A CENTURY OF PROGRESS
IN THE NATURAL SCIENCES

^ '

A CENTURY OF

PROGRESS IN THE

NATURAL SCIENCES

1853-1953

PUBLISHED IN CELEBRATION OF

THE CENTENNIAL OF

THE CALIFORNIA ACADEMY OF SCIENCES

/

California Academy of Sciences
SAN FRANCISCO

1955

COPYRIGHT, 1955. BY THE
CALIFORNIA ACADEMY OF SCIENCES

Dedicated to the memory of
JOHN WARD MAILLIARD, JR.

m appreciation of his long and faithful service as

a Trustee and as Chairman of the Board, of his

many benefactions to the Academy, and of

his stimulating faith in its future

COMMITTEE ON PUBLICATION

Dr. Robert C. Miller, Chairman

Dr. Edward L. Kessel, Editor

Dr. G. F. Papenfuss

FOREWORD

This volume of essays has been prepared as part of the recognition of the
Centennial of the California Academy of Sciences. In May, 1951, three mem-
bers of the Council were authorized by the Trustees of the Academy to make
plans for a volume of scientific papers appropriate to the occasion. After care-
ful consideration the committee decided that a most appropriate central theme
for the volume would be the historical treatment of biosystcmatics, using this
term in the literal sense, namely, the systematic treatment of living things and
with emphasis on developments since the founding of the Academy a cen-
tury ago.

This theme appealed to the committee as especially appropriate since it
was during this period, from the middle of the nineteenth to the middle of the
twentieth century, that the basic principles underlying our present concepts
and aims in the classification and systematic treatment of organisms were clearly
enunciated and definitely accepted among biologists. The nineteenth century
brought to biology two all-important contributions, Darwin's and Wallace's con-
ception of organic evolution and Mendel's principles of heredity. Recognition
of the doctrine of organic evolution led directly to the working concepts of the
continuity of species and the transformation of old species into new ones. Recog-
nition of the basic laws of heredity has led, in the twentieth century, to very
great progress in the development of our concepts of the nature of the evolu-
tionary processes.

It was inevitable that these tremendous forward steps should have a pro-
found impact on the thinking and practices of those systematists who recognize
the significance of the facts, not only of comparative morphology, but also of
variation and heredity and of the contributory disciplines of cytogenetics, physi-
ology, biochemistry, serology, biometry, ecology, and biogeography. Inevitable
too was the apathy shown toward these epoch-making advances by many taxono-
mists who were content to pile up new names of species and genera without
critical study of all available criteria of relationship, thus creating a maze of
names rather than systematics. Although some taxonomists are still littering
the waj^sides of biological literature with unnecessary names, there is a growing
tendency among systematists to bring to bear upon problems of classification
and nomenclature all of the various categories of evidence that are available in
order that the decisions reached shall represent as nearly as possible the true
state of nature. This modern viewpoint and aim is the culmination of many
experiments in the systematic treatment of organisms prior to and extending
throughout this ''Darwinian" century.

It is only in recent decades, however, that the advantages of the many-sided
attack on problems of relationship and phylogeny have been realized. Many ob-
scure problems in the relationship of organisms have been cleared up by the
evidence from cytology, genetics, and biochemistry, not to mention other con-
tributor}^ disciplines; and, in many instances, such evidence has resulted in radi-
cal changes in older taxonomic treatments. At the same time, it has been clearly

vii

Viii A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

demonstrated that the evidence on relationship provided by the "newer" dis-
ciplines corroborates in the main the earlier systematic treatments that were
devised by taxonomists who based their schemes primarily on comparative mor-
phology. Certainly due credit should be given to the many "specimen taxono-
mists" who have labored through the centuries, often without fair recognition
from other biologists and under great difficulties, in their conscientious efforts
to bring hitherto unknown organisms into some sort of classificatory system.
Without their invaluable services the general advance of biology would not
have been possible.

Most of the essays in this volume attempt to review the progress made dur-
ing the past century in the classification of organisms. The original plan of the
volume included all the major groups of organisms. It was found impossible to
achieve this degree of completeness; but except for a few gaps the earth's organic
life is well represented and the committee consider it a great honor to be able to
present to the biological world this series of authoritative historical reviews.

In the exploratory phase of plant and animal classification the services of
field workers, especially of trained naturalists, are indispensable. Much of the
activity of the California Academy of Sciences has been concerned with the
collection and preservation of specimens. It seemed appropriate, therefore, that
the first essay should deal with naturalists and the early days of the Academy.
The following chapter presents a review of the beginnings of geodesy and astron-
omy in California because this Academy was so closely tied in with those events;
and the third essay is a stimulating contribution by a philosophically minded
biosystematist. Then follows the series of systematic reviews, together with four
essays which do not treat of major groups of organisms — one on invertebrate
paleontology, two on biogeography, and one on wildlife conservation. In all of
these essays the disciplines represented are largely, but with some additions,
those which have come within the purview of the California Academy of Sciences.

The committee are confident that this volume will long serve as a most valu-
able source book in the history of science.

ERNEST B. BABCOCK
J. WYATT DURHAM
GEORGE S. MYERS

CONTENTS

San Francisco as a Mecca for Nineteenth Centurv Naturalists

Joseph Ewan 1

A Century of Astroxo.^iy and (Jioodesy in California . . .Ericin (i. Giulde 65

The Contribution of Natural History to Human Progress . .G.F. Ferris 75

Classification and Taxonoimy of the Bacteria and BluegreexV Algae

C. B. Van Niel 89

Classification of the Algae George F. Papenfuss 115

Mycology Ernst Athearn Bessey 225

Bryology WiUimn C. Steere 267

Pteridology Irc7ie Manton 301

The Systematics of the Gymnosperms Rudolf Florin 323

The Systematics of the Angiosperais Lincoln Constance 405

Systematic Entomology Edward S. Ross and Collaborators 485

Introduction Edward S. Ross 485

The "Apterygota" Charles L. Remington 495

Odonata Leonora K. (}lo\jd 506

Ephemeroptera George F. Edmunds, Jr. 509

Plecoptera Per Brinck 512

Embioptera Edward S. Ross 515

Zoraptera Ashley B. Gurney 516

SoAiE Minute Insects: Anoi'luka, Mvllophaga, and the Scale

Insects G. F. Ferris 517

Psocoptera ( Corrodentia, Copeognatha)

K. M. Sommerman and J. V. Pearman 523

Thysanoptera Stanley F. Bailey 525

IIOMOPTERA AUCHENORHYNCHA Z. P. Metcolf 527

IIemiptera Robert L. Usinger 534

Neuroptera and Mecoptera F. M. Carpenter 536

ix

Systematic Entomology (Continued)

Trichopter.v Herbert H. Ross 538

Lepidoptera William T. M. Forhes 540

CoLEOPTERA Melville H. Hatch 555

Strepsiptera R. M. Bohart 566

Ant Taxonomy W. L. Brown, Jr. 569

The Aculeate Wasps Paul D. Hurd, Jr. 573

The Apoidea Charles D. Michener 575

Diptera Charles P. Alexander 578

SiPHONAPTERA George P. Holland 585

Fossil Insects F. M. Carpenter 588

Herpetology Karl P. Schmidt 591

Ornithology Charles G. Sibley 629

Mammalogy in North America W.J. Hamilton, Jr. 661

Invertebrate Paleontology and Historical Geology from 1850 to 1950

Charles E. Weaver 689

Plant Geography Ronald Good 747

Animal Geography Karl P. Schmidt 767

The Conservation op Wildlife A. Starker J^eopold 795

ALBERT KELLOGG

JOHN B. TRASK

HENRY GIBBONS

THOMAS J. NEVINS

SAN FRANCISCO AS A MECCA FOR
NINETEENTH CENTURY NATURALISTS

With a Roster of Biographical References
to Visitors and Residents

By JOSEPH EWAN

Tulane University

As THE Genus is first identified by the distinctness of its species, so the country
is first distinguished by its most prominent city. Charleston served as the germ
of Carolina, New Orleans of Louisiana, Lima of Peru, Montevideo of Uruguay,
and San Francisco of California. California, a vast and diversified country, was
an island on the edge of El Dorado, said to be fabulous and fortunate, sought by
many, reached only with difficulty, and San Francisco was her heart. Even before
the Gold Rush, to come to California from European cities amounted to a journey
half way "round the world. And for the American back in the "States" coming
to the City of the Golden Fifties was not just going across the Shenandoah to a
frontier valley, or just setting out west from Albany, or even the equivalent of
taking a clipper ship out of a New England port or New York for Charleston
or Apalachicola or New Orleans, but a voyage to a land far away, hemmed in by
the Humboldt Sink and the Sierra Nevada, and peopled by men and women who
had a different derivation and who spoke a different language. Very early in the
history of California reports came back of giants and riches, where ordinary
things were extraordinary, and superlatives were elementary parts of speech.
Great flocks of wildfowl in the marshes, grizzly bears that challenged the bravest
men, giant birds (the California condor), giant trees, and giant seaweeds. Even
the slugs in settlers' gardens were enormous ! But it was those giant nuggets of
gold ! The spirit of the Seven Cities of Cibola lives on.

Naturalists have always been in the vanguard of explorers : so it was in Cali-
fornia. With a party of prospectors who took the Gila Trail came Audubon's
son, John Woodhouse Audubon,^ and with a party of trappers following the
trail west from Santa Fe, came William Gambel. Most of these naturalist adven-
turers in the Great West were young men between the ages of nineteen and thirty
years. Some were serious naturalists trained in the essentials of the natural
sciences, either with field experience or with training in medicine, apprentices
to an apothecary or a taxidermist's helper. A few, like John Woodhouse Audubon,
Isaac J. Wistar, Titian Ramsey Peale, and John Lawrence LeConte, were scions
from old naturalist rootstocks. Some of these emigrant naturalists would cast
their lot to stav in California — and California meant in the cultural sense San

1. For biographical notices of naturalists mentioned in this account see the appended
roster.

[1]

2 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Francisco — to share in the founding and the support of the California Academy
of Sciences.

Naturalists in San Francisco Before 1853

In 1939 Alice Eastwood summarized the history of botanical exploration on
the Pacific Coast and four years later Eoland H. Alden and John D. Ifft pub-
lished in the Occasional Papers of the Academy a review entitled "Early Natural-
ists in the Far West." For this reason the notice given here to naturalists active
before 1853 will be brief.

The first naturalists to visit San Francisco were French explorers under
Comte de La Perouse who made a landfall there in 1786. La Perouse was com-
mander of the Boussole, and with him was the gardener and botanist, Jean
Nicholas Collignon, while the corps of the second vessel, Astrolabe, included
De Boissieu la Martiniere, "doctor of physic and botanist," and the naturalist,
Louis Dufresne. Six years later, in November, 1792, Captain George Vancouver
visited both San Francisco and Monterey and Archibald Menzies, surgeon-
naturalist to the expedition, took back to England the California condor (per-
haps taken along the lower Columbia River) but was able to collect only a few
plants. In 1806 another flag entered San Francisco Bay, representing a nation
that had as yet not challenged the Spanish supremacy in California. On March
28, 1806, the Russian ship Juno sought supplies for Russia's stricken colony
at Sitka, the base of her fur seal operations in the North Pacific. Langsdorff, an
officer on board the Juno, has left us a detailed account of the forty-four days
at anchor here.

The Russian settlement was established at Fort Ross in 1812, primarilj^ to
supply fresh vegetables for the scurvy-cursed men plying the boats in the Behring
Sea for seals. Trading vessels were not allowed to enter any port of California
at this time and Russians from Fort Ross who ventured into San Francisco were
held prisoners there by the Spanish for violations of the laws. It is unlikely,
therefore, that the Russians were able to collect many specimens in the region
at this time.

Ten years passed before a second Russian vessel, the Rurik, carrying an-
other surgeon-naturalist, Johann Friederich Eschscholtz, entered San Francisco
harbor on October 1, 1816. Captain Kotzebue carried with him on the Rurik
the well known poet and naturalist Adelbert von Chamisso. Though the visit of
the Rurik was made during the late fall dry season the expedition collected
a large number of novelties because of unusual rains.

Kotzebue visited San Francisco for the second time in 1824 and Dr. Esch-
scholtz again accompanied Kotzebue. The Russian ship spent nearly two months
in California, leaving San Francisco on November 25, 1824. The captain opined :
"I confess I could not help speculating upon the benefit this country would derive
from becoming a province of our powerful empire, and how useful it would prove
to Russia." Eschscholtz 's collections were exclusively zoological on this second
voyage. He died in 1831 before the completion of his Zoologischer Atlas, in which
he published his Calif ornian discoveries.

During the last years of the Russian occupation several Russian naturalists
visited northern California. These included Governor Ferdinand P. Wrangell;
Dr. F. Fischer and Dr. Edward L. Blaschke, of the Russian American Company;

fVVAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 3

the agriculturist, George Tsehernikh, and I. G. Vosnesensky, curator of the
Zoological Museum of St. Petersburg. The plant collections of Vosnesensky
came back to San Francisco, to the Academy, after lying in their herbarium
covers for nearly a hundred years in Russia. The collections were returned to
California for identification by John Thomas Howell, and then sent to Lenin-
grad's national herbarium.

In 1824 hide ships began operating along the California coast. These vessels
were the source of introduction of many organisms, some injurious: insects,
weeds, and rodents. This traffic in hides marked the reintroduction of some weeds
earlier introduced with the Mission Period which began in 1769 with the founding
of Mission San Diego. One of these ships took on a little piece of immortality,
for it was the Alert that carried Thomas Nuttall from California around the
Horn, with that commentator of the day, Richard Henry Dana.

The British expedition under Captain Beechey visited California in 1827.
The natural history collections were made on the voyage of H.M.S. Blossom
by the ship's surgeon, Dr. Alexander Collie, assisted by George Tradescant Lay,
and Lieutenant Belcher. The Blossom was in port twice, from November 7 to
December 28, 1826, and November 19 to December 3, 1827. Dr. Collie collected
the t5T)e specimens of thirteen species of birds either at San Francisco or Monte-
rey, both ports having been visited twice on the voyage.

The French sailing vessel Heros put in at San Francisco on January 26.
1827, with a surgeon on board, Dr. Paolo Emilio Botta, who was then twenty-one
years of age. Botta collected both birds — including the roadrunner — and plants.
The Heros spent nearly two years intermittently on the coast, from Fort Ross
to San Diego, finally departing on July 27, 1828. The California buckeye, named
Calothyrsus calif ornica by Spach, was one of Botta 's collections.

David Douglas, "Douglas of the Fir," arrived in San Francisco in 1831, fol-
lowing his first highly successful visit to America. His California visit introduced
dozens of species to horticulture and to systematic botany. Douglas botanized as
far south as Santa Barbara, making the Franciscan missions his lodging places
along the route. It is unfortunate that his fieldbooks were lost for few explorers
in California natural history would have had so much to tell. "Douglas, no mere
collector, was a skilled natural scientist in his own right. Of his character and
personality, what more need we say than that he courageously faced adversity
for the science he loved, and died in pursuit of knowledge?"

The Irish naturalist. Dr. Thomas Coulter, first served as a physician to a
mining company in Mexico before coming to Monterey in 1831, where he met
David Douglas in November. Coulter spent nearly three years on the Coast,
including a trip to the Colorado Desert, but did not remain on the Coast to meet
Nuttall, who closely followed him. Coulter may have met Ferdinand Deppe, a
professional collector from Berlin, at jMonterey but we have only fragmentary
knowledge of Deppe, save that he arrived in California during the winter of
1831-1832, possibly from the Mexican port of Loreto. David Douglas had met
Deppe in California sometime prior to October 24, 1832, and Deppe was at
Monterey as late as December, 1834, when he shipped bird skins to Lichtenstein,
then director of the Zoological jVIuseum of Berlin. The beautiful endemic ]\Iatilija
poppy, Romneya coulteri, was one of Coulter's discoveries in southern California.

Thomas Nuttall and John Kirk Townsend crossed the continent together witli

4 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Captain Nathaniel Wyeth, setting out from Independence, Missouri, on April 28,
1834, as members of an original party of seventy men with 250 horses. The
Townsend narrative, a model of forthright reporting and a treasure for the
serious student of the American West, makes some mention of the events of a
botanical and ornithological nature along the way, and gives us concrete evidence
of the devotion of Nuttall to science. Finally Nuttall returned 'round the Horn
in 1836 but To^vnsend remained another year on the Coast. Both of their collec-
tions ultimately reached Philadelphia, Nuttall dividing his plant specimens be-
tween the Philadelphia Academy and his personal herbarium, which ultimately
came to rest at the British Museum (Natural History). Audubon purchased
Townsend 's bird skins and enriched his own ornithological writings thereby.
Nuttall "raised himself from a penniless orphan to a highly respected man of
science," joining the era of B. S. Barton, his one-time patron, with that of Asa
Gray and Elias Durand. Nuttall's travels in America have been delineated by
Pennell with documentation, and his California visit has been fraternally told
by Jepson.

The London Horticultural Society, which first sponsored David Douglas in
America, sent twenty-four-year-old Karl Theodore Hartweg, of Karlsruhe, to
Mexico in 1836, and to California in 1846. He arrived in Monterey on June 7
and proceeded north to San Francisco and Chico late that year. His plant col-
lections in the northern Sierra Nevada were particularly valuable. Hartweg's
botanical collections fared better than most in that the British systematist George
Bentham handled them and published a commentary upon them entitled Plantae
Hartivegianae. Hartweg's companion on his visit to Bear Valley in the Sierra
Nevada was Theodor Cordua, "pioneer of New Mecklenburg," whose account of
the trip has recently been translated.

The French frigate La Vhius, under command of Admiral Abel du Petit-
Thouars, arrived at Monterey, October 18, 1837, and departed November 14.
Both zoological and botanical collections were made then and a description of the
California visit appears in Thouars' Voyage autour du monde sur la frigate "La
Venus" (Paris, 1840-1843, 2 : 77-142). The surgeon on the La Venus was Adolphe
Simon Neboux, who most likely made the natural history collections. A dexterous
piece of detective work involving this French expedition is John Thomas Howell's
story "Sea-gulls and Tarweeds: a Distributional Mix-up" (Leafl. West. Bot.,
1:189-191, 1935.).

Richard Brinsley Hinds, surgeon on H.M.S. Sulphur, visited the California
coast in 1836 and 1839. Hinds was assisted by Barclay and Dr. Sinclair. Their
collections on the coast of Baja California were particularly important. Captain
Edward Belcher's narrative (London, 1843) contains Hind's report on the
"Regions of vegetation ... of the globe in connexion with climate and physical
agents," a rather commonly overlooked essay of considerable interest for the
plant geographer.

The six ships that set sail as the United States Exploring Expedition — our
first Government expedition — under Captain Charles Wilkes on August 18, 1838,
carried six scientists. (There had not been such a concentration since the "Boat-
load of Knowledge" set off down the Ohio for New Harmony!) The six scientists
with Wilkes' Expedition were : Pickering, Brackenridge, Couthouy, J. D. Dana,
Titian Peale, and William Rich. The expedition was surveying the Pacific Coast

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 5

between April 6 and November 1, 1841, and the results were eventually published
after prolonged disaffection between Captain Wilkes on one side and the staff
and authors who prepared the texts of the various departments of science on the
other. That the publication of the results depended on Congressional approval
was no small discouragement. Titian Peale reported on the vertebrate collections
and Torrey and Gray on the plants, with Pickering publishing a remarkable
omnibus volume entitled a Chronological History of Plants: Man's Record of His
Own Existence Illustrated Through Their Names, Uses, and Companionship,
based in some considerable part on his travels with the Wilkes' Expedition.

Captain John Charles Fremont, the "Pathmarker," entered California in 1844
on his first overland expedition. In his diaries he noted trees and items of natural
history — he had been instructed by Dr. John Torrey to take dried plants along
the route — but in the end he did not bring back many specimens, partly owing
to the misfortune of having hard rains ruin his collection. On his expedition of

1846 Fremont paid closer attention to collecting and these specimens were the
subject of a memoir by John Torrey.

Keenly aware of the attractions of California as a potential colony for the
Crown, H.M.S. Herald arrived in Monterey during these days of contested
Spanish rule. But the American chronicler Stillman sums up that story in a
sentence: "Monterey had already fallen into the hands of the Americans, and
she sailed away disgusted." Berthold Seemann, who was later to distinguish
himself in the botany of Fiji and other tropic lands, accompanied the Herald.

Historian John Walton Caughey says, "Take away the initial bonanza of
gold and how much less rapid and how different would the state's rise have
been." James Wilson Marshall's discovery of gold on the American River in

1847 set off "one of the most articulate migrations in history," drawing shiploads
of emigrants from virtually every country of the world. During the year 1849
several visitors with some interest in natural history arrived in California, some
of them members of emigrant parties lured by the activity in the goldfields.

On April 5, 1849, William Gambel, a protege of Nuttall, who had made the
overland trip to California in 1841 via the Gila Route and had returned to Phila-
delphia with 176 species of birds, joined a party of adventurers bound for the
goldfields. The original party divided and Gambel joined those who followed
Hudspeth's trail but they were caught by snow in the mountains and only
Gambel and a few others reached Rose's Bar on the Feather River. Gambel,
sick and exhausted, died of typhoid fever on December 13, 1849. Joseph Grinnell
remarked to this writer that Gambel 's bird skins — the ones taken on the earlier
trip of 1841, the collection of 1849 being lost — were among the best skins he had
ever handled. The ornithologist Cassin described Gambel's skins many years
ago as "some of the most magnificent specimens I ever saw." Witmer Sitone says
that Gambel "in the short space of eight years demonstrated that he was possessed
of remarkable ability both as an explorer and field naturalist and as a student
of natural history."

The New York taxidermist, John Graham Bell, who accompanied Audubon
up the Missouri in 1843, reached California in 1849 via the Central American
isthmian route. He visited Sutter's Mill and localities from Sonoma to San Diego;
considering the short duration of Bell's visit he made a notable collection, taking
the types of four birds described as new by Cassin. Bell himself described the

6 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Pacific Coast towliee. It is interesting to contemplate what might have been the
history of California ornithology had Bell decided to stay in the State rather than
return to New York ! He died at Sparkhill, New York, in October, 1889.

John Woodhouse Audubon, son of the famous ornithologist, came overland
across Texas and northern Mexico, arriving in San Diego, November 4, 1849.
He evidently proceeded to the Sierra diggings directly. The Academy has a
direct connection with John Woodhouse Audubon through the late Leslie Simson,
mining engineer and sportsman who collected specimens of African big game in
Kenya and was also the donor of the California Academy's Simson African Hall.
Simson learned taxidermy as a lad from his father, who in turn had been in-
structed by John W. Audubon.

With Audubon came Dr. John Boardman Trask, cofounder with Dr. David
Wooster of California's first medical journal. He was the first resident naturalist
to describe the State's recent and fossil shells. His work appeared in Volume
One of the Academy's Proceedings. Trask, one of the seven founders of the
Academy in 1853, later became distinguished as physician, chemist, mineralogist,
seismologist, geologist, paleontologist, and botanist.

Particularly versatile was Dr. Jacob Davis Babcock Stillman, perhaps best
known for his association with Senator Leland Stanford, whom he served as per-
sonal physician. Dr. Stillman was a writer of some merit, and his book entitled
Seeking the Golden Fleece (San Francisco, 1877) is highly readable for its per-
sonal approach. He arrived in San Francisco on August 5, 1849, after 194 days'
passage on the ship Pacific; the fare from New York was $300. Upon his
arrival at Sacramento Stillman began collecting plants, ranging as far afield as
Marysville and Long Bar in 1850. Some of this material he sent to John Torrey,
and Asa Gray subsequently based Leptosyne stillmanii on part of it. Stillman was
a classmate friend of Dr. Charles Christopher Parry at Union College, and they
worked together occasionally on the smaller problems of the California flora.
Stillman refers to "my old college friend, Charley Parry, botanist [of the Mex-
ican Boundary Survey]. Charley is now [1877] on the Gila River." Stillman 's
friendship for Parry certainly stood Parry in good stead in securing such favors
as railroad passes for his collecting trips and the like. Within the pages of the
Overlayid MontJily, dear to the heart of the antiquarian, are buried some spark-
ling paragraphs, and not a few were written by naturalists ! One of these stories
is "Old Fuller," a vignette of the Day of Resurrection, written by Dr. Stillman.

The Reverend Augustus Fitch was in southern California between 1846 and
1849 and sent a few plants to John Torrej^ perhaps through the suggestion of
Dr. Parry, but we lack exact knowledge of this fact. There is a note in the
Torrey correspondence of the Reverend Fitch finding Ahronia umbellata at San
Francisco and IMonterey and pointing out its technical characters.

W^illiam Lobb, employee of the large nursery firm of James Veitch, of Exeter,
England, arrived in 1849. He had left England at the age of thirty-one and
collected seeds and plants in South America before his arrival in California,
but his story properly falls a little later in connection with the Big Tree. George
Black collected on the Yuba River in 1850; he may have been associated with
Lobb but I find no evidence that he was employed by a foreign seed house, and
we can only surmise that he may have turned (perhaps unsuccessfully?) from
the mines to work with Lobb in the Sierra foothills.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 7

Much less known than Lobb is Dr. Timothy Langdon Andrews, physician and
botanical collector, who reached San Francisco in November, 1849, and after a
month in the Bay city went to Monterey, the capital of the American colony.
There Andrews opened a school and in his leisure time made a large collection
of plants in the vicinity. In the summer of 1850 he made a two weeks' horseback
trip with William Lobb to the Mission San Antonio de Padua and into the
adjacent Santa Lucia Mountains. It is certainly possible that Dr. Andrews met
Dr. Parry during his stay at IMonterey, but in any event Dr. Andrews made
contact with Torrey and Gray, w^ho studied his collections. Gray named the
endemic tufted Galium of the Coast Ranges for him as a pleasant gesture of one
botanist to another. Later Andrews was an inspector of customs in San Francisco
and a newspaper journalist, and there he met Dr. Albert Kellogg of the Academy,
who must have been delighted w^th Andrews' wide experiences. Both had lived
and traveled in the South before reaching California, Kellogg being a brief resi-
dent of Charleston and Andrews of New Orleans.

In the fall of 1850 two great figures in American science arrived in California
together: James Graham Cooper, the zoologist, and John Lawrence LeConte,
renowned student of beetles and cousin of Professor Joseph LeConte. Dr. Cooper,
son of William Cooper of New York, later became prominent in the history of
the West as an Army surgeon attached first to the Northern Pacific Railway
Survey, then to Mullan's Expedition. Between 1860 and 1862 Cooper was sta-
tioned at Fort Mojave, and from there he explored the almost unknown north
slope of the San Bernardino Mountains. In 1864 he served with the California
Volunteers. After the Civil War came a period as naturalist with the California
State Geological Survey. Brewer, whose judgments wxre generally fair, wrote
of him, upon the occasion of his first meeting in 1861 as "a man of more than
ordinary intellect and zeal in science, but I fear not a very companionable fellow
in camp." Cooper contributed to the text of T. F. Cronise's popular Natural
Wealth of California, published in 1868. From 1875 until his death in 1902 he
lived at Hayward, and his name is commemorated in that of the Cooper Ornitho-
logical Club, now "Society." Cooper was interested in mollusks and general
zoology, ethnology, and kindred subjects, several of which were the topics of
papers contributed to the early volumes of the American Naturalist.

In the early days of California's statehood probably every tenth man was a
Frenchman. This was owing to two reasons : first, the natural attraction of gold
and the untried opportunities in new lands, and, second, the unsettled homeland
conditions of France resulting from the revolutionary movements of 1848 on
the Continent. One of the Frenchmen who left Paris then was Pierre Joseph
Michel Lorquin, pioneer collector of butterflies in California. He said that he
came in 1850 for "the number of new things he would be sure to get" ! Lorquin
traversed much of the State on foot from Plumas County to San Diego, wielding
his net and sending the specimens to J. A. Boisduval, who described 83 butter-
flies and twelve moths from Lorquin's collections. In 1852 Lorquin met Dr. 11. II.
Behr, who later presented Lorquin's duplicate butterfly types to the Academy,
but these were destroj-cd in tlie fire of 1906. The Lorquin's admiral, Basilarchia
lorquini (Boisduval), generally distributed throughout California, is a living
memento of this zealous collector of the 'fifties.

The German physician, Frederick Adolphus Wislizenus, came to America

8 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

in 1835 and is best remembered for his pioneer explorations in Chihuahua. He
visited California in 1851, collecting some plants on the American River. Dr.
Samuel Washington AVoodhouse, surgeon-naturalist Avith Lieutenant Sitgreave's
Zuiii River Expedition of 1851, paused in San Francisco before returning home
via Nicaragua. Woodhouse's article on ornithology in Sitgreave's Report includes
field notes on 219 species of birds. The territory covered is actually greater than
the title of the expedition would suggest, since it covered Indian Territory and
Texas to California.

The Swedish frigate Eugenie paused at San Francisco in 1852 on lier
voyage around the world. Aboard was the botanist, Nils Johan Andersson, then
thirty-one, who later became the most prominent contemporary authority on wil-
lows. Dr. Eric Hulten tells me that Andersson's narrative. En V erldsomsegling
(Stockliolm, 1854), which was based closely on his existing diary, contains a de-
scription of San Francisco (pp. 98-180), his journey to Sacramento, and the
goldfields. But Hulten says Andersson does not record having met any natural-
ists in California.

The year 1852 saw California's maximum gold production: $81,294,700 that
year. San Francisco's part was integral in the State's prosperity and, in the
words of Robert Glass Cleland, the historian, "many cities in the United States
boast a more ancient lineage than that of San Francisco; but none can look back
to a more vigorous, boisterous or interesting youth." From a town of nine hun-
dred souls in the spring of 1848 San Francisco became a bustling market place
where "speculation, open-handedness, startling success or equally swift failure,
hurry, rush and disregard of caution" were characteristics. A decline in business
values set in in 1853, following the boom year in the Mother Lode, but shipping
was on the upswing and approximately five hundred vessels were employed in
the whaling industry by 1855. Ten years later San Francisco was the headquar-
ters for the whale-oil industry. Significant in the cultural sense was Edwin
Booth's playing at the San Francisco Theatre to an appreciative audience.

In the national perspective 1853 saw the beginning of the Pacific Railroad
Surveys under Secretary of War Jefferson Davis. For two years these surveys
reconnoitered so thoroughly and efficiently that the railroad routes of today
were laid out along essentially their original markers. These surveys covered
the five transcontinental routes traversed today from the Northern Pacific Rail-
road to the Southern Pacific Railroad via the Gila Route. Each of the five field
parties included a surgeon-naturalist, who collected objects as opportunity
afforded. The published reports arising from these surveys served as reference
Avorks for the first residents of California, as many well-worn copies of the Pacific
Railway Reports to be seen in second-hand bookshops today will attest. W. P.
Blake was geologist and mineralogist to Williamson's Expedition. "The party
will rendezvous at Benicia" were Lieutenant Williamson's instructions. Blake's
own papers dealt among other topics with Tertiary Infusoria and "observations
on the extent of the gold region." Among other specialists who reported on the
results of the expedition were T. A. Conrad on the fossil shells; A. A. Gould on
recent shells; Louis Agassiz on fossil fishes; and S. F. Baird on mammals. Four
physicians attached to these various surveys, John ]\Iilton Bigelow, Thomas
Antisell, Adolphus L. Ileermann, and John Strong Newberry, all visited San
Francisco during this period and must have been welcome wayfarers for Dr.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 9

Kellogg in the city. Dr. Bigelow's collections were the most extensive for central
California and more than 1,100 collections were enumerated in Volume Four
alone of the Beports. Though Dr. Heermann collected in nearly all fields, he
was particularly interested in birds and birds' eggs. He introduced, in fact, the
word "oology" into ornithological literature. Heermann came to California in
1849, but his activities prior to the Pacific Railway Surveys are unknown. The
beautiful Heermann gull places his name in California skies.

AVhat appears to be wholly sound scientific progress was the subject of satire
by Lieutenant George Horatio Derby, graduate of West Point in the class of
1846, who wrote a book, Phoenixiana or Sketches and Burlesques, under the nom
de plume of John Phoenix (New York, 1903). Derby's burlesque on the surveys
is entitled "Official Report of Professor John Phoenix, A.M., of a Military Survey
and Reconnaissance of the Route from San Francisco to the Mission of Dolores,
made with a view to ascertaining the practicability of connecting these points
by a railroad." In the same volume appears "The San Francisco Antiquarian
Society and California Academy of Arts and Sciences." In this sketch Derby
patently parallels the founding of the Academy, beginning with a committee to
draw up the constitution consisting of "Dr. Keensarvey, A. Cove, and James
Calomel, M.D." Who these characters equate to in real life may test the historic
senses !

Founding of the Academy

When the five doctors, a real estate man, and a school superintendent met
informally on April 4, 1853, to consider organizing an academy to bring together
persons with a collecting urge, or a curiosity to know the singular forms of life
that they noticed were different from those "back home," there could have been
little notion of the expeditions, comprehensive collections, and reference libraries
in the natural sciences that would follow. Though, to speak quite honestly, we
know little about some of the men who met that day, they must have had some-
thing of the spirit of the Salem merchants who, while they spent most of their
time vending staples and making money, always took time to remind their friends,
the sea captains, to watch for big conch shells on the next voyage, a nice perfect
shell of a Galapagos tortoise, or a better tail feather of the Australian lyre bird
than Nicholas Titcomb down the way had just acquired.

Lewis W. Sloat, the real estate man in whose office the "founders" met on
old Montgomery Street, was an amateur conchologist and had a cabinet of shells
in his office. He does not, however, seem to have been in contact with Eastern
naturalists.

Colonel Thomas J. Nevins must certainly have been an idealist, for it was
Nevins who, against considerable opposition, persuaded the Common Council of
San Francisco to establish a free public school system. This was in 1851. After
the first meeting the Academy repaired to Colonel Nevins' office on Clay Street,
and they continued to meet there for many years. It was not until 1874 that the
Academy moved to larger quarters in Dr. Stone's old brick church at California
and Dupont streets. Of two of the five physicians we have little knowledge.
Dr. Andrew Randall was selected chairman of the first meeting, and elected
president of the Academy three successive years. He was shot by a gambler
on .luly 24, 1856, and the murderer was hanged five days later by the Vigilance

10 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Committee. But what may have been Dr. Randall's natural history interest I do
not know. Nor do I know the interests of Dr. Charles Farris, who attended the
first and third meetings of the Academy but left the state in the summer of 1853
and was lost track of. The other three physicians were well known citizens of
the city and left distinguished records. The youngest of the three when the
Academy was founded was Dr. Trask, twenty-nine, then Dr. Kellogg, forty, and
Dr. Gibbons, forty-one.

John Boardman Trask came to California overland with John W. Audubon,
as related before this, and his interests seem to have been perhaps the broadest
of any of the seven founders. It was doubtless to Dr. Trask that each of the
Academy members turned for that sympathetic interest in the individual special
studies that so often isolate members of a scientific society. Perhaps Trask's
particular interest was that of the potential use of native plants for medicinal
purposes. E. E. C. Stearns, who knew him as a close friend, spoke of Trask's
"genial qualities, untiring energy and all-around ability" and said that he was
"the leader, closely followed by Dr. Albert Kellogg." Complementing the gentle-
ness of Kellogg, Trask's calm assurance in the face of difficulties must have been
a staying power in the survival of the Academy during its insecure years. John
Xantus, when in San Francisco on his way to Lower California for birds for
Baird and plants for Gray, wrote to Baird at Washington that "Dr. Trask is
particularly kind to me, and so is Dr. Ayres, who both told me to consider their
houses as my own, and command their services no matter how."

Dr. Henry Gibbons, the first of four generations of physicians, was particu-
larly interested in meteorology and kept weather records of such accuracy that
the Smithsonian Institution was happy to publish them.

Naturalists in California After 1853

Born in New Hartford, Connecticut, educated in medicine at Charleston,
South Carolina, and Transylvania College, Lexington, Kentucky, Albert Kellogg
came to California in 1849 and evidently first engaged in business. He had
practiced in the South but those who knew him say he was never known to
request a payment. Never blessed with a strong constitution, Dr. Kellogg re-
turned to his New England home and soon joined a party bound for California
by way of the Horn. He arrived at Sacramento on August 8, 1849. The plant
collections he had made along the west coast of South America at ports of call
were destroyed in a flood at Sacramento soon after his arrival. He was associated
in Sacramento with the Connecticut Mining and Trading Company, but removed
to San Francisco about the year of the Academy's founding and established a
pharmacy business there with some medical practice on the side. He entered into
the spirit of the Academy from its very inception, and seems to have especially
stimulated the members and visitors to the city to communicate specimens to the
Academy for study and identification. One of the most prominent of these par-
ticipants was Dr. John A. Veatch, of whom we shall have more to tell directly.
Dr. Kellogg's personal botanizing began in earnest in the summer of 1867 when
he accompanied Professor George Davidson of the United States Coast Survey
and W. G. W. Harford to Alaska. Several hundred species were collected in
triplicate, one specimen going to the National Herbarium at Washington, one to

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 1 ]

the Philadelphia Academy, and one remaining in the growing collection of the
Academy. George Davidson described this Alaskan trip thus :

We lived in the same contracted temporary deck cabin for four or five months
under many trials and inconveniences, and the sweetness of [Kellogg's] character was as
pervading and refreshing as the beauty and fragrance of the flowers he gathered. . . .
He was completely absorbed in his duties; he knew no cessation to the labor of col-
lection and preservation; his genial nature attracted assistance from every one, and
all learned to admire and to love him.

Davidson continues :

[Kellogg] worked for the [Academy] and believed in its success when the number
of members could have been counted on one's fingers, and when the means of sup-
porting such an institution and publishing its results came wholly from their pro-
fessional earnings.

From 1867 to 1870 Dr. Kellogg visited localities from Donner and Cisco to Ukiah,
Red IMountain, Cahto, and Santa Cruz Island. Scwne of his local trips recall the
days when the geography of California was quite different from today: "Lobos
Creek, near San Francisco"! These collections often, though not always, carried
collection numbers but a new series was evidently initiated every year. His last
decade was pretty constantly spent drawing trees and shrubs. More than four
hundred of these drawings "including all the oaks, all the coniferous trees,
poj^lars, many of the willows and ceanothi, dogwoods, and many herbaceous
species" were left with his friends. Dr. W. P. Gibbons and Mr. Harford, to be
disposed of as they might think best. The oak drawings were published with
commentary by Professor E. L. Greene as West American Oaks, under a sub-
vention from Captain James Monroe McDonald, 1825-1907, pioneer capitalist
and philanthropist. Captain McDonald was one of the three donors of the Rick-
secker Collection of Coleoptera to the University of California in 1881. Kellogg's
drawings showed "the very faithfulness of detail with the taste of an artist,"
yet "the botanist may rely upon the scrupulous exactness of every minute line
and dot." Kellogg would not have claimed the rank of scientific botanist but
rather a nature lover in the true and full sense. Kellogg lived in the early years
at San Francisco with Harford in a small place on Telegraph Hill where they
kept "batchelor's hall." He never married and died at the home of his very dear
friend Harford in Oakland in 1887. William H. Brewer tersely summarized his
role when he wrote, "no name is more intimately associated with the botany of
the state during this period" than Kellogg's.

John Allen Veatch was one of those early collectors whose specimens engaged
Kellogg's attention. Veatch lived in Texas from 1836 until 1845, during which
years he had met the enthusiastic botanical collector, Charles Wright. Veatch
left a wife and five children in Texas to join the Gold Rush, and when his wife
Ann failed to hear from her husband as the months stretched into years she filed
a petition for divorce on the grounds of continued abandonment. It is not certain
just when Veatch first got in touch with the Academy but in 1855 he was elected
a corresponding member and he later served as Curator of Conchology. During
these years Dr. Veatch — for he had certified for practice in the custom of those
days — traveled from Red Bluff to the Salton Sea, where he carefully inspected
the mud volcanoes and wrote his observations. In 1858 Veatch was on Cedros
Island [written "Cerros Island" in contemporary accounts], where lie was pre-
ceded only by the surgeon aboard H.M.S. Herald, Mr. J. Goodridge. Veatch "s

12 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

collections were by far the most extensive yet made on the island, though often
scrappy specimens by our standards, and Dr. Kellogg published his discoveries
in the San Francisco weekly The Hesperian, illustrating many of his novelties
with drawings. Kellogg's poetic soul is laid bare in the vernacular names that
he gave the new species. One of Veatch's plants appeared, for example, as the
"hummingbird's dinner horn." Kellogg's scientific names were not infrequently
hyphenated words of curious construction that some botanists felt obliged later
to edit or disregard altogether.

Though not a "founder" in the strict sense of being present at the meeting
of April 4, Dr. H. H. Behr joined the Academy on February 4, 1854, to launch
a lifetime of service to the young organization. Dr. Behr was thirty-six when
he joined the Academy; he was born at Colthen, Duchy of Anhalt, Germany,
and took his medical degree in Berlin in 1843. His coming to the feverish San
Francisco of 1850 was the outcome of his participation in the Revolution of 1848.
In temperament, then, Behr easily adjusted to the rough manners of the frontier
city, and took up practice at once. But he allowed plenty of time to collect plants
and these he sent to Hamburg, St. Petersburg, and elsewhere. Fortunately Dr.
Behr has narrated his experiences of these early years in an article entitled
"Botanical Reminiscences of San Francisco" {Erythea, 4:168-173, 1896). Behr's
copy of Endlicher's Genera plantarum was the chief resource for the study of the
troublesome specimens that were brought to the Academy at this time. He taught
classes at the California College of Pharmacy and prepared his Flora of San
Francisco, a rare book today, for the use of the pupils. But Behr's interests were
much broader than botany alone. He wrote poetry, humor, and travelogues — ^his
account of two years spent in the Philippine Islands appeared in the Atlantic
Monthly. His writings were warmly acclaimed in Germany. It is natural that
his spiritual link was with Alexander von Humboldt, Schlechtendahl, Ferdinand
von Mueller, Hillebrand, Louis Agassiz, and Max Miiller. Those who came to
San Francisco from afar were sure to find Dr. Behr a hearty host, and it would
be difficult to know how important was his influence in the lives of the many
scientists and others that he chanced to meet. A man of good will and generous
spirit, he died at the age of eighty-six at his home at 1215 Bush Street, in the
city with which he had been identified for fifty-four years.

Dr. William Peters Gibbons had taken his ^I.D. degree in 1846 and sailed
from New York in 1852 for California via Panama. While crossing the Isthmus
he fell a victim to cholera and would likely have perished there, had not W. C.
Ralston carried him in his arms aboard the vessel bound for San Francisco. This
is the Ralston who later directed the Bank of California, was a steamship owner,
and enterprising capitalist. Dr. Gibbons arrived in San Francisco in January,
1853, and at once began to practice medicine in the city. Quite certainly he met
Dr. Behr early that year, as well as Dr. Kellogg. He became active, not only in
the Academy, but in the California State Medical Society as well, serving as
chairman of the committee on medical botany and as a contributor to its Trans-
actions. He was particularly interested in fishes and J. G. Cooper named the
genus Gihhonsia in his honor. Dr. Gibbons was the son of William Gibbons
(1781-1845), Quaker physician and friend of the Pennsylvania botanist. Dr.
William Darlington. W. P. Gibbons collected plants in California at least as
late as 1874, as represented by sheets in the Torrey Herbarium. He mentions

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 13

makiiio; an herbarium on one occasion but whetluM' tliis fell to the Academy and
in 1906 to destruction I do not know. Prom 186)} until his death at the age of
eighty-five Gibbons was a resident of Alameda. We will quote from his writings
later in our chronicle when he considers the State Geological Survey.

Hiram G. Bloomer first set out for California in 1849 but had to turn back
on reaching Panama because of sickness; he tried again, successfully, in 1850.
I have no information on his principal occupation but he was devoted to botany
from the first of his California residence, and participated in the life of San
Francisco, serving as a member of the Committee of Vigilance and of the Fire
Department. He was active, too, in the Lincoln presidential campaign. He was
generous in presenting books to the Academy's library in its early years. It is
important to recognize that Bloomer introduced James Lick, the philanthropist,
to the needs of the Academy. It will be remembered that the Academy built new
quarters on Market between Fourth and Fifth streets in 1891 upon property
deeded to it by James Lick. Lick also made the Academy one of two residuary
legatees, to receive one half of his estate after all other bequests had been paid.
Bloomer's botanical interests centered around the Liliaceae, and he grew many of
the native species in his garden. Kellogg named a flower found by Dr. Veatch
at New Idria Bloomeria. in Bloomer's honor. Bloomer's herbarium of several
thousand sheets was evidently lost soon after its presentation to the Academy
but duplicates had been sent to Asa Gray and others during the State Survey
period.

William G. W. Harford was one of those Academy members who could be
expected at every meeting. "Six feet in height, of a Lincolnian gauntness, with a
pioneer style of luxuriant beard and bushy eyebrows," he was even more shy
and retiring than his friend, Kellogg. Like Kellogg, he was of a simple manner,
of a deeply religious nature, and devoted to the beautiful. Concliology was per-
haps Harford's special interest, and he served as the Academy's curator in that
field in 1867, 1868, 1874, and 1875. He was Director of the Academy from 1876
to 1886. Spiders and beetles also interested Harford, along with botany. He
and Kellogg made up sets of Oregon and California plant collections in 1868
and 1869 and these reached the herbaria of Europe, as well as the herbaria of
Englemann, Torrey, and Gray. Greene and Parry dedicated the polygonaceous
genus. HorforcUa, to his memory in 1886. He was a close associate of George
Davidson, with whom he traveled to Alaska in 1867 as naturalist on the United
States Coast Survey. Like so many of his cronies at the Academy, Harford
lived to be an octogenarian.

Colonel Leander Ransom was an engineer before he came to California by
sea in 1852. He was then fifty-three years of age, and had served the previous
thirteen years as President of the Public Works of Ohio. He was sent to Cali-
fornia by the Federal Government to establish a United States Surveyor Gen-
eral's office in San Francisco and, finding the city to his liking, he became a per-
manent resident. Always interested in geography and land forms, he is remem-
bered for establishing two of the most important meridian lines on the North
American continent, the Mount Diablo base and meridian lines, on July 17, 1851.
For many years Colonel Ransom served as the Academy's president, and Dr.
Kellogg remembered him in the name of a native oak, but Quercus Ransomi is
hard to find todav even in the svnonjnnies of the oaks!

14 A CENTURY Of PROGRESS IN THE NATURAL SCIENCES

Three botanical explorers, Archibald Menzies, David Douglas, and John
Jeffrey, were born only a few miles apart, in the county of Perth, in England.
The last of the trio, John Jeffrey, collected plants and seeds in northern California
and Oregon during 1852-1853, sponsored by the "Oregon Committee" of Edin-
burgh, which had raised money by subscription for what is generally called the
■'Oregon Expedition." Each member was to receive a portion of the seeds col-
lected. Jeffrey was chosen and contracted to keep a diary on the trip, but no seeds
ever reached Scotland. Of perhaps ten boxes of seeds and specimens sent, five
reached England but they contained relatively few herbarium collections. Jeffrey
botanized in the Salmon River Mountains and on the south slope of Mount Shasta,
and reached San Francisco on October 7. He was ill in San Francisco that
winter, and did not write his sponsors in Edinburgh or even call for his mail at
the British Consulate. Mr. William Murray, of Henderland, who was in San
Francisco during the fall of 1853, and Andrew Murray, brother of the secretary
of the Committee, could not locate Jeffrey in the city. Jeffrey, perhaps through
a friend, dispatched a final small box of tree seeds early in January of 1854.
Sometime in the spring of that year Jeffrey is said to have left with an American
party for Yuma, with the intention of collecting on the Colorado Desert. He
was never heard from again and only conjectures surround his death. "Bearing
in mind that Menzies and Douglas went to a virgin country, [Jeffrey's] collec-
tions [after them] do him no discredit, even as compared with theirs."

Jeffrey's unfinished work was carried on by William Murray, accompanied
by A. F. Beardsley, "a gentleman from whose energy and knowledge of the
mode of life in the regions they traversed, he derived much assistance." They
collected conifers, so much in demand in British gardens, in the Sierra Nevada,
including Pinus Beardshyi, later considered a synonym of Pi7ius ponderosa.
Beardsley visited the Santa Lucia range in 1856 for seeds of Abies venusta,
which had been recently introduced into England by William Lobb. But evi-
dently neither Murray nor Beardsley were employees of Peter Lawson and
Company, Scottish seedsmen. William IT. Brewer, wlio reenters our chronicle
later, met Beardsley in October, 1861, at a tavern in Napa Valley whence Brewer,
then with the State Geological Survey, had repaired "to read the news." Brewer
says:

While there, a rough but intelligent looking man entered into conversation and
invited me to his house a few rods distant for a "glass of good cider." I went, got the
cider, the best I have tasted in the state, and went into his house. I found him an
intelligent man, quite a botanist, and even found that he had some rare and expensive
illustrated botanical works, such as Silva Americana, worth sixty to eighty dollars — the
last place in the world I would have looked for such works. He does not own the ranch,
is merely a hired man. having charge! There is an oichard of ten or twelve thousand
trees and a vineyard — he makes wine and cider and sells fruit.

Brewer returned the next day for more cider :

Mr. Beardsley came to camp and invited us to his house for more cider. We went,
spent an hour, when it cleared up, and we started for a peak seven or eight miles
northeast.

Just as Douglas and Jeffrey collected seeds and plants in California for
English horticulture, William Lobb spent seventeen years with the nursery firm
of James Veitch of Exeter, going first to South America to collect orchids and
new plants for the "stoves." Lobb reached San Francisco in the hectic summer

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 15

of 1849 but he turned from the lure of the bonanza road to complete immediate
plans for the exploration of southern California! His first season included a trip
into the Santa Lucia Mountains, whence he was able to introduce the bristle-cone
fir successfully into England. During the spring of 1850 Lobb was joined by
Dr. C. C. Parry, then sojourning in Monterey, on a trip south at least as far as
Mission San Antonio de Padua. The 1851 season he spent north of San Francisco,
and in the following year he reached the Columbia River, collecting all the while.
Perhaps it was during the winter of 1852-1853 that he learned of the fabulous
"Big Tree" through the testimony of a hunter, Mr. Dowd by name. In any event,
Lobb set off directly for the Calaveras Grove early in 1853 and, finding the trees
and collecting the foliage, cones, and seeds, hastened back to England as the
scientific herald of the greatest tree on earth. The apogee of Lobb's career came
perhaps, not in California, where he was hardly known, but at Sydenham at the
exposition put on in 1857 in the Crystal Palace ! There a section of a Big Tree
was exhibited, standing 116 feet high — as high as the bark had been stripped from
a living tree — in all its majesty, bearing the name Wellingtonia which had been
given it in December, 1853, by England's excellent botanist, Professor John
Lindley. Some saw in it proof again that Britain was still the general in the
vanguard of discovery, with Wellingtonia her latest conquest! It was called the
"Mammoth Tree," and public interest ran high on both sides of the Atlantic,
although Americans were not a little piqued at the "scoop"! But history takes
some sharp and unexpected turns. A decade later William Lobb was lowered
into an unmarked grave in the Public Lot at Laurel Hill cemetery, deserted and
forgotten, a victim of paralysis at fifty-five.^ If we are to believe Parry's report,
Dr. Kellogg thought that Lobb took unwarranted license with the information
that he had wrested from Mr. Dowd. But though William Lobb did first make
known the Big Tree in a formal way, the American name. Sequoia, has found a
secure place in our literature and language.^

Julius Froebel and H. H. Behr were both "Forty-eighters," that is, members
of the "group of German idealists who fought to establish a liberal and unified
Germany and then came to the United States as refugees from the reaction."
Froebel had founded a radical opposition newspaper, the Siviss Republican, in
1839, and subsequently participated in the 1848 Revolution. He was arrested,
condemned to death, pardoned, and returned to Switzerland, but he left for
America and arrived in New York in 1849. In all, Froebel made four different
trips to Central America and the Southwest. It was toward the close of his third
trip that he visited San Francisco in the fall of 1854, arriving by coastwise boat
from San Pedro. He wrote :

On the morning of October 3rd, we entered the Golden Gate. Much had I heard of
the grand scenery of the Bay of San Francisco, and I can only state that reality sur-
passed my expectations. . . . Whatever splendid sites of cities other parts of the world
may have to boast of, in North America the palm will never be disputed to San
Francisco.

Froebel comments further:

Every European, many Asiatic, and some American languages, meet the ear while

2. Lobb's grave was moved and appropriately marked years later by San Fran-
ciscan garden lovers under the aegis of Miss Eastwood.

3. Buchholz's segregate genus Sequoiadendron for the Sierran tree as distinct from
the coastal redwood happily carries on the historic connotation.

16 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

you are walking in the streets. This apparent chaos of heterogeneous elements has
been brought together, and is kept in motion, under the great form and system of
Americanism, with its restless labour, its ever-active spirit of speculation, and its de-
votion to utilitarian purposes.

His two-volume narrative Aus Amerika. Erfahrungen, Reisen und Studien
(1857-1858) was abridged as Seven Years' Travel in Central America, Northern
Mexico, and the Far West of the United States (1859). He contributed an article
on the physical geography of North America, dated "San Francisco, Dec. 8, 1854"
to the Ninth Annual Report of the Smithsonian Institution (1855).

Emanuel Samuels was sent to California jointly by the Smithsonian Institu-
tion, the Boston Society of Natural History, and Academy of Natural Sciences
of Philadelphia to collect birds. He arrived in 1855 and most of his collections
were made in the vicinity of Petaluma. What relation, if any, Emanuel Samuels
may have borne to "Rev. Mr. Samuels" mentioned by Sereno Watson when he
described Chorizanthe valida collected at the Russian Colony in Sonoma County
I have not been able to determine.

General Amos Beebe Eaton, the father of the distinguished Professor of
Botany at Yale, Daniel Cady Eaton, collected a few ferns about Carquinez Strait
in 1855.

January 27, 1855, saw the completion of the Panama Railroad from Panama
City on the Pacific to Navy Bay, or Aspinwall, on the Atlantic. Its construction
had employed in all some seven thousand men drawn from all over the world,
some from the mines of California a few years before. Daily service was estab-
lished both ways, the fare for adults being set at $25. The running time at first
was from five to six hours but was later cut to three hours, with as many as
fifteen hundred passengers carried in a single half-day. And, you will be right
when you predict : most of the passengers were en route to California !

Coming by boat from across the Pacific, Ezechiel Jules Remy, French natu-
ralist and explorer, traveled under the nominal auspices of the Natural History
Museum of Paris. Remy had been collecting in the Hawaiian Islands intermit-
tently between 1851 and 1855 before he arrived in San Francisco in the summer
accompanied by the Reverend Julius Brenchley. Brenchley will be remembered
for his placing a plaque at the site of David Douglas' grave on the island of
Hawaii. Remy and Brenchley left San Francisco on July 18, 1855, for Salt Lake
City via Carson Valley. From their extended visit in the Mormon city they pub-
lished an illustrated two-volume account of the geographic and social features
of the communit3^ Leaving on October 26 Remy traversed the Great Basin to
St. George and went on to Las Vegas and Los Angeles, which he reached Novem-
ber 29. Returning to San Francisco, Remy took passage for Central America.
Parry refers briefly to Remy's few plant collections reaching the Natural History
Museum at Paris.

Thomas Bridges, British naturalist and horticultural collector, a Fellow of
the Linnaean and Zoological societies of London, had been in South America
before coming to San Francisco in November, 1856. There is substantial evidence
that he was an enthusiastic collector and he proved to be California's first resi-
dent ornithologist. One obituary noted that "few, if any, more useful lives have
passed away as martyrs to science during the present century." Bridges' prin-
cipal field of collecting was the Sierra Nevada. There he collected seventy-five

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 17

bulbs of the lily, Lilium ivashingtonianum, for his English employer but the
steamer Central America, which carried them, was lost at sea. lie wrote W. -I.
Hooker that he was going to make an effort to replace them. Evidently he visited
the Academy often, and in 1858 he wrote Hooker of his pleasure at finding
Beechey's Voyage, Torrey's works, and other books in the Academy's library.
He lived in "Chinese House" on Eleventh Street between Market and Mission
streets, and may have associated with William Lobb, then a resident of the city,
but of that friendship we have no hint. One of Bridges' most profitable trips
w^as to the mining town of Silver Mountain on the east slope of the Sierra Nevada
near Ebbetts Pass in 1863. There he met William II. Brewer and Brewer wrote :

It was a relief to meet Mr. Bridges, an old rambler and botanical collector, well
known to all botanists. ... It was a relief to meet him and talk botany; yet, even
he is affected — he has dropped botany and is here speculating in mines. "Mining
fever" is a terrible epidemic; when it is really in a community, lucky is the man
who is not affected by it. Yet a feiv become immensely rich.

In April, 1865, Bridges set out on a collecting trip to Nicaragua but was stricken
with malaria and died at sea, September 9, 1865, en route back to San Francisco
on the steamer Moses Taylor. Captan Blethen, Bridges' friend, brought his
body to San Francisco and he was carried to the ultima thule of the city. Lone
Mountain Cemetery.

One of the most colorful figures in the history of California's progress in
science was Andrew Jackson Grayson. Born at the Grayson cotton plantation
on the Ouachita River in northern Louisiana, August 20, 1819, he traveled widely,
won and lost, and died three days short of his fiftieth birthday at the Mexican
port of San Bias. Grayson made the overland trip from Independence, Missouri,
in 1846, with his young wife and child, and reached California in October. The
Donner party traveled with them as far as Fort Bridger, when the emigrants
separated, the Donner party pushing on to tragic death, the Graysons to some
considerable fortune in the "diggins," followed by a less fortunate venture into
the mercantile business. Finally Grayson tried his hand at trapping, and it was
during this period, when he occasionally visited the Mercantile Library in San
Francisco, that he chanced upon Audubon's Birds of America. He was so deeply
thrilled with the paintings that he determined to match them for the birds of the
Pacific slope. So ardently did he adopt Audubon's flamboyant style, sketching
the birds in stiff or unnatural postures, that he quite aptly may be called the
"Audubon of the Pacific." Grayson also gave his bird portraits backgrounds of
quite accurate, if occasionally mixed, delineations of the native plants. From
1855 to 1857 Grayson made sketches of the birds about San Jose and the Napa
Valley, and in 1857 sailed for Tehuantepec on the Mary Taylor. But his plan
to include the Mexican fauna in his opus was dealt a blow by the wreck of the
schooner in the bay of Ventosa, when his books, drawings, paper stock, and colors
were ruined. Penniless, he took up a job as surveyor to recover his funds, but
he found drawing paper impossible to procure and he turned to the preparation
of bird skins. Some of these reached S. F. Baird, who was most enthusiastic about
them. After a visit to San Francisco, Grayson returned to Mexico in company
with J. M. Hutchings, of "Yo-Semite Valley" fame, determined to settle at
Mazatlan and sketch the local birds for his book. During this period he wrote
travel articles for the Overland 3Ionthly and the press. John Xantus was his

18 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

correspondent at Cape San Lucas. A hearing was effected with Emperor
Maximilian and Empress Carlotta but the collapse of their regime brought
an early end to Grayson's support for a projected Birds of Western 3Iexico.
It was while on an expedition to Isabel Islands for nesting sea birds that he
was taken sick with the "coast fever." The journal Condor has been currently
publishing his beautiful drawings of Mexican birds. Grayson's notes for many
of these will be found in Bryant's article published in Zoe for April, 1891.

Robert Edward Carter Stearns came to San Francisco in 1858 at the age
of thirty-one to become a partner in the large printing establishment of his
brother-in-law. This firm published the influential Pacific Methodist and, in
the absence of the editor, Stearns took over. This journal was instrumental in
keeping California in the Union during the Civil War. Always interested in
zoology, Stearns made a trip to Florida in 1863 for invertebrate collections
for the Smithsonian Institution. In the Proceedings of the Academy for 1868
Stearns treated the mollusks of Bolinas Bay. The University of California
made important advances under President Gilman, and during this period
Stearns served as secretary to the University, beginning in 1874. He launched
a plan for developing the plantings on the campus in 1882 which was carried
forward by Professor Greene when he came in 1885. In turn Stearns was
U. S. Fish Commissioner, paleontologist under John Wesley Powell, and assist-
ant curator of mollusks under S. F. Baird at the Smithsonian. Stearns often
contributed articles on marine life to Charles Russell Orcutt's West American
Scieiitist, as well as to Brandegee's Zoe. Through the years away from Cali-
fornia Stearns kept in touch with his friends Trask, Kellogg, Harford, Dr.
Wesley Newcomb, and others at the Academy.

Particularly interesting was Dr. Newcomb 's cabinet of shells. Josiah
Whitney remarked in a letter to his brother Will on June 2, 1862, that he had
examined Newcomb's "superb collection of shells — one of the best in the coun-
try, especially in the department of land shells. He has in all between 10,000
and 11,000 species." Stearns and Newcomb were brought into close friendship
by their common interest in conchology and it was a bitter loss to Stearns on
his return to California in 1892, to learn of Newcomb's death. Newcomb had
been a practicing physician in the Hawaiian Islands for five years and had
become an authority on the land shells of the islands.

It was in 1859 that Dr. Veatch set out for Cedros Island to verify the
rumors of mineral wealth there. Whalers, seal hunters, and fishermen visited
Sebastian Viscaino Bay and brought out wealth in furs and oil, but few
persons paid much attention to the volcanic soil itself. Since there was a high
point on the island which might yield plants characteristic of northern lati-
tudes. Dr. Veatch was eager to examine its flora. He brought back only about
two dozen specimens for Dr. Kellogg to study, but they proved almost with-
out exception to be undescribed! Of course one of them became Veatchia!

In 1859 Louis Agassiz' son, Alexander Agassiz, twenty-four, came to San
Francisco to take a position with the Coast Survey a§ engineer to survey the
Gulf of Georgia and was assigned to the Fauntleroy. Returning to the city,
Agassiz applied himself to the medusae and viviparous "perch" (Embiotocidae)
of San Francisco harbor, making drawings and notes for his father. Alexander
Agassiz later invested over a million dollars, made in the Calumet and Hecla

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 19

Copper Mine on Lake Superior, in Harvard's Museum of Comparative Zool-
ogy, which his father had founded. "The Bismarck of American Science,"
"fearless, resolute, quick to anger, definitely purposeful and full of resource,"
Alexander proved a "colossal leader of great enterprises, fully as much as he
was a man of science."

The California State Legislature created the office of State Geologist and
authorized a geological survey of the State on April 21, 1860. Josiah Whitney
was selected as State Geologist and William Henry Brewer, Botanist. Rather
later, J. G. Cooper was prominent as a zoologist. William More Gabb joined the
Survey in 1862 as paleontologist, and was described in Brewer's words as "young,
grassy green, but decidedly smart and well posted in his department." Thus
just seven years to the month came the second organized institution for the pro-
motion of natural sciences on the Pacific Coast. It was fortunate, too, that
Whitney and Brewer were destined to work together on this survey for they
proved a well matched team.

Whitney was forty-one when he took over the leadership of California's geo-
logical survey. Schooled at the Round Hill School, founded at Northampton,
Massachusetts, by George Bancroft and J. G. Cogswell, and subsequently at
Yale under Benjamin Silliman, whose chemistry lectures excited him, AVhitney
managed the Iowa Geological Survey before taking over the California job. The
State Survey proceeded well enough at first, but met with little sympathy from
the legislature after it failed to lead a waning mining industry to a new bonanza
at home and halt the loss of men to the Pikes Peak gold rush. But Whitney was
thorough in his prosecution of the Survey and by the end of the first year of his
work he had already visited iovty of the then forty-six counties of the State.
Brewer, his first assistant, had traveled 2,600 miles on muleback, a thousand more
on foot. The age of the auriferous gravels had been determined as Jurassic; the
coal of the Coast Ranges, Cretaceous; about two hundred species of fossils had
been discovered and a "great many new animals and plants." In the personal
sense Whitney was less the State Geologist to his scientific associates "than the
gay Apothecarius of Clover Den. He was kindly, just, unsparing of himself;
and his associates gave him not merely esteem but affection." Dr. Trask turned
over his geology notes and fossil collection for the use of the Survey but Brewer
found Blake "distinctly less friendly." Whitney was influential in the life of the
Academy and in matters of publications was ever a driver for accuracy and thor-
oughness. In a letter to his brother, William Dwight Whitney, he reported :

... of late I have been much engaged with the the affairs of the California Acad-
emy, as we have had to move into and fit up new rooms [this was January, 1867], and
have tried to resuscitate in general. We seem now to be in a fair way to live; but
when I came back last year, it seemed as if it was as dead as a doornail. We have
now a pleasant reading room with a goodly number of scientific periodicals; and we
are fitting up our meeting room and collections in a respectable manner. The last
sheets of the Proceedings . . . will tell you what we have been doing, and you will
notice my account of the [Calaveras] skull, etc.

But the State Survey issued only three of its final reports, the other volumes being
published through outside resources, including Whitney's personal funds. Brewer
brought out the botany volume by means of a $5,000 private subscription, "engi-
neered by Judge S. C. Hastings of San Francisco and helped on by Gilman,

20 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Leland Stanford, and D. 0. Mills," with "AVhitney's help." But it cost Brewer
"two years' unpaid labor, $2,000 of [my] pocket, and the accompanying loss of
[my] salary at Yale." The botany volume sold to the public for four dollars. The
three volumes on birds were printed largely at the expense of Alexander Agassiz,
with Baird contributing a thousand dollars on his portion. The geological ma-
terials were published largely through the "M.C.Z." of Cambridge: "the gravel
volume will form one of the Memoirs of the Zoological Museum."

Whitney stayed on with the State Survey until 1874, the next year taking
the Sturgis-Hooper Professorship of Geology at Harvard, which he held until
his death in 1896. "Honors did not come to him as abundantly as to many per-
haps less worthj^," concludes the historian of geology, G. P. Merrill. Some strong-
worded opposition to the State Survey came even from scientists. Dr. "William
P. Gibbons wrote in the Overland Monthly:

... as to any report on botany, or any collection of California plants, three sets
have been made up: one for the California Academy of Sciences; one for the University
of California; while one has been sent out of the State, and eastern botanists have
the credit of devoting their time to working it up, in occasional paroxysms, without
remuneration. It would have been far better for the interests of the State and of
science had this [California Geological] commission never existed.

Dr. Gibbons evinced more local pride than imagination when he said :

California scientists would have accomplished more work, without aid from the
State, than has thus far, to all practical purposes, been achieved by the commission.

Gibbons' assessment appeared in August, 1875. The first volume of the "Botany
Report" was published the following year, and the second volume, in a neces-
sarily smaller edition, four years later. Kellogg, Bolander, Behr, and perhaps
a few others, might have eventually described the greater part of the California
flora, but the number of avoidable synonyms may well have increased thereby
because of the inability of the resident botanists to check against the existing
specimens in Eastern herbaria.

Thlrty-two-year-old AYilliam Henry Brewer accompanied Whitney and his
family from Massachusetts to California via Aspinwall. When the party stepped
ashore from the Golden Age on November 14, 1860, they were greeted by Mr.
S. Osgood Putnam, of the California Steam Navigation Company, who had backed
the State Survey appropriation in the legislature. Brewer had finished at the
Sheffield Scientific School at Yale in 1852 — a member of its first class — and had
studied abroad under the chemists Liebig and Bunsen. Along the academic way
he had acquired a lively taste for botany and a near dead-shot judgment in geol-
ogy. He had applied for a post on Captain Gunnison's expedition but had been
turned down; Gunnison and his party, it will be remembered, were massacred
by a band of Indians in Utah. Brewer was "fond of travel, not for rest, but for
the recreation which he found in careful observation and record of facts in all
departments of human interest." No botanical collector in California up to his
time made as careful field tickets as did Brewer; fortunately, too, his field book
is preserved at the Gray Herbarium. His journal, edited by F. P. Farquhar and
first published in 1930 under the title Up and Down California in 1860-1864, is a
rich but unscheduled dividend of the State Survey !

William More Gabb of Philadelphia was the same age as Brewer when he
joined the State Survey but there the likeness breaks, for hardly could two men

EVVAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 21

have been more contrasting: where Brewer was modest, Gabb was bumptious;
Brewer was resilient in the face of inevitable adjustment, Gabb, reluctant. Gabb
came as an acknowledged authority on Cretaceous fossils. He is described as a
"distinctly loquacious person." Brewer was pleased when the serious, unbending
Dr. J. G. Cooper saw fit to name a new species of brachiopod Lingula gahhii! A
close friend of Gabb's in Philadelphia was George II. Horn, the entomologist,
who came to California the next year.

Dr. George Henry Horn came to Camp Independence in Owens Valley in
1862, as a member of the Survey, after graduating from "Penn" the year before.
But the doctor soon turned from medicine to beetles, a field in which he became
a recognized authority. While in California Dr. Horn collected actively about
Fort Tejon, Fort Yuma, Surprise Valley, Warner's Ranch, and many other
localities. He occasionally made plant collections, particularly in the Owens
Vallc}^, and these may be found cited in the "Botany Report" of the Survey. The
year 1862 brought the establishment of the Department of Entomology at the
Academy, with Dr. Behr as Curator. He served first for five years and then a
second term from 1881 until 1904.

A little known figure of this period was Dr. Charles Austin Stivers, U. S.
Army, who interested himself in collecting plants about the post in Mariposa
County. He brought his specimens to Dr. Kellogg and among them was the
remarkable endemic lupine which bears his name today. There is a record of
Dr. Stivers' interest in marine algae, too.

The Prussian expedition to East Asia in 1860-1862 had as its geologist
and geographer Freiherr Ferdinand Paul Wilhelm von Richthofen. When the
expedition set out on its return voyage to Germany from China in 1862, Baron
Richthofen parted from the corps and sailed for San Francisco. He arrived in
California, "a modest, sincere, affectionate" man about thirty years old, intent
on studying volcanic phenomena. Having some private means, he worked only
intermittently for the State Survey in those fields that appealed to him. But for
Whitney he had a "worshipful admiration," and the two geologists fitted as neatly
as pick-head and tool handle. It was Whitney w^ho conceived the idea of a
geological survey for China and, indeed, the China survey was planned by the
tw^o men on New Year's Eve of 1868. During the subsequent years in China
Richthofen wrote long letters to Whitney, which Whitney edited and transmitted
to the American Academy of Arts and Letters at Boston for publication. Richt-
hofen evidently made some botanical collections in California, but it is difficult
to discover the extent or the destiny of them. He returned to Germany after
twelve years of travel to teach first at Berlin, then at Bonn, Leipzig, and finally
again at Berlin. From the clues I have seen the as yet unwritten biography of
"Life and Times of Baron Richthofen" could be a warm and gracious tale.

Behr's friend, Dr. William Hillebrand, went to the Hawaiian Islands in 1844
for his health, stayed twenty-eight years and identified himself as the leading
authority on the flora of the islands. He visited California in 1863 and made
some collections about the Yosemite Valley, Big Tree grove, and Mount Dana,
as a part of the State Survey.

Brewer mentions William Holden's collecting about a hundred species of
plants in the vicinity of Oakland in 1863. These were included in the State
Survey, but Holden evidently did not continue his scientific interests.

22 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

English-born Shakespearean tragedian, Henry Edwards, traveled with a
theatrical company from Australia to Peru and California in 1853, and wrote
of his impressions in a slender volume called A Mingled Yarn (1883). In 1865
Edwards came back to San Francisco and was associated wdth the old California
Theatre. During all these j^ears on the stage, traveling and as a San Franciscan,
he collected butterflies at every opportunity. His collection grew by his own
takes and through exchanges until it was one of the finest ever assembled in this
country, numbering some 250,000 specimens. In addition, Edwards found time
to collect beetles, plants, and shells for his friends in the Academy, of which he
was a faithful associate. There and at the Bohemian Club he found a congenial
friend in Dr. Belir. Edwards made plant collections at Sausalito, in March, 1877:
Summit, on the Central Pacific Railroad, July, 1877; Knights Valley and Skaggs
Springs, Sonoma County, in 1877, and in Santa Clara Valley — all of these are
represented in the liarbarium of the New York Botanical Garden. There's a hint
of the actor in his locality on one label "San Leander"!

John Torrey, the senior associate of Asa Gray in midcentury botany, visited
California on two occasions. His trip of 1865, made via the Isthmian passage,
included a short stay in San Francisco, but he took the Revenue steamer Shuhrick
for Santa Barbara on business for the U. S. Treasury as inspector of banks.
Writing in his usual buoyant mood, he told Asa Gray that he was "high admiral
of the expedition." He made sure to save some time from the inspection of ledgers
and balances to devote to the plants growing around the towns visited : from
Borax Lake and vicinity to Donner Lake, Bear Mountain, and the Yosemite.

One of the collectors well known to Torrey and Gray for his valued specimens
was Dr. Charles Lewis Anderson, who moved from Minneapolis in 1862 to Carson
City, Nevada, and four years later to Santa Cruz. At Santa Cruz his name became
synonymous with natural history since there for forty years Dr. Anderson
engaged, not in botanical, zoological, and geological investigations for himself,
but generously answered various questions for others. In botany he devoted him-
self especially to marine algae about the bay, to grasses in the hills and valleys
of the county, and the willow species along the stream courses.

Edward Tuckerman was a genial, if meticulous, professor at Amherst, and
one of the students there in the 1850's was George Lincoln Goodale. Goodale took
the medical degree at both Harvard and Bowdoin, and then set up practice at
Portland, Maine. From all of this close application his health broke and the year
1865 found him in California trying to find a cure in tramping the hills and
collecting the plants about which Professor Tuckerman had talked back at
Amherst. The cure must have been complete, and more and more botany sup-
planted medicine until he settled as Curator of the Botanical Museum at Harvard
and for thirty years taught and studied the economic plant collections that came
to him. He is remembered as one of the first professors to use lantern slides to
illustrate his lectures. Goodale possessed a fine historical sense, too, and pre-
served mementoes of our botanical past for Harvard's "glory hole," as Thomas
Barbour would say.

Less honored but perhaps more influential was the author of the botany best
seller that sold 800,000 copies, Professor Alphonso Wood. First a student of
theology, then a practicing civil engineer, a teacher of Latin and natural history
in the Kimball Union Academy near Hanover. Alphonso Wood found it difficult

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 23

to teach botany to students from the existing texts prepared by Professor Asa
Gray and ventured to discuss the matter with him. Wood suggested that Gray
prepare a text better suited to the secondary schools, but the titular head of
botany in this country denied the need was a valid one. Professor Wood ap-
proached Gray a second time but was again refused, whereujjon he set out to
prepare a "Class book of Botany" of his own. The first edition of Wood's Clnss-
hook appeared in 1845 in an edition of 1,500 copies and met with some consider-
able success. Those faithful to Gray disjiaragod Wood's intrusion on Gray's
established precincts, bolstering their opposition chiefly with the premise that
Professor Wood was ill-trained and had an inadequate background to undertake
the text. But the Class-hook was accepted more and more widely among the
academies and Wood kept pace with the trend by widening the scope of the
text with each new printing until in 1855— only ten years after its first publica-
tion — forty-one such "editions" had been issued! With ambition reminiscent of
that other challenging professor, Amos Eaton, Alphonso Wood determined to
extend his book to include the growing frontiers of America. So he made field
trips to Ohio and into the southern states, and in 1865-1866 to the Pacific Coast.
It is unfortunate that the details of his Western journey have not survived;
suffice to say that he traveled from San Diego to Oregon. Plagued with poor
health, limited funds, and the general insecurity attendant on the Civil War, he
found it difficult to make headway in his chosen field, but he devoted his last
years to botany from the year of his settling at West Farms, New York. The
student of California history would like to know more of the association of
Alphonso Wood with the person he commemorated in the naming of the endemic
mariposa of San Diego County, Calochortus Weedii. In the tradition of William
Young, w^lio contested the field with John and William Bartram in the early
years of the nation, and John Linnaeus Shecut, who nettled Stephan Elliott in
the description of the botany of the Carolinas, Alphonso Wood stood against
Asa Gray, not so much as a serious challenge to the supremacy of the leaders
but to remind us of the impossil^ility of establishing a monopoly in knowledge.
John Gill Lemmon was an ardent Abolitionist and, as in all the events of his
lifetime, turned a loyalty into action and enlisted in the Union Army. But he
was taken prisoner iand placed in the largest and best, if infamously known, of the
Confederate military prisons, at Andersonville, Georgia. It was a log stockade
of sixteen and a half acres holding within its pickets 31,678 prisoners in the
summer of 1864. Corn meal and beans, with a little meat, was the diet; respira-
tory diseases, diarrhoea, and scurvy were rami)ant in the ranks. John Lemmon "s
health was broken but he escaped interment with the 12,912 men left in the
National Cemetery there. He went to California as soon as possible, first to Sierra
Valley in 1866, and from eight years of tramping the meadows and slopes in pine-
scented air regained his health. ]\Ieanwhile, he discovered a world of plant life
about him and early in his Sierran residence sent some of his specimens to Asa
Gray for their names. With Gray's encouraging letters he continued the search,
and paused now and then to write homespun letters to the local newspapers on
plant lore. It was a high point when in 1876 he met Asa Gray personally. By
1880 Lemmon was devotedly wedded to botany and so it was with a kind of
bigamy prevalent among naturalists that he married Sara Allen Plummer of
Santa Barbara. They moved to Oakland and set up a botanical establishment at

24 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

5985 Telegraph Avenue, more or less completely filling a large frame house with
their plants, books, and the kind of enthusiasm that effectively combats the
paralysis of poverty, which followed them from the beginning. The house was
easily identified by a large wooden sign bearing in capital letters the words "Lem-
mon Herbarium." Taking every travel opportunity that presented itself the
couple managed to reach distant points in Arizona, the Mount Shasta region, and
the San Bernardino Mountains of southern California, taking specimens in sets
for sale and exchange. With near idolatrous devotion they sent the first specimen
to Professor Gray, and as soon as he responded with the identifications they so
eagerly awaited they distributed the duplicates, printed up circulars, and sent
off scripts to the press of wild potatoes, resurrection ferns, and outsize records
of California trees. From 1888 to 1892 Lemmon was botanist to the California
State Board of Forestry and during this period published Pines of the Pacific
Slope and Cone-hearers of California. This last duodecimo handbook was a kind
of forerunner of the popular pocket guides of today. The Lemmon collection,
rich in isotypes and early records, ultimately came to the University of California
but the transcription of the data, written hastily on the margins of the news-
papers, suffered somewhat in the curating process. It is unfortunate that the
specimens lacked original labels bearing Lemmon 's own record of the data for
some facts may be learned from a comparative study of the labels made at dif-
ferent times in his lifetime. "J. G. Lemmon and wife" (as the labels read in the
older herbaria, bearing witness to a marital warmth that they shared in adversity)
were self-sacrificing bearers in the caravan of botanical discovery.

Three women who lived in northeastern California and were enthusiastically
interested in plant study were Rebecca Merritt Austin, her daughter, Mrs. C. C.
Bruce, and Mary E. Pulsifer Ames. Better known to Asa Gray and Eastern
botanists than to most at the California Academy, their plant collections and
field notes gave the foundation to our knowledge of the vegetation of that region.
"R. M. Austin," as she labeled her collections, came with her husband and children
to the gold mines of Black Hawk Creek of Plumas County in 1865. There she
began collecting plants and other objects of natural history with no thought of
the particular value of her "hobby." Early in 1872 John Gill Lemmon, while
peddling books in the mining towns of Sierra Valley, visited her. We may imagine
Lemmon showed Mrs. Austin Hittell's Resources of California, Scott's Wedge of
Gold, and perhaps Mrs. Clarke's Teaching of the Ages, but it would be sport to
know if he took orders for Bret Harte's Luck and Stoddard's South Sea Idylls.
But we do know that Lemmon was exultant when he saw Mrs. Austin's specimens
displayed in a "cabinet" made from a soap box. Jepson says that "those who
knew the exuberant Lemmon will readily credit the story as related by Mrs.
Austin" that "he took off his hat and gave three cheers for the woman who was
cooking for miners and at the same time trying to study nature under such
adverse circumstances." The Austins removed in 1875 to Butterfl}^ Valley and
there she carried out her studies on the pitcher plant Darlingtonia known to the
local residents as the "cobra plant." Mrs. Austin observed that the amount of
fluid increased in the pitchers when they were stimulated by the introduction of
bits of meat. One of her earliest correspondents was William M. Canby, of Wil-
mington, Delaware, to whom she wrote no less than twenty letters on the Dar-
Ungto7iia studies. She was also in touch with C. Keck, an Austrian botanist and

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 25

dealer in natural history specimens, who reported her recent findings in the
organ of the Austrian botanical society. About this time Asa Gray mentioned
Mrs. Austin's observations in his book Barwiniana, and she published a short
note in Coulter's Botanical Gazette for 1878. That year the Austins moved to
Big Meadows, Plumas County, where Canby paid her a visit. But in 1881 they
moved again, this time to Modoc County, where she made some of the first collec-
tions for the county, alone or in company with her daughter. The pages of
Pittonia carry frequent mention of the Austin and Bruce collections, some sin •
gled out for such recognition as Scutellaria austinae and Collinsia hruceae.

Comparatively little is known of Mary E. Pulsifer Ames of Auburn, whose
plant collections, like those of Mrs. Austin, are occasionally cited in the Botany
of California, particularly the second volume. She was evidently at one time a
resident of Taylorsville, Indian Valley, a correspondent of C. Keck of Austria,
as Avas Mrs. Austin, and a contributor to the California Horticulturist and Floral
Magazine. Astragalus pulsiferae of Plumas County was named in her memory
by Asa Gray. She died at San Jose, at the age of fifty-seven.

Gulian Pickering Rixford, the son of a scythe-maker, born in East Highgate,
Vermont, came to San Francisco in 1867. Rixford's real interest was evidently
horticulture and applied entomology, but he worked as a journalist "to pay ex-
penses." For eight years he was on the editorial staff of the Evening Bulletin and
its business manager for thirteen years. An ingenious plan to finance the intro-
duction of the Smyrna fig from Asia Minor was put up to the proprietor of the
Bulletin. Cuttings w^ere to be distributed to three thousand subscribers to the
paper as a sort of premium, and gratis to nurserymen and fruit growers. Seventy
thousand cuttings were distributed in 1880 by this device. In April, 1892, Rix-
ford made incidental collections of some interest in Owens Valley of Inyo
County, including Eremolithia Rixforclii, named by Brandegee. In 1913 Rixford
was chosen Director of the Academy and in 1930 awarded the Frank N. Meyer
Medal for distinguished services in plant introduction.

English-born Richard Harper Stretch, engineer and entomologist, visited
America first in 1861 and finally settled in California in 1867. Educated in
Quaker schools abroad and apprenticed to a draper, he became enthusiastic about
natural history as a boy. He joined the Academy as a resident member the year
he came to California and devoted his time particularly to moths and their
taxonomy. Fine drawings of moths executed by him were published in 1874, and
later his collection of about five thousand specimens was given to the University
of California. Stretch was a close friend of Dr. Behr and of Henry Edwards,
following whose death he lost interest in entomology and devoted his time more
wholly to engineering. Stretch was the first to call attention in official circles to
the presence of the cottony cushion scale in California. He spent his later years
in the Puget Sound region.

Of Henry Nicholson Bolander, Asa Gray wrote in 1868 that "for the last few
years no one has done so much as Mr. Bolander for developing the botany of
his adopted State, and perhaps no one is likely to do so much hereafter." At that
time he dedicated the pretty genus Bolandra of the Saxifrage family to him.
Bolander came to Columbus, Ohio, at the age of fifteen, from Schleuchtern, near
Frankfort, Germany, his birthplace. In Columbus he came under the influence
of Leo Lesquereux, the bryologist, and from this early contact persisted a life-long

26 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

interest in mosses. Bolander arrived in San Francisco December 5, 1861, to find
the State Survey staff assembled in the city. Dr. Kellogg, and other members of
the Academy, became his intimate friends. It is singular that there is not a single
mention of Bolander in Brewer's letters, at least in so far as edited by Farquhar.
Bolander became State Botanist at the close of the State Survey late in 1864, on
the resignation of Brewer. Between 1864 and 1873 Bolander botanized over
nearly all parts of the State, his ramblings being exceeded perhaps only by those
of Brewer himself : from Ukiah and Red Mountain to Mount Dana, Mono Lake,
and south to Cuyamaca Mountains and San Felipe Caiion. Bolander's most seri-
ous interest was in grasses, about which he wrote briefly in the Academy's Pro-
ceedings. Lesquereux wrote in 1869 that Bolander had in less than one year
collected as many species of mosses as all the other collectors together. The San
Francisco publishing firm of Anton Roman and Company published Bolander's
small quarto volume in 1870 entitled Catalogue of the Plants Orowing in the
Vicinity of San Francisco, Embracing the Flora within 100 Miles of the City.
Between 1871 and 1875 he served as State Superintendent of Schools, and during
this period his botanical activities began to wane. Plis plant collections were
well known in Europe, De Candolle reporting the herbarium at Geneva as con-
taining 1,156 species of his gathering, and his specimens were also received at
Kew and Leipzig. His death occurred at Portland, Oregon, August 28, 1897, by
which time his name had quite disappeared from current botanical literature.

On the morning of October 21, 1868, a destructive earthquake shook the city
of San Francisco. As Bret Harte remarked, "Enough that w^e know that for the
space of forty seconds — some say more — two or three hundred thousand people,
dwelling on the Pacific slope, stood in momentary fear of sudden and mysterious
death." Bret Harte chastises the citizens for trying to hide the seriousness of
the earthquake lest the reports have an unfavorable eft'ect on tourist interest in
the city, and adds :

It is surprising liow little we know of the earth we inhabit. Perhaps hereafter
we in California will be more respectful of the calm men of science who studied the
physique of our country without immediate reference to its mineralogical value. We
may yet regret that we snubbed the State Geological Survey because it was impractical.

The earthquake and its economic reverberations threatened the Academy's income
at this time, and it was Stearns and Whitney, in particular, who stood behind its
survival.

Though not realized at the time, an important stimulus to the promotion of
the natural sciences in California at this time was the formal charter granted
the University of California on March 23, with Henry Durant installed as its
first President. Practically from the beginning the University worked along
with the Academy across the Bay in many matters of mutual scientific interest.

An obscure visitor to Califoi-nia at this time was Heinrieh Sylvester Theodor
Tiling, from Livonia, a physician at the hospital at Sitka, who collected at
Unalaska in 1851 and at Sitka between 1866 and 1868. He visited Nevada City
about 1869 and collected the type there of Horkelia Tilingi described by Regel.
Tiling died in 1871 and it seems fairly certain that the visit of Benedict Roezl
to America in 1872 was a follow-up of Tiling's brief visit.

Samuel Brannan, Jr., accompanied Dr. Kellogg on his trips botanizing
in the Sierra Nevada in 1869 and 1870. Brannan collected insects as well,

fWAN: SAN FRANC/SCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 27

and the agaristid moth, AndroJoma bninnani, was named for him ])y Stretch.

The year 1869 was a critical one in California history, for it brought the
completion of the transcontinental railroad. "Sir: we have the honor to report
that the last rail is laid, the last spike is driven. The Pacific Railroad is finished"
read the telegram sent from Promontorj^ Point, Utah, to President Grant, on
May 10, 1869. It was not long before there set in a growing feeling against the
large land holdings under the monopolistic control of the few wealthy men or
corporations, such as the very group that had won the railroad triumph. "Out
of three drops of rain which fall in the San Joaquin Valley, two are owned by
Collis P. Huntington." The big strikes of the early years of the Gold Rush were
stories now, the whale oil industry began its steady decline. New industries came
with the advent of the railroad. Fruit culture was soon the first agricultural
interest of the State. This period of economic transition, like the earthquake of
1868 and its consequences, brought financial restrictions on the Academy.

The newly chartered University of California began classes on September 20
at its old Oakland campus — it was not until 1873 that the move "five miles to
the north to the site christened Berkeley" was made — and a man with scientific
traditions, John LeConte, served as its third president jrro tern. His brother,
Joseph LeConte, arrived that month to lecture on geology, zoology, and botany;
he re-enters our narrative again very soon.

"When I set out on the long excursion that finally led to California I wandered
afoot and alone, from Indiana to the Gulf of Mexico, with a plant-press on my back,
holding a generally southward course, like the birds when they are going from
summer to winter.

So wrote John Muir. After a near-fatal siege of fever in Florida and a short stay
in Cuba, Muir arrived in San Francisco by way of the Panama steamer. He
soon set out on foot for the Yosemite. My First Summer in the Sierra was his
diary of 1869. For the next six years Muir — "the wiry young man with auburn
hair, full beard, and electric blue eyes had one trait that outweighed all other
elements in his nature, the trait of persistence" — absorbed the geology, zoology,
and botany of the region and became in turn guide for geologist Joseph LeConte,
lepidopterist Henry Edwards, and, in 1872, botanist John Torrey on his second
visit to California. Muir wrote "Harry" Edwards under date of June 6, 1872 :

Your bundle of butterfly apparatus is received. You are now in constant remem-
brance, because every flying flower is branded with your name. I shall be among the
high gardens in a month or two and will gather you a good handful of your favorite
painted honeysuckers and honeysuckles. I wish you all the deep far-reaching joy you
deserve in your dear sunful pursuits.

On February 22, 1873, Muir wrote Asa Gray :

Our winter is very glorious. January was a block of solid sun-gold, not the thin
frosty kind, but of a quality that called forth butterflies and tingled the fern coils
and filled the noontide with dreamy hum of insect wings.

Eventually Muir moved down to the big city to write up his Sierra experiences,

which appeared first in such journals as the Overland Monthly.

Some of my grandfathers must have been born on a muirland, for there is heather
in me. and tinctures of bog juices, that send me to Cassiope, and oozing through all
my veins impel me unhaltingly through endless glacier meadows, seemingly the deeper
and danker the better.

28 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

In the summer of 1870 Joseph LeConte, Professor Frank Soule, Jr., and eight
students camped in the Sierras for six weeks. LeConte said, "I never enjoyed
anything else so much in my life — perfect health, the merry party of young men,
the glorious scenery, and, above all, the magnificent opportunity for studying
mountain origin and structure." This summer's foray was the theme of his
Journal of Ramhlmgs through the High Sierras of California by the University
Excursion Party, published in 1875.

The third of the trilogy of mountain essays was Clarence King's Mountaineer-
ing in the Sierra Nevada, published in 1871. Clarence King, Sheffield Scientific
graduate, was twenty years old when, almost providentially, he met Brewer, a
Sheffield alumnus, on the steamer plying between Sacramento and San Francisco
on August 31, 1863. King w^as traveling with his college chum James Terry
Gardiner, and in a letter to his mother Gardiner described Brewer in these words :

. . . nothing peculiar about him, yet liis face impressed me. . . . the roughest
dressed person on the steamboat [with] an old felt hat, a quick eye, a sunburned face
with different lines from the other mountaineers, a long weather-beaten neck pro-
truding from a coarse grey flannel shirt and a rough coat, a heavy revolver belt, and
long legs, made up the man; and yet he is an intellectual man — I know it.

Three days after meeting Brewer, Clarence King was made an assistant geologist
of the State Geological Survey. He lived to climb many of the highest peaks of
the Sierra Nevada ahead of others, but King "was an amateur, not a scientific
climber, and he delighted in thrills." By his thirtieth birthday he was in charge
of the Fortieth Parallel Survey, and soon afterwards he became the first director
of the U. S. Geological Survey.

Louis Agassiz visited San Francisco in 1872 en route home from Brazil by
way of Cape Horn aboard the Hassler. Agassiz visited Joseph LeConte in
Oakland on this trip.

During September (or October ?), 1872, Benedict Eoezl, native of Bohemia,
passed through the city on a plant-collecting foray for European horticultural
firms, en route from Panama via Acapulco. The details of Roezl's visit, which
must have been brief, as Tiling's was before him, are confused in the few accounts
in the literature. The beautiful dull-red flowered gooseberry of middle elevations
in the Sierra Nevada, Rihes roezlii, was named for him by the botanist Kegel.

Gustavus Augustus Eisen, born in Stockholm, Sweden, came to tlie United
States in October, 1872, after taking his Doctor of Philosophy degree at L'psala
earlier that year. He apparently headed for California, for he soon settled at
Fresno, then a pioneer community. Eisen 's most important work was in horti-
culture. By lectures and pamphleteering he fostered the introduction of the
Smyrna fig and avocado into the State. He joined the Academy in 1874 and
served as curator from 1892 to 1900. From time to time he collected plants in
Fresno County; for example, Phacelia eisenii, named by Brandegee. Eisen must
be credited, too, for his part in the creation of Sequoia National Park by execu-
tive decree. Mount Eisen, elevation 12,000 feet, in the Park, perpetuates his
name. Dr. Eisen led Academy expeditions — apparently the first under the Acad-
emy's sponsorship — to Lower California in 1892, 1893, and 1894. During those
years his interests included helminthology, archaeology, and geology, in addition
to botany.

In the 1870's one of the leading taxidermists in San Francisco was Saxon-born

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 29

Ferdinand Gruber, the Academy's one-time Curator of Birds. He assisted in the
arrangement of the collection of mounted birds at Woodward's Gardens, one of
the city's earliest and much-beloved pleasure resorts. The statues and urns that
once graced the Gardens may be seen today at Sutro Heights. Gruber invented
a rotating tableau of natural history called the "Zoogeographicon," exhibited at
the Gardens from 1874 to 1889. Xantus called Gruber "a very excellent taxi-
dermist, and [a man] who sells at a very high figure his birds for drawing room
ornaments . . . Mr. Gruber is a very honest man, but a very strict commerciante
also." It was Gruber who collected birds on the Farallones for Xantus in exchange
for skins from Cape San Jose del Cabo, and hereby hangs a tale. Xantus, whose
veracity seems to have eroded pretty far on other occasions as well, wished to
swell the collection of Cape birds to be sent to Spencer F. Baird at the Smith-
sonian, so he took Farallon skins of Tufted Puffin and Pigeon guillemot collected
by Gruber and attached labels reading "Sandoval point, 1860" and "Cape Los
Martires, 1861" to tliem. These are birds not otherwise known from Lower Cali-
fornia and when Joseph Grinnell was preparing his Distributional Summation
of the Ornithology of Lower California he remarked, without knowing of the
switch perpetrated by Xantus or, indeed, of Xantus' exchange contacts with
Gruber, that the skins showed a remarkable resemblance to Gruber's well kno^vn
specimens! There are still unsolved problems of this nature, as witness the hawk
Onychotes gruheri. It is supposed to have a California origin but is now regarded
as a later name for an Hawaiian hawk. Gruber was in touch with Dr. Frick,
French Consul General in Honolulu — can this be a clue to the mystery of
Onychotes gruheri f

Dr. Kellogg found a sympathetic colleague in Dr. Arthur Wellesley Saxe, who
came to California in 1850 and worked in the mines until 1852. In 1854 he took
up residence as practicing physician at Santa Clara, where he lived until his death
in 1891, with one visit to the Hawaiian Islands to study leprosy. Dr. Howard A.
Kelly says he was President of the California Horticultural Society and had "one
of the largest collections of roses and rare bulbs in the state." Dr. Kellogg named
Rumex saxei for his friend in 1879, and Professor Greene named Clarkia saxeana
in 1887, but Saxe's collections at the Academy, which were perhaps never exten-
sive, were lost in the fire of 1906.

A close friend of Harford at the Academy was George Washington Dunn,
who came to California in 1850 and worked in the placer mines. Along with many
another miner Dunn left the placers penniless, whereupon he determined to
devote his life to professional collecting, which seems to have been his first love
all along. Taking up residence in San Diego, he ranged far and wide for speci-
mens to sell. He was described as "a genial sort, always on his uppers, who col-
lected insects, plants, shells, and anything else he could sell. Like IMicawber, he
waited for something to 'turn up'." An acquaintance relates how he would climb a
couple of hundred feet up pine trees when he was past eighty, and put lengths of
stove pipe on his legs when collecting in rattlesnake-infested areas. He was elected
a resident member of the Academy March 16, 1874, and it was at this time that
Dunn, along with Harford and some other Academy members, organized the in-
formal Arthrozoic Club. He was admitted into the San Francisco almshouse in
his ninety-first year but left of his own accord four months later and died the
following year. In all, lie made twelve trips to Lower California, including one

30 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

in 1885 to Guadalupe and Cedros islands with Professor E. L. Greene, and several
to Cantillas Canyon, which he was the first naturalist to explore, once with
Edward Palmer.

Associated with Harford and Dunn in the Arthrozoic Club was James John
Rivers, a broadly trained Eng-lish biologist and an acquaintance of T. H. Huxley,
Charles Darwin, A. R. Wallace, and others. He came to this country in 1867 and
arrived in California between 1875 and 1880, having made the friendship of Pro-
fessor Francis Huntington Snow, of the University of Kansas, in the interim.
Rivers was Curator of Organic Natural History at the University of California
from 1881 to 1895, when he removed to Santa Monica. His biological interests
included insects, shells, spiders, and reptiles, as well as botany.

It was during late February or March in 1874 that the Reverend Edward
Lee Greene first came to California from Colorado. An enthusiastic field collector,
his coming rather initiated a botanical revival. In Colorado his duties as Episco-
palian rector were light and he had filled his days with botany. "But my new
parish at Vallejo is too much for me," he wrote Ludwig Kumlein back in Wis-
consin. "I have a large congregation and good salary, but with all that, so much
pastoral work, that my scientific studies are interfered with not a little." Napa
Valley in the spring ! — it must have set Greene's botanical senses atingle. Always
aware of the importance of the written record against which discoveries must be
checked, he repaired to the Academy across the Bay and conferred with Dr.
Kellogg. Greene stayed at Vallejo about a year, then returned to Colorado in
1875. He filled the pulpit at Georgetown until March, 1876, then returned to
California, this time to Yreka. Along with his shepherding, he found time to
botani/e on the Humbug Plills that first year and in other directions away from
town. On January 21, 1877, he set off for New Mexico and another charge at
Silver City, taking his time along the way to collect plants. For the next few
years he explored the mountains of western New Mexico and in 1882 returned
to California as pastor of St. Mark's Episcopal Church on Bancroft Way in
Berkeley. From this time forward Greene took an intense interest in the Cali-
fornia flora, and it is agreed that his best work was done with that subject. He
spent much of his time at the Academy both while he was at St. Mark's and after
becoming the first Professor of Botany at the University of California. It was
during this period that he founded the botanical journal Pittonia. He continued
his field work in California and in Lower California, and from his own and the
collections of others described hundreds of new species. The pages of the Acad-
emy's BuIJetm bear witness to his driving capacity for work. The appearance of
the Botany of California posed a challenge for Greene and some other resident
botanists like him to extend the boundaries of our knowledge. Greene's coming
to the University as Professor of Botany initiated a program of local exploration
into the more remote parts of the State by his students and correspondents. Some
of these will be briefly noticed at a later point in our chronicle.

The Centennial Exposition of 1876 called for nation-wide exhibits, including
forestry and horticulture, and George Richard Vasey, son of the Washington
agrostologist. Dr. George Vasey, came to California for w^ood exhibits in 1875.
He also made general collections of vascular plants as far north as Mendocino
County, but his labels liave caused some serious confusion from a lack of careful
localitv data.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 31

The Russian diplomat, Carl Robert Osten Sackcn, visited California first in
1875-1876 as a private citizen interested in collecting Diptera. Previous to this
he had served as Secretary of the Russian Legation and Consul General of Russia
in New York City.

In many ways he constituted the beau ideal of a scientific entomologist: absolute
master of numerous languages, independence of means, social rank, retentive memory,
accurate observation, possessor of an almost perfect library of works upon Dipterology,
and polished manners — these qualities all combined enabled him to hold the highest
rank in his special branch of science.

LjTuan Belding, "the Nestor of California ornithologists," knew the passenger
pigeon in Pennsylvania's Wyoming Valley, and after he came to Stockton in 1854
the elk of the tule marshes and beaver and otter about the valley town were
familiar sights to him. In 1862 Belding moved to Marysville, but it was not until
the publication of Cooper's OrnitJioIogy of California in 1876 that his interest
took a serious turn. He was no doubt encouraged by S. F. Baird and Robert
Ridgway, who guided his collecting energies. They suggested that Belding make
a trip to Guadalupe Island in the spring of 1881, but this was abandoned in favor
of a visit to Cedros Island. Belding made several trips to Lower California; he
made especially notable collections about Cape San Jose del Cabo, where, to his
wonderment, Xantus had missed certain common birds. But the high Sierras of
central California drew his closest scrutiny, for neither Heermann, Gambel, nor
Xantus explored them and Bell may well not have reached much above the foot-
hills. Belding's 274-page account of Birds of the Pacific District was published
in 1890 by the Academy. He sent several papers to the West American Scien-
tist and to Zoe.

The lepidopterist, William Greenwood Wright, author of the Butterflies of
the West Coast (San Francisco, 1905) — a rare book because of the destruction
of the warehouse stocks in the fire of 1906 — v^as a well-known figure about the
Academy. Henry Edwards, Dr. Behr, R. H. Stretch, and others at the Academy,
as well as Dr. Parry, who botanized in Wright's territory about San Bernardino,
were all his friends. He was a largel}^ self-educated man, who came to California
shortly after the Civil War. For twenty years he operated a planing mill at San
Bernardino, devoting his leisure to collecting insects, especially butterflies, and
plants. George II. Horn characterized Wright as "a zealous botanist, for whom
neither the privations incident to an exploration of the Mojave Desert nor the
jealous watchfulness of the Indians, seemed to have held any terrors." In June,
1888, he botanized in the Greenhorn Mountains; in January, 1889, about the
Mexican port of San Bias; at Sitka, Alaska, in July, 1891; and in Mendocino
County, in May, 1894. His later years were passed at San Bernardino, where he
was a familiar figure because of his natural history interests and his fondness
for instructing children in the subject, and there he died in 1912, at the age of
eighty-three.

Charles Christopher Parry is well known as a botanical explorer of Colorado,
and before that as a member ol; the ^lexican Boundary Survey, but he also made
several botanical visits to California. Sargent has remarked on the zeal, industry,
and intelligence with which he botanized for a period of more than forty-eight
years in the West. The winter of 1880-1881 Dr. Parry spent in and around San
Francisco, with nominal headquarters at the Academy. Returning in the spring

32 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

of 1883, he spent that season on excursions both to the north and south of the
city. During those years he was able to secure a pass on the Southern Pacific
Railroad through the offices of his good friend Dr. Stillman, Leland Stanford's
personal physician. He stayed ten months during 1886-1887 investigating the
genera Ceanothus, Arctostaphylos, and Alnus. Though Parry did not write any
manuals or even extensive revisions of genera, aside from the synopses of the
ceanothi, ehorizanthes, and manzanitas, he wrote a fair stream of chatty articles
to the local newspapers, as well as to his home-town Davenport Gazette. Some
of these sketches demonstrate a fine command of English and a poetic quality
not often found in such ephemera. He was fond, too, of writing terse messages
to his botanical cronies, Englemann, Gray, and to Canby, Eedfield, ^nd just about
all the contemporary botanical figures of the day, for Parry was friendly and
communicative. Typical of these short letters is the following to Samuel Bonsall
Parish of San Bernardino, here quoted in part :

Since leaving your dry region for pastures green, I have been able to see some
things that may be of interest to you — at least you deserve an attempt to make them
so. Among other things I made a short trip to lone in Amador Co to look up an
anomalous Arctostaphylos collected in leaf only by Mrs Curran last year — I found it
on her directions abundant and in full flower Feby 1st of which I secured plenty
of specimens — (one of course for you). On subsequent examination I conclude that
it is a good n sp — nearly allied to A. nummularia — but abundantly distinct. To which
I gave the provisional name A. myrtifolia n sp. I shall wait to get mature fruit be-
fore publishing, and will probably offer it for publication in Cal Acad Bull — when I
shall try & tell the whole story.

Another thing that may interest you is an investigation I have been making of
our Pacific Coast Alnus. ... So you see there is plenty to be done in studying common
things — Greene is busy in his revisions is now at Boraginaceae Dr. Gray I hear has com-
menced printing Polypetalae. now in Papaveraceae. Will accept most of Greene's Escholt-
zias [sic.], quite a triumph for Greene. Acad [em] y affairs as you will infer are run a la
Curran and nobody else has anything to say in the matter — Greene draws off to Berkeley —
how long this state of things may last qiiien sabe. I enclose Harkness's inaugural
written as I understand by Curran. Let us hear from you. Mrs. P joins in regards
to yourself & Mrs. Parish.

Dr. Parry's last visit to California was made in the spring of 1889. For forty
years he was a "familiar figure to hunters, prospectors, mountaineers, and all
sorts of outdoor people, from the Arizona deserts to the Siskiyou pine forests."
Sargent remarked that "no other botanist of his generation . . . revealed so many
undescribed North American plants."

During the decade of 1875-1885, with its delays in the publication of the
Academy's Proceedings, internal dissensions raked the organization. Joseph
LeConte said:

It might be supposed that the Academy of Sciences was an important element
in my career [in California] but not so. It had little effect in determining my scien-
tific activity. I read many papers there, to be sure, and several of them were pub-
lished in their Proceedings, but I always reserved the right to publish them elsewhere
also.

He remarked further that

. . . under the presidency of J. D. Whitney the Academy was prosperous and held
a high position among the scientific institutions of our country; but from that time,
"because of internal dissensions, it dropped lower and lower.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 33

The "internal dissensions" of wliieh LeConte speaks were compounded of petty
jealousies and institutional politics. Jepson contended tliat these dissensions
were "engineered" by Mrs. Mary K. Curran. Harford served as Director of the
Museum from 1876 to 1886, but he "resigned" in altercation. The able Professor
George Davidson was replaced as President by Dr. H. W. Harkness. It is clear
from Setchell's biography of Mary Katharine Layne Curran Brandegee that he
admired her generous qualities and judged her actions disinterested. Professor
Jepson, on the contrary, looked upon her activities as scheming and vindictive.
In the professional sense Mrs. Brandegee showed penetrating insight in
botanical judgment, as abundantly demonstrated in reviews she prepared for the
journal Zoe. Though she recorded only the briefest data on her collection labels
— as if she intended to stymie another collector revisiting her station! — she
made excellent series of specimens illustrating the ecologic variations to be found
within a species. She joined the Academy about 1880, after taking her M.D.
degree two years before at the University of California, and began studying
botany under Dr. Behr. As Mary K. Curran, a widow, without heavy financial
obligations, she was able to devote her time and resources to the Academy's
Department of Botany fully, and she was made Curator of the Herbarium in
1883. There is no doubt but that she did important spade work for the herbar-
ium, which she described as "in a shocking condition" when she assumed the
curatorship. She also became acting Editor of the Academy's Bulletin. Katharine
Layne 's second marriage was felicitous for botany, as for the couple. Marcus
Jones remarked to me on one occasion, "Brandegee should have been born a
woman and Mrs. Brandegee should have been a man. So their marriage could
hardly help being a success!"

Townshend Stith Brandegee came into the Academy's orbit soon after his first
visit to California in 1886-1887. It was the winter he came to collect tree trunks
for the Jesup collection of woods at the American Museum of Natural History,
A student of Daniel Cady Eaton in botany at Yale, where he graduated in engi-
neering, Brandegee went as a young man to Colorado to carry on surveying. He
took the opportunity to botanize widely over southern Colorado, as his surveying
duties took him to remote districts, and what is more important he had the
acumen to recognize the value of his discoveries and to communicate them to
Eastern botanists who were in the best position to assist him, Brandegee's self-
effacing reticence won him warm friendship from Asa Gray, C. S. Sargent, and
others, though his increasing deafness isolated him more and more after he came
to live in California. From 1884 to 1890 Mr, Brandegee visited several of the
Santa Barbara Islands, one of the most ambitious trips being that to Santa Cruz
and Santa Eosa islands in 1888, In 1889 the Academy sent its Curator of Birds,
Walter Pierce Bryant, and an assistant, Charles Haines, to Magdalena Bay, and
Brandegee joined the party at his own expense, collecting a large series of plants
in Lower California that season. It was following this first trip to Lower Cali-
fornia that the Brandegees were married, on May 29 in San Diego, after which
they set out on foot overland to San Francisco on a botanical honeymoon! For
five years thereafter the Brandegees made their headquarters at the Academy,
until 1894 when they moved to San Diego. A modest and unassuming man,
Brandegee expressed himself crisply on occasion. On one of the several trips to
San Jose del Cabo, when he attended the church there more out of deference to

34 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

the prevailing mores than to his own beliefs, he quipped: "Religion sits very
lightly on the males — they think it good for women and children."

The notable event of 1877 was the visit of the botanists Hooker and Gray to
California. Traveling together they both recorded their impressions and from
their letters, fortunately rather fully published, we can gain some first-hand
knowledge of California in that era. In San Francisco they stopped at the Palace
Hotel, went to Mount Shasta with a pause at Chico : "the trip to Shasta involved
long stagecoach journey's, but they were most interesting. Returning to Sacra-
mento w^e went on to Truckee, where Lemmon joined us by appointment. We
gave one day to Mount Stanford and one to Tahoe, then took the overland train
as it came on at midnight." Hooker was alarmed at the destruction of the
sequoias in the Calaveras grove which they visited: "the doom of these noble
groves is sealed." Hooker also decried the wasteful lumbering practices that he
saw. After the trip. Gray put it succinctly when he wrote : "we should like to
do it all over, and more."

There is no set of chaps so unblushing as naturalists; they are always wanting
something that the other party don't care a straw about.

Thus wrote Alexander Agassiz, from Cambridge, Mass., April 9, 1879, to William
Sillern. Agassiz continues:

Nevertheless, I am going to ask you to put yourself out for me and get me one
of the large Cuttle Fish which used to be so common in San Francisco market when
I was there. The room in the Museum [of Comparative Zoology at Cambridge] de-
voted to the beast and its nearest allies is nearly ready, and I am greatly in want of
a large Cuttle Fish to scare small boys and frighten women. I don't want him too
big, say not more than five feet when fully expanded. The Chinamen used to get them
very often, of all sizes, in their nets and then cut them up and sell them to unsuspect-
ing Frenchmen who mistook the species for frogs' legs. Now if Ralston^ has left any
Chinamen in San Francisco, can you speak to a promising specimen of Mongolian and
ask him to cling to a good specimen, if the species does not freeze to him. Then by
a judicious cutting open of his lower side, so as to let alcohol into his insides, put him
into a keg of alcohol and ship him, via Panama, to your humble servant, who will
receive him with open arms.

The next time you visit the Blaschka glass flowers at Cambridge remember
Agassiz' cuttle fish in the next room !

A zoologist who was to figure prominently in the Academy's history later on
was Barton Warren Evermann, whose first California appointment was as super-
intendent of public schools at Santa Paula, in Ventura County, from 1879-1881.
He was interested in birds and plants at that time, especially birds. On his
twenty-second birthday, October 24, 1875, Barton Evermann married Meadie
Hawkins and she assisted him in preparing bird skins, and in collecting plants.
They assembled a good library but this was lost by fire in 1889 at Indiana State
Normal School. After his return to Indiana State University for advanced studies,
Evermann came under the lasting influence of David Starr Jordan, to weld a
friendship that was to yield rich rewards in scientific authorship. He was special
lecturer at Stanford in 1893-1894, and in the years between 1896 and 1902,

4. William C. Ralston of Bank of California fame. The thousands of Chinese em-
ployed in the construction of the transcontinental railroad flocked to San Francisco
and by 1872 they constituted about half the factory workers in the city. The Chinese
Exclusion Act of 1880 was the result of the campaign to rid the state of Chinese
cheap labor.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 35

alone or in collaboration with Jordan, he published works of classic importance
on North American fishes. Evermann published in all 387 books and articles, of
which about half are devoted to fishes.

It was natural that exploration of Alaska often involved San Francisco, for
the scientific corps commonly assembled there before departure. Charles Haskins
Townsend acted as naturalist on the Revenue Steamer Corivin in 1885, and
on the U. S. Fish Commission Steamer Albatross in 1886-1896. Townsend first
came to California in 1884 as a field naturalist to collect zoological specimens
for the U. S. National Museum. But the Albatross expedition was the most
important trip for on it he collected some plants, along with mostly vertebrate
material along the Alaskan coast. In other years he visited the Marquesas,
Paumotu, Society, Cook, Tonga, and Fiji archipelagoes. Then for thirty-five
years he served as Director of the New York Aquarium, and his conservation
efforts to save from extinction the Alaskan reindeer, Pribiloff fur seal, and Gala-
pagos tortoise earned for him the true gratitude of thoughtful citizens everyAvhere.

We have remarked on the part that Professor Greene played in stimulating
botanical exploration among his students at Berkeley. One of them, Frederick
Theodore Bioletti, tells it this way :

We belonged perhaps to the romantic school of botany. We used the field of botany
not as a laboratory but as a playground. Our heroes were not De Bary, nor Stras-
burger nor Zimmerman, not even Prantl and Engler, but Theophrastus, Rafinesque,
and Edward Lee Greene.

Bioletti came to be best known as a viticulturist and professor of that subject at
his alma mater. In Professor Greene's class with Bioletti were W. L. Jepson,
Victor King Chesnut, Walter Blasdale, and Bioletti's particular chum and com-
panion on field trips, Charles A. Michener. Of one of these Bioletti writes :

Victor Chesnut we looked upon as an enemy and outlaw. He had collected a Rihes
and a Trifolium in the Napa-Sonoma Mountains in the heart of our main hunting
grounds. If we had known his territory we would have invaded it without scruple.
To capture a beautiful and apparently new Ribes in a remote gorge on the slopes of
Hood's Peak, to bring it back to camp in triumph and then to find that it had already
been branded Ribes victoris was intolerable.

Professor Greene as the Great Chief was of course free from all restrictions. We
had too much to gain from his friendship to object to his hunting on our grounds. It
was Professor Greene who used the names Michener and Bioletti several times in
christening some of our discoveries. For this we were deeply grateful.

Chesnut entered the United States Department of Agriculture in 1894 in charge
of poisonous-plant investigations, his previous instruction in chemistry at Berke-
ley serving him well as a background. His Principal Poisonous Plants of the TJ. S.
was one of the most popular publications ever issued by the Government, widely
copied in the press of the day and translated into French, German, and Bohemian.
Elmer Reginald Drew, with whom Chesnut often botanized in the north Coast
Ranges, became Professor of Physics at Stanford. Edwin C. Van Dyke, M.D.,
Assistant Professor of Entomology at Berkeley, was another student of Professor
Greene's, and the botanist, Ivar Tidestrom, one of his last before he left Berkeley
for Washington.

Greene himself botanized on San Miguel Island during the summer of 1886,
leaving Santa Barbara August 19 and landing at Cuylers Harbor nine days later.
The island had been visited by Cabrillo in the winter of 1542-1543, and liis ex-

36 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

hausted body had been lowered into an unmarked grave. Greene did not find the
treasure chest perennially sought by the Conquistadores but he did discover some
remarkable endemic plants on the island. In 1887 Joseph LeConte published in
the Academy's Bulletin a paper entitled "The Flora of the Coast Islands of Cali-
fornia in Relation to Recent Changes of Physical Geography" from the data
supplied by Professor Greene, "though the interpretation of [the data] was
entirely my own/' says LeConte.

In addition to Greene's students there was an array of country school teachers
and ranchers, wives of miners, and travelers, who corresponded with the Berkeley
professor and sent him notable collections. C. C. Marshall was a teacher who
collected around Eureka in the mid-1880's. J. B. Hickman taught school at
Carneros, in Carneros Canyon, on the Natividad road in the San Miguel Hills
and spent his Saturdays and vacations searching the countryside for new plants.
Andrea Massena Norton was born at Lanesboro, Susquehanna County, Pennsyl-
vania, September 7, 1853, and taught school for twelve years at Gonzales, Monte-
rey County, beginning in 1880. He was for part of that period also a member of
the County Board of Education. It was J. B. Hickman, a fellow teacher and
close friend, who introduced Andrea Norton to the scientia amahilis. The very
restricted Eriogonum of the Pinnacles region, and the Monterey County Chori-
zanthe that bear his name were but two of his botanical discoveries.

Some day a historian will tell the story of California's natural history from
the vantage point of the ranches where the naturalists foregathered as field bases.
There will be Talley's ranch in San Diego County, and Warner's ranch; the
Parish ranch near San Bernardino; Duffield's ranch in the Sierra foothills; and
the Ricksecker farm in Sonoma County, to mention a few. Lucius Edgar Rick-
secker was an entomological collector and a propagator of insects for specialists
and cabinet collectors. When not employed as surveyor for Sonoma County, he
lived on his farm at Sylvania near the present site of Camp Meeker. He came
to California in 1873 after serving as a corporal in the Civil War and maintaining
a short residence in Salt Lake City. The insects associated with the sand dunes
of Marin County and about San Francisco interested Ricksecker, and he found
that his talents for netting unusual forms was profitable. Except for a short
residence at Spokane, he lived continuously in the State from 1873 until his death
in 1913. To his farm at Sylvania came many Academy members, including Hark-
ness, to search for truffles and other fleshy fungi; Harford, for spiders; Rivers,
for Lepidoptera and Coleoptera; and Mary Katharine Curran, for plants.

William C. Bartlett of the San Francisco Bulletin remarked in an article
published in the Overland Monthly for December, 1875, that "through the
munificence of a single citizen, the Academy of Sciences has been handsomely
endowed, and will soon be equipped for effective work." The benefactor will be
recognized as James Lick, who gave the property for the erection of the new
museum building for the Academy on Market Street, between Fourth and Fifth
streets. This new center of activity, with its fine display features for museum
exhibits, was the parent of the California Botanical Club, founded on March 7,
1891, "in response to a call" from seven Academy members— something still
miraculous about that number seven! — Harkness, Behr, Eisen, the Brandegees,
To^vnshend and Kate, Mrs. Mary W. Kincaid, and Miss Agnes M. Manning, to
bring the Pacific Coast botanists closer together. Ninety-nine signatures appeared

fWAN: SAN FRANC/SCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 37

on the charter roll, from Carl Purdy on the north to Cleveland, Parish, and
Hasse, from southern California, to mention only a few well kno^vn figures.
C. F. Sonne, G. P. Rixford, (Mary) Elizabeth Parsons, and Alice Eastwood were
among the charter members resident in San Francisco. Miss Eastwood was
leader of the Club after Mrs. Brandegee moved to San Diego, the meetings being
held nearly every week "to study living plants, both native and exotic." From
this more or less informal study group have come valuable collections for the
Academy's herbarium. In this connection the collections of Evelina Cannon,
Caroline L. Hunt, Mary C. Bowman, Mrs. E. C. Sutliffe, Ella Dales Cantelow,
and others across the years, are notable.

In the fall of 1895 David Starr Jordan was elected President of the Academy
and in his autobiography. Days of a Man, he inventories his impressions :

This useful institution struggled on for years with inadequate support until en-
dowed by James Lick in 1876. Its funds were then mainly invested in a large office
building in San Francisco, the museum occupying cramped quarters at the rear. For
some time previous to my election [Jordan continues] the Academy membership had
been divided into two warring factions — one led by Dr. Davidson, the other by Dr.
Harkness, a physician of prominence and an expert in the study of fungi, especially
of the group known as truffles. Both men were vigorous and rather intolerant, a com-
bination of qualities which was not rare in pioneer days, and disrupted more than
one California organization even as it affected the famous "society on the Stanislow."
Indeed, it is reputed that the discords in the institution furnished the motive for Bret
Harte's satirical verse.6

Harkness expressed a desire to retire in Jordan's favor, and Jordan says, "I
then endeavored, with fair success, to put an end to the old feud." Between
1898 and 1911, during Jordan's intermittent presidency, he remarks:

[The] Academy publications were raised to a very high standard as to number,
scientific value, and typographical appearance. For this, special credit was due Dr.
Ritter, the editor; and it should be added that the same level of excellence has been
continuously maintained by our successors.

During these years the Academy's library and collections were growing stead-
ily. To select one of many areas of activity for illustration, we note that the
botanical department acquired the George Thurber herbarium, rich in the Gov-
ernment Railroad Survey collections, and a good set of those of the Death Valley
Expedition. Fifty years after the Academy's founding. Professor T, D. A.
Cockerell wrote in the Popular Science Monthly for April, 1903 :

The civilization of the West is so young that perhaps we ought not to expect much
of the native-born therein . . . indeed a very good crop of young men and women,
who will be prominent in the next twenty years. Everything shows that California,
in particular, will be the center of great biological activity.

Coekerell's prophecy was amply borne out, though interjected in those years was
the destruction of the most valuable collection center in the "West by the fire of
1906 when "a single day saw the destruction of a museum and a librarj^ that had
been fifty years in building. Of thousands of books and specimens of almost
priceless value, nothing was saved except what could be loaded into one spring
wagon and carted to safety ahead of the fire." As Dr. Robert C. Miller, present
Director of the Academy, continues :

5. The "warring factions" of the 1890's postdated the publication of Bret Harte's
verse, which perhaps rests on the altercations of the 1860s.

38 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

That anything at all was saved was due especially to Miss Eastwood, then as
now [1942] the Academy's curator of botany, who lost all of her own possessions
while attempting to save those of the Academy. ... It was justice in the most poetic
sense that more than half a century after the Academy had voted to admit women to
its activities, the book of minutes containing the record of that action, along with
other documents and specimens of inestimable value, should have been saved through
the energy and resourcefulness of a woman curator.

Alice Eastwood first visited California in 1890 as a tourist, then returned the
next year for a brief but active visit engaged in Academy affairs. In 1892 she
joined the Academy staff as joint Curator of Botany with Mrs. Katharine
Brandegee. Following Mrs. Brandegee's taking up residence in San Diego in
1894, Miss Eastwood became the Academy's Curator and head of the Department
of Botany. She struck her characteristic stride in a series of papers published
in the Botanical Gazette, the Bulletin of the Torrey Botanical Chib, and the
Proceedings of the Academy on the California flora. She prefaced A Handbook
of the Trees of California with the statement that "the pressing need of a popular
manual of the trees of California is the reason for this little book." "Throughout
the work the aim has always been brevity and clearness — the desire to help rather
than to shine." Endowed with unusual energy, she rebuilt the Academy's botani-
cal resources and initiated many worth-while activities. These ranged from the
around-the-year "live exhibit" of named flowering specimens in the Academy's
foyer for the instruction of visitors to the republication of Lindley's useful
glossary of botanical terms and the initiation of the Leaflets of Wester7i Botany,
a periodical founded jointly with Jolm Thomas Ilowell, the present Curator of
Botany. For Alice Eastwood, as for Sir ChristoiDher AVren, we may well recall
his motto. Si monumentum requiris, circumspice.

The Academy's first salaried director was B. AV. Evermann, whose California
residence from 1879 to 1881 as a school superintendent has been mentioned.
Beginning in 1914 Dr. Evermann served the Academy for eighteen years. In
1915 he reported 20,000 specimens in the Department of Birds; 31,500 reptiles
and amphibians, including 266 specimens of the Galapagos land tortoises; and
the recent acquisition of the Hemphill conchological collection of over 60,000
specimens. At that time Evermann reported that the Academy's herbariiun
contained more than 18,000 sheets. The collections were then temporarily housed
at 343 Sansome Street, but soon were installed at the new quarters in Golden
Gate Park. Under Dr. Evermann's direction the Academy grew in prestige and
importance. A hard-driving worker for himself as for others, he introduced the
punching of time clocks on one occasion! Evermann made capital gains during
his years at the Academy. In addition to his own research studies on fishes and
the bringing of the Eigenmann South American fish collections to the Academy
as the nucleus of its ichthyological department, he implemented the Steinhart
Aquarium in 1921 and eight years later the Leslie Simson habitat groups of
African wild life. During his directorship the Academy published twenty-five
volumes of scientific reports. His energies were so thoroughly dedicated to the
Academy and the natural sciences that it is doubtful if he gave more than
passing thought to the amenities of social living. Certainly the awesome severity
he evinced toward his Academy associates was more defensive than real.

During Evermann's directorship John Van Denburgh served as Curator of
Reptiles and his assistant was the present citrator, Joseph R. Slevin, most widely

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 39

known for liis detailed knowledge of Galapagos rejitilcs, who will have com-
pleted fifty years of service to the Academy in 1954. Leverett Mills Loomis, who
served as director before Dr. Evermann, was later curator of sea birds. Though
a competent ornithologist, Loomis' stern, uncompromising opinions ruffled other
feathers from time to time. There was no question, however, but that Loomis
was an able "museum man."

In entomology the Academy's collections and reputation grew under the
curatorship of a coleopterist, E. C. Van Dyke, who served from 1904 until 1916,
assisted by Carl Fuchs. Later E. P. Van Dazce, a hemipterist, became curator
of the collections and edited the Pan-Pacific Entomologist, a periodical aided
financially by the Academy.

"History itself," writes Professor Frederick J. Teggart, "does not seek to
elucidate the future; it takes account only of the steps by which the present
situation has come to be as it is." Prophecy, then, has no proper place in this
sketch. The emphasis has been rather on the character of the naturalist, his
sources and resources, his efforts to found an Academy of Sciences devoted first
to the descriptive fields of the natural sciences and more recently metamorphosing
into an interpretative effort where the accumulated facts may be fitted into a
possible pattern.

Dr. Stillman, the pioneer naturalist-physician of San Francisco, wrote a bit
wryly :

Of those who returned to their old homes [from California] to enjoy the fruits
of their enterprise we know but little, we pity them much. ... To them and our
children we leave this beloved land. . . . We have not all realized the hopes that
made radiant the morning of our lives and sustained us through so great hardships; —
fortune was ever a capricious goddess. . . . Our brethren told us fin 1S49] to go in
freedom's name and possess the land — "to read no more history until you have made it."

Crescit sciential

Roster of Biographies

This roster is planned as a guide to biographical references to persons, both
visitors and residents, who have become associated with San Francisco, a contri-
bution toward some ultimate "IVIeisel" for California natural history. "San
Francisco" as used in the title is inclusive and refers to the general San Francisco
Bay region but does not extend south of the Stanford habitat nor north of Marin
County. "Naturalist" herein accents the natural history collector but includes
resident persons who have been traditionally associated with such collections as
descriptive biologists. The time limits extend from the earliest contacts subse-
quent to the purely historic figures whose role was merely incidental (and are
thus not included) to the present time, but no effort has been made to include
all the contemporaries since to do so would amount to reproducing membership
lists of local organizations and to throwing the whole portrait of the growth of
San Francisco natural history out of focus.

The plan of this roster follows certain other bibliographic tools of this nature,
provided by Britten and Boulgcr in England, ])y Ignatz Urban for the West
Indies, and by the author for the Rocky IMountain region. Code words in italics
used to abbreviate sources wherein biographical materials may be found are
explained in the introductory list of ablu-ovintions. Ancillary references to the

40 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

usual sources are given; indeed, many of the better known references are omitted
for the more prominent persons in the interest of saving space in favor of over-
looked commentary. Particular effort has been made to list less familiar sources
of information. These sometimes include commentary of a very incidental nature
in autobiographies and the like where persons may be succinctly evaluated as
well as identified,

A few important general accounts of reference value to anyone concerned
with the San Francisco region are indicated by an asterisk prefixed to the code
word in the following list.

ABBREVIATIONS

The following biographical directories, dictionaries, and various published sources
of Information on the life, travels, and collections of naturalists associated with San
Francisco are referred to by the italicized abbreviation explained here:

ACAB Appleton's Cyclopedia of American Biography, ed. by J. G. Wilson and John
Fiske with rev. supp. New York, 1887-1924.

*Alden Alden, Roland H., and John D. Ifft, Early naturalists in the Far West. Occ.
Pap. Calif. Acad. Sci., 20:i-v, 1-59. 1943.

ADB Allgemeine Deutsche Biographic. Leipzig, 1875-1912.

Amsci American Men of Science, ed. by Jacques and J. McKeen Cattell. Ed. 1 (1906);
ed. 2 (1910) ; ed. 3 (1921) ; ed. 4 (1927) ; ed. 5 (1933) ; ed. 6 (1938) ; ed. 7 (1944).

Bade Bade, William Frederic, Life and Letters of John Muir. Boston and New York,

1923-1924.
Blankins?iip Blankinship, Joseph William, A century of botanical exploration in

Montana, 1805-1905: collectors, herbaria and bibliography. Montana Ag. Coll.

Sci. Studies Bot., 1:1-31. 1904.

Bradley Bib. Rehder, Alfred, Bradley Bibliography. A Guide to the Literature of the
Woody Plants of the World Published Before the Beginning of the Twentieth
Century. 5 Vols. Cambridge, Mass., 1911-1918.

*Breicer Brewer, William H., List of persons who have made botanical collections in
California. In Sereno Watson, Botany of California, 2:553-559. 1880.

Brewster Brewster, E. T., Life and Letters of Josiah Dwight Whitney. Boston, 1909.

Britten Britten, James, and George S. Boulger, A Biographical Index of Deceased
British and Irish Botanists. Ed. 2. London, 1931.

Butler Butler, Ruth Lapham, A Check List of Manuscripts in the Edward E. Ayer Col-
lection. Newberry Library, Chicago, 1937.

Candolle de Candolle, Alphonse, La Phytographie. Paris, 1880, esp. pp. 383-462.

Carpenter Carpenter, Mathilde M., Bibliography of biographies of entomologists. Amer.
Midi. Nat., 33:1-116. 1945.

DAB Dictionary of American Biography, ed. by Allen Johnson and Dumas Malone.
New York, 1928-1937, and Suppl. I, 1944.

Dall Dall, William Healey, Spencer Fullerton Baird, a Biography. Philadelphia and

London, 1915.
DNB Dictionary of National Biography, ed. by L. Stephen and S. Lee. London. 1885-

1901, and supplements.
Dean Dean, Bashford, A Bibliography of Fishes. 3 vols. Amer. Mus. Nat. Hist.. New

York, 1916-1923.
*Eastwood Eastwood, Alice, Early botanical explorers on the Pacific Coast and the

trees they found there. Calif. Hist. Soc. Quart. 18(4) :335-346. 1939.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 41

Emhacher Embacher, Friedrich, Lexikon der Reisen und Entdeckungen. Leipzig, 1882.

*Essig Essig, Edward Oliver, A History of Entomology. New York, 1931.

Ewan Ewan, Joseph, Rocky Mountain Naturalists. Denver, 1950.

Farquhar's Brewer Farquhar, Francis P., ed., Up and Down California, the Journal
of William H. Brewer. New Haven, 1930. Reissued, Berkeley, Univ. Calif.
Press, 1949.

Farquhar's Yosemite Farquhar, Francis P., Yosemite, the Big Trees and the High
Sierra, a Selective Bibliography. Berkeley and Los Angeles, 1948.

Geiser-two Geiser, Samuel Wood, Naturalists of the Frontier. Ed. 2. Dallas, 1948.

Gray Gray, Jane Loring, Letters of Asa Gray, 2 vols. Boston, 1893.

Harshberger Harshberger, John William, Botanists of Philadelphia and Their Work.
Philadelphia, 1899.

Howell Howell, John Thomas, Marin Flora. Berkeley and Los Angeles, 1949.

Hughes Hughes, Katherine Whipple, A Contribution Toward a Bibliography of Oregon
Botany with Notes on the Botanical Explorers of the State. Oregon State College
Thesis Ser. no. 14 (mimeographed). 1940.

Hult^n Hulten, Oskar Eric Gunnar, History of botanical exploration in Alaska and
Yukon territories from the time of their discovery to 1940. Bot. Not., 1940:
289-346. Map. 1940.

Hume Hume, Edgar Erskine, Ornithologists of the United States Army Medical Corps.
Baltimore, 1942.

Huxley's Hooker Huxley, Leonard, Life and Letters of Sir J. D. Hooker. 2 vols. Lon-
don, 1918.

*Jepson Jepson, Willis Linn, Botanical explorers of California — I, Madrono 1:67-170
(1928).— n, 1:175-177 (1928).— Ill, 1:183-185 (1928).— IV, 1:188-190 (1928).- V,
1:214-216 (1929).— VI, 1:262-270 (1929).— VII, 2:25-29 (1931).— VIII, 2:83-88
(1933).— IX, 2:115-118 (1934).— X, 2:130-133 (1934).— XI, 2:156-157 (1934).

Kelly Kelly, Howard A., Some American Medical Botanists, Commemorated in Our Bo-
tanical Nomenclature. Troy, N. Y'., 1914.

Lasegue Lasegue, A. Musee Botanique de M. B. Delessert. Paris, 1845.

Lemmon Lemmon, John Gill, Oaks of the Pacific Slope, pp. 1-19. 1902. Reprinted from
Trans. Pac. States Floral Congress, Oakland, 1902. Copy examined in the Ref. Lib.
of Univ. Calif. Herb., Berkeley.

Liverpool Stansfield, H., ed.. Handbook and Guide to the Herbarium Collections in
the Public Museums, Liverpool, 81 pp. 1935.

*Meisel Meisel, Max, Bibliography of American Natural History, 3 vols. New York,
1924-1929.

Moldenke Moldenke, Harold N. and Alma L., A brief historical survey of the Verbe-
naceae and related families, Pt. II (Biographical). Plant Life (H. P. Traub, ed.),
2:46-98. "1946," 1948.

Musgrave Musgrave, Anthony, Bibliography of Australian Entomology, 1775-1930,
with Biographical Notes on Authors and Collectors. Sydney, 1932.

NBG Hoefer, J. Ch. F., Nouvelle Biographie Universelle (later, Gen^rale). Paris,
1852-1866.

NCAB National Cyclopedia of American Biography, ed. by "distinguished biogra-
phers," Ainsworth R. Spofford, advisory ed. New York, 1898-1947. Current vols.
1930-1948.

Ostorn Osborn, Herbert, Fragments of Entomological History. Including Some Per-
sonal Recollections of Men and Events. Pts. I and II. Columbus, Ohio, 1937-1946.

42 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

*Palmer Palmer, Theodore Sherman, Notes on persons whose names appear in the
nomenclature of California birds. A contribution to the history of West Coast
ornithology. Condor, 30:261-307. 1928.

Pennell Pennell, Francis Whittier, Botanical collectors of the Philadelphia local area.
Bartonia, 21:38-57, 1942; 22:10-31, 66. 1943.

Piper Piper, Charles Vancouver, Flora of Washington, Contr. U. S. Nat. Herb. 11:10
20. 1906.

Rodgers' Gray Rodgers, Andrew Denny, III, American Botany, 1873-1892, Decades of
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Rodgers' Torrey Rodgers, A. D., Ill, John Torrey. a Story of North American Botany.
Princeton, 1942.

Rydberg Rydberg, Per Axel, Scandinavians who have contributed to the knowledge of
the flora of North America. Augustana [College, Rock Island, 111.] Library Publ.
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Sargent Sargent, Charles Sprague, Silva of North America. 14 vols. Boston, 1891-1902.

Sherhorn Sherborn, Charles Davies, Where is the Collection? An Account of the

Various Natural History Collections Which Have Come Under the Notice of
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*8tiUman Stillman, J. D. B., Seeking the Golden Fleece; a Record of Pioneer Life in
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*Stone Stone, Witmer, Philadelphia to the coast in early days, and the development,
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Sivainson Swainson, William, Taxidermy, with the Biography of Zoologists, and
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Urban Symb. Ant. Urban, Ignatz, in Symbolae Antillanae, 3:1-158. Berlin, 1898.

Urban Fl. Bras. Urban, Ignatz, Flora brasiliensis. Vol. 1, pt. 1. 1906.

Van Steenis Van Steenis-Kruseman. M. J., Flora Malesiana. l:i-clii, 1-639. 1950.

Wagne7--th7-ee Wagner, Henry R., The Plains and the Rockies. Ed. 3. by Charles L.

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EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 43

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Eioan, 155; cf. Madrono, 4:283, 1938.

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Beechey, Frederick William, 1796-1856
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Behr, Hans Hermann, 1818-1904

Brewer, 556; Britten, 339; Carpenter, 7; Essig, 553-556 (portr.) ; cf. Geiser-tico.
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Belcher, Edward, 1799-1877

Britten; DNB; Stillman. 325.

44 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Belding, Lyman, 1829-1917

Palmer, 268; autobiog. notes in Condor, 2:1-5, 1900; A. K. Fisher in Auk, 37:33-45
(portr.), 1920; W. K. Fisher in Condor, 20:51-61 (portr.), 1918.

Bell, John Graham, 1812-1889

Ball, 89-91; Palmer, 268; Stone, 12-13; "Scientific Arena" for Aug., 1887; F. M.
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BiDWELL, Annie E. Kennedy (Mrs. John Bidwell)

Bade, 2:72 et passim; R. D. Hunt, John Bidwell, Prince of California Pioneers, Cax-
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Bigelow, John Mutton, 1804-1878

Brewer, 557; Ewan, 164; Geiser-two, 271; Meisel, 1:165, 3:194, etc.; Howell, 30;
Sargent, 1:88; Wagner-three, 265; A. E. Waller in Ohio Arch, and Hist. Quart., 51:313-
331, 1942.

BiOLETTi, Frederick Theodore, 1865-1939

Bradley Bib., 1:320; Howell 31; autobiog. in Sci. Mo., 29:333-339, 1929; obit, in
Science, 90:364, 1939.

Black, George, fl. 1850-1855

Brewer, 557. An associate of William Lobb; a San Franciscan?

Blaisdell, Frank Ellsworth, 1862-1946

Avisci, ed. 7; Hulten, 316; cf. A. Eastwood in Bot. Gaz., 33:126-149, 199-213; 284-
291, 1902.

Blake, James

Meisel, 3:539; C. D. Leake in Calif. Monthly, 38:22 et seq., 1937.

Blake, William Phipps, 1825-1910

DAB; Farquhar's Brewer, passim; Geiser-two, 271, where dates given as "1828-
1910"; NCAB, 25:202-203, as 1826-1910"; R. W. Raymond in Amer. Inst. Mining En-
gineer. Trans., 4:851-864, 1910.

Blaschke, Edtjard Leontjevitch

Bradley Bib., 1:453; Essig, 557-558; Hulten, 300.

Blasdale, Walter Charles, 1871-
Amsci.

Bloomer, Hiram G., 1821-1874

Brewer, 557; W. L. Jepson in Erythea, 7:163-166 (portr.), 1899; Woodcock & Stearn,
232.

BOLANDER, Henry Nicholas, 1831-1897

Brewer, 558; Candolle, 397; Howell, 31; Woodcock d Stearn, 160, as "Henry Nichol-
son Bolander"; W. L. Jepson in Erythea, 6:100-107 (portr.), 1898; "Directions for bot.
collecting" in Calif. Teacher, 1:131-132, Dec, 1863; C. Purdy in Madroiio, 2:33-34, 1931.

BoTTA, Paolo Emilio, 1802-1870

Alden, 31-32; Brewer, 555; Emhacher, 44-45; Palmer, 269; Stone, 7; IslBG; T. S.
Palmer in Condor, 19:159-161, 1917; J. Grinnell in Univ. Calif. Publ. Zool., 38:275 et
passim, 1932.

Boucard, Adolphe, 1839-1905

Sherhorn, 21; ed. note in Nature, 4:50, May, 1871; T. S. Palmer in Condor, 19:168,
1917; C. A. Kofoid in Condor, 25:85-89, 1923; W. F. H. Rosenberg in Condor, 26:38-39,
1924; J. Grinnell in Pac. Coast Avifauna, 16:12, 1924, for description of Boucard's

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 45

"Travels of a Naturalist," 1894, comprising a series of collected chapters first published
in The Humming Bird [London], Vol. 3, Mar. 1893, to Vol. 4, Dec. 1894. Boucard landed
in San Francisco Aug. 15, 1851, and remained a year, collecting birds and insects.

Bbackenridge, William Dunlop, 1810-1893

Alden, 53; Brewer, 555; Britten, 42; DAB; Eioan. 168; Hughes, 15; Meisel, 1:167
and 3:542; Van Steenis, 74-75; D. C. Haskel, U. S. Exploring Expedition, 1838-1842,
and its publications, 1844-1874. New York, 1942 A. B. Maloney in Calif. Hist. Soc.
Quart, 24:321-325 (portr.), 1945; A. Eastwood, ihid., 337-342, 1945 H. H. Bartlett in
Proc. Amer. Philos. Soc, 82:673-679, 1940.

Bbandegee, Mary Katherine Layne Cx:rran, 1844-1920

DAB; W. A. Setchell in Univ. Calif. Publ. Bot., 13:165-168 (portr.), 1926; M. E.
Jones, Contr. West. Bot., 18:12-18 (portr.), 1933,

Bbandegee, Townsuend Stith, 1843-1925

DAB; Ewan, 170; NCAB, 23:366-367 (portr.); Piper, 18; W. A. Setchell in Univ.
Calif. Publ. Bot., 13:155-178 (portr.), 1926; M. E. Jones, Contr. West. Bot, 15:15-18,
1929; J. Ewan in Amer. Midi. Nat, 27:772-789, 1942.

Brannan, Samuel, Jr.

Cf. A. Kellogg in Proc. Calif. Acad. Sci., 5:16 (Feb. 3) and 5:39 (Mar. 3), 1873.

Brewer, William Henry, 1828-1910

Bade, 2:321; Brewer, 558; Brewster, 190-192; DAB; Ewan, 170; Farguhar's Brewer,
introd., xv-xxx; Howell, 31; Hulten, 315; NCAB, 13:561 (portr.); Sargent, 8:28; E. H.
Jenkins in Amer. Jour. Sci., ser. 4, 31:71-74, 1911; R. H. Chittenden in Nat. Acad. Sci.
Biog. Mem., 12:289-323 (portr.), 1929.

Bridges. Thomas, 1807-1865

Brewer, 558; Britten, 45; Jepson, 2:84-88, 1933; Meisel, 3:726 s.v. "Brydges, Thomas";
A. Gray in Amer. Jour. Sci., ser. 2, 41:265, 1866; I. M. Johnston in Contr. Gray Herb.,
81:98-106, 1928; W. H. Dall, "Memorial sketch. . . read before the California Academy
of Natural Sciences, Jan. 8th, 1866," pp. n.d., examined in N. Y. Bot. Garden Library.

Bruce. (Mrs.) C. C.

Cf. A. Eastwood in Bull. Torrey Bot Club, 30:494, 1903.

Bryant, Harold Child, 1886-
Amsci.

Bryant. Walter [Pierce] E., 1861-1905

Palmer. 271; W. K. Fisher in Condor, 7:128-131 (portr.), 1905, and in Auk, 22:439-
441, 1905; H. H. Bailey in Auk, 23:369-376, passim, 1906; H. W. Henshaw in Condor,
22:59, 1920.

BuRBANK. Luther, 1849-1926

DAB: Fairchild, 263-265 et jmssim (portr.); NCAB, 33:149-150; Woodcock d Steam,
171; J. Y. Beaty in Nature Mag., 42:309-311 (portr.), 1949, and in Flower Grower, 36:363
et seq. (portr.), April, 1949; W. L. Howard in Science Digest 20:9-11, Nov. 1946; H. De
Vries in Pop. Sci. Mo., 67:329-347, 1905; W. L. Howard, Luther Burbank: a victim of
hero worship, Chron. Bot., 9:300-508 (portr.), 1946.

Burke, Joseph

Britten. 55; Ewan. 175; Sargent, 9:4; Wagner-three, 144 refers to Dr. Elijah White
meeting [erroneously "Dr."] Burke near Ft. Hall; Burke evidently shipped his colls,
to England from San Francisco.

Burtt-Davy, Joseph, 1870-1940

Bradley Bib., 5:213 s.v. Davy, Joseph Burtt; Howell, 31; J. Hutchinson in Nature,
146:424. 1940.

46 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Butler, George Dexter, 1850-1910

Jepson, 1:188-190 (portr.) ; S. B. Parish, Biog. Bot., Vol. 1, Ms in Dept. Bot. Library,
Pomona College.

Campbell, Doitglas Houghton, 1859-1953

Amsci; D. S. Jordan, Days of a Man, 1:294, 398 (portr.), 1922; I. L. Wiggins in
Amer. Fern Jour., 43:97-103, 1953.

Cannon, Evelina

Cf. A. Eastwood in Leafl. West. Bot, 4:154, 1945 and 5:45-46, 1947.
Carrigek, Henry Ward

Cf. Pac. Coast Avifauna, 16:175, 1924 for his papers.

Chamisso, Adelbert Ludwig von, 1781-1838

Alden, 21-27; Breiver, 554; Eastwood, 338; Embacher, 73; Essig, 619-620 (portr.);
Hulten, 298; Lasegue, 371-373; Urban Fl. Bras., 1:11-12; Yan Steenis, 104 (portr.) ; A. C.
Mahr, Visit of the Rurik to San Francisco in 1816, Stanford Univ. Publ. Hist. Econ. and
Pol. Sci., 2(2):15-18 et jmssim, 1932; A. Eastwood in Leafl. West. Bot., 4:17-21, 1944;
W. E. Safford in U. S. Nat. Herb. Contr., 9:28-29, 1905; T. I. Storer in Univ. Calif. Publ.
Zool., 27:47-48 and 79-80, 1925; autobiog. notes in Reise um die Welt mit der Roman-
zofiischen Entdeckungs-Expedition in 1815-18, auf der Brigg Rurik, Capt. Otto Kotzebue,
2 vols., Leipzig, 1846, and later eds., e.g., Eutdeckungsreise um die Welt, pp. 103-118,
Munich, 1925.

Chesnut, Victor King. 1867-1938

Amsci, ed. 6; Bradley Bib., 5:173; Hoivell, 31; NCAB, 13:295; Who Was Who in
Amer.

Clark, Howard Walton, 1870-1941

Dean, 1:381; edit. obit, in Copeia, 1941:278-279 (portr.), 1941.

Cleveland, Daniel, 1838-1929

Brewer, 559; Jepson, 1:267-268 (portr.), 1929; H. L. Mason in Madrono, 4:67, 1937;
cf. San Diego [Calif.] Union for July 22, 1921.

Collie, Alexander, (?)-1835

Alden, 30; Breiver, 554; Britten. 70; Huxley's Hooker, 1:106; Lasegue. S4-S5;
Palmer, 273; J. Grinnell in Univ. Calif. Publ. Zool., 38:303 et passim, 1932.

Collignon, Jean Nicholas, 1762-1788

Yan Steenis, 113, 602; cf. NBG. s.v. Cels, J. M.; Monterey colls, made by Collignon
grown at garden of Jacques Martin Cels, 1743-1806, near Paris (cf. Ventenat, Hort.
Cels, pref., p. 2, 1800).

Cooper, James Graham, 1830-1902

Blankinship, 6; Breiver, 558; DAB: Essig, 588; Ewan, 187; Hume. 38-51; Meisel,
1:174; Palmer, 273; Piper, 17; Sargent. 1:30; Wagner-three, 262; anon, in Auk, 19:421-
422, 1902; cf. Condor, 53:194 for overlooked papers in Calif. Farmer and Journ. Useful
Sciences; W. H. Dall in Science, n.s. 16:268-269, 1902; H. W. Henshaw in Condor. 22:59,
1920; contributed chapter on zoology to Titus Fey Cronise's Natural Wealth of Cali-
fornia, 1868.

Cordua, Theodor, 1796-1857

Cf. J. T. Howell in Leafl. West. Bot., 1:180-181, 1935; Memoirs of Theodor Cordua,
the pioneer of New Mecklenburg in the Sacramento Valley, ed. and transl. by E. G. Gudde,
Calif. Hist. Soc. Quart., 12(4):l-33, Dec. 1933.

CoRMACK, William Epps, 1796-1868

Bradley Bib., 1:309; F. A. Bruton in Journ. Bot., 66:175-176, 1928; cf. Narrative

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 47

Of a Journey Across the Island of Newfoundland in 1822 St. Johns, 1824, which con-
tains a list of about 170 plants taken in Newfoundland; map of route publ. in 1824 in
Edinburgh Philos. Journal, 10:156-162; Cormack was a tobacco grower in Australia,
forester in New Zealand, collector in California, and founder of agricultural society in
British Columbia.

Coulter, Thomas. 1793-1843

Alden, 38-39; Brewer, 74; Britten, 74; Easttvood, 341; Stillman, 319; F. V. Coville in
Bot. Gaz., 20:519-531, map, 1895; R. McVaugh in Journ. Wash. Acad. Sci., 33:65-70,
1943; R. Lloyd Praeger, Some Irish Naturalists, 68, 1949; E. P. Wright in Notes from
the Botanical School of Trinity College, Dublin, no. 1, pp. 3-4, 1896.

CoTTTHOUY, Joseph Pitty, 1808-1864

Dall, 72 and 80; Meisel, 1:175 and 3:557; H. H. Bartlett in Proc. Amer. Philos. Soc,
82:650-655, 1940.

Ceum, Ethel Katherine, 1886-1943

H. L. Mason in Madrono, 7:33-35, 1943.

CuRRAN, M. K., see Brandegee, M. K. L. C.

Dall, William Healey, 1845-1927

H. A. P[ilsbry] in Nautilus, 41:1-6, 1927; C. H. Merriam in Science, 65:345-347, 1927.

Dana. James Dwight, 1813-1895

DAB, Meisel, 1:176-177 and 3:560; NCAB, 6:462 (portr.) ; Van !?teenis 129; H. H.
Bartlett in Proc. Amer. Philos. Soc, 82:655-663, 1940; F. S. Collins in Rhodora, 14:66
et passim, 1912; D. C. Oilman, Life of James Dwight Dana, 1899; L. V. Pirsson in Nat.
Acad. Sci. Biog. Mem., 9:41-92 (portr.), 1919.

Davidson, George, 1825-1911

DAB, NCAB. 7:227 (portr.) ; C. B. Davenport in Nat. Acad. Sci. Biog. Mem., 18:189-
217 (portr.), 1938.

Delattre, Pierre Adolphe. 1805-1854

J. Grinnell in Univ. Calif. Publ. Zool., 38:262. 1932; T. S. Palmer in Condor, 20:123-
124, 1918.

DE Mofras, Eugene Duflot, see Dhflot de Mofras, Eugene

Deppe, Ferdinand, 1794-1867

Alden, 39; Brewer, 555; Lasegue, 468; H. Harris in Condor, 43:23-27, 1941; autobiog.
notes in Reisen in Kalifornien, Liidde, Zeitschr. f. Erdkunde, 7:383-390, 1847, not seen.
The death year "1828" cited by Chittenden, Diet, of Gardening, 1951, is an error.

DoANE, Rennie Wilbur, 1871-1942

Carpenter, 24; W. M. Mann, Ant Hill Odyssey, 68, 1948.

Douglas, David, 1799-1834

Alden. 32-37 (portr.); Blankinship, 5; Brexoer, 554; Britten, 94; Candolle, 408;
Douglas, 9; Easttcood, 339; Ewan, 197; Geiser-two, 273; Howell. 29; Lasegue. 193-196;
Meisel 1:179 and 3:729; Palmer, 276; Piper, 12-13; Stillman, 317-319; Wagner-three,
60; cf. L. Constance in Leafl. West. Bot., 2:21-22, 1937; H. Harris in Condor, 43:19-21,
1941; J. T. Howell in Leafl. West. Bot, 3:160-162, 1942; W. L. Jep.son in Madrofio,
2:97-100, 1933; E. L. Little, Jr., in Phytologia, 2:485-490, 1948; portr. in Appalachia,
13:54, 1913.

Drew, Elmer Reginald, 1865-1930
Amsci, ed. 4; Howell, 31.

Dudley, William Russel. 1849-1911

Bradley Bib.. 5:240; DAB; various authors in Dudley Memorial Volume, Stanford
Univ. Publ.. 5-28 (portr.), 1913; D. S. Jordan, Days of a Man, passim. 1922.

48 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

DUFLOT DE MOFBAS, EUGENE, 1810-1884

Hughes, 17; NBG; SaUn, 21144; Stillman, 325; cf. M. E. Wilbur, Duflot de Mofras
Travels on the Pacific Coast, 2 vols., Santa Ana, Calif., 1937.

DuFKESNE, Louis

Sherborn; cf. M. Deleuze, Histoire et Description du Museum Royal d'Historie Na-
turelle, 1:169-170, 1823.

Dunn, George Washington, 1814-1905

Essig, 605 et passim; Jepson, 2:156-157 (portr.), 1934.

Eastvtood, Alice, 1859-1953

Ewan, 200; Fairchild, 444; Howell, 31; L. R. Abrams in Pac. Discovery, 2(1): 14-17
(portrs.), 1949; cf. Acad. News Letter no. 36 (portr.), Dec, 1942; E. Crura in Madrono,
5:74 (portr.), 1939; M. E. Jones, Contr. West. Bot, 18:8, 1933 (portr.); cf. Leafl. West.
Bot., 4:153-156, 1945 for recollections; R. C. Miller in Golden Gardens, 9(12) :3-4, 15
(portr.), 1941; N. Valjeans in Nature Mag., 42:361-362 (portr.), 1949; editorial in Sun-
set for Feb., 1938, pp. 13-15 (portr.); San Francisco Chronicle for Oct. 31, 1953 (portr.).

Eaton, Amos Beebe
Brewer, 558.

Edwards, Henry, 1830-1891

Badd 1:262-264 et passim; Carpenter, 27; Essig, 611-613 (portr.); Ewan, 201; Mus-
grave, 76; Osdorn, 1:162; edit. obit, in Entom. News, 2:129-130 (portr.), 1891; J. S.
Wade in Sci. Mo., 30:240-250, 1930.

Eisen, Gustavus Augustus, 1847-1940

Bradley Bib., 5:254; Carpenter, 28; Essig, 615-617 (portr.) ; A. B. Benson and N.
Hedin, Americans from Sweden, 295, 1950; L. H. Miller, Lifelong Boyhood, 77, 1950;
Edgar Swenson in Amer. Swedish Mo. for Nov., 1935.

Emerson, William Otto, 1856-1940

T. S. Palmer in Auk, 65:492-493, 1948; portr. in Condor, 39:46, 1937.

EscHSCHOLTZ, JoHANN Fbiedricii, 1793-1831

Alden, 27-28; Brewer, 554; Carpenter, 29; EmbacJier, 108; Essig, 617-622 (portr.);
Howell, 29; Lasegue, 212-213; Musgrave, 83; NBG; Van Steenis, 157, as "Eschscholz";
A. Eastwood in Leafl. West. Bot, 4:17-21, 1944; W. L. Jepson in Madrono, 1:253 (portr.),
1929. W. E. Safford in U. S. Nat. Herb. Contr., 9:28-29, 1905.

Evermann, Barton Warren, 1853-1932

DAB, Suppl. One; Dean, 1:377-383; Eioan, 206; Hulten, 309; NCAB, 13-570 (portr.);
D. S. Jordan, Days of a Man, 1:169 et passim, 1922; T. S. P [aimer] in Auk, 50:465-466,
1933; G D. Hanna in Copeia, no. 4:161-162 (portr.), 1932; San Francisco Chronicle for
Sept. 28, 1932; autobiog. note in Proc. Indiana Acad. Scj., 1916:209-210, 1916.

Farris, Charles

R. C. Miller in Pacific Discovery, 6(2):19-20, 1953.

Peilner, John

Cf. J. Grinnell in Pac. Coast Avifauna, 5:20, 1909 ; cf. Nineteenth Ann. Rept. Smithson.
Inst, (for 1864), 421-430, 1865.

Fischer. Friedrich Ernst Ludwig von, 1782-1854
Bradley Bib., 5:281; Essig, 630.

Fisher. Walter Kenrick, 1878-1953

Amsci: D. S. Jordan, Days of a Man, 2:130 et passim, 1922; autobiog. notes In Con-
dor, 42:35-38, 1940.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 49

Fitch, Augustus

Bradley Bib., 5:282; Meisel, 3:574; cf. J. Torrey in Pac. RR. Repts., 4:109, 1857.

Fkemont, John Chakles, 1813-1890

Brewer, 556; DAB; Ewan, 211; Huglies, 17; Meisel, 1:184-185 and 3:577, 730; NCAB,
4:270-272 (portr.); Wagner-three, 95, 115; G. A. Zabriskie in N. Y. Hist. Soc. Quart.,
31:4-17 (portr.), 1947.

Fboebel, Julius, 1805-1893

Embacher, 122; Geiser-two, 274; Meisel, 3:577; NBG, b. "1806"; Wagner-three, 292;
A. E. Zucker, Forty-eigliters, 295, 1950; Aus Amerika. Erfahrungen, Reisen und Studien,
2 vols., Leipzig, 1857-1858, in transl. as Seven Years' Travel in Central America, North-
ern Mexico and the Far West of the United States, London, see esp. 570-578, 1859, and
as A Travers I'Amerique, 3 vols., Brussels, 1861.

FucHS, Carl, 1839-1914

Carpenter, 34; Essig, 635-637 (portr.); W. M. Mann, Ant Hill Odyssey, 79, 1940.

FUNSTON, Frederick, 1865-1917

DAB; Hulten, 309; NCAB, 11:40-41 (portr.); D. S. Jordan, Days of a Man, 1:317
and 2:177, 1922; autobiog. Memories of Two Wars: Cuban and Philippine Experiences,
1911, "a vigorous and unconventional narrative"; cf. "V. Bailey, Into Death Valley fifty
years ago, Westways, 32 (no. 12, pt. 1):8-11 (portrs.), Dec, 1940.

Gabb, WiLLLiM More, 1839-1878

Brewster, 239 and 256; DAB; Essig, 638; W. H. Dall in Nat. Acad. Sci. Biog. Mem.,
6:347-361 (portr.), 1909; edit. obit, in Amer. Nat, 12:494-495, 1878.

Gambel, William, 1821-1849

Brewer, 556; Ewan, 213; Harshberger, 231-233; Meisel, 1:185; Palmer, 278; Sargent,
8:35; Stone, 11-12; J. Grinnell in Univ. Calif. Publ. Zool., 38:316 et passim, 1932; H.
Harris in Condor, 43:35, 1941; C. F. Millspaugh and L. W. Nuttall, Flora of Santa Cata-
lina Island, 28, 1923; W. Stone in Cassinia, 14:1-8, 1910; D. B. Woods in Amer. Journ.
Sci., ser. 2, 11:143-144, 1851.

Gardner, Nathaniel Lyon, 1864-1937

Amsci, ed. 5; D. S. Jordan, Days of a Man, 1:302, 1922; W. A. Setchell in Madrofio,
4:126-128 (portr.), 1937.

Gabman, Samuel, 1843-1927

DAB; Ewan, 214; NCAB, 10:294.

Garvitt, ?

Brewer, 557.

Gibbons, Henry, 1808-1884.

Meisel, 1:186; NCAB, 7:287-288 (portr.) ; R. C. Miller in Pacific Discovery, 6(2) :18-
25 (portr.), 1953.

Gibbons, William Peters, 1812-1897

Bradley Bib., 5:320; Bade, 2:70; Dean, 1:457; Kelly, 216; Meisel, 1:186; W. L. Jep-
son in Erythea, 5:74-76, 1897; author of Waysides of Nature, I, II, and III, Overland Mo.,
Aug. 1870 and Aug., 1875.

GiBBs, C. [or G?] D.

Bradley Bib., 4:527; Brewer, 557; cf. P. A. Munz in Leafl. West. Bot., 7:69, 1953, as
"C. D. Gibbes at Stockton."

GiBBs, George, 1815-1873

ACAB, s.v. Geo. Gibbs, his father; DAB; Dall, 338; Meisel, 1:187; S. F. Baird in

50 -^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Ann. Rec. Sci. and Indust. for 1873, 683, 1875; Biog. sketch in folder Oregon Biog. (A-Z)
at Bancroft Library, Berkeley; G. P. Fisher, Life of Benj. Silliman, 1:214 et passim, 1866.

Gilbert, Charles Hexry, 1859-1928

Amsci, ed. 4; Dean, 1:459-461; D. S. Jordan, Days of a Man, 1:201-229 et passim
(portr. opp. p. 140), 1922; W. M. Mann, Ant Hill Odyssey, 69-70 and 81, 1948.

GoDDAKD, Pliny Earle, 1869-1928

Amsci, ed. 4; Who Was Who in Amer.; F. Boas in Science, 68:149-150, 1928; A. L.
Kroeber in Amer. Anthro., 31:1-8 (portr.), 1929.

GooDALE, George Lincoln, 1839-1923

Amsci, ed. 3; Bradley Bib., 5:331; DAB; Meisel. 3:730; B. L. Robinson in Pop. Sci.
Mo., 39:691-694 (portr.), 1891; W. Trelease in Science, n.s., 57: 654-656, 1923; L. H.
Bailey in Rhodora. 25:117-120 (portr.), 1923; W. J. V. Osterhout, B. L. Robinson, and
M. L. Fernald in Amer. Journ. Sci., ser. 5, 6:275-276. 1923.

GOODRIDGE, J.

Cf. B. Seemann, Botany of the "Herald," 286, 1852; Frederick Scheer named a cactus
of Cedros Island Mamillaria goodridgii for the ship's surgeon attending the "Herald"
but his collecting activities in this region were evidently negligible.

GORDON-CUMMING, CONSTANCE FREDERICA, 1837-1924

Bradley Bib., 1:320, s.v. "Gumming, C. F. G."; cf. her autobiog. acct. Granite Crags.
Edinburgh and London, 1884.

Gray, Asa, 1810-1888

ACAB ; DAB; Eivan. 218; HarsTiberger, passim; Huxley's Hooker, 2:210-218; Kelly.
165-177 (portr.); Meisel, 1:188-189; IslCAB, 3:407-408 (portr.); Rodger's Gray. 131-143
et passim; Bade, passim; G. Bradford in N, Amer. Rev., 215:99-108, 1922; H. H. Bart-
lett in Proc. Amer. Philos. Soc, 82:664-673, 1940.

Grayson, Andrew Jackson, 1819-1869

Palmer. 279-280; W. E. Bryant in Zoe, 2:34-68, 1891; L. C. Taylor in Condor, 53:194-
197, 1951; H. Harris in Condor, 43:31-32, 1941.

Greene, Edward Lee, 1843-1915

Brewer, 559; DAB; Ewan, 219; IsfCAB, 19:332-333; K. B(randegee) in Zoe, 2:88-89.
1891; P. L. Ricker in Science, n.s., 39:109-112, 1914; A. K. Main in Trans. Wis. Acad.
Arts and Letters, 24:147-185, 1929; H. H. Bartlett in Torreya, 16:151-175 (portr.), 1916;
W. L. Jepson in Newman Hall Rev. for Oct. 1918; Amer. Catholic Who's Who, 256, 1911;
M. E. Jones, Contr. West. Bot., 14:49-50, 1912, and 15:2.5-27, 1929; Jack Barber in Catho-
lic AVorld, 160:444-449, Feb., 1945.

Grinnell, Joseph, 1877-1939

Palmer, 280-281; J. M. Linsdale in Auk, 59:269-285 (portr.), 1942; Hilda Wood Grin-
nell in Condor, 42:3-34 (portrs.), 1940; W. K. Fisher, ibid. 35-38 (portr.), 1940; F. B.
Sumner, Life History of an American Naturalist, 213-217, 1945; J. Mailliard in Condor,
26:16, 1924; A. H. Miller in Joseph Grinnell's Philosophy of Nature, pref., vii-x,
(frontis. portr.), 1943.

Griber, Ferdinand, 1830-1907

PaZmer, 281; H. W. Henshaw in Condor, 22:59, 1920: cf. J. Grinnell in Univ. Calif.
Publ. Zool., 38:263, 315 et passim, 1932; cf. Condor, 53:194 for overlooked papers in serial
Calif. Farmer and Journ. Useful Sci.

Haenke, Thaddeus, 1761-1817

Alden, 13; Breiver, 553; Eastwood, 336; Hulten, 297; Lasegue, 451; NBG; Van
Steenis. 209-210; W. L. Jepson in Erythea, 7:129-134, 1899; E. C. Galbraith in Calif. Hist.
Quart., 3(3):215-237, 1924; W. E. Safford in U. S. Nat. Herb. Contr., 9:25-28. 1905.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 51

Hall. Carlotta Case, 1880-1949

Cf. Amer. Fern. Journ., 40:192, 1950; Madrono, 4:283, 1938.

Hall, Harvey Monroe, 1874-1932

Eican. 222; E. B. Babcock in Univ. Calif. Publ. Bot., 17:355-368 (portr.), 1934; W. L.
J(epson) in Madrono, 2:63, 1932; portr. in Madrono, 1:12, 1916.

Hanna, G Dallas, 1887-

Ajiisci: Hulten, 331; D. S. Jordan, Days of a Man, 1:551 et passim, 1922; initial does
not stand for a name, teste Condor, 33:210.

Hansen. George, 1863-1908

Bradley Bib., 5:364; DAB; Jepson, 1:183-185 (portr.), 1928.

Harford. William G. W., 1825-1911

Breicer, 556; Essig, 650; Jepson, 2:83-84 (portr.), 1933; D. S. Jordan, Days of a
Man, 1:218, 1922; W. H. D[all] in Nautilus, 25:8, 1911.

Harkness, Harvey Willson, 1821-1901

Cf. Essig, 740, s.v. Ricksecker; Who Was Who in Amer.; [T. S. Brandegee in] Zoe,
2:1-2 (portr.), 1891.

Hartweg, Karl Theodore, 1812-1871

Alden, 47-48; Brewer, 556; Britten, 141; Eastwood. 342; Hoioell. 29: Lasegue. 207-
209; Sargent. 2:34; Urban Symb. Ant., 57; J. T. Howell in Leafl. West. Bot.. 1:180-181.
1935; W. L. Jepson in Erythea, 5:31-35, 51-56, 1897.

Heath. Harold. 1868-

Amsci: Dean, 1:555; W. M. Mann, Ant Hill Odyssey, 69, 1948.

Heermann. Adolphus Lewis, 1827-1865

Brewer, 557; Dall. 280-281; Geiser-iwo. 275; Hume. 190-205 (portr.), best account;
Meisel. 3:732; Palmer, 282; Stone, 13; H. Harris in Condor, 43:35-36, 1941.

Heller. Edmund, 1875-1939

Excan. 226; Hulten, 322; Who Was Who in Amer.; D. S. Jordan, Days of a Man,
2:421, 1922.

Hemphill. Henry, 1830-1914

W. H. Dall in Science, 40:265-266, 1914, and in Nautilus, 28:58-59, 1914; cf. B. W.
Evermann in Nature and Science on the Pacific Coast, 208, 1915; obit, in Trans. San
Diego Soc. Nat. Hist, 2:58-60 (portr.), 1914.

Henshaw. Henry Wetherp.ee, 1850-1930

Palmer. 282; D. S. Jordan, Days of a Man, 2:90, 1922; E. W. Nelson in Auk. 49:399-
427 (portr.), 1932; T. S. Palmer in Auk, 47:600-601, 1930; autobiography in Condor.
21:102-107 (portr.), 165-171 (portr.), 177-181, 217-222, 1919, and 22:3-10, 55-60, 95-101,
1920.

Hepblrn. James, 1811-1869

Cf. T. S. Palmer in Condor, 33:221, 1931; H. S. Swarth in Condor, 28:249-253, 1926.

Herre, Albert William Christian Theodore, 1868-
Amsci: Who's Who in Amer.

Hickman. John Bale, fl. 1880-1900
Bradley Bib., 5:391.

Hilgard, Eugene Woldemar, 1833-1916

Bradley Bib.. 5:392-393; DAB; Fairchild, 444; R. M. Harper in Bull. Torrey Club.
43: 389-391. 1916; F. Slate in Nat. Acad. Sci. Biog. Mem., 9:95-155 (portr.). 1919.

52 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

HiLLEBEAND, WILLIAM, 1821-1886

ADB; Brewer, 558; Van Steenis, 232 (portr.); A. Gray in Amer. Journ. Sci., ser. 3,
33:164-165, 1887; H. St. John in Chron. Bot., 7:69-70, 1942; cf. E. T. Allen in Science,
74:60-62, 1931; autobiog. notes in Fl. Haw. Isl., pref., vii-xii, 1888.

Hinds, Richard Brinsley, 1812?-1847

Alden, 46; Brewer, 555; Britten, 149; Shertorn; Van Steenis, 232.

HOLDEN, E. S.

J. Grinnell in Univ. Calif. Publ. Zool., 38:273, 300-301, 1932, refers to Stockton colls.;
Dean, 1:594, refers to "E. C. Holden" who may be same person but D. S. Jordan, Days
of a Man, 1:392, 1922, refers to E(dward) S(ingleton) Holden, 1846-1914, astronomer
of Lick Observatory (cf. Who Was Who in Amer. and Amsci, ed. 2), who can scarcely
be same person though his interests were diverse.

Holder, William

Brewer, 558; see C. F. Holder's Holders of Holderness (n.d.).

Holmes, Frank Henry, 18 — ?-1924

T. S. Palmer in Condor, 33:221, 1931.

Hooker, Joseph Dalton, 1817-1911.

Badd, passim; Britten, 152-153; DNB, suppl. 2, 2:294; Ewan, 233; Gray, 672-675 for
Calif, trip of Aug., 1877; Huxley's Hooker, 205-218; Rewa Glenn, Botanical Explorers of
New Zealand, 81-86, 1950; D. Prain in Ann. Kept. Smith. Inst., for 1911, 659-671 (portr.),
1912; B. L. Robinson in Proc. Amer. Acad. Arts and Sci., 62:257-266, 1928.

Horn, George Henry, 1840-1897

Brewer, 558; Carpenter, 46; DAB; Essig, 654-658 (portr.); Meisel, 1:196; NCAB,
7:502-503; J. B. Smith in Science, n.s., 7:73-77, 1898, and in Pop. Sci. Mo., 76:468-469,
1910; edit, obit, in Entom. News, 9:1-3 (portr.), 1898.

Howell, John Thomas, 1903-
Amsci; Ewan, 236.

Hudson, Charles Bradford, 1865-

Artist of Academy's diorama backgrounds; Benezit, Dictionnaire critique . . . pein-
tres, 1952; D. S. Jordan, Days of a Man, 2:87, 1922, as "Charles Bradley Hudson"; Who's
Who in American Art, A. C. McGlauflin, ed., 1:211, 1935.

HtTTCHiNGs, James Mason, 1818-1902

Bade, passim; Farquliar's Yosemite, 18-21, 73-77; F. Walker, San Francisco's Lit-
erary Frontier, 28 et liassim, 1939.

Jeffrey, John, 1826-1854

Brewer, 557; Britten, 165; Eastioood, 343; Hughes, 19; Meisel. 3:733; F. V. Coville
in Proc. Biol. Soc. Wash., 11:57-60, 1S97; J. T. Johnstone in Notes from Roy. Bot. Gard.
Edinburgh, 20:1-53, 1939.

Jepson, Willis Linn, 1867-1946

D. D. Keck in Madrono, 9:223-228, 1948, where a bibliog. of biog. refs, is given. To
Keek's list may be added: H. D. Carew in Touring Tropics, 20(12) :32-34, 50 (portr.),
Dec, 1928; obit, in Carnegie Found, for Adv. Teaching, 42nd Ann. Rept. (1946-47), pp.
79-80, 1947; H(elen) M(arr) Wheeler in Desert Plant Life, 19:43-45, March, 1947. Cf.
also San Francisco Examiner for Nov. 8, 1946, p. 15; San Francisco Chronicle for Nov.
8, 1946, p. 11 and Nov. 9, 1946, p. 7; Hoicell, 31; autobiog. notes in Madrono, 4:276-286
(frontis. portr.), 1938.

Jones, Katherine Davies, 1860-1943

M. Symmes in Madrono, 8:184-187, 1946.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 53

JOEDAN, David Starr, 1851-1931

DAB; Dean, 1:643-661; Ewan, 241; NCAB, 22:68-70 (portr.) ; B. W. Evermann in
Proc. Indiana Acad. Sci., 1916:205-207, 1916; B. W, Evermann in Condor, 34:6-7, 1932,
on his interest in birds; H. Zinsser, As I Remember Him, the Biography of R. S., 188-
194, 1940; autobiog. Days of a Man, 2 vols., 1922; W. M. Mann, Ant Hill Odyssey, 67-81,
1941; T. D. A. Cockerell in Pop. Sci. Mo., 62:516, 1903.

Jordan, Eric Knight, 1903-1926

Anon, in Nautilus, 40:33-34, 1926.

Kaeding, Henry Barroilhet, 1877-1913

Palmer, 283; J. Mailliard in Condor, 15:191-193 (portr.), 1913.

Keep, Josiah, 1849-1911

Author of Common Seashells of California, ed. 1, 64 pp., 1881, and West Coast Shells,
230 pp. 1887; W. H. Dall in Science, 34:371, 1911, and Nautilus, 25:61-62 (portr.), 1911.

Kelley, Lynwood J.

Woodcock d Steam, 244.

Kellogg, Albert, 1813-1887

Bade, 2:70 et passim; Bradley Bib., 5:449-450; Brewer, 556; DAB; Essig, 650 et
passim; Geiser-Uco, 276; Hulten, 302; Meisel, 1:200 and 3:734; NCAB, 25:205-206; Wag-
ner-three, 274a; Woodcock d Steam, 245; A. Gray in Amer. Journ. Sci. ser. 3, 35:261-262,
1888; E. L. Greene in Pittonia, 1:145-151, 1887; D. S. Jordan, Days of a Man, 1:218,
1922; R. C. Miller in Pacific Discovery, 6(2):18-25 (portr.), 1953; P. A. Munz in Leafl.
"West. Bot, 7:70-71, 1953; cf. Leafl. West. Bot, 7:101 (pi. 5), 1953, for his handwriting;
C. H. Shinn in Garden and Forest, 2:298, 1889.

Kellogg, Veenon Lyman, 1867-1937

Amsci, ed. 5; Carpenter, 51; L. 0. Howard, Fighting the Insects, 188, 1933; D. S.
Jordan, Days of a Man, passim, 1922; C. E. McClung in Nat. Acad. Sci. Biog. Mem.,
20:245-257 (portr.), 1939; W. M. Mann, Ant Hill Odyssey, 68-69, 1948; Who was Who
in Amer.

/

Kennedy, Patrick Beveridge, 1874-1930

W. L. Jepson in Madrono, 2:34-35 (portr.), 1931; cf. Bot. Soc. Amer. Publ. 105,
19-20, 1931.

King, Clarence, 1842-1901

Bade, passim; DAB; Farquliar's Brewer, passim; Farguhar's Yosemite, 49-53; Mei-
sel, 1:201; S. F. Emmons in Nat. Acad. Sci. Biog. Mem., 6:25-55 (portr.), 1909; cf. his
Mountaineering in the Sierra Nevada, 1871.

Knoche, Edward Louis Herman, 1870-

Dudley Memorial Volume, 31, 1913, in list of Dudley's students; cf. Madroiio, 4:283,
1938.

Kofoid, Charles Atwood, 1865-1947

C. Dobell in Nature, 160:115-116, 1947; R. B. Goldschmidt in Nat. Acad. Sci. Biog.
Mem., 26:121-151 (portr.), 1951; H. Kirby in Sci. Mo., 61:415-418 (portr.), 1945 and in
Science, 106:462-463, 1947; portr. in Fortune, 33(6) :157, 1946.

KoTZEBtJE, Otto von, 1787-1846

Brewer, 554; Embaclier, 176; Lasegue, 371; NBG; Palmer, 284; Stillman, 310-316.

Laglaize, Leon

A. Boucard, Travels of a Naturalist, 50, 1894; "grandson of Lorquin" who collected
insects in San Francisco region during the 1850's.

54 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Langsdorff, Georg Heineicii von, 1774-1852

Alden, 19-21; Brewer, 554; Hoicell, 29; HuUcn, 297; KBG: Stilhnan. 308; Swain-
son, 231.

Lanszweeet, L.

Dean, 2:12, who cites one paper.

La Pebouse, Jean Francois Galavp de. 1741-1788

Alden, 9-12; Eastwood, 335; Embacher, 182; A'^BG; Stone, 4; G. Chinard, Le Voyage
de Laperouse sur les cotes d I'Alaska et de la Californie (1786), esp. p. 106. 1937; cf. M.
Gabriel Marcel, bibliog. of La Perouse in Bull. Soc. Geog. France for 1888.

Lathrop, Barbour, 1846-(?)

Fairchild, 104, 302, et passim (portr., 84A) ; D. Fairchild, Exploring for Plants, 328
et passim, 1930; and World Grows Round My Door (portr.), 73 et jxissim, 1947; M. S.
Douglas in Reader's Digest, 53:67-71 (portr.), Nov., 1948.

Lay, George Tradescant, (?)-1845.

Alden, 30-31; Brewer, 534; Britten, 182; Lasegue, 84-85; Van Steenis, 315-316; cf.
Notes and Queries, ser. 1, 5:386, 1852.

LeConte, John Lawrence, 1825-1883

Carpenter, 58; DAB: Essig, 680-685; Eivan, 248; Meisel, 1:203-204; Palmer, 285;
G. H. Horn in Science, 2:783-786, 1883; S. H. Scudder in Nat. Acad. Sci. Biog. Mem.,
2:261-293, 1886; J. B. Smith in Pop. Sci. Mo., 76:468-469 (portr.), 1910.

LeConte, Joseph, 1823-1901

Bade, passim; DAB; Farguhar's Yosemite, 58; Meisel, 1:204; E. W. Hilgard in Nat.
Acad. Sci. Biog. Mem., 6:147-218 (portr.), 1909; D. S. Jordan, Days of a Man, passim,
1922; L. H. Miller, Lifelong Boyhood, 104-105, 1950: cf. Autobiography of Joseph LeConte,
ed. by W. D. Armes, 1903; cf. his A Journal of Ramblings Through the High
Sierras of California by the University Excursion Party (1875), reprinted by Sierra
Club, 1900; cf. his Flora of the Coast Islands of California in Relation to Recent Changes
of Physical Geography, Bull. Calif. Acad. Sci., 8:515-520, 1887.

Lemmon, John Gill, 1832-1908

Brewer, 558; DAB: Eivan, 249; H. F. Copeland in Madroiio, 5:77 (portr.); mss.
notes in Ewan files; Harold St. John is preparing an account of J. G. Lemmon (cf.
Berkeley Gazette for June 9, 1941) the "Professor" in Mabel Craft Deering's story "Kid-
naping the Casting Vote" (Sunset Mag., 16:371-378. Feb., 1906) is Lemmon, fide S. B.
Parish in mss. Biog. Bot., Vol. 2, Dept. Bot. Lib. Pomona Coll.

Letcher, Beverly, 1864-1905
Essig, 636 (portr.).

Loi:b, William, 1809-1863

Brewer, 557; Britten, 191; Eastivood, 343; Sargent. 10:60; Veitch's account re-
printed by A. Eastwood in Muhlenbergia, 7:100-103, 1911; cf. A. Eastwood in Leafl. West.
Bot., 5:155-156, 1949; cf. Farquhar's Yosemite, 5-13. for survey of early literature on
Big Tree but no mention of Lobb.

LocKiNGTON, William Neale, 1842(?)-1902

Dean, 2:52-53; D. S. Jordan, Days of a Man, 218, 1922.

LooMis, Leverett Mills, 1857-1927

L. B. Bishop in Auk, 46:1-13 (portr.), 1929; T. S. Palmer in Auk, 45:263-264, 1928;
H. S. S(warth) in Condor, 30:194-195, 1928.

LoRQUiN, Pierre Joseph Michel, 1797-1873

Carpenter, 62; Essig, 694-697 (portr.); F. Grinuell, Jr.. in Eiitom. News, 15:202-204,
1904. as "ca. 1800-1877"; cf. J. Grinnell in Univ. Calif. Publ. Zool., 38:318. 1932. and H.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 55

Harris in Condor, 43:44, 1941; cf. H. W. Henshaw in Condor, 22:59, 1920, on Ernest F.
Lorquin of "410 Kearney St., San Francisco."

LoTSY, Johannes Pauli^s, 1867-1931

Van Steenis. 330-331 (portr.); A. D. Rogers, III, Erwin Frink Smitli, 220-221, 1952;
autobiog. notes in Van Den Atlantischen Oceaan naar de Stille Zuid Zee, Dagboek van
een botanicus, die niet alleen naar planten keek. s-Gravenhage, esp., 288-294, 1930.

McLaren, John, 1846(?)-1943

Bradley Bib., 5:534; Fairchild. 444, and World Grows Round My Door, 46 and 146,
1947; Samuel Dickson, San Francisco Is Your Home, 215-221, 1947; Frank J. Taylor in
St. Eve. Post for July 29, 1939.

McLean, F. P.

Collected plants on "stream of Tamalpais" in 1873 (cf. Psoralea fruticosa Kell.) ;
(?) relative to Miss K. D. McLean of Oakland (cf. Cassino, Nat. Direct, for 1890),

Mackie, William Wylie, 1873-

Cf. W. L. Jepson in Madrono, 4:276, 1938.

Mailliard, Joseph, 1857-1945

R. C. Miller in Auk, 64:300-302 (portr.), 1947; autobiog. in Condor, 26:10-29
(portr.), 1924.

Mann, Horace, Jr., 1844-1868

Brewer, 558; Wm. T. Brigham in Boston Soc. Nat. Hist. Proc, 12:152-155, 1868;
"Friend and Associate" in Essex Inst. Bull., 1:25-31, 41-50, 1869.

Mann, William M., 1886-

Amsci; autobiog. Ant. Hill Odyssey, 1948 (portr.); Sci. Mo., 63:358 (portr.), 1946.

Martiniere, De Boissieu la

AUen, 11; Yan Steenis, 350; cf. Vellozo, PI. Flumin., 232, 1825, and Antoine Guille-
min in Delessert, Icon, select., 3:23, t. 49, 1837.

Mason, Herbert Louis, 1896-

Amsci; Howell, 31; Hulten, 338.

McDonald, James Monroe, 1825-1907

Essig, 61 et passim; San Francisco Call for Feb. 28, 1892, and June 9, 1907; San
Francisco Chronicle for Dec. 17, 1921.

McGregor, Richard Crittenden, 1871-1936

Dean, 1:657; Palmer, 287; obit, in Auk, 54:234, 1937; J. Grinnell in Auk, 55:163-175
(portr.), 1938; J. G(rinnell) in Condor, 39:45, 1937; D. S. Jordan, Days of a Man. 1:709,
1922.

Menzies, Archihald, 1754-1842

Alden, 14-18 (portr.); Brewer, 553; Britten, 213; Dean, 2:129; D^SB; Hughes, 7;
Hulten, 297; Lasegue, 366; Liverpool, 61; Jepson. 1:262-266 (portr.), 1929; Rtonr, 4;
A. Eastwood in Leafl. West. Bot., 2:92-94, 1938; J. Grinnell in Condor, 34:243-252, 1932;
E. S. Meany, Vancouver's Discovery of Puget Sound, 295-297 et passim (portr.), 1915;
Geo. Godwin, Vancouver: a Life, 1757-1798, 134-143 et passim, 1930; A. Eastwood in
Calif. Hist. Soc. Quart, 2:265-340, 1924; Rewa Glenn, Botanical Explorers of New Zea-
land, 42-44, 1950.

Merriam, John Campbell, 1869-1945

Amsci, ed. 7; NCAB, Current Vol. A, 485-486 (portr).; Palmer, 288; Chester Stock
in Science, 103:470-471, 1946, and in Geol. Soc. Amer. Proc, 1946:183-197 (portr.), 1947,
and in Nat. Acad. Sci. Biog. Mem., 26:209-232 (portr.), 1951.

56 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Mexia, Ynes Enriquetta Julietta Reygadas [nee Mexia], 1870-1938

N. Floy Bracelin (Mrs. H. P.) in Madrono, 4:273-275 (portr.), 1938; cf. Madrono,
4:284, 1938; H. N. Moldenke in Plant Life, 2(1-3) :78, "1946" 1948; San Francisco News
for Mar. 6, 1937 (portr.).

Michenek, Charles A.

Howell, 31.

MuiB, John, 1838-1914

DAB; HulUn, 304; W. F. Bade, Life and Letters of John Muir, 2 vols., 1923-1924;
L. M. Wolfe, John Muir, 1838-1914, 15 pp. (n.d.) (brochure publ. by H. Mifflin Co.) ; D. S.
Jordan, Days of a Man, 1:217, 1922; s.v. Edwards, Henry, ante.

Neboxtx, Adolphe Simon, fl. 1836-1840

Palmer, 289; T. S. Palmer in Condor, 20:114-116, 1918; cf. J. Grinnell in Univ.
Calif. Publ. Zool., 38:319-320, 1932.

Nelson, Edward William, 1855-1934

W. S(tone) in Auk, 51:431-432, 1934; E. A. Goldman in Auk, 52:135-148 (portr.),
1935; V. Bailey in Westways, 32 (no. 12, pt. 1):8-11 (portr.), Dec, 1940; cf. Sci. Mo.,
1:232-234, 1876, for birds of Oakland, Calif., teste E. Coues.

Nevins, Thomas J.

R. C. Miller in Calif. Hist. Soc. Quart., 21:364, 1942, and Pacific Discovery, 6(2) :18-
25, 1953.

Newberry, John Strong, 1822-1892

BlankinsMp, 10; Brewer, 557; DAB; Dean, 2:179-181; Eivan, 272; Hughes, 20;
Meisel, 1:214; C. A. White in Nat. Acad. Sci. Biog. Mem., 6:3-24 (portr.), 1909; N. L.
Britton in Bull. Torrey Club, 20:89-98 (portr.), 1893.

Newcomb, Wesley, 1808-1892

Brewster, 217; Meisel, 3:736; R. E. C. Stearns in Nautilus, 5:121-124 (portr.), 1892,
and in Science, 28:243, 1908.

Norton, Andrea Massena, 1853-1930

Badd, 2:71, perhaps A. M. Norton (?) ; cf. J. T. Howell in Leafl. West. Bot., 2:99, 1938.

Nunenmacher, Frederick William, 1870-
Essig, 717-719 (portr.).

NuTTALL, Thomas, 1786-1859

Alden, 42-46 (portr.); BlankinsMp, 5-6; Brewer, 555; Britten, 231; Candolle, 437;
DAB; Dall, 47; DNB; Eastwood, 341; Ewan, 273; Gray, 1:326; Harshberger, 151-159,
with some errors (portr.) ; Hughes, 12; Lasegue, 464; Liverpool, 55 et passim (portr.);
Meisel, 1:215-216 and 3:737; l^CAB, 8:374 (portr.); Palmer, 289 (portr.); Piper, 14-15;
Sherborn; Stone, 7-9; F. V. Coville in Proc. Biol. Soc. Wash., 13:109-121, 1899; F. W.
Pennell in Bartonia, 18:1-51, map (portrs.), 1936, the most complete and accurate acct.;
W. Brewster in Mem. Nuttall Ornith. Club, 4:73-81, et passim (portr.), 1906; W. L.
Jepson in Madrono, 2:143-147 (frontis. portr.), 1934; W. C. Coker in Elisha Mitchell Sci.
Soc. Journ., 57:102-104, 1941.

Osgood, Wilfred HtrDsoN, 1875-1947

Ewan, 274; Who's Who in Amer. for 1946; C. C. Sanborn in Journ. Mammal. 29:95-
112 (portr.), 1948.

Ostensacken, Carl Robert Romanovich von der, 1828-1906

Carpenter, 76; DAB; Essig, 724-727 (portr.); Meisel, 3:737; Sherborn: autobiog.
Record of My Life Work in Entomology, Cambridge, Mass., 1903, pts. 1 and 2, and Heidel-
berg, 1904, pt. 3. Only 225 copies printed; copy no. 138 examined at John Creiar Library,

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 57

Chicago; J. M. Aldrich in Entom. News, 17:269-272 (portr.), 1906; C. W. Johnson, ibid.,
273-275, 1906; J. B. Smith in Pop. Sci. Mo., 76:468 and 473 (portr.), 1910.

Palmer, Elizabeth Day, 1872-1945

T. S. Palmer, her brother, in Auk, 67:429, 1950; M. A. thesis, Univ. Calif., 1909: A
taxonomic revision of the genus Chorizanthe R. Br. ms.

Pabker, Hubert G., (?)-1888

Dean, 2:232; H. W. Henshaw in Condor, 22:8-9, 1920.

Parry, Charles Christopher, 1823-1890

Bade, 1:343 and 2:242-243; Blankinship, 8; Breiver, 556 and 559; Britten. 237;
Candolle, 439; DAB; Ewan, 278; Geiser-two, 279; Harshherger, passim-; Kelly. 180-186
(portr.); Lemmon, 11-12; Meisel. 1:217-218 and 3:737 NCAB, 13:228; Sargent. 7:130;
Stillman, 167; Urban Symb. Ant. 98; M. E. Jones, Contr. West. Bot, 17:3-6, 1930; J. G.
Lemmon in Pac. Rural Press, 39:385 (portr.), Apr. 12, 1890; N. L. Britton in Bull.
Torrey Bot. Club, 17:74-75, 1890; Woodcock & Steam, 305.

Parsons, Mary Elizabeth

Author of highly popular Wild Flowers of California, San Francisco, 1897.
Paulsen, Ove

Cf. Madrono, 1:12-18 (portr.), 1916.
Peabody, a.

Brewer, 557.

Peale, Titian Ramsey, 1799-1885

Alden, 51-52; ACAB; Carpenter, 78; DAB; Ewan, 281; Meisel, 1:218 and 3:738;
NCAB, 21:170-171, portr. as "1800-1885"; Stone, 6-7; P. P. Calvert in Entom. News,
24:1-3 (portr.), 1913; H. H. Bartlett in Proc. Amer. Philos. Soc, 82:640-644, 1940.

Pickering, Charles, 1806-1878

Alden, 49-51; Brewer, 555; Carpenter, 79; DAB; Eivan, 283; Harshberger, 190-
193; Hughes, 15; Kelly, 151-153; Meisel, 1:219; NCAB, 13:176; Piper, 15; Van Steenis,
406-407; J. H. Barnhart in Mem. Torrey Club, 16:298, 1921; F. S. Collins in Rhodora,
14:57-68, 1912; W. W. Diehl in Mycologia, 13:38-41, 1921; C. S. Sargent, Sci. Papers
Asa Gray, 2:406-410, 1889; F. W. Pennell in Bartonia, 21:53, 1942; H. H. Bartlett in
Proc. Amer. Philos. Soc, 82:646-650, 1940.

Plummer, Sara Allen

Brewer, 558; s.v. J. G. Lemmon, her husband, ante.

Pratten, Henry

Meisel. 3:640; W. C. Coker in Elisha Mitchell Sci. Soc. Journ., 57:154, 1941.

Price, William Wightman, 1871-1922

Bradley Bib., 5:689; L. H. Miller, Lifelong Boyhood, 73-103, 1950; E. W. Nelson in
Mem. Nat. Acad. Sci., 16:145, 1921; W. K. Fisher in Condor, 25:50-57 (portr.), 1923;
relationship, if any, to forester Overton Westfeldt Price (cf. Quercus pricei Sudworth),
not determined.

Randall, Andrew

R. C. Miller in Pacific Discovery, 6(2): 20, 1953.

Ransom, Leander, 1800-1872

Bradley Bib., 1:210; Brewer, 557; W. C. Ransom, Hist. Outline of the Ransom
Family of America, 1903; D. A. R. Records of the Families of Calif. Pioneers, 12:374, 376.

Rattan, Volney, 1840-1915

Brewer, 558; Jepson, 1:168-170 (portr.), 1928.

58 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Ready, Geokge Henry, 1858-1903

T. S. Palmer in Condor, 33:221, 1931.

Remy, Ezechiel Jules, 1826-1893

ACAB; Bradley Bib., 5:716; Embacher, 246; Ewon, 288; NBG: Wagner-three, 364;
V. MacCaughey in Hawaiian Forester and Agric, 16:26-27, 1919; assoc. in liis travels
with Rev. Julius Lucius Brenchley, 1817( ?)-1873, English missionary; ms. Vocabulaire
Havaiien-Frangais, 167 pp. in Ayer Coll., Newberry Library (Butler, 1768) and another
MS. Vocabulaire Frangais-Havaiien. Recuelli dans I'Archipel de Hawaii pendant les
annees 1852-1855, 250 pp. (Butler, 1769).

Rich, William

Dull, 106; Meisel, 3:644; Van Steenis, 434.

RiCHTHOFEN, FERDINAND PAUL WiLHELM VON, 1833-1905

Brewster; Emhacher, 247; Sher-born ; Van Steenis. 435; Bretschneider, Botanical
Discoveries in China, 943, 1898; Poggendorff, Biog. Liter. Handworterbuch, 3:1121, 1898,
and 5:1048, 1926.

RiCKSECKER, Lucius Edgar, 1841-1913

Carpenter, 85; Eivan, 289; Essig, 738-741 (portr.) ; H. C. Fall in Entom. News,
24:239-240, 1913.

RiTTER, William Emerson, 1856-1944

Avisci, ed. 7; Dean, 2:350; T. S. Palmer in Auk, 64:665-666, 1947; F. B. Sumner
Life History of an American Naturalist, 198-209, et jmssim. 1945; L. H. Miller, Lifelong
Boyhood, 28-32, 104, et i)assim, 1950; D. S. Jordan, Days of a Man, 1:541, 1922; autobiog.
notes in California Woodpecker and I. 315-318, et passim (portr.), Berkeley, Calif.. 1938.

Rivers, James John, 1824-1913

Carpenter, 86; Essig, 746-747 (portr.); Sherborn; Ewan, 290.

RiXFORD, Emmet, 1865-1938

Amsci, ed. 5; anon, in Nautilus, 51:141, 1938.

RlXFORD, GULIAN PICKERING, 1838-1930

Amsci, ed. 4; NCAB, Vol. B:172 (portr.) and 35:537-538 (portr.); W. C. Tesche in
Journ. Heredity, 21:98-106 (portr.), 1930; Millspaugh and Nuttall, Field Mus. Nat. Hist.
Publ. Bot, 5:33, 1923.

RoEZL, Benedict, 1823-1885

Bradley Bib., 5:734; Ewan, 291; Vi^oodcock d Steam, 231, 302; S. B. Parish in Bot.
Gaz., 44:414, 1907, and 48:462-463, 1909; autobiog. in Card. Chron., ser. 2, 2:73 (portr.),
1874, reprinted, ibid., ser. 2, 24:521-522 (portr.), 1885; E. Regel in Gartenflora, 21:369,
1872, and 34:330-331, 1885; E. Morren in Belg. Hort., 30:5-12 (portr.), 1880; A. East-
wood in Leafl. AVest. Bot., 5:103, 1948.

Rose, Lewis S.

J. T. Howell in Leafl. West. Bot., 7:91, 1953.

RxJBEL, Eduard August, 1876-

Cf. Madrono, 1:12-18 (portr.), 1916.

Samuels, Emanuel, 1816-1886

Palmer, 294 (portr.); H. W. Henshaw in Condor, 21:106-107, 1919.

Saxe, Arthur Wellesley, 1820-1891

Dean, 2:396, where no initials given; Kelly, 178-179 (portr.).

ScAMMON, Charles Mellville
Dean, 2:396.

EV/AN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 59

ScHROTER, Carl Joseph, 1855-1939

Cf. Madrono, 1:12-18 (portr.), 1916; Van Steenis, 476-477 (portr.).

Seemanx. Bebthold Carl, 1825-1871

Britten, 271; DNB ; Embacher, 267; Hulten, 300; Van Steenis, 481.

Sessions. Kate Olivia, 1857-1940

Bradley Bib., 5:795; T. D. A. Cockerell in Bios, 14:167-179 (portrs.), 1943; cf. L. H.
Bailey in Gentes Herbarum, 4:99-105 (portr.), 1937.

Setchell. William Albert, 1864-1943

Amsci. ed. 6; T. H. Goodspeed in Essays in Geobotany. In Honor of William Albert
Setchell, xi-xxv (frontis. portr.), 1936; L. Constance in Journ. Wash. Acad. Sci., 33:288,
1943; H. L. Mason in Madroiio, 7:91-93 (portr.), 1943; C. R. Ball, ibid., 5:231-232
(portr.), 1940; D. H. Campbell in Nat. Acad. Sci. Biog. Mem., 23:127-147 (portr.), 1945.

SiLLEBN, William

Cf. G. R. Agassiz, Letters and Recollections of Alexander Agassiz, 161, 1913.

Simpson, George, -1860

Embacher, 271 s.v. Thomas Simpson; StillTnan, 325; Wagner-three, 140; A. S. Mor-
ton, A History of the Canadian West to 1870-71, passim, n.d.

Sinclair, Andrew, 1796-1861

Britten, 276; DNB; Van Steenis, 485; Rewa Glenn, Botanical Explorers of New Zea-
land, 107-114, 1950; H. F. von Haast, Life and Times of Sir Julius von Haast, 173 et pas-
sim, 1948.

Skottsberg, Carl Johan Fredrik, 1880-

Cf. Madrono, 1:12-18 (portr.), 1916; Van Steenis, 486-487.

Slevin, Joseph Richard, 1881-

[Calif.] Academy News Letter no. 164 (portr.), 1953; E. W. Nelson in Mem. Nat.
Acad. Sci., 16:144, 1921; I. M. Johnston in Proc. Calif. Acad. Sci., ser. 4, 20:13, 1931.

Slevin. Thomas Edwards, 1871-1902

Palmer, 296; L. M. L(oomis) in Auk, 20:326-327, 1903.

Sloat, Lewis W.

R. C. Miller in Calif. Hist. Soc. Quart., 21:364, 1942, and Pacific Discovery, 6(2) :18-
25, 1953; evidently Sloat's dupls. did not reach the National Museum teste H. A. Rehder,
who checked the records for me there.

Smith, Charles Piper, 1877-

Cf. Madrono, 4:283, 1938; Eican, 306.

Snodgrass, Robert Evans

Dean, 2:465-466; D. S. Jordan, Days of a Man, 1:577, 1922.

Snyder, John Otterbein, 1867-

Dean, 2:466; D. S. Jordan, Days of a Man, passim, 1922.

Sonne, Charles Frederick, 1845-1913

Bade. 2:308; Jepson, 2:115-116 (portr.), 1934.

Starks, Edwin Chapin, 1867-1932

Aynsci, ed. 4; Dean, 2:478-480; W. M. Mann, Ant Hill Odyssey, 64-71, 1948.

Stearns. Robert Edward Carter, 1827-1909

Bradley Bib., 5:821; Dean, 2:481; W. H. Dall in Science, 30:279-280, 1909; Mary R.

60 -A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Stearns in Smithson. Misc. Coll., 56(18) :1-15, 1912, bibliog. (portr.); H. W. Henshaw in
Condor, 21:107, 1919; autobiog. notes in Amer. Nat, 13:141-154, 1879.

Stillman, Jacob Davis Barcock, 1819-1888

Brewer, 556; J. D. B. Stillman, "Old Fuller," Overland Mo., 14:557-559, June, 1875;
ms. notes in Ewan files; cf. Calif. Med. Gazette, 2:152-153, 1870, for unsigned edit, con-
cerning the work of the State Geol. Survey.

Stewart, Alban, 1875-1940
Amsci, ed. 6.

Stivers, Charles Austin

Jepson, 2:28 (portr.), 1931.

Stomps, Theodoor Jan, 1885-

Cf. Madrono 1:12-18 (portr.), 1916; Van Steenis, 508-509 (portr.).

Stout, Arthur B.

Though initial is generally given as "B" the physician who was one of the original
members of the Academy may be "A. A. Stout, M.D., U.S.N.," elected to the New York
Academy (Lyceum of Nat. Hist.) in 1847.

Street, Joseph A.

Cf. A. Eastwood in Occ. Pap. Calif. Acad. Sci., 9:3, 1905.

Stretch, Richard Harper, 1837-1923

Carpenter, 101; Essig, 767-770 (portr.), who cites 1926 as death year; K. R. Coolidge
and H. H. Newcomb in Entom. News, 31:181-185 (portr.), 1920.

Sumner, Francis Bertody, 1874-1945

Dean, 2:517-518; NCAB, 34:333-334 (portr.); R. R. Heustis in Jouru. Mammal.,
27:1-3 (portr.), 1946; C. M. Child in Nat. Acad. Sci. Biog. Mem., 25:147-173 (portr.),
1949; autobiog, Life History of an American Naturalist, 1945.

Swarth, Harry Schelwaldt, 1878-1935

Amsci, ed. 5; Palmer, 298; J. Mailliard in Auk, 54:127-134 (portr.), 1937; J. M.
Llnsdale in Condor, 38:155-168 (portr.), 1936.

Tansley, Arthur George, 1871-

Cf. Madrono, 1:12-18 (portr.), 1916.

Taylor, Henry Reed

H. Harris in Condor, 43:51, 1941; cf. Pac. Coast Avifauna, 5:153, 1909, for his papers.

Thouars, Abel Aubert du Petit, 1793-1864

Lasegue, 385-386; cf. J. T. Howell in Leafl. West. Bot, 1:189-191, 1935; J. Espasa,
Enciclopedia Univ. Ilustrada; C. Nissen, Bot. Buchillustration, 2:54, 1951.

TiDESTROM, IVAR, 1865-

Amsci; Ewan, 321; Rydberg, 45; M. E. Jones, Contr. West. Bot, 15:20-24. 1929;
autobiog. notes in I. Tidestrom and Sister T. Kittell, Flora of Arizona and New Mexico,
ix-x, 1941.

Tiling, Heinrich Sylvester Theodor, (?)-1871

Hulten, 301; cf. Bradley Bib., 1:455 for Florula ajanensis, in collaboration with E.
Regel. Regel described Horkelia tilingi and Mimulus tilingi from his collections taken
in vicinity of Nevada City.

TORREY, Harry Beal, 1873-

Amsci; cf. H. Kirby in Sci. Mo., 61:416, 1945.

EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 61

ToBRET, John, 1796-1873

Brewer, 558; DAB; Ewan, 322; Kelly, 136-144; Meisel, 1:234 and 3:666-667;
Rodgers' Torrey, passim; Torrey's visit to the Academy seems not to have been
chronicled.

TowNSEND, Charles Haskins, 1859-1944

Dean, 2:549-550; Hulten, 306; NCAB, 32:37 (portr.) ; Palmer, 298; T. S. Palmer
in Auk, 64:349-350, 1947; D. S. Jordan, Days of a Man, passim, 1922; autobiog. in Con-
dor, 29:224-232 (portr.), 1927.

TowNSEND, John Kirk, 1809-1851

Alden, 40-42; DAB; Dall, 41 et passim; Dean, 2:550; Ewan, 323; Meisel, 1:235;
Palme?; 299; Stone, 7-11; Wagner-three, 79; W. Stone in Cassinia, 7:1-5 (portr.), 1903;
F. W. Pennell in Bartonia, no. 18, 35, et passim, 1936; H. Harris in Condor, 43:21-23,
1941; cf. Stone in Auk, 47:414-415, 1930; cf. J. Grinnell in Univ. Calif. Publ. Zool.,
38:269-270, 1932; though Townsend is intimately associated with California natural
history he did not visit the State.

Trask, John Boardman, 1824-1879

Jepson, 2:117-118, 1924; Meisel, 1:235; A. W. Vodges in Trans. San Diego Soc. Nat.
Hist, 1:27-30, 1907; R. E. C. Stearns in Science, 28:240-243, 1908.

Trowbridge, William Petit, 1828-1892

DAB; Dall, 299; Palmer, 300; C. B. Comstock in Nat. Acad. Sci. Biog. Mem., 3:363-
367, 1895.

Tschernikh, George, fl. 1835-1841
Essig, 772-773.

TUBEIUF, KaEL von

Cf. Madrono, 1:12-18 (portr.), 1916.

Vancouver, George, 1758-1798

Brewer, 553; DNB; Eastwood, 336; E. S. Meany, Vancouver's Discovery of Puget
Sound, 7-21, et passim (portr.), 1915; Geo. Godwin, Vancouver: a Life, 1758-1798, 1930.

Van Denbitegh, John, 1872-1924

Amsci, ed. 3; edit, note in Condor, 27:83, 1925; D. S. Jordan, Days of a Man, 1:541
and 710, 1922. .

Van Duzee, Edward Payson, 1861-1940

Amsci, ed. 6; H. Osborn, Fragments of Entom. Hist., 1:234 (portr., pi. 5), 1937.

Van Dyke, Edwin Cooper, 1869-

Amsci; Hulten, 323; H. Osborn, Fragments of Entom. Hist, 1:284 (portr., pi. 28),
1937; W. M. Mann, Ant Hill Odyssey, 79-80, 1948.

Vasey, George Richard

Brewer, 559; Ewan, 327; HarsWberger, 385; Piper, 18; cf. J. T. Howell in Amer.
Midi. Nat, 30:33-35, 1943.

Veatch, John Allen, 1808-1870

Bradley Bib., 5:876; Geiser-two, 282; cf. mimeo. letter addressed to W. P. Webb, ed..
Southwestern Hist. Quart., dated 18 Sept., 1942, circularized by S. W. Geiser, relating
to Veatch 's genealogy; P. A. Munz in Leafl. West. Bot., 7:70, 1953; cf. Amer. Journ. Sci.,
ser. 2, 26:288-295, 1858, for Veatch's account of mud volcanoes of Salton Sea; Hesperian,
2:21-26, 1859, for his account of Clear Lake, Calif., and ibid., 3:529-534, 1860, for his
account of Cerros (i.e., Cedros) Island.

Vollmer, Albert Michael
Woodcock d Steam, 361.

62 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

VosxESENSKY, Ilya Gavriloch, 1816-1871

Brewer, 555, as "Wosnessensky"; Eastwood, 338; Essig. 777-789 (poitr.); Hulten.
300; cf. J. Grinnell in Univ. Calif. Publ. ZooL, 38:321, 1932.

Vries, Hugo de, 1848-1935

A. D. Rogers, III, Liberty Hyde Bailey, jyassim, 1949; E. Nordenskiold, Hist. Biol. (A.
Knopf, ed.), 587-588, 1928, and other general histories of science; portr. in Chron. Bot..
9(5-6), pi. 22, 1946, at Burbank's garden: autobiog. notes in Naar Californie, Amster-
dam, 1905, and transl. in Pop. Sci. Mo., 67:329-347, 1905.

Wallace, Alfred Russel, 1823-1913

Britten, 314; Carpenter, 109; Dean, 2:599-600; DNB. 20th Cent. Suppl. 546; Eican,
330; Musgrave, 337; Urban Fl. Bras., 130-131; Van Steenis. 555-557; L. O. Howard,
Fighting the Insects, 303-305, 1933; W. S(tone) in Auk, 31:138-141, 1914; D. S. Jordan,
Days of a Man, 1:303, 1922; T. D. A. Cockerell in Pop. Sci. Mo., 62:517-518, 1903.

Walther, Eric

V. Reiter, Jr., in Leafl. West. Bot., 7:82-83, 1953.

Wei-.ber, David Gould, 1809-(?)

J. Ewan in A. S. Hitchcock, Man. Grasses U. S., ed. 2 (U.S.Dept. Agri. Misc. Publ.
200), 989, 1951.

Webber, Herbert John, 1865-1946

Fairchild, 444, et passim, and World Grows round my Door, passim. 1947; H. S.
Reed in Madroiio, 8:193-195, 1946, portr. as frontis. to Vol. 8.

Whipple, Amiel Weeks, 1816-1863

DAB; Howell, 30; Etvan, 335; G. Foreman, A Pathfinder in the Southwest, Norman,
Okla., 1941.

Whitney, Josiah Dwigiit, 1819-1896

ACAB; Bade, imssim; Brewster. 206-207, 239-240, 272-273, et passim: DAB; Etvan.
336; Meisel, 1:239; Palmer, 302; brief acct. of Calif. Geol. Surv. in No. Amer. Rev.,
121:63-64, 1875.

WiCKSON, Edward James, 1848-1923

DAB; Fairchild, 302; W. L. Howard in Chron. Bot., 9:314-316, et passim (portr.,
pi. 22), 1946.

Wilkes, Charles, 1798-1877

Brewer, 551; DAB; Dall. 71-72, et passim; Eivan. 337; Hughes. 15; Meisel, 1:240
and 3:678; Urban Fl. Bras., 1:144-145; Van Rteenis. 575-577; J. D. Hill. Sea Dogs of
the Sixties, 88-127 (portr.), 1935; H. H. Bartlett in Proc. Amer. Philos. Soc, 82:601-
705, 1940; M. E. Cooley, ibid., 82:707-719, 1940.

Williams, Francis Xavier, 1882-

Amsci; Musgrave, 354; portr. in Pacific Discovery, 6(2): 22, 1953.
Williamson, Rouert Stockton, 1824-1882

Hume, 195, et passim; Palmer, 303.

Wislizenus, Frederick Adolph, 1810-1889

Brewer, 557; Dall, 180, 258, 265; Ewan, 338; Geiser-two, 283; Meisel. 1:242 and
3.680; Sargent, 6:94; Wagner-three, 83; Geo. J. Engelmann in St. Louis Acad.
Sci. Trans., 5:464-468, 1892; F. Starr in Pop. Sci. Mo., 52:643-644 (portr.), 1898; P.
Spaulding, ibid., 74:244-246 (portr.), 1909; biog. sketch by his son in Mo. Hist. Soc. ed.
of W's Journey, 5-13 (portr.), 1912.

WiSTAR, Isaac Jones, 1827-1905

NCAB, 12:359 (portr.); see Autobiography, Phila. 1914, reissued in 1937 and again
in 1938.

fW-AN: SAN FRANC/SCO -AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 63

Wood. Alphonso, 1810-1881

ACAB; Brewer, 558; Meisel, 1:242; 1\CAB, 14:278; O. R. Willis, A Biographical
Sketch of Dr. Alphonso Wood, of West Farms . . ., 6 pp. (n.d.), examined in N. Y. Bot.
Gard. Library; anon, in New York [Times?] for April 19, 1903 (portr.); C. J. Lyon in
Dartmouth Alumni Mag., 31:18, 81-82, March, 1939, and in Science, 101:484-486, May
11, 1945; Woodcock d- Steam, 364.

WooDHOUSE, Samuel Washington, 1821-1904

Eioan, 340; Geiser-tivo, 283; Hume. 469-509 (portr.), the fullest act.; Palmer, 303;
Sargent, 8:88; Wagner-three, 230; W. Stone in Cassinia, 8:1-5 (portr.), 1904; W. S(tone)
in Auk, 22:104-106, 1905.

W^ooDWORTH, Charles William, 1865-1940
Carpenter, 114; Essig, 800-802 (portr.).

Wrangell, Ferdinand Petrovich, 1794-1870

Brewer, 554, as Wrangel; Embacher, 299, as "1795-1870"; Essig, 802; Hulten, 300.

Wright, Charles, 1811-1885

Brewer, 558; DAB; Eicon, 342; Geiser-tioo, 283; Urban Symb. Ant., 141; C. S. Sar-
gent, Sci. Pap. of Asa Gray, 2:468-474, 1889.

Wright, William Greenwood, ca. 1830-1912

Carpenter, 115; Essig, 802-804 (portr.); Hulten, 308; J. D. Gunder in Entom. News,
40:33-34 (portr.), 1929; F. Grinnell, Jr., in Entom. News, 24:91-92, 1913, and Bull. So.
Calif. Acad. Sci., 12:19-21, 1913; his Butterflies of the West Coast, San Francisco, 1905,
is a rare book from the destruction of the stock in the 1906 fire.

Xantus de Vesey, Lons John, 1825-1894

Breioer, 558; Embaclier. 300; Essig, 804-808, map; Geiser-tico, 283, s.v. "Wiirttem-
berg"; Hume, 510-532 (portrs.), useful acct.; Meisel, 1:244 and 3:743; Palmer, 304;
Wagner-three, 316 ; Henry M. Madden, Xantus, Hungarian Naturalist in the Pioneer
West, Palo Alto, 1949, the latest full-length biog.; J. Grinnell, J. S. Dixon, and J. M.
Linsdale, Fur-bearing Mammals of California, 1:76-77, 1937.

A CENTURY OF ASTRONOMY AND GEODESY

IN CALIFORNIA

By ERWIN G. GUDDE

University of California

Until 1769 California remained a geographical conception. Navigators — with
the exception of Francis Drake, all Spanish or in Spanish service — had sailed
up and down the coast, but they had come, not for scientific observation, but in
search of fabulous rich lands, of booty on the high sea, of harbors in which the
Manila galleon could find safety. Their observations of latitude and longitude
were completely inadequate and caused cartographers for two centuries to indulge
in imaginary geography of the somewhat mythical land, "California."

In 1769 the land route to California was opened up by the Portola expedi-
tion and during the next half-century the Spaniards made California into a
Spanish colony. The representatives of Spanish imperialism who created the
new province were the officers of the military detachments and the Franciscan
fathers. There were among the latter some personalities — Crespi, Garces, Palou
— who left their mark upon California history because in the vastness and new-
ness of the territory they were the only ones who could read, write, and observe.
They lacked, however, the scientific fervor which, in addition to the religious
fervor, had distinguished their predecessors on the American continent — the
Jesuits. Hence the few astronomical and geodetic data left by these padres are
negligible and unimportant. After California became a Mexican province and
until the occupation by the United States not even a trace of scientific activity
existed in California.

Whatever scientific work was done before United States scientists began
their task was accomplished, not by the Spaniards or Mexicans, but by the for-
eign navigators and explorers: La Perouse, Vancouver, Kotzebue, Belcher,
Beechey, Wilkes, Fremont. Indeed, Beechey's geodetic and hydrographic work
of San Francisco Bay was so accurate that the United States Coast Survey, when
it started its work in 1850, allowed the resurvey of the harbor to wait and under-
took other tasks which seemed more pressing.

Real astronomic and geodetic work began with the end of the Mexican War,
It was mainly army engineers who began the great task of establishing the
boundaries, surveying the land, and examining and evaluating its resources and
possibilities. Greatly accelerated was the progress of these tasks when California
suddenly moved into the center of world interest after the discovery of gold.
Next to gold-seekers, traders, and lawyers (who reaped a rich harvest in con-
nection with the disposal of the land grants), the engineers formed the largest
contingent of professions that descended upon California. The coast had to be
made safe for navigation, base lines had to be established, land grants measured,

[65]

66 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

transportation established, minerals assayed, resources investigated — tasks for
every type of engineering. All scientific knowledge available at that time was
used for practical purposes. Science for science's sake was unknown in those
hectic years following the Gold Rush. Astronomy played a role only in so far
as the elements of the science were essential to the geodetic work necessary to
create the basis for material culture.

In 1848 the United States Coast Survey, one of the most efficient Federal
agencies, then under the direction of Alexander Bache, Benjamin Franklin's
grandson, decided to start the survey of the Pacific Coast in the following year.
A hydrographic and a geodetic party, both well equipped, arrived in California
in 1849. Both came to naught; the lure of the goldfields proved to be too strong
for the underpaid employees of the Government.

In 1850 George Davidson and a group of stalwart young members of the
Coast Survey arrived in San Francisco. They had volunteered to go to the
Pacific Coast out of cheerful, youthful exuberance. For almost half a century
Davidson was one of the leading figures in the evolution of the State. In the
development of the sciences of astronomy, geodesy, geography, and seismology
in California he dominated the scene. The Coast and Geodetic Survey, the Cali-
fornia Academy of Sciences, and the University of California owe much to this
indefatigable, universalistic, and, above all, practical scientist.

The auguries, to be sure, were not very encouraging. The journey of the four
young geodesists — Lawson, Harrison, Rockwell, and Davidson — consumed one-
fourth of the year's allotment for the Pacific Coast work. In San Francisco they
soon realized that their salary of $800 per annum would not last very long if
they had to pay $7.50 for room and board per diem. They had to bivouac with
their 2,500 pounds of instruments in a 12 by 12-foot room. The water for ablu-
tion and for washing their shirts they carried from a spring four blocks away.
The only mechanic in the city charged them $900 for making four larue foot
screws and for tapping the cast-iron frame of the large transit instrument.

Davidson resisted the temptation to start the survey of the Golden Gate and
San Francisco Bay. He realized that Beechey's survey was good and that other
points along the coast needed urgent attention. The islands of the Santa Bar-
bara Channel were badly located, the position of Point Conception was in error,
and thither the party embarked the end of June, 1850. There, at El Cojo. the
real hardships began. The Mexican cook promptly absconded with their horse
and the party had to cook their steaks and fiapjacks over a fireplace made of
three whale vertebrae and fed by dry cattle chips, and to do all other chores
necessary to maintain the most primitive essentials of human existence.

But the work was done. Three months and a half were spent in astronomical
observations for the latitude and longitude of the station. The observations
included lunar transits, occultations of stars by the moon, and one solar eclipse.
In spite of the fog, Davidson could observe for sixty nights until he was "heartily
sick of starlight." Returning to San Francisco in October, the party worked
systematically on the reductions of the field observations. Their preliminary
work proved so satisfactory to the Superintendent of the Survey that he pro-
cured an extra appropriation for the party. Assistants and laborers could be
hired and the work at the second station. Point Pinos, could be carried on under
more agreeable circumstances during January and February, 1851.

GUDDE: A CENTURY OF ASTRONOMY AND GEODESY IN CALIFORNIA 67

As the third station, Davidson selected San Diego, because its latitude on
the existing charts was completely erroneous.

At this port [he wrote], I made the usual astronomical observations of lunar transits,
occultations of stars by the moon, latitude observations, azimuth observations for the
triangulation, determination of the magnetic elements, etc., working the greater part
of the night and computing the greater part of the day. I had undertaken work on this
coast to make a record in a new field, and therefore labored nearly to the utmost strain
of my energies, never less than eighteen hours a day.

After this first year of astronomic observation and determination the work
of the United States Coast Survey was carried on with ever-increasing speed,
volume, and variety. Until his retirement in 1895, except for the years during
and after the Civil War, which were spent chiefly in war work at Philadelphia,
Davidson was in charge of the astronomic, geodetic, topographic, and hydro-
graphic work of the Pacific Coast and later also of the coast of Alaska.

Besides the practical work the members of the Coast Survey inaugurated
astronomical observation on the Pacific Coast. While at Monterey Bay in the
winter of 1850-1851, Davidson began his computation of the star factor tables,
which were later published. In 1852 he discovered and observed a brilliant comet
at Astoria on the Columbia Eiver. The solar eclipse of May 26, 1854, was
observed by members of the Survey at Benicia, Loma Prieta, and Humboldt Bay.
Davidson also observed the solar eclipse of March 25, 1857, in San Francisco.
In 1856 he published the "Occultation of Stars by the Moon on Western Coast of
the United States," and in the following year "The Occultation of 22 Stars of
the Pleiades, and Solar Eclipse of 1857."

The crowning achievement of Davidson during his first phase of Pacific
Coast Survey was a practical work, the Directory of the Pacific Coast, first pub-
lished in 1857. This work, republished at irregular intervals and later called
Coast Pilot of California, Oregon and Washington, systematized the astronomic,
geodetic, hydrographic, and topographic work of the Coast Survey and became
the bible of the mariners who sailed up and down the Pacific Coast.

The year in which Davidson left San Francisco, 1860, witnessed the first
attempts of astronomical observations by agencies other than the United States
Government. To the University of Santa Clara belongs the honor of being the
first educational institution of the State to acquire a telescope. The 4-inch
refractor with altazimuth mounting, installed in 1860, was the nucleus of an
observatory which in later years became well known, especially through Jerome
Eicard's observations of sun spots and faculae.

In the same year an amateur astronomer, George Madeira, started observing
with a 3-incli refracting telescope with equatorial mounting at Volcano, Amador
County. According to Campbell, on June 30, 1861, Madeira discovered the
brilliant Comet 1861 II only a few hours after its discovery in Europe.

In the meantime other agencies were at work surveying the State. A United
States Commissioner of the General Land Office was sent to California shortly
after the treaty of Guadalupe Hidalgo had been signed. The principal tasks
of his office were the establishment of the extent of the Spanish and Mexican
land grants and the division of the newly acquired territory into townships.
The commissioner established the three township base lines and meridians: the
Mount Diablo, the San Bernardino, and the Humboldt, which have formed the

58 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

skeleton for land-measuring purposes ever since. While the Land Office did
extremely valuable work for the future development of the State, unlike the
Coast Survey it did not contribute to the advancement of astronomy and scien-
tific geodesy.

Another Federal project consisted of the explorations and surveys to ascer-
tain the most practical route for a railroad from the Mississippi to the Pacific,
undertaken in 1853-1854 under the direction of the United States War Depart-
ment. The result of this well-equipped project was published in a Report of
thirteen imposing volumes — a great contribution to the geography and cartog-
raphy as well as to the natural conditions and resources of the American West.
At no less than 174 stations, including many in California, astronomical obser-
vations were made and the latitude, longitude, and magnetic declinations of
many places were determined. The tables of these observations were published
in the second volume of the Report and formed a valuable basis for future sur-
veys, especially for the heretofore neglected mountainous and desert regions of
the State.

The government of the State likewise participated in the geodetic delineation
of California. The office of Surveyor General of the State of California, founded
in 1850, published annual reports. In 1860 the legislature established the State
Geological Survey, which carried on its tasks for fourteen years until a new
political constellation put a sudden end to its work, so that not even its valuable
maps could be completed. The principal work was carried on by four great men
in the fields of geodesy, geology, and topography, Josiah D. Whitney, Clarence
King, Charles F. Hoffmann, William H. Brewer.

The work of the Coast Survey continued, and its annual reports bear witness
to the excellent achievements of its members. It received a new impetus when
in 1868 Davidson was again put in charge of the survey on the Pacific Coast, an
assignment which he continued uninterruptedly until 1895.

During the eight years of absence from San Francisco, Davidson had achieved
national recognition. He had participated in the War between the States in
various capacities, had been the engineer of a party sent to Panama to examine
the possibility of a canal through the isthmus, and had been sent to Alaska by
the State Department to make a survey of the territory preliminary to the con-
summation of its purchase by the United States.

With renewed vigor Davidson took up his various tasks. Soon after his return
he became intimately connected with two California institutions to which he
remained devoted till the end of his life : the University of California and the
California Academy of Sciences. In 1870 he was elected Professor of Astronomy
and Geodesy, in 1877 he became a Regent of the University, and after his retire-
ment from the Coast Survey he was appointed Professor of Geography; a year
before his death he received the degree of Doctor of Laws.

His first contribution to the Proceedings of the California Academy of
Sciences was a report on the "Observations of the Meteors of November 14, 1869,
at Santa Barbara." In the course of years he contributed about thirty papers
on astronomy and geodesy alone to the periodical publications of the Academy.
In 1872 he was elected President of the Academy, an office which he held for
fifteen years.

In his capacity as President he visited James Lick to convey the thanks of the

GUDDE: A CENTURY OF ASTRONOMY AND GEODESY IN CALIFORNIA 69

institution for a most munificent endowment, the valuable corner lot of Fourth
and Market Streets, which Lick had deeded to the Academy on February 15, 1873.

Of the many strange characters who had come to California in the early days,
James Lick was perhaps the most peculiar. Whereas thousands rushed to Cali-
fornia to make a fortune, Lick arrived in the early part of 1848 bringing with
him a handsome capital, acquired through twenty years of hard work as a cabinet-
and piano-maker in South America. In another twenty years he greatly increased
this fortune and decided to spend it for the benefit of his adopted state and for
the glorification of his name.

Upon Lick's request Davidson repeated his visits and was finally let in on
a secret : Lick wanted to create a new world wonder by erecting a telescope much
larger and much more powerful than any in existence. The somewhat conserva-
tive Davidson soon realized that Lick had strange ideas about such a telescope,
that he expected that it would provide spectacular discoveries in the universe, and
that it would be a world-wide attraction. Davidson's first task was to guide Lick's
enthusiasm in the right direction. He did this with tact and understanding. If
in the end he did not succeed entirely, it was not his fault.

Before the location of the observatory was discussed by the cautious Davidson,
a mutual friend. Dr. Frederick Zeile, the pioneer of the bathtub in San Francisco,
informed him that Lick had made up his mind to build the observatory at Fourth
and Market Streets in San Francisco, between the sites he had given to the
Academy of Sciences and the Pioneer Society. In front of the observatory he
planned to erect three statues: one for Francis Scott Key (the one now stand-
ing in front of the Academy of Science buildings in the Golden Gate Park), one
for Thomas Paine, the pioneer of atheistic thought in America, and one for Lick's
own grandfather, who had once shared the trials of Washington's revolutionary
army in Valley Forge. It took Davidson several months of diplomatic and per-
sistent argument to convince Lick that, though downtown San Francisco would
doubtless be the ideal spot to attract tourists to his spectacular show piece, it left
much to be desired as a site for scientific research in astronomy. Gradually he
guided Lick's judgment to place the observatory in the Sierra Nevada — not on
one of the high peaks where conflicting upper air currents would be detrimental
to astronomical observation but near the summit of Donner Pass.

On October 20, 1873, Davidson announced at the monthly meeting of the
Academy that Lick had agreed to his proposals and to the erection of an observa-
tory with "a telescope superior to and more powerful than any telescope yet
made." The next morning the Alta Calif ornian, in a three-column spread on the
front page, imparted the news to the world. Since the announcement of the dis-
covery of gold no more exciting intelligence had come from California, and the
names of Lick and Davidson were as much in the mouth of the people as the
names of Sutter and Marshall had been twenty-five years before. The young
state, which many still associated with lawlessness, fraudulent land grants, and
unscrupulous lawyers, was suddenly to take the lead in the study of an important
field of human knowledge.

Davidson's task, however, was not yet done. Next he had to dissuade Lick
from building a reflector telescope. This type of telescope, an invention of Isaac
Newton, had just then been greatly improved and was especially favored in
England. Davidson, however, as well as the majority of American astronomers,

70 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

considered the refractor type superior. When we realize the marvelous results
obtained by reflector telescopes at the Lick, Mount Wilson, and Palomar ob-
servatories we have to admit that James Lick, the half-educated Pennsylvania-
Dutch piano-maker, had the right instinctive vision and that Davidson and the
other American astronomers were wrong in their conviction that a refractor of
limited size would be superior to the immense reflector Lick had proposed.

After the latter had agreed to a refractor telescope he wanted one six feet in
diameter, and Davidson had to convince him that a 40-inch objective would be
the maximum possible size of a refractor. The question of the amount of money
necessary caused more difficulties, because Lick could not see that an observatory
needed other equipment besides a giant telescope. He believed that Davidson's
figure of $1,500,000 was too high but finally agreed to spend $1,200,000 on the
project.

In May, 1874, Davidson went East to confer with astronomers about the
preparations for the observation of the transit of Venus in December. During
his absence other influences gained the confidence of Lick, who decided to build
the observatory on the shores of Lake Tahoe, where the name Observation Point
still marks the chosen site. (As it turned out, Lick's advisor owned a quarter-
section of land adjacent to the Point.) Davidson succeeded in convincing Lick
of the unsuitability of this site but his patience was by this time rather taxed
by the donor's constant vacillations. He made no further attempts to influence
Lick when the latter cut down the endowment to $700,000 and chose Mount Ham-
ilton, 4,209 feet elevation, as the site for the observatory. Mount Hamilton, named
in 1861 for an Oakland independent clergyman, the Eeverend Laurentine Hamil-
ton, was, according to some astronomers, much better suited than Davidson's
favorite spot near Donner Pass.

The work on the observatory could not begin until the Lick estate was liqui-
dated in 1879. In 1888 the great project was completed and was given to the
University of California, as provided by Lick in his final deed of trust. The
36-inch equatorial refractor was at that time the largest in the world and the
general equipment of the observatory was second to none. Within a few years
the fifth satellite of Jupiter, the revolving sun of the Procyon, and a large num-
ber of comets and double stars were discovered. For the first time the angular
diameter of a fixed star was measured and epoch-making work was done by spec-
troscopic observation of stars, nebulae, and comets. This is not the place to
attempt to enumerate the achievements of the distinguished astronomers con-
nected with the Lick Observatory. Its various periodical publications give the
record.

There is no question that the project of an observatory of the size and equip-
ment of the Lick Observatory was a healthy stimulus to astronomical interest in
the world. In California itself observatories began to mushroom even before the
Lick Observatory was completed.

The first scientifically constructed observatory was erected by George David-
son in San Francisco for special study of the physical features of the planets,
and later for observing the variations of latitude and determining the constant
of aberration. Davidson had made astronomical observations on Washington
Plaza since 1870. In 1879 he removed his station to Lafayette Square, equipping
it with a 6.4 Clark refractor, a chronograph, and a telegraphic apparatus. Here

GUDDE: A CENTURY OF ASTRONOMY AND GEODESY IN CALIFORNIA 71

Davidson often observed until the small hours in the morning, and that after
his strenuous duties with the Coast Survey.

As a labor of love [says Campbell J, Professor Davidson undertook the observations
of latitude pairs of stars at his observatory. Between May, 1891, and August, 1892, he
secured for this purpose, 5.308 observations on 283 stars. . . . His results were in good
agreement with those obtained at European, Atlantic coast, and Hawaiian stations.

The results of his observations he published in luunerous articles in the publica-
tions of the California Academy of Sciences, the Royal Astronomical Society,
and tlie United States Coast Survey. The observatory remained on Lafayette
Square until ]902. Its principal instrument is now at Chabot Observatory.

The sudden interest in astronomy naturally also had great influence upon
astronomy as a subject of instruction in our schools. Davidson himself again
took the lead by inviting high school students and their teachers to his observa-
tory, and thus he aroused in the young intellects an interest in the wonders of
the universe. In 1883 Anthony Chabot presented to the Oakland School Depart-
ment his well-known observatory with an 8-inch refractor, to which the Board
of Education added in 1913 a 20-inch refractor. In 1885 the College of the Pacific
received an observatory with a 6-inch Clark equatorial. The Students' Observa-
tory of the University of California was erected in 1886, and in 1892 was placed
in charge of Armin 0. Leuschner. It has since been the elementary training
ground for many astronomers who have achieved fame in their profession. Only
a year later Mills College received its observatory with a 5-inch refractor and an
8-inch reflector, and in 1890 Napa College started its astronomy department with
an 8-inch Clark-Saegmuller refractor, which was later acquired by the University
of Santa Clara.

However, the hopes of the University of Southern California to outdo the
Lick Observatory by having an observatory with a 40-inch refractor telescope
were shattered. The donor died shortly after the discs were given and insufficient
funds prevented the University from erecting the observatory. The discs were
purchased in 1893 by C. T. Yerkes and became the nucleus of the famous observa-
tory of the University of Chicago ! The chief factor in this move was no other
than George Ellery Hale, destined to play a most important role in the develop-
ment of astronomy in California. Since then astronomy has become a subject
generally taught, and most colleges and many high schools have their own observ-
atories.

George Davidson continued to play an important role in the geodetic work
of the State, as in the field of astronomy. Between 1875 and 1879 Captain George
M. AVheeler, Corps of Engineers, United States Army, had carried on the "Geo-
graphical Surveys AYest of the One Hundredth ]\Ieridian." On March 3, 1879,
the United States Geological Survey was established under the Department of the
Interior and began its great work of creating the topographical atlas of the
United States. Important as were the Wheeler Survey and the Geological Survey
— and later the United States Forestry Survey and the United States Corps of
Engineers — for the scientific delineation of California, the extension of the scope
of the Coast Survey was of much greater value in the line of applied astronomy.
The Coast Survey, heretofore responsible for the survey of our coasts, was
assigned in 1879 the tremendous task of the trigonometrical survey of the United
States.

72 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Davidson, who was, so to speak, at the western end of the arc of the 39th
parallel, which extends 2,825 miles from the Atlantic Coast, entered upon his new
duties with renewed vigor. Observation lines of triangulations used by him
reached the length of almost 200 miles — a feat at that time "unique in the history
of geodesy," as the Superintendent of the Coast and Geodetic Survey approvingly
stated.

The crowning achievement of Davidson's career was the measurement of the
two base lines upon which the triangulation of California rests. In 1881 he meas-
ured the Yolo Base Line twice, with the result that the probable error, as com-
puted by his collaborator C. A. Schott, was 9.57 millimeters on a line measuring
17,486.5 meters — a minimum of error probably never equaled under similar cir-
cumstances. The story of this unusual feat may be found in the U. S. Coast and
Geodetic Survey Reports of 1882 and 1883. In 1888-1889 Davidson repeated this
performance by measuring the Los Angeles Base Line three times.

The final achievement of the Coast and Geodetic Survey during Davidson's
incumbency was the definite establishment of the California-Nevada boundary.
California's boundaries with Oregon and Mexico had been established without
difficulty, though not without error. The Nevada line remained for several decades
a problem. Captain Sitgreaves began the survey in 1852, G. H. Goddard con-
tinued it in 1855, J. F. Haughton ran the line from Lake Tahoe to a point east
of Mono Lake in 1863, and James Lawson extended it to beyond AVhite Moun-
tain. In 1872-1873, San Francisco's pioneer engineer, A. W. von Schmidt, finally
ran through the entire line. After the Coast Survey had been placed in charge
of the inland triangulation, an error was discovered in checking the initial start-
ing point at Lake Tahoe. In the final survey, begun in 1893, von Schmidt's line
south of Lake Tahoe was moved west several miles.

The total solar eclipse of January 1, 1889, helped to augment the interests of
Californians in astronomy. About six scientifically equipped parties and numer-
ous amateurs observed the phenomenon. The Astronomical Society of the Pacific
was organized the same year. Well supported, it soon became one of the strongest
organizations devoted to science.

In 1894 the second mountain observatory was erected north of Pasadena on
Echo Mountain, a shoulder of Mount Lowe, at an elevation of about 2,500 feet.
The telescope was a 16-inch refractor, with which Lewis Swift, its owner, had
discovered 960 nebulae and nine comets in Rochester, New York. During the
next six years, as director of the Mount Lowe Observatory, Swift discovered 230
additional nebulae and five other comets.

The third observatory to be established on a California mountain is on Mount
Wilson, 5,710 feet above sea level. It was upon S. P. Langley's recommendation
that the Carnegie Institution of Washington provided the funds for the estab-
lishment of this observatory. Langley, the director of the Allegheny Observatory,
had done extensive work in solar radiation and wished to check the influence of
the vapor and dust content at low altitude as compared to conditions at very
high altitude. "A southern latitude," he wrote to Davidson on May 30, 1881,
"dry climate, and above all, clear deep blue sky. Another important thing is the
provision of an adjacent station having great difference of altitude. All these
conditions seem to meet at Whitney." From July to September, 1881, Langley's
party, among them James Keeler, subsequently Director of the Lick Observatory,

CUDDE: A CENTURY OF ASTRONOMY AND GEODESY IN CALIFORNIA 73

observed from three stations, Mount Whitney, 14,496 feet, Mountain Camp,
11,600 feet, and Lone Pine, 3,727 feet high. The success of Langley's party led
to a number of other observations on Mount Whitney, especially after the Smith-
sonian Institution had erected a suitable building on the summit, the lack of
which had been felt by the Langley party.

It was in 1902 that the Carnegie Institution of Washington was founded. In
1904 steps were taken by the Institution and by Dr. George E. Hale (who nego-
tiated the first lease) preliminary to the actual establishment of an observatory
on Mount Wilson. In 1905 the Carnegie Institution made the first grant for the
building and maintenance of the Mount Wilson Solar Observatory. The usual
controversy among astronomers had arisen about the desirability of altitude for
astronomical observation. A committee of leading astronomers arrived apparently
at a compromise, suggesting Mount Wilson, which had already been occupied by
a Harvard University party from 1889 to 1891. At the same time the committee
recommended a 60-inch reflector telescope as most suitable. Hale, the chief
advocate of the reflector telescope, was appointed director. With that a new
phase in the history of astronomy was ushered in, and California was again
in the lead.

In 1898, James E. Keeler, Director of the Lick Observatory, had already
shown the superiority of the reflector for discovering nebulae and star clusters
by means of photography. With the comparatively small auxiliary reflector at
Lick Observatory hundreds of new nebulae were discovered in a small section
of the sky, which led to the conclusion that hundreds of thousands of nebulae
existed and awaited discovery.

This method of observation was now employed at Mount Wilson on a larger
scale and with more powerful telescopes : first a 60-inch, and then, since 1918, a
100-ineh reflector. It is, of course, here impossible even to summarize the spec-
tacular results obtained on Mount Wilson in solar research, stellar distances and
velocities, spectroscopy, compositions of star clusters and nebulae, and so forth.
New was Hale's idea of considering an observatory as a huge physical laboratory
of which the telescope forms only one part — the most essential, to be sure.

Even before the installation of the 100-inch reflector Hale had visions of a
more powerful telescope and with his energy and perseverance he w^ent about to
make his dream come true. After the usual trials and tribulations the trustees
of the Rockefeller Foundation in 1928 voted the sum of $6,000,000 for the erec-
tion of a 200-inch reflector. The marvelous results on Mount Wilson had shown
that the peaks of the Southern California mountain ranges offered the best atmos-
pheric conditions for astronomical observation in the United States. Palomar
Mountain, in San Diego County, 6,126 feet above sea level, was selected as the
site for the new telescope, which was to penetrate more deeply into space.

Palomar, "place of the pigeons," is a remarkable orographic feature for
which even the Indians had a name, "Paauw." American surveyors named it
Palomar after a Mexican land grant, but generally it was known as Smith Moun-
tain until the Board on Geographic Names restored the beautiful old Spanish
name in 1901.

Hale died ten years before the completion of the great work of which he had
been the chief mover. When the observatory was dedicated in 1948, the immense
instrument was named Hale Telescope in his memory. The Palomar Observatory

74 -A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

is operated jointly with the Mount Wilson Observatory by the Carnegie Institu-
tion and the California Institute of Technology. The Hale telescope together
Avith its essential auxiliary, the 48-inch Schmidt telescope, will continue in the
lead of exploring the mysteries of the universe.

Astronomy may be said to blend more with the main stream of general culture
than any other science. Many of our great astronomers were not professional
men but began as amateurs who took up the search into cosmic existence as a
hobby, and there are innumerable other laymen who are interested in the vari-
ous phases of astronom^^ California has not been remiss in satisfying this interest.
At Lick, Griffith, and some smaller observatories special nights are set aside
when the general public may get a glimpse of the heavenly bodies and their
motions. The most suitable invention to arouse the public's interest in astronomy,
the planetarium, is represented in California by the Griffith Planetarium in
Griffith Park, Los Angeles, and the Morrison Planetarium, a unit of the Cali-
fornia Academy of Sciences in San Francisco. California is thus the only state
in the Union which possesses two planetariums, one of which, although based on
the principle developed by the famous Zeiss Works, was entirely constructed,
assembled, and mounted in the shops of the California Academy of Sciences.

The great variety of California topography as well as its historic background
made the State in its infancy a successful testing ground for geodetic work;
climatic and atmospheric conditions, the generosity of its citizens, and the
enthusiasm of its people have contributed significantly toward making California
the leading commonwealth in the science of astronomy.

SOURCES

The George Davidson papers, Bancroft Library, University of California, Berkeley.

The reports and publications of the Government agencies and the institutions men-
tioned in the text.

W. W. Campbell, "A Brief History of Astronomy in California" in The History of
California (1914). New York: The Century History Company.

Helen Wright, Palomar (1952). New York: Macmillan Co.

THE CONTRIBUTION OF NATURAL HISTORY
TO HUMAN PROGRESS

Btj G. F. FERRIS

Stanford University

The ^Ieanings of words quite commonly change over a period of time and a
meaning- that may have been current a hundred years ago may now be obsoles-
cent or even obsolete. So with the meaning of the words "natural history." If
we look back at the history of the development of biology, those words carried a
meaning a little over a hundred years ago that subsumed almost everything that
was then known about plants and animals, since what was then known, apart
from some small amount al)out human anatomy as a subject entirely by itself,
was mostly concerned with the questions of how many and how different were
the various kinds of organisms on the earth. It considered to some small degree
the manner in which those organisms were grossly put together, for a knowledge
of this was involved in determining how varied they might be. Work had been
done also in what we now call comparative anatomy, but this comparative anat-
omy, lacking the stimulating influence of the idea of evolution, really involved
nothing much more, even in the work of Cuvier, than a recital that in certain
kinds of animals certain structures were to be seen and in other kinds of animals
other structures were to be seen, together with the idea that an animal could be
identified merely by its bones or even by a single bone.

It is quite true that some other things were included merely on the fringe of
natural history as thus conceived. Such was the knowledge of the cell and an
appreciation of its significance, which dates only from 1839. Such was the knowl-
edge of paleontology, which, long ago kidnaped by geology, is actually an aspect
of natural history and has its beginnings in the work of this same Cuvier, who
died in 1832. Such was the very slight knowledge of physiology that was all
there was of this now mighty branch of biology. Some of the subjects which now
occupy our attention had not yet been born. There could have been no cytology
until the knowledge of the cell had been developed beyond the point of its mere
recognition. There could consequently have been no such thing as histology until
the aggregation of cells into tissues had been grasped. There was a ]:)it of embry-
ology, going as far as macroscopic examination could carry observers, but the
real development of embryology had still to come. Genetics was not then even
conceived. Biochemistry was undreamed of and the various inferences to be
drawn from the knowledge of how many and how various are the forms of organ-
isms were just beginning to germinate in the minds of naturalists.

So the naturalist as he existed, at least almost to the middle of the nineteenth
century, was primarily, if not almost exclusively, a man who had a knowledge
of as many different kinds of animals or plants as possible and who knew some-

[75]

Tt A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

thing of what could be learned about these organisms by observations made in
the field. He was a man characterized above all by the range of his interests,
which might encompass the entire field of natural history. He was the man who
is referred to now, sometimes with respect, sometimes with a sort of envy, and
sometimes with a slightly condescending air, as the "Old Time Naturalist." The
race e:^isted until well into the early years of the present century; some of its
members have died only within the last few years. Now with the passing of the
last few stragglers it is extinct or so nearly extinct that at the most it constitutes
a "relict species." The intellectual climate has changed and it is perhaps as well
for their own sakes that the "Old Time Naturalists" are gone. They would not
be comfortable in the present climate! The environmental pressures are too great!

Here is an example of the alteration of a species brought about by changes
in the environment. Natural history has changed to meet the demands of the
new environment and naturalists have either disappeared or altered their out-
look to meet the new conditions.

Continuing this method of nomenclature, these "Old Time Naturalists" have
been replaced by what, at the best, might be called the "New Time Naturalist."
He is a modification of the earlier form, a derivative of it, but modified to succeed
in this new climate. He has of necessity become a specialist in some one or more
of the many subdivisions into which the old field of natural history has been
fragmented; but he retains something of the spirit of his predecessor and some
vision of the freedom with which that predecessor roamed at will over his domain.
There are a few men still who deserve the distinction of being thus listed in the
line to which the "Old Time Naturalist" gave rise. But alas! Even they are now
relatively few and perhaps lonely. They have, of necessity, largely themselves
been superseded by the "narrow specialist," whose interest is bounded by a fence
surrounding one of these fields or fragments of the fields into which natural
history has been shattered and subdivided, fields that all too often are surrounded
by a fence "hog tight, bull strong, and horse high"^ through which they cannot
escape, even if they would. They have been conditioned to accept their fate and
seek for no other.

But there are signs that these fences may be in part crumbling and of recent
years there have been indications that still another breed is rising, a second gen-
eration in which the recessive or suppressed characteristics of the Fi generation
are now reappearing in the F2 generation. There are now an increasing number
of men in biology who recognize that restriction to these narrow fields is neither
comfortable nor desirable and who have begun the task of reintegrating them
into fields of larger dimensions. Perhaps those reintegrated fields are not yet as
large as was the old natural history, but there are indications that in time they
may become even larger and more productive. Here, as is the nature of wheels,
the wheel begins to come back full circle but farther along.

So it is perhaps a propitious time at which to consider what the contribution
of natural history to human progress has been in the past, in part as an aid to
developing an appreciation of what was done and in part as an aid to the appre-
ciation of what may still be done by one who refuses to be confined within a
narrow specialty.

i

1. A characterization derived from advertising contemporaneous with the last days
of the Old Time Naturalists and the early days of the author as a farm boy.

ferris: the contribution of natural history to human progress 77

The Legacy of the Old Natural History

AVhat of the old natural history was there that may be carried over and
legitimately included within the field of consideration of the new natural history?
Shall we limit the applicability of the term itself to the activities of the period
up to roughly 1900, when it had a certain generally accepted meaning, or shall
we extend it to include at least some of the derivatives that have developed during
the last part of the nineteenth century and the first half of the twentieth? On
the one hand, we risk limiting it too much; on the other hand, we risk extending
it beyond any acceptable limits. For one thing, the earlier natural history was
certainly not co-extensive with all of what we now call biology and even many
of the special fields of the present day are certainly not entirely devoid of what
we might call natural history. If we search for the common element, we may at
last come to the solution that what we wish to find is to be sought for not so much
in content as in an attitude of mind.

This attitude of mind has been discussed by Marston Bates in his delightful
book The Nature of Natural History. It is in brief, the attitude of mind which
displays interest primarily in the organism as a functioning whole and as a part
of the living world. With such a conception, the person who is interested only
in the permutations and combinations of the chromosomes within cells may call
himself a biologist, but he is certainly not a naturalist — a fact upon which he
would probably pride himself. But as soon as he begins to think about these
chromosomes and their permutations and combinations in conjunction with the
influences from the world around them, his thoughts begin to impinge upon
natural history, upon the fate of the organism which contains the chromosomes
as it has to accommodate itself to the facts of life. He begins to think of the
organism as a whole. The physiologist who is interested only in the processes
which go on within the membrane that surrounds a cell is certainly not a natu-
ralist and — if my observation of such individuals is at all correct — is not at all
disturbed by that fact. But when he begins to think about these cells as organized
into a complete plant or a complete animal, he must begin to think at least a
little about how this plant or this animal is going to live in company with and
in competition with other plants or animals. He begins to show some faint indi-
cations of the mental processes of a naturalist.

On the other hand the thoroughgoing naturalist of the old style suffered cer-
tain limitations. His interest may have been confined entirely to the organism
as a whole, to the complete ignorance of the processes going on within the organ-
ism and upon which its outward functioning as a whole depends. He accepted
the fact that there is such a thing as heredity but was not much concerned with
just what heredity implies concerning the processes by which a character is passed
on from one generation to another. Concepts of processes being involved in this
functioning — processes of respiration, processes of the utilization of food, proc-
esses of excretion, processes of nervous stimuli and the transmission of those
stimuli, processes by which cells arise and divide and tissues are formed, proc-
esses by which substances are transferred from one cell to another — these were
entirely beyond his ken and hence beyond his interest.

So. as knowledge of these processes began to appear and to increase and the
need for a detailed factual understanding of them became apparent, the naturalist
commenced to lose his hold upon the body of knowledge that was developing. It

78 -A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

began to go beyond his immediate horizon. He became more and more restricted
to observation of what can be seen or inferred only from the complete organism,
without any regard to what goes on within it or to the way in which what s'oes
on within it conditions its activities. He himself began to build a fence around
his own thinking as it were and finally to lose all connection with the workers in
these special fields.

Conversely, the knowledge of these special fields ultimately came in many
instances to be so detailed that it seemed almost beyond the range of any one
person to grasp more than one of them. Not only was natural history crowded
out but also the specialized fields began to elbow each other. Witness what hap-
pened to comparative anatomy, which is the term generally used if one is study-
ing the structure of vertebrates, and comparative morphology, a term that has
come to have the same meaning if one is studying certain invertebrates. The
great era of comparative anatomy began with Cuvier in the early years of the
nineteenth century and lasted to about 1900. During its early stages it could
very well be included under natural history, but it developed into a specialty by
itself and in turn came into competition with the rising studies of cytology and
histology and physiology, which last was more concerned with what goes on
within the tissues than in how they are put together. And at last, coincident
with the rise of genetics, comparative anatomy almost faded from the scene.

During the rise of the various specialties in biology great masses of detailed
information have been accumulated, making it difficult for anyone not immedi-
ately concerned with these specialties to master their content. This has been
seemingly inevitable, for the first necessity in the development of any field is
merely to accumulate facts. Eventually, however, these facts lead to the devel-
opment of theories and principles and then to a degree of simplification. The
pertinent facts are sorted out, the principles are established, and in time a stage
is reached when it is no longer necessary to have all the details at one's fingertips
in order to appreciate the bearing of a particular discipline upon other disci-
plines. When that stage has been reached, the general student does not need to
know all the details that have been worked out about the physiology of the cell,
but he does need to know the principles involved. And we are coming to the
point where those principles are being formulated in such a way that it is possible
to grasp them and their implications for workers in other fields. In other words,
we are coming to the point where the general student can begin to get an under-
standing of the principles that are involved in many fields and which have a
bearing upon the special field in which he is engaged.

Thus the principles of genetics have a very profound Ijearing upon the work
of the systematist especially as it concerns species. They even have a bearing
upon the work of the student of comparative morphology. Conversely, the work
of the systematist has a profound bearing upon the broader problems of genetics
and I feel sure that the comparative morphology of the arthropods, for example,
Avill, when well enough developed, have a profound bearing upon conclusions
derived from genetics.

So we are coming once more to a situation in which the person of broad inter-
est need not necessarily have to be a master of all the details of all these spe-
cialties. He need concern himself only with the principles — with perhaps enough
knowledge of details to understand those principles. He is to a degree freed irom

FERRIS: THE CONTRIBUTION OF NATURAL HISTORY TO HUMAN PROGRESS 79

slavery to detail. And if that be true, the naturalist can arise again and con-
tribute as a naturalist to the progress of biology and Ihi'ongh the progress of
biological understanding to the progress of man.

There is here, however, one disturbing thought. The progress of biology has
been coincident with the rise and recognition of the professional biologist. The
Old Time Naturalist was in many instances a man who did not earn his living
through his knowledge of natural history. The present-day biologist is generally
employed in a professional capacity. Now a professional position demands pro-
fessional competence and professional competence demands something more than
acquaintance merely with principles. So the professional biologist who wishes
to compete within his profession is forced to consider and become proficient in
details as well as principles. And there is many a professional position which
demands nothing more — and frequently does not encourage anything more —
than competence in details. How that difficulty is to be resolved is not immedi-
ately apparent. But we may hope that the genuinely competent man -who has it
in him to extend the bounds of knowledge wdll also have it within him to triumph
over difficulties and eventually to emerge from the forest of details into the high
places where his view is unobstructed and far-ranging.

So as one approaches the story of the contributions of natural history to
human progress it is desirable to remember something of the history which we
have been discussing. Natural history has given us some great things; in the
hands of real naturalists it can still give us great things. Let us consider how
natural history has expanded our range of thought and how it has contributed
to human progress, by this and by other means.

There are tAvo aspects of these contributions which need to be considered.
One has to do with philosophical matters, the other has to do with material or
practical considerations.

Tfieoretical Aspects

First as to philosophical matters. Out of the work of the Old Time Naturalists
came the beginnings of most of the great ideas that not only dominate biology
today but reach far beyond.

Consider the concept of evolution, the men from whom it came and the men
who first of all rose to its support and establishment. This was purely a contri-
bution from natural history; physiology had nothing at all to do with it. Experi-
mental biologj' had only an infinitesimal connection. Biochemistry had nothing
to do with it. Cytology had nothing to do with it. Comparative anatomy had only
a small part. Genetics had nothing to do with it, for genetics was not yet con-
ceived, even less born. Natural history, in its purest form, was almost all that
existed of w^hat w^e call biology at the time when the idea of evolution was accu-
mulated in the minds of Darwin and "Wallace and their predecessors. The idea
of evolution arose in the minds of men whose knowledge of any aspect of what
we now call biology except' what was included in natural history was almost nil.
Their i»redecessors. Buff on, Lamarck, and Erasmus Darwin, were purely natural-
ists. Wallace, who shares with Charles Darwin the honor of first formulating a
definite and intelligible concept of how evolution could have been brought about,
was a field collector of insects. Charles Darwan himself was the purest of pure
naturalists, whose ideas concerning evolution were first developed in the course

80 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

of his voyage about the world collecting' and observing objects and phenomena
with the eve and the interests of a naturalist. He was indeed a naturalist in the
oldest and most uncontaminated meaning of the word. His interest was in animals
and plants as complete and functioning wholes, living with other animals and
plants, themselves complete and functioning wholes. The impact of this idea of
evolution has been felt not only in biology, of which it is the great and unifying
idea — indeed the greatest idea that has been contributed to human thought —
but it extends into every field — philosophy, theology, sociology, even politics. Its
influence extends indirectly even into the newest of all fields, nuclear physics;
indirectly into this last field, since the idea of organic evolution has broadened
into a concept of inorganic evolution as well, and nuclear physics has contributed
to the idea of the transformation of one element into another as an accepted and
established process. The idea of a physical world which is not static but is for-
ever changing and evolving is made possible by the prior establishment and
acceptance of an organic world that is changing and evolving.

The evolution of life and the evolution of nonliving matter are no longer
separate and distinct things but are merely parts of a continuum. The idea of
organic evolution gave cogency to the thought that for this evolution time was
needed and the realization of the need for this time undoubtedly influenced the
thought that time must be found. From a world that was perhaps a little more
than flve thousand years old to a world that is probably two billion years old
and on which life has existed for probably five hundred million years — that is
the measure of the influence of an idea which sprang from the activities of a
naturalist! That is the measure of the foundations which natural history of the
nineteenth century laid down for us.

What difference does it make that Darwin knew no physiology, no c}i:ology,
no histology, no chemistry, no experimental biology, no biochemistry, no genetics,
no nuclear physics ? All that counted for the development of his great idea was
the fact that he had some appreciation of the richness of life upon the earth and
some appreciation of the fact that all these organisms live in a world of other
organisms with which they must compete. The remainder is the development of
inferences to be drawn from this recognition and the development of the tech-
niques necessary to investigate the facts. Most of this work would probably have
been done even if the idea of evolution had never been brought forth, but the
idea of evolution gave a guidance and a direction to the whole process that would
otherwise have been lacking. Without this central theme one can conceive only
of confusion resulting from all this uncoordinated activity.

In the ancient religions of the Mediterranean world and the Near East there
recurs time after time the concept of the "Great Mother" and we see this concept
continued today in what some students regard as a lineal succession in one of
the predominant religions of the western world. Cybele, she was once called,
this "Great Mother." If a biologist were to accept this idea as having had an
influence on the development of biological thought and were to seek her name it
might justifiably be accepted as natural history, which was the great mother of
all the branches of investigation and thought which we now place under biology.
These branches are her children and her grandchildren, and we can even see
something of the gestation, at least, of her great-grandchildren.

It is perhaps here that natural history has its chief claim to our respect. As

FERRIS: THE CONTRIBUTION OF NATURAL HISTORY TO HUMAN PROGRESS 81

the great mother she was the founder of a dynasty. "With all her wealcnesses,
with all her deficiencies, with all her naivete, with all her actual ignorance of
many things, she was still great. She is now old and feeble and condemned to
withdraw from the main stream of activity, but the memory of her former great-
ness still remains. It is in her children and grandchildren — by direct descent
and as they have been hybridized with other lines — that we must seek to continue
this resume of her influence upon human progress. That lineage is beginning to
become involved, somewhat like the lines of descent of ancient royal families.

Practical Aspects

Of her children the one which most closely resembles its parent is ecology.
In fact, there are those who would say that ecology is merely natural history
under another name. "Were that entirely true, we would have something analo-
gous to the history of the gods and goddesses of mythology, many of whom
changed their names but not their attributes. But natural history lived and
flourished before the days of fingerprints and so a positive identification of
ecology with natural history cannot very well be established. AVe may make a
concession to the desires of ecologists who wish their subject to have the dignity
of an identity all its own. Let it rest. Let them have that dignity, but let them
not forget who was their maternal parent.

Here, if amnvhere, the need for considering the organism as a whole, living
in a world of other organisms functioning as whole, still remains. In fact that
is what ecology is by definition, "the relation of an organism to its environment"
both living and physical. True, there is a branch of experimental ecology which
follows the experimental technique of bringing the subject of study into the
laboratory, dissociating it into its component parts and studying each of these
parts — temperature, moisture, pressure, light — as a thing by itself with the hope
eventually of combining these things in various degrees and then submitting the
combinations to similar study. This branch of experimental ecology is almost a
grandchild of natural history, for it is a hybrid involving elements from physics,
chemistry, and statistics. It displays something of that ''hybrid vigor" that is
often talked about, but as yet it is merely a strong and active child. Ecology in
general is still based upon the necessity for actually going out into the fields and
the woods and the waters and observing what is going on. The ecologist may at
times don his white jacket, retire to his laboratory and listen to the music of a
computing machine, but by and large, withal, he will be working up the data
that were initiallj' obtained while he wore a pair of field boots and was engaged
with the activities of plants and animals as they live in company with each
other, subjected to the wind and the rain, to heat and frost, and to the rolling
seasons. The ecologist of today may use registering thermometers and improved
rain gauges and barographs, improved methods of obtaining population counts
— and above all, improved means of transportation that prevent blisters on the
feet — but the objective and the outcome are in spirit very much the same as
they were years ago in the days of natural history. I go along with IMarston Bates
who remarks that "both labels apply to just about the same package of goods."

Even if an ecologist might object to being called a naturalist, he would surely
not object to being included in a survey of what natural history has done for
human progress. There is here, however, actually a defining line to be drawn.

82 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Natural history, as has been pointed out, made some great contributions to
philosophy. Ecology has made, and above all has the potentiality for making,
some great practical contributions. There are two aspects in which this last is
clearly apparent. One of these is connected with the conservation of renewable
resources. The other aspect concerns the application of ecology to medicine.

We are confronted at the present time with a growing realization that our
renewable resources need to be studied. Our forests are beginning to show signs
of wear from use. Our wild food animals from sardines to ducks and trout — if
we may by courtesy include the last two as ''food animals"^ — are showing signs
of depletion. Our soil is suffering from improper handling, which, at least at
times, implies improper treatment of the natural covering of grass and woodland.
Any solution of these problems depends in the first instance upon a basic knowl-
edge of the plants and animals involved, how they maintain themselves, how
they reproduce, their requirements, how they fit into an environment that can
maintain a balance between their numbers, the food supply that they themselves
must have and the food supply that they may yield.

It is only within recent years that any appreciation of the idea that these
problems are fundamentally problems in ecology has begun to develop even among
biologists. This is because they have very commonly been approached from some
other point of view, such as that of the commercial fisherman, the lumberman or
the farmer desirous only of obtaining an immediate return from his activities.
But the idea that any proper approach to such problems must rest upon a knowl-
edge of the organisms involved is beginning to grow and eventually must become
dominant if these problems are to be solved in any satisfactory way. In this lies
one of the greatest contributions to human welfare that are still to be made by
any subdivision of biology.

In the relation of ecology to medicine we have a very special situation. A
physician is of course primarily concerned with what goes on in the human body
and the relation of the doctor's activities to ecology is in many instances more
or less remote. It is in connection with diseases of parasitic origin or diseases
for which transmission is dependent upon other organisms that his activities
come into contact with ecology. Now it so happens that the physician was at
one time solely responsible for the development of our laiowledge concerning
these diseases. He was concerned with the effects of such diseases as malaria
upon the human body, and it was entirely natural, in fact, inescapable, that he
should search for the pathogen and explore the problem of how that pathogen
gets into the human body. But, since he was the first to inquire into these ques-
tions, he quite naturally took over first of all a consideration also of the organ-
isms which act as vectors for these diseases. Since it is hardly compatible with
human nature to let go a hold that has once been established, the physician con-
tinued for some time to include these vectors within the range of his special
domain, although he was scarcely qualified by his training to maintain this hold.
In fact, the problem of the relation of these vectors to the pathogen and to man
is not a medical problem at all except as medical men may be interested in pre-
ventive medicine. If I may employ an analogy, consider the instance of injuries
from automobile wrecks. The doctor has to treat these injuries and he may be-
come impressed, in the course of his duties, by the need for some procedure whicli
will reduce the incidence of wrecks. But the prol)lems involved in handlino

FERRIS: THE CONTRIBUTION OF NATURAL HISTORY TO HUMAN PROGRESS 83

traffic, designing- Iiighwa.ys to minimize accidents, formulating and administering
laws which will aid in doing so — these arc not problems for the doctor at all.

So with these diseases of parasitic origin or parasitic transmission. Until the
parasite is present in the body of man, it is beyond the range of the physician's
activities and even beyond the range of his proper interest except in so far as a
knowledge of this sort broadens the scope of his understanding. How to control
these parasites is properly no part of his concern, for it embodies problems that
are not within the range of a hospital-trained medical man. These problems are
actually those of ecology, of an understanding of the insect vectors and parasites
themselves, their ways of life and their relations to other organisms.

This idea has finally begun to penetrate even into the minds of doctors, and
there is a growing body of men whose training fits them especially to deal with
these organisms. They have no collective or corporate name at the present time,
but one may safely predict that such a name will finally appear. They are
scarcely to be called sanitarians. They are not strictly parasitologists. They are
not necessarily medical entomologists. Just what are they ? That remains to be
determined, but in time some term will inevitably appear that properly indicates
the range of their activity. They are actually naturalists. Personally, I am
inclined to the opinion that the term "environmental medicine" could in some
way be employed for their field.

But, regardless of what they may eventually be called, it is clear enough
that their activities have a large part to play in the future story of human prog-
ress. The activities in which they engage have already almost eliminated from
some parts of the world diseases which once made those areas relatively unhabit-
able by man — witness especially yellow fever — and they promise to do the same
for even greater areas. In fact, it seems reasonable to predict that the control of
parasites and their vectors will eventually lead to making habitable and useful
to mankind those great areas of the tropics which now maintain but a scanty
population and contribute but little to the commerce of the world. Whether or
not this is actually a consummation devoutly to be hoped for is another matter.

Systematics

Another child of the first generation derived from the Great Mother, natural
history, is biological systematics, which, as I have pointed out, at one time con-
stituted a very large part of natural history. It was the question "How many and
how varied are the kinds of organisms ?" with which the naturalist was concerned.
Now, however, it has become merely a section — sometimes a strongly fenced-off
section — of the activities which we have inherited. It has a rather peculiar his-
tory. Originally, in a rapidly expanding world, it amounted to but little more
than an expression of curiosity aroused in large part by the great numbers of
previously unheard-of kinds of plants and animals that were discovered and it
became to a large extent merely an attempt to give these plants and animals
names and to arrange them into some sort of system by which knowledge con-
cerning them could be handled. From this there grew what became at times
almost a cult, embodying the idea that it was the sole purpose of the systematist
or taxonomist to find and name as many as possible of these animals and plants
and to fit them into the system. In fact, it became somewhat the idea — although
perhaps never clearly expressed — that this goal extended to naming all the ani-

84 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

mais and plants of the world. This was contributed to by the circumstances that
systematics lends itself nicely to the gratification of that instinct for collecting
which is so deeply embedded in our minds. What collector of postage stamps has
never dreamed of possessing a complete collection of all the postage stamps that
have ever been issued ? Or, if the impossibility of achieving this goal is too evi-
dent, has not relished at least the possibility of obtaining a complete collection of
those stamps within the specialized field to which he restricts himself?

So this cult of finding and naming all the kinds of plants and animals of
the world and of squirreling them away in collections drifted away from any
special thought about the bearing of these activities upon biology. It drifted
away from the desire to know anything much about these subjects of its interests.
In the desire to possess collections it became concerned primarily with the col-
lections themselves and their possession and in so doing it became at least as
detached from biology as the collecting of postage stamps is detached from the
primary functions of the Post Office Department. It became a subject that could
be engaged in without previous training by children, retired army officers, police-
men, janitors, street-sweepers, preachers, medical men, and perhaps even poli-
ticians. Some of the objects of its interest became objects of commercial enter-
prise. One could purchase a collection of butterflies or beetles or shells as one
can purchase a collection of stamps and there are instances on record of insects
having been described merely in order to increase the list of collectors' desiderata.
And yet even this expression of the collector's passion was not without its
influence upon the development of natural history for, through it, men came to
know something of and to appreciate the richness and the variety of life. Inci-
dental this may in part have been, yet the indirectly beneficial result is clear.
Linnaeus, the patron saint of biological systematists, knew less than ten thousand
kinds of animals for the whole world. Today we know — or it may be more truthful
to say know of — something up toward one million and we have reason to suspect
the existence of as many as perhaps ten million kinds of animals alone, not to
mention the kinds of plants.

Reflect for a moment! A biology based upon the existence of but ten thousand
kinds of animals in the world would be on a very different philosophical basis
from a biology based upon a concept that allows for the existence of ten million
kinds. With only ten thousand kinds in the world one could almost accept the
literal truth of the story of the Ark! With only ten thousand species of animals
in the world one would not be confronted with the necessity of examining the
multiplicity of physiological processes and phenomena that we know to exist.
With only ten thousand species of animals in the world the concept of a special
creation for each might well be acceptable. In other words, the idea of evolution
is necessary because this multiplicity of forms demands it and makes it the only
idea that the reason of a scientist can accept as offering any basis for some
final understanding of the facts.

But after all, not all systematic biology has been entirely motivated or lim-
ited strictly by the mere gratification of the collector's instinct. After all, men
had to go out into the world to collect these animals and in doing so they became
at least to some degree acquainted with their ways of life. And so a knowledge
of the occurrence and the habits of animals and plants grew up along with —
possibly to some degree merely as a by-product of — this search for new species.

FERRIS: THE CONTRIBUTION OF NATURAL HISTORY TO HUMAN PROGRESS 85

Above all was this true of the earlier explorer naturalists. So a very large body
of information that went into the development of early natural history grew up
in this way. In fact, all of these things really went together, for a person finding
a strange plant or animal naturally wished to talk about it and he could not
very well do so with any definiteness unless he had some sort of name for it. It
was only later that a knowledge of the kinds of plants and animals moved to the
laboratory and became at times completely detached from the natural world.

It was out of this combination of the knowledge of plants and animals as
things living in the natural world and the describing and naming of them by
what came to be called the "closet naturalist" that there came the ideas which
led to that great philosophical concept, evolution. Darwin himself was a great
field naturalist, but he did not disdain the work that had to be performed, for
example, on barnacles in his study. He was that very desirable combination, a
field naturalist and a closet naturalist.

So the mere describing and naming of the different kinds of animals had its
place in the development of those concepts which, broadened and deepened, led
to biology as we Imow it.

But apart from these philosophical concepts biological systematics has had
a profound effect in the development of other aspects of biology. After all, it is
at least intellectually satisfying to know what the world was like in past ages
and our knowledge of what the world was like depends upon historical geology.
Historical geology in turn rests upon paleontology and paleontology rests upon
a study of the kinds of animals and plants that existed in the past and have
come down to us as fossils. Here the recognition of the various kinds is nothing
more than an extension of the knowledge of present-day species embodied in
systematic biology. Any conclusions as to what the world was like when these
fossils lived must be based upon observations of how similar kinds now live. If
fossil plants are found which are known only from tropical regions, it is a fair
assumption that these fossils must have been laid down under tropical conditions.
So, reasoning from the conclusions concerning the kinds of organisms involved
and field natural history concerned with the habits of similar organisms, we
come finally to some understanding of the climates of the past. Thus another
step is taken in broadening our outlook on the world.

BiogeograpJiy : Another matter that has at least an intellectual interest as well as
some practical concern is the problem of how animals and plants are arranged
naturally about the world. This is what is known as biogeography. It depends
entirely upon the results of systematics. The data utilized are merely those of
systematics, further systematized by embodying them in maps of the world or
portions of the world. The validity of its conclusions depends, then, upon how
well the world has been explored and how well the systematic work has been done.
The practical aspect of this may be indicated by examples from economic
entomology. Let us say that a hitherto unknown pest is found in the United
States — as has happened many times. For various reasons we wish to know
where that pest came from. Some of these reasons are merely concerned with
satisfying curiosity, others with practical considerations. In entomology that
practical consideration has to do with the question of what we call "biological
control," which is an aspect of applied ecology. We know that in its natural habi-

86 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

tat an insect has certain enemies which control its numbers and that, if we could
introduce those natural enemies into the area now infested by the insect, we
might be able to restore the balance that existed in the land of their origin. But
to search blindly for these natural enemies with no idea of where they are to be
found is a wasteful business. Sometimes that has been done, and on a few occa-
sions the search has, by great good fortune, been successful. On some other
occasions it has failed. Thus expeditions searching for natural enemies of the
"red scale" — an insect of much economic importance to the citrus growers of
California — were sent to South America, to Australia, to Africa. They secured
no parasites that were effective against the red scale. Why? The red scale, we
now are quite sure, is a native of southeastern Asia.

How do we determine purely from sytematics where an animal came from?
First of all, there should be a study of the great group to which the animal
belongs — let us say, in this case, scale insects. By this study we arrive at an idea
of the minor groupings that exist. Next by a study and a mapping of the dis-
tribution of the species of a minor group we determine what part of the world
it belongs to. Then, by a more detailed study of all the species contained in this
minor group, we arrive at an idea of where a particular species naturally belongs.
Finally we can put our finger on the map of the world and say, "This is the most
probable place in which a search for parasites would be profitable." A study of
this sort indicated that parasites of the "olive scale," an insect of importance
in California, would most probably be found in northwestern India or Persia.
A search guided by this information found parasites in India, which have been
introduced into California and promise to be of value.

In the field of that "environmental medicine" already discussed the syste-
matics of mosquitoes has demonstrated its value. Only certain species of mosqui-
toes carry malaria, while different species carry yellow fever and other diseases.
It is useless to spend money for the control of these diseases by attempting to
control "the mosquito." There are hundreds of kinds of mosquitoes, and the
recognition of the particular mosquito concerned is essential if our efforts to
control the disease by controlling mosquitoes are to be properly directed.

Paleontology : There is one other field worthy of some special consideration in
which biological systematics has a very practical contribution to make. That is
the field of paleontology, which is fundamentally the recognition of the different
kinds of animals and plants that have lived in past times and that have left
fossil remains in the rocks. Paleontology could possibly be regarded as purely
a consideration of these fossils, but that would be relatively unprofitable and
it is well that it merges with, and is united with, information from other fields
to become historical geology. Historical geology has had not only a profound
influence upon the development of the idea of evolution but also upon many
practical matters. Time was when this was about all the explorer searching for
oil had to depend upon, although it is now aided and abetted by other methods,
but the practical aspects of historical geology still exist.

These are merely pertinent examples, which could be multiplied many times,
to show that biological systematics has a proper place and at times is essential
to the development of a proper understanding of the world in which we live.

In the end the systematist, if he is to fulfill his possibilities of being helpful

FERRIS: THE CONTRIBUTION OF NATURAL HISTORY TO HUMAN PROGRESS 87

to mankind, must think of his specimens as being merely samples of great popu-
lations living out of doors under natural conditions. This systematist may sit
at his microscope or his desk working only with the variously preserved remains
of his specimens, but if he has any vision of his place in tlie great endeavor to
improve the world, that vision must reach far outside the walls of his study or
laboratory — and does.

So biological systematics still has a place as a part of the great endeavor that
has as its goal human progress — progress intellectually and progress in more
immediatel}" applicable things. It still maintains its former place of importance
in natural history, for it furnishes the material with which a naturalist must
work. The ecologist, the student of geographical distribution, the student of
biological control, and even the student of genetics — especially with reference
to the origin of species — must make use of its findings. Systematics may change
— and it is to be hoped that it does change — ^from concentrating its attention so
much upon ''new species" to concentrating primarily upon the problems of clas-
sification and upon its liaison with other branches of biology and the contribu-
tions that it may make to such general problems as those having to do with the
mechanism of evolution, but its continuing place is secure.

Genetics: One of the most interesting developments of systematic biology is its
liaison with genetics. During the years in which Mendelian genetics was strug-
gling to establish its body of ascertained fact there was but little opportunity and
little time to consider the relation of the implications of genetics to other fields
of biology. But, with this basic body of fact quite well determined and with the
underlying principles established, the opportunity has finally come and to some
extent has been grasped to explore connections with other fields. One of the
most fruitful of those fields is biological systematics. In the problem of how the
members of a single interbreeding population become differentiated into two or
more distinct and finally non-interbreeding populations genetics and systematics
reach a common ground, for both have here their common interest in the matter
of evolution. Thus at last there has arisen by hybridization between the offspring
of natural history and that relatively recent, apparently quite unrelated disci-
pline, genetics, a new way of approach to these common problems. This too is
at present a field without an accepted name, although there is some reason to
think that the name now used by some of those who are interested in such matters
— biosystematics — may eventually receive a wide acceptance.

We could explore these matters further and call attention to other ways in
which natural history and her lineal descendants, "bone of her bone and flesh of
her flesh" — if we may revert to an ancient phrase — -have contributed their share
to human knowledge, to the advance of biology, and to the practical affairs of
life. AVe have, for example, not mentioned the bearing of a knowledge of the
fungi which is involved in the development of what the medical man calls the
"antibiotics." AVe have not mentioned agriculture, which involves certain aspects
of ecology and which will do so more and more as the needs of the world for an in-
creased food supply becomes more manifest. We have not mentioned — but let it rest !

In the words attributed to the mother of the Gracchi in referring to her dis-
tinguished sons, "these are my jewels," natural history has been the Great Mother
of them all.

CLASSIFICATION AND TAXONOMY OF THE
BACTERIA AND BLUEGREEN ALGAE

By C. B. VAN NIEL

Hopkins Marine Station of Stanford University
Pacific Grove, California

Introduction

The early ISSO's as a starting point for the examination of the development of taxo-
nomic theory are appropriate not only because of the centenary aspect of the Edinburgh
meeting of the British Association but also because they have an intrinsic importance
as the culminating point of pre-Darwinian taxonomy, when the natural system had
triumphed completely over the Linnean. — Gilmour, 1951, p. 400.

It might be suggested that a few simple changes in the quotation above, such
as the substitution of "California Academy of Sciences" for "Edinburgh meet-
ing," would render it applicable to the present chapter. This, however, is far from
true. The fact is that a century ago there did not exist even a rudimentary
taxonomic theory for the bacteria. And it is highly questionable whether at
present we have advanced much beyond the equivalent of a Linnean system.
Nevertheless, advances there have been, though hardly in the sense meant by
Professor Gilmour. Rather have they been concerned with a clearer appreciation
of the problems inherent in the classification and taxonomy of the bacteria and
bluegreen algae.

Tlie following essay is intended as a sketch of the main trends of these devel-
opments. It does not contain a detailed description and discussion of the various
systems of classification of these organisms that have been proposed in the course
of the past century. Information of this sort can be found in various text- and
handbooks; Migula's System der Bakterien (1897-1900) and his contribution to
Lafar's Handhuch (1904—1907), Buchanan's General Systematic Bacteriology
(1925), and Bergey's Manual of Determinative Bacteriology (6th ed., 1948)
trace them satisfactorily for the bacteria, and Geitler's extensive treatise on the
bluegreen algae (1932) comes close to performing this task for the latter group.

The Natural Affinities of Bacteria and Bluegreen Algae

Quoi qu'il en soit, les Schizomycetes ne sont point une classe. Une classe de quoi?
ai-je demande au Comite International de Nomenclature a New York en 1939; et aucun
des nombreux delegues representant le monde bacteriologiste n'a pu repondre. C'est au
moins un embranchement, mais un embranchement autonome, intermediaire entre les
regnes animal et vegetal et nettement separ^ d'eux. Pourquoi ne pas avoir le courage
de dire: le Regne Bact^rien? — Pr6vot, 1940, p. 10.

Although first seen and described nearly three hundred years ago by Antonie
van Leeuwenhoek, bacteria could not be adequately studied, for lack of an appro-

[89]

90 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

priate methodology, until the second half of the nineteenth century. Nevertheless,
some of the general features of the organisms, such as the occurrence of motile
forms, and multiplication by transverse fission, had been established, and the
discovery of the bacteria had raised the question whether they ought to be
regarded as plants or animals.

Prior to 1854 their animal nature had been taken for granted, locomotion
probably being the chief criterion on which this belief was based. But in that
year Colin (1854) argued in favor of a close relationship with plants, especially
with the bluegreen algae. Following Nageli's introduction (1857) of the name
"Schizomycetes" (''fission fungi") it became customary to use this term, with
the ending appropriately modified to indicate the status as a family, order, or
class, for the collective designation of the bacteria. Along with this practice the
notion of the plantlike nature of the organisms gradually won ground.

It cannot be denied that there are good reasons for subscribing to this view.
Especially the existence of an autotrophic mode of life among the bacteria may
be considered a strong point in its favor. The chemo-autotrophic sulfur bacteria
of the Beggiatoa-Thiothrix-Thioploca group in particular form a striking ex-
ample because also from a morphological-anatomical point of view they show
their plantlike nature; the structural similarity with the bluegreen algae of the
family Oscillatoriaceae is great indeed (Pringsheim, 1949). The green and purple
sulfur bacteria, and the brown and red nonsulfur bacteria resemble the plants
even more closely in physiological respect by virtue of their photosynthetic
ability. Recently it has been proposed that the chemo-autotrophic mode of life
can be envisaged as a precursor of the photosynthetic one, and that such processes
as characterize the photosynthetic bacteria would represent a logical link between
chemo-autotrophy and green plant photosynthesis (van Niel, 1949a).

In spite of these rather compelling considerations, doubts as to the exclusively
plantlike nature of the bacteria have also been expressed, and this with increasing
frequency. It should be emphasized that Niigeli had not in the least committed
himself concerning the general relationships of his Schizomycetes ; this is evident
from the statement (Nageli, 1857, p. 760) :

Ueber die Bedeutung der Gruppe Schizomycetes, ob es Pflanzen, Thiere, oder krank-
hafte thierische oder vegetabilisclie Elementartheile seien, dariiber giebt die anatomische
Struktur keinen Aufscliluss, dass es Pflanzen und keine Thiere sind, dafiir liegen wenig
Grijnde vor.

The vast increase in our knowledge of "the bacteria" gained during the past cen-
tury has not made Nageli 's statement obsolete. This must in part be ascribed to
the difficulty of finding close affinities of certain bacteria with specific taxonomic
groups among the plants. While F. W. Andrewes, for example, states (1930,
p. 298) :

... It was not till the middle of the nineteenth century that first Naegeli and then
Cohn proclaimed the vegetable nature of the bacteria. So gradual is the transition from
the mould-fungi, through the streptothrix group and the acid-fast bacteria, to ordinary
bacteria, that there are few who do not agree with Naegeli.

it is equally true that relationships with bluegreen algae and with other groups
of organisms can also be defended on reasonable grounds. The quotation from
Prevot at the beginning of this section clearly reveals this difficulty. And from a

VAN NIEL SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 91

phylogenetic standpoint it is hardly surprising that a major problem would
exist; it is, in fact, inherent in the concept of evolution itself.

Acceptance of the doctrine of organic evolution implies that the clearly
recognizable forms of plant and animal life must have had a beginning in some
far more primitive ancestry. It does not appear unreasonable to envisage the
evolution of an elementary "molecrobe" to typical plants and animals, respec-
tively, as having passed through intermediate stages of increased complexity
which, in a number of respects, would have the characteristics of "bacteria."
Such intermediate stages are themselves neither plants nor animals; they occupy
a position in the realm of living organisms that is antecedent to the emergence
of the later developmental stages, and display characteristics of both major
kingdoms. It is not the contention of this argument that the present-day bacteria
are, in effect, such intermediate stages; it is easily conceivable that they might
represent organisms that have evolved from the same precursors from which also
the typical plants and animals, by different routes, originated.

As early as 1866 this situation was clearly recognized by Haeckel, who wrote
(1:202-203) :

Wir finden in den bekannten Thatsachen durchaus keine Nothigung fiir die An-
nahme, dass alle Organismen-Stamme entweder Thiere oder Pflanzen sein miissen. Viel-
mehr miissen wir die bisher giiltige exclusive Zweitheilung in Thier- und Pflanzenreich
in dieser Beziehung fiir niclit begriindet eracliten. Es ist schon von verschiedenen
Seiten darauf aufmerksam gemacht worden, dass es sowohl fiir die Zoologie als fiir die
Botanik ein grosser Gewinn sein wiirde, wenn man die vielen zweifelhaften Lebewesen,
die weder echte Thiere noch eclite Pflanzen sind, in einem besonderen Mittelreiche oder
Urwesenreiche vereinigen wiirde; doch hat unseres Wissens noch Niemand den Versuch
gemacht, ein solches neues Reich der Urwesen nach Inhalt und Umfang fest zu bestim-
men, und seine Begrenzung wissenschaftlich zu begriinden und zu rechtfertigen. Wir
wagen hier diesen Versuch auf Grund der obigen Deductionen und schlagen vor, alle
diejenigen selbststandigen Organismen-Stamme, welche weder dem Thier- noch dem
Pflanzenreiche mit voller Sicherheit und ohne Widerspruch zugeeignet werden konnen,
unter dem Collectivnamen der Protisten, Erstlinge oder Urwesen, zusammenzufassen.

In this new kingdom the bacteria, along with such dubious organisms as
Protogenes and Protamoeba, were allocated to the first phylum, Moneres, com-
prising, in Haeckel's words, "the completely structureless and homogeneous or-
ganisms which consist solely of a bit of plasma (a mucoid protein compound),
obtain their nutrients simply by endosmosis, and reproduce by schizogony or
.sporogony" (1866, 2:20).

Unquestionably there is much that can be said in favor of Haeckel's third
kingdom. Nevertheless, its acceptance raises a new problem to which P. W.
Andrewes (1930), following Kent (1880-1882) and Biitschli (1880-1889), has
called attention^n the statement (p. 298) :

To revive Haeckel's third kingdom of "Protista" for organisms so low down in the
scale that they cannot definitely be assigned to either of the other kingdoms, may be a
useful expedient, but it is a doubtful gain, for it necessitates two arbitrary lines of
demarcation in place of one.

The seriousness of this problem becomes at once apparent when one considers
the extreme paucity of characteristics which one is compelled to associate with
the early forms of life, the pre-plant and pre-animal organisms for which the
kingdom Protista was proposed. Morphological and developmental features must

92 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

here be so primitive that they can hardly be expected to serve as a useful
guide in determining phylogenetic trends and relationships. Haeckel, realizing
this, had had recourse to physiological properties as well, a practice which led
him to incorporate the bluegreen algae, as photosynthetic organisms, in the plant
kingdom. As a result, the views of Cohn in respect to the close affinity between
the bacteria and the bluegreen algae did not come to a clear expression in
Haeckel's system.

Since it was Cohn who, in 1872, took the most significant steps toward the
development of a more detailed classification of the bacteria, it is understandable
that in these attempts he adhered to his notion that the bacteria are bona fide
members of the plant kingdom. And Cohn's influence has been so great that for
a long time Haeckel's proposal was not seriously considered, at least by the bac-
teriologists.

But Copeland, in an important contribution, reexamined the arguments in
favor of Haeckel's ideas and conceded their soundness (1938, p. 384) :

It is an ancient and familiar hypothesis, too widely accepted as a law of nature, that
every living creature is and must be either a plant or an animal. Judged by knowledge
and theory which were available to Linnaeus, this hypothesis is sound; judged by mod-
ern knowledge and theory, it seems untenable.

As he further pointed out (ibid.) :

Various authors more recent than Haeckel have shown a disposition to recognize
more kingdoms than two, but none of them, apparently, has formulated a system includ-
ing all organisms. Pending such an accomplishment, the old system of two kingdoms
has persisted for want of a workable substitute.

With a view to improving this situation Copeland developed a substitute in which
four kingdoms were recognized: Monera, Protista, Plantae, and Animalia. The
first phylum of Haeckel's Protista was here raised to the rank of an independent
kingdom, the criterion for inclusion in this taxon being "organisms without
nuclei, the cells solitary or physiological (ly) independent. Groups included,
bacteria and bluegreen algae" (p. 416). In this manner a seemingly unambigu-
ous separation of the bacteria and bluegreen algae from all other organisms w^as
achieved, while at the same time justice was done to Cohn's concept regarding
the close relationship between the two major groups of the Monera.

Several years later Copeland returned to the problem of basic classification.
At this time he stated the phylogenetic significance of the first kingdom more
clearly, as follows (1947, p. 342) :

The most profound of all distinctions among organisms is that which separates those
without nuclei from those which possess them. The foi'mer are the bacteria and blue-
green algae. . . . Whether or not life originated more than once, it is certain that life
possessing nuclei came into existence once only, by evolution from "tionnucleate life.
This conclusion is as certain as any which can be based on induction: it is established
by the uniformity of the nucleus, in its structure and in its behavior, in mitosis, in
sexual reproduction, and as the vehicle of Mendelian heredity, wherever it occurs.

He also recognized that his designation of the kingdom as Monera was invalid
because Enderlein (1925) had earlier used the name Mychota for just such a
taxon. Meanwhile, the proposition of uniting the bacteria and bluegreen algae
in a separate kingdom had found favor with Stanier and van Niel (1941), who
had, furthermore, seen fit to expand the characterization of this unit by the

VAN NIEL SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 93

incorporation of two additional, and equally negative, criteria, viz., the absence
of plastids in the cells, and the absence of sexual reproduction.

However attractive Copeland's system may have appeared a decade ago,
recent developments have raised difficulties great enough to threaten the very
basis of the characterization of the kingdom. The most important of these deal
with the problem of the "bacterial nucleus."

Even in 1938 there were some indications that bacteria contain discrete struc-
tures that might be considered, on the basis of their behavior and chemical nature,
as nuclei (Badian, 1933; Stille, 1937; Piekarski, 1937). Studies of this sort have
been continued, with improved methods and instruments, especially by Delaporte
(1939), Eobinow (1944, 1945), Knaysi (1947, 1951), Boivin (1948), Welsch and
Nihoul (1948), Tulasne and collaborators (1947, 1949), and DeLamater (1952);
the results support the previous allegations. Even though a convincing demon-
stration of nuclei has not yet been accomplished for more than a few bacterial
and myxophycean types, it may be confidently expected that future work will
fill the existing gap. It is thus becoming increasingly clear that these organisms
cannot be incorporated into Copeland's kingdom of "microorganisms without
nuclei."

Similar remarks, while not yet as definitive, may well apply to the two addi-
tional criteria mentioned above. The finding in cells of the photosynthetic bac-
terium, RJio do spirillum ruhrum, of uniform spherical particles in which all the
pigment is contained (Schachman, Pardee, and Stanier, 1952) indicates that plas-
tidlike elements are not lacking in the bacteria; according to Calvin and Lynch
(1952) a, very similar situation is apparently encountered in the bluegreen
alga, Synechococcus.

Last, there is the matter of sexual reproduction in these organisms. While
there are some published reports alleging the occurrence of fusion of individual
cells in bacterial cultures (Potthoff, 1922, 1924), these had not been taken too
seriously, and it is fair to state that the actual conjugation of two cells with the
formation of a zygote has yet to be observed by continuous microscopic examina-
tion. But the startlingly novel report by Lederberg and Tatum (1946; see also
Tatum and Lederberg, 1947, Lederberg, 1947) of the occurrence of "recombina-
tion effects" in mixed cultures of bacterial mutants has changed the picture. The
observed phenomena cannot be ascribed to "back mutations"; they are, however,
readily interpretable on the basis of a postulated conjugation, followed by recom-
bination of genetic factors during the mitotic division of the nucleus of the con-
jugant. It is true that the recent studies of Hayes (1952) have shown that similar
recombinations occur in mixed cultures of mutants in which one of the partners
has been rendered nonviable. This suggests that an unequivocal interpretation
of the recombination effect as the result of a primary conjugation is not possible.
On the other hand, there exists at present a healthy skepticism with regard to
the earlier belief that sexual phenomena do not occur among the bacteria.

Thus it is clear that the criteria for a kingdom of organisms without nuclei
do not apply to the bacteria and bluegreen algae. This does not mean, however,
that the notion of establishing a separate kingdom for these organisms should be
abandoned. As mentioned before, there are good reasons for subscribing to the
idea that we must reckon with the existence of organisms that are neither plants
nor animals and represent the descendants of precursors of both these groups.

94 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

The difficulty will be to devise adequate criteria for such a taxon; this remains a
task for the future.

The Species Concept in Bacteriology

These two criteria — practical expedience in the interpretation of biological phe-
nomena, and the application of an effective system of nomenclature — are the elements
from which the systematist must fashion his concept of species. — Camp and Gilly, 1943,
p. 381.

The peculiar difficulties encountered in attempts to give formal expression
to the general relationships of the bacteria and bluegreen algae to other living
organisms can evidently be referred to the paucity of salient characteristics
among the former. This same feature is responsible for the fact that also at the
other extreme end of the classification problem, concerned with the species con-
cept, no clear-cut solution within the framework of accepted taxonomic procedure
has been possible.

Until 1872, advances in this field had been greatly handicapped by the pre-
vailing notion, purportedly based on unambiguous experimental results, that
bacteria exhibit an enormous range of variability. It stands to reason that one
can hardly expect to "classify" organisms that behave in the manner claimed
for them by the early protagonists of the doctrine of pleomorphism, according
to whom practically any bacterium could assume the shape of any other, depend-
ing largely on the conditions under which it had developed.

There had been some responsible claims and observations to the contrary.
Going back to the pioneering studies of Louis Pasteur, one can find considerable
evidence in favor of the view that the transformations claimed by the pleo-
morphists were, to say the least, not always observed. The experienced eye of
the great French chemist-turned-microbiologist, together with his uncanny ability
to devise experimental methods apt to give clearly interpretable results, soon
convinced him, as they should have convinced others, that there is often a close
and consistent correlation between the chemical changes brought about in a par-
ticular environment by the organisms growing therein and the microscopic
aspects of the cultures. Pasteur had unhesitatingly taken this to mean that there
are different and recognizable types among these microorganisms and had pro-
ceeded to describe and name them. But some later workers insisted on the occur-
rence of drastic transformations in the appearance of the organisms themselves
with changes in environmental conditions. It was, however, not always appre-
ciated that their observations might equally well be interpreted as resulting from
the use of impure cultures, by the mechanism of preferential development of
different organisms elicited by modifications of the external milieu. As long as
this fundamental ambiguity had not been resolved, the picture remained too
confused to permit serious attempts at classification.

It must have been with much relief that bacteriologists who had learned from
Lister and Koch how pure cultures could be procured and who had started experi-
menting with such material became increasingly convinced that the concept of
pleomorphism was untenable. Their results clearly indicated that, provided
pure cultures, sterile media, and aseptic techniques were employed, transforma-
tions of the sort claimed by the pleomorphists simply did not occur. With the
gradual development of rigorous techniques and criteria for work with pure cul-

VAN NIEL: SYSTEMATICS OF THE BACTERIA AND RLUEGREEN ALGAE 95

tures, experimental evidence tended more and more to favor the view that even
bacteria display a remarkable constancy in both morphological and physiological
respects. This further implied the existence of numerous intrinsically different
types of bacteria.

At this stage the needs for methods of differentiation and recognition became
apparent, and it was Cohn who early made some notable contributions towards
filling this need. As one of the leaders in the fight against pleomorphistic dogma,
Cohn (1872, p. 133) had raised the question:

. . . ob es denn bei den Bacterieu iiberhaupt Arten in dem namlichen Sinne giebt
wie bei den hoheren Organismen. Selbst wer von der Metamorphosenlehre jener Myko-
logen nichts wissen will, die Alles aus Allem entstehen und zu Alles sich entwickeln
lassen, wird docli beim Anblick eines Bacterienhaufens oft verzweifeln, unter diesen
zahlreichen Korperchen von alien moglichen Formen eine Sondeiung natiirlicher Arten
vorzunehmeu.

Cohn's conclusion was in the affirmative, as follows from the statement {ibid.) :

Gleichwohl bin ich zu der Ueberzeugung gekommen, dass die Bacterien sich in eben so gute
und distincte Arten gliedern, wie andere niedere Pflanzen und Thiere, und dass nur
ihre ausserordentliche Kleinheit, das meist gesellige Zusammenwohnen verschiedener
Species so wie die Variabilitat der Arten die Unterscheidung in vielen Fallen fiir unsere
heutigen Mittel unmoglich macht.

In the same paper a beginning was made with the systematic differentiation and
naming of bacterial "species." Differentiation was based on morphological char-
acteristics exclusively. This does not mean, however, that Cohn was not aware
of the existence of physiological dift'erences as well. He clearly recognized that
two morphologically indistinguishable organisms might yet be found to exhibit
clear-cut and constant physiological differences. But he found it difficult to deter-
mine how far such differences should be accepted as grounds for species differen-
tiation. The pertinent passage in Cohn's paper is, it appears to me, so significant
that it is worth quoting in full; a free translation follows. After pointing out
that perhaps physiological differences may later be correlated with morphological
ones, he stated {ibid., pp. 135-136) :

But, on the other hand, I suspect that in the class of bacteria similar conditions ob-
tain as found in higher animals, and particularly among cultivated plants. Of two almond
trees which cannot be distinguished by their growth, their leaves, blossoms, and fruits,
not even by the external and microscopic aspects of their seeds, one produces only bitter
seeds that contain amygdalin and emulsin and produce toxic hydrocyanic acid, whereas
the other always yields sweet almonds. We assume that these two trees belong to the
same species and originated from a common ancestor from which the two, physiologi-
cally so different, came about through variation. . . . Perhaps there exist also among
the bacteria which are morphologically indistinguishable, yet exhibit differences in
chemical and physiological activity, similar varieties or races which, initially derived
from a common germ, always produce the corresponding products through continued,
natural or artificial, cultivation under identical conditions and on the same medium.
With various yeast types Rees has demonstrated the formation of special races through
artificial cultivation. Just as summer rye is unsuitable for winter seed, though initially
both races have the same origin and can be interconverted by prolonged cultivation,
so Is a top yeast unsuitable for the production of a Bavarian type beer, and nearly every
kind of wine or beer is made with its own special yeast. Nonetheless, it is most prob-
able that many alcohol-producing yeasts belong to only one species, comprising nu-
merous "cultured races." I suspect that also among the bacteria, which act as ferments
in totally different chemical and pathological processes, there occur, besides a small

96 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

number of independent species, a far greater number of natural and "cultivated" races,
the latter tenaciously retaining their individual physiological particularities because
they multiply exclusively by asexual means.

So keen an appreciation of the value of physiological and biochemical char-
acteristics for systematic purposes inevitably led Cohn to refrain from using
them. Nor did this practice cause, at the time, serious inconveniences. In 1872
knowledge of the bacteria was still so rudimentary that the twenty-one species
which Cohn proposed satisfactorily consolidated the existing information.

But not for long did this state of sophomoric bliss persist. With the rapidly
growing interest in Pasteur's "infiniment petits" as biological agents of economic
and particularly sanitary importance, it was only a matter of years before the
accumulated information led to the realization that an enormously larger num-
ber of "different" bacteria existed, and it thus became necessary to devise more
adequate methods for systematizing this Imowledge. The approach generally
adopted was the creation of a new "species" for every organism that in some
respects differed from the previously proposed ones, generally without the least
attempt at formulating what was to be understood by a "species" of bacteria. Not
until 1912 was this matter clearly discussed by Benecke, and his answer to the
question "What is a bacterial species?" was far from reassuring to those who
might have felt that it should be possible to establish definite criteria for such
entities. With considerable candor Benecke (1912, p. 212) stated: "Die Antwort
lautet: Das, was der Forscher, welcher die Art aufstellt, nach seinem 'wissen-
schaftlichen Takt' darunter zusammenfasst." This statement bears a striking
resemblance to Dobzhansky's remembrance of a definition by "an affable sys-
tematist": "A species is what a competent systematist considers to be a species."
Dobzhansky, however, continued (1941, p. 372) :

The cause of this truly amazing situation — a failure to define species which is sup-
posedly one of the basic biological units — is not too difficult to fathom. All of the at-
tempts, mentioned above have striven to accomplish a patently impossible task, namely
to produce a definition that would make it possible to decide in any given case whether
two given complexes of forms are already separate species or are still only races of a
single species. Such a task might be practicable if species were separate acts of crea-
tion or arose through single systematic mutations. If species evolve rather than sud-
denly appear, there will necessarily be a residue of situations intermediate between
species and races. This need not, however, deter biologists from attempting to elucidate
the nature of species, provided it is clearly realized that no rigid standard of species
distinction can be secured.

Even at the time Dobzhansky wrote this passage new concepts had been devel-
oped which render the systematic treatment of special groups of higher plants
and animals much less arbitrary than the quotations above would seem to imply.
Elsewhere in this volume a discussion of such developments may be found; suffice
it here to refer to the important contributions by Babcock and Stebbins (1938),
Dobzhansky (1941), Petrunkevitch (1952), and Camp (1951). Unfortunately,
in the realm of bacteria and bluegreen algae no comparable advances have been
made. In large part this is connected with the lack of conclusive evidence for
the occurrence of sexual reproduction in these organisms, and Dobzhansky has
concisely treated this aspect in the last chapter of his book (1941, p. 379), con-
cluding that "the species as a category which is more fixed and therefore less
arbitrary than the rest is lacking in asexual and obligatorily self-fertilizing

VAN NIEL: SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 97

organisms. All the criteria of species distinction utterly break down in such
forms." A similar verdict was rendered earlier by Babcock and Stebbins (1938,
p. 64) : "The species, in the case of a sexual group, is an actuality as well as a
human concept; in an agamic complex it ceases to be an actuality." Even if
future investigations were to reveal a more or less common and frequent sex-
uality in bacteria and bluegreen algae, a phenomenon which at present is sus-
pected to characterize some actinomycetes (Lieske, 1921; Stanier, 1942; Bisset
et al., 1951), and perhaps some few strains among the eubacterial groups (Leder-
berg et al., 1951), the situation hardly warrants the hope that the modern tax-
onomic concepts of the botanists and zoologists will soon be successfully applied
to these microorganisms so as to render the bacterial and myxophycean species
"actualities" rather than merely "human concepts."

The arbitrariness of such "species" is now generally conceded. Also, it is
well-nigh impossible to escape the conclusion that "scientific tact" in delineating
these taxa must carry different connotations for different investigators. This is
quite understandable if we realize that it is often imperative, even for no other
than strictly practical purposes, to distinguish between individual strains (pure
cultures), differing from one another with respect to only one type of property,
such as pathogenicity, serological reactions, growth factor requirements, or utili-
zation of special carbohydrates. As has been pointed out in more detail elsewhere
(van Niel, 1946) the relative weight given to various possible differential charac-
teristics thus depends to a large extent on the nature of the investigation in which
the organisms in question play a role.

In this respect there has been a shift in emphasis in the direction of physio-
logical and biochemical studies. Consequently there has also developed a tendency
to use physiological and biochemical criteria for the delineation of species among
the bacteria ; studies on the physiology of the bluegreen algae have not progressed
far enough to include them in the present argument. But this departure from
Cohn's approach has rarely been justified, except perhaps on the basis of the
consideration that the paucity of morphological characteristics makes it inevitable
to resort to the use of differential properties other than morphological ones, and
that physiological differences can be regarded as the detectable expressions of
differences in submicroscopic morphology (Winslow, 1914; Kluyver and van Niel,
1936). The implications of this procedure have, however, become very clear and
very disturbing during the past decade as a result of the important investiga-
tions with naturally occurring or artificially induced "mutants" of bacterial
cultures. Apart from demonstrating that the properties of a pure culture are not
firmly and irrevocably fixed, many of these studies have also indicated that
especially the biochemical characteristics of the "mutant strains" show the same
sort of relationship to those of the "wild type" as those that have been recognized
as the result of single-gene differences in organisms in which the occurrence of
sexualitj^ has permitted a genetic analysis. This very fact has sharply raised the
question as to how far strains exhibiting such differences should be regarded as
distinct species. AYhat Cohn, without benefit of genetic knowledge, had intuitively
grasped and clearly expressed, has now once more become a point that has to be
seriously analyzed; and it is not an easy problem.

Few taxonomists will challenge the opinion that a series of mutants, produced
by the action of mutagenic agents from a pure culture of bacteria, should still

98 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

be regarded as distinct clones of the same species. But the problem is an alto-
gether different one when the question is transferred to a number of isolates from
natural sources showing similar differences. Here the practice has been to indi-
cate such differences by the use of different specific designations. Hence the
bacteriological literature is replete with descriptions of "species" that differ no
more from one another (as far as the actual characterization has proceeded) than
by properties that might well be the result of "single gene" differences. And it
should be remarked that it is not only biochemical characters, but also morpho-
logical ones that may be aft'ected in a similar manner. The now widely recognized
"smooth-rough" variation, determining the appearance of bacterial colonies, may
well be a case in point.

In consequence of this situation some students of microbial genetics have
expressed the view that the separation of species among the bacteria cannot be
taken seriously. And, admittedly, the evidence for the occurrence of variation,
even in pure cultures, is so overwhelming that its implications have to be con-
sidered. Naively, one might formulate the problem in some such form as : How
many differences, equivalent to single-gene differences, shall one accept as justi-
fication for the establishment of a species 1 It will be clear that even this formu-
lation is hardly conducive to a solution of the problem. The geneticist will counter
that, by the use of an appropriate methodology, it is easy to produce from a pure
culture offspring that differ from it by one-, two-, three-, four-, etc., gene char-
acters. Where, then, shall one draAV the line?

The developments sketched in the above paragraphs seem to lead to the con-
clusion that the problem of speciation in bacteria — and, by a similar reasoning,
this would apply equally to the bluegreen algae — has not been solved, and that
the recent work on variability and induced mutations has led us back to the stage
before Cohn's contributions, when an almost unlimited variability was accepted.
Obviously, this new emphasis on variation is not the result of "inadequate tech-
niques"; it is well established, and it is also in much closer agreement with the
Darwinian approach to biology. In a sense, one would call Cohn's ideas on clas-
sification of bacteria the outcome of the Linnean philosophy; this now has to be
abandoned.

It is an interesting problem to consider how far the "evolutionary" approach
can ever render service in reaching a more satisfactory basis for establishing some
rationale in clarifying the meaning of a bacterial species. Is it really true tliat
we have now to admit that Cohn's predecessors and antagonists have "won," and
that an unlimited variability or mutability has to be reckoned with, thus invali-
dating any and all attempts to arrive at an acceptable concept of a bacterial
species! This I do not believe; it will be necessary to recognize, not merely that
Cohn's ideas on the constancy of characters was based on inadequate informa-
tion, but also that his insistence on "constancy" had an equally sound basis in fact.
As happens so often in scientific and other controversies, the ultimate answer is
not to be found by application of the "either-or" approach, but by synthesis. It
is in this respect that the recent contributions of the botanists and zoologists
have done so much in bringing about a considerable clarification in problems of
"biosystematy," as Camp (1951) calls this branch of science, and the question
arises how far similar approaches are possible as a means of reaching the same
level with respect to the classification of bacteria and bluegreen algae.

VAN NIEL: SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 99

The quotations from Babcock and Dobzliansky show that we cannot expect
that the same methods now so successfully used elsewhere will soon solve the
problem. But it is important to point out that much can be done, and that a
great deal of the present confusion in our thinking is the result of an utterly
inadequate appreciation of the truly "biological" possibilities that the bacteria
and bluegreen algae still offer. ]\Iost of the present difficulties have resulted from
studies with isolated, pure cultures, often grown under extremely artificial con-
ditions, having little if anything in common with those that have permitted the
persistence of various types of these microbes in nature. No one has realized
this fallacy better than Winogradsky who, about thirty years ago, started to inject
the notion that pure cultures may be necessary for an adequate study of certain
physiological problems, but that an understanding of the role of these organisms
in nature cannot be gained exclusively by this methodology (see Winogradsky,
1949). It is from investigations on their behavior in competition with others that
we may expect advances which will ultimately be of the greatest significance
for gaining a better perspective also concerning the systematics of the organisms.
It is quite possible that many of the artificially produced mutants of bacteria
can be maintained only under the abnormal conditions provided by the use of
pure cultures and culture media that bear no resemblance whatever to the envir-
onments in which the organisms are naturally found. For the development of
sound principles of bacterial classification it is of the utmost importance that
this criticism be heeded; it is a serious one, and suggests at the same time an
approach that is far better suited to the problem.

Just as the modern taxonomists of the higher plants and animals have come
to insist on the need for far more than the detailed examination of a few museum
specimens and have stressed the importance of field studies on naturally occur-
ring populations, amplified by cytological and genetic investigations, bacteriolo-
gists must realize that bacterial systematics will not be greatly advanced so long
as it remains based largely on routine examination, by standard methods, of pure
cultures. In spite of the fact that those pure cultures are "living," they are in
some ways not much better than museum specimens; and their continued propa-
gation on the customary nutrient media all too often is apt to induce changes
in the organisms which make their recognition as offspring of the initial isolate
difficult, if not downright impossible. Numerous are the instances in which a
special feature that provided the first impetus to a detailed study of a bacterial
culture, be it a characteristic pigmentation, pathogenicity, or biochemical prop-
erty, such as the ability to live autotrophically as a hydrogen bacterium, or to
carry out a vigorous denitrification, was lost on continued cultivation, and the
evidence is strong indeed that the use of the routine meat extract-peptone-agar
media, on which,' to be sure, good growth of the pure culture could be secured,
must be held responsible for the changes in characteristics.

It should be self-evident that these remarks are not intended to advocate that
pure cultures are useless for taxonomic purposes. AYere this implied, the devel-
opments would soon lead us back to the pre-Cohn era of experimentation, with
results so equivocal that their interpretation would become impossible. No; they
are meant to stress the necessity of learning more about the factors that operate
in maintaining the various types of bacteria and bluegreen algae in nature. In
the elective or enrichment cultures we possess a simple and powerful methodology

100 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

for achieving this very end. Such cultures permit us to determine which among
the vast diversity of germs present in a rich inoculum can successfully compete
with the others under the specific environmental conditions, determined and im-
posed by the investigator, so that gradually they become the predominant micro-
organisms in the culture. This proced^ire, chiefly initiated by Beijerinck and
Winogradsky (see their Collected Works, published in 1921 and 1949, respec-
tively) is pre-eminently suited to determine by direct experiment what particular
features of the environment are responsible for the abundant or exclusive devel-
opment of special types and, by inference, to clarify the "natural" conditions for
their existence. Furthermore, the results provide the information necessary for
studies on the behavior of pure cultures under such conditions. And last but not
least, they can be used to isolate at will from natural sources representatives of
those types whose ecological relationships have been sufficiently established. This,
in turn, makes it possible to conduct comparative studies with several strains
isolated from different localities in order to elucidate the normal range of varia-
tion displayed by the "wild types." Amplified with investigations on the competi-
tive value of observed differences in characteristics the accumulated knowledge
promises to be far more significant for reaching a satisfactory solution of tax-
onomic problems than are the results of those "standard tests" which at present
are the chief basis of our methods of differentiation, and which are generally
performed under conditions and with media utterly at variance with the "natural"
ones. (See, in this connection, e. g., van Niel, 1949b; Winogradsky, 1952.)

But however much the approach outlined above may contribute to a better
understanding of the microorganisms in question, we should not anticipate that
it will solve the "species problem," and this for the reasons already mentioned.
Once this is recognized, the question arises whether a more promising attack can
be suggested. In this connection I believe that Winogradsky's latest publication
(1952) has opened up prospects for sound developments. In essence he proposes
the establishment of "biotypes," rather than species, genera, etc., for those groups
of bacteria that are easily recognizable and accessible and that represent special
and distinctive patterns of characteristics which can be related to the normal role
of the organisms in nature. Around these "biotypes" are to be grouped the
numerous "satellites," comprising the strains that differ from the "types" only
with respect to some secondary details, these to be indicated simply by numbers.
Abandoning all attempts at further classification, Winogradsky concludes (1952,
pp. 130-131) :

. . . je ne pense pas que ce travail [i.e., to reconstruct present systems of classifica-
tion along these lines] puisse etre entrepris avant longtemps; je crois neanmoins, que
mes suggestions se montreront utile du jour ou les bact^riologistes, fatigues par I'aspect
touffu de la systematique bact^rienne, songeraient a la reformer en faveur d'un mode
plus simple et, a mon avis, plus rationnel.

II se peut que certains microbiologistes soient cheques par I'idee de supprimer la
classification Linneenne dans le cas des bacteries, habitues qu'ils sont de s'en servir
pour toute classification.

Or, tout travail 6tabli selon les regies de cette classification devrait etre base avec
quelque precision sur le principe philogenetique, qu'il est impossible d'appliquer aux
bacteries. II serait done plus correct de nous borner a I'appliquer au regne animal et
au regne v^g^tal, 6u il est bien k sa place, sans chercher a englober dans sa sphere les
formes plus el^mentaires de la vie.

VAN NIEL: SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 101

On devrait se contenter de ce que les bacteries se laissent tout de meme systematiser,
sous forme de groupes representes par des Biotypes, qui sont, eux, bion differenciables.

At first sight this approach may appear simply to avoid the species problem
by substituting for it the new one of what shall be considered the criteria for a
biotype. Yet this mere substitution may exert a healthy influence because the
name is still untinged by connotations such as those that have come to be asso-
ciated with the term "species." Also in connection with the problems to be dis-
cussed in the next section, acceptance of Winogradsky's proposal would go far
in removing obstacles that must otherwise be faced.

The validity of these statements is well illustrated by the following example.
It can be reasonably expected that some of the "biotypes" established in the course
of time would correspond more or less closely with now accepted "true species"
of bacteria. The use of the latter term has, however, been restricted and is gen-
erally applicable only to the first described species of a genus, a situation that
results from the virtually complete acceptance by bacterial systematists of the
rules of nomenclature adopted by the botanists. Now, this inevitably entails the
consequence that a number of "type species" represent bacteria that have not
been studied in sufficient detail to make them acceptable as biotypes in the sense
in which I have interpreted this expression in the preceding pages, and which
would definitely include the availability of specific elective culture procedures
for the organism in question. Adherence to the present code of bacterial nomen-
clature would make it difficult to change a large number of "type species"; but
when "biotype" is used instead, no one is hampered by "rules and regulations"
that have not yet been formulated.

Winogradsky's suggestions therefore appear to me worthy of careful consid-
eration and strong support; in a sense they represent a logical development of
my own ideas, expressed some years ago as follows (van Niel, 1946, pp. 297-298) :

Discontinuation of the terms species and genus for bacteria, along with the introduc-
tion of multiple keys, would eliminate some of the difficulties now encountered, because
it would insure a far greater autonomy to specialists in dealing with their own groups
and problems, unencumbered by the exigencies of different groups. There would be no
need for the sort of consistency required as the foundation of a single system of clas-
sification. Whether the further elaboration of a rational nomenclature along the lines
laid down by Orla-Jensen, and further expanded by Kluyver and van Niel, would prove
adequate, or whether it might even be preferable to drop the use of Latin names with
their taxonomic implications, is a matter for future developments. And, while I am
fully in agreement with the opinion that stability in nomenclature is of great importance,
I must once more insist that, in the long run, it may turn out to be easier to gain adher-
ence to a more rational, modernized system than to the current one.

The Genera, Families, and Orders of the Bacteria and Bluegreen Algae

In the development of our system of classification the discovery and naming of spe-
cies with a generic and specific name came first. Grouping into Genera was followed
by grouping of Genera into Tribes and Tribes into Families and Families into Orders.
In developing the key in the reverse order, the authors of the keys in the Manual were
forced to use initially for identification characters which by their very nature are
largely indeterminable. — V. B. D. Skerman, 1949, pp. 177-178.

What made Winogradslvy (1952) grant that the systematics of plants and
animals on the basis of the Linnean system is defensible, while contending that

102 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

a similar classification of the bacteria is out of the question? The answer must
be obvious to those who recognize in the former an increasingly successful attempt
at reconstructing a phylogenetic history of the higher plants and animals, based
on comparative-anatomical, embryological, distributional, ecological, and paleon-
tological studies and who feel that comparable efforts in the realm of the bacteria
(and bluegreen algae) are doomed to failure because it does not appear likely
that criteria of truly phylogenetic significance can be devised for these organisms.
Forty-five years ago Orla- Jensen (1909) believed that it was possible to formu-
late an acceptable phylogeny of the bacteria by means of physiological-biochemical
considerations. But it has since been shown that there are compelling reasons
for doubting the validity of Orla- Jensen's premises (Oparin, 1938; van Niel,
1946; 1949a).

Nevertheless, systems of classification of these organisms, complete with
genera, families, and orders have been developed in the course of the past century;
they have become more and more elaborate and complicated, and seem to be taken
seriously in at least some quarters. The simplest explanation for this attitude is
that classifying organisms in this manner has become an accepted habit, so
ingrained that one just kept on doing it, to paraphrase the last verse of Paul
Geraldy's "Meditation"^ :

On prend I'habitude, vite,
d'echanger de petits mots.

Quand on a longtenips dit les memes,
on les redit sans y penser.
Et alors, mon Dieu, Ton aime
parce qu'on a commence.

When Cohn (1872) first proposed his six bacterial genera he was, however,
quite explicit in stating that these units did not have any phylogenetic signifi-
cance. They were simply "form-genera," providing descriptive names for groups
of bacteria possessing similar shapes. Though useless as guides to "natural rela-
tionships," these categories greatly facilitated the naming and identification of
bacteria. Once a newly isolated culture had been characterized as composed of
short rods, for example, it was thereby fixed as a Bacterium species, and the
establishment of its possible identity with earlier described bacteria could be
restricted to a comparison with the known members of this genus.

Cohn subsequently (1875) expanded his system considerably, integrating the
( form- ) genera of the bluegreen algae with those of the bacteria as components
of the class or family of the Schizomycetes. With further increase in our knowl-
edge of these microorganisms, owing largely to advances in microscopic tech-
niques, additional differential properties were discovered. Incorporation of such
characteristics in the descriptions consequently led to modifications of the diag-
nosis of several genera, and to the proposal of many new ones. During this period
a number of more or less "private" systems of classification were developed, such
as those of Zopf, Marpmann, de Bary, Fischer, Lehmann and Neumann, Migula,
Kruse, Orla-Jensen, and Chester, each one commanding a certain number of
adherents, with the result that various authors might refer to one and the same
organism by several different names. An extensive study of this somewhat con-

1. Paul Geraldy, Toi et Moi, Paris: Stock, 1922.

VAN NIEL: SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 103

fusing situation was made by a Committee of the Society of American Bacteri-
ologists whose members published reports and recommendations (Winslow et al.,
1917, 1920) for the development of a more uniform system of classification of
the bacteria, largely based on Buchanan's proposals (1916-1918). This became
the nucleus from which originated in due course Bergey's Manual of Determina-
tive Bacteriology (1923-1948), prepared by an ever-increasing number of spe-
cialists with expert knowledge of various groups of bacteria (Breed et al., 1948).
The classification followed in this handbook has been more and more generally
adopted and is today the most widely used.

But in spite of the growing recognition afforded the painstaking efforts rep-
resented by this collaborative enterprise, the end result has never been wholly
satisfactory, and each successive edition has come in for a certain amount of
criticism. Objections have been raised to the inclusion of a vast array of poorly
characterized species, for example by Winogradsky (1952) and Skerman, the
latter presenting a well-reasoned argument (Skerman, 1949, p. 175) :

Many of the descriptions of bacteria in Bergey's Manual of Determinative Bac-
teriology are decidedly poor when viewed from present-day standards. Some will be
difficult to improve since a number of the original cultures have probably been lost.
The original descriptions which still remain on record present us with an awkward
problem in establishing priorities. Some of these descriptions are so inadequate that
one description could be equally well applied to many new isolates. The original authors
cannot be blamed for the inadequacy of these descriptions which no doubt conformed
to the standard of the day and it would be a breach of ethics to refuse recognition of
these descriptions. Nevertheless present-day workers cannot regain the original cultures
in some instances to subject them to further examination and would-be key formers are
handicapped by the lack of this information. Thus one cause of the chaotic state of
bacterial nomenclature is the lack of "type" specimens regarded as essential by syste-
matic botanists. There is only one remedy for this, namely the redescription of all
available cultures according to a certain code which should be applied to all bacteria
alike. On the basis of these descriptions the organisms should be renamed, for the
most part with the names they now possess. Priorities should be based on these names
and all descriptions and names for which there are no procurable cultures should, by
common consent, be discarded.

Besides, the characterization of many of the genera has been found wanting, and
again I quote from Skerman (1949, p. 176) :

There is also need for more precise definitions for genera. In the hands of the
authors of most of our textbooks the term "definition" has entirely lost its meaning.
Many of the definitions contain very little which is definite. They approach more to-
wards condensed, and often confusing, descriptions which attempt to embrace all the
possibilities which one may encounter among the species in the genus rather than a
precise statement of the characters which can be uniformly found among all or the
majority of species within that genus to be distinguished from other genera.

And finally the taxa of higher order suffer from the same deficiency, here even
more aggravated because, as Skerman remarks (ibid., p. 177) :

A close study of the number of determinable characters which could represent all
species within a genus would reveal this number to be very small. The number of
characters which are common to all genera within a tribe must inevitably be smaller,
and would continue to diminish as groupings become broader.

These remarks should suffice to indicate that the satisfactory demarcation of
systematic units above the rank of species is beset with even greater difficulties

104

A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

than that concerned with mere species. Occasionally, however, a particular
property has been encountered whicli is qualitatively so striking that it would
appear suitable as the special distinguishing character of a family or order. This
happened, for example, when Migula (1897) introduced the order Thiobacteria
for those microbes which Winogradsky had called "sulfur bacteria." It was the
first time that the bacteria were divided into two separate orders; and Migula
justified the procedure by emphasizing that both the cellular organization and
the physiology of the sulfur bacteria were clearly distinct from those of the
"true" bacteria, or Eubacteria.^ Morphologically the former are conspicuous on
account of their relatively large size and their content of sulfur globules ; physio-
logically they represent the prototype of the autotrophic bacteria ; they can grow
in strictly mineral media, and are dependent on an external supply of sulfide
which is oxidized to sulfate.

It was also the first time that a physiological property was used for the
establishment of a large systematic group of the bacteria. Coupled as it was
in this case with some morphological peculiarities, this may have appeared de-
fensible. But later developments have shown how much confusion was created
by this ostensibly simple expedient.

Elsewhere I have sketched these developments in some detail (van Niel,
1944) ; suffice it here to recapitulate the major aspects. The Thiobacteria, in
1900, comprised two subgroups, viz., the colorless, filamentous organisms which,
except for lack of pigmentation, closely resemble the bluegreen algae of the
family Oscillatoriaceae (see, e.g., Pringsheim, 1949), and the red-colored, so-
called purple sulfur bacteria which are much more "bacteria-like," though gen-
erally much larger. Within a decade, however, two more groups of organisms
were discovered with characteristics that made their incorporation into one or
the other of Migula 's orders largely a matter of personal preference. Tliese
were the small, colorless Thiohacillus species, physiologically typical sulfur bac-
teria, but morphologically in no way distinguishable from many eubacterial
types, and the small purple bacteria that are physiologically not sulfur bacteria,
though their pigment system, composed of chlorophyllous and carotenoid com-
ponents, closely resembles that of the purple sulfur bacteria.

The properties of these four groups obviously show "interrelationships"
which can best be presented in the form of a diagram, as follows :

Thiobacillus species

(similar ptiysiology)

(morpliologically "true bacteria")

Colorless, filamentous
sulfur bacteria

Nonsulfur purple bacteria

(intracellular

sulfur globules)

(similar pigment
systems)

Sulfur purple bacteria

2. In an earlier publication (van Niel, 1944) I erroneously stated: "One looks in
vain, however, for an exposition of the reasons which had induced Migula to create the
new orders" (p. 71). A vague attempt at rationalizing this measure can be found in
the brief section on the sulfur bacteria at the end of Vol. 1 of Migula's System.

VAN NIEL: SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 105

This diagram shows that the new situation called for a decision as to the rela-
tive importance of the characters that can be used to link the different groups.
Obviously, a combination of morphological and physiological properties, once
justified because "intermediate" gi'oups were not known, was no longer ade-
quate. The formulation of a diagnosis of separate orders had, from now on, to
be based on either morphological or physiological features. Even this could
not provide a fully satisfactory solution to the problem of establishing larger
systematic units, however. For, when "morphology" was given preference, there
would still be the question whether the occurrence of sulfur globules, the indi-
vidual cell size, or the presence or absence of the special pigment system was
considered the most significant, while preferential use of physiological charac-
ters would imply the need for "grading" the respective values of sulfide oxi-
dation and pigment formation.

Of course, the very admission of physiological characters in bacterial sys-
tematics might be blamed for the confused situation here discussed. Would it
not have been better if such criteria had been left out altogether in the crea-
tion of the two orders? In that event the filamentous colorless sulfur bacteria
could have been neatly segregated from the Thiohacillus group and from the sul-
fur and nonsulfur purple bacteria, regarding the latter assemblage as members
of the order Bubacteriales. While this may be considered a great improvement,
it nevertheless serves merely to shift the basic problem to the question of how
families should be defined. It can still be maintained that there would be ample
justification for the creation of a large systematic group of all the purple bac-
teria, especially because it is now known that the pigment system of these or-
ganisms confers upon them the ability to carry out an "aberrant" photosynthetic
mode of life (Molisch, 1907; Buder, 1919; van Niel, 1931, 1941, 1952). And
many arguments could be advanced to defend the thesis that such a unit, which
would also accommodate the green sulfur bacteria, has considerably greater
phylogenetic significance than, for example, groups comprising all Gram nega-
tive, nonsporeforming, polarly flagellated rod-shaped bacteria, regardless of
their physiological properties.

The preceding discussion of the systematic status of the sulfur- and purple
bacteria may have served to illustrate the difficulties inherent in attempts to
accomplish primary divisions in the realm of the bacteria. Similar difficulties
are encountered at lower levels, and here, too, the problem must be faced
whether physiological characters are admissible. In some circles the idea that
they are not still prevails; on the other hand, the large number of generic
names with definite physiological connotations {Thiohacillus, Acetohacter, Lac-
tohacillus, Projnonihacterium, Hydrogenomonas, Nitrohacter, Methanococcus,
Photohacterium, etc.) testifies that this attitude is not universal,

Manj^ of these names were introduced by Beijerinck and Winogradsky, and
it is clear that the ecological-physiological approach to general microbiology of
these two masters was largely responsible for the practice. The discovery that
a particular tj^pe of metabolism (sulfur oxidation, acetic acid production, lactic
or propionic acid formation, hydrogen or nitrite oxidation, methane production,
or ability to luminesce) seemed to be closely associated with certain types of
bacteria that were both easily procurable and readily distinguishable, compris-
ing relatively small groups of organisms with many common morphological

106 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

characteristics in eacii group, naturally suggested the existence of a high degree
of specificity which was reflected in both physiological and morphological prop-
erties. Since each group contained representatives exhibiting minor differences,
one from the other, in shape, size, color, or physiology, it must have seemed
eminently rational to consider these as species and the entire group as a genus.
A logical consequence of this approach was Orla-Jensen's classification (1909)
in which the bacteria were assigned to genera that were defined by a combi-
nation of morphological and physiological characters. By considerably extend-
ing the number of differential morphological traits and incorporating the newer
concepts of the mechanisms of biochemical processes, derived from studies on
the comparative biochemistry of microorganisms, Kluyver and van Niel (1936)
sought to provide a more up-to-date system along the same general lines.

Some systematists have, however, consistently condemned the use of physio-
logical criteria for the definition of even such small taxonomic units as genera.
They seem to agree with Lehmann and Neumann (1927, 2:190) who wrote:

Dass die Systematik der Spaltpilze und der ihnen nahestehenden Mikroorganismen
genau so wie die aller anderen Lebewesen zunachst nacti morpliologischen Grundsatzen
(Form, Begeisselung, Sporenbildung) versucht werden muss, ist klar, trotz aller oben
angegebenen Schwierigkeiten.

Statements to this effect can be found, for example, in Prevot's extensive paper
on the classification of the anaerobic cocci (1939, p. 50) :

. . . nous pensions qu'il est possible aujourd'hui de chercher a adapter au monde bac-
terien les doctrines classiques qui ont reuissi pour le regne vegetal et le regne animal
entre les mains des freres de Jussieu, de Cuvier, de Geoffrey Saint-Hilaire, etc., et des
modernes: il existe une relation enti'e la valeur des characteres et le determinisme du
groupement des Bacteries, et cette relation est commune au trois mondes, vegetal, ani-
mal et bacterien: les characteres morphologiques ont la priorite sur les characteres
physiologiques.

On the basis of such considerations Prevot has even developed a set of rules for
the delineation of taxa of higher order, as follows (ibid., p. 61) :

Les characteres de morphologie generale sont des characteres de classe.

Les characteres de reproduction (simple, par spore, par conidie) sont des characteres
d'ordre.

Les characteres de structure cytochimique (coloration de Gram) sont des characteres
de famille.

Les characteres de morphologie speciale (ectoplasme, biometvie, directions de division,
arrangement cellulaire) sont des characteres de genre.

Les characteres physiologiques (culturaux, pathogenes, biochemiques) sont des charac-
teres d'espece.

Les characteres physiologiques secondaires et serologiques (agglutination) sont des
characteres de variete ou race.

From a scientific viewpoint it is, however, astonishing that the validity
of such verdicts generally seems to have been taken for granted; rarely, if ever,
has an attempt been made to .justify the belief that for the purpose of classifi-
cation of the bacteria morphological characters are more significant than physi-
ological or biochemical properties. Occasionally it is possible to infer from the
context the reasons for this notion. The reference to Jussieu, Cuvier, and Saint-
Hilaire in the above quotation from Prevost, for example, indicates the trend of
thought. And Kluyver and van Niel ( 1936, p. 370) expressed this still more directly:

VAN NIEL: SYSJEMATICS OF THE RACTERIA AND BLUECREEN ALGAE 107

... It cannot be denied that the studies in comparative morphology made by botan-
ists and zoologists have made phylogeny a reality. Under these circumstances it seems
appropriate to accept the phylogenetic principle also in bacterial classification.

The question then arises in what characters phylogeny expresses itself. There is no
doubt that in this respect morphology remains the first and most reliable guide.

But is this inference concerning the superior value of morphological prop-
erties actually applicable to the bacteria and bluegreen algae ? It has been used
to justify the establishment of taxa above the rank of species for organisms
with similar outward shape, and the tacit implication has been that such taxa
reflect truly "natural relationships." This, however, is open to serious doubt,
as illustrated by the genus Sarcina, comprising bacteria of spherical shape, di-
viding in two or three perpendicular directions, thus producing squares, flat
sheets, or cubical packages. It would not be surprising to find that bacteriolo-
gists familiar with these organisms balk at the notion that the aerobic ;S^. lutea,
the anaerobic S. ventriculi, S. maxima, and S. methanica, exhiljiting an alco-
liolic, butyric acid, and methane fermentation, respectively, the lialophilic ;S'.
f/igantea, and the motile, sporeforming S. ureae represent a group of phylo-
genetically closely related types.

It seems to me that the most important reason for much confused thinking
about bacterial classification is that Cohn's careful appraisal of the meaning
of his "form genera" has not been given the attention it deserves. Proponents
of the view that morphological characters are of primary importance for the
establishment of natural relations appear often to have failed to realize that
only those associated with the developmental history or embryology of a higher
plant or animal have served to trace its phylogeny. Even though a sufficiently
advanced knowledge of the various types of organisms may sometimes permit
the use of a special shape as the only character needed for the determination
of relationships, this approach can be very precarious, as shown, for example
by Ginkgo hiJoha and the whales. Now, most bacteria and bluegreen algae do
not exhibit the kind of developmental history that can be useful in reconstruct-
ing phylogeny. Once this is recognized, genera such as Sarcina stand revealed
as signifying no more than the "form genera" of Cohn.

It should thus be evident that many of the morphological features used in
the past as differential characters in the classification of bacteria and blue-
green algae cannot be depended upon as guides to phylogeny. Is there any
reason to believe that physiological and biochemical properties are more sig-
nificant in this respect? A priori this possibility cannot be dismissed; there
does not seem to be any valid basis for Prevot's insistence that these can be
used only for the differentiation of species but not of higher taxa. In fact, the
group of photosynthetic bacteria (green and purple sulfur bacteria, and non-
sulfur purple and brown bacteria), as also that of the lactic acid bacteria in the
sense of Orla- Jensen can easily be regarded as phylogenetically much more
homogeneous than the Sarcina group, in spite of a considerably diversified
morphology among the organisms comprising the first two assemblages. In
the photosynthetic bacteria the cell shapes range from small spheres and short
rods to large vibrios, rods, and spirals, and the lactic acid bacteria include strep-
tococci, tetraeocci, short rods, and long rods, even to the point of becoming
filamentous.

108 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

But, while discrediting Prevot's contention, this argument does not mean that
a particular type of metabolism is a more reliable index of phylogeny than is the
gross morphology of the cells. The ability to carry out a lactic acid fermentation,
for example, is not the prerogative of the "lactic acid bacteria"; it has been
found also in some members of the facultatively anaerobic sporeformers. Simi-
larly, a typical alcoholic fermentation is produced by Sarcina ventriculi and by
Pseudomonas lind^ieri, and a propionic acid fermentation by Propionihacte-
rium species as well as by some anaerobic micrococci, anaerobic sporeformers,
and facultatively anaerobic myxobacteria of the Cytophaga type. In these
cases it is as difficult to find convincing grounds for the claim that the organisms
characterized by similarity in metabolism are phylogenetically closely related as
it is to assign natural relationships primarily on the basis of cell shapes.

Awareness of this situation led Kluyver and van Niel (1936) to propose that
■a bacterial genus be defined both morphologically and biochemically. In this
manner cross-relations in these two respects could find adequate expression, and
homogeneity in the composition of the individual genera was insured. However,
it did not solve the problem of a phylogenetic classification; once more it was
necessary to make a choice between morphological and physiological characters,
now for delineating families, and from the foregoing discussion it would appear
that a decision in this respect had to be an arbitrary one.

Besides, another difficulty presents itself, even on the genus level, because not
all biochemical properties appeared equally suitable as generic characters. In
some cases a guiding principle can be found to aid in evaluating various fea-
tures. Thus, the lactic acid fermentation brought about by the lactic acid bac-
teria, the mixed acid fermentation of Escherichia coli and its relatives, the
ethanol-butanediol fermentation of Aerohacter and Aerohacillus, the propionic
acid fermentation, the butanol-acetone fermentation, the ethanol-acetone fermen-
tation of Bacillus macerans, the alcoholic fermentation of Sarcina ventriculi and
Pseudomonas lindneri, represent as many distinctive metabolic patterns. It was
therefore felt that they provide legitimate criteria for separate biochemical
genera, while the differential utilization of some particular members of the
class of carbohydrates, presumably depending merely on the presence or ab-
sence of specific carbohydrases, was deemed useful only for the demarcation of
species. There are, however, many instances in which the situation is more com-
plicated because one and the same bacterium may exhibit a number of different
metabolic patterns, each one of which would be suitable for the definition of a
"biochemical genus." This again implies the need for making a choice. As a
way out of the dilemma Kluyver and van Niel (1936, p. 389) suggested:
... In those cases it is, of course, desirable to classify the organism in question
according to its most characteristic type of katabolism, that is, the type which permits
the distinction from otherwise related organisms. This implies that for organisms capable
of development under anaerobic conditions the katabolic process involved in this mode
of life has been determinative, regardless of the question whether or not the organism
also possesses a respiratory mechanism. If two different types of anaerobic katabolism,
e.g., saccharolytic and proteolytic, are represesnted, the latter, as being the rarer, has
been decisive.

It will be superfluous to belabor the point that this passage contains nothing
to suggest a phylogenetic basis for the choice, nor does it seem likely that a
sound one can be discovered. Nevertheless, the classification proposed has much

VAN NIEL SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 109

to recommend it, because it permits the ready assignment of a particular bac-
terium to a specific and small group as soon as its general morphological and
biochemical characters are known. Final identification then requires compari-
son with other members of only this assemblage. The advantage is, therefore,
of the same kind as that offered by Cohn's "form genera," and the categories
resulting from the combination of morphological and biochemical properties are,
in a sense, quite comparable though more numerous. In view of the great in-
crease in the number of different types of bacteria discovered in the course of
time this is a distinct benefit. Undoubtedly, such strictly utilitarian considera-
tions were responsible for the application of biochemical criteria in the manner
outlined above, as shown especially by the decision to use the "rarer" of two
otherwise equivalent characters.

But if the homogeneous, morphologically and biochemically defined genera
cannot lay claim to phylogenetic significance, the superstructures of tribes, fami-
lies, and orders can do so even less. It follows that the existing systems of clas-
sification of the bacteria and bluegreen algae should not be considered "natural"
ones. If this be granted, the question whether retention of such systems is ad-
visable can be examined more critically.

At first sight the now more or less generally accepted genera and families
of these organisms, even if devoid of phylogenetic meaning, might appear to
serve as a fully satisfactory framework for purely determinative purposes. This,
however, can be contested on the ground that they are too rigid, because the
families, tribes, and orders represent collections of genera grouped together on
the basis of only one set of arbitrarily chosen "primary" characters. While these
may be the most useful ones as determinative aids in some instances, in others a
different set of primary divisions would be preferable, thereby yielding a super-
structure of different composition. It is obviously inadmissible to include a par-
ticular "genus" in two or more different families, tribes, or orders. But if these
larger groups are considered as no more than convenient contrivances for rapid
identification, there is no need to insist on an "either-or" approach. By discon-
tinuing the use of families, tribes, and orders it becomes possible to construct a
diversity of groupings in which all the different opportunities for emphasizing
similarities in various respects can be expressed. It seems to me a dubious gain
to have all the photosynthetic bacteria assembled in a suborder, Khodobacteri-
ineae, if this practice eliminates the possibility of recognizing the existence of
the large group of "sulfur bacteria" comprising only some of the photosynthetic
bacteria in addition to organisms now incorporated in the orders Eubacteriales
(genus ThiohaciUus) and Chlamydobacteriales " (families Beggiatoaceae and
Achromatiaceae). Such an entity as the sulfur bacteria remains an extremely
useful assemblage, since it represents an ecological-physiological community of
all the conspicuous inhabitants of natural environments in which hydrogen
sulfide is present.

It is not hereb}^ intended to dispute the probability that the photosynthetic
bacteria actually represent a phylogenetically related group, nor that the Beg-
giatoaceae might be similarly regarded. But the phylogenetic relationships of
the other "sulfur bacteria" are far less certain. Clearly, it is not imperative
that even the probable affinities of the first-mentioned organisms be given recog-
nition by uniting them into a family, tribe, suborder, or order; and if doing so

no A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

implies that bacteria with doubtful phylogeny must then be treated likewise,
there seems to be much in favor of abandoning the practice. If and when the
natural relationships of a large number of bacteria have been unambiguously
established, it would become advisable to consider the construction of a system
of classification based on phylogeny.

As long as this remains a pious hope for the future, one might do well to
approach the problem of the classification of bacteria and bluegreen algae in the
manner suggested by Winogradsky's latest recommendations. Substitution of
"biotypes" for genera and species, and the use of common names, such as "sulfur
bacteria," "photosynthetic bacteria," "chemoautotrophic bacteria," "denitrifying
bacteria," "nitrogen-fixing bacteria," etc., instead of the Latin names represent-
ing taxonomic units with definite phylogenetic implications, would permit the
development of more rational arrangements for the rapid identification and com-
parison of the organisms. This problem calls for an elaborate system of cross-
indexing of their properties, and the present organization, based on the Linnean
approach, not only is unjustifiably pretentious, but also impedes the best utiliza-
tion of established characteristics because they are employed for the construction
of mutually exclusive combinations. While much can be done to remedy the re-
sulting situation through the preparation of mechanical keys, such as the emi-
nently useful one developed by Skerman (1949), a more radical departure from
accepted procedure remains desirable in the opinion of the writer.

In this connection attention should be called to the ideas recently expressed
by C. H. Andrewes concerning the classification and nomenclature of viruses
(1952, p. 136) :

The nomenclature of plants and animals has been the subject of much controversy
and change, owing largely to the fact that the earlier names were bestowed without
understanding of the principles of taxonomy as we now know it, often without reference
to type material, and on the basis of very inadequate descriptions. In the reviewer's
opinion, such troubles would be avoided in the virus field by dating valid nomenclature
in this group not from the time of Linnaeus 200 years ago, but from a date to be de-
cided upon in the future. . . .

A very few descriptions of viruses published hitherto would satisfy those who are
seriously considering the matter today. Binomials are not in common use for any viruses,
and there seems therefore everything to be gained by starting with a clean sheet. . . .
Such virus names already published as seem suitable would also be validated, but virus
nomenclature need not be forever overlaid by the dead hand of bad naming, linked to
descriptions which are hard to interpret and are based on unsuitable guiding principles.
If, however, students of viruses take thought in time and base their classification and
nomenclature on solid foundations with reference from the very beginning to type mate-
rial, they can forever be free from the nightmares of change and contentiousness which
bedevil nomenclature in other fields.

In contrast to the quotation at the start of this paper, the above, with a few
minor modifications, seems eminently applicable to the problems presented by
the classification of the bacteria and bluegreen algae.

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CLASSIFICATION OF THE ALGAE'

By GEORGE F. PAPENFUSS

University of California, Berkeley

Introduction

A GENERAL TREATMENT of sucli a hetcrogeneoiis assemblage of organisms as the
algae may fittingly be introduced with a statement of the criteria used in the
delimitation of the group.

These plants are readily separated from those next above them in the evo-
lutionary scale, the archegoniate plants, by the fact that their reproductive or-
gans lack a primarily produced sterile jacket of cells. (The antheridium of the
Charophycophyta is an exception.) The separation of some algae from certain
members of the other groups of simple organisms, such as the bacteria, the
fungi, and the protozoa, is much more difficult and not infrequently the as-
signing of an organism to the algae or to one of these groups is a purely arbi-
trary procedure.

Although the major taxa of algae show little or no relationship to one an-
other, the group as a whole is clearly distinguished from other simple organ-
isms by the ability of a great majority of the species to synthesize organic
compounds by the process of photosynthesis. There are many exceptions to this
rule but the saprophytic, parasitic, or holozoic forms usually reveal their al-
liance to autotrophic types by their structure, life history, and storage products.
In very many instances the heterotrophic forms appear to have been derived
from photosynthetic types. The autotrophic bluegreen algae may be distin-
guished from the autotrophic bacteria by their possession of chlorophyll a and
the evolution of oxygen as a by-product of photosynthesis.

In modern systems of classification the algae comprise more than half the
number of plant phyla. Of the known species, however, they constitute less
than 10 per cent. The disproportionately large number of major algal taxa
reflects the great diversity in the structure, reproduction, and metabolism of
these plants as contrasted with the remainder of the plant kingdom.

Within the confines of this brief treatment, my review of the history of the
classification of the group of necessity will be confined to the broad outlines of
the system. Attention will also be given to the history of the discovery of sex
in the algae and to the growth in knowledge of their life histories since ad-
vances in these aspects of phycology have almost always contributed to a better
understanding of the interrelationships and phylogeny of the groups concerned.

The nomenclature of the majority of algae, like that of most plant groups.

1. I am deeply indebted to Dr. Johannes Proskauer for critically reading the manu-
script and for his many constructive suggestions. I should also like to thank Professor
G. M. Smith and Dr. T. V. Desikachary for kindly reading the manuscript and making
helpful suggestions.

[115]

116 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

begins with Linnaeus' (1753) Species pl<intarum. It is of interest to note that
in Linnaeus' system the algae were grouped along with the pteridophytes, mosses,
and fungi in a single class, the Cryptogamia, whereas the phanerogams were
divided into 23 classes. Linnaeus recognized 14 genera of algae but only four
of them {Conferva, TJlva, Fucus, and Char a) comprised algae in the current
sense and two others {Byssus and Tremella) included a few species of algae.
During the following fifty years botanists were content to accept the classifi-
cation of Linnaeus and, with very few exceptions, always referred their algal
species to his genera. Usually, Conferva received the filamentous, TJlva the mem-
branous, and Fucus the fleshy forms, and treatises were written on the largest
of these three genera, namely, Fucus and Conferva (e.g., by Gmelin, 1768; Es-
per, 1797-1808; Dillwyn, 1802-1809; Vaucher, 1803; Lamouroux, 1805; Turner,
1808-1819). Chara was frequently excluded from the algae.

Stackhouse was the first to break away completely from the custom of recog-
nizing only the Linnean genera of algae. He concerned himself especially with
the British species of the marine genus Fucus, and his study of them brought him
to the realization that this comprehensive genus comprised a large number of
distinct taxa which he accordingly removed to new genera. In a series of three
works (1795-1801, 1809, 1816) he divided Fucus into 67 genera (including one,
Pygmaea, now known to be a lichen). Papenfuss (1950a) has typified Stack-
house's genera and has discussed their fate.

In 1813 Lamouroux published his Essai sur les genres de la famiUe des thaJ-
lasssiopJiytes non articulees, which formed an important advance in that he,
in addition to establishing a number of new genera, mostly as segregates from
Fucus, proposed a system of classification of the algae into major taxa (orders)
based in part on color.

C. Agardh (1817, 1824) and especially Harvey (1836) made significant
modifications in Lamouroux' system. Harvey divided the algae with which he
was concerned into the four divisions (phyla) Melanospermeae (brown algae),
Rhodospermeae (red algae), Chlorospermeae (green algae), and Diatomaceae
(diatoms and desmids). Without realizing it, Lamouroux and Harvey thus in-
troduced a biochemical character into the classification of the algae. Subse-
quent investigations have firmly established the soundness of this character as
an indicator of phylogenetic afiinit}^ On the basis of pigment composition and
other characters of seemingly comparable merit several major taxa, in addition
to Harvey's original four, have now been recognized. In the present treatment
the assemblage is considered as comprised of seven phyla and in addition three
classes, one of which, the Schizophyceae (bluegreen algae), is regarded as con-
stituting along with the class Schizomyceteae (bacteria) the phylum Schizo-
phyta, and two of which are composed of forms of uncertain affinity. The his-
tory of these major groups is reviewed in the separate sections into which the
body of this chapter is divided.

In 1836, Endlicher divided plants into two kingdoms — Thallophyta and Cor-
mophyta. The designation Thallophyta was later used as a phyletic name for
all plants below the level of the Bryophyta. Although the term still has merit
as denoting plants of a certain morphological type, it is now generally recog-
nized that the various classes of algae and fungi can no longer be regarded as
belonging to a single phylum.

PAPENFUSS: CLASSIFICATION OF THE ALGAE . 117

For a long time all algae exhibiting movement — diatoms, desmids as well as
flagellated green, yellow-green and golden-brown forms — were automatically re-
garded as animals. Although the zoologists in the course of time relinquished
the diatoms and desmids, the flagellated forms and tlieir amoeboid relatives
(but frequently not their nonmotile unicellular and multicellular relatives) are
still considered as belonging to the artificial phylum Protozoa.

Among the early biologists who regarded some of the flagellated organisms
as algae rather than protozoa are Siebold (1848, 1849), Braun (1851), and Colin
(1852). The bulk of the so-called Flagellata, however, for a long time remained
the exclusive domain of zoologists. Klebs (1883) was the first to emphasize the
plantlike nature of many of these forms and in the ensuing years the group
Flagellata received increasing attention from botanists and was included among
the algae in treatises on the plant kingdom (Warming, 1890; Engler, 1898;
Senn, 1900; and others). Epoch-making contributions that revealed tlie inti-
mate relationship of some of these forms to nonflagellated unicellular and multi-
cellular or so-called "algal" types were made by Bohlin (1897a, 1897b), Luther
(1899), Klebs (1912), and Pascher (1912b, 1914). The history of this change
in the outlook of botanists in regard to the systematic position of the flagel-
lates is reviewed in the appropriate sections below, especially in that on the
Euglenophycophyta.

The discoveries and growth in knowledge of sex in plants form an extremely
interesting chapter in the history of botany. The algae provide particularly
favorable material for the study of sex and have been used to advantage in this
connection since the middle of the last century (cf. Kniep, 1928).

Although numerous observations had been made on the reproduction of algae
before 1853 and many botanists had come to believe that some algae exliibited
a sexual process, the established facts in support of such a view were extremely
meager. Some forms appeared in places and under circumstances that neces-
sitated the assumption of spontaneous generation. In this way, Meyen (1827)
explained the appearance of small algae, known as "Priestley's matter," in stag-
nant water and even in closed vessels. Kiitzing (1833a) and others put forth
the view that the simplest algae once produced spontaneously could develop
according to circumstances into a variety of algae. When the zoospores of an
undisputed alga were seen in the process of liberation from the thallus, the
phenomenon was interpreted as a changing of the plant into an animal. The
remarkable thing is not so much that such views were entertained but that the
majority of biologists of the time combined with them a belief in the immut-
ability of species.

The first alga suspected of showing a sexual process was Spirogyva. Hedwig
(1798) was of the opinion that the zygospores, which were discovered by 0. F.
Miiller in 1782, were formed as the result of a sexual act. Vaucher (1803) also
studied conjugation in Spirogyra and related forms but he, like many later
botanists, was not fully convinced of the sexual nature of this process since it
was difficult to conceive of a sexual process without the criterion of a morpho-
logical difference between the sex organs; and furthermore, it was evident that
one and the same filament could both deliver and receive substance from a
neighboring filament. During the first half of the nineteenth century and for
some time afterward (as recently as 1916 and 1926 by West [1916, p. 135]

118 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

and Schiffner [1926]) a sharp distinction was made between fertilization (the
union of morphologically different cells) and conjugation (the union of cells
of like form). Thwaites (1848), the discoverer of conjugation in diatoms, was
the first to regard conjugation as a sexual process without qualification.

Vaucher (1803) also studied the genus that was later named for him {Vau-
cheria) and designated as sex organs the structures now referred to as anthe-
ridia and oogonia. Vaucher was far in advance of his time, however, and his
conclusions were not accepted. As late as 1847 Nageli, for instance, thought
that red algae were tlie only algae that reproduced sexually but even in this
assumption he was entirely wrong inasmuch as he believed the tetrasporangia
to be the female sex organs.

The turning point in the outlook of botanists occurred with the appearance
of Thuret's papers of 1853 (a and b) and 1854. He established that in Fucus
only eggs to which sperms had had access would germinate. Shortly afterward,
Pringsheim (1855, 1856) observed the penetration of the sperm into the egg of
Vaucheria and Oedogonium. At first the exact function of the sperm in sexual
reproduction was not understood, but it seemed doubtful that it actually fused
with the egg. Pringsheim's observations were quickly confirmed, however, by
a number of workers. Cohn (1855, 1856) established the occurrence of sexuality
in Spkaeroplea and Volvox, De Bary (1858a) confirmed the findings of Prings-
heim in regard to Vaucheria and Oedogonium, and Pringsheim (1860) pub-
lished his observation on Coleochaete.

The majority of forms studied during this early period showed oogamous
sexual reproduction. The first report of a conjugation of motile isogametes was
by Pringsheim in 1870 in regard to Pandorina. This genus thus bridged the
gap between the oogamous types and the condition as shown by Spirogyra and
diatoms. Shortly afterward isogamous sexual reproduction was discovered in
a number of other algae and sufficient evidence was brought to bear to dispel
the old belief that conjugation and fertilization were different processes.

Hertwig in 1876, working on a species of sea urchin, showed for the first
time that a significant feature of sexual reproduction was the fusion of the
gamete nuclei. Schmitz (1879c) observed a similar fusion of nuclei for the
first time in plants in Spirogyra and Berthold (1881) next saw it in the brown
alga Ectocarpus. In the words of Mobius (1937, p. 345) :

Die Kryptogamen waren somit mehr phanerogam geworden als die Phanerogamen,
t'iir die man zwar die Differenzierung der Geschlecliter und die Notwendigkeit der Bestau-
bung erkannt hatte, die aber in Hinsicht auf den eigentliclien Vorgang der Befruchtung
noch ganz kryptogam geblieben waren.

Phylum Chlorophycophyta

Characterization: This phylum is comprised of a large and diversified assemblage of
algae, ranging from motile and nonmotile unicellular forms to massive coenocytic types,
such as certain species of Codium. The majority of species are aquatic. Some genera
(e.g., Pleurococcus, Trentepohlia. Fritschiella) are terrestrial or subaerial in occurrence.
Certain orders (Siphonocladales, Dasycladales) are wholly marine, others (Zygnematales,
Oedogoniales) are freshwater in distribution. Several orders and even some genera
(e.g., Chlaviydoiiionas. Chulophora) have both marine and freshwater representatives.

In the majority of species the cell is provided with a definite wall which is usually
composed of an inner, and often stratified, cellulosic layer and an outer pectic layer. In

PAPENFUSS: CLASSIFICATION OF THE ALGAE 119

the Siphonales the cellulose is frequently replaced by callose. In the Dasycladales and in
many Siphonales the pectic layer of the wall is impregnated with calcium carbonate. (In
some seas the lime-incrusted fronds of Halimeda form an important constituent of coral
reefs.) In the desmids and a few other forms the wall consists of two overlapping pieces.

The chloroplasts are usually well defined and ordinarily lie in the peripheral cyto-
plasm, but axile plastids are not uncommon and are especially characteristic of the des-
mids. According to the genus or species, the cells are provided with one or more plastids
of varying form and size. The pigment complex is essentially the same as in higher
plants, consisting of chlorophyll a, chlorophyll b, xanthophylls, and carotenes. The Sipho-
nales contain two xanthophylls (siphonein and siphonaxanthin) that are peculiar to
them (Strain, 1951). A few forms (e.g., Polytomella) are colorless. The customary food
reserve is starch. In many species the chloroplasts contain pyrenoids. Usually, the
pyrenoid is enclosed by a starch envelope consisting of separate plates of starch.

In the majority of green algae the vegetative cells are uninucleate. The Sphaeropleales,
Cladophorales, Siphonocladales, and many Chlorococcales have multinucleate cells, and
the nonseptate filaments of the order Siphonales are, of course, also multinucleate.

Except in the Polyblepharidaceae of the order Volvocales, the Oedogoniales, and
Derbesia (Siphonales) the motile stages are provided with two or four terminal fiagella
that are of equal length and devoid of cilia. In the Polyblepharidaceae the cells bear
two, three, four, five, or more fiagella of equal length and in the Oedogoniales and in
Derbesia the reproductive cells bear a subterminal collar of many equal fiagella.

Asexual reproduction by ordinary cell division is of common occurrence in the unicel-
lular forms. Many species reproduce by zoospores. In most instances the cells in which
the zoospores are produced are not differentiated as specialized sporangia. The zoospores
are formed singly or in numbers l)y the cell contents. Nonmotile spores (aplanospores,
akinetes) are produced in a number of genera.

Sexual reproduction has been established for a large number of genera representative
of all the orders, with the possible exception of the Schizogoniales (cf., however, Fuji-
yama, 1949). The species may be monoecious or dioecious and isogamous, anisogamous,
or oogamous. In isogamous and anisogamous forms the gametangia may or may not be
morphologically differentiated structures. Oogamous forms ordinarily produce differen-
tiated oogonia and antheridia — SphaerojUea is an exception to the rule. Except in Sphaer-
oplea, only one egg is formed in each oogonium. The egg is ordinarily fertilized in posi-
tion within the oogonium. In a few instances (CJilorangium oogamum, Chaetonema ir-
regular e) it is extruded prior to fertilization. There is clear evidence that oogamy has
evolved independently in several of the orders.

In almost all freshwater species, the zygote is a thick-walled resting cell. In marine
species, on the contrary, it is a thin-walled cell which ordinarily germinates directly. It
thus Seems reasonable to assume that those freshwater species in which the zygote is
not a resting cell are derived from marine species or conversely that the marine species
in which the zygote is a resting cell are derived from freshwater species.

A large majority of the green algae and almost all the freshwater representatives of
the phylum apparently are haploid, with meiosis occurring in the germinating zygote.
Certain representatives of the orders Volvocales, Ulotrichales, Cladophorales, and possibly
some Chlorococcales and Siphonales, show an alternation of generations. The Siphono-
cladales, the majority of the Siphonales and the Dasycladales are diploid as far as known,
with meiosis occurring at gametogenesis. The Volvocales constitute a basic group from
which the other orders apparently have evolved.

History: Recognition of the Chlorophycophyta as an autonomous group begins
with Lamouroux (1813) who, largely on the basis of color, established an "ordre"
Ulvacees to receive certain genera of green algae, Viva, Bryopsis, Caulerpa, and
Asperococcus. Asperococcus, however, was later shown to belong to the brown
algae. Harvey (1836) erected the "division" Chlorospermeae and assigned to it
not only the green algae but also the bluegreen algae and a few genera that
were later found to be red or brown algae. The currently accepted class name
Chlorophyceae was proposed by Kiitzing (1845).

120 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

By the middle of the last century, a comparatively large number of genera
of unicellular green algae — both flagellated and nonfiagellated forms — had been
described, mostly by Ehrenberg (1838 and earlier), Nageli (1849), and others.
Many of these genera, however, were regarded as animals, the gutless stomach
animalcules of Ehrenberg.

In 1819, Lyngbye described the genus Palmella and in 1824 C. Agardh de-
scribed Protococcus. Since these two genera of unicellular algae were comprised
of nonmotile forms (cf., however, Silva and Starr, 1953, regarding Protococcus)
they were accepted as algae rather than animals from the beginning. Recogniz-
ing the distinctiveness of PahneUa and Protococcus as contrasted with the gen-
era that constituted the Ulvaceae (which received mostly membranous forms)
and the Cpnfervaceae (which received mostlj^ filamentous forms), Endlicher
(1843) created for them and various other unicellular genera the suborder
Palmellae, which he placed in the "order" Confervaceae. The Palmellae were in
turn subdivided by Endlicher into the two tribes Protococcoideae (which re-
ceived Protococcus) and Coccochloreae (which included PahneUa).

Kiitzing (1833b) was the first to recognize the differences between the dia-
toms and the desmids, for the latter of which he established the family Des-
midiaceae. He (1843, 1845, 1849) placed this family in a separate group Cha-
maephyceae (dwarf algae), which he (1845) assigned to the Chlorophyeeae.
The Chamaephyceae also received, among others, the Palmellae, which Kiitzing
regarded as a family. Hassall (1845) erected a separate family Protococceae
for Protococcus and several other genera.

Since these early beginnings, the Palmellaceae and Protococcaceae have had
an extensive and involved taxonomic history. For a long time they functioned
as a catch-all for many different kinds of unicellular and colonial algae.

Siebold (1849), Nageli (1849), Braun (1851), and Colin (1852, 1854) re-
garded as algae certain unicellular and colonial motile green organisms such as
Chlaynyclomonas, Gonium (first recognized as an alga by Turpin, 1828a, pp. 322-
329), Panclorma, StepJianosphaera, and Volvox (all members of the Volvocales).
Cohn at first (1852) placed these organisms in the "order" Palmellaceae but in
1856 considered them as constituting the family Volvocaceae. Rabenhorst ( 1863,
1868) grouped the Volvocaceae along with the Palmellaceae and Protococcaceae
in a common order which he (1868) called Coccophyceae.

The relationship between the Zygnemataceae and the Desmidiaceae was recog-
nized by Nageli (1849) and others but it was De Bary (1858b) who first fur-
nished conclusive proof of this. He united these two groups in a "family"
Conjugatae.

In his important work on the freshwater algae of Europe, Rabenhorst
(1868) divided the green algae in accordance with the system of Stizenberger
(1860) into four orders: (1) Coccophyceae, with the families Palmellaceae, Pro-
tococcaceae, and Volvocaceae; (2) Zygophyceae, with the families Desmidieae and
Zygnemeae; (3) Siphophyceae, with the families Hydrogastreae (^Botrydia-
ceae) and Vaucheriaceae; and (4) Nematophyceae, with the families Ulvaceae,
Sphaeropleaceae, Confervaceae, Oedogoniaceae, Ulotrichaceae, Chroolepidiaceae
(=Trentepohliaceae), and Chaetophoraceae. The separation of the Chlorophy-
cophyta into the four orders recognized by Stizenberger (but later, for example,
by De Toni [1889] and Wille [1890-1891], usually called Protococcoideae, Con-

PAPENFUSS: CLASSIFICATION OF THE ALGAE 121

jugatae, Siphoneae, and Confcrvoideae, respectively) remained in force until
the beginning of the present century.

The classification of the unicellular and colonial green algae (other than the
desmids, whose relationshij) with the Zygnemataceae is clear) and of the siphon-
ous forms has always been fraught with many difficulties. It is the sorting
out of these algae that will be considered especially in the following pages.

In agreement with Kabenhorst, Kirchner (1878) grouped the families Volvo-
caceae, Protococcaceae, and Palmellaceae in a common order which he named
Protococcoideae. Ilansgirg's (1888a) and Wille's (1890-1891) circumscriptions
of the order Protococcoideae were much the same as that of Kirchner, except
that Hansgirg merged the Protococcaceae in the Palmellaceae whereas AVille
recognized a larger number of families.

The far-reaching contributions by Klebs (1883, 1892) and L^^ther (1899),
which resulted in the removal of the euglenids from the Protococcoideae and the
establishment of a separate class Heterokontae for those "green" algae with two
unequal flagella, are considered under the Euglenophycophyta and Chrysophy-
cophj^ta, respectively.

Blackman (1900) and Blackman and Tansley (1902) arrived at the signifi-
cant conclusion that the order Protococcoideae comprised families representa-
tive of three divergent vegetative tendencies which furnished the phylogenetic
lines on which the different types could be arranged. (1) Those forms in which
the plant body is motile by means of flagella during the vegetative phase. (2)
Those in which the plant body is not motile and in which the cells are uninu-
cleate and divide during the vegetative phase. (3) Those in which the plant
body is nonmotile and in which the cells do not undergo vegetative division but
the nucleus divides and the cells consequently become multinucleate.

As delimited in the arrangement at the end of this section, these three lines
are represented by various orders and families as follows. The first group cor-
responds to the Volvocales. The second group is represented by the suborder
Tetrasporineae of the order Volvocales and the family Pleurococcaceae of the
Ulotrichales. The third group is represented by the order Chlorococcales. (In
the breaking up of the heterogeneous order Protococcoideae [= Protococcales
Kirchner orth. mut. Engler, 1892], the ill-fated designation Protococcales has
fortunately been relegated to synonymy. Silva and Starr, 1953, have produced
convincing evidence that the type species of Protococcocus C. Agardh is actu-
ally a species of Haematococcus rather than of the plant that is commonly known
as Protococcus but which probably should be known as Pleurococcus. Further-
more, it is now known that the unicellular habit of this genus is a derived rather
than a primary condition and that it belongs in the Ulotrichales.)

The classification of the multinucleate segmented (exclusive of the Ilydro-
dictyaceae) and the siphonous Chlorophycophj-ta has also been a matter of much
confusion and disagreement. Egerod (1952) has recently given an excellent
treatment of the taxonomic history of these algae; a brief review will conse-
quently suffice here.

Greville (1830) established the "order" (family in the modern sense) Siphon-
eae to accommodate certain green algae {C odium, Bryopsis, Vaucheria, Botry-
dium) that possess a tubular, nonseptate thallus. (He placed Caiderpa in its
own "order.") In the course of time a number of additional families were erected,

122 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

especially by Kiitzing (1843, 1849), to receive various genera belonging to the
siphonous complex, among which are the Sphaeropleaceae, Cladophoraceae, Va-
loniaceae, Caulerpaceae, and Dasycladaceae (a group comprised of calcified
plants that for a long time were thought to be corals). Stizenberger (1860) ar-
ranged the algae belonging to this complex in his order Siphophyceae (except
for the Sphaeropleaceae and Cladophoraceae, which he placed in the Conferva-
ceae of his order Nematophyceae). This order was comprised of the families
Valoniaceae (in which he included Caulerpa), Vaucheriaceae (now placed in
the Xanthophyceae of the Chrysophycophyta), Codiaceae, and Dasycladaceae.

Schmitz (1879a) created the group Siphonocladaceae to receive his new genus
Siphonocladus and a number of other multinucleate septate or saccate genera
(e.g., Valojiia, Microdictyon, CludopJiora, and Botrydium, which genus is now
known to belong to the Xanthophyceae).

Starting with Bohlin's paper of 1901, the complex of siphonous algae has
been segregated into six orders. Bohlin erected the order Vaucheriales and re-
moved it to the class Heterokontae. Blackman and Tansley (1902) substituted
the ordinal name Siphonales for the designation Siphophyceae or Siphoneae and
divided the order into the two suborders Siphoneae (which received nonseptate
genera) and the Siphonocladeae (which received septate forms such as Siphono-
cladus, CladopJwra, Yalonia, SpJiaeroplea, and many other genera). Oltmanns
(1904) elevated the suborder Siphonocladeae to the rank of order (Siphonocla-
dales) and placed in it the five families Siphonocladaceae, Cladophoraceae,
Sphaeropleaceae, Valoniaceae, and Dasycladaceae.

West (1904) removed the Cladophoraceae and Sphaeropleaceae to an autono-
mous order Cladophorales. As characterized by him this order conformed closely
to Oltmanns' Siphonocladales and in 1916 West merged the Cladophorales in
the Siphonocladales. Heering (1921) and Oltmanns (1922a) not only followed
West but in conformity with Oltmanns' classification of 1904 extended the con-
cept of the Siphonocladales to include even the nonseptate Dasycladaceae. In
the intervening period, however, B0rgesen (1905, 1913) had discovered in Si-
phonocladus and related genera the peculiar method of cell division termed
segregative division by him. Despite the distinctiveness of this character dis-
agreement has persisted with respect to the autonomy of one or the other of
these two orders. Some authors (e.g., Fritsch, 1935, 1947) have accepted the
Cladophorales but place the genera comprising the Siphonocladales in the Si-
phonales, whereas others (e.g., Feldmann, 1938) have recognized the Siphonocla-
dales as a distinct order but have included in it the genera constituting the
Cladophorales. Egerod (1952) has given a careful analysis of this confusing
state of affairs and has detailed the evidence favoring recognition of all three
orders.

As has been pointed out by Fritsch (1944) and Egerod (1952) the Anadyo-
menaceae (at least as far as Anadyomene and Microdictyon are concerned) de-
part from other Siphonocladales in not exhibiting segregative division. Both
these authors have also brought attention to the correspondence between this
family and the Cladophoraceae. The Anadyomenaceae are here transferred from
the Siphonocladales to the Cladophorales.

In 1931 Pascher removed the Dasycladaceae from the Siphonocladaceae and
erected for them the order Dasycladales, a group distinguished by the formation

PAPENFUSS: CLASSIFICATION OF THE ALGAE 123

of operculate cysts and certain other features. It is now known ( Ilammerling,
1931) that the members of this order are actually uninucleate during their vege-
tative phase and become multinucleate only when they become fertile.

The status of the monogeneric family Sphacropleaceae remains to be consid-
ered. Owing to the multinucleate condition of the septate thallus, Sphaeroplea
was for a long time associated either with the Siphonocladales or the Cladopho-
rales. As has been emphasized by Fritsch (1929; 1935, pp. 224-225; 1947, pp.
43-48), however, SphacropJca possesses a number of characters that appear to
ally it with the Ulotrichales rather than the complex of siphonous orders con-
sidered above, which Egerod (1952) would derive from the Chlorococcales.
Fritsch (1935) considers Sphaeroplea as constituting a suborder in the Ulotri-
chales. Prescott (1951) has elevated this suborder to the rank of order. I con-
cur with the view of Prescott.

In addition to the orders which have so far been considered, namely, the Vol-
vocales (including the Tetrasporales), Chlorococcales, Cladophorales, Siphono-
cladales, Siphonales, Dasycladales, and Sphaeropleales, the Chlorophycophyta
have been credited in recent times with the following orders: Zygnematales
(=the order Zygophyeeae of Stizenberger), Ulotrichales, Chaetophorales, Ul-
vales, Microsporales, Cylindrocapsales, Oedongoniales, and Schizogoniales. These
orders, with the exception of the Zygnematales, constitute the bulk of the Nema-
tophyceae of Stizenberger.

The Zygnematales (Conjugales of some authors) embrace a well-marked
group of algae characterized by the conjugation of nonflagellated gametes. The
relationship of the essentially unicellular desmids with the filamentous Zygne-
mataceae (which family includes the classical Spirogyra) has been generally
recognized since the time that De Bary (1858) pointed to their alliance. The
Zygnematales occupy an isolated position in the Chlorophycophyta. At one
time the order was regarded by some authors (e.g., Engler and Gilg, 1924) as
sufficiently distinct from the other green algae to merit recognition as an inde-
pendent phylum. There seems to be little justification, however, for considering
the Zygnematales as distinct from the Chlorophycophyta, and in recent systems
of classification the assemblage has usually been treated as an order of the
Chlorophycophji;a. It is not inconceivable that the Zygnematales evolved, in the
distant past, from the Volvocales. Monographic treatments of the order or of
some of its families have in recent times been published by Czurda (1932, 1937),
Krieger (1933-1937, 1939), Kolkwitz and Krieger (1941-1944), and Transeau
(1951). The classification of the Zygnematales adopted in the arrangement at
the end of this section is essentially that of Fritsch {in West, 1927).

The Ulotrichales as here accepted include the Chaetophorales, Ulvales, Micro-
sporales, and Cylindrocapsales. The order Chaetophorales was established by
Wille (1901) to receive essentially the same algae, including Ulothrix, that Borzi
(1895) assigned to his order Ulotrichales. In current systems of classification
the Chaetophorales are either accepted as an autonomous order (e.g., by Fritsch,
1935) or they are merged in the Ulotrichales (e.g., by Smith, 1950). Fritsch
(1935; 1939; 1944, p. 245) especially, has argued for the retention of the order,
primarily because of the heterotrichous habit of the majority of the forms. How-
ever, in some genera of the Chaetophorales either the prostrate or the erect
system may be absent or poorly developed (cf. Papenfuss, 1951b). Heterotrichy

124 A CENTURY Of PROGRESS IN THE NATURAL SCIENCES

has evolved independently in a number of different groups of algae — simple
forms as well as advanced types— and it is doubtful that any great weight should
be placed on this character in the delimitation of major taxa. In the family
Aerochaetiaceae of the red algae, for instance, some species of a genus have a
single basal cell whereas others produce an extensive prostrate system.

Although the majority of recent systematists have accepted the order Ul-
vales, proposed by Blackman and Tansley (1902), there is a great deal of
justification for the view of Fritsch (1935, 1944) that they are advanced Ulo-
trichales. The possession by the Ulvales of a parenchymatous thallus does not
distinguish them from all Ulotrichales since some of the latter (e.g., Fritscliiella)
also form parenchymatous thalli. Some Ulvales show an alternation of isomorphic
generations but this is now believed to be true also of certain Ulotrichales (e.g.,
DraparnaldioiJsis and FritschieUa; ^ingh, 1945, 1947).

The order Microsporales, which was established by Bohlin (1901) to accom-
modate the genus Microspora, has never received wide acceptance. Eecently, how-
ever, it was resurrected by Prescctt (1951), who in the same work also erected
an order Cylindrocapsales for the genus C ylindrocapsa. Although these two
genera, especially the oogamous C ylindrocapsa, occupy a somewhat isolated po-
sition among the Ulotrichales it seems best to regard them in agreement with
Fritsch (1935) and Smith (1950) as constituting well-defined families within
the Ulotrichales. Printz (1927) does not even recognize the family Microspora-
ceae; he places Microspora in the Ulotrichaceae.

The Oedogoniales occupy an isolated position in the Chlorophycophyta. The
peculiar method of cell division shown by the three genera comprising the order
is not met with am^vhere else and the collar of subterminal flagella present in
the sperms and zoospores is encountered elsewhere only in Derhesia (Sipho-
nales). Monographic treatments of the order have been presented by Him
(1900), Tiffany (1930), and Gemeinhardt (1938-1940).

The Schizogoniales, established by West (1904), comprise a small group of
terrestrial, freshwater, and marine algae placed by some (e.g., Fritsch, 1935)
in one, by others (e.g., Knebel, 1935) in two. genera. The group is character-
ized by the formation of parenchymatous thalli, stellate plastids, and the ap-
parent lack of motile reproductive cells (cf., however, Fujiyama, 1949). The
order has recently been the subject of a monograph by Knebel (1935).

The more important early discoveries relating to sexuality in the green algae
have been considered briefly in the introduction to this chapter. In the present
section attention will be focused especially on some of the more recent work
on the life histories and the associated nuclear phenomena.

A fusion of the gamete nuclei in the zygote was observed by Schmitz
(1879c) in Spirogyra. Klebahn (1888, 1891, 1892) observed it in desmids and
Oedogonium, and Goroschankin (1890) saw it in ChJamydomonas.

Following the postulate of Weismann (1887) that the doubling of the chro-
matin mass at syngamy must be followed by a regulatory reducing process,
many investigations were undertaken with the view of testing this hypothesis
and of determining the place in the life history where the reduction may occur.
The first observations on meiosis in the green algae were by Allen (1905) in
Coleochaete. The life history of Coleochaete had previously been investigated
by Pringsheim (1860) who showed that the contents of the zygospore at germi-

PAPENFUSS: CLASSIFICATION OF THE ALGAE 125

nation divide into a number of cells, each oi which later gives rise to a zoospore.
Pringsheim and others regarded this structure as the sporophyte of Coleochaete.
Allen established, however, that meiosis occurs during the first two divisions of
the germinating zygote and that the cellular structure produced by the zygote
is haploid. Thus Allen eliminated the only green alga that till then was re-
garded as exhibiting an alternation of generations. Cytological studies by vari-
ous later workers on diverse green algae confirmed the observations of Allen
and until 1925 it was generally held that green algae are haploid, with meiosis
always occurring during the germination of the zygote.

Since 1925 when Miss AVilliams showed that Codiuni is a diploid alga, our
concept of the life histories of the green algae and the associated nuclear cycles
has undergone a profound change. It is now known that the Siphonocladales
(Schechner-Fries, 1934; Schussnig, 1938), Siphonales (Williams, 1925; Schuss-
nig, 1930, 1932, 1939, 1950; Zinnecker, 1935), and Dasycladales (Schulze, 1939)
are diploid algae, at least those that have been studied cytologically, and that it
is the diploid soma that in them, as in the Fucales, diatoms, and animals, func-
tions as the gamete producing generation. (See, however, the statements below
on Derhesia and Halicystis.)

The occurrence of an alternation of isomorphic generations in the green algae
was first demonstrated by Hartmann (1929) and F0yn (1929, 1934a, 1934b) in
members of the orders Cladophorales {Cludophora, Chaetomorphn) and Ulo-
trichales {Enteromorpha, Viva). Singh (1945, 1947) has reported the occur-
rence of a similar cycle in Draparnaldiopsis and FritscJiiella (both Ulotrichales),
and Iyengar and Ramanathan (1940, 1941) have established its occurrence in
Anadyomene and Microdictyon (both here placed in the Cladophorales).

Juller (1937) has shown that Stigeodonium subspinosum (Ulotrichales) pos-
sesses an alternation of heteromorphic generations, with the diploid asexual gen-
eration smaller than the haploid one. Jorde (1933) has brought forth fairly
convincing evidence indicating that certain species of the unicellular Codiolum,
a genus of the family Chloroeoccaceae in the order Chlorococcales, represent the
diploid asexual generation of species of the filamentous Urospora, a member of
the family Cladophoraceae in the Cladophorales. These observations still re-
quire corroboration, but if they should prove to be correct, we would have here
an alternation of heteromorphic generations, representing two kinds of green
algae which for a long time have been regarded as phylogenetically very far
apart.

Kornmann (1938) and Feldmann (1950) have made observations suggesting
that Halicystis and Derhesia also constitute phases in the life history of one
and the same alga, with Halicystis representing the gametophytic and Derhesia
the sporophytic generation. These two genera have been regarded as the type
representatives of two distinct families, Ilalicystidaceae and Derbesiaceae, of
the order Siphonales. In view of the fact that Derhesia is the only genus of
green algae outside the order Oedogoniales that is known to possess swarmers
with a subterminal collar of flagella arid that Halicystis forms terminall}^ bi-
flagellate gametes, the apparent existence of an intimate relationship between
these two kinds of plants is a matter of far-reaching signifleanco.

Although the majority of Chlorococcales appear to be haploid, there is some
indication that CMorocliytrium Lemnae shows an alternation of generations

126 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

(Kurssanov and Schemakhanova, 1927) and that Apiococcus consociatus is
diploid (Korshikov, 1926).

Among the more remarkable discoveries of recent times are those showing
that even the Volvocales include forms exhibiting an alternation of generations.
Thus it has been shown by Strehlow (1929) and Belilau (1935) that ChJoro-
brachis gracilUma is the diploid motile stage of Pyrohotrys gracilis. Two other
species of Pyrobotrys likewise have a motile zygote (Behlau, 1935). Behlau
(1939) has also shown that Carteria ovata is the diploid motile stage of Chlamy-
domonas variabilis.

Judged by their morphology, especially that of the motile cell, the Chloro-
phyeophyta show affinities only with the Charophycophyta. The possession by
the Euglenophycophyta of chlorophyll a and chlorophyll b suggests that this
phylum is related to other green plants. Morphologically the euglenid cell is
very different, however, from the motile cell of the green algae, and whatever
relationship there may be between these two groups probably is extremely re-
mote. Fossil Chlorophycophyta are known from the Ordovician onward.

The following synoptic outline of the Chlorophycophyta is adapted largely
from Fritsch (1935), Smith (1950), and Egerod (1952).

Phylum CHLOROPHYCOPHYTA Papenfuss (1946, p. 218)

Syn.: Chlorophyta Pascher (1914, p. 158); Glaucophyta Skuja (1948, p. 63)
Class Chloropiiyceae Kiitzing (1843, p. 118)

Syn.: Chlorospermeae Harvey (1836, p. 163); Glaucophyceae Bohlin (1901, p. 16);
Chlorophyllaceae Rabenhorst (1863, p. 117) ; Chlorophyllophyceae Rabenhorst (1868,
p. 1) ; Zygophyceae Rabenhorst (1868, pp. 1, 101) ; Isokontae Blackman et Tansley
(1902, p. 20) ; Akontae Blackman et Tansley (1902, p. 168) ; Stephanokontae Black-
man et Tansley (1902, p. 166); Prasinophyclnees Chadefaud (1950b, p. 988); Eu-
chlorophycin^es Chadefaud (1950b, p. 988); Pocillophycinees Chadefaud (1950a,
p. 788)

Order VOLVOCALES Oltmanns (1904, p. 133)

Syn.: Chlamydomonadales Fritsch, in West (1927, p. 67); Chlorodendrales
Fritsch, in West (1927, p. 67) ; Pyramidomonadales Chadefaud (1950b, p. 988) ;
Tetrasporales Lemmermann (1915, p. 21)
Suborder Volvocineae West (1916, p. 161)

Syn.: Chlamydomonadineae Fritsch (1935, p. 78)
Family Polyblepharidaceae (Blackman et Tansley) Oltmanns (1904, p. 135)
Syn.: Polytomellaceae (Blackman et Tansley) Skuja (1930, p. 158)
Family Pedinomonadaceae Korschikov (1938; not seen, cited from
Skuja, 1939b)
Family Nephroselmidaceae Pascher (1913b, p. 110)

(See Skuja, 1948, pp. 65, 66, 367)
Family Chlorovittaceae Schiller (1925b, p. 104)

Family Chlamydomonadaceae Stein orth. mut. G. M. Smith (1920, p. 90)
Syn.: Carteriaceae G. M. Smith (1920, p. 92); Sphaerellaceae
(Schmidle) West (1916, p. 166)
Family Haematococcaceae (Trevisan) Marchand orth. mut. G. M. Smith
(1950, p. 109)
Syn.: Protococcaceae (Endlicher) Hassell orth. mut. Nageli (1847, p.
153; as to type only cf. Silva and Starr, 1953)
Family Spondylomoraceae (Ehrenberg) Korschikov (1923, p. 178)
Family Astrephomenaceae Pocock (1954, p. 126)

Family Volvocaceae Ehrenberg orth. mut. Cohn (1856, p. 323, as Volvo-
cin^es)

Syn.: Pandorinaceae Luerssen orth. mut. Eichler (1880, p. 4)

PAPENFUSS: CLASSIFICATION OF THE ALGAE 127

Family Phacotaceae (Biitschli) Oltmanns (1904, p. 147)
Suborder Tetrasporineae West (1916, p. 182)

Family Palmellaceae (Endlicher) Kiitzing orth. mut. Nageli (1847, p.
123)

Family Coccomyxaceae (Cliodat) G. M. Smith (1933, pp. 350, 366)
Family Tetrasporaceae (Nageli) Klebs orth. mut. Wille, in Warming
(1884, p. 23)
Syn.: Gloeochaetaceae Bohlin (1901, p. 25), nomen nudum; Chaeto-
peltidaceae (Borzi) West ortli. mut. Wille (1909b, p. 98)
Family Chaetopediaceae Skuja (1948, p. 121)
Suborder Chlorodendrineae Fritsch (1935, p. 130)

Family Chlorodendraceae Oltmanns (1904, p. 136)
Family Clilorangiaceae Lemmermann (1915, p. 25)
Order ZYGNEMATALES Borge et Pascher (1913, p. 1, as Zygnemales)

Syn.: Conjugales (De Bary) Rabenhorst orth. mut. G. M. Smith (1920, p. 183);
Mesotaeniales Fritsch, in West (1927, p. 225); Desmidiales Krieger (1933, p.
173), nomen nudum

Suborder Zygnematineae Papenfuss, nom. nov.

Syn.: Euconjugatae (Fritsch) Fritsch (1935, p. 311)
Family Mesotaeniaceae Oltmanns (1904, pp. 52, 53)

(According to the current Code, the nomenclature of this family starts
with Ralfs, 1848.)
Family Zygnemataceae (Meneghini) Kiitzing orth. mut. Engler (1898,

p. 11)

Syn.: Spirogyraceae Palla (1894, pp. 234, 235, as Spirogyraceen)
Family Mougeotiaceae Palla (1894, pp. 234, 235, as Mougeotiaceen)
Syn.: Mesocarpaceae (De Bary) Wittrock orth. mut. Wille, in Warm-
ing (1884, p. 29) ; Temnogametaceae West et West (1897, p. 37)
Family Gonatozygaceae (West et West) Fritsch, in West (1927, pp.
239, 240)

Syn.: Archidesmidiaceae Blackman et Tansley (1902, p. 189)
Suborder Desmidineae (Fritsch) Fritsch orth. mut. Papenfuss

(According to the current Code, the nomenclature of this suborder starts
with Ralfs, 1848.)

Family Desmidiaceae Kiitzing (1833b, p. 591) ex Ralfs orth. mut. Stizen-
berger (1860, p. 27)

Syn.: Eudesmidiaceae Blackman et Tansley (1902, p. 189, p.p.)
Order ULOTRICHALES Borzi (1895, p. 348, as Ulothrichiales)

Syn.: Chaetophorales Wille (1901, p. 13); Protococcales (Meneghini) Kirchner
orth. mut. Engler (1892, p. 9, p.p.); Pleurococcales Chodat (1909, p. 149);
Chroolepoidales Chodat (1909, p. 155) ; Microsporales Bohlin (1901, pp. 19, 25);
Cylindrocapsales Prescott (1951, pp. 66, 109); Ulvales Blackman et Tansley
(1902, p. 136)

Family Ulotrichaceae Kiitzing orth. mut. Rabenhorst (1868, pp. 298, 360)

Syn.: Stichococcaceae Bohlin (1901, p. 19)
Family Microsporaceae Bohlin (1901, pp. 19, 25)
Family Cylindrocapsaceae Wille, in Warming (1884, p. 30)
Family Chaetophoraceae Harvey orth. mut. Stizenberger (1860, p. 34)
Syn.: Aphanochaetaceae Oltmanns (1904, pp. 197, 240); Herposteira-
ceae (Hasen) West (1904, p. 70); Protodermaceae Kiitzing (1849, p.
471)
Family Pleurococcaceae Klebs (1883, p. 342)
Family Chlorosphaeraceae Klebs (1883, p. 343)

Family Coleochaetaceae (Nageli) Pringsheim (1860, p. 32, as Coleochae-
teen)

Family Chaetosphaeridiaceae Blackman et Tansley (1902, p. 143)
Family Chaetosiphonaceae (Huber) Blackman et Tansley (1902, p. 142)
(To judge from the classical account of Huber, 1892, it is not improb-

^28 A CENTURY OF PROGRESS IN THE NATURAL SCIINdS

able that future work will show that this family or some of its members
actually belong in the Siphonocladales, a group with which he had
allied them.)
Family Trentepohliaceae Hansgirg (1886, p. 85)

Syn.: Chroolepidaceae Rabeuhorst (1868, pp. 287, 300, 371); Mycoidea-
ceae (van Tieghem) De Toni (1888, p. 447) ; Hansgirgiaceae (De Toni)
De Toni (1889, p. 262); Ctenocladaceae Borzi (1895, p. 353); Phyco-
peltidacees Marchand (1595, p. 14)
tFamily Microthamniaceae West (1904, p. 89)
Family Wittrockiellaceae Wille (1909a, p. 222)
Family Coelodiscaceae Jao (1941, p. 294; see also 1947, p. 255)
Family Schizomeridaceae G. M. Smith (1933, p. 452)
Family Monostromaceae Kunieda ex Suneson (1947, p. 245)
Family Ulvaceae Lamouroux orth. mut. Dumortier (1822, p. 72)
Syn.: Capsosiphonaceae Chapman (1952, p. 55)
Order SPHAEROPLEALES (Fritsch) Prescott (1951, p. 110)

Family Sphaeropleaceae Kiitzing (1849, p. 362)
Order OEDOGONIALES Blackman et Tansley ex West (1904, p. 55)

(According to the current Code, the nomenclature of this order starts with
Hirn, 1900.)

Family Oedogoniaceae (Thuret) De Bary orth. mut. Stizenberger ex Hirn
(1900, p. 1)
- Order SCHIZOGONIALES West (1904, p. 56)

Syn.: Prasiolales Fritsch, in West (1927, p. 164)

Family Prasiolaceae (Rabenhorst) Borzi orth. mut. Blackman et Tansley
(1902, p. 138)
Syn.:, Schizogoniaceae Chodat (1902, p. 341, as Schizogoniacees) ;
Blastosporaceae Jessen orth. mut. Wille (1909b, p. 73)
Order CHLOROCOCCALES Marchand orth. mut. et emend. Pascher (1915, p. 2)
Syn.: Protococcales (Meneghini) Kirchner orth. mut. Engler (1892, p. 9, p.p.)
(See Silva and Starr, 1953, on the nomenclature of Chlorococcum.)
Family Chlorococcaceae Blackman et Tansley (1902, p. 95)

Syn.: Planosporaceae West (1916, p. 209); Chlorochytriaceae (West)
Setchell et Gardner (1920, p. 146) ; Endosphaeraceae Klebs (1883, p.
344); Nautococcaceae Korshikov (1926, p. 491)
Family Chlorellaceae (Wille) Brunnthaler (1913, p. 86)

Syn.: Micractiniaceae (Brunnthaler) G. M. Smith (1950, p. 232)
Family Dictyosphaeriaceae (De Toni) West (1916, p. 190)
Family Characiaceae (Nageli) Wille, in Warming (1884, p. 23)

Syn.: Characiochloridaceae Skuja (1948, p. 99), nomen nudum
Family Characiosiphonaceae Iyengar (1936, p. 317)
Family Gomontiaceae Bornet et Flahault ex De Toni (1889, p. 389)
(This family is placed in the order Chlorococcales on the strength of
Kylin's [1935] observations. It has been pointed out to me by Dr. J.
Proskauer and Dr. R. H. Thompson [personal communications] that
Gomontia may possibly be intimately connected with such forms as
Kentrosphaera, Excenti'osphaera, and Chlorochytrium.)
Family Protosiphonaceae Blackman et Tansley (1902, p. 115)
Family Hydrodictyaceae (S. F. Gray) Dumortier orth. mut. Cohn (1880,
p. 289)

Syn.: Pediastraceae Wille, in Warming (1884, p. 23)
Family Coelastraceae (West) Wille (1909b, p. 64)
Family Botryococcaceae Wille (1909b, p. 32; see Blackburn, 1936)
Family Oocystaceae Bohlin (1901, pp. 17, 25)

Syn.: Glaucocystidaceae Bohlin (1901, p. 25), nomen nudum; Eremo-
sphaeraceae (Wille) Brunnthaler (1913, p. 86); Selenastraceae (Black-
man et Tansley) Fritsch, in West (1927, p. 127)
Family Scenedesmaceae Oltmanns (1904, p. 183)

PAPENFUSS: CLASSIFICATION OF THE ALGAE 129

Order CLADOPHORALES West (1904, p. 56)

Family Cladophoraceae (Hassell) Cohn (1880, p. 289)

Syn.: Pithophoraceae Wittrock (1877, p. 47)
Family Arnoldiellaceae Fritsch (1935, p. 246)
Family Anadyomenaceae Kiitziug orth. mut. Hauck (1884, p. 420)

Syn.: Microdictyaceae (De Toni) Setchell (1929, p. 584)
Order SIPHONOCLADALES (Blackmail et Tansley) Oltmanns (1904, p. 134)

Syn.: Valoniales Pascher (1931, p. 327), vonien nudum
Family Valoniaceae Nageli (1847, p. 154)
Family Siplionocladaceae Schmitz (1879a, p. 20)

Syn.: Apjohniaceae Setchell (1929, p. 584), nomen nudum
Family Boodleaceae (Bdrgesen) Brirgesen (1925, p. 19)
Order SIPHONALES Wille, in Warming (1884) orth. mut. Blackman et Tansley
(1902, p. 114)

Syn.: Codiales Setchell (1929, p. 584); Caulerpales Setchell (1929,

p. 584); Pascher (1931, p. 327); Feldmann (1946, p. 753), nomen

nudum; Eusiphonales Feldmann (1946, p. 753)
Family Derbesiaceae (Thuret) Kjellman (1883, p. 316)

Syn.: Halicystidaceae G. M. Smith (1930, p. 227; see also 1944, p. 69)
Family Dichotomosiphonaceae Chadefaud ex Feldmann (1946, p. 753)
Family Caulerpaceae Greville ex Kiitzing orth. mut. Cohn (1880, p. 288)
Family Bryopsidaceae Bory orth. mut. De Toni (1888, p. 449)
Family Codiaceae (Trevisan) Zanardini (1843, table opposite p. 171)

Syn.: Udoteaceae (Endlicher) J. Agardh (1887-1888, p. 12; see also

Feldmann, 1946, p. 752) ; Siphonaceae Greville orth. mut. Harvey

(1849, p. 190); Spongodiaceae Lamouroux orth. mut. De Toni (1888,

p. 449)
Order DASYCLADALES Pascher (1931, p. 328, fn. 37)

Family Dasycladaceae Kiitzing orth. mut. Stizenberger (1860, p. 32)

Syn.: Acetabulariaceae (Endlicher) Hauck (1884, p. 421)

Phylum Chakophycophyta

Characterization: Members of this isolated group of only seven living genera have
erect, whorled, equisetoid, haploid, branched thalli that grow by means of a conspicuous
dome-shaped apical cell and are attached by multicellular, branched rhizoids. By trans-
verse division, the apical cell cuts off segments proximally. Each segment divides trans-
versely into an upper nodal and a lower internodal cell. The internodal cell elongates
greatly but undergoes no further septation. The nodal cell divides by vertical and curved
walls to form a nodal tissue, certain peripheral cells of which become apical cells and
give rise to a whorl of laterals of limited growth, the "leaves." The "leaves" are likewise
differentiated into nodes and internodes and may be simple or branched. Indeterminate
branches when formed, are produced in the axils of the laterals of limited growth.

The internodal cells of the axes and of the short laterals may or may not be ensheathed
by a layer of cortical filaments that are produced by the basal nodes of the whorl of short
branches. Some grow upward and ensheath the basal half of the internodal cell next
above, others grow downward and cover the upper half of the internodal cell below. The
cortical filaments grow by means of an apical cell and are also differentiated into nodal
and internodal cells.

Young and small cells are uninucleate, whereas the large internodal cells become
multinucleate by amitosis. The cells contain many small discoid chloroplasts which lack
pyrenoids. As far as known, the pigment complex does not differ from that of other green
plants, and food is stored as starch. The wall consists of an inner cellulosic and an outer
gelatinous layer of unknown composition. In many species the thallus becomes calcified.

Vegetative multiplication is of common occurrence. Secondary protonemata may
develop from the primary rhizoid or the primary protonema or the nodes of plants that

130 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

have passed through a period of hibernation. Adventitious long branches may develop
from the nodes of hibernating plants or parts that have become detached. Frequently,
tuberlike organs of vegetative propagation are formed on the rhizoids or at the nodes of
buried parts of the long branches.

Sexual reproduction is oogamous. The plants may be monoecious or dioecious. The
oogonia and antheridia are produced at the nodes of the primary laterals of limited
growth.

The antheridia are much more complex than those of other thallophytes. The anther-
idial initial divides by two intersecting longitudinal and a median transverse wall. Each
of the octants so formed then divides by two periclinal walls. The eight peripheral cells,
known as the shield cells, constitute the antheridial wall, the middle series of cells form
what is known as the manubrium, and the innermost eight cells constitute the primary
capitulum. Through the elongation of the manubrium cells and the enlargement of the
capitulum cells the manubria become laterally separated from one another in the mature
condition. The shield cells and the manubrium cells do not undergo division during the
further development of the antheridium. Through division each primary capitulum pro-
duces six secondary capitulum cells which may or may not produce tertiary and quater-
nary capitula. The secondary capitula, but at times also those of lower and higher order,
cut off initials which give rise to branched or unbranched septate spermatogenous fila-
ments, each cell of which eventually forms a single, elongated, anteriorly biflagellate
sperm.

The oogonia are like those of thallophytes in being single-celled and at first naked
structures. During their development the cell below the oogonium produces five corti-
cating filaments that invest the oogonium. These filaments remain undivided except at
their apices, where each cuts off by transverse division one or two coronal cells. The
single living family (Characeae) is divided into two subfamilies on the basis of the
formation of one (Charoideae) or two (Nitelloideae) tiers of coronal cells. Only one
egg is formed in each oogonium. In the mature condition the cortical filaments are
spirally twisted (clockwise in all living species, counterclockwise in some fossil forms)
about the oogonium.

Meiosis occurs during germination of the zygospore. At first there is produced a
protonema, from the primary branch of which the mature plant arises as a lateral branch.

The living Characeae are essentially freshwater in occurrence. Many of the fossil
forms were marine in distribution (Peck, 1934, p. 101).

History: The first published record of the designation Chara in its present
sense apparently is that by the herbalist Daleehamps (1587, p. 1070) who gave
it as the popular name of an Equisetum-like aquatic plant used by the inhabi-
tants of Lyons to scour plates and other utensils. Vaillant in 1721 formally
erected the genus Chara. During the following one hundred and fifty years these
plants had an extremely checkered systematic history. Many of the early botan-
ists regarded them as species of Equisetum or Hipimris. Linnaeus (1753) con-
sidered Chara a genus of algae. Adanson (1763, p. 472) placed it in family 56,
Ara (aroids) of the flowering plants. Others, as for example De Candolle (1805,
p. 584), associated the genus with the Naiadaceae.

In 1815 Richard {in Bonpland and Humboldt) proposed (as a nonien nu-
dum) the family Characeae and regarded it as belonging to the angiosperms.
Many early botanists, however, such as C. Agardh (1824), who established the
genus Nitella, Wallroth (1833), Endlicher (1836), Kiitzing (1843, 1849), and
others, had no hesitation in according these plants a place among the algae. On
account of the spirally arranged ensheathing filaments of the oogonium, Wallroth
(1833) erected for them (and a number of unrelated genera) the order Gyrophy-
kea (one of four which he recognized in the algae), and this name was later used
for the group by Rabenhorst (1847).

PAPENFUSS: CLASSIFICATION OF THE ALGAE 131

For a long time a great deal of misunderstanding existed regarding the na-
ture of the reproductive organs of these plants and this more than anything
else was responsible for the differences of opinion among early botanists as re-
gards the systematic position of the group. Some authors, as for example De
Jussieu (1789), regarded the antheridium as an anther and the oogonium as a
pistil and consequently placed these plants among the phanerogams. The first
to disagree with such an interpretation of the reproductive organs were Wallroth
(1815) and Bischoff (1828) , although neither of them understood their true nature.

Vaucher (1821), Kaulfuss (1825), and Bischoff (1828) studied the germi-
nation of the zygospore and thereby threw light on one phase of the reproduc-
tion of these plants. The function of the antheridium remained doubtful until
Thuret in 1840 (see also his paper of 1851) discovered that flagellated sperms
were produced in it. Braun (1852, 1853), Pringsheim (18G3a, 1863b), Sachs
(1874) and De Bary (1875) contributed further to the knowledge of the struc-
ture of the thallus, the development of the reproductive organs, and the germi-
nation of the zygospore, and De Bary (1872) studied in detail the process of
fertilization. Oehlkers in 1916 produced convincing evidence that the thallus
is haploid, meiosis occurring at the germination of the zygospore. (The belief
of Tuttle [1926] that the plants are diploid, with meiosis occurring during the
early development of the sex organs, appears to be based upon inaccurate
observation. )

In addition to contributing a great deal to knowledge of the structure of
the thallus and the reproductive organs, Braun in a long series of publications,
especially those from 1849 onward, laid the foundation upon which the present
classification of the Charophycophyta is based. At the time of his death he had
in an advanced stage of preparation a monograph of the species of the world,
which was completed by Nordstedt (see Braun and Nordstedt, 1883). In more
recent times important contributions to the taxonomy of the group have been
made especially by Migula (1890-1897, 1925), Groves and Bullock- Webster
(1920, 1924) and Zaneveld (1940). Wood (1952) has given a list of the described
species. In addition to the family Characeae, to which are referred all the liv-
ing species as well as certain fossils. Peck (1946) recognizes three fossil fami-
lies— Clavatoraceae, Trochiliscaceae, and Sycidiaceae. Miidler (1953) divides
the fossil charophytes into three orders and a total of six families as follows :
Sycidiales, with the family Sycidiaceae ; Trochiliscales, with the family Trochilis-
caceae; Charales with the families Palaeocharaceae, Clavatoraceae, Lagynopho-
raceae, and Characeae.

Although knowledge of the Charophyceae has progressed far beyond the stage
when there was disagreement as to whether these very ancient plants were flow-
ering plants, vascular crytogams, or nonvascular crytogams, a great deal of un-
certainty still exists as to the exact phylogenetic position of the group. Cohn
(1872a, 1872b, 1880) placed them as an order, Phycobryae, in the Bryophyta,
to which phylum they were also referred by Bennett (1878, 1879), who appar-
ently was not aware of Cohn's classification, and by various other botanists of
the last century and by Hy as recently as 1913.

Current opinion is divided on whether the Charoj)hyceae belong with the green
algae or constitute an autonomous pliylum, and, if so, whether this phylum be-
longs in the algae or occupies a position higher than the thallophytes.

132 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

Harvey (1849, p. 2), De Bary (1872, p. 238), and Sachs (1875, p. 278) were
the first to consider the Charophyceae as forming a group removed from both the
thallophytes and the bryophytes. They were followed by Migiila (1890-1897),
who erected the phylum Charophyta. Migula thought that both the Bryophyta
and Charophyta probably had evolved from green algae but had developed
along different lines. Many botanists, including a large number of students of
the Charophyceae, agree with Migula in regarding the assemblage as constituting
a distinct phylum, although it is not always clear from their writings whether
they consider this phylum as belonging with the algae (thallophytes) or not.
Groves and Bullock-Webster (1920, p. 1) remark: "The Charophyta are a small
group of Cryptogams, and occupy a peculiarly isolated position, having no clear
affinity with any other plants." Oltmanns (1922a, p. 457) sa^'s that he at times
was doubtful whether he should include the group in his book on the algae.

Fritsch (1935, pp. 447, 465-466) although admitting (p. 447) that "the sex
organs, and in particular the antheridium, though quite unparalleled among the
algae, are equally unique when considered in relation to other groups of plants,"
nevertheless places the group as an order in the Chlorophyceae. Smith, who
omitted them from the first edition (1933) of his Fresh-ivater Algae of the
United States later (1938, 1950) considered them as constituting a separate
class among the green algae.

Although the Charophyceae differ from green algae in a number of features,
the most important single character which removes them from this group or,
for that matter, perhaps from all thallophytes lies in the structure of the anthe-
ridium. As a primary and integral part of its development, this organ cuts off
an outer series of sterile cells, the shield cells, which function as a protective
layer to the later produced inner fertile cells. Since the exposed nature of the
reproductive organs remains as one of the few clear-cut characters whereby thal-
lophytes may be separated from plants of a higher evolutionary level, it may
thus even be questioned, as have Migula, Oltmanns, and many others, whether
the Charophyceae should be classified as algae.

In this connection it is of interest to consider Goebel's (1930) ingenious
interpretation of the antheridium. He regards it as a compound structure con-
sisting of eight congenitally fused short branches, each composed of three cells
— an apical cell (corresponding to a shield cell) and a segment cell which has
divided into two cells, the basal of which has become a capitulum cell and the
other a manubrium cell.

On this interpretation of Goebel each cell of the spermatogenous filaments
of the compound antheridium is an antheridium, as in many algae. It should
be remembered, however, that in many plants above the level of thallophytes
(e.g., bryophytes) the sperms are also produced in individual cells and yet
the antheridia are not considered compound structures. It may be emphasized,
furthermore, that the method of initiation of the eight short branches through
longitudinal division of an initial cell does not conform to the usual method
of branch initiation in the Charophyceae. But even if Goebel's interpretation
should be correct, the fact remains that a reproductive structure is formed in'
which the fertile cells are protected by a primarily produced sterile covering.
Fritsch retains the Charophyceae in the Chlorophyceae largely because they
have green plastids, produce starch, are haploid, and the Nitelleae have a simple

PAPtNFUSS: CLASSIFICATION OF THE ALGAE 133

vegetative organization; and he suspects that, if tlie fossil record were known,
all transitions to the green algae would be found. It is quite probable that the
Charophyceae evolved from the Chlorophyceae. But this is probably true also of
other green plants above the level of the algae and, if all the intermediate types
had persisted or were known from fossils, it would be impossible to separate the
angiosperms from the green algae. The presence in the Charophyceae of green
plastids, their storage of starch, and the fact that they are haploid can hardly be
considered valid criteria for retaining them among the Chlorophyceae.

The classification below is adapted from the systems of Pia (1927), Peck
(1946), andMadler (1953).

Phylum CHAROPHYCOPHYTA Papenfuss (1946, p. 218)
Syn.: Charophyta Migula (1890, p. 60)

Class Charophyceae G. M. Smith (1938, p. 127)

Order CHARALES Llndley (1836, p. 414, as "alliance")

Syn.: Sycidiales Miidler (1952, p. 14); Trochiliscales Madler (1952, p. 14)
Family Palaeocharaceae Pia (1927, p. 90)
Family Characeae Richard ex C. Agardh (1824, p. xxvii)

Syn.: Lagynophoraceae Stache (1880, not seen, cited from Madler, 1953)
Family Clavatoraceae Pia (1927, p. 91)

Family Trochiliscaceae Karpinski orth. mut. Peck (1934, p. 104)
Family Sycidiaceae Karpinski orth. mut. Peck (1934, p. 116)

Phylum Euglenophycophyta

Characterization: This phylum comprises both green and colorless, naked, and often
spirally twisted unicellular, flagellated, or rarely palmelloid organisms with a complex
vacuolar system. In some forms (certain species of Euglena) the periplast is soft and
the individuals thus show marked metaboly; in others (Pfiacus) it is rigid and the cells
consequently do not change in shape. Depending upon the genus, the individuals usually
possess one or two or occasionally three flagella that arise from the invaginated anterior
end of the cell, the reservoir. The available information on the structure of the flagella
of members of this phylum has been summarized by Vlk (1938), Brown (1945), Pitelka
(1949), Pringsheim and Hovasse (1950), and Jahn (1951). The flagella are apparently
provided with a single row of cilia along their entire length (but see the review by Pring-
sheim and Hovasse, 1950). Certain of the colorless forms, the Peranemaceae, are equipped
for the ingestion of particulate food as contrasted with the green and saprophytic species,
which apparently are unable to ingest solid food. In the green species the pigment
complex consists of chlorophyll a, chlorophyll b, beta-carotene, and several unidentified
xanthophylls (Strain, 1951, p. 253). Typically, food is stored in the form of the polysac-
charide paramylum. No coccoid or filamentous types have become known in this phylum.
The ordinary method of reproduction is by cell division. In some forms cysts are formed,
the contents of which divide into a number of cells. Various instances of gametic union
have been reported but none of these is entirely convincing.

History: The early history of the classification of the euglenids is inextricably
linked with that of many other groups of microorganisms, or Infusoria as they
were named by Ledermueller in 1763 (according to Kent, 1880-1881, p. 14).
Hence, in reviewing the classification of the euglenids the history of the entire
complex must be taken into account and the steps traced that led finally to their
separation as an autonomous phylum.

Although a few representatives of the Euglenophycophyta had already been
described before the end of the eighteenth century, especially by 0. F. Miiller
(1786) in his Animalcula infusoria . . .(the first comprehensive work on the
Infusoria), it was Ehrenberg who in 1838 in his volume Die Infusionsthier-

134 -A CENTURY Of PROGRESS IN THE NATURAL SCIENCES

chen als vollkommene Organismen laid the foundation upon which the structure
of knowledge of the euglenids rests. Ehrenl^erg erected for them the family
Astasiaea, wliich he placed along with several other families, comprising uni-
cellular and colonial forms such as volvocines, dinoflagellates, desmids, diatoms,
and amoebae, in his "Polygastrica anentera," the gutless stomach animalcules.
Dujardin (1841) showed that the "Polygastrica anentera" were not perfect
miniature replicas of the Metazoa, pointing out among other things that the so-
called stomach of these beings was a vacuole. He proposed a system of classifi-
cation based primarily on the various means of locomotion. His third order (p.
270) comprised Infusoria ". . . pourvus d'un ou plusieurs filaments flagelli-
formes servant d'organes locomoteurs. — Sans bouche." It received a number of
flagellated forms such as monads, volvocines, euglenids, and dinoflagellates. By
uniting these organisms in a single group, Dujardin became the founder of
the assemblage for which Cohn (1853, p. 273) later proposed the name Flagellata.

Siebold (1848, 1849) extended the observations of Dujardin and in con-
formity with Schleiden and Schwann's new cell theory pointed out for the first
time that the Infusoria of Ehrenberg were single-celled beings. He abandoned
Du jar din's group since he believed that the flagellated organisms were either
plants or animals and there were no intermediates. Siebold erected a class Rhizo-
poda for the nonflagellated (amoeboid) members of this complex and placed it
along with the amended class Infusoria (which included among others the eu-
glenids) in a major group for which he adopted (with altered circumscription)
the designation Protozoa of Goldfuss (1820, p. 57). Siebold excluded from his
group Protozoa organisms that were unable to change their body form through
contraction and expansion, such as the volvocines, desmids, and diatoms, which he
regarded as plants. He was inconsistent in this, however, for he retained the
peridinians as a family of animalcules in the Infusoria.

Following Siebold's establishment of the unicellular nature of the Flagellata,
little of major importance to our knowledge of the group (as circumscribed by
Siebold) was published until the appearance of the papers by Cienkowski (1865a,
1865b, 1870), who made the first detailed observations on the life histories of
various members of the group and brought light and clarity into the chaotic state
of affairs as regards knowledge of the reproduction of these organisms.

In 1878 appeared the first part of Stein's classical work on the natural his-
tory of the Flagellata. Stein regarded the Flagellata (including the Volvoca-
ceae), in agreement with Dujardin, as belonging to the Infusoria (that is, as
animals), since they possessed flagella, nuclei, and contractile vacuoles, appar-
ently overlooking the fact that a nucleus and a contractile vacuole had previ-
ously been shown to be present in the motile reproductive cells of certain typical
algae. He gave an excellent historical review of the advances in knowledge of
these organisms up to the time of his writing and his illustrations of many of
them still rank among the best that have been produced of the forms in question.

Comprehensive treatises covering much the same field were published shortly
afterward by Kent (1880-1881) and Biitschli (1883-1887). Both these authors
also gave excellent reviews of the history of the group in the broad sense.

In proclaiming that some of Ehrenberg's Infusoria were plants rather than
animals, Siebold (1848) started a long-continuing dispute as regards the nature
of many of the flagellated microorganisms. At first botanists were not particu-

PAPENFUSS: CLASSIFICATION OF THE ALGAE 135

larly disturbed by these beings but in the course of time more and more of
them became involved in the argument. By 1850, zoologists had already con-
ceded that desmids and diatoms were plants but few of them were prepared to
relinquish the flagellates, including the Volvoeaceae. Haeckel (1866) attempted
to resolve the problem by erecting for the flagellates and various other micro-
organisms a kingdom Protista, which he considered intermediate between plants
and animals. Although the concept of a separate kingdom Protista was for a
time accepted by some biologists, it was later found untenable and has been
abandoned.

A major advance toward an understanding of the morphology and the inter-
relationships of the flagellatas, with special reference to the euglenids, was made
by Klebs in 1883. Those biologists (Carter, 1856; Bcrgmann and Leuckart, 1852,
pp. 132-133; Cienkowski, 1870, p. 426; Hofmeister, 1867, p. 29; Schmitz, 1882,
p. 13, fn. 2) who had held that the euglenids, especially the green ones, were
algae had usually related them to the Palmellaceae. Since this family encom-
passed a very heterogeneous assemblage of organisms such as Protococcus (at
times referred to a separate family Protococcaceae), members of the Chlamydo-
monadaceae and Tetrasporaceae as now understood, and a number of other
types, Klebs decided to study the structure and reproduction of various mem-
bers of the Palmellaceae in order to obtain a sound basis for their comparison
with the euglenids. He found that the euglenids differed from the Palmella-
ceae in such important points as the structure of the limiting membrane of the
cell, the structure of the anterior end of the cell, the storage products, and the
method of division of the cell. Consequently, Klebs concluded that the Palmel-
laceae were typical algae and that the Euglenaceae constituted a sharply defined
group which bore no relationship to typical algae but perhaps was related, by
way of the Peranemeae, to the ciliates in the Infusoria. This possible alliance
was sufficiently remote, however, to justify recognition of the Euglenaceae and
Peranemeae as an assemblage distinct from the ciliates. He suggested that the
old designation Flagellata be employed for this group and was of the opinion
that probably the monads should also be retained in the Flagellata. In exclud-
ing the Euglenaceae from the algae, Klebs was not particularly perturbed by
their possession of chloroplasts since he erroneously believed that these struc-
tures were comparable to the cells of Zoochlorella in Paramaecium hursaria.

Klebs (1883) also presented a systematic arrangement of the genera and
species of the family Euglenaceae, which served as the basis of later classifica-
tions of the assemblage. He divided the family into the two groups Euglenae
and Astasiae, the Euglenae receiving primarily photosynthetic forms whose cells
contained an eyespot and which went into a state of rest before dividing, and
the Astasiae receiving saprophytic forms which lacked plastids and an eyespot
and divided while in a motile state.

Klebs believed that certain other organisms, typified by the genus Peranema,
represented a second natural group in the euglenid alliance. Among other dis-
tinguishing features, the members of this group possessed an oral apparatus.

In a later monograph, Klebs (1892) maintained that the argument whether
the Flagellata were thallophj'tes or protozoa had lost significance and that it
was best to look upon them as a group intermediate between plants and ani-
mals and one from which various other microorganisms had evolved. At this

136 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

time, he divided the Flagellata into the five subgroups Protomastigina, Poly-
mastigina, Eugienoidina, Chloromonadina, and Chromomonadina. The Eu-
glenoidina he divided much as he had in 1883 except that he elevated the group
Astasiae to the rank of family and he now definitely accepted the Peranemeae
as a third family in the assemblage.

As a group of plants, the Flagellata were treated by Senn (1900) in Engler
and Prantl's Pflanzenfamilien. He divided the Flagellata into seven subgroups
as follows: Pantostomatineae, Protomastigineae, Distomatineae, Chrysomona-
dineae, Cryptomonadineae, Chloromonadineae, and Euglenineae.

In accordance with the classification of Klebs (1892), but in conformity with
botanical nomenclature, Senn divided the Euglenineae into the three families
Euglenaceae, Astasiaceae, and Peranemaceae. In 1903 {in Engler, 1903), he
treated the Flagellatae as a division and the seven groups named above as orders.
In his treatment of the euglenids in Pascher's Silsswasser-Flora . . . , Lemmer-
mann (1913) accepted the three families proposed by Klebs.

Pascher in 1931 (p. 322) formally recognized the Euglenineae as an autono-
mous phylum of plants, the Euglenophyta. Smith (1933) established the family
Colaciaceae for the genus Colacium and later (1938) created for it the order
Colaciales. According to Jahn (1951) the removal of Colacium to a group of
its own is well warranted.

The three families (Euglenaceae, Astasiaceae, Peranemaceae) that comprise
the Euglenales are largely separated on the basis of method of nutrition al-
though morphological characters (especially plastid structure, presence or ab-
sence of pyrenoids and nature of flagellar apparatus) are also utilized (cf.,
Pringsheim, 1948a; Jahn, 1951). The family Euglenaceae includes all the chloro-
phyll-containing genera and those colorless forms that appear to be derived
from green species. The Astasiaceae are saprophytic and the Peranemaceae are
holozoic. It is generally agreed by students of the group that the classification
of the Euglenales is artificial but for practical reasons the separation into three
families has been adhered to pending further knowledge of the complex. The
autonomy of Astasia and certain other colorless (saprophytic) forms is espe-
cially doubtful (cf., Pringsheim, 1948b, 1952).

The order Colaciales embraces the single chlorophyll-containing genus Cola-
cium. The individuals are nonmotile in the vegetative phase and are surrounded
by a gelatinous sheath affixed to components of the freshwater zooplankton.
Usually the individuals produced by division secrete a stalk of their own and
these stalks remain attached to the stalk of the parent cell. As a result of re-
peated cell division there is thus formed a dendroid colony with the cells at
the terminations of the dichotomously branched stalk system.

As a group, the Euglenophycophyta constitute a highly specialized and seem-
ingly isolated assemblage with no clear alliance to other flagellated organisms.
The literature on the phylum may be traced through the bibliographies of
Fritsch (1935), Jahn (1946, 1951), Pringsheim (1948a), and Pringsheim and
Hovasse (1950).

A synoptic arrangement of the orders and families follows.

Phylum EUGLENOPHYCOPHYTA Papenfuss (1946, p. 218)
Syn.: Euglenophyta Pascher (1931, p. 322)

Class EuGLENOPHYCEAE G. M. Smith (1933, pp. 4, 607)

PAPENFUSS: CLASSIFICATION OF THE ALGAE 137

Order EUGLENALES Engler (1898, p. 7, as "Reihe")

Syn.: Euglenocapsales Pascher (1931, p. 326), 7wmc7i nudum, based on Eugleno-
cajisa Steinecke (1932); Euglenomonadales Chadefaud (1950a, p. 789)
Family Euglenaceae Stein ortli. mut. Klebs (1883, p. 296)

Syn.: Eutreptiidae Hollande (1942, p. 168); Distigmidae Hollande
(1942, p. 168) ; Euglenocapsaceae Pascher (1931, p. 326), nomen nudum
Family Astasiaceae Ehrenberg orth. mut. Senn (1900, pp. 174, 177)
Family Peranemaceae Klebs orth. mut. Senn (1900, pp. 174, 178)

Syn.: Menoidiidae Hollande (1942, p. 168)
Family Rhynchopodaceae Skuja (1948, p. 233)
Family Rhizaspidaceae Skuja (1948, p. 235)
Order COLACIALES G. M. Smith (1938, p. 148)

Family Colaciaceae G. M. Smith (1933, p. 612)

Phylum Chrysophycophyta

This phylum (as Chrysophyta) was established by Pascher in 1914 (cf. also
Pascher, 1921) to encompass the three classes Xanthophyceae, Chrysophyceae,
and Bacillariophyceae. Although the Bacillariophyceae appear to be only re-
motely allied to either the Xanthophyceae or the Chrysophyceae, a good deal
of evidence is at hand that points to a close relationship between the latter two
classes (Pascher, 1911, 1921, 1932, 1937).

The more important features of correspondence between the three classes
as stressed by Pascher (1914, 1921, 1924, 1937, pp. 155-173) and other authors
are: (1) storage of leucosin or oil as food reserves in members of all three
classes; (2) formation of a distinctive type of endoplasmatic spore (cyst) with
a usually silicified wall of two pieces; (3) possession by the vegetative cells of
some Xanthophyceae {OpJiiocytium, Tribonema) of a wall of two pieces com-
parable to that of the diatom frustule and that of the cysts of all three classes;
(4) growth in length of the. cell wall by the deposition of thimblelike segments
or intercalary bands in certain members of all three classes.

Although the pigmented members of these three groups usually have yellow-
green or golden-brown chromatophores and it has consequently been assumed
that they possess similar pigment complexes, it is now known that there are some
significant differences (Strain, 1951, p. 253). The three classes are considered
separately below.

CLASS XANTHOPHYCEAE

Characterization: This class is comprised of forms which in the vegetative condition
are: (1) unicellular, naked, and terminally biflagellate, excepting Nephrocliloris which
appears to be uniflagellate; (2) unicellular and amoeboid; (3) unicellular, nonmotile,
and in the form of gelatinous aggregates of various shapes and sizes (palmelloid types) ;
(4) unicellular, nonmotile, provided with a firm cell wall, and usually attached by a short
mucilaginous stalk (coccoid types); (5) simple or branched septate filaments which may
or may not be attached; or (6) vesicles or nonseptate branched filaments.

In the flagellated species and in the zoospores of the nonflagellated members of the
class, the two flagella are of unequal length and the longer flagellum is beset with cilia.
The majority of the species are pigmented, being yellow-green owing to a preponderance
of carotenoid pigments. Depending upon the species, the cells have one to many plastids
which are usually of a discoid shape. The pigment complex consists of chlorophyll a,
chlorophyll e (in Tribonema) , beta-carotene, and xanthophyll. Pyrenoids are only rarely
present and are of the naked type. Reserve food is stored as oil or leucosin. Rarely cer-

138 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

tain species of a genus are colorless. These forms, and also some of the pigmented
species, ingest solid food or are saprophytic.

The majority of the species are uninucleate; a few are multinucleate. "Where present,
the cell wall is composed of pectic compounds, and it frequently consists of two equal or
unequal overlapping pieces.

Multiplication is by cell division, akinetes, aplanospores, or zoospores. Cysts with
a silicified wall composed of two pieces have been observed in a number of species (cf.
Pascher, 1937, pp. 71-78). Sexual reproduction has been observed with certainty (cf.
Pascher, 1937, pp. 150-154) only in Trilionema (Scherffel, 1901, p. 149), Botrydium (Rosen-
berg, 1930; Moewus, 1940) and YaucUeria. Vaucheria is oogamous, Trihonema is isogamous,
and Botrydium is isogamous or anisogamous (Moewus, 1940).

History: Despite the fact that at least one member of this class has been known
since the time of Linnaeus (1753), who described Ulva granulata {^^^ Botry-
dium granul-atum) , the bulk of our knowledge of the group has been acquired
in the course of the present century, mainly through the efforts of Pascher. The
distinguishing characteristics of the class remained unrecognized until the lat-
ter part of the past century. Owing to their greenish color, the few species
which had become known previous to 1899 were regarded as Chlorophyceae.

Alexander Braun in 1855 (p. 49) recognized certain features of correspond-
ence between Opliiocytiiim, Sciadium, and Tribonema; but it was Borzi who in
1889 (see also 1895, p. 199) first convincingly pointed to the alliance of sev-
eral genera which had been placed in widely separated families of the Chloro-
phyceae. He erected an order Confervales for these algae and credited it with
the three families Sciadiaceae, Confervaceae, and Botrydiaceae. In his work
of 1895 Borzi considered the Confervales as comprising nine genera, all of which
are still regarded as belonging to the Xanthophyceae. These forms were brought
together by Borzi mainly on the basis of three characters: (1) they possessed
discoid, yellowish-green plastids; (2) they did not store starch; and (3) their
zoospores had only one flagellum ( as he believed ) .

Some j^ears later, Bohlin (1897a) published a significant study of certain
cytological characters of members of the Confervales, pointing out that the two
overlapping pieces that form the lateral wall of the cells have a layered structure,
that the wall is not composed of cellulose but of a pectic acid derivative, that
the plastids contain a preponderance of yellow pigments and that the storage
product is not starch (as Borzi had already established) but a fatty substance.

In this paper Bohlin also described a remarkable amoeboid flagellate {ChJor-
amoeha) which recalled the zoospores of Conferva sensu Lagerheim (^ Tribo-
nema). He regarded it as the progenitor of the genera comprising the Confer-
vales. In a footnote (p. 48) Bohlin remarked that Chloramoeha at times possessed
two flagella — one much shorter than the other. In a later paper Bohlin (1897b)
described Chloramoeha in some detail. It was found that if the cell lay in a cer-
tain position, two flagella — one much shorter than the other — could be observed
in the majority of instances, that the cells possess two to six discoid plastids of
a yellow-green color, and that the assimilatory product is stored as oil.

Two years later, Luther (1899) described another remarkable genus {Chloro-
saccus) belonging to this complex. This genus was palmelloid in habit, its zoo-
spores were provided with two flagella of unequal length, and the cells contained
several yellow-green, discoid plastids. In connection with his work on Chloro-
saccus Luther also investigated the zoospores of Trihonema and Botrydiopsis

PAPENFUSS: CLASSIFICATION OF THE ALGAE 139

and made the important discovery tliat the zoospores of these genera likewise
possessed two unequal flagella instead of one flagcllum as had been believed.
This character was thus found to exist in several genera of the Confervales
showing various levels of thallus specialization — flagellated, palmelloid, coccoid,
and filamentous types.

In evaluating the phylogenetic implications of the facts contributed by Borzi,
Bohlin, and himself, Luther arrived at the far-reaching conclusion that the
characters whereby these algae differed from the Chlorophyceae were of such
magnitude that it was no longer possible to retain them in this alliance. Con-
sequently, he erected for them a separate class which he named Heterokontae.
Luther's class, interestingly enough, corresponded very closely to Borzi's Con-
fervales, except that he included in it the newly erected order Chloromonadales
(as exemplified by Vacuolaria), as well as the genera Chloramoeha and Chloro-
saccus. The Chloromonadina, as typified ]:>y Vacuolaria, had previously been
established as an autonomous group of flagellates by Klebs (1892). It is now
known that the Chloromonadales w^ere misplaced in the Heterokontae.

The views of Luther as regards the autonomy of the Heterokontae were
quickly adopted by a number of students of the algae, including especially
Blackman (1900), Bohlin (1901), Blackman and Tansley (1902), Oltmanns
(1904), West (1904), and Heering (1906). Heering gave a comprehensive treat-
ment of the forms represented in the flora of Schleswig-IIolstein and a full his-
torical review of the class. He also pointed to (as Blackman, 1900, p. 671, had
previously done) the striking parallelisms in thallus types between the Heter-
kontae and the Chlorophyceae.

Blackman (1900, p. 674) brought attention to the fact that Vaucheria a;p-
peared to be the only "green" alga outside the Heterokontae which had chloro-
phyll possessing the same characters as in members of the Heterokontae and
wondered what the phylogenetic significance of this would prove to be. A year
later, Bohlin (1901) removed the Vaucheriaceae to the Heterokontae and estab-
lished for the family the order Vaucheriales. The transfer was made on the
basis of the same pigment reaction he had obtained in Tribonema, the presence
of discoid plastids, the storage of food as oil, and the observation by Walz (1866-
1867, p. 134, pi. 12, fig. 4) that the sperms had two unequal flagella.

Blackman and Tansley (1902) followed Bohlin in the inclusion of Vaucheria
in the Heterokontae and, what is important in the light of Mangenot's (1948)
recent corroborative conclusion, they also removed the Phyllosiphonaceae from
the Chlorophyceae to the Heterokontae, presumably on account of the storage
of oil as a food reserve in this family.

Following the pioneering studies of Borzi, Bohlin, and Luther, numerous
workers, but more especially Pascher, have contributed materially to our knowl-
edge of the Xanthophyceae. In 1912 (b) Pascher elaborated upon the earlier
systems of classification of the group and in accordance with the morphology
of the thallus established orders which paralleled certain chlorophycean orders.
He erected the order Heterochloridales to receive the flagellated members, the
Heterocapsales for the palmelloid types, the Heterococcales for the coccoid genera,
the Heterotrichales for the filamentous forms, and the Ileterosiphonales for the
siphonous representatives. As mentioned above, Pascher in 1914 and 1921 brought
the Xanthophyceae in alliance with the Chrysophyceae and Bacillariophyceae.

140 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

In 1925 (a) Pascher gave a treatment of the class in his Siisswasserflora
Beutschlands. . . At this time he established the order Rhizochloridales to receive
the amoeboid forms. More recently Pascher (1937-1939) has produced, as a
volume in the second edition of Rabenhorst's Kryptogamen-Flora von Deutsch-
land . . . , a monumental work of 1092 pages on the morphology and taxonomy of
the known Xanthophyceae of the world. In this work Pascher recognized some
89 genera of which he alone authored 60.

In 1930 Allorge proposed the designation Xanthophyceae as a substitute for
Heterokontae, and since this appellation conforms to the majority of class names
of algae in connoting color and in terminating in -phyceae, it has met with fa-
vor in many quarters.

Significant evidence supporting Pascher's (1914, 1921) conclusions of a re-
lationship between Xanthophyceae and Chrysophyceae was furnished in 1931
and 1938 by Vlk who established that the biflagellate motile cells of Xantho-
phyceae agreed with those of Chrysophyceae in that the long flagellum is of
the tinsel type, being beset with two rows of delicate cilia, whereas the short
flagellum lacks cilia.

Further facts favoring this alliance were brought to the foreground by Pascher
in 1932. He pointed out that the bivalved endogenously produced cysts which
he had discovered in certain Xanthophyceae in 1930 (1930a, p. 406, fig. 3c; 1930c,
pp. 332-335, fig. 17; see also Pascher, 1937, pp. 71-78, figs. 56-63) were similar
to the bivalved cysts characteristic of the Chrysophyceae.

Of especial interest is the abundant evidence brought forth in recent years
indicating that the classical Yaucheria actually belongs in the Xanthophyceae
rather than in the Chlorophyceae (Seybold, Egle, and Hiilsbruch, 1941; Chade-
faud, 1945; Strain, 1948; Koch, 1951). It will be recalled that Bohlin (1901) and
Blackman and Tansley (1902) had placed Vaucheria in the Xanthophyceae. In
general, however, phycologists have preferred to retain the genus in the order
Siphonales of the green algae. Egerod (1952, p. 336) has assembled the facts
in support of the inclusion of the Vaucheriales in the Xanthophyceae, the most
important of which are: (1) the unequal length of the flagella of the sperm
(Pringsheim, 1855, p. 142; AValz, 1866-1867, p. 134, pi. 12, fig. 4; Woronin,
1869, p. 156; Strasburger, 1887, p. 396; Koch, 1951); (2) the ciliated condition
of the shorter flagellum of the sperm (Koch, 1951) ; and (3) a pigment complex
comparable to that of Xanthophyceae (Seybold, Egle, and Hiilsbruch, 1941;
Strain, 1948). It is to be noted, however, that Vaucheria is reported as possessing
only chlorophyll a whereas Trihonema, the only other member of the class whose
green pigment has been analyzed (Strain, Manning and Hardin, in Strain, 1951,
p. 247 and table 1), possesses chlorophyll a and e.

In 1948 Mangenot produced evidence for the removal of PhyUosipJwn from
the Chlorophyceae to the Xanthophyceae, where Blackman and Tansley (1902)
had once accorded it a position.

The following classification of the Xanthophyceae is largely based on that
of Pascher (1937-1939).

Class Xanthophyceae Allorge (1930, p. 230)
Syn.: Heterokontae Luther (1899, p. 17)

Order HETEROCHLORIDALES Pascher (1912b, p. 10)

Syn.: Series Chloramoebales Fritsch, in West (1927, pp. 300, 301); Xantho-
monadales Chadefaud (1950a, p. 790)

PAPENFUSS: CLASSIFICATION OF THE ALGAE 141

Family Chloramoebaceae Luther (1899, p. 19)
Syn.: Heterochloridaceae Pascher (1925a, p. 22)
Order RHIZOCHLORIDALES Pascher (1925a, p. 26)

Family Rhizochloridaceae Pascher (1925a, p. 26)

Family Stipitococcaceae Pascher ex G. M. Smith (1933, p. 144)

Family Chlorarachniaceae Pascher (1937, p. 251)

Family Chlamydomyxaceae Hieronymus, in Engler (1897, p. 570; cf.

Hieronymus, 1905, p. 156)

Syn.: ?Myxochloridaceae Pascher (1937, p. 256)
Order HETEROCAPSALES Pascher (1912b, p. 13)

Family Chlorosaccaceae Bohliii ex Blackman et Tansley (1902, p. 217)

Syn.: Heterocapsaceae Pascher (1912b, pp. 13, 21)
Family Malleodendraceae Pascher (1937, p. 301)
Order HETEROCOCCALES Pascher (1912b, p. 14)

Syn.: Mischococcales Fritsch, in West (1927, pp. 300, 302) ; Xanthococcales
Chadefaud (1950a, p. 790)

Family Pleurochloridaceae Pascher (1937, p. 333)

Syn.: ?Halosphaeraceae Oltmanns (1904, p. 181); cf. Pascher (1925a,
p. 41; 1939, p. 910)
Family Chlorobotrydaceae Pascher (1925a, p. 48)
Syn.: Gloeobotrydaceae Pascher (1938, p. 632)
Family Botryochloridaceae Pascher (1938, p. 661)
Family Gloeopediaceae Pascher (1938, p. 696)
Family Mischococcaceae Pascher (1912b, p. 14)
Family Characiopsidaceae Pascher- (1938, p. 718)

(Pascher [1938, pp. 718, 800-812] includes Harpochytrium in the Char-
aciopsidaceae. Wille [1900, p. 371] had proposed, as a nomen nudum,
the family Harpochytriaceae for this genus. According to Jane [1946]
Harpochytrium, as to type, may have to be removed to the fungi.)
Family Chloropediaceae Pascher (1938, p. 812)
Family Trypanochloridaceae Geitler (1935, p. 146)
Family Centritractaceae Pascher (1938, p. 830)
Family Sciadiaceae Borzi (1889, p. 68)

Syn.: Chlorotheciaceae Bohlin (1897a, p. 48); Ophiocytiaceae Wille
(1909, p. 49)
Order HETEROTRICHALES Pascher (1912b, p. 18)

Syn.: Tribonematales Pascher (1939, p. 915); Confervales Borzi (1889, p. 68),
not including Conferva L. (cf. Silva, 1952, p. 271); Xanthotrichales Chadefaud
(1950a, p. 790)

Family Heterotrichaceae Pascher (1939, p. 916)

Family Tribonemataceae West orth. mut. G. M. Smith (1933, p. 157)
Syn.: Confervaceae sensu Borzi (1889, p. 69); non Confervaceae
(S. F. Gray) Dumortier (1822, pp. 71, 96)
Order HETEROCLONIALES Pascher (1939, p. 991)

Family Heterodendraceae Pascher (1937, p. 992)
Family Monociliaceae West (1916, p. 414)

Syn.: Heteroclonlaceae Pascher (1931, p. 324)
Order VAUCHERIALES Bohlin (1901, p. 14)

Syn.: Heterosiphonales Pascher (1912b, p. 21); Botrydiales Pascher (1939,
p. 1023) ; Xanthosiphonales Chadefaud (1950a, p. 790)
Family Botrydiaceae Rabenhorst (1863, p. 219)

Syn.: Hydrogastraceae (Endl.) Rabenhorst orth. mut. De Toni (1889,
p. 527)
Family Phyllosiphonaceae Frank orth. mut. De Toni (1888, p. 449)
Family Vaucheriaceae (S. F. Gray) Dumortier (1822, p. 71)

CLASS CHRYSOPHYCE.VE
Characterization : This class embraces forms which are attached or free-floating,

142 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES

unicellular, colonial, or filamentous. The unicellular species may be naked or provided
with a wall — usually of unknown composition but in some instances known to be composed
of pectin and rarely also containing cellulose — or the naked cell may be enclosed in a
capsule (lorica) which is open at one end. In many, if not all, the Mallomonadaceae and in
Aurosphaera (Chrysosphaerales) siliceous scales are embedded in the pectic wall and
the scales may bear delicate, hinged silicified needles. In the Coccolithophorineae and in
Achrosphaera (Chrysosphaerales) the pectic wall contains discoid bodies of calcium
carbonate (coccoliths) which in some instances are provided with spinelike processes.
In the Silicoflagellatophycidae the naked cell contains an internal skeleton consisting of
a framework of variously arranged siliceous rods.

The unicellular forms are either flagellated, or are consistently rhizopodial, or occur
as gelatinous aggregations of cells (palmelloid types) or as nonmotile cells enclosed by
a wall (coccoid types). Depending on the species, the flagellated cells have one, two equal
(isokont), two unequal (heterokont), or one short and two long flagella. As far as known
(Petersen, 1918, 1929; Vlk, 1938) the flagellum of the uniflagellate Chrysomonadales is
of the tinsel type whereas in the isokont Isochrysidales one of the flagella is of the tinsel
type and in the heterokont Ochromonadales the long flagellum is of the tinsel type. The
structure of the flagella in the triflagellate Prymnesiales has not yet been determined.

The filamentous forms are simple or branched and have a firm cell wall which, at least
in Phaeothavuiion, is known to be composed of cellulose.

The majority of the Chrysophyceae are photosynthetic. Some are colorless and are
either saprophytic or engulf solid food. Some of the pigmented forms also ingest solid
food. Food is stored as leucosin, a substance of unknown chemical composition (prob-
ably a carbohydrate), and oil. The cells usually contain only one or two chromatophores
which are parietal in position, and in some instances naked pyrenoidlike bodies are
present. The pigmented species have a golden-brown color owing to a preponderance of
carotenes and xanthophylls. As far as known the pigment complex consists of chlorophyll
a. beta-carotene, lutein, and fucoxanthin (Strain, 1951, p. 253).

Contractile vacuoles are of common occurrence either in the vegetative stages or in
the reproductive cells of species representative of all the orders.

The ordinary method of reproduction is by vegetative cell division. Some species
also produce zoospores. Sexual reproduction appears to be of extremely rare occurrence
and is isogamous. Up to the present a union of gametes has been observed with certainty
only in Ochrosjihaera (Schwarz, 1932) and Dinobryon Borgei (Skuja, 1950). The report
by Schiller (1926) of a fusion of gametes in Dinohryon sertularia is not entirely convinc-
ing and the observations by Mack (1951) with respect to Chrysolykos require confirma-
tion. Many of the species are known to produce cysts.

The cysts constitute one of the most distinctive features of the class. They were first
observed by Cienkowski (1865b) and have since been studied in a large number of species
by Scherffel (1911, 1924), Conrad (1927, 1928), Doflein (1923), Pascher (1924, 1932) and
others. These resting stages are formed endoplasmatically and have a wall consisting of
two pieces which are usually of a different size. The larger piece is formed first and is
composed of cellulose which is impregnated with silica; and the outer surface is often
elaborately sculptured. The smaller piece is ordinarily in the form of a plug which seals
from the inside the terminal opening left in the larger piece. The plug usually contains
little or no silica and is dissolved at germination of the cyst or is separated from the wall
around the pore. These cysts contain almost all the original protoplasm of the cell and
leucosin, and at germination the contents ordinarily divide to form a number of motile
cells which escape through the pore.

History: Hydrurus foetidus (Villars) Trevisaii is the first member of this chiss
to have been described with sufficient accuracy to be recognized by later investi-
gators. It was described by Villars in 1789 as Conferva foetida. C. Agardh in
1824 (p. xviii) erected the genus Hydrurus. It was not until the latter part of
the nineteenth century, however, that the relationship of Hydrurus with the
Chrysophyceae was established by Klebs (1892, pp. 283-285, 420-427) and

PAPENFUSS: CLASSIFICATION OF THE ALGAE 143

others. Because of its brown color, the genus had for a long time been classified
with the Phaeophyceae (cf. Ilansgirg, 1886; De Toni, 1895).

Long before Hydrunis had been recognized as a member of the Chrysophy-
ceae, a number of other genera of the class had become well known as animals.
The first of these (e.g., Syncrypta, Synura, Uroglena, Dinohryon) were de-
scribed by Ehrenberg who established the family Dinobryina for Dinohryon
and Epipyxis (cf. Ehrenberg, 1838).

Stein (1878) not only added to knowledge of the genera of Ehrenberg but
described and illustrated several new genera belonging to this complex, includ-
ing the genus Chrysomonas {= ChromuUna Cienkowski, 1870). Stein was the
first to recognize the inten'elationship of the majority of the forms known at
the time of his writing. He placed the genera in the two families Dinobryina
(Dinohryon, Epipyxis) and Chrysomonadina (p. 152), to the latter of which
he referred (p. x) ten genera, eight of which are still regarded as representa-
tive of the Chrysophyceae.

Biitschli (1883-1887) appears to have had little appreciation of the signifi-
cance of Stein's classification for he placed the genera in a number of widely
separated families of flagellates. Only in his assigning of Monas, Dinohryon,
Epipyxis, and Uroglena to a family Heteromonadina, characterized by flagella
of unequal length, did he attain a natural grouping.

The greatest advance during this early period in the delimitation of the
group as a natural assemblage was made by Klebs (1892, pp. 394-427). He re-
garded the genera known in his time (including Dinohryon and Epipyxis) as
constituting a single family Chrysomonadina in his newly established group
Chromomonadina (which also included as a second family the Cryptomona-
dina). Klebs remarked (p. 278) that one could refer to the Chromomonadina
as chrysophytes, a designation which was later formally adopted by Pascher
(1914) as the phyletic name for the chrysomonads, heterokonts, and diatoms.

Klebs clearly recognized the salient features w^hieh characterized the group :
(1) the golden-brown color of the organisms; (2) the characteristic storage
products leucosin (named by him, 1892, p. 395) and oil in both the pigmented
and the colorless members; (3) the three types of flagellation — one, two un-
equal flagella, or tw^o more or less equal ones; and (4) the formation of endo-
plasmatic cysts of a unique type such as had been observed in a number of
forms since they were first seen by Cienkowski (1865b).

Although various authors (e.g., Schmitz, 1882; Rostafinski, 1882; Hansgirg,
1886; De Toni, 1895) before the turn of the century had regarded some of
the Chrysophyceae as algae (usually as Phaeophyceae), general acceptance of
them as a group of plants begins with the works of Engler (1898) and Senn
(1900).

In agreement with the classification of Engler, Senn divided the chrysomo-
nads according to the number and length of the flagella into three families:
Chromulinaceae (with one flagellum), Hymenomonadaceae (with two equal or
more or less equal flagella), and Ochromonadaceae (with two unequal flagella).

A very significant advance in the classification of the chrysomonads was
made by Pascher in 1910. He elevated the three groups (families) recognized
by Engler and Senn to the rank of order (Chromulinales, Isochrysidales, Ochro-
monadales) and segregated the genera into seven families. (At this time Pascher

144 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

also erected an order Phaeochrysidales which included organisms with two later-
ally inserted flagella; this group was subsequently shown to belong to the
Cryptophyceae. )

In various later contributions Pascher (1912a, 1913a, 1914, 1925b, 1931)
elaborated upon his classification of this group. In addition to his three original
orders, he established among others the orders Rhizochrysidales, Chrysocapsales,
Chrj^'sosphaerales, and Chrysotrichales to receive the amoeboid, palmelloid, coc-
coid, and filamentous types, respectively. As is true of the Xanthophyceae, the
bulk of our knowledge of the Chrysophyceae has been acquired during the past
forty years, mostly through the investigations of Pascher. At the time of his
death in 1945 he was engaged with a monograph on the group, which was to have
appeared as a volume in Rabenhorst's Kryptogamen-Flora. . . Through his death
phycology has lost its foremost student of the Chrysophyceae and the present
gap in organized knowledge of this group of algae may remain unfilled for a
long time. For an autobiography and bibliography of Pascher, see Pascher
(1953).

In addition to Pascher, various authors (Lohmann, 1902; Scherffel, 1911,
1924, 1927; Petersen, 1918, 1929; Doflein, 1922, 1923; Schiller, 1925a, 1925b,
1930; Conrad, 1914, 1926, 1927, 1928, 1933; Kamptner, 1928; Gemeinhardt, 1930;
Vlk, 1938; Huber-Pestalozzi, 1941; and others) have made significant contribu-
tions to knowledge of the Chrysophyceae during the present century. Petersen
(1918, 1929) and Vlk (1938) have investigated the structure of the flagella.
Scherffel (1911, 1924), Korshikoff (1929), Pascher (1916a, 1917, 1930b) and
others have brought to light abundant evidence pointing to a relationship be-
tween various colorless flagellates and certain pigmented Chrysophyceae. Huber-
Pestalozzi (1941) has contributed a great deal to knowledge of the freshwater
planktonic forms but his work is of less value than it might have been because
of the omission of a bibliography.

Brief mention should be made of the main steps in the growth of knowledge
concerning the Coccolithophorineae and the Silicoflagellatophycidae which are
now generally regarded as Chrysophyceae but have an interesting history of
their own.

COCCOLITHOPHORINEAE

The history of our knowledge of these organisms begins with Ehrenberg
(1836, 1839) who discovered in cretaceous deposits large numbers of circular
and elliptic carbonate disks, wliich he believed had an inorganic origin.

New information as to the origin of these bodies was not forthcoming until
the survey work in the North Atlantic preparatory to the laying of the first
cable between Europe and America. Huxley and "VVallich found in the ooze
brought up from the sea bottom many carbonate bodies that resembled the disks
of Ehrenberg. Huxley (1858), like Ehrenberg, believed the disks had an inor-
ganic origin, and because of their resemblance to Protococcus cells he named
them coccoliths.

In addition to many coccoliths, Wallich (1860a, 1861) also found in the
ooze spherical bodies to whose surface adhered such coccoliths. He regarded
the spherical bodies as cells of living organisms and the chalk disks as part of

PAPENFUSS: CLASSIFICATION OF THE ALGAE 145

the skeleton of the cells. The isolated eoccoliths occurring in the ooze repre-
sented, in his opinion, the remains of disintegrated cells. Wallich called the
cells coccospheres and thought they were developmental stages of Foraminifera.

A few years later, Wallich (1865, p. 81, fn.; 1869) announced that he had
obtained living coccospheres in surface waters of the sea. But it was not until
1877 that he proposed a generic name {Coccosphaera) for his coccospheres and
credited the genus with two species.

"Wallich and various early authors believed that these organisms Avere color-
less. J. Murray (1891, p. 257) and Haeckel (1894, p. 110) considered them
algae, although they had no adequate foundation for their belief. G. Murray
and Blackman (1898) observed that the coccospheres contained a yellow-green
pigment and thus furnished the first proof of their algal nature. They believed
that the cells possessed a single chromatophore, but it was later shown by Loh-
mann (1902) and others that two plastids were present.

On the basis of a study of living material from the Mediterranean, Loh-
mann (1902) gave the first monographic treatment of the group, together with
an account of the history of the complex up to the time of his writing. He was
the first to observe that the cells were provided with one (as he believed) or two
equal flagella. (Schiller, 1925a, p. -42, later found that all the flagellated species
possess two equal flagella.)

Lohmann (1902, p. 125) concluded that the Coccolithophorineae shared
more characters with the chrysomonads than with any of the other large groups
of flagellates, and he had little hesitation in placing them in this group. Since
the name Coccosphaera, proposed for the flrst genus by Wallich, was preempted
by Coccosphaera Perty, Lohmann (p. 93) substituted the very appropriate ge-
neric name Coccolithophora and erected the family Coccolithophoridae, by which
designation the group as a whole has since been known.

Although Lohmann was aware of the long known freshwater genus Hymeno-
monas Stein (1878), which also forms calcium carbonate plates on the cell sur-
face, he failed to recognize it as a member of the Coccolithophorineae. The
relationship between this genus and the marine representatives of the group
was first pointed out by Conrad (1914).

The majority of more recent students of the Coccolithophorineae (e.g.,
Conrad, 1926; Kamptner, 1928; Schiller, 1930; Huber-Pestalozzi, 1941) have
regarded the group as belonging to the Chrysophyceae, although Schiller (1930,
p. 147), in agreement with Schussnig (1925), considers them sufficiently dis-
tinct from other Chrysophyceae to warrant placing them in a separate subclass.
In agreement with Conrad (1926), Fritsch (1935) and Iluber-Pestalozzi (1941),
the group is here considered as representative of the order Isochrysidales, which
is comprised of motile unicellular forms with two equal flagella. It should be
pointed out, however, that Schiller (1926) has shown that a few genera ap-
parently lack flagella.

The most comprehensive monograph of the group is that bj^ Schiller (1930)
which appeared as part of a volume in Rabenhorst's Kryptogaynen-Flora . . . Al-
though a great majority of the species are marine in occurrence, forming a very
important component of the phytoplankton, a number of freshwater species
have become known. The classification of the complex here adopted is essen-
tially that of Schiller.

146 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

SILICOFLAGELLATOPHYCIDAE

This subclass includes only six clearly defined genera of marine flagellates.
The first representatives of the group to be described were fossil forms that
were found by Ehrenberg in 1839 in cretaceous marls from Oran and Sicily.
He erected the genus Dictyocha for these fossils and two years later (Ehren-
berg, 1841) observed the first living specimens of this genus in water from the
North Sea. In subsequent years he described a large number of additional spe-
cies as well as a second genus {Mesocena) .

Ehrenberg believed these organisms to be diatoms. Haeckel (1862) placed
them with doubt with the Radiolaria. The group retained its doubtful alliance
with the Radiolaria until 1891 when Borgert showed, as a result of a detailed
study of living specimens of DistepJianus speculum, that they differed strikingly
from Radiolaria. He observed the occurrence of brown plastids in the cells and
also established for the first time that the cells owed their motility to the pres-
ence of one (Distephanus) or two {Ehria) flagella. Borgert consequently con-
sidered these organisms as an autonomous group of flagellates for which he
(1891, p. 661) proposed the name Silicoflagellata.

On the basis of Borgert's findings, Haeckel in 189-4 (p. 126) classified these
organisms with the algae. Engler (1903) considered them (with a query) as
constituting an independent phylum of thallophytes.

Lemmermann (1901a, 1901b) gave the first systematic treatment of the group
and the present system is still essentially that proposed by him. Largely on
the basis of skeletal structure he divided the group into two orders: (1) the
Siphonotestales, which are uniflagellate and in which the skeleton is composed
of hollow siliceous beams, and (2) the Stereotestales, which are biflagellate and
in which the siliceous framework of the skeleton is solid. Each of these orders
received a single family. Although the,y appeared to constitute a clearly de-
marked group, Lemmermann (1901b, p. 254) thought the silicoflagellates might
be related to certain of the other groups of flagellates.

Pascher (1912a, p. 193) brought attention to the correspondence between
the skeletons of silicoflagellates and the cysts of Chrysophyceae and hence allied
these groups. With the notable exception of Schulz (1928) and Gemeinhardt
(1930, 1931), who believe that the silicoflagellates constitute an autonomous
class, the majority of students of the group concur with Pascher in relating
them to the Chrysophyceae. Hovasse (1932) is of the opinion that the Ebria-
ceae (which are heterotrophic) may be more nearly related to the Radiolaria
or certain Dinophyceae than to the Silicoflagellatophycidae.

The most comprehensive treatment of the group is that given by Gemein-
hardt (1930) in Rabenhorst's Kryptogamen-Flova . . . Almost half the known spe-
cies of the world are known only from fossils. In 1931 Gemeinhardt published
a valuable account of the silicoflagellates collected during the German South
Polar Expedition of 1901-1903.

The systematic arrangement of the Chrysophyceae presented below departs
in certain respects from that of Pascher (1931). The present arrangement is
a synthesis of the systems of Pascher (1931), Fritsch (1935), Huber-Pestalozzi
(1941), and Smith (1950). It should be emphasized, however, that our knowl-

PAPENFUSS: CLASSIFICATION OF THE ALGAE 147

edge of the Chrysophyceae is still extremely fragmentary and any systematic
arrangement adopted at tliis time is unavoidably artificial. Thus, for instance,
the orders Chrysomonadales, Isochrysidales, Ochromonadales, and Prymne-
siales- are based largely on the possession by the component forms of one, two
equal, two unequal, or one short and two long flagella, respectively, whereas dif-
ferences in flagellation are not considered a valid criterion for the segregation
into separate orders in the Chrysocapsales (in which the motile stages of some
genera possess one and of others two flagella of equal or unequal length), Chry-
sosphaerales (some contain one and some two flagella of imequal length), and
Chrysotrichales (some with one and some with two flagella of unequal length).

Class Chrysophyceae (Pascher) Fritsch, in West (1927, p. 22)
Syn.: Chrysophyceae Pascher (1914, p. 143, as "Reihe")
Subclass CIIRYSOPHYCIDAE Papenfuss, nom. nov.
Syn.: Chrysomonadineae Senn (1900, pp. iv, 152)
Order CHRYSOMONADALES Engler (1898, p. 8)

Family Chrysomonadaceae Stein orth. mut. De Toni (1895, p. 598)

Syn.: Chromulinaceae Engler (1897, p. 570); Chrysapsidaceae Pascher
(1910, p. 11); Euchromulinaceae Pascher (1910, p. 15); Chromo-
phytonaceae Hansgirg orth. mut. De Toni (1895, p. 599)
Family Oicomonadaceae Senn (1900, p. 118)

Cf. Scherffel (1911, p. 329) ; Pascher (1912a, p. 190)
Family Mallomonadaceae Pascher (1910, p. 31)
Family Pedinellaceae Pascher (1910, p. 8)
Syn.: Cyrtophoraceae Pascher (1911a, p. 122)

Order ISOCHRYSIDALES Pascher (1910, p. 36)

Syn.: Hymenomonadales Fritsch, in West (1927, p. 315)
Suborder Isochrysidineae G. M. Smith (1933, pp. 170, 174)
Family Isochrysidaceae Pascher (1910, p. 36)

Syn.: Syncryptaceae G. M. Smith (1933, p. 174)
Family Synuraceae G. M. Smith (1933, p. 175)
Suborder CoccoUthophorineae Papenfuss, nom. nov.'

Syn.: Coccolithineae (Lohmann) Kamptner (1928, p. 23); Family Cocco-
lithophoridae Lohmann (1902, p. 127); Class Coccosphaerales Lemmer-
mann (1908, p. 24); Class Coccolithophorales Lemmermann (1908, p.
33); Order Coccosphaerales Haeckel (1894, p. 110)

Family Syracosphaeraceae (Lohmann) Lemmermann (1908, p. 35)

Syn.: Pontosphaeraceae Lemmermann (1908, p. 33)
Family Halopappaceae Kamptner (1928, p. 24)
Family Deutschlandiaceae Kamptner (1928, p. 27)
Family Hymenomonadaceae Senn (1900, p. 159)

Syn.: Euhymenomonadaceae Pascher (1910, p. 41); Thoracosphaera-
ceae (Kamptner) Schiller (1930, p. 156)
Family Coccolithophoraceae (Lohmann) Lemmermann (1908, p. 38)
Syn.: Coccosphaeraceae G. Murray et Blackman (1898, p. 439) ; Rhabdo-
sphaeraceae Lemmermann (1908, p. 39); Coccolithaceae Kamptner
(1928, p. 25)
Order OCHROMONADALES Pascher (1910, p. 47)

Family Monadaceae Stein orth. mut. Engler (1898, p. 7)

Syn.: Ochromonadaceae Senn (1900, p. 163); Dendromonadaceae Stein

2. Pascher (1929a, p. 271, footnote) is inclined to think that a second short flagellum
may have been overlooked in Prymnesium (cf., however. Carter, 1937, pp. 40-43). The
only other genera in the order, Platychrysis and Chrysochromulina, contain one short and
two long flagella, according to Carter (1937) and Lackey (1939), respectively.

3. The classification of this suborder is based on the systems of Kamptner (1928)
and Schiller (1930).

148 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

orth. mut. Engler (1898, p. 7) ; Euochromonadaceae Pascher (1910, p.
47); Physomonadaceae G. M. Smith (1933, p. 182, cf. Korshikov, 1929,
pp. 253-261)
Family Dinobryaceae Ehrenberg orth. mut. Engler (1897, p. 570)
Syn.: Lepochromonadaceae (Pascher) Fritsch (1935, p. 555)

Order PRYMNESIALES (Fritsch) Papenfuss, stat. nov.
Syn.: Series Prymnesieae Fritsch (1935, p. 512)

Family Prymnesiaceae Conrad (1926, pp. 219-221, as Prymnesiac^es)

Syn.: Chrysochromulinidae Lackey (1939, p. 138)
Family Platychrysidaceae Carter (1937, p. 47)

Order RHIZOCHRYSIDALES* Pascher (1925b, pp. 497, 561)
Syn.: Myxochrysidales Pascher (1931, p. 323)

Family Rhizochrysidaceae (Pascher) Doflein orth. mut. G. M. Smith
(1933, p. 183)

Syn.: Chrysarachniaceae Pascher (1931, p. 323)
Family Chrysothecaceae Pascher (1931, p. 323; cf. Huber-Pestalozzi,
1941, p. 241)

Family Stylococcaceae Huber-Pestalozzl (1941, p. 242)
Family Lagynionaceae Fritsch ex Huber-Pestalozzi (1941, p. 242)
Family Myxochrysidaceae Pascher ex Huber-Pestalozzi (1941, p. 242)
Order CHRYSOCAPSALES Pascher (1912a, p. 175)
Syn.: Hydrurales Pascher (1931, p. 323)

Family Chrysocapsaceae Pascher (1912a, p. 175)
Family Naegeliellaceae Pascher (1925b, pp. 559, 561)

Family Hydruraceae (Rostafinski) Hansgirg orth. mut. De Toni (1895,
p. 596)

Family Celloniellaceae Pascher (1931, p. 323)
Family Ruttneraceae Geitler (1943, p. 108)
Order CHRYSOSPHAERALES Pascher (1914, p. 143)

Syn.: Silicococcales Schiller (1925b, p. 67); Pterospermales Schiller (1925b, p.
72) nomen nudum; Ochrosphaerales Schwarz (1932, p. 459)
Family Chrysosphaeraceae Pascher (1914, p. 159)

Syn.: Aurosphaeraceae Schiller (1925b, p. 67)
Family Chrysostomataceae Chodat (1921, p. 83, as Chrysostomatac^es)
This provisional family is probably based, according to Pascher (1925b,
pp. 546-548) and Scherffel (1927, pp. 355-356), on the cysts of members
of the Chrysomonadales.
Family Pterospermaceae Lohmann (1904, p. 39, as Pterospermaceen)
See the remarks of Schiller (1925b, p. 72) and Fritsch (1935, p. 550)
regarding the status of this group.
Family Chrysopediaceae Pascher (1931, p. 323)
Family Stichogloeaceae Wille ex Huber-Pestalozzi (1941, p. 263)
Order CHRYSOTRICHALES Pascher (1914, p. 143)

Syn.: Cryptotrichales Pascher (1914, p. 150); Chrysothallales Huber-Pestalozzi
(1941, p. 14)

Family Chrysotrichaceae Pascher (1914, p. 143)

Syn.: Nematochrysidaceae Pascher (1925b, p. 498)
Family Phaeothamniaceae (Lagerheim) Hansgirg orth. mut. De Toni
(1888, p. 448)

Family Thallochrysidaceae Conrad, in Pascher (1914, p. 143)
Syn.: Chrysothallaceae Huber-Pestalozzi (1941, p. 14)
Subclass siLicoFLAGELLATOPHYCiDAE (Borgert) Papenfuss, stat. nov.
Syn.: Order Silicoflagellatae Borgert (1891, p. 661)

4. As various authors (Pascher, 1913a; G. M. Smith, 1920; Doflein, 1928, p. 461;
Huber-Pestalozzi, 1941, p. 241) have remarked, this is an artificial order since the major-
ity, if not all, the forms placed here may have been derived from or represent the non-
flagellated stages of various flagellated members of the class.

PAPENFUSS: CLASSIFICATION OF THE ALGAE 149

Order SIPHONOTESTALES Lemmermann (1901a, p. 92)

Family Dictyochaceae Lemmermann (1901a, p. 92)

According to Gemeinhardt (1930, pp. 22, 77), Scluilz established a fam-
ily Cornuaceae for the monotypic genus Cormia Schulz (1928, p. 285),
but I can find no mention of such a family in Schulz's writings.
Order STEREOTESTALES Lemmermann (1901a, p. 93)
Family Ebriaceae Lemmermann (1901a, p. 93)

APOCHROMATIC GROUPS OF UNCERTAIN SYSTEMATIC POSITION

Klebs (1892, pp. 282-283) and Semi (1900, p. 152) even in their time al-
ready suspected a relationship between certain colorless flagellates belonging
to the family ]\Ionadaceae and certain pigmented chrysomonads of the family
Ochromonadaceae. Subsequent work by a number of investigators (Scherffel,
1911, 1924; Pascher, 1916a, 1917, 1930b; Korshikoff, 1929, among others) have
amply substantiated the suspicions of Klebs. It is generally agreed today that
many of the colorless species are derived from pigmented species or are perhaps
only colorless forms of pigmented species. These forms not only agree with their
pigmented counterparts in the general morphology of the cell, type of flagel-
lation, and kind of food reserve but they also produce cysts of the same kind.
Consequently the families Oicomonadaceae and IMonadaceae have in the pre-
ceding treatment of the Chrysophyceae been accorded positions in the Chryso-
monadales and Ochromonadales, respectively.

Klebs (1892) recognized two groups of colorless flagellates, the Protomas-
tigina and the Polymastigina. Senn (1900) distributed these colorless forms
among the three groups Pantostomatineae, Protostomatineae, and Distomati-
neae. As mentioned above, some of these organisms (e.g., members of the Mona-
daceae and Oicomonadaceae) have been shown to be colorless Chrysophyceae.
The systematic position of the majority of the forms, however, is still uncertain.
Since at least some of them possess features that suggest an affinity with the
Chrysophyceae, the three groups recognized by Senn and many subsequent
authors are here appended to the Chrysophyceae.

The history of these groups is briefly considered below.
Pantostomatineae: This group was established by Kent (1880-1881, pp. 211,
229, as Flagellata-Pantostomata) to embrace a heterogeneous assemblage of
flagellated organisms that engulf food by pseudopodia. Its present circiimscrip-
tion is that given by Senn (1900, pp. 110, 111). He assigned to it a number of
genera, belonging to the two families Holomastigaceae and Rhizomastigaceae,
which share certain features, especially the absence of a differentiated oral ap-
paratus, solid food being engulfed by pseudopodia that form at any point on the
cell surface. Since the time of Senn, treatments of the group have been given by
Lemmermann (1907-1910, 1914), Doflein (1928), Fritsch (1935) and Huber-
Pestalozzi (1941). The complex comprises only the two families assigned to
it by Senn.

Family Holomastigaceae (Lauterborn) Senn (1900, p. 112)
Family Rhizomastigaceae Biitschli orth. mut. Senn (1900, p. 113)

Protomastigineae: This group was established by Klebs (1892, p. 293) to
include a number of families characterized by the fact that food is taken in at

150 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES

a specific place on the cell. The present circumscription of the assemblage is
essentially that given by Senn (1900, pp. 117-118), who removed the Rhizo-
mastigaceae (placed here by Klebs) to the Pantostomatineae and added certain
families, among others the Tetramitaceae, which Klebs had placed in his group
Polymastigina. Since the time of Senn, Lemmermann (1914, pp. 52-121) and
Huber-Pestalozzi (1941, pp. 280-301), among others, have given treatments of
the group. It is comprised of the following families.

Family Trypanosomaceae Doflein orth. mut. Lemmermann (1914, p. 64)

Family Bicoecaceae Stein orth. mut. Senn (1900, p. 121)

Family Craspedomonadaceae Stein orth. mut. Senn (1900, p. 123)

Family Phalansteriaceae Senn (1900, p. 129)

Family Bodonaceae Biitschli orth. mut. Engler (1898, p. 7)

Family Cryptobiaceae Lemmermann (1914, p. 107)

Family Amphimonadaceae Kent orth. mut. Engler (1898, p. 7)

Syn.: Spongomonadaceae Stein orth. mut. Engler (1898, p. 7)
Family Trimastigaceae Kent orth. mut. Senn (1900, p. 141)
Family Tetramitaceae Kent orth. mut. Engler (1898, p. 7; see Skuja,
1948, p. 68)
?Family Paramastigaceae Skuja (1948, p. 68)

Distoniatineae : This group was first established by Klebs (1892, p. 329) as a
subgroup Distomata of his group Polymastigina. The present circumscription