OCCASIONAL PAPERS

OF THE

California Academy of Sciences

No. 46, 27 pages, 2 figures, 2 plates.

July 1, 1964

PLEISTOCENE DIATOMS

FROM

MONO AND PANAMINT LAKE BASINS,

CALIFORNIA

By
Walter W. Wornardt, Jr.

Marine Bioiogica! Laboratory

LIBRARY

JUL 1 319B4 •

WOODS HOLE, MASS.

SAN FRANCISCO

PUBLISHED BY THE ACADEMY
1964

OCCASIONAL PAPERS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

No. 46, 27 pages, 2 figures, 2 plates. July 1, 1964

PLEISTOCENE DIATOMS
FROM

MONO AND PANAMINT LAKE BASINS,

CALIFORNIA *

By

Walter W. Wornardt, Jr.

Introduction

The purpose of this investigation is to study and describe two diatom
florules from ancient lake beds in California, each representing a different
type of environment, and to compare the similarities and differences between
them. It should be noted that the author has not visited either locality.

Beds of Mono Lake Basin

A diatomaceous clay was collected at California Academy of Sciences,
Department of Geology, Locality Number 37113, on October 20, 1960, from an
outcrop on the shore of Mono Lake (fig. 1) by G D. Hanna and C. W. Chesterman.
The sampled locality is an exposed bank approximately 25 feet in height and
about 100 feet above the surface of the lake. The beds are composed of a
light gray, silty and ashy clay, horizontally bedded in thin layers up to two
inches thick. The location is in Sec. 6, TIN, R28E, M.D.B. & M., as shown
on the U. S. Geological Survey topographic map, Mt. Morrison quadrangle (30

* The results reported herein were made possible through National Science Foundation Grant
No. 2958-C1, N.S.F. 11083-

CALIFORNIA ACADEMY OF SCIENCES

(Occ. Papers

minute series, 1914. Reprinted, 1950). The locality is just below an old aban-
doned cabin beside a seldom traveled dirt road.

Much has been written on the geology of the Mono Lake area (Russell,
1885) but no mention of fossil diatoms has been found in the literature except
as pertains to Paohoa Island, which is located in the lake. There are exten-
sive deposits of diatomaceous sediments on this island but the diatoms have
not been studied in detail. A brief examination of them by G D. Hanna (person-
al communication) suggests that their age is much older than the material con-
sidered in this paper, perhaps as old as Pliocene.

The water in Mono Lake is saline at present and supports a very limited
fauna and flora. A minute green alga was noted in one place but no living dia-
toms were found in a brief examination by Hanna (personal communication).
The larvae of one or more species of flies were abundant and the brine shrimp,
Artemia, has been reported by members of the Steinhart Aquarium of the Cali-
fornia Academy of Sciences.

, .

Figure 1. A portion of U. S. Geological Survey Topographic Map, Mt. Morrison
Quadrangle, June, 1914 edition, Mono County, California, showing California Academy
of Sciences locality no. 37133.

Lake Beds of Panamint Valley

A diatomaceous ash was collected at California Academy of Sciences,
Department of Geology, Locality Number 37055, on December 26, 1960, by F.

No. 46)

WORNARDT: PLEISTOCENE LAKE DIATOMS

Weidenbenner, G D. Hanna, and M. M. Hanna. The elevation of the outcrop is
1840 feet, or 800 feet above the present floor of Panamint Valley. These beds
are located in Sec. 20, T21S, R43E, M.D.B. & M., U. S. Geological Survey top-
ographic map, Maturango Peak quadrangle (15 minute series), Inyo County,
California, 1951, (fig. 2).

Figure 2. A portion of U. S. Geological Survey Topographic Map, Maturango
Peak, California, Quadrangle, edition of 1951, showing California Academy of Sciences
fossil collection locality no. 37055 in Panamint Lake basin.

Panamint Valley formed one of the many sumps in the Pleistocene chain
of lakes that probably started with Lake Lahontan to the north. It received
overflow drainage directly from Searles Lake basin to the west (Blanc and
Cleveland, 1961, p. 5, map) and there is a possibility that it in turn overflowed
into Death Valley. A complete and satisfactory drainage pattern has not been
worked out because of changes in topography since Pleistocene time and lack
of fossil evidence. Whatever the drainage pattern may have been, there can be
no doubt that a very large and relatively pure freshwater lake occupied Pana-
mint Valley during late Pleistocene time. The Pleistocene sediments which
accumulated are largely covered with detrital outwash so that outcrops are
extremely limited in the vicinity of the collecting station of this report. The
total thickness of these sediments cannot be measured. Associated with the
diatom-bearing strata are some soft, sandy beds containing a rather extensive
fauna of well preserved freshwater mollusks. Hanna (1963, p. 5) stated that

4 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

the molluscan species there are almost exactly the same as those found in
the basin of SearlesLake. He therefore considered the fossils in the two lakes
to be contemporaneous, and believed that they furnish evidence of a former
connection between the two bodies of water. No diatoms were found in the
Searles Lake basin. Scholl (1960) has published a general account of events
pertaining to the geology of Searles Lake and an extensive bibliography of
the area. A formal name for the lake beds in Panamint Valley has not been
proposed.

Methods Used for the Identification of Species

The species described in this report were identified by comparison with
the reference slides of G D. Hanna and H. E. Sovereign. Many identifications
were made from published plates in the extensive diatom library at the Cali-
fornia Academy of Sciences. The magnifications given for the diatom species
on plates 1 and 2 have been given in round figures for the purpose of utility,
and maybe in error by a few per cent. Inasmuch as exact measurements of the
illustrated diatoms are given, the precise magnifications may be obtained if
so desired.

Acknowledgments

The writer is extremely grateful to Dr. G Dallas Hanna of the California
Academy of Sciences, for suggesting this investigation, for critically reading
the manuscript, and for aid and advice throughout this project. The writer is
also indebted to Dr. Hanna for his instruction in preparing, mounting, and pho-
tographing these minute organisms. His type diatom slides, many strewn slides,
and unpublished manuscripts were also made available.

Acknowledgements are likewise due to many others: Dr. Leo G. Hert-
lein, California Academy of Sciences, for his excellent advice on numerous
editorial and systematic problems; Dr. R. M. Kleinpell of the Department of
Paleontology, University of California, Berkeley, for his valuable advice on
general systematics; the late Ignatius M'Guire, Librarian, California Academy
of Sciences, for his assistance in locating numerous obscure publications,
many of which were cited under inadequate references; Dr. George F. Pappen-
fuss, and Dr. Paul C. Silva, Department of Botany of the University of Cali-
fornia, Berkeley, for their valuable discussions on plant taxonomy and classi-
fication; Dr. Taro Kanaya and Dr. Reimer Simonsen for their advice and dis-
cussions concerning diatoms. Finally, the writer would like to record his in-
debtedness to the California Academy of Sciences for the facilities provided,
especially the use of the vast diatom library. Additional references were ob-
tained from the Library of the University of California, Berkeley.

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS 5

Class BACILLARIOPHYCEAE Fritsch

Order B ACILL ARIALES Schiitt

Suborder Discineae Schiitt

Family Coscinodiscaceae Kiitzing

Subfamily Melosiroideae Kiitzing

Genus Melosira (Ehrenberg) Kiitzing, 1844

Melosira undulata (Ehrenberg) Kiitzing.
(Plate 1, figure 1.)

Under high magnification, the valve surface is characterized by radiat-
ing lines of punctae that disappear toward the center. Under low magnifica-
tion, the valve surface has a somewhat wavy appearance. This species is
found in short cylindrical tubes. The valves range from 0.0590 to 0.650 mm.
in diameter. The size ranges of the valves are taken from different individuals
seen in the Panamint Valley samples.

Tempere and Peragallo (1915, p. 59) reported Melosira undulata from
Tacoma, Washington, living. Sovereign (1958, pp. 105-106) reported it from
Crater Lake and Emerald Pool, Oregon, and stated that Hustedt has said: "..
M. undulata as a recent species is known only from the tropics but that it oc-
curs as a fossil in Tertiary deposits in the whole of Europe. This species
now has been found as recent in eight locations in the Pacific Northwest in
waters varying from pH 7.1 to 8.8."

This species was not only abundant in Europe during theCenozoic, but
it was also very abundant in North America through the same interval of time.
It has been reported from the followingdeposits of many different ages. Ehren-
berg (1870, table 12, opposite p. 68), Truckee River, Nevada; Fall River and
Columbia River, Oregon. Tempere and Peragallo (1915), Reno, Nevada. Mann
(1926, pp. 51-52), Mica, near Spokane, Washington, Miocene, Latah formation,
probably a shallow inland freshwater lake with increasing amounts of mineral
constituents. Lohman (1934, p. 10), Tipton, Oregon, Miocene, Mascall forma-
tion. Lohman (1936, p. 98), 49 Camp, Nevada, Miocene, Upper Cedarville for-
mation. Lohman (1937), Kettleman Hills, Kings County, California, Tulare
formation, fresh to brackish water. Sovereign (1958, pp. 105-106), stated that:
"It also occurs as a fossil in Upper Miocene deposits of central Washington
in approximate stratigraphic position with fossil trees of the Ginkgo State Park
near Vantage, Washington. These tree fossils indicate a temperate zone for-
est and it is thought that no considerable climatic change occurred up to late
Pliocene time when the Cascade Range was elevated, producing arid but still
temperate conditions in central Washington. Melosira undulata has a long his-
tory as a temperate zone species in this area."

6 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

Subfamily Coscinodiscoideae Schu'tt

Genus Cyclotella O) Kiitzing, 1833

Cyclotella compta (Ehrenberg).
(Plate 1, figures 6, 9.)

Discoplea comta EHRENBERG, 1844. Beitrage zur Kenntniss des kleinsten Lebens im

agaischen Meere am Euphrat und auf den Bermuda-Inseln. Monatsber. Berlin Akad.

Wissen., p. 267. "Kurdistan."
Cyclotella comta (Ehrenberg), Kctzing, 1849- Spec. Algarum, p. 20. VanHeurck, 1882.

Syn. Diat. Belgique, pi. 92, figs. 16-23. De Toni, 1894. Syl. Algarum, p. 1353-

Fricke in Schmidt, 1900. Atlas Diat., pi. 224, figs. 1-5, 13-25. Elmore, 1921.

Diat. Nebraska, p. 40, pi. 1, fig. 18. BOYER, 1926. Syn. N. American Diat., p. 40.

Hustedt, [1928] . Kieselagen, p. 354, fig. 183, 127-159; 1930, Baeillariophyta,

p. 103, fig. 69.
Cyclotella comta var. radiosa Grunow in VAN HEURCK, 1892. Syn. Diat. Belgique, pi.

92, fig. 23; pi. 93, figs. 1-9.
Cyclotella comta var. affinis GRUNOW in Van Heurck, 1892. Syn. Diat. Belgique, pi.

93,, figs. 11-13, 21.

Valves with regular radiating costae which become a series of radiating
beads as they approach the center of the valve. This species is found as very
short cylinders, the diameter of which ranges from 0.0590 to 0.0650 mm.

Cyclotella comta is one of the most common species in the Panamint
Valley deposit. It has been reported living in British Columbia, Wolle (1890,
pi. 110); Bailey (1921, p. 6), Little and Big Quill Lakes, Saskatchewan, Can-
ada, brackish water.

In the fossil record, it has been reported by Patrick (1936, p. 160), from
Great Salt Lake, Utah. Lohman (1937, p. 48), Sprague River, Klamath County.
Van Heurck (1892, pi. 93, figs. 11-13), Carson City, Nevada. Tempere and
Peragallo (1915, p. 227), Orcas Island, Puget Sound, Washington.

Genus Stephanodiscus Ehrenberg, 1845
Stephanodiscus niagarae Ehrenberg.

(Plate 1, figure 2.)

Stephanodiscus niagarae Ehrenberg, 1845- Neue Unterschungen iiber das kleinste
Leben als geologisches Moment, p. 80. Niagara Falls, Ontario, Canada. EHREN-
BERG, 1854- Mikrogeologie, pi. 35A, group 7, figs. 21, 22- Van Heurck, 1882.
Syn. Diat. Belgique, pi. 95, figs. 13, 14- DE TONI, 1894. Syl. Algarum, p. 1 152-
Wolle, 1890. Diat. N. America, pi. 66, figs. 28, 29. FRICKE in Schmidt, 1901.

(1) This genus originally appeared as a subgenus in Kiitzing (1833b, p. 535). Most
workers refer to the reprint (1834, p. 7) of the original article and not to the original itself.

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS

Atlas Diat., pi. 227, figs. 1-9- ELMORE, 1921. Diat. Nebraska, p. 41, pi. 1, figs.
24,25. BOYER, 1926. Syn. N. American Diat., pp. 60-61. HANNA, 1933- Diat. Flor-
ida Peat Dep., p. 90, pi. 2, fig- 3- Lohman, 1937- Diat. East. Oregon, pi. 4,
figs. A-2, A-5, C-2. Okuno, 1952. Atlas Fos. Diat. Japan, p. 34, pi. 3, fig. 1;
pi. 4, fig. 6; pi. 7, figs. 1,2; 1956a, Electron-Micro. Fos. Diat., p. 133, pi. 21,
figs, la, lb; 1956b, Diat. Earth, p. 345, pi. 1, figs. 1-3- Sovereign, 1958. Diat.
Crater Lake, Oregon, p. 106.

The valves are usually elevated at the center, with a depressed area
between that and the margin. Radiating rows of costae are distinct as seen
under high magnification. The marginal zone is characterized by fine beads
which intersect at various angles. Spines are present on the outer margin of
the various valves. The valves of this diatom from the Panamint Valley de-
posit range from 0.0600 to 0.0870 mm. in diameter.

Sovereign (1958, p. 106) reported the species in Diamond Lake and Up-
per Klamath Lake, Oregon, and stated that it is fairly common in the larger
lakes in the Pacific northwest and may be found in streams and ditches with
slowly moving water, usually having a pH of 6-2 to 9-5.

This species has been found as a fossil at the following localities:
By Lohman (1937, p. 22), Klamath Lake, Oregon, Holocene; (p. 26), Davis
Lake district, Deschutes County, Oregon, Holocene; (pp. 21, 31), Terrebonne,
Deschutes County, Oregon, Pleistocene; and (p. 42-43), Klamath Falls, Klam-
ath County, Oregon, Pliocene?.

Suborder Araphidineae Smith, H. L,
Family Fragilariaceae Kutzing
Subfamily Fragilarioideae Schutt

Genus Synedra Ehrenberg, 1830

Synedra species.

(Plate 2, figure 7.)

Characters used for specific identification were not found on any speci-
mens of this genus from the Mono Lake assemblage. All forms were badly
corroded, probably owing to the leaching activity of alkaline solutions after
deposition, or the contact of the frustules with alkaline waters while the dia-
toms were living. This corrosion is very common in fossil diatoms. This form
ranges from 0.1905 to 0.2150 mm. in length and from 0.0150 to 0.0195 mm. in
breadth.

8 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

Suborder Monoraphidineae Ostrup
Family Achnanthaceae Kiitzing
Subfamily Cocconeioideae Kiitzing

Genus Cocconeis Ehrenberg, 1833

Cocconeis placentula Ehrenberg.

(Plate 2, figures 8, 11.)

Cocconeis placentula Ehrenberg, 1838. Infusionsthierchen, p. 194- Van Heurck,
1880. Syn. Diat. Belgique, p. 133, pi. 30, figs. 26, 27. DeToni, 1891- Syl. Al-
garum, p. 454. SCHMIDT, 1894. Atlas Diat., pi. 192, figs. 38-44, 46, 47, 49, 50.
BOYER, 1927. Syn. N. American Diat., p. 244. HUSTEDT, 1930. Bacillariophyta,
p. 189, fig- 260; 1933, Kieselalgen, p. 347, fig. 802, a. b. Lohman, 1938. Plio-
cene Diat. Kettleman Hills, California, p. 83.

Cocconeis placentula var. lineata (Ehrenberg), CLEVE, 1895- Naviculoid Diat., p. 169.
Meister, 1912. Kieselalgen Schweiz, p. 94, pi. 12, fig. 6, 7. Hustedt, 1930,

Bacillariophyta, p. 190, fig. 262; 1933, Kieselalgen, p. 348, fig. 802 d.

Cocconeis euglypta Ehrenberg, 1854. Mikrogeologie, pi. 34, gr. 6 A, fig. 2-

Cocconeis lineata var. euglypta, Van Heurck, 1880. Syn. Diat. Belgique, pi. 30, figs.
33, 34.

Cocconeis placentula var. euglypta, CLEVE, 1895. Naviculoid Diat., p. 170. HUSTEDT,
1930. Bacillariophyta, p. 190, fig. 261; 1933, Kieselalgen, p. 349, fig. 802 c.

Cocconeis pellucida var. lineata Grunow, PERAGALLO, 1897. Diat. France, pi. 3, figs.
22, 23.

Cocconeis bonnieri HERIBAUD AND PERAGALLO, 1903. Diat. Fos. d'Auvergne, pi. 11,
Figs. 24, 25.

These forms from Panamint Valley deposit are characterized by a dis-
tinct linear, somewhat lanceolate medial line. Under low magnification, the
valve surfaces appear to have many wavy lines, but properly resolved, fine
puncta appear in their place (as shown in the illustrations on plate 2). These
forms range from 0.0201 to 0.0360 mm. in length and from 0.0135 to 0.0270 mm.
in breadth.

Many diatomists may be inclined to call the illustrated form on plate 2,
figure 11, Cocconeis placentula var. lineata or C. lineata, while other diatom-
ists may refer the illustration on plate 2, figure 8, to Cocconeis placentula
var. euglypta. It is doubtful, however, that these "varieties" really have tax-
onomic and therefore phylogenetic significance.

Cocconeis placentula has been reported from many freshwater fossil de-
posits. Ehrenberg (1870, table 12 opposite p. 68), reported it from Fall River
and Columbia River, Oregon and from the Great Salt Lake area in Utah. Pat-
rick (1936, p. 160) also reported the species from the Great Salt Lake area and
stated that Cocconeis placentula is a euryhaline species. Boyer (1926b, table
opposite p. 26, column E) recorded the species from deposits from Pukaist
Creek, Kamloops District, and from Vancouver Island, British Columbia. Tern-

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS 9

pere and Peragallo (1915) reported it from many western North American de-
posits such as Orcas Island, Puget Sound, Washington (p. 179); "Washington
County" (p. 311); Pit River, California (originally reported as "Pitt River,
Oregon" (p. 312). Lohman (1937) reported the species from a deposit at Klam-
ath Lake, Oregon (p. 22); from Burns, Harney County, Oregon (p. 34), and Ter-
rebonne, Deschutes County, Oregon (p. 31). Hasler and Crawford (1938) p. 25)
reported a marl deposit south of Sevier Lake, Millard County, Utah, Pleisto-
cene. Lohman (1935, p. 456) reported it from a Quaternary deposit at Clovis,
New Mexico, and stated that the species is now "...living sparingly in fresh-
water lakes, but more frequently in saline lakes and brackish estuaries." Han-
na and Grant (1929, p. 91) reported the species from the Etchegoin formation,
Kettleman Hills, Fresno County, California.

Suborder Biraphidineae Hustedt

Family Naviculaceae Kiitzing

Subfamily N aviculoideae Schiitt

Genus Navicula Bory, 1822

Navicula oblonga (Kiitzing).

(Plate 2, figure 5.)

Frustulia oblonga KUTZING, 1834. Syn. Diatomearum, p. 20, pi. 2, fig- 24.

Navicula oblonga KUTZING, 1844- Bacillarien, p. 97, pi. 4, fig. 21- Schmidt, 1876.
Atlas Diat., pi. 47, figs. 63-68. Van Heurck, 1880. Syn. Diat. Belgique, p. 81,
pi. 7, fig. 1. DETONI, 1891- Syl. Algarum, p. 37. Cleve, 1895- Naviculoid Diat.,
p. 21. BOYER, 1916. Diat. Philadelphia, p. 97, pi. 27, fig. 21; 1927, Syn. N. Amer-
ican Diat., p. 395. Hustedt, 1930. Bacillariophyta, p. 307, fig. 550.

P innul aria oblonga VI M. Smith, 1853- British Diat., p. 55, pi. 18, fig. 165.

This diagnostic species is characterized by terminal convergent striae
that are angularly bent. There are about seven striae in 0.01 mm. This form
ranges from 0.0800 to 0.1100 mm. in length and from 0.0100 to 0.0150 mm. in
breadth.

A very striking but not very abundant species found in the Panamint Val-
ley beds is Navicula oblonga. Tempere and Peragallo (1915, p. 180) reported
it in Big Lake, Arlington, Washington, living. Bailey (1921, p. 7) found it living
in brackish water at Little and Big Quill lakes, Saskatchewan, and (p. 8) at
Airdrie, Alberta. Sovereign (1958, p. 20) gave the location of Emerald Pool,
Crater Lake, Oregon and stated that this seems to be a common species in
the Pacific northwest, especially in more alkaline waters with a pH of 7.0 to 9.0.

In the fossil record, Navicula oblonga has been reported by Patrick
(1936, p. 163) from Great Salt Lake, Utah, in fresh to brackish water. Ehren-

10 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

PLATE 1

Figure 1. Melosira undulata (Ehrenberg) Kiitzing. Hypotype no. 3460- (Calif.
Acad. Sci., Dept. Geol. Type Coll.), from locality 37055 (CAS), Panamint Valley, Cal-i
ifornia, Pleistocene. Diameter 0.0640 mm. x 735.

Figure 2. Stephanodiscus niagarae Ehrenberg. Hypotype no. 3461. (Calif . Acad.
Sci., Dept. Geol. Type Coll.) from locality 37055 (CAS), Panamint Valley, California,
Pleistocene. Diameter 0.0870 mm. x 400.

Figure 3- Campy lodiscus noricus var. hibemica (Ehrenberg) Grunow. Hypotype
no. 3535 (Calif. Acad. Sci., Dept. Geol. Type Coll.), from locality 37055 (CAS), Pan-
amint Valley, California, Pleistocene. Diameter 0.0715 mm. x 615.

Figure 4. Surirella ovata var. utahensis Grunow. Hypotype no. 3536. (Calif.
Acad. Sci., Dept. Geol. Type Coll.), from locality 37113 (CAS), Mono Lake, California,
Pleistocene. Length 0.0903, width 0.0920 mm. x 450.

Figure 5. Surirella ovalis Brebisson. Hypotype no. 3537. (Calif. Acad. Sci.,
Dept. Geol. Type Coll.), from locality 37113 (CAS), Mono Lake, California, Pleisto-
cene. Length 0.0870, width 0.0590 mm. x 550.

Figure 6. Cyclotella compta (Ehrenberg) Kiitzing. Locality 37055 (CAS), Pana-
mint Valley, California, Pleistocene. Diameter 0.0190 mm. x 2250.

Figure 7. Surirella striatula var. testudo Ehrenberg. Hypotype no. 3538. (Calif.
Acad. Sci., Dept. Geol. Type Coll.), from locality 37113 (CAS), Mono Lake, California,
Pleistocene. Same specimen as figure 8 enlarged to show details of the valve surface,
x 2000.

Figure 8. Surirella striatula var. testudo Ehrenberg. Hypotype no. 3538- (Calif.
Acad. Sci., Dept. Geol. Type Coll.), from locality 37113 (CAS), Mono Lake, California,
Pleistocene. Length 0.1400, width 0.1060 mm. x 400.

Figure 9. Cyclotella compta (Ehrenberg) Kiitzing. Hypotype no. 3448. (Calif.
Acad. Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Valley, Cal-
ifornia, Pleistocene. Diameter, 0.0480 mm. xlOOO.

OCC. PAPERS CALIF. ACAD. SCI., NO. 46 (WORNARDT) PLATE 1

12 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

berg (1870, table 12 opposite p. 68) reported the species from a Great Salt
Lake deposit and also from "Fall River, Oregon." Boyer (1926b, table oppo-
site p. 26, column E) listed Pukaist Creek, Kamloops District and Deadman
River, north of Kamloops Lake, British Columbia. Hasler and Crawford (1938,
p. 26) reported it from a marl south of Sevier Lake, Millard County, Utah, Plei-
stocene. In Mexico, Ehrenberg, (1869, table opposite p. 60) reported it from
Barranca Honda [Freshwater fossiT] .

Genus Pinnularia Ehrenberg, 1840
Pinnularia viridis (Nitzsch),Ehrenberg.

(Plate 2, figure 4.)

Bacillaria viridis NlTZSCH, 1817. Beitrag Infusorienkunde, p. 97, pi. 6, figs. 1-3.

Pinnularia viridis (Nitzsch), EHRENBERG, 1841 [T843] • Verbreitung und Einfluss des
mikroskopischen Lebens in Sud- und Nord-Amerika, pi. 1, gr. 1, figs. 4, 7; gr. 3,
fig. 3; Pi- 2, gr. 1, fig. 22 [see Mills, 1934, p. 1309]- Cleve, 1895- Naviculoid
Diat., p. 91. DeTONI, 1891- Syl. Algarum, p. 11. Boyer, 1927- Syn. N. American
Diat., p. 446. Hustedt, 1930. Bacillariophyta, p. 334, fig. 617a; 1949- Susswas-
ser-Diat. Albert Park, p. 109, pi. 6, fig. 21. Patrick, 1936. Diat. Siam and Ma-
lay States, p. 427.

Navicula viridis (Nitzsch), Ehrenberg, 1876. Infusionsthierchen, p. 182, pi. 13, fig.
16; pi. 21, fig- 12- Kutzing, 1844. Bacillarien, p. 97, pi. 30, fig. 12- Schmidt,
1876. Atlas Diat., pi. 42, figs. 11-14-

It is important to call attention to the complex undulating raphe, and the
terminal comma-shaped fissure so common in this species. There are about
six striae in 0.01 mm. This form ranges from 0.2200 to 0.2300 mm. in length
and from 0.0330 to 0.0395 mm. in breadth and is therefore somewhat larger
than those recorded by Cleve (1895, p. 91) and Hustedt (1930, p. 334). It dif-
fers from Pinnularia flexuosa Cleve (1895, p. 93), which has 4.4 to 4.5 striae
in 0.01 mm. This species is not common in the Panamint Valley deposit.

Although P innularia viridis is very commonly found in freshwater lakes
in western North America, it was rare in the "Panamint Valley" samples. It
has been reported living from the following localities: Ehrenberg (1841, p. 36),
Kotzebue Sound, Alaska; Cleve (1895, p. 91), Santa Rosa Mountains and Sier-
ra Nevada; Bailey (1921, p. 8), Airdrie, Alberta, brackish water; Tempere and
Peragallo (1915, p. 59), Tacoma, Washington; (p. 166, Orcas Island, Puget
Sound, Washington; and (p. '80), Big Lake, Arlington, Washington.

P innularia viridis is one of the more common species in the western
North America Cenozoic diatomaceous formations. It has been reported from
the following localities: Patrick (1936, pp. 160, 162), Great Salt Lake, Utah,
freshwater; Boyer (1926b, table opposite p. 26, column E), north end of Pro-
spect Lake, Lake District, and Quamichan Lake, Vancouver Island, British

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS 13

Columbia; six miles south of Chilliwack, and Pukaist Creek, Kamloops Dis-
trict, British Columbia; Ehrenberg (1870, table 12 opposite p. 68), Salt Lake,
Utah; Humboldt Valley, Nevada; Truckee River, Nevada; Fall River and Co-
lumbia River, Oregon; Tempere and Peragallo (1915, p. 228), Orcas Island and
Puget Sound, Washington; (p. 280), Lost Spring Ranch, California; (p. 310),
Carson City, Nevada; (p. 312), Pit River, California; and (p. 328), Reno, Nev-
ada. Ehrenberg (1869, p. 11), Nahe von Regla, 6114'; (p. 12), Istalahuaca,
7740'; and (p. 18), Guadalupe Hidalgo, Mexico. Lohman (1935, p. 456), Clovis,
New Mexico, Quaternary. Boyer (1926b, p. 9), Blackwater River, Cariboo and
Coast Districts, British Columbia, Tertiary. Lohman (1937, p. 84), Kettleman
Hills, Kings County, California, Pliocene, Tulare formation. Mann (1926, p. 53),
Mica, near Spokane, Washington, Miocene, Latah formation.

Genus Anomoeoneis Pfitzer, 1871
Anomoeoneis costata (Kiitzing), Hustedt.

(Plate 2, figures 1, 2.)

Navicula costata KUTZING, 1844. Bacillarien, p. 93, pi. 3, fig. 56-

Anomoeoneis costata (Kiitzing), HUSTEDT, 1959- Kiesselalgen, p. 744, fig. 1111.

Anomoeoneis polygramma Ehrenberg, CLEVE, 1895. Naviculoid Diat., p. 6. BOYER, 1927.
Syn. N. American Diat., p. 324.

Anqmoeoneis polygramma var. pannonica, FRENGUELLI, 1934. Diat. Plioceno Superior,
p. 355, pi. 2, fig. 9.

Anomoeoneis sculp ta (Ehrenberg), HANNA and GRANT, 1931- Diat. Pyramid Lake, Nev-
ada, p. 285, pi. 25, figs. 7, 8.

Anomoeoneis spbaeropbora (Ehrenberg), HANNA, 1932. Pliocene Diat. Kansas, p. 373 ,
pi. 31, fig. 3.

Anomoeoneis sphaerophora var. polygramma (Ehrenberg), O. Muller, HUSTEDT, 1930,
Bacillariophyta, p. 262, fig. 425.

Navicula bobemica Ehrenberg, 1854. Mikrogeologie, pi. 10, gr. 1, fig. 4a, b. Schmidt,
1877. Atlas Diat., pi. 49, figs. 43-45- Peragallo, 1897. Diat. France, p. 62,

pl. 8, fig. 1.

Navicula perdurrans PANTOCSEK, 1902. Balaton Floraja, p. 58, pl. 5, fig- 120.
Navicula rostrata Ehrenberg, Elmore, 1921. Diat. Nebraska, p. 86, pl. 11, figs.

392.

390-

This robust species, abundant in the Mono Lake assemblage, is char-
acterized by obtuse apices. The broad axial area is bordered by a row of punc-
ta that more or less converge from the transverse central area toward the api-
ces. There are 13-16 transverse striae in 0.01 mm. Size ranges from 0.1060
to 0.2070 mm. in length and from 0.0295 to 0.0560 mm. in breadth. An outline
of history of this "Anomoeoneis sphaerophora- Anomoeoneis costata " complex,
can be found in Hanna (1932, p. 373-375), and Hustedt (1959, pp. 746-747).

14 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

This is the most abundant and characteristic species in either of the
two deposits. It was originally described from Saxony, "Im Bergmehl von St.
Fiore," by Kutzing (1844, p. 93). Tempere and Peragallo (1915, p. 180) re-
ported the species living from several localities: Big Lake, Arlington, Wash-
ington Originally recorded as "Big Lake, Arlington - Washington Co.'J], and
(p. 165) Orcas Island, Puget Sound, Washington [originally as "Climacum Val-
ley - Orcas Islands '3; (p. 193) [^'Swan Lake, Klamath City [County], Oregon '3-
Boyer (1927 p. 324) reported it from the Gulf of California, and considered it
to be characteristic of brackish water.

This species has been reported from the following fossil deposits: Ehr-
enberg(1870, table 12 opposite page 68, pi. 1, gr. 1, fig. 2; pi. 2, gr. 1, fig. 34);
Tempere and Peragallo (1915, p. 328), Reno Nevada; Lohman (1935, p. 456),
Clovis, New Mexico. He stated that individuals of this species, "...are living
sparingly in freshwater lakes, but more frequently in saline lakes and brack-
ish estuaries." Lohman (1937, p. 34), "freshwater pond deposits, Burns, Har-
ney County, Oregon, Pleistocene." According to Hanna (1932, p. 376), "At the
present time it \A. costata\ is a definite indicator of more or less saline waters."

Subfamily Pleurosigmoideae Heiden and Kolbe
Genus Gyrosigma Hassal, 1845
Gyrosigma attenuatum (Kutzing), Rabenhorst.

(Plate 2, figures 6, 10.)

Frustulia attenuata Kutzing, 1833- Algarum aquae dulcis Germaniae, Decas 9.
Gyrosigma attenuatum (Kutzing), Rabenhorst, 1853- Siisswasser-Diat., p. 47, pi. 5,

fig. 2. Cleve, 1894. Naviculoid Diat., p. 115. Elmore, 1921- Diat. Nebraska, p.

106, pi. 23, fig. 857. Boyer, 1927. Syn. N. Amer. Diat., p. 455. Hustedt, 1930.

Bacillariophyta, p. 224, fig. 330. Lohman, 1938. Pliocene Diat. Kettleman Hills,

California, p. 84, 1852-
Pleurosigma attenuatum WM. SMITH, 1852. Notes on Diat. p. 11, pi. 2, figs. 11-13, 18-

1853, British Diat., vol. 1, p. 68, pi. 22, fig. 216. Peragallo, 1891, Mono.

Pleurosigma, p. 16, pi. 7, fig. 9. De Tom, 1891- Syl. Algarum, p. 248-
Pleurosigma attenuatum var. caspium Grunow, PERAGALLO, 1891. Mono. Pleurosigma,

p. 18, pi. 7, fig. 8.
Pleurosigma attenuata Kutzing, VAN HEURCK, 1880. Syn. Diat. Belgique, p. 117, pi.

21, fig. 11.
Pleurosigma hippocampus Wm. Smith, PERAGALLO, 1891- Mono. Pleurosigma, p. 18, pi-
7, figs. 4-7. Van Heurck, 1880. Syn. Diat. Belgique, p. 117, pi. 20, fig. 3.

The central or nearly central median line, is slightly sigmoidal in out-
line. The transverse striae are not as well developed or as close together as
the longitudinal striae. The transverse striae number about 12 in 0.01 mm.,
while the longitudinal striae about 14 in 0.01 mm. This form ranges from
0.2090 to 0.2250 mm. in length, and from 0.0220 to 0.0270 in breadth.

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS 15

Only a few individuals of Gyrosigma attenuatum were found in the Pan-
amint Valley samples. Tempere and Peragallo (1915, pp. 144, 203) reported
it from "Yagiana [^Yaquina] Bay, Oregon"; and (p. 180) from Big Lake, Arling-
ton, Washington. Bailey (1921, p. 7) reported it from the Little and Big Quill
lakes, Saskatchewan, living in brackish water.

Gyrosigma attenuatum has been reported from the following fossil de-
posits: Lohman (1935, p. 456) Clovis, New Mexico, Quaternary; Lohman(1937,
p. 84), Kettleman Hills, Fresno County, California, Pliocene, Tulare formation;
Lohman (1934, pp. 10-11), Tipton, Oregon, Miocene, Mascal formation.

Family Cymbellaceae Agardh
Subfamily Cymbelloideae Karsten

Genus Cymbella Agardh, 1830
Cymbella mexicana (Ehrenberg) Cleve.

(Plate 2, figure 7-)

Cocconema mexicanum EHRENBERG, 1844. Einfluss des unsichtbar kleinen organischen
Lebens aus die Bildung von Bimestein, Tuff und anderen vulkanischen Gestein-
en, p. 342; 1854, Mikrogeologie, pi. 33, gr. 7, figs. 6, 7. Schmidt, 1875- Atlas
Diat., pi. 10, figs. 32, 33? Wolle, 1890. Diat. N. America, pi. 6, fig. 4.

Cymbella mexicana (Ehrenberg) Cleve, 1894- Naviculoid Diat., p. 177. Hanna and
GRANT, 1931- Diat. Pyramid Lake, Nevada, p. 287, pi. 25, fig. 3. Hustedt in
SCHMIDT, 1931- Atlas Diat., pi. 376, figs. 1,2. Hanna, 1938- Pliocene Diat. Kan-
sas, pi 379, pi- 32, fig. 4. Lohman, 1938. Pliocene Diat. Kettleman Hills, Cali-
fornia, p. 84, pi. 23, fig. 14. OKUNO, 1956. Elect.-Micro. Fos. Diat., p. 136, pi.
21, figs. 2 a-c. Sovereign, 1958. Diat. Crater Lake, Oregon, p. 125.

Cymbella kamtschatica Grunow in Schmidt, 1875- Atlas Diat., pi. 10, fig. 31.

Cymbella gastroides Kiitzing, Hanna and Grant, 1929- Brackish-water Pliocene Diat.,
Etchegoin formation, California, p. 92, pi. 12, fig. 3-

This conspicuous diatom is characterized by a distinct stigma in the
middle of the central nodule. The costae, about six to eight in 0.01 mm., are
coarsely punctate, with 10 to 12 puncta in 0.01 mm., and are usually alter-
nately long and short in the central portion of this particular form. Valves
range in length from 0.0143 to 0.01780 mm. and in breadth from 0.0320 to
0.0360. It is common in the Panamint Valley assemblage.

Cymbella mexicana was first described by Ehrenberg (1844) from a dia-
tomite locality in Mexico. Cleve (1894, p. 177) reported it living in fresh wat-
er in Washington and Vancouver Island. Tempere and Peragallo (1915) p.
194) reported the species from Swan Lake, Klamath County, Oregon; (p. 59),
Tacoma, Washington; (p. 165), Orcas Island, Puget Sound, Washington; (p. 179),
Big Lake, Arlington, Washington. Hanna and Grant (1931, pp. 287-288), re-
ported it from Pyramid Lake, Nevada, living, brackish water, and stated that

16 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

Cymbella mexicana is very common in many lakes in the western part of North
America, especially those slightly saline. Sovereign (1958, p. 125), reported
it from Crater Lake, Emerald Pool, Diamond Lake, and Upper Klamath Lake,
Oregon, living in waters with a pH of 7.0 to 9.0.

This species has been found in the following fossil deposits in western
North America: Boyer (1926, table opposite p. 26, column E), Deadman River,
north of Kamloops Lake, British Columbia. Ehrenberg (1869, p. 11), near Tul-
ancingo, Stadt Mexiko A. 6840'; (p. 12), Istlahuaca, 7740'; (p. 13), San Andres
Chalchicomula, 7100', Stadt Mexiko B. 6480'; Hacienda Escalera A.; (p. 20),
Stadt Mexiko K. 6840', (p. 21), Stadt Mexiko L. 6840', and Hacienda Escalera
D., all in Mexico. Tempere and Peragallo (1915) (p. 279), Lost Spring Ranch,
(p. 312), Pit River, California. Okuno (1956, p. 139), Terrebonne, Oregon.
Lohman (1937, p. 84), Kettleman Hills, Kings County, California, Pliocene,
Tulare formation, fresh to brackish water, and the upper part of the Etchegoin
formation, fresh to brackish water. Hanna and Grant (1929, p. 92), Kettleman
Hills, Fresno County, California, Pliocene, Etchegoin formation.

Family Epithemiaceae Grunow
Subfamily Epithemioideae Hustedt

Genus Epithemia Brebisson, 1838
Epithemia argus (Ehrenberg) Kiitzing.
(Plate 2, figure 12.)

Eunotia arcus EHRENBERG, 1838. Infusionsthierchen, p. 191, pi. 21, fig. 22-
Epithemia argus (Ehrenberg) KiJTZING, 1844- Bacillarien, p. 35, pi- 29, fig. 55. Fricke

in Schmidt, 1904. Atlas Diat., pi. 251, figs. 12, 14. Boyer, 1916. Diat. Phila-
delphia, pi. 31, figs. 15, 21; 1927, Syn. N.American Diat., p. 489. HUSTEDT,

1930. Bacillariophyta, pp. 383-384, fig. 727a.
Epithemia argus (Ehrenberg) Kiitzing var. alpestris GRUNOW, I860. Ueber neue oder

ungenugend gekannte Algen, pi. 3, fig. 28. Grunow, 1862. Osterreichischen

Diat., p. (329) 15-
Epithemia argus (Ehrenberg) var. amphicephala, GRUNOW in VAN HEURCK, 1881- Syn.

Diat. Belgique, pi. 31, fig. 19.
Epithemia argus (Ehrenberg) var. angusta, FRICKE in SCHMIDT, 1904. Atlas Diat., pi.

251, fig. 5.
Epithemia argus (Ehrenberg) var. capitata, FRICKE in SCHMIDT, 1904. Atlas Diat., pi.

251, fig. 14.
Epithemia alpestris Wm. SMITH, 1853- British Diat., pi. 1, fig. 7.
Eunotia argus EHRENBERG, 1854. Mikrogeologie, pi. 15, gr. A, fig. 59.
Eunotia comta Ehrenberg, 1840. Characteristic von 274 neuen Arten von Infusorien,

p. 210; 1854, Mikrogeologie, pi. 6, gr. 2, fig- 17 e-f.
Cystopleura argus (Ehrenberg) KUNTZE, 1891. Rev. Gen. Plant., p. 891- DE TONI, 1892.

Syl. Algarum, p. 782- Elmore, 1921. Diat. Nebraska, p. 129, fig- 661; pi. 23,

figs. 851, 854.

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS 17

Several varieties of this species have been named. After searching the
literature and different diatom strewn slides, all of them, with the exception
of Epithemia argus var. longicornis, seem to grade into one another.

In zone (edge) view the frustule is rectangular and characterized by two
rows of large nodules which are formed from the thickening of robust costae.
Thes'e are about one to two costae in 0.01mm. A well developed canal raphe ex-
tends almost to the dorsal margin of the valve. This species varies considerably.

Although this species is abundant in the Panamint Valley fossil assem-
blage, it has been reported living from only one western locality: Tempere and
Peragallo (1915, p. 179), Big Lake, Arlington, Washington.

It has been reported from many fossil deposits, such as: Patrick (1936
p. 164), Great Salt Lake, Utah. Ehrenberg (1870, table opposite p. 68), has
also reported this species from the same area. Hasler and Crawford (1938, p.
25), south of Sevier Lake, Millard County, Utah, Pleistocene. Boyer (1926b,
table opposite p. 26, column E), north of Kamloops Lake, British Columbia.
Ehrenberg (1870, table 12 opposite p. 68), Fall River, Oregon. Tempere and
Peragallo (1915, p. 179), Lost Spring Ranch, California. Lohman (1935, pp.
455, 457), Clovis, New Mexico, Quaternary. He stated that Epithemia argus
is, "...living in fresh-water lakes, but more frequently in saline lakes and
brackish estuaries." Ehrenberg (1869, p. 13), San Andres Chalchicomula,
7200'; (p. 14), Stadt Mexiko C. 6840'; and (p. 20), Stadt Mexiko K. 6840'. Loh-
man (1937, p. 84), Kettleman Hills, Fresno County, California, Pliocene, Tu-
lare formation.

Epithemia argus var. longicornis (Ehrenberg), Grunow.

(Plate 2, figure 13.)

Epithemia longicornis (Ehrenberg), Wm. SMITH, 1853- British Diat., p. 13, pi. 30, fig. 247.

Epithemia argus var. longicornis GRUNOW, 1862. Osterreich Diat. p. (329) 15. FRICKE
in Schmidt, Atlas Diat., 1904, pi. 251, figs. 1, 6, 9. Van Heurck, 1881. Syn.
Diat. Belgique, p. 139, pi. 31, figs. 15, 16. HUSTEDT, 1930. Bacillariophyta, p.
384, fig. 727.

This form is characterized by obtuse extremities (not capitate or con-
stricted in anyway). As such it seems to be a good variety. There are four or
more striae (12-14 in 0.01 mm.) between two consecutive costae. The length
of the valves range from 0.0550 to 0.0680 mm. while the breadth ranges from
0.0100 to 0.0120. Only a few individuals of this species were found in the
Panamint Valley assemblage.

The variety longicornis, has not been reported living in western North
America. It has been noted in fossil deposits, probably of Holocene age, from
the Great Salt Lake area by Ehrenberg (1870, table 12 opposite p. 68). It has
also been recorded from "Utah," probably near Great Salt Lake, by A. Schmidt
(1904, pi. 251).

18 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

PLATE 2

Figure 1. Anomoeoneis costata (Kiitzing) Hustedt. Hypotype no. 3449. (Calif.
Acad. Sci., Geol. Dept. Type Coll.), from locality 37113 (CAS), Mono Lake, California,
Pleistocene. Length 0.2070 breadth 0.0560 mm. x 400.

Figure 2- Anomoeoneis costata (Kiitzing) Hustedt. Locality 37113 (CAS), Mono
Lake, California, Pleistocene. Length 0.1860, breadth 0.0390 mm. x 475.

Figure 3. Rhopalodia gibba (Ehrenberg) O. Muller. Hypotype no. 3450. (Calif.
Acad. Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Valley, Cal-
ifornia, Pleistocene. Length 0.1340, breadth 0.0270 mm. x 700.

Figure 4. Pinnularia viridis (Nitzsch) Ehrenberg. Hypotype no. 3451. (Calif.
Acad. Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Valley,
California, Pleistocene. Length 0.2430, breadth 0.0400 mm. x 375.

Figure 5. Navicula oblonga (Kiitzing). Hypotype no. 3452- Calif. Acad. Sci.,
Geol. Dept. Type Coll.), from locality no. 37055 (CAS), Panamint Valley, California,
Pleistocene. Length 0.0805, breadth 0.0104 mm. x 600.

Figure 6. Gyrosigma attenuatum (Kiitzing) Rabenhorst. Hypotype no. 3453-
(Calif. Acad. Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Val-
ley, California, Pleistocene. Length 0.2140, breadth 0.0250 mm. x 20.

Figure 7. Dynedra sp., Hypotype no. 3454. (Calif. Acad. Sci., Geol. Dept. Type
Coll.), from locality 37113 (CAS), Mono Lake, California, Pleistocene. Length 0.2040,
breadth 0.0180 mm. x 470.

Figure 8. Cocconeis placentula Ehrenberg. Hypotype no. 3455. (Calif. Acad.
Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Valley, California,
Pleistocene. Length 0.0280, breadth 0.0170 mm. x 1570.

Figure 9. Cymbella mexicana (Ehrenberg) Cleve. Hypotype no. 3456. (Calif.
Acad. Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Valley,
California, Pleistocene. Length 0.1510, breadth 0.0320 mm. X1200.

Figure 10. Gyrosigma attenuatum (Kiitzing) Rabenhorst. Same specimen as in
figure 6, enlarged to show details of valve, x 1000.

Figure 11. Cocconeis placentula Ehrenberg. Hypotype no. 3457. (Calif. Acad.
Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Valley, California,
Pleistocene. Length 0.0260, breadth 0.0190 mm. x 2000.

Figure 12. Epithemia argus (Ehrenberg) Kiitzing. Hypotype no. 3458. (Calif.
Acad. Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Valley, Cal-
ifornia, Pleistocene. Length 0.0665, breadth 0.0125 mm. x 1000.

Figure 13. Epithemia argus var. longicornis Grunow. Hypotype no. 3459. (Calif.
Acad. Sci., Geol. Dept. Type Coll.), from locality 37055 (CAS), Panamint Valley,
California, Pleistocene. Length 0.0550, breadth 0.0100 mm. x 1365.

OCC PAPERS CALIF. ACAD. SCI., NO. 46

(WORNARDT)

PLATE 2

20 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

Subfamily Rhopalodoideae Hustedt
Genus Rhopalodia O. Muller, 1897
Rhopalodia gibba (Ehrenberg) O. Muller.

(Plate 2, figure 3.)

Navicula gibba EHRENBERG, 1830 [1832] . Beitrage zur Kenntniss der Organisation der
Infusorien und ihrer geographischen Verbreitung, besonders in Siberien, p. 64.

Rhopalodia gibba (Ehrenberg), O. MULLER, 1897. Rhopalodia, neues Genus Bacill.,
p. 65, pi. 1, figs. 15-17. Hustedt in Schmidt, 1905- Atlas Diat., pi. 253, figs. 1-
13. SCHONFELDT, 1907. Deutschen Diat., p. 206, pi. 14, fig. 14- MEISTER, 1912.
Kieselalgen Schweiz, p. 200, pi. 35, fig. 6. Boyer, 1916. Diat. Philadelphia, p.
112, pi. 31, fig. 23; 1927, Syn. Diat. N.America, p. 481. HUSTEDT, 1930. Bacil-
lariophyta, p. 390, fig. 740. HANNA, 1932. Pliocene Diat. Kansas, p. 386, pi. 34,
figs. 2, 3- PATRICK, 1936. Diat. Siam and Malay States, p. 428; 1936, Diat. Lin-
seley Pond, Connecticut, p. 86. LOHMAN, 1937. Pliocene Diat. Kettleman Hills,
California, p. 84, pi. 23, figs. 1, 2- KRASSKE, 1939- Kieselalgenflora Sudchiles,
p. 408.

Epithemia gibba Kutzing, 1844. Bacillarien, p. 35, pi. 4, fig. 22- Van Heurck, 1881.
Syn. Diat. Belgique, p. 139, pi. 32, figs. 1, 2. WOLLE, 1890. Diat. N. America,
pi. 35, figs. 1, 2, 8, 9.

Cystopleura gibba (Ehrenberg) KUNTZE, 1891- Rev. Gen. Plant., p. 891. ELMORE, 1921.
Diat. Nebraska, p. 127, pi. 17, fig. 654.

Eunotia gibba Ehrenberg, BAILEY in Fremont, 1842-1844 [l845J • Rept. Explor. Exp., p.
302, pi. 5, figs. 4, 5.

The raphe is characteristically found along the convex edge or keel of
the linear valves in this genus (not present in view shown on plate 2, figure 3).
The central portion of the frustule is gibbous, while the ends tend to be slight-
ly reflexed. There are about seven costae in 0.01mm. The length ranges from
0.1240 to 0.1310 mm. and in breadth from 0.0280 to 0.3050 mm.

A few individuals of Rhopalodia gibba were found in the Panamint Val-
ley samples. The species was originally described by Ehrenberg (1830 [1832],
p. 64) from a locality in Siberia. Tempere and Peragallo (1915, p. 59) have
reported it from Tacoma, Washington; (p. 165), Orcas Island, Puget Sound,
Washington; (p. 179), Big Lake, Arlington, Washington; and (p. 194), Swan
Lake, Klamath County, Oregon. Sovereign (1958, p. 127) reported the species
living in waters with a pH of 6.0 to 11.0 but having its optimum in water with
a pH of 9.0, in Emerald Pool and Diamond Lake, Oregon.

Rhopalodia gibba has been reported in fossil deposits from Canada to
Mexico, such as: Boyer (1926b, table opposite p. 26, column E), near Cobble
Hill, Vancouver Island, British Columbia. Patrick (1936, p. 162), Great Salt
Lake, Utah [Holocene) , fresh to brackish water. Ehrenberg (1870, table 12 op-
posite p. 68), Truckee River, Nevada. Tempere and Peragallo (1915) (p. 280),

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS 21

Orcas Island, Puget Sound, Washington; (p. 328), Reno, Nevada. Bailey (1845,
p. 302), Deschutes Rives, Oregon, freshwater, fossil. Lohman (1935, pp. 455-
457), "...living sparingly in fresh-water lakes, but more frequently in saline
lakes and brackish estuaries."; Lohman (1938, pp. 84, 90), Kettleman Hills,
Kings County, California, Pliocene, Tulare formation, fresh to brackish water.

Suborder SURIRELLINE AE West

Family Surirellaceae Kiitzing
Subfamily Surirelloideae Schutt

Genus Surirella Turpin, 1928
Surirella ovalis Brebisson.
(Plate 1, figure 5.)

Surirella ovalis BREBISSON, 1838. Considerations sur les Diatomees et essai d'une
classification des genres et des especes appartenant a cette famille, p. 17.
Kutzing, 1844. Bacillarien, p. 61, pi. 30, fig. 64- Rabenhorst, 1853- Susswas-
ser-Diat., p. 30, pi. 3, fig. 24- WM. SMITH, 1853- Syn. British Diat., p. 33, pi. 9,
fig. 68. Pritchard, 1861. Infusoria, p. 796. Schmidt, 1885 [l 875], pi. 24, figs.
1-4. Van HeuRCK, 1881. Syn. Diat. Belgique, p. 188, pi. 73, figs. 2-4- Boyer,
1927. Syn. N. American Diat., p. 541- HUSTEDT, 1930. Bacillarophyta, p. 441,
figs. 860, 861.

Valves ovate and somewhat lanceolate. The apices of the valves are
both cuneate or rounded or one may be rounded and the other cuneate. One
end of the valve is usually somewhat broader than the other. The costae are
short, marginal, and radiate. The central portion is indistinctly striated. This
form ranges from 0.0710 to 0.0870 mm. in length and from 0.0410 to 0.0590 mm.
in breadth. Hustedt (1930, pp. 440-445), has unraveled most of the confusion
concerning the identification of Surirella angusta, S. crumena, S. minuta, S.
ovalis, S. ovata, S. pinnata, and S. salina.

A few individuals of Surirella ovalis were found in the sample from the
Mono Lake beds. Although the species has been reported as living, it is
probably fairly common in only one area of western North America; viz., in
the brackish water of the Little and BigQuill lakes, in Saskatchewan, Canada,
(Bailey, 1921, p. 7). It has been found in several fossil deposits. Lohman
(1935, p. 457) reported the species in Quaternary lake beds from Clovis, New
Mexico. He stated that the species is "...living sparingly in fresh-water lakes,
but more frequently in saline lakes and brackish estuaries." Hasler and Craw-
ford (1938, p. 26) noted its occurrence in a marl deposit of "Bonneville Age"
(Pleistocene) from south of Sevier Lake, in Millard County, Utah.

22 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

Surirella ovata var. utahensis Grunow.

(Plate 1, figure 4.)

Surirella ovata Kl'TZlNG, 1844. Bacillarien, p. 62, pi. 7, figs. 1-4- BOYER, 1927. Syn.
N. American Diat., p. 541-

Surirella ovata var. utahensis GRUNOW in SCHMIDT, 1885 [l875] • Atlas Diat., pi. 24, figs.
11-13. Patrick, 1936. Diat. Great Salt Lake, p. 163.

Surirella ovalis Brebisson, Wolle, 1894 [T890]. Diat. N.America, pi. 53, figs. 7, 8.

Not Surirella utahensis Grunow, Hanna and GRANT, 1929- Brackish-water Diat., Etche-
goin formation, California, p. 100, pi. 14, fig. 6; 1931, Diat. Pyramid Lake, Nev-
ada, p. 290, pi. 26, figs. 5, 6.

The distinguishing character of this variety is given in Grunow's origi-
nal definition, "S. ovata K. var. utahensis Grunow. hat zu ausgepragte Eigen-
thumlichkeit, als dass ich sie unter S. ovata stellen mochte; man beachte
die bogenformige Falte in der Mittel!" (Schmidt's Atlas Diat., 1885, pi. 24,
figs. 11-13.)

This transverse fold is present on all forms of the variety, and is locat-
ed about two-thirds the distance from the acute apex. The submarginal mod-
erately long costae (four in 0.01 mm.) with fine striae between, radiate from
an indistinct medial line. The forms range in length from 0.0820 to 0.1100 and
in breadth from 0.0900 mm.

Surirella ovata var. utahensis is found in the lake beds at Mono Lake,
but was first obtained in material from the Great Salt Lake area, Utah. It was
also reported from a Holocene deposit in Great Salt Lake area by Patrick (1936,
p. 163), who stated that the species is commonly found in fresh and brackish
water.

Surirella striatula Turpin var. testudo Ehrenberg.

(Plate 1, figures 7, 8.)

Surirella Testudo EHRENBERG, 1840. Characteristik von 274 neuen arten vonlnfusorien,
p. 215; 1869, Uber machtige Gegirgs-schichten vorherrschend aus Mikro. Bacill.
unter und bei der StadtMexiko, pi. 1, gr. E, fig. E-4; 1870, Uber die wachsende
Kenntnis des unsichtbaren Lebens als felsbildende Bacillarien in Californien,
pi. 1, figs. 1, 2; pi. 2, gr. 1, fig. 6. HANNA and GRANT, 1931- Diat. Pyramid Lake,
Nevada, p. 290, pi. 27, figs. 2-4, 1931-

Surirella striatula Turpin, Schmidt, 1875- Atlas Diat., pi. 24, fig. 17. Wolle, 1894
[1890]. Diat. N. America, pi. 55, fig. 6.

Not Surirella striatula TURPIN, 1828. Observations sur le nouveau genus Surirella, p.
362, pi. 15.

Surirella baileyana Mackay or 5" striatula var. baileyana McKay, Bailey, 1921 [1922].
Diat. Saskatchewan, Albe a, p. 158, pi. 1, figs. 3-6. BOYER, 1927- Syn. N.Amer-
ican Diat., p. 539.

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS 23

The diagnostic features of this variety are that the parallel valves of
the frustule are twisted in the same plane, sometimes at both ends or some-
times just at one end. The pseudoraphe is therefore asymmetrical. Canaliculi
are distinct, about 7-15 on each side of the valve surface; striae range be-
tween 15-20 in 0.01 mm. There is a distinct row of beads along the margin of
each lobe. The diatom was fairly common in the Mono Lake bed sample.

This variety was originally described from the Great Salt Lake area,
Utah, by Ehrenberg (1871, table 12 opposite p. 68), but it has never been de-
termined whether or not these specimens were living or fossil. I think it is
reasonable to assume that the sample was from a fossil deposit because Pat-
rick (1936, p. 165) stated, "No diatom flora is found living in the lake proper
today." She has, however, reported many individuals of this species from de-
posits in and around the Great Salt Lake area. These deposits are probably of
Holocene age. According to L.W.Bailey (1921, p. 4), the species is living
in the brackish waters of the Little and Big Quill lakes in Saskatchewan,
Canada. Hanna (1939, p. 290) also has reported it living in the brackish wat-
ers of Pyramid Lake, Nevada.

In 1926 Mann (p. 53) reported the species from fresh to brackish water
sediments of the Latah Formation, of Miocene age, exposed near the town of
Mica, near Spokane, Washington. He made the following statement:

"This species seems to flourish in waters high in mineral content. It
often holds its place in lakes that show a gradual increase in salinity after
other species have been exterminated by the high percentage of salts. Thus
in samples of borings made at the site of a prehistoric lake in the Black Rock
district of Nevada, which slowly changed from fresh water to salt water, this
species was the last diatom to succumb to the growing salinity. It is also
sometimes the dominant species in such saline water as Devils Lake, N. Dak.,
and Salt Lake, Utah."

This variety, therefore, would be considered a euryhaline form.

Subfamily Campylodiscoideae Heiden and Kolbe
Genus Campylodiscus Ehrenberg, 1838
Campylodiscus noricus var. hibernica (Ehrenberg) Grunow.

(Plate 1, figure 3.)

Campylodiscus noricus var. bibemicus GRUNOW, 1862. Osterreichischen Diatomace'en,
p. (439) 125. MAYER, 1912. Bacill. Regensburger Gewasser, p. 347, pi. 26, fig.4.
HUSTEDT, 1914. Bacill. aus den Sudeten, p. 127; 1930, Bacillariophyta, p. 446,
fig. 872. Dahm, 1956. Diatomeenuntersuchungen zur Geschichte der westlichen
Ostsee, pi. 6, fig. 5. Sovereign, 1958- Diat. Crater Lake, Oregon, p. 132.

Not Campylodiscus noricus var. bibemicus Grunow, MEISTER, 1912. Kieselalgen
Schweiz, p. 231, pi. 43, fig. 2.

24 CALIFORNIA ACADEMY OF SCIENCES (Occ. Papers

Campylodiscus costatus Wm. SMITH, 1851. Notes on the Diatomaceae; Campylodiscus,
Surirella, and Cymatopleura, p. 6, pi. 1, fig. 1; 1853, Brit. Diat., p. 29, pi. 6,
fig. 52; pi. 7, fig. 52. Meister, 1912. Kieselalgen Schweiz, p. 230, pi. 47, fig. 3.

Campylodiscus hibernicus EHRENBERG, 1842. Samples of Infusorial Earths from the
Mourne Mountains, Ireland, p. 337; 1845, Vorlaufige zweite Mittheilung iiber die
weitere Erkenntniss der Beziehungen des kleinsten organischen Lebens zu den
vulkanischen Massen der Erde, p. 154; 1854, Mikrogeologie, pi. 15, gr. A, fig. 9.
SCHMIDT, 1878 [I877J. Diat., pi. 55, fig. 13. Wolle, 1894 [[890]- Diat. N. Amer-
ica, pi. 69, fig. 13.

Campylodiscus noricus Ehrenberg, MEISTER, 1912. Kieselalgen Schweiz, p. 230.

Campylodiscus noricus var. costatus Grunow, MEISTER, 1912. Kieselalgen Schweiz,
p. 230, pi. 47, fig. 3.

Campylodiscus noricus var. sublaevis MEISTER, 1912. Kieselalgen Schweiz, p. 230,
pi. 47, fig. 2.

This variety is characterized by robust costae, two in 0.01 mm., with
many fine striae between. The central portion of the valve is usually finely
punctate. Valves are usually bent or saddle-shaped. Size ranges from 0.0715
to 0.0750 mm. in diameter. Most of the confusion concerning Campylodiscus
costatus, C. noricus, and their varieties has been resolved by Hustedt (1914,
pp. 127, 128; 1930, pp. 446-448). Grunow originally spelled the name of this
variety "hibernicus" (1860, p. 439), as did Hustedt (1914, p. 127). Hustedt
then changed the spelling to "hiberica'" (1930, p. 446), and was followed by
Dahm (1956, pi. 6, fig. 5), and Sovereign (1958, p. 132).

One of the least common but most diagnostic species found in the Pana-
mint Valley sample was Campylodiscus noricus var. hibernica. It has been re-
ported in Big Lake, Arlington, Washington, oy Tempere and Peragallo (1915,
p. 179), and probably was living there. In 1958, Sovereign (p. 132) noted its
occurrence in Diamond Lake and Upper Klamath Lake, Oregon, and he stated
that the species was fairly common in the Pacific Northwest. He also noted
that it was common in waters with a pH between 7.0 and 9.0 but that it had
its optimum in waters with a pH of 8.0 to 9.0-

Campylodiscus noricus var. hibernica has been reported from only two
fossil localities in western North America, both being noted by Tempere and
Peragallo (1915). The first was from the Lost Spring Ranch, California, prob-
ably from a freshwater fossil deposit (p. 279). The second is from Conrad's
Red Bluff, California, and is probably also from a freshwater fossil deposit
(p. 426). The exact location of these places has not been found.

No. 46) WORNARDT: PLEISTOCENE LAKE DIATOMS 25

REFERENCES

BAILEY, J. W. (in Fremont, J. C.)

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BAILEY, L. W.

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26 CALIFORNIA ACADEMY OF SCIENCES (Oc:c. Papers

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MBI. WHOI I.IBKAKV

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