Member: collections 75 (1), 77 (1), 80 (2), 

105 (1), Lincoln County; 122 (1), 126 (1), Teton 

County, Wyo. 

Specimens.-Holotype, USNM 165177; paratypes, 

USNM 165178-184. 

Family THECOSTEGITIDAE Fromentel 

Genus DUNCANOPORA n. gen. 

Type species.-Duncanopora duncanae n. sp. 

Diagnosis.-Corallum pha.celoid and cerioid, composed of cylindrical corallites that are joined at 

regular intervals in the corallum by lateral expansions or connecting structures. Regular connections 

within the corallum may be by simple increase in 

diameter of corallites, which crowds the corallites 

together to produce polygonal cross sections, or by 

connecting tubules or encircling deposits of sclerenchyme perforated by pores connecting adjacent corallites. Increase lateral. Tabulae sparse, ordinarily 

complete, horizontal, and slightly concave upward. 

Septal spines rare. 

Discussion.-This new genus is referred to the 
C26 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

z 

0 

1-4 

E-1 

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r:il 

00 

r:il 

00 

~ 

r:il 

> 36 00 

z .. < 

~ 

E-1 

~ 

0 

~ 

r:il 32 

E-1 

r:il 

~ 1-4 

E-1 30 

z 

r:il 

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r:il 28 ~ 

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00 26 

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A-t 

00 24 

r:il 

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...::l 22 < 

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20 

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r:il 12 

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1.0 1.5 2.0 2.5 

AVERAGE CORALLITE DIAMETER, IN MILLIMETERS 

FIGURE 6.-Corallite diameter and frequency in syringoporoid corals from the Amsden Formation. Each diameter value 

represents the average of 20 measurements in a transverse section of a single corallum. Frequency values represent the average of 2 to 10 measurements in transverse sections of coralla. 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C27 

Family Thecostegitidae because of its regularly connected corallites, a feature characteristic of the family. It thus becomes the youngest representative of 

the Thecostegitidae, which was known previously 

only from the Silurian and Devonian. It appears to 

be most similar to Thecostegites Milne-Edwards and 

Haime but is distinguished from the latter by its 

more restricted communication between corallites 

through the connecting processes and by its sparse 

tabulae. It also differs in having offsets that arise 

laterally rather than from connecting structures. 

The tabulation of Duncanopora is similar to that of 

the Multithecoporidae as interpreted by Sokolov 

(1950), but the me.mbers of this family have extremely thickened walls and lack abundant and regular connections between corallites. 

The Early Carboniferous genus Fuchungopora 

Lin (1963, p. 587, 594) has connecting structures 

similar to those of Duncanopora but has complex 

and abundant infundibular tabulae which sharply 

contrast with the simple tabulation of the Amsden 

forms. In North America, corals similar to Fuchungopora were described as Syringopora mccutcheonae 

by Wilson and Langenheim (1962, p. 515-516, pl. 

89, figs. 11-13) on specimens from the Permian of 

Nevada. 

Duncanopora is presently known only from its 

type species, which is described below. This species 

occurs in the Amsden Formation of western Wyoming, the Monroe Canyon Limestone of southeast 

Idaho (recorded as Tetraporinus? sp. by Sando in 

Dutro and Sando, 1963, p. 1982), the Surrett Canyon 

Formation of south-central Idaho, and the Green 

Ravine Formation of west-central Utah. The species 

was recorded as H ayasaka1:a? sp. by Sando in Sando 

and others (1969, p. Ell, fig. 3). All known occurrences are in Zone K of Sando and others, (1969), 

an interval of Chesterian age (equivalent to V3cs of 

the Visean through E2 of the Namurian in Europe). 

Foraminifera in three of the Amsden samples indicate Foraminiferal Zones 17 and 18, which suggests 

that the species may be restricted to lower N amurian (E1 and E2) equivalents. 

Duncanopora duncanae n. sp. 

Plate 10 

1963. Tetraporinus? sp. Sando in Dutro and Sando, p. 1982. 

1969. Hayasakaia? sp. Sando in Sando and others, p. Ell, 

fig. 3. 

Descn:ption.-The Amsden specimens are all fragments of coralla ; the largest and most complete 

specimen in the collections suggests that the species 

is characterized by a discoidal corallum about 5 to 

6 em high and as much as 18 em in diameter. Corallites are subparallel; they are subcylindrical except 

for regular layers in the corallum where they are 

expanded and cerioid. Cross sections of corallites 

range from circular to polygonal. Walls of individual corallites are distinct in cerioid levels of the 

corallum. Corallites are connected by mural pores 

in the cerioid layers of the corallum; these layers 

are commonly 2 to 5 mm thick. In the intervening 

phaceloid layers of the corallum, the corallites are 

connected by means of abundant short connecting 

tubules that have a vertical spacing of 1 to 3 mm. 

Connecting tubules arise at regular levels in the 

corallum; they are augmented by sclerenchyme that 

forms imperfect platforms in most specimens. There 

are commonly two to four connecting canals in a 

single corallite at a given connecting tubule level. 

Increase is commonly multiple; as many as three 

offsets may arise at one level in a corallite. 

Corallite diameters range from 1.1 to 1.7 mm in 

the phaceloid layers of the corallum; the variation 

within a given corallum is 0.3 to 0.5 mm. The average diameter of phaceloid corallites ranges from 1.3 

to 1.6 mm, based on measurements of transverse 

sections of eight coralla (fig. 6). Corallite diameters 

in cerioid layers of the corallum range from 1.5 to 

2.5 mm; the variation within a given corallum is 

0.5 to 0.9 mm. The average diameter of cerioid corallites ranges from 1.7 to 2.0 mm, based on measurements of transverse sections of four of the cor~:tlla 

in which phaceloid measurements were made (fig. 

6). Average corallite frequency in phaceloid layers 

ranges from 18 to 43 corallites per square centimeter, measured in transverse sections of 8 coralla 

(fig. 6). In cerioid layers, the average corallite 

frequency is about the same as in phaceloid layers; 

it ranges from 21 to 36.6 corallites per square centimeter, measured in 4 coralla (fig. 6). Wall thickness is extremely variable, ranging from 0.1 to 0. 7 

mm; the walls of most corallites are 0.3 mm or less 

in thickness. Concentric layers of stereoplasm are 

common in thickened walls of some corallites, but 

the diameter of the visceral cavity is ordinarily at 

least one-half to three fourths diameter of the 

corallite. 

Tabulae are ordinarily complete, horizontal, and 

gently concave upward, although some may be 

slightly inclined from the horizontal and flat or 

gently convex upwards. They may be erratically 

distributed or in groups of 2 to 20, separated by 

nontabulate intervals of 1 to 4 mm; they appear 

to be concentrated at connecting tubule levels in 
C28 THE AMSDEN FORMATION (MISSISSIPPIAN AND PENNSYLVANIAN) OF WYOMING 

some specimens. In transverse sections the tabular 

intercepts are concentric or nonconcentric, and 

where concentric the centers of concentricity range 

from axial to excentric in position. 

Septal spines are ordinarily rare, no more than 

0.1 mm in length, and arise from the walls of the 

corallites. 

Occurrence and abundance.-Moffat Trail Limestone Member: collections 76(1), 77(3), 80(2), 

88(1), 91(1), 92(1?), 93(1?), 104(2), 109(1?), 

Lincoln County; 126 (1), Teton County, Wyo. 

Specimens.-Holotype, USNM 165184; para types, 

USNM 165185-197. 

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Chudinova, I. I., 1970, Novyye tabulyaty iz paelzoya Zakavkaz'ya [New tabulate corals from the Paleozoic of 

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126 p., 14 pis., 1 text fig. 

Fischer von WaJdheim, G., 1829, in Eichwald, C. E. von, 

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the fossil faunas of USSR] tom 5, Sredniy i verkhniy 

otdely kamennougol'noy sistemy [Middle and Upper Carboniferous], Tsentral'nyy Nauchno-issledovatel'skiy Geologo-razvedochniy Institut, Leningrad, p. 50-64, pis. 

6-11. 

Fromentel, E. de, 1861, Introduction a I'etude des polypiers 

fossiles : Paris, 357 p. 

Hill, Dorothy, 1940, A monograph of the Carboniferous rugose corals of Scotland, pt. 3: Palaeontographical Soc. 

London, p. 115-204. 

--- 1956, Rugosa in Moore, R. C., ed., Treatise on invertebrate paleontology, pt. F, Coelenterata: Geol. Soc. 

America and Kansas Univ. Press, p. 233-324. 

Hill, Dorothy, and Stumm, E. C., 1956, Tabulata in Moore, 

R. G., ed., Treatise on invertebrate paleontology, pt. F, 

Coelenterata: Geol. Soc. America and Kansas Univ. 

Pres~s, p. 444-477. 

Hudson, R. G. S., 19,41, On the Carboniferous corals-Zaphrentis carruthersi sp. nov. from the Mirk Fell beds and 

its relation to the Z. delanouei species-group: Yorkshire 

Geol. Soc. Proc., v. 24, pt. 4, p. 290-311, pls. 21, 22, 1 fig. 

Kachanov, Ye. I., 1967, Novyye dannyye o korallakh roda 

N eomultithecopora iz nizhnego karbona N ovoy Zemli i 

Urala [New data on corals of the genus Neomultithecopora from the lower Carboniferous of N ovaya Zemlya 

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ill us. [Am. Geol. Inst. translation]. 

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571-630, 3 pls., 19 text figs. 

--- 1966, Note on some Carboniferous coral generaClisaxophyllum, Clisiophyllum (Neoclisiophyllum), Zaphrentoides, Stylidophyllum, and Actinocyathus: J apanese Jour. Geology and Geography, v. 37, nos. 2-4, p. 

93-104, pl. 3. 

Lane, N. G., and Martin, R. A., 1966, Redescription of 

Chaetetes milleporaceus Milne-Edwards and Haime, 

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corals of Russia in Murchison, R. I., Verneuil, Edouard, 

and Keyserling, Alexander, The Geology of Russia in 

Europe and the Ural Mountains, v. 1, p. 591-634, illus., 

London. 

Macqueen, R. W., Bamber, E. W., and Mamet, B. L., 1972, 

Lower Carboniferous stratigraphy and sedimentology of 

the southern Canadian Rocky Mountains: 24th Internat. 

Geol. Cong., Montreal, 1971, Guidebook Excursion C17, 

62 p., 7 figs. 

Martin, W., 1809, Petrificata derbiensa; or, figures and 

descriptions of petrifactions collected in Derbyshire: 

102 p., illus., Wigan. 

McCoy, F., 1849, On some new genera and species of Paleozoic corals and Foraminifera: Annals and Mag. Nat. 

History, 2d. ser., v. 3, p. 1-20. 

Michelin, J. L. H., 1840, in Gervais, P., Astree, Astraea, 

Diet. Sci. Nat. Paris, Supplement 1, p. 481-487. 

Minato, M., and Kato, M., 1965, Waagenophyllidae: Hokkaido Univ. Fac. Sci. Jour., ser. 4, v. 12, nos. 3-4, 241 p., 

ill us. 
COELENTERATA OF THE AMSDEN FORMATION OF WYOMING C29 

Moore, R. C., and Jeffords, R. M., 1945, Description of Lower 

Pennsylvanian corals from Texas and adjacent states: 

Texas Univ. Pub. 4401, p. 63-208, figs. 1-214, pl. 14. 

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southern Canadian Rocky Mountains: Jour. Paleontology, v. 34, no. 1, p. 107-126, pls. 21-25, 3 text figs. 

--- 1962, Analysis of Mississippian Syringopora from 

the southern Canadian Rocky Mountains: Jour. Paleontology, v. 36, no. 3, p. 442-460, illus. 

Orbigny, Alcide d' 1849, Note sur des polypiers fossiles: 

Paris, 12 p. 

Phillips, J., 1836, Illustrations of the geology of Yorkshire, 

pt. 2, The Mountain Limestone district: London, 253 

p., illus. 

Pickett, John, 1966, Lower Carboniferous coral faunas from 

the New England district of New South Wales : New 

South Wales Geol. Survey Mem., Paleontology, no. 15, 

38 p., ill us. [1967]. 

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a new Mississippian coral genus: Jour. Paleontology, v. 

35, no. 1, p. 65-81, pis. 17, 18, 13 text figs. 

--- 1965, Revision of some Paleozoic coral species from 

the western United States: U.S. Geol. Survey Prof. 

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--- 1967, A technique for studying early ontogeny in 

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p. 1291-1293. 

Sando, W. J., Gordon, Mackenzie, Jr., and Dutro, J. T., Jr., 

197 4, Stratigraphy and geologic history of the Amsden 

Formation (Mississippian and Pennsylvanian) of Wyoming: U.S. Geol. Survey Prof. Paper 848-A. (In press.) 

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v. 71, p. 218-272, pls. 17-21. 

Sokolov. B. S., 1939, Stratigraphical value and types of 

Chaetetidae of the Carboniferous of the USSR: Akad. 

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--- 1950, Khetetidy karbona severo-vostochnoy Ukrainy 

i sopredel'nykh oblastey [Carboniferous chaetetids of 

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Neft. Nauchno-Issled. Geol.-Razved. Inst. Trudy, nov. 

ser., vyp. 27, 143 p., illus. 

--- 19·55, Tabulyaty Paleozoya Evropeiskoy Chasti SSSR 

[Paleozoic tabulates from the European USSR]: Trudy 

VNIGRI, nov. s1er., vyp. 85, 525 p. illus. 

--- editor, 1962, Osnovy paleontologii, spravochnik dlya 

paleontologov i geologov SSSR; [t. 2], Gubki, arkheotsiaty, kishechnopolostnye, chervi [Fundamentals of paleontology, v. 2, sponges, archeocyathids, coelenterates]: 

Moscow, Akad. Nauk SSSR, 485 p., illus. 

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--- 1966, Geological Survey Research 1966: U.S. Geol. 

Survey Prof. Paper 550-A, 385 p., illus. 

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tabulate corals from Permian rocks in the Ely quadrangle, White Pine County, Nevada: Jour. Paleontology, 

v. 36, no. 3, p. 495-520, pls. 86--89, 4 text figs. 

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Geol. Soc. Ch~na Bull., v. 5, p. 291-293, pl. 1. 

INDEX 

[Italic page numbers indicate descriptions and major references] 

Page 

Actinocyathus ----------------------- C20, 21 

(Actinocyathus) Lonsdaleia ----------- 1 

stelcki, Lonsdaleia ---------- 2, 4, 21; pl. 8 

Amplexidae 7 

Amplexus ----------------------------- 7 

sp ---------------------------- 2, 7; pl. 3 

amsdenense, Barytichisma ----- 1, 2, 3, 8; pl. 3 

amsdenensis, Multithecopora ___ 2, 4, 25; pl. 10 

Zaphrentis ----------------------- 8 

Anthozoa ___ --------------------------

Anthracospirifer welleri-shawi Zone ____ .1, 1iQ 

arcuatum, Neokoninckophyllum ------- 15 

Aulophyllidae -------------------- 10 

Aulophyllinae 10 

Barytichisma ---------------------- 3, 7, 8, 9 

amsdenense --------------- 1, 2, 3, 8; pl. 3 

callosum -------------------------- 9 

crassum --------------------------

repletum -------------------------

zaphrentiforme -------------------

Big Snowy Group ---------------------

Bryozoa ------------------------------

callosum, Barytichisma 

9 

9 

9 

4 

4 

9 

Caninia ------------------------------ 18,20 

Caninia enormis --------------------- 4, 19 

excentrica ----------------------- 4, 19 

montanensis ---------------------- 4, 19 

nevadensis ----------------- 2, 4, 18; pl. 7 

Zone ----------------------------- 1, 4, 5 

Carlinia amsdeniana Subzone --------- 10 

Chaetetes -------------------------- 1. 3, 4, 5 

eximius ------------------- 2, 3, 6, 7; pl. 1 

favosus --------------------------- 6 

milleporaceus --------------------- 5, 6 

subtilis --------------------------- 6 

wyomingensis ----------- 2, 4, 5, 6, 7; pl. 1 

sp ------------------------------- 2, 7 

Chaetetida --------------------- 4 

Chaetetidae -------------------------- 4 

Chaetetiform bryozoan genus A -------- pl. 2 

Chaetetiform bryozoan genus B -------- pl. 2 

Chainman Shale ----------------------- 4 

Coelenterata 4 

Columnariina 20 

C01nposita poposiensis Subzone ------- 1, 10 

crassum, Barytichisma ----------------

crenulare, Cyathophyllum ------------- 20 

crustosa, Pleurosiphonella ------------- 23 

Cyathaxoniicae -----------------------

Cyathophyllum crenulare -------------- 20 

Cyathopsidae ------------------------- 18 

Darwin Sandstone Member -----------

Dibunophyllum ------------------------ 20 

drummondi, Pleurosiphonella ___ 2, 4, 24; pl. 9 

duncanae, Duncanopora ----- 2, 4, 25, 27; pl. 9 

Page 

Duncanopora ------------------------ 025,27 

duncanae -------------- 2, 4, 25, 27; pl. 10 

enormis, Caninia ---------------------

Erismatolithus Madreporites 

(floriformis) -------------

4, 19 

20 

Etherington Formation --------------- 4, 24 

excentrica, Caninia ------------------- 4, 19 

eximius, Chaetetes ------------ 2, 3, 6, 7; pl. 1 

Fasciculiamplexus --------------------- 7 

Fasciculophyllus ------------------- 7 

favosus, Chaetetes --------------------

( floriformis) , Erismatolithus 

Madreporites --------------

Fuchungopora --- _ --------------------

6 

20 

27 

gracile, Neokoninckophyllum ---------- 15, 18 

Great Blue Limestone --------------- 4 

Green Ravine Formation 4,27 

Halls Creek Limestone --------------- 24 

hamatilis, Neokoninckophyllum -- 2, 3, 10, 15, 

18; pls. 4, 5 

7 

27 

Hapsiphyllidae ------------------------

Hayasakaia sp ----------------------

Horseshoe Shale Member ------------- 1, 7, 10 

Hydrozoa 4 

inconstans, 

lpciphyllum 

Neokoninckophyllium 2, 3, 15; pl. 6 

20 

Koninckophyllum --------------------- 20 

Lithostrotionella ---------------------- 21 

stelcki 

Livingstone Formation ---------------

Lonsdaleia _ ---------------------------

(Actinocyathus) ------------------

21 

24 

20 

stelcki ----------------- 2, 4, 21; pl. 8 

Lonsdaleiidae 

Lonsdaleiinae 

M adreporites { ftoriformis) , 

Erismatolithus 

magnussoni, Pleurosiphonella ----------

milleporaceus, Chaetetes --------------

Moffat Trail Limestone Member ______ _ 

Monroe Canyon Limestone -----------

montanensis, Caninia -----------------

Mount Head Formation --------------

20 

20 

20 

24 

5, 6 

1, 5, 6 

4, 24 

4, 19 

24 

Multithecopora ------------------------ 25 

amsdenensis ---------------- 2, 4, 25; pl. 9 

Multithecoporidae --------------------- 25, 27 

Nemistium ---------------------------- 20 

Neokoninckophyllum ----------------- 1, 3, 10 

arcuatum ------------------------- 15 

gracile --------------------------- 15, 18 

Page 

Neokoninckophyllum-Continued 

hamatilis -------- C2, 3, 10, 15, 18; pis. 4, 5 

Zone 1 

inconstans ----------------- 2, 3, 15; pl. 6 

simplex --------------------------- 5, 18 

sp A ---------------------------- 15 

Neomultithecopora -------------------- 25 

nevadensis, Canini!L ------------ 2, 4, 18; pl. 7 

Ochre Mountain Limestone -----------

Pleurosiphonella ----------------------

crustosa --------------------------

4 

29 

23 

drummondi ----------------- 2, 4, 24; pl. 9 

magnussoni 

virginica 

Ranchester Limestone Member -------

repletum, Barytichisma ---------------

Rugosa -------------------------------

24 

24 

6, 7 

9 

4, 7 

St. Louis Limestone ----------------- 24 

simplex, Neokoninckophyllum --------- 15, 18 

stelcki, Lithostrotionella -------------- 21 

Lonsdaleia (Actinocyathus) -- 2, 4, 21; pl. 8 

Streptelasmatina ----------------------

Stylidophyllum ------------------------

7 

20 

subtilis, Chaetetes -------------------- 6 

Surrett Canyon Formation ---------- 4, 27 

Syringopora distans ------------------ 24 

drummondi --------------------- 24; pl. 9 

magnussoni -----------------------

mccutcheonae __ -------------------

pachytheca -----------------------

virginica -------------------------

Syringoporella ------------------------

Syringoporidae -----------------------

Systematic paleontology ---------------

Tabulata 

Tabulatesare 

24 

27 

24 

24 

25 

23 

4 

23 

4 

27 

27 

Tetraporinus sp ----------------------

Thecostegites -------------------------

Thecostegitidae ----------------------- 25, 27 

Turbinatocaninia. ---------------------- 19 

sp ------------------------ 2. 4,19; pl. 7 

virginica, Pleurosiphonella ------------ 24 

Wentzelophyllum ---------------------- 20 

wyomingensis, Chaetetes ---- 2, 4, 5, 6, 7: pl. 1 

Yokoyamaella ------------------------ 20 

Zaphrenticae --------------------------

zaphrentiforme, Barytichisma ---------

Zaphrentis amsdenensis --------------

Zaphrentites --------------------------

sp --------------------------------

Zoantharia ---------------------------

Zone K ------------------------------

C31 

10 

9 

8 

7 

2, 7 

7 

4 

PLATES 1-10 

Contact photographs of the plates in this report are available, at cost, from the U.S. 

Geological Survey Library, Federal Center, Denver, Colorado 80225. 
PLATE 1 

FIGURES 1-3. Chaetetes wyomingensis n. sp. (p. 5). 

Holotype, USNM 165080, collection 78. 

1. Transverse thin section (X 10); USNM 165080b. 

2. Longitudinal thin section (X 10); USNM 165080a. 

3. Transverse thin section (X 100); USNM 165080b. 

4-7. Chaetetes cf. C. eximius Moore and Jeffords (p. 6). 

USNM 165085, collection 17. 

4. Transverse thin section (X 100); USNM 165085e. 

5. Longitudinal thin section (X 10); USNM 165085f. 

6. Transverse thin section (X 10); USNM 165085e. 

7. Longitudinal thin section (X 10); USNM 165085c. 
GEOLOGICAL SURVEY 

1 

5 4 3 

6 7 

CHAETETES 
PLATE 2 

FIGURES 1-3. Chaetetiform bryozoan genus A 

USNM 165087; collection 76. 

1. Transverse thin section (X 10); USNM 165087a. 

2. Transverse thin section (X 100); USNM 165087a. 

3. Longitudinal thin section (X 10); USNM 165087b. 

4-6. Chaetetiform bryozoan genus B 

USNM 165086, collection 9·4. 

4. Transverse thin section (X 10); USNM 165086a. 

5. Transverse thin section (X 100); USNM 165086a. 

6. Longitudinal thin section (X 10); USNM 165086b. 
PROFESSIONAL PAPER 848-C PLATE 2 

GEOLOGICAL SURVEY 

1 

4 

CHAETETIFORM BRYOZOANS 
PLATE 3 

FIGURES 1-25. Barytichisma amsdenense (Branson and Greger) (p. 8). 

1-18. Lectotype, UM 2663, collection 36. 

1-5. Drawings of serial transverse peel sections (X 

10); dark stippling represents sparry calcite; 

light stippling represents stereoplasm. 

6, 7. Drawings of serial transverse peel sections (X 2) 

continuous with sections in figs. 1- 5. 

8-14. Serial transverse thin sections (X 2) continuous 

with sections in figs. 6 and 7. 

15. Calicular view (X 1). 

16. Cardinal side view (X 1). 

17. Counter side view (X 1). 

18. Alar side view (X 1) . 

19. Longitudinal thin section (X 2); paralectotype, UM 2663, 

collection 36a. 

20-25. Serial transverse thin sections (X 4); topotype, USNM 

165091a-f, respectively, collection 27. 

26-29. "Amplexus" sp. indet. (p. 7). 

Serial transverse thin sections (X 2), USNM 165088a, c-e, respectively, collection 41. 
GEOLOGICAL SURVEY 

A~A 1), K 

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c 22 I 

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26 

I 27 c 

c 

A 

PROFESSIONAL PAPER 848-C PLATE 3 

K 

A 

c 

c 

17 

6 

K 

4~~ ~WA c 7 

16 

BARYTICHISMA AMSDENENSE (BRANSON AND GREGER) AND "AMPLEXUS" SP. INDET. 
PLATE 4 

FIGURES 1-19. Neokoninckophyllum hamatilis n. sp. (p. 10). 

Holotype, USNM 165095, collection 3. 

1- 8. Drawings of s·erial transverse peel s·ections (X 10); USNM 

165095a-h, respectively. Figs. 1 and 2, brephic stage; figs. 

3-8, neanic stage. 

9-15. Serial transverse thin sections (X 2) continuous with sections shown in figs. 1-8, USNM 165095i-o, respectively. 

Figs. 9-11, neanic stage; figs. 12-15, ephebic stage. 

16. External view of counter side (X 1). 

17. External view of right alar side (X 1). 

18. External view of left alar side (X 1). 

19. External view of cardinal side (X 1). 
GEOLOGICAL SURVEY 

K 

~ 1 I 

c 

c 

c 

NEOKONINCKOPHYLLUM HAMATILIS N. SP. 

PROFESSIONAL PAPER 848-C PLATE 4 

7 l 

c 
PLATE 5 

FIGURES 1-12. Neokoninckophyllum hamatilis n. sp. (p. 10) . 

1-3. Paratype, USNM 165097; collection 3. 

4-7. 

1. Longitudinal thin section (X 2), neanic stage; 

USNM 165097a. 

2, 3. Transverse thin sections (X 2)' neanic stage; 

USNM 165097b, c, respectively. 

Paratype USNM 165102; coHection 1. 

4, 5. Transverse thin sections (X 2)' ephebic stage; 

USNM 165102a, b, respectively. 

6. Longitudinal thin section (X 2)' ephebic stage; 

USNM 165102c. 

7. External view (X 1), ephebic stage. 

8, 9. Right and left alar side (X 1) views, paratype; USNM 

165096. Collection 3. 

10. Galicular view (X 2); paratype. USNM 16M06; colle~ 

tion 2. 

11, 12. Paratype. USNM 165098. CoUection 3. 

11. Longitudinal thin section (X 2), ephebic stage; 

USNM 165098a. 

12. Transverse thin section (X 2) , ephebic stage; 

USNM 165098b. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 848-C PLATE 5 

• c 

1 

c 

c 

7 

11 c 

8 

c 

NEOKONINCKOPHYLLUM HAMATILIS N. SP. 
PLATE 6 

[All figures, collection 7 

FIGURES 1-33. Neokonincl,ophyllum inconstans n. sp. (p. 15). 

1-7. Holotype. USNM 165109. 

1-6. Serial transverse thin sections (X 2). USNM 

165109a-f, respectively. 

7. External side view (X 1). 

8-17. Paratype. USNM 165114. 

8-11. Serial transverse peel sections (X 10). USNM 

165114a-d, respectively. 

12-16. Serial transverse thin sections (X 3) continuous 

with sections shown in figs. 8-11. USNM 

165114e-i, respectively, 

17. External side view (X 1). 

18-24. Paratype USNM 165115. 

18, 19. Serial transverse thin sections (X 10) . USNM 

165115a, b, respectively. 

20-24. Serial transverse thin sections (X 3) continuous 

with sections shown in figs. 18 and 19. USNM 

165115c-g, respectively. 

25. Transverse thin section (X 2) paratype. USNM 165117a. 

26, 27. Paratype. USNM 165118, (X 2). 

26. Longitudinal thin section. USNM 165118a. 

27. Transverse thin section. USNM 165118b. 

28. External oblique side view (X 2), para type. USNM 165113. 

29. External side view (X 1), paratype. USNM 165111. 

30. External side view (X 1), para type. USNM 165112. 

31. Transverse thin section (X 2), para type. USNM 165l20a. 

32, 33. Paratype. USNM 165116 (X 2). 

32. Transverse thin section. USNM 165116a. 

33. Longitudinal thin section. USNM 165116b. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 848-C PLATE 6 

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28 26 31 

NEOKONINCKOPHYLLUM INCONSTANS N. SP. 
PLATE 7 

FIGURES 1-8. Caninia cf. C. nevadensis (Meek) (p. 18). 

1, 2. USNM 165123, collection 104. 

1. Transverse thin section (X 1); ephebic stage; USNM 

165123a. 

2. Longitudinal thin section (X 1); ephebic stage; USNM 

165,123b. 

3. Transverse thin section (X 1); ephebic stage; USNM 

165128a, collection 105. 

4, 5. Transverse thin sections (X 2) ; early ephebic and neanic 

stages, respectively; USNM 165125a, b, collection 76. 

6, 7. USNM 165126, collection 105. 

6. Transverse thin section (X 1); ephebic stage; USNM 

165126a. 

7. Longitudinal thin section (X 1); ephebic stage; USNM 

165126b. 

8. Transverse thin section (X 2); early ephebic stage; USNM 

165132a, collection 109. 

9-19. Turbinatocaninia? sp. (p. 19). 

9. Longitudinal thin section (X 2); USNM 165147a; collection 

105. 

10, 11. USNM 165148; collection 105. 

10. Longitudinal thin section (X 2); USNM 165148a. 

11. Transv·erse thin section (X 2); USNM 165148b. 

12. Transverse thin section (X 2); USNM 165145a; collection 

105. 

13. Transverse thin section (X 2) ; USNM 165151; collection 

105. 

14. Transverse thin section (X 2); USNM 165150a; collection 

105. 

15. Longitudinal thin section (X 2); USNM 165144a; collection 

105. 

16. Transverse thin section (X 2); USNM 165149a; collection 

105. 

17. Transverse thin section (X 2); USNM 165143a; collection 

105. 

18. Transverse thin section (X 2); USNM 165142a; collection 

105. 

19. Longitudinal thin section (X 2); USNM 165159a; collection 

73. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 848-C PLATE 7 

c 

0 1 14 

c 

13 

c 

c 

c 

15 

c 

CANINIA CF. C. NEVADENSIS (MEEK) AND TURBINATOCANINIA? SP. 
PLATE 8 

FIGURES 1-4. Lonsdaleia (Actinocyathus) stelcki (Nelson) (p. 21). 

USNM 165171, collection 104. 

1. Transverse thin section ( X 2) ; USNM 165171a. 

2. Longitudinal thin section ( X 2) ; USNM 165171b. 

3. Enlarged view of some of the corallites shown in fig. 2 (X 4). 

4. Enlarged view of some of the corallites shown in fig. 1 (X 4). 
GEOLOGICAL SURVEY 

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PROFESSIONAL PAPER 848-C PLATE 8 

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4 

LONSDALE/A (ACTINOCYATHUS) STELCKI (NELSON) 

2 
PLATE 9 

FIGURES 1-4. Pleurosiphonella drummondi (Nelson) (p. 24). 

USNM 165174, collection 109. 

1. Transverse thin section (X 4). USNM 165174a. 

2. Longitudinal thin section (X 1). USNM 165174b. 

3. Transverse thin section (X 1). USNM 165174a. 

4. Longitudinal thin section (X 4). USNM 165174c. 

5-9. Multithecopora? amsdenensis n. sp. (p. 25). 

5- 8. Holotype. USNM 165177, collection 105. 

5. Transverse thin section (X 4). USNM 165.!77a. 

6. Longitudinal thin section (X 4). USNM 165177b. 

7. Transverse thin section (X 1). USNM 165177a. 

8. Longitudinal thin section (X 1). USNM 165177b. 

9. Side view of silicified specimen ( X2); paratype; USNM 

165178, collection 80. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 848-C PLATE 9 

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4 

2 3 

6 

PLEUROSIPHONELLA DRUMMONDI(NELSON) AND MULTITHECOPORA? AMSDENENSIS N. SP. 
PLATE 10 

FIGURES 1- 6. Duncanopora duncanae n. sp. (p. 27). 

1-4. Holotype. USNM 165184, collection 80. 

1. Transverse thin section (X 1). USNM 165184a. 

2. Longitudinal thin section ( X 1) . USNM 165184b. 

3. Enlarged view ( X 4) of some of the corallites shown in 

fig. 1. 

4. Enlarged view ( X 4) of some of the corallites shown in 

fig. 2. 

5, 6. Paratype. USNM 165187, collection 77 (X 1). 

5. Longitudinal thin section. USNM 165187a. 

6. Transverse thin section. USNM 165187b. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 848-C PLATE 10 

1 

3 4 

DUNCANOPORA DUNCANAE N. SP. 

* u.s. GOVERNMENT PRINTING OFFICE: 1975 0-585-473/79