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Middle Triassic Pteriomorphian Bivalvia (Mollusca) from the New Pass Range, West-
Central Nevada: Systematics, Biostratigraphy, Paleoecology, and Paleobiogeography

Author(s): Thomas R. Waller and George D. Stanley Jr.
Source: Memoir (The Paleontological Society) , Jan., 2005, Vol. 61, Supplement to Vol. 79,
no. 1 of the Journal of Paleontology (Jan., 2005), pp. 1-64
Published by: Cambridge University Press
Stable URL: https://www.jstor.org/stable/4095819
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Copyright ? 2005, The Paleontological Society
MIDDLE TRIASSIC PTERIOMORPHIAN BIVALVIA (MOLLUSCA) FROM THE

NEW PASS RANGE, WEST-CENTRAL NEVADA: SYSTEMATICS,
BIOSTRATIGRAPHY, PALEOECOLOGY, AND PALEOBIOGEOGRAPHY
THOMAS R. WALLER' AND GEORGE D. STANLEY JR.2
'Department of Paleobiology, National Museum of Natural History, Washington, District of Columbia 20013-7012, USA,
<wallerthomas @nmnh.si.edu>,
2Department of Geology, University of Montana, Missoula 59812, USA, <fossil @ selway.umt.edu>
ABSTRACT-The parautochthonous Anisian and Ladinian rocks of the New Pass Range previously referred to as the Augusta Mountain
Formation have a unique, endemic assemblage of pteriomorphian bivalves with Triassic Northern Hemisphere biogeographic relation-
ships, especially to the Carnian cratonal faunas of northeastern British Columbia as well as to Ladinian faunas of the Shoshone
Mountains and the Gillis Range. A new suborder Prospondyloidina is described, as well as three new genera, Promysidiella, Loxoch-
lamys, and Nevadapecten, a new subgenus, Gervillaria (Baryvellia), and 17 new species either named or referred to by letters: Pro-
mysidiella planirecta, P. desatoyensis, Bositra favretensis, Enteropleura species a, Gervillaria (Baryvellia) ponderosa, Atrina sinuata,
Antiquilima ladinica, Palaeolima newpassensis, Plagiostoma acutum, Leptochondria shoshonensis, Asoella? species a, b, and c, Oxytoma
(Oxytoma) grantsvillensis, Pleuronectites newelli, Loxochlamys corallina, and Nevadapecten lynnae. Promysidiella aff. otiosa is a new
combination for Mytilus otiosus McLearn; Mysidiella newtonae n. sp., Norian, Wallowa terrane, is based on Krumbeckiella cf. timorensis
(Krumbeck) of Newton, 1987. Triassic "Posidonia" is placed in Bositra de Gregorio, 1886, regarded as a senior synonym of Peribositria
Kurushin and Trushchelev, 1989. The morphological and stratigraphic sequence Bositra-Enteropleura-Daonella-Aparimella-Halobia is
rooted in Paleozoic Caneyella and Posidonia and indicates declining dependence on and eventual loss of byssal attachment as well as
a possible shift from low to high dispersal ability (between Bositra and Enteropleura). There is no evidence for trans-Panthalassan
dispersal of bivalves in low latitudes within the interval of Ladinian coral beds. So-called Tethyan and Muschelkalk species are endemics
in slowly evolving genera or are phylogenetically convergent on west Tethyan species.
Shoshone Mountains near Berlin, "Excelsior Formation" in the

INTRODUCTION
Gillis Range; see Fig. 1 and Appendix), and in some cases ho-
FOSSILIFEROUS MARINE Middle and Upper Triassic (Anisian
lotypes to designated from these localities because of bet-
have been
Carnian) rocks exposed in South Canyon, New Pass Range,
ter states of preservation.
west-central Nevada (Fig. 1), have long been of biostratigraphic
The ammonoid
interest, because they display one of the most complete present study concentrates on the superorder Pteriomorphia
successions across the Ladinian-Carnian boundary because the great
known in majority of bivalves that we collected belong
North America (Johnston, 1941; Silberling, 1956; in this group.and
Silberling Protobranchs are very rare and not identifiable, and
Tozer, 1968). These rocks are also of biogeographic the few palaeoheterodonts
interest be- (trigonioids) that we found are not well
cause unlike truly allochthonous terranes they are parautochthon- enough preserved to describe. A single heterodont valve possibly
ous with respect to the craton. They were depositedrepresenting on an island- a new species of the cardiniid genus Minepharus
arc terrane after it had accreted to the craton before the end of Tokuyama, 1958, and an anomalodesmatan valve possibly rep-
Early Triassic time (Speed, 1979, fig. 1; Tozer, 1982, p. 1085resenting
and a new genus in the Modiomorphoidea (sensu Fang and
references therein; see Silberling and Wallace, 1969 and Nichols Morris, 1997) were found in Unit E (see explanation of unit let-
and Silberling, 1977, for regional geology). The fossil assem- tering below), but we prefer to delay their description and illus-
blages in this section include the earliest colonial scleractiniantration until more material becomes available. Megalodontids pre-
corals known from the Americas (Stanley, 1979, p. 37; Roniewicz viously reported from the New Pass Range by Stanley (1979, p.
and Stanley, 1998, p. 246). The biotas thus give precious insight 54) apparently were misidentified with the thick-shelled bakev-
to the Triassic marine biogeography of the continental shelf alongelliid described herein as Gervillaria (Baryvellia) ponderosa n.
the western margin of Pangea, much of which was subsequently subgen. and sp. We also exclude from our study species of Dao-
obscured by the complex accretionary tectonics of later Triassic nella Mojsisovics, 1874 that occur in the Favret Formation (Unit
terranes (Tozer, 1982). B) in the lower part of the section. These species were described
The present study is the first to examine South Canyon bivalvesand illustrated by Silberling and Nichols (1982) from the Fossil
in detail. Fourteen of the 20 species that we describe from South Hill Member of the Prida Formation in the Humboldt Range
Canyon (Table 1) are from the interval containing colonial corals (Fig. 1).
(mostly the endemic Ceriostella variabilis Roniewicz and Stanley, What emerges from this study is the description and bio-
1998) in about the middle of the approximately 1,160 m-thick geographic interpretation of a unique western Pangean cratonal
section (Stanley, 1979; Roniewicz and Stanley, 1998). In seeking (parautochthonous) pteriomorphian bivalve fauna that contains
to understand the biogeographic significance of the coral-associ- co-occurring low- and midpaleolatitude elements. The low-paleo-
ated bivalves, however, we have also collected from the interval latitude elements have Tethyan affinities at the generic level. In
spanning from near the base of the section (middle Anisian) to
contrast, the midpaleolatitude elements are endemic at the generic
the level of a lens of ammonoids and large bivalves (late Ladi- and species levels with their closest relatives, either ancestral or
nian) about 20 m stratigraphically above the coral beds. Higher descendant, being either to the north on the Triassic cratonal shelf
in the section bivalves decrease in species richness. Although (northeastern British Columbia, Canada) or in the Muschelkalk
these bivalves are also worthy of study, particularly an assem- faunas of the Germanic Basin and elsewhere in Europe.
blage of halobiid bivalves associated with the Carnian ammonoids
described by Johnston (1941), they are considered beyond the SOUTH CANYON BIOSTRATIGRAPHY AND AGE CONSTRAINTS
scope of the present work. Some of the species that we describe
occur at other localities in west-central Nevada (Favret Formation South Canyon (Locality 1 in Appendix) is at lat 39036'42"N;
in the Wildhorse mining district, Grantsville Formation inlong the 117029'31"W on the western flank of the New Pass Range,
1

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
2 WALLER AND STANLEY
PERSHING30
39 FH' / 305
Lovelock
80
NEVADA AM e/
.Humboldt
Humboldt
Sink Humboldt
Salt WM 0. LANDER
North Marsh
Nutgrass
95Lake
CHURCHILL/ SC *k
Fallon 11r 5
116 <Stillwater 50
Point Austin
Carson so
Reservoir
95 Lake Eastgate
- - 722
361 ....
L...... 839
MINERAL / 376
Lake
Walker . Gabb
gWC
31 361
NYE
0 10 20 mi
Hawthorne 0 10 20 km
~ Luning I I I
FIGURE 1-Index map showing principal Middle Triassic sites
Range; GF, Grantsville Formation, Shoshone Mountains; SC,
Canyon, Gillis Range.
Desatoya Mountains, Churchill County,

Silberling west
(1956, central
fig. 1) hasNev
be
and northwest
proximately 43 km [27 miles] is used herein (Fig. and
of Austin 2).
west-northwest of New Pass names
Mineof(Fig.
formations
1, SC). (Refer
on Silbe
to
minute Mount Airy and the quently found
15-minute to be Creek
Edwards erroneou
Va
pographic Quadrangle Maps.)60), we have removed these n
graphic
The generalized stratigraphic columnunitsof beginning
Middle and with
Upp
Unit G
assic rocks exposed on the north at the
wall top, just
of South below
Canyon d
TABLE 1-Stratigraphic distribution of bivalves from South Canyo

in Figure 2. Bl = Unit B, Hyatti Zone; B2 = Unit B, Shoshonen
lens; EF = Excelsior Fm., Gillis Range; GF = Grantsville Fm., S
Stratigraphic Occurrence
Species B1 B2 D El E2 GF EF
Promysidiella aff. otiosa (McLearn) x
P. planirecta n. sp. x
P. desatoyensis n. sp. x
Bositra favretensis n. sp. x
Enteropleura species a x
Gervillaria (Baryvellia) ponderosa n. sbgen. and sp. x x x x
Atrina sinuta n. sp. x
Liostrea? jungi n. sp. x
Antiquilima ladinica n. sp. x
Palaeolima newpassensis n. sp. x
Plagiostoma acutum n. sp. x
Leptochondria shoshonensis n. sp. x x
Asoella species a x
Asoella species b x
Asoella species c x
Oxytoma (Oxytoma) grantsvillensis n. sp. x x x
Pleuronectites newelli n. sp. x
LoxochIamys corallina n. gen. and sp. x
Nevadapecten lynnae n. gen. and sp. x
Pseudoplacunopsis aff. fissistriata (Winkler) x x

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
TRIASSIC MARINE BIVALVIA FROM NEVADA 3
Tertiary base of Unit E. He reported no age-diagnostic fossils from the

volcanics -7 "Coral zone" but referred to its age as "Ladinic (or Karnic?),"
- FAULT apparently because of its stratigraphic position between known
upper Anisian and lower Carnian rocks. Johnston's "Joannites
zone (or zone of Trachyceras aon)" coincides with Unit F (Fig.
2). All but one of the ammonoids described by him are from this
Upper G
Camian? zone, and they establish that this part of the South Canyon section
is "approximately equivalent to the zones of Trachyceras aon or
Trachyceras aonoides" (early Carnian: Cordevolian and Julian)
? in the Alpine region of Europe. Johnston referred to Unit G (Fig.
2), which forms the cliffs at the mouth of South Canyon, as the
Lower r"Trachyceras beds" "Red Rock limestone" and speculated that its age is "Karnic?,"
Carnian F (Desatoyense Zone) but he found no age-diagnostic fossils in the unit.
with Halobia Silberling (1956, fig. 1) provided a measured stratigraphic col-
umn for the north wall of South Canyon, described lithologies,
and indicated the positions of biostratigraphically important fos-
sils. One, Daonella dubia (Gabb, 1864), occurs at the top of Unit
Beds with ammonoids B. It is now known to be restricted to the Occidentalis Zone in
the Humboldt Range of Nevada (Silberling and Nichols, 1982,
(Fig. 3) (Meginae? to
E Sutherlandi Zones) and 68), this zone being the uppermost chronozone of the Anisian
(following Silberling and Nichols, 1982, fig. 5). Another, "Pro
Gervillaria (Baryvellia)
trachyceras aff. archelaus," shown by Silberling as occurring i
Coral beds Unit E well above the coral beds, is a clear indicator of Ladinia
Ladinian ., with bivalves age. In Europe the stratigraphic range of Protrachyceras arch
.?i Gervillaria laus (Laube, 1870) spans all but the lowest part of the Ladinia
D .. . o (Baryvellia) bed (Silberling and Nichols, 1982, fig. 5), and in North America spe
cies close to P. archelaus have a similarly long range (Tozer
1994, p. 201).
Silberling and Tozer (1968, p. 35) renamed Johnston's (1941
Daonella beds "Joannites zone" the "Desatoyense Zone," noting that most of
the ammonoids described by Johnston occur in the "lowest fe
9 i(Rotelliformis to
tens of feet" of Unit F (Fig. 2) and that this assemblage sugges
Md I de Occidentalis Zones)
Middle B "an earliest Karnian age, equivalent to that of the Aon Zone i
to Enteropleura horizon the Alps." They also reported (p. 36) the discovery of Daonell
Upper (Favret (Shoshonensis
Anisian Fm.)
Zone)
elegans McLearn, 1947b in this assemblage, this species being a
indicator of the Sutherlandi Zone in northeastern British Colum-
Bositra beds bia. The Sutherlandi chronozone is close to the Ladinian-Carnian
(Hyatti Zone) boundary and was arbitrarily assigned to the Ladinian side of the
boundary by Tozer (1967, p. 29; 1994, table 1).
A In Unit E of the South Canyon section, "several hundred feet
below the Desatoyense Zone," Silberling and Tozer (1968) noted
Paratrachyceras cf. sutherlandi McLearn, 1947b (which now is
FIGURE 2-Stratigraphic column modified from Silberling (1956, fig. 1)
placed in the genus Frankites Tozer, 1971, 1994, p. 165) in as-
measured on north wall of South Canyon, New Pass Range, Nevada. sociation with Protrachyceras cf. archelaus and Hungarites sp.
Letters refer to stratigraphic units described in text. Lithologic symbols A position "several hundred feet" below the Unit-E/-F boundary
are from Silberling's original section; see text for new lithologic de- (Fig. 2) would place this level close to but above the Unit E coral
scriptions. beds. It is only the report of Frankites cf. sutherlandi (McLearn,
1947b) that indicates a latest Ladinian (or possibly earliest Car-
nian) age for this level. The associated ammonoid Protrachyceras
There is as yet no international agreement on the placement of cf. archelaus allows for a much broader age range, from late early
the Anisian/Ladinian and Ladinian/Carnian boundaries relative to Ladinian to latest Ladinian (Silberling and Nichols, 1982, fig. 5).
ammonoid chronozones. The biostratigraphic zonal terms usedHungarites Mojsisovics, 1879 ranges broadly in the Middle Tri-
herein and their relationship to European stages and substages assic in Europe. In the Humboldt Range of Nevada, Silberling
follow Silberling and Tozer (1968) and Silberling and Nichols and Nichols (1982, p. 11, 38) reported Hungarites in the upper
(1982). member of the Prida Formation associated with ammonoids char-
Past work.-Johnston (1941, p. 448) described the stratigraphic acteristic of the Meginae Zone of middle late Ladinian age in
section in South Canyon, reviewed earlier work, and commented Canada (Tozer, 1994, table 1) as well as with "Daonella cf. D.
on stratigraphically useful bivalves and ammonoids in the lower lommeli," a late Ladinian indicator in Europe. In Canada, Tozer
(Anisian) and upper (lower Carnian) parts of the section. He de-(1994, p. 3) listed Hungarites only in the Maclearni Zone, of late
scribed a transitional change about midway through Unit B (Fig. Ladinian age between the Meginae and Sutherlandi Zones.
2) from a lower "Acrochordiceras zone" containing abundantNichols and Silberling (1977), in an extensive review of the
ammonoids and the bivalve "Posidonia" (Bositra de Gregorio, stratigraphy of the Star Peak Group in regions to the north of our
1886 herein) into an upper "Daonella zone" of late Anisian age.section, provided new comparisons and correlations with the New
Based on his lithologic description and thicknesses, what he de- Pass Range. They concluded that the Favret Formation of the Star
scribed as the "Coral zone" encompasses all of Units D andPeak E Group in the Augusta Mountains is the only formation name
(Fig. 2), with the corals themselves in about the position of the that can be appropriately applied in the New Pass Range, where

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
(1982,
it is represented by Unit p.2).
B (Fig. 22-23)
Most (USNM
of the t
B in South Canyon corresponds
of bivalvesto reported
the upper
bymeN
Hill Member, with only the the Fossilbasal Hill few meters of
Member o
sponding to the informallyMountains. named Silberling "lower memb a
these taxa in
section of the Favret Formation as the indicators Augusta of
in Fig. 1). The Favret Formation
Anisian age. ranges The presence in age f
Anisian at its base to latest
greatlate interest,Anisian at its to
because th
Stanley (1979, p. 54) described
lutionarythe link partbetween of the T
section containing coral the beds genus (Unit E, Fig. see
Bositra; 2) in Sy
part of the section again rope. was presented in
Mojsisovics (1874, the f p
graphic column in Gaidzicki
guembeli, and whichStanley later (1983 was
Roniewicz and StanleySilberling
(1998, fig.and 2). Stanley Tozer (19 (1
marked, "the occurrence (1982, of Paratrachyceras
p. 49) referred suth to
er characteristic ammonites
E. bittneri above and below
Kittl, 1912. W th
limestone [near base of it Unit below E, Fig. as Enteropleura
2] is indicat
Ladinian," and this statement
The Daonella was beds repeated contai
identifiedthere
Stanley (1983, p. 87). However, and illustrated is no publ f
of any ammonoids indicative
Formation of at Ladinianits type age areb
beds, nor were we able to and Nichols
verify any (1982).
such occurr These
and D. cf.
field work in South Canyon. sturi (Benecke,
Roniewicz and St
246) recognized this errorbedsand (USNM referred 526388, to the 5263
occurring "below Protrachyceras
B (USNM 526390). ammonoids Ammo n
belonging to the Sutherlandi
respond Zone." to those They describe also
the only age-diagnostic fossils
the Fossil Hill Member of below the
the Prida Formation. Near the coral
base of h
lower in the section, at the
the Daonella bedstop of Unit
in South Canyon, B. inThes
D. americana occurs as-
monoids and Daonella of sociationthewith the ammonoid
OccidentalisGymnotoceras cf. rotelliformis
Zone,
and Tozer (1968, p. 12) Meek, 1877 (cf. Silberling and
regarded as Nichols,
being 1982) (USNMarbitrar
526391).
top of the Anisian rather than
The presence at
of Daonella the
americana base
indicates the lowerof
part ofthe
Gaidzicki and Stanley (1983)
the Rotelliformis Zonedescribed involut
of Silberling and Nichols (1982, table 1),
which they regarded
fers found in thin sections of the as early late Anisian in age and correlated
carbonates in t
Unit E. These involutinids, which
with the Illyrian were found as
substage of Europe.
colonial corals and also Inoccurring
the middle Daonella beds wejust collected the ammonoids
above the
long-ranging Tethyan Eutomoceras
species dunni(Early
Smith, 1904 (USNM 526392), Frechites
Triassic tonev C
do not serve any useful adanus
purpose(Mojsisovics, 1888)
for (USNM 526393), possible Parafr
refinement o
part of the section. chites meeki (Mojsisovics, 1888) (USNM 526394), and the
Roniewicz and Stanley bivalve
(1998) Daonella reported on
cf. sturi (USNM 526389), four
indicating cn
the upper
the coral beds near the base of
most Rotelliformis Unit
and lower E and
Meeki Zones in South
an early late A
also gave detailed descriptions
isian age (Silberling andof associated
Nichols, lit
1982). Our collecting confirm
that the coralline interval lies
the presence some
of Daonella tens
dubia of (USNM
(Gabb, 1864) meter
52639
careous shale that contains
near the top of "Paratrachyceras
Unit B as previously indicated by Silberling (195 am
ferring to the association
fig. 1). Theof Paratrachyceras
occurrence of this species indicates the Occidenta cf
McLearn, Protrachyceras cf.
Zone, regarded P. archelaus
by Silberling and Nichols (1982, table(Laub1) as lat
ites reported by Silberling
Anisian in age. and Tozer, 1968, p. 3
(p. 250) also gave the stratigraphic occurrence
The Occidentalis Zone is the youngest identifiable zone belo o
as "Upper Ladinian (Sutherlandi Zone
the Unit E coral beds. A possible or
exception is alower)
stringer of b
New biostratigraphic valves
observations.--We
in gray limestones in upper Unit D that confirm
contains Gerv
rence of the bivalve "Posidonia"
laria (Baryvellia) ponderosa (=Bositra
n. subgen. and sp. and herein
Leptochon
B and Daonella in upper Unit
dria shoshonensis B. taxa
n. sp. These These
reoccur in the "flat
ammonoid b
of Unit E 20 m stratigraphically
typical monospecific assemblages on above the coral beds as
bedding plawell
shales, siltstones, andin the
silty limestones.
Grantsville Formation of the Shoshone MountainsThe
and in "Po
the "Excelsior Formation"
of a single species, described herein on the upperasplate Bositra
of the Gillis thrust f
associated with the ammonoids
in the Gillis Range (Fig. 1). Acrochordicera
Associated ammonoids in the am-
1877 (USNM 526383),
monoidCuccoceras
bed and in the Grantsville Formationbonaevista
indicate a late Lad-
Smith, 1905) (USNM 526384), Hollandites?
inian age, but no such age-diagnostic sp.
fossils were found with the (
and Intornites sp. (USNM
Unit D bivalves. 526386). These am
which also occur in the The
Fossil Hill
contact between Unit D andmember of
Unit E is conformable and in the
in the Humboldt Range, are
the study area isconsistent
placed at a change from black, with a m
barren limestones
and placement in the Hyatti Zone
to lighter gray bioclastic (Silberling
limestones with some thin interbedded an
p. 5). No Daonella were shales.found
The base of the coral
in beds of Unit E is about lower
these 2 m above b
pears that the stratigraphic
the base of the unit.occurrences
The coral beds measure about 4 m inofthick- "Pos
nella are mutually exclusive.
ness near the base of the north slope of the Canyon and thin
In about the middle of Unit
gradually Bupthere
along strike the slope. Onlyis a thina very
remnant of the n
between the upper limit coral bedsof Bositra
is present on the opposite and
(south) sidethe fir
of the canyon.
Daonella containing a In new
an unnamed species
canyon (informally ofcalled the
"Brachiopodbivalv
Canyon")
Kittl, 1912. It is associated
about 2.5 km with
north of South the
Canyon on ammonoid
the west flank of the
cf. A. carolinae Mojsisovics,
New Pass Range ("SC" in1882 s. a Silberlin
Fig. 1), we found thin (1 m) layer

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
containing colonial corals and the Hungarites

pectinidinermis tobivalves
the preceding Maclearni Subzone 3, and
Pleuronectites
newelli n. sp. and LoxochlamysNitanoceras corallina selwyni n. to the older Meginae and
subgen. Subzone sp.
2. Protrachy-
that
characterize the coral beds in South ceras sikanianum
Canyon. rangesWe broadly through the
regard Upper layer
this Ladinian,
as a northward extension of the from Unit E coral
the Poseidon into thebeds
Maclearniof South
Zones, Can-
and this occurs with-
yon. out apparent morphologic differences (Tozer, 1994, p. 146). If our
The fauna of the coral beds in South determinations are correct, then
Canyon the upper age limit
comprises for the Unit
a dense
assemblage of colonial scleractinian E coral beds is late Ladinian
corals and(Meginae Subzone 2), and the
hydrozoans (Stan-upper
ley, 1979; Roniewicz and Stanley, 1998),
age limit for the involutinid
ammonoid bed is Sutherlandi foraminifera
Subzone 2, latest
(Gaidzicki and Stanley, 1983), and Ladinian. bivalves (Waller, herein) as-
sociated with ooids and bioclasts indicating a shallow-water, high-
MATERIALS AND METHODS
energy environment. Many of the corals and some of the epifaunal
bivalves are in situ and in life position; The bivalve specimens others of haveSouth been
Canyon re- do not
worked, presumably by storm the waves.original The shellcoral beds ofOnly
microstructure. Unit E
originally
are overlain by about 20 m of gray shellthin-bedded
layers remain, usually in a recrystallized
limestones, poorly state
fossiliferous but with abundant aragonitic bioclastsinner andshell thelayersspherical are either red completel
alga
"Solenopora." [See Riding (2004) on calcite
sparry the inappropriate
or are missing altogether. applica- The best p
tion of this name to post-Ordovician the surface ornament
fossils.] is in molds of outer shell surfaces. Most
On the lower part of the north slope
specimens of fragmented
are either South or Canyon about
partially exposed in a hard
20 m stratigraphically above the coral
limestone matrixbeds is a poorly
that is difficult exposed,
to remove without losing parts
approximately 2 m-thick bed of of the specimen.
shaly dark A vexing
gray problem has been the tracing
limestone weath- of shell
ering brown or reddish brown. outlines Thisinlithology
a rock matrix that in some cases contains
contrasts withclosely the
predominantly gray limestones packed on either
fragments ofside. The
other shells. bed contains
Observations were made with a
rich assemblage of ammonoids aand Wild M-5 largestereo microscope
bivalves, with magnifications
including up to X50.
the
enormously thick-shelled bakevelliid, Specimens selected Gervillaria
for photography were (Baryvellia)
coated with ammonium
ponderosa, specimens of which reach
chloride at least
and photographed with23 cm
oblique light in length.
to enhance contrast.
Except for these large bakevelliids, which
With the exception appear
of Figure to be concen-
9.7, all photographs are digital, taken
trated at the very base of thewith unita Nikon and Coolpixare4500 four-megapixel
commonly digitalaligned
camera.
parallel to one another, fossils have a chaotic
All bivalve orientation
identifications are by Waller. Coral within a
identifications
calcareous mudstone matrix. are by Stanley. N. J. Silberling assisted Waller with some of the
The ammonoids present in this bed are critical for providing ammonoid identifications, but the responsibility for these identi-
an upper age constraint for the coral beds. Silberling (1956, p.fications rests with Waller.
1151, and personal commun., 1997) reported "Protrachyceras Abbreviations.--CAS, California Academy of Sciences, San
aff. P. archelaus" from this level. It is also the site (USGS Me-
Francisco; GSC, Geological Survey of Canada, Ottawa; Ht, shell
sozoic Locality M2559) from which Silberling and Tozer (1968, height; L, shell length; MCZIP, Museum of Comparative Zoology
p. 36) reported the discovery of "Paratrachyceras cf. P. suther-(Invertebrate Paleontology), Harvard University, Cambridge,
landi McLeam [now in the genus Frankites Tozer, 1971] in Massachusetts; UMIP, Invertebrate Paleontology Collections, Uni-
association with Protrachyceras cf. P. archelaus (Laube) and versity of Montana, Missoula; USNM, Department of Paleobiol-
Hungarites sp." These specimens have been moved from the col- ogy, National Museum of Natural History, Smithsonian Institu-
lections of the U.S. Geological Survey in Denver into the Ceph- tion, Washington, D.C.
alopod Type Collection of the Department of Paleobiology,
PALEOECOLOGICAL AND BIOGEOGRAPHIC SIGNIFICANCE
Smithsonian Institution, and are illustrated herein for the first time
(Fig. 3). We collected additional specimens of these taxa from the The 20 species of pteriomorphian bivalves describe
ammonoid bed and have assigned USNM catalogue numbers from Units B, D, and lower Unit E of South Canyon
(USNM 526395-526409) to the entire suite of specimens. three distinct paleoecological settings:
Although these ammonoids clearly require further study by spe- Setting A is a basin margin represented by the dark sa
cialists, we are impressed by the variation present among the stones and shales of Unit B (= Favret Formation). It is
specimens of Protrachyceras Mojsisovics, 1893, some of which ized by a stratigraphic succession (from bottom to to
(Fig. 3.6-3.10) appear to us be morphologically closer to P. si- "flat-clam" genera Bositra, Enteropleura, and Daonella
kanianum McLearn, 1930 than to Protrachyceras archelaus. sons detailed in the Systematics section, these genera r
Specimens of Hungarites that we collected resemble Hungarites succession of morphological grades that are the proba
inermis Tozer, 1994 on the basis of shell shape, a shallowly fas- of a phylogeny rooted in Caneyella Girty, 1909 and
tigate venter, and a very close correspondence of the exposed Bronn, 1828 of the late Paleozoic and ending in Halob
1830 in the Late Triassic. The four Triassic genera ar
parts of the suture line (Fig. 3.3-3.5) to that illustrated by Tozer
(1994, fig. 63, pl. 80). On the same block that contains a specimenrecliners, with Bositra probably retaining facultative
of Protrachyceras aff. sikanianum (USNM 526407) is a very tachment inherited from an ancestor in Posidonia. Bys
small, smooth, round-shouldered ammonoid (USNM 526408). Itsment was lost in Enteropleura and Daonella, with th
exposed part is only 5 mm in length and 2.5 mm in width (Fig. developing broadly flaring thin shells (the "snow-sh
3.12) and displays well-preserved ceratitic suture lines. These su-tion") for life on generally low-oxygenated soft sedimen
tures are nearly identical to those of Nitanoceras selwyni (Mc-thic opportunists. On phylogenetic grounds, the so-cal
Learn, 1930) as figured by Tozer (1994, fig. 57j). tube" of the later-occurring (Carnian) genus Halobia
In summary, the ammonoids from the ammonoid bed of Unit morphic feature probably unrelated to any byssal atta
E have their closest counterparts in the Ladinian of northeastern discussed in more detail below.
British Columbia. Furthermore, they indicate the presence of more Setting B is a shallower, nearer-shore, muddy-bottom environ-
than one ammonoid zone with reference to the Canadian Triassic ment represented by lenses of lighter gray sandy limestone in Un
zonation of Tozer (1994, p. 14). In that zonation, Frankites suth- D and in the ammonoid bed of Unit E. It is characterized by th
erlandi was thought to be restricted to Sutherlandi Subzone 2,enormously thick-shelled bakevelliid, Gervillaria (Baryvellia)

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
" el:
Vi
8-w-
17rk
I M
bai~
~iip
Le
q!,.m
2~aXXF?x
y7..
?t~i7''=';-?-?~;40
7p- :iCi,~~3% 1~
FIGURE 3-Late Ladinian ammonoids present in the ammonoi

1947b): 1, USNM 526396; 2, USNM 526397, external mold
and detail of venter; 5, USNM 526400. 6-10, Protrachyce
USNM 526407, exterior composite mold. 11, Protrachyceras
obliquely. 12, Nitanoceras selwyni (McLearn, 1930), USNM
Silberling and Tozer; 10-12 collected by Waller and Stanley

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
ponderosa n. subgen. and sp., interpreted

In Settings B and C, noneas of the
a species
benthic
appear to haverecliner
en-
larged prodissoconchs
that is morphologically convergent on but that not
would suggest expanded larval
closely dis-
related to
Gervillia joleaudi Schmidt, 1936, from
persal stages, and the Muschelkalk
these assemblages (Anisian
show a higher species-level
and Ladinian) of the Mediterranean
endemism thanregion.
that in Setting A.The large
Many of the Nevada
genera in Settings
B and C have long histories,
species probably lived in an environment in some
which extendingsedimentation
back into the Pa-
was slow, an interpretation thatleozoic,
is and wide geographic distributions.
supported by the Speciation occurred
presence of
in areas thatThe
cemented pteriomorphs on its shells. became widely separated, perhaps of
alignment driven Baryvellia
by the geo-
graphicof
shells and the chaotic arrangement complexity
other of nearshore basins and submarine
bivalves, platforms.
particularly
in the ammonoid bed of Unit E, Two examples are discussed
indicate in detail in the Systematics
short-distance section.
transport
and disturbance. The presence of Pleuronectites
two Schlotheim,
stringers 1820, a pectinid
orgenus common in of
lenses the this
assemblage in the South Canyon Muschelkalk,
section apparently
andunderwent
the much more rapid evolution
more in thecom-
Germanic Basin
plete development of this assemblage inasthethat basin became progressively restricted
Grantsville Formation in
Anisian time
about 80 km (50 miles) to the south is than in the more open-ocean
consistent with habitatSilberling's
of a younger
(late Ladinian) age represented in Setting
(1959, p. 13) interpretation of paleogeography. C in South Canyon.
Silberling recog-
Pleuronectites newelli
nized the association of a thick-shelled n. sp. of the coral beds of
"Gervilleia" Unit E, even
with species
of Protrachyceras in both the NewthoughPassyounger in age than the and
Range Muschelkalk
inspecies
the Pleuronec-
Grants-
tites laevigatus
ville Formation. Because the marine Middle Schlotheim, 1820, exhibits ais
Triassic moremore
plesiomorphiccom-
morphology.
pletely represented in the New Pass A similar comparison
Range thancan inbe made
the between the mys-
Grantsville
Formation and has abundant algal idiellids ofand
South Canyon and their westernlimestone,
coralline Tethyan counterparts. Sil-
Plesiomorphic
berling reasoned that the New Pass mysidiellids, here
section mightplaced in Promysidiella
have been n. gen., de-
were Formation
posited seaward of the Grantsville present in both the western Tethys, including
located tothethe Germanicsouth
Basin, and eastern
("GF" in Fig. 1). He further suggested Panthalassa
that the(Nevada, Oregon, northeastern
presence of con-
British Columbia).
temporaneous "reefs" in the seaward New Evolution
Passon the section
western side of Panthalassa
may have
led toin
restricted open marine circulation moreGrantsville
morphological divergencetime,
than on the eastern
thus side,giving
the Grantsville its "Muschelkalk"with characteristics
a succession of genera from Promysidiella
(Hagdorn,to Botulopsis 1985,
p. 239). This "Muschelkalk" character Reis, 1926 (emended herein) to Mysidiella
is thus the result Cox, 1964 occurring
of sim-
ilarity in habitat rather than the in Europe but with speciation
influx ofonly within Promysidiella occurring
Muschelkalk species
from the actual Germanic facies. on the eastern side. By the beginning of the Late Triassic (Car-
Setting C is a shallow, clear-water platform removed from thenian), Mysidiella, which apparently had a European origin, ap-
peared in cratonal areas of northeastern British Columbia while
influences of terrigenous clastics. It is represented by the colonial
coral beds of Unit E in light gray bioclastic and algal limestones. it appeared later in the early Norian assemblage of the Wallowa
The bivalves of Setting C comprise a unique assemblage that isterrane of Oregon (Newton et al., 1987, p. 27, referring to "Krum-
dominated by epifaunal byssate pteriomorphs, particularly mysi-beckiella cf. timorensis").
diellids, pectinids, and limids, all of which are endemic. The pres- It thus appears that the apparent "Tethyan" affinities of the
South Canyon bivalves of Setting C are the result of widely dis-
ence of some articulated shells suggests little transport other than
by current winnowing among the corals. Deep-burrowing heter- tributed genera that have undergone independent evolution on the
odonts and deposit-feeding protobranchs are notably absent, thustwo sides of a vast Panthalassa Ocean. In general, Setting C is
reinforcing the concept of a hard-substrate biostromal environ- inhabited by a midpaleolatitude northeastern Panthalassan com-
ment. munity that contained northern species near their southern limit
andset-
The fossils occurring in each of these paleoenvironmental some "tropical" species, including the corals, that probably
are at
tings have different zoogeographic implications as suggested by or near their northern limit in terms of northern midpaleo-
latitudes.
modes of larval development and by the distributions of nearest
relatives, particularly whether these distributions appear to be per-
SYSTEMATIC PALEONTOLOGY
icratonal or transoceanic. In Setting A, the Bositra-Enteropleura-
Daonella succession is accompanied by ammonoids, indicating
THOMAS R. WALLER
environments that were either on the edge of the open ocean or
that had intermittent broad connections with the open ocean. Introduction.-There
Bos- is as yet no universally accepte
itra favretensis n. sp., so far as can be determined from thefication
avail- of the Bivalvia above the level of superfamily
able material, had normal planktotrophic larval development,eries
sug- of fossil and Recent taxa, new morphological and
gesting normal rather than teleplanic dispersal ability. The nearest
characters, and new data on the stratigraphic ranges o
relatives are the many Lower and Middle Triassic species of Bos-all contribute to new ideas about phylogenetic rela
taxa
itra. These are present along the cratonal margins of northernThe higher level classification used herein is an outgro
Pangea, particularly in northeastern Russia. It would appear that
number of papers that have examined various aspects o
dispersal was mainly pericratonal from the north, with speciation
phylogeny on a broad scale, e.g., Waller (1978, 1990, 19
occurring among isolated suitable habitats along the way. In con-
ter (1990), Carter et al. (2000), Steiner and Hammer (20
trast, both Enteropleura and Daonella appear to have largerother
larval sources cited in the systematic section.
(prodissoconch-II) shells that suggest greater larval dispersal abil-
Word endings for taxa above the family group are unre
ity. The fact that many species of Daonella have broad geographic
by the International Commission of Zoological Nomen
but narrow stratigraphic ranges is well known and suggests andtrans-
there is at this writing no international agreement on
oceanic dispersal ability, perhaps facilitated by distantly placed
endings for the names of orders and suborders. In the
archipelagoes that served as stepping stones (Stanley, 1994). theThesuffix "-oida" has come into fairly common usage
names of orders. The suffix "-ina," commonly used for t
genus Enteropleura also has a broad geographic and narrow strati-
graphic range. Although the species in South Canyon apparentlyof suborders, is in conflict with the same suffix now man
is new, it is closely related to species in the Alpine region theofInternational Code of Zoological Nomenclature, fou
Europe. tion, Article 29.2, for the names of subtribes (in the fami

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Waterhouse (2000, 2001) used

study, the family "-idina"
Mysidiellidae for th
is revised by emending Mysidiel-
ders in classifications that
la, removing Protopis he presented.
and Tommasina, and adding Botulopsis and I u
"-oidina" for names in the
Promysidiella suborder
n. gen. rank
All of the genera now included on
in the Mys-
is a more logical and idiellidae
easily recognized
have morphological dimin
features that are regarded as apo-
"-oida" used for orders. morphic for the Mytiloidea: a bimineralic shell with an outer cal-
citic layer of an antimarginal fibrous microstructure (Waller, 1978,
Class BIVALVIA Linnaeus, 1758
p. 350), an opisthodetic ligament supported by pseudonymphae
Subclass AUTOLAMELLIBRANCHIATA Grobben, 1894 (Waller, 1990, p. 59), a monomyarian (or possibly strongly ani-
Superorder PTERIOMORPHIA Beurlen, 1944
somyarian) adductor musculature, and an equivalved shell with
Order MYTILOIDA Fdrrusac, 1822
the beaks nearly or actually terminal except in very early ontog-
Superfamily MYTILOIDEA Rafinesque, 1815 eny. Characters that are autapomorphic for the Mysidiellidae are
Family MYSIDIELLIDAE Cox, 1964 [emended] curved beaks, inflated umbones, a nonmyophoric anterior septum
Type genus.-Mysidia Bittner, 1891, non Westwood, 1840, nec on which neomorphic hinge teeth may be elaborated, and a var-
Dana, 1850; renamed Mysidiella Cox, 1964, p. 44. iably developed byssal invagination of the anterodorsal shell mar-
Original diagnosis.--"Shell of medium size, ovate or trape- gins, these margins overlapping along the dorsal side of the in-
zoidal, equivalve or slightly inequivalve with the right valve thevagination. Hinge teeth, when present, show signs of transposition
larger; dorsal margin straight or arcuate, umbones more or less indifferently on one valve or the other (Hautmann, 2001b). The
byssal invagination was called a lunule by Cox (1969a, p. N280).
anterior; no differentiated posterior wing; anterior auricle present
or absent; byssal gape probably present in all genera; ligament It may have facilitated strong byssal attachment in turbulent shal-
internal, elongated, extending posteriorly in a groove from near low-water environments (Newton, 1987, p. 18). Details of the
the beak; hinge edentulous; probably monomyarian; shell struc- muscle-scar pattern and hinge of most mysidiellids are still poorly
ture unknown" (Cox, 1964, p. 44). known.
Emended diagnosis.--Mytiloidea with prosogyrous beaks ter- The family Mysidiellidae, as now understood, differs from the
minal at or near anterior end, either narrow and strongly project-pteriomorphian family Myalinidae in shell microstructure [outer
ing anteriorly or broad and coiled inward toward anterior endcalcitic
of layer of mysidiellids antimarginal fibrous rather than
dorsal margin; anterior shell lobe minor or absent except in earlycolumnar prismatic, see Newell (1942, p. 44)] and in ligament
dissoconch; shape of shell variable, from mytiliform to pergami- structure (opisthodetic mytiloid type in mysidiellids compared to
diiform; anterior margins beneath beak steep or strongly inturned duplivincular ligament in myalinids). In all of the mysidiellid
to form a narrow, deep invagination enclosing a narrow byssal specimens examined in this study, the originally aragonitic inner
gape; hinge plate narrow, forming a nonmyophoric septum be- shell layers are either not preserved or are recrystallized to sparry
neath the beaks, edentulous except for interlocking anterior shellor coarsely crystalline calcite. In contrast, a thin outer layer re-
margins at anterior end and, in some species, one or two neo- tains traces of an antimarginal fibrous microstructure. The calcitic
morphic hinge teeth transposable on either valve. Shell thin and outer layer is much thinner than the recrystallized inner layers
bimineralic; outer layer calcitic with a distinctive antimarginal and apparently was originally secreted only in a very narrow band
fibrous structure enhanced by diagenesis; inner layer aragonitic along the shell margin. Although I have not examined the micro-
with original microstructure unknown. Ligament semi-internal, structural details of this layer with the scanning electron micro-
opisthodetic, of the mytiloid type occupying a single groove bor- scope, it superficially appears very similar to the microstructure
dered ventrally by a pseudonymph. Musculature probably mon- of the outer shell layer of many modem mytilids. This was re-
omyarian; anterior adductor insertions not observed. ferred to as "finely prismatic" by Taylor et al. (1969, p. 80),
Discussion.-Cox (1964, p. 44) included three genera in his "fibrillar" by Waller (1978, p. 350), and "fibrous prismatic" by
new family Mysidiellidae: Mysidiella Cox, 1964, Protopis Kittl, Carter (1990, p. 277).
1904, and Tommasina Cox, 1964 (new name for Mytiloconcha Comparison of the Mysidiellidae with the Pergamidiidae is
Tommasi, 1911 non Conrad, 1863). This brought together taxa more difficult, because the latter family is likely a polyphyletic
that previously had been placed in disparate systematic positions.assemblage. Pergamidia Bittner, 1891, originally described from
For example, Kittl (1904, p. 718) introduced Protopis as a sub- the Upper Triassic (Norian) of Anatolia, Turkey, has a large byssal
genus of Opis Defrance, 1825, a genus now placed in the heter- gape between very short but high anterior auricles that are clearly
odont family Astartidae (Chavan, 1969, p. N571), but Krumbeck demarcated from the body of the shell (Cox, 1969c, fig. C44.2).
(1924, p. 261) raised Protopis to the rank of genus and left Its itsligament system is oblique alivincular, and at least in the type
systematic position undetermined. Waagen (1907) placed Joan- species, Pergamidia eumenea Bittner, 1891, the ligament area ex-
nina Waagen, 1906 in the Myalinidae Frech, 1891, but Cox (1964, tends on both sides of the beaks (Cox, 1969c, fig. C44.2b). The
p. 44) determined that this genus is a junior synonym of Protopis.shell microstructure of Pergamidia is unknown, but the genus is
Tommasi (1911, p. 35) concluded that his new genus Mytilicon- almost certainly of a different clade than the genera Krumbeck-
cha (=Tommasina) may belong in the Mytilidae Rafinesque, iella Ichikawa, 1958 and Manticula Waterhouse, 1960. The latter
1815, but his closest comparison was to Dreissena Beneden, two genera, placed in the Pergamidiidae by Cox (1969c, p. N314),
1835, a genus that is now placed in the veneroid family Dreis- have a small, skewed anterior auricle and a horizontally striated
senidae Gray in Turton, 1840 (Keen, 1969b, p. N643). Cox (1964) ligament area with a very shallow, broad ligament pit (Crame,
1995). These features, as well as the high southern paleolatitude
noted that Mysidiella, based on its type species Mysidia orientalis
Bittner, 1891, has a shape resembling that of members of the occurrences of Manticula and some Krumbeckiella, suggest re-
lationship to the Permian genus Eurydesma Morris, 1845 (Crame,
family Pergamidiidae but has a very different hinge structure (ali-
vincular in Pergamidiidae, elongate opisthodetic in Mysidiella). 1995, and references therein). This relationship is strengthened by
Subsequently, Cox (1969a, p. N280) placed the Mysidiellidae similarities in shell microstructure. Eurydesma and Manticula
within the superfamily Mytiloidea, a position that most but not have similar calcitic microstructures, including those that are cal-
all later workers have followed. citic cross-foliated, and a similar innermost aragonitic microstruc-
The study of new Middle Triassic mysidiellids from the New ture described by Carter (1990, p. 203). The shell microstructure
Pass Range and elsewhere in the western Americas has led to aand mineralogy of the shell of Krumbeckiella is unknown. Se-
reevaluation of the generic content of the family. In the presentmuridia Melville, 1956, from the Jurassic of England, was also

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
included in the Pergamidiidaemost by Cox

probably from (1969c,
Promytilus Newell, p.1942,N314) but may
an Early Carbon-
represent yet another clade. Its iferousanterior
to Early Triassic formauricles
that had a prominentdiffer
antimarginal in shape
from those of Pergamidia, Krumbeckiella, fibrous outer shell layer but lackedand Manticula,
a byssal invagination and an and
Semuridia also differs from umbonal these septum genera
(see Dagis et al.,in possessing
1989, p. 84, for occurrence of a weak
posterior lateral tooth beneath Promytilus
the in Lower Triassic and Newell,
posterior end1942, of pl. 2, fig. 4b ligamen
the
area. The inner shell layer of Semuridia
showing is As
lack of umbonal septum). said
discussedtoin thebe nacreous
following
(Cox, 1969c, p. N314); its outer shell
systematic layer,is although
section, Promysidiella not de-
the geologically oldest mys-
scribed, appears to be columnar idiellid genus. Its oldest species,
prismatic based originally
on named Mytilus eduli- of the
figures
left valve of Semuridia dorsetensis formis Schlotheim, (Cox, 1926)
1820, was already present in
in the Cox
Rdt of Ger-(1969c,
fig. C44.4a). many (Schmidt, 1928, p. 168) by the beginning of the Middle
Protopis is removed from the Mysidiellidae for the following Triassic (Hagdorn, 1991, fig. 3, showing Bithynian age of the
reasons: 1) Its shell form, as shown by Kittl's (1904, pl. 23, fig. R6t). By Anisian time, Promysidiella had diversified in the Ger-
15a-e) drawings of the type species, P. triptycha Kittl, 1904, is manic Basin (Frech, 1904, describing variation of Mytilus eduli-
that of a probable dimyarian bivalve in the Heteroconchia. The formis) and was present in the eastern Tethyan region (Mytilus
shell shape is rectangular, with a very sharp angle formed between eduliformis in the Anisian of Jordan, reported by Cox, 1932, p.
the posterior and ventral margins at the point where the ventral- 108). In northwestern Pangea, Promysidiella is unknown before
most of three distinct dorsal radial keels intersects the margin. the Ladinian, but this may be an artifact of the lack of suitable
The beaks, although highly enrolled, are not terminal with respect facies and lack of taxonomic recognition rather than the effect of
to the hinge and are overlapped by extension of the anterior mar- an eastward dispersal across Panthalassa. Occurrences of Pro-
gin of the shell. 2) There is no byssal invagination, because the mysidiella in higher paleolatitudes in western Pangea are later, as
lunule is merely the spiral trace of the concavity between the exemplified by P. americana (K6rner, 1937) in the Carnian of
coiled beak and the anterior margin. The nature of the lunule is northern Peru and P. otiosa (McLearn, 1947a) in the late Carnian
particularly well shown by Krumbeck (1924, pl. 197, fig. 18b) of northeastern British Columbia. Promysidiella is thus far un-
for P. timorensis Krumbeck, 1924, and the lack of a byssal open- known from Norian rocks in the Tethyan region, but on the east-
ing is shown by Waagen (1907, pl. 34, fig. 12c, 12d) for Protopis ern side of Panthalassa it persisted at least until the early Norian
joannae (Waagen, 1906). 3) There is no indication that the shell (Mysidiella cordillerana Newton, 1987, herein selected as the
of Protopis was originally bimineralic, and antimarginal fibrous type species of Promysidiella from the Wallowa terrane of north-
shell fabric was not reported by Kittl (1904), Waagen (1906, eastern Oregon).
1907), Krumbeck (1924), or Cox (1964, 1969a). 4) The nature of Botulopsis and Mysidiella share blunt, medially inturned beaks
the ligament of Protopis is unknown. However, the escutcheon- that do not project anteriorly as far as the most anterior point on
like space between the dorsalmost carinae of opposite valves the shell margin, and both have a rounded, tumid, egg-shaped
(Kittl, 1904, pl. 33, fig. 15b) suggests the presence of nymphae form. On the basis of these characters, they are derived relative
and a true parivincular ligament system rather than a submerged to Promysidiella, which is closer in both beak form and shell form
mytiloid-type system (s. Waller, 1990) between nearly or actually to the outgroup genus Promytilus. Mysidiella, in turn, is derived
closed dorsal margins. relative to Botulopsis because of the deepening and narrowing of
The monotypic genus Tommasina is poorly known, and con- its byssal invagination. This phylogenetic relationship is corrob-
cepts of it rest entirely on the original description and illustrations orated by first occurrences. Botulopsis is known only from the
of the type species, Mytiloconcha orobica Tommasi, 1911, from Ladinian of western Tethys; the oldest-known Mysidiella occurs
the Ladinian of Lombardy, Italy (Tommasi, 1911, p. 35, pl. 3, in the Carnian of the same region.
figs. 21-23; figures in part reproduced by Cox, 1969a, figs. Mysidiella, as herein emended (see below), persisted and
C22.2a-d). Like Protopis, its beaks are nonterminal, with the an- evolved in Tethys until the Norian-Rhaetian. The earliest member
terior margin of the shell extending anteriorly. It has dorsal radial of the genus, M. orientalis, has only very weakly developed ne-
carinae in a position similar to those of Protopis but not so strong- omorphic hinge teeth on its umbonal septum. The latest-occurring
ly developed. Tommasi's plate 3, figure 21a suggests, however, member, Mysidiella imago Hautmann, 2001b, from the Norian-
that there may be three carinae, as in Protopis. Because of frac- Rhaetian of Iran, has strongly developed teeth. Mysidiella is not
turing of the anterior margin, the lunule is poorly shown in Tom- known to occur in the Americas before the Carnian. The first
masi's figures, but his plate 3, figure 21a suggests that it is like known American occurrence is Cardium? saxitilis McLearn,
that of Protopis. As for shell ornament, Tommasi specifically stat- 1947a, a poorly known species herein transferred to the genus
ed that only rough, uneven, concentric (i.e., commarginal) growth Mysidiella on the basis of its large size, egg-shaped form, and
bands are present. Furthermore, if Tomassi's (1911, pl. 3, figs. diagenetically enhanced antimarginal fibrous shell microstructure.
21c, 22b) interpretive drawings are correct, Tommasina orobica It occurs in the cratonal Lima? poyana beds of the Peace River
lacks an umbonal septum, hinge teeth, a byssal invagination, and foothills of northeastern British Columbia, regarded as Carnian in
overlapping anterior shell margins on the anterior side of the age (McLearn, 1953, p. 1221; Tozer, 1967, p. 8). The only other
hinge plate. Although the hinge features of Protopis are unknown, Mysidiella thus far recognized is the species that Newton (1987,
Tommasina is regarded as a junior subjective synonym of Pro- p. 27) described as Krumbeckiella cf. K. timorensis (Krumbeck,
topis and therefore is also removed from the Mysidiellidae. 1924) from the early Norian of the Wallowa terrane of Oregon.
With removal of Protopis and Tommasina from the Mysidiel- It is shown herein to be a Mysidiella and is described below as
lidae, the only genus remaining among those originally placed in Mysidiella newtonae n. sp.
the family by Cox (1964, 1969a) is Mysidiella. However, in the The diversification of Promysidiella on the two sides of Pan-
present study (see following systematic sections), Mysidiella is thalassa would appear to be separate events stemming from a
emended and subdivided into two genera, Mysidiella and Pro- trans-Panthalassan Early Triassic ancestral stock rather than in-
mysidiella, and another Triassic genus, Botulopsis, is added to the dicating trans-Panthallasan dispersal during the Anisian and Lad-
family. This systematic revision of the Mysidiellidae permits a inian. In contrast, Mysidiella indeed seems to have a Tethyan or-
tentative reconstruction of mysidiellid phylogeny that in turn has igin, having evolved from a Ladinian genus, Botulopsis, which is
interesting biogeographic implications. unknown in the Americas. If this is the case, then it would appear
Promysidiella is probably descended from within the Mytilidae, that Mysidiella spread eastward across Panthalassa in the Carnian,

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
perhaps facilitated by theBased

spread
on this
of planktonic
putative an
land terranes. There is, shape of the
however, shell
little of Prom
support fo
from the records of Mytilus,
first where
occurrences. the
Thus shell
far, the
known from an island terrane
turned isbutM.not
newtonae
greatlyfro
pr
rian of the Wallowa terrane,
dient and it is preceded
directed more posteby
nian) occurrence, M. saxitilis
partures (McLearn, 1947a),
of Promysidiella
tonal rocks of Carnian to
age
bein British
in Columbia.
three trends: 1)
shape remains mytiliform
Genus PROMYSIDIELLA
remainsnew genusIn wester
shallow.
the series
Type species.--Mysidiella Promysidiella
cordillerana Newton, a
fig. 12, Early Norian, Wallowa terrane,
respectively Oregon.
present in ro
Other species.-Mysidia Norian americana age. Based Korner, on the 1
Peru; Mytilus eduliformis umbonal Schlotheim, angle in 1820, this sequ An
Mytilus otiosus McLearn, otiosa 1947a, through Carnian, an averag nort
Columbia; Promysidiella about planirecta 45?in P. n. cordilleran
sp., Ladini
Promysidiella desatoyensis a similar n. sp., trend Ladinian, in Promys Nev
some species described as "Mytilus"
chelkalk, specimens from fromthe E
chelkalk may prove to er be umbonalmembersangles of this than genu t
Diagnosis.-MytiliformLower to isognomoniform Muschelkalk. 2) MysTh
narrow, anteriorly projecting
growth beaks, along abyssal posteroven inva
able depth but commonly length shallow, of theand anteroventra a short u
lacking hinge teeth. perpendicular to plane of
Etymology.-Prefix "pro-" from combined the flattened with remain gen
diella, indicating probableonly known in
ancestral the uniqu
relationship
Occurrence.--Lower to Upper Triassic
planirecta. 3) A mytilifor (Spathian
low to middle paleolatitudes
present in in Eurasia
early ontogeny and wes
South America (Nevada, occur Oregon, in later British ontogeny: Columb t
Discussion.-Promysidiella shifts differs ventrally, from Botulop
leading to
diella in having a mytiliform
sal invagination to isognomoniform deepens a
row, prosogyrous beaks dimension.
that project The beyond end result the
of the shell. In Botulopsisrow, and strongly Mysidiella anteriorly the bea p
do not project anterior to observed the anteriormost in Promysidiell point
South Canyon.
margin. The byssal invagination of Promysidiella It is also o
from very shallow and nian of Peru.
scarcely differentiated as
otiosa (McLearn, 1947a) to narrow and deep as in
desatoyensis. Even in the PROMYSIDIELLA
latter aff. OTIOSA (McLearn, 1947a)
case, new combination
however, t
narrow and strongly project anteriorly. Figure 4.1 Unlike
mysidiella lacks neomorphic hinge
Mytilus otiosus MCLEARN, 1947a, Appendix, teeth.
p. 2, pl. 1, fig. 2.
Mysidiella cordillerana is selected as the type s
its silicified state of preservation
Description.-Mytiliform Mysidiellidae allowed
of small size (maxi- Newto
trate and describe both mum exterior
dimension less than 10 mm),and with umbonal angle of about
interior fea
the umbonal septum and 660, heightligament
and length about equal, overall insertion.
shape trigonal, with In
type species and the three species
anteroventral margin described
tending to be straight in lateral view; beak belo
inian of South Canyon, slightly Promysidiella
prosogyrous; lunule slightly concave, not set off probably
from
European and Asian Triassic species
lateral surface by a distinct carina, height of lunulethat about 38% of have p
placed in Mytilus Linnaeus,
maximum dimension 1758.
of shell; byssal Based
gape not observed;on small speci
Muschelkalk of Germany septum presentexamined
beneath beak, other internal featuresinunknown; Ingelfin
(Hagdorn Collection, Muschelkalk
exterior surface smooth except for oneMuseum),
to several unevenly spaced Myt
eduliformis Schlotheim, commarginal
1820 growth ledges;
and thin outer shell layer has a micro-edulif
Mytilus
Renz in Frech, 1904 belong
structure with distinctin Promysidiella.
antimarginal lineations; inner shell layer S
tilus eduliformis described
recrystallized. by Cox (1932, p. 108
probable Anisian age in Material examined.--UMIP 18048-B,
Jordan, based RV exterior on on same my e
Cox's specimens at The pieceNatural
of rock as the holotype History
of Loxochlamys corallina Museum n. sp.,
As discussed above, a andlikely
USNM 526410, aancestor
posterodorsal fragment for Promy
of LV exterior, both
Permian-Lower Triassic from genus
the coral beds of Promytilus.
Unit E, Locality 1. The la
diate between ModiolusOccurrence.-Known
Lamarck, only from1799 the coral bedsand of Unit E,Myti
mytiliform except for theSouth Canyon,retention
New Pass Range, Nevada, of a small lob
late Ladinian.
beak, and at least some species
Discussion.-This have
species resembles an antimar
Promysidiella otiosa from
the Lima? poyanaouter
probably originally calcitic, beds of Carnian shellage in the Peace River foot-
layer com
of mysidiellids (Newell, hills, British 1942,
Columbia (McLearn, p. 1947a,33; Appendix, Nakazaw
p. 2, pl. 1,
1968, p. 50; but see Carter,
fig. 2; Fig. 4.4) in1990,
having a trigonal, p. 277).
Mytilus-like shape. Furthe
The New
a relatively small time Pass
gap specimensbetween
differ, however, in having the last
an umbonal know
angle great-
Promytilus in the early er thanEarly
that of 10 of theTriassic
11 specimens of Promysidiella (Promyti otiosa
rushin in Dagis et al., that
1989 ofin the
were examined the collectionsTaimyr
of the Geological Survey Penin
the oldest known Promysidiella,
of Canada, Ottawa. Umbonal anglesP. of theseeduliformi
10 specimens ranged
from 430 to 630 with
1820) from the early Middle a mean of 52.50. The lunule of P. aff. otiosa
Triassic.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
is difficult to observe, but it layer

appears to be less
comprising inturned
about (less o
one-third
concave) and with a relatively higher
form,dorsoventral
with nearly dimension thanan
straight dorsal
that of true P. otiosa. The antimarginal
ing at an fibrous
umbonal outer
angle shell layer 3
of about
of the Canadian specimens of P. a radius
otiosa near anteroventral
is not margin
as well preserved
as that of the New Pass specimens and is visible
anteroventral in only meeting
valve surfaces a few
small patches. I attribute this difference to taphonomic
along a sharp curvature conditions,
of the shell
the limestone matrix of the Canadianregion long, specimens extending having from a much near
more coarsely crystalline texture nature indicatingof lunule and byssal
a higher degree gape of u
recrystallization and/or dissolutionsmooth of shells.except for faint growth lin
Promysidiella aff. otiosa also ginalresembles injuries. another
Internal mytiliform
features unk
species from western North America, Etymology.-The Promysidiella prefix cordillerana
"plani-" re
from the early Norian of the the lateral and anteroventral
Wallowa Terrane valve surfaces,
of Oregon.combined with The
umbonal angle of the latter, however, is to
"-recta," referring even narrower
the right-angle intersection ofthan that
these surfac-
of true P. otiosa, averaging only es. 450.
Promysidiella aff. otiosa differs from from
Types.--All co-occurring
the coral beds of adult
Unit E,Pro- South Can
mysidiella planirecta n. sp. (Fig. cality 4.2, 4.3) in
1. Holotype: USNM having
526411, aapair convex
of matching b
rather than nearly planar lateral valves;shell paratypes,
surface USNMand 526412, ina lacking
single rock the specim
right-angle bend that separates thethree
taining lateral
fragments shell of rightsurface
valves, and fromUSNM 5264 a
ment of
long, steep, flattened anteroventral a left region.
shell valve on block I do containing
not have paratypes any of
early ontogenetic stages of P. mys planirecta
corallina n. that gen. and are of a 526485).
sp. (USNM size com-
parable to that of P. aff. otiosa, butOccurrence.-Known
somewhat later only from stages the coralsuggest beds of
that juvenile P. planirecta wouldSouthstill have New
Canyon, a morePass Range, flattened
Nevada, late lateral
Ladinian
Discussion.-None
shell surface and a steeper right-angle inturning of the fiveof specimens
the anterov-of this specie
entral region than in P. aff. otiosa
plete, but of in comparable
combination they provide size. a fairly consisten
Promysidiella desatoyensis n. of sp.shell (Fig.
shape.4.5, 4.6), which
The holotype (Fig. 4.2, 4.3) co-oc-is the only
curs in the Unit E coral beds ofwith South paired valves. Its with
Canyon right valve is nearly
P. aff. complete
otiosa,
is isognomoniform with its maximum except for thegrowth beak and the gradient
anterior part tendingof the flatten
ventrally or anteroventrally ratheroventral region,
than which are obscured by matrix.
posteroventrally and has The le
a strong prosogyrous curvature.offset,The distorted,
early growth and crushed with stage onlyof its P.posterior
depar
satoyensis has a narrower umbonal angle the
ing. By comparing than trends P.ofaff.growth otiosa
lines on the of t
comparable size and, unlike P.itaff. can be otiosa,
seen that the has left a valvenarrow carina
has been slightly rotat
setting off the anteroventral region.
to the right when the specimen is viewed from the left
The only species to which McLearnrotation has moved thecompared
(1947a) flattened anteroventral his My- region o
tilus otiosus is Modiola klipsteini Bittner,
valve forward, where 1895, it isfrom
out of sightthebeneathSan Cas- rock ma
siano Formation (upper Ladinian specimen
to Lower well demonstrates
Carnian) the ofthinness
theof the shell r
Alpine
size and species
region. McLearn said that Bittner's is the source for hasthe measurement
a narrower on shell thickness
outline
and shorter hinge margin than givendoes in theM. description.
otiosus. In fact, the single
specimen that Bittner (1895, p. The 47) newdescribed
species is unique among and illustrated
mytiliform Promysidiellahas in
a maximum shell dimension of its only 4 mm
cross-sectional shape. Its and apparently
lateral valve lacks
surfaces are much flatter
than those of
antimarginal fibrous microstructure. ItP. isotiosa, P. aff. otiosa, and P.
doubtfully a cordillerana,
Mysidiella. and
This is the first time that the thosespecies
species lack the described
extensive flattened by McLearn
anteroventral region. P.
(1947a) as Mytilus otiosus has been
planirectaassigned
also has a much thinner
to the shell than
Mysidiellidae,
those of the other
and this assignment has important mytiliform biogeographic implications.
species. The decreased shell thickness is due primarily
Newton (1983, p. 17) thought that to a much the
thinner species
recrystallized,that
presumably she described
originally aragonitic,
inner shellthe
as Mysidiella cordillerana constituted layer. Compared
first to isognomoniform
report of mysidiellids,
this genus the
anteroventral report
in North America, the only previous carina of P. planirecta
being is nearly
that straight
of rather
Mysi-than
diella americana (K5rner, 1937) from
anteriorly concavethe Carnian
in lateral view. of Peru. The
determination that McLearn's species, Like otherfrom mysidiellids, P. planirecta
known was likely byssate.
cratonal rocks Its
flattened anteroventral
of Carnian age in British Columbia, surface suggests that it wascloser
is morphologically epifaunal and to
Mysidiella cordillerana than is likely
theattachedPeruvian species
to a hard substrate. lends shape,
Its cross-sectional support with
to the idea that Wrangellia (whichthe most has inflated apartbiota
of the shell similar
adjacent to the to that
flattened anter-of
the Promysidiella cordillerana-bearing
oventral surface,stratasuggests that of the did
individuals Wallowa
not live together ter-
in
clusters as do some
rane) occupied a Northern Hemisphere positionmodern mytilids,
close suchtoas Mytilus
the edulisNorth Lin-
American continental margin rathernaeus, 1758.than a Southern Hemisphere
position such as was indicated by Newton (1983, fig. 5).
PROMYSIDIELLA DESATOYENSIS new species
PROMYSIDIELLA PLANIRECTA new species Figure 4.5, 4.6
Figure 4.2, 4.3
Diagnosis.-Medium-sized, thin-shelled, isognomoniform,
Diagnosis.-Large, thin-shelled, mytiliform
quadrangular Promysidiella
Promysidiella n. gen. with greatest growth gradient n.
ventral to slightly
gen. with nearly planar lateral valve posteroventral;
surfaces beaks narrow, strongly
turning proso-
sharply in-
ward about 900 to a broad, flattened anteroventral
gyrous, anteriorly region.
pointing; dorsal third of anterior margin strong-
Description.-Shell large, reaching
ly concave in a maximum
lateral view, turning inward dimension
to form a deep byssal of
about 50 mm, but very thin-shelled
invagination. (0.3 mm at distance of about
35 mm from beak), with the antimarginal fibrous,
Description.-Shell with maximum dimension calcitic
about 30 to 35 outer

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
;,I
ul-
~?~h~:
II:i
1.
~1'-; ? 1
L :si
'~ -?:-;??:?::?-::~~
"it `:??;~
;"I-'-?i-:- .. ?id~~F~
,r4?'
it:?? ?i ~ P"~IT'~
I
~h,:~i~~4SS
" ~~?I?
-:~:_ ??~:u i:? a~:: "
r?::-
:: r, ~rf?
~?.s1~5~fi
I~ d ;,.
55~1
;n ,: X~? ~:
.i
1~ ?'-
?-: i: sii~'??'-:::;?? :~?~~i~?::
c-~-,.~~~a
t~~:?~?l? ;,.-:?n~Z:
:~
c;~t: k--
.il :~i~~i~p~i
~,,i:t?
u:?r "-
,i: ?I,-.?
ri~S:~~r~t-
1~1~4 `?;.?c: ~~2~5~i~-~ jll?
:~ I: ?:g~ _;i.? ~~ ~r~Til r;~:-~?l??~-;?~?:?i-:?
~iiS ;P.
i :i3:"~:.?,r
...-.::
~~i~ ~ ~tlil
~I:i?i
it~i
~T~
~n?-r
i~i~r~
i I
I?Y1~
;.c
~i~?l;~~,~,~i---- ---???--??~:;i~i~~
~p~888~8~8~
-Z: ?'
~?c:i
..I?. bi
~ ..-~?. ? ..lt.~s4
~%:;i?-~~
?,?i?-;r
iC
~7 CI ?:::i?-?:
S~5~ ? :2a~ ~ia~ :?
~~:P T\
,.?
ji'.
.tc.? .?:~?!\ :
FIGURE 4-Mysidiellidae. 1, Promysidiella aff. otiosa (McLearn, 1947a), UMIP 18048-B, exterior of fragmented right valve showing fibrous shell
microstructure, coral beds of Unit E, Locality 1. 2, 3, Promysidiella planirecta n. gen. and sp., holotype, USNM 526411, exterior of right valve
and oblique view of inturned anteroventral margin, coral beds of Unit E, Locality 1. 4, Promysidiella otiosa (McLearn, 1947a), holotype, GSC
9507, exterior of left valve, "Grey beds," Upper Triassic, Peace River, Pardonet Hill, British Columbia. 5, 6, Promysidiella desatoyensis n. gen.
and sp., holotype, USNM 526414, exterior of right valve and oblique view of byssal invagination, coral beds of Unit E, Locality 1. 7, Botulopsis
cassiana (Bittner, 1895), reproduction of Bittner's plate 5, figure 18, San Cassiano Formation, Italy, natural size. 8-10, Botulopsis reisi n. name,

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
the concave
mm, with thin antimarginal fibrous anterior margin,
calcitic outer deeplayer
byssal invagination,
and some- and sub-
what thicker inner recrystallizedstantial byssal gape
layer thatof M. wasdesatoyensis suggest strong
probably byssal at-
origi-
tachment but tending
nally aragonitic. Shape isognomoniform possibly a suspended
toward living habit. The species may
quadrate,
equiconvex, moderately inflated, have
with been beaks
gregarious,narrow
living in clusters
and in anteriorly
the manner of some
pointed; anterior margin sigmoidal in lateral
living Mytilus. This is in factview, with
suggested by dorsal
the close proximity of
third deeply concave and turned numerous
inward disarticulated
medially valves (e.g.,
to USNM 526415).
meet a narrow
byssal gape, posterior two-thirds of anterior margin convex, ven-
Genus BOTULOPSIS
tral margin less convex, posteroventral margin Reis, 1926 [emended]
more strongly
Figure 4.7-4.10
curved, posterior margin nearly straight with overall trend form-
ing a 900 or slightly obtuse angle Type
withspecies.--Botulopsis
the slightly reisi new name
curved[=Botulopsis
(dor- cas-
sally convex) dorsal margin; umbonal siana Reis, 1926
anglenon Botulopsis
of maturecassiana (Bittner, 1895), by mon-
specimens
very broad (800 to over 1200) due to strong
otypy, Ladinian prosogyry,
(Wettersteinkalk), much
Hochalpe, Karwendel, Austria
narrower in juvenile specimens.(Fig. 4.8-4.10)].
External surface marked by nu-
merous, irregular commarginal growth Other species.-Botula?
lines showing cassiana Bittner, 1895, San
signs ofCassiano
re-
peated marginal chipping and repair Formation, during
Upper Triassiclife; traces
(Carnian), Italy. of anti-
marginal microstructure best developed Original diagnosis.--Original
inward from description
what of Botulopsis
appears cassi-
to be a homogeneous outermostana Reis, 1926,
shell p. 124 (my Small
fabric. translation from German): "The
internal sep- shape
tum beneath beak; other internal features
is rounded oblong, unknown.
highly coiled; the beak is strongly in-curved;
Etymology.--Named after the the Desatoya
anterior umbonal Mountains, Nevada.
area is probably separated by two blunt edg-
Types.-Holotype: USNM 526414 (Fig.
es. The hinge margin 4.5, 4.6),
is short and a in
straight, nearly
figure 1 [Fig. 4.8]
complete right valve with outer sideonly
extending exposed.
posterior to theParatypes:
beaks; the shell inUSNM
figure 2 [Fig.
526415, a single block of rock containing 4.9] should be oriented twosomewhat
rightmore perpendicular
valves, to twoits lon-
left valves, and several fragments; gitudinalUSNM
axis, so that it would thenabelarge
526416, homogeneously
exfo- set up
liated right valve; USNM 526417, a partial
with figures right
1 and 3 [Fig. valve;
4.8, 4.10]. USNM
The shell margin has an
526418, juvenile left valve, and on
otherwise the
evenly same
pointed block,
egg-shaped outline. USNM
526419, a juvenile articulated shell On the shell surface
with its only
right some incremental
valve exposed. bands (growth
All from the coral beds of Unitlines) E, appear
South with the
Canyon,
naked eye, but Locality
with the magnifying
1. glass
Measurements.-Height and length of holotype,
one recognizes many more numerous, 34.3 even,mm and
longitudinal striae in
28.0 mm. the background; with fractured thin peels (tangential fractures),
Occurrence.-Known only from the coral beds of Unithowever, E, a shiny surface appears, which is covered with extreme-
South Canyon, New Pass Range, Nevada, late Ladinian. ly fine and narrowly delineated longitudinal striae."
Discussion.-The early growth stage of the holotype at a length Emended diagnosis.-Mysidiellidae moderately inflated and
of about 5 mm is modioliform, with a very narrow umbonal angle rounded-oblong in shape, with umbones broad and rounded, beaks
and a short anterior lobe anterodorsal to the byssal gape (Fig. medially inturned and not extending beyond anteriormost point
4.6). The actual byssal gape of the specimen is well formed and of anterior shell margin; byssal invagination very shallow.
Occurrence.--Ladinian (Botulopsis reisi) to Carnian (B. cas-
nearly semicircular, with a dorsoventral height parallel to the shell
margin of about 5 mm and a transverse dimension of about 2
siana), Alpine Europe.
mm. Because no articulated specimens with an exposed byssalDiscussion.-Botulopsis has generally been placed in or close
gape were found, it is not possible to say whether the gape toasthe Mytilidae, possibly in the subfamily Crenellinae Adams
seen in the right valve is mirrored by a comparable gape on and the Adams, 1857 (e.g., Bittner, 1895, p. 49; Reis, 1926, p. 124;
left valve. The hinge is not exposed or preserved on any of Soot-Ryen,
the 1969, p. N275). Reis (1926, p. 124) introduced the
specimens, and it is not possible to reconstruct the manner genus in by means of a combined genus and species description
which the shell margins in the dorsal part of the byssal invagi- under the heading, "Botulopsis Cassiana Bittn. spec. nov., gen.
nation may have overlapped and articulated, if at all. nov." Although Reis provided an adequate description of the new
Among the three species of Promysidiella that occur in South species and distinguished it from true Botula? cassiana Bittner,
Canyon, this is the only species that is isognomoniform rather 1895, he unfortunately did not name it. According to the rules of
than mytiliform. The type species of Mysidiella, M. orientalis, nomenclature Reis's new species must therefore be referred to as
from the Carnian and the Norian of Europe and Turkey (see Botulopsis cassiana Reis, 1926, as was done by Kutassy (1931,
above), is similar in shape but has beaks that are inturned towards
p. 353). However, because it is clear from Reis's short discussion
that he also regarded true Botula? cassiana Bittner, 1895, to be
the hinge line and do not project anteriorly beyond the anterior-
most point of the shell margin (see Cox, 1969a, p. N281). Fur- another member of the new genus, the name Botulopsis cassiana
thermore, the byssal invagination of M. orientalis is deeper Reis,
and 1926, is a junior secondary homonym of Botulopsis cassi-
dorsoventrally shorter. In Mysidiella newtonae n. sp., from ana the (Bittner, 1895). I therefore rename Reis's species, which is
early Norian of the Wallowa Terrane of Oregon (described the as type species of Botulopsis, Botulopsis reisi Waller. The basis
Krumbeckiella cf. K. timorensis by Newton, 1987, p. 27), the for regarding this as the type species by original designation by
lunule is even shorter and more deeply incurved than in M.Reis or- is Article 68.2.1 of the International Code of Zoological
ientalis. Nomenclature (fourth edition).
Unlike Promysidiella planirecta n. sp., where the broad, flat- There is another nomenclatural task that is necessary to fix both
tened anterior region suggests attachment to a hard, flat substrate,
the genus name Botulopsis and the name of its type species. Reis
reproduction of figures of "Botulopsis cassiana Bittner spec. nov., gen. nov." of Reis (1926, pl. 8, figs. 1-3), Wettersteinkalks, Germany, natu
size. 11-13, Mysidiella newtonae n. sp., Norian, Hells Canyon, Wallowa terrane, northeastern Oregon, reproduction of figure 21.3, 21.5, and 21
respectively, of Newton (1987): 11, USNM 416395, interior view of fragmented right valve; 12, 13, holotype, USNM 416396, anterior and inter
views of right valve. Scale bars: 1, 3-6, 5 mm; 2, 10 mm; 11-13, 20 mm.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
(1926, p. 124) based his Genusnew

MYSIDIELLA Cox, 1964 [emended] both
species
closed, articulated specimen (pl. 8, figs. 1-
Mysidia BITTNER, 1891, p. 113, pl. 2, fig. 10.
on a specimen figuredMysidiellaby Bittner
Cox, 1964, (1895,
p. 44, pro Mysidia BITTNER, pl
1891 non WEST-
under the name Botula?WOOD, 1840. cassiana. Bittner (1
that his Botula? cassiana consists of two mo
illustrated, respectively, on orientalis
Type species.-Mysidia his Bittner, plate 5, figu
1891, by original
even suggested that these
designation, Norian,morphotypes
Anatolia, Turkey. possi
ent species. He did not Otherspecify aMcLearn,
species.--Cardium? saxitilis holotype,
1947a; Mysidiel- h
both morphotypes comprise
la imago Hautmann, 2001b; the type
Mysidiella newtonae n. sp. series
cies that he named Botula? cassiana. In order to stabilize Botu-
Original diagnosis.-The original diagnosis is the full descrip-
lopsis reisi, new name, I hereby designate the specimen tion
repre-of the type species, Mysidia orientalis Bittner, 1891, p. 113
sented by Bittner's (1895) plate 5, figure 18 (Fig. 4.7) as(my
the translation from German): "Only four right valves are pre-
lectotype of Botula? cassiana. I also designate as lectotype
sent.of
The shell is moderately curved, thin, very finely striated, the
Botulopsis cassiana Reis, 1926 (renamed Botulopsis reisi striae
herein)becoming clearer in exfoliated specimens. The anterior is
the specimen represented by Reis's (1926) plate 8, figures 1-3 broad, without wings or ear, the rear wing in contrast
sloping,
(Fig. 4.8-4.10). gradually grading into the posterior margin. The turned-in part of
Bubnoff (1921, p. 335) placed Botula? cassiana in Cardi-
the anterior edge is substantially broad, somewhat withdrawn
omorpha de Koninck, 1841, a Lower Carboniferous genus fromnowthe plane of commissure, so that a byssal opening was prob-
regarded as a member of the Anomalodesmata, superfamily ablyPho-
present; under the beak a tooth ascends very steeply, behind
ladomyoidea, family Edmondiidae (Newell, 1969b, p. N818; which
Run- a socket follows, which in turn is followed by the long
negar and Newell, 1974, p. 7). Bubnoff's placement of therear edge of hinge which, in turn, is hollowed out by a ligament
genus
was apparently overlooked by Reis (1926) but was repeated groove bylimited rather sharply on its inward side by a raised lon-
Kutassy (1931, p. 415). A position in the Anomalodesmatagitudinal
would border. Lateral teeth are absent."
Revised diagnosis.-Revised diagnosis by Hautmann (2001b,
be highly unlikely, not only because of the apparent bimineralic
shell with antimarginal fibrous microstructure, but also p. 42; my translation from German): "Valves distinctly higher
because
than long,
of its apparent monomyarian condition and the likelihood that the with prosogyrous umbones situated at the extreme an-
opisthodetic ligament is not parivincular. terior end. Below the umbones, the shell margin is folded inward
forming
Botulopsis is placed in the Mysidiellidae for the following rea- a deep lunula and gaping between both valves. The rel-
atively short, slightly convex hinge margin is fitted with a rim for
sons: 1) Both Botulopsis reisi and Botulopsis cassiana (Bittner)
have an antimarginal shell fabric, as clearly described by the attachment of a submarginal ligament. Hinge with well de-
both
Reis (1926, p. 124) and Bittner (1895, p. 49). 2) Both veloped specieshinge plate, carrying a prosocline ridge in one valve
appear to have a shallow byssal invagination. Bittner (1895, whichp. does not project beyond the commissural plane. On the
49) mentioned that the blunt valve edges in this area of the opposite
shellvalve, a strong tooth is developed in a dorsal position
form the only hinge articulation, with the margin of one with regard to this ridge, clearly projecting beyond the commis-
valve
fitting behind the margin of the opposite valve. 3) Although sural the
plane and articulating above (behind) the ridge of the op-
posite valve. The mentioned hinge elements are not fixed to one
ligament structure is not described for either species, the original
valve, i.e., there are individuals with the tooth in the right valve
illustrations of each species show that the dorsal margins in the
and the groove in the left and vice versa. Outer shell layer formed
region of the hinge are closed. There is no lenticular opening such
by foliated calcite. Middle and inner shell layers only known re-
as might be expected for a parivincular ligament. It is therefore
crystallized, probably because of the originally aragonitic com-
possible that the ligament is a sunken mytiloid type. 4) The over-
position."
all shell shape and the nature of the strongly inwardly coiled
Emended diagnosis.-Mysidiellidae with shell inflated, round-
beaks and inflated umbones are similar to these features in Mys-
ed, and egg-shaped; umbones broad, not projecting anteriorly
idiella. The main difference is that the byssal invagination of Bo-
beyond anteriormost point of shell margin, beaks anterior and
tulopsis is very shallow, whereas that of Mysidiella is very deep.
medially inturned; byssal invagination deep and narrow. Hinge
In addition to Botulopsis reisi and B. cassiana, another species
articulation either limited to overlapping shell margins on dorsal
that probably belongs in this genus is Mysidia taramellii De Toni,
side of byssal invagination or consisting of a transposable tooth
1913, an Alpine species from the late Ladinian (Longobardian) and socket developed on the umbonal septum.
of Italy. Based on De Toni's illustrations, its beak is not so strong-
Occurrence.-Eurasia: Greece (Corfu), Turkey (Anatolia), Iran,
ly medially inturned as in Mysidiella, but neither is it strongly
China (east Xizang and west Sichuan), Carnian to Norian-
anteriorly projecting as in Promysidiella n. gen. It has aRhaetian.
tumid North America: northeastern British Columbia, Canada
rounded shape, as in both Botulopsis and Mysidiella, but [Lima? it ap-poyana beds?], Carnian, and Wallowa terrane, northeast-
pears to have a very broad and shallow byssal invagination, ern thus
Oregon, lower Norian.
resembling the other species of Botulopsis. Discussion.--Mysidiella as herein emended differs from Pro-
The shallow byssal invagination of Botulopsis suggests that this n. gen. in having wide umbones (in lateral view) that
mysidiella
genus is derived from a primitive Promysidiella such as Promy- do not project anteriorly beyond the anteriormost point of the
sidiella eduliformis. Similarities in shape between Botulopsis shell and
margin. In Promysidiella the umbones are narrow and
Mysidiella suggest a close relationship between them, but curvedthe and project well anterior to the anteriormost point of the
much deeper and narrow byssal invagination of Mysidiella indi-
shell margin. Mysidiella differs from Botulopsis in the nature of
cates that it is the more derived genus and may have evolved its byssal invagination, deep and narrow in Mysidiella, broad and
from a member of Botulopsis. This interpretation is consistent scarcely developed in Botulopsis. Contrary to Hautmann's
with stratigraphic first occurrences. Botulopsis reisi occurs(2001b,
in thep. 42) diagnosis, no traces of foliated calcite have been
Ladinian in the eastern Alpine region of Europe (Reis, 1926), detected in the Mysidiella examined for this study, and it is un-
whereas the earliest Mysidiella, M. orientalis, first appears likely
in the that such a microstructure would yield the antimarginal
Carnian (in Corfu, according to Diener, 1923, p. 134). fibrous pattern that appears especially on slightly exfoliated and

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
recrystallized specimens. Mysidiellaby an outer lacks shell layer."

anThese outer"riblets" are of very low colum-
simple relief
nar prismatic shell layer of the andtype appear tothatbe highest is at theprominent
anterior and ventral margins in(New- many
pterioidan groups, including the Ambonychioidea
ton, 1987, fig. 21.5). In the center of the shell Miller,
(Newton, 1987,1877
(Carter, 1990; Waller, 1998), where Nevesskaya
fig. 21.4), however, they appear to be ofet al.scale
the same (1971,
and relief p.
150) and Hautmann (2001b, p. 42) as thesuggested
antimarginal fibrous striae that of the the Mysidiel-
outer calcitic shell layer
lidae may belong. Both the fibrillarof typicalmicrostructure
Mysidiella. Newton also described "a and ligament
triangular liga-
ment pit oriented
structure of Mysidiella, most similar to types obliquely." present
I would describe the inligament
thenot My-
tiloida (Waller, 1990), contradict asplacement
a pit but as an elongatein opisthodetic
the ligament Ambonychioi-supported by
dea. pseudonymphae as in modem Mytilus (Waller, 1990). What New-
Mysidiella in this emended sense ton described
is known as "a ligament area relatively narrow
mainly on the but long"basis
is
of its well-known type species distributed
the groove that was occupied inbythe Upper
the opisthodetic ligament.Triassic
New-
ton noted the absenceMysidiella
of Eurasia and by the recently described of striations on this "ligament
imago area," from
but
the Norian-Rhaetian of Iran. Mysidiella
no strong striations would sichuanensis
be expected in a ligament Chen
groove. A (in
Zhang et al., 1985, p. 105) and M. smallyunnanensis
septum that does not appear to Zhang
be myophoric (in Zhang
occurs beneath
et al., 1985, p. 105) from the Upperthe beak (Newton,
Triassic 1987, fig. of21.3, 21.6).
China A narrow are marginalpoorly
band
illustrated and difficult to evaluate
that on
is darkerthe in colorbasis of the
than the interior region of illustrations.
the shell appears
With the transfer of the American in Newton's figure 21.6. This
species may indicate that the
Mysidiella shell was
americana
and M. cordillerana into Promysidiella,
originally bimineralic,
no species with a thin outerthat calcitic and
were thicker inner
orig-
inally described under the namearagonitic
Mysidiella layers. (or its earlier name
Mysidia) remain in previous publications
Recently, Crame (1995) onbroughtTriassic
to light newbivalves
information on of
either North or South America. Inthethe morphologypresent of Krumbeckiella
study, Ichikawa, 1958 and related two
however,
North American species originally generaassigned
in the families Pergamidiidae
to other Cox, 1969c and Eurydes- have
genera
midae Reed, 1932. TheOne
been found to represent true Mysidiella. ligamentis of Krumbeckiella
the species is indeed a that
Newton (1987, p. 27) described as Krumbeckiella
resilium in an oblique, shallow triangular cf. timorensis
pit, the ventral edge of
from the lower Norian of the Wallowawhich projects very terrane, Oregon.
little into the shell interior (Crame, It1995,
is re-
described here as a new species text-fig.
of Mysidiella
3c). The ligament area, caused(see by thebelow).
ventral migration The
other is Cardium? saxitilis from of thethetriangular Carnian
fibrous ligament and "Lima?
the bordering poyana
lamellar lig-
zone" of the Peace River foothills, northeastern
aments, is conspicuously longitudinallyBritish striated. ThisColumbia.
ligament
True Mysidiella is unknown from system older
is unlike that Triassic
of Newton's (1987) rocks Krumbeckiella in cf. North
ti-
America and has not yet been found morensis. in the South American Tri-
assic. The "lunule" and associated features of Krumbeckiella and
related genera in the Pergamidiidae (Crame, 1995, pl. 2, text-fig.
MYSIDIELLA NEWTONAE new species 3) also differ in detail from those of Mysidiella and Krumbeckiella
Figure 4.11-4.13 cf. timorensis of Newton (1987). In true Krumbeckiella, a small
Krumbeckiella cf. timorensis (KRUMBECK, 1924). NEWTON, 1987, anterior
p. 27, auricle-like projection, not the beak, extends anterodor-
fig. 21. sally upward along the anterior end of the ligament area. The
auricular lobe is bordered anteriorly by what may be a byssal
Diagnosis.-Strongly inflated Mysidiella with height to length groove, and this in turn is bordered anteroventrally by an inturned
ratio in mature shells estimated at 1.24; umbonal septum with a shell margin. There is no septum in the vicinity of the beak as in
single, low, weakly developed tooth posterior to the interlocking Mysidiella.
margins on the dorsal side of a deep and narrow byssal invagi- Lastly, although the shell microstructure of Krumbeckiella is
nation. still incompletely known, the possibly related genera Eurydesma
Description.-See Newton (1987, p. 27, fig. 21) for description Morris, 1845 and Manticula have outer calcitic layers that are
and illustrations of "Krumbeckiella cf. timorensis." foliated or cross-foliated, not antimarginally fibrous (see Carter,
Etymology.-Named after Dr. Cathryn R. Newton, who dis- 1990, p. 201, 203, and references therein). There may be inner
covered this species in Hells Canyon in the Wallowa terrane of aragonitic shell layers, but apparently the calcite-aragonite bound-
northeastern Oregon. ary does not closely border the shell margin. In Newton's Mysi-
Types.--Known only from the specimens described by Newton diella and possibly in Newton's Krumbeckiella cf. K. timorensis,
(1987), consisting of four right valves and one left valve. I des- the secretion area of the calcite outer layer appears to be limited
ignate a right valve, USNM 416396 (Newton, 1987, fig. 21.4- to a very narrow band on the inner shell surface along the shell
21.6; Fig. 4.12, 4.13), as the holotype. Paratypes are the other margin.
specimens figured by Newton: a fragmentary right valve (USNM Mysidiella newtonae differs from the type species of the genus,
416395, Fig. 4.11) and a fragmentary left valve (UMIP 7366). Mysidiella orientalis from the Norian of Turkey, in having a great-
Measurements.-Measurements of the holotype, a right valve, er height-to-length ratio, with less ontogenetic expansion of the
USNM 416396, based on a reconstructed outline: height 57 mm, shell on the posterior side, as well as a relatively broader umbonal
length 46 mm, maximum convexity 16 mm. septum. M. newtonae differs from M. imago from the Norian-
Occurrence.-Silicified beds of the "Martin Bridge Limestone, Rhaetian of Iran in having a much weaker hinge tooth on the
near Spring Creek, on the western flank of Hells Canyon of the umbonal septum. Compared to Mysidiella saxitilis from the late
Snake River," Oregon (Newton et al., 1987, p. 2). According to Carnian Lima? poyana Zone of northeastern British Columbia,
Newton (1987, p. 5), the ammonoid Tropiceltites cf. columbianus Canada, M. newtonae has a higher height-to-length ratio (1.29
(McLearn, 1940) occurs in these silicified beds and is "charac- compared to about 1.12) and somewhat narrow umbones in lateral
teristic of the Mojsisovicsites kerri Zone, the lowermost biostrati- view.
graphic interval within the lower Norian (Tozer, 1967, 1982; Sil- Reassignment of Krumbeckiella cf. K. timorensis of Newton to
berling and Tozer, 1968)." the Mysidiellidae and its description as a new North American
Discussion.-Newton (1987) described "extremely regularly endemic species has important biogeographic implications. The
spaced radial riblets that appear to be ornamental features formed genera of the Eurydesmidae and Pergamidiidae mentioned above

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
have mainly Hemisphere Southern

p. 362; Carter, 1990, p. 247), clearly distributions
indicate that monotoidians in
Jurassic 1995). (Crame,
In
represent contrast,
a different Triassic m
clade from that of the Posidonioidea.
mainly Northern Hemisphere Scarlato and Starobogatov
forms. (1979) elevated the
The family Posidon-
new sys
of this species removes iidae toanother
superfamily rank but providedSouthern
no precise diagnosis. They Hemi
merely noted thatof
from the bivalve assemblage genera with
the short hinges
Wallowa(relative to shell terra
an additional indication length)
that customarily
this assigned terrane
to the Posidoniidae, such as Posi-
originated
Hemisphere probably closedonia Bronn, 1828,
to Posidonotis
the Losacco, 1942, and Aulacomyella
North Americ
Furlani, 1910, are distinct from other posidoniid genera that have
Order PTERIOIDA Newell,
relatively 1965
long hinges, such as Daonella and Halobia, which they
Suborder PTERIOIDINA Newell, 1965
placed in a family Halobiidae. However, hinge length relative to
Superfamily POSIDONIOIDEA Frech,
shell length is not by itself a sufficient 1909
criterion to differentiate [eme
Original diagnosis.-Frech the two families Posidoniidae and Halobiidae.
(1909, p. 8, The Early
for Jurassic the su
idoniinae; my translation (late Toarcian
from to early Pliensbachian)
German): genus Posidonotis has dis-
"Thin-sh
ably fully planktonic shells tinct asymmetric with indistinctly
anterior and posterior auricles, with the right set-
vonian to Jurassic, with peak
anterior auricle set off fromin the
the disk Triassic."
by an incised suture (Hayami,
Emended diagnosis.-Thin-shelled, 1988, fig. 1), suggesting that it is a pleurotheticvery pectinoidean.
low-co
ioidina having only very Another genussmall with the sameauricles
narrow stratigraphic range or in the none
lacking hinge teeth; ligament Early Jurassic, Pectinula Leanza,area 1943, is duplivincular
also auriculate. Dam-
horizontally striated; musculature borenea (1987, p. 190) regarded it asanisomyarian
a member of the pectino- o
in monomyarian forms, adductor
idean family Entoliidae and placed it in scar
the synonymy oval
of Posi- or ci
centic, and without an adjacent donotis, but Hayami (1988,prominent
p. 567) disagreed, noting differencesposterio
scar.
in the shapes of the auricles between Posidonotis and Pectinula.
Discussion.--Frech (1909, p. 8) was the first to The base Late
a Jurassic
family- (Tithonian, Kimmeridgian) genus Aulacomyella
group name on Posidonia, as a subfamily Posidoniinae does have a relatively
within the short hinge and is nonauriculate, but its
"Aviculidae," a family that then included a great variety ligament of structure
pter- is unfortunately unknown (Kelly and Doyle,
iomorphian bivalves ranging from inoceramids to pectinids.
1991). Its much later He stratigraphic occurrence suggests that it may
included in the subfamily the genus Posidonia, broadly be merely convergent on the posidoniid form, but there is still not
interpret-
ed to include species ranging from Devonian through enoughJurassic
morphological ininformation to determine its systematic po-
age, as well as the Triassic genera Daonella and Halobia. sition.
Newell (1938, p. 37) placed Posidonia in the Pterinopectinidae, The families that Scarlato and Starobogatov (1979) included in
a group of pleurothetic Paleozoic pectinoids having the Posidonioidea
a duplivin- are a polyphyletic mixture: Posidoniidae Frech,
cular ligament as well as a right anterior auricle separated 1909, Halobiidae Kittl, 1912, Pterinopectinidae Newell, 1938,
from
the disk by a distinct suture and a deep byssal notch. However,
Deltopectinidae Dickins, 1957, and Oxtomidae Ichikawa, 1958.
no such clearly demarcated auricle and byssal notch The lastarethree of these are distinctly pleurothetic, inequivalved
present
at any growth stage of Posidonia. Rather, the right pectinoids anterior with well-demarcated auricles and a byssal notch on
auricle
of Posidonia, as shown by Weigelt (1922), is poorly demarcated,
the right valve. They belong in the order Pectinoida.
with only an incipient byssal sinus. These characters H. are more(1994,
J. Campbell in p. 64), in a useful review of the taxo-
line with placement near the Pterineidae rather than nomicthe Pterino-
history of the Halobiidae, raised the family to the rank of
pectinidae. superfamily containing only the single family Halobiidae. He
Newell (1965, p. 18) elevated the Posidoniinae to family rank chose not to associate the Halobiidae with the Posidoniidae be-
within the superfamily Pectinacea, and in the Treatise, Cox cause, in his words, the latter "has to some extent become a
(1969e) included in the Posidoniidae 11 genera, some of which convenience family to accommodate thin-shelled forms of uncer-
are only questionably phylogenetically related to Posidonia. The tain character and affinity." Nevertheless, the association is al-
superfamily Pectinacea was later restricted to a subset of pecti- ready in place under the prior name Posidonioidea (Scarlato and
noids having a resilium with a central organic, nonmineralized Starobogatov, 1979), and, as noted above, the taxonomic scope
core (Waller, 1978, p. 253), which, so far as known, is not the of the group has already been narrowed, many of the problematic
case for any member of the Posidoniidae. taxa no longer being regarded as posidonioidians.
More recent studies have placed the Posidoniidae in the Pter- In the present study, the scope of the superfamily Posidonioidea
ioida with a likely derivation from a duplivincular pterineoidean is limited to two families, Posidoniidae and Halobiidae, which
stem group, most likely the Lower Carboniferous genus Caneyella collectively are worldwide in occurrence and have a stratigraphic
Girty, 1909 (Carter, 1990, p. 211; McRoberts, 2000, p. 599). Wei- range from Lower Carboniferous to Upper Jurassic (Oxfordian;
gelt (1922, p. 129; 1927, p. 76) had earlier regarded Caneyella see following discussion of Bositra). On opposite sides of Pan-
and Posidonia as so close to suggest that they are the same genus. thalassa, in Eurasia and western Laurasia, there is a stratigraphic
In contrast, Johnston and Collom (1998, table 1) included the succession of first occurrences of the genera Posidonia (Lower
family Posidoniidae, with a query, in a revived and reconstituted Carboniferous), Bositra [=Peribositria Kurushin and Trushchel-
subclass Cryptodonta Neumayr, 1884 and superfamily Inocera- ev, 1989 (Lower Triassic)], Enteropleura (early Middle Triassic),
moidea Giebel, 1852. By their own criteria, however, this place- Daonella (early Middle Triassic, but later than Enteropleura),
ment fails, because Posidonia has a duplivincular ligament (Wei- Aparimella H. J. Campbell, 1994 or comparable transitional forms
gelt, 1922, 1927; confirmed by Amler, personal commun., 2003). between Daonella and Halobia (late Middle Triassic, near the
Rathmann and Amler (1992, p. 51) placed the Posidoniidae in Ladinian-Carnian boundary), and Halobia (early Late Triassic).
the superfamily Buchiacea Waller, 1978 (later corrected on the Based on current knowledge of morphology, this succession of
basis of priority to Monotoidea P. Fischer, 1887; see Begg and broadly interpreted genera represents a series of increasingly de-
Campbell, 1985, p. 727). Monotoidians, however, are pleurothetic rived grades of evolution of ligament and shell morphology. Some
pectinoids with a distinctive right anterior auricle that is clearly of the steps in the succession may involve the polyphyletic ac-
demarcated from the disk and a distinct byssal notch. These fea- quisition of the next morphological grade, for example in the
tures, as well as their calcitic shell microstructures (Waller, 1978, evolution of Halobia from Daonella as recently suggested by

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Bositra
Jur. Halobia
Halobia
U. Tr.
U. Tr.Daonella
Aparimella 6
S 5 Daonella
Enteropleura 4
M. Tr. 3L
Enteropleura
..2
L. Tr.
Posidonia
Bositra
Perm.
Posidonia
Carb. Caneyella
Dev. Caneyella
FIGURE 5-Schematic phylogeny an
Gregorio, 1886, Enteropleura Ki
Paleozoic ancestors. Stratigraphic
McRoberts (2000,
2. The principal apomorphies p.at Node 2 are600).
the loss of the du- Fig
morphological changes
plivincular in
ligament system and the anterior adductor. Begin- this
ogram. The synapomorphies
ning in the lower Olenekian (base of the upper Lower Tri- fo
assic), bivalves previously referred to Posidonia, then placed
1. Posidonia: Posidonia is rega
in a new genus Peribositria by Kurushin and Trushchelev
superfamily Posidonioidea. Its
(1989), and herein regarded as belonging to Bositra, have an
bimineralic prismato-nacreou
alivincular ligament with a single, rather broad resilifer and
ductor musculature (Weigel
a single adductor muscle (the posterior adductor) that is shift-
morphic characters, probably
ed more centrally and dorsally than the corresponding muscle
cestor such as the Lower Carb
apomorphic of Paleozoic Posidonia. (Compare figures
features of in Kurushin Posido
and
Trushchelev, 1989, with reconstructions of muscle scars
with a slightly sinuate or flat
early ontogeny shown by and Weigelt, 1922; seeornamen
also Fig. 5.) Bositra continues
ridges that the rounded, oval, commarginally
corrugate the corrugated thin shell of
ultra-
byssal gape and Posidonia,
sinus the principal differencein in shape
matur appearing to be
an obligate the absence of any anterior sinus
byssate or flattening in early ontog-
ancestor
Roberts and eny. In terms of function, these
Stanley, changes probably signify
1989, forthe
tached specimens further decline or
in loss of obligate
the byssal attachment
Missis in early
with obligate ontogeny and a recliningattachmen
byssal mode of life. Etter (1996) recently
later ontogeny reviewed the controversies regarding the lifestyle
Posidonia of Bositra
was
substrates with and providedonlyconvincing evidence that it was benthic, not
occasion
ment. A vexing pseudoplanktonic
problem or nektoplanktonic. in p
rise to Lower Triassic Bositra is the absence of authentic 3. At Node 3, Enteropleura retains an alivincular ligament sys-
specimens of Posidonia in the Permian (Dickins, 1963, temp. very similar to that of Bositra (based on the only known
71). illustration of its ligament, by Arthaber, 1896, p. 194, fig. 12)

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
as well as a single adductor

function muscle
of the in a position
shell tube o
that of Bositra, Daonella,
gape and
relatedHalobia. The shel
to feeding by
of Enteropleura is intermediate
possibly to between
bring in that
hydroof
teria
Daonella in the sense that in(Seilacher,
Enteropleura 1990, p. 30
comm
rugations are retained in early
byssal ontogeny,
gape but fine
that served to
tellae begin later, increasing
bivalve by intercalation
attached to a benth on
through the remainder J. of Campbell,
ontogeny. 1994, The mostp. 59, 1
sign
morphy at this point of is the
thiselongation
controversy of is
the
beyohi
expanded and flattened D-shaped
former shell, characte
hypothesis for the
shared by Enteropleura, not Daonella,
a locus for and Halobia.
chipping or
flat shell is regarded bebyexpected
many as ifa it
"snowshoe
were the s
(Thayer, 1975) to prevent sinking
functional into soft
byssus. sedi
b) There
Wignall, 1993, p. 195;oration
1994, p. of 37-38).
the mantle edg
4. At Node 4 there is another
that appearsshift to in beligament
well re
Daonella, Aparimella, and Halobia1994,
Campbell, the single
p. 59). resi
Fu
from most or all of ontogeny.
interior of(None of the
the byssal tubesp
figures of these generathethatattachment
I have seen sites for r
are suffic
similar
preserved in early ontogeny toattachment sites in
permit determinin
resilium was present beneath
retractorthe beak
scars in would
that earliestb
before the onset of thedition, would more
horizontally likely
striated lig
ligament areas of theseto genera
the beak.
are c) The evoluti
horizontally
grooved, and the entire of length decreasing of these dependence areas w
occupied by a thin lamellar
tachment ligament in Caneyella, or an onto to
cession of lamellar ligaments.
ontogeny Local
in Posidonia, ligament to stra
lifestyle
occurred either at the ends of the in ligament
Bositra, Apari area o
of "knots of ligament," there probably wouldlamellar be a reversal ligam
than fibrous ligament, evidence
in more central in modern positions pterio (
bell, 1994, p. 57). The formed
important from point apposed is that inner th
systems of these halobiidsgape, were can be not formedduplivincula to acco
mantle margins
they multivincular as sometimes stated near (H. J.the Cam h
p. 57; McRoberts, 2000, figs. p. 599). 60, 63, Rather showing thesethe lig
pear to be neomorphic, having
Iredale, evolved
1939). in concer
e) Taphonom
creased shell surface areathe and modes elongation
of occurrence of theof h
particular group of bivalves.
reviewed by Kauffman
5. At Node 5, Aparimella Kauffman
and Halobia andshare Sageman ligam (1
are somewhat raised above Etter the (1996),
dorsal and margins others. of In t
of benthic
means of small indentations of the opportunism
shell margin i
of the ligament area (H. McRobertsJ. Campbell, (1997, 1994, p. 199), p. 65
ring to these indentationslife habit as "auricles"). for Halobia Aparime shou
a stratigraphic positiontiny,that lies twisted between threads that on of t t
nella and the first abundant
Halobia Halobia. plicosa Mojsisovi There is s
however, whether Aparimella
Campbellis a true
(1994, p.phylogen
48) as po
mediary (McRoberts, It 2000, is equally p. 599). likely, There however is the
perhaps associated
complication that Enteropleura, as shown with in the lif
description of Nevada phur-rich
material, also environment,has a hinge as
dered at each end by a slightIn South indentationCanyon the of the strati do
of the genera
6. The main derived character of Halobia Bositra that (regdis
from the other genera Peribositria),
in this succession Enteropleur is a nar
anterodorsal shell tubefollows
that the succession
begins outlined above: Bositra occurs
near thein the basal
beak a
part of Unit Bmargin.
in diameter to the anterior in the Hyatti Zone, Thereof middle Anisian the age (Sil-
rais
berling and Nichols,
on opposite valves produce a 1982,
small fig. 5). Enteropleura
intervalve occurs in a nar- g
dition, some Halobia haverow zone between
multiplethe beds occupied small
by Bositra below and Dao-
muscle
the dorsal edge of thenella above. This tube
shell is inferred to(H. be the Shoshonensis
J. Campbell, Zone, of late
and pl. 7, fig. 6). The amplitude of the shell restricted
middle Anisian age, because Enteropleura is apparently tube v
ly in the genus, from to this zone in Nevada (Silberling
scarcely developed and Tozer, 1968, p. 37; Nichols
and with
entiated ornament to highand Silberling, 1977, p. 21; Silberling and
amplitude and Nichols,lacking
1982, p. 49). r
A poorly preserved
that are elsewhere present on specimen
the tentatively
shell. identifiedSome
as an Apari- au
referred to the radial mella (USNM 526420)
sector ofwas found
the in the ammonoid that
shell bed above int
tween the shell tube the andcoral beds of Unit Edorsal
the (Fig. 2) with associated
margin ammonoids thatas
(e.g., H. J. Campbell, indicate
1994, a late Ladinian
p. 50; age. TheMcRoberts,
lowest specimens of Halobia 20
This is, however, not thus far found in the section occur
homologous in the Desatoyense
with the Zone,anter
Unit
F, of early Carnian age. The Rather
of alate pterioids and pectinoids. following descriptionsit of South
is Can-mere
of the shell between the yon Posidonioidea
tube are only
and for species
thein the genera
hinge Bositra and tha
guishable only because Enteropleura.
the Daonellatube and Aparimella?
is present. are not well enough rep-
resented by in-place collections to merit further description here.
Two competing hypotheses forcollecting
They require concentrated the mode
through the section.of for

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Family POSIDONIIDAE Frech,Other 1909 [emended]

species.-Probably most of the Lower and Middle Tri-
Type genus.-Posidonia Bronn, 1828 (=Posidonomya assic species that have been referred to Posidonia belong in Bos-
Bronn,
1834). itra. Included among these are the many species described from
Original diagnosis.-Frech (1909, p. 8), as for the superfamily northeastern Russia and Svalbard by Kurushin and Truschchelev
Posidonioidea. (1989) in their new genus Peribositria, regarded herein as a junior
Emended diagnosis.-Subcircular to obliquely oval Posidon- synonym of Bositra (see below).
ioidea with ligament either duplivincular or alivincular; ornament Original diagnosis of Bositra.-De Gregorio (1886, p. 11; my
mainly commarginal; radial ornament, when present, much weak- translation from French): "I propose this subgenus for Posidon-
er than commarginal ornament; hinge line generally much shorter omya without the subauricles like the type (i.e., P. becheri Bronn),
than shell length. but deprived of any cardinal appendage, or at most with a small
Discussion.-As presently construed, three genera are included prolongation on only one side."
in the family Posidoniidae: Posidonia (Late Paleozoic), Bositra Emended diagnosis.-Posidoniidae with a single adductor and
(Triassic and Jurassic), and Lentilla Conti and Monari, 1992 an alivincular
(Ju- ligament.
rassic). As discussed above, the Jurassic genus Posidonotis isOriginal now description of Peribositria.-Kurushin and Trushchel-
regarded as a pectinoidean, and the family placement ofevAula- (1989, p. 59; translated from Russian by R. D. Johnson): "The
comyella is unclear. The family placement of Silberlingia shell Imlay, is small to large in size, thin-walled, equivalved, with an-
1963, which occurs in the Jurassic (mainly Callovian) in Califor- terior and posterior gapes, from rounded to obliquely elongated,
nia, Chile, Peru, and Japan (Imlay, 1963; Damborenea, 1987, weaklyp.to strongly convex with greatest convexity in the beak
163), is possibly another member of the Posidoniidae, where area, middle,
it or ventral parts, and usually with a keel-shaped fold
was placed by Imlay (1963, p. 100). It is distinguished bydelimited its by a gently sloping depression. The beaks are small,
peculiar ornament of coarse commarginal ridges and short, slightly
dis-pointed, swollen, central or off-center, either orthocline
continuous, irregular radial costellae on the external surfaces or somewhat
of prosocline, and weakly projecting over the hinge
the commarginal ridges. The nature of its ligament, however, is margin. The outer surface is with concentric, radial or combined
unknown. Examination of Imlay's type material of Silberlingia ornament.
in The auricles are small, clearly separated from the an-
the Smithsonian collections revealed characters that suggest terior and posterior margins of the shell and are found in the plane
that
the affinities of this genus may lie elsewhere. Silberlingiaofisthe closing of the valves.
dis-
tinguished from Posidonia and Bositra by its large size, a thickerhinge margin is straight and comprises 0.2-0.6 of the
"The
outer calcitic columnar prismatic shell layer with larger length prismof the shell. A wide triangular ligament pit, sloping pos-
diameters on both valves, and a hinge that appears to be entirely teriorly [i.e., prosocline] is found under the beak. The muscle
opisthodetic. The thick prismatic calcite of the type specimens impression
of is of average size, oval and slightly recessed, of vary-
Silberlingia is more like that of associated inoceramids than ing degrees
like of relief, with longitudinal grooves and is situated in
that of associated Bositra. the section around the beak. In the upper half of the shell, up to
The Lower Jurassic genus Steinmannia Fischer, 1886 (pro Au-3 or 4 subconcentric beak-area crura of varying lengths, inter-
lacomya Steinmann, 1881) resembles Bositra in shell form and rupted at the muscle impression, are developed. Sometimes, short
commarginal ornament. Guillaume (1928), however, discovered crura of radial direction are noted in the middle part of the valve
clear evidence for multiple ligament pits on the ligament areas of and on the muscle impression."
three specimens of Steinmannia bronni (Voltz in Zieten, 1833) Occurrence.-Lower Triassic (Lower Olenekian) to Middle Ju-
from the Toarcian of France and suggested that this species may rassic (Lower Oxfordian, Quenstedtoceras mariae Zone according
have more in common with the Inoceramidae than with the Pos- to Duff, 1978, p. 53); mainly or entirely in Boreal realm of Laur-
idoniidae. Assuming that the multivincular ligament of Steinman- asia in Early Triassic, spreading southward on both sides of Pan-
nia is homologous with that of the Inoceramidae, subsequent au- gea and into Tethys by Middle Triassic, becoming nearly cos-
thors have placed Steinmannia in the Inoceramidae (e.g., Cox, mopolitan by the Early Jurassic. Two reports of "Posidonomya"
1969d, p. N320; Duff, 1978, p. 51; Milova, 1988, p. 62). This from more southerly localities in the Lower Triassic are not pos-
assumption, however, may be questioned. Recent studies have idoniids. "Posidonomya angusta Hauer, var." reported from the
shown that the ligament area of inoceramids is underlain by hy-Otoceras beds of the Himalayas by Bittner (1899, p. 2) was re-
pertrophied calcitic prisms that are an extension of the outer pris- identified by Bittner (1899, p. 2) as a new species, Pseudomonotis
matic calcitic shell layer and that the ligament structure is sub- griesbachi, and is a claraiid, not a posidoniid. Species of "Posi-
stantially different from the well-known multivincular ligaments donia" described by Waterhouse (2000, p. 181) from the Lower
of the Isognomonidae (Crampton, 1988; Johnston and Collom, Triassic of the Himalayas also appear to be claraiids.
1998). It seems unlikely that the Steinmannia ligament is really Discussion.-Many authors have lamented the difficulty of dis-
of the inoceramid type. Rather, the relatively few ligament pits criminating posidoniid species, because their ultrathin bimineralic
on the ligament area of Steinmannia bronni (three or four ac-shells are generally poorly preserved, and they are subject to dia-
cording to Guillaume, 1928, p. 221) may be a phylogenetically genetic distortion, particularly where they occur in shaly facies.
independent multiplication of the simple alivincular ligament of Taxonomic difficulty is compounded by apparent high natural var-
Bositra possibly functionally associated with increase in size and iability in shape and ornament and by the tendency for ligament
convexity. areas and internal features to be destroyed during diagenesis.
Although seldom preserved, ligament structure is of great im-
Genus BOSITRA de Gregorio, 1886 [emended] portance for the discrimination of genera. It is duplivincular in
Posidonia, alivincular in Bositra, and multivincular in Steinman-
Posidonomya (Bositra) DE GREGORIO, 1886, p. 11.
Peribositria KURUSHIN AND TRUSHCHELEV, 1989, p. 59, type species nia. The ligament structure of the Jurassic genus Lentilla is un-
Posidonia mimer OBERG, 1877, by original designation, Lower Triassic known. Lentilla is distinguished mainly on the basis of its orna-
(Smithian), Spitsbergen. ment, which consists of thin commarginal flanges that are not
reflected on the shell interior in contrast to the deep commarginal
Type species.-Posidonia ornati Quenstedt, 1856, by subse- corrugations of the other posidoniid genera. It is hypothesized
quent designation (Cox, 1964, p. 47; see also Damborenea, 1987, here that the major shift in ligament structure between Posidonia
p. 163), Middle Jurassic, Braunjura 5, Gammelshaufen, Germany. and Bositra occurred during the Permian-Triassic transition. At

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
this point, however, posidoniids

is a stratigraphic gap. In of any
the Triassic, Bositra type are rar
(=Peribositria) ex-
ian and lowermost Triassic,
tends into the and the
Carnian in the hypothesis
western Tethys (species of "Posi- th
be tested until more material becomes
donia" from the Halobia limestones of Sicilyavailable.
listed by Frech,
The most extensive documentation 1909, p. 461), but no authenticated specimens
of areligament
known from st
muscle scars in Bositra is that of Kurushin and Trushchelev the higher Triassic. In the Jurassic, the oldest Bositra is B. buchi
(1989), who introduced the genus name Peribositria for the Tri- reported from the upper Pliensbachian of the Yukon Territory of
assic species that they studied. They stated that their new genus Canada by Aberhan (1998, p. 78).
name combines the prefix "peri-," meaning near, with the genus The presence of shell gapes in Bositra remains a controversial
name Bositra, with reference to the Jurassic posidoniid that issue. has Jefferies and Minton (1965, p. 164) calculated that anterior
a similar external form. They did not explain, however, why they and posterior gapes are present in Jurassic Bositra buchi based
chose to insert an "i" before the final "a" in the new name. on plaster models trimmed along growth lines. This interpretation
Regardless of their intentions, this insertion avoided the creation contributed to their conclusion, now discredited (e.g., by Kobay-
of a junior homonym of Peribositra Chen, 1981, type species ashi et al., 1966, p. 109; Kauffman, 1981, p. 311; Conti and Mon-
Peribositra baoqingensis Chen, 1981, from the Lower Triassic ari, 1992, p. 205; Hollingworth and Wignall, 1992, p. 25; Wignall,
(basal part of Yinkeng Formation) of China. Chen (1981, p. 81)1993, p. 195; Etter, 1996, p. 335; Etter and Tang, 2002, p. 285),
did not venture to place Peribositra in a higher classification butthat Bositra buchi was capable of swimming and was probably
noted its similarity to the Jurassic posidoniid Bositra, the Paleo-nektoplanktonic rather than benthic. Likewise, Kurushin and
zoic posidoniid Posidonia, and especially to Claraia bioni Nak- Trushchelev (1989, p. 59) stated in their diagnosis of Peribositria
azawa, 1977, from the Permian-Triassic boundary in Kashmir.(see above) that anterior and posterior shell gapes are present in
The last, however, is a pterinopectinoidean in the family Claraii-Triassic species. However, others have rejected the presence of
dae and has a small but distinctly set-off right anterior auricle andgapes in Bositra (e.g., Kauffman, 1981, p. 340; Conti and Monari,
a deep byssal notch. The shell shape of Peribositra baoqingensis,1992, p. 205), and no convincing evidence for their presence was
as well as its commarginal ornament that consists of flanges ratherfound among the Triassic specimens from the New Pass Range
than corrugations, are characters that suggest that it is a claraiid,described below.
not a posidoniid, but the triangular ligament described by Chen
distinguishes Peribositra from other claraiids, which have dupli- BOSITRA FAVRETENSIS new species
vincular ligaments. Chen did not mention whether the valves of Figure 6.1-6.3
Peribositra are equal or unequal in convexity. His figures, how- Diagnosis.--Oblique-oval Bositra with beaks anterior of cen-
ever, suggest that the right valve is less convex than the left. If
ter; fine, regular, nonsinuate, commarginal ribbing beginning early
so, this is another indication that this genus is a claraiid, not a
in ontogeny; lacking radial striae.
posidoniid.
Description.--Shell of small size, reaching a maximum dimen-
Kurushin and Trushchelev (1989, p. 59) said that Peribositria
sion of about 20 mm, equivalved and inequilateral, with beaks
differs from Bositra in having a distinct muscle impression, crura about 30%-49% of shell length from anterior; shape generally
in the beak region, and a ligament pit that is narrower relative to obliquely oval throughout ontogeny with maximum growth gra-
length of hinge. In the present study, Peribositria is regarded as dient tending posteroventrally; height and length about equal in
a junior synonym of Bositra, because none of these differentiating early ontogeny (where shell length is about 3 or 4 mm), generally
characters can be supported. The distinctiveness of the muscle becoming longer than high at shell lengths above 8 mm (ratios
impression depends on taphonomy and diagenesis. The shells of of Ht/L ranging from 0.64 to 1.12, averaging 0.88, n = 12); less
both genera are bimineralic and very thin, and the muscle scar commonly transverse oval, with maximum dimension parallel to
disappears with the disappearance of the inner aragonitic shell hinge. Rounded anterior and posterior margins intersecting hinge
layer in which the aragonitic myostracum of the muscle scar is at oblique angles without auricles or indentations, anterior angle
embedded. By virtue of the several ligament areas preserved in about 130?-140?, posterior angle more oblique, about 150'-160'.
the Russian specimens and illustrated by Kurushin and Trush- Hinge line short, about 40% of shell length and asymmetric, with
chelev (1989), the state of preservation of this material is excep- ratio of anterior hinge to posterior hinge ranging from 0.6 to 0.9,
tional. Such muscle scars are not preserved, or at least were not forming a low, broad ligament area between straight hinge line
described, in Triassic "Posidonia" from other regions, including and sloping dorsal margins. Beaks slightly prosogyrous to ortho-
the New Pass Range. The "beak-area crura" of Peribositria are gyrous, not greatly inflated, and extending only slightly above
almost certainly another diagenetic effect and appear to be merely dorsal margins; prodissoconch not preserved but probably small
irregular wrinkles caused secondarily by compaction and distor- (less than 1 mm in height). Byssal notch or sinus absent; shell
tion. They are not present in the New Pass material, nor are they gapes not evident, probably absent.
present on the Siberian posidoniids illustrated by Dagys and Ku- Ornament consisting of regular closely spaced commarginal
rushin (1985, pls. 19, 20, including the type species Posidonia ridges that affect entire thickness of the thin shell; first ridge form-
mimer). Finally, the idea that the ligament pit of Peribositria is ing at height of 0.5-1.5 mm.; earliest ribs closely spaced (about
narrower than that of Bositra is not supported by sufficient evi- three to six ribs per mm in central sector of shell at shell heights
dence. Kurushin and Trushchelev (1989, p. 58) stated that in Bos- of 1-3 mm), becoming more widely spaced in later ontogeny
itra the length of the ligament pit is 45% of the length of the (about 1-2.5 ribs per mm in central sector at shell height of about
hinge, while in Peribositria this ratio is only 35%, but they gave 10 mm), commonly increasing in number by intercalation in cen-
no statistical evidence. Apparently the only available illustration tral part of shell, rarely increasing by branching and rarely or-
of a Bositra ligament area is that of Jefferies and Minton (1965, dered, with higher amplitude first-order commarginals separated
fig. 1 and pl. 19, fig. 9), and it shows a ligament pit of about the from one another by one to four second-order commarginals.
same dimensions as in the types species of Peribositria as illus- Interior features: ligament area broadly triangular; ligament pit
trated by Kurushin and Trushchelev (1989, fig. 2), both sketches and muscle scars not preserved.
indicating that the length of the ligament pit is on the order of Shell microstructure not preserved but preservational style is
47% of hinge length (restored in the case of the figure of Bositra). consistent with an ultra thin bimineralic shell; outer layer presum-
The only remaining reason to separate Peribositria from Bositra ably originally calcitic but trace of prisms or laths not detected

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Triassic.
or not preserved; inner layer generally Halobia
absent, rarely developed
represented from D
ically or polyphyletically.
by recrystallized material and presumably originally aragonitic. In additio
Etymology.-Named after the Favret teropleura Formation. and Dipleurites, of whic
Types.--Holotype, USNM 526421 (Fig. 6.1).
Germanic Paratypes:
Triassic. about
Both represent o
60 valves and numerous fragments Daonella."occurring on same block as
holotype and in clusters on rock Emended pieces numbered
diagnosis.-Monomyarian USNM
Posidonioidea with broad
526422-526433. All from lowerlyUnit expandedB "Posidonia
shells of low-convexity,beds," hinge long,South commonly near
Canyon (Locality 1). ly as long as shell length, ligament alivincular or horizontall
Measurements.--Holotype height 17.0
striated, mm,plicae
commarginal length limited to 15.5 mm. radial cos
early ontogeny,
Occurrence.-Known only from tellae,Favret
if present, Formation
stronger than commarginal (=Unit B in late on
ornament
in Fig. 2), Hyatti Zone, middle togeny.
Anisian, of the New Pass Range
and Humboldt Range, Nevada. Discussion.-The emended diagnosis for the family Halobiidae
Discussion.--As previously discussed,
accommodates Enteropleura posidoniids in shales
as well as Daonella and Halobia in
and mudstones are subject to distortion of
the broad sense (see form.
preceding discussionWhen rock
of the superfamily Pos-stress
idonioidea).
is perpendicular to specimen height, the shells are compressed
along an anterior-posterior axis, leading to a deceptively high oval
form; just the opposite occurs when Genusstress
ENTEROPLEURA is parallel
Kittl, 1912 [emended] to shell
height, the result being a transversely elongate oval form. The
Enteropleura KITTL, 1912, p. 162.
description of Bositra favretensis is based on specimens that ap-
pear to be the least distorted.
Type species.-Daonella
The two species of Triassic "Posidonia" orguembeli Mojsisovics, 1874, by sub-that
"Peribositria"
sequent designation
resemble Bositra favretensis most closely occur (Diener, 1923, p. 52),inLowerthe
Muschelkalk of
Triassic
Germany and Hungary, Middle
Boreal region of the Svalbard Archipelago and Triassic.
Siberia. Posidonia
backlundi Wittenburg, 1910, which Other species.-Enteropleura
occurs in bittneri
the Kittl, 1912 and E. spe-
Lower Olene-
cies a (described
kian substage of the Lower Triassic in below).
both Siberia and the Sval-
bard Archipelago (Kurushin andOriginal diagnosis.-Kittl (1912, p. 162;
Trushchelev, 1989, my translation
p. 62),from has
German): "This genus
regular commarginal ridges throughout shows generallybut
ontogeny in its shape the outline from
differs
and the flat shape
B. favretensis in that its commarginal ridgesof Posidonia.
are The surface
coarser ornament isand
some- more
widely spaced and its shape more times weak, sometimes strong.
circular. In Somewhat
addition,behind the shellP.
centerback-
lies an internal short shell
lundi reaches a larger size (50 mm) and has fine radial striae ridge extending from the beak. This on
latter is recognizable
the distal part of its shell. Posidonia sibirica on steinkerns as a furrow. The posterior
Kurushin, 1980 is
triangular field
also similar to B. favretensis. Like P. isbacklundi,
somewhat swollen and clearly P. defined.
sibiricaThe ge- is
nus Enteropleura certainly resembles
more circular and has coarser commarginals than Daonella,B.fromfavretensis.
which it dif-
fersstriae
In addition, P. sibirica has radial by the internal shell ridge.
that areSo far more
only two forms belonging
extensively
developed than in P. backlundi to and this genus
its are commarginals
well known. Both originate fromtend the Alpine to be
Muschelkalk."
sinuous (Dagys and Kurushin, 1985, pl. 21, figs. 6-10). P. sibirica
occurs in the Olenekian stage inEmended
central diagnosis by Ichikawa.-Ichikawa
Siberia and the (1958, p. 190; my
early
translation from
Spathian of arctic Canada and British German): "A subgenus
Columbia of Daonella with
(Kurushin and the fol-
Trushchelev, 1989, p. 65). lowing characteristics: Radial ornament of the Daonella type only
Posidonomya stella Gabb, 1864weakly
wasdeveloped,
describedthreadlike,from
and appearing
Staronly Can-
in later growth;
yon (or Star Creek Canyon) on shell
thesurface
east from the juvenile
flank of stage
the to Humboldt
the early mature stage
Range of west-central Nevada butonly concentrically
from anornamented unspecified and similarhorizon.
to Posidonia; the
Although Smith (1914, p. 9-11)straight edge of hinge
included relatively short; beak stella
Posidonomya among most inspecies
approximately
his list of Middle Triassic fossils from at Star
the frontCanyon,
end of the edge he
of thelater
hinge margin."
(1927, p. 113) gave the occurrenceEmended
of thisdiagnosis herein.-Halobiidae
species with an alivincular lig-
as "presumably
ament, hingeunknown,
in the Upper Triassic, definite horizon shorter than shellof
length,
Starand ornamentation
Canyon,that is
intermediate between that of Bositra and Daonella.
West Humboldt Range, Nev." He repeated Gabb's original figure
Occurrence.-Apparently
of the species and stated in the explanation of the limited
figure to the (pl.
Middle Triassic
104,
fig. 10), "Pseudomonotis zone of (Shoshonensis
Star Peak Zone, formation,
middle Anisian) in Europe
Star andCan-
Nevada.
yon," thereby implying that the Discussion.-Kittl
age of P. (1912) placed two
stella is species
Norian, in his new
not genus,
Middle Triassic. The type specimen"Enteropleura
(or type Giimbeli (Mojs.)"
lot) ofand "Enteropleura Bittneri
Posidonomya
n.n.," and of
stella is presumably at the Academy suggested
Naturalthat a third species that he
Sciences of described,
Phil-Dao-
adelphia, because Richards (1968, nella lamellosa
p. 86) Kittl, listed
1912, may alsothe belong in this genus.
species asHe
"30789-Holotype? Type lot." As regarded
of thisthe internal shell ridge as an
writing, important defining
however, character
this
specimen or lot cannot be located of Enteropleura, allowing it to be and
at the academy, distinguished from Daonella,
a compar-
but did not mention
ison of P. stella to Bositra favretensis must the important
depend findingonof Arthaber
Gabb's (1896, p.
194,that
rather stylized figure. It indicates fig. 12) that
in "Posidonomya
comparison nov. spec."
to (the
thespecies
Newthat Kittl
named Enteropleura
Pass species, P. stella is substantially more bittneri)
gibboushas an alivincular ligament.
and inequi-
lateral, with a more inflated beak and much
Krumbeck broader
(1924, p. 213-214) determinedand thatlower
the internal
commarginal ridges. ridge described by Kittl (1912) as characteristic of Enteropleura
is likely homologous with the anterior of two ridges present in
Family HALOBIIDAE Kittl,both 1912 [emended]
Daonella and Halobia. The two ridges arise on the anterior
Original definition.-Kittl (1912, p. 42;
and posterior my
edges of thetranslation
impressed adductor scarfrom beginning in
early ontogeny
German): "This family includes three and track the muscle
apparently insertion as it moves distally
phylogenetically
related genera, Posidonia, Daonella, and
with ontogeny. He Halobia.
surmised that the The first
posterior ridge is rep-
also present
resents the long-lived stem from which but
in Enteropleura Daonella
was missingbranched
or indistinct in thein steinkerns
the

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
4.4
4 Z.
:pit
rV~i
P-4"
Itti
a?22
4:: ~
;,!"V _R n;
NMN~?
?' Y~Pl
FIGURE 6-1-3, Bositra favretensis n. sp., Unit B (Favret Formation), Hyatti Zon
of left valve; 2, paratype, USNM 526423, composite exterior mold of left
Enteropleura species a, Shoshonensis Zone, Wildhorse mining district, Locality
in 5, 6, and Figure 7, and (on opposite side of block) Figure 8.1; 5, 6, com
diagenetically raised area of adductor scar, arrows point to posterior auricles.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
examined by Kittl. For this reason, Krumbeck

curved and passing tangentially into hinge advised placing
margin, posterior mar-
Enteropleura in the synonymy gin of Daonella.
straight or truncated throughout ontogeny where it crosses the
posteriorhave
Some subsequent studies, however, radial trough, then rounding sharply
continued to andrecognize
passing tan-
Enteropleura as a valid taxon, either as adorsal
gentially into posterior subgenus of absent;
margin. Anterior auricles Daonella
(e.g., Ichikawa, 1958, p. 190; De Capoa
small Bonardi,
posterior auricles 1970,
present, very narrow, p. disk
separated from 43) or
as a full genus (e.g., Cox, 1969e,
by narrowp.
sulcus,N344).
posterior marginIchikawa
of auricle forming very(1958,
obtuse p.
190-191) and De Capoa Bonardi angle
(1970, p.margin,
with its dorsal 43)angle included in
subtended by dorsal Dao-
margin
nella (Enteropleura) the entire group and base of auricleof only"daonelliform
3o-7', surface lacking ornament butposidon-
bear-
iids" of Kittl (1912, p. 31), its ing members distinguished
a radial fold on its dorsal only by
side that probably reflects an internal
their weak radial ornament. linear outer-ligament groove. Hinge line short, its length about
In the present study, Enteropleura is regarded as a valid taxon one-third of shell length, and commonly asymmetric, with pos-
at genus rank primarily because of the presence of an alivincular terior hinge longer than anterior hinge (ratio of anterior hinge
ligament, as demonstrated by Arthaber (1896) and overlooked by length to posterior hinge length ranging from 1.02 to 1.24). Prod-
subsequent authors. This and other characters present in Entero- issoconch large and triangular, length about 2 mm, with probable
pleura fit logically into the phylogeny discussed above and shown small Prodissoconch I stage and large Prodissoconch II stage. Ex-
in Figure 5. The muscle-bordering internal shell ridges shared by ternal surface of dissoconch, not including posterior auricle, di-
Enteropleura, Daonella, and Halobia comprise a synapomorphyvisible into three contiguous radial sectors ("triangular fields"):
for these taxa, but the neomorphic horizontally striated ligamenta) central sector, bounded by anterodorsal sector and posterodor-
is a synapomorphy only for Daonella and Halobia. The alivin- sal trough, having fine, cancellate commarginal lirae and radial
cular ligament of Enteropleura, as well as its Posidonia-like com- costellae over most of its surface except near the trough, sides of
marginal ornament in early ontogeny, are carryovers from ansector forming angle of 1059-1160; b) anterodorsal sector distin-
ancestor in the genus Bositra. A word of caution is in order, guished only by lack of radial costellae, its ventral margin sub-
however, because thus far Arthaber's illustration of an alivinculartending an angle of 22'-29' with dorsal margin; and c) postero-
ligament in Enteropleura bittneri is the only observation that this dorsal sector slightly inflated (outwardly convex), lacking radial
type of ligament is present. No ligament observations are avail-ornament but having commarginal lirae, and bordered ventrally
able for the type species, Enteropleura guembeli. by a broad, shallow radial trough that diverges from dorsal margin
Enteropleura, as emended herein, includes three species: E. at an angle between 360 and 43'.
guembeli, E. bittneri, and E. species a, described below. These Ornament: Fine sharply rounded but not lamellose commargin-
species all occur in a narrow stratigraphic interval, the Shosho- al lirae beginning near the prodissoconch/dissoconch boundary,
nensis Zone, of early Middle Anisian age (Bucher, 1992). Untilabout seven to nine per millimeter in first few millimeters of
detailed population studies of these species are carried out, how- ontogeny of central sector, increasing in spacing to two or three
ever, it is not possible to say whether there is any restriction of per millimeter at shell heights between 5 and 12 mm, and finally
particular species to particular subzones. Bucher (1992, p. 428)to a spacing of 0.7 mm near the margin at height of about 20
observed that Enteropleura cf. bittneri (=Enteropleura species amm. Radial costellae beginning gradually in early ontogeny, be-
herein) occurs throughout the four subzones that he recognizedcoming well developed by height of 5 mm, increasing in number
in the Shoshonensis Zone in Nevada. However, in its type area at distally mainly by intercalation but with some branching to pro-
Gro8reifling, Austria, E. bittneri occurs in the older of two faunasduce a density of radials of about 4 per mm along midventral
in the Shoshonensis Zone (the Rahnbauerkogel fauna) and is ap-margin at shell height of 15 mm. Commarginals and radials about
parently not present in the younger (Tiefengraben) fauna, which equally developed in middle to late ontogeny, producing a fine
correlates with the latest Shoshonensis Zone (Bucher, 1992, p.reticulation. Broad, shallow, commarginal undulations also pre-
429). E. guembeli is also reported from the Shoshonensis Zonesent, irregular in extent but generally of greater amplitude on
(the Balatonites beds of western Hungary; Mojsisovics, 1874, p.posterior half of shell than on anterior, and variably spaced.
8; Kittl, 1912, p. 163), but the species is based on poorly pre- Internal shell features: Monomyarian, with posterior adductor
served material and may prove to be synonymous with E. bittneri.scar oval in shape, slightly higher than long, about 5 mm in di-
The systematic position of Daonella lamellosa, which Kittlameter in holotype, its dorsal margin only 3.5 mm from hinge
thought may possibly be an Enteropleura, has not been reevalu-line and its center slightly posterior to a line perpendicular to
ated. It was described from "Hang des Orlovica, Ost von Plavnohinge and passing through beak. The scar originates in a distinctly
bei Knin, Dalmatien" but its stratigraphic position within the An-posterior position on early dissoconch just inside boundary of
isian is not known. prodissoconch and when first formed is bordered along its dorsal
side by a variably developed shell ridge that is inclined antero-
ENTEROPLEURA species A dorsally and forms a distinct short groove near beak of most
Figures 6.4-6.6, 7, 8.1-8.3 steinkerns and composite exterior molds; adductor scar migrates
ventrally and anteriorly during ontogeny and comes to occupy a
Enteropleura cf. bittneri KITTL, 1912. SILBERLING AND NICHOLS, 1982, position that is relatively more central and dorsal; track of ad-
p. 49.
ductor scar bordered on both its anterior and posterior sides by a
single low, narrow, radial ridge, the anterior one more prominent
Diagnosis.--Enteropleura with finely cancellate ornament and than the posterior, both ridges terminating distally along anterior
a truncated (straight or slightly insinuated) posterior margin and posterior borders of adductor scar. Anterior adductor scar ab-
throughout ontogeny. sent; other muscle scars not detected.
Description.--Shape: Shell up to about 30 mm in length, D- Shell microstructure: Outermost shell layer very thin, probably
shaped, longer than high (height/length ratio 0.7-0.8), of low con- originally calcitic, represented in composite molds only by im-
vexity; beaks orthogyrous or slightly prosogyrous, projecting pression of exterior surface, this suggesting a very fine columnar
slightly above dorsal margins and positioned posterior to center prismatic structure with prism diameters less than 20 Im. Ad-
point of shell length in early ontogeny but anterior to center point ductor scar differentially dissolved, suggesting an originally ara-
in late ontogeny; dorsal margins descending slightly from beak, gonitic myostracum. Recrystallized inner shell layer thicker than
ventral margin broadly curved, anterior margin more sharply outermost layer in umbonal region, covering umbonal anterior and

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Institute of Paleontol
to locate it. Based on A
the new species are si
session of a small, na
there are distinct dif
are less prosogyrous a
Throughout dissoconc
vada species is more r
truncated or even sli
margin is evenly roun
sector, on which radi
species, is broader in
Kittl (1912, p. 164) r
losa, from the Dinari
semblance to E. bittn
shapes, posterior tr
spaced commarginal li
principal difference i
costellae, a feature th
cies. Unfortunately, K
on incomplete specim
species remains poorly
cies to Daonella rather
establish the presenc
above, the presence o
FIGURE 7-Enteropleura garded species as a, important
Shoshonens i
of repaired injury of left?Enteropleura valve shownguembe in Figu
dimples on edge of repaired
Nevadamargin. speciesScale in hav bar
a more circular shape
The description of En
thinning distally around on specimens dorsal and present latera
ductor scar, fading out "lithographic" toward shell limestmarg
structure detected on (USNM inner surface 526434, of Fig. this 6
solution of the adductor this scar block relative are ammon to th
gests that the innerBalatonites
shell layer was Mojsisov origin
Material examined.-The another best-preserved
is a partial cm
valves, three left valves,
Assereto, and numerous 1966, proba fr
block of black fine-grained
1905), with limestone a whorl (U h
6.6, 7, 8.1) from themonoid Wildhorse conchs mining (2-6 di m
southern Augusta Mountains.
also present. Other Only specisin
MCZ Invertebrate Paleontology
present, with Catalogue
right va
which is a left valve terior adhering orientation to a parat of
nevadanus Hyatt and ative to Smith,
the substrate is not. Almost all of the valves said
1905, have their to
see discussion of this specimen
outer surfaces facing in the same direction,bybutSilberl because the ori-
p. 49), and USNMentation 526435 of the block was not andrecorded, it526436,
is not known whether
between the Hyatti and
the valves Rotelliformis
were concave-up or convex-up on the sea bottom. The Zo
Canyon (Locality 1) (Fig.
valves lack epifauna8.3).
and only one specimen displays a substantial
Occurrence.-Favret Formation,
repaired injury (Fig. 7). A piece of shell was broken Shosh from its
middle Anisian, Nevada.ventral margin, probably by a biting predator with cone-shaped
Discussion.-The new species
sharp teeth or denticles, because is left
an arcuate row of unna
small conical
the preparation of this
dimples is present on manuscript
the shell just above the part that was itre-
McRoberts and a studentmoved. The mantle submitted
was also damaged as indicated for by the direc-p
describing the same tionstaxon
of shell regeneration,basedwhich were from on new
the sides of the in- m
lected from the Wildhorse
jury toward its center, miningthe newly formed shelldistric meeting medially
sonal commun., 2003). to form a radial suture.
The new species is closest to Enteropleu
from the Rahnbauerkogel Superfamily BAKEVELLIOIDEA
level of King,the1850 An
at GroBreifling, Austria Family(seeBAKEVELLIIDAE King, 1850
Summesber
for description of this section). Genus GERVILLARIAThe Cox, 1954 holoty
was figured by Arthaber Subgenus (1896,
BARYVELLIA new subgenus p. 194, f
found and is possibly lost because
Type species.-Gervillaria (Baryvellia) ponderosa of n.wa sp., by
H. Summesberger, monotypy.
Natural History Muse
commun., 2003), Arthaber's collection
Diagnosis.-Elongate, scimitar-shaped, nearly equivalved, very w
university collections, with
thick-shelled Gervillaria with longone
finger-shaped part
anterior auricle of t
at the University comprising
of one-fourthVienna. K.
to one-third of total shell Rauch
length; posterior
2003) kindly searched for
auricle prominent, the
extending specimen
nearly to posterior margin of shell in

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
and having a posterior margin that is outwardly

denticulations are apparently concave missingwith on theaanterior
of the
pointed dorsal extremity; ligament anteriorprosocline,
pits auricle. In Gervillanceabroadly
the teeth beneath the beak
trian-
are enlarged
gular or rhombic, widely separated, and anduneven
resemble cardinalin teeth,
size andand
the posterior
spac- den-
tition
ing on both anterior and posterior consists of long,areas,
ligament linear teeththe
nearly pit
parallelbeneath
to the hinge line
(Skwarko, throughout
beak larger than others and persisting 1967, p. 54). ontogeny.
Etymology.--Prefix baro from Greek There is a strong
barys,superficial resemblance between
meaning heavy, Gervillaria
combined with -vellia, common (Baryvellia) suffixand in names
Gervillia ofa Middle
joleaudi, bakevelliid
Triassic western
genera. Tethyan species known from southern Spain, Algeria, and Jordan
Occurrence.--Known only from the Ladinian of Nevada in (Schmidt, 1936, p. 53; Lerman, 1960, p. 34). This Tethyan species
Units D and E of South Canyon, New Pass Range, the Grantsville has a similar narrow, elongate shape and large size (up to 20 cm
Formation in the Shoshone Mountains, and the "Excelsior For- in length), a large narrow anterior auricle, and a bimineralic pris-
mation" on the upper plate of the Gillis thrust, Gillis Range. mato-nacreous shell that reaches enormous thickness (see shell
Discussion.--In his original description of Gervillaria, Cox microstructural study by Martinez and Mirquez-Aliaga, 1994).
(1954, p. 49) emphasized the oblique form, high inequality ofThe similarity extends to stratigraphic occurrence, because G. jo-
valves, obtuse posterior wing, pointed anterior wing, greater in- leaudi is known mainly from the Ladinian. Schmidt (1936, p. 55),
curvature of the beak of the left valve compared to that of the in fact, regarded it as a good marker fossil for the middle of the
right, and a mature hinge with numerous uneven transverse cren- Longobardian stage, late Ladinian, in southern Spain and northern
ulations. In an emended diagnosis, Muster (1995, p. 50) added Africa, although Lerman (1960, p. 34) later found this species in
that the valves are twisted, the posterior auricle large, and thethe Anisian (Beneckeia Zone) and lower Ladinian of Ramon, Is-
transverse crenulations continuous on the hinge plate from onerael. There are, however, important differences between G. jo-
end to the other. Twisting of the plane of commissure is well leaudi and the type species of Gervillaria (Baryvellia). The an-
marked, however, only in the type species, Gervillaria alaeformisterior auricle of G. joleaudi is prominent but not as long, being
(J. Sowerby, 1819) (see Muster, 1995, pl. 8, fig. 2a-c). This also only one-fifth to one-sixth the total length of the shell (Schmidt,
applies to valve inequality, which again is well marked only in 1936, p. 54; Lerman, 1960, p. 34). The dentition of G. joleaudi
the type species (see figures in Muster, 1995). Although the pos- consists of inclined dental ridges beneath the umbo that become
terior auricle was said to be "oblique" by Cox (1954, p. 49),longitudinal elongate teeth posteriorly, and there are several lig-
referring to the trend of the posterior margin of the auricle, thisament pits of more or less equal size in the umbonal region rather
applies only to the ventral part of this margin. As this margin than a single very large pit.
inclines dorsally, it sweeps around in a distally concave arc to In spite of the shared presence of a giant anterior auricle in
produce a rounded, distally pointing extremity adjacent to theGervillaria (Baryvellia), Gervillancea, and Gervillia joleaudi,
dorsal margin. This feature is even more accentuated in earlytheir morphological differences and widely separated geographic
ontogeny, where this auricle is distinctly pointed in most members occurrences (Nevada, Papua New Guinea, western Tethys) sug-
of Gervillaria. The narrow byssal gape mentioned by Cox (1954, gest evolutionary convergence. Gervillaria (Baryvellia) is prob-
p. 49) applies to a shallow, obscure sinus on the anteroventralably endemic to eastern Panthalassa and evolved as a sister group
margin posterior to the point where the anterior auricle joins theof Gervillaria s.s., which was already present in the Middle Tri-
body of the shell. assic, e.g., Gervillaria hartmanni (Miinster in Goldfuss, 1834)
The new subgenus Baryvellia shares a number of characters(Muster, 1995, p. 58). Gervillancea coxiella and Gervillia joleau-
with Gervillaria (Gervillaria) Defrance, 1820. The shape and sizedi, because of their well-developed posterior longitudinal teeth,
of the posterior auricle of Baryvellia is remarkably like that ofare probably both derived from Bakevellia (Bakevellia) King,
Gervillaria s.s. Both also show differences in the convexity of1848. Schmidt (1936, p. 55) regarded Gervillia joleaudi to be
the valves in early ontogeny, with the left valve more strongly closely related to Gervillia albertii Credner, 1851, of the Lower
convex than the right and with strong interumbonal growth pro-Muschelkalk of Dalmatia, where the anterior auricle is already
ducing broad ligament areas. In both taxa, the left ligament area somewhat enlarged, the shell substantially thickened, and the
is broader and meets the plane of commissure at a steeper anglebody shape elongated. The generic assignment of these Tethyan
than the right ligament area. Both also share a similar pattern ofspecies, however, remains uncertain. Muster (1995, p. 44) regard-
hinge denticulation, with relatively short transverse denticulationsed G. albertii as a junior synonym of Pterinea polyodonta Strom-
along the entire hinge plate of mature individuals. beck, 1849, which she placed in the subgenus Bakevellia (Bak-
Gervillaria (Baryvellia) differs from Gervillaria s.s. in havingevelloides) Tokuyama, 1959, but it seems unlikely that Gervillia
a much longer anterior auricle, the midventral margin of this au- joleaudi belongs in this subgenus.
ricle nearly parallel to its dorsal margin rather than inclined. The
shell body of G. (Baryvellia) is much narrower and more elon- GERVILLARIA (BARYVELLIA) PONDEROSA new species
gate, and rather than having multiple ligament pits in the umbonal Figures 8.4-8.9, 9.1-9.4
region, there is a single, large pit that is larger than the unevenly Diagnosis.-As for subgenus.
spaced pits that form on both the anterior and posterior ligament Description.-Shell of large size, narrow, and greatly elongat-
areas.
ed, scimitar-shaped, up to 23 cm in length, with line of maximum
Gervillancea Skwarko, 1967, a monotypic genus based
convexity on beak
from Ger- to posterior margin forming an angle of 100-
villancea coxiella Skwarko, 1967, from the Upper 200 with the horizontal
Triassic (Car-axis and beaks at about one-third of total
nian-Norian) of Papua New Guinea, also has a giant anterior
shell length au-
from anterior margin; anterior auricle very large, fin-
ricle, which may reach a length of 6 cm in matureger-shaped, with nearly parallel dorsal and ventral margins and
specimens.
Other features of Gervillancea, however, clearly distinguish
rounded it
anterior margin; posterior auricle also large, with its pos-
from Gervillaria (Baryvellia): 1) both the plane of margin
terior commissure
dorsally pointed, centrally concave toward the pos-
and the ligament areas are highly twisted; 2) rather
terior, andthan being
ventrally slanted posteroventrally to merge with pos-
widely separated by interumbonal growth, the umboterior margin
of the of shell body. Mature shells with valves of nearly
right
valve contacts the umbo of the left valve, creating
equalaconvexity,
shallow withfur-maximum convexity in dorsal third of shell
row; 3) the right valve is smaller than the left valve,
body; beaksits distal
prosogyrous, left beak more inturned than the right,
margins nesting inside those of the left valve;beaks
and separated
4) transverse
by extensive interumbonal growth that produces

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
4A, pf
.yx, '?gf
TZ'R
'i iO'i
:?
?qii8'7rm
7,1-
? W. 1NO
PIN,?-
IN I
r'-i?_rvn
-'4, P, B
?:-:~q77
ME IM:??
~C::.:L' .':'~tl; 41;
4 `1 IF It11 ? ? :-- ?
;vi
. e?E
?/41
4 tt;
- 4 (t,
?kA
?IN' 34, e`:?
we ?,?
"V .t
Lii9?~rS$
11 on
4N 10M
FIGURE 8-1-3, Enteropleura species a: 1, interio

showing well-preserved adductor scar and ridge
valve impressed on flank of a paratype of Intor
scale bar 5 mm; 3, USNM 526435, exterior of l
ponderosa n. subgen. and sp., Unit E ammonoid
cm) and interior (scale bar 2 cm) views showing
6, paratype, USNM 526438, left valve steinkern w
to shell interior, scale bar 2 cm; 7, paratype,

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Occurrence.-Known
wide ligament areas, the left ligament area forming only from thea Middle
higherTriassicangle
(Ladinian)
with the plane of commissure than of western
the Nevada.
right; umbonal ridge sharp-
Discussion.-No otherlower
ly rounded in young individuals, becoming bakevelliidsand
are known to attain com-
fading out
along posteroventral trend toward parable size and shell thickness
posterior shell margin. while having such Byssalan elongate
gape not evident except for a shallow form with both sinus in growth
an enormous anterior auriclelines on pointed
and a large
anteroventral side of shell body posterior well posterior
auricle. The preceding to the sulcus
discussion that com-
of the subgenus
separates the anterior auricle from pared the"Gervillia"
shell joleaudi
body. of the European, North African, and
Ligament multivincular with large, Israel Middle Triassic, as well
widely and as Gervillancea
irregularly coxiella from the
spaced, broadly triangular to rhombic, Upper Triassicprosocline
of Papua New Guinea, ligament
concluding that pits,
these forms
the largest beneath the beak, two or three
represent evolutionarymore on anterior
convergences from disparateand bakevelliid
posterior ligament areas. Hinge plate ancestries. narrow along anterior auri-
cle, with short, transverse, irregular The morphological
crenulations description of commonly
the new species is based in- on
clined anterodorsally at a high aangle set of specimens,
to hinge no one of which
line; is complete.
hinge The specimen
plate
beneath beak enlarging ventrally with in the most complete
a short, shell outline is USNM
ventrally curving526438 (Fig.
arc8.6),
that accommodates longer and more linear
but it is mainly crenulations
a poorly preserved steinkern with that in-
the posterior
tersect hinge line at a lower angle; posterior
auricle obscured by matrix. The hinge
best-preservedplate again
exteriors of nearly
with high-angle short crenulations complete anterior auricles to
continuing are on USNM 526439 and
posterior end;USNM
posterior longitudinal hinge teeth526445 (Fig. 8.7, 8.9), but these specimens, a right valve and a
absent.
Ornament consisting of irregular, left valve, commarginal,
are missing the greater part flat-lying
of their shell bodies. The
growth lamellae that continue onto maximum estimated length
auricles; of 23 cm andauricle
posterior maximum shellwith thick-
low, rounded, distally concave, ness widely
is based on spaced
USNM 526441, rugae;
an articulatedradial or-
specimen fractured
nament absent throughout ontogeny. longitudinally and obliquely through both valves and with its dor-
Posterior adductor scar large, oval, sal side eroded (Fig. 9.1). The
slightly descriptionlocated
raised, of differential at valve
about two-thirds the distance from convexity, beakbeak coiling, and differing angles
to posterior of interumbonal
margin of
shell, with long axis of scar inclined growthanterodorsally,
of ligament areas is based on USNM 526450 (Fig.
possibly con- 9.2),
joined with a small muscle scar (posterior representing only a pedal
small part of retractor?) on its
a large articulated specimen. One
dorsal side; another smaller dorsal of the muscle scar
planes of fracture, at distal
however, provides a end of per-
cross section a
low rounded ridge that arises inpendicular umbonal cavity
to the hinge line and planeand fades that
of commissure out passes
about midway between beak and posterior
through both ligament adductor
areas. The description scar; two
of muscle scars is
small muscle-attachment pits (raised based mainly
nodes on theon holotype, USNM 526437. This
steinkerns) specimen, a
present
beneath beak (Fig. 9.3), the ventral right valve,
pitwas chosen to be the
slightly holotype because
larger than it displays
the
dorsal one. Pallial line pitted along both theits inner and outer shell
entire surfaces and
length andnearlyparallel
the complete
to but well inset from ventral margin length of the of hingeshell;
plate as well as the ligament area,
posterior end but of
it is
pallial line joins posterior adductor scar
missing at itspart
the posterior posteroventral
of the shell posterior to the mar-
adductor
gin; anterior end of pallial line joins scar (Fig.
the 8.4, more
8.5). USNM ventral
525454, a steinkern
of the of a right
two valve,
anterior muscle pits. Differences in sizes
displays the pair of and
muscle positions
pits (nodes on the of muscle
steinkern) at the
scars between the two valves not observed. anterior end of the umbonal cavity (Fig. 9.3).
Shell microstructure bimineralic, with a calcitic columnar pris- Many of the shell characters of Gervillaria (Baryvellia) pon-
matic outer layer and a much thicker laminar recrystallized inner derosa invite reconstruction of its living habit following criteria
layer, probably originally nacreous aragonite; maximum shell set forth by Aberhan and Muster (1997) in their study of Jurassic
thickness typically 4-5 mm in mature individuals but increasingbakevelliids from western Canada. The thick shell, nearly equi-
to as much as 15 mm in gerontic individuals. valved form, and ventrally curved ventral margins that are some-
Etymology.-From the Latin ponderosus, meaning heavy or what inturned but not flattened suggest that Gervillaria (Baryvel-
weighty, referring to the large size and thick shell of this species. lia) ponderosa lived on firm but soft sediment in an orthothetic
Types.-Holotype, USNM 526437, a partial right valve with position, that is with the plane of commissure nearly or actually
both inner and outer surfaces exposed, Unit-E ammonoid bed,vertical. As in most bakevelliids, the species lacked a significant
South Canyon (Locality 1). Paratypes: USNM 526438-526448,byssal gape, but a slight bend in growth lines suggests that a
ammonoid bed of Unit E, South Canyon (Locality 1); USNMbyssus was present and anchored the shell along the ventral mar-
526449, Unit D, South Canyon (Locality 1); and USNM 526450- gin at a point lying somewhat behind the beaks. This would sug-
526454, upper plate of Gillis thrust, Locality 4. gest a very low angle between the anterior-posterior axis of the
Collectively this assemblage of USNM specimens catalogued shell and the sediment surface, and also that the long anterior
as holotype and paratypes consists of parts of 12 right valves, twoauricle served as an anterior stabilizing device to keep the shell
left valves, and five conjoined valves. Additional material, from oriented at this angle rather than as a device to penetrate the
the limestone member of the Grantsville Formation in the Union substrate. The considerable biconvexity of this auricle is also in
district, Shoshone Mountains (Locality 4), was examined in the accord with this orientation.
collections of the U.S. Geological Survey in Denver and remains Muscle scars provide further support for this orientation and
in those collections. living position. Although there is still uncertainty, the material
Measurements.-Holotype, unrestored length 166 mm, height examined suggests the presence of three sites for the attachment
59 mm. of pedal/byssal retractor muscles: 1) on the dorsal side of the
hinge crenulations (arrows), scale bar 1 cm; 8, 9, paratype, USNM 526445, fragmented anterior part of left valve, oblique interior view
ligament area with three ligament pits and exterior view, scale bars 1 cm.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
t;S a8dt;
rat:
MNXI,(
llco ts,
A di~J
W-C'A. . . . . . . 11" ??l r
A?
A-~: ?
.......
:i4i
:"": r Owl
ul, 47~
~i~ci .f? a'A
;;? 50
? X~
,?f k;? T',4C
eW,
re?,:
df?z
?f, .47
.?r A
FIGURE 9-1-4, Gervillaria (Baryvellia) ponder

viewed obliquely from right ventral side, am
specimen viewed from dorsal side with anter
mold with umbo and pair of umbonal muscle p
4, scale bar 2 cm; 4, paratype, USNM 526446,
and on anterior beneath beak, a second alig

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
ATRINA SINUATA newof

posterior adductor, interpreted to be the attachment species
a posterior
Figure
pedal retractor; 2) mid-dorsally at the end of the 9.5 low umbonal
ridge, interpreted to be the attachment of a byssal retractor muscle
Diagnosis.-Small Atrina with a broad, shallow sinus in pos-
as in mytiloid and mytiliform terior
bivalves (Yonge,
margin; low radial 1976,
costae and commarginal fig.
ridges 1.4);
strongest
and 3) in the pit or the pair of on
pits
ventral on
half ofthe
shell. anterior end of the
umbonal cavity, interpreted to be the insertion comprising
Description.-Fragment of an anterior pedal
the posterior half of a left
retractor muscle. The more ventral of the pair of pits is possibly
valve preserved as a composite mold of the exterior with some
the insertion of a vestigial anterior
adheringadductor, commonly
shell material. Dorsal margin straight;present
ventral margin
in bakevelliids (Cox, 1969b). A similar
concave towardpairventralofside umbonal pits oc-
in plan view and subtending an apical
curs in Middle Triassic Gervillia joleaudi
angle of about 420 (well
with dorsalshown by margin
margin; posterior Nie- sharply
meyer, 2002, fig. 12) as well as rounded
in Upper Triassic
on ventral side, then passing Gervillancea
into a shallow broad sinus,
coxiella. Skwarko (1967, p. 56) interpreted the posterodorsally
then straightening, becoming muscle attach- inclined and inter-
ments in Gervillancea in the samesecting
way. hingeThe strongest,
at an acute mid-dorsal
angle of about 700. Very weakly inflated,
pedal/byssal muscles would have without
beenmedialcapable
flexure zone andof probably
drawingwithout significant
the pos-
shell downward against an infaunalterior byssal
gape. Ornament weak and restricted with
attachment, to ventral sed-
half, con-
sisting of fine radial
iment penetration aided by the broadly roundedstriae and ventral
rounded, regularly spaced com-
margin
marginal ridges
but with depth of penetration limited by that the parallel the rounded shape
increased of the ventral part
convexity
of the shell in its dorsal third. of the posterior margin. Posterior end of ligament trace preserved,
In the ammonoid bed of Unit indicating
E, South insertionCanyon,
in a groove. Anterior end of shell and muscle
Gervillaria
(Baryvellia) ponderosa is abundantscars not preserved.
near the Remnants
base of outer
of the calcitic shell
unit, layer show
where it occurs mainly as single a valves
polygonal pattern
stackedindicatingbeside
original presence
and of on
columnar
top pris-
matic parallel
of one another with their long axes structure; inner aragonitic
(Fig. layersThese
9.4). not preserved.
occur
Etymology.-From
in a terrigenous sandy matrix containing the Latin
a hash ofadjective sinuatus, with
randomly reference
ori-
to the shallow sinus in the posterior margin.
ented shell fragments that do not appear to be size sorted. The
external surfaces of the Baryvellia Type.-The
shells species
inis the
represented only by the holotype,
ammonoid bed an asex-
ternal mold of an incomplete left valve, USNM 526455, near top
well as at other localities display abundant in-life repair of
of coral beds of Unit E, South Canyon, Locality 1.
chipped shell margins on both the shell body and auricles. The
Measurements.-Holotype, height 22 mm, restored length 35
presence of epifaunal bivalves (unidentified
mm.
pteriomorphs) on the
inner surfaces of both valves of articulated specimens suggest that
Occurrence.-Known only from the Unit E coral beds
these epifaunal taxa preferred the protected environment of gap-
Canyon, New Pass Range, Nevada, late Ladinian.
ing shells for settlement, and their size indicates slow sedimen-
Discussion.-There is some confusion regarding the
tation and long exposure of the Baryvellia shells on the sea bot-
tom.
stratigraphic ranges of the pinnid genera Atrina and Pi
Linnaeus, 1758. Cox and Hertlein (1969, p. N283) l
Collectively, these observations suggest that Baryvellia n. stratigraphic
sub- range of Atrina as Middle Jurassic to R
gen. lived in turbulent water with frequent dislodgement from a
that of Pinna (Pinna) as Lower Carboniferous to Recen
shallow semi-infaunal living position. Sedimentation was slow contradiction, Turner and Rosewater (1958, p. 297) said t
enough to permit epifaunal settlement and growth in the protectedna is known from the Carboniferous but that the earliest record
interiors of dead, gaping shells. Both living and dead shells were
of Pinna s.s. is from the Jurassic. As Turner and Rosewater noted,
occasionally transported by storms and buried in deeper, less thetur-younger occurrence of the oldest Pinna is in accord with its
bulent environments together with the shells of ammonoids. more derived morphology, particularly the presence of a medial
shell carina that is associated with the deep division of its internal
Suborder PINNOIDINA Waller, 1978 nacreous layer, producing a zone of shell flexure. Hyatt (1892)
Superfamily PINNOIDEA Leach, 1819 speculated that the early ontogenetic stages of the two genera
Family PINNIDAE Leach, 1819 would show little difference in morphology with respect to the
Genus ATRINA Gray, 1842 carina and division of the nacreous layer, and Turner and Rose-
water (1958) found this to be true. The contradiction seems to
Type species.-Pinna nigra Dillwyn, 1817, p. 325, by subse- stem from the uncritical use of the genus name Pinna by pale-
quent monotypy (=Pinna vexillum Born, 1778), Recent, Indo- ontologists, who have applied this name to fossils even though
Pacific.
they have no demonstrable carina. No such carina is visible on
Diagnosis.-Ham-shaped, rapidly distally expanding pinnids the illustration of "Pinna (Pinna) costata Philipps" that Cox and
that lack division of the inner nacreous layer into two lobes andHertlein (1969, fig. C23.2c) gave as a Carboniferous example of
also lack an exterior flexure zone. Pinna, suggesting that it is not of that genus.
Occurrence.--Triassic? to Recent; at present cosmopolitan in Atrina is probably derived from Carboniferous to Permian pin-
warm to temperate seas. nids that also lacked a flexure zone, such as Aviculopinna Meek,
5, Atrina sinuata n. sp., holotype, USNM 526455, external mold of posterior fragment of left valve, coral beds of Unit E, Locality 1, scale bar 1
cm. 6, 7, Liostrea? sp., holotype, USNM 526456, exterior of left valve and detail of antimarginal striae at site shown by arrow, coral beds of Unit
E, Locality 1, scale bars 3 mm and 1 mm. 8, Antiquilima ladinica n. sp., holotype, USNM 526457, exterior mold of left valve, coral beds of Unit
E, Locality 1, scale bar 2.5 mm. 9, Palaeolima newpassensis n. sp., holotype, USNM 526458, exterior of left valve, ammonoid bed of Unit E,
Locality 1, scale bar 5 mm. 10, 11, Plagiostoma acutum n. sp., holotype, UMIP 18048A, exterior of left valve and detail of dorsal region showing
posterior auricle, coral beds of Unit E, Locality 1, scale bars 10 mm and 3 mm.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
1864a or Meekopinna Yancey,

irregularities1978, both well
suggesting xenomorphic growth, repres
and 3) its sh
Permian of western North America
correctly (Yancey,
interpreted as retrocrescentic 1978).
(i.e., with the beaks
na, these Paleozoic genera
ing are narrower
toward the in with
posterior), is consistent form, its being com
an ost
apical angles of only 150,
left and
valve. It have
is placed insubterminal
Liostrea and not in Gryphaea beak La
the described specimen does not
1801 because have
of its preserved
shallow curvature bea
in a dorsal-ventra
ham-shaped outline suggests
and lackthat
of a strongitposterior
belongs in Atrina
groove, although clearly a
of the Paleozoic genera.groove
Modern
and posterior species
marginal sulcus of Atrina
are present. If an atta
faunal in soft, generallyscarsandy
was present sediments,
near the beak, it is notin warm
preserved, beca
Triassic species probablybeak
had a similar
and umbonal living
region are missing. Queries habit,
for both the
pockets of sediment between
and family colonial corals.
placements are necessary, because interior fe
including the hinge structure and the position and shape
Suborder OSTREOIDINA Ferrusac, 1822
adductor muscle scar, are not exposed. Furthermore, the fi
Discussion.--The placement of occurring
timarginal striae true within oysters
the ostracumas a ante
on the su
the Pterioida is based onsalstrong evidence
side of the holotype have not tofromour knowledge rece prev
been observed
genetic studies (18S rDNA) indictingin any Liostreathator Gryphaea.
oysters The antima ar
rived from pterioids rather
orientationthan from
of these striae, as well pectinoids
as their apparent great
Hammer, 2000; D. C. Campbell,
tance from the shell2000). The and
margin, shortening, largely
apparent disa
citic shell microstructure
ance of true
in a ventral oysters,
direction, suggest that once
they may be take
choma
indication of a pectinoid relationship
chomata (Waller,
are unknown in the Gryphaeinae, 1978)
including both L
dependently of that of pectinoids,
and Gryphaea (Stenzel,as 1971, indicated
p. N1097, N1103). by C
Liostrea? sp. resembles species of Liostrea from the L
Superfamily OSTREOIDEA Rafinesque,
Europe, including 1815
the type species Liostrea hisingeri (N
Family OSTREIDAE Rafinesque,
1832), in overall shape 1815 (s. Malchus,
(see Stenzel, 1971, fig. J61.2a,
Subfamily LIOSTREINAE Vialov, 1983
differs in being much smaller and in having a more pron
Genus LIOSTREA Douvill6, 1904
radial posterior depression and posterior marginal sulcus. Li
Type species.--Ostrea sublamellosa
newelli Newton, 1987, a Dunker,
small oyster from1846, the early Norb
designation, Jurassic (Lias), Halberstadt,
the Wallowa terrane of northeasternGermany.
Oregon, differs greatly
our specimen in shape. The Wallowa specimens have a
LIOSTREA? sp. lower height-length ratio, have an apparently more deeply
Figure 9.6, 9.7 and more extensively cemented left valve, and lack rounded
Description.--Left valve strongly
marginal retrocrescentic
undulations. They are possibly early members w o
mum curvature in anterior-posterior genus Catinula Rollier, 1911. rather than d
plane. Distance from preserved Liostrea (Catinula)
dorsalmostshiraiwensis Tokuyama,
point 1960, to fromve
10.0 mm, maximum transverse Carnian of Japan, distance
was reassigned6.0 mm,
by Hayami (1975,estp. 8
imum convexity 1.5 mm measured
Liostrea s.s. It differs fromat theabout midheig
New Pass specimen in being
margin broadly and evenly circular in shape and lacking
rounded, the anterodorsal
ventral margin striae. Ostr m
rounded, posterior margin nearly
tetiana Mortillet in straight posterovent
Stoppani, 1863, originally described fro
ing concave within broad Rhaetianposterodorsal
of Lombardy and later identified sulcus, by Wohrmann the
strongly convex along margin p. 201) in the of Carnianposterodorsal
(Raibl) of Germany, is possibly lobe; a L
gin and beak not exposed. Exterior
(see following discussion). worn
Like Liostrea? butsp. it posse
is a high
broad, unevenly spaced, form, irregularly commarginal
but it differs in having a more broadly rounded riv
strongest in ventral third ofa scarcely
margin, valve particularly
distinguishable on
posterior sulcus, andpoa siz
side, and some irregularis rugosities, about three times that suggesting
of the Nevada specimen. a cem
and xenomorphic growth; Ifradial the New Pass ornament
specimen is indeed absent
a Liostrea, itson Ladie
fine, antimarginal striaeage visible
makes it the within
oldest membershell fabric
of this genus ever to bea
dorsal margin, about 10/mm, extending
and it contributes significantlyinward from
to ongoing debates about
1.3 mm, shortening ventrally and and
cestors of Gryphaea becoming
the early evolution inset
of the Ostrefr
and no longer visible at Paleontologists 5.5 mm beginning
from withexposed
Douvill6 (1910, p. 118; 19
dors
of valve. Interior not accessible; hinge
635) have long regarded structure
Liostrea as ancestral to and
Gryphaea m
unknown. Shell microstructure, zel (1971, p. 1068) examined
questioned this idea,onlystating by
that "it ref
wa
very fine with apparent more antimarginally
than an assumption without oriented
good proofs" and lath
sugge
compact lathic or foliated structure;
N 1103) cross-foliated
that Liostrea "is perhaps a descendant of Gryphaeas
sent; outermost prismatic layer
than not
its ancestor, detected
as commonly and
assumed." prob
Stenzel's skep
served.
was based mainly on the relative stratigraphic positions
Material examined.-USNM 526456,
earliest occurrences of each ofa nearly
these com
genera. His review (S
valve, co-occurring on the 1971, p. N1069)rock
same of occurrences
specimenof Gryphaea in the Triass
with L
corallina n. sp. and Pleuronectites gests that the earliest recordnewelliis Gryphaean. sp., U
arcuataeformis
beds, Locality 1. The matrix sova, 1936, is a bioclastic
in rocks of late Carnian to limestone
Norian age in far e
dant echinoderm debris. Siberia. Stenzel (1971, p. N1055) concluded that the oldest
Measurements.-USNMtrea 526456,
were of Norian in height
age and therefore8.9 mm,
inferred that L
mm.
may well have originated after Gryphaea. Because he cou
Occurrence.--Known only from the Unit E coral confirm beds, Southearlier occurrences of Liostrea in the Tethyan re
Canyon, New Pass Range, late Ladinian. Europe, Stenzel further assumed that Liostrea, like Gry
Discussion.---The specimen is regarded as an oyster because 1)
originated in the Boreal region and that it did not become
it appears to be a calcitic shell with a fine microstructure
spread insug-lower latitudes until the Rhaetian.
gesting foliated calcite, 2) its exterior has indentations and other
Recently McRoberts (1992), in a review of the distribut

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Late Triassic Gryphaea in the North shell; ornament consisting

American of three orders of radial
Cordillera, chal- costae, with
lenged the Boreal-origin concept by about 12 unevenly spaced
showing that first-order
Late costae on disk.
Triassic
gryphaeids occur in lower paleolatitudes Description.-Known
than previously only from a single thought.left valve. Small,
Indeed they were widely distributed moderately along convex, or close slightly toopisthocline, with umbo extending
the western
cratonal margin of North America from above dorsalAlaska
margins and to asmargins
dorsal far sloping
south slightlyas ventrally
Nevada as well as in western Southaway America.
from beak; length McRoberts
of hinge line relatively (1992,
long (3.8 mm),
p. 31) also determined that the oldest beak posterior to centerof
specimens of hinge situated at 41%then
Gryphaea of total hinge
known are Gryphaea arcuataeformis (which
length from posteriorheendregarded
of hinge; anterior as auricleaprominent
se- and
nior synonym of G. chakii McLearn, well demarcated
1937) from from disk, itstheanterior margin slightly
lower Car- convex
nian of the eastern Brooks Range of and sinuous
Alaska and extending
in a ventrally
suspect to half terrane
the height of shell;
that may originally have been at a posterior lower latitude.
auricle indistinguishableThe new
from disk, data
the posterior margin
indicated to McRoberts (1992, p. 37) of diskthat "anmargin
and posterior Arctic of auricle beginning
forming a single uniform
for Gryphaea cannot be ruled out, but curve with only slight
neither can recurvature
origin closeat
to posterior
a lower dorsal mar-
paleolatitude in eastern Panthalassa." gin, meeting dorsal margin at a high angle. Ornament consisting
The idea of Boreal origins for both of three orders of radialand
Gryphaea costae on disk, the first-order
Liostrea has about 12
to some extent been buttressed by discrediting in number and unevenly orspaced on disk, continuing
ignoring reports onto posterior
of occurrences of the genus in the auricular Tethyan region, originating
region at shell
inheight
the of about
Late 3 mm, the
Triassic. Stenzel (1971, p. N105d3), second-order for example, unevenly intercalatedaccepted
in late ontogeny, the and the third-
several species from Sicily described orderby Scalia
consisting of very (1912) as belong-
fine striae covering interspaces; first- and
ing to Gryphaea, but doubted that they second-order radials absent
actually from anterior
came from auricle,Upper
third-order stri-
Triassic rocks as originally reported ae not detectable. Traces
because of ofstructural
muscle scars and ligamentcom- structures
plexities in the area. McRoberts (1992, p. 29)
not preserved. discounted
Shell apparently earlier
bimineralic, with very thin outer
reports of low-latitude Gryphaea in calcitic
the layerUpper
having a very fine antimarginal
Triassic asfabric;
being inner layer
questionable in terms of taxonomy not or age.orHallam
preserved represented by(1982,
recrystallizedp. 363),
material, probably
however, acknowledged the occurrence originally of Late Triassic Gryphaea
aragonitic.
in the Tethyan realm but emphasized its Etymology.-From
rarity,the Ladinian Stage,
citing oneto of emphasize
Scal- that this
ia's (1912) species from Sicily and a species similar
is the firstundescribed
definite Antiquilima to be species
found in the Middle
in Sarawak as the only examples. No Triassic.
authors, to our knowledge,
have reexamined the "Ostrea Pictetiana Type.-Holotype,
(Mort.)" USNMdescribed
526457 (Fig. 9.8),by a nearly complete
W6hrmann (1889, p. 201) from the Carnian external mold of of a the
left valveAlpinewith some adhering recrystallized
region,
although W6hrmann's figures (pl. 6, shell fig. material,
6, 6a) coral
showbeds ofan Unitunribbed,
E, Locality 1.
high-oval, retrocrescent form that could Measurements.-Holotype,
well be a Liostrea. height 6.8 Sten-mm, length 5.5 mm,
zel (1971, p. N1054), in fact, assigned to this
hinge length 3.8 mm, genus
height of a species
anterior auricle, 3.4 mm.
from Siberia identified by Kiparisova Occurrence.-Known
(1938a, pl. only 7, fig. from the 16a, coral b) beds of Unit E,
as "Ostrea aff. pictetiana." South Canyon, New Pass Range, Nevada, late Ladinian.
The New Pass specimen is older than Discussion.-The
any of new species isreported
these assigned to the genus Anti-
occurrences of either Liostrea or Gryphaea. quilima, because it possesses characters
Furthermore, the that as-Cox (1943, p. 180)
sociation of the New Pass specimen andwithCox and warm,Hertlein (1969, p. N386) regarded as characteristic
shallow-water
indicators, particularly colonial corals, of the genus, suggests
viz. prominent that the
anterior cool,
auricles that are dorsoven-
deepwater habitats regarded as typical trally for elongate
later and Liostrea
enclose a correspondingly
and Gry- elongate byssal
phaea may be the result of later (Late gape, and ornament consisting
Triassic) range ofextension two or more orders of radial
and an evolutionary shift in temperature costae that are tolerance.
commonly sinuous Another
where interrupted in-by irregularly
triguing possibility is that the Middle spacedTriassic
commarginal Liostrea?
growth stops.from An additional thefeature of An-
New Pass Range may well represent the common
tiquilima, first observed by ancestor
Hayami and Maeda (stem (in Hayami et al.,
group) for the Gryphaeinae, which evolved 1977, p. 211-212),
greater concernsleft-valve
the inequality of coil- the anterior auricles
ing and a reduced attachment area while of the tworemaining
valves. The anteriorstenohaline,
margin of the right anterior auricle
and the Ostreinae, which became adapted is deeply concave to and euryhaline
intersects the dorsal condi- margin at an acute
tions. In fact, Hudson and Palmer angle; (1976) havemargin
the anterior already of the leftsupplied
anterior auricle is slightly
compelling arguments for the derivation convex (or sigmoidal)
of the andeuryhaline
intersects the dorsal Os- margin at a high
treinae from within the Liostreinae by angle. The resultJurassic
middle of this discordance,(Bathonian)as noted by Hayami and
time. Maeda, is an asymmetric byssal gape that allowed byssal protru-
sion to be directed to the right side. The new species of Anti-
Order LIMOIDA Waller, 1978 quilima described herein, although based only on a single left
Superfamily LIMOIDEA Rafinesque, valve, possesses 1815 a left anterior auricle with an anterior margin of
Family LIMIDAE Rafinesque, a shape
1815 known to occur in other members of the genus.
Genus ANTIQUILIMA Cox, 1943 So far as known, only four species of Antiquilima have been
identified or described from the Triassic: Antiquilima praelonga
Type species.--Lima antiquata J. Sowerby,
(Martin, 1860) from the Upper 1818, Triassicp. 25
(Norian-Rhaetian) of east-
(=Chamites succinctus Schlotheim, 1813), by original designa-
ern Siberia and the northern Prookhotsk region (Kiparisova et al.,
tion (Cox, 1943, p. 179), Jurassic (Lias), Great Britain.
1966; Milova, 1976); A. atacamensis Hayami and Maeda in Hay-
ami et al., 1977 from the Upper Triassic (probably Norian) of
ANTIQUILIMA LADINICA new species
northern Chile; A. vallieri Newton, 1987, from the early Norian
Figure 9.8
of the Wallowa terrane in eastern Oregon; and A. hians Hautmann,
Diagnosis.-Antiquilima of very small size, slightly opisthoc- 2001b, from the Norian-Rhaetian of Iran. In addition, Lima (Rad-
line, with posterior auricle not set off from disk and anterior au- ula) baliana Bittner, 1891, from the Norian (Diener, 1923, p. 108)
ricle well demarcated, extending ventrally to half the height of of Anatolia, Turkey, appears to belong in Antiquilima. Although

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC34:56 UTC
study of limids from the than the posterodorsal

European margin (560), ventralTriassic
margin gently curved, may
species of Antiquilima,then atcurvingthis
more sharply point
around anteroventraltheand posteroven-
new spe
is the oldest known. tral extremities; sides of umbo gently
Compared tosloped, Cox's
lacking sharp um- (1943
that the genus is exclusively Lower
bonal ridges. Hinge short, and
estimated to be less Middle
than half length of J
graphic range now extends from
shell; beak about central. Shape of anteriorupper
auricle unknown; pos-Ladini
taceous (Aptian, based terior on auricleAntiquilima
poorly set off from disk, the ventral partultima
of its pos- Ha
west Japan). terior margin indicating that this margin is slightly concave, its
Antiquilima ladinica differs from all known species in its very intersection with dorsal margin not exposed. Ornament of disk
small size, very unevenly spaced first-order costae, uneven sub- consisting of radial costae uneven in spacing and size, tending to
medial intercalation of second-order costellae, and a posterior aube in three orders, about 20-25 major costae separated by from
ricle that is weakly demarcated, if at all, from the disk. one to three finer costae; prominent commarginal growth lamellae
present in interspaces, forming small scales or nodes where in-
Genus PALAEOLIMA Hind, 1903
tersecting radial costae. Interior characters not exposed. Shell mi-
Type species.-Pecten simplex Phillips, 1836, p. 212, by sub- crostructure not preserved.
sequent designation (Cox, 1952, p. 48), Carboniferous, England. Etymology.-Named for the New Pass Range, Nevada.
Discussion.--Newell (1999, p. 4) rejected the genus Palaeoli- Type.-Known only from the holotype, USNM 526458, from
ma "as being unrecognizable" on the grounds that the original the ammonoid bed of Unit E, Locality 1, a left valve with disk
specimens of its type species, Pecten simplex Phillips, 1836, have nearly complete but auricles and exterior surface in poor condi-
disappeared and that the characters of the genus therefore cannot tion.
be assessed. Dickins (1963, p. 91), however, had earlier addressed Measurements.-Height 15.9 mm, length 18.3 mm.
this problem and designated a neotype of Phillips's species, spe- Occurrence.-Known only from the ammonoid bed above the
cifically the specimen figured by Hind (1903, pl. 19, fig. 26) from coral beds of Unit E, South Canyon, New Pass Range, Nevada.
Little Island, County Cork, Ireland. This specimen, with Hind's Discussion.-The uneven but ordered ribbing in combination
other specimens of the species, is in the collections of The Natural with the elongate shape and sharply rounded posterior margin set
History Museum, London, and bears the catalogue number the new species apart from all other species of Palaeolima that
L24706. Shape and ornament vary among the four specimens of were examined in the literature. Among the many limids de-
Palaeolima simplex illustrated by Hind (1903, pl. 19, figs. 24- scribed by Bittner (1895) from the San Cassiano Formation of
27). With the exception of the neotype, all are longer than higher Alpine Italy, some, such as Palaeolima cancellata (Bittner, 1895),
and have substantial anteroventral extension of the disk. All are P. austriaca (Bittner, 1895), and P. angulata (Mtinster, 1841)
radially costate, with costae varying from simple and evenly dis- have costae of at least two orders but their detailed ribbing pat-
tributed to variously branched and unevenly spaced. All indicateterns are different from that of the new species, and they lack
the lack of infolding of the anterodorsal margins and a lack ofsharply rounded anterior and posterior disk margins.
strong umbonal folds. Both auricles are visible in lateral views.
One of two other species described by Hind, Palaeolima laevis, Genus PLAGIOSTOMA J. Sowerby, 1814
is smooth and rounded; the other, P. obliquiradiata, has bifurcat- Type species.-Plagiostoma giganteum Sowerby, 1814, by sub-
ing and intercalated costae and a highly oblique oval shape ex-sequent designation of Stoliczka (1871, p. xxii), Jurassic (Lias),
tending anteroventrally. England.
In general, many of the species that have been assigned to
Palaeolima have the anteroventrally extended shape and lack of PLAGIOSTOMA ACUTUM new species
infolding in the byssal region. As far as can be determined, the Figure 9.10, 9.11
stratigraphic range of the genus is from the uppermost Devonian Lima cf. striata (SCHLOTHEIM, 1820). STANLEY, 1979, p. 18.
of Germany (Amler et al., 1990, p. 51, Palaeolima? sp.) to Upper
Triassic (Norian) (e.g., Lu, 1981). The genus apparently achieved Diagnosis.-Small to medium-sized, strongly opisthocline Pla-
its greatest geographic distribution before the end of the Paleo-giostoma with posterior margin of disk extending only a short
zoic, when it occurred in Tethyan regions as well as in the Boreal distance posterior to extremity of posterior auricle, disk with
region [Permian of Spitsbergen (Nakazawa, 1999, p. 15)], North about 47 low costae, and posterior auricle prominent, its concave
America [e.g., Palaeolima retifera (Shumard in Shumard and posterior margin meeting hinge line at an acute angle.
Swallow, 1858) from the Upper Carboniferous of Kansas and Description.-Plagiostoma of small to medium size, height and
Ohio (Morningstar, 1922, p. 233)], South America [latest Car- length about equal, highly opisthocline with straight anterodorsal
boniferous to earliest Permian of Argentina (Sterren, 2000, p.margin subtending angle with hinge of about 135', umbonal angle
435)], and Australia [Permian of Western Australia (Dickins, about 900, and only moderately inflated with beak extending only
1963, p. 92)]. Palaeolima survived the great extinction at the endslightly dorsal to hinge line; ventral margin highly asymmetric,
of the Permian, but its geographic range was restricted to the with anteroventral extremity sharply curved. Posteroventral ex-
Tethyan region of Eurasia. It is apparently absent from Triassic tremity shallowly curved, and posterior of disk extending only
faunas of northeastern Asia and the Boreal region, and until now slightly beyond extremity of posterior auricle; anterior umbonal
has not been reported from the Triassic of the Americas. ridge absent; lunule and anterior auricle obscured by matrix; pos-
terior auricle large, poorly demarcated from disk by gradual
PALAEOLIMA NEWPASSENSIS new species change in curvature, dorsal margin of auricle straight and not
Figure 9.9 declining away from beak in lateral view, posterior margin of
Diagnosis.-Palaeolima with length greater than height; disk auricle shallowly sigmoidal with overall trend slightly acute to
costae irregular in spacing and size, with 20-25 major costae dorsal margin.
separated by from one to three minor costae. Ornament of disk consisting of about 47 simple, unbranched
Description.--Small, longer than high (height 15.9 mm, length radial costae that are narrow, rounded, and only slightly wider
18.3 mm), moderately convex, distinctly opisthocline, anteroven- than interspaces in anterior and posterior quarter sectors of disk;
trally extended, with nearly straight anterodorsal margin of disk costae in central two quarter sectors of disk broader, flattened, and
longer and descending at a lower angle from the hinge line (320) much wider than interspaces; intracostal spaces occupied by

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
modern
closely spaced commarginal lirae, pterioid genus
giving a Malleus Lamarck, 1799.
punctate The hypertro-
appearance
in the central part of disk; costae phied point of its posteriorwithout
continuing auricle is also a pterioidan feature not
interruption
onto posterior ear. Ligament area known
andto interior
be present in any features
demonstrable limoidan. McLearn's
concealed.
species should
Shell thin with only the thin calcitic probably retain
outer his initial generic
ostracum assignment to
remaining,
its microstructure fine and with Aviculomyalina
an antimarginal Assmann, 1916, a genus
fabric.that we would place in
Etymology.--Name formed from the Pteriidae
the or Malleidae
Latin (s. Waller, 1978) rather than
adjective in the
acutus,
meaning sharp, pointed, with reference Myalinidae. to the sharply rounded
anteroventral margin of the disk. Species of Plagiostoma from the Triassic of Europe and North
Type.--Holotype, UMIP 18048A, America that havebeds
coral straight anterodorsal
of Unit marginsE,and strong
Local-op-
ity 1, a left valve. isthoclinity differ from the new species in details of ornament and
Measurements.-Height 31.3 mm, shape.length
For example, Lima
31balatonica
mm, Bittner, 1901, of Arthaber 5
convexity
mm. (1915, p. 192), from the Middle Triassic of Anatolia (possibly
Occurrence.-Known only from the Unit E coral frombeds,
beds equivalent
South to the late Ladinian Sutherlandi Zone of
Canyon, New Pass Range, Nevada, late Ladinian.Canada; see Tozer, 1967, p. 30), has higher, more rounded radial
Discussion.-The New Pass specimen is unique amongcostae and known
a smaller posterior auricle, and the posterior auricular
Triassic and Jurassic Plagiostoma in combiningmargin 1) anforms an obtuse angle with the hinge. "Lima (Plagios-
oblique,
toma) sp. near
highly opisthocline shape with height equal to or slightly rigidula Phil.," described by Lees (1934, p. 39)
greater
from the Lewis
than length, 2) no significant projection of the posterior River Formation of the Laberge area, Yukon, is
margin
highly oblique
of the disk beyond the posterior limit of the hinge, and 3) a prom- and has a prominent posterior ear with a concave
posterior
inent posterior auricle having a sigmoidal posterior margin, but
margin, thethe overall trend of this auricular margin is
overall trend of which forms a slightly acute angleat with
an obtuse
theangle to the dorsal margin. Further, the Yukon species
dorsal
margin. is about twice the size of that from New Pass, and its posterior
The new species is based on the same specimen listed by Stan- shell region projects further beyond the posterior extremity of the
ley (1979, p. 18) as "Lima cf. striata." Plagiostoma striatum posterior auricle.
(Schlotheim, 1820), and its probable conspecific variant, Plagios- Some species of Plagiostoma from the Jurassic of Europe and
toma lineatum (Schlotheim, 1820), are forms that are particularly Great Britain (Dechaseaux, 1936; Cox, 1943) also resemble the
common in the Lower Muschelkalk (Anisian) of Europe. They New Pass species in shape [e.g., Plagiostoma annonii Merian in
differ from the New Pass specimen in being more inflated, cir- Greppin (1899) of Dechaseaux (1936, pl. 1, fig. 11) from the
cular, and equilateral and in having a smaller posterior auricle, Bajocian of France and P. schimperi (Branco, 1879) of Cox
the posterior margin of which intersects the hinge in an obtuse (1943, pl. 11) from the Aalenian of Great Britain]. In general,
rather than acute angle. In contrast to the distinctly concave pos- however, the Jurassic species have more circular ventral margins,
terior margin of the posterior auricle of the new species, that of posterior disk margins that project well beyond the end of the
the Muschelkalk species is commonly nearly straight to only very posterior auricle, and obtuse rather than acute posterior auricles.
slightly concave or convex. Bittner (1895, p. 177) and Cox (1943, p. 153) reasoned that
The specimens figured by Newton (1987, fig. 22.12-22.15) as Plagiostoma is probably derived from Early Triassic Mysidiop-
tera. As they observed, an important clue is the morphology of
"?Plagiostoma sp." from the early Norian Martin Bridge For-
Plagiostoma lineatum of the Lower Muschelkalk, which retains
mation of the Wallow terrane of Oregon have a small posterior
some of the equilateral, circular disk shape and asymmetric, pos-
auricle with a concave posterior margin and a dorsal point. Their
teriorly directed ligament area of Mysidioptera while displaying
overall form, however, is much more equilateral than that of P.
some of the features more typical of Plagiostoma, particularly the
acutum. Furthermore, the hinge view in Newton's figure 22.13
strongly excavated lunule and flattened costation with "punctate"
indicates that the ligament area is highly inequilateral, with its
interspaces. As indicated in the above comparisons, Plagiostoma
principal development posterior to the beak and its resilifer having
from the Upper Triassic and Jurassic are commonly strongly op-
a strong migration toward the posterior throughout ontogeny.
isthocline, but they still retain obtuse posterior auricles and, like
These features suggest to us that these specimens may belong in
Mysidioptera and Plagiostoma lineatum, the posterior margin of
Mysidioptera Salomon, 1895, rather than in Plagiostoma.
the disk commonly extends well beyond the end of the posterior
A second so-called limid in Newton (1987, p. 29) from the auricle. Against this background, Plagiostoma acutum appears to
Martin Bridge Formation is "Mysidioptera" williamsi (McLearn, represent an endemic lineage that evolved in the Anisian or Lad-
1941 a), originally described from the "Lima" poyana beds of the inian and remained outside the mainstream of Late Triassic and
Peace River foothills of northeastern British Columbia (late Car- Jurassic evolution of the genus.
nian according to Tozer, 1994, p. 16). It has a large posterior
auricle that has a concave posterior margin meeting the hinge at Order PECTINOIDA H. Adams and A. Adams, 1858
an acute angle. In early ontogeny, the angle is even more acute, Suborder PECTINOIDINA H. Adams and A. Adams, 1858
producing an elongate point along the dorsal margin. The main Superfamily PSEUDOMONOTOIDEA Newell, 1938
part of the shell is coarsely commarginally lamellose and lacks Family LEPTOCHONDRIIDAE Newell and Boyd, 1995
radial costae. The disparity in shape, size, and ornament between Genus LEPTOCHONDRIA Bittner, 1891
this and the other Martin Bridge species that are assigned to
Type species.--Pecten (Leptochondria) aeolicus Bittner, 1891,
Mysidioptera Salomon, 1895 results from the fact that "Mysi- p. 101 by monotypy, Triassic (Norian), Anatolia, Turkey.
dioptera" williamsi is not a limoid at all, but a pterioid. This is
evidenced by the presence of coarse columnar prismatic micro- LEPTOCHONDRIA SHOSHONENSIS new species
structure in the outermost shell layer of both valves in the holo- Figure 10.1-10.5
type and paratypes as well as by the alivincular ligament system Pecten (Velopecten) albertii (GOLDFUSS, 1835). MULLER AND FERGUSON,
illustrated by Newton (1987, fig. 25.4). Rather than a resilifer that 1939, p. 1593; SILBERLING, 1959, p. 13.
continuously migrates toward the posterior as in Mysidioptera,
that of "Mysidioptera" williamsi turns from posterior migration Diagnosis.--Leptochondria of moderate size and circular to
in early ontogeny to directly ventral migration later, as in the high-oval form; left valve moderately inflated, with densely

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
packed, unordered, wavy There costellae ondensity

are also differences in costal both disk
and morphology. In an
ricles obtuse and poorly setspecies
the Nevada off from
the costae disk,
are more dense, the
with narrower in- an
with a convex anterior margin. terspaces and less of a tendency for medial intercalation. The only
Description.-Represented only
other Eurasian finely andby densely left valves.
costate species known to me M
size (up to 25 mm), orthocline that is closely comparable
to slightlyis Leptochondria minima (Kiparisova,
opisthoclin
with height generally equal 1938b) fromto the Lower
orTriassic of Russia (see also
greater thanGavrilova, leng
to moderate convexity; 1995, beak p. 88). Thebarely
new species differsprojecting
from it in being narrower abov
gin; hinge line short (about and higher athalf a comparable of growth stage and in
shell having less
length);
and weakly set off fromoblique disk auricles.if at all, anterior marg
auricle slightly convex with Previous reports
overallof Leptochondriatrend
in North America
forming have been a
with dorsal margin and from withthe Permian and Lower Triassic.
only a very Leptochondria curtocar-
shallow b
change in curvature into dinalis
the (Hall and Whitfield, 1877), illustrated
anterior disk by Newell and Boyd
margin, p
gin of posterior auricle more (1995, fig. 51.1,obtuse
51.2) from the Permian
except of Pakistanforand Lower very
vature just below dorsalTriassic margin.of Idaho (Thaynes Formation, Nammalian, Meekoceras
Ornament of dense, fine Zone),antimarginal
has a shape very similar to that of costellae
European L. inaequis-on d
ricles, originating at less triata,
than but the costellae
1 mm of L. curtocardinalis
from are beak
finer and farand
both branching and intercalation weaker on the auricles than in either L. inaequistriata or the
throughout new
ontoge
costellae variable, ranging Nevadafromspecies. Leptochondria
fiveoccidaneus to nine (Meek, 1877), which mil
per
sured in central sector at occurs in the Lowerheight
valve Triassic (Nammalian)
of Moenkopi
10 Formation
mm; co
ing direction or becoming wavy
(Virgin Limestone) adjacent
of southern to growt
Nevada and the Thaynes Forma-
paired injuries; weak commarginal tion of northern Nevada and Utah lirae(Newell and Boyd, 1995, p. in i
visible
Shell thin and presumed to be
71), is markedly originally
different biminer
in having coarse, more widely spaced
of differential dissolution costae with
ofsecond-order
inner costaeshell
medially intercalated
layer; (Newell and
outer
very thin and subject toBoyd, diagenetic
1995, fig. 51.3-51.9). distortion, its m
not resolved. The abundance of Leptochondria shoshonensis in the Grants-
Etymology.-Named after the Shoshone Mountains, west-cen- ville Formation of the Shoshone Mountains compared to its
tral Nevada. paucity in the New Pass Range may have a paleoecological ex-
Types.-Holotype: USNM 526459, a left valve (Fig. 10.1, planation. Silberling (1959, p. 13) observed that the Grantsville
10.2), from Locality 5, Grantsville Formation, Shoshone Mts., Formation, consisting of a lower clastic member and an upper
Nevada. Paratypes from Locality 5: USNM 526460, left valve limestone member, represents a transgression. The clastic member,
(Fig. 10.3); USNM 526461, left valve (Fig. 10.4); and USNM with abundant conglomerate, grades into the basal impure silty
526462, 34 left valves. Paratype from Unit D, South Canyon, limestone beds of the limestone member, which in turn grade into
Locality 1: USNM 526463, inner surface of small partial left the purer limestones of the upper part of this member. A common
valve. fossil in the basal impure limestone beds is a bivalve that Silber-
Measurements.-Holotype, height 17.7 mm, length 16.1 mm, ling referred to as "an unidentified concavo-convex radially
hinge length 6.9 mm. ribbed oysterlike pelecypod." It is in fact a species (probably
Occurrence.-Known only from the limestone member of the new) of Newaagia Hertlein, 1952, that lived attached by its very
Grantsville Formation in the Shoshone Mountains and from Unit convex right valve to hard substrates, commonly to shell frag-
D in South Canyon, New Pass Range, Nevada, late Ladinian. ments of other bivalves. That it lived in a highly turbulent envi-
Discussion.-The preceding description is based on well-pre- ronment is indicated by the repeated damage sustained by its thick
served specimens, all left valves, from the Grantsville Formation shell throughout life. Many of the specimens were apparently dis-
in the Shoshone Mountains. Specimens (also left valves only) lodged from a hardground and buried, as articulated shells, in the
from Unit D of South Canyon, New Pass Range, are distorted basal limestone beds. In contrast, Silberling noted that Pecten
and/or fragmentary, but appear to represent the same species (Velopecten) albertii (=Leptochondria shoshonensis) is most
based on features of shape and costal density. The occurrence of abundant in the higher part of the limestone member, suggesting
Leptochondria only as left valves is a common phenomenon, that it lived in deeper, more offshore waters, an interpretation
probably because of the flatness and inherently weaker structure consistent with its thinner shell and less common repair of shell
of its right valve (Newell and Boyd, 1995, p. 69, 71). margins fractured during life. Silberling regarded the coral-algal
Muller and Ferguson (1939, p. 1593) and Silberling (1959, p. limestones at New Pass (Unit E herein), only 50 miles to the
13) referred to the Grantsville specimens as Pecten (Velopecten) north, to be correlative with the Grantsville Formation and to
albertii. This common Muschelkalk species, originally described represent a more offshore depositional site. He speculated that the
as Monotis albertii Goldfuss, 1835, p. 38, is now generally re- coral reefs possibly acted as a barrier that delimited a more re-
garded as a junior synonym of Pecten inaequistriatus Miinster instricted Muschelkalk-like sea that bordered the mainland. Support
Goldfuss, 1833, p. 42 (e.g., Diener, 1923, p. 65; Cox, 1924, p. for this onshore-offshore gradient is supported by the lack of any
68) and is well established in the genus Leptochondria (e.g., Nak-trace of the Newaagia at New Pass and by the rare occurrence of
azawa and Newell, 1968, p. 73; Allasinaz, 1972, p. 229). The Leptochondria.
species is very broadly distributed in the Lower and Middle Tri-
assic, occurring from Europe to more eastern and northern regions Superfamily MONOTOIDEA P. Fischer, 1887 (s. Begg and
including Jordan (Cox, 1924, p. 69; Lerman, 1960, p. 40), north- Campbell, 1985)
eastern Russia (Bychkov et al., 1976, p. 77, as Leptochondria aff. Family ASOELLIDAE Begg and Campbell, 1985
albertii), and Spitzbergen [Bi5hm, 1913, as "Pseudomonotis (Eu- Discussion.-Begg and Campbell (1985, p. 727) erected the
microtis?) arctica nov. sp."]. family Asoellidae to accommodate monotoideans having a rela-
Compared to typical Muschelkalk Leptochondria inaequistria- tively unreduced right anterior auricle, an amphidetic ligament
ta, the new species from Nevada attains larger size, has a narrow- system with a subcentral resilifer developed beneath the beaks, a
er form in early ontogeny, and has a less oblique left anterior lack of accessory hinging structures, and shell length commonly
auricle that has a convex rather than straight anterior margin. exceeding height. They included three genera in the family:

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
ASOELLA? species A1985; and

Asoella Tokuyama, 1959; Etalia Begg and Campbell,
(tentatively) Aucellina Pompeckj, 1901. Unlike Figure
the10.6 other genera,
the left valve of Asoella is generally only moderately inflated, so
Description.-Left valve small, thin-shelled and distorted,
that the left beak projects relatively less beyond the hinge. In
equilateral and nearly circular, with height and length equal, and
comparison to both Etalia and Aucellina, the right anterior auricle
moderately convex; beak narrowly inflated, barely extending
of Asoella has a much shallower byssal notch, particularly in late
above dorsal margin and centered on a hinge margin that is about
ontogeny.
half the length of valve. Auricles small and poorly set off from
The systematic position of the Asoellidae was discussed by
disk in mid- to late ontogeny; anterior margin of anterior auricle
Waterhouse (2001, p. 124), who doubted its position within the
convex toward anterior with overall trend forming slightly acute
Monotoidea. The specimens on hand, however, are too poorly
angle with dorsal margin in early ontogeny, becoming obtuse in
preserved to shed any new light on this matter.
later ontogeny, lacking a byssal sinus; posterior auricle even more
poorly set off from disk than the anterior one, its posterior margin
Genus ASOELLA Tokuyama, 1959 convex toward posterior and forming an obtuse angle with dorsal
Type species.-Eumorphotis (Asoella) confertoradiata Toku- margin. Exterior smooth except for low, irregularly spaced, com-
yama, 1959, by original designation, Carnian (or lower Norian), marginal growth rugae in late ontogeny, accentuated in anterior
Mine Group (Aso Formation), Mine area, west Japan (Hayami, and posterior auricular regions. Right valve unknown.
1975, p. 60). Material examined.-USNM 526464, a left valve, coral beds
Discussion.-Tokuyama (1959, p. 3) introduced Asoella as a of Unit E, South Canyon, Locality 1.
subgenus of Eumorphotis Bittner, 1901, noting strong similarity Measurements.--Height 14.9 mm, length 14.8 mm, hinge
to that genus in ornament and shape. Newell and Boyd (1995, p. length 8.0 mm.
37) remarked on the strong similarity and the possibly arbitrary Occurrence.-Known only from the Unit E coral beds, South
distinction between Eumorphotis of the Lower Triassic and some Canyon, New Pass Range, Nevada, late Ladinian.
members of the Upper Paleozoic aviculopectinoidean genus Het- Discussion.-Most species of Asoella have radial ornament, at
eropecten Kegel and da Costa, 1951. The principal morphological least in early ontogeny. A noncostellate exception is Asoella lae-
trend among these genera involves auricular shapes. In Hetero- vigata (Tokuyama, 1959), from the Carnian or lower Norian (Aso
pecten both auricles of the left valve tend to be set off from the Formation) of western Japan (Hayami, 1975, p. 61). As shown
disk by low disk flanks. Eumorphotis retains the clearly demar- particularly by plate 1, figure 8 in Tokuyama (1959), this species
cated and sinuate left anterior auricle of Heteropecten, but the differs from ours in having a broader, more inflated umbo that
posterior auricle of Eumorphotis, although remaining acute as in extends much further above the dorsal margin and a more pro-
Heteropecten, loses its demarcation from the disk with the dis- socline shape with the anteriormost point of the disk extending
appearance of the posterior disk flank. In Asoella, the left anterior only slightly anterior to the anterior end of the hinge. Although
auricle also becomes less clearly differentiated from the disk, and it is difficult to determine from Tokuyama's figures, it appears
the left anterior byssal sinus becomes shallower, disappearing in that the left anterior auricle loses its byssal sinus fairly early in
some species. Some species of Asoella show reduction in strength ontogeny so that the anterior margin of the disk sweeps dorsally
or loss of radial costae, becoming smooth throughout ontogeny. with even curvature to meet the hinge at an obtuse angle. In
Etalia, a smooth form that is thus far known only from the Asoella? species a, the byssal sinus persists on the left valve,
Anisian (Etalian) of New Zealand, possibly has an independent becoming obsolescent only very late in ontogeny. Compared to
origin in Eumorphotis. This is suggested by Etalia's retention of Asoella? species b, A.? species a is less high and more circular
a deep byssal notch, the shape of its radially costate early growth and has smaller, more poorly demarcated auricles with more ob-
stage, and its posteriorly extended resilifer. tuse auricular margins. Compared to Asoella? species c, A.? spe-
cies a lacks radial ornament.
In this study, three left valves, all from talus adjacent to the
Unit E coral beds of South Canyon, are tentatively assigned to
Asoella because many of their features conform to those included ASOELLA? species B
by Tokuyama (1959, p. 2) in his original diagnosis of the genus, Figure 10.7
particularly the small, "roundly quadrate" acline to slightly pro- Description.-Left valve small, thin-shelled, somewhat higher
socline shape, orthogyrous beaks, small, poorly demarcated au- than long, slightly opisthogyrous and opisthocline, and moderate-
ricles, and subdued ornament. The generic assignment remains ly convex; ventral margin of disk broadly rounded, anterior mar-
questionable, however, because right valves are unknown, and gin more sharply rounded merging into anterior margin of anterior
because the specimens appear to lack the prismatic and crossed- auricle without a byssal sinus; posterior margin broadly rounded,
foliated calcitic microstructures that are broadly present in the becoming nearly straight where merging into straight posterior
family. Because each of the three specimens differs from the oth- margin of posterior auricle; beak moderately broadly inflated,
ers in details of shape and ornament, they are tentatively described overhanging the dorsal margin, and centered on hinge margin that
as three species. Open nomenclature is used, however, because is somewhat greater than half the length of valve. Large anterior
they are the only specimens collected, and there remains a pos- auricle high but poorly set off from disk, anterior margin of au-
sibility that they all lie within the range of variation of a single ricle straight or very slightly concave, meeting dorsal margin at
species. slightly obtuse angle; posterior auricle also poorly set off from
If these specimens are correctly assigned to Asoella, they are disk but lower than anterior auricle, with a nearly straight pos-
the first representatives of the genus to be described from the terior margin that meets the dorsal margin at a right or slightly
Americas. They may also be the oldest members of the genus. obtuse angle. Exterior smooth except for low, irregularly spaced,
The only Ladinian species that Tokuyama (1959, p. 2) placed in commarginal growth rugae in late ontogeny, accentuated in an-
Asoella is "Pseudomonotis illyrica Bittner from the Ladinian and terior and posterior auricular regions. Right valve unknown.
Carnian of Asia Minor." Bittner (1902, p. 227), however, de- Material examined.-USNM 526465, a left valve, coral beds
scribed this species from Krain (in Slovenia), and Diener (1923, of Unit E, South Canyon, Locality 1.
p. 41) gave its stratigraphic occurrence as Ladinian only. It ap- Measurements.-Height 15.7 mm (restored), length 16.2 mm,
pears more likely that this species is a Leptochondria, not Asoella. hinge length 9.5 mm.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
f PIN.
;Y.;
wit"
tr;
ii'iIP~;s~S~',~"%l_~~P1 1~8o~"'fit If t-i~
f1_1 ZZi~
lip I IWOP~g~l
CM 411-iVF
:?,:;e~ -: ?kT ?-
%W*17
i~i~8ikw t
V Ak gz
14?
uaP~r'ib i~~ ~&~1
i ;Id
ITT3L
USA EIVIO 7-1.~~5
FIGURE 10-1-5, Leptochondria shoshonensis n. sp.: 1, 2, holotype, USNM 526459, exterior of left valve and detail
Locality 5; 3, paratype, USNM 526460, exterior of left valve, Grantsville Fm., Locality 5; 4, paratype, USNM 5
Grantsville Fm., Locality 5; 5, paratype, USNM 526463, composite exterior mold of left valve, Unit D, Locality 1. 6
from coral beds of Unit E, Locality 1: 6, Asoella? species a, USNM 526464, exterior of left valve; 7, Asoella? spe
of left valve; 8, Asoella? species c, USNM 526466, exterior of left valve. 9-12, Oxytoma (Oxytoma) grantsvillen
526467, external mold of left valve, Grantsville Fm., Locality 5; 10, paratype, USNM 526470, external mold of left

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Occurrence.--Known only from(e.g., the Unit density E coralabout beds,

28/mm South at a dist
Canyon, New Pass Range, Nevada, thelate dorsal Ladinian.
margin). The boundaries be
Discussion.-The auricles of thisauricle are difficult in
species to setlate
because of the lack of demarcation
ontogeny re- and
semble those of Asoella? species the
a steady
in early ontogeny.
curvature from the ventral margin Theof the final
disk onto this
growth forms differ substantially, however,
auricle. The boundary was setwithsomewhat A.? species
arbitrarily a
as the border
becoming more circular with much smaller
of the anteriormost and rib
first-order more
borderingobtuse au-
the anterior disk sec-
ricles.
tor. The boundary between disk and left posterior auricle is easier
to discern, because the posteriormost disk rib is commonly a first-
ASOELLA? species C order rib of high amplitude.
Figure 10.8
Description.--Left valve small,OXYTOMA thin-shelled, equilateral,
(OXYTOMA) GRANTSVILLENSIS new species
obliquely ovate, slightly prosocline, and somewhat Figure 10.9-10.12 higher than
long; beak narrowly inflated and slightly overhanging dorsal mar-
Diagnosis.--Very small, nearly equilateral, upright, narrow Ox-
gin; disk moderately inflated with greatest convexity at about
ytoma with posterior auricles meeting dorsal margin at a right or
midheight; hinge length about half of shell length, with beak ap-
slightly acute angle, not sharply pointed, and not extending pos-
proximately in center. Auricles poorly set off from disk; anterior
teriorly as far as disk margin.
auricle high and triangular in early ontogeny, with anterior margin
Description.-Left valve small (heights ranging from 4.0 to 9.5
shallowly sigmoidal with overall trend intersecting dorsal margin
mm), somewhat higher than long, prosocline, nearly equilateral,
at an acute angle, becoming more obtuse and convex in later
ontogeny; posterior auricle smallermoderately
andconvex,more with poorly
narrow umbonal angle (70o-80')its
defined, and
beak centrally positioned and only slightly projecting above dor-
posterior margin slightly concave and forming a more obtuse an-
sal margin. Left auricles about equal in size, both poorly set off
gle with dorsal margin. Ornament consisting of about 30 low
from disk; anterior
wavy radial costae on disk, with uneven auricle with outwardly
intercalation of convex
new anterior
cos- margin
intersecting dorsal margin
tellae in late ontogeny; costae narrower than interspaces, fading
at an obtuse angle, lacking a byssal
out towards auricles, with tendencysinus, and to
meeting
formanterior disk
small margin scales
with only a where
slight change
in curvature; posterior auricle with an outwardly slightly concave
intersected by poorly defined commarginal growth stops; com-
marginal growth rugae accentuated posterior margin
on that intersectshalf
anterior dorsal margin
of at a slightly
disk andacute
or 900
in auricular regions. Right valve unknown. angle, not prolonged into a sharp point and not projecting
Material examined.-USNM 526466, beyond the posterior
a left limit valve,
of the disk margin.
coral beds
Ornament
of Unit E, South Canyon, Locality 1. of disk consisting of from six to eight first-order ribs;
second-order
Measurements.-Height 13.3 mm, ribs medially intercalated
length 12.0 mm, in central and anterior sec-
hinge
length 6.1 mm. tors, submedially and multiply intercalated in posterior sectors;
Occurrence.-Known only from third-order
Unit E verycoral
fine costellae coveringSouth
beds, interspaces. Can-
Anterior au-
yon, New Pass Range, Nevada, late ricle with five or six evenly spaced radial costae and without
Ladinian.
noticeable intercalations
Discussion.-Asoella? species c differs from ofA.? lower-order
species costae; posterior
a and auricle
b in having radial ornament and with
a less uniformly distributed
higher obliquely radial costae
ovalof at least two orders
prosoc-
line form. Its sparse wavy costae with tendency for first-order
distinguish it from costae to other
be concentrated in middle
species
of Asoella having radial ornament.of auricle. Sharp, evenly spaced commarginal growth lamellae
present in interspaces on both disk and auricles.
Family OXYTOMIDAE Ichikawa, 1958
Shell apparently very thin; shell microstructure not determined
Genus OXYTOMA Meek,but 1864b
probably originally calcitic, without evidence of a differen-
Subgenus OXYTOMA Meek, tially 1864b
recrystallized inner layer. Interior features not preserved.
Known
Type species.-Avicula muensteri Bronn only fromin left Goldfuss,
valves. 1835, p.
Etymology.-Named
131 (=Avicula inequivalvis J. Sowerby, 1819), for the
p. Grantsville
78, by Formation,
original Shoshone
Mountains, Nevada.
designation, Braunjura 8, Thurnau (Oberfranken), Germany. [See
Cox, 1940, p. 96, concerning Bronn's Types.-Holotype:
nomina USNM nuda
526467, external
and mold Gold- of a left
fuss's (1835, p. 131) validation ofvalveBronn's
(Fig. 10.9), limestone
speciesmembername.]
of Grantsville Formation,
Discussion.-Up to four orders of Shoshone
radialMountains,
ribs Locality
occur 5. Paratypes from same locality:
in Oxytoma.
USNM 526468, partial
For purposes of the following descriptions ofleftleft
valve exterior
valves, (Fig. 10.11);
first- USNM
526469, in
order ribs are those that are highest moldamplitude
of partial left disk(or
exterior (Fig. 10.12). with
deepest Paratypes
reference to molds of the external from coral beds of and
surface) Unit E, traceable
South Canyon, Locality
onto 1: USNM
the
526470, mold of LV
beak. The interspaces between first-order exterior
ribs are(Fig.referred
10.10) and USNMto 526471,
as
exterior mold of sectors
sectors. Along a growth line or margin, a left-valve fragment.
differ in width
and are variably subdivided by the Other material examined.-Twoof
intercalation very lower-order
small partial external
molds of left
ribs, each successive order beginning atvalves and an even smaller
a greater left anterior
distance fromfragment
with external surface
the beak. The intercalations are generally medial exposed, all on the blockhigher-
between containing the
ammonoid
order ribs in the central part of the disk Hungarites
but inermis
may(USNM 525404) but not separate-
be off-center in
interspaces or multiple in lateral ly catalogued. particularly on the pos-
sectors,
terior side. All interspaces on bothMeasurements.-Holotype,
disk and auricles height 9.0
of mm,thelength 8.2 mm.
species
described below are covered by exceedingly
Occurrence.--Grantsvillefine radial
Formation, costellae
Shoshone Mountains;
Locality 1; 11, paratype, USNM 526468, exterior of fragmented left valve, Grantsville Fm., Locality 5; 12, paratype, USNM 526469, external
mold of fragmented left valve, Grantsville Fm., Locality 5. Scale bars 5 mm, except for 3, 1 cm, and 10, 2 mm.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
coral beds and ammonoid bed and

Waller of Unit E, South
Stanley (1998
Pass Range, Nevada, late Ladinian.
Triassic bivalves from th
Discussion.--The new ment
species
ofisa assigned to Ox
small pectinoi
(Bittner,
toma) because of its broad 1903)
posterior from
wing, the
poorly
anterior auricle lackingwas based on
a byssal a fragment
sinus, and orn
of three distinct ordersregarded
of radialasribs,
Oxytoma (Ox
the first
narrow, widely separated, and possibly projec
Superfamily
midventral margin, the third-order ribs PECTINO
being mu
costellae. The auricular configuration,Waller,
[emend. particula1
deep byssal sinus in the Family PECTINIDAE
left anterior auricle,Rafd
toma from Eumorphotis. Discussion.-Waller
Two other features (1993
tha
acceptance of
ytoma are great valve discordance, Rafinesque
with the righ
fitting within the disk dae in favor
margins of left
of the Wilkes (1
valve
structure, with the shellto his attentionhaving
of Oxytoma by E. Coa
foliated calcitic inner shell layer.
Wilkes's Neither
work of thes
is incompl
ever, can be determined in the
terests new species, b
of nomenclatural s
valves are available, and recrystallization
menclatural or dis
purposes. I
stroyed or obscured shell microstructure.
(2000, p. 211) in returnin
In its nearly acline andof equilateral
the family. disk, as wel
left posterior auricle that is only weakly pointed
resembles Upper Triassic Subfamily CHLAMYDI
(Carnian) species of O
ma) that Kobayashi and Genus PLEURONECTITE
Ichikawa (1950, p. 2
primitive in the sense thatType they species.-Pleuronec
reflect a postulat
subsequent
Lower Triassic Eumorphotis. Oxytoma designation
species of (
and Jurassic tend to have
of Germany. more acutely pointed
Emended diagnosis.-Prosogyrous,
extended posterior auricles with aacline to opisthoclinedeeply
more Pec-
margin. As an example of
tinidae a transitional
with height commonly greater than length. Left valve species
more convex than right, commonly
photis and Oxytoma, Kobayashi and dome-shaped with the free
Ichikawa (1
0. (0.) sedaka Kobayashi margins of and
the auricles Ichikawa,
oblique and the auricles not separated 1950,from p.
species from Japan. [Hayami,
disk by distinct disk flanks. 1975,
Right anterior auricle p. 62,
with deep byssal regar
as a junior synonym of notch, O.prominent raised byssal
(0.) facsciole, persistent functional
kashiwaiensis Kob
kawa, 1950, p. 223.] ctenolium, and commonly deeply scrolled dorsal margin. Weak
radial costae rarely appearing
Similar so-called primitive features over all or parts ofare disk after presen
an
Ladinian species O. (0.) gussevi
early ontogenetic smooth phase; Kurushin,
auricular costae and antimarginal 1987
microornament
et al., 1996, pl. 17, figs. 7-9), absent.which
Exterior calcitic shell layer of both valves from
differs
in being broader, more prosocline,
antimarginally fibrous; prismatic phase and more
on the right valve absent post
as well as having a less ordered
or very ribbing
brief; inner aragonitic layer pattern
extending nearly to margins
more closely resembles of both disk and auricles and comprising the
specimens hinge. Hinge den-
described b
(1950, p. 86-88) as "Oxytoma sp. weak,
tition poorly known but apparently aff. with only0. inaequ
the resilial
intermedia Emmr." and teeth moderately
"Oxytoma developed. Zigzag and/or sp. radial color
aff.patterns Oxyt
Teller" from the Triassic of the
common, transformed Fujian
into pseudo-ornament Province
by diagenesis (Hag-
China. Chen's specimens, dorn, 1995).all left valves, resemb
villensis in their very smallDiscussion.-The presence
size of a and
triangular resilifer
similarlythat lies be- sh
terior auricles. The Chinese
low the hinge axis specimens
is evidence that Pleuronectites is adiffer,
member of h
of disk ribbing and in the thesuperfamily Pectinoidea as defined byof
presence Waller (1978, p. 353),
a slightly
margin on the anterior and theauricle. The
presence of a true ctenolium places itage of
in the family Pec- the C
tinidae (Waller, 1984).
is apparently Early Triassic becauseThese features make it very unlikely
they that
occur in
Eumorphotis, which isPleuronectites
regarded is derived from or as an to index
closely related Paleozoic Av- foss
and upper parts of theiculopectinoideans
Lower such Triassic
as Streblopteria M'Coy, 1851 in (e.g.,the
New- sout
lio Loriga and Mirabella, ell, 1969a, p. 1986,
N339), which have ap. 245).
different type of resilifer and
Oxtoma (Oxytoma) grantsvillensis
lack a ctenolium. is the oldest
most member of the genus Placement of Pleuronectites
to be within reportedthe Pectinidae is more from prob-
lematic. Many authors have
Although the genus apparently thought that the genusin
originated is closely
the re-
southeastern Asia [based on to
lated or ancestral the specimens
Upper Triassic-Cretaceous Camptonectes ofAg-
(1950) discussed above], assizitsin Meek, 1864b (Allasinaz,
main 1972, p. 323; Johnson, 1984, p.
evolutionary ra
in the Carnian in Japan 113; Waller,
and 1991,eastern
p. 13; Waller and Marincovich,
Siberia 1992, p. (Ko
219),
kawa, 1950, p. 219; see and Waller alsoand MarincovichDagis etPleuronectites
(1992) included al., 199 in the
tending during the late pectinid Carnian
subfamily Camptonectinae and Habe,Norian
1977. It now seems int
chipelago of Canada (Tozer,
likely, however, that 1961,Camptonectesp. 100).
originated from a member Spec of
Oxytoma from the Permian the Triassic Chlamys do group,not perhaps belong
from a species of Praechla-to th
ashi and Ichikawa, 1950, mys Allasinaz,
p. 219-220).1972 in the Late Triassic and that Pleuronectitessc
Oxytoma
1936) from the Loweroriginated Triassic independentlyof from the China,
Chlamys group said in the earlyb
Ichikawa (1950, p. 217) Middleto Triassic.be the oldest repres
toma in the Triassic, isThe nowtype species of Pleuronectites is
assigned to not monomorphic
the fam but
genus Towapteria Nakazawa shows a substantial andrange of Newell,
variation from its earliest 1968, to lates
as characteristic of the occurrences
lower in the Muschelkalk of the Germanic Basin. Based on
Griesbachian (Yan

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
specimens examined in the Hagdorn Stoppani Collection, Muschelkalk

(1858, pl. 21, fig. 5, Pecten Mu-
schmiederi Giebel, 1856, a
seum, Ingelfingen, Germany, those occurring
right valve) and Schmidt (1928,inp. the Lower
128, Pleuronectites Mus-
laevigatus,
chelkalk are of only moderate size, both valves).
notThese show a simple hinge
exceeding apparatus 50
about with only
mm
in height. Their shape is acline to slightly opisthocline,
resilial and infraresilial teeth (terminology ofwith both
Waller, 1986). Al-
valves convex, the convexity of the LV
lasinaz not
(1972, pl. 43, more than
fig. 6b, c) showed about
a similar dentitiontwice
but of
that of the right valve. Some specimens
higher relief inin samples
a left from
valve he identified the Low-
with Pleuronectites bal-
er and Upper Schaumkalkbank of the
atonicus Lower
(Bittner, 1901). TheMuschelkalk (Mu2,
specimen, however, is actually Pec-
Anisian-latest Pelsonian and earliest Illyrian
ten inornatus in (cf.
Stoppani, 1858 age; see
Allasinaz, 1972,Hagdorn,
pl. 36, fig. 5a-
1991, fig. 3) rarely may show low radial
c, and costae
Zardini, 1981, pl. 19, fig.on
4a, b),the disk after
a Ladinian-Carnian species
an initial smooth phase that ranges from
that is herein assignedabout 14genus
to the entoliid toCrenamussium
25 mm New-in
height. These costae, when present, ton, 1987.are usually confined to the
anterior and posterior parts of the disk of identified
Specimens either withvalve, more
Pleuronectites rarely
laevigatus from the
extending onto the bases of the left auricles of the left valve and
Tethys outside the Germanic Basin are poorly known, but those
even more rarely distributed across
that havethe entire
been illustrated disk. Such
have a morphology speci-
resembling the strat-
mens are consistent with the descriptions and stratigraphic occur-
igraphically lower Muschelkalk specimens. Pleuronectites cf. lae-
rences of Pecten (Streblopteria) laterestriatus Philippi, 1899 and
vigatus of Lerman (1960, p. 30), from the late Anisian to early
Pecten (Pleuronectites) schmiederi Giebel,
Ladinian Lower Member1856. Because
of the Saharonim Formation they
of Ramon,
appear to co-occur with noncostate forms that are similar in all
southern Israel, was described and illustrated only from right
other respects, I regard these as junior synonyms of Pleuronectites
valves. (See Druckman, 1974a, 1974b, and Parnes, 1986, for up-
laevigatus.
dated stratigraphy of the Ramon section.) These range from 40 to
Beginning approximately at the stratigraphic level of the Ro-
50 mm in height, appear to be moderately convex, and are only
bustus Zone in the lower Upper Muschelkalk (top of mol 1p, latest
Anisian age; see Hagdorn, 1991, fig. 4) specimens of Pleuronec- slightly opisthocline. Pleuronectites laevigatus of Farsan (1972,
tites laevigatus tend to be larger in size, commonly with heights p. 160), from the Lower Khenjan Series of Afghanistan, was de-
of 100 mm or greater, and have greater differential convexity of termined by Farsan to be of early Ladinian age. Farsan com-
valves with the left valve greater than twice the convexity of the mented that although the right valve is similar to typical Mus-
right valve. Right valves are more commonly nearly flat and left chelkalk specimens, the left valve is smaller, inferred herein to
mean less convex.
valves are deeply domed, with the left umbonal region pushed
dorsally over the hinge line by the deeper coiling of this valve. Until recently, the oldest known representatives of both Pleu-
It also seems that in the highest parts of the Muschelkalk com- ronectites and Praechlamys were of Middle Triassic (Anisian) age
pared to the lowest parts, shells are more opisthocline and more (Allasinaz, 1972, p. 228), but it was not known which genus may
prosogyrous, leading to greater development of the anterior part have appeared first. Recently, Pleuronectites has been reported
of the disk relative to the posterior part. The radial costae that from the Lower Triassic, but these assignments to the genus are
appear on some specimens of Pleuronectites in the Lower Mus- uncertain or incorrect. Pleuronectites puerensis Guo, 1985, from
chelkalk are not known to occur in the Upper Muschelkalk. Well- the Lower Triassic (upper Induan and lower Olenekian) of China,
preserved specimens are not numerous enough to permit a quan- is probably a member of the Lower Triassic aviculopectinoidean
titative analysis of stratigraphic trends, and it is unknown whether genus Eumorphotis, indicated by its shape and ligament impres-
these general trends in the Muschelkalk are stepped or continuous. sion [compare to Eumorphotis multiformis (Bittner, 1899) figured
Hagdorn (1995) described a process whereby zigzag, radial, or by Newell and Boyd, 1995, fig. 28]. Pleuronectites? falcaurita
commarginal color patterns in the shells of Pleuronectites laevi- Guo, 1985, from the Lower Triassic of China, also appears to be
gatus are brought into relief by diagenesis, thereby producing a an aviculopectinoidean, judging from the impression of the liga-
pseudo-ornament in the outer calcitic shell layer or in the im- ment area shown in Guo's plate 23, figure 8a. Pleuronectites mee-
pressions of this layer in the rock matrix. As noted by Hagdorn ki Newell and Boyd, 1995, from the Lower Triassic (Spathian)
(1995), past observers of such "sculpture" used it as an erroneous Virgin Limestone of Nevada, is more likely an entoliid similar to
basis for naming new taxa, examples being Pecten vestitus Gold- the genus Pectinella Verrill, 1897. It apparently lacks a ctenolium
fuss, 1844, Pleuronectites laevigatus derognati (P. H. Fischer, and clearly does not have the prominent byssal fasciole that is
1925), and P. laevigatus zonatus (P. H. Fischer, 1925), all of present in Pleuronectites. Pleuronectites pulchrus Yin and Xu,
which are junior synonyms of Pleuronectites laevigatus. Although 1992, from the early Anisian of China, is too poorly known to
this feature is particularly common in the Upper Muschelkalk, its
be certain of its generic assignment. However, its unusually great-
limits depend on diagenetic conditions. As reported below, such
ly flaring right anterior auricle, very deep and narrow byssal notch
pseudo-ornament also occurs in the new species of Pleuronectites
without a clear fasciole, and its sharp posterior disk flank suggest
described from the New Pass Range of Nevada. It is likely a
that it is not a member of this genus. Although specimens have
feature of limited taxonomic value at the species level.
not been illustrated, the oldest true Pleuronectites is apparently
In many specimens of Pleuronectites laevigatus that I have
the species identified by Assmann (1944, p. 14, 22) as P. laevi-
examined, there is a shallow commarginal step on the inner sur-
gatus from the lowest part of the Gogolin Formation of Poland
face of the outer calcitic layer very close to the margins of disk
in the so-called Pecten and Dadocrinus limestones (Assmann,
and auricles. This likely represents the distal edge of an originally
relatively thick aragonitic inner layer. The hinge structure of Pleu- 1944, p. 13). This horizon is of early Anisian (Bithynian) age
ronectites is poorly known, because it was composed of aragonite, (Hagdorn, 1991, p. 15, fig. 3). In the Muschelkalk of Germany,
and so few specimens have ever been found that have the inner, the oldest known Pleuronectites have been found in the Upper
aragonitic shell layers preserved. After the differential solution of Terebratelbank of middle Pelsonian age (H. Hagdorn, personal
aragonite compared to calcite, what remains of the shell is only commun., 2001; see stratigraphic chart in Hagdorn, 1991).
the thin outer calcitic layer devoid of muscle scars and hinge Praechlamys may first appear in the fossil record earlier than
structure, including the resilifer and hinge teeth. Rare specimens does Pleuronectites. Yin et al. (1992, p. 183) listed Praechlamys
that apparently retain either the original inner shell layer or its weiyuanensis (author and date not given) as "a regional index of
replacement and that show hinge structure were illustrated by the uppermost Jialingjiang Formation in southwest China, thus

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
upper Spathian in age." It would

very close to margins; then
original hinge structures, muscle scars,seem
and
ronectites originatedinternal from shell buttresses lost through differential solution of
Praechlamys ara-
thro
costae on the disk. gonite relative to calcite; outer calcitic layer thin and consisting
These morphological differences within and among speciesofofdistally diverging antimarginal fibers or laths. Presence of an
outer columnar prismatic layer in early ontogeny of right valve
Pleuronectites suggest that the origin of the genus was somewhere
in the tropical Panthalassa Ocean. Pleuronectites laevigatus then not determined, but if present not extending more than a few
originated from a population in the Tethys Ocean, with the rela- millimeters from origin of growth.
tively isolated westernmost Tethyan populations in the Germanic Etymology.-Named in honor of Prof. Norman D. Newell, who
Basin evolving to the highly inequivalved and opisthocline end- has been contributing to our knowledge of the Bivalvia of the
members of the species. The Panthalassan stock probably retained Late Paleozoic and Triassic for nearly 70 years.
its relatively primitive, more equivalved, and occasionally radiallyTypes.-Holotype: USNM 526472 (Fig. 11.1, 11.2), a fractured
costate morphology. left valve with the outer surface of the outer shell layer exposed,
colonial coral beds, "Brachiopod Canyon," New Pass Range, Lo-
PLEURONECTITES NEWELLI new species cality 2. Paratypes: USNM 526473-USNM 526481, three right
Figure 11.1-11.9 valves, five left valves, and one fragment, coral beds of Unit E,
Diagnosis.--Pleuronectites of moderate size and nearly equi- South Canyon, Locality 1; USNM 526482 and USNM 526483:
one right valve and one fragment, coral beds of "Brachiopod
lateral form; left valve only slightly greater in convexity than right
valve, with left beak umbo not inflated and beak not projecting Canyon," Locality 2.
dorsally beyond dorsal margins of shell. Radial costae absent on Measurements.-Holotype, estimated restored height, 36 mm,
both disk and auricles; very low, barely perceptible, broad radial restored length 30 mm.
undulations commonly present on disk; zigzag pseudo-ornament Occurrence.-Known only from South Canyon and "Brachio-
present on both valves. pod Canyon" in the New Pass Range, in association with the coral
Description.--Shell outline: Shell of moderate size, with beds of Unit E in South Canyon and its presumed extension into
heights ranging from about 34 to 65 mm and with height generally "Brachiopod Canyon," late Ladinian.
exceeding length (Ht/L 0.96 to 1.33, x = 1.12, n = 8). DiskDiscussion.-The New Pass specimens are more equilateral
and equivalved than any specimens of Pleuronectites laevigatus
nearly equilateral, commonly acline, but varying from distinctly
observed from the Muschelkalk of Germany. The German spec-
prosocline to slightly opisthocline. Anterodorsal margin of disk
imens commonly have left valves that are at least twice the con-
outwardly slightly concave to straight in lateral view; postero-
dorsal margin straight. Umbonal angle 80'-98'. Left valve only vexity of right valves and have beaks that project beyond the
slightly more convex than right, the convexity-to-height ratios dorsal margins of the auricles, whereas in the Nevada specimens,
varying in both valves from about 0.1 and 0.2 (articulated spec- valve convexities are nearly equal and the left beak is on about
imens not found). Disk flanks low and indistinct on left valve, the same level as the dorsal margins of the auricles. The distinct
somewhat more clearly demarcated on right posterior; right doublean- sinuosity of the anterior margin of the left anterior auricle
of the new species was not observed in P. laevigatus, where this
terior disk flank low and steep. Presence of disk gapes not deter-
mined. Hinge line short, about half the length of disk. Auricles margin is generally sigmoidal.
unequal in length, the anterior ranging from 1.33 to 1.9 times the The two right valves of Pleuronectites cf. laevigatus from
southern Israel illustrated by Lerman (1960, pl. 4, figs. 11, 12)
length of the posterior measured at the hinge line. Right anterior
auricle with deep byssal notch and a prominent wide, raised bys- resemble the Nevada species in shape of auricles but differ in
sal fasciole; functional ctenolium persistent, with a few active being more opisthocline and prosogyrous, leading to a greater
curvature of the anterodorsal margin of the disk in lateral view.
teeth in the byssal notch that are closely spaced in early ontogeny
but become widely spaced and relatively larger later; anterior The valves may also be flatter than are right valves of comparable
margin of this auricle appearing truncated and nearly straight, size in the new species.
with small concavity adjacent to ventral edge of a deep dorsal Pleuronectites hirabarensis Amano, 1955, from the lower Car-
nian Hirabara Formation of Japan, differs from P. newelli in being
scroll. Left anterior auricle with only a very shallow byssal sinus,
anterior margin doubly sinuate with shallow concavities on the highly inequivalved, with the left valve more convex than the
right. The Japanese species is also more opisthocline, with its
dorsal and ventral sides of a central larger outward convexity (Fig.
anterior disk margin extending much further anteriorly than does
11.2, 11.4), the overall trend of this margin nearly perpendicular
to hinge in small specimens, becoming oblique to hinge later; this margin in the Nevada species, and the anterodorsal disk mar-
gin of the Japanese species is more outwardly concave in lateral
dorsal margin not scrolled. Posterior margins of posterior auricles
varying from nearly straight to slightly concave, commonly meet- view.
ing the dorsal margin at an obtuse angle and with overall trendPleuronectites newelli is the first definite member of the genus
forming an angle of 115'-120' with the dorsal margin; posterior to be described from the Triassic of the Americas.
dorsal auricular margins not scrolled.
Ornament: Disks of both valves nearly smooth, with only flat- Genus LOXOCHLAMYS new genus
lying shingled growth lines accentuated by weathering; auricles
with fine, regular, close-set commarginal growth lines except on Type species.-Loxochlamys corallina n. sp., Middle Triassic
right anterior auricles, where coarser commarginal ornament(late is Ladinian), South Canyon, New Pass Range, Nevada (Fig.
accentuated; radial costae absent, but very low, broad, regular 12.1-12.6).
undulating radial ribs visible under oblique light on some speci- Included species.-Pecten chiwanae McLearn, 1941a (Fig.
mens; all specimens observed have strong diagenetically accen- 12.7-12.9), and Pecten sasuchan McLearn, 1941a (Fig. 12.10),
tuated pseudo-ornament consisting of zigzag or diagonal ridges both from the Upper Triassic (Carnian) of the Peace River foot-
with apical angles tending to be 450 or greater (see Hagdorn, hills, northeastern British Columbia, Canada.
1995, for explanation of this structure). Diagnosis.-Equivalved, slightly opisthocline Pectinidae with
Shell interior and shell microstructure: Presence of extensive a persistent byssal notch of moderate depth; ornament of disk
original aragonite nearly extending to the margins of disk and consisting of noncarinate radial plicae or radial undulations over-
lain by a distinctive shagreen or near-shagreen microornament
auricles indicated by a shallow commarginal step on shell interior

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
-4; P. I .-,,, ,
'al , .. ? , a okk
.? ? - 1. I I 0 - ..1
I' ?.' -
?. . r
" _1?
k ` ??n~n~anu?,~
. ? _r khl5j~:;;
"
". I I , 1 .1 i : ?1? ? ? .??1 7 - - 1,?,p~~l ~ ~hk~ ~-;rT3i~l'l , " ,_?A ?_ .:" ,- ,
:?i,?,, I '?.? :
?V -.4 .,
, -`%
" . ;I .
." , , ?-_ ? - -w,,

1'?. ?-7? ,
. - k , ? .dr 1~~B~lg~-a'b~
? .
_"'?. :. , , ,??_:s
"jf?* , ,,,4
-",: ??;j ,. 1 -" ?7_,??,i'?,__,? - ?,-?? ?
,- ?11??. M?,_ ? -
I- .
,, ? , 4,
11,? , ? ,4
, _..
1 1I? - . _ " Z . , ? - - - - , - 1 ; Z _ .
. _ ,
-?,..:",? .1.,, ,_?? A,,_z_,; ,?.?, `,',,
FIGURE 11--Pleuronectites newelli n. sp.: 1, 2, holotype, USNM 526472, exterior of fragmented left valve and d
paratype, USNM 526473, interior of outer shell layer of nearly complete right valve and detail of right anter
(arrow); 4, paratype, USNM 526475, exterior of left anterior auricle; 6, paratype, USNM 526477, interior of ou
right valve showing faint radial undulations on disk; 7, 8, paratype, USNM 526479, exterior of left valve showi
and detail of zigzag pseudo-ornament; 9, paratype, USNM 526478, external mold of zigzag pseudo-ornament of f
All from Unit E coral beds at Locality 1 except for 1 and 2 from probable extention of these beds at Locality
3, 6, 7, 1 cm, 2, 5 mm, and 8, 2 mm.
that merges into antimarginal ridgelets near and interspaces

on disk and dorsally
flanks across plicae, becoming replaced by
and is replaced by coarse commarginal lamellae near coarse commarginal
ventral lamellae
mar-in late ontogeny; shagreen or near-
gin. shagreen microornament present through most of ontogeny, giv-
Description.-Shape equivalved, procrescentic, and slightly ing way to antimarginal ridgelets in anterior and posterior sectors
opisthocline. Disk ornament consisting of simple radial plicae that of disk and extending onto disk flanks. Byssal notch of moderate
vary in amplitude among species from obscure radial undulations to substantial depth, with a fine but persistent functional cteno-
to high-amplitude plicae but in all cases lacking internal carinae, lium. Auricles commonly non-costate, rarely with a few weak
and fine commarginal growth lines that tongue ventrally in plical costae. Outer shell layer of mainly antimarginal fibrous calcite

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
I
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1; ;?;t ?V "Iwj!,
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,,? M ..Ik-.,?-4.
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?51 I I
,?tkl?"i '?kl-', ?,-'Akl m:.1 .
FIGURE 12-1-6, Loxochlamys corallin

valve: 1, whole valve; 2, detail of orna
paratype, USNM 526484, left valve,
left anterior auricle, scale bar 2 mm;
mm. 7-9, Loxochlamys chiwanae (McL
GSC 8786, left valve, view of entire v

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
and an extensive inner aragonitic as is also the

layer opisthocline shape,
extending apparent to
nearly lack mar-
of typical anti-
gins on both disk and auricles; presence of an and
marginal microornament, outer prismatic
the antimarginal or radial fibrous
layer in early ontogeny of right valve microstructure
not of the outer calcitic shell layer.
observed.
Etymology.-The new genus name combines the Greek adjec-
tive, loxos, with the pectinid genus name LOXOCHLAMYS
Chlamys.CORALLINA new species
The Greek
adjective, meaning slanting or crosswise, calls Figure 12.1-12.6
attention to the
peculiar shell microornament present in all members of the new
Diagnosis.-Loxochlamys having 13-15 (commonly 14) prom-
genus.
inent radial rounded-trigonal plicae that begin early in ontogeny
Occurrence.-The new genus is known only from the Triassic
and remain of moderate height throughout ontogeny.
of Nevada and British Columbia, with a stratigraphic range from
Description.-Shell outline: Shell of small size, with heights
Ladinian (Nevada) to Carnian (British Columbia).
ranging from about 9-25 mm and with height consistently ex-
Discussion.--The combination of characters described above
ceeding length (Ht/L 1.02 to 1.41, R = 1.20, n = 16); acline to
sets Loxochlamys apart from all other known pectinid genera. slightly opisthocline, with anterior half-diameter of disk com-
Among Triassic Pectinoidea, a near-shagreen microornament is
monly exceeding posterior half-diameter. Anterodorsal margin of
known only in Pecten rosaliae Salomon, 1895, from the Ladinian
disk straight to slightly outwardly concave; shorter posterodorsal
of Italy (Allasinaz, 1972, p. 307). The holotype of this species in straight to slightly convex. Umbonal angle about 900, with
margin
Miinich (Bayerische Staatssammlung ftir Paliontologie undanterodorsalhis- margin of disk forming a greater angle with the hinge
torische Geologie, Catalogue No. 1892.X.32), which I examined, line than does the posterodorsal margin. Left valve slightly more
is a right valve that bears a ctenolium associated with a deep convex than right valve, the convexity of the left about 14% of
byssal notch and a distinct byssal fasciole. Its outer shell layer valve is height and convexity of right valve about 12% of valve
of undetermined microstructure but does not appear to be simple height. Disk flanks low and with shallow slope toward plane of
columnar prismatic. The presence of a ctenolium and lack commissure.of a Disk gapes not observed directly but probably ab-
persistent columnar prismatic shell layer contradict Allasinaz's sent. Hinge line short and asymmetric, commonly about 50%-
(1972, p. 307) placement (with a query) of this species in 60% Filo-of shell length. Auricles unequal in length, the anterior com-
pecten Allasinaz, 1972, because Filopecten is an entoliid genus monly 1.5 to nearly twice the length of the posterior measured
that lacks a ctenolium throughout ontogeny and has a persistent along the hinge line. Right anterior auricle with acute to rounded
columnar prismastic outer shell layer on its right valve. Although byssal notch of moderate depth; byssal fasciole clearly demarcat-
clearly a member of the Pectinidae, Pecten rosaliae differs ed, from moderately broad, slightly outwardly convex and lamellose;
members of Loxochlamys in lacking both radial and commarginal fine functional ctenolium present; anterior margin of this auricle
ornament and in having broad, low commarginal undulations. with shallow outwardly convex curvature meeting hinge line at
Other aspects of the shell microstructure of P. rosaliae areabout un- 900, dorsal margin straight or with a shallow scroll. Left
known.
anterior auricle with only a very shallow byssal sinus or none at
The antimarginal fibrous calcitic outer shell layer, configuration
all, the free margin of this auricle commonly with a central out-
of commarginal growth lines that parallel a dentate disk margin,wardly convex part bordered dorsally and ventrally by a shallow
and presence of simple plicae that commonly are trigonal in crosssinus, the overall trend of the margin being oblique to hinge line.
section are features that Loxochlamys shares with Pseudopecten Posterior margins of posterior auricles outwardly concave, with
dentatus (J. de C. Sowerby, 1827), a species that is distributed dorsal part of this margin intersecting hinge line at about 90' or
mainly in the Lower and Middle Jurassic (lowest Hettangian to less and overall trend of margin oblique to hinge line; posterior
highest Bajocian) of the western Tethys (Johnson, 1984, p. 75).dorsal auricular margins straight, not scrolled.
Pseudopecten dentatus, however, is prosocline, more equilateral Ornament: Both valves with from 13-15, most commonly 14,
and flaring, and lacks shagreen or near-shagreen microornament.simple interlocking radial plicae that originate less than 2 mm
Furthermore, the plicae of P. dentatus are internally carinate, itsfrom beak and continue to ventral margin; plicae rounded to
ctenolium becomes overgrown and ceases to be functional in late broadly trigonal in cross section, with shallowly sloping flanks,
ontogeny, its antimarginal fibrous calcite is limited to only the narrowly rounded crests, lacking carinate edges on shell interior,
outermost part of the outer shell layer, which otherwise is foliatedand interplical spaces rounded in cross section. Shell micro-or-
calcite, and its ligament system migrates ventrally during on-nament of disk surface complex, resulting from interplay of
togeny (new observations). A possible ancestor of P. dentatusoblique microridges and commarginal lamellae. In proximal
postulated by Johnson (1984, p. 77), Pseudopecten coronatiformis region, oblique microridges dominate. In midontogeny, oblique
(Krumbeck, 1924), from the Upper Triassic of Timor, has the microridges and commarginal lamellae interact to produce a quin-
sculptural features of Pseudopecten (Pseudopecten) Bayle, 1878cuncial pattern of tiny cusps or nodes, recalling shagreen microor-
and not of Loxochlamys. The same applies to Chlamys coronatanament of some modern pectinids (Waller, 1972a, fig. 12), but
(Schafhliutl, 1851) from the uppermost Norian of Europe (John- adjacent cusps not merging to form typical shagreen pattern; be-
son and Simms, 1989, p. 192). ginning at shell height between 10 and 19 mm, commarginal la-
On both stratigraphic and morphological grounds, the origin of mellae become more lamellose and dominate the ornament pattern
Loxochlamys is probably from an ancestral species in Praechla-(Fig. 12.1, 12.2); commarginal lamellae shallowly tongued dis-
mys, a genus that was already well diversified by Ladinian time tally in plical interspaces and deeply tongued or "V'd" dorsally
in Europe (Allasinaz, 1972). If so, the lack of well-developed over tops of plicae. Disk flanks either smooth except for fine
auricular costae in Loxochlamys is a carryover from Praechlamys,commarginal growth lines or crossed by continuations of oblique
2 mm; 9, paratype, GSC uncatalogued, left valve, GSC Locality 9512, Mahaffy Cliffs, north side of Peace River between Folded Hill and Schooler
Creek, British Columbia, scale bar 10 mm; 10, Loxochlamys sasuchan (McLearn, 1941a), holotype, GSC 8787, Lima? poyana fauna, east of
Pardonet Hill, Peace River foothills, northeastern British Columbia, right valve, scale bar 1 cm.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
L. sasuchan
microridges of disk, these becoming is antimargina
also a strat
found
disk flanks and on adjacent that Loxochlamys
surfaces of posterio
Cliffs fauna of
ment on auricles consists in the Peace River foothills of British Columbia,lame
commarginal
absent except on right whereas L. sasuchan occurs
anterior in the "Lima? poyana
auricle, fauna" in the th
where
weakly developed and samefew in
area. Later number.
McLearn (1941b, p. 99; 1941c, p. 96) described
the "Lima?
Shell interior and shell poyana fauna" as stratigraphically succeeding
microstructure: Inner the
accompanying muscle Mahaffy
scars not
Cliffs fauna which, preserved, but
in turn, occurs well above the "Na-
thorstites
ginal ridge near the shell fauna." The Nathorstites
margin Zone was later subdividedthe
indicating into d
the Meginae, inner
was probably an extensive Maclearni, and Sutherlandi
aragonitic Zones (Tozer, 1967,shel
p.
64). Because
prised the shell interior Loxochlamys corallina
including the in South
hingeCanyon occursstru
citic shell layer recrystallized
stratigraphically lower
but
than the Unit
displaying
E ammonoid bed containing dist
fibrous microstructure; evidence
ammonoids oftheregularly
of these zones, it is clearly oldest member of this f
sent; columnar series.
prismatic outer shell layer in e
right valve not preservedLoxochlamys
but corallina
if is the most abundant bivalve
present in the coral
probably
beds. Although no
in height based on position ofarticulated
originspecimens wereof found,radial
right and p
left valves occur in nearly equal
1972b, p. 52, on the relationship numbers (27 right, the
between 21 left), in-
ter
matic microstructure and
dicating a the onset
lack of extensive postmortemof ribbing).
transport. Fragments of
Etymology.-The speciesthe pectinidname refers
are common bioclasts to
in the coral beds the
but are rare c
of the new above the
withspecies coral beds and unknown
colonial below them. The
corals in persistent
the N
Types and other
material examined.-The
byssal notch and ctenolium indicate that L. corallina was byssally h
18048B, is a right valve
attachedwith the
throughout life, concave
probably to dead coral or shells oninner
bioh-
erm surfaces.
outer shell layer facing upward from the rock m
from the coral beds of Unit E, South Canyon, L
Subfamily TOSAPECTININAE
its entire outline is preserved, with Trushchelev,
some1984 [emended]
detai
nament preserved as an Original diagnosis.-Trushchelev
impression in (1984,
the p. 65; translation
matrix
from Russian by R. D. Johnson, with additions
antimarginal fibrous microstructure presentor clarifications
(Fig
in brackets): "The
from Locality 1 that show shell reaches largemorphologica
certain dimensions, is sometimes
ularly clearly are catalogued
thick-shelled, of rounded as USNM
outline, 526484-
almost equilateral, equivalved
ples shown in Fig. 12.4-12.6).
or with one of the valvesOther USNM
being strongly convex. The auricles are par
cality 1 are uncatalogued
usually largebut and well bear
delimited from USGS locality
the disk. The anterior au-
34046, 34048, 34056, 34058,
ricle is somewhat larger and than the34067.
posterior. A byssalIn notch istheal-
ways present. The ends
leontology of the University ofof the Montana,
auricles are drawn upward parat from
on rock pieces bearing the
the hinge catalogue
[referring numbers
to deep scrolls] in both valves. The left valve
18029, 18029A, 18035,has18035B,
two, more rarely, one,18048B lateral swelling [of the(which
external disk a
holotype), and 18048-C.surface],In total,
separated from the [centralthe part of] material
the disk by [radial] e
of 24 partial or nearlydepressions.
whole Sometimes right
there is one largevalves,
lateral swelling on the 20 p
whole left valves, and right
about 150
valve on the anterior part fragments.
of the shell. The ornament consists
Measurements.-Holotype, height
of distinct radial ribs, differing on (restored
the right and left valves. Some- estim
length 14.9 mm. times fine radial lines are observed. Concentric [commarginal]
Occurrence.-The holotype
growth lines are and most of the para
usually distinct.
the coral beds of Unit E "Pseudo-hinge
in South apparatus is well Canyon
developed and consistsof of sim- the
A few specimens were also
ple cardinal found
crura [hinge teeth] divergingin alongSouth
both sides from Ca
in Unit E about 12 m theabove
resilium [resilifer],the the lateral bolsters [resilial
coral teeth] on the i.e
beds,
between the coral beds right
and valve fitting
the into overlying
lateral sockets on the left valve,ammon and of
auricular crura [basal
chiopod Canyon" (Locality 2), auricular
3.9 buttresses]
km with north
distal teeth [tu- of
on the western flank bercles].
of Anteriorthe distal
Newteeth [dorsalPassteeth?] are absent.
Range, p
fragments of the species "The inner were found
ligament is situated in associat
in the triangular-oval resilifer.
The inner newelli
corals and Pleuronectites surface [of the disk] isn. ribbed,sp.
with a large
This oval muscleasse
[adductor] scar displaced
a continuation of the South Canyon toward the posteriorcoralmargin. The pallial
beds i
Discussion.-The three line known
is whole [i.e., entire, species
not discontinuous], becoming
of Loxoc more
similar shape and micro-ornament,
deeply impressed and consisting of pits in including
the dorsal part of the a
shell."
come commarginally lamellose late in ontogeny
differs from Loxochlamys Emended diagnosis.-Pectinoidea
chiwanae of medium to large size,
(McLearn,
12.9) and L. sasuchan (McLearn,
having distinct radial plicae that lack 1941a)
internal carinae and (Fig.
begin
a greater modal rib count (14 Left
in very early ontogeny. asvalve compared
with length at maturity com- to 1
of the new species distinctly
monly exceeding height, higher
the rapid ontogeneticand increasebeginn
in length
togeny. In the series L.
producingcorallina-L.
disk flanks that are outwardly chiwanae-L
concave in plan view.
is a stepwise decrease in rib
Anterior and/orheight
posterior sectors of and a lengthe
left valve commonly flattened
or inflated, during
of appearance of the ribs the plicae on these sectors weaker and commonly th
ontogeny,
uchan being so low in more closely spaced than on theand
amplitude central sector. Strong valve in-
beginning
eny (at a distance from beak
equality common, eitherof about
the right or left being12the moremm)
convex. t
ficult to count. In the same
Auricles series,
noncostate or there
only weakly and sparsely is
costate, with freeals
marginsright
relative convexity of the of auricles commonly
valve meetingrelative
the hinge line at acute
to t
two Canadian species areangles; anterior
much auricle slightly
closerlonger than posterior;
to one dorsal mar-ano
ically than either is toginsL. corallina.
of right auricles commonly with hypertrophied scrolls, es-
The morphological series
pecially on theLoxochlamys corallin
anterior; auricles of left valve with straight or only

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
None has ever

very weakly scrolled dorsal margins; been described,
byssal and the published
notch figures of to-
of variable
depth, commonly persisting throughout ontogeny;
sapectinine species are of insufficientctenolium ab-
quality to determine wheth-
er onein
sent in late ontogeny, not observed is present.
early The presence of a true ctenolium,
ontogeny. at least in
Antimar-
ginal microornament obscure or early
absent;
ontogeny, isdisk
a definingflanks
autapomorphynon-costate
of the family Pectin-
idae (Waller,
and without cross-ridging; regular 1984). Among pectiniform
commarginal lirae andcommonly
especially amusi-
present on disk and auricles. Hinge iform dentition
pectinids, the ctenolium becomes obsolete
simple, in very early
dominated
by resilial teeth and sockets; dorsal ontogeny
teeth but remains
weak detectible
orin absent.
mature specimens by close
Discussion.--The subfamily Tosapectininae examination of the suture asbetween the disk and theunder-
currently right anterior
stood is known only from the Middle auricle (Waller,and 1984). Although
Upper disappearance
Triassic of the ctenolium
and
consists of three genus-group taxa: even in very early ontogeny is theoretically
Janopecten Arkhipov possible, this has
and not
Trushchelev, 1980, of the upperyet been observed in
Anisian to anylower
taxon knownCarnian
to be derived phyloge-
of
northeastern Russia and central Siberia; netically from a ctenolium-bearing
Tosapecten ancestor.
(Tosapecten)
Kobayashi and Ichikawa, 1949, of The the Tosapectininae,
lower however,
Carnian may be oneto groupupper
in which a
Norian of Asia and western North ctenolium has disappeared and
America; during the course of evolution through
Tosapecten (In-
digiropecten) Trushchelev, 1984, of heterochrony.
the early That Tosapecten
and is peramorphic
middle (s. McNamara,
Norian
of central Siberia. All of these reach 1986) relative
a large to Janopecten
sizeis (60indicated
mmnot onlyorby the shallow-
larger)
and are radially plicate. The main ing of its byssal notch in early ontogeny,
differentiating but also by the short-
characters
among the genera involve differences ening and possible disappearance of thebetween
in convexity prismatic stage of thethe
two valves, the extent of scrolling rightof valve the
and corresponding
dorsalpreradial marginsstage of theof left valve.
the
right auricles, the relative size of This is evidenced by
auricles inspection of figures
relative to sizeof tosapectinine
of disk, genera
and the degree to which the byssal in Bychkov
notch et al. (1976),
becomesArkhipov and Trushchelev (1980),
shallow in lateand
ontogeny. Trushchelev (1984). If a ctenolium is indeed present in the early
Arkhipov and Trushchelev (1980, p. 12) observed that early ontogeny of the earliest Tosapectininae and is secondarily absent
representatives of Janopecten in the Anisian have a left valve that in later representatives, then the subfamily belongs in the Pectin-
is only slightly more convex than the mildly convex right valve. idae. The first occurrence of Janopecten in the Upper Anisian
Beginning in the Ladinian and extending into the early Carnian, would suggest that the Tosapectininae result from an early branch
however, species of Janopecten developed strongly inequivalved of the Chlamydinae.
shells with the right valve flat or even concave and the left valve However, Hertlein (1969, p. N371) regarded Tosapecten as a
strongly inflated. The auricles are relatively large and nearly equal subgenus of Weyla Bihm, 1922, and placed it in the "Neithea
in size, the right auricles having only weakly developed scrolls Group" within the Pectinidae. Damborenea (1987, p. 167) re-
or none at all. The byssal notch remains deep throughout ontog- garded Tosapecten and Weyla as separate genera but regarded
eny. Weyla and Neithea Drouet, 1824 as closely related and placed
them both in the family Neitheidae Sobetsky, 1960, in which So-
Tosapecten s.s., in contrast, has a mildly to strongly convex
right valve and a flat to concave left valve. Its right anterior au-
betsky (1960) had included only Neithea. Weyla is restricted to
the Lower Jurassic (Damborenea and Mancefiido, 1988) and Nei-
ricles are commonly deeply scrolled, and there is a tendency for
thea to the Cretaceous (Dhondt, 1973).
the byssal notch to become shallow in later ontogeny. Arkhipov
A close relationship between Tosapecten and Weyla is unlikely
and Trushchelev (1980, p. 12) thought that Tosapecten subpolaris
for several reasons. Damborenea and Mancefiido (1979) showed
Polobotko in Bychkov et al., 1976 may exemplify a transitional
that Weyla originated in western Pangea in the Hettangian (low-
state between Janopecten and Tosapecten s.s. This species occurs
ermost Jurassic) and by at least Pliensbachian time had spread
in the lowermost Carnian (the Protrachyceras omkutchanicum
eastward into the Tethyan region with the opening of the Hispanic
Zone) of northeastern Russia (Bychkov et al., 1976, p. 11, 74). Corridor between North and South America (Damborenea, 2000;
Tosapecten (Indigiropecten) resembles Tosapecten s.s. in its
Aberhan, 2001). Although the first occurrence of Weyla closely
basic shell outline and ribbing pattern, the difference being that
follows the last occurrence of Tosapecten, the morphology of
here the valves become nearly equiconvex, because the left valve
members of the Weyla Group does not corroborate phylogenetic
becomes secondarily convex after a flatter stage in early ontogeny. descent. Among the oldest members of the Weyla clade to occur
The auricles of T. (Indigiropecten) are somewhat shorter relative in western North America is a nearly biconvex specimen (CAS
to shell length than are those of Tosapecten s.s., and the byssal 61459) identified by S.W. Muller as "Pecten acutiplicatus Meek"
notch of T. (Indigiropecten) becomes very shallow by maturity. (Meek, 1864c, p. 46) and collected from near the base of the
The right auricles of T. (Indigiropecten) are very deeply scrolled, Sunrise Formation in the Gabbs Valley Range of Nevada (Muller
producing a distinctly V-shaped dorsal margin. Trushchelev and Ferguson, 1939, p. 1611). This specimen is unlike the type
(1984) claimed that the left auricles may also be deeply scrolled, of the species, which was determined by Aberhan (1998, p. 124)
but his photographs do not indicate the very deep scrolling on the to be piano-convex and assignable to Weyla (Weyla). On the basis
left valve that is shown in his drawing (fig. 1). Tosapecten (In- of ammonoids associated with the nearly biconvex specimen,
digiropecten) is known only on the basis of its type species, T. Muller and Ferguson (1939, p. 1612) correlated this part of the
(I.) mirabilis Trushchelev, 1984, which occurs in the Lower No- section with the Arietites ["Ammonites"] bucklandi Zone, of early
rian Pinacoceras verchojanicum Zone and lower Middle Norian Sinemurian age. They reported Weyla alata (von Buch, 1838) and
Otapiria ussuriensis Zone of eastern Yakutiya in central Siberia W. cf. W. bodenbenderi (Behrendsen, 1891) higher in the same
[Trushchelev, 1984; see Dagis and Tozer (1989, p. 4) for strati- section. These species are now assigned to Weyla (Weyla) by
graphic placement of ammonoid zones]. Tosapecten (Indigiropec- Damborenea (1987) and Aberhan (1998).
ten) is probably derived from Tosapecten s.s., because its distinc- All of these species have features that have never been ob-
tive characters can all be derived from character states already served in either the Tosapectininae or in Neithea. Most important
present in that subgenus. are a well-formed ctenolium in early ontogeny, prominent cross-
The family-level placement of the Tosapectininae within the ridging on the disk flanks, a broad hinge plate that is completely
superfamily Pectinoidea is still an open problem, in part because crossed by transverse ridgelets, and a distinctly foliated calcitic
it is still unknown whether any tosapectinine has a ctenolium. outer shell layer. It is also unlikely that Neithea evolved from

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Weyla because of substantial

they differences in the mo
are acutely triangular
early ontogenetic stages.
erWeyla has a rather
Tosapectininae, typic
Nevadap
early ontogeny with a onstrong ctenolium,
the left valve thata deep
are d
ricles having rounded or sigmoidal
a change margins, an
in transverse cu
prismatic stage on the and/or
right valve
width andofpreradial
radial ri
valve. New observations 11)onand
Neithea, in contrast,
Trushchelev (1984
swellings
has a highly unusual early ontogeny separated from
that is distinc
The prismatic/preradial tion]
stageand
is commented
absent, and on d
radi
tors among
the edge of the prodissoconch. tosapectinin
The auricles in ear
relatively large and distinctly trigonal,
closely resemble withof
those th
meeting the hinge line ferentiated
at acute angles.sectorA ctenoliu
is the le
the right anterior auricle
Withlacks a clearly
regard defined
to the mor
A more likely possibility is that Weyla
genus-group is des
taxa describe
lineage of Pseudopecten close to
opecten andPseudopecten
Tosapecten de s.
scrolling
on, common in the Hettangian ofof auricles,
Europe but and
poss
in the earliest Jurassicdapecten
of North America,
also occupies share
an
probable
early ontogenetic characters with late Ladinian
Weyla. Furtherage
of Pseudopecten dentatus,
of the based on new
Anisian dataea
and the f
28178 from the Middle nian.
Lias of England, has ex
crossing the entire hinge plate and only very w
NEVADAPECTEN LYN
resilial teeth, features that are nearly identical to
Figure and
and differ from those of Tosapecten 13.1-13.4
Neithe
plano-convex shape, pointedDiagnosis.-As
auricles, smoothfor thedis ge
Description.-Shell
sence of antimarginal microstructure outlin
in Neithea
Tosapecten, there are no mumknown shell length ofspeci
transitional abou
ed in the time interval L between
0.8 for the theholotype);
disappearan s
pectininae at the end of nearly acline and
the Triassic andorthogy
the fir
of disk broadly
Neithea in the Early Cretaceous. circular, t
Nonetheless,
and posterior disk
trends present in the Tosapectininae pointflanks.
towa
possible descendant. of disks of both valves str
in lateral view. Umbonal a
Genus NEVADAPECTEN a right valve,
new and 1400 on
genus
with the anterior umbonal
Type species.-Nevadapecten lynnae n.
equally convex, withsp.,ma M
(Ladinian) of the New Pass Range, Nevada,
height; maximum inflation by m
Diagnosis.-Biconvex Tosapectininae
vature of right with
disk broad
flatte
gle and nearly equilateral form; byssal
gins, especially toward notch the p
auricles and right posterior
flattened auricle
toward with conca
anterior
meeting hinge at acute anterior
angle but andwith overall sec
posterior tr
forming an obtuse angle with
part ofhinge line; flanks
disk. Disk disk fla o
steep and with ornament not preserved;
setting off auricles ornam
from
sisting of low, rounded, linesimple plicae of
symmetric, only
with hingon
valves; dorsal margins of right auricles
anterior auricle not prese stra
scrolled, those of left auricles
persistent straight
byssal and not sc
notch o
ornament not preserved except for the
served or detected. Left right post
wavy commarginal growth dorsal lines
marginbut lacks radia
and obtus
entiated anterior and posterior
lacking a byssal sinus.pres
disk sectors Rig
valve, where the posterior sector
of dorsal is wellstraigh
margin develo
terior sector only weakly developed.
posterior margin outward
Etymology.-The name an combines
obtuse angle the with
name hingof
vada, with that of the to scallop,
form Pecten.
an acuteNo close
point. Le
lationship with true Pecten
margin, is implied.
posterior margin
Occurrence.-Known obtuse only from angle the withcoral
dorsal bem
South Canyon, New Pass Range, Nevada,
Ornament: Right disklate L w
Discussion.--The new genus
broader resembles other to
than interspaces
reaching a size that is large
toward for anterior
Triassic pectinids
and post
and in having radial plicae that simple
rounded, lack internal
plicae na ca
from Janopecten and Tosapecten
appearing on (Tosapecten)
the anterior in
equally convex valves, plicae
with and a simpler
interspacespatternof b
plicae. It differs from marginal
Tosapecten lirae(Indigiropect
with spaci
flattened early stage in about 35 mm. These
the development of itscomm
lef
longer auricles relative to length
across plicae and of disk,
ventral in
scrolled dorsal auricularmarginal
margins,microornament
and in having aa
notch. Although the auricles
auricleand with growth lines on
commargina
poorly preserved, another significant
costae apparently difference
absent o
ricles appear to be obtusely
cle, not triangular
determined in Nevad
on rig

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
:Y..
. . . . . . . . ......I
??' X1' i
Al"r
'w"s; : ,
iv: c:
I'':-~~?c~. $ .a
x w x.:"i ? r?4
2!wo'I5~S~d?- wnE?
W7 li~lt- -Fr
w" ,.tm
zf~?: ??~~iz
4 iM
ft7, K':5bf~
FIGURE 13-Nevad
of posterior auri
internal mold of
Shell interior Occurrence.-Known only from the coral beds of Unit E, a

apparatus South Canyon, New Pass not Range, Nevada, late Ladinian. ac
recrystallized Discussion.-The new species differs from all other members
an
inal calcitic of the Tosapectininae in having a combinationcom of uniform bicon-
crystallized vexity and simple radial plicae of a single order throughout on-
with
aragonitic togeny. Compared to Pseudopecten of the Lower Jurassic, the
comp
served or detected. new species has typical tosapectinine flattened sectors on the left
Etymology.-The new species is named in honor of my daugh- valve, much lower disk flanks lacking cross-ridging, and a thinner
outer calcitic shell layer that does not appear to be foliated.
ter, Lynn Waller Fegley, who assisted with field work in South
Canyon and collected the specimen that is now the holotype.
Suborder PROSPONDYLOIDINA new suborder
Types.-All from the coral beds of Unit E, South Canyon, Lo-
cality 1. Holotype: USNM 526497, a right valve exterior (Fig. Definition.-Pectinoida directly or indirectly descended from
Prospondylus Zimmermann, 1886, which in turn is derived from
13.1-13.3). Paratypes: USNM 526498, a left valve exterior (Fig.
13.4); USNM 526499, fragment of disk interior of right valve,the
on paraphyletic family Pseudomonotidae in the suborder Pecti-
same block as a paratype (USNM 526477) of Pleuronectites new-noidina. Included families are Prospondylidae Pchelintseva, 196
elli n. sp.; USNM 526500, fragment of posterior disk of right(as emended by Hautmann, 2001a, p. 341), Plicatulidae Watson,
1930, Anomiidae Rafinesque, 1815, and Dimyidae Fischer, 1886
valve; USNM 526501, external mold of fragment of right? valve.
Measurements.-Holotype, height 46 mm, length 56.5 mm, Shells are either monomyarian or secondarily dimyarian, inequi
length of posterior hinge line 14 mm. valved, with the right valve generally cemented to a substrate

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
early in ontogeny by Placunopsis means is more derivedof pallial

than Eonomia secretio
in displaying over-
fibrous resilium, but the form
arching of the ligament by theof resilifers
dorsal margins. Furthermore, Pla- varies
triangular (the plesiomorphic condition)
cunopsis has an internal to but
shell ridge similar to that in Eonomia deeply su
row, or to ventrally shifted one that is moreand
strongly developed
dorsally and that extends further
overarched
pos-
gins. teroventrally. This internal ridge indeed appears to be the precur-
Discussion.-Bringing together the Prospondylidae (=Terque- sor of the stalklike resilifer support of more advanced anomiids.
miidae Cox, 1964; see Repin, 1996 and Hautmann, 2001a), Pli- As Todd and Palmer (2002, p. 507) noted, there are abundant
catulidae, Anomiidae, and Dimyidae is supported by recent dis- small bivalves cemented by their right valves in the Triassic of
coveries in paleontology and molecular genetics. Newell and southern Europe that have been referred to Placunopsis, generally
Boyd (1970, p. 253) suggested, on the basis of the shell mor- to Placunopsis ostracina (Schlotheim, 1820) or P. plana (Giebel,
phology of Permian specimens, that there is a morphological and 1856). They were unable to find a byssal foramen in any of the
possibly phyletic series from Pseudomonotis (Pseudomonotis) Triassic specimens that they examined and thus rejected an as-
Beyrich, 1862 through Pseudomonotis (Trematiconcha) Newell signment to Placunopsis, suggesting that these Triassic species
and Boyd, 1970, Prospondylus, Paleowaagia Newell and Boyd, may instead belong to the Prospondylidae.
1970, and Newaagia, to Spondylus Linnaeus, 1758. Spondylus, The position of the Dimyidae within the Pteriomorphia is still
however, is no longer regarded as an end-member of this series, being debated. Yonge (1975, 1978) thought that ligament structure
because its resilium, which has a nonfibrous core, places it of Plicatulidae and Dimyidae is so close that the two should be
squarely in the superfamily Pectinoidea (Waller, 1978, p. 354). combined in a superfamily Plicatulacea (but according to the Prin-
Paleontological discoveries from the Upper Triassic of Iran by ciple of Coordination mandated for family-group names by the
Hautmann (2001a) tied the Plicatulidae to Prospondylus via an International Code of Zoological Nomenclature, fourth edition,
intermediate ancestor similar to the Upper Triassic genus Persia Article 36, and the now mandatory superfamily word-ending, the
Repin, 1996, through the intermediate genera Eoplicatula Carter, superfamily name should be Dimyoidea). The specific structure
1990 and Pseudoplacunopsis Bittner, 1895, to Plicatula Lamarck, in common is the anterior-posterior abbreviation of primary an-
1801. terior and posterior outer ligaments to a central position above
The relationship of the family Anomiidae to Prospondylidae the dorsal side of the fibrous resilium. In addition, each group
and Plicatulidae is less straightforward on paleontological lacks a foot. In the context of placement in an order Pectinoida
grounds. Newell and Boyd (1970, p. 275) described a new genus that excludes true oysters (see preceding discussion of the pter-
Permanomia based on left valves from the middle Permian (Leon- ioidan suborder Ostreoidina), this absence is a unique synapo-
ardian, early Guadalupian) of Texas. Although these have a du- morphy for these two groups.
plivincular ligament and a musculature of a type unknown in Me- Yonge (1975, 1978) regarded the dimyarian condition of the
sozoic and later anomioidans, they regarded Permanomia as the Dimyidae to be a primitive character and the monomyarian state
"first acceptable record of Paleozoic anomiaceans." Previously of the Plicatulidae to be derived. He therefore supposed that right-
the earliest accepted Anomioidans were of Cretaceous age (Keen, valve cementation, present in both dimyids and plicatulids, pre-
1969a, p. N383; see Fiirsich and Palmer, 1982, p. 902 for a dis- ceded the monomyarian condition in this clade, despite the earlier
cussion of so-called anomiids of the Jurassic). However, Fiirsich occurrence of plicatulids compared to dimyids in the fossil record
and Palmer (1982) then discovered a more likely candidate for (Middle Triassic and Middle Jurassic, respectively). Given the
the oldest true anomiid, which they described as a new genus universal presence of two adductor muscles in bivalve larvae,
Eonomia based on right valves from the Middle and Upper Ju- however, the dimyarian condition may have evolved from a mon-
rassic (upper Bathonian and Oxfordian) of England and Norman- omyarian ancestor through paedomorphosis and would therefore
dy. Among the typical anomiid features that they described, the be secondary. This would be more in accord with Hautmann's
ligament structure is less modified than in modern forms, over- (2001a) demonstration of descent of Plicatula from a monomy-
arching of the ligament system has not yet occurred, and the bys- arian prospondylid ancestor that had a normal alivincular ligament
sal foramen is primitive in the sense that it is still connected by structure.
a suture to the anterodorsal margin of the shell. None of these Recent independent molecular genetic studies based on 18S
characters provides a convincing morphological tie between this rDNA (Steiner and Hammer, 2000; D. C. Campbell, 2000) pro-
early anomiid and the Prospondylus-Plicatula clade, but another vide strong support for a sister-group relationship of the Plicatu-
feature of Eonomia described by Fiirsich and Palmer (1982, text- lidae and Anomiidae and establish that these groups are more
fig. 1) possibly does. It is a "tooth-like process" extending pos- closely related to the order Pectinoida than to the order Pterioida,
teroventrally from the posterior part of the ligament structure. the latter now including the Ostreoidina as a suborder. DNA se-
This ridge may have been incorporated into the stalklike support quences of dimyids have not yet been analyzed.
of the ligament structure in later anomiids, and it may also be
homologous with an interior shell ridge that extends posteroven- Family PLICATULIDAE Watson, 1930
trally from the posterior part of the hinge structure in Eoplicatula. [emended Hautmann (2001a)]
Another possible feature that may indicate an anomiid-plicatulid Genus PSEUDOPLACUNOPSIS Bittner, 1895
relationship is the lack of auricles in anomiids and early plicatu- [emended Hautmann (2001a)]
lids. There is no sign of an anterior auricle bordered dorsally by Type species.-Plicatula (Pseudoplacunopsis) affixa Bittner,
an outer ligament in Eonomia or in any other anomiid. Rather, 1895, p. 215, by monotypy, San Cassiano Formation, Middle Tri-
the byssal opening seems to be the result of extension of the assic (Carnian), Italy.
anterior shell margins around an ontogenetically very early strong
byssal attachment. PSEUDOPLACUNOPSIS aff. FISSISTRIATA (Winkler, 1861)
Todd and Palmer (2002) presented convincing evidence that Figure 14.1-14.4
another Jurassic bivalve genus, Placunopsis Morris and Lycett,
Anomia fissistriata WINKLER, 1861, p. 467, pl. 5, fig. 10a-c.
1853, from the Middle and Upper Jurassic of Europe (Aalenian-
Plicatula archiaci STOPPANI, 1863, p. 140, pl. 33, figs. 1-6; p. 158, pl.
Portlandian), is also an anomiid, thereby rendering the monoge- 34, figs. 4, 5; Cox, 1932, p. 103.
neric family Placunopsidae Freneix in Freneix et al., 1986 unnec- Placunopsis fissitriata (WINKLER). WOHRMANN, 1889, p. 201, pl. 6, figs.
essary. In spite of its earlier appearance in the stratigraphic record, 7, 7a, 8.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Plicatula (Pl
1932, p. 103.
Plicatula (Pseudoplacunopsis) fissistriata (WINKLER). COX, 1924, p. 67,
We pl. 1, figs. 13, 14; LERMAN, 1960, p. 28, pl. 2, figs. 17, 18.
ow- WR --,r. - MIN

Description.-Shell of medium size, typically with maximum
diameter of 20-30 mm, rarely reaching 50 mm; outline typically
nearly circular to oval with height somewhat greater than length
and maximum growth gradient posteroventral, more rarely highly
irregular in outline with length greatly exceeding height; nonaur-
iculate and inequivalved; hinge line short. Cemented right valve
more or less flat, its entire surface conforming to topography of
substrate that is typically a shell or shell fragment in early on-
togeny and a coarse bioclastic sediment surface in later ontogeny;
ventral margins rarely slightly upturned, more typically flat
against substrate throughout growth. Left valve typically convex
in early ontogeny, becoming flat or concave later, but with irreg-
ularities due to xenomorphic growth.
i k%
40?~~" Shell originally bimineralic, with very thin, finely textured

61 _ Sm;, ? (granular to very weakly and finely radially fibrous) light gray,
1?7 3 C~i~l:' ?
probably originally calcitic outer layer and much thicker, coarsely
recrystallized but apparently nonlaminate, dark gray, probably
aet
originally aragonitic inner layer. The thick inner layer closely ap-
~~?l A
proaches shell margins, its convex leading edge descending steep-
ly near shell margin to meet the flat, narrow, marginal band of
outer layer, the two layers interleaving along shell margins to
produce a thick shell rim. Columnar prismatic microstructure not
observed at any stage of ontogeny and probably absent.
Interior features including hinge structures commonly not pre-
served due to differential solution of inner shell layer. Hinge
structure unknown, but a fracture section through one left valve
and accompanying mold of inner surface shows presence of an
inner shell ridge directed from hinge region posteroventrally to-
ward posterior side of presumptive site of posterior adductor scar.
?,'e Adductor scar not observed; pallial line consisting of a series of
deeply inset, closely spaced, radially elongate pits.
Radial ornament present, generally restricted to exterior of left
valve and consisting of irregular, wavy, rather coarse costellae
developed primarily in thin outer shell layer but imprinted also
onto outer surface of thick inner layer; radial ornament also pre-
sent on right valves that have upturned ventral margins; in some
right valves internal costellae, conforming in size and spacing to
external costellae of left valve, occur on inner surface of inner
recrystallized shell layer. Outer surface of right valve commonly
coarsely dimpled, the dimples reflecting large sedimentary parti-
cles on the substrate, this topography commonly transmitted
through to the left-valve surface. Inner surface of inner shell lay-
ers of both valves generally commarginally rugose, with com-
marginal growth stops on left valve interrupting the wavy radial
costellae.
Material examined.-Locality 1, South Canyon: upper Unit D,
10 right valves and five left valves; coral beds of Unit E, three
right valves. Figured specimens have been catalogued as USNM
526502-526506.
Occurrence.-Known only from upper Unit D and the c
FIGURE beds of Unit E, South Canyon, New Pass Range, Nevada, Lad
14-Pseudoplacu
USNM nian.
526502, fragment
of inner surface and
Discussion.-Winkler's (1861) original o
description of Anom
coral beds fissistriata
of fromUnit E,
the Rhaetian Kissen Formation of theLo
Bavar
Unit D, Locality 1; 4, U
Alps referred only to the exterior, and thus there was little
of Unit E, Locality 1. A
for his generic assignment. Two years later, however, Stop
(1863) described Plicatula archiaci from the Avicula contorta
beds of Lombardy (Rhaetian). Based on Stoppani's figures of the
exterior, it is nearly identical in size and shell outline to Winkler's
species. Stoppani also figured the hinge, and, as noted by Cox
(1924, p. 67), it is clearly a plicatulid type, and, correcting for

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Stoppani's apparent reviewed

confusion the manuscript and of provided
rightuseful suggesti
wi
1932, p. 103), the relative
improvement. convexity of the
that attributed to the genus Pseudoplacunop
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ABERHAN,
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Lerman
of a specimen from theSubclasses Palaeotaxodonta,
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ABERHAN, M. 2001. Bivalve palaeobiogeography and the Hispan
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of a proto-Atlantic
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affix
The specimens from Arranged
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Similarities pl
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doplacunopsis fissistriata among
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145.
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ZIETEN, C. H. DE. Locality 3.-Wildhorse

1830-1833. Les mining district, Augusta Mountains, about 750 d
P6trifications
gart. ft west of McCoy Mine, Gilbert Creek SW Quadrangle, Churchill County,
ZIMMERMANN, E. H. 1886. Ein neuer Monomyarier aus dem ostthiirin- Nevada, lat 39.8680N, 117.490'W. The specimen is from the USGS Tri-
gischen Zechstein (Prospondylus liebeanus). Jahrbuch der Preussischen assic collections in Denver. Collector: S. W. Muller, 1940.
Geologischen Landesanstalt zu Berlin, 1885:105-119. Locality 4.--(=SU 1319) upper plate of Gillis thrust, Gillis Canyon
(Wildhorse Canyon), Gillis Range, Hawthorne Quadrangle, Mineral
ACCEPTED 10 SEPTEMBER 2004
County, Nevada. Collectors: S. W. Muller and H. G. Ferguson, 1928. See
Muller and Ferguson (1939, fig. 2, pl. 1) for maps of Wildhorse Canyon
APPENDIX and vicinity. These show the approximate location as 5 km east of Walker
Localities Lake at approximately 38o46.3'N, 118'36.0'W. Material from this local-
ity, removed from the USGS collections in Denver, bears the Stanford
Locality .--South Canyon, on the western flank of the New University Pass locality number SU1319. Rocks on both sides of the Gillis
Range, Desatoya Mountains, Churchill County, west central Nevada, ap- were once regarded as belonging to the Excelsior Formation (Mull-
thrust
proximately 43 km [27 miles] northwest of Austin and 1.3 km wester and Ferguson, 1939, p. 1589). Silberling and Roberts (1962, p. 32)
northwest of New Pass Mine (Fig. 1), lat 39036'42"N; long 117029'31"W. excluded the rocks on the upper plate of the Gillis thrust from this for-
(Refer to the 7.5-minute Mount Airy and the 15-minute Edwards Creek mation.
Valley Topographic Quadrangle Maps.) Collected by T. R. Waller, G. Locality
D. 5.--Limestone member of the Grantsville Formation as ex-
Stanley Jr., and assistants in 1997, 1999, 2000, and 2002. posed in faulted segments from Shamrock Canyon south to Grantsville
Locality 2.-Second canyon north of South canyon, informally called Canyon, Union district, Shoshone Mountains, lone (15') quadrangle, Nye
"Brachiopod Canyon," on west flank of New Pass Range 2.25 km north County, Nevada, approximately lat 38051'to 38056'N, long 117035'W.
of South Canyon (Locality 1) and 3.9 km north of New Pass Mine, Mount Collector: N. J. Silberling, 1950-1954. Equivalent to USGS M76 and to
Airy NW (7.5") quadrangle, approximately lat 38030'N, long 117030'W. Stanford University localities SU734 and SU 2109. See Silberling (1959,
pl. 10) for geologic map of area.
Collected by T. R. Waller, G. D. Stanley Jr., and assistants, June 7, 1999.

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
INDEX
Acrochordiceras Aulacomyella ........................ 16, 19

cf. carolinae ..................... ............ 4
austriaca, Palaeolima ............ .........
Avicula contorta beds ............................ 49
hyatti ......................... ............. 4
Zone ............... ....................... 3 Aviculidae ............ ................. . 16
acutiplicatus, Pecten ...................... ......... 45 Aviculomyalina williamsi ......................... 33
acutum, Plagiostoma ............ 1, 32, 33, Fig. 9.10, 9.11 Aviculopectinoidea ........................ 35, 38, 39
aeolicus, Leptochondria ............................ 33 Aviculopinna .................................. 29
affixa, Pseudoplacunopsis ...................... 48-50 backlundi, Bositra ............................... 21
alaeformis, Gervillaria (Gervillaria) ................... 25 Bakevellia (Bakevellia) ............................25
alata, Weyla .................................. 45 Bakevelliidae ...............................24, 38
albertii Bakevellioidea ................................ 24
Gervillia ....................... ............ 25 (Bakevelloides), Bakevellia ..................... 25
Leptochondria ........................ 33, 34 balatonica, Lima ............... .............. 33
Monotis ............ ....................... 34 balatonicus, Pleuronectites ......................... 39
Pecten (Velopecten) ......................... 33, 34 Balatonites ...................................24
Ambonychioidea .............................. 15 beds ...................................... 23
americana baliana, Lima (Radula) ............................ 31
Daonella ......................... ............ 4 baoqingensis, Peribositra .......................... 20
Mysidiella .............................. 10, 11, 15 (Baryvellia), Gervillaria ...................... 1, 24-29
Promysidiella ................................9, 10 ponderosa ......... 1, 4, 5, 7, 25-29, Figs. 8.4-8.9, 9.1-9.4
amphidoxa, Antijanira .............................38 becheri, Posidonomya ............................ 19
angulata, Palaeolima .............................. 32 Beneckeia Zone ................................ 25
angusta var., Posidonomya ......................... 19 bioni, Claraia ................................... 20
annonii, Plagiostoma .............................. 33 bittneri, Enteropleura ....................... 4, 22, 24
Anomalodesmata .............................. 1, 14 bodenbenderi, Weyla .............................. 45
Anomiidae ................................... 47
bonaevistae, Cuccoceras ............................
Antijanira amphidoxa ............................. 38 borealis, Promytilus ................ .. ............. 10
antiquata, Antiquilima ............................. 31 Bositra ....................... 1, 3-5, 7, 16-23, Fig. 5
Antiquilima .............. ...................... 31 backlundi ..................................... 21
antiquata ..................................... 31 buchi ............................................. 20
atacamensis ............................. 31 favretensis ................. 1, 4, 7, 20-22, Fig. 6.1-6.3
baliana ....................................... 31 mimer ..................................... 19, 20
hians ................. ........................ 31 ornati ........................................ 19
ladinica ......................... 1, 29, 31, Fig. 9.8 sibirica ................................... 21
praelonga ..................................... 31 Botula? cassiana ........................... 13, 14
ultim a ........................................ 32 Botulopsis .............................. 7-10, 13, 14
vallieri .................................... 31
cassiana ......................... . 12-14, Fig.
reisi .......................... 12-14, Fig. 4.8-4.10
aon, Trachyceras ................... ............... 3
Aon Zone ............ ....................... 3 taramellii .................................. 14
aonoides, Trachyceras .............................. 3 Braunjura .........................
bronni, Steinmannia ............................. 19
Aparimella .............................. 1, 16, 17, 18
archelaus, Protrachyceras ......................... 3, 5 buchi, Bositra .................................. 20
aff. archelaus ................................. 3, 5 Buchiacea ....................................... 16
cf. archelaus ............................... 3, 4, 5 Camptonectes ............................... ...38
archiaci, Plicatula ............................ 48, 49 Camptonectinae ................................ 38
arctica, Leptochondria ............................. 34 cancellata, Palaeolima ............................. 32
arcuataeformis, Gryphaea .......................... 30 Caneyella ......................... 1, 5, 16-18, Fig. 5
Arietites bucklandi Zone ........................... Carboniferous
45 ...................... 9, 14, 16, 17, 29, 32
Aso Formation .................. .... ......... 35 Cardiomorpha .................................. 14
Asoella .................................... 35-37 Cardium? saxitilis ....................... 9, 10, 14, 15
confertoradiata ................................. 35 cf carolinae, Acrochordiceras ....................... 4
laevigata .................................... 35 cassiana
speciesa ...................... 1, 35, 36, 37, Fig. 10.6 Botula? ................................. 13, 14
species b ...................... 1, 35, 36, 37, Fig. 10.7 Botulopsis ........................... 13, 14, Fig. 4.7
species c ...................... ..1, 36, 37, Fig. 10.8 Catinula ........................................ 30
Asoellidae .......... .................. 34 Ceriostella variabilis ............................. 1
Astartidae ................. ................... 8
chakii, Gryphaea ................................ 3
atacamensis, Antiquilima ........................... 31
Atrina ................. .. ........... 29, 30
Chamites succinctus ..............................
chiwanae
nigra ....................................... 29 Loxochlamys .................... 40, 44, Fig. 12.7-12.9
sinuata ............................. 1, 29, Fig. 9.5 Pecten ...................................... 40
Chlamydinae ............................... 38, 45
Aucellina ..................................... 35
Augusta Mountain Formation ...................... 1 Chlamys ....................................... 43
Aulacomya . ..................................... 19 coronata ...................................... 43
59

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Group ..................... .................... 38 falcaurita, Pleuronectites? .......................... 39

Claraia bioni ................ .................... 20 Favret Formation ..................... 1, 3-5, 21, 22, 24
Claraiidae ........................................ 20 favretensis, Bositra ............. 1, 4, 7, 20-22, Fig. 6.1-6.3
cf. columbianus, Tropiceltites ........................ 15 Filopecten ...................................... 43
confertoradiata, Eumorphotis (Asoella) ................ 35 fissistriata, Pseudoplacunopsis .................... 48-50
corallina, Loxochlamys ..... .1, 5, 10, 11, 30, 40, 43, 44, aff.
Fig. fissistriata, Pseudoplacunopsis .......... 48-50, Fig. 14
12.1-12.6 Fossil Hill Member .............................. 1, 4
cordillerana Frankites ................. ..................... 3, 5
Mysidiella .............................. 9, 10, 11, 15 sutherlandi .......................... 5, Fig. 3.1, 3.2
Promysidiella .......................... 9, 10, 11, 15 cf. sutherlandi ................................ 3, 5
coronata, Chlamys ............ .................... 43 Frechites nevadanus ................................ 4
coronatiformis, Pseudopecten ........................ 43 Gervillancea .................................. 25, 29
costata, Pinna ................................... 29 coxiella ................................. 25, 27, 29
coxiella, Gervillancea ........................ 25, 27, 29 Gervillaria (Baryvellia) ....................... 1, 24-29
Crenamussium ................................... . 39 ponderosa ......... .1, 4, 5, 7, 24-29, Figs. 8.4-8.9, 9
Crenellinae ................... .................... 13 Gervillaria (Gervillaria) ............................ 25
Cryptodonta .................. .................... 16 alaeformis..................................... 25
Cuccoceras, bonaevistae ............................ 4 hartmanni ................. ..................... 25
curtocardinalis, Leptochondria ....................... .34 Gervillia
Daonella .............. 1, 3-5, 7, 16-18, 21, 23, 24, Fig. 5 albertii .................... .................... 25
americana ................... ................... 4 joleaudi .............................. 7, 25, 27, 29
dubia ................... .................... 3, 4 giganteum, Plagiostoma ............................ 32
elegans ..................... .................... 3 Gogolin Formation ................................ 39
guembeli ................. .................... 4, 22
lamellosa ............... ................... 22-24
Gloripallium ...................................... 18
Grantsville Formation ........... 1, 4, 7, 25, 27, 34, 37, 58
cf. lom m eli ..................................... 3 grantsvillensis, Oxytoma (Oxytoma) 1, 37, 38, Fig. 10.9-10.12
cf. sturi .................... .................... 4 griesbachi, Pseudomonotis .......................... 19
Deltopectinidae .................................... 16 Gryphaea .............. ...................... 30, 31
dentatus, Pseudopecten.......................... 43, 46 arcuataeformis .................................. 30
derognati, Pleuronectites laevigatus ................... 39 chakii .................... ..................... 31
Desatoyense Zone .............................. 3, 18 Gryphaeinae ............. ..................... 30, 31
desatoyensis, Promysidiella ...... 1, 10, 11, guembeli 13, Fig. 4.5, 4.6
Devonian ................. .................... 16, 32 Daonella .................................... 4, 21
Dimyidae ................. .................... 47, 48 Enteropleura ........................... 4, 21, 23-24
Dimyoidea .................... ................... 48 gussevi, Oxytoma (Oxytoma) ........................ 38
Dipleurites ................... .................... 21 Gymnotoceras
dorsetensis, Semuridia .............................. 9 cf. rotelliformis ................... ............. 4
Dreissena ..................... .................... 8 Halobia ..................... ..1, 5, 16-18, 21, 23, Fig. 5
Dreissenidae .................. ..................... 8 plicosa .................... .................... 18
dubia, Daonella ............. .................... 3, 4 Halobiidae ................................... 16, 22
dunni, Eutomoceras ................................ 4 hartmanni, Gervillaria (Gervillaria) ................... 25
Edmondiidae ................ .................... 14 Heteroconchia ................ .................... 9
eduliformis heterodont .................................. 1, 7, 8
Mytilus .................. .................... 9, 10 Heteropecten ............... ..................... 35
Promysidiella ............................... 10, 14 hians, Antiquilima ................................ 31
eduliformis eduliformis, Mytilus ...................... 10 Hirabara Formation ............................... 40
eduliformis praecursor, Mytilus ...................... 10 hirabarensis, Pleuronectites ......................... .40
edulis, Mytilus ................ ................... 10 hisingeri, Liostrea ................................ 30
elegans, Daonella .................................. 3 Hispanic Corridor ................................ 45
England ................................ 8, 32, 46, 48 Hollandites sp................. .................... 4
Enteropleura ........... 1, 4, 5, 7, 16-18, 21-24, Figs. ..................
Hungarites 5-8 .................... 3-5
bittneri ............................... 4, 21, 23-24 inermis .............. ............ 5, 37, Fig. 3.3-3.5
guembeli .............................. 4, 21, 23-24 hyatti, Acrochordiceras ............................ 4
species a ............ 1, 4, 22-24, Figs. 6.4-6.6, 7, 8.1-8.3 Hyatti Zone .......................... 4, 18, 21, 22, 24
Entoliidae ........... ...................... 16 illyrica
Eonomia ........... .................... 48 Leptochondria................................. 35
Eoplicatula ........... ..................... 48 Pseudomonotis ................................. 35
Etalia ................................ 35 imago, Mysidiella ............................ 9, 14, 15
eumenea, Pergamidia ............................... 8 inaequistriata, Leptochondria ........................ 34
Eumorphotis .............................. 35, 38, 39 inaequistriatus, Pecten ............................. 34
multiformis .................................... 39 (Indigiropecten), Tosapecten ..................... 45, 46
Eumorphotis (Asoella) confertoradiata ................. .35 inermis, Hungarites .................. . 5, 37, Fig. 3.3-3.5
Eurydesma ................................. 8, 15 inequivalvis, Oxytoma (Oxytoma) ..................... 37
Eurydesmidae .......... .................... 15 Inoceramidae .......... .................... 19
Eutomoceras dunni................................ 4 Inoceramoidea .......... .................... 16
"Excelsior Formation" ................... 1, 4, 25, 28, 58 inornatus, Pecten .............................. 39

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Intornites ................ ..................... 4, 24 meeki

nevadanus .............................. 24, Fig. 8.2 Parafrechites ................................... 4
sp......................... .................... 4 Pleuronectites ................................... 39
Isognomonidae ................ .................... 19 Meeki Zone .................. .................... 4
Janopecten .................................... 45, 46 Meekoceras Zone ............. .................... 34
Jialingjiang Formation ............................. 39 Meekopinna ................. .................... 30
joannae, Protopis ............. ..................... 9 megalodontids ................ .................... 1
Joannina .................... .................... 8 Meginae Zone ................................ 3, 5, 44
Joannites Zone ................ .................... 3 mimer, Bositra ............... .................... 20
joleaudi, Gervillia ........................ 7, 25, 27, 29 Mine Group ................. .................... 35
kashiwaiensis, Oxytoma (Oxytoma) ................... 38 Minepharus .................. .................... 1
kerriZone ...................................... 15 minima, Leptochondria ............................. 34
Khenjan Series ............... .................... 39 Mississippian .................................... 17
klipsteini, Modiola ................................ 11 moabensis, Plicatula (Plicatula) ...................... 49
Krumbeckiella ............................... 8, 9, 15 M odiola klipsteini................................. 11
cf. timorensis ............. ........... 1, 7, 9, 13, 15 Modiolus ................... .................... 10
ladinica, Antiquilima .................... 1, 31, Fig. 9.8 Modiomorphoidea ................................. 1
laevigata, Asoella ............. .................... 35 Moenkopi Formation .............................. 34
laevigatus, Pleuronectites ................... .7, 38, 39, 40 mojsisovicsi, Oxytoma (Oxytoma) ..................... .38
laevis, Palaeolima ................................ 32 Monotoidea ............................... 16, 34, 35
lamellosa, Daonella ......................... 21, 23, 24 muensteri, Oxytoma (Oxytoma) ...................... 37
laterestriatus, Pecten (Streblopteria) .................. 39 multiformis, Eumorphotis ........................... .39
Lentilla ................... ....................... 19 Muschelkalk ............ 1, 7, 10, 21, 25, 33, 34, 38-40, 50
Leptochondria ................................ 33-35 Myalinidae ................ .................... 8, 33
aeolicus .................. .................... 33 Mysidia ................................... 8, 14, 15
albertii ................ .................... 33, 34 americana .................. .................... 10
arctica .................... ................... 34 orientalis ............... .................... 8, 14
curtocardinalis .................................. 34 taramellii ................. ..................... 14
illyrica .................... .................... 35 Mysidiella ............................. 7, 8-11, 13-16
inaequistriata ................................... 34 americana................................... 11, 15
minima .................... .................... 34 cordillerana ........................... 9, 10, 11, 15
occidaneus .................................... 34 imago ................................... 9, 14, 15
shoshonensis ................ .1, 4, 33, 34, Fig. 10.1-10.5
newtonae ............... . 1, 9, 10, 13-16, Fig. 4.11-4.13
Leptochondriidae ............. .................... 33 orientalis ............................... 6, 9, 13-15
Lewis River Formation.............................. 33
Lima
saxitilis .................... .......... 9, 10, 14,
sichuanensis .................................... 15
balatonica................. .................... 33 yunnanensis ............... .................... 15
cf. striata .............. .................... 32, 33 Mysidiellidae ................................... 8-16
Lima? poyana Mysidioptera ................. ................... 33
beds ................................. 9, 10, 14, 33 "Mysidioptera" williamsi ........................... 33
fauna .................... .................... 44 Mytilidae ............... .................... 8, 9, 13
zone ..................... ..................... 15 Mytiloconcha ...................................... 8
Lima (Radula) baliana ............................. 31 orobica ..................... ................... 9
Limidae .................... .................... 31 Mytiloida ................. .................... 8, 15
Limoida ..................... .................... 31 Mytiloidea ................... .................... 8
Limoidea ................ ................... 31 Mytilus ................. ..................... 10, 13
lineatum, Plagiostoma ............................. 33 eduliformis ................................. 9, 10
Liostrea eduliformis eduliformis ........................... 10
hisingeri ................... .................... 30 eduliformis praecursor ............................ 10
newelli ................... .................... 30 edulis .................... ..................... 11
shiraiwensis .................................... 30 otiosus .............. .................... 1, 10, 11
Liostrea? sp......................... 30, 31, Fig. 9.6, 9.7 Nathorstites
Liostreinae................................... 30, 31 fauna ........................................ 44
ef. lommeli, Daonella ............................... 3 Zone ........... ..................... 44
Longobardites nevadanus ........................... 24 Neithea ..................................... 45, 46
Loxochlamys ................................ 1, 40-44 Neithea Group ............................... 45
chiwanae ...................... 40, 44, Fig. 12.7-12.9 Neitheidae .......... ..................... 45
corallina ...... 1, 5, 10, 11, 30, 40, 43, 44, Figs. 12.1-12.6 nevadanus
sasuchan .......................... 40, 44, Fig. 12.10 Frechites .......... ......... ........... 4
lynnae, Nevadapecten .................. 1, 46, 47, Fig. 13 Intornites .............................. 24, Fig. 8.2
Maclearni Zone .............................. 3, 5, 44 Longobardites ............................... 24
Mahaffy Cliffs fauna .............................. 44 Nevadapecten ................................ 1, 46
Melleidae .......... ..................... 33 lynnae ............................ 1, 46, 47, Fig. 13
Malleus ................... ........... 33 Newaagia .......... ..................... 34, 48
Manticula ........... ..................... 8, 15 newelli
Martin Bridge Limestone ........................ 15, 33 Liostrea .......... ......... ........... 30

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Pleuronectites ............ 1,...............

Pectinula 5, 7, 30, 40,
..................... 16 44,
newpassensis, Palaeolima ................. 1, 32, Fig. 9.9 Pergamidia .................................. 8, 9
newtonae, Mysidiella ........ 1, 9, 10, 13-16, Fig. 4.11-4.13 eumenea .............. ...................... 8
nigra, Atrina ................................. 29 Pergamidiidae ..............
Peribositra
Nitanoceras selwyni ........................ ...................................
5, Fig. 3.12 20
baoqingensis ................................
obliquiradiata, Palaeolima ......................... 32 20
Peribositria ........................... 1, 16-21, 22
occidaneus, Leptochondria ........................ 3, 4
Permanomia .................................. 48
Occidentalis Zone .............................. 3, 4
Opis .................. ....................... 8 Permian ................. 8, 10, 17, 1
ornati, Bositra ................................ 19
Permian-Triassic
orientalis, Mysidiella ........................ 9,Persia .............
13-15 ............... ....................... 48
orobica, Mytiloconcha ............................ 9 Pholadomyoidea ............................... 14
ostracina, Placunopsis ........................... 48 pictetiana, Ostrea ............................ 30, 31
Ostrea pictetiana ............................ 30, 31 Pinacoceras verchojanicum Zone .................... 45
Ostreidae .............. ...................... 30
Ostreinae ................................... 31
Pinna ................. ...................... 29
costata .............. ...................... 29
Ostreoidea . o . ........ o ................... 30 nigra............... ...................... 29

vexillum .................................. 29
Ostreoidina ................... .... ........ 30, 48 Pinnidae ............... ...................... 29
Otapiria ussuriensis Zone .......................... 45Pinnoidea .............. ...................... 29
otiosa, Promysidiella ..................... 9-11, Fig. 4.4Pinnoidina
aff. otiosa, Promysidiella .............. 1, 10, 11, Fig. 4.1 .............. ..................... 29
otiosus, Mytilus ........................... 1, 10, 11 Placunopsidae ................................. 48
Placunopsis
Otoceras beds ................... ............ 19 .................................. 48
Oxytoma (Oxytoma) .......................... 37, 38
ostracina .................................. 48
plana .............. ....................... 48
grantsvillensis ........... .... 1, 37, 38, Fig. 10.9-10.12
gussevi ......... ........ ....... ............ 38
inequivalvis................................. 37
Plagiostoma ................................ 32, 33
acutum ..................... 1, 32, 33, Fig. 9.10, 9.11
kashiwaiensis ............................... 38 annonii ........................................ 33
mojsisovicsi ................................ 38 balatonica..................................... 33
muensteri .................. .................. 37 giganteum ..................................... 32
scythicum .................. .................. 38 lineatum ....................................... 33
rigidula................ ....................... 33
sedaka ...........
Oxytomidae .... ..... ... .............
................................... 3738 schimperi ...................................... 33
striatum ............... ....................... 33
palaeoheterodonts
Palaeolima ........ ...
..................... ........................
........... 32 plana,1 Placunopsis ................................ 48
planirecta, Promysidiella ........ 1, 10, 11, 13, Fig
angulata ....................................
austriaca ........................ ............
32 32
Pleuronectites ............................... 7, 38, 39
cancellata .................................. 32 balatonicus ..................................... 39
laevis ....................... ....... .......... 32 falcaurita ...................................... 39
newpassensis ........................ 1, 32, Fig. 9.9 hirabarensis .................................... 40
obliquiradiata ............................... 32 laevigatus .............................. 7, 38, 39, 40
retifera .................... ................ 32 laevigatus derognati ............................. 39
simplex ............. .. ... .... ............ 32 laevigatus zonatus .............................. 39
Paleowaagia.................................... 48 laterestriatus................................... 39
Paleozoic ..... 1, 5, 7, 16, 17, 19, 20, 30, 32, 35, 38, 40,
meeki 48
......................................... 39
Parafrechites meeki ..............................
Paratrachyceras ............................... 4
newelli 4
................. 1, 5, 7, 30, 40, 44, 47, Fig. 11
puerensis ............... ...................... 39
sutherlandi .................................. 4 pulchrus ....................................... 39
cf. sutherlandi ............................ 3-5 schmiederi ..................................... 39
Pecten vestitus ........................................ 39
acutiplicatus ................................ 45 Plicatula .................. ...................... 48
chiwanae .................................. 40 archiaci .................................... 48, 49
inaequistriatus ............................... 34
inornatus ............... .. ....Plicatulacea moabensis
.. ........... 39 .......................
........... ..................... 48 ........
rosaliae ............ ........... ............ 43 Plicatulidae ..................... ........... 47, 48
simplex ......................... ........... 32 plicosa, Halobia ............................... 18
vestitus ................... ...... ........... 39 polyodonta, Pterinea ............................ 25
Pecten (Pleuronectites) schmiederi ponderosa, Gervillaria
.................... 39 (Baryvellia) ... 1, 4, 5, 7, 24-29,
8.4-8.9, 9.1-9.4
Pecten (Streblopteria) laterestriatus ................... 39
Poseidon Zone
Pecten (Velopecten) albertii ........................ ................................. 5
34
Pectinacea .................................. 16
Posidonia ...................... 1, 3-5, 16-21,
Pectinella ............ ...................... 39 Posidoniidae ............................... 16, 19
Posidoniinae ........... ..................... 16
Pectinidae .................
Pectinoida................... ........... 16, ...............
33, 48 38, 43,..............................
Posidonioidea 45 16-18
Posidonomya ................... ........... 19, 21
Pectinoidea ............................ 38, 43, 45, 48
Pectinoidina ................................ angusta
33, 47 var. .......... ......... ........... 19

f:ffff:ffff:ffff:ffff:ffff:ffff:ffff on Thu, 01 Jan 1976 12:34:56 UTC
TRIASSIC MARINE BIVAL VIA FROM NEVADA 63
becheri .................. ...................... 19 Robustus Zone ................................ 39

stella ................. ........................ 21 rosaliae, Pecten .................................. 43
Posidonotis ............. ...................... 16, 19 R6t ......................................... 9
poyana, Lima? cf. rotelliformis, Gymnotoceras ........................ 4
beds .............................. 9, 10, 14, 33 Rotelliformis Zone ............................ 4, 24
fauna ............... ...................... 44 Saharonim Formation ......................... 39,50
zone .................. .................... 15 San Cassiano Formation ................... 11, 13, 32, 48
Praechlamys ................................ 38, 39, 43 sasuchan, Loxochlamys ................ 40, 44, Fig. 12.10
weiyuanensis ................................... 39 saxitilis
praecursor, Mytilus eduliformnis Cardium?
...................... 10
............................. 9, 10, 14, 15
praelonga, Antiquilima ............................. 31 Mysidiella ............................. 9, 10, 14, 15
Prida Formation ............................. 1, 3, 4 Schaumkalkbank............................... 39
Promysidiella ................................. 1, 7-15 schimperi, Plagiostoma ............................ 33
americana ................................... 9, 10 scleractinian ................................. 1, 5
cordillerana ............................. 10, 11, 15 schmiederi, Pecten (Pleuronectites) ................... 39
desatoyensis ................ 1, 10, 11, 13, Fig. 4.5, 4.6
scythicum
eduliformis ................................. 10, 14 Oxytoma (Oxytoma) ............................. 38
otiosa ............................... 9-11, Fig. 4.4 Towapteria ..................................... 38
aff. otiosa ............ ............ 1, 10, 11, Fig. 4.1 sedaka, Oxytoma (Oxytoma) ......................... 38
planirecta .................. 1, 10, 11, 13, Fig. 4.2, 4.3 selwyni, Nitanoceras........................ 5, Fig. 3.12
Promytilus .................................... 9, 10 Semuridia ................. ..................... 8, 9
borealis ....................................... 10 dorsetensis ...................................... 9
Prospondylidae ..............................
Prospondyloidina ............................ 1, 47-48
47, 48
shiraiwensis, Liostrea .............................. 30
shoshonensis, Leptochondria ..... .1, 4, 33, 34
Prospondylus ................................. 47, 48 Shoshonensis Zone ..................... 4, 18, 21, 23, 24
protobranchs ................................. 1, 7 sibirica, Bositra .................................. 21
Protopis ................. ...................... 8, 9 sichuanensis, Mysidiella ............................ 15
joannae ........................................ 9 sikanianum, Protrachyceras .......................... 5
timorensis ...................................... 9 aff. sikanianum, Protrachyceras ............ 5, Fig. 3.6-3.
triptycha ................. ....................... 9 Silberlingia ................. ...................... 19
Protrachyceras ................................ 4, 5, 7 simplex
archelaus .............. ...................... 3, 5 Palaeolima ..................................... 32
aff. archelaus ................................. 3, 5 Pecten ................. ...................... 32
cf. archelaus ............................... 3, 4, 5 sinuata, Atrina ......................... 1, 29, Fig. 9.5
omkutchanicum Zone ........................... 45
"Solenopora".....................................
sikanianum ..................................... 5
5
species, Liostrea? ................... 30, 31, Fig. 9.6, 9.7
aff. sikanianum ....................... 5, Fig. 3.6-3.10 species a, Asoella? ................ 1, 35, 36, 37, Fig. 10.6
sp . ........... .......................... Fig. 3.11 species a, Enteropleura .. 1, 4, 22-24, Figs. 6.4-6.6, 7, 8.1-8.3
Pseudomonotidae ................................. 47
species b, Asoella? ................ 1, 35, 36, 37, Fig. 10.7
Pseudomonotis ................................. 21, 48 species c, Asoella? ................... 1, 36, 37, Fig. 10.8
griesbachi ..................................... 19 Spondylus ....................................... 48
illyrica ....................................... 35 Star Peak Group ................................ 3
(Pseudomonotis) ................................. 48 Steinmannia ...................................... 19
(Trematiconcha) ................................. 48 bronni................... ..................... 19
Pseudomonotoidea .............................. 33 stella, Posidonomya ............................... 21
Pseudopecten ............................... 43, 46, 47 Streblopteria .................................. 38, 39
coronatiformis ................................. 43 cf. striata, Lima ............................... 32, 33
dentatus ...................... .... ...... 43, 46 Plagiostoma .............................. 33
striatum,
Pseudoplacunopsis .............................. 48-50
cf. sturi, Daonella ................................. 4
affixa ..................................... 48-50 subpolaris, Tosapecten ............................. 45
fissistriata .................................. 48-50 succinctus, Chamites ............................. . 31
aff. fissistriata ......................... 48-50, Fig. 14 Sunrise Formation ............................... 45
Pteriidae .. ................................... 33 sutherlandi, Frankites .................... 5, Fig. 3.1, 3.2
Pterinea polyodonta ............................... 25 cf. sutherlandi, Frankites .......................... 3, 5
Pterineidae ........... ..................... 16 Sutherlandi Zone ........................ 3, 4, 33, 44
Pterinopectinidae ................... ........... 16 taramellii
Pterioida ............................... 16, 30, 48 Botulopsis ................................... 14
Pterioidina ........... ..................... 16 Mysidia ........... .......... ............ 14
puerensis, Pleuronectites ........................... 39 Terebratelbank ..................... ........... 39
pulchrus, Pleuronectites ............................ 39
Quenstedtoceras mariae Zone ...................... 19 Terquemiidae ..............................
Rahnbauerkogel fauna ........................ 23, 24 Thaynes Formation .........................
Red Rock limestone .............................. Tiefengraben
timorensis, 3 Protopisfauna ..........................
................................ 9
reisi, Botulopsis .................... . 12-14, Fig. 4.8-4.10 cf. timorensis
retifera, Palaeolima ............................... 32 Krumbeckiella ........................ 1, 7, 9, 13, 15
rigidula, Plagiostoma ..............................33 Tommasina .................................... 8, 9

182.66.141.128 on Fri, 15 Apr 2022 11:35:20 UTC
Tosapecten ................. ................... 45, 46 Pecten ................... .................... 39

(Indigiropecten) ............................. .45, 46 Pleuronectites .................................. 39
(Tosapecten) ................................ 45, 46 vexillum, Pinna .................................. 29
Tosapectininae ................................ 44-47 Virgin Limestone .............................. 34, 39
Towapteria scythicum .............................. 38 weiyuanensis, Praechlamys .......................... 39
W ettersteinkalk ................................... 13
Trachyceras
aon ....................... ..................... 3 W eyla ....................................... 45, 46
alata ......................................... 45
aonoides ........................................ 3
(Trematiconcha), Pseudomonotis ..................... .48
bodenbenderi ...................................45
G roup ........................................ 45
trigonioids ....................................... 1 Wildhorse mining district ...................... 1, 24, 58
triptycha, Protopis ................................. 9 williamsi
Tropiceltites cf. columbianus ........................ 15 Aviculomyalina ................................. .33
ultima, Antiquilima ................................ 32 "Mysidioptera" .............. ................... 33
vallieri, Antiquilima ............................... . 31 Yinkeng Formation ................................ 20
variabilis, Ceriostella ............................... 1 yunnanensis, Mysidiella ............................ 15
vestitus zonatus, Pleuronectites laevigatus .................... 39

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Middle Triassic Pteriomorphian Bivalvia (Mollusca) from the New Pass Range, West-

Central Nevada: Systematics, Biostratigraphy, Paleoecology, and Paleobiogeography

Stable URL: https://www.jstor.org/stable/4095819

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MIDDLE TRIASSIC PTERIOMORPHIAN BIVALVIA (MOLLUSCA) FROM THE

Shoshone Mountains near Berlin, "Excelsior Formation" in the

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Desatoya Mountains, Churchill County,

TABLE 1-Stratigraphic distribution of bivalves from South Canyo

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Tertiary base of Unit E. He reported no age-diagnostic fossils from the

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containing colonial corals and the Hungarites

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FIGURE 3-Late Ladinian ammonoids present in the ammonoi

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ponderosa n. subgen. and sp., interpreted

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Waterhouse (2000, 2001) used

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included in the Pergamidiidaemost by Cox

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perhaps facilitated by theBased

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is difficult to observe, but it layer

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-:~:_ ??~:u i:? a~:: "

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(1926, p. 124) based his Genusnew

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recrystallized specimens. Mysidiellaby an outer lacks shell layer."

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have mainly Hemisphere Southern

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as well as a single adductor

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Family POSIDONIIDAE Frech,Other 1909 [emended]

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this point, however, posidoniids

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examined by Kittl. For this reason, Krumbeck

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and having a posterior margin that is outwardly

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~C::.:L' .':'~tl; 41;

?IN' 34, e`:?

FIGURE 8-1-3, Enteropleura species a: 1, interio

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W-C'A. . . . . . . 11" ??l r

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FIGURE 9-1-4, Gervillaria (Baryvellia) ponder

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ATRINA SINUATA newof

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1864a or Meekopinna Yancey,

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