The Brachiopod Antiquatonia coloradoensis (Girty)

from the Upper Morrowan and Atokan

(Lower Middle Pennsylvanian) of the United States

U.S. Department of the Interior

U.S. Geological Survey

U.S. Geological Survey Professional Paper 1588
M A R C H 3, 18 4 9

U.S. DEPARTMENT OF THE IN

The Brachiopod Antiquatonia coloradoensis (Girty

from the Upper Morrowan and Atokan

(Lower Middle Pennsylvanian) of the United State

By Thomas W. Henry

U.S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 1588

Taxonomic and biostratigraphic analysis of a widespread and 

stratigraphically restricted, semireticulate productid brachiopod

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1998
For sale by U.S. Geological Survey, Information Services

Box 25286, Federal Center

Denver, CO 80225

Any use of trade, product, or firm names in this publication is for descriptive purposes only and

does not imply endorsement by the U.S. Government

Library of Congress Cataloging-in-Publication Data

U.S. DEPARTMENT OF THE INTERIOR

BRUCE BABBITT, Secretary

U.S. GEOLOGICAL SURVEY

Mark Schaefer, Interim Director

Henry, Thomas W.

The brachiopod Antiquatonia coloradoensis (Girty) from the upper Morrowan and 

Atokan (lower Middle Pennsylvanian) of the United States / by Thomas W. Henry.

p. cm.—(U.S. Geological Survey professional paper ; 1588)

Includes bibliographical references (p. – ).

1. Antiquatonia coloradoensis—United States. 2. Paleontology—Pennsylvanian. 

3. Animals, Fossil—United States. I. Title. II. Series.

QE797.S89H46 1988

564′.68—dc21 97–42005

CIP
CONTENTS

Abstract ................................................................................................................................ 1

Introduction ......................................................................................................................... 1

Repositories and Locality Register............................................................................... 1

Acknowledgments ....................................................................................................... 2

Stratigraphy and Setting of Type Locality .......................................................................... 2

Type Locality and Collections ..................................................................................... 2

Tectonic Setting............................................................................................................ 5

Stratigraphy ................................................................................................................. 5

Measured Sections in Fourmile Creek Area................................................................. 6

Macrofauna .................................................................................................................. 8

Correlation and Age ..................................................................................................... 10

Systematic Paleontology ...................................................................................................... 10

Genus Antiquatonia Miloradovich, 1945 [emend. Sarycheva, 1949, and

Sarycheva and Sokolskaya, 1952].................................................................. 10

Diagnosis .............................................................................................................. 10

Discussion ............................................................................................................ 10

Range ................................................................................................................... 12

Antiquatonia coloradoensis (Girty)...................................................................... 12

Description Based on Type Specimens ........................................................ 13

Measurements............................................................................................... 14

Cataloged Specimens.................................................................................... 14

Discussion of Type Material......................................................................... 14

Inclusion of Antiquatonia portlockiana var. quadratia in

A. coloradoensis ............................................................................... 16

Discussion of Supplemental Brachial-Valve Interiors ................................. 19

 Inferred Phylogeny and Comparison with Other Species ................................... 19

Inferred Phylogeny ....................................................................................... 19

Comparison with Antiquatonia morrowensis............................................... 20

Comparison with Antiquatonia hermosana.................................................. 21

Comparison with Antiquatonia portlockiana ............................................... 24

General Discussion............................................................................................... 24

Occurrences and Ranges....................................................................................... 26

Southern Rocky Mountains .......................................................................... 26

Eastern Great Basin ...................................................................................... 26

Midcontinent................................................................................................. 28

Appalachian Basin........................................................................................ 28

Summary of Occurrences and Ranges.......................................................... 29

References Cited .................................................................................................................. 29
IV CONTENTS

PLATES

[Plates 1–5 follow References Cited]

1. Primary type specimens of Antiquatonia coloradoensis (Girty) from lower part of Belden Shale, Park County, 

2. Primary type specimens of Antiquatonia coloradoensis (Girty) from lower part of Belden Shale, Park County, 

and hypotype of A. coloradoensis from Burgner Formation, Jasper County, Mo. 

3. Primary type specimens of Antiquatonia coloradoensis (Girty) and primary type specimens of A. portlockiana 

quadratia Sturgeon and Hoare, herein referred to A. coloradoensis.

4. Primary type specimens of Antiquatonia portlockiana var. quadratia Sturgeon and Hoare, herein referred to A. 

coloradoensis (Girty), and representative species of brachiopods from type locality of A. coloradoensis (Girty)

5. Characteristic elements of brachiopod fauna from type locality of Antiquatonia coloradoensis (Girty) from low

of Belden Shale, Park County, Colo.

FIGURES

1. Map of Colorado showing locations of Park and Lake Counties and study area ..................................................

2. Composite topographic map of parts of Mount Sherman and Fairplay West 7.5-minute quadrangles, showing 

type locality of Antiquatonia coloradoensis (Girty) and approximate locations of measured sections ................

3. Correlation of measured stratigraphic sections on both sides of Fourmile Creek in vicinity of type locality of 

Antiquatonia coloradoensis (Girty) .......................................................................................................................

4. Sketches identifying characters measured on specimens of Antiquatonia ............................................................

5. Graphs showing number of costellae and costae in 10 mm at 20 mm surface length for Antiquatonia 

morrowensis (Mather) and its probable descendants, A. coloradoensis (Girty), A. hermosana (Girty), and 

A. portlockiana (Norwood and Pratten).................................................................................................................

6. Chart showing lineage of Antiquatonia in Lower and Middle Pennsylvanian strata in the United States ...........

7. Map of the United States showing locations of verified or probable occurrences of Antiquatonia coloradoensis

(Girty) ....................................................................................................................................................................

TABLES

1. Invertebrate macrofauna from collections containing primary and secondary type specimens of Antiquatonia 

coloradoensis (Girty) from lower part of Belden Shale on ridge south of Fourmile Creek, near Fairplay, Colo. 

2. Measurements of figured and (or) measured primary type specimens and topotypes of Antiquatonia 

coloradoensis (Girty) from the upper Morrowan part of the Belden Shale, Park County, Colo., and of a 

hypotype from the upper Atokan Burgner Formation, Jasper County, Mo. ..........................................................

3. Measurements of primary type specimens of Antiquatonia portlockiana var. quadratia Sturgeon and Hoare 

(referred herein to A. coloradoensis (Girty)) from upper Atokan strata of the “lower Mercer limestone and shal

of the Pottsville Formation, southeastern Ohio. ....................................................................................................

4. Measurements of specimens identified as Antiquatonia coloradoensis (Girty) and A. hermosana (Girty) by Rag

(1959) from the Burgner Formation (upper Atokan), Jasper County, Mo., and referred herein to A. coloradoens

(Girty) ....................................................................................................................................................................

5. Index to original figures and replication of same in publications for lectotype and two paralectotypes of 

Antiquatonia coloradoensis (Girty) from the lower part of the Belden Shale (upper Morrowan equivalents) 

from U.S. Geological Survey colln. GR2281–PC, from near Fairplay, Park County, Colo. .................................
CONTENTS 

CONVERSION FACTORS

[Specimen measurements are in millimeters and centimeters, whereas other measurements are in feet and miles]

Multiply By To obtain 

millimeter (mm) 0.0394 inch

 centimeter (cm) 0.3937 inch

foot (ft) 0.3048 meter

 mile (mi) 1.609 kilometer
The Brachiopod Antiquatonia coloradoensis (Girty) 

from the Upper Morrowan and Atokan

(Lower Middle Pennsylvanian) of the United States

By Thomas W. Henry

ABSTRACT

Primary type specimens and topotypes of the large,

semireticulate productid, Productus inflatus var. coloradoensis (Girty), collected near Leadville, Colo., form the

basis for the redescription of this species. The type locality,

for the first time, is firmly established stratigraphically in

upper Morrowan (lower Middle Pennsylvanian) rocks of the

lower part of the Belden Shale. 

The identification and clarification of several previously poorly known physical features affirm assignment of

Girty’s species to the genus Antiquatonia Miloradovich, as

currently restricted and understood. Detailed analysis of

other collections of brachiopods assigned to A. coloradoensis and to related taxa confirms that the probable ancestor of

A. coloradoensis is the lower and middle Morrowan A. morrowensis (Mather), which was described from the Ozarks

region of northwestern Arkansas. One descendant of A. coloradoensis is A. hermosana (Girty), which is common in

lower Desmoinesian strata of the southern Rocky Mountains and Paradox Basin. A descendant of A. hermosana

may be A. portlockiana (Norwood and Pratten), which is

common in uppermost middle and upper Desmoinesian

strata in the Rocky Mountains and which is reported from

Desmoinesian strata in the Midcontinent, Illinois Basin, and

Appalachian Basin. However, the variability, taxonomy, and

stratigraphic ranges of the species of Antiquatonia in the

Desmoinesian of the eastern United States have not been

resolved satisfactorily. 

The stratigraphic range of Antiquatonia coloradoensis

extends from the upper part of the Morrowan Series through

the Atokan Series. The known geographic range of this species extends from the Eastern Great Basin, through the

southern Rocky Mountains, the southern and central Midcontinent area, to the eastern and southern Appalachian

Basin. It is a moderately common to rare component of

open-bay, shelf-lagoon, and shelf-margin marine facies.

INTRODUCTION

Studies of the Mississippian and Pennsylvanian b

opod faunas in the United States routinely deal with 

taxonomic and stratigraphic problems. However, rare

difficulties encountered that are as intricate as those 

fronted in the investigations of the large, semireticulat

ductids of the Lower to Middle Pennsylvanian Morr

Atokan, and Desmoinesian Series. Specifically, I am 

ring to the confusion in the literature about three th

(1) the exact geologic age and geographic site of the 

locality of Productus inflatus var. coloradoensis 

(1910), now referred to the genus Antiquatonia Milo

vich (1945) [emend. Sarycheva, 1949, and Sarychev

Sokolskaya, 1952]; (2) the stratigraphic range and 

graphic distribution of this distinctive, biostratigraph

diagnostic productid; and (3) the proper taxon

disposition and assignment of numerous collections 

the United States of Morrowan, Atokan, and lower 

moinesian specimens assigned to Antiquatonia. Like 

productid brachiopods, A. coloradoensis is a highly va

taxon. A lack of understanding of its breadth of varia

and lingering tendencies toward typologic application 

contributed significantly to widespread misinterpretat

this species. This paper clarifies the situation. 

REPOSITORIES AND LOCALITY REGIST

All primary and secondary type specimens of An

tonia coloradoensis illustrated or measured in this pap

reposited at the U.S. National Museum of Natural H

(Smithsonian) in Washington, D.C., and are identified 

the prefix “USNM.” All of the type specimens describ

Antiquatonia portlockiana var. quadratia by Sturgeo

Hoare (1968) in their paper on the Pennsylvanian bra

pods of southeastern Ohio are included in the synony

A. coloradoensis, are reposited in the paleonto
2 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

collections of the Orton Geological Museum at Ohio State

University, and are identified by the prefix “OSU.” Additional paratypes of the variety quadratia formerly reposited

at Bowling Green State University now have been reposited

at Ohio State University and given new specimen numbers

(R.D. Hoare, oral commun., July 21, 1993) used in the current paper. Specimens of Antiquatonia from southwestern

Missouri studied by me from the University of Missouri at

Columbia are prefixed “UM.” 

The U.S. Geological Survey (USGS) collections

studied for this report are cataloged in the so-called “PermoCarboniferous” Fossil Locality Register. Collection numbers

are suffixed by “–PC” and are prefixed by either “GR” (designating the green subregister) or “BL” (blue subregister). 

ACKNOWLEDGMENTS

I am indebted to my colleague, Patrick K. Sutherland,

University of Oklahoma, for providing additional data on

the range of Antiquatonia coloradoensis in southern Oklahoma, Texas, and New Mexico and for enabling me to

examine the collections of this and related species at the

Oklahoma Museum of Natural History, University of Oklahoma, Norman. I also appreciate his thorough and constructive review of the manuscript. Robert B. Blodgett, USGS,

also reviewed this manuscript. 

Primary type specimens of Productus inflatus var. coloradoensis (Girty) were made available by Jann W.M.

Thompson, Division of Paleobiology, Smithsonian Institution, for additional preparation and photography. Stig M.

Bergstrom and Gale Gnidovec, Ohio State University,

loaned me one set of Sturgeon and Hoare’s (1968) type

specimens of Antiquatonia portlockiana var. quadratia from

southeastern Ohio. Richard D. Hoare, Bowling Green State

University (BGSU), provided the additional paratypes of

this named variety that were reposited previously at BGSU.

Raymond L. Ethington, University of Missouri at Columbia, loaned additional specimens of Antiquatonia cataloged

in the University of Missouri collections. 

My appreciation is extended to Richard De Voto, formerly of the Colorado School of Mines, with whom I had

several conversations about the stratigraphy and tectonic

setting of the type area for Girty’s species. Charles L. Rice,

USGS, and Donald R. Chesnut, Jr., Kentucky Geological

Survey, provided useful information on Appalachian Basin

stratigraphy. T.L. Thompson, Missouri Geological Survey,

discussed current information on the age of the Burgner

Formation. C. Howard Brunton, British Museum (Natural

History), provided an extensive written discussion of the

genus Antiquatonia and its type species Productus antiquatus J. Sowerby. The late Mackenzie Gordon, Jr., USGS,

encouraged my interest in pursuing this study and guided

me through some of the intricacies of the taxonomic problems encountered. I also appreciate constructive comments

by J. Thomas Dutro, Jr., USGS, Washington, D.C. Many of

the photographs were made by Eric M. Fisher, USGS, and

computer-graphic drawings were produced by Patricia

Holroyd, USGS. 

STRATIGRAPHY AND SETTI

OF TYPE LOCALITY

TYPE LOCALITY AND COLLECTI

The type locality of Antiquatonia colorado

the high country of the southern Rocky Mountai

Leadville and Fairplay, Colo. (figs. 1 and 2). 

above timberline in the Mosquito Range in the Al

district, a historic gold-producing area.

The primary type specimens of Productus i

coloradoensis Girty were collected in 1873 by A

who was the geologist in charge of the South Par

of the Hayden Survey of parts of the Colorado 

When he collected this material, Peale was co

geologic reconnaissance of the mining district

accompanying the topographers who surveyed 

piled the first topographic maps of the region. H

section 18 (Peale, 1874, p. 229–233) was made o

crest on the south side of the glaciated valley o

Fourmile Creek, extending on the east from just 

of Sheep and Lamb Mountains to “the Horse Sho

ques east of Horseshoe Mountain) near the apex o

quito Range on the west. Peale’s panoramic ske

pl. IX) of this ridge was made looking southwar

north side of the broad valley (see fig. 2 of curr

This sketch identifies several features keyed to t

section, and the approximate geographic position 

lection can be plotted from these. The pertine

Peale’s traverse section (in descending stratigrap

is quoted below: 

According to this traverse section (Pe

p. 231–232), the top of unit 38 lies stratigraphica

imately 170 ft above a thick, light-colored “

volcanic rock.” 

S.F. Emmons conducted a detailed stu

Leadville-Fairplay area 9 years later in the summ

and produced the first geologic map of the area (

following observations were made by Emmo

p. 162) about the stratigraphic section on the ri

south side of Horseshoe Gulch:

Directly above the White Porphyry is a bed of black carbona

from one hundred to one hundred and fifty feet above it is 

dark, impure limestone, from which were obtained a large n

sils, among which the following were recognized: Chonete

Productus cora, Productus nodosus (variety of Productus cor

semireticulatus, Myalina perattanuata, fragment of Pinna sp

Aviculopecten, Phillipsia sp., Phillipsia major, fragment of L

37. Greenish-gray micaceous sandstones. Toward the top the 

very coarse, 15 feet.

38. Limestones and bluish argillaceous shales with sandstone

portion of the bed contains in blue shaly limestones fine 

Productus semireticularis, P. nebrascensis, Spirifer opimu

prattenanus, and a Pleurotomaria, 57 feet. 

39. Black shaly limestones, in the lower part of which we find 

fossils, Productus, Spirifer, and fragments of Trilobites, 3

40. Quartzitic sandstone laminated and micaceous, 15 feet.
STRATIGRAPHY AND SETTING OF TYPE LOCALITY 

Denver 40°

38°

108° 106° 104° 102°

LAKE

PARK

Leadville Fairplay

Study area

of figure 2

0 10 20 30 40 50 MILES

0 10 20 30 40 50 KILOMETERS

0 100 MILES

0 100 KILOMETERS

COLORADO

The “White Porphyry” to which Emmons referred is

the commonly applied, local name for the intrusive porphyry (mainly granodiorite and quartz monzonite) of Late

Cretaceous-early Tertiary age that Chronic (1964) and

Tweto (1974) mapped as a thick sill cropping out on the

south side of the Horseshoe amphitheater (cirque) and

extending to the ridge top west of the type locality of

P. inflatus var. coloradoensis. 

Girty’s figured specimens (1903, pl. 3, figs. 1–3) of

Productus inflatus McChesney (later designated the types of

P. inflatus var. coloradoensis) were cited as from the “Weber

formation (?): Leadville district (sta. 2281)” (Girty, 1903, p.

490). In his description of this station, Girty (1903, p. 525)

conveyed the following information: “Leadville district;

Weber shales, base of Weber formation (?). Near mouth of

canyon of Fourmile Creek. The locality is bed 38 (and

possibly 39) of section 18” of Peale (1874, p. 231). E

tially the same information was given by Girty (190

245) in his discussion of the faunas from the South 

area. However, the handwritten label accompanyin

USNM 8258 (from which Girty’s primary types, U

colln. GR2281–PC, were drawn) reads: “Carbonife

Ridge E. of Horseshoe Mt., S. Park, Colo.” Thus, G

original descriptions of this locality are misleading i

the collection was not taken from near the mouth of the 

yon of Fourmile Creek but rather on the ridge crest so

the glacial valley occupied by Fourmile Creek (ac

south of Horseshoe Gulch, a tributary of Fourmile C

and southeast of Horseshoe Mountain (see fig. 2). 

Gordon (1975, p. D48) concluded that the type lo

of Antiquatonia coloradoensis (USGS colln. GR2281

probably was on “the crest of the ridge south of Four 

(Horseshoe) Creek, about 2 miles east of Horseshoe M

tain and 8 miles west-southwest of Fairplay, Colo.” H

mised that this locality would lie approximately in the 

sec. 18 (unsurveyed), T. 10 S., R. 78 W., at an al

slightly greater than 12,400 ft and cautioned that this n

field verification. 

Four collections of fossils from the ridge on the 

side of Fourmile Creek were examined in addition to th

set for this study. I consider that all four of these (USG

lns. BL6852–PC, BL6852A–PC, BL12096–PC, 

BL31394–PC) contain topotypes of Antiquatonia col

oensis. The first two (BL6852–PC and BL6852A–PC) 

made by J.H. Johnson in July 1930 from “near top of 

south of Horseshoe cirques” (see fig. 2). Colle

BL6852–PC was reported to occur “about 150 ft abov

phyry,” and BL6852A–PC was made about “50 ft 

graphically higher” and slightly to the east of his 

collection (Johnson, 1934, p. 33). Despite Johnson’s U

collns. BL6852–PC and BL6852A–PC being separat

about 50 ft of section in this immediate area, both are co

ered herein to contain topotypic material of Antiqua

coloradoensis. This is justified because Girty’s ma

(1903, p. 525) may have come from both beds 38 and 

Peale’s traverse section 18 (1874, p. 231), represent

composite thickness of 91 ft. Both collections were 

lated by Johnson (1934) with a section that he measure

north of the old townsite of Leavick (see B in fig. 2 an

tion below on “Measured Sections in Fourmile Creek A

The third collection that I examined (BL12096–PC

made in 1953 by Ogden Tweto from the “Alma dist.

Co., Colo.; ridge on south side Horseshoe Gulch.” The f

(BL31394–PC) was collected in 1991 by E.M. Fisher an

in an effort to locate the original site and obtain addi

fossils from it. The locale of USGS collection BL3139

is plotted in figure 2, and its description follows: 

Quadrangle.—Mount Sherman 7.5 minute. 

Description.—Site on open swale on rounded, 

alpine ridge south of Horseshoe Gulch, between Hors

Mountain and Lamb Mountain, at point 1.65 mi 

southeast of vertical angle elevation bench mark V

Figure 1. Location of Park and Lake Counties, Colo., communities of Fairplay and Leadville, and study area. Mosquito Range

separates Park and Lake Counties. West and east parts of study

area are Mount Sherman and Fairplay West 7.5-minute quadrangles, respectively. 
4 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

Figure 2. Composite map (scale 1:48,000) taken from adjacent parts of Mount Sherman and Fairplay West 7.5-minute-quadrangle 

maps (U.S. Geological Survey, scale 1:24,000, 1961 and 1960, respectively), showing geographic features mentioned in text. Type lo

tiquatonia coloradoensis (Girty) is depicted by filled circle, and approximate estimate of type locality by Gordon (1975, p. D48) is 

open circle. Zigzag lines indicate approximate traverses for measured sections: “A” indicates Kelly’s (1983) Lamb Mountain section

dicates Johnson’s (1934) Horseshoe Mountain section, which equals Kelly’s (1983) White Ridge section. Approximate positions of 

section lines were added to pertinent parts of Mount Sherman sheet. Contour interval equals 20 ft on Fairplay West quadrangle and 40 

Sherman quadrangle.
STRATIGRAPHY AND SETTING OF TYPE LOCALITY 

13,898 ft (on Horseshoe Mountain) and 1.5 mi southwest of

abandoned ore mill at old Leavick townsite; site is 0.35 mi

north-northeast of temporary bench mark TBM 13,128 ft

(marked by three large cairns on small knoll held up by

platy-weathering, coarse-grained quartzarenite). 

Geographic coordinates.—Approx. C., SW1/4 sec. 18

(unsurveyed), T. 10 S., R. 78 W., Park County, Colo.;

lat 39°10'35'' N.; long 106°09'23" W. 

Altitude.—12,805 ft, measured barometrically on highest fossiliferous bed on ridge crest. 

Attitude.—Strike N. 40° W., dip 16° NE. 

Stratigraphy.—Belden Shale (lower part); collected

free-weathering and “crack-out” macrofossils from approximately 8 to 10 ft of alternating, nodular, micritic limestone and shale. Lower limestone: skeletal carbonate

mudstone, medium-dark-gray to dark-gray, weathering

medium light gray to yellowish gray. Shale: medium-darkto dark-gray, slightly calcareous. Upper limestone (skeletal, brachiopodal, bryozoan wackestone to packstone):

becoming more evenly bedded, slightly more skeletal,

coarser grained, more sandy (containing quartz and feldspar

grains), and thicker bedded upward, grading at top into

fine-grained, pale-yellowish-brown-weathering, calcareous, skeletal, subarkosic sandstone; top beds make small

ledges at ridge crest and on hillside. 

Date collected.—Sept. 4, 1991.

The top of the beds from which USGS colln.

BL31394–PC was obtained is approximately 160 ft stratigraphically (determined trigonometrically) above the top of

the “White Porphyry” sill and probably is exactly equivalent

to Johnson’s (1934) colln. BL6852–PC. The site is about

400 ft higher on the ridge and about 0.75 mi southwest from

where Gordon (1975, p. D48) inferred that Peale’s original

collection might have been made. 

TECTONIC SETTING

Most of central and western Colorado, including Park

and Lake Counties, was broken into a mosaic of fault

blocks during the Pennsylvanian and Permian, when the

ancestral Front Range and Sawatch and Uncompahgre

Uplifts and the Central Colorado Trough were formed (De

Voto and others, 1986). Syndepositional tectonism resulted

in abrupt thickness and facies changes of alternating marine

and nonmarine upper Paleozoic strata in the fault-bounded

basins in many areas, including the Leadville-Fairplay area.

There, the strata that are now referred to the Belden Shale

and that contain the types of Antiquatonia coloradoensis

were deposited in the South Park Subbasin of the Central

Colorado Trough.

STRATIGRAPHY

The stratigraphic name Weber was applied initia

the Leadville-Fairplay area by Emmons (1883, 

p. 219) for the sequence of generally interbedded 

shale, limestone, sandstone, and conglomeratic sand

that now compose the Belden Shale, the Molas Form

and tongues of the Minturn Formation (Middle Penn

nian). Emmons’ application of the name Weber form

was based on the assumption that these beds correlated 

the Weber quartzite of the Uinta Mountains of northe

Utah. Girty (1903, p. 171) questioned this early corre

in his discussion of what became the type locality for 

quatonia coloradoensis, as noted above, and utilize

name “Weber formation” with a query for these str

practice followed by many subsequent workers, incl

Johnson (1934) and Read (1934). 

Brill (1942, p. 1385) proposed the name “Belden 

member of the Battle Mountain formation” for 

sequence in central Colorado, later raised the Belden t

mation rank (Brill, 1944, p. 624), and subsequ

extended its use throughout the Central Colorado T

(Brill, 1952, 1958). For a detailed discussion of the 

graphic nomenclature in the Mosquito Range, see De 

(1972, p. 142–145). 

The Belden Shale (Morrowan and lower Atokan) 

Mosquito Range is normally underlain by the Molas F

tion (Upper Mississippian? and Lower Pennsylvania

clastic red-bed sequence of silty, variegated shale and 

stone and other lithic types, interpreted in part as a re

soil blanketing the Leadville Dolostone (Lower Miss

pian, Osagean), which was subaerially exposed and e

sively karsted during the Late Mississippian and ea

Pennsylvanian (De Voto, 1972). In the Colorado M

Belt, of which the Mosquito Range is a part, the Lea

Limestone was originally deposited in intertid

supratidal environments, extensively dolomitized durin

Mississippian and (or) Pennsylvanian, and then recr

lized by mid-Cenozoic hydrothermal brines derived fro

igneous source. Armstrong, Mamet, and Repetski (

proposed that the name Leadville Dolostone be appl

this formation in the current study area, and I follow 

practice in this paper.

Tweto and Lovering (1977) described the Molas 

Mosquito Range as filling in caves and channelways 

Leadville and ranging in thickness there from a few i

to as much as about 40 ft. In the Fourmile Creek area

thin, extensively bleached by hydrothermal action (T

and Lovering, 1977; Armstrong, Mamet, and Rep

1992), and generally covered. 
6 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

The Belden Shale on the south side of Fourmile Creek

is about 670 ft thick (Kelly, 1983) and contains several

wedges of coarse-grained, conglomeratic, terrigenous clastic deposits, interpreted as tongues of the overlying Minturn

Formation (Chronic, 1964, p. 109; De Voto, 1972). These

wedges pinch out to the south, and the Belden becomes

thicker and more shaly. Farther northward in the Mosquito

Range, many additional coarse terrigenous clastic deposits

are present in the same correlative interval, and it is not feasible to differentiate the two formations. 

Tweto (1974) mapped the ridge top south of Horseshoe

Gulch as Minturn and Belden Formations undifferentiated

for his geologic map of the area west of Fairplay. However,

on notes with his field labels accompanying his USGS colln.

BL12096–PC, Tweto indicated that it was from the Belden

Shale and probably from its lower part. Unfortunately, no

more light is shed by Tweto’s notes regarding a more precise geographic locality for his collection other than that

given in the preceding sentence.

MEASURED SECTIONS 

IN FOURMILE CREEK AREA

The invertebrate fossil collections studied came from

the lower part of the Belden Shale slightly above the horizon

where Read (1934) reported the “middle Pottsville” compression and impression flora from sections a short distance

to the west of the current study area. It is possible to more

precisely document the stratigraphic position of the invertebrate collections from two measured sections in the vicinity

by Kelly (1983) and from one by Johnson (1934) on the

north side of Fourmile Creek. Johnson’s section on the north

side of the glacial valley—his Horseshoe Mountain section—was remeasured by Kelly as his White Ridge section

(plotted as sec. “B” in fig. 2). Kelly’s other section, the Lamb

Mountain section (sec. “A” in fig. 2), apparently follows the

ridge crest on the south side of the glacial valley where the

type locality of Antiquatonia coloradoensis is located. 

Kelly’s (1983) Lamb Mountain section was measured

above timberline mainly in “secs. 17 and 18, T. 10 S., R. 78

W.” The lower part of the Lamb Mountain section and his

correlation of it with his White Ridge section based on

detailed petrographic and stratigraphic analysis and environmental interpretations are depicted in figure 3. The

White Ridge and Lamb Mountain sections are separated by

a major fault (south side upthrown) that Kelly (1983) interpreted as having been active during deposition of the Belden

and Minturn sediments, thus partly explaining many of the

major changes in interval thicknesses and lithofacies that

occur in the short distance across the glacial valley occupied

by Fourmile Creek. 

Exactly where was the Lamb Mountain section measured? The ridge crest on the south side of Fourmile Creek

(Horseshoe Gulch) affords the most accessible traverse and

the best exposures of the Belden Shale, and the more resistant strata of the Belden, dipping gently eastward, also crop

out on the north side of the ridge and extend eastward down

the side of the glacial valley. The ridge top was also certainly the line of traverse made by Peale (1874). In this section, the Molas Formation is absent, and the Belden is

underlain by a thick sill (the “White Porphyry” of local

usage), intruded approximately along the contact between

the Leadville and the Belden. TBM 13,474 ft in figure 2 is

on the promontory formed by about the middle part of this

sill. The Lamb Mountain section probably extends from just

east of that point along the ridge crest east-northeastward to

some point northeast of TBM 12,478 ft. The lower tongue

of the Minturn Formation (fig. 3) forms the knob at TBM

13,128 ft in the line of the measured section, and the type

locality of Antiquatonia coloradoensis occurs at the filled

circle marking USGS colln. BL31394–PC. 

Johnson (1934) collected and identified invertebrate fossils from several stratigraphic sections and localities throughout the Mosquito Range. Although J.H. Johnson identified

most of the fossils himself, G.H. Girty corroborated and

assisted in the identifications (Johnson, 1934, p. 16). Girty’s

distinctive handwriting appears on many of the collection

labels, including those identifying specimens of Productus

coloradoensis from collns. BL6852–PC and BL6852A–PC.

Johnson’s (1934, p. 37–39) Horseshoe Mountain section was measured “uphill northward from old town of

Leavick, following around hill and through first cirque.” The

approximate position of Johnson’s Horseshoe Mountain

section, plotted as “B” in figure 2, is compatible with

Tweto’s (1974) geologic map and Kelly’s (1983) description. Johnson’s (1934) designation of this measured section

as the Horseshoe Mountain section is confusing. Horseshoe

Mountain is on the crest of the Mosquito Range approximately 3 mi southeast of White Ridge (fig. 2).

Figure 3 (following page). Measured stratigraphic sections on

both sides of Fourmile Creek in vicinity of type locality of Antiquatonia coloradoensis (Girty). Kelly’s (1983) White Ridge section is

same as Johnson’s (1934, p. 37–39) Horseshoe Mountain section.

Approximate locations of measured sections are shown by zigzag

lines in figure 2. Data for Lamb Mountain and White Ridge sections, including correlation lines, are from Kelly (1983, pl. 1, meas.

secs. 6 and 7); however, a different horizontal datum was used

(Kelly’s T–2 datum). Kelly’s unit numbers are shown on sides of

measured sections, and upper parts of both measured sections are

not shown. Coarse-grained conglomeratic units (shown in gray on

cross section) are interpreted as tongues of Minturn Formation that

pinch out southward. See text for explanation of arrow on left of

figure.
STRATIGRAPHY AND SETTING OF TYPE LOCALITY 

       





































































 























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2A

1

0

2B

3A

3B

USGS 6852A-PC

USGS 6852 and

31394-PC

"White porphyry"

USGS 6851A-PC

DATUM

USG 6851-PC

4A

5A

5B

5C

5D

6A

6B

7B

4B

7A

8A

1

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2A

2B

3A

3B

4A

4B

5A

5B

6A

6B

7A

7B

7C

8A

8B

8C

8D

9A

9B

9C

10A

LAMB

MOUNTAIN

WHITE

RIDGE

BELDEN SHALE BELDEN SHALE

BELDEN SHALE

TONGUE OF MINTURN FORMATION

LOWER

TONGUE

OF

MINTURN

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Limestone

Dolomitic limestone

Fine-grained sandstone

Coarse-grained sandstone

Shale

Covered

Conglomerate and

conglomeratic sandstone

Porphyritic igneous rock

EXPLANATION

METERS FEET

0

50

100

0

10

20

30
8 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

This section includes several fossiliferous limestone

and shale beds, the lower two of which are especially significant. Johnson’s (1934, p. 39) lowest fossiliferous bed in the

Horseshoe Mountain section, his unit 8, is described as an

11-ft-thick, dark-gray, sparsely fossiliferous, thin-bedded

limestone from which he made colln. BL6851–PC. According to his measurements, the base of this unit is approximately 200 ft above the top of the Leadville Dolostone. It is

this unit with which Johnson (1934, p. 33) correlated colln.

BL6852–PC, which contains topotypes of Productus inflatus var. coloradoensis Girty. The next higher fossiliferous

unit described in Johnson’s (1934, p. 38) Horseshoe Mountain section, his unit 13, is a 13-ft-thick, highly fossiliferous,

slightly calcareous, black shale, from which colln.

BL6851A–PC was derived. The base of Johnson’s unit 13 is

roughly 440 ft above the top of the Leadville Dolostone.

Johnson (1934, p. 33) correlated the strata from which he

made colln. BL6852A–PC (the second topotypic collection

of coloradoensis) with his unit 13. Thus, according to his

measured section and correlation of collns. BL6852–PC and

BL6852A–PC with it, these marine beds are separated by

about 229 ft of unfossiliferous sandstone and cover on the

north side of the broad glacial valley and by only about 50 ft

of strata on the south side of the valley, where the types of

Antiquatonia coloradoensis originated. A problem exists,

however, with the upper of Johnson’s two correlations. 

As stated above, Kelly’s (1983, pl. 1, meas. sec. 7)

White Ridge section (fig. 3) is essentially the same as

Johnson’s Horseshoe Mountain section, just discussed.

Kelly’s section was measured in “secs. 4 and 5 (unsurv.), T.

10 S., R. 78 W.,” just north and northeast of Leavick (fig. 2).

By comparing interval (rock-unit) thicknesses and similarities of the two sets of lithic descriptions, Johnson’s data can

be interpreted quite reasonably in terms of Kelly’s White

Ridge section. Johnson’s USGS collns. BL6851–PC and

BL6851A–PC are plotted against Kelly’s White Ridge section in figure 3. 

From analysis of these sections and my personal observations in the field, I have made the following two conclusions. (1) Johnson’s colln. BL6851–PC from the Horseshoe

Mountain-White Ridge section correlates with the relatively

thin, fossiliferous, alternating limestone and shale sequence

in the top of Kelly’s (1983) unit 2A of the Lamb Mountain

section (fig. 3), which yielded Johnson’s colln. BL6852–PC

and my colln. BL31394–PC. Therefore, Johnson was correct in his correlation of the lower fossil-bearing unit in his

Horseshoe Mountain section. (2) Johnson’s BL6852A–PC

probably came from Kelly’s unit 3A, which he noted contains brachiopods but which I was unfortunately not able to

recollect due to a sudden afternoon thunderstorm. Therefore, Johnson’s colln. BL6851A–PC does not correlate with

BL6852A–PC as he contended, but rather with the lower

part of the limestone bed in unit 4A in the same section (see

arrow in fig. 3). 

MACROFAUNA

Table 1 lists the total macrofauna from US

GR2281–PC, BL6852–PC, BL6852A–PC, BL

and BL31394–PC. Plates 1–3 show Antiquaton

doensis from these and other collections. The oth

pod taxa from this fauna are illustrated in plates 4 

Girty (1903, p. 243, table 12) identified the 

brachiopods from USGS colln. GR2281–PC: 

inflatus McChesney, Productus cora d’Orbigny, 

nebrascensis Owen, and Spirifer rockymontanu

The eight specimens from this collection that he 

as P. cora (Girty, 1903, p. 364–367) are in the pa

collections of the U.S. National Museum of Natu

(lot USNM 8259). His P. cora is the highly distin

productus nodosus (Newberry). A slab contain

specimens of this species and two free-weathe

mens (one from GR2281–PC) are illustrated he

figs. 13-17). The specimens from GR2281–PC 

referred to Productus nebrascensis and Spirifer ro

nus could not be located in the USNM collect

examination of the other related collections, I co

the small Anthracospirifer that Girty (1903, p

identified as Spirifer rockymontanus is A. tanoen

land and Harlow (see pl. 4, figs. 18–20). Gir

p. 364–376) Productus nebrascensis is probably 

with the specimens that I tentatively identify as P

pustulosa Sutherland and Harlow (see pl. 5, fig

My additional identifications from four other 

from the type locality are shown in table 1, and 

these specimens are figured in plate 5.

Linoproductus nodosus is the most commo

pod in the collections from the type locality of An

coloradoensis, as can be seen in table 1 (130 spec

fragments). Although the shape is highly 

L. nodosus is easily identified by a single row 

almost recumbent body spines emerging from 

valve exterior at or near the plane of symmetry o

This is a common, geographically widespread lin

that has been identified from rocks of late Mor

early Atokan age from the Eastern Great Basin, G

yon area of Arizona, northern Sonora (Mexico

Rocky Mountains, Ozarks, frontal Ouachitas

Basin, Llano area of Texas, central and southe

chian Basin, and northern South America. Like 

productids, L. nodosus was gregarious, most 

growing in closely packed clusters on and partial

in argillaceous- and carbonate-mud bottoms. The 

illustrated in plate 4, figure 15, contains a cluster 

imens from USGS colln. BL31394–PC. Note t

one of the specimens is oriented with brachial v

growth position). The specimen identified by the 

one of the body spines. The large specimen above 
STRATIGRAPHY AND SETTING OF TYPE LOCALITY 

the same figure is an exfoliated interior of a brachial valve;

the entire pedicle valve, the muscle fields and other interior

features of the brachial-valve interior, and much of the cardinal area have been worn away. Note that the bumps forming a row along the midline of the brachial-valve interior

correspond to the body spines on the pedicle-valve exterior. 

I strongly suspect that Emmons’ collection (1886,

p. 162), cited above in the section “Type Locality and Collections,” was from the same beds and locality as the type

specimens of Antiquatonia coloradoensis, and I con

that his Chonetes granulifera is Neochonetes cf. N. w

his Productus semireticulatus is A. coloradoensis, an

P. cora (together with his P. nodosus) is Linoprod

nodosus. However, the disposition of his collectio

unknown, and, because of apparent differences in the f

lists, it is not likely that specimens in Emmons’ list 

from Peale’s 1873 collection (USGS colln. GR2281

described in 1874. 

Table 1. Invertebrate macrofauna from collections containing primary and secondary type specimens of Antiquatonia

coloradoensis (Girty) from lower part of Belden Shale on ridge south of Fourmile Creek, near Fairplay, Colo.

[Presence denoted by “+” or by a number; specimens identified by Girty (1903, p. 243, table XII), interpreted in terms of modern taxonomy,

that were not examined (or found) are identified by asterisk; absence denoted by dash]

Taxon U.S. Geological Survey colln. no.

GR2281–PC BL6852–PC BL6852A–PC BL12096–PC BL31394–PC

ECHINODERMATA:

Pelmatozoan columnals......................... * + + + +

Crinoid plates ........................................ – + – + +

Archaeocidaris...................................... – – + – +

BRYOZOA:

Rhombopora spp. .................................. * – + + +

Fenestellid ............................................. – – + + +

Other bryozoans .................................... – – + + +

COELENTERATA:

Lophophyllid coral................................ – 4 – – 1

Conularid............................................... – 1 – – –

BRACHIOPODA:

Neochonetes cf. N. whitei...................... – 3 18 1 6

Linoproductus nodosus 8 14 33 29 46

Antiquatonia coloradoensis .................. 19 8 4 18 11

Sandia welleri ....................................... – – – 2 1?

Desmoinesia nambeensis...................... – 43––3

Pulchratia? pustulosa(?) ....................... * – 2 1 –

Productid, gen. & sp. indet.................... – – – – 1 

Spirifer goreii........................................ – – – 1 2

Anthracospirifer tanoensis.................... * – 4 4 2

Composita ovata ................................... – 1 3? 1 2

Composita gibbosa................................ – 1 1? 1 1?

PELECYPODA:

Lima? sp. indet. ..................................... – 1–––

Myalinid ................................................ – – 1 – 1

Pectenid (Deltopecten?) ........................ – – 1 – –

GASTROPODA:

Meekospira sp. ...................................... – 1 – – 1

Sphaerodoma sp. ................................... – 1–––

Euconospira taggarti ............................ *––––

CEPHALOPODA:

Orthoceratid sp. indet............................ – 3 – – 1

ARTHROPODA:

Paladin? sp. (trilobite)........................... – 1 1 – 1
10 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

CORRELATION AND AGE

The fauna documented from the lower part of the

Belden Shale south of Fourmile Creek (table 1) is closely

similar to faunas of the following four units: (1) the lower

part of the La Pasada Formation (Lower and Middle Pennsylvanian) in the southern Sangre de Cristo Mountains of

northern New Mexico (Sutherland and Harlow, 1973),

(2) the basal 2.5 ft (upper Morrowan) of the type “Derryan”

Series in southern New Mexico (Sutherland and Manger,

1984; Sutherland, 1991), (3) the middle slope unit of the

Watahomigi Formation (Lower and Middle Pennsylvanian)

of northern Arizona (Gordon, 1982), and (4) the upper part

of the type Morrowan Series of the Ozark Mountains region

(Henry, 1973; Sutherland and Henry, 1975, 1980; Henry

and Sutherland, 1977). I would correlate the type beds of

Antiquatonia coloradoensis with the Zia novamexicana

Brachiopod Subzone (upper Morrowan equivalents) of the

lower part of the La Pasada Formation, which is part of the

Linoproductus nodosus Brachiopod Zone of the Midcontinent and the eastern United States.

The Belden Shale in this part of the Central Colorado

Trough has been correlated with the upper Morrowan

through most of the Atokan Series (lower and middle Middle Pennsylvanian) (De Voto, 1972, 1980; Mallory and others, 1972; Chronic, 1979). My data confirm that the lower

part of the Belden Shale is indeed late Morrowan in age,

although certainly some of the individual components of the

Linoproductus nodosus Zone range upsection into generally

accepted Atokan equivalents. The thin sections examined by

me from colln. BL31394–PC contain only arenaceous foraminifers, and I do not have corroborative data from other

faunal groups, particularly conodonts, because of the limited amount of matrix material packed out by us in 1991. 

SYSTEMATIC PALEONTOLOGY 

Characters measured on Antiquatonia coloradoensis

and related taxa are shown in figure 4. In the succeeding text

and tables, the following terms are used:

PV = pedicle (ventral) valve; 

BV = brachial (dorsal) valve;

length of pedicle valve;

length of brachial valve (same as length from hinge to

anterior margin, HAM); 

hinge width = maximum width of hingeline of pedicle

valve; 

body width = maximum width of pedicle valve; 

height = height of pedicle valve; 

DBV = depth of brachial valve; 

SL = surface length of pedicle valve from beak to anterior margin measured on surface of valve in plane

of symmetry (also called “surface length,” “surface

measure,” or “arc length” by some authors); 

SLR = surface length to end of reticulation 

valve; and 

SLG = surface length to end of geniculation 

of trail) on brachial valve. 

The standard statistical parameters (descrip

tics) include the following: M=mode; =arithm

s=standard deviation; n=sample size. The statist

tions and formulae are given in Miller and Kahn 

Sokal and Rohlf (1969). 

Phylum BRACHIOPODA Duméril, 18

Class ARTICULATA Huxley, 1869

Order STROPHOMENIDA Öpik, 193

Suborder PRODUCTINA Waagen, 18

Superfamily PRODUCTACEA Gray, 1

Family PRODUCTIDAE Gray, 1840

Subfamily RETARIINAE Muir-Wood and Co

Genus ANTIQUATONIA Miloradovich, 

[emend. Sarycheva, 1949, and

Sarycheva and Sokolskaya, 1952]

DIAGNOSIS

Small to moderately large productids, subq

subrectangular in outline, generally having larg

ears with maximum width at ears; flanks of pe

steep, ears commonly set off from visceral area b

preauricular ridge bordered laterally by narrow s

valves generally geniculate, pedicle valve gene

trail, brachial valve with smaller trail; both valv

costellate, and rugose, strongly reticulate posteri

cle valve with three types of body spines: (1) gen

spines scattered on costae, (2) smaller spines on 

low angle to hinge margin, and (3) row of large

teroid spines generally set on curved preauricul

lateral slopes overhanging ears; spines rare o

valve. Brachial-valve interior with small triloba

process, median lobe of process dorsally directed

lophidium; lateral ridges diverging from hinge

not extending down lateral margins. Interior of pe

possessing transverse ridge across ears in som

dendritic adductor muscles set on median rid

grooved diductor muscle scars. 

DISCUSSION

Antiquatonia was split off of the genus Di

Muir-Wood (1930) by Miloradovich (1945), wh

the type species Productus antiquatus J. Sowe

p. 15, pl. 317, figs. 1, 5, 6), from the Lower Car

(Viséan) of Derbyshire, England. Sowerby’s sp

redescribed and refigured by Muir-Wood (192

pl. 6, fig. 7; pl. 7, figs. 1–4c; text-fig. 22), and 

was selected but not figured. Most of the ori

x
SYSTEMATIC PALEONTOLOGY 

P P

P

V V V

A A A

Ventral view of pedicle valve Dorsal view of brachial valve Lateral view with trace of brachial

valve shown by dotted line

H H H H B

B Body width

C

DBV

Length

Length BV

Height

Hinge width

Surface length

SLG

specimens of Productus antiquatus figured by Sowerby are

lost, and no interiors are available. For a more complete discussion of the type material, refer to Muir-Wood and Cooper (1960, p. 270–271). 

Antiquatonia was not fully described by Miloradovich

(1945) and was emended first by Sarycheva (1949, p. 167)

and subsequently by Sarycheva and Sokolskaya (1952,

p. 145). Lazarev (1990, p. 104, 125) assigned Antiquatonia

to the Family Productidae Gray, 1840, and the Subfamily

Retariinae Muir-Wood and Cooper, 1960, rather than to the

Dictyoclostidae Stehli (1954) and Dictyoclostinae Stehli

(1954), as proscribed by Muir-Wood and Cooper (1960) and

Muir-Wood (1965). 

As emended, Antiquatonia is a highly variable taxon in

the development of most external features. This variability

includes the size of the shell, extent of development of the

trails, degree of development of a sulcus, coarseness of

radial ornamentation, and size and density of body spines.

To me and most workers who study North American upper

Paleozoic brachiopods, one of the most salient characteristics of the species commonly referred to Antiquatonia is the

conspicuous row of large spines that crosses the flanks of

the pedicle valve; these spines are most commonly set on a

ridge bordering a shallow sulcus that separates the flanks of

the pedicle valve from the ears. Girty (1935, p. 7), in one of

the last papers that he published, called these ridges “preaural ridges.” Corresponding to the spine-bearing ridge on

the exterior is a small groove on the interior of the pedicle

valve and a corresponding ridge or flange on the interior of

the brachial valve that fitted into the groove, thus separating

the ears from the visceral area. These related features 

weak points on the valves, and the ears of both valves 

monly are broken off here. Girty (1935, p. 7) suggeste

“the function of the ridge [on the brachial-valve interi

not probably to shut off the auriculations from the inter

the shell but to serve as a hinging structure.” Girty (19

9) also suggested separating Productus hermosanus Gi

coloradoensis Girty, P. morrowensis Mather, and othe

cies of the “hermosanus Group” from the rest of the se

ticulate productids on the basis of this featur

Miloradovich’s (1945) original description of Antiqua

and in Sarycheva’s (1949) and Sarycheva and Sokolsk

(1952) emendment of that genus, references are ma

Girty’s preauricular ridges, and it is clear to me tha

structure was intended as a major generic entity. 

Sarycheva (1949) noted that a similar spine-be

ridge on the pedicle valve and the related ridges o

brachical-valve interior that separate the auricular cham

from the visceral area are not present in the type speci

Antiquatonia. Indeed, neither J. Sowerby’s (1821, p. 1

317, figs. 1, 5, 6) original illustrations nor Muir-W

(1928, pl. 7, figs. 1, 3) photographs of P. antiquatus 

these features, nor does Muir-Wood’s description me

them. The absence of these features in P. antiquatus is 

firmed by C. Howard Brunton (written commun., 199

is significant, however, that all of the species illustrat

Muir-Wood and Cooper (1960, pl. 94, figs. 1–11; p

figs. 4, 5) as examples of Antiquatonia do exhibit the p

ricular ridge on the brachial-valve interior. My investig

of A. coloradoensis shows that the degree of developm

Figure 4. Characters measured on specimens of Antiquatonia. Abbreviations: A=anterior, B=beak, P=posterior, H=hingeline, V=venter. Measured characters in text and tables: length (maximum length of shell)=VA, body width=maximum

width of pedicle valve (not hinge width), hinge width=maximum width of hingeline of pedicle valve, surface

length=BVA=length along arch, measured on surface of pedicle valve in plane of symmetry (also called “surface measure”

or “arc length from beak to anterior margin” by some authors), DBV=depth of brachial valve, length BV=length of brachial

valve, SLG=surface length to end of geniculation (beginning of trail), height=height of pedicle valve, c=number of costae

or costellae in 10 mm, measured at specified surface lengths.
12 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

the preauricular ridges and related features is an intraspecific feature, at least in this species. Furthermore, the

description for A. coloradoensis given herein of low internal

flange around the anterior visceral margin of the pedicle

valve (that rests in a groove on the brachial-valve interior) is

a feature not reported previously in Antiquatonia, although

it is developed in several other productid genera. 

RANGE

Species currently assigned to the genus Antiquatonia

are widely reported from the Lower Carboniferous (upper

Tournaisian and Viséan) of Europe, Asia, Africa, and Australia and from the Upper Carboniferous (Namurian and

higher strata) of Europe, Asia, and Australia. However, Lazarev (1990, p. 104) assigned the Tournaisian species to

Avonia Thomas (1914), emend. Muir-Wood (1928). In

North America, Antiquatonia is known from the lower part

of the Upper Mississippian (Chesterian) presumably

through the Lower Permian (Wolfcampian). However, Lazarev (1990, p. 104–106) would place the two species

described as Antiquatonia by Cooper and Grant (1975) from

the Lower Permian of Texas in a yet-unnamed genus more

closely related to the genus Reticulatia Muir-Wood and

Cooper (1960).

Antiquatonia coloradoensis (Girty)

Plates 1–3; plate 4, figures 1–12

Productus inflatus Girty (not McChesney, 1859, 1867),

1903, p. 359, pl. 3, figs. 1–3.

Productus inflatus? Girty (not McChesney, 1859, 1867),

1904, p. 52, pl. 11, figs. 5, 6; Boutwell (not McChesney,

1859, 1867), 1912, pl. 6, figs. 3, 4. 

Productus inflatus var. coloradoensis Girty, 1910, p. 216

(part); Girty, 1915a, p. 65 (not pl. 8, figs. 1, 2); 1920, pl.

55, figs. 3, 4.

Productus inflatus coloradoensis Girty. Plummer and

Moore, 1921, p. 48, 79, pl. 13, fig. 24.

Productus coloradoensis Girty. Girty, 1927, pl. 27, fig. 17

(not p. 73); 1935, p. 7–9, pl. 2, figs. 1–4. 

Not Productus coloradoensis Girty. Snider, 1915, p. 80. 

Not Productus cf. coloradoensis Arnold and Sadlick (not

Girty, 1910), 1962, p. 248.

Productus morrowensis Morgan (not Mather, 1915), 1924,

pl. 44, fig. 6. 

Productus semireticulatus Smith and Siebenthal (not Martin, 1809), 1907, p. 7; Price (not Martin, 1809), 1916, p.

699 [part]; Morningstar (not Martin, 1809), 1922, p. 181

[part], pl. 8, fig. 6; Butts (not Martin, 1809), 1926, p. 213

(not pl. 67, fig. 12); Morse (not Martin, 1809

311–312 [part], pl. 49, fig. 5(?), 6–8 (probably 

Productus [Dictyoclostus] semireticulatus Ch

Martin, 1809), 1981, p. 25; Chesnut (not Mar

p. 19.

Productus semireticulatus var. Morningstar (n

1809), 1922, p. 181–182 [part], pl. 8, figs. 4, 5

Dictyoclostus coloradoensis (Girty). Ragan, 1959

pl. 8, figs. 1–10.

Dictyoclostus inflatus var. coloradoensis (Girty)

1950, pl. 12, fig. 9.

Dictyoclostus [Antiquatonia] coloradoensis (Gi

and Ketner, 1975, p. 53. 

Not Dictyoclostus inflatus var. coloradoensis (G

rig, 1958, p. 7–8, 18–19, pl. 1, figs. 1–10; equa

tonia hermosanus (Girty). 

Dictyoclostus morrowensis (Girty). Gehrig, 195

19, pl. 1, figs. 11–15.

Not Dictyoclostus coloradoensis (Girty). Sut

p. 563. 

Antiquatonia coloradoensis (Girty). Stehli, 195

Muir-Wood and Cooper, 1960, p. 272–273; H

p. 57–58, pl. 7, figs. 6–9; Stevens, 1962 [at 

p. 623, (?) pl. 95, figs. 2, 3, 5; Chronic, 196

Sutherland and Harlow, 1973, p. 51, pl. 11, 

Sutherland, 1991, p. 188; Sutherland and Gray

p. 88–89, pl. 3, figs. 10–12; Rice and others, 1

table 2.

Not Antiquatonia cf. A. coloradoensis (Girty

1975, p. D48, pl. 5, figs. 1–9. 

Dictyoclostus hermosanus Searight and Palmer 

1903), 1957, p. 2131. Ragan (not Girty, 1903

69–72, text figs. 1–6, pl. 9, figs. 1–6.

Antiquatonia hermosanus [sic] Hoare (not Gi

1961, p. 56–57, pl. 6, figs. 1–3. 

Part Antiquatonia portlockiana (Norwood an

1855). Hoare, 1961, p. 53–54 [not pl. 5, figs. 1

material from Burgner Formation is in

A. coloradoensis.]

Antiquatonia portlockiana var. quadratia Stu

Hoare, 1968, p. 47–48 [middle and upper A

probably not Desmoinesian specimens], pl

1–11. 

Antiquatonia cf. A. portlockiana quadratia Stu

Hoare. Chesnut, 1981, p. 25, 57–66, pl. 9, figs

Neospirifer cameratus Ragan (not Morton, 18

pl. 3, fig. 10.
SYSTEMATIC PALEONTOLOGY 

DESCRIPTION BASED ON TYPE SPECIMENS

Shell strongly concavo-convex, subcircular in ventral

view, subtrapezoidal in anterior view; species medium to

large for genus; maximum width generally at about

midlength; largest relatively complete specimen (paratype

USNM 121482) measures 73 mm SL, 44.7 mm length of

pedicle valve, 41.5 mm maximum body width, 28.9 mm

hinge width, 31.0 mm height. 

Pedicle valve strongly convex near beak, becomes

progressively, uniformly less convex anteriorly, without

geniculation, almost planar on trail; trail variably developed, generally long, flaring slightly on some mature specimens, not distinctly set off from rest of valve; beak large,

slightly overhanging, slightly incurved; umbo inflated,

broad, prominent; steep lateral margins of umbo diverge at

100° to 110°, flare slightly anteriorly; very shallow, relatively narrow, rounded sulcus normally present on umbo,

extends to anterior margin without deepening or broadening; single specimen with mere flattening across umbo;

ears (generally broken off or chipped) moderately large,

depressed, blunt (nonalate) posterolateral extremities; ears

set off from venter by narrow, generally shallow groove

developed just laterally to auricular ridges; auricular ridges

subangular to subrounded, begin generally within about 12

mm of beak, enlarge slightly anteriorly, transect costellae,

normally extend onto trail (where developed), becoming

less conspicuous. 

Surface ornamentation of pedicle valve consists of

strong, fine, subrounded to subangular radial costellae on

beak and umbo; costellae become slightly less elevated,

broader anteriorly, increase irregularly by bifurcation; number of costellae varies considerably at given surface lengths

across venter from specimen to specimen, ranges from 14 to

19 in 10 mm at 20 mm SL (M=15 and 16, =16.8, n=13),

13 to 17 (M= about 16) at 30 mm SL, 13 to 16 (M=14) at 40

mm SL, 13 to 16 at 50 mm SL; costellae tend to fuse anterior to base of large body spines, particularly on trail, and

extend anteriorly from spine base as slightly raised ridges,

imparting irregular, unequally costellate surface on trail;

umbo has strong, concentric wrinkles, subequal in strength

to radial costellae, imparting strong semireticulate pattern to

umbonal area; surface length to end of reticulation ranges

from 29 to 36 mm, with M around 30 or 31 mm; wrinkles

extend onto ears, although not as prominently; ears ornamented with weaker radial costellae compared with umbo

and trail; entire valve has fine, irregular concentric growth

lamellae. Suberect body spines scattered across valve tend

to be arranged in crude rows radiating from beak area;

spines are small, spaced less than 5 mm apart on umbo,

become larger, spaced as far as 10 mm apart on trail, have

bases up to 1.1 mm in diameter. Single row of suberect halteroid spines initiates on either side of beak; spines are very

small near beak, spaced as close as 1.5 mm apart, enlarge

anteriorly, become more widely spaced, mounted on curved

x

ridge located ventrally to groove demarcating ears, 

monly not extending onto trail and to anterior marg

valve; largest spine base on anterior part of row mea

around 2.1 mm on single specimen. Distinct row o

hinge spines located on ears, extends laterally along h

line or diverges from hingeline at about 5°; largest obs

base of hinge spine 1.0 mm in diameter. 

Brachial valve gently concave across visceral are

slightly depressed, triangular area immediately anter

beak of pedicle valve; valve geniculate anteriorly, ante

erally, with short trail; angle of geniculation approxim

75° to 85°; line of geniculation somewhat rounded; vi

area generally has slight, broad fold, continuing ante

onto trail to anterior margin; ears slightly concave, s

from visceral region by slight fold. Valve reticulate a

visceral area, has strong concentric rugae subequally p

nent as radiating costellae; fine growth lamellae 

present; only costellae and fine growth lines conti

anterior to geniculation; ears ornamented with fine g

lines; brachial valve aspinose, with dimples correspo

to body spines on opposite valve; row of dimples on 

fold defining inner margin of ears corresponds to ro

spines on preauricular ridge on opposite valve. 

Interior of pedicle valve has strongly elevated, elo

adductor-muscle platform, strongly impressed did

scars, both posteriorly placed; adductor platforms slop

teriorly, have steep anterior margins, apparently sm

diductor scars are well defined anteriorly, laterally, 

deep, strong longitudinal striae; ears separated from vi

area by distinct, strong ridge that begins near posterior 

gin of valve, extends anteriorly to anterolateral extrem

visceral disc, becoming very weak, indistinct around an

margin of visceral disc; ridge corresponding in positi

preauricular groove on valve exterior; ridge slopes grad

posteriorly toward visceral area, has stronger, almost ve

anterior margin; ridge attains observed maximum wid

2.5 mm, height of about 3 mm near posterior extremity, 

mented with fine, rather widely spaced endospines, 

striae. Trail of valve with strong, anteriorly dir

endospines, arranged in rough, concentric rows; rema

of valve interior with ornamentation reflecting ornam

tion on valve exterior; auricular chamber smooth. 

Visceral area of brachial-valve interior poorly k

from type material. Cardinal process small, short, rou

sessile, has small, medially grooved median lobe cu

dorsally, flanked by two curved lateral lobes surrou

distinct pits; process supported by cardinal ridges, de

ing at low angle from hingeline; median septum, thin

extending to point just posterior of geniculation; add

scars moderately deeply impressed; brachial ridges 

weak flange apparently present as continuation of l

ridges near posterolateral margin of valve on many 

mens; where present or better developed, flange fit

posterior to ridge on opposite valve interior, continues 

riorly to about midlength, apparently generally disapp
14 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

weak groove located just anterior to ridge (shelf or flange on

pedicle-valve interior fits into this groove); trail ornamented

with coarse, anteriorly directed endospines crudely arranged

in concentric rows, bunched irregularly, producing slight,

discontinuous ridge across anterior margin of visceral disc. 

MEASUREMENTS

Measurements for the type material are given in

table 2. 

CATALOGED SPECIMENS

This paragraph lists reposited specimens of Antiquatonia coloradoensis (Girty) that are referenced in this paper.

Specimens reposited in the U.S. National Museum of Natural History:

figured lectotype (USNM 35345)

figured paralectotypes (USNM 8258a–c, 11660a,

121482, 121483, 208827–208830)

unfigured measured paralectotypes (USNM 8258d,

11660b)

unfigured measured topotypes (USNM

476630–476634)

unmeasured topotypes (USNM 476635–476639;

USNM 476661, 8 specimens)

figured hypotype (USNM 208805). 

Measured specimens reposited at the University of

Missouri (stars indicate specimens from the Burgner Formation, southwestern Missouri, originally identified by Ragan

(1959) or Hoare (1961) as Antiquatonia hermosana (Girty): 

UM *13192 *13664a–h 

*13193 *13665a–j 

*13196 13671a–f 

*13197 13672a–g

*13657a–e, g 13673a–f

*13662a–g 14353

*13663a–f, h 14354. 

Specimens of Antiquatonia portlockiana var. quadratia

Sturgeon and Hoare (now referred to A. coloradoensis)

reposited at Ohio State University: 

figured holotype (OSU 26020)

figured paratypes (OSU 26022–26026, 47102)

unfigured measured paratypes (OSU 26021, 26027,

47103–47109). 

DISCUSSION OF TYPE MATERIAL

My redescription of Antiquatonia coloradoensis is

based on 19 specimens from the original lot from USGS

colln. GR2281–PC and an additional 41 specimens plus

fragments from the four collections considered 

topotypic material (USGS collns. BL

BL6852A–PC, BL12096–PC, and BL31394–PC

mary types originally were identified by Girty 

359) as Productus inflatus McChesney (1859, 1

(1903, pl. 3, figs. 1–3) figured three of these sp

engraved drawings (see table 5). Later, in a pape

dedicated to naming species from the Fayette

(Chesterian) of the Ozarks, Girty (1910, p. 216

following statement: “In 1903, I referred to McC

inflatus a group of shells from the Pennsylvania

rado, expressing at the time a certain doubt wh

were actually identical with it [McChesney’s t

this group I would now suggest the varietal nam

oensis.” To me, Girty clearly was designating th

material the primary types of Productus inflatus v

doensis, and, because the three figured specim

from colln. GR2281–PC, it becomes the type 

Gordon (1975, p. D48) selected as lectotype 

specimens (USNM 35345) that Girty (1903, p

1–1b) initially illustrated and designated the rem

specimens that Girty figured (USNM 121482 an

and his 19 unfigured specimens (divided into l

8258 and 11660) as paralectotypes. In the curre

am illustrating photographically for the first time 

type (pl. 1, figs. 1–4) and the two previously fig

lectotypes (pl. 1, figs. 5–13) and an additi

previously unfigured paralectotypes (pl. 1, figs. 14

figs. 1–15; pl. 3, figs. 1–15). 

Most specimens in the five type collections 

what decorticated, cracked, and at least partial

The most complete specimens are the lectotype 

herein as pl. 1, figs. 1–4) and two of the par

(pl. 1, figs. 5–8; pl. 2, figs. 5–14). The ears are sl

ken on all three specimens, and the trail is bro

right side of the first-cited paralectotype. The de

of a trail in both valves seems to be a variable fe

specimen figured as plate 1, figures 5–8, has m

trail preserved, and the internal mold figured 

figures 5–7, has the complete pedicle-valve trai

the true shape of the slightly flaring trail, althoug

imen is crushed posteriorly. In other specimens 

ple, that shown in pl. 1, fig. 6), the trail seems to 

much shorter, although the trail, like the ears, is 

most fragile portions of the shell and is easily bro

The brachial-valve exterior is shown on fo

paralectotypes (pl. 1, figs. 13, 16; pl. 2, figs. 4, 

eight unfigured paralectotypes, although the trai

ken on most of these. Portions of the interior of t

valve can be seen on several of the specimens 

exteriors are partially exfoliated (decorticated) 

away or where the most of the shell was etche

weathering. Plate 1, figure 13, reveals the obscu

flange that, on some specimens, extended along 

olateral extremity of the visceral disc, fitted into 
Table 2. Measurements (in millimeters) of figured and (or) measured primary type specimens and topotypes of Antiquatonia coloradoensis (GMorrowan part of the Belden Shale, Park County, Colo., and of a hypotype from the upper Atokan Burgner Formation, Jasper County, Mo. [Whole shells are figured and (or) measured primary type specimens and topotypes from the Belden Shale, Park County, Colo. The disarticulated brachial valve Burgner Formation, Jasper County, Mo. Specimens are reposited at the U.S. National Museum of Natural History (USNM), Washington, D.C. Asterisk indicatecle valve; dash means data not available. Characters are defined in figure 4. Abbreviations: BV, brachial valve; HAM, hinge to anterior margin; PV, pedicle vaalong venter; SLG, surface length to geniculation (beginning of trail); SLR, surface length to end of reticulation. Types: h, hypotype; L, lectotype; P, figured paraparalectotype; t, measured topotype. Designations for dimensions that could not be measured exactly: a, approximate; e, estimated; m, minimum; t, estimated 

Measurements by T.W. Henry]

1Excluding cardinal process.

USNM catalog 

number Type

Length

SL

Width

Height Depth SLR (PV) SLG (BV) Trail length

Costellae in 

PV HAM (BV) Hinge Body 20 mm 30 mm 

Whole shells (articulated) from Belden Shale, Park County, Colo.

208830 *........... P 25 m 19 m 38 m 29.0 e 29.7 m 14.4 m 12 m – 18 a – –––476630.............. t 27 m 20 m 40 m – – – 8 m – – – –––476631.............. t 27 m 24 m 42 m – 35 m – 10 m 28 a 19 – 14 a 13 a 1476632.............. t 27 m – 45 m 22.0 a 31.1 a – 14.7 28 a 20 – –––8258c................ P 28 m – 45 m 25 m 33 m – 14.5 29 20 – 16 17 1208829.............. P 30 m 24 m 52 m 31.0 t 32.9 m – 16.0 32 20 – 19 16 1121483.............. P 35 m 25 m 63 m 33.4 39 m 21.5 m 16.9 32 20 – 15 ––11660b.............. p 35 m 27 m 65 m 30.0 e 39.6 a 21.8 m – 32 – – 16 14 1476633.............. t 36.3 a – 58 – 36.0 a 21.0 e – 28 – – 14 14 18258d................ p 37 m 28 m 62 m – 42.6 a – 17.0 32 22 – 17 a 16 1208827.............. P 37.7 a 28.4 a 68 32.8 t 39.7 a 22.7 15.8 32 21 – 15 14 18258b................ P 38.1 a 30.3 a 68 25 m 38.6 a 21.9 a – 29 – – 16 15 1208828.............. P 38.1 30.3 a 68 36.0 t 46.3 a 24.5 a 16.0 30 21 – 15 17 18258a................ P 40 m 30.3 a 70 m 38.6 t 46 e 26 e – 31 – – 15 a 15 1476634.............. t 40.5 a – 80 – 48.8 a 26.6 a – 30 a – – –––35345................ L 41.7 a 31.0 a 72 34.3 39.7 25.1 a – 34 – – 16 17 1121482.............. P 44.7 – 73 28.9 a 41.5 a 31.0 a – 33 – – 14 13 111660a*............ P 47.5 e 34.4 a 75 e 33.9 a 47.2 a 21 m – – – – –––

Brachial valve (disarticulated) from Burgner Formation, Jasper County, Mo.

208805.............. h – 133.6 157 42.4 – – – – 123 34 –––
16 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

on the opposite valve, and helped seal off the auricular area

from the body chamber or visceral cavity. The distinct lateral flange and groove also can be seen clearly in plate 2,

figures 6, 8–10, which show a specimen that has a portion

of the brachial valve broken away; the weak extension of

this ridge or flange across the anterior part of the valve also

is shown in plate 1, figure 19. The large internal mold (pl.

3, figs. 5–7) is of a complete pedicle valve, although the

beak area is crushed, and the trail is long and slightly flaring at its extremities. This specimen had only a slight shelf

extending around the anterior and anterolateral parts of the

visceral cavity (see pl. 3, fig. 5). None of the specimens

from the type locality reveal a strong flange or ridge

extending from the auricular ridge distinctly and completely around the anterior portion of the visceral area of

the brachial-valve interior, and, in some of the specimens,

this ridge and groove apparently were not developed at all.

The flange on the interior of the pedicle valve (pl. 2, fig. 9)

resting in a groove in the interior of the brachial valve (pl.

2, figs. 6 and 11) is a distinctly different pattern from that

developed in the genus Tesuquea Sutherland and Harlow

(1973), in which a flange on the interior of the brachial

valve (Sutherland and Harlow, 1973, pl. 6, figs. 9, 17, and

18) is opposite a thickened low ridge on the interior of the

pedicle valve (their pl. 6, fig. 18). 

The details of the musculature and cardinal process of

the interior of the brachial valve are not preserved on any of

the primary types, but some of the internal features can be

seen on four of the unfigured topotypes from colln.

BL12096–PC (USNM 476630–476632 and 476636), and

on two uncataloged specimens from colln. BL31394–PC.

The description of the musculature of the pedicle-valve

interior is based on two internal molds (paralectotypes) figured herein (pl. 2, fig. 15; pl. 3, figs. 5–7), and eight unreposited topotypes. 

The halteroid spines on the ridge on the pedicle-valve

exterior were very large and long. Although not collectable,

a specimen at the type locality, apparently in growth position, was observed by me that had slightly curving and

tapering spines that on the anterior part of the valve attained

a length of at least 4.5 cm. 

INCLUSION OF ANTIQUATONIA PORTLOCKIANA VAR. 

QUADRATIA IN A. COLORADOENSIS

Sturgeon and Hoare (1968, p. 47–48, pl. 15, figs. 1–11)

described Antiquatonia portlockiana var. quadratia from

16 specimens from strata that they called the “Lower Mercer limestone and shale” from southeastern Ohio. The holotype (OSU 26020) and 13 paratypes (OSU 26021–26025;

OSU 47102–47109) are from a locality in Hocking County.

The two remaining paratypes (OSU 26026–26027) are

reportedly from the same stratigraphic units in nearby areas.

These “Lower Mercer” strata correlate with the upper part

of the Atokan Series (Zone of Fusulinella), ac

Douglass (1987, p. 12, 13). Sturgeon and Ho

p. 48) stated that their variety is “rare to comm

Lowellville, Boggs, Upper Mercer, and Vanpor

rare to abundant in the Lower Mercer and Pu

units.” In other words, according to their identific

variety quadratia ranges from potentially upper 

strata through upper Atokan equivalents of the 

Formation and into the lower Desmoinesian equ

the Allegheny Formation. They included in their 

the material identified by Morningstar (1922) bo

ductus semireticulatus Martin and P. semireticula

The “taxon” Antiquatonia portlockiana var

was essentially typologically defined. Little of th

ity inherent in the type collections and discussed 

included in either the description or discussion in 

nal reference by Sturgeon and Hoare (1968, p

have examined the type specimens of this varie

measurements of the type material are presented i

include the upper Morrowan and Atokan specime

quatonia portlockiana var. quadratia Sturgeon an

the synonymy of A. coloradoensis (Girty) in s

minor differences discussed below. At best, A. po

var. quadratia should be considered a named 

Girty’s A. coloradoensis. 

Sturgeon and Hoare (1968, p. 48) contend

coloradoensis (Girty) is less convex posteriorly 

quadratia.” The differences in external shape and 

tation and in degree of development of intern

between the types of var. quadratia and the pr

secondary types of A. coloradoensis are insigni

range of variation in size and shape exhibited b

of quadratia is encompassed by the observed ex

ation of A. coloradoensis. For example, Stu

Hoare’s holotype (OSU 26020, refigured herei

figs. 16–20) is almost identical in size and shape 

the degree of convexity of the posterior part of 

valve) to the paralectotype that I figure as pl. 2

(USNM 208829). The newly figured paratype (O

pl. 4, figs. 10–12) has its analog in the large, bro

totype (USNM 208828) figured as plate 2, fig

The only difference in these two specimens is th

totype USNM 208828 has no sulcus, an atypic

unknown situation for the primary and seconda

A. coloradoensis. 

One observable difference between Antiqua

lockiana var. quadratia and A. coloradoensis (se

is in the radial ornamentation, duly noted by Stu

Hoare (1968, p. 48). However, both the types of A

oensis and those of quadratia have about the sam

of costellae, measured at comparable surface len

exterior of the pedicle valve (see tables 2, 3). For 

the number of costellae measured in 10 mm at 

ranges from 15 to 17 (M=16, n=6), well within th

A. coloradoensis. Both sets of types contain spe
Table 3. Measurements (in millimeters) of primary type specimens of Antiquatonia portlockiana var. quadratia Sturgeon and Hoare 

coloradoensis (Girty)) from upper Atokan strata of the “lower Mercer limestone and shale” of the Pottsville Formation, southeastern Ohio. 

[Specimens are reposited in the Orton Geological Museum at Ohio State University (OSU). Asterisk indicates internal mold of pedicle valve; dash means data not defined in figure 4. Abbreviations: BV, brachial valve; HAM, hinge to anterior margin; PV, pedicle valve; SL, surface length along venter; SLG, surface length to g

trail); SLR, surface length to end of reticulation. Types: H, holotype; P, figured paralectotype, p, measured paralectotype. Designations for dimensions that could no

approximate; e, estimated; m, minimum; t, estimated from twice half-width. Measurements by T.W. Henry]

OSU catalog 

number Type

Length

SL

Width

Height Depth SLR (PV) SLG (BV) Trail length

Costellae i

PV HAM (BV) Hinge Body 20 mm 30 mm 

Whole shells (articulated) or disarticulated pedicle valves

26027* ............. p 25.5 e – 45 e 35 e 34 e 21 e – – – – –––26021 ............... p 30.8 a 21.9 a 60 23.3 a 34.8 20.5 – 35 – – 16 16 – 47102 ............... P 32 m 24.5 m 50 m 42 t 46 e 18 m – 25 – – 17 16 126023 ............... P 33.0 25 e 55 e 36 t 36.1 a 20 e – 27 – – –––26026 ............... P 33.2 a 22.3 a 62 32.4 a 22.3 a 20.2 a – 27 – – 16 15 – 26020 ............... H 33.5 27.0 a 64 32.3 35.5 21.7 – 27 – – 15 a – – 47105 ............... p 34.1 a 26.5 a 65 a 28.7 a 36.7 a 20.9 a – 25 a – – 16 15 147104 ............... p 35 m 29 m 52 m 36.5 e 36.5 m – – 25 e – – –––47103 ............... p 40 e 32.5 e 65 e 38 t 44 e 21.6 a – 32 – – 16 a – – 

Brachial valves (disarticulated)

26024 ............... P – 124.6 130 31.8 33.8 – – – 123 8 – – – 

47107 ............... p – 125.0 127 a 28.3 a 34.8 a – – – 126 7 a – – – 26022 ............... P – 125.2 a 128 21 e 36 e – – – 121 7 – – – 26025 ............... P – 126.3 130 32.7 a 37.3 a – – – 123 6 – – – 47106 ............... p 127e 128 e 26.6 t 33.2 – – – 123 e – – – – – 47108 ............... p – 128.8 a 129 e 26.4 36 e – – – 124 a 8 a – – – 47109 ............... p – 128.9 a 128 34.7 a 40.8 a – – – 124 8 a – – – 
18 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

which the costellae fuse anteriorly to the bases of the large

body spines and form “ribs,” particularly on the anterior part

of the pedicle valve over the body cavity and on the trail.

These “ribs” extend anteriorly from the spine bases as a

cluster of costellae on the slightly raised ridges, imparting

an irregular, unequally costellate surface, similar to that of

A. portlockiana (Norwood and Pratten, 1855). However, a

few of the paratypes of quadratia do have only one large,

raised costa extending anteriorly from the body spine bases

rather than a cluster of “fasciculate” costellae (for example,

OSU 26026, pl. 3, figs. 24, 25). Nonetheless, the right side

of the anterior view of this specimen (pl. 3, fig. 25) shows

the costa splitting into two costellae farther anteriorly on the

trail. A small proportion of the types of A. coloradoensis

have this single larger, higher costa anterior to the large

body spines. This is considered a trivial difference. 

One other characteristic of the radial ornamentation that

Sturgeon and Hoare (1968, p. 47–48) did note was that the

costellae tend to become “obsolete” anteriorly, particularly

on the trail. However, all of the shells of the types of A. coloradoensis are at least slightly decorticated. The types of

quadratia that show the costellation extending to the anterior

extremities of the valve are also decorticated or worn. All of

the shells that have the ornamentation becoming indistinct or

subdued are well preserved and not decorticated at all. The

strength of radial ornamentation is therefore a function of

preservation and is not a valid taxonomic discriminator. This

effect of preservation on the strength of the radial ornamentation has been observed by me with many different taxa of

productid (and chonetid) brachiopods. 

The primary types of A. portlockiana var. quadratia

differ from the types of A. coloradoensis in having a generally relatively long trail and in having a generally weak

flange and groove anteriorly and anterolaterally ringing the

visceral area on the interiors. All of the disarticulated brachial-valve interiors from Sturgeon and Hoare’s collection

show at least a faint extension from the auricular ridges of a

raised area or flange delimiting the visceral cavity anteriorly

and anterolaterally. The disarticulated brachial valve figured

as plate 4, figure 9 (OSU 26025), seems to have come from

the same individual as the disarticulated pedicle valve figured as plate 3, figures 21 and 22 (OSU 26023). The ridge

that continues anterolaterally from the cardinal ridges on the

brachial-valve interior, separating the visceral chamber

from the auricular area, fits neatly inside the ridge on the

pedicle-valve interior that also separates the auricular area

from the visceral chamber. This brachial-valve interior

exhibits the weakest development of this ringing feature of

the three refigured interiors and has the least amount of

thickening of the valve at the point of geniculation. In fact,

the shell material of this large valve is very thin. Three of

the unfigured paratypes (OSU 47107–47109) have at best a

weak extension and very thin valves also like OSU 26025. 

In contrast to the situation just described, t

relatively complete specimens (OSU 26020, 2

26026) possess valves that are thickened at the d

of the trail anterior to the visceral area. The holo

26020; see pl. 3, figs. 16–20) has a ragged, lamel

opment of shelly material at this point. The dark 

anterior margin of the specimen shown in plate 3

is actually material from the pedicle valve. One o

two articulated paratypes (OSU 26026, pl. 3, fi

shows similar but less drastic lamellation of the s

point, as does the other paratype that I did not fi

26021). Two of the other figured disarticulate

valves (pl. 4, figs. 6–8 and figs. 1–5, OSU 26022 

respectively) and one of the unfigured interiors (O

are also thickened near the anterior margin in a si

ner. The first of these disarticulated valves (OSU 

strongly thickened and heavily lamellose. It can 

reasonably that the trails of all of these specimens 

and extended some distance above the substrate. 

repeatedly broken and regenerated. At any time

were probably relatively short. To me, these thick

imens are merely reflecting the variability inher

sample of a population in response to environme

tions and do not have any taxonomic significance

The paratype (OSU 26024) of quadratia ill

plate 4, figures 1–5, and discussed above has 

coarsely developed internal features. At the poin

ridge delimiting the auricular chamber extends a

ally beyond the ears, it deflects somewhat abruptl

at this point is a rather deep, concave “socket-like

that is not as clearly seen on any other specimen o

tonia that I have examined. The muscle platform

brachial ridges are raised strongly on this specim

anterior portion of the median septum, normally 

mm high and less than 1 mm thick, is raised app

3.5 mm above the floor of the valve and is 3.0 m

its lateral extremity where it bluntly terminates. T

surface of the septum is slightly sulcate at its term

The large collections of Antiquatonia from 

graphically higher Putnam Hill Limestone Mem

Allegheny Formation that Sturgeon and Hoa

included in this variety and that Mornings

identified from this member as Productus sem

Martin or her undescribed variety should be exam

fully, because I believe that it is not likely that 

coloradoensis. Douglass (1987) reported the 

Beedina leei (Skinner) in the Putnam Hill Limes

ber and correlated these strata with the basal 

Desmoinesian Series. 

Chesnut (1981, p. 25, 56–57, pl. 9, figs. 4–6

Antiquatonia cf. A. portlockiana quadratia Stu

Hoare three specimens from the Magoffin Mem

kan) of the Breathitt Formation from Perry Co

also have large collections of this taxon from road

the Daniel Boone Parkway in the same area, an
SYSTEMATIC PALEONTOLOGY 

reservation, I include the Magoffin species in Girty’s Antiquatonia coloradoensis. The specimen figured by Chesnut

(1981) as his plate 9, figure 6, is almost identical to the lectotype of Girty’s species (see pl. 1, figs. 1–4, of current

paper). I also include in A. coloradoensis specimens from

my collections from the Winifrede Shale Member (formally

named by Blake, Keiser, and Rice, 1994, p. 47–48) of the

Kanawaha Formation in southern and central West Virginia.

The Winifrede is the same marine band as the Magoffin

Member of Kentucky and northern Tennessee.

DISCUSSION OF SUPPLEMENTAL 

BRACHIAL-VALVE INTERIORS

Well-preserved, complete interiors of Antiquatonia

coloradoensis are rare. Among the best brachial-valve interiors available, at least in terms of the details of the visceral

area and trail, are (1) the upper Atokan paratypes of var.

quadratia discussed in detail in the preceding section,

(2) several specimens from the Burgner Formation (upper

Atokan; see discussion of age and notations about the specimens from the “E.J. Palmer collection” in the section below

on “Comparison with Antiquatonia hermosana”) of Missouri, and (3) several specimens from upper Morrowan and

lower Atokan strata in the La Pasada Formation of the Sangre de Cristo Mountains in northern New Mexico.

Among the exceptionally well preserved specimens

that I examined from USGS collns. BL8113–PC and

BL25128–PC from the Burgner Formation are three relatively complete brachial-valve interiors, the best one of

which is illustrated herein (USNM 208805, pl. 2, figs.

16–21). This specimen is an almost complete valve with relatively large, potentially pointed ears and a long trail. The

maximum width of the shell for this specimen may have

been at the hingeline rather than farther anteriorly. The lateral ridges supporting the cardinal process deflect anteriorly

near the posterolateral extremities and form a ridge that separates the auricular chambers from the visceral disc (see pl.

2, fig. 17). However, these marginal ridges continue only a

few millimeters anteriorly before disappearing and clearly

do not continue across the anterior portion of the valve (see

pl. 2, figs. 16–18). Rather than a flange delimiting the visceral chamber anteriorly and anterolaterally, at best, only a

rough, weakly and irregularly developed thickening is

present. The other two interiors are very similar in this

respect, and this is also true for the fragmentary brachialvalve interior of Antiquatonia coloradoensis (UM 13193)

that was figured by Hoare (1961, pl. 6, fig. 3) as A. hermosana (Girty) (see point 5 of “General Discussion” for amplification). Note on the hypotype that I am illustrating

(USNM 208805, pl. 2, fig. 16) that the coarser endospines

are confined to the auricular chambers and to a band about

10 mm wide around the anterior and anterolateral portions

of the valve just anterior to the geniculation. The interior of

the trail of the brachial valve is ornamented only with 

tellae, fine scattered endospines, and very fine g

lamellae. The pedicle valve corresponding to this in

would probably have been about the same size and sha

the paralectotype from Colorado illustrated in pla

figures 5–14 (USNM 208828), the maximum wid

which is anterior to the hingeline. 

Sutherland and Harlow’s (1973) collection (it

above) has a few brachial-valve interiors, but neither 

two specimens that reveal the anterior margin and tra

any suggestion of the internal groove around the an

margin of the visceral disc.

It would appear from the preceding analysis of the 

plemental material that both the development of the l

of the trails and the degree of development (if any) o

internal flange and groove in the anterior and anterol

parts of the shell of Antiquatonia coloradoensis are va

characteristics within the species. Where developed

structure of a flange on the interior of the pedicle 

(pl. 2, fig. 9) resting in a groove in the brachial valve 

figs. 6, 11) is a distinctly different pattern from that fou

Tesuquea, in which a flange on the interior of the br

valve (Sutherland and Harlow, 1973, pl. 6, figs. 9, 17, 

opposite a thickened low ridge on the interior of the p

valve (1973, pl. 6, fig. 18). 

 INFERRED PHYLOGENY AND

COMPARISON WITH OTHER SPECIES

INFERRED PHYLOGENY

Antiquatonia coloradoensis probably evolved fro

morrowensis (Mather, 1915), a robust form, the typ

which are from a bioclastic, non-oolitic limestone b

the basal part of the Prairie Grove Member of the Hal

mation (lower middle Morrowan) in northwestern A

sas. There, Antiquatonia morrowensis ranges throug

rest of the Hale Formation, through the Brentwood L

stone Member of the Bloyd Shale, and apparently in

basal part of the Dye Shale Member of the Bloyd (

Morrowan). In turn, A. coloradoensis gave rise to A. h

sana (Girty), common in lower Desmoinesian strata 

southern Rocky Mountains and Paradox Basin. The 

species possibly gave rise to Antiquatonia portloc

(Norwood and Pratten), which is common in uppe

middle and upper Desmoinesian strata in the Rocky M

tains and which is reported from throughout Desmoin

strata in the Midcontinent, Illinois Basin, and Appala

Basin. The trend in coarsening of the radial ornamen

of the lineage of Antiquatonia in North America fro

morrowensis through A. coloradoensis, through A. h

sana, and into A. portlockiana is striking and is sho

figure 5. Ages of A. morrowensis and its descendan

shown in figure 6.
20 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

A. portlockiana (Norwood an

New Mexico

x = 11.5, n = 19

A. hermosa

New Mexic

x = 13.2, n 

A. coloradoen

Missouri

x = 16.3, n = 4

A. coloradoe

Colorado

x = 16.8, n = 

A. coloradoensis (Girty)

New Mexico

x = 17.4, n = 19

A. morrowensis (Mather)

Arkansas (type specimens)

x = 21.6, n = 43

NUMBER OF SPECIMENS

NUMBER OF COSTAE/COSTELLAE 

10 MILLIMETERS AT 20 MILLIMETER

12

10

8

6

4

2

0

8

10

6

4

2

0

16

12

14

10

8

6

4

2

0

8

6

4

2

0

6

4

2

0

0 9 10 11 12 13 14 15 16 17 18 19 20 21 

4

2

0

COMPARISON WITH ANTIQUATONIA MORROWENSIS

The types of Antiquatonia coloradoensis (Girty) exhibit

about the same size and shape as the types of A. morrowensis

(Mather, 1915) and also possess a row of robust spines on the

preauricular ridge along the flanks of the pedicle valve. Antiquatonia coloradoensis has a slightly deeper and more consistently developed sulcus, somewhat broader umbo, lower

and more uniform convexity, a larger body cavity than

Mather’s species, and coarser radial ornamentation. The

larger body cavity of A. coloradoensis is reflected by the

greater distance to the end of reticulation on the pedicle valve

(contrast a range from 28 to 34 mm SLR, =32.4, to a range

from 18 to 28 mm SLR, =22.6). Perhaps the most conspicuous difference between the two species is in the relative

coarseness of the radial ornamentation (see fig. 5). Antiquatonia coloradoensis is conspicuously more coarsely ornamented, the type collection having between 14 and 19

costellae ( =16.8, M=15 and 16, n=13) in 10 mm at 20 mm

SL (fig. 5). Type specimens of A. morrowensis from the basal

beds of the Prairie Grove Member of the Hale Formation

range between 17 and 25 costellae in 10 mm at 20 mm SL

( =21.2, M=21 and 22, n=43). Collections of A. morrowensis from middle and basal upper Morrowan strata in the

Ozarks (the Brentwood Limestone Member and the basal

part of the Dye Shale Member of the Bloyd Shale, respectively, in northwestern Arkansas and equivalent strata in

northeastern Oklahoma) do show a slight but statistically

nonsignificant increase in the coarseness of radial ornamentation. The differences in this factor between the types of A.

morrowensis and the type specimens of A. coloradoensis, in

spite of the relatively small sample size of the latter species,

are statistically highly significant. 

x

x

x

x

Figure 5. Number of costellae and costae in 10 mm across venter

at 20 mm SL (surface length) for Antiquatonia morrowensis

(Mather) and its probable descendants, listed from oldest to youngest: (1) A. morrowensis (Mather) from its type locality (early middle Morrowan) in Ozark Mountains, northwestern Arkansas; (2) A.

coloradoensis (Girty) from upper Morrowan rocks of northern New

Mexico, (3) A. coloradoensis (Girty) from its type locality (late

Morrowan) in Colorado, (4) A. coloradoensis (Girty) from upper

Atokan rocks of Missouri, (5) A. hermonsana (Girty) from lower

Desmoinesian rocks of northern New Mexico, and (6) A. portlockiana (Norwood and Pratten) from lowest middle and lower upper

Desmoinesian rocks of northern New Mexico. Note general progressive increase in coarseness of radial ornamentation (decrease in

number of costellae and costae in 10 mm) in younger species. Differences between mean numbers of costae and costellae ( ) are statistically highly significant between all species. No significant

differences exist between means of the samples of A. coloradoensis

from the type locality in Colorado, from New Mexico, or from Missouri. Data for New Mexico material from Sutherland and Harlow

(1973, fig. 24); other data from current study.

x
SYSTEMATIC PALEONTOLOGY 

Desmoinesian

Atokan

Morrowan

Upper

Middle

Lower

A. morrowensis

A. coloradoensis

A. hermosana

A. portlockiana

Upper

Middle

Lower

COMPARISON WITH ANTIQUATONIA HERMOSANA

Antiquatonia hermosana also was described initially

by Girty in his 1903 paper (p. 358, pl. 2, figs. 1–4) as a variety of Productus semireticulatus Martin (1809) from lower,

middle, and upper Desmoinesian strata of the Hermosa Formation (Middle and Upper Pennsylvanian) in southwestern

Colorado. Girty (1903, p. 358) noted considerable overlap

in external features between these two taxa. Although the

primary type material and other collections from these strata

show considerable physical variation, A. hermosana differs

from A. coloradoensis (Girty) in being generally larger and

more robust and in having a broader and longer body cavity

(thus continuing a trend from A. morrowensis), a flatter venter, a broader and shallower sulcus, slightly more enrolled

ears, thus imparting a less angular appearance to the lateral

extremities of the valves, and coarser radial ornamentation

(see fig. 5). Sutherland and Harlow (1973, p. 51) made the

following statement about the overlap and differences in

radial ornamentation between A. coloradoensis and A. hermosana in their collections from the southern Sangre de

Cristo Mountains, New Mexico: 

Our New Mexico specimens [identified as A. coloradoensis] from the

[upper] Morrowan and lower Atokan intervals are closely similar to the

Colorado specimens of A. coloradoensis, described by Girty, and are distinctly separable from our large collections of A. hermosana in the lower

Desmoinesian. The lower Atokan specimens of A. coloradoensis have

slightly coarser costae on the average than those in the [upper] Morrowan,

and the lowest Desmoinesian specimens of A. hermosana have finer costae

on the average than typical for that species. Although higher in the lower

Desmoinesian a few specimens have finer costae approaching those of A.

coloradoensis, the average specimen is much coarser.

Sutherland and Harlow (1973, p. 52) noted that the

upper Atokan sequence in the southern Sangre de Cristo

Mountains is sparsely fossiliferous but observed that their

specimens of Antiquatonia hermosana from the lowermost

Desmoinesian strata appear to be transitional forms from A.

coloradoensis. Significantly, A. portlockiana occurs in

northern New Mexico only in the highest middle Desm

sian and lower upper Desmoinesian (Sutherland and 

low, 1973, p. 52). I examined three well-pres

collections of specimens of Antiquatonia from sin

deposits constituting the Burgner Formation from (1

E.J. Palmer collection, University of Missouri (UM) lo

(2) USGS collections BL8113–PC and BL25128–PC (

to G.H. Girty from the Palmer collection); and (3) H

(1961) Eldon collection from UM loc. 37. The Palme

lection (see Searight and Palmer, 1957, p. 2127–

Ragan, 1959) consists of material acquired from old 

mine dumps near the center of sec. 20, T. 28 N., R. 3

Jasper County, Mo. Hoare’s (1961) collection (UM lo

was from material exposed on the dump of an old coal 

on the Neville farm about 2.5 mi southwest of Eldon, 

NW1/4 SE1/4 sec. 6, T. 41 N., R. 16 W., Miller County

These sinkhole deposits are dated as Atokan on the ba

cephalopods (Unklesbay and Palmer, 1958) and middl

late Atokan on the basis of associated fusulinids and 

odonts (see M.L. Thompson, 1953; T.L. Thompson, 1

R.C. Douglass (written commun., January 1989) rep

that the matrix from USGS colln. BL25128–PC contai

foraminifer Bradyina sp. and an advanced form 

Fusulinella, indicating a definite late Atokan age. Th

USGS lot (BL8113–PC) is part of Palmer’s collection, 

which Girty (1935, pl. 2, figs. 1–4) figured a superbly 

served pedicle-valve exterior as Productus colorado

The repository was not mentioned in Girty’s paper, 

could not find the specimen in the USNM or USGS c

tions, and R.L. Ethington (written commun., July 15, 

could not locate it in the University of Missouri collec

This specimen showed the acute and distinctly angula

terolateral extremities that characterize this species. 

and Siebenthal (1907, p. 7) earlier had listed a collect

invertebrate fossils from this same mine dump o

Burgner mine property and identified the large Antiqua

as Productus semireticulatus (Martin). 

My measurements of the large collection of gen

well preserved material from UM loc. 47 are presen

table 4. Searight and Palmer (1957) and Ragan (

reported both Antiquatonia coloradoensis and A. herm

from this and other Burgner collections in southw

Missouri, as did Hoare (1961) from other localities. St

cally, no significant difference exists (at 95 percent 

dence limits) in the coarseness of radial ornamen

between the Burgner set that Ragan identified as co

doensis and the set from this locality that he identifi

hermosanus (the latter marked by two asterisks besid

catalog number in table 4). Furthermore, because co

ous variation characterizes all of the other significant 

nomic characters noted between the two sets, I conc

that only one variable species (A. coloradoensis) is pr

in the Burgner collections.

Figure 6. Lineage of Antiquatonia in Lower and Middle Pennsylvanian strata in the United States.
Table 4. Measurements (in millimeters) of specimens identified as Antiquatonia coloradoensis (Girty) and A. hermosana (Girty) by Ragan (1Formation (upper Atokan), Jasper County, Mo., and referred herein to A. coloradoensis (Girty). [Specimens are reposited at the University of Missouri (UM) at Columbia. Specimens are from the E.J. Palmer collection from mine dumps at UM locality 47 in SE

W., 1.5 mi south of Carterville, Jasper County, Mo. Asterisk in first column indicates internal mold of pedicle valve; double asterisks after type identification indicaRagan (1959) as A. hermosana; dash means data not available. Characters are defined in figure 4. Abbreviations: BV, brachial valve; HAM, hinge to anterior marg

surface length along venter; SLG, surface length to geniculation (beginning of trail); SLR, surface length to end of reticulation. Types: R, specimen illustrated by R

not illustrated by Ragan (1959). Designations for dimensions that could not be measured exactly: a, approximate; e, estimated; m, minimum; t, estimated from tw

ments by T.W. Henry] UM catalog number Type Length SL Width Height Depth SLR (PV) SLG (BV) Trail length Costellae i

PV HAM (BV) Hinge Body 20 mm 30 mm 

Whole shells (articulated) or disarticulated pedicle valves 13665g........... r ** 24.2 a 21.4 37 – 32.7 a 11.5 7.5 28 – – 16 16 – 

13664g........... r ** 27.7 a 26.0 a 50 a – 35.5 a 18.9 a – 30 – – 13 ––13663a........... r ** 28.6 a 25.2 a 46 40 m 37.0 a 16.7 a – 31 – – 16 18 113671d........... r 29 m 23 m 53 m – 41 e – 16.5 a 30 23 – 15 13 – 13664h........... r ** 29.2 m – 50 m – 39.0 a – – 32 – – 15 16 113665j............ r ** 30 m 21 m 55 m – 37.8 a – 13.4 35 – – 15 16 113672d........... r 30 m 25 e 65 m 44.0 t 37.1 a – – 33 – – 17 14 113664a........... r ** 31 m 25 m 55 m – 34 m – 15.0 33 a – – 15 16 113664c........... r ** 31 m 20 m 52 m – 35.0 a – 14.2 32 – – – 12 113665c........... r ** 31 m 26 m 45 m 40.0 e 38.6 e – – 28 – – 19 15 113665a........... r ** 31 m 26 m 53 m – 39 e – 14.0 31 21 – 19 18 113662d........... r ** 31.4 m 22.5 m 58 m – 38.5 m 21 m 16.9 30 18 – – 13 – 13673d........... r 31.4 a 25.3 a 52 – 39.6 a 25 e – 32 – – 14 14 a – 13663c........... r ** 31.5 a – 60 44.4 a 38.5 a 20.5 a – 25 – – 16 14 113665h........... r ** 32 m 22 m 58 m – 39.2 a – 16.6 29 – – 18 18 113664d........... r ** 32 m 24 m 51 m – 33.8 a – 15.2 28 – – 14 15 – 13664b........... r ** 32 m – 53 m 42.4 t 42.0 t – – 32 – – 14 14 – 13662e........... r ** 32.3 a 26.8 a 52 39.3 a 37.8 a 20.7 a – 25 – – –––13665f ........... r ** 32.5 a 22.4 a 60 a 43.6 a 40.7 a 23.9 a – 34 – – 15 14 113665b........... r ** 33.4 a 23.8 a 55 40.0 t 56.0 t1 23.2 a – 33 – – 19 19 1

13673c *........ r 33.5 m 25.5 m 60 m 44.8 t 32.9 21.5 m 20.0 34 21 – –––13672e........... r 33.5 m 26 m 60 m – 38.2 a 19 m 15.3 30 22 – – – 113665d........... r ** 34.0 a 24.4 a 55 – 41.4 e – – 28 – – 15 14 113673e........... r 34.0 27.0 a 59 42.1 a 40.1 21.6 a – 33 a – – – – – 13672f ........... r 34 m 28 m 58 m – 35.0 a 19 m 13.2 29 22 – –––14353............. R 34 m 25 m 60 m 39.2 t 40.0 a 20 m 14.4 31 16 – 16 14 113663d........... r ** 34.2 a 26.2 a 58 – 37.9 a – – 29 – – 15 14 – 13663b........... r ** 34.4 a 25.5 a 69 – 41.0 a 22.9 a – 35 – – 18 15 113665e........... r ** 34.8 a 26.5 a 60 53.8 t 40.8 e – – 35 – – 15 14 113672c........... r 35 m 24 m 62 m – 38.5 20 m 15.0 38 23 – 18 14 114354............. R 35 m 25 m 60 m – 43.6 24 m 14.5 40 21 – 15 a 15 a 1
Table 4. Measurements (in millimeters) of specimens identified as Antiquatonia coloradoensis (Girty) and A. hermosana (Girty) by Ragan (1

Formation (upper Atokan), Jasper County, Mo., and referred herein to A. coloradoensis (Girty)—Continued.

1Specimen has flaring trail. 2Excluding cardinal process.

UM catalog 

number Type

Length

SL

Width

Height Depth SLR (PV) SLG (BV) Trail length

Costellae i

PV HAM (BV) Hinge Body 20 mm 30 mm 

Whole shells (articulated) or disarticulated pedicle valves

13663f ........... r ** 35.0 a 25.6 a 60 – 39.3 a 22.2 a – 32 – – 18 17 13663e........... r ** 35.2 a 24.9 a 70 48.4 t 38.5 t 22.7 a – 32 – – 16 15 13662a........... r ** 35.5 26.5 a 67 54.6 t 39.2 a 26.2 a – 28 a 17 a – –––13672b........... r 35.6 a – 60 e 37.5 e 35.5 22.0 a – 28 – – –––13673a........... r 35.9 27.7 a 66 43 e 39.0 24.6 a – 32 – – 13 12 13671e........... r 36.0 a 26.6 a 70 – 41.5 a 26.7 a – 34 – – 18 16 13662b........... r ** 36.0 a 27.1 a 66 – 40 e 21.7 e – 29 19 – 17 15 a – 13664f ........... r ** 36.0 m 29.5 m 63 m – 39.9 a – – 34 – – 15 – – 13671b........... r 36.1 a 27 e 70 49.0 t 43.8 29 e – 33 – – 15 14 – 13662c........... r ** 36.2 28.8 a 68 – 39.2 a 26.3 a – 31 – – 15 15 13664e........... r ** 36.7 28.6 a 65 46.2 a 46.2 a1 23.6 a – 32 – – 17 17 

13665i ........... r ** 36.8 a – 62 57.2 t 39.5 a – – 27 – – 15 17 13662g........... r ** 37 m 24.4 m 55 m – 35.0 a 23 m 13.7 32 – – 17 – – 13657a........... r ** 38.9 28.3 a 73 48.7 46.5 25.1 a – 35 – – 16 a 16 – 13671c........... r 39.6 m 25.5 m 68 m 39.0 t 34.5 a 22.0 m 17.5 23 20 – 17 14 13657c........... r ** 39.7 a – 60 a 47.7 e 45.0 e – – 32 a – – 15 14 13672a........... r 39.9 28.5 a 78 48.8 a 43.1 28.1 a – 34 – – 17 16 13662f ........... r ** 40.1 a 30.5 a 70 – 41.4 a 23.0 a – 32 a – – – – – 13673b........... r 40.7 a 28.6 a 72 – 42.2 a 24.8 a – 35 – – –––13657b........... r ** 42.5 a 33.5 a 80 51.5 a 48.8 a 28.3 a – 25 – – 15 16 13657d........... r ** 44.3 a 34.4 a 78 a 47.7 a 51.8 a1 26.6 a – 29 – – 15 – – 

13671a........... r 47.0 27.5 a 78 52.0 t 43.0 a 28.0 a – 31 – – 17 16 13657e........... r ** 48 e – 74 a 48.2 43.4 a – – 28 – – 14 16 

Brachial valves (disarticulated)

13663h........... r ** – 222.8 228 37.0 t 33.2 – – – 220 8 – – – 

13672g........... r – 225.3 237 38.3 a 36.3 a – – – 222 15 – – – 13671f ........... r – 226.7 a 240 44.3 a 38.1 a – – – 220 20 – – – 13673f ........... r – 230.3 251 48.6 t 47.2 – – – 221 30 – – – 13657g........... r ** – 235 e 250 e 53.4 t 67.8 t1 ––– 221 29 e – – – 
24 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

The number of costellae in 10 mm measured at 20 mm

SL for the composite set from the Burgner ranges between

13 and 19 (M=15 and =16.3; see fig. 5). This range therefore overlaps heavily with that of the types of A. coloradoensis (fig. 5) from the Belden Shale, the range of which is

from 14 to 19 (M=15 and 16 and =16.5). No statistically

significant difference (99.5 percent confidence limits) exists

for this feature between the Burgner sample and the types of

A. coloradoensis from Colorado.

I also examined the University of Missouri’s collection

of specimens of Antiquatonia, including Hoare’s figured

specimens, from the Burgner Formation from Hoare’s

(1961, p. 18) UM loc. 37, mentioned above, from central

Missouri. According to T.L. Thompson, Jr. (oral commun.,

November 15, 1991), UM loc. 37 near Eldon is about the

same age (late Atokan) as the type Burgner in the southwestern part of the State. Hoare’s Eldon collection consists

of 11 relatively complete specimens or valves plus fragments. Hoare (1961, p. 56–58) typologically assigned specimens from this collection from the Burgner Formation to

three Antiquatonia species: A. hermosana, A. coloradoensis,

and A. portlockiana. He illustrated specimens as the former

two species (UM 13192–13194 and UM 13195–13197,

respectively), but his figured specimens assigned to A. portlockiana (Hoare, 1961, pl. 5, figs. 1–7) were from younger

Pennsylvanian (middle and upper Desmoinesian) strata. The

range of variation in the Burgner collection assigned by

Hoare to these three discrete species is completely encompassed by the large collection of primary and secondary

types of A. coloradoensis from the Mosquito Range. Size

alone does not constitute a valid criterion on which to discriminate among these species. 

COMPARISON WITH ANTIQUATONIA PORTLOCKIANA

Girty (1935, p. 9) suggested, with strong caveats, that

A. hermosana might be a junior synonym of A. portlockiana

(Norwood and Pratten). Sutherland and Harlow (1973,

p. 32) concluded, however, that the two are discrete species

and that A. portlockiana succeeds A. hermosana stratigraphically in the upper middle and upper Desmoinesian of northern New Mexico. I agree with their conclusion based on the

New Mexico material. However, to fully resolve this question, large collections of topotypes of A. portlockiana need

to be examined to determine more fully its characteristics

and range of variation. Dunbar and Condra (1932, p. 216)

noted that Norwood and Pratten (1855) had only two specimens when they described A. portlockiana, one from the St.

Louis, Mo., area and the other from Illinois. The disposition

of these two cotypes is not known. Dunbar and Condra also

noted, however, that they had available large collections of

topotypes from Charbonniere Bluffs, St. Louis County, Mo.,

from strata that they reported as “lower Fort Scott limestone” (middle Desmoinesian). They illustrated one of these

x

x

topotypes (Dunbar and Condra, 1932, pl. 33, fig

the absence of material with which to redescrib

lockiana, suffice it to say that it would appear th

cies differs from A. coloradoensis in being sligh

having a proportionally less inflated umbo, pro

smaller ears, perhaps a more strongly incurved 

coarser radial ornamentation (see fig. 5).

GENERAL DISCUSSION 

The confusion that exists in the literature 

nature and stratigraphic range (geologic age) of 

nia coloradoensis (Girty) (see Carter and Ca

p. 218, 219) stems mainly from the following six 

1. When Girty (1910, p. 216) formally pr

varietal name Productus inflatus var. coloradoen

in a paper that was not illustrated and that was 

primarily with naming the components of the i

macrofauna of the Fayetteville Shale (Chester

Mississippian) of the Ozark Mountains. In the 1

although it was clear that the primary types were 

Middle Pennsylvanian strata of Colorado (see 

section), he also provisionally referred to the ne

variety, Productus inflatus var. coloradoensis, a la

ficially similar, semireticulate productid from the 

of the Fayetteville Shale. This material, also as

the genus Antiquatonia (as currently understoo

from A. coloradoensis in a number of important 

cifically, the Fayetteville material is larger and p

ally broader, has a consistently deeper sulcus on 

valve, and possesses both fewer and smaller body 

the pedicle valve. It is not conspecific with A. col

and represents an unnamed, undescribed specie

have been ancestral to A. morrowensis (Mather). 

2. Girty, in later publications, applied (agai

ervation) the name Productus inflatus var. colora

other Mississippian collections as well. The first t

publications (Girty, 1911, p. 42; 1915b, p. 46) w

faunas of the Moorefield Formation (Meramecian 

terian) and Batesville Sandstone (Chesterian) of t

To further complicate matters, in a subsequently 

paper (1915c) primarily on the fauna from “th

Boone chert,” from near Batesville, Ark., Girty (19

referred material from the chert to the same variety

var. coloradoensis. Girty’s fauna was mainly from 

age beds of the Reeds Spring Chert Member of 

Formation (Kinderhookian through lower Me

However, the fragmentary productids in question p

were collected (USGS colln. GR1237B–PC) from 

Creek Member (Meramecian and lower Chester

Moorefield Formation and represent an unnamed 

semireticulate productid that is likely not an Antiq

all. Girty (1915a, p. 65) later recognized that none 

sissippian material was referable to P. coloradoe
SYSTEMATIC PALEONTOLOGY 

fairness to Girty, who published over 40 major papers

between 1900 and 1920, the order of publication of the papers

was not the same as the order in which they were written. For

example, the manuscript on the faunas from the Wewoka Formation (Girty, 1915a) was written after those on the Mississippian faunas of northern Arkansas (1915b, d).

Snider (1915, p. 80), in his study of the Chesterian faunas of northeastern Oklahoma, followed Girty’s example

and used the name Productus coloradoensis for Upper Mississippian forms, although he did not illustrate the species

from the “Mayes limestone” (probably Moorefield Formation) thus referred. These Moorefield and Batesville forms,

from strata slightly older than the Fayetteville Shale, probably belong to the unnamed Antiquatonia present in the Fayetteville. Many other authors (Croneis, 1930; Sutton, 1938;

Nelson, 1961) also mistakenly assumed that P. coloradoensis is based on material from the basal part of the Fayetteville and used the name widely and indiscriminately for

Mississippian antiquatonids. Carter and Carter (1970,

p. 218–219) recognized that at least two taxa were involved. 

3. Although Girty initially proposed the name for the

Pennsylvanian specimens in the 1910 paper for the Colorado material, he later (1915a, p. 64, 65—the paper on the

fauna from the Wewoka Formation) again presented the

same name Productus inflatus var. coloradoensis, based on

the same material, and treated it as though his earlier proposal (Girty, 1910) had not been made. I emphasize that

Girty (1915a)—the Wewoka paper—is not the publication

in which the name coloradoensis was first applied to the

Pennsylvanian species, although it has been cited thusly by

several workers. To quote Girty (1915a, p. 65): 

The best and most characteristic specimens referred to this variety [from

the Wewoka Formation] resemble the form from Colorado identified by us

as P. inflatus rather than the one described as P. semireticulatus var. hermosanus. Having studied what are probably characteristic specimens of P.

inflatus, we have reached the conclusion that the shell from Colorado is a

distinct form and propose for it the varietal name coloradoensis. 

Contrary to Girty, I conclude that the material from the

Wewoka Formation (upper Desmoinesian, Middle Pennsylvanian) that Girty (1915a, p. 64, 65, pl. 8, figs. 1, 2) referred

to P. coloradoensis does not belong to this species at all. I

examined both imperfect specimens (USNM 120914,

192095) that he figured from the lower part of the Wewoka

Formation and sizable collections from throughout this formation. The Wewoka specimens are large and generally

more coarsely costellate than A. coloradoensis and in many

other respects are similar to specimens normally assigned to

A. hermosana (Girty), which, however, are generally in

lower and middle Desmoinesian strata.

4. The primary type specimens of Productus inflatus

var. coloradoensis were reillustrated (as engraved figures)

several times in publications for Carboniferous strata for

other geographic areas (see table 5). Such reillustration was

a widespread practice at the time, following the concept of

“index fossils” and the convention of illustrating “typical

fossils” for a given age, strata, and region. However

practice created confusion in that the illustrated specim

are indeed Antiquatonia coloradoensis (and therefore 

be included in the synonymy of the species) but the ma

ostensibly referred to that species in a cited report m

may not be that taxon. Three examples follow. 

Boutwell (1912) reported Productus inflatus from 

“Weber quartzite” (part of the Morgan Formation of cu

usage) of the Park City mining district of the Uinta M

tains, northern Utah. He illustrated several primary 

specimens of Girty’s Productus inflatus var. colorado

from the Fairplay-Leadville, Colo., area. Second, 

(1920, pl. 55, figs. 4, 5) likewise reillustrated two of h

mary types for Butler and others’ “The Ore Depos

Utah” (1920) as one of the “index fossils” for the Carb

erous of that region, but neither Girty (1920) nor Butle

others (1920) listed the species from any specific 

graphic or stratigraphic horizon in those papers. Third, 

(1927) refigured one of his primary types as P. colora

sis in Mansfield’s (1927) monograph on the geolo

southeastern Idaho. In this monograph, Mansfield (

p. 73) cited Girty’s identification of this taxon in the “

formation.” However, it is extremely doubtful that the 

productid thus reported in the Utah or Idaho sequen

Girty’s species at all. 

5. The Burgner and Riverton Formations of south

ern and central Missouri consist of a series of dark shal

carbonate units deposited in sinkholes developed o

karsted Mississippian limestone surfaces and in othe

regions on an irregular, undulating erosional surface o

flanks of the Ozark dome during the later Morrowa

early Atokan (see Seawright and Palmer, 1957; Hoare, 

p. 14; Thompson, 1979, p. N13–N16). As stated above 

section, “Comparison with Antiquatonia hermosana,” 

Douglass concluded that the fusulinids from these sin

deposits indicate a late Atokan age for these strata. How

Girty (1935, p. 8) referred the upper Atokan Burgner str

the Palmer locality (discussed above) to the “Cheroke

mation,” a stratigraphic term now reserved for the basa

moinesian (Middle Pennsylvanian) coal-bearing seq

overlying the Burgner Formation. Girty’s reference o

Atokan material to the Cherokee undoubtedly has led t

ther uncertainty about the upper extent of the range of 

quatonia coloradoensis. As indicated above, I conclude

a variable form more reasonably identified as A. colora

sis is present in the Burgner collections, not both A. co

doensis and A. hermosana. 

6. Antiquatonia coloradoensis is present in the 

part of the Minturn Formation in the McCoy area of c

Colorado. This taxon was reported by Stevens (1962) 

present in the USGS collections from this area. The Mi

strata in which this productid was reported by St

(1962, p. 618–620, unit 3) also contain Fusulinella 

(Thompson), indicating equivalence with the upper p

the type “Derryan” Series (Atokan) in New Mexico an
26 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

upper part of the Atokan Series of the Midcontinent. The

three specimens that Stevens (1962, pl. 95, figs. 2, 3, and 5)

illustrated as A. coloradoensis are captioned (1962, p. 628)

as from Stevens’ unit 9 of the Minturn Formation, reported

to contain early Desmoinesian fusulinids and other brachiopods characteristic of that part of the Middle Pennsylvanian.

He listed A. coloradoensis in his discussion of unit 3 (see

Stevens, 1962, p. 623) but cited only A. hermosana in his

discussion of the fauna of unit 9. Furthermore, the two large

specimens that he illustrated as A. hermosana (1962, pl. 95,

fig. 6; pl. 96, fig. 4) are also captioned as from unit 9. Without having looked at the Stevens collections from units 3

and 9, and assuming that the plate caption is correct, I suggest that the specimens that Stevens figured as A. coloradoensis in this paper are the rarer, smaller, more finely costellate variants of A. hermosana and should not be identified

as A. coloradoensis. This question cannot be resolved by the

USGS collections from the Minturn area, because most of

these do not have corroborative biostratigraphic control

from other faunal groups.

OCCURRENCES AND RANGES

Antiquatonia coloradoensis is reported from the southern Rocky Mountains, the Eastern Great Basin, the southern

and central Midcontinent area, and both the eastern and

southern parts of the Appalachian Basin. This productid is a

component of the marine faunas interpreted to represent

open-bay, shelf-lagoon, and shelf-margin facies. 

SOUTHERN ROCKY MOUNTAINS

Antiquatonia coloradoensis occurs in the northern part

of the southern Rocky Mountains in the following strata and

areas: (1) in the upper Morrowan strata of the lower part of

the Belden Shale in the Fairplay, Colo., area (typ

loc. 10 of fig. 7); (2) in the Belden Shale (upper 

and Atokan equivalents) of the Eagle Basin ne

Butte, Colo. (loc. 11 of fig. 7); (3) in the midd

Atokan parts of the Minturn Formation in centra

(loc. 5 of fig. 7); (4) in strata probably assign

Pinkerton Trail Formation (Atokan equivalents) i

part of the Hermosa Group in the San Juan regio

western Colorado (loc. 12 of fig. 7); and (5) i

upper Morrowan and lower and middle Atokan 

of the La Pasada Formation in northern New Me

erland and Harlow, 1973, p. 51) (loc. 13 of fig. 7)

In southern New Mexico, Antiquatonia col

is present in the upper Morrowan part of the type 

Series (Sutherland and Manger, 1984; Sutherl

p. 188) (loc. 14 of fig. 7) along with 11 species o

late Morrowan brachiopods. Sutherland assig

strata to his Zia novamexicana Brachiopod Subz

Linoproductus nodosus Zone. Gehrig (1958) als

Dictyoclostus inflatus var. coloradoensis from 

bearing Desmoinesian strata of the Armendaris 

southern New Mexico (Mud Springs Mountains

fig. 7). However, Gehrig’s specimens were ex

P.K. Sutherland (oral commun., Nov. 1, 1991), wh

that his Desmoinesian specimens identified as 

var. coloradoensis are, in fact, typical A. hermosa

and his Atokan specimens of D. morrowensis (M

A. coloradoensis. 

EASTERN GREAT BASIN

The material reported by Boutwell (1912

“Weber quartzite” (Weber? Formation) in the Par

trict in northeastern Utah (loc. 15 of fig. 7) as 

inflatus?, and later included in synonymy of P. c

sis by Girty, needs to be examined for possible 

Table 5. Index to original (engraved) figures and replication of same in publications for lectotype and two paralectotypes of A

coloradoensis (Girty) from the lower part of the Belden Shale (upper Morrowan equivalents), from U.S. Geological Su

GR2281–PC, from near Fairplay, Park County, Colo. 

Publication Figured or identified as: USNM 35345 USNM 121482 USNM 12

Girty (1903) .......... Productus inflatus McCh. Pl. 3, figs. 1–1b, figured 

specimen.

Pl. 3, figs. 2–2a, figured 

specimen.

Pl. 3, fig. 3, figur

specimen.

Girty (1910) .......... Productus inflatus var. 

coloradoensis n. var.

Not illustrated, designated 

cotype.

Not illustrated, designated 

cotype.

Not illustrated, d

cotype.

Boutwell (1912) .... Productus inflatus? McCh. Pl. 6, figs. 3–3a, figured 

specimen.

– Pl. 6, fig. 4, figur

specimen.

Girty (1920) .......... Productus inflatus var. 

coloradoensis Girty.

Pl. 55, fig. 3, cotype – Pl. 55, fig. 4, coty

Girty (1927) .......... Productus coloradoensis 

Girty.

Pl. 27, fig. 17, cotype – –

Gordon (1975)....... Antiquatonia coloradoensis 

(Girty).

Not illustrated, designated 

lectotype.

Not illustrated, designated 

paralectotype.

Not illustrated, d

paralectotype.

This paper.............. Antiquatonia coloradoensis 

(Girty).

Pl. 1, figs. 1–4, lectotype Pl. 1, figs. 5–8, 

paralectotype.

Pl. 1, figs. 9–13, 

paralectotype.
SYSTEMATIC PALEONTOLOGY 

19

16

21

15

20 5

11

12

10

13

14

6

8

7

2 1

3

4 9

18

17

1

2

3

4

6

7

8

Northwestern Arkansas

Northeastern Oklahoma

Jasper County, Missouri

Miller County, Missouri

Ardmore Basin, southern

Oklahoma

Northeastern flanks of the

Arbuckle Mountains,

southern Oklahoma

Llano area of Texas

9

17

18

Floyd County, eastern

Kentucky

Southern West Virginia

in general vicinity of

Boone County

Warrior coal field,

southern Appalachian

5

10

11

12

13

14

Central Colorado

Fairplay, Colorado

Crested Butte, Colorado

San Juan region,

southwestern Colorado

Northern New Mexico

Mud Springs Mountians,

southern New Mexico

15

16

19

20

21

Park City district, northeastern

Utah

Northern Nye County, Nevada

Carlin, Nevada

Wasatch Mountains, near

Provo, Utah

Arrow Canyon Range, Clark

County, Nevada

LOCALITIES

EASTERN GREAT BASIN MIDCONTINENT SOUTHERN ROCKY MTNS. APPALACHIAN BASIN

0 100 200 300 400 500 MILES

0 100 200 300 400 500 KILOMETERS

to Antiquatonia coloradoensis. Just south of this area, I have

identified A. coloradoensis from several collections from

the Wasatch Mountains near Provo, Utah (loc. 20 of fig. 7),

associated with Atokan Profusulinella-bearing rocks of the

Bear Canyon Member of the Oquirrh Formation. 

This species is known from three areas in Nevada. I

have examined a small collection from the Ely Limestone

(Upper Mississippian to Lower Permian) in northern Nye

County (loc. 16 of fig. 7). These specimens are tentatively

identified as A. coloradoensis and occur with a moderately

diverse brachiopod fauna strongly suggesting an Atoka

for the strata. A second, recently obtained collection 

the Bird Spring Formation (Pennsylvanian and Lowe

mian) in the Arrow Canyon Range, Clark County (loc. 

fig. 7), also contains A. coloradoensis along with a m

ately diverse late Morrowan or early Atokan brach

fauna. These brachiopods occur above the stratigraph

lowest Eoschubertella but below the lowest Profusul

(Langenhein, Webster, and Weibel, 1984; Webster and 

genheim, 1984). Smith and Ketner (1975, p. 53) repor

Figure 7. Localities of verified or probable occurrences of Antiquatonia coloradoensis (Girty) in the United States. Numbers 1–21 

spond to areas identified in text.
28 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

coloradoensis from the upper Morrowan part of the Moleen

Formation (Lower and Middle Pennsylvanian) near Carlin

(loc. 19 of fig. 7). 

MIDCONTINENT

Antiquatonia coloradoensis first appears in the Midcontinent in upper Morrowan strata and ranges apparently

through the Atokan Series. The oldest confirmed appearance of Antiquatonia coloradoensis is in the Kessler Limestone Member of the Bloyd Shale in the upper part of the

“type” Morrowan Series in northwestern Arkansas (loc. 1

of fig. 7) and in the physically correlative Greenleaf Lake

Limestone Member of the McCully Formation in northeastern Oklahoma (loc 2. of fig. 7). Its youngest confirmed

occurrence is in upper Atokan strata of the Burgner Formation of Jasper and Miller Counties, Mo. (locs. 3 and 4,

respectively, of fig. 7). 

Other Midcontinent occurrences of Antiquatonia coloradoensis follow. This species is rare in the Trace Creek

Member of the Atoka Formation in northwestern Arkansas

(loc. 1 of fig. 7). Sutherland and Grayson (1992, p. 84,

88–89) reported A. coloradoensis with other “typical” late

Morrowan brachiopods in the Ardmore Basin of southern

Oklahoma (loc. 6 of fig. 7) in the Golf Course Formation

(upper Morrowan)—specifically in its Jolliff Member, in the

lower parts of their unnamed units 1 and 2, and in the Otterville Member. They (Sutherland and Grayson, 1992, pl. 3,

figs. 10–12) figured three well-preserved specimens from

the lower part of the Bostwick Conglomerate Member of the

overlying Lake Murray Formation. These strata contain

Fusulinella dakotaensis, the lower of two zones of

Fusulinella in the basin, and are correlated with the middle

part of the Atokan Series. This species also occurs in the

upper Morrowan part of the Wapanucka Limestone of

southern Oklahoma on the northeastern flanks of the

Arbuckle Mountains (loc. 7 of fig. 7) (P.K. Sutherland, oral

commun., Nov. 1, 1991). The specimen that Morgan (1924,

pl. 44, fig. 6) illustrated as Productus morrowensis Mather

from the Wapanucka from this area belongs to Girty’s species. Antiquatonia coloradoensis is in the lower Atokan part

of the Marble Falls Limestone (Lower and Middle Pennsylvanian) of the Llano area of Texas, where it occurs abundantly with species of Profusulinella (P.K. Sutherland, oral

commun., Nov. 1, 1991) (loc. 8 of fig. 7). It is most likely

that the specimens figured by Plummer and Moore (1921, p.

79, pl. 13, fig. 24) and Plummer (1950, pl. 12, fig. 9) are

from these lower Atokan strata. Antiquatonia coloradoensis

does not occur in the Fusulinella-bearing beds in the upper

part of the Marble Falls Limestone (P.K. Sutherland, oral

commun., Nov. 1, 1991). 

APPALACHIAN BASIN

Antiquatonia coloradoensis has been iden

three general areas in the Appalachian Basin, from 

vanian strata that range in age from late Morrow

Atokan. The Morrowan and lower Atokan collect

ally also contain Linoproductus nodosus (Newb

and other faunal elements characteristic of the 

Brachiopod Zone. The stratigraphically highest 

(for example, those of the Winifrede Shale Mem

Kanawha Formation of West Virginia and the 

Magoffin Member of the Breathitt Formation of 

contain A. coloradoensis, but the other brachiopo

teristic of the L. nodosus Zone are not present.

The first of these areas and the oldest kno

rence of Antiquatonia coloradoensis in the A

Basin is in southern West Virginia (loc. 17 of fig. 

have collected several specimens of this species 

the Eagle limestone of I.C. White (1891) and 

shale of Hennen and Reger (1914) in the lower m

of the Kanawha Formation. The strata of this m

are now assigned to the Betsie Shale Memb

Kanawha Formation (Rice and others, 1987) and 

Morrowan correlatives (Henry and Gordon, 

101–102). Antiquatonia coloradoensis also occu

in this general vicinity in an overlying Kanawha m

in Boone County, W. Va., in what Price (1916) 

Seth limestone of Krebs (in Krebs and Teets, 1

marine band is the one that occurs just above 

Grove coal bed in Kanawha and Fayette Counti

and has been correlated more recently with the 

Creek Limestone of I.C. White (1885) by Blake, 

Rice (1994, fig. 2). I have collected both L. nodo

coloradoensis from this marine unit in all thre

Price (1916, p. 699) referred his Boone County 

from the Seth and from the stratigraphica

Winifrede Shale Member to Productus sem

(Martin, 1809). Both Price’s Seth and Winifrede 

are definitely Girty’s species. 

The second area where Antiquatonia colora

found is in eastern Kentucky (loc. 9 of fig. 7) in t

graphic units in the Breathitt Formation. The low

rence is in the upper Morrowan Elkins Fork Sha

in Floyd County. This species also was reported b

others (1994, table 2) from the Kendrick Shal

from strata that Rice (1980) identified as “open-b

in Floyd and Pike Counties. On the basis of the 

goniatite faunas described by Furnish and Kna

the Kendrick beds correlate with the Trace C

Member, now considered the basal member of 

Formation in northwestern Arkansas (see Rice 

1994, p. 91–93, for detailed discussion). All of th

described also bear L. nodosus and other compon

L. nodosus Zone. The stratigraphically higher 

Member contains a medium to large sem
REFERENCES CITED 

productid and a species of Linoproductus that is not Newberry’s (1861) L. nodosus. Douglass (1987) placed the

Magoffin Member of the Breathitt Formation in eastern

Kentucky in the lower part of the Atokan Series, but this

member is higher stratigraphically than the lowest strata

correlated with the Atokan of northwestern Arkansas (see

discussion of the Trace Creek Shale Member above). Morse

(1931, p. 311–312, pl. 49, fig. 5, probably not figs. 6–8)

identified the semireticulate productid as Productus semireticulatus (Martin, 1809) and probably also lumped specimens into Martin’s taxon that I would identify as Sandia sp.

Specimens of Sandia were identified as Productus [Dictyoclostus] semireticulatus by Chesnut (1981). As I mentioned

above, in the section “Inclusion of Antiquatonia portlockiana var. quadratia in A. coloradoensis,” Chesnut (1981,

p. 25, pl. 9, figs. 4–6) referred specimens from the Magoffin

to Antiquatonia cf. A. portlockiana quadratia Sturgeon and

Hoare (1968). I would place this Atokan material from the

Magoffin in A. coloradoenis. Probably, some of the middle

and upper Atokan specimens from southeastern Ohio

assigned to A. portlockiana var. quadratia by Sturgeon and

Hoare (1968) are A. coloradoensis as well (see discussion in

the section just cited).

The third area of occurrences of Antiquatonia coloradoensis is in the Warrior Coal Field of the southern Appalachians (loc. 18 of fig. 7); I have identified material from several cores from upper Morrowan strata of the Pottsville

Formation from some of the collections made by Culbertson

(1964). These cores also contain Linoproductus nodosus.

Antiquatonia coloradoensis was identified by Butts (1926,

p. 213) as Productus semireticulatus (Martin, 1809) from

the Pottsville of the Warrior Coal Field. The specimen that

he (Butts, 1926, pl. 67, fig. 12) figured as Martin’s species,

however, is from younger Pennsylvanian rocks of Illinois

and is not assignable to Girty’s species.

SUMMARY OF OCCURRENCES AND RANGES

It appears highly likely that Antiquatonia coloradoensis

is restricted to the upper Morrowan and Atokan (lower Middle and middle Middle Pennsylvanian) strata in the Midcontinent, the southern Rocky Mountains, the Eastern Great

Basin, and the Appalachian Basin. Its range in the Appalachian Basin is restricted to the marine bands of late Morrowan and Atokan age. Its highest known occurrence in the

Appalachians is in the Magoffin Member of the Breathitt

Formation and the Winifrede Shale Member of the Kanawha

Formation in southern and central West Virginia. Antiquatonia coloradoensis does not appear to range upward into the

Desmonesian Series and equivalent strata, although smaller,

more finely costellate variants (morphotypes) form minor

percentages in large collections of A. hermosana. Collections in which both A. coloradoensis and A. hermosana have

been identified from other Atokan strata (for example,

collections described by Brill, 1942, p. 1388; 1944, p

and certainly from Desmoinesian rocks need to be re-e

ined in light of interspecific variation in populations of 

quatonia. The Desmoinesian specimens of Antiqua

portlockiana var. quadratia Sturgeon and Hoare from s

eastern Ohio need to be carefully re-examined, beca

believe that it is not likely that they are A. coloradoens

The Mississippian collections of Antiquatonia ar

assignable to A. coloradoensis. 

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Duméril, A.M.C., 1806, Zoologie analytique ou méthode naturelle

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Emmons, S.F., 1883, Atlas to accompany a Monograph [No. 12] on

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Geological Survey Atlas, 34 maps [sheets], 1 table. [First geologic map of area is sheet 7.]

———1886, Geology and mining industry of Leadville, Colorado,

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24 p., 6 pls., 1 fig.

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546 p., 10 pls. 

———1904, Note on the Carboniferous fossils, in Ra

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———1910, New genera and species of Carbonife

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———1911, The fauna of the Moorefield Shale of Ar

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———1915a, Fauna of the Wewoka Formation of Okl

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———1915b, The fauna of the Batesville Sandstone 

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———1915c, Fauna of the so-called Boone Chert nea

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———1915d, Faunas of the Boone Limestone at St. 

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———1920, Carboniferous and Triassic faunas, in B

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———1927, Descriptions of new species of Carbon

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———1935, An unrecorded structure in certain sem

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Gordon, Mackenzie, Jr., 1975, Brachiopoda of the Am

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Geological Survey Professional Paper 848–D, 86 

3 figs.

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Henry, T.W., 1973, Brachiopod biostratigraphy and f

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Arkansas and northeastern Oklahoma: Norman, 

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pocket). 

Henry, T.W., and Gordon, Mackenzie, Jr., 1979, La

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Henry, T.W., and Sutherland, P.K., 1977, Brachiopod 

phy of Morrowan Series (Pennsylvanian) in n

Arkansas and northeastern Oklahoma: Oklahoma 

Survey Guidebook 18, p. 107–115, 2 pls., 3 figs. 
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32 BRACHIOPOD ANTIQUATONIA COLORADOENSIS (GIRTY) FROM UPPER MORROWAN AND ATOKAN

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letin 65, 212 p.
PLATES 1–5
PLATE 1

Primary Type Specimens of Antiquatonia coloradoensis (Girty) from 

Lower Part of Belden Shale, Park County, Colo. (USGS Colln. GR2281–PC)

[All figures natural size]

Figures 1–4. Lectotype, USNM 35345; ventral, anterior, posterior, and lateral views of typical 

mature specimen; ears slightly broken; anterior portion of trail decorticated. 

Specimen was shown in Girty’s (1903) plate 3, figures 1–1b, as Productus inflatus

McChesney. 

5–8. Paralectotype, USNM 121482; ventral, anterior, posterior, and lateral views of large 

specimen; ears broken, trail slightly broken; anterior view showing trail of brachialvalve interior; note trail partially broken, slightly flaring. Specimen was shown in 

Girty’s (1903) plate 3, figures 2, 2a, as Productus inflatus McChesney.

9–13. Paralectotype, USNM 121483; ventral, anterior, posterior, lateral, and dorsal views 

of coarsely costellate specimen; trail broken, anterior portion of pedicle valve 

broken away, revealing part of trail of brachial-valve interior; note row of halteroid 

spines on arcuate ridge and row of smaller hinge spines on posterior view (fig. 11); 

flange on pedicle-valve interior (marked by “s”) on dorsal view (fig. 13), 

continuing posteriorly and separating auricular chamber (ears broken away) from 

visceral chamber in dorsal view in area where brachial valve is missing. Specimen 

was shown in Girty’s (1903) plate 3, figure 3, as Productus inflatus McChesney. 

14–19. Paralectotype, USNM 208827; ventral, anterior, dorsal, posterior, lateral, and 

canted posterior views of typical specimen; note repaired injury to pedicle-valve 

beak area, shown particularly on posterior view (fig. 17); anterior part of trail 

decorticated; note weak flange (marked by “s”) at edge of valve on canted posterior 

view (fig. 19) where brachial valve is broken away. 
U.S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 1588 

ANTIQUATONIA COLORADOENSIS (GIRTY)

1

2

3

4

14

15

16

17

19

18

13

11

10

6

5 9

7

8

s

s
PLATE 2 

Primary Type Specimens of Antiquatonia coloradoensis (Girty) from 

Lower Part of Belden Shale, Park County, Colo. (USGS Colln. GR2281–PC), and 

Hypotype of A. coloradoensis from Burgner Formation, Jasper County, Mo. 

(USGS Colln. BL25128–PC)

[Figures natural size, unless otherwise indicated on plate]

Figures 1–4. Paralectotype, USNM 208829; ventral, anterior, posterior, and dorsal views of 

broken specimen; note hinge spines and row of large spines on arcuate ridge on 

posterior view (fig. 3). 

5–14. Paralectotype, USNM 208828; nearly complete specimen with separable valves. 

Note lack of sulcus on pedicle valve (figs. 5, 6).

5–7. Ventral, anterior, and posterior views of large specimen with pedicle-valve 

trail partly broken off, revealing portion of interior of brachial-valve trail; 

note weak groove (“g”) on figure 6, into which flange (“s”) of figures 8, 9, 

and 10 fits when valve is closed. 

8, 9. Views (×1 and ×2, respectively) of anterior portion of pedicle-valve 

interior showing flange (“s”), taken from portion of valve that was broken 

away just above arrow marked by “g” on figure 6.

10. Dorsal view of shell, showing flange (“s”) around pedicle-valve interior 

where brachial valve is broken away; ear missing on specimen where ridge 

separates auricular area from visceral chamber.

11–13. Anterior, lateral, and ventral views of decorticated brachial-valve interior 

with pedicle valve removed; note groove (“g”) in figure 11.

14. Lateral view of complete shell; pedicle-valve trail slightly broken.

15. Paralectotype, USNM 208830; ventral view of internal mold of pedicle-valve 

interior.

16–21. Hypotype, USNM 208805; complete brachial-valve interior; note long trail, ridge 

attached to cardinal buttresses and separating auricular chambers from visceral 

area; ears slightly more extended than typical.

16–18. Anterior, ventral, and lateral views; compare lateral view of brachial-valve 

interior with lateral view of specimen shown in plate 2, figure 14, to get 

sense of size of body cavity. 

19, 20. Posterior view (×4.5 and ×1, respectively) of cardinal area, showing 

cardinal process.

21. Ventral view (×4.5) of cardinal area, showing cardinal process, buttress 

plates, and median septum.
U.S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 1588 

ANTIQUATONIA COLORADOENSIS (GIRTY)

1

2

3

4

14

15

16

17

19

20

21

18

13

1

11

1

6

9

5

7

8

s

s

g

s

×4a

×4a

×2
 PLATE 3

Primary Type Specimens of Antiquatonia coloradoensis (Girty) and

Primary Type Specimens of A. portlockiana var. quadratia Sturgeon and Hoare,

Herein Referred to A. coloradoensis

[All figures natural size]

Figures 1–15. Paralectotypes of Antiquatonia coloradoensis (Girty) from USGS colln. 

GR2281–PC, lower part of Belden Shale, Park County, Colo. 

 1–4. USNM 8258b; ventral, anterior, posterior, and lateral views of mature 

specimen having trail partly broken and ears missing. 

5–7. USNM 11660a; canted ventral, anterior, and lateral views of internal mold 

of mature specimen; beak area crushed; note flaring margin of trail and 

very subdued flange, marked by “x.” 

8–11. USNM 8258a; ventral, anterior, lateral, and posterior views of large, broad 

specimen; note ears missing and anterior part of pedicle valve broken off 

showing part of interior of brachial valve and trail; note groove (“g”) on 

brachial-valve interior into which flange (not preserved) of pedicle-valve 

interior fit; in lateral view (fig. 10), note row of spines sitting on external 

ridge (“r”) of pedicle valve and aligning with groove (“g”) in brachialvalve interior. 

12–15. USNM 8258c; ventral, posterior, lateral, and dorsal views of small, broken 

specimen.

16–25. Primary type specimens of Antiquatonia portlockiana var. quadratia Sturgeon and 

Hoare, from “Lower Mercer limestone and shale,” Pottsville Formation, Hocking 

County, Ohio. 

16–20. Holotype, OSU 26020, loc. Hfg–5; ventral, dorsal, posterior, anterior, and 

lateral views of specimen with crushed brachial valve; note subdued ridges 

anterior to body spines, relatively short trail characterized by repeated 

breakage and thickening of both valves anteriorly, and pronounced, spinebearing preauricular ridges on exterior of pedicle valve. 

21, 22. Paratype, OSU 26023, loc. Hfg–5; slightly canted dorsal and lateral views 

of broken, disarticulated pedicle valve; note strong ridge separating ear 

from visceral chamber on interior, subdued ridge bearing row of body 

spines on exterior of valve; valve may be from same specimen as OSU 

26025 (see pl. 4, fig. 9). 

23–25. Paratype, OSU 26026, loc. Hs–3; posterior, ventral, and anterior views of 

slightly crushed articulated specimen with broken trail; note that exterior 

of pedicle valve has ridges extending anteriorly from bases of body spines.
U.S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 1588 

ANTIQUATONIA COLORADOENSIS (GIRTY)

1

2

3

14

15

16

17

19

20 21

22

23

24 25

18

13

12

6

9

5 7

8

x

x

g

g s
PLATE 4

Primary Type Specimens of Antiquatonia portlockiana var. quadratia Sturgeon 

and Hoare, Herein Referred to A. coloradoensis (Girty) [above line], 

and Representative Species of Brachiopods from Type Locality of Antiquatonia 

coloradoensis (Girty) [below line]

[Figures natural size, unless otherwise indicated on plate]

Figures 1–12. Antiquatonia portlockiana var. quadratia Sturgeon and Hoare, loc. Hfg–5, from 

"Lower Mercer limestone and shale," Pottsville Formation, Hocking County, Ohio. 

 1–5. Paratype, OSU 26024; complete, disarticulated brachial valve showing 

well-preserved interior; enlarged dorsal and posterior views (×3.5) of 

cardinal process; posterior, ventral, and lateral views (×1) of interior; note 

atypical, club-shaped boss at anterior end of median septum, pronounced 

ridges separating visceral chamber from auricular chamber.

 6–8. Paratype, OSU 26022; anterior, ventral, and dorsal views (×1) of broken, 

disarticulated brachial valve; note flange around anterior of visceral area, 

thickening of anterior margin of shell at geniculation. 

 9. Paratype, OSU 26025; dorsal view (×1) of well-preserved brachial valve 

showing interior, ears slightly broken; note subdued flange around anterior 

margin; this valve may be from same specimen as OSU 26023 (see pl. 3, 

figs. 21, 22).

10–12. Paratype, OSU 47102; ventral, lateral, and posterior views of large, broken, 

disarticulated brachial valve; note row of hinge spines and row of large 

body spines on preauricular ridge. 

13–17. Linoproductus nodosus (Newberry), from lower part of Belden Shale, Park County, 

Colo. Note single row of large body spines down venter of pedicle valves.

13, 14. Referred specimen, USNM 476640, USGS colln. BL12096–PC; lateral 

and ventral views of cracked, incomplete, mature pedicle valve. 

 15. Referred specimens, USNM 476641, USGS colln. BL31394–PC; slab 

showing cluster of generally decorticated specimens mostly in growth 

position (pedicle valve down), viewed from bottom side of bed; note large 

body spine on venter on specimen marked by “x”; on specimen above “y,” 

pedicle valve and most of brachial-valve muscle area have been worn 

away, revealing row of “bumps” corresponding to dimples on exterior of 

valve and spine row on opposite valve. 

16, 17. Referred specimen USNM 476642, USGS colln. GR2281–PC; ventral and 

lateral views of broad, cracked, incomplete, mature pedicle valve. 

18–20. Anthracospirifer tanoensis Sutherland and Harlow, from lower part of Belden 

Shale, Park County, Colo. Note three costae in sulcus of pedicle valve. 

18, 19. Referred specimen USNM 476643, USGS colln. BL6852–PC; ventral and 

anterior views of uncrushed, articulated specimen with right ear missing.

 20. Referred specimen USNM 476644, USGS colln. BL12096–PC; ventral 

view of crushed and broken articulated specimen; ears missing.
U.S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 1588 

ANTIQUATONIA, LINOPRODUCTUS, AND ANTHRACOSPIRIFER

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PLATE 5

Characteristic Elements of Brachiopod Fauna from Type Locality of Antiquatonia 

coloradoensis (Girty) from Lower Part of Belden Shale, Park County, Colo.

[All specimens are referred specimens. Figures natural size, unless 

otherwise indicated on plate]

Figures 1, 2. Spirifer goreii Mather.

 1. USNM 476645, USGS colln. BL12096–PC; ventral view of crushed, 

complete young adult; note absence of fasciculation of costellae.

2. USNM 476646, USGS colln. BL31394–PC; ventral view of broken, 

slightly decorticated, disarticulated pedicle valve of a large specimen.

3–8. Neochonetes cf. N. whitei Sutherland and Harlow. 

3–5. USNM 476647, USGS colln. BL31394–PC; ventral view ×1 and enlarged 

(×3) ventral and anterior views of articulated, slightly decorticated 

specimen; note extremely weak mesial fold in sulcus; gastropod(?) boring 

in pedicle valve. 

6–8. USNM 476648, USGS colln. BL12096–PC; ventral view ×1 and enlarged 

(×3) ventral and anterior views of complete, slightly decorticated, 

articulated specimen; note absence of mesial fold in sulcus. 

9–14. Composita ovata Mather. 

9, 10. USNM 476649, USGS colln. BL12096-PC; ventral and anterior views of 

large, complete, articulated specimen.

11–14. USNM 476650, USGS colln. BL6852–PC; ventral, anterior, lateral, and 

posterior views of well-preserved articulated specimen with slightly 

broken beak. 

15–23. Composita gibbosa Mather.

15–18. USNM 476651, USGS colln. BL6852–PC; ventral, lateral, dorsal, and 

anterior views of mature specimen.

19–23. USNM 476652, USGS colln. BL12096–PC; ventral, posterior, dorsal, 

anterior, and lateral views of mature specimen; beak slightly broken. 

24–26. Pulchratia? pustulosa Sutherland and Harlow. 

24, 25. USNM 476653, USGS colln. BL12096–PC; dorsal and ventral views of 

complete specimen, beak area slightly crushed.

 26. USNM 476654, USGS colln. BL6852A–PC; dorsal view of complete, 

crushed specimen.

27–31. Sandia welleri (Mather). 

 27–29. USNM 476655, USGS colln. BL12096–PC; ventral, posterior, and lateral 

views of mature specimen; trail broken off.

 30, 31. USNM 476656, USGS colln. BL12096–PC; posterior and ventral views of 

decorticated specimen; trail broken.

32–40. Desmoinesia nambeensis Sutherland and Harlow. 

 32. USNM 476657, USGS colln. BL6852–PC; dorsal view of incomplete 

specimen; ornamentation slightly coarser than typical.

33, 34. USNM 476658, USGS colln. BL6852–PC; enlarged (×2) dorsal and 

anterior views of internal mold of brachial valve; note prominent margin 

around edge of visceral disc.

 35. USNM 476659, USGS colln. BL6852–PC; enlarged (×2) posterior view of 

internal mold of brachial valve.

36–40. USNM 476660, USGS colln. BL31394–PC; ventral view (×1) and 

enlarged (×2) lateral, ventral, dorsal, and posterior views of slightly 

broken, decorticated, articulated specimen. 
U.S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 158

SPERIFER, NEOCHONETES, COMPOSITA, PULCHRATIA?, SANDIA, AND DESMOINESIA 

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