Bulletin 39

New Mexico Museum of Natural History & Science

A Division of the
DEPARTMENT OF CULTURAL AFFAIRS

Upper Aptian-Albian Bivalves of Texas and Sonora.

Biostratigraphic, Paleoecologic and Biogeographic Implications

edited by
Robert W. Scott

Albuquerque, 2007

Aiapmoltsgeilroaientcg-iaroAtalipobhginacs,
Upper
Scott
R.W.
Sonora:
Texas
of
BPBivalves
Iand
39
Bulletin
NMMNH&S

Bulletin 39
New Mexico Museum of Natural History & Science
A Division of the
DEPARTMENT OF CULTURAL AFFAIRS

Upper Aptian-Albian Bivalves of Texas and Sonora:

Biostratigraphic, Paleoecologic and Biogeographic Implications

edited by

Robert W. Scott

Albuquerque, 2007

Bulletin 39
New Mexico Museum of Natural History & Science
A Division of the
DEPARTMENT OF CULTURAL AFFAIRS

Upper Aptian-Albian Bivalves of Texas and Sonora:

Biostratigraphic, Paleoecologic and Biogeographic Implications

edited by

Robert W. Scott

New Mexico Museum of Natural History & Science

Printed with the support of the

University of Texas, University of Tulsa, Precision Stratigraphy
Associates, and the U.S. Bureau of Land Management
Albuquerque, 2007

STATE OF NEW MEXICO

Department of Cultural Affairs
Stuart A. Ashman, Cabinet Secretary
NEW MEXICO MUSEUM OF NATURAL HISTORY AND SCIENCE
Adrian P. Hunt, Ph.D., Executive Director
BOARD OF TRUSTEES
Bill Richardson, Governor, State of New Mexico, ex officio
Adrian P. Hunt, Ph.D., Executive Director, ex officio
Gary Friedman, President
Mary B. Gavin, Emerita
Peter F. Gerity, Ph.D.
Robert A. Jung II
Laurence Lattman, Ph.D.
Morton Lieberman, Ph.D.
Imogene Lindsay, Emerita
Viola Martinez
John Montgomery, Ph.D.
Joseph Powell, Ph.D.
Dennis P. Trujillo, Ph.D.
Alexa Tysseling
Steve West
Ron Wilmot

Cover illustration: Scabrotrigonia [Trigonia] Clavigera Cragin, 1893 (from Cragin, F.W., 1893,
Geological Survey of Texas, 4th Annual Report)

EDITORIAL BOARD
Spencer G. Lucas, Ph.D., Managing editor
John R. Foster, Ph.D.
Jerald D. Harris, Ph.D.
Adrian P. Hunt, Ph.D.
Gary S. Morgan, M.S.
Justin A. Spielmann, B.A.
Roxanne Witt, Production editor
Original Printing

ISSN:1524-4156
Published by Authority of the State of New Mexico
Available from the New Mexico Museum of Natural History, 1801 Mountain Road NW, Albuquerque, NM 87104
Telephone (505) 841-2800; Fax (505) 841-2866; www.nmnaturalhistory.org
Published as Public Domain, therefore reproducible without permission, Source credit requested.

iii

CONTENTS
Preface
Chapter 1
1 Late Aptian Bivalve Assemblage of the Transgressive Hensel Sandstone,
Central Texas...............................R. W. Scott, D. L. Amsbury A. Molineux, and W. C. Ward 1
Chapter 2

2 Late Aptian-Early Albian Mollusks of the Comanchean and Sonora Shelves...R. W. Scott 7

vii

PREFACE
The two reports that comprise this Bulletin are complementary. The first report describes one
of the best exposed and complete outcrops of the transition between the Upper Aptian Hensel Sand
stone and the mainly Lower Albian Glen Rose Formation in Texas. This important stratigraphic
interval records the transgression that resulted in widespread drowning of the Comanche Shelf along
the northern margin of the Proto-Gulf of Mexico basin. The geologic significance of the outcrop at
Edgefalls was recognized by David Amsbury and Bill Ward, who promoted its study. Subsequently,
the well-preserved molluscan fauna preserved as dolomite casts was collected by paleontologists at
the Texas Natural Science Center of The University of Texas at Austin.
The second report presents updated systematics, taxonomy, and nomenclature of the diverse
bivalve fauna collected at Edge Falls and in northwestern Sonora. The research in Sonora was conduct
ed by Dr. Carlos M. Gonzlez Len and Hannes Lser of ERNO, Instituto de Geologa, Universidad
Nacional Autnoma de Mxico in Hermosillo, Sonora, Mxico. The bivalve biota consists principally
of oysters and trigoniids that are well known in northern Mexico, Arizona, New Mexico, and Texas.
Not surprisingly, many of the species are either con-specific with Mediterranean species or are closely
related. Together these reports provide the stratigraphic control for significant Lower Cretaceous
megafauna having worldwide paleoecologic and paleobiogeographic significance.

R. W. Scott

Scott, R.W., 2007, Upper Aptian-Albian Bivalves of Texas and Sonora: Biostratigraphic, Paleoecologic and Biogeographic Implications. New Mexico Museum of Natural History and Science Bulletin 39.

LATE APTIAN BIVALVE ASSEMBLAGE OF THE TRANSGRESSIVE HENSEL

SANDSTONE, CENTRAL TEXAS
ROBERT W. SCOTT1, DAVID L. AMSBURY2, ANN MOLINEUX3, AND WILLIAM C. WARD4
1Precision

Stratigraphy Associates and Tulsa University, RR3 Box 103-3, Cleveland, Oklahoma 74020 USA, [email protected]; 2deceased, Ker
rville, TX; 3Texas Memorial Museum, Austin, TX; 426328 Autumn Glen, Boerne, TX, 78006, wcwkaw@ aol.com

AbstractThe Hensel Sandstone was deposited during the latest Aptian transgression onto the Co
manchean Shelf in Central Texas. A unique assemblage of bivalves and gastropods is preserved as
dolomitized shells in the uppermost bed of the Hensel Sandstone at Edge Falls, central Texas. Trigoniid
bivalve taxa are abundant and diverse, and other bivalve and gastropod groups are represented by one
or few species. This assemblage is similar to that in the Aptian Lower Mural Formation of southern
Arizona and in northern Sonora. Some genera are also members of the South American assemblage.
The nearshore biota characterizes a transgressive system tract facies.

INTRODUCTION
Exceptionally well-preserved molluscan megafossils that
show exquisite detail weather out of the upper bed of dolomitic
calcareous sandstone in the Upper Aptian Hensel Sandstone at
Edge Falls south of Kendalia, Central Texas (Fig. 1A). This type of
preservation is rare for Early Cretaceous fossils in Texas. The nor
mal preservation of aragonitic mollusks is as internal or external
molds and casts. In fact, the type specimens of some Texas species
are internal casts that show very few critical biological features
needed to define species accurately, to understand their phylo
genetic history, and to interpret how organisms lived. So, these
dolomite specimens are important to understand Cretaceous pale
ontology in North America. The objective of this contribution is to
provide new stratigraphic data that: 1) support regional sequence
stratigraphic analyses of Cretaceous strata and 2) document the
stratigraphic and lithologic framework of nearshore marine mol
lusks. Study of the Edge Falls assemblage will interest specialists of
Cretaceous mollusks everywhere because this material will clarify
not only many species concepts and their age ranges, but also will
complement understanding of the suprageneric classification and,
hence, evolution of this important group of organisms.
Lower Cretaceous strata of Texas were deposited on the
Comanche Shelf, a mixed carbonate-siliciclastic sedimentary
province. Older regional structures greatly influenced thicknesses
and facies. The section thickens eastward into the East Texas
Basin where shale, sandstone and carbonate are about of equal
volumes. The Central Texas area was a positive feature where
thin carbonates accumulated. In southwestern Texas, the section
thickens into the Chihuahua Trough. Paleoclimate modeling sug
gests that Tethyan sea-surface currents flowed westward across
the Comanche Shelf, and surface winds varied seasonally from
the southwest in summer to the west and northwest in winter
(Glancy et al., 1993).
Edge Falls is an intermittent waterfall on Curry Creek, a
tributary of the Guadalupe River, about 4 mi (6.4 km) south of
Kendalia, Kendall County, Texas (eastern edge of the Kendalia,
Texas USGS 71/2 minute topographic sheet, UTM 14547692E
3309307N [WGS84/NAD83], (2954.81N, 98 30.35W) (Fig. 1A).
Erosion has undercut the resistant beds of the basal Glen Rose
Formation exposing uppermost dolomitic and sandy beds of the
Hensel Sandstone. The site has been a recreational attraction for
many years and is now administered by the Nature Conservancy
(Fig. 2A). A second section spans the Cow Creek Limestone and

Hensel Sandstone and was measured in the Edge Falls Ranches

development 0.7 mi northeast of the Guadalupe River crossing
about 6 mi (~10km) south of Kendalia (UTM 14546331E 3306524N
[WGS84/NAD83], 29 53.31N, 98 31.21W) (Fig. 1B).
STRATIGRAPHY
Regional widespread unconformities on the Comanche Shelf
divide the Cretaceous into three long-term depositional cycles that
are provincial time-rock units. The second long-term cycle is the
Comanchean Series (Hill, 1901) that spans the interval from the
intra-Aptian unconformity at the top of the Sligo Formation to
the unconformity between the Washita Group and the Woodbine
Formation (McFarlan, 1977; Scott et al., 2003). The Comanchean
correlates with mid-Aptian to mid-Cenomanian and was about
22 m.y. in duration.
The oldest transgressive-regressive cycle of the Comanchean
Series is represented in outcrop by the Sycamore Sandstone,
the Hammett Formation and the Cow Creek Limestone (Lozo
and Stricklin, 1956; Loucks, 1977). In the subsurface this cycle is
comprised of the Pine Island Shale and James Limestone of the
Pearsall Formation. The base of this cycle is Ap SB SL 1 (Scott et
al., 2000), and the revised age of the overlying strata is 122 Ma.
The second cycle begins with the Hensel Sandstone, and in the
subsurface the Bexar Shale, and grades up into the Glen Rose
Formation (Amsbury, 1974; Loucks, 1977, fig. 4; Scott, 1993, fig. 3).
The base of this cycle is Ap SB PR 2, and basal beds are now dated
at 118 Ma. Locally, lowstand facies may underlie transgressive
facies of the Hensel, which was deposited along the shore of the
Llano Uplift (Loucks, personal communiction, 2006). These units
crop out south of Kendalia, Texas, where two outcrop sections
were measured and a well-preserved, diverse molluscan fauna
was collected (Fig. 3).
The Cow Creek Limestone gradationally overlies the
Hammett Formation and disconformably underlies the Hensel
Sandstone. It is the shallow water deposit of a single transgressive
regressive cycle that spans the boundary of the Lower and Upper
Aptian (Young, 1974, 1986). In the outcrop at Edge Falls Ranches,
8.8 m of the Cow Creek are exposed (Fig. 3). The facies succession
grades up from wavy nodular-bedded, soft, argillic limestone to
medium-bedded, resistant, well-sorted grainstone. Crossbedding
dips at 15-20S45W. The topmost bed is caliche breccia with chert
crusts. The basal part of the Hensel is covered.
The Hensel Sandstone was named for outcrops of coarse,

FIGURE 1. Location maps. Topographic maps of Kendalia Quadrangle 1:24,000 modified from Topozone.com used with permission. Shaded areas
are covered by vegetation.

arkosic sand overlying the Cow Creek Limestone and underlying

the Glen Rose Formation in central Texas. Exposures are along
the Colorado River and its tributaries in Travis County and east
ern Burnet County (Lozo and Stricklin, 1956). The Hensel crops
out widely in Kendall and Comal counties (Stricklin et al., 1971;
Barnes, 1983; Collins, 1994) and it has been cored near the outcrop
belt (Amsbury, 1996a, b). The base of the overlying Glen Rose is
placed at the base of resistant, medium-bedded limestone (19.9 m
in the Edge Falls Ranch section and 5.9m in the Edge Falls Nature
Conservancy section) (Figs. 2A, 3). This bed is an oolitic-mollusk
grainstone, in which most bivalve shells have been leached (Ap
pendix 1). The biota consists of trigoniids and thick-shelled oysters;
wedge-planar cross stratification is developed near the falls. The
lithology of the basal 10 m of the Glen Rose is similar, but the
abundance of large bivalves decreases up section.
The uppermost 6 m of the Hensel Sandstone are a mixed
siliciclastic-carbonate lithology. The unit grades up from clay and
siltstone to calcareous sandstone to sandy limestone at the top.
Some beds are highly bioturbated and locally cross bedded. The
abundance and diversity of mollusks increase based on visual ob
servations (Fig. 2C, D). The uppermost bioturbated, sandy, calcare
ous siltstone bed at the water falls section has oysters, trigoniids,
turritellid gastropods, and crab claws (Fig. 3) (Appendix 2). The
underlying bed in this section (between 3.7 and 4.9 m) is the fossil
rich, bioturbated, silty dolomite from which the dolomite-replaced
shells were collected. Nodules of milky quartz or, locally, white
calcite may have replaced gypsum nodules (Fig. 2B). The bed has
many vugs, some of which are up to a meter in diameter. The top
contact of this bed with the overlying calcareous siltstone bed is
sharp and highly irregular; it may represent a hiatus formed at a
ravinement surface during relative sea-level rise.
BIOSTRATIGRAPHY
The Hammett, Cow Creek, Hensel, and Glen Rose forma
tions span the Lower Aptian to Lower Albian. The Hammett
Formation is in the uppermost Lower Aptian Dufrenyoia rebeccae
Zone, and the Cow Creek Limestone is in the basal Upper Ap

tian Dufrenoyia justinae (Hill) Zone (Young, 1974, 1986). A new

ammonite was collected from the lower part of the Cow Creek
Limestone, Eodouvielliceras sp. cf. E. horridum Riedel (L. Bulot, per.
comm., December, 1999, February, 2006, Appendix 3). This species
ranges from uppermost Aptian to lowermost Albian. The Hensel
Sandstone is in the Kazanskyella spathi Zone. The lowest occurrence
of the Lower Albian Hypacanthoplites cragini Scott Zone is about 3
m above the base of the Glen Rose (Young, 1974, 1986).
The Late Aptian age, about 118-112 ma, of the Hensel
Sandstone at Edge Falls is based on these ammonites and on
the stratigraphic relations of the overlying Glen Rose Formation
with other localities that contain ammonites. The upper Upper
Aptian ammonite, Kazanskyella spathi, has been found at three
localities of the lower Glen Rose in central Texas (Young, 1974).
The lower Lower Albian ammonite, Hypacanthoplites cragini Scott,
has been collected in the lower 90 m of the Glen Rose. The middle
Lower Albian ammonite, Douvilleiceras sp. cf. D. mammillatum
(Schlotheim), spans the Corbula marker bed about 100 m above
the base of the Glen Rose.
Some of the trigoniid species found in the top of the Hensel
Sandstone at Edge Falls are known from Arizona with ammo
nites (Stoyanow, 1949), but one, Pisotrigonia sp., is new to North
America. Rutitrigonia weaveri (Stoyanow) is with Upper Aptian
ammonites in Arizona. Scabrotrigonia stolleyi (Hill) occurs in
Arizona with Upper Aptian ammonites, and in Texas it is found
throughout most of the Glen Rose Formation, ranging into the
Lower Albian.
Three non-trigoniid bivalves have been identified to date
(Scott, this volume). Cucullaea (Idonearca) gratiota Hill ranges from
the uppermost Upper Aptian Kazanskeylla spathi Zone into the
Lower Albian above the Douvilliceras mammillatum Zone in Texas
and Arizona. Anatina simondsi Whitney (1952a) is an elegant,
elongate shell that until now was represented only by composite
molds (UT40578); it is known from the middle Glen Rose zone
of Salenia texana Credner. A perfectly preserved entire free shell
shows the internal features and is identified as Epicyprina sp. cf. E.
[formerly Corbis] hamiltonae Whitney (1952b), which was defined
upon internal molds. This species originally was reported from

FIGURE2. Close-up photos of Edge Falls. A, Permanent pool with water line as of December, 1999 at scoured recessed face near top of Hensel Sandstone
below cliff cut in basal Glen Rose Formation; above is the very white travertine lip of the wet-weather falls. B, View of top of fossiliferous Hensel bed
showing mainly disarticulated, concave down, transported shells and fragments; these originally aragonitic shells are now replaced by dolomite.
C, Close-up of several shells of trigoniids (note coin for scale). D, Edge-on view of bed showing abundant milky quartz concretions that possibly
replaced original gypsum nodules that grew in the sediment soon after deposition; sample EF-4.

the lower Glen Rose. A fourth specimen is tentatively identified

as Venericardia sp. Gastropods in this collection await study.
PALEOECOLOGY
The depositional environment of the upper Hensel Sand
stone at Edge Falls was a nearshore, mixed carbonate-terrigenous,
shallow marine shelf of normal oceanic conditions within the zone
of wave or current energy. The overlying basal Glen Rose Forma
tion consists largely of fully marine limestones that contain abun
dant mollusks including marine clams, snails, and ammonites,
echinoderm fragments, calcareous algae, miliolid foraminifera,
and ooliths (Stricklin et al., 1971). The more lagoonal upper
Hensel dolomite beds also contain abundant fragmented echinoid
spines and plates, and ophiuroid ossicles, as well as the marine
clams described here (Amsbury, 1996b).
The bivalve shells in bed 3 between 3.7 and 4.9 m (Fig. 3) are
replaced by dolomite, and the original ornamentation and internal
features are very well preserved (Fig. 2C). The diagenetic condi
tions of this preservational phenomenon extended for several km
because dolomitic shells occur near the Guadalupe River Resort
east of Ammans crossing on state road FM 474. The greatest majority of specimens from the uppermost 30 cm of bed 3 near the
top of the Hensel Sandstone in the measured section at Edge Falls
(Fig. 3; Appendix 1) are bivalves, and eight species are identified
(Appendix 2); gastropods are much less abundant. Most of the
species are trigoniids, which generally are strongly ornamented,
triangular-shaped clams. Most of the specimens can be assigned

to previously named species; one appears to be new to Texas.

The molluscan deposit in the uppermost 30 cm consists
mainly of shell fragments and disarticulated shells suggesting
transport either by currents or waves (Fig. 2C, D). Many of the
shells are concave down, which is the most stable position instrong
currents or waves. The apex of the snails is not in a consistent di
rection because they are surrounded by so much debris that they
could not respond to the water flow. Some of the shell breakage
could have been done by scavenging or predaceous animals like
crabs. Large-diameter burrows are common in the upper 1 to 1.3
m of sandy dolomite at the top of the Hensel Sandstone. Crab
claws and other fragments are abundant in the basal bed of the
Glen Rose Formation. This deposit may have accumulated near
where the mollusks lived, but shells have been winnowed and
subsequently re-oriented by bioturbation.
The abundance of molluscan shells and debris increases
upwards in the uppermost bed of the Hensel Sandstone, suggest
ing that many of these species lived in this sandy substrate. The
species recognized to date lived as shallow infaunal suspension
feeders that fed upon suspended and mobile organic particles in
the sea water. The uppermost 30 cm of sediment probably was
winnowed at the end of deposition and possibly subaerially ex
posed prior to the change to a limestone depositional regime of
the Glen Rose Formation.
The basal 4.6 m of the Glen Rose Formation contains molds
of disarticulated, concave-up and down trigoniid bivalves similar
to that in the topmost bed of the Hensel Sandstone. This suggests
that environmental conditions such as temperature, salinity, water

FIGURE 3. Measured sections at Edge Falls Ranches and Edge Falls measured by D.L. Amsbury, R.W. Scott, and W. Ward, December 13, 1999, Kendalia
Quadrangle, 4 mi S20E of Kendalia and downstream from Edge Falls Road crossing of Curry Creek.

depth, water energy and turbidity, nutrients, and water chemis

try were similar. The major difference between deposition of the
Hensel and the Glen Rose was in the virtual absence of a quartz
sand supply during deposition of the Glen Rose and subsequent
dolomitization of the Hensel beds. The sharp contact between the
Hensel and Glen Rose at 5.1 m shows evidence of re-working of the
Hensel shells and erosion of the top of the bed before deposition of
the Glen Rose sediments. This high-energy event may have been
the result of a storm or a set of storms or by ravinement during
sea-level rise. Possibly sealevel fell and shallow waves eroded the
bivalve beds; perhaps even the beds were subaerially exposed
prior to flooding and carbonate deposition.
CONCLUSIONS

ing transgression of the second cycle of the Comanchean Series

in Central Texas. These shells were replaced by dolomite and are
now exposed beneath the carbonate shelf Glen Rose Formation.
These shells reveal for the first time important external and internal
morphological features of species that prior were known only from
composite molds. This assemblage has genera found both in the
Mediterranean Tethys and in southern South America.
ACKNOWLEDGMENTS
Dr. Joe Yelderman and family, landowners, graciously per
mitted access to the outcrop at Edge Falls and permission to collect
by the staff of the Texas Natural Science Center, The University
of Texas at Austin. We appreciate the perceptive and constructive
review by Bob Loucks that focused this contribution.

An assemblage of abundant and relatively diverse mol

lusks occupied the nearshore, high energy sandy substrate dur
REFERENCES
Amsbury, D.L., 1974, Stratigraphic petrology of Lower and Middle Trinity
rocks on the San Marcos Platform, South-Central Texas, in Perkins, B.F.,
ed., Aspects of Trinity Division geology: Geoscience and Man, v. VIII,
Louisiana State University, p. 1-35.
Amsbury, D.L., 1996a, Pearsall (Aptian Cretaceous) subsurface to outcrop

sequence stratigraphy, central Texas: Transactions of the Gulf Coast

Association of Geological Societies, v. 47, p. 1-7.
Amsbury, D.L., 1996b, Observations of Hensel outcrops and cores, central
Texas, in Amsbury, D.L., and Jones, J.O., eds., Classic outcrops of the
Trinity Group (Lower Cretaceous) of south-central Texas: Sequence

5
stratigraphic units of marine and non-marine stratigraphic packages,
a field book and related papers: Fieldbook 99-2, South Texas Geological
Society, p. 69-99.
Barnes, V.E., 1983, Geologic Atlas of Texas San Antonio Sheet: Bureau of
Economic Geology, University of Texas Austin.
Collins, E., 1994, Geology of the Kendalia Quadrangle, Texas, Revision
2 (1995): Bureau of Economic Geology, University of Texas Austin,
Open-file map.
Glancy, Jr., T.J., Arthur, M.A., Barron, E.J., and Kauffman, E.G., 1993, A
paleoclimate model for the North American Cretaceous (Cenomanian
Turonian) epicontinental sea, in Caldwell, W.G.E., and Kauffman, E.G.,
eds., Evolution of the Western Interior Basin: Geological Society of
Canada Special Paper 39, p. 219-241.
Hill, R.T., 1901, Geography and geology of the Black and Grand Prairies,
Texas: U.S. Geological Survey, 21st Annual Report, pt. 7 Texas, 292
p.
Loucks, R.G., 1977, Porosity development and distribution in shoal-water
carbonate complexes-subsurface Pearsall Formation (Lower Creta
ceous) South Texas, in Bebout, D.G., and Loucks, R.G., eds., Cretaceous
carbonates of Texas and Mexico: Applications to subsurface explora
tion: Bureau of Economic Geology, University of Texas at Austin, Report
of Investigations No. 89, p. 97-126.
Lozo, F.E., and Stricklin, Jr., F.L., 1956, Stratigraphic notes on the outcrop
basal Cretaceous, central Texas: Gulf Coast Association of Geological
Societies, v. 6, p. 67-78.
McFarlan, E., Jr., 1977, Lower Cretaceous sedimentary facies and sea level
changes, U.S. Gulf Coast, in Bebout, D.G., and Loucks, R.G., eds.,
Cretaceous carbonates of Texas and Mexico: Applications to subsur
face exploration: Bureau of Economic Geology, University of Texas at
Austin, Report of Investigations No. 89, p. 5-11.
Scott, R.W., 1993, Cretaceous carbonate platform, U.S. Gulf Coast, in
Simo, J.A.T., Scott, R.W., and Masse, J.-P., eds., Cretaceous carbonate
platforms: American Association of Petroleum Geologists, Memoir
56, p. 97-109.

Scott, R.W., 2007, Late Aptian-Early Albian bivalves of the Comanchean

and Sonoran shelves, in Scott, R.W., ed., Upper Aptian-Albian Bivalves
otTexas and Sonora: Biostratigraphic, Paleoecologic and Biogeographic
Implications: New Mexico Museum of Natural History Bulletin, v.
39, p.5-37.
Scott, R.W., Benson, D.G., Morin, R.W., Shaffer, B.L., Oboh-Ikuenobe,
F.E., 2003, Integrated Albian-Lower Cenomanian Chronostratigraphy
Standard, Trinity River Section, Texas, in Scott, R.W., ed., Cretaceous
Stratigraphy and Paleoecology, Texas and Mexico: Perkins Memorial
Volume: GCSSEPM Foundation, Special Publications in Geology, No.
1, CD book, p. 277-334.
Scott, R.W., Schlager, W., Fouke, B., and Nederbragt, S.A., 2000, Are
Mid-Cretaceous eustatic events recorded in Middle East carbonate
platforms?, in A.S. Alsharhan and R.W. Scott, eds., Middle East Models
of Jurassic/Cretaceous Carbonate Systems: SEPM Special Publication
No. 69, p. 77-88.
Stoyanow, A., 1949, Lower Cretaceous stratigraphy in southeastern Ari
zona: Geological Society of America, Memoir 38, 169 p.
Stricklin, F.I., Jr., and Amsbury, D.L., 1974, Depositional environments
on a low-relief carbonate shelf, middle Glen Rose Limestone, Central
Texas: Geoscience and Man, v. VIII, p. 53-66.
Stricklin, F.I., Jr., Smith, C.I., and Lozo, F.E., 1971, Stratigraphy of Lower
Cretaceous Trinity deposits of Central Texas: University of Texas at
Austin, Bureau of Economic Geology, Report of Investigations No.
71, 63 p.
Whitney, M., 1952a, Some zone marker fossils of the Glen Rose Formation
of central Texas: Journal of Paleontology, v. 26, p. 65-73.
Whitney, M., 1952b, Some new Pelecypoda from the Glen Rose Formation
of Texas: Journal of Paleontology, v. 26, p. 697-707.
Young, K., 1974, Lower Albian and Aptian (Cretaceous) ammonites of
Texas: Geoscience and Man v. 8, p. 175-228.
Young, K., 1986, Cretaceous, marine inundations of the San Marcos plat
form, Texas: Cretaceous Research, v. 7, p. 117-140.

APPENDIX 1
The Edge Falls section is a Nature Conservancy site in Kendall
County, Texas. Description of Edge Falls section measured beginning
about 100 yards downstream-southeast from the ladder at Edge Falls by
D. Amsbury, R. Scott, and W. Ward, December 13, 1999.
Unit 9 was measured in creek above waterfall. Total thickness from
base of Glen Rose Formation unit 4 to car park hand-leveled at ladder
was 13.7 m. Location UTM 14547692E 3309307N (WGS84/NAD83), 29
54.81N, 98 30.35W.
Bed No./Thickness m/Cumulative Thickness m
Glen Rose Formation (lower part)
9
4.0/17.8 Limestone, light yellow gray, fine grained,
bioturbated.
8
0.8/13.8 Limestone, bioclast-peloid grainstone, poorly sorted
7
3.5/13.0 Limestone, oolitic-bioclast-peloid grainstone,
abundant oysters & trigoniids, abundance decreases upward.
Sample 7 at base, sample 8 at top.
3.4/9.5 Limestone, oolitic grainstone, cross bedded; very
6
abundant molds of whole shells in lenses. Sample 6 in
upper part.

0.5/2.0 Sandstone, light brown, thin shell lag at base,

mid part bioturbated, upper part a shell concentration,
trigoniids & turritellids. Sample 2 in lower part.
1.5/1.5 Sandstone, light brown, medium to coarse grained,
small-scale trough cross beds, sparse burrows cemented with
calcite or dolomite; grades down into siltstone & sandy shale,
medium gray. Sample 1 in lower part.

The Edge Falls Ranches section was measured in shallow gully 0.7 km
east of the low water crossing of Edge Falls Road on Guadalupe River
by D.L. Amsbury, W. Ward, R. W. Scott, E. Robert, and A. Molineux,
September 18, 1999.
Bed No./Thickness m/Cumulative Thickness m
Glen Rose Formation (lower part)
7
6.0/25.9 Grainstone, few large mollusks, large exogyrids.
Hensel Sandstone (at Guadalupe Dam site Hensel is 11.6 m thick)
6
3.4/19.9 Grainstone to packstone, glauconitic, coarse quartz
sand, leached large concretions, whole leached mollusks.
7.8/16.5 Covered interval.

Hensel Sandstone
1.0/6.1 Siltstone, calcareous, abundant oysters, clam & snail
5
molds, & crab claws. Sample 5 in upper part.
1.4/5.1 Sandy dolomite, light gray, glauconitic, bioturbated,
4
sparse large concretions. Sample 4 in mid part.
1.7/3.7 Sandy dolomite, light brown, fine to coarse grained,
3
poorly sorted, glauconitic, in upper 0.75 m shells increase to
about 60% of grains, disarticulated, concave down, rarely
articulated & concave up or oblique; TMM collection; large,
ovate leached concretions. Sample 3 at top.

Cow Creek Limestone

5
1.4/8.7 Grainstone, fine grained, chert nodules.
4
2.1/7.3 Grainstone, well sorted, medium to coarse grained,
accretion cross bedded; whole mollusk shells.
3
1.5/5.1 Grainstone, fine grained, whole mollusks;
2
2.0/3.6 Grainstone, as above, less resistent.
1
1.6/1.6 Mudstone, calcareous, soft; base covered by alluvium.

6
APPENDIX 2
Classification of bivalves from uppermost Hensel Sandstone at Edge
Falls. Repository at the Texas Memorial Museum (TMM), Texas Natural
Science Center, Austin.
Class Bivalvia Linn, 1758
Subclass Pteriomorphia Beurlen, 1944
Order Arcoidea Stoliczka, 1871
Superfamily Arcacea Lamarck, 1809
Family Cucullaeidae Stewart, 1930
Genus Cucullaea Lamarck, 1801
Subgenus Idonearca Conrad, 1862
Cucullaea (Idonearca) gratiota Hill, 1888
Material. One disarticulated RV, TMM1994TX2.

Tribe Scabrotrigoniini Cooper, 1989

Genus Scabrotrigonia Dietrich, 1933
Scabrotrigonia stolleyi (Hill, 1893)
Material. One articulated specimen from Edge Falls locality is TMM
1994TX13, and two disarticulated specimens TMM1994TX9, -14.
Family Rutitrigoniidae van Hoepen, 1929
Rutitrigonia van Hoepen, 1929
Rutitrigonia weaveri (Stoyanow, 1949)
Material. TMM1994TX4, -5, -6, one RV, two LV.

Subclass Palaeoheterodonta Newell, 1965

Order Trigonioida Dall, 1889
Suborder Myophorellina Cooper, 1991
Superfamily Myophorellacea Kobayashi, 1954
Family Vaugoniidae Kobayashi, 1954
Subfamily Quadratotrigoniinae Kobayashi, 1954
Genus Quadratotrigonia Dietrich, 1933
Quadratotrigonia guildi (Stoyanow, 1949)
Material. Two nearly complete disarticulated RV and LVTMM1994TX7, -8.
Superfamily Megatrigoniacea van Hoepen, 1929
Family Megatrigoniidae van Hoepen, 1929
Subfamily Pterotrigoniinae van Hoepen, 1929
Tribe Pterotrigoniini van Hoepen, 1929
Genus Pisotrigonia van Hoepen, 1929
Pisotrigonian. sp.
Material. Five disarticulated valves either partly buried by matrix or
with the rostrum broken TMM1994TX10, -11, -12, -15, -16.

Order Veneroida H. Adams & A. Adams, 1856

Superfamily Carditacea Fleming, 1820
Family Carditidae Fleming, 1828
Subfamily Venericardiinae Chavan, 1969
Genus Venericardia Lamarck, 1801
Venericardia sp.
Material. A single LV showing the strong radial ribs but mainly covered
with matrix, TMM1994TX18.
Superfamily Arcticacea Newton, 1891
Family Arcticidae Newton, 1891
Genus Epicyprina Casey, 1952
Epicyprina [Corbis] hamiltonae (Whitney, 1952b)
Material. One exquisitely preserved entire LV free of matrix,
TMM1994TX1.
Taxonomic Position Indeterminate
Anatina simondsi Whitney, 1952a
Material. A single articulated entire shell with posterior broken,
TMM1994TX3.

APPENDIX 3
NEW AMMONITE SPECIMEN IN COW CREEK LIMESTONE
LUC G. BULOT
Chemin des Iscles - Quartier de lAbb Bressy, 13660 Orgon, France, [email protected]
The specimen was an external mould and we made a latex cast
of it. It is clearly an Eodouvilleiceras. Preservation prevents a 100% sure ID
at the specific level but it is closer to horridum Riedel than to any other
species described in the literature. The known range of Eodouvilleiceras
is uppermost Aptian (Nolani Zone) to lowermost Albian (Tardefurcata
Zone) but the maximum abundance is in the lower part of the range. The
exact range of E. horridum is still unknown and disputed. The specimen is

in my collection in the Museum of Paleontology at Marseille University.

The latex cast is in Grenoble with Emmanuel Robert.
Riedel, L, 1938, Ammonites del Cretcico inferior de la Cordillera
Oriental: Estudios geolgicos y paleontolgicos sobre la Cordillera Ori
ental de Colombia, v. 2: Departamento de Minas y Petrleos, Republica
de Colombia, Ministerio de Industrias y Trabajo, p. 7-78.

Scott, R.W., 2007, Upper Aptian-Albian Bivalves of Texas and Sonora: Biostratigraphic, Paleoecologic and Biogeographic Implications. New Mexico Museum of Natural History and Science Bulletin 39.

LATE APTIAN-EARLY ALBIAN BIVALVES OF THE COMANCHEAN AND SONORAN SHELVES

ROBERT W. SCOTT
Precision Stratigraphy Associates and Tulsa University, RR3 Box 103-3, Cleveland, Oklahoma 74020 USA, [email protected]

AbstractA diverse assemblage of bivalves and fewer species of gastropods, ammonites, corals, sponges,
and algae characterize Upper Aptian and Lower Albian strata in the southwestern United States and
northwestern Mexico. Mixed carbonate and siliciclastic strata of the Comanchean Shelf in central Texas
yield numerous oyster and trigoniid bivalves. At one new locality complete shells are preserved in the
uppermost Hensel Sandstone. Additional new collections in coeval strata in Sonora corroborate the
well known assemblages in southern Arizona. The taxonomy is updated so that biostratigraphic and
biogeographic hypotheses can be developed. The mainly infaunal species represent both Tethyan and
south Temperate components.

INTRODUCTION
Early Cretaceous bivalve mollusks are a diverse and abun
dant part of marine paleocommunities on terrigenous clastic and
carbonate substrates. Such substrates were widespread during
the Early Cretaceous on the Comanche Shelf in southwestern
United States and on the Sonoran Shelf in northwestern Mexico
(Figs. 1-2). Many bivalve and gastropod taxa have been defined
in monographs (among others, see Roemer, 1852; Bse, 1910;
Stanton, 1947; Stoyanow, 1949; Perkins, 1961; Gonzlez Lenet al.,
2007). The purpose of this contribution is to review selected Lower
Cretaceous bivalves including trigoniids of the Comanchean and
Sonoran shelves in northwestern Mexico and southwestern U.S.,
to revise their taxonomy, and to define their stratigraphic ranges
relative to guide fossils that correlate these species with stadial
stratotypes. New collections from excellent outcrops in Texas
and Sonora have produced many fine specimens that enable
comparison with previous studies in North America and Europe.
Thus, a comprehensive taxonomic evaluation of the assemblage
is possible. However, the collections do not meet the standards
for statistical treatment.
The Early Cretaceous was a time of rapid diversification of
trigoniid genera (Stanley, 1978), particularly in the Superfamily
Megatrigoniacea van Hoepen, 1829. The classification of the Order
Trigonioida Dall, 1889 is maturing but not yet settled. Cooper
(1991) recognized two suborders, six superfamilies, and ten fami
lies. Leanza (1993), on the other hand, recognized 14 subfamilies
within the Family Trigoniidae Lamarck, 1819. The basic goals of
each author were fundamentally different. Cooper attempted to
show cladistic and phylogenetic relationships. Leanza attempted
to refine taxonomy for future discussions.
Since 1852, 22 trigoniid species have been identified from
Lower Cretaceous strata in Texas, New Mexico, Arizona, and
northern Mexico by nine authors (Table 1). In order to identify
the new material and to assess its significance, it was imperative
to evaluate these species. In 1852, Roemer applied the European
taxon, Trigonia crenulata Lamarck, 1819, to specimens he collected
in central Texas. I have seen Roemers specimens in the Paleonto
logical Institute in Bonn, Germany, and they are like the species
common in the Glen Rose and Walnut formations, Trigonia stolleyi
Hill, 1893. In 1857, Conrad erected the taxon Trigonia emoryi for
specimens collected near the U.S.-Mexican border by the Emory
boundary survey, presumably from near Cerro de Cristo Rey at
El Paso, Texas. Gabb (1869) defined several molluscan species
from outcrops of Albian strata north of Hermosillo. Cragin (1893)
created four species from new collections by a subsequent border
survey from Upper Jurassic and Lower Cretaceous strata. Hill

(1893) named a new trigoniid taxon in the Glen Rose Formation

in central Texas. An Upper Jurassic species was named by Castillo
and Aguilera (1895) from Mexico. The extensive collections from
the Upper Jurassic Malone Formation near the Malone Mountains
in West Texas were described by Cragin (1905). In 1910 Bse stud
ied a diverse collection from Lower Cretaceous rocks in Chihua
hua, Mexico that included a new trigoniid. The Upper Aptian to
Lower Albian Mural Formation in southeastern Arizona yielded
excellent specimens that were named by Stoyanow (1949). Finally,
Marion Whitney (1952b) created three more trigoniid taxa from
the Glen Rose in central Texas. Trigoniids are also common in the
diverse molluscan fauna of the Aptian-Albian Alisitos Formation
in Baja California (Allison, 1974).
A new outcrop in Kendall County, Texas has produced the
complete
first
shells of many bivalves common in Lower Creta
ceous rocks in North America. South of Kendalia the waterfalls at
Edge Falls is a natural feature that exposes the transitional interval
between the Upper Aptian Hensel Sandstone and the uppermost
Aptian-Lower Albian Glen Rose Formation. A large collection of
shells replaced by dolomite were deposited in the Texas Museum
of Natural History (formerly Texas Memorial Museum-TMM).
Complementary collections were made from the Mural Formation
in northern Sonora by Carlos M. Gonzlez Len and Hannes Lser,
Instituto de Geologa, UNAM, Hermosillo, Sonora, Mxico.
Molluscan megafossils weather out of the upper bed of
dolomitic sandstone in the Hensel Sandstone and show exquisite
detail, which is rare preservation for Early Cretaceous fossils in
Texas. The normal preservation of mollusks is as internal or ex
ternal molds and casts. In fact, the type specimens of some Texas
species are internal casts that show very few critical biological
features needed to define species accurately, to understand their
evolutionary history, and to interpret how the organisms lived.
So, these new specimens are important to the understanding of
Cretaceous paleontology in North America.
The greatest majority of specimens from the uppermost 30
cm of bed three in the measured section at Edge Falls (Scott et al.,
this volume; Appendix 1) are bivalves; gastropods are much less
abundant. At this time seven species of bivalves are described and
named; future studies will identify additional species. Most of the
species are in the group of trigoniids, which generally are strongly
ornamented, triangular-shaped clams. Most of the specimens can
be assigned to previously named species; one appears to be new to
Texas. Study of the Edge Falls specimens will interest specialists of
Cretaceous molluscs everywhere because this material will clarify
not only many species concepts and their age ranges, but also will
complement understanding of the suprageneric classification and,

8
TABLE 1. Lower Cretaceous trigoniid species described from Texas and Mexico.
Species

Author/date

Types

Synonyms

Trigonia crenulata
Scabrotrigonia emoryi
Scabrotrigonia mooreana
Scabrotrigonia clavigera
Trigonia concentrica
Quadratotrigonia taffi
Trigonia stolleyi
Trigonia quadalupe
Q. maloneana
Quadratotrigonia guildi

Lamarck in Roemer, 1852

(Conrad, 1857)
(Gabb, 1861, 1869; Bse, 1910)

Bonn
USNM

S. mooreana

(Cragin, 1893)
Cragin, 1893
(Cragin, 1893)
Hill, 1893

unknown
UT 17243
BEG 36912, 36917, 21756
BEG 19723(509), 21758(509)
USNM

T. resoluta
T. saavedra

Bse, 1910
Stoyanow, 1949
(Stoyanow, 1949)
Stoyanow, 1949
Stoyanow, 1949

Quadratotrigonia mearnsi

(Stoyanow, 1949)

unknown
USNM 1807/29020, 28967
USNM 91205,91413
USNM 91418
USNM 91333,91320, 91321,
91334
USNM GU46, GU42, -45, -47,

Apiotrigonia cragini

(Stoyanow, 1949)

-50
USNM 91429,91430, 91434,

Apiotrigonia kitchini

(Stoyanow, 1949)

Buchotrigonia reesidei
Rutitrigonia weaveri
T. wendleri
T. whitneyi
T. gordoni
Pisotrigonia amsburyi

(Stoyanow, 1949)
(Stoyanow, 1949)
Whitney, 1952b
Whitney, 1952b
Whitney, 1952b
Scott, this publication

91445,91409
USNM 91499, 91505, 91507,
91603
USNM 91049, 91050, 91051
USNM 91997, 91200, 91245
UMMP 53653
UMMP 53654
UMMP 53655, UMMP 53656
UTTM1994 TX10-12,15,16

crenulata? fide Bse

crenulata? fide Gabb

S. mooreana
S. emoryi
vyschetzkii
Q. guildi
Q. guildi

S. mooreana
S. mooreana
Q. taffi

hence, evolution of this important group of organisms. Morpho

logical nomenclature follows that of Leanza (1993).
STRATIGRAPHY

FIGURE 1. Map showing locations of sections studied and new fossil sites on the
Comanche Shelf. BB = Bisbee Basin; CP = Coahuila Platform; CT = Chihuahua
Trough; CTP = Central Texas Platform; DRT = Devils River Trend; ETB = East
Texas Basin; FSB = Fort Stockton Basin; MB = Maverick Basin; SS = Sonoran
Shelf. Reproduced from Scott (2003) with permission of the Gulf Coast SEPM
Foundation.

On the Comanchean and Sonoran shelves, regional wide

spread unconformities divide the Cretaceous into three long-term
depositional cycles that are provincial time-rock units. The Hensel
Sandstone is in the lower part of the Comanchean Series (Hill,
1901), which is the second of the three cycles. The Comanchean
spans the interval from the intra-Aptian unconformity at the top
of the Sligo Formation below up to the unconformity between
the Washita Group and Woodbine Formation. The Comanchean
correlates with mid-Aptian to mid-Cenomanian, which was about
22 m.y. in duration.
The oldest transgressive-regressive cycle of the Comanchean
Series is represented by the Hammett Formation and Cow Creek
Limestone (Lozo and Stricklin, 1956; Loucks, 1977). The second
cycle begins with the Hensel Sandstone, which grades up into the
Glen Rose Formation (Amsbury, 1974). These units crop outsouth
of Kendalia, Texas where two outcrop sections were measured,
and a well-preserved, diverse molluscan fauna was collected (Scott
et al., this volume).
Edge Falls is an intermittent waterfall on Curry Creek, a
tributary of the Guadalupe River, about 4 mi (6.4 km) south of
Kendalia, Kendall County, Texas (eastern edge of the Kendalia,
Texas USGS 71/2 minute topographic sheet, UTM 14547692E
3309307N [WGS84/NAD83], 29 54.81N, 98 30.35W). Erosion
has undercut the resistant beds of the basal Glen Rose Formation
exposing uppermost dolomitic and sandy beds of the Hensel Sand

stone. The site has been a recreational attraction for many years
and is now administered by the Nature Conservancy. A second
section spans the Cow Creek Limestone and Hensel Sandstone
and was measured in the Edge Falls Ranches development 0.7 mi
northeast of the Guadalupe River crossing about 6 mi (~10 km)
south of Kendalia (UTM 14546331E 3306524N [WGS84/NAD83],
29 53.31N, 98 31.21W).
The Mural Formation in northern Sonora, Mexico, correlates
with the Upper Aptian and Lower Albian and is comprised of
the Fronteras, Rancho Bufalo, Cerro La Ceja, Tuape Shale, Los
Coyotes, Cerro La Puerta, Cerro La Espina, and Mesa Quemada
members in ascending order. These units were deposited during
three transgressive-regressive depositional cycles (Gonzlez Len
et al., 2007).
BIOSTRATIGRAPHY
The Late Aptian age, about 118-112 Ma, of the Hensel
Sandstone at Edge Falls is based on these bivalves and on the
stratigraphic relations with the overlying Glen Rose Formation at
other localities that contain ammonites. The upper Upper Aptian
ammonite, Kazanskyella spathi (Stoyanow, 1949), was found at three
localities of the lower Glen Rose in central Texas (Young, 1974). The
lower Lower Albian ammonite, Hypacanthoplites cragini Scott, 1930
has been collected in the lower 300 ft (95 m) of the Glen Rose. The
middle Lower Albian ammonite, Douvilleiceras sp. cf. D. mammil
latum (Schlotheim, 1813), spans the Corbula marker bed about
330 feet (100 m) above the base of the Glen Rose (Fig. 2).
Some of the trigoniid species found in the top of the Hensel
Sandstone at Edge Falls are known from Arizona with ammonites
(Stoyanow, 1949), but one, Pisotrigonia n. sp., is new to North
America. Quadratotrigonia guildi (Stoyanow, 1949) occurs with
Lower Aptian ammonites in Arizona, so this occurrence in the
Hensel extends its range into the Upper Aptian. Rutitrigonia weaveri
(Stoyanow, 1949) is with Upper Aptian ammonites in Arizona.
Scabrotrigonia mooreana (Gabb, 1861) occurs in Arizona with Upper
Aptian ammonites, and in Texas it is found throughout most of
the Glen Rose Formation, ranging into the Lower Albian.
Three non-trigoniid bivalves have been identified to date.
Cucullaea (Idonearca) inermis (Gabb, 1869) ranges from the up
permost Upper Aptian zone of Kazanskeylla spathi into the Lower
Albian above the zone of Douvilliceras mammillatum in Texas and
Arizona. Platymyoidea [formerly Anatina] simondsi (Whitney,
1952a) is an elegant elongate shell that until now was represented
only by composite molds (UT40578); it is known from the middle
Glen Rose zone of Salenia texana Credner. A perfectly preserved
entire free shell shows the internal features and is identified as
Epicyprina [Corbis] hamiltonae (Whitney, 1952b), which was defined
upon internal molds. This species originally was reported from
the lower Glen Rose. A fourth specimen is tentatively identified
as Venericardia sp. Gastropods in this collection await study.
PALEOECOLOGY
Two distinct latest Aptian-earliest Albian bivalve assem
blages in the northwestern end of the Chihuahua Trough were
dominated by: 1) epifaunal, suspension feeding exogyrid oysters
in the transgressive systems tracts and 2) semi-infaunal to in
faunal, suspension feeding trigoniids in deeper shelf mudrocks.
Assemblages of abundant Amphidonte obliquata (Pulteney, 1813)
populated the shallow shelf substrates of the Fronteras Formation
in Sonora, and Aetostreon latissimum (Lamarck, 1801) is abundant
in the Cerro La Ceja Formation in Sonora, parts of the Lower
Member of the Mural Formation in SE Arizona, and the Cow Creek
Limestone in Texas. The Cerro La Ceja represents transgressive
deposits of a depositional cycle, and the Cow Creek west of Austin,

9
Texas, is the nearshore, prograding systems tract.
Trigoniid bivalves were efficient shallow burrowers adapted
to muddy and unstable sandy substrates (Stanley, 1978). The
steinmanellids with knobby flank costae and coarse areal or
namentation were semi-infaunal in muddy substrates oriented
with the posterior elevated above the substrate at a steep angle
(Villamil et al., 1998) with the current flowing across the valve
from ventral to dorsal margins for efficient waste removal (Saul,
1978). The presence of a gape at the posterior margin indicates
that the mantle was without siphons so that the posterior had to
be elevated above the sediment-water interface (Stanley, 1978;
Saul, 1978; Kelly, 1995; Checa and Jimnez-Jimnez, 2003; Lazo,
2003). The species from the Comanchean and Sonoran shelves in
the SW U.S. and NW Sonora have such a posterior gape. Most of
these species also possess symmetrical, oblique costae that migrate
along the commissure during growth; the Apiotrigonia Cox, 1952
and Buchotrigonia (Dietrich, 1938) have asymmetric, concentric
costae that are adaptive to the efficient rocking motion during
burrowing and to securing the valves in the sediment. Oblique
costae are adaptive to both efficient burrowing and shell reinforce
ment (Checa and Jimnez-Jimnez, 2003). The Late Jurassic-Early
Cretaceous diversification of the Order Trigonioida (Stanley, 1978,
fig. 29) coincided with the diversification of bivalves having
oblique ribs, including Trigonioida (Checa and Jimnez-Jimnez,
2003, text-fig. 6).
A dilemma arises from the late Mesozoic diversification
of trigoniids followed by their near extinction at the end of the
Cretaceous. Generalized patterns of first and last appearances of
genera from the Treatise of Invertebrate Paleontology (Cox et al.,
1969) (Fig. 3) show that the maximum diversification occurred
in the Late Jurassic followed by a two-step reduction of genera
by the end of the Early Cretaceous and finally at the end of the
Late Cretaceous. A number of environmental changes during the
Cretaceous seemingly affected bivalve evolution and specifically
trigoniid evolution (Checa and Jimnez-Jimnez, 2003). Predation
in marine benthic communities increased because of the diver
sification of gastropods and vertebrates, the Mesozoic Marine
Revolution (Vermeij, 1977; Harper and Skelton, 1993; Checa
and Jimnez-Jimnez, 2003). Trigoniids developed thick, strongly
ornamented valves during the approximately 100 m.y. interval
from Late Jurassic to end Cretaceous that may have been adap
tive to defending against predators. Such features are some of the
potential defensive strategies to predation (Harper and Skelton,
1993). So, if the trigoniids became better adapted to increasing
predation pressure, why did the clades become extinct during and
at the end of the Cretaceous? A number of abiotic environmental
crises recurred during the Cretaceous and affected other benthic
paleocommunites (Scott, 1995) that may have stressed the trigoniid
populations. Perhaps a combination of factors, such as repeated
bottom-water anoxia on the shelves and increasing temperature
and salinity weakened these semi-infaunal, gaping suspension
feeders so that their reproductive abilities were impaired. Detailed
studies of clades in narrowly delimited stratigraphic intervals and
facies would test such speculation.
PALEOBIOGEOGRAPHY
The Texas outcrops were part of the Comanchean Shelf that
bordered the northern continental shelf of the proto-Gulf of Mexico
(Fig. 1). The Sonoran outcrops were part of the Bisbee Basin and
its southern Sonoran Shelf (Fig.1). The biota comprises the Carib
bean Province and has both endemic and cosmopolitan species.
The trigoniid taxa represent a mainly eastern Pacific assemblage
found in both Argentina and British Columbia. The oyster taxa are
common in Temperate Europe and the Mediterranean Province.

10
The caprinid rudists are mainly endemic as are some ammonites,
although others occur in South America, Antarctica, and even in
Europe.
The southwestern North American trigoniid assemblage
shares taxa with the west coast of North America. A distinct Early
Cretaceous trigoniidassemblage occurs in California, Oregon and
British Columbia of the northeastern Pacific region. Species of
Yaadia Crickmay range from Valanginian to Maastrichtian, Qua
dratotrigonia (Dietrich, 1938)is in Albian to Turonian strata, and
Litschkovitrigonia Saveliev is Turonian (Saul, 1978); Pterotrigonia
Van Hoepen ranges from Albian to Campanian (Jones, 1960).
In British Columbia, in addition to Yaadia, the genera include
Apiotrigonia Cox, Pterotrigonia, Quoiecchia Crickmay, Steinmanella
Crickmay, Heterotrigonia Cox, and Columbitrigonia Poulton (Poul
ton, 1979). Some of these genera are known mainly from the west
coast of North and South America, some also occur in eastern and
central Asia, some are found in South Africa and Europe, and three
are cosmopolitan. Steinmanella species are diverse and common in
Argentina from Valanginian to Hauterivian deposits of a muddy
shelf and shoreface (Lazo, 2003).
REPOSITORIES
Specimens are housed at the U.S. National Museum, Smith
sonian Institution in Washington, D.C. (USNM); at the Texas
Memorial Museum (TMM, UT, BEG); at the Estacin Regional
del Noroeste, Instituto de Geologa, UNAM, Hermosillo, Sonora,
Mxico (ERNO); at the Instituto de Geologa, Ciudad Universita
ria, Mexico City (IGM); and at the Los Angeles County Museum
of Invertebrate Paleontology (LACMIP); and the Museum of
Paleontology, The University of Michigan, Ann Arbor, Michigan
(UMMP). Roemers (1852) type specimens are at the Palontolo
gisches Institut der Rheinischen Friedrich-Wilhelms-Universitt,
Bonn. Specimens in the personal collection of the author are
indicated by RWS.
A taxonomic nomenclatural issue requires clarification. In
her 1937 dissertation Marion Whitney named new species from the
Glen Rose Formation. However, she understood that this did not
constitute publication as defined in Article 8 of the the Interna
tional Code of Zoological Nomenclature (1999). Subsequently she
published 30 names in the Journal of Paleontology (1952a, 1952b).
In 2002 the names in Whitneys dissertation (1937) were included
in a compilation of Cretaceous bivalves in Texas (Akers and Ak
ers, 2002). Unfortunately, the inclusion of Whitneys unpublished
names does not constitute publication because those names are
nomina nuda, that is, they do not conform to Articles 11 and 13 of
the Code. Consequently, none of Whitneys unpublished names
are included in the following synonomies.
SYSTEMATIC PALEONTOLOGY
Subclass Pteriomorphia Beurlen, 1944
Order Arcoida Stoliczka, 1871
Superfamily Arcacea Lamarck, 1809
Family Cucullaeidae Stewart, 1930
Genus Cucullaea Lamarck, 1801
Type species: Cucullaea auriculifera Lamarck, 1801, SD
Children, 1823
Subgenus Idonearca Conrad, 1862
Type species: Cucullaea tippana Conrad, 1858, SD Dall, 1898
Cucullaea (Idonearca) terminalis Conrad, 1857
Figures 4 C-E, J
Cucullaea (Idonearca) terminalis Conrad, 1857, p. 148, pl. 4, figs. 2a,
b; Hill, 1893, p. 26; Cragin, 1893, p. 174-175; Adkins, 1928, p. 88;

Cucullaea Terminalis
gracilis Cragin,
var. recedens
1893, p.Cragin,
173; Adkins,1928,p.88;
1894;
Cucullaea recedens Cragin, Adkins, 1928, p. 88; Perkins, 1961, p.
14, 19, 24; Scott, 1970, p. 60.
Types: Syntypes of C. terminalis, USNM 9890, both internal
molds; the specimen figured here is the lectotype. Numerous
topotypes are in TMM. Syntypes of Cucullaea terminalis var.
recedens are USNM 32681. Syntypes of C. gracilis are UT 17298
and UT 21700.
Type locality: Conrad did not list the locality of the syn
types; Cragin (1894, p. 3) stated that the species comes from
the alternating beds of the Upper Member of the Glen Rose
Formation in Texas; Adkins (1928) listed it as being widespread
in central Texas. Cragins specimens of C. gracilis were collected
in Burnet and Gillespie counties, Texas, from the alternating
beds of the Upper Member of the Glen Rose. The type locality
of Cucullaea terminalis var. recedens is the lower part of the Kiowa
Formation in Clark County, southern Kansas.
Original diagnosis: Ovate-triangular, ventricose; buccal
margin in the cast almost direct; anal side produced, cuneiform,
extremity angulated; basal margin straight; summit profoundly
prominent.
Revised diagnosis: Triangular valves, beak and umbo
steeply inclined towards anterior margin, beak very near anterior
margin, the ratio of the beak distance to total length about 3.0,
angle between posterior umbonal ridge and anterior margin
between 50-65.
Description: Moderate to large bivalve shell, triangular,
beak inclined toward anterior margin and close to it; umbonal
ridge arched, posterior surface steep, posterior margin truncated,
short, posterior-ventral corner very abruptly rounded. On interior
casts interior umbonal carina well defined between inhalant and
exhalent siphonal areas. Interior valve surface has numerous fine
radial mantle striae. Dentition consists of 3-5 long subparallel
lateral teeth, the dorsal ones with hooked ends parallel to 10-12
vertical short denticles.
Discussion: Conrad differentiated C. terminalis from Cucul
laea vulgaris Morton (1830) by its more prominent beak that is
more anteriorly placed than in C. vulgaris. Cucullaea gracilis (Cragin,
1893, p. 173) is defined from internal casts from the Upper Member
of the Glen Rose Formation. Cragin differentiated it from C.
terminalis by its more compressed and slender beaks, although its
shape and size are similar and its beaks are inclined towards its
anterior margin. It is a junior synonym of C. terminalis.
The Middle-Upper Albian Cucullaea terminalis recedens (Cra
gin, 1894) occurs in the Walnut Formation, the Marys Creek Marl
Member of the Goodland Formation, and the Kiamichi Formation
in Texas (Perkins, 1961), and throughout the lower Upper Albian
Kiowa Formation in Kansas (Scott, 1970). Cragin distinguished
C. recedens from C. terminalis by the formers more central beak
placement and by their different stratigraphic positions. Although
Stephenson (1952, p. 63) placed the Kiowa specimens in terminalis,
Perkins and Scott used recedens for the younger specimens without
making full comparisons. Most specimens from the Glen Rose and
the Walnut-Goodland-Kiamichi-Kiowa formations are internal
molds, and the ornamentation is unknown. The beaks of the Glen
Rose specimens tend to be slightly closer to the anterior margin
than the Kiamichi specimens, about 26 percent of the total length
compared to about 30 percent; this small difference hardly seems
to be a reliable characteristic for species differentiation. Therefore,
the name, recedens, should be considered as a junior synonym of
terminalis (Table 2).
The Mediterranean Albian species, Arca (Idonearca) orientalis
Douvill (1916), also has an anteriorly inclined umbo with beaks
close to the anterior margin (Abbas, 1961, pl. 2, fig. 7) so that an

11

FIGURE 2. Aptian-Albian biostratigraphy in Sonora, Arizona, and Texas. Solid range bar is based on Young (1974, 1986); dashed bar is based on
Stoyanow (1949); dash-dot bar is based on Lawton et al. (2004) and new data herein. Numbers are keyed to the names of the ammonite zones above.
The bars show the ranges of the species not of the zones. Two radiometric dates bracket the Rancho Bufalo Member of the Mural Formation (Gonzlez
Len et al., 2007); ages in Texas are interpolated.

internal cast would be very similar to those of C. terminalis.

Occurrence: The range of C. terminalis is Upper Aptian
Dufrenyoia justinae Zone to lower Upper Albian Hysteroceras
varicosum Subzone of the Mortoniceras inflatum Zone. C. terminalis
occurs in the Cow Creek Limestone and is common in the Salenia
interval and the alternating beds of the Glen Rose Formation.
C. gracilis is reported from the Cow Creek, Upper Member of the
Glen Rose, Walnut, and Comanche Peak formations based on
specimens cataloged at TMM. It is also common in calcareous
shale of the Kiamichi Formation in north-central Texas where it
occurs in place with its posterior margin oriented up. It is also in
the Kiowa Formation in Kansas.
Cucullaea (Idonearca) inermis Gabb, 1869
Figures 4A, B, F-I, K-M
Cucullaea inermis Gabb, 1869, p. 271-272, pl. 3, figs. 18, 18a;
Cucullaea (Idonearca) inermis Gabb, Almazan-Vazquez, 1990, pl.
3, figs. 8, 8b;
Arca gratiota Hill, 1888, p. 133, pl. 14, figs. 2, 2a;
Cucullaea gratiota Hill, 1893, p. 25; Cragin, 1893, p. 173-174; Ad
kins, 1928, p. 88 (as C. gratiola);
Idonearca stephensoni Stoyanow, 1949, p. 63-64, pl. 8, figs. 9-12; pl.
9, fig. 1.

Types: Holotype of C. inermis unknown. Holotype of C.

gratiota is USNM 145630. Syntypes of C. stephensoni are LACMIP
10664, 10665, 10666, 10667, 10668 (= UCLA 28510, 28511, 28512,
28513, 28514, respectively), Natural History Museum of Los
Angeles County, California (Saul, 1991). Specimen LACMIP 10664
is here designated as lectotype, and the other specimens become
paralectotypes. Topotype specimens are UT21701 and 1999TX2.
Type localities: Gabbs specimens of C. inermis are from
the Albian section in the Cerro las Conchas near Arivechi,
Sonora, Mexico (Almazan-Vazquez, 1990). The type locality of C.
gratiota is Plaster Bluff (= Gypsum bluff of Hill, 1888) southwest
of Murfreesboro, Pike County, Arkansas in the Lower Albian De
Queen Formation (Ham and Landrey, 1983). The type locality of
C. stephensoni is Ninety One Hills, Cochise County, Arizona, from
Stoyanows Lowell Formation, Pacheta Member, division 9g,
Lancha limestone bed in the Kazanskyella spathi Zone, uppermost
Aptian.
Material: One disarticulated RV, TMM1994TX2, Hensel
Sandstone, Edge Falls section, Texas. Three articulated external
casts from the Tuape Shale, Mural Limestone, Santa Ana section,
Sonora. One partial LV silicified is from the Upper Member of the
Mural Limestone in southeastern Arizona.
Diagnosis: Subquadrate valve with erect beak, angle of
umbonal ridge to anterior margin 80-90; few fine radial costae

12
TABLE 2. Comparison of selected Comanchean species of Cucullaea. Beak length is measured from anterior margin to midpoint of
beak. All measurements in mm. T indicates type specimens.
Inermis-1
Species

Gabb
Number

55.9
Height

76.2
Length

50.8
Width

L/HRatio
1.36

Beak Length

L/B Ratio

Umbonal Angle

Inermis-2
Inermis-3
Inermis-4
Gratiota
Gratiota-T

Gabb
Gabb
pl. 3-8b
USNM
UT
Almazan
21701
145630

53.3
38.1
72
53
55.5
36

63.5
45.6
75
67
45
71.0

50.8
33.0
-30
40

1.19
1.20
1.04
1.24
1.28
1.25

27
20

2.6
2.25

90
90
85

Gratiota
Gratiota
Gratiota
Gratiota

UT-Hensel
UT - Hensel
UT - Hensel
ERNO

80
~55

67
75
65
~63
107

33
~17

1.3
1.1

25
30
28
18

2.7
2.5
2.3
3.5

Gratiota

ERNO

71

100

35

1.4

Stephensoni-T
Stephensoni-T

LACMIP 10664
LACMIP 10668
10665

44.8
36.0
38.2

57.8
~49
46+

~23
~15

1.3
1.3
~1.3

25
19

2.3
~2.6
2.4+

Terminalis-T
Gracilis-T
Terminalis-T

USNM 9890
USNM
UT
17298
9890

34
40

50
61
42

22
32
16

1.5
1.5
1.1

14

>3
3.0

Gracilis-T

UT 21700
UT Glen Rose
UT Glen Rose
UT Glen Rose
USNM
UT
- Goodland
32681

31

24

1.5

29
36

45
57
59
55
57
36
50

1.2
1.4

15
15
15
12
15

3.8
3.9
3.7
3.8
3.0
3.3

37
44.5
53

72
46
47
72
57.5

1.3
1.4

23
14
13
12
20

3.1
3.3
4.8
3.6

Terminalis
Terminalis
Terminalis
Terminalis
Recedens-T

Terminalis
Terminalis
Recedens

UT - Kiamichi
UT - Kiamichi
RWS-Kiamichi

1.0-1.2

14
20

16
25
21

on exterior antero-dorsal valve, and median carina on postero

umbonal slope.
Description:Shell equivalved, subequilateral, beaks inflated
extending beyond hinge margin, slightly opisthosogyrate; cardinal
margin broadly arched forming an angle of 155 with the broadly
curved anterior valve margin, antero-ventral corner more tightly
curved, ventral margin broadly curved, postero-ventral corner
very narrowly curved at an angle of 45-60 with the nearly straight,
steeply inclined posterior margin that forms an angle of 140 with
the postero-cardinal margin. The umbonal profile viewed from the
anterior is a broadly expanding spiral with beaks forming a right
angle with the cardinal margin. Valve surface ornamented by fine
to moderate concentric growth lines; antero-dorsal valve surface
with four to six radial costae of varying widths and heights that
converge and weaken towards beak (Cragin, 1893); rarely faint
radial costae are present on the posterior flank anterior to the
umbonal ridge (Stoyanow, 1949, p. 8, fig. 11); radial ornamentation
may not be preserved on external casts. Umbonal ridge narrowly
rounded forming an angle with cardinal margin slightly greater
than 90, postero-umbonal slope wide, expanding from beak,

80

55

>3

60
50

65
50
60

divided by a low rounded median carina across which growth

rings deflect slightly towards commissure, postero-umbonal sur
face on either side of median carina slightly concave; the median
carina on the interior valve surface forms the myophore ridge that
extends from umbo to posterior margin and impresses a groove
into the internal cast.
Hinge structure consists of cardinal area and taxodont
dentition; ligament area a narrow triangular flat surface below
beaks with three chevron ligament grooves; two types of teeth,
six to eight short central transverse denticles and three longer
subhorizontal anterior and posterior lateral teeth, the dorsal
most has tips deflected forming short transverse denticles (see
Stoyanow, 1949, pl. 8, figs. 10, 12). This pattern of dentition and
ligament is similar to those of Cucullaea terminalis variety recedens
(Cragin, 1893), Cucullaea blanpiedi (Stephenson, 1952), and Cucul
laea vulgaris (Morton, 1830).
Some specimens in the collections of the Texas Memo
rial Museum are distorted by compaction and are much higher
than long and are trigonal; a compressed internal cast of these
specimens would be very similar to Cucullaea terminalis Conrad

13
possesses fine radial striae on the anterior slope and an umbonal
carina like C. gratiota. C. castilloi has about the same form and
length/height ratio but it does not have the umbonal carina, and
its posterior slope is ornamented by fine radial striae. According
to Cragin (1905), C. catorcensis is very massive and ventricose; it is
more inflated than C. gratiota, and it does not posses an umbonal
carina or anterior striae.
Several species of Cucullaea have been described from both
the Aptian and the Albian sections in England (Woods, 1899-1903).
C. fittoni (Pictet and Campiche, 1866) has radial ornament on the
anterior margin and a carina on the umbonal slope similar to C.
gratiota; it is about half the size of the North American species,
but it is similar in most other ways.
Occurrence: The occurrence of C. inermis in the Tuape Shale
Member of the Mural Formation extends its range into the lower
part of the Upper Aptian in the Dufrenoyia justinae Zone. Its upper
limit is above the Lower Albian Douvilleiceras mammillatum Zone
in Texas and Arizona and with Middle Albian species in Cerro
las Conchas (Almazan-Vasquez, 1990). C. stephensoni (Stoyanow,
1949) is from the uppermost Aptian ammonite zone of Kazanskyella
spathi (Young, 1974; Scott, 1987).
Order Pterioida Newell, 1965
Suborder Pteriina Newell, 1965
Superfamily Pteriacea Gray, 1847
Family Bakevelliidae King, 1850
Genus Gervillaria Cox, 1951
Type species: Modiola? alaeformis J. Sowerby, 1819
Gervillaria alaeformis (J. Sowerby, 1819)
Figure 5A-C

FIGURE 3. Plot showing numbers of first and last occurrences of

Trigonioida genera (from Cox and others, 1969).

(Table 2). One articulated specimen was collected in place in shale

oriented with its posterior up in the upper part of the Tuape Shale
at the Santa Ana section.
Discussion: Gabbs species, C. inermis, seems to have been
overlooked by subsequent workers. However, topotype specimens
are well illustrated by Almazan-Vasquez (1990). It is indistinguish
able from Hills species. Although the type specimens of Cucullaea
(Idonearca) gratiota are internal casts, and the types of C. stephensoni
are whole shells, the shapes and proportions of both species are
similar (Table 2). C. inermis and C. stephensoni have faint radial cos
tae on the posterior flank (Stoyanow, 1949, pl. 8, fig. 11). However,
these costae are not consistently present, and this species should
be treated as a junior synonym of C. inermis.
Cucullaea inermis differs from Cucullaea terminalis (Conrad,
1857, syntypes USNM 9890; topotype UT 21702) by the subcentral
position of its beaks and its more upright umbo compared to the
anteriorly inclined umbo of C. terminalis.
Cucullaea comanchensis (Hill, 1893; Adkins, 1928, p. 88) in
the basal Glen Rose in north-central Texas is higher than long,
its length/height ratio being 0.93 compared to a range from 1.0
to 1.5 for C. inermis (Table 2). This ratio, of course, can be altered
by compaction and fracturing of the shells. C. inermis also has a
distinct carina on the umbonal slope and several radial striae on
the anterior slope, which seem to be absent in C. comanchensis.
Cragin (1905) described three species of Cucullaea from the
Upper Jurassic Malone Formation in West Texas (Albritton, 1938):
C. transpecosensis Cragin, C. castilloi Cragin, and C. catorcensis Cas
tillo and Aguilera. According to Cragin (1905), C. transpecosensis
(UT 21704) is more ventricose than C. gratiota, although it also

Modiola? alaeformis J. Sowerby, 1819, p. 93, pl. CCLI;

Gervillaria alaeformis (J. Sowerby, 1819), Dhondt and Dieni, 1988,
p. 18-19, Pl. 4, figs. 2-4 (provides detailed synonmy);
Gervillia heinemani Stoyanow, 1949, p. 64, pl. 9, figs. 6-7;
Gervillia cholla Stoyanow, 1949, p. 64-65, pl. 10, fig. 1;
Gervillia rasori Stoyanow, 1949, p. 65, pl. 10, figs. 2.
Types: The type of Gervillaria alaeformis is lost (Woods, 1905,
p. 83). The holotype of G. heinemani is LACMIP 10669 (UCLA
28515); the holotype of G. cholla is LACMIP 10670 (UCLA 28516);
and the holotype of G. rasori is LACMIP 10671 (UCLA28517), and
the paratype is LACMIP 10672 (UCLA 28518).
Type locality: The type of Gervillaria alaeformis was collected
in the Lower Greensand at Sandown, Isle of Wight, England. G.
heinemani is from the Saavedra Member, unit 7i, Lowell Formation
at Ninety One Hills. G. cholla and G. rasori are from the next over
lying Cholla Member of the Lowell Formation, Upper Aptian at
the same locality.
Material: Two nearly complete specimens with up to 8090% shell and partly encrusted by serpulids and an oyster are
from the Tuape Shale, Santa Ana section, sample 3-19-05-1 (ERNO
8501, 8502).
Discussion: Stoyanow (1949) distinguished Gervillaria heine
mani from G. alaeformis by its nearly straight umbonal arch, and
by its very short posterior sulcus below the beak. These features
are subject to preservational differences and are not considered
to be of taxonomic significance. G. cholla is a smaller species that
has a small umbonal angle, a nearly straight umbonal arch, and
a reduced posterior sulcus. However, the beak and the posterior
umbonal surface are eroded. The types of G. rasori are even smaller
than those of G. heinemani and G. cholla; the umbonal arch is less
curved than in G. cholla, and the umbonal angle is similar; these
differences are minor and insignificant; they are a result of

14
preservation and, perhaps, intrapopulation variability.
Dimensions: Most Arizona and Sonoran specimens are
distorted to some degree by compaction and are incomplete
because of weathering, so some dimensions may not be accurate
(Table 3).
Occurrence: Gervillaria alaeformis is widespread in the
Hauterivian to Aptian of Tethyan and southern Temperate Realms
of Europe, southwest Asia, North and East Africa, and Argentina
(Dhondt and Dieni, 1988, text-fig. 7), and now in Arizona and
northwestern Mexico.
Superfamily Pectinacea Rafinesque, 1815
Family Plicatulidae Watson, 1930
Genus Plicatula Lamarck, 1801
Type species: Spondylus plicatus Linne, 1758, SD Schmidt,
1818
Plicatula (Plicatula) sonoranesis n. sp.
Figure 5D-G
Types: Holotype ERNO 8522 articulated shell; paratypes
ERNO 8523, 8524, 8525.
Type locality: The Cerro La Ceja Member of the Mural
Formation at the Rancho Bufalo section, sample 1-8-06-4.
Diagnosis: A moderate-sized, ovate shell with strong
growth squamae and discontinuous, irregularily-spaced plicae.
Description: Moderate-sized ovate shells, nearly equilateral,
inequivalved, RV more inflated than LV, which may be slightly
concave or reflexed; dorsal margins forming nearly a right angle
or slightly less than 90, anterior and posterior margins broadly
curved, ventral margin rounded. Umbonal arch on RV very low,
broad; attachment scar at beaksmall. No auricles. Ornamented by
coarse concentric growth squamae and short, radial plicae discon
tinuous between growth bands, stronger and more consistently
developed on RV than on LV.
Valve interior is unknown. The valve height is 35.8 to 45.7
mm, length is 30.8 to 39.4 mm, and width of articulated shells is
12.1 to 15.6 mm.
Comparisons: These Sonoran specimens are larger than
Plicatula incongrua Conrad (1857), which is a triangular shell
with strong costae that extend the valve height; the type locality
of P. incongrua is at El Paso, Texas, in Albian strata. The Sonoran
specimens are also larger than the Upper Albian Plicatula denton
ensis Cragin (1893) and Plicatula senescens Cragin (1894), and the
concentric growth rings are stronger, the radial ornament is more
discontinuous, and the margins are not thickened. Plicatula subgur
gitis Bse (1910) from the Upper Albian at El Paso, Texas, is smaller
than the new species and has continuous plicae. Plicatula gurgitis Pictet
and Roux (Woods, 1899-1903) from the Upper Albian at Folkestone,
England, is triangular, and the plicae are continuous from the beak,
forming small spines crossing the growth squamae.
Occurrence: The genus Plicatula ranges from Middle Triassic
to Holocene and is cosmopolitan. The new species is known from
the Sonoran Shelf of the Bisbee Basin in Upper Aptian strata.
Suborder Ostreina Frussac, 1822
Superfamily Ostreacea Rafinesque, 1815
Family Gryphaeidae Vyalov, 1936
Subfamily Exogyrinae Vyalov, 1936
Tribe Exogyrini Vyalov, 1936
Genus Aetostreon Bayle, 1878
Type species: Gryphaea latissima Lamarck, 1801, 1819; SD
Douvill, 1879
Aetostreon latissimum (Lamarck, 1801)

TABLE 3. Dimensions of studied specimens of Gervillaria in

mm.
Height

Length Width

Angle

Alaeformis X

Heinemani
Cholla
Rasori
Rasori

~130
~117
~80
91+

83
~62
~58
~55

53
36
25
~34

65
50
50
compressed

122
82

85
>50

~35
~28

~65
~45

Species

Number

LACMIP10669
LACMIP 10670
LACMIP 10671
LACMIP 10672
Alaeformis ERNO 8501
Alaeformis ERNO 8502

Figure 6A-F
Gryphaea latissima Lamarck, 1801, 1819, p. 199;
Gryphaea couloni Defrance, 1821, p. 534;
Gryphaea sinuata Sowerby, 1822, pl. LXI, fig. 13;
Exogyra sinuata (Sowerby), Woods, 1913, p. 395-404, text-figs.
194-214; (synonomies to 1908);
Exogyra couloni (Defrance), Weaver, 1931, p. 229-231, pl. 19, figs.
88-91;
Exogyra couloni dOrbigny var. leufuensis Weaver, 1931, p. 228
229, pl. 19, figs 93a, b;
Aetostreon latissimum (Lamarck, 1801), Pugaczewska, 1975, p. 5153, pls. VII-IX; Dhondt and Dieni, 1988, p. 38-40, pl. 8, figs. 1a-7,
pl. 9, figs. 1-6 text-fig. 18; complete synonomy;
Exogyra americana Marcou var. quitmanensis Cragin 1893, p. 183184, pl. 31;
Exogyra quitmanensis Adkins, 1928, p. 110; Stanton, 1947, p.36,
pl. 26, figs. 1, 2; pl. 27, fig. 5; pl. 28, fig. 6;
Exogyra aquila (Brongniart), Stanton, 1947, p. 31-32, pl. 19, fig.
13; pl. 20, figs. 1, 12.
Type: Types unknown. Plesiotypes of E. quitmanensis are
USNM 103205a-d and 103206a-b. Specimens figured by Stanton
are USNM 9609 and 103194.
Type locality: The types of Gryphaea latissima Lamarck are
from the Aptian of France. The types of G. couloni Defrance are
from Lower Cretaceous strata near Neuchtel, Switzwerland. The
types of G. sinuata Sowerby are from Aptian beds in England.
The type of E. aquila is from the Upper Aptian at Perte du Rhne,
France (Woods, 1913, p. 404). The type locality of E. quitmanensis
is in the Upper Aptian to Lower Albian Quitman Formation in
the Quitman Mountains, Hudspeth County, Texas.
Material: Illustrated specimens are from the lower part of
the Tuape Shale in the El Ocuca section, sample 4-11-05-7, speci
men ERNO 8504, and in the Rancho Bufalo section, sample 9-16-F,
specimens ERNO 8505, 8506.
Description: Medium to large exogyrid, quite inequivalved;
LV very convex, elongate, gently curved, with abruptly rounded
umbonal ridge, angle between anterior and posterior slopes less
than 90; beak in smaller specimens in a tight 360 spiral, in larger
specimens the beak is blunt; anterior slope flat, posterior slope
concave from umbo and flattening out toward posterior-ventral
margin; LV ornamented by smoothly curved to wrinkled growth
lines, in some specimens few, very faint, low rounded, radial
costae on anterior slope; RV ovate to gently arcuate, slightly con
cave on posterior side and slightly reflexed on anterior margin,
beak spiral indistinct, ornamented by irregular growth lines.
Muscle scar large, semicircular, small oval pedal muscle pit below

15

FIGURE4. A, B, F-I, K-M, Cucullaea inermis Gabb; A, B, holotype of C. gratiota (Hill, 1888) USNM 145630, internal mold, LV and posterior views, X1.1.
C, E, Cucullaea terminalis var. recedens Cragin, 1894, syntype USNM 32681, X1.1, C, RV exterior, E, LV interior. D, Cucullaea terminalis Conrad, 1857,
syntype USNM 9890, LV,X1.3. F-I, Cucullaea (Idonearca) inermis RV, posterior, dorsal, lateral, and anterior views respectively, X0.8, UTMM 1999TX2. J,
Cucullaea terminalis Conrad, LV, X0.75, Tuape Shale, Santa Ana section, sample 3-19-05-1, specimen ERNO 8503; K-M, C. inermis two specimens from
the Glen Rose Formation, Wise County, Texas (BEG 21701, BEG 248-T-21), K, L, RV of articulated specimen, lateral and anterior views respectively,
X1.1, M, LV, posterior view, X1.2.

16

FIGURE 5. A-C, Gervillaria alaeformis Sowerby, RV, LV respectively, specimen ERNO 8501, X0.8, C, G. alaeformis RV specimen ERNO 8502, X1.0; both
specimens from Tuape Shale, Mural Formation, Santa Ana section, Sonora, sample 3-19-05-1. D-G, Plicatula sonoraensis n. sp., Cerro La Ceja Member
at Rancho Bufalo (sample 1-8-06-4); D, E, holotype ERNO 8522, RV and LV; F, G, paratype ERNO 8523, RV and LV. Scale bar = 1 cm.

beak; resilifer a shallow groove beneath beak of LV.

Discussion: Woods (1913) included a wide variety of mor
photypes under the name of Exogyra sinuata (Sowerby), ranging
from the large trigonal, ovate, and subquadrate forms to the arcu
ate forms that are higher than long. This highly variable taxon is
known from the Valanginian Claxby Ironstone, the Hauterivian
Tealby Clay, the Lower Aptian Atherfield Clay, and the Lower
Albian Lower Greensand (Woods, 1913).
Cragin (1893) erected Exogyra americana Marcou var. quit
manensis for the basally produced or caudate form from
outcrops south of the Quitman Mountains. He believed it to be
intermediate between Exogyra americana and Exogyra ferox. Stan
ton (1947) described this species fully and differentiated it from
E. americana.
Weavers (1931) description of Exogyra couloni is quite simi
lar to Woods description of Exogyra sinuata as noted by Weaver.
This species is common in Neocomian to Aptian strata in the
Neuquen Basin of Argentina. Weaver (1931) distinguished Exogyra
couloni dOrbigny var. leufuensis by its smaller size; many speci
mens also have a bilobed umbonal ridge. Because these arcuate
specimens are like the arcuate forms of A. latissimum, this taxon
is synonomized with the latter taxon. This morphotype occurs in
lower Neocomian strata in the Neuquen Basin of Argentina.
Imlay (1939) reported common oysters in Canyon Santa
Rosa east of El Tigre in northeastern Sonora that he compared
with Exogyra latissima variety aquila (Brongniart). He differentiated
them from E. quitmanensis by the weak umbo and angular umbonal
ridge. Associated ammonites are of the Upper Aptian Dufrenoyia
justinae (Hill) [formerly texana Burckhardt] Zone in the Cow Creek
Limestone in Texas.
Dhondt (1982) maintained the distinction between Aetostreon
couloni, an elongate and narrow oyster compared with Aetostreon
latissimum, which is a more subtrigonal, subquadrate, or ovate
morphotype. But, later Dhondt and Dieni (1988) synonymized the
two. In Spain, A. couloni occurs in Neocomian strata, and A. latis
simum is in Aptian beds (Dhondt, 1982). A. latissimum also occurs
in uppermost Valanginian to lower Hauterivian strata in eastern
Sardinia with Ceratostreon boussingaulti (Dhondt and Dieni, 1987).
These taxa are members of a northern Tethys Mediterranean to
western European province.
Dimensions: El Ocuca section, 4-11-05-1, ERNO8504: height
(H) = 14.4 cm, length (L) = 8.0 cm, width (W) = 5.2 cm; Rancho
Bufalo section, 9-16-F, ERNO 8505: H = 8.3 cm, L = 5.2 cm, W = 2.6
cm; ERNO 8506: H = 6.3 cm (incomplete), L= 4.7 cm, W= 3.0 cm;
ERNO 8507: H = 6.2 cm, L = 4.2 cm, W = 2.7 cm. Stanton (1947)
reported a specimen 20 cm high, 15.7 cm long and 9 cm wide, and
a second specimen 14.2 cm high, 9.8 cm long, and 6.5 cm wide.
Occurrence: Aetostreon latissimum is common in the Tethyan
area of Europe and North Africa to the Caucasus in Valanginian to
Albian strata. E. quitmanensis is in the Quitman Formation and in
the lower part of the Bluff Formation (Underwood, 1963) with the
uppermost Aptian to Lower Albian foraminifer Orbitolina (Mesor
bitolina) texana. In New Mexico, Exogyra quitmanensis occurs with
Ostrea franklini Coquand, and Ctenostreon cumminsi Stanton, 1947
in beds that elsewhere have ammonites of the uppermost Aptian
Kazanskyella spathi Zone (Lucas and Lawton, 2000). Stanton (1947)
reported E. aquila from the Upper Aptian Travis Peak Formation
at Cow Creek in northwestern Travis County, Texas.
Genus Amphidonte Fischer de Waldheim, 1829
Type species: Amphidonte humboldtii Fischer de Waldheim,
1829
Amphidonte obliquata (Pulteney, 1813)
Figure 7A-F

17
Chama obliquata Pulteney, 1813, May, p. 108, figs. 8, 8*, 8** of
plate;
Chama haliotoidea J. Sowerby, 1813, June, p. 67, pl. XXV, figs. 1-5;
Chama conicaJ. Sowerby, 1813, June, p. 69-70, pl. XXVI, figs. 2-4;
Exogyra conica (J. Sowerby, 1813), Woods, 1913, p. 407-413, figs.
215-242;
Exogyra obliquata (Pulteney), Cox, 1940, p. 125, pl. 7, fig. 8;
Amphidonte conica (Sowerby, 1813), Freneix, 1972, p. 85-87, pl. 5,
figs. 1a-b, 2a-b; provides older synonomies;
Amphidonte conicum (Sowerby), Sobetski, 1977, p. 158-159, pl.
14, figs. 13, 14;
Exogyra obliquata (Pulteney, 1813), Castell and Cox, 1975, pl. 55,
figs. 4, 5.
Types: The types of Chama conica Sowerby are in the British
Museum of Natural History.
Type locality: C. obliquata Pulteney is from the Upper
Greensand in Dorset, England. The types of C. conica Sowerby
are from the Upper Albian Upper Greensand near Warminster
in England.
Material: Fourteen free, articulated shells from the Cerro
La Ceja Formation at Rancho Bufalo, sample 1-8-06-4; numerous
small, juvenile specimens are also in the Fronteras Member at
Rancho Bufalo (sample 1-29-06-1) and in the Los Coyotes Member
at Tuape (sample 1-28-06-6).
Description: Shell small, ovate to elongate, very inequi
lateral, inequivalved, tending to be taller than long; beak and
umbo of LV coiled at least 360; umbonal ridge angular to sharply
rounded forming an acute angle to nearly 90 between anterior and
posterior slopes; concentric growth lines distinct wrinkles; a few
specimens have short, closely spaced radial striae either on part
of anterior LV margin or on posterior surface (e.g., Woods, 1913,
figs. 240-242). RV ovate, slightly concave, beak tightly coiled up
to 360, distinct growth wrinkles. Interior commissure margins of
both LV and RV are lined with very fine chomata to ventral margin;
adductor scar ovate, located close to postero-dorsal margin.
Discussion: Shells from the Upper Greensand (Upper
Albian) in southern England have fine chomata completly en
circling the LV margin, and the ovate adductor scar is situated
very close to the posterior margin at about mid height (Castell
and Cox, 1975, pl. 55, fig. 5). Woods (1913) and Freneix (1972) al
lowed for wide polymorphic variation within the species concept,
which seems fitting.
Amphidonte [first named as Chama] obliquata (Pulteney,
May 1813) is the senior synonym of Amphidonte [Chama] conica
(J. Sowerby, June 1813). However, Freneix (1972) concluded that
A. obliquata was a nomen oblitum because it had not been used
until 1940, when Cox resurrected it and placed it in Exogyra.
Furthermore, Exogyra obliquata was used in 1975 by Castell and
Cox. According to the International Commission on Zoological
Nomenclature (1999) Article 23.9, two conditions must be met in
order to continue usage of the younger name, i.e., A. conica. First,
the senior synonym must not have been used as a valid name
after 1899 (Art. 23.9.1). Clearly this condition is not met. Second,
the junior synonym has been used as the valid name in at least 25
publications by ten or more authors in the immediately preceding
50 years and encompassing a span of not less than 10 years (Art.
23.9.2). This condition seems to have been met because between
1813 and 1937 at least 37 authors used the name E. conica according
to the synonymy lists of Woods (1913) and Freneix (1972). Unless
action is taken by the Commission to establish A. conica as the
valid name, A. obliquata should be used (Art. 23.9.3).
Dimensions: The specimens from the Cerro La Ceja Member
of the Mural Formation at Rancho Bufalo average 3.4 cm high,
2.4 cm long, and 1.6 cm wide (seven specimens). The Upper

18
Albian specimens illustrated by Woods (1913, Figs. 215-226) are
approximately the same size and proportions, but the specimens
from the Cenomanian Chalk Marl (Woods, 1913, Figs. 227-231)
are significantly larger.
Occurrence: Woods (1913) reported the species from the
Aptian to Cenomanian beds in England. Amphidonte obliquata
(=conica) and its polymorphs are reported from the Albian to
Lower Cenomanian in Europe, Caucausus, Middle East, and North
Africa (Freneix, 1972) and the Ukraine (Sobetski, 1977). This report
extends its geographic range to southern North America.
Genus Ceratostreon Bayle, 1878
Type species: Exogyra spinosa Matheron, 1843
Ceratostreon tuberculiferum (Koch and Dunker, 1837)
Figure 6G-I
Exogyra tuberculifera Koch and Dunker, 1837, p. 54, pl. vi, fig.
8; Woods, 1913, p. 404-406, pl. LXI, figs. 7-11; provides older
synonomy;
Exogyra lancha Stoyanow, 1949, p. 66-67, pl. 10, figs. 4-6;
Ceratostreon tuberculiferum (Koch and Dunker), Dhondt and
Dieni, 1988, p. 42.
Types: The type of C. tuberculiferum is unknown. The holotype
of E. lancha is LACMIP 10676 (UCLA 28502), paratypes LACMIP
10677 and 10678 (UCLA 28503, 2850).
Type locality: Koch and Dunker evidently selected a RV
from Neocomian strata as the type specimen (Woods, 1913). The
types of E. lancha were collected in the Ninety One Hills, south
eastern Arizona, Lowell Formation, Pacheta Member, division 9g,
now Lower Member, Mural Formation, Upper Aptian.
Diagnosis: A small to moderate sized exogyrid with a dis
tinct arched umbonal carina with few, low radial costae on the
dorsal-anterior margin.
Description: Small to medium exogyrid, inequilateral, quite
inequivalved; LV very convex to flattened, elongate, gently curved,
with abruptly rounded umbonal carina, angle between anterior
and posterior slopes less than 90 in younger stage and tends to
be greater in later stage; beak in smaller specimens in a tight 360
spiral, in larger specimens the beak is overgrown; anterior slope
flat, posterior slope concave from umbo and flattening out toward
posterior-ventral margin; LV ornamented by smoothly curved to
wrinkled growth lines, on anterior marginal slope in younger stage
few, very faint, low rounded, radial striae, and in later stage short
radial plicae developed on the commissure, seven in the holotype
of E. lancha; RV ovate to gently arcuate, slightly concave on pos
terior side and slightly reflexed on anterior margin, beak spiral
indistinct, ornamented by irregular growth lines and regular rows
of fine pustules that may be the imprint of attachment substrate;
chomata developed on interior valve margin; adductor scar ovate
and slightly offset toward posterior margin.
Discussion: Many Nineteenth Century authors con
fused Ceratostreon tuberculiferum with Ceratostreon boussingaulti
(dOrbigny), as is evident in the synonomy lists of both Woods
and Dhondt and Dieni. However, C. tuberculiferum has fewer and
weaker radial costae on the anterior margin than C. boussingaulti.
Stoyanow (1949) described Exogyra lancha as small to medium,
elongate and crescentic having on the left valve an obtuse
angulation. He also noted plicae on the anteroventral margin of
the umbo. He differentiated E. lancha from Ceratostreon boussin
gaulti (dOrbigny), Exogyra minos (Coquand), Ceratostreon tuber
culiferum (Koch and Dunker) (formerly Exogyra), and Exogyra
laevigata Sowerby by the rounded carina of the left valve of E.
lancha versus the angular carina of the other species. Dhondt and
Dieni (1988) synonymized E. minos with C. boussingaulti.

Dimensions: Measurements of E. lancha: holotype LAC

MIP 10676: height 6.1 cm, length 3.8 cm, width 1.0 cm; paratypes
LACMIP 10677: height 1.6 cm, length 1.0 cm, width 0.5 cm, and
10678: height 2.1 cm, length 1.6 cm, width 0.7 cm.
Occurrence: C. tuberculiferum ranges from Valanginian
to Upper Aptian in northwestern Europe (Woods, 1913) and in
Valanginian rocks on eastern Sardinia (Dhondt and Dieni, 1988).
Exogyra lancha is from the Lower Member of the Mural Limestone
in Arizona.
Family Ostreidae Rafinesque, 1815
Subfamily Ostreiniae Rafinesque, 1815
Genus Ostrea Linn, 1758
Type species: Ostrea edulis Linn, 1758
Ostrea riograndensis Stanton, 1947
Figure 7G-I
Ostrea riograndensis Stanton, 1947, p. 24, pl. 5, figs. 1, 2.
Types: Cotypes USNM 103183a-b.
Type Locality: The cotypes were collected from the Upper
Aptian-Lower Albian Quitman Formation along the trail between
Quitman Arroyo and Hot Springs south of the Quitman Moun
tains, southern Hudspeth County, Texas. An associated oyster
is Aetostreon latissimum (= Exogyra quitmanensis Cragin). Other
specimens were collected from the Glen Rose Formation near
Decatur, Wise County, Texas.
Material: Mainly articulated shells were collected in Sonora
from the Fronteras Member (sample 1-28-06-3), from Cerro La Ceja
Member (sample 1-8-06-3), and from the Mesa Quemada Member
at Ocuca (sample 12-08-05-5).
Diagnosis: Valves are large, thick, subovate, dorsal margin
narrowly rounded, ventral margin broadly rounded, slightly re
flexed, anterior and posterior margins are subparallel; RV flat to
very slightly convex, LV slightly convex, exterior of both valves
with coarse to weak growth lines spaced irregularly.
Description: LV is gently arched in umbonal area and flat
tens ventrally, slightly reflexed; RV is gently convex to slightly con
cave towards umbo. Adductor muscle outline is ovate to arcuate,
slightly longer than high (~1.3X), close to ventral margin; resilifer
on RV is broad, wider than high, bounded by anterior and pos
terior bourrelets; no umbonal cavity; no chomata along margins.
Commissure margins smooth, arched with valve convexity.
Dimensions: Height ranges from 10.9 to 4.8 cm; length from
to
3.7
6.8
cm; width of articulated specimens from 3.1 to 1.8 cm.
Stantons specimen USNM 103183a is 10.8 cm high and 7.7 cm
long. A specimen from the Carbonate Hill Member of the U-Bar
Formation in Doa Ana County, New Mexico is more than 16 cm
high and 10 cm long (New Mexico Museum of Natural History and
Science locality 4262, specimen P-30038: Lucas and Estep, 2000).
Occurrence: This species is uncommon in the Cerro La Ceja
Member at the Rancho Bufalo and Tuape sections and in the Mesa
Quemada Member at the Ocuca section. It also occurs in the up
permost Aptian to lower Albian in southwestern New Mexico.
Subclass Palaeoheterodonta Newell, 1965
Trigonioida Dall, 1889
Suborder Myophorellina Cooper, 1991
Superfamily Myophorellacea Kobayashi, 1954, emended
Cooper, 1991
Family Myophorellidae Kobayashi, 1954
Subfamily Steinmanellinae Cooper, 1991

19

FIGURE 6. A-F, Aetostreon latissimum (Lamarck), Tuape Shale Member, Mural Formation, Sonora. Figures A, B, LV, RV respectively, X0.6, El Ocuca
section, sample 4-11-05-7, specimen ERNO 8504; C, D, LV, RV respectively, X1, Rancho Bufalo section, sample 9-16-F, specimen ERNO 8506; E, LV,X0.8,
Rancho Bufalo section, sample 9-16-F, specimen ERNO 8505; F, LV, X0.6, Rancho Bufalo section, sample9-16-F, specimen ERNO 8507. G-I, Ceratostreon
tuberculiferum (Koch and Dunker); G, LV, LACMIP 10678; H, LV LACMIP 10677; I, LV, holotype LACMIP 10676. Scale bar = 1 cm.

20
Genus Steinmanella Crickmay, 1930
Type species: Trigonia holubi Kitchin, 1908
Steinmanella vyschetzkii (Cragin, 1893; emend. Stoyanow, 1949)
Figure 10O-R
Trigonia vyschetzkii Cragin, 1893, p. 215; Cragin, 1897, p. 816-817;
Cragin, 1905, p. 56-58, pl. 8, figs. 1, 2; pl. 9, figs. 1-3; Adkins, 1928,
p. 118-119, includes additional synonomies; Imlay, 1940, p. 402,
pl. 54, fig. 9; Stoyanow, 1949, p. 70-74; Saul, 1978, p. 8, table 1;
Trigonia sp. Stoyanow, 1949, p. 72;
Trigonia maloneana Stoyanow, 1949, p. 74;
Trigonia maloneana variety Stoyanow, 1949, p. 74-75;
Steinmanella vyschetzkii (Cragin), Saul, 1978, p. 8, 11;
Trigonia vyschetzkii Cragin, Leanza, 1993, p. 41.
Types: USNM 28967: three specimens, USNM 28967
lectotype of T. vyschetzkii selected by Stoyanow (in Cragin, 1905,
pl. 8, fig. 2; Fig. 10R; Stoyanow designated this specimen
#2) (plaster cast UCLA 48291); paralectotype USNM 28967 is
Stoyanows specimen #1 (plaster cast UCLA 48290; (Fig. 10O);
paralectotype USNM 29020 is Stoyanows holotype of T. malonea
na (plaster cast UCLA 48749; Fig. 10P, Q); paralectotype USNM
28967 is Stoyanows specimen #4 (plaster cast UCLA 48291).
Additional specimens identified by Cragin are in the Texas
Memorial Museum, UT 19736.
Type locality: Malone Mountains, 1.5 miles northeast of
Malone Station (3115W, 10531N), USGS locality 506, about 5.5
miles west of Sierra Blanca, Hudspeth County, Texas.
Diagnosis: A large, subquadrate moderately convex valve;
carinae tuberculate along most of their length; escutcheon crossed
by irregular nodate costellae; area divided by a median groove,
crossed by narrow, discontinuous costallae; flank costae gently
concave anteriorly, topped by large, low, rounded nodes.
Description: Valve subquadrate, moderately inflated; beaks
subterminal, incurved, slightly opisthogyrate; anterior and ventral
margins broadly curved semicircular, posterior margin truncated
slightly inclined into straight dorsal margin. Flank a very gently
arched surface, ornamented by about fifteen gently curved, noded
costae that merge with the marginal carina at an acute angle; the
costal nodes are large, rounded, elongated subparallel to growth
lines, 12 to 16 nodes per costa; intercostal spaces flat, widening
to anterior and ventral margins. Valve crossed by numerous fine
growth lines.
Area triangular, quite wide, flat to slightly concave; mar
ginal carina a row of large rounded nodes elongated parallel to
growth lines; median carina developed in the anterior part of the
area as a row of very small, inconsistently developed nodes that
fade posteriorly and bounded dorsally by a shallow sulcus that
persists to the posterior margin; the nepionic area ornamented by
rows of small nodes, and on the adult portion by wide rounded
costellae that cross carinae at nodes.
Escutcheon wide, elongate, ornamented by straight to
curved costellae slightly oblique to commissure, topped by low
nodes; escutcheon carina a row of rounded nodes elongated with
growth lines. Lunate ligamental fossette is directly posterior to
beak, about one-fourth the length of the dorsal margin.
Discussion: Stoyanow (1949) used rather minor characters
to separate Cragins (1893, 1905) types of T. vyschetzkii into three
additional taxa: Trigonia maloneana, T. maloneana var., and Trigonia
sp. I agree with Saul (1978) in re-uniting these specimens into
one slightly variable species, Steinmanella vyschetzkii. Cragins
syntype specimens are incompletely preserved and altered by
compaction. Stoyanow (1949, p. 73) selected as lectotype of T.
vyschetzkii Cragins (1905) specimen in pl. 8, fig. 2 (USNM 28967;

Fig. 10R). Stoyanow (1949, p. 74) defined T. maloneana (Fig. 10P,

Q; Cragins specimen, pl. 9, fig. 1, 1905, USNM 29020) by (1) its
...very concave, nearly crescentic dorsal margin with rapidly
rising umbo and posterodorsal extremity; (2) the angle between
the area and the escutcheon is more obtuse than in T. vyschetzkii
and T. guildi; (3) the area is concave posteriorly; (4) the costellae
on the escutcheon and area are broadly V-shaped; and (5) the shell
anterior is strongly inflated. Stoyanow (1949) differentiated T.
maloneana variety (Cragins specimen in pl. 9, fig. 2, 1905, USNM
28967) by the fine tubercles on the posterior part of the area and
by oblique costellae and tubercles on the escutcheon. Stoyanow
(1949) set aside Cragins specimen (1905, pl. 8, fig. 1; pl. 9, fig. 3,
same specimen, also USNM 28967) and four other specimens be
cause: (1) the anterior part of the area is horizontal, (2) the marginal
carina is close to the dorsal commissure, (3) the flanks are strongly
inflated, and (4) the escutcheon costellae have tubercles.
Steinmanella vyschetzkii is most similar to Quadratotrigonia
guildi, but Stoyanow differentiated vyschetzkii from Q. guildi by its
somewhat longer shape, its more gently curved ventral margin,
and its posterior flattening. He noted that S. vyschetzkii has 10
flank ribs compared with 11 to 12 on Q. guildi, and the nodes of
S. vyschetzkii are larger than those on Q. guildi. However, Cragin
(1905) counted up to 15 costae on S. vyschetzkii. The median carina
of nodes in S. vyschetzkii is shorter than in Q. guildi.
The holotype of Trigonia dumblei Stoyanow (1949, p. 77-78,
pl. 12, fig. 4; UTMM BEG19723; Figs. 7A, B) is from the Upper
Jurassic Malone Formation one mile northeast of Malone, Hud
speth County, Texas, in the Malone Formation (Cragin, 1905;
Albritton, 1938). This specimen was one of two that Cragin used
to define Trigonia taffi (Cragin, 1893, p. 214). Stoyanow correctly
recognized that Cragins type specimens of T. taffi included two
species. Stoyanow defined Trigonia dumblei on Cragins specimen
No. 511/214 (now TM BEG 19723) and designated it as holotype
(Fig. 8A, B). This specimen consists only of the dorsal part of a
LV with matrix covering part of the umbo and the junction of the
area with the umbo as illustrated by Stoyanow (1949, pl. 12, fig.
4). Stoyanow compared T. dumblei with T. resoluta and differenti
ated T. dumblei by its greater number of areal costellae and by
rather minor differences in the posterior inclination of its flank
ribs. However, because the anterior flank ornamentation of the
valve is not preserved, this specimen cannot be assigned to either
Steinmanella or Quadratotrigonia, although Nakano (1960, p. 265;
1968, p. 34) and Saul (1978, p. 7) placed it in Quadratotrigonia. Its
presence in the Malone Formation with specimens of S. vyschetzkii
tempts one to compare it with this Jurassic species. If this specimen
is identified as vyschetzkii, then dumblei becomes a junior synonym.
Thus, it is recommended that the taxon, T. dumblei, be restricted to
its holotype and not used for any other specimens.
Steinmanella vyschetzkii is quite similar to the earliest Cre
taceous Argentinian species Steinmanella transitoria (Steinmann)
and Steinmanella quintucoensis (Weaver). Cragin distinguished his
species by its less elongate shape and its finer ornamentation on
the area and escutcheon. Leanza (1993, p. 41) noted that vyschetz
kii has a different shape than quintucoensis, its marginal carina is
tuberculated, and its antecarinal sulcus is less well developed.
Occurrence: Cragins specimens of Steinmanella vyschetzkii
(1905) are from outcrops in low hills about 1.5 miles east of the
Malone Mountains. These interbedded sandstones, limestones
and clays are mapped as the Malone Formation (Albritton, 1938;
Barnes, 1968), and yield Kimmeridgian ammonites. Imlay (1940)
reported S. vyschetzkii from the Upper Jurassic La Casita Forma
tion near Las Cuevas, Durango and the La Gloria Formation in
Sierra Jimulco, Coahuila, Mexico. Imlay (1945) also identified a
specimen in core of the Upper Jurassic Cotton Valley Formation
in southern Louisiana as T. sp. aff. T. vyschetzki (USNM 103627).

21

FIGURE 7. A-F, Amphidonte obliquata Pulteney; middle part of Cerro La Ceja Member, Mural Formation at Rancho Bufalo, sample 1-9-06-4; LV and
RV pairs: A, D, ERNO 8516, B, E, ERNO 8517, C, F, ERNO 8518. G-I, Ostrea riograndensis Stanton; G, ERNO 8526, LV, Cerro La Ceja Member, Tuape
section, sample 1-28-06-3; H, I, ERNO 8527, LV exterior and interior, Fronteras Member, Rancho Bufalo section, sample 1-08-06-3. Scale bar = 1cm.

22
This specimen is a very incomplete external cast of a LV with flank
and areal ornament, and its specific assignment is uncertain.
Family Vaugoniidae Kobayashi, 1954
The family Vaugoniidae is distinguished from other families
of Myophorellacea by nodate, chevron-shaped flank costae near
the umbo (Cooper, 1991). All members of Myophorellacea have
the trigonian hinge grade (Boyd and Newell, 1997).
Subfamily Quadratotrigoniinae Kobayashi, 1954
This subfamily is separated from the Subfamily Vaugoniinae
by its large areal carinae that divide into rows of nodes or even
become obsolete in the adult stage and by transverse growth
rugae on the area in the mature stage (Cooper, 1991). This mainly
Tethyan subfamily is convergent with Steinmanellinae but differs
by its V-shaped, umbonal costae and its strongly ornamented area
(Nakano, 1968; Cooper, 1991).
Genus Quadratotrigonia Dietrich, 1933
Type species: Trigonia nodosa J. de C. Sowerby, 1826
Quadratotrigonia taffi (Cragin, 1893)
Figure 8C
Trigonia taffi Cragin, 1893, p. 214; Cragin, 1905, p. 10, footnote b;
Adkins, 1928, p. 120; Stoyanow, 1949, p. 76-77;
Trigonia gordoni Whitney, 1952b, p. 702-703, pl. 87, fig. 2;
Quadratotrigonia taffi (Cragin), Saul, 1978, p. 8; Saul, 1991, p. 65;
Lucas and Estep, 2000, p. 83.
Types: Holotype TMM BEG 19723(509) disarticulated RV,
length estimated about 82 mm, height about 78 mm, width about
24 mm; width of area at posterodorsal corner about 18 mm; para
type 21758(509). The two cotypes of Trigonia gordoni Whitney are
UMMP 53655 and UMMP 53656.
Type locality: Bluff Mesa, USGS locality 509, three miles
southwest of Sierra Blanca Station, Hudspeth County, Texas.
Diagnosis: A subquadrate shell having a curved anterior
margin and a narrow area; flank costae closely spaced and topped
by closely spaced small nodes.
Description: Valve subquadrate, moderately inflated, beaks
subterminal, slightly opisthogyrate; anterior margin forms a semi
circle with ventral margin; posterior margin apparently truncate;
dorsal margin straight; area without median sulcus.
Flank a gentle curved surface ornamented by strongly
arched rows of nodes concave posteriorly, nodes are low rounded,
ovate becoming elongate near ventral and anterior margins;
interspaces very narrow, much less than the width of the costae;
towards anterodorsal margin on the umbo the costae are V-shaped
to strongly arcuate; a line bisecting the arc of the costae is steeply
inclined to the dorsal margin and subparallel with the anterior
margin.
Width of area at posterodorsal corner is about 24% of valve
height; length/height about 1.05. Escutcheon of holotype is cov
ered by matrix. Area partly exposed showing ornament of fine
costellae, at least two connecting to each node on the marginal
carina; marginal carina a row of low rounded nodes.
Discussion: The overall shape of Q. taffi is very similar to
that of Q. mearnsi; however, the area of Q. taffi is much narrower
than that of Q. mearnsi, about 24% of its height versus about 43%.
Furthermore, the nodes on the flank costae of Q. taffi are much
smaller, and the interspaces are much narrower than in Q. mearnsi.
The costae of Q. taffi are also more strongly arcuate than on the
other species.
Whitney (1952b) suggested that T. gordoni was closely related
to Q. taffi, but could not decide because taffi was not illustrated. In

comparison with the holotype of Q. taffi, T. gordoni shows the same

style of nodate flank costae in which the interspaces are about as
wide as the costae; in addition, the proportions of the valve and
the area are close to those of Q. taffi. However, Saul (1978, p. 9)
referred T. gordoni to Q. mearnsi because the ornament on the area
of both taxa ...fades toward the posterior valve margin.
Saul (1978, 1991) placed taffi in the genus, Quadratotrigonia,
which is justified by both the areal and flank ornament. Although
only the adult portion of the area is exposed on the holotype, its
fine, closely spaced costellae are more similar to those of Quadra
totrigonia nodosa (Sowerby) than to any of the steinmanellids, in
which the adult costae are much coarser. Also, the flank costae of
Q. taffi have a very oblique to arcuate bend bisected by a line to
the postero-ventral corner imparting a chevron pattern. The costae
of steinmanellids tend to be broadly arcuate and widely-spaced
ridges topped by nodes.
Occurrence: Quadratotrigonia taffi is reported at Bluff Mesa
about 5 km southwest of Sierra Blanca, Hudspeth County, Texas.
Presumably the species occurs in the Bluff Formation, which is
equivalent with the lower part of the uppermost Aptian to Lower
Albian Glen Rose Formation in central Texas (Underwood, 1963).
Underwood listed Trigonia mearnsi and Trigonia stolleyi from the
Bluff in the Eagle Mountains. Q. taffi is also found at Red Bull
Canyon at the southern end of the Quitman Mountains (Cragin,
1905). The holotype of T. gordoni comes from the lower part of
the Glen Rose Formation at Cow Creek, Burnet County, Texas,
which is approximately coeval with the stratigraphic position
of taffi in the Bluff Formation and within the range of Orbitolina
texana Roemer. Q. taffi is reported in the Carbonate Hill Member
of the U-Bar Formation in the East Potrillo Mountains, Doa Ana
County, southwestern New Mexico (Lucas and Estep, 2000). This
unit is correlated with the Kazanskyella spathi Zone and co-occurs
with Orbitolina texana.
Quadratotrigonia guildi (Stoyanow, 1949)
Figure 8D-G
Trigonia guildi Stoyanow, 1949, p. 75, pl. 12, figs. 1-2;
Quadratotrigonia guildi (Stoyanow), Saul, 1978, p. 8, table 1;
Jacques-Ayala, 1990, p. 72; Saul, 1991, p. 63;
Trigonia resoluta Stoyanow, 1949, p. 76, pl. 12, fig. 3;
Quadratotrigonia resoluta (Stoyanow), Saul, 1978, p. 8, table 1;
Saul, 1991, p. 65.
Trigonia saavedra Stoyanow, 1949, p. 78, pl. 13, fig. 5;
Quadratotrigonia saavedra (Stoyanow), Saul, 1978, p. 8, table 1;
Saul, 1991, p. 65.
Types: T. guildi: UCLA28520 (Stoyanows No. 91205); para
type 91413; T. resoluta: holotype UCLA 28521 (91418); T. saavedra:
UCLA 28522 (91333); location of un-illustrated paratypes 91320,
91321, 91334, unknown.
Type locality: T. guildi is from the Cholla Member bed 6a,
Lowell Formation, Ninety One Hills; T. resoluta is from the Cholla
Member bed 6a, Lowell Formation, Ninety One Hills; T. saavedra
is from the Saavedra Member, division 7n, Lowell Formation,
Ninety One Hills, Cochise County, Arizona.
Material: Two nearly complete disarticulated RV and LV
from the Hensel Sandstone at Edge Falls TM1994TX7-8. Poorly
preserved specimens from the Tuape Shale at the Santa Ana sec
tion are tentatively assigned to this species (sample 3-19-05-1).
Other specimens are in the Canova Member at Cordon Caloso
sample 4-21-05-1.
Diagnosis: A subovate to subquadrate shell having a
broadly curved anterior margin; flank costae strong and topped
by large nodes; area with median sulcus.

Description: Valve subovate to subquadrate, slightly

inflated, beaks terminal, opisthogyrate. Anterior margin very
gently curved, inclined posteriorly relative to dorsal hinge 10 to 20
degrees; ventral margin broadly rounded, inclined into truncated
posterior margin; dorsal margin relatively straight.
The escutcheon poorly exposed, tends to be narrow and
tuberculate. The area is trianglar, widening posteriorly, and its
surface is moderately inclined from the dorsal margin towards the
ventral margin. Marginal carina low, rounded, tends to be topped
with nodes in the more adult portion; in the nepionic portion
carina crossed by angular costae with posteriorly oriented Vs.
Median carina is a row of low conical nodes merging anteriorly
with marginal carina near beak. Median sulcus (supramedian
furrow of Stoyanow) is shallow, directly dorsal to median carina.
Escutcheon carina is a row of low rounded nodes merging with
umbo. Carinal nodes tend to diminish in size near the posterior
corner. Area is ornamented by fine costellae oriented ventral
dorsally between marginal and escutcheon carinae; on nepionic
portion costellae widen and are the same size as the flank costae;
posteriorly the area costellae are much finer than the flank costae
and more numerous than the carinal nodes; on some specimens
the costellae become obscured and undeveloped.
Flank in nepionic portion is ornamented by 3-4 angular
costae, steeper dorsally than ventrally, with a V flexure pointed to
wards postero-ventral corner; adult portion of flank is ornamented
by strongly arcuate to V-shaped costae topped by large nodes
tending to be rounded on antero-ventral margin and elongated
towards postero-ventral corner on dorsal margin.
Discussion: Stoyanow (1949) distinguished T. resoluta from
T. guildi by ...the areal costellae resolved into tubercles in the
anterior part of the outer area. However, this slight difference is
within the variation of the areal ornamentation, and it rarely is
observed on most specimens that have matrix around the umbo.
He also used the relative spacing of the flank costae and the size
of their nodes to differentiate these species. These differences are
difficult to quantify and appear to be in the range of variability.
T. saavedra was defined by its smaller, subquadrate outline and
the postero-ventral inclination of the areal costellae relative to the
escutcheon (inner) carina. However, this orientation is present
in the types of T. guildi (Stoyanow, pl. 12, fig. 1) and T. resoluta
(Stoyanow, pl. 12, fig. 3).
Occurrence: This species and its synonyms occur in the Up
per Aptian Cholla and Saavedra members, beds 7n and 6a, Lowell
Formation, now known as the Lower Member, Mural Formation
in Arizona and Texas.
Quadratotrigonia mearnsi (Stoyanow, 1949)
Figure 8H
Trigonia mearnsi Stoyanow, 1949, p. 78-79, pl. 12, figs. 5, 6; pl. 13,
figs. 1-4;
Quadratotrigonia mearnsi (Stoyanow), Saul, 1978, p. 8, table 1;
Quadratotrigonia cf. Q. mearnsi (Stoyanow), Jacques-Ayala, 1990,
p. 71-72, lamina 1, fig. 1;
Myophorella mearnsi (Stoyanow), Leanza, 1993, p. 30.
Types: Holotype UCLA 28523 (GU46; pl. 13, fig. 1); para
types UCLA28524 (GU42; pl. 13, fig. 4), 21525(GU45; pl. 13, fig. 3),
28526 (GU47; pl. 12, figs. 5, 6), and 28527 (GU50; pl. 13, fig. 2).
Type locality: Perilla Member, bed 2b, Lowell Formation,
Guadalupe Canyon, Cochise County, Arizona.
Material: One articulated complete specimen is from the
Lower Member, Mural Formation, Sierra Anibacachi, nine miles
southwest of Agua Prieta, Sonora, Mexico, collected by E.R. War
zeski; length 93 mm; height 79 mm; width of articulated valves
about 41 mm. A second articulated, nearly complete specimen is

23
from the Lower Member, Mural Formation, Mexican Saddle Horn
east of Douglas, Cochise County, Arizona, collected by H. Lser;
length (posterior margin incomplete) about 105 mm; height 90mm;
width of articulated valves 55 mm. One nearly complete articulated
specimen and three fragments are from the Tuape Shale Member
at El Pimiento section sample EPI-1-79. Another articulated but
poorly preserved specimen is from the Lower Member at Ninty
One Hills, Arizona.
Diagnosis: Asubquadrateshell having a broadly curvedan
terior margin; flank costae widely spaced topped by large elongate
nodes; area with median carina of large nodes, no median sulcus.
Description: Valve is large, subquadrate, moderately inflat
ed, beaks subterminal, very slightly opisthogyrate; anterior margin
merges with ventral margin in a semicircle; posterior margin is
truncate, forming an obtuse angle with the straight dorsal margin.
Length/width ratio is about 1.15. Width of area at posterodorsal
corner to median carina is about 43% of the valve height.
Escutcheon is a narrow rectangular area ornamented by
growth lines subparallel with commissure and sparse nodes;
ligamental fossette is lanceolote, about one-fourth the length of
the margin directly posterior to beaks.
Area is a flat triangular surface, ornamented by fine growth
lines deflected slightly across median carina, and irregularly ar
ranged small nodes on the anterior portion posterior to umbo;
marginal carina is a row of rounded nodes elongated with growth
lines, angle between carina and dorsal margin about 40 degrees;
median carina is a row of small, low rounded nodes that tends
to disappear towards posteroventral corner; escutcheon carina is
a row of large, high rounded nodes elongate with growth lines,
tending to be smaller in nepionic portion.
Flank is a gently arched surface with fine concentric growth
lines; costae are rows of strong subrounded nodes that become
elongate near commissure margin; in nepionic stage first several
costae are angular ridges steeper towards beak that extend onto
area as costellae (Stoyanow, 1949, pl. 12, fig. 6; pl. 13, fig. 3); next
several costae in posteroventral direction are V-shaped rows of
nodes with apex towards posteroventral margin; the final six-eight
costae are curved from ventral margin to marginal carina, convex
posteriorly; interspaces are as wide as or somewhat wider than
costae and subparallel or slightly widening ventrally.
Discussion: Q. mearnsi is very similar in overall shape and
ornamentation to S. vyschetzkii, but differs in subtle but distinct
ways: (1) the areal median carina of Q. mearnsi continues nearly
to the posterior margin, whereas in S. vyschetzkii it fades out; (2)
Q. mearnsi has no median sulcus on the area, but it is distinct
in S. vyschetzkii; (3) the width of the area from median carina to
posterodorsal corner is greater in Q. mearnsi than in S. vyschetzkii,
about 43% of the valve heigth versus about 27%; and (4) the ju
venile costae of Q. mearnsi change from small ridges to V-shaped
rows of small nodes, but in S. vyschetzkii the juvenile costae seem
to be rows of small nodes.
Q. mearnsi differs from Q. guildi by its subquadrate anterior
margin compared to the posteriorly inclined anterior margin of Q.
guildi, and the flank costae of Q. mearnsi change from V-shaped to
curved much earlier in the growth history than in Q. guildi.
Leanza (1993) compared Q. mearnsi to the Argentinian Titho
nian species Myophorella schulzi Leanza and differentiated them by
the more subquadrangular shape of Q. mearnsi, by its larger size,
and by the strong tubercles of the marginal carina.
Occurrence: Q. mearnsi ranges from Upper Aptian to Lower
Albian and is present in the Lower Member of the Mural Forma
tion or the Perilla Member of Stoyanows Lowell Formation at
numerous localities in southeastern Arizona and northern Sonora.
It is also present in the Quitman Mountains at Mayfield Canyon
in the Cuchillo Formation with Trinitoceras (Stoyanow, 1949),

24

FIGURE 8. A, B, Trigonia dumblei Stoyanow, topotype, LV, BEG 19723, lateral and dorsal views. C, Quadratotrigonia taffi, RV, holotype BEG 18723.
D-G, Quadratotrigonia guildi, TMM 1994 TX7, LV lateral, anterior and dorsal views respectively; G, RV lateral view, TMM un-numbered specimen.
H, Quadratotrigonia mearnsi (Stoyanow), RV, Lower Member, Mural Formation, Sierra Anibachi, 5km SW of Agua Prieta, Sonora, RWS collection. I,
Apiotrigonia cragini (Stoyanow), LV, ERNO 2760, La Colgada Member, Tuape section. Scale bar = 1 cm.

which is a synonym of Douvilleiceras sp., Lower Albian (Casey,

1962). Quadratotrigonia cf. Q. mearnsi is known from northwestern
Sonora in the middle member of the Arroyo Ssabe Formation,
which is Upper Aptian (Jacques-Ayala et al., 1990) and at Tuape
in the Cerro La Puerta Member of the Mural Formation (Lawton
et al., 2004). Saul (1978) reported Q. mearnsi from the Alisitos
Formation in Baja California.
Superfamily Megatrigoniacea Van Hoepen, 1929
Family Megatrigoniidae Van Hoepen, 1929
Diagnosis: Commonly pyriform to subovate megatrigoni
aceans that are generally weakly inflated anteriorly; carinae lacking
or generally restricted to the nepionic stages; escutcheon smooth or
with transverse costellae; area smooth or with transverse or radial
costellae; flank costae V-shaped or oblique, weakly nodate, entire
or tuberculate (Cooper, 1991). All members of Megatrigoniacea
have the trigonian hinge grade (Boyd and Newell, 1997).
Discussion: Cooper (1989, 1991) defined three phyletic
groups within the Family Megatrigoniidae based on the ornament
of the escutcheon, the character of the flank costae, the degree of
valve inflation, and the shape of the posterior margin. The large,
pyriform shells of Megatrigoniinae Van Hoepen (1929) have a
rounded posterior margin without an escutcheon carina; anterior
flank costae are oriented differently than posterior costae. Valves
of Pterotrigoniinae Van Hoepen (1929) are generally smaller, club
shaped to ovate having a rostrate posterior margin and without
costae on the escutcheon; flank ornament is generally differenti
ated into anterior and posterior costae. Apiotrigoniinae Tashiro
(1979) has transverse costellae on the escutcheon. The Subfamily
Buchotrigoniina Leanza (1993) is defined by the same characterset
as Apiotrigoniinae except that Buchotrigoniina has more distinct
marginal and escutcheon carinae, and thus seems synonymous
with Apiotrigoniinae.
Subfamily Apiotrigoniinae Tashiro, 1979
Tribe Apiotrigoniini Cooper, 1991
Genus Apiotrigonia Cox, 1952
Type species: Trigonia sulcataria Lamarck, 1819
Apiotrigonia cragini (Stoyanow, 1949)
Figure 8I

25
wider escutcheon by the inner furrow; escutcheon ornamented by
fine costellae subparallel to postero-dorsal commissure.
Discussion: A. cragini posseses the characteristic two sets
of flank costae, the concentric costae on the umbonal and ante
rior part of the flank, and the subvertical costae on the posterior
part of the shell that form a V where they join with the apex
pointing toward the posteroventral corner. The narrow area is
delineated by the low marginal carina and median furrow. The
wider escutcheon has fine radial costellae subparallel with the
commissure margin.
Stoyanow (1949) noted that the valve of A. cragini is very
similar to that of Anditrigonia calderoni Castillo and Aguilerae,
however the escutcheon of calderoni is smooth, and the area is
ornamented by fine radial costellae (Cooper, 1991, p. 26).
Occurrence: The cosmopolitan distribution of the genus
includes western Australia, Japan, Kamchatka, western North
America, Chile, northwestern Europe, Turkestan, and South Af
rica, as well as in the Caribbean Province in Arizona. The genus
ranges from Tithonian to Maastrichtian. A. cragini is known only
in the Upper Aptian of the Bisbee Basin in Arizona and Sonora.
Apiotrigonia kitchini (Stoyanow, 1949)
Trigonia kitchini Stoyanow, 1949, p. 82, pl. 14, figs. 4-10; Saul,
1991, p. 64;
Apiotrigonia kitchini (Stoyanow), Tashiro, 1979, p. 183.
Types: Syntypes UCLA 28533 (=91449), 28534 (=91505),
28535 (=91506), 28536 (=91507), 28537 (=91603) (Stoyanows
numbers). Specimen UCLA 28536 (pl. 14, figs. 9, 10, Stoyanow,
1949) is designated lectotype and the other syntypes become
paralectotypes (pl. 14, figs. 4-8, Stoyanow, 1949).
Type locality: The Ninety One Hills, Cochise, Arizona in
the Cholla Member, division 6a of the Lowell Formation (now the
Lower Member of the Mural Formation).
Diagnosis: An Apiotrigoniid characterized by zigzag costel
lae on the posterior part of the escutcheon; in other characteristics
similar to A. cragini.
Occurrence: The species is known only in the Upper Aptian
of the Bisbee Basin.
GENUS Buchotrigonia Dietrich, 1938
Type species: Trigonia abrupta von Buch, 1839
Buchotrigonia reesidei (Stoyanow, 1949)

Trigonia cragini Stoyanow, 1949, p. 80-81, pl. 13, figs. 6-10; pl. 14,
fig. 3; Saul, 1991, p. 62;
Apiotrigonia cragini (Stoyanow), Tashiro, 1979, p. 183.

Trigonia reesidei Stoyanow, 1949, p. 83-86, pl. 14, figs. 11-14; pl.
15, figs. 1-3; Saul, 1991, p. 64-65;
Buchotrigonia reesidei (Stoyanow), Leanza, 1993, p. 57.

Types: Syntypes UCLA 28528 (=915429), 28529 (=91430),

28530 (=91434), 28531 (=91445), 28532(=91509) (Stoyanows num
bers in parens). Specimen UCLA 28531 (pl. 13, fig. 8, Stoyanow,
1949) is designated the lectotype because it displays ornament on
both the area and the flank; the other specimens become the para
lectotypes (pl. 13, figs. 6, 7, 9, 10; pl. 14, fig. 3, Stoyanow, 1949).
Type locality: Ninety One Hills, Cochise County, Arizona
in the Cholla Member, division 6a of the Lowell Formation (now
the Lower Member of Mural Formation).
Material: A single articulated specimen was collected from
the La Colgada Formation in the Tuape area in northeastern Sonora
(ERNO 2760). Dimensions of ERNO 2760: height=2.5 cm, length
= 3.3 cm, width of articulated shell = 2.0 cm. It has 20 concentric
costae that are slightly steeper on the dorsal slope.
Diagnosis: Area narrow, in mature stage divided by a median
furrow into narrow outer and inner areas and separated from the

Types: Holotype UCLA 28538 (=91049), paratypes 28539

(=91050), 28540 (=91051) (Stoyanows numbers).
Type locality: Ninty One Hills, Cochise County, Arizona,
the Lowell Formation (now the Lower Member of Mural Forma
tion).
Material: A single external cast of a disarticulated RV is
the
from
Tuape Shale at Rancho Bufalo, sample 9-16-F, specimen
ERNO 8508, and one incomplete external cast is from Santa Ana
sample 3-19-05-1 and another from the Los Coyotes Member at
El Ocuca.
Diagnosis: A Buchotrigoniid with diagonal costellae on
the area and escutcheon that meet the dorsal commissure at an
angle.
Discussion: The valve flank ornamentation of Buchotri
gonia reesidei consists of oblique costae on the umbo, and on the
anterior region of the mature part concentric costae join subver

26
tical costae on the posterior, forming a V or zigzag pattern. The
marginal carina is a distinct ridge, and the area is divided by a
shallow median furrow; a second shallow furrow separates the
area and escutcheon. In contrast to Apiotrigonia, however, the area
and escutcheon have thin diagonal costellae that meet the dorsal
commissure at an angle.
Occurrence: B. reesidei occurs in the Pacheta Member, divi
sion 9g, Lowell Formation (now the Lower Member of the Mural
Formation). It also is present in the Upper Aptian Tuape Shale at
Rancho Bufalo and Santa Ana.
Subfamily Pterotrigoniinae Van Hoepen, 1929
Diagnosis: Small to moderately large megatrigoniids,
ovate to club-shaped and posteriorly rostrate; umbones prominent,
generally with strongly incurved, opisthogyrous beaks; escutch
eon sunken, lanceolate, with transverse costellae; escutcheon
carina obsolete; marginal carina generally restricted to umbonal
region; area crossed by concentric ribs in nepionic stages, later
ribbing effaced, oblique, V-shaped or radial; flank costae oblique,
prominently tuberculate or finely crenulated, generally differentiated
into anterior and posterior sets (Cooper, 1991).
Discussion: Cooper (1989, 1991) separated the genera
of Pterotrigoniinae into two phyletically significant tribes. The
Pterotrigoniini are club-shaped, rostrate, and strongly inflated
genera having a steep anterior face and strongly opisthogyrous
beaks. Anterior ribs are coarser than posterior ribs, and the mature
area is smooth. The taxa included are Pterotrigonia Van Hoepen
(Valanginian-Turonian), Pisotrigonia Van Hoepen (Tithonian
Maastrichtian), and Notoscabrotrigonia Dietrich (Neocomian
Campanian). In contrast, the Scabrotrigoniini Cooper are lunate
to crescentic, less inflated shells having a more rounded anterior
face and weakly opisthogyrous beaks. Flank costae are generally
similar anterior to posterior, and the mature area is ornamented.
Genera included are Scabrotrigonia Dietrich (Aptian-Maastrictian),
Acanthotrigonia Van Hoepen (Coniacian-Maastrichtian), Ptilotrigo
nia Van Hoepen (Albian-Maastrichtian), Linotrigonia Van Hoepen
(Coniacian-Maastrichtian), and Arabitrigonia Nakano (Aptian). The
pterotrigoniines range from the Lower Tithonian to Maastrichtian,
and the scabrotrigoniines range from Aptian to Maastrichtian.
Cooper (1989, 1991) proposed that the Pterotrigoniini evolved
from the Subfamily Apiotrigoniinae in the Early Tithonian by
transition from Paranditrigonia Reyes and Prez to Pisotrigonia Van
Hoepen. In the Valanginian Pisotrigonia gave rise to Pterotrigonia,
which, in turn, was the ancestor of Scabrotrigonia and the Tribe
Scabrotrigoniini Cooper that appeared in the Aptian.
Tribe Pterotrigoniini Van Hoepen, 1929
Diagnosis: Shell commonly club shaped and rostrate
posteriorly, strongly inflated anteriorly. Anterior face broad,
flattened. Beaks strongly opisthogyrous; area smooth in matu
rity. Ribs coarser anteriorly than posteriorly and with a strong
tendency to separate into distinct anterior and posterior sets
(Cooper, 1989).
Genus Pisotrigonia Van Hoepen, 1929
Type species: Pisotrigonia salebrosa Van Hoepen, 1929
Diagnosis: Like Pterotrigonia but as high as long, often
extremely inflated anteriorly and with broadly flattened anterior
face. Posterodorsal margin deeply excavate. Umbones very con
spicuous, subterminal, with beaks exceptionally incurved, opis
thogyrous; escutcheon with transverse costellae, which may be
very weak or absent in early representatives; area smooth except

in nepionic stages; flank costae markedly discrepant, with gener

ally thick, robust, distant, coarsely tuberculate, anterior costae and
narrow, straight crowded, finely serrated posterior costae. Age:
Tithonian-Cenomanian (?Maastrichtian) (Cooper, 1991).
Discussion: Pisotrigonia and Rinetrigonia Van Hoepen are
synonyms (Cooper, 1989; Leanza, 1993), however, these authors
disagree as to which has priority. Van Hoepen (1929) erected
Pisotrigonia to include the type species, P. salebrosa Van Hoepen
and Trigonia ventricosa Krauss, both of which occur in Kutch, In
dia (Kitchin, 1903). On a subsequent page, Van Hoepen created
Rinetrigonia for the original specimens of T. ventricosa Krauss. Ko
bayashi and Nakano (1957) tentatively treated Rinetrigonia as the
senior synonym of Pisotrigonia but without explanation. Cooper
(1989, 1991) reviewed the history of these names and designated
Pisotrigonia the senior synonym because it has page priority.
Subsequently, Leanza (1993) interpreted Kobayashi and Nakano
to be the first revisers, by designating Rinetrigonia as the senior
synonym. So, the issue is what constitutes first reviser? The
International Code of Zoological Nomenclature (1999), Article 24,
is very specific in defining actions that constitute revision. An au
thor must state that the two names are synonyms, and must select
one as the taxon name. Recommendation 24A adds that the first
reviser should select the name that ensures nomenclatural stability
or appropriateness, but if neither name has such an advantage,
the name having page priority should be selected. Because the
action by Kobayashi and Nakano does not meet the standards
of first reviser, page priority should take precedence as clearly
stated by Cooper (1989).
Pisotrigonia amsburyin. sp.
Figure 9A-L

Pisotrigonia n.sp. in Gonzlez-Len et al., 2007, p. 22, fig. 5F.

Types: Holotype is UTTMM 1994TX16, paratypes are
UTTMM 1994TX10, 11, 12, and 15, which are disarticulated valves
either partly covered by matrix or with the rostrum broken.
Type locality: Edge Falls Nature Conservancy site, Bandera
County, Texas, in the uppermost beds of the Hensel Sandstone,
Upper Aptian.
Material: Nine partially exposed external casts from the
Fronteras Member at Rancho Bufalo, samples 1-8-06-1, 1-8-06-2,
ERNO 8528, and 1-30-06-1.
Diagnosis: Moderate-sized, ovate, valves moderately
inflated, beaks subterminal; two types of flank costae, anterior
set wide, topped by elongate tubercles and grooves that cross
intercostae, posterior set narrow and steeply inclined dorsally,
meeting commissure at oblique angle; marginal and escutcheon
carinae consisting of rows of low nodes fading posteriorly; area a
narrow arcuate surface with median sulcus, crossed by fine costel
lae; escutcheon with few widely spaced costellae.
Description: Valve ovate, longer than high, rostrate; umbo
moderately inflated, beak nearly terminal, incurved, opistho
gyrate. Anterior margin steeply curved from beak, inclination
of anterior face varies from steep to very steep depending upon
degree of inflation. Ventral margin broadly rounded, posterior
margin truncate. Dorsal margin is slightly concave.
Escutcheon narrow, elongate, concave, separated from area
by a low, noded carina. Ornamented by 13 or more narrow, trans
verse costellae that connect with carinal nodes; nepionic costellae
gently curved concave posteriorly.
Area a rapidly widening arc from beneath beak to posterior
truncation; marginal carina a distinct row of nodes that become
smaller posteriorly; median sulcus shallow; area ornamented by

27

FIGURE 9. A-L, Pisotrigonia amsburyin. sp. A, Holotype, RV lateral view, UTMM 1994 TX.16; B, same specimen rotated to show area and escutcheon;
C-F paratypes, C, LV, UTMM 1994TX15; D, RV, UTMM 1994TX11; E, F, RV dorsal and lateral views, UTMM 1994TX10, G, I, LV dorsal and lateral
views, RWS collection; H, J, K, LV anterior, lateral, and dorsal views, UTMM 1994TX12, L, Pisotrigonia amsburyin. sp. Fronteras Member, Rancho
Bufalo section (ERNO 8528, sample 1-8-06-2), Sonora.

28
narrow transverse costellae that connect with flank costae and
escutcheon costellae; low nodes present where costellae cross the
marginal and escutcheon carinae.
Flank ornamented by two types of costae: (1) anterior and
ventral costae are broadly curved from ventral to dorsal margin,
strongly oblique to growth lines, wide, rounded at ventral margin,
bearing low-relief ridges and grooves transverse to crest that con
tinue across intercostae as low striae; and (2) these taper dorsally
into narrow, straight costae at dorsal margin that are inclined
steeply forward bearing very low-relief beads on crest. Costae
about as wide as intercostae; up to six costae on nepionic stage,
steeper antero-dorsally and with nearly imperceptible nodes; up
to 13 costae on remainder of valve, nearly symmetrically rounded,
with 5-6 nodes per 10 mm length.
Discussion: Pisotrigonia amsburyin. sp. is completely un
like any trigoniid previously recognized in Texas, New Mexico,
Arizona, and Mexico. It is less inflated and more broadly ros
trate than Pterotrigonia (Pisotrigonia) coihuicoensis (Weaver) from
the Hauterivian in Nequn, Argentina (Leanza, 1993). It is also
wider posteriorly than Pterotrigonia (Pisotrigonia) windhauseniana
(Wilckens) from the Maastrichtian of Argentina (Leanza, 1993),
and has more numerous coarse flank costae. The costae of the
Upper Albian Pisotrigonia laevicosta Cooper (1989) of Zululand are
wider and smoother than those of P. amsburyi. Pisotrigonia parva
Van Hoepen (1929) from the Tithonian in India and Pisotrigonia
ventricosa (Krauss), which is common in the Valangian of southeast
Africa (Cooper, 1991), are greatly inflated species with very coarse
costae and nodes. P.amsburyi is similar to Pterotrigonia pocilliformis
(Yokohama) and Pterotrigonia hokkaidoana (Yehara) from the Lower
to middle Cretaceous of Japan (Hayami, 1975). Neither species
however, shows the dual flank costae. The escutcheon and area
of Pterotrigonia (Pterotrigonia) hottingeri Freneix (1972) from upper
Albian beds in Morocco are very similar, suggesting a convergent
relationship.
Pisotrigonia amsburyi appears to be a homeomorph of the
mainly Jurassic species plexus of Myophorella (Haidaia). The shape
and ornament of these shells are similar to Myophorella (Haidaia)
volkheimeri Leanza and Garate (1987) from the Hauterivian of
Neuqun, Argentina. The primary difference is that the flank cos
tae of P. amsburyi are oriented more steeply relative to the anterior
commissure and define abroader arc than those of M. volkheimeri.
The dorsally-tapered costal segment of P. amsburyi is narrower
and has much smaller nodes than does M. volkheimeri. Also, the
costal beads of P.amsburyi are smaller than those of M. volkheimeri.
The ornamentation of P. amsburyi is similar to that of Myophorella
dawsoni from the Middle Bajocian of Canada. However, the area
of P. amsburyi is much narrower, and its marginal carinal nodes
are much smaller than those of M. dawsoni (Whiteaves).
Specimens of the Fronteras Formation are partially exposed
as molds and casts in calcareous sandstone. The shells are more
than 4.5 cm long, more than 1.5 cm wide and more than 3 cm high.
The anterior part is strongly inflated, and the costae are much
narrower than the interspaces and very slightly asymmetric with
steeper anterior-dorsal. The costae are capped by low linear ridges
subparallel to the commissure and they continue across the costal
interspaces as thin striae. The area is not exposed. Specimens
identified as Trigonia ornata dOrbigny by Rosales-Domnquez et
al. (1995) from the middle part of the Aptian Morita Formation at
Rancho Culantrillo, northern Sonora, are similar to P. amsburyi.
T. ornata is probably a Pisotrigonia because it is produced pos
teriorly, has two styles of flank costae, and the marginal carina
fades posteriorly. It occurs in the Aptian in England, France and
Bulgaria (Dimitrova, 1974, p. 95-96, pl. 45, figs. 5-7, as Linotrigo
nia (Oistotrigonia) ornata). It differs from P. amsburyi by its wider
area. These Texas and Sonoran trigoniid specimens suggest that

the eastern Pacific biota migrated into the Bisbee Basin sometime
during the Late Aptian.
Occurrence: Pisotrigonia amsburyi is in the Upper Aptian
Hensel Sandstone in Texas and in the Fronteras Member of the
Mural Formation in the Rancho Bufalo section, northwestern
Sonora.
Tribe Scabrotrigoniini Cooper, 1989
Diagnosis: Shell lunate to crescentic and oval, weakly
inflated anteriorly and produced but not rostrate posteriorly.
Anterior face narrow relatively rounded. Beaks moderately opis
thogyrous. Area generally ornamented in maturity. Flank costae
more or less unform and equidistant in most forms (Cooper,
1989).
This tribe is separated from Pterotrigoniinae by its lunate
to crescentic shape, by its less inflated anterior, its less extended
posterior margin, its less opisthogyrate beaks, by its areal orna
ment in the mature stages, and by its uniform and equidistant flank
costae (Cooper, 1991). Three Texas species are placed in the genus,
Scabrotrigonia, because each is lunate and posteriorly truncated,
the entire area is costellate, and the flank costae are subequal.
These three species, mooreana, emoryi, and clavigera, appear to have
formed a lineage during the Aptian to Early Cenomanian in the
Caribbean Province.
Genus Scabrotrigonia Dietrich, 1933
Type species: Trigonia scabra Lamarck, 1819
Diagnosis: Shell lunate, posterodorsal margin shallowly
concave. Transverse costellae of escutcheon continuous onto or
across area, forming chevrons with the flank costae; in some, area
may become smooth in maturity (Cooper, 1989).
Discussion: Leanza (1993) treated Scabrotrigonia as a subgenus
of Pterotrigonia. He considered the narrow area ornamented by
oblique costae that form chevrons with the flank costae to be
the most diagnostic feature of Scabrotrigonia (Leanza, 1993, p. 60).
He gave the range of Scabrotrigonia as Tithonian to Maastrichtian
in North America, Argentina, and Europe.
Scabrotrigonia mooreana (Gabb, 1861)
Figure 10A-F
Trigonia aliformis Parkinson, 1811; Roemer, 1849, p. 404;
Trigonia crenulata Lamarck, 1819; Roemer, 1852, p. 51-52, pl. 7,
fig. 6; Shumard, 1853, p. 206, pl. 4, fig. 1; Hill, 1893, p. 27, pl. 3,
fig. 4; Adkins, 1928, p. 120; not Gabb, 1869, p. 269;
Trigonia mooreana Gabb, 1861, p. 176; Gabb, 1869, p. 269; Bse,
1910, p. 123-124, pl. 25, figs. 2, 9; pl. 26, fig. 2; Anderson, 1958, p.
110; Almazan-Vazquez, 1990, pl. 3, fig. 1a-c;
Trigonia stolleyi Hill, 1893, p. 26-27, pl. 3, figs. 3, 5; Adkins, 1928,
p. 120; Stoyanow, 1949, p. 88-89, pl. 15, figs. 9-11;
Trigonia wendleri Whitney, 1952b, p. 701, pl. 88, fig. 2;
Trigonia whitneyi Whitney, 1952b, p. 701-702, pl. 86, figs. 7, 8;
Pterotrigonia (Notoscabrotrigonia) whitneyi (Whitney, 1952b),
Cooper, 1989, p. 243;
Pterotrigonia (Notoscabrotrigonia) stolleyi (Hill), Leanza, 1993, p.
60.
Types: The current location of Gabbs types is unknown.
Syntypes of T. stolleyi are USNM 316193, 145634; Hills specimen
in plate 3, figure 4 (1893) is USNM 145635. Stoyanows plesiotypes
No. GU20, GU33 are curated in the Los Angeles County Museum
of Invertebrate Paleontology as LACMIP10706 and 10707 (UCLA

28545 and 28546), respectively (Saul, 1991). Whitneys holotype

of Trigonia wendleri is UMMP 53653, and the holotype of Trigonia
whitneyi is UMMP53654.
Type Locality: Gabbs specimens of T. mooreana are from
the Albian section in the Cerrolas Conchas near Arivechi, Sonora,
Mexico (Almazan-Vazquez, 1990). Hills specimens are from the
Glen Rose Formation, Plant beds on Paluxy Creek 3 mi west of
Glen Rose, Somervell County, Texas, which is in the vicinity of Dino
saur State Park. Hills internal mold (USNM 145635) is from the
bluffs of the Colorado near Bull Creek, Travis County. This area is
the type locality of the Bull Creek Member of the Walnut Formation
(Moore, 1964). Possibly the specimen was from this unit.
Material: New specimens from the Hensel Sandstone at
Edge Falls are UTTM1994TX9, -13, -14, -17, disarticulated and
articulated shells. Numerous partial specimens, some with
shell remaining, are found in Sonora in the Tuape Shale at Santa
Ana, sample 3-19-05-1, in the Los Coyotes Member at El Ocuca,
samples 4-12-05-1 and 4-12-05-1A, in the Cerro La Ceja Member at
Cerro Pimas sample 3-16-05-1, in the Canova Member at Cordon
Caloso, sample 4-21-05-CL3, and in the Tuape Shale Member at
El Pimiento samples EPI-1-79, 3-15-05-5, 3-16-05-5, and 3-17-05-6.
Specimens in the authors collection are from the Glen Rose and
Walnut formations in Texas.
Diagnosis: Valve arcuate, ovate, strongly inflated anteri
orly; escutcheon with widely-spaced fine costellae; area wide
with widely spaced, strong costellae; anterior flank costae wide,
concave anteriorly, dorsally tapering with closely spaced, low
elongate nodes.
Original descriptions: Shell elongate subtrigonal, beaks
small, terminal, incurved; anterior end slight convex, rapidly
sloping backwards; base forming a regular curve, continuous
with the anterior margin; posterior cardinal margin broadly and
slighty concave; posterior end unknown. Surface ornamented by
about twenty-five moderately large and slightly sinuous ribs, very
slightly crenulated, with broad concave interspaces. These ribs be
come obsolete on the border of the corselet, or cross it transversely
as fine lines, continuing across the central area as sharply defined,
linear ribs, in a transverse direction, slighty curved forwards at
their extremities. Corselet broad, nearly flat, with a narrow border,
marked by a deep longitudinal groove (Gabb, 1869).
Semilunate in general outline, beaks well forward and
stongly recurved; anterior and pallial margin a strong continu
ous curve; posterior portion elongated with truncated posterior
margin; cardinal area compressed. Surface marked by flexuous,
noduled costae, about twenty-two in number, narrow and high,
separated by broad intercostal areas as in T. alformis Lmk.;
depressed cardinal area bordered on its outer side by a long nar
row groove and marked by cross-ribs, flexing anteriorly (Hill,
1893).
Description: Valve outline ovate-triangular, rostrate; beaks
moderately inflated, incurved, opisthogyrate, nearly at anterior
margin. Valves strongly inflated, in dorsal view maximum width
slightly posterior of beaks, flanks taper in a wide arc to posterior
margin. In lateral view beaks curve abruptly into anterior valve
margin, anterio-ventral margin broadly arcuate to a more narrowly
arcuate ventral margin, posterior margin truncate; maximum
height about mid-flank; dorsal margin gently concave. Anterior
margin is very steep, nearly normal to commissure.
Escutcheon wide, elliptical, slightly concave; separated from
the area by distinct escutcheon carina with nodes on anterior half
and posterior half with growth lines; antecarinal sulcus shallow,
narrow, ornamented by fine growth striae. Escutcheon ornamented
by narrow beaded costae aligned approximately with flank costae
and inclined anteriorly. Area is a very narrow sulcus merging
anteriorly with escutcheon carina. Ligamental fossette is directly

29
posterior of umbo, short, not well developed.
Flank ornamented by 24 or more concentric, wide costae
bearing low nodes transverse to costae and subparallel with
growth lines; nodes stronger on anterior costae and weaker on
posterior; costae taper dorsally up to marginal carina. Intercostal
spaces much wider than costae, diverging and becoming wider
from dorsal to ventral. Costae on anterior flank tending to be
concave, midflank costae tending to be slightly flexuous with
dorsal end curved posteriorly and ventral end curved anteriorly,
posterior costae straight.
Discussion: Roemers (1849, 1852) composite molds were
collected from Lower Cretaceous rocks near Fredericksburg, Gil
lespie County, Texas. Rocks of the Trinity, Fredericksburg and
Washita groups crop out in this area, so his specimens could have
come from any of these units. Roemer (1849) first identified these
specimens as the Lower Albian Trigonia aliformis Parkinson. In his
1852 monograph Roemer changed the identification to Trigonia
crenulata Lamarck, which is a Cenomanian species that has a very
wide area and very small nodes on the costae (Woods, 1899-1903).
The ornament and narrow area of Roemers specimens are like
those of Scabrotrigonia mooreana; the ventral margin of Roemers
figure appears to be more gently rounded than that of S. mooreana.
Bse (1910) identified Roemers specimens as Trigonia emoryi and
discussed the differences.
Gabb (1869) contrasted his specimens of Trigonia mooreana
at Arivechi with Trigonia crenulata Lamarck; Scabrotrigonia mooreana
has a more slender form, its escutcheon is wider and bounded by
a deep sulcus, and the ribs are less strongly crenulate. He distin
guished S. mooreana from S. emoryi by the more elongate shape of T.
mooreana and its narrower area and escutcheon. Bse (1910) noted
that the flank costae of S. mooreana joined the marginal carina verti
cally and on S. emoryi they joined obliquely; also the escutcheon
of S. mooreana was wider and shorter than on S. emoryi.
Hill (1893, pl. 3, fig. 4) identified specimen USNM 145635 as
Trigonia crenulata but wondered if it was T. stolleyi. This specimen
is a weathered, articulated composite mold from the Colorado
River bluffs in Travis County where the Walnut Formation crops
out. Adkins (1928) considered that Hills specimen was like
Roemers specimens and differentiated it from T. emoryi by its
sparser ribs.
Whitney differentiated Roemers specimens from the con
cept of T. crenulata by the fewer number of costae on the Texas
material, their more gently concave dorsal margin, and by the
size, shape and ornament of the escutcheon. T. crenulata from the
Cenomanian of England (Woods, 1899-1903, pl. 19, fig. 14) has a
wide area unlike that of either S. mooreana or S. emoryi.
Trigonia wendleri Whitney is from the upper part of the
Lower Member of the Glen Rose Formation, the Salenia texana
interval, on the Colorado River at the west end of Lake Travis,
Travis County, Texas. This species is based on incomplete com
posite molds that show up to 20 costae on an incomplete valve;
its height measured from the umbo to the ventral margin is only
slightly less than the length.
Trigonia whitneyi Whitney is from the lower part of the Glen
Rose Formation on Cow Creek, Burnet County, Texas, and is like
S. mooreana in all critical characters. Only 20 costae are counted
because the nepionic stage is not present. It is slightly smaller and
its height is incomplete because the umbo is broken.
Both Cooper (1989) and Leanza (1993) placed Scabrotrigonia
stolleyi in the taxon Pterotrigonia (Notoscabrotrigonia), which is a
member of the Tribe Pterotrigoniini Van Hopen (1929). The shell of
this group is strongly inflated anteriorly and rostrate posteriorly;
beaks are very opisthogyrate. The mature area is smooth, and the
anterior ribs are stronger than the posterior ribs. However, the
Texas species has a lunate, less produced and less inflated shape

30

FIGURE 10. A-F, Scabrotrigonia mooreana; A, LV dorsal view syntype of T. stolleyi USNM 145634; B, LV lateral view, syntype USNM 316193; C, D,
LV dorsal and lateral views UTM 1994TX13; E, F, RV dorsal and lateral views UTMM 1994TX9. G-J, Scabrotrigonia emoryi, G, H, RV dorsal and lateral
views holotype USNM 9849; I, J, RV dorsal and lateral views, Kiowa Formation, Saline County, Kansas, latex cast, RWS collection. K, L, Scabrotrigonia
clavigera, RV dorsal and lateral views UT 17243, Washita Group, Weno Formation?, Browns Ferry, Cooke County, Texas. M, N, Scabrotrigonia emoryi
labeled as Trigonia mooreana variety roemeri (nomen nudum), LV lateral and dorsal views UT 37828. O-R, Steinmanella vyschetzkii, O, LV lateral view
paralectotype USNM 28967 (Cragins pl. 8, fig. 1); P, Q, RV dorsal and lateral views paralectotype USNM 29090 (Cragins pl. 9, fig. 1); R, LV lateral
view lectoholotype USNM 28967 (Cragins pl. 8, fig. 2). Scale bar = 1 cm.

and the flank ornament is uniform. S. mooreana, S. emoryi, and S.

clavigera are very similar, comprise a clade and should be grouped
in the same genus. S. mooreana differs from Scabrotrigonia emoryi
(Conrad) by its wider escutcheon with coarser costellae and by its
wider costae with elongate ridge-like nodes. Scabrotrigonia clavigera
has very large rounded costal nodes.
Occurrence: Scabrotrigonia mooreana ranges from uppermost
Aptian to Middle Albian. The type is from the Albian section
near Arivechi, Sonora, Mexico, where it co-occurs with species
typical of the Middle Albian Walnut Formation in central Texas:
Texigryphaea mucronata (Gabb), Ceratostreon texanum (Roemer),
and several species of Engonoceras and Metengonoceras (Alma
zan-Vasquez, 1990). Cragins (1893, p. 212) type specimens were
collected in marl below the highest bed of Gryphaea pitcheri
var. washitaensis. Hills type material is from the middle part
of the Glen Rose Formation, the plant beds three miles west of
Glen Rose, Somerville County, Texas (Hill, 1893, p. 14). This is in
the area of the present-day Dinosaur Valley State Park.
Hill listed S. stolleyi from all beds of the Trinity Division in
Arkansas and Texas (1893, p. 16). S. mooreana shells are common in
the Walnut Formation in Bosque County, Texas, and it is the abun
dant trigoniid in the Whitestone Lentil of the Walnut Formation. S.
stolleyi is reported in the upper part of the Carbonate Hill Member
of the U-Bar Formation in the East Portrillo Mountains with Q. taffi
in the K. spathi Zone (Lucas and Estep, 2000). In southern Arizona,
S. stolleyi is reported from the Saavedra and Perilla members of
the Lowell Formation (Stoyanow, 1949). In Sonora this species
is in the Cerro La Ceja, Tuape, and Los Coyotes members of the
Mural Formation. Known range is Upper Aptian to Middle Albian.
The species characterizes a partial-range zone in the upper part
of the Hensel Sandstone and lower part of the Glen Rose Forma
tion in north-central Texas with Orbitolina (Mesorbitolina) texana
Roemer (Scott, 1930).
Scabrotrigonia emoryi (Conrad, 1857)
Figure 10G-J, M, N
Trigonia emoryi Conrad, 1857, p. 148, pl. 3, figs. 2a-c; Cragin,
1895b, p. 376; Shattuck, 1903, p. 23, pl. 8, figs. 6-8; Bse, 1910, p.
121-124, pl. 24, figs. 1-5; pl. 25, figs., 1, 3, 5; pl. 16, fig. 1; Adkins
& Winton, 1920, p. 73, pl. 17, figs. 4-6; Twenhofel, 1924, p. 85, pl.
13, figs. 1-3; Adkins, 1928, p. 121;
Scabrotrigonia emoryi (Conrad), Scott, 1970, p. 70-71;
Trigonia quadalupe Bse, 1910, p. 124-125, pl. 23, figs. 11-16; Ad
kins, 1928, p. 121.
Types: Holotype USNM 9849, paratype USNM 316171. Rep
resentative specimens are KU500189,500190. Roemers specimens
are in the Paleontological Institute, Bonn, Germany. The syntypes
of T. guadalupe are IGM 364, pl. 23, figs. 13, 15, 16.
Type locality: El Paso, El Paso County, Texas along the U.S./
Mexico international border. The type locality of T. guadalupe is at
La Encantada near Placer de Guadalupe, Chihuahua, Mexico.
Diagnosis: A moderate-sized, elongate ovate valve, moder
ately inflated anteriorly; escutcheon wide with fine radial costel
lae; area narrow with closely spaced, nodate costellae; anterior
flank costae S-shaped, posterior costae straight, inclined posteri
orly, slightly tapering dorsally with closely spaced, distinct, low,
rounded nodes.
Description: Valve outline trigonal, very inflated, beaks
subterminal, distinctly opisthogyrate; anterior margin steep and
broadly curved, ventral margin more tightly curved sloping into
posterior truncation, dorsal margin concave.
Escutcheon ovate, wide, concave, ornamented by fine radial
costellae that join the dorsal commissure at a steep angle anteri

31
orly, bearing fine nodes; escutcheon carina a row of small nodes.
Ligamental fossette is ovate, short, slightly posterior of beaks.
Area is arcuate, slightly concave, ornamented by fine costel
lae bearing small nodes; marginal carina a very subtle row of
nodes at the point where the flank costae bend into area. Median
sulcus is a very shallow line where costellae deflect from anterior
to dorsal orientation.
Flank very convex, steep anterior face into commissure,
ventral surface curved more gently, posterior surface tapers
into narrow posterior margin. Up to 26 transverse costae, gently
convex posteriorly; capped by low rounded nodes aligned with
growth lines.
Discussion: S. emoryi has a more inflated umbo than Trigonia
crenulata Lamarck. S. emoryi has a wider posterior margin and the
escutcheon is smaller and narrower than Trigonia scabra according
to Conrad (1857). Cragin (1895b) separated emoryi from clavigera by
its ridge-like radial costae with low, compressed, elongate nodes;
clavigera, instead, has costae formed by rows of elevated nodes that
are expanded, triangular, clavate, or tubercular forms. S. emoryi
is more inflated and less produced posteriorly than S. mooreana,
and the costal nodes are oval shaped.
Bse (1910) noted that Trigonia guadalupe was quitesimilar to
Trigonia emoryi. He characterized T. guadalupe by its more extended
and narrower posterior and by its fewer number of costellae on
the escutcheon. These differences are within the range of species
variability. Two incomplete specimens in the Texas Memorial
Museum are labeled as Trigonia mooreana var. roemeri Bse from
the upper part of the Georgetown Formation at Shakespeare Tank,
location unknown. The articulated specimen is UT37828 (Figs.
10M, N). However, Bse (1910) never published this name, and,
thus, it is a nomen nudum. Here these specimens are identified
as Scabrotrigonia emoryi (Cragin).
Conrad (1857, p. 148, pl. 3, figs. a-c) created the name, Trigo
nia texana, for an internal cast of a large, triangular, very inflated
bivalve collected at Leon Springs (holotype USNM 9843). This
taxon was placed in the genus, Arctica, by Adkins (1928). It is not
a trigoniid.
Occurrence: This species ranges from Middle to Upper Al
bian. It is common in the lower Upper Albian Kiowa Formation
in Kansas (Cragin, 1895a, 1895b; Twenhofel, 1924), in the partly
coeval Tucumcari Formation in New Mexico, and in the Middle
to Upper Albian Walnut, Goodland, and Fort Worth formations
in north-central Texas (Shattuck, 1903; Perkins, 1961; Scott, 1970).
T. guadalupe is from lower Upper Albian strata in Chihuahua with
the ammonite Venezoliceras chihuahuensis (Bse, 1910).
Scabrotrigonia clavigera (Cragin, 1893)
Figure 10K-L
Trigonia clavigera Cragin, 1893, p. 212-213, pl. 46, figs. 12-13;
Cragin, 1895b, p. 377; Adkins & Winton, 1920, p. 73, pl. 17, figs.
4-6; Adkins, 1928, p. 121, pl. 16, fig. 4.
Types: Cragin based his species on two specimens, one of
which is designated the neotype, UT 17243. Other well preserved
topotypes are BEG 34107 and BEG 34108.
Type locality: Cragins specimens were collected at Browns
Ferry on the Red River, Cooke County, Texas (BEG locality 49-T12).
This is the approximate crossing of U.S. I-35. Adkins and Winton
(1920) reported the specimens from the Weno Formation.
Material: One excellently preserved articulated specimen,
TMM UT17243, from the middle of the Washita Group at Browns
Ferry on the Red River, downstream from the I-35 Bridge, Cooke
County, Texas. Length 52 mm, height 43 mm, width of articulated
valves 26 mm.

32
Diagnosis: An elongate ovate valve, moderately inflated an
teriorly; escutcheon wide with noded costellae; area narrow with
closely spaced, coarse costellae; flank costae concave anteriorly,
tapering dorsally, with widely spaced, large, rounded nodes.
Description: Valve triangular, very inflated at umbo, com
pressed at posterior margin, beaks subterminal, incurved, opis
thogyrate; anterior margin steeply curved below beaks, ventral
margin broadly curved, posterior margin truncate and steeply
inclined into dorsal margin that is very slightly concave. Valve
is strongly arcuate, umbo concave, posterior margin truncate;
anterior face beneath beaks nearly vertical at commissure.
Escutcheon lanceolate, about half the length of the dorsal
margin, commissure forms an acute ridge that descends steeply
into a sulcus that separates escutcheon from area, the sulcus fades
out near posterodorsal corner; surface ornamented by up to 13
costellae of very small nodes inclined anteriorly towards beaks,
the costellae become longer at the mid point of the escutcheon
where they consist of 5 nodes, the shorter posterior costellae
consist of 1-2 nodes.
Area an arcuate surface that changes from convex directly
behind the beaks to steeply inclined near the posterior margin;
marginal carina in the nepionic region a sharp flexure capped by
small acute nodes; posteriorly the flexure broadens, the nodes
become smaller, and the flank costae bend sharply from a dorsal
ventral orientation to directed anteriorly; costellae ornamented
by small nodes.
Flank costae are rounded ridges topped by numerous large
nodes, up to 28; nodes on umbonal surface small, sharp, elongated
in direction of growth lines, at midflank nodes become taller and
elongated into bladed or clavate form, along ventral margin nodes
are very tall, elongate with enlarged tips, towards the dorsal and
posterior margins the nodes become smaller; costae are widest at
ventral margin and taper dorsally; interspaces about as wide as
costae, narrowing dorsally. Costae on nepionic part of beak very
low, bearing minute pointed nodes.
Discussion: S. clavigera has wider flank costae bearing larger
nodes than does Scabrotrigonia scabra, which is an Upper Albian
species in Europe and North Africa (Abbass, 1962).
Occurrence: The species ranges from the upper part of
the Upper Albian to Lower Cenomanian. At the type area strata
exposed in bluffs on the south bank of the Red River range from
the Duck Creek to the Weno formations. Perkins (1961) reported S.
clavigera from the Weno, Main Street, and Grayson formations.
Family Rutitrigoniidae van Hoepen, 1929
This family shares several characters with Iotrigoniidae and
Megatrigoniidae according to Cooper (1991): (1) the escutcheon
carina is obsolete or present only in the nepionic stage; (2) the
marginal carina is generally restricted to the umbonal region; (3)
the umbo generally is not subterminal and the valve is extended
anteriorly; (4) the area is relatively narrow with concentric nepionic
ornament and smooth in the mature stage; and (5) flank costae are
subconcentric. Rutitrigoniidae differ from the other two families,
which have V-shaped flank costae that tend to form nodes at the
anterolateral shoulder.
Genus Rutitrigonia van Hoepen, 1929
Type species: Rutitrigonia peregrina van Hoepen, 1929
Rutitrigonia weaveri (Stoyanow, 1949)
Figure 11 A-D
Trigonia weaveriStoyanow, 1949, p. 87-88, pl. 15, figs. 4-8; Saul, 1978, p. 9.

Types: Syntypes UCLA 28541 (fig. 4), 28542 (fig. 6), 28543
(fig. 5), and 28544 (figs. 7-8).
Type locality: Quajote Member, division 3d, Lowell Forma
tion, Ninety One Hills, Cochise County, Arizona.
Material: Three partial external casts were collected from
the Hensel Sandstone at Edge Falls, TMM1994TX4, 5, 6, one RV,
two LV. An additional specimen was collected from the Tuape
Shale at Santa Ana, Sonora, sample 3-19-05-1.
Diagnosis: A moderately inflated, broadly rounded Rutitri
goniid with short costae on the anterior and mid-flank; posterior
margin truncate.
Description: Valve subovate, rostrate, umbo moderately
inflated, opisthogyrate; beaks about one-third of valve length
from anterior margin. Anterior margin more narrowly curved
than ventral margin, posterior margin truncate, dorsal-posterior
margin behind umbo gently concave.
Up to 28 subconcentric flank costae subparallel with
valve length, straight or with ventrally pointed, low-amplitude
Vs in one or two adjacent costae; rounded costae in the nepionic
stage, and mature-stage costae asymmetrical with dorsal side
steeper than ventral side. Posteriorly the costae fade into growth
lines along an arcuate line from the umbo to the posteroventral
margin.
Escutcheon is elongate, lanceolate, concave, with fine growth
lines; escutcheon carina obsolete. Marginal carina indistinct fold
in nepionic area, on posterior dorsal shoulder a shallow sulcus in
place of carina; ornamented by short costellae in nepionic region
directly behind beaks, costellae join nepionic costae in a sharp
flexure with apex towards beak.
Discussion: None of the Hensel Sandstone specimens show
the ...low, narrow, and ill-defined ridge noted by Stoyanow on
some specimens extending from the beak to the anterior margin
of the umbo.
The genotype, Rutitrigonia peregrina, is more narrowly
rounded and extended posteriorly, and the costae are relatively
much longer extending nearly to the area compared with R.
weaveri.
Stoyanow (1949) distinguished R. weaveri from European
Albian-Cenomanian species of Trigonia excentrica Parkinson, Trigo
nia laeviscula Lycett, and Trigonia dunscombensis Lycett by its overall
shape and proportion of the posterior margin. He separated R.
weaveri from the Argentinian species, Trigonia agrioensis Weaver,
by the relative length and character of ornament.
Pholadomya lerchi (Hill, 1893, p. 30-31, pl. 4, fig. 3, USNM
145645) is very similar to R. weaveri and may be a senior synonym;
however, Stoyanows types of R. weaveri are more numerous,
complete and representative of the species concept than the single
known specimen of P. lerchi. The type of P. lerchi is an incomplete
composite mold of a RV that bears the distinctive subparallel,
subconcentric costae similar to those of R. weaveri. Although it is
larger than R. weaveri, its overall shape and position of beak in the
anterior third of the valve compare very closely with R. weaveri.
Its stratigraphic position is Aptian in ...the heavy conglomerate
which marks the base of the Comanche Series at its contact with
the Carboniferous formation... (Hill, 1893, p. 31). The Cow Creek
Limestone is the only fossiliferous marine unit at the type locality
at the Sycamore Creek crossing of State Road 1431 about 4.5 mi
east of Marble Falls, Burnet County, Texas (Amsbury, 1988). Gillet
(1924), Adkins (1928) and Stoyanow (1949) considered P. lerchi to
be a trigoniid in the excentrica group. Saul (1978, p. 9) equated T.
weaveri Stoyanow with T. lerchi (Hill). This species is placed in
Rutitrigonia, and it is recommended that the name be restricted to
the type specimen, which is validly named but inapplicable to the
more completely defined and represented R. weaveri.
Dimensions: Four specimens collected by Stoyanow are: LV

33

FIGURE 11. A-D, Rutitrigonia weaveri (Stoyanow, 1949). A, B, RV, dorsal and lateral views UTMM 1994TX5; C, D, LV dorsal and lateral views UTMM
1994TX4. E-G, Platymyoidea simondsi (Whitney), articulated shell, UTMM 1994TX3, posterior, lateral RV, and dorsal views. H-K, Epicyprina hamiltonae
Whitney, LV, UTMM 1994TX1, interior, lateral, dorsal, and anterior views. Scale bar = 1 cm; scales of E-K are the same.

50 mm high, >58 mm long, 14 mm wide; LV 49 mm high and 28

mm wide; LV 48 mm high, 61 mm long, 23 mm wide; and RV 51
mm high, >66 mm long, 21 mm wide. The holotype of Rutitrigonia
lerchi (Hill) is a RV >65 mm high, 85 mm long, and 25 mm wide.
Occurrence: This species is abundant in shale and limestone
of the Upper Aptian Quajote Member, bed 3d, Lowell Formation
in southeastern Arizona (Stoyanow, 1949). Saul (1978) reported R.
weaveri from the Alisitos Formation in Baja California. Rutitrigonia
lerchi, which appears to be conspecific with R. weaveri, is in basal
Lower Cretaceous rocks in central Texas, presumably from the
Cow Creek Limestone.
Subclass Heterodonta Neumayr, 1884
Order Veneroida H. Adams and A. Adams, 1856
Superfamily Lucinacea Fleming, 1828
Family Fimbriidae Nicol, 1950

Genus Sphaera Sowerby, 1822

Type species: Sphaera corrugata Sowerby, 1822
Sphaera roblesi (Bse, 1910)
Figure 12A-C
Corbis (Mutiella) roblesi Bse, 1910, p. 127-128, pl. 27, figs. 1-3;
Adkins, 1928, p. 149;
Unicardium sp. Stoyanow, 1949, p. 95, pl. 16, fig. 4.
Types: Syntypes IGM 372, pl. 27, fig. 1, and IGM 7216, pl.
27, figs. 2, 3.
Type locality: La Encantada near Placer de Guadalupe,
Chihuahua, Mexico.
Material:Asingle articulated external cast with some replaced
shell partly abraded on the umbo of both valves and on the middle
of the left flank is from the Tuape Shale at the Santa Marta section

34
Type species: Crassatella martinicensis dOrbigny in Sagra, 1853
TABLE 4. Comparison of dimensional ratios of species of
Sphaera.
in Woods
Corbis
Taxon
Tuape Shale
Sphaera
roblesi
hamiltonae
banderaensis
corrugata
specimen
in Bse
Dhondt

L/H
1.20
1.11
1.10
1.03
1.01
1.07

L/W
1.33
1.21
1.28
na
1.12
1.31

H/W
1.11
1.09
1.15
na
1.10
1.23

# Specimens
3
1
4
4
1
1

sample 3-23-05-F, specimen ERNO 8509: height = 75 mm, length

= 76 mm, width = 68 mm. One incompletely preserved, internal
mold.
Diagnosis: This large, greatly inflated bivalve has low,
subcentral, prosogyrate beaks, the lunule projects forming an
angular dorsal margin that merges with the straight anterior
margin; posterior margin curved; strong concentric ribs wider
than interspaces, steeper side faces the dorsal margin; faint, closely
spaced radial striae on mid flank.
Discussion: This strongly inflated species differs from
Sphaera corrugata Sowerby (1822; Woods, 1907, p. 157-159, pl. XXIV,
fig. 24, pl. XXV, figs. 1, 2; Dimitrova, 1974, p. 101, pl. 48, figs. 3a,
b, 4a, b; Dhondt and Dieni, 1988, p. 46-47, pl. 10, figs. 7-10) by
its sub-central beaks and strong, concentric asymmetric ribs. Its
shape varies because of preservation as internal molds and as a
result of compaction.
Bse (1910) defined Corbis roblesi from the Middle to basal
Upper Albian strata; he differentiated his species from Sphaera
rotundata (dOrbigny) by the larger size and coarser concentric
costae of S. roblesi.
The proportions of articulated valves of the new Mexican
specimen are similar to those of the specimens from Placer de
Guadalupe, Chihuahua, Mexico (Bse, 1910) and to the Sardinian
specimens of S. corrugata (Dhondt and Dieni, 1988) (Table 4).
Two species from the Glen Rose Formation, Corbis hamiltonae
and Corbis banderaensis (Whitney, 1952b), are similarly greatly
inflated species. However, these species have strongly prosogyrate
beaks, and occur in the Salenia texana Assemblage Zone in the
upper part of the Lower Glen Rose Formation in the Lower Albian
Douveilliceras mammillatum Zone.
Occurrence: Sphaera roblesi is known in the Arizona and
northern Mexican Aptian-Albian Comanche Shelf of the Caribbean
Province. The new specimen is from the Tuape Shale Member of
the Mural Limestone at the Santa Marta section, Sonora, Mexico
(3-23-05-F). Stoyanows specimens are from the Upper Aptian
Quajote Member, interval 3b, which also has the ammonites Acan
thohoplites, Beudanticeras, and Colombiceras. Stoyanow also collected
specimens from the Saavedra Member, interval 7i, of the Lowell
Formation, which underlies the Upper Aptian Dufrenoyia justinae
Zone. Both intervals are now part of the Lower Mural Member
of the Mural Limestone. Sphaera corrugata is a Tethyan species
known from Valanginian to Aptian strata in the Mediterranean,
Eastern Europe, southwestern Asia, Mexico, Peru, and Trinidad
(Dhondt and Dieni, 1988).
Superfamily Crassatellacea Frussac, 1822
Family Crassatellidae Frussac, 1822
Subfamily Scambulinae Chavan, 1952
Genus Crassinella Guppy, 1874

Crassinella semicostata Scott, 1970

Figure 12D
Crassinella semicostata Scott, 1970, p. 72-73, pl. 5, figs. 3-6.
Types: Holotype KU500268, paratypes KU500262,500263a
b, 500264a-b, 500265-500267,500269-500280.
Type locality: Locality M3, McPherson County, Kansas,
NE SE, section 1, T17S, R5W, clay pit north of Kansas Highway
4 about 6 mi west of Lindsborg.
Material: Numerous external molds are on a sandstone bed
from the Los Coyotes Member at the Sierra San Jose section sample
9-14-6F, ERNO8510. The largest specimen is 9mm long, 6 mm high,
smaller specimens in the range of 5 mm long by 4 mm high.
Diagnosis: Asmall triangular valve with strong asymmetric
concentric ribs.
Discussion: These specimens have all the exterior
features of this lower Upper Albian species in the Southern
Western Interior of Kansas. This occurrence extends the range
considerably downward into the Upper Aptian.
Superfamily Arcticacea Newton, 1891
Family Arcticidae Newton, 1891
Genus Epicyprina Casey, 1952
Type species: Venus angulata J. Sowerby, 1814
Epicyprina hamiltonae (Whitney, 1952b)
Figure 11H-K
Corbis hamiltonae Whitney, 1952b, p. 706, pl. 89, figs. 8, 9.
Types: The holotype of Corbis hamiltonae is UMMP 53663.
Type locality: On Texas State 46 12 misouth of Boerne, Texas
in Bandera County near the town of Pipe Creek.
Material: One exquisitely preserved entire LV free of matrix
is from Edge Falls, TMM1994TX1.
Diagnosis: Valve inflated, uniformly ovate, with subcentral
beaks; concentric ribs low and evenly rounded.
Discussion: This valve is of the genus Epicyprina because it
is oval and has the typical arcticoid dentition type: AII2b 4b PII. A
small swelling between the anterior lateral tooth AII and 2b may
be 2a. It has a simple pallial line, but the nymphs do not show
rugosities; perhaps these are worn away?
Although the holotype of Corbishamiltonae is an internal cast,
as are most specimens collected from the Glen Rose Formation,
the new valve can be identified with the holotype by the same
shape, size, denticulate ventral margin, and pallial line style.
Whitney named a second cast Corbis banderaensis (p. 706, pl. 89,
figs. 10, 11) and distinguished it from C. hamiltonae by the more
terminal beaks and the less ventricose shape. However, these dif
ferences may be the result of compaction. The Albian-Cenomanian
species, Epicyprina quadrata (dOrbigny) and Epicyrprina oblonga
(DOrbigny) in Ukraine illustrated by Sobetski (1977) are quite
similar in size and shape to the Glen Rose species.
Occurrence: E. hamiltonae is known from the uppermost
Hensel Sandstone and the Lower Member of the Glen Rose For
mation, ranging from the uppermost Aptian Kazanskyella spathi
Zone to the Douvielliceas mammillatum Zone.
Subclass Anomalodesmata Dall, 1889
Order Pholadomyoida Newell, 1965
Superfamily Pandoracea Rafinesque, 1815
Family Laternulidae Hedley, 1918

35

FIGURE 12. Bivalves, Mural Formation, Sonora. A-C, Sphaera roblesi (Bse), A, RV, B, anterior view, C, posterior view, X1, Tuape Shale, Santa Marta
section, sample 3-23-05-F, specimen ERNO 8509. D, Crassinella semicostata Scott, cast of external molds, X2, Los Coyotes Member, San Jose section,
sample 9-14-6-F, specimen ERNO 8510. Scale bar = 1 cm.

36
Genus Platymyoidea Cox, 1964
Type species: Platymya dilatata Agassiz, 1843
Platymyoidea simondsi (Whitney, 1952a)
Figure 11 E-G
Anatina simondsi Whitney, 1952a, p. 70-71, pl. 15, figs. 5, 6;
Platymyoidea simondsi (Whitney), Freneix, 1972, p. 185.
Types: The holotype illustrated by Whitney is UT 40578;
two un-illustrated paratypes are UMMP 53642.
Type locality: Six miles southeast of Bandera, Texas, from
the Salenia Assemblage Zone, Lower Member of the Glen Rose
Formation.
Material: A single articulated entire shell with posterior
broken is from Edge Falls, TMM1994TX3.
Diagnosis: Valve oblong, compressed, posteriorly expand
ed, gaping; beaks on anterior half of valve length; faint concentric
growth rings.
Discussion: The new shell from the Hensel Sandstone
compares very well with Whitneys holotype, which is an internal
cast. However, the beaks of the new shell are closer to the anterior
margin than those of the holotype. The umbonal ridge posterior
to the beaks of the new valves corresponds with the umbonal
grooves described by Whitney on the internal cast; these grooves
reflect internal ridges on the inside of the valves. The species has
a wide posterior gape.
Although Whitney placed this species in Anatina, its ge
neric status has been unclear because the hinge was unknown.
However, certain external features seen on the new shell align
it with the genus Platymyoidea: (1) the overall shape is oblong,
compressed, not tapering, gaping posteriorly; (2) the beaks are
positioned between one-third and one-half of the length from the
anterior margin; (3) the internal umbonal plate is indicated by the
umbonal grooves on the cast, and (4) the shell is ornamented by
concentric growth rings. Freneix (1972) referred A. simondsi to the
genus Platymyoidea while distinguishing it from the Upper Albian
Platymyoidea africana Freneix (1972).
Whitney (1952a) named two other species of Anatina in the
Lower Member of the Glen Rose Formation that differ in degree
of elongation and position of the beaks relative to the anterior
margin: Anatina hanseni (holotype UT40579) and Anatina beckleyi
(holotype UT40557). These differences appear to be distinct, and
the casts also have grooves formed by internal umbonal ridges,
so these species may also be placed in Platymyoidea. A biometrical

analysis of a large collection of specimens would test whether these

are distinct species or part of a variable population.
Cragin (1893, p. 168, pl. 41, fig. 5; plastosyntype UCLA
32240 fide Saul, 1991) based Anatina texana on internal casts
from the Middle-Upper Albian Comanche Peak Formation near
Georgetown, Texas, on the San Gabriel River. This species is more
than twice the size of P. simondsi, 13.1 cm long versus 6.6 cm long,
but should also be placed in Platymyoidea. Interestingly, Cragin
compared A. texana with Platymya rostrata Agassiz, which is now
placed in Platymyoidea (Keen and Cox in Cox et al., 1969, p. N845).
Cragin (1893, p. 168) named a second species Anatina tosta that
has both concentric and radial ornament. The type of this species
is from the Comanche Series in West Texas.
Occurrence: P. simondsi is known from the uppermost
Hensel Sandstone and the Lower Member of the Glen Rose For
mation, ranging from the uppermost Aptian Kazanskyella spathi
Zone to the Douvielliceas mammillatum Zone.
ACKNOWLEDGMENTS
Dr. David Amsbury and Dr. William Ward first recognized
the geologic significance of the Edge Falls locality and promoted
its study. Dr. Joseph Yelderman gave permission to access the
property, study the outcrop and collect geological samples. Dr.
Ann Molineux, Texas Natural Science Center at the University of
Texas Austin, directed the recovery and preparation of the speci
mens. Mexican collections were made by Dr. Carlos M. Gonzlez
Len and Hannes Lser, both at Estacin Regional del Noroeste,
Instituto de Geologa, UNAM, Hermosillo, Sonora, Mxico. Type
specimens and reference collections were made available at these
institutions: the U.S. National Museum by Ms. Jann Thompson,
at the Instituto de Geologa, Ciudad Universitaria by Dra. Mara
del Carmen Perrilliat, Coordinadora Coleccin Nacional de Pale
ontologa, at the Los Angeles County Museum of Natural History
by Dr. Harry Filkorn. Prof. Dr. M. Sander permitted me to examine
Roemers collection at Bonn. Discussions with Dr. Michael R.
Cooper, University of Durban-Westville, South Africa, Dr. Hector
Leanza, Buenos Aires, Argentina, and Dr. Donald Boyd, Univer
sity of Wyoming, Laramie, clarified trigoniid morphology and
taxonomy. LoElla Saul, Spencer Lucas and Charles Newsom pre
pared constructive and thorough reviews. Dr. Nikolaus Malchus,
Autonoma University, Barcelona, provided a key reference and
critical observations on Ceratostreon tuberculifera. Larry F. James,
DDS, Tulsa, prepared latex casts of selected specimens.

REFERENCES
Abbass, H. L., 1962, Amonograph on the Egyptian Cretaceous pelecypods:
Geological Survey and Mineral Research Department, Palaeontological
Series, Monograph 1, 224 p., 24 pl.
Adkins, W.S., 1928, Handbook of Texas Cretaceous fossils: University of
Texas Bulletin 2838, 385 p.
Adkins, W.S., and Winton, W.M., 1920, Paleontological correlation of the
Fredericksburg and Washita formations in north Texas: University of
Texas, Bulletin 1945, 128 p.
Akers, R.E. and Akers, T.J., 2002, Texas Cretaceous Bivalves 2: Houston
Gem and Mineral Society, Paleontology Section, Texas Paleontology
Series Publication Number 7, 516 p.
Albritton, C.C., Jr., 1938, Stratigraphy and structure of the Malone
Mountains, Texas: Geological Society of America Bulletin, v. 49, p.
1747-1806.
Allison, E.C., 1974, The type Alisitos Formation (Cretaceous, Aptian-Al
bian) of Baja California and its bivalve fauna (edited by Durham, J.W.,
and Peck, J.H., Jr.), in Gastill, G. and Lillegraven, J., eds., A Guidebook

to the Geology of Peninsular California: 49th Annual Meeting of the

Pacific Sections of the American Association of Petroleum Geologists,
SEPM, and the Society of Exploration Geophysicists, p. 21-59.
Almazan-Vazquez, E., 1990, Fauna Aptiano-Albiana del Cerrolas Conchas,
Sonora centro-oriental: Actas Facultad Ciencias Tierra UANL Linares,
v.4, p. 153-173.
Amsbury, D.L., 1974, Stratigraphic petrology of lower and middle Trinity
rocks on the San Marcos Platform, South-Central Texas, in Perkins,
B.F., ed., Aspects of Trinity Division Geology: Geoscience and Man,
v. III, p. 1-35.
Amsbury, D.L., 1988, The middle Comanchean section of central Texas:
Geological Society of America Centennial Field GuideSouth-Central
Section, p. 373-376.
Anderson, F.M., 1958, Upper Cretaceous of the Pacific Coast: Geological
Society of America, Memoir 71, 378 p.
Barnes, V.E., 1968, Geologic Atlas of Texas Van Horn-El Paso Sheet: Bureau
of Economic Geology, University of Texas Austin.

37
Bse, E., 1910, Monographa Geolgico y Paleontolgica del Cerro de
Muleros, Estado de Chihuahua: Instituto Geolgica de Mxico Boletn
25, 193 p., 68 plates.
Boyd, D.W., and Newell, N.D., 1997, Areappraisal of Trigoniacean Families
(Bivalvia) and a description of two new Early Triassic species: American
Museum Novitates, No. 3216, 14 p.
Casey, R., 1962, Amonograph of the Ammonidea of the Lower Greensand,
Pt. 4: Palaeontographical Society, v. 116, p. 217-288.
Castell, C.P., and Cox, L.R., 1975, British Mesozoic fossils: British Museum
(Natural History), Publication 703, 207 p.
Castillo, A.D., and Aguilera, J.G., 1895, Fauna fsil de la Sierra de Catorce,
San Lui Potos: Comisin Geologa, Boletn 1.
Checa, A.G., and Jimnez-Jimnez, A.P., 2003, Evolutionary morphology
of oblique ribs of bivalves: Palaeontology, v. 46, p. 709-724.
Conrad, T.A., 1857, Descriptions of Cretaceous and Tertiary fossils, p. 141
174, in Emory, W.H., Report of the United States and Mexican boundary
survey: U.S. 34th Congress, 1st session, Senate Executive Document
108, House Executive Document 135, v.1, pt. 2, 174 p.
Cooper, M.R., 1989, The Gondwanic bivalve Pisotrigonia (family Trigoni
idae), with description of a new species: Palontologische Zeitschrift,
v. 63, p. 241-250.
Cooper, M.R., 1991, Lower Cretaceous Trigonioida (Mollusca, Bivalvia)
from the Algoa Basin, with a revised classification of the order: South
African Museum, Annals, v. 100 (pt. 1), p. 1-52.
Cox, L.R., 1940, Cretaceous Mollusca described by R. Pulteney in the
second edition of Hutchins history of Dorset: Proceedings of the Mala
cological Society of London, v.24, pt. 3, p. 121-127, pl. 7, fig. 8.
Cox, L.R., 1952, Notes on the Trigoniidae with outlines of a classification
of the family: Proceedings of the Malacological Society of London, v.
29, p.45-70.
Cox, L.R., and others, 1969, Mollusca 6, Bivalvia, in Moore, R.C., ed., Trea
tise on Invertebrate Paleontology: The Geological Society of America
and The University of Kansas, Part N, v.1, p. N1-N489.
Cragin, F.W., 1893, A contribution to the invertebrate paleontology of the
Texas Cretaceous: Geological Survey of Texas, 4th Annual Report, Pt.
2, p. 141-294.
Cragin, F. W., 1894, New and little-known Invertebrata from the Neoco
mian of Kansas: The American Geologist, v. 14, p. 1-12.
Cragin, F.W., 1895a, The Mentor beds: a central Kansas terrane of the
Comanchean Series: The American Geologist, v.16, p. 162-165.
Cragin, F.W., 1895b, Astudy of the Belvidere beds: The American Geolo
gist, v.16, p. 357-385.
Cragin, F.W., 1897, Notes on some fossils of the Comanche Series: Science,
new series, v. 6, p. 134-136.
Cragin, F.W., 1905, Paleontology of the Malone Jurassic formation of Texas:
U.S. Geological Survey, Bulletin 266, 172 p.
Crickmay, C.H., 1930, Fossils from Harrison Lake area, British Columbia:
National Museum of Canada, Bulletin 63, p. 33-113.
Dall, W.H., 1889, On the hinge of pelecypods and its development, with
an attempt toward a better subdivision of the group: American Journal
of Science, v. 38, p.445-462.
Defrance, M.J.L., 1821, Dictionnaire des sciences naturelles, tome 22, 570
p., Paris.
Dhondt, A.V., 1982, Some Spanish Cretaceous bivalves: Cuadernos Ge
ologa Ibrica, v. 8, p. 847-865.
Dhondt, A.V., and Dieni, I., 1987, The Sardinian Early Cretaceous Bivalves
and their paleobiogeographic affinities, in Wiedmann, J., ed., Creta
ceous of the Western Tethys: Proceedings 3rdInternational Cretaceous
Symposium, Tbingen 1987, p. 281-297.
Dhondt, A.V., and Dieni, I., 1988, Early Cretaceous Bivalves of Eastern
Sardinia: Memorie di Scienze Geologiche, v. XL, p. 1-97, 25 figs.
Dietrich, W.O., 1938, Lamelibranquios cretcicos de la Cordillera Oriental:
Estudios Geolygicos y Paleontolygicos Cordillera Oriental, Colombia, p.
81-108.
Dimitrova, N., 1974, The fossils of Bulgaria, IVb, Lower Cretaceous
(Gastropoda and Bivavlvia): Bulgarian Academy of Sciences, 258 p.
(in Russian).
Douvill, H. 1916, Les Terrains Secondarie dans le massif du Moghara:

Paris Academie Sciences, Mmoire v. 54, third series, 184 p., 21 pl.
Freneix, S., 1972, Les mollusques bivalves crtacs du Bassin ctier de
Tarfaya (Marocmridional): Notes and Memoires Service gologique
Maroc 228, p. 49-255.
Gabb, W.M., 1861, Synopsis of the Mollusca of the Cretaceous formation,
including the geographic and stratigraphical range and synonomy:
American Philosophical Society, Proceedings, v. 8, p. 57-257.
Gabb, W.M., 1869, Cretaceous and Tertiary fossils: California Geological
Survey, Paleontology, v. 2, 299 p.
Gillet, S., 1924, tudes sur les lamellibranches nocomiens: Societe
Gologique France, Mmoires, series, n., v.1, Mmoire 3, p. 5-224.
Gonzlez Len, C.M., Scott, R.W., Lser, H., Lawton, T.F., Robert, E., and
Valencia, V.A., 2007, Upper Aptian-Lower Albian Mural Formation:
Stratigraphy, biostratigraphy and depositional cycles on the Sonoran
Shelf, northern Mxico: Cretaceous Research, in press.
Ham, T.L., and Landrey, R.J., 1983, A suggested nomenclatural change
and a new reference locality for the De Queen Formation, Pike County,
Arkansas: Gulf Coast Association of Geological Societies, Transactions,
v. 33, p. 109-114.
Harper, E.M., and Skelton, P.W., 1993, The Mesozoic marine revolution and
epifaunal bivalves: Scripta Geologica, Special Issue 2, p. 127-153.
Hayami, I., 1975, Asystematic survey of the Mesozoic Bivalvia from Japan:
The University Museum, University of Tokyo, Bulletin No. 10, 249 p.
Hill, R.T., 1888, The Neozoic geology of southwestern Arkansas: Arkansas
Geological Survey Annual Report, v. 2, p. 1-319.
Hill, R.T., 1893, Paleontology of the Cretaceous formations of Texasthe
invertebrate paleontology of the Trinity division: Proceedings of the
Biological Society of Washington, v. 8, p. 9-40.
Hill, R.T., 1901, Geography and geology of the Black and Grand Prairies,
Texas: U.S. Geological Survey, 21st Annual Report, pt. 7, 292 p.
Imlay, R.W., 1939, Paleogeographic studies in northeastern Sonora: Geo
logical Society of American Bull., v. 50, p. 1723-1744.
Imlay, R.W., 1940, Upper Jurassic Pelecypods from Mexico: Journal of
Paleontology, v. 14, p. 393-411.
Imlay, R.W., 1945, Jurassic fossils from the Southern States, No. 2: Journal
of Paleontology, v. 19, p. 253-276.
International Commission on Zoological Nomenclature, 1999, International
Code of Zoological Nomenclature, Fourth Edition: The International
Trust for Zoological Nomenclature, The Natural History Museum,
London, 306 p.
Jacques-Ayala, C., Alencaster, G., and Buitron, B.E., 1990, Macrofauna
marina del Aptiano-Albiano en el area de Caborca, Sonora: Revista
Sociedad Mexicana Paleontologica, v. 3, p. 67-77.
Jones, D.L., 1960, Pelecypods of the genus Pterotrigonia from the west coast
of North America: Journal of Paleontology, v. 34, p. 433-439.
Kelly, S.R.A., 1995, New trigonioid bivalves from the Albian (Early
Cretaceous) of Alexander Island, Antarctic Peninsula: Systematics,
paleoecology, and Austral Cretaceous paleobiogeography: Journal of
Paleontology, v. 69, p. 264-279.
Kitchin, F.L., 1903, The Jurassic fauna of Cutch. The Lamellibranchiata.
Genus Trigonia: Memoirs of the Geological Survey of India, Palaeon
tologica Indica, ser. 9, v. 3, pt. 2, 122 p.
Kitchin, F.L., 1908, The invertebrate fauna and paleontolgical relations
of the Uitenhage Series: Annals of the South African Museum, v. 7,
p. 21-250.
Kobayashi, T., and Nakano, M., 1957, On the Pterotrigoniinae: Japanese
Journal of Geology and Geography, v. 28, p. 220-238.
Koch, Fr.C.L., and Dunker, W., 1837, Beitrge zur Kenntnis des nordde
utschen Oolithgebildes und dessen Versteinerungen: Verlag Oehme
und Mller, Braunschweig, Germany, 64 p., 7 pl.
Lamarck, J.B. de, 1801, Systme des animaux sans vertbras: Paris, v. 8,
432 p.
Lamarck, J.B. de, 1819, Historie des animaux sans vertbras: Paris, v. 6,
part 1.
Lawton, T.F., Gonzlez Len, C.M., Lucas, S.G., and Scott, R.W., 2004,
Stratigraph and sedimentology of the upper Aptian-upper Albian Mu
ral Limestone (Bisbee Group) in northern Sonora, Mexico: Cretaceous
Research, v. 25, p. 43-60.

38
Lazo, D.G., 2003, The genus Steinmanella (Bivalvia, Trigonioida) in the
Lower Member of the Agrio Formation (Lower Cretaceous), Neuqun
Basin, Argentina: Journal of Paleontology, v. 77, p. 1069-1085.
Leanza, H.A., 1993, Jurassic and Cretaceous trigoniid bivalves from west
central Argentina: Bulletins of Paleontology, Paleontological Research
Institution, v. 105, No. 343, 95 p.
Leanza, H.A., and Garate Zubillaga, J.I., 1987, Faunas de Trigonias
(Bivalvia) del Jursico y Cretcico inferior del Neuqun, Argentina,
conservadas en el Museo Juan Olsacher de Zapala, in Volkhemer, W.,
ed., Bioestratigrafa de los Sistemas Regionales del Jursico y Cretcico
de Amrica del Sur: Mendoza, v.1, p. 201-255.
Loucks, R.G., 1977, Porosity development and distribution in shoal-water
carbonate complexes-subsurface Pearsall Formation (Lower Creta
ceous) South Texas, in Bebout, D.G. and Loucks, R.G., eds., Cretaceous
carbonates of Texas and Mexico, Applications to subsurface explora
tion: Bureau of Economic Geology, University of Texas Austin, Report
of Investigations No. 89, p. 97-126.
Lozo, F.E., and Stricklin, Jr., F.L., 1956, Stratigraphic notes on the outcrop
basal Cretaceous, central Texas: Gulf Coast Association of Geological
Societies, v. 6, p. 67-78.
Lucas, S.G., and Estep, J.W., 2000, Stratigraphy, paleontology and corre
lation of the Lower Cretaceous Carbonate Hill Member of the U-Bar
Formation, East Portrillo Mountains, New Mexico, in Lucas, S.G., ed.,
New Mexicos Fossil Record 2: New Mexico Museum of Natural His
tory and Science Bulletin No. 16, p. 81-86.
Lucas, S.G., and Lawton, T.F., 2000, Stratigraphy of the Bisbee Group (Juras
sic-Cretaceous), Little Hatchet Mountains, New Mexico: New Mexico
Geologic Society Guidebook, 51st Field Conference, p. 175-194.
Moore, C.H., Jr., 1964, Stratigraphy of the Fredericksburg Division, south
central Texas: University of Texas, Bureau of Economic Geology, Report
of Investigation 52, 48 p.
Morton, S.G., 1830, Synopsis of the organic remains of the ferruginous
sand formation of the United States with geological remarks: American
Journal of Science, 1st series, v. 17, p. 274-295.
Nakano, M., 1960, Stratigraphic occurrences of Cretaceous trigoniids in the
Japanese Islands and their faunal significances: Hiroshima University,
Journal of Science (C), v. 3, p. 215-279, pls. 23-30.
Nakano, M., 1968, On the Quadratotrigoniinae: Japanese Journal of Geol
ogy and Geography, v. 39, p. 27-41.
Parkinson, J., 1811, Organic remains of a former world: An examination
of the mineralized remains of Vegetables and Animals of the Antedi
luvian World generally termed Extraneous Fossils. Pelecypods, v. III,
p. 165-226, London.
Perkins, B. F., 1961, Biostratigraphic studies in the Comanche (Cretaceous)
Series of northern Mexico and Texas: Geological Society of America,
Memoir 83, 138 p.
Pictet, F.J., and Campiche, G., 1866, Description des fossils du Terrain
crtac des environs de Sanite-Croix, Part III: Matrielpour la Palon
tologie, Suisse, Series 4.
Poulton, T.P., 1979, Jurassic trigoniid bivalves from Canada and western
United States of America: Canadian Geological Survey Bulletin 282,
60 p.
Pugaczewska, H., 1975, Neocomian oysters from central Poland: Acta
Palaeontologica Polonica, v. 20, p. 47-93.
Pulteney, R., 1813, Catalogues of the birds, shells, and some of the more
rare plants of Dorsetshire, in Hutchins, J., History and antiquities of
the County of Dorset, 2nd edition, vol. 3, 110 p., 23 pl.
Roemer, F., 1849, Texas: Adolph Marcus, Bonn, 464 p.
Roemer, F., 1852, Die Kreidebildungen von Texas and ihre organischen
Einschlsse: Adolph Marcus, Bonn, 100 p.
Rosales-Domnquez, M.C., Grajales-Nishimura, J., Snchez-Ros, M.A.,
Gmez-Luna, M.E., and Dueas, M.A., 1995, Biostratigraphy of the
Lower Cretaceous Bisbee Group, Rancho Culantrillo area, northeastern
Sonora, in Jacques-Ayala, C., Gonzlez-Len, C.M., and Roldn-Quin
tana, J., eds., Studies on the Mesozoic of Sonora and adjacent areas:
Geological Society of America, Special Paper 301, p. 49-57.
Saul, L., 1978, The North Pacific Cretaceous trigoniid genus Yaadia: Univer
sity of California Publications in Geological Sciences, v. 119, 65 p.

Saul, L. R., 1991, Type and referred specimens of fossil Invertebrata in

the Natural History Museum of Los Angeles County, Supplement
3: Mollusca formerly conserved at the University of California, Los
Angeles: Natural History Museum of Los Angeles County, Technical
Report 6, 173 p.
Scott, G., 1930, The stratigraphy of the Trinity Division as exhibited in
Parker County, Texas: University of Texas Bulletin 3001, p. 37-52.
Scott, R. W., 1970, Paleoecology and paleontology of the Lower Cretaceous
Kiowa Formation, Kansas: University of Kansas Paleontological Con
tributions, Article 52 (Cretaceous 1), 94 p.
Scott, R. W., 1987, Stratigraphy and correlation of the Cretaceous Mural
Limestone, Arizona and Sonora: in Dickinson, W.R. and Klute, M.A.,
eds., Mesozoic rocks of southern Arizona and adjacent areas. Arizona
Geological Society, Digest, v. 18, p. 327-334.
Scott, R.W., 1995, Global environmental controls on Cretaceous reefal
ecosystems: Palaeogeography, Palaeoclimatology, Palaeoecology, v.
119, no. 1-2, p. 187-199.
Scott, R.W., 2003, Introduction, in Scott, R.W., ed., Cretaceous Stratigra
phy and Paleoecology, Texas and Mexico: Perkins Memorial Volume:
GCSSEPM Foundation, Special Publications in Geology, No. 1, CD
book, p. vii-xiv.
Scott, R.W., Amsbury, D.L., Molineux, A., and Ward, W.C., 2007, Late
Aptian bivalve assemblage of the transgressive Hensel Sandstone,
Central Texas, in Scott, R.W., ed., Upper Aptian-Albian Bivalves ot
Texas and Sonora: Biostratigraphic, Paleoecologic and Biogeographic
Implications: New Mexico Museum of Natural History Bulletin 39,
this volume.
Shattuck, G.B., 1903, The Mollusca of the Buda limestone: U.S. Geological
Survey Bulletin 205, 94 p.
Shumard, B.F., 1853, Paleontology; description of the species of Carbon
iferous and Cretaceous fossils collected: Appendix E, in Marcy, R.B.,
Exploration of the Red River of Louisiana in 1852: U.S. Congress,
Washington, 310 p.
Sobetski, V.A., 1977, Bivalve Mollusks of Late Cretaceous marine plat
forms: Akademia Nauk SSSR, Trudy Paleontological Institute, v. 159,
256 p. (in Russian).
Sowerby, J., 1812-1822, The mineral conchology of Great Britian; or co
loured figures and descriptions of those remains of testaceous animals
or shells which have been preserved at various times and depths in
the earth, 7 volumes.
Stanley, S.M., 1978, Aspects of the adaptive morphology and evolution
of the Trigoniidae: Philosophical Transactions of the Royal Society of
London, B, v. 284, p. 247-258.
Stanton, T.W., 1947, Studies of some Comanche Pelecypods and Gatropods:
U.S. Geological Survey Professional Paper 211, 256 p., 66 pl.
Stephenson, L. W., 1952, Larger Invertebrate fossils of the Woodbine Fo
mation (Cenomanian) of Texas: U.S. Geological Survey, Professional
Paper 242, 226 p, 59 pl.
Stoyanow, A., 1949, Lower Cretaceous stratigraphy in southeastern Ari
zona: Geological Society of America, Memoir 38, 169 p.
Tashiro, M., 1979, Astudy of Pennatae Trigoniids from Japan: Transac
tions and Proceedings of the Paleontological Society of Japan, N.S., v.
116, p. 179-222.
Twenhofel, W.H., 1924, The geology and invertebrate paleontology of the
Comanchean and Dakota formations of Kansas: Kansas Geological
Survey, Bulletin 9, 128 p.
Underwood, J.R., Jr., 1963, Geology of Eagle Mountains and vicinity,
Hudspeth County, Texas: The University of Texas, Bureau of Economic
Geology, Geologic Quadrangle Map 26, 32 p.
Van Hoepen, E.C.N., 1929, Die krytfuana van Soeloeland. 1. Trigoniidae:
Paleontologiese Navorsing van die Nasionale Museum van Bloem
fontein, v.1, p. 1-38.
Vermeij, G.J., 1977, The Mesozoic marine revolution: evidence from snails,
predators and grazers: Paleobiology, v. 3, p. 245-258.
Villamil, T., Kauffman, R.G., and Leanza, H.A., 1998, Epibiont habitation
patterns and their implications for life habits and orientation among
trigoniid bivalves: Lethaia, v. 31, p. 43-56.
Weaver, C.E., 1931, Paleontology of the Jurassic and Cretaceous from west

39
central Argentina: University of Washington Memoir 1,496 p.
Whiteaves, J.F., 1878, On some Jurassic fossils from the Coast Range of
British Columbia: Canadian Naturalist and Quarterly Journal of Sci
ence, new series, v. 8, p. 400-410.
Whitney, M.I., 1937, Fauna of the Glen Rose Formation: University of Texas
Doctor of Philosophy Thesis, 222 p., 59 pls.
Whitney, M.I., 1952a, Some zonal marker fossils of the Glen Rose Formation
of Central Texas: Journal of Paleontology, v. 26, p. 65-73.
Whitney, M.I., 1952b, Some new Pelecypoda from the Glen Rose Formation
of Texas: Journal of Paleontology, v. 26, p. 697-707.
Woods, H., 1899-1903, The Cretaceous Lamellibranchia. Vol. I: A Mono
graph of the Cretaceous Lamellibranchia of England, Palaeontographi
cal Society, 232 p., 42 plates.
Woods, H., 1905, The Cretaceous Lamellibranchia. Vol. II, Part II: A Mono

graph of the Cretaceous Lamellibranchia of England, Palaeontographi

cal Society, p. 57-96.
Woods, H., 1907, The Cretaceous Lamellibranchia. Vol. II, Part IV: Mono
graph of the Cretaceous Lamellibranchia of England, Palaeontographi
cal Society, p. 133-180.
Woods, H., 1913, Ostreidae, Radiolitidae, Additions, Distribution, Bibli
ography, Index: A Monograph of the Cretaceous Lamellibranchia of
England, Vol. II, Part IV, Palaeontographical Society, p. 341-473.
Young, K., 1974, Lower Albian and Aptian (Cretaceous) ammonites of
Texas: Geoscience and Man, v. 8, p. 175-228.
Young, K., 1986, The Albian-Cenomanian (Lower-Upper Cretaceous)
boundary in Texas and northern Mexico: Journal of Paleontology, v.
60, p. 1212-1219.

ADDENDUM
Barremian to Lower Aptian bivalves and gastropods are
diverse and well preserved in the Zapotitln and San Juan Raya
formations in the state of Puebla, Mexcio between Puebla and
Veracruz (Alencaster, 1956). Unfortunately, this publication came
to my attention after completion of this manuscript. Several spe
cies are similar to those described above and deserve comment
and comparison.
Isognomon lamberti (Mllerried); Alencaster, 1956, p. 8-10,
1,
pl. figs. 2, 3, 9; pl. 2, fig. 9.
This species in the San Juan Raya Formation is similar to
Gervillaria alaeformis Sowerby in the Mural Formation in Arizona
and Sonora, but differs by its greatly arched dorsal margin and
wide ligamental area.
Exogyra tuberculifera Koch and Dunker; Alencaster, 1956, p.
12, pl. 1, figs. 7, 8.
The specimen from the Zapotitln Formation is indeed
Ceratostreon tuberculiferum, although the radial costae are more

distinct than on the Arizona specimens. Alencaster is the first to

report this species in North America.
Pterotrigonia plicatocostata (Nyst and Geleotti); Alencaster,
1956, p. 14-15, pl. 3, figs. 1, 2.
This species is abundant in the San Juan Raya Formation
(Escalante-Ruiz and Quiroz-Barroso, 2006), and it differs from
Pterotrigonia amsburyi by the uniform width of the posterior flank
costae and by the wide non-costate posterior flank marginal to
the marginal carina and sulcus. It is more similar to Scabrotrigo
nia mooreana, which has wider flank costae even on the posterior
flank.
Corbis (Sphaera) corrugata (Sowerby); Alencaster, 1956, p.
17-18, pl. 4, figs. 1-3.
This species is common in the San Juan Raya Formation and
differs from Sphaera roblesi by its more anterior umbo and finer
concentric growth rings.

REFERENCES
Alencaster de Cserna, Gloria, 1956, Pelecpodos y gasterpodos del
Cretcico Inferior de la regin de San Juan Raya-Zapotitln, Estado
de Puebla: Universidad Nacional Autnoma de Mxico, Instituto de
Geologa, Paleontologa Mexicana, No. 2, 61 p.
Escalante-Ruiz, A.R., and Quiroz-Barroso, S.A., 2006, Paleobiologa

de las trigonias de la Formacin San Juan Raya, Estado de Puebla,

Mexico (Abst.): Memoria del X Congreso Nacional de Paleontologa,
Universidad Nacional Autnoma de Mxico, Instituto de Geologa,
Publicacin Especial 5, p. 35.