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Upper Aptian Albian Bivalves of Texas
Upper Aptian Albian Bivalves of Texas
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
edited by
Robert W. Scott
Albuquerque, 2007
Bulletin 39
New Mexico Museum of Natural History & Science
A Division of the
DEPARTMENT OF CULTURAL AFFAIRS
edited by
Robert W. Scott
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.
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
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.
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.
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.
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.
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.
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.
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.
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
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.
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
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
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
S. mooreana
S. emoryi
vyschetzkii
Q. guildi
Q. guildi
S. mooreana
S. mooreana
Q. taffi
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.
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.
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
80
55
>3
60
50
65
50
60
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
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)
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.
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.
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;
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
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.
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.
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
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
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.
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.
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
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
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
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.
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
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
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