IS THE CENOZOIC AN “ALL YOU

CAN EAT SEAFOOD BUFFET”?

by

Ms. Emily Feldman

A Thesis Submitted in Partial Fulfillment of Requirements for

the Degree of Bachelor of Science Department of Geological
Sciences, California State University, Fullerton

Approved, Thesis Committee

Dr. Nicole Bonuso

Assistant Professor of Geology
Thesis Advisor

August 2008

*Undergraduate Thesis Contribution Number 1 to Cal-State

Fullerton Evolutionary Paleoecology Laboratory

1
Abstract:

Researchers argue that throughout the Phanerozoic, primary productivity in the

seas, and therefore, biomass, has increased. Archaic mode shell beds tend to be thin, and
usually contain Paleozoic fauna dominated by brachiopods, while modern mode beds
usually are thicker and tend to contain Modern fauna such as bivalves and gastropods. As
bivalves tend be fleshier and more massive than brachiopods, it is inferred that abundance
and biomass have also increased through time. This thesis examines Late Triassic shell
beds from central Nevada to determine whether the mode of the studied shell beds is
archaic or modern by examining the geometry and the taxonomy of the deposits. I found
the shell beds to be composed of chiefly Schwagerispira brachiopods in a complex
micrite matrix, implying the beds are archaic in mode. The results indicate that 1) the
archaic mode continued into the Late Triassic and 2) in general biomass has not increased
in the Late Triassic compared to Paleozoic marine shell beds.

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 TABLE OF CONTENTS
Hypothesis 6

Introduction 6

Background and Study Site 7

Methods 8

Rock preparation 8
Taxonomic composition 8
Biofabric 9
Geometry 9
Biostratinomic classification 9
Internal structure 10

Results 10

Taxonomic composition 10
Biofabric 11
Geometry 11
Biostratinomic classification 12
Internal structure 12
Comparative anatomy 12

Discussion 13

Conclusion 14

References 15

Figures 18

Tables 28

Graphs 30

LIST OF FIGURES

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Figure 1 – Diversity curve 18

Figure 2 – Study location map 10

Figure 3 – Site stratigraphy 20

Figure 4 – Brachiopod morphology 21

Figure 5 – Schwagerispira specimen cross-sectional photographs 22

Figure 6 – Schwagerispira specimen photographs on bedding 23

surface

Figure 7 – Schwagerispira specimen photographs from Bittner, 24

1890

Figure 8 – Paleogeographic map of study site during Late Triassic 25

Figure 9 – Partial paleogeographic world map of study site 26

Figure 10 – Paleogeographic world map of study site 27

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 LIST OF TABLES

Table 1 – Measurement data of Schwagerispira specimens 28

Table 2 – statistical analysis of measurement data 29

LIST OF GRAPHS

Graph 1 – Plot of Schwagerispira shell length vs. thickness 30

Graph 2 – Sorting of Schwagerispira shell sizes 31

I. Hypothesis

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 I plan to test the hypothesis that the 1) Late Triassic shell beds sampled at the
South Canyon locality are archaic, as defined by (Kidwell, 1990); and 2) that the shell
beds are event deposits (Kidwell, 1991). The purpose of this research is to determine
whether or not biomass has increased through time.

II. Introduction

Researchers argue that increased primary production through out the Phanerozoic
has lead to an increase in biomass through time (Bambach, 1993). Bambach’s (1993)
classic paper on seafood through time bases this argument on numerous observations
within the Phanerozoic fossil record. The argument discusses the increase in diversity,
ecosystem complexity, and utilization of ecospace during the Phanerozoic. At present,
researchers indicate that diversity increased, as well as the dominant structure of faunas,
through time (Sepkoski, 1976, 1981, 1991; Sepkoski et al., 1981; Sepkoski and Miller,
1985) (Fig. 1). Three major evolutionary faunas emerged from the compilation of
diversity data, each possessing distinct faunal characteristic (Sepkoski et al., 1981).
During the Early Cambrian and until the Ordovician, trilobites maintained dominance
among the “Cambrian fauna” through their explosive abundance and diversification.
Beginning in the Ordovician, Rynchonellid brachiopods, crinoids, ostracods, and
cephalopods dominated benthic communities thus defining the “Paleozoic fauna”. After
the end Permian mass extinction, the “Modern fauna” comprising of such organisms as
gastropods, bivalves, bony fish, and echinoids rose to dominance.
In conjunction with the changes in faunal diversification explained above,
research indicates that major expansion of ecosystem space occurred as well as an
increase in fleshiness (Bambach, 1983; Bambach, 1985). Research suggests that each
evolutionary fauna was possibly displaced by its successor – with the appearance of each
new dominant fauna coinciding with the decline of the preceding fauna (Sepkoski, 1981).
The less fleshy, primarily epifaunal, trilobites gave way to the rise of epifaunal
brachiopods, which transitioned into fleshier, infaunal bivalves dominating the seas
(Bambach, 1993). As the Modern fauna progressed, pelagic modes of life increased
along with increased predation in the seas. This radiation of predators with new shell-
crushing, -destroying and -boring adaptations in the marine realm during the Mesozoic
accompanied, and possibly was reason for, the increased shell strength in marine
gastropods (Vermeij, 1977). Vermeijj (1977) coined this time interval as the Mesozoic
marine revolution. Bambach cites evidence of increases in the biomass of marine
consumers and in energetics during the Phanerozoic. He claims this evidence of change
in the food supplies in marine realms can be seen all the way down the food chain to
primary producers (Bambach, 1993). Bambach also reports an increase in utilized marine
ecospace during the Phanerozoic, in accordance with the diversification that added new
species with new modes of life to the marine realm (Bambach, 1993). Researchers argue
that this expansion of ecological guilds implies that an increase in total resource supply
occurred throughout the Phanerozoic thus equating to an increase in biomass though time
(Bambach, 1993).
Although researchers predict an increase in biomass through time, biomass has
not been quantified throughout the Phanerozoic in a rigorous manner because it is often

6
difficult to obtain precise measurements. One possible solution to this problem is
examining the abundance and degree of fleshiness through time. This thesis aims to
quantify abundance by comparing the thicknesses and frequencies of fossil
concentrations from the Late Triassic of Nevada with known fossil concentration from
the Paleozoic and Cenozoic. Then, once I quantify abundance, comparative anatomy
will be used to determine the fleshiness of the individuals comprising the fossil
concentrations. If the data indicate abundance and fleshiness are constant through time
then we could conclude that biomass does not increase through time. If the data reveals
that abundance is consistent but fleshiness increases through time then we could conclude
that biomass does increase through time. Lastly, if both abundance and fleshiness
increase through time then it is safe to assume biomass increased through time.
Kidwell (1990) defines two different modes of shell bed concentrations - archaic
mode and modern mode – using physical dimension, taxonomic composition and
taphonomic attributes. The archaic mode, common in the Paleozoic and the Triassic, is
characterized by thin concentrations of predominantly brachiopods. The modern mode,
found preferentially in Cretaceous and Cenozoic deposits, is characterized by thin
pavements and thicker, bioclastic concentrations, that are dominated by mollusks.
The archaic mode was dominated by less fleshy brachiopods than the modern
mode deposits which were dominated by mollusks, especially bivalves, with much
fleshier bodies. Thus, along with the transition from the archaic mode to the modern
mode among marine life, it is thought that a shift in biomass would have also occurred
simultaneously (Bambach, 1993; Bottjer, 1996). I used Kidwell’s classification scheme
to characterize the shell beds, thus determining if abundance has changed through time.
Once the shell beds were characterized and the dominant organisms were identified,
biomass was able to be determined based on modern anatomical studies.

III. Background and Study Site

The South Canyon locality is of particular interest to biostratigraphers because it

contains one of the most complete ammonoid successions across the Ladinian-Carnian
boundary in North America (Waller and Stanley, 2005). Accordingly, it has been
extensively studied for the ammonoids and the bivalves it contains. In researching
Middle Triassic fossils all over North America, Smith identified ammonoids, bivalves,
and brachiopods at the South Canyon locality (Smith, 1914). Some have published
research consisting solely of data on the ammonoids contained there (Balini and Jenks,
2007; Johnston, 1941; Silberling, 1956). Silbering and Wallace explored the stratigraphy
of the entire Star Peak Group and interpreted the paleogeography of the area (Silberling
and Wallace, 1969). Waller and Stanley examined the bivalves from South Canyon in
detail (Waller and Stanley, 2005). Balini (2007) described the Ladinian-Carnian
boundary at South Canyon in detail by extensively sampling bed-by-bed for ammonoids,
bivalves and conodonts (Balini et al., 2007).
The shell beds sampled in this study are from the Upper Triassic (Carnian) at
South Canyon, located in the New Pass Range in present-day central Nevada, near
Austin, NV (39º36’42”N, 117º29’31”W) (Silberling, 1956; Waller and Stanley, 2005)
(Fig. 2).
The section of the Star Peak Group exposed at South Canyon consists of the

7
Augusta Mountain Formation (Silberling, 1956), underlain by the Koipato Group
(conglomerates and siltstones containing volcanic and tuffaceous rocks) and the Favret
Formation (shelf limestones), and overlain by Tertiary volcanics, and ranges from late
Early Triassic (Anisian and Ladinian) to Late Triassic (Carnian) (Balini et al., 2007;
Nichols and Silberling, 1977; Waller and Stanley, 2005). The samples used in this study
were collected from the lower member of the Augusta Mountain Formation (Fig. 3). The
lower member of the Augusta Mountain Formation consists of a massive gray limestones
interspersed with small beds of siltstones or conglomerates that grades into a massive
gray limestones that are often bioclastic and sometimes oolitic. The uppermost section of
the member contains the brachiopod shell beds occurring in coquina-like layers (Balini et
al., 2007; Waller and Stanley, 2005).

IV. Methods

Rock Preparation:
I examined three rock samples from two beds from the Late Triassic of samples at
the South Canyon locality for their biotic signature and their environmental context. One
sample was collected from each bed in the field, the two beds separated by no more than
one vertical meter. The shell beds were described in field and in the laboratory using
methods previously published in Kidwell et al. (1986), Kidwell (1991) and Kidwell and
Holland (1991). A total of three thin sections were prepared, one from each rock sample
collected from the study site. The faces of the cut slabs were also polished to gain a more
clear view of the internal fabric of these beds and the fossils they contain.
To acquire a more detailed look at the internal structure of the shell beds, acetate
peels were be made of the polished cross-sectional, using instructions by Mark A. Wilson
and Thomas J. Palmer (Wilson and Palmer, 1989). The polished cut surfaces were etched
for 20 seconds each in a 2% hydrochloric acid solution, and left to air dry untouched.
After completely dry, the etched surface was flooded with 100% acetone, and then the
acetate peel plastic sheet placed quickly onto the surface, flattening out all air bubbles,
before the acetone dried. After being left to dry 24 hours, the acetate peel sheets were
peeled off, and paper-thin pictures of the internal structure of the rocks were left behind
on the clear plastic sheets.

Taxonomic Composition:
To determine the taxonomic composition of the beds, I examined the bedding
plane surfaces, the polished faces, the thin sections and the acetate peels of my rock
samples. Fossils were identified to genus level. If the shell beds contained only one
genus, they were categorized as monotypic; if the shell beds contained multiple genera, I
categorized them as polytypic (Fagerstrom, 1964; Kidwell et al., 1986). In the process of
classifying the shell beds, I was able to determine the paleoecology and the taphonomy of
the deposits (Kidwell, 1991). I determined the number of species, the relative abundance
of each species and the life habits of those species through this research.

Biofabric:
To determine the biofabric, the three-dimensional arrangement of skeletal

8
elements in the matrix, I examined the orientation, packing and sorting of the samples. To
determine orientation, I observed and measured the angles that the long axes of the
hardparts make to the bedding plane. If close to 0º, the orientation was called
concordant, if ~45º, it was known as oblique, and perpendicular orientation if the hard
parts were found to be aligned about 90º (perpendicular) to the bedding plane (Kidwell et
al., 1986). The terms imbrication, edgewise, stacking, nesting, and telescoping can also
be used to describe some additional particularly characteristic styles of orientation
observable in cross section (Kidwell et al., 1986).
To determine packing, I used both quantitative and qualitative techniques to
ensure complete analysis. I performed point-counts in the thin sections to determine the
hardparts by volume (cross-sectional area). Point counts were based on a 1” x 2” (thin
section sized) quadrant. I also visually determined close-packing: If the hardparts
appeared to “float” in the matrix, the sample was called matrix-supported, and it was
called bioclast-supported if the hardparts appeared to be physically touching in the matrix
(Kidwell et al., 1986).
Furthermore, using the polished surfaces, I measured the shells (length-wise)
under magnification to determine the size sorting of the shell accumulations. If the
central 80% of the samples fell within one phi size column, then the deposit would have
been considered well-sorted, that is, essentially unimodal; if two phi size models, then
essentially bimodal (Kidwell and Holland, 1991). The lengths and thicknesses of the
shells were measured, via the standard convention (See Fig. 4), using a computer
equipped with a microscope and with SPOT Advanced software.

Geometry:
The geometry of a fossil concentration is based on many factors, including the
depositional surface, the mode of life of the fossilized organisms, the activities of other
organisms and the physical processes the deposit underwent (Kidwell et al., 1986).
Extremely thin shell concentrations are called pavements, or stringers if they’re laterally
discontinuous. The geometry of a fossil concentration can also be classified as a clump,
pod, lens, wedge, or bed (Kidwell et al., 1986). I determined the geometry of the samples
using these guidelines.
I also characterized the mineralogy of the beds employing similar laboratory
techniques. The matrix, the fossils and their mineralogy were identifiable in thin section
using a petrographic microscope.

Biostratinomic Classification:
After gathering all of the information about the shell beds I was able to determine
their biostratinomic classification. A biogenic concentration is the result of biologic
processes alone, while a sedimentologic concentration is produced by the physical
(usually hydraulic) reworking of the hardparts, and a diagenetic concentration is created
or enhanced significantly by physical or chemical process after burial (which includes
compaction). Concentrations can also be produced from a combination of two or more of
these processes (Kidwell et al., 1986).

Internal Structure:
The internal structure of a skeletal concentration can be called simple (internally

9
homogenous, exhibiting no trend, or just one unidirectional trend) or complex (having
more complicated patterns of variability, such as multiple horizons and multiple and/or
bi-directional variability patterns) (Kidwell et al., 1986). Also according to Kidwell,
internally simple concentrations tend to be deposited during a single event, and complex
concentrations tend to form from a multiple-event background (Kidwell et al., 1986). I
was also able to classify the taphonomic signature as event, composite, or hiatal, as
defined by (Kidwell, 1991).
Kidwell described two main modes (styles) of marine deposits throughout the
Phanerozoic – archaic mode and modern mode. The archaic mode is typical of Paleozoic
and Triassic strata, and is characterized by thin (several 10’s of centimeters or less)
bioclastic deposits dominated primarily by brachiopods and other epifaunal and semi-
infaunal groups. The modern mode, typical of Cretaceous and Cenozoic strata, is
characterized by thin pavements and thick (1 meter or greater) bioclastic deposits
dominated by mollusks and other epifaunal and fully infaunal groups. Modern mode
concentrations exhibit a greater diversity of habitats, shell mineralogies, and sizes and
thicknesses than archaic mode concentrations. Kidwell says that throughout the Jurassic,
brachiopod concentrations tend towards the archaic mode of deposition, typically being
small in scale and taphonomically simple. Kidwell’s data suggests the Jurassic record is
biostratinomically mixed – that is, typified by concentrations of both archaic and modern
modes. While the data shows bivalve concentrations from the Jurassic to be modern
mode, it also shows brachiopod concentrations of the same period to be archaic in mode
(Kidwell, 1990). I was able to biostratinomically classify the mode of the Late Triassic
shell beds of the South Canyon locality using Kidwell’s descriptions of shell bed
depositional styles.

V. Results

Taxonomic Composition:
A challenging task in analyzing these shell beds was determining the taxonomic
composition. Under magnifying microscope and in thin section, two different genera of
brachiopods could be identified. One of these genera was only represented by one large
specimen, and due to the nature of the rock, could not be identified from only one view. I
have identified the second genera of brachiopod as Schwagerispira (Dagys, 1972), also
called Retzia schwageri (Bittner, 1890).
The specimens, numerous and visible on the weathered top bedding surface of the
shell bed and in cross section on the polished surfaces, are small, almond-shaped,
biconvex brachiopods. They appear to have a very subdued fold and sulcus, thus
appearing almost spherical, and have a slightly curved (hooked) “beak” in cross section
(See Fig 5). They also have a relatively small number of wide, angular costae, which
radiate from the umbo (see Fig. 6).
I have classified these brachiopods as Schwagerispira because they are described
as small (~5mm), having angular costae, being subequally biconvex, having subcircular
shells, and are found in rocks of the Middle Triassic (Anisian) to Upper Triassic
(Carnian) from Slovakia, Hungary, Bulgaria, Romania, Primorya, and many parts of
China (See Fig. 7) (Boardman and Nestell, 2000).

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 Only one specimen of an unidentified brachiopod genus was found, and at least
50 of the genus Schwagerispera (See Table 1, measurement data for 50 specimens). This
makes the shell beds taphonomically uniform. There was a wide range in the sizes of
Schwagerispira specimens (Standard deviation in the lengths of the shells of
1.829860751, See Table 2), and therefore a large range of inferred ages and stages of life
in the population of that one genus. Brachiopods with rounded, biconvex shells tend to be
epifaunal, being raised above the sediment by their more 3-dimensional shells. Because
of the brachiopods’ mode of life as filter feeders and the morphology of their shells, these
organisms must have been epifaunal, living on top of the substrate on the sea floor.

Biofabric:
The biofabric of the beds was classified by looking at the orientation of the long
axes of the shells and the packing of the hardparts within the matrix. The orientations of
the shells within the bed were a mix of concordant, oblique and perpendicular, with
angles to the bedding plane ranging from approximately 0º to 90º, thus it cannot be
described by any one of the terms; The orientations were a combination of all three.
I also characterized the biofabric of the beds by quantitatively and qualitatively
evaluating the packing of the bioclasts. In one 2” x 1” thin section through rock sample
090707-1, 1 fossil is seen in the matrix. In a second 2” x 1” thin section through a
different area of the same rock sample 090707-1, 7 fossil specimens can be seen. In
another 2” x 1” thin section through rock sample 090707-2 from the other shell bed, 8
fossil specimens can be seen in the matrix. Examination of the internal structure of the
beds on the polished cuts revealed that no bioclasts appear to be touching, thus the shell
beds must be matrix-supported.
Sorting of the deposit was determined after measuring the length and thickness of
50 specimens in cross section (See Table 1). The lengths ranged from 2.13 mm to 11.02
mm, and width ranged from 0.74 mm to 6.84 mm. The mean length was 5.8 mm, and the
mean thickness of the shells was 3.5 mm. The median length was 6.0 mm, and the
median thickness was 3.5 mm (See Table 2). The mean and median thicknesses/lengths
being the same or almost the same indicates that there are not many outliers in the data.
A clear linear relationship (R2 value = 0.8164) is seen when plotting the lengths of the
Schwagerispira specimens against their thicknesses (See Graph 1). The size-sorting of
the deposit was characterized as bimodal, as 74% of the data (37/50 data points) fell into
one phi size column, and 94% of the data (47/50 data points) fell into two phi size
columns. These beds could not be classified as unimodal, as less than 80% of the
measurement data fell into one phi size column (See Graph 2).

Geometry:
I have determined these shell beds should indeed be called beds because of their
thickness of only about 8-10 cm. Furthermore, the beds themselves are not internally
homogenous and contain thin stringers of shell material that pinch out laterally and
pavements.
Under a petrographic microscope I determined the matrix of the deposits was
composed primarily of micrite, mud-sized crystals of calcite that appear as tiny brown
specks on lower magnification and on higher magnification show high order
birefringence colors. The fossils brachiopod shells must have originally been composed

11
of low Mg calcite, since dissolved and the void spaces left behind filled in with sparry
calcite, large coarse-grained calcite crystals.

Biostratinomic Classification:
I found the biostratinomic classification of the concentrations to likely be a mix of
biogenic, sedimentologic and diagenetic processes. This was due to the complex nature
of the beds, containing both skeletal hardparts and grains of lime mud, indicating
biogenic origins, the replacement of the shells by calcite after deposition, indicating
diagenetic processes, and the development of hummocky and discontinuous bedding,
which would indicate hydraulic reworking.

Internal Structure:

The structure and fabric of the beds show the deposit must be complex in nature,
because the beds display no real organization, grading, or unidirectional trend.
“Bedding” within the concentrations is incomplete, messy, sub-parallel at best, and
contains random broken fragments of shells and thin, intensely brick-red mud layers.
Further, I categorized this deposit as an event deposit based on the lack of any
visible repeated pattern or laminations within the beds, and a general lack of basic
structure. Taxonomy and preservation are uniform throughout the bed, and the size
variance in the specimens was well-mixed.
Furthermore, I have characterized the shell beds as archaic, because their
contained biota is relatively uniform, they are not thick beds, and they contain only
brachiopods, which are all attributes of typical archaic mode deposits.

Comparative Anatomy:

Evidence shows that articulate brachiopods have much lower energy life habits
than those of suspension-feeding bivalves (Rhodes and Thayer, 1991). A hypothesis
called escalation states that competing taxonomic groups force each other to adapt
resulting in an increases specializations and energy flux through communities (Vermeij,
1987). Brachiopods, with their low energy lifestyles, were driven into habitats with
lower nutrient supplies, by comparatively high energy bivalves (Vermeij, 1987). In one
study, articulate brachiopods and bivalves with body masses of 50 mg, 100 mg, and 350
mg were selected to compare their clearance rates. Clearance rate refers the rate a filter
feeder can move water through its body, which is an indication of how much food they
are consuming/filtering out of that water. In this study, there was only one articulate
brachiopod out of five that reached 350 mg, and almost every (eleven of twelve) bivalve
specimen reached 350 mg (Rhodes and Thompson, 1993). In addition, there was no data
for bivalve specimens weighing 50 mg for half of the bivalve specimens, indicating that
bivalves tend to eat more than brachiopods (Rhodes and Thompson, 1993). It takes more
energy for fleshier organisms with more muscle mass to function, so it would follow that
fleshier bivalves require more energy (calories) to survive than less fleshy brachiopods,
and need to consume more at a faster rate.
In addition to requiring more calories than brachiopods, bivalves would also need
to take in a larger oxygen supply to support the number of cells in their bodies. James et

12
al., (1992) reports a correlation between both polar and temperate 50 mg (dry mass)
bivalve mollusks and 50 mg (dry mass) brachiopods. Oxygen consumption rates
averaged about 10 to 20 micro-liters/hour higher for the bivalve mollusks than the
brachiopods (James et al., 1992). Based on these data, along with the decline of
brachiopod diversity and expansion of bivalve genera through the Phanerozoic, and the
increase in the relative abundance of bivalves in relation to brachiopods through time, we
can assume the hypothesis that biomass has increased through time is correct (Bambach,
1993).

VI. Discussion

With this research, I have concluded that these beds are archaic type deposits, and
that they are event deposits based on all the evidence.
These 8-10 cm thick shell beds have been described as coquina-like by other
studies, however, after investigating the internal fabric of the beds I believe these
findings are incorrect. I found that these shell beds should be described as matrix-
supported, and classified as a wackestone or a biomicrite, rather than a coquina, which is
a soft, rather unconsolidated, poorly-cemented biogenic form of limestone composed of
carbonate fragments. This evidence suggests they are archaic style, as archaic mode
deposits are typically characterized as being thin (generally <10 cm) and bioclastic
(Kidwell, 1990). The depositional environment of this location in central Nevada would
have been a back-arc basin on the continental shelf off of the west coast of Laurasia,
during the Late Triassic. This area developed behind the volcanic arc caused by the
convergent boundary (See Fig. 8). This geographical setting is consistent with the
micrite (lime mud) grains working as the supporting matrix of the shell bed samples.
The lack of any visible pattern in shell orientation signifies chaotic and non-
uniform deposition, which would indicate the shells are not fossilized and preserved in
their life positions. Within the shell beds, the packing of the fossils was equally as non-
uniform, with number of specimens varying within the thin sections from one to eight
specimens. The bi-modal size-sorting of the beds, which contain a wide range of shell
lengths and thicknesses that align along a linear relationship, indicate a rapidly-deposited,
moderately well-mixed concentration. This indicates heterogeneity in the sorting of these
deposits, also affirming the hypothesis that these beds are event deposits.
The significance of identifying specimens of the genus Schwagerispira in these
beds is great. Foremost, finding only two genera of brachiopods affirms the simple
taxonomy of the beds. Less diverse fossil concentrations, dominated by brachiopods, are
typical of the archaic mode of deposition (Kidwell, 1990). Also, those areas where
Schwagerispira specimens were previously found to have lived during the Carnian
include locations on the east coast of Laurasia, on the Tethys Ocean (See Fig. 9). The
Tethys Ocean was connected to the western coast of Laurasia, where today’s Nevada was
then located, via the Pacific Ocean. This could possibly provide a route for the genus to
migrate and inhabit both coasts of Pangaea, connected by a very large ocean (See Fig.
10).
Finding deposits displaying the archaic mode from this time period indicates
biomass is still lower in the Late Triassic than today. At this time of transition, during

13
and after the recovery period from the P-T extinction, the switch from archaic to modern
modes had not yet occurred, meaning the shift in biomass must have occurred later than
the Late Triassic.

Conclusion:

It is the conclusion of this paper that the hypotheses are supported. It is

confirmed that the Late Triassic shell beds sampled at the South Canyon locality are
archaic, as defined by (Kidwell, 1990), and that the shell beds are event deposits
(Kidwell, 1991). As the archaic mode of deposition transitioned to the modern mode,
leaner brachiopods gave way to more fleshy bivalves, and biomass in the seas increased.
However, this transition had not yet occurred when the shell beds from this study were
deposited, due to their archaic mode character.

VII. References

Balini, M., and Jenks, J.F., 2007, The Trachyceratidae from South Canyon (Central

14
 Nevada): Record, taxonomic problems and stratigraphic significance for the
definition of the Ladinian-Carnian boundary: New Mexico Museum of Natural
History and Science Bulletin, v. 41, p. 14-22.
Balini, M., Jenks, J.F., McRoberts, C.A., and Orchard, M.J., 2007, The Ladinian-Carnian
Boundary Succession at South Canyon (New Pass Range, Central Nevada):
Bulletin - New Mexico Museum of Natural History and Science, v. 40, p. 127-
138.
Bambach, K., 1983, Ecospace Utilization and Guilds in Marine Communities Through
the Phanerozoic, in Tevesz, M.J.S., and McCall, P.L., eds., Biotic interactions in
Recent and fossil benthic communities, p. 719-746.
Bambach, R.K., 1985, Classes and adaptive variety; the ecology of diversification in
marine faunas through the Phanerozoic, in Valentine, J.W., ed., Sixty-third annual
meeting of the Pacific Division, American Association for the Advancement of
Science: Phanerozoic diversity patterns: Princeton, NJ, Princeton Univ. Press.
—, 1993, Seafood Through Time: Changes in Biomass, Energetics, and Productivity in
the Marine Ecosystem: Paleobiology, v. 19, p. 372-397.
Bittner, A., 1890, Brachiopoden der Alpinen Trias.: Abhandlungen der kaiserlich-
koniglichen geologischen Reichsanstalt v. 14, p. 1-325.
Boardman, D.R., II, and Nestell, M.K., 2000, Outcrop-based sequence stratigraphy of the
Council Grove Group of the Midcontinent, in Johnson, K.S., ed., Platform
carbonates in the southern Midcontinent, 1996 symposium Circular, Volume 101,
Oklahoma Geological Survey, p. 275-306.
Boardman, R.S., Cheetham, A.H., and Rowell, A.J., 1987, Fossil Invertebrates, Blackwell
Science, 728 p.
Bottjer, D.J., 1996, Comparative evolutionary palaeoecology; assessing the changing
ecology of the past, Biotic recovery from mass extinction events, Volume 102,
Geological Society Special Publications, IGCP Project No. 335, p. 1-13.
Dagys, A.S., 1972, Morfologiia i Systematika Mezozoiskikh Retsiodynkh Brakhiopod:
Morfologicheskie i Filogeneticheskie Voprosy Paleontologii, v. 112, p. 94-105.
Fagerstrom, J.A., 1964, Fossil Communities in Paleoecology; Their Recognition and
Significance: Geological Society of America Bulletin, v. 75, p. 1197-1216.
James, M.A., Ansell, A.D., Collins, M.J., Curry, G.B., Peck, L.S., and Rhodes, M.C.,
1992, Recent advances in the study of living brachiopods: Advances in Marine
Biology Review, v. 28, p. 175-387.
Johnston, F.N., 1941, Trias at New Pass, Nevada (New lower Karnic ammonoids):
Journal of Paleontology, v. 15, p. 447-491.
Kidwell, S., M., 1990, Phanerozoic Evolution of Macroinvertebrate Shell Accumulations:
Preliminary Data from the Jurassic of Britain, in Miller, W., III, ed.,
Paleocommunity temporal dynamics: the long-term development of multispecies
assemblages., Volume 5, The Paleontological Society Special Publication, p. 309-
327.
Kidwell, S.M., 1991, The Stratigraphy of Shell Concentrations, in Stehli, F.G., Briggs,
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Fossil Record, Volume 9: Topics in Geobiology.
Kidwell, S.M., Fursich, F.T., and Aigner, H., 1986, Conceptual Framework for the
Analysis and Classification of Fossil Concentrations: Palaios, v. 1, p. 228-238.

15
Kidwell, S.M., and Holland, S.M., 1991, Field Description of Coarse Bioclastic Fabrics:
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VIII. Figures

17
Figure 1: Diversity curve illustrating the number of families through time, from the
Cambrian to the Tertiary (From Raup and Sepkoski, 1982)

18
Figure 2: Basic map of Nevada, USA, showing the location of the thesis study area at
South Canyon

19
Figure 3: Generalized stratigraphic section of the strata at the South Canyon
locality. Samples studied in this research were taken from beds labeled
"brachiopod beds with echinoids" in green.

20
Figure 4: Basic brachiopod morphology, adapted from Boardman et al. (1987)

21
Figure 5: Photos taken of cross-sections of Schwagerispira specimens showing
almond shape in lateral view and slightly curved beak (red arrow). Measured and
photographed specimens were labeled A to Z and then AA through ZZ – above are
specimens M and PP.

22
Figure 6: Photos of several Schwagerispira specimens from the shell beds at South
Canyon, illustrating their wide, angular costae.

23
Figure 7: *Schwagerispira schwageri specimens (2x magnification) from Bittner
1890, in (Williams et al., 2002)

24
Figure 8. Map of the western United States (then, west coast of Laurasia) during the
Late Triassic, indicating the South Canyon study area with a yellow star (Image by
Dr. Ron Blakey, http://jan.ucc.nau.edu/~rcb7/)

25
Figure 9. Partial paleogeographic world map (during Late Triassic) with locations
where Schwagerispira specimens have been found today mapped as yellow dots, with
location of South Canyon study area where the specimen was also found marked
with a yellow star (from http://paleodb.org)

26
Figure 10. Paleogeographic world map (during Late Triassic) with locations where
Schwagerispira specimens have been found today mapped as yellow dots, with
location of South Canyon study area where the specimen was also found marked
with a yellow star. Note that the Tethys Ocean and the Pathalassa are connected via
the South Pole (from http://paleodb.org).

27
IX. Tables

Length Thickness BB 6.22 3.53

Specimens (mm) (mm) Length Thickness
A 8.13 4.88 Specimens (mm) (mm)
B 7.15 3.63 CC 3.89 1.58
C 4.97 3.31 DD 4.86 2.7
D 7.23 3.67 EE 6.53 3.5
E 5.09 3.51 FF 7.92 4.62
F 6.87 4.82 GG 4.33 1.36
G 6.2 4.98 HH 3.68 3.23
H 6.29 3.34 II 6.57 3.11
I 5.5 3.47 JJ 5.91 4.2
J 7.95 5.07 KK 6.08 4.38
K 6.93 3.66 LL 6.06 3.15
L 10.65 6.63 MM 3.67 2.44
M 6.52 3.68 NN 6.17 4
N 6.02 3.52 OO 5.52 3.43
O 3.43 1.73 PP 11.02 6.84
P 6.07 3.79 QQ 6.39 4.49
Q 5.68 3.34 RR 7.51 3.89
R 4.56 2.74 SS 7.24 5.12
S 2.55 1.43 TT 4.11 1.9
T 2.94 1.35 UU 5.58 2.17
U 3.52 1.67 VV 4.01 2.76
V 7.86 6.55 WW 3.36 1.7
W 7.4 5.15 XX 3.87 2.07
X 2.13 0.74
Y 5.77 3.95
Z 5.88 3.82 Table 1: Measurement data of fifty
AA 6.27 3.42
Schwagerispira specimens.

28
 Lengths Thicknesses

Mean 5.8012 Mean 3.4804

Median 6.04 Median 3.505
Mode #N/A Mode 3.34
Standard Standard
Deviation 1.829860751 Deviation 1.373973769
Range 8.89 Range 6.1
Minimum 2.13 Minimum 0.74
Maximum 11.02 Maximum 6.84
Count 50 Count 50

Table 2: Statistical analysis of length and thickness data on Schwagerispira

specimens.

X. Graphs

29
 Specimen Lengths vs. Thicknesses

8 y = 0.6784x - 0.4554

7 R2 = 0.8164

3
Thickness (mm)

0
0 2 4 6 8 10 12
Length (mm)

Graph 1: Plot of Schwagerispira specimens’ lengths versus their thicknesses,

illustrating the clear linear trend of data, and the equation and R2 value of the
trendline.

30
Graph 2: Graph showing sorting of Schwagerispira shells by organization into Φ
(phi) size columns, where -1Φ is 2 mm, -2Φ is 4 mm, -3Φ is 8 mm, and -4Φ is 16
mm.

31